Silas Little and Peter W. Garrett
The species name of pitch pine (Pinus rigida) means rigid or stiff and refers to both the cone scales (17) and the wide-spreading, sharply pointed needles (5). It is a medium-sized tree with moderately strong, coarse-grained, resinous wood that is used primarily for rough construction and where decay resistance is important. One tree in Maine measured 109 cm (43 in) in d.b.h., 29 m (96 ft) tall, with a crown spread of 15 m (50 ft) (11).
In the Northeast, pitch pine is most common on the sandy soils of Cape Cod, Long Island, and southeastern New Jersey, and in some sections of sandy or shallow soils in Pennsylvania (19).
- The native range of pitch pine.
Aboveground description: Pitch pine is a hard pine with highly variable growth forms [44,86]. It grows as a prostrate shrub on Fire Island, New York . On the Mt Everett ridgetop in the Taconic Mountains of Massachusetts, pitch pine occurs as a low-growing, 1-foot (0.3 m) mat and as a single-stemmed tree, 10 feet (3 m) tall. The largest prostrate mats measure over 10 mÂ² . Dwarf-stature pitch pines ranging from 20 inches (50 cm) to 13 feet (4 m) tall occur in the in plains area of the New Jersey Pine Barrens. In an area 22 miles (35 km) from the Pine Barrens, pitch pine trees reach 100 feet (30 m) tall . Studies revealed that dwarf and tall-tree pitch pine populations in the Pine Barrens were almost genetically identical .
As a tree, pitch pine is medium sized and rarely grows beyond 82 feet (25 m) tall and 3 feet (1 m) DBH. Branching is often irregular [13,23,39,53]; branches can be twisted or gnarled , and self-pruning is poor . Crown and branch forms can be affected by growing conditions. Suppressed trees, fire-damaged trees, or trees released through heavy logging usually have drooping, slender branches along the lower trunk. Dead branches are persistent and contain more resin than live branches . Trunks are straight to somewhat curved  and covered with thick, large, rough, irregular plates of bark [13,44]. Often there are stubby branches or needle bundles on the trunk [39,70]. In Great Smoky Mountains National Park, bark thickness increased with increasing DBH until trees reached 9 inches (25 cm) in DBH, when bark thickness decreased with increased DBH . Pitch pine is not considered long lived. On the Mt Everett ridgetop, pitch pine averaged 78 years old, but age ranged from 12 to 170 years . For more on pitch pine growth and longevity, see Growth.
Pitch pine needles are most often in bundles of 3. Needles measure up to 5.9 inches (15 cm) long and are stiff and straight to twisted. Pitch pine retains needles for 2 to 3 years [23,44,45,112]. Mature pitch pine cones measure 1 to 3.5 inches (3-9 cm) long and wide and often occur in clusters. Cone scales are thick with stout spines [23,38,112,159]. Male cones are produced at the base of the current year's growth and are often more abundant on low branches. Female cones are more common on upper branches and mature in the fall of the second year following pollination. Pitch pine produces serotinous and nonserotinous cones. Nonserotinous cones shed seed soon after they mature but persist a long time [44,45,53,86,135]. For more on factors that affect serotinous cone production, see Serotinous and nonserotinous cone production. Both cone and seed size increase from northeastern to southwestern populations, as does the number of viable seeds produced per cone . Pitch pine produces smooth, winged seeds. Seeds are typically 4 to 5 mm long, and wings measure 15 to 20 mm [23,44,45].
Belowground description: While Hosie  suggests that pitch pine produces short lateral and taproots, others indicate that the pitch pine root system may reach 10 feet (3 m) deep , and that lateral root extension may exceed 6 feet (2 m) in 6 years of growth . It is likely that site conditions affect root growth and structure.
In the Lebanon State Forest in New Jersey, roots of 1- to 30-year-old pitch pine trees were excavated and studied. Trees grew in partial shade, with "moderate competition", and on well-drained Lakewood sand soils. First-year seedlings had taproots that ranged from 3 inches (8 cm) to 1 foot (0.3 m) deep. Elongation of the taproot was best for seedlings growing in clean, loose sand in full sun. Elongation decreased with shading and increased humus. Mycorrhizae were found on the roots of 2-month-old seedlings. Trees beyond the sapling stage had definite and "fairly strong" taproots that typically divided at 2- to 3-foot (0.6-0.9 m) depths. Taproot prominence decreased as trees aged. The researcher described the taproot of an 85-year-old pitch pine as weak, but noted that some long roots reached 8 feet (2 m) deep. Three trees had roots extending and growing into the water table. There was also some root grafting observed. In only one case was grafting with another species, shortleaf pine. The Table below summarizes some root system characteristics for young pitch pine trees .
|Root system characteristics of pitch pine trees from 1 to 30 years old excavated from the Lebanon State Forest, NJ|
|Tree age (years)||Tree size||Maximum taproot depth||Maximum length of primary lateral root||Other notes|
|1||3 inches||8 inches||3 inches|
|4||11 inches||18 inches||15 inches|
|8||22 inches||27 inches||33 inches||10 or more primary laterals|
|12||4 feet||4 feet||8 feet||3 other laterals measured 4 feet; primary laterals to 3.5 feet deep|
|17||14 feet||5 feet||19 feet|
|30||22.5 feet; |
3.5 inch DBH
|9 feet||31 feet||5-inch-diameter taproot at 6 inches below soil surface, 3-inch diameter at 2 feet; 20 large laterals, 13 at less than 6 inches and 7 at up to 2 feet|
Habitat and Ecology
Climate: A humid climate prevails throughout pitch pine's range. Annual precipitation averages between 37 to 56 inches (940-1,420 mm) and is well distributed throughout the year. The frost-free season typically lasts 112 to 190 days. Low temperatures can reach -40 Â°F (-40 Â°C) in the northern part of pitch pine's range, and highs of 100 Â°F (40 Â°C) occur in the southern range . Pitch pine is considered tolerant of cold, drought, and salt spray, and while persistent in a variety of climates, humid climates with well-distributed rainfall are preferred . Some report that pitch pine is susceptible to sea spray damage and may be restricted from extreme coastal locations .
Seventy-five years of records show that pitch pine habitats in the southern Piedmont average 47 inches (1,200 mm) of evenly distributed precipitation/year and a frost-free period of 196 days . New Jersey Pine Barrens average 45 to 50 inches (1,100-1,300 mm) of evenly distributed precipitation/year. Growing seasons are long, winters are mild, and summers are hot in the Pine Barrens .
At the Pitch Pine Ecological Preserve of Haut-Saint-Lauret, near pitch pine's northernmost extent, January temperatures average 14 Â°F (-10 Â°C), and July temperatures average 69.4 Â°F (20.8 Â°C). Average annual precipitation is 38 inches (975 mm)/year. Precipitation is delivered throughout the year, with slight increases in the summer. The number of frost-free days averages 159. Pitch pine occupies a wide range of edaphic conditions on the Preserve, and after an in-depth study of the area, researchers concluded that it is not climate but a lack of suitable habitat that restricts pitch pine's distribution in the area .
Cold tolerance: Pitch pine needles are tolerant of very cold temperatures, and cold tolerance increases as climates become cooler. One-year-old pitch pine seedlings grown from seed collected in the northern half of pitch pine's range had an average stem and needle tissue cold tolerance of 21.7 Â°F (-5.7 Â°C) in October and -25.1 Â°F (-31.7 Â°C) in January. Stems were less cold-tolerant than secondary needles in midwinter . Needles collected from trees in Acadia National Park were tolerant to at least -85 Â°F (-65 Â°C). Freezing tests on entire 1-season-old pitch pine seedlings revealed no needle damage after exposure to -48 Â°F (-55 Â°C) .
Drought tolerance: Drought conditions are tolerated by pitch pine, but insect outbreaks coupled with very dry conditions may produce mortality. Researchers observed no pitch pine mortality in an oak-pine forest during drought conditions from 1984 to 1991 in the Coweeta Basin watershed. The drought was severe both in terms of duration and accumulated precipitation deficit, which was 24% to 31% below normal precipitation from 1985 to 1988. Measurements in 1985 and in 1991 revealed no change in pitch pine density, and basal area increased from 1985 to 1991 . Smith  observed pitch pine mortality in the Coweeta Basin during this time in a southern pine beetle outbreak area.
Flood tolerance: Experimental studies found that young pitch pine seedlings are better able to survive root flooding than older seedlings and saplings and that "flood hardening" can occur. Researchers compared the growth and survival of 3-month-old seedlings, 15-month-old seedlings, and 5-year-old saplings in flooded conditions. Seedlings exposed to flooding in their first growing season were more flood tolerant than those not exposed to flooding. Mortality was 3 times more likely in nonexposed than flood-exposed seedlings. Nonexposed seedlings experienced at least 50% mortality after 10 weeks of root flooding, while flood-exposed seedlings experienced 50% mortality after 16 weeks of root flooding. Flooded seedlings developed expanded lenticels along the stems and produced roots near and above the soil surface .
Elevation: Pitch pine occupies habitats from sea level to over 5,600 feet (1,700 m) throughout its range. Generally, higher-elevation habitats are occupied in the southern than northern part of pitch pine's range [39,45,97]. In the Adirondack Uplands, pitch pine occurs between 100 and 1,040 feet (30-317 m) .
Soils: Soils in pitch pine habitats are often dry, thin, infertile, and sandy or gravelly in texture ; however, soils from rapidly draining to swampy are tolerated . Sopodosol, Alfisol, Entisol, and Utisol soil orders are common in pitch pine habitats . Pitch pine occupies limestone and sandstone soils in the Adirondack Uplands  and "poor" soils along the Atlantic Coast from Delaware to Maine . In the Harvard Forest of north-central Massachusetts, cluster analyses orient pitch pine at the "nutrient-impoverished" end of the fertility gradient . In the Black and Craggy mountains of North Carolina, pitch pine occurs in xeric pine forests on southern slopes with low-nutrient soils having pH levels of 3.4 to 4.5 . In xeric rock outcrops in the Shawangunk Mountains, pitch pine trees occur on soils that range from 3 to 14 inches (8-35 cm) deep .
Soils in New Jersey's pine barrens and plains may contain high levels of aluminum (>500 ppm possible). These levels do not, however, affect pitch pine seedling growth or the relative distributions of dwarf and tree-size pitch pine . Soils in the pine barrens are acidic with leached A2 horizons, and pitch pine occupies sites with excessively drained, imperfectly drained, very poorly drained, and mucky swamp soils . In the Pitch Pine Ecological Preserve of Haut-Saint-Lauret, pitch pine occurs on thin (≤8 inches (20 cm) of organic or mineral deposits), dry, acidic (pH <4) soils overlaying bedrock  but is also the most abundant species in bogs .
Burned soils: For information on the effect of fire on soils within the oak-pitch pine forests in the New Jersey Pine Barrens, see Burns . Changes in soil nutrients were measured periodically for up to 5 years after felling and burning in mixed oak-pine (pitch pine dominant) in the Nantahala National Forest in western North Carolina. For results, see Knoepp and others .
Key Plant Community Associations
Pitch pine is often dominant in pine barrens vegetation in the northeastern
United States. On extremely harsh sites it may represent a climax vegetation
type , but most often pitch pine forests are early seral and are replaced by
oaks (Quercus spp.) and other hardwoods in the absence of fire
[25,96,171]. In the Appalachian region, pitch pine is common on ridges with Virginia pine (Pinus virginiana)
and Table Mountain pine (P. pungens) [22,106]. Pitch pine is a
dominant species in the following vegetation types and communities.
General/eastern United States:
pitch pine forests, cover type 45 recognized by the Society of American
northern pine barrens on the Atlantic Coast Plain north of Delaware Bay 
pitch pine and pitch pine-oak communities in central New Jersey, central Long
Island, near Albany in upstate New York, and on Cape Cod, Massachusetts 
oak-pine forests dominated by scarlet oak (Q. coccinea), chestnut oak
(Q. prinus), pitch pine, Virginia pine, and Table Mountain pine in the Blue
Ridge Province from Pennsylvania to northern Georgia 
pitch pine-oak forests throughout New England, but best
represented on Cape Cod and Martha's Vineyard, Massachusetts 
mixed deciduous forests of pitch pine, red maple (Acer rubrum), gray
birch (Betula populifolia), and quaking aspen (P. tremuloides)
open-canopy pitch pine forests
pitch pine/heath communities
pitch pine/bear oak (Q. ilicifolia) communities in Waterboro Barrens 
pitch pine vegetation associations on Mt Desert Island off the southern coast;
may contain red pine (Pinus resinosa) and/or eastern white pine (P.
strobus) in the canopy; understory dominants are common juniper (Juniperus communis)
in open stands and black huckleberry (Gaylussacia baccata) in closed
pitch pine/kinnikinnick (Arctostaphylos uva-ursi)
pitch pine/wavy hairgrass (Deschampsia flexuosa)
pitch pine-black oak/yellow sedge (Q. velutina/Carex pennsylvanica)
pitch pine/bear oak/broom crowberry (Corema conradii)
pitch pine-white oak (Q. alba)-black oak/black huckleberry
pitch pine/bear oak-northern bayberry (Myrica pennsylvanica) on Cape Cod National Seashore 
pine plains communities or plains vegetation in the Pine Barrens are described by several
authors; plains communities are dominated by pitch pine
trees <10 feet (3 m) tall and by shrubby bear and blackjack oak (Q. marilandica) [55,101,104]
barrens communities support pitch pine trees >25 feet (7.6 m) tall 
pitch pine-post oak (Q. stellata) forests 
pitch pine-black oak forests in the Pine Barrens 
pitch pine-bear oak communities
pitch pine-swamp doghobble (Eubotrys racemosa) in the Pine Barrens 
pitch pine lowlands forest; canopy dominated by 15- to 20-foot (4.6-6.1 m)-tall
pitch pine-oak (black oak, blackjack oak, post oak)
oak-pitch pine (black oak, chestnut oak, white oak and/or scarlet oak) in the
Pine Barrens 
mesic pitch pine-bear oak forest types
dry pitch pine lowland forest types
wet pitch pine lowland forest types
pitch pine-red maple swamps in the Pine Barrens 
pitch pine-bear oak communities in High Point State Park 
pine barrens; nearly pure pitch pine with scattered white and
scarlet oak and a dense bear oak shrub layer
pine plains; tall shrub communities dominated by dwarf pitch pine and bear oak on Long Island 
closed-canopy pitch pine forests; few shrubs and a well-developed herbaceous
open-canopy pitch pine forests; stunted trees and an extensive bear oak or
ericaceous (Ericaceae) shrub understory
dwarf-pitch pine communities; dense 7- to 10-foot (2-3 m)-tall pitch pine in canopy, black huckleberry and
Blue Ridge blueberry (Vaccinium pallidum) dominate the understory in the Hudson Valley 
scarlet oak-pitch pine forests on low southern slopes
pitch pine-scarlet oak forests on
upper southern slopes of the Thompson Gorge in the southeastern Blue Ridge
xeric pine forests; dominated by pitch and Table Mountain pine in Black and Craggy
pine-oak/heath (Ericaceae) vegetation types in the Wine Spring Creek watershed of the Nantahala
National Forest; scarlet oak, chestnut oak, and pitch pine dominate
the canopy and mountain-laurel (Kalmia latifolia) the understory 
Carolina hemlock (Tsuga caroliniana) bluff communities scattered throughout
the Blue Ridge Mountains and in the upper Piedmont
low elevation rocky summit communities, rare in the Piedmont and the Blue
ultramafic outcrop barren communities, scattered in the Piedmont and the Blue Ridge
yellow pine (Virginia pine and pitch pine, with eastern white pine) forest types
scarlet oak-yellow pine (Table Mountain and pitch pine) forest types in
western Great Smoky Mountains National Park 
shortleaf pine-pitch pine forests
chestnut oak-pine (pitch and Virginia pine) forests
pine forests dominated by pitch and shortleaf pine with an open heath layer on Pine
yellow pine (Virginia pine and pitch pine, with eastern white pine) forest types
scarlet oak-yellow pine (Table Mountain and pitch pine) forest types in western Great Smoky Mountains
National Park 
Soils and Topography
Generally, pitch pine grows on Spodosols, Alfisols, Entisols, and Utisols. In southern New Jersey, the pH of the A and B horizons range from 3.5 to 5. 1, and in northern New Jersey, from 4 to 4.5 (9).
Pitch pine grows on sites with a wide range of moisture conditions. In southern New Jersey it is found on excessively drained, imperfectly drained, and poorly drained sands and gravels, as well as on muck soils in the white-cedar swamps. Even in the hilly regions it grows on both well drained and excessively drained slopes and in the swamps (9).
In New England it is most common in the coastal districts and in river valleys. In New York it is not common above 610 m (2,000 ft), but in Pennsylvania it grows at all elevations up to the highest point in the State, 979 m (3,213 ft) (13). In the Great Smoky Mountains and vicinity, pitch pine is found at elevations between 430 and 1370 m (1,400 and 4,500 ft). In hilly sections, pitch pine often occupies the warmer and drier sites, those facing south or west (9).
Habitat & Distribution
Habitat & Distribution
aecium of Coleosporium asterum parasitises live Pinus rigida
Associated Forest Cover
According to the Forest Survey, pitch pine types cover 9 900 ha (24,500 acres) in New Hampshire, 85 400 ha (211,000 acres) in Massachusetts, 1200 ha (3,000 acres) in Rhode Island, possibly 44 500 ha (110,000 acres) in New York, more than 121 400 ha (300,000 acres) in Pennsylvania, and more than 283 300 ha (700,000 acres) in New Jersey. Other Northeastern States contain fewer hectares of this species, though about 187 400 ha (463,000 acres) in Maryland and 346 000 ha (855,000 acres) in West Virginia were classified in the Forest Survey as having pitch pine-Virginia pine-hardwood stands. However, in these two States, the Virginia pine (Pinus virginiana) component greatly exceeds pitch pine in most stands (9).
Usually, the most common shrubs growing with pitch pine on upland sites are lowbush blueberries (often Vaccinium vacillans or V. angustifolium) and black huckleberry and dangleberry (Gaylussacia baccata and G. frondosa). Some stands include bear oak (Quercus ilicifolia), dwarf chinkapin oak (Q. prinoides), and mountain-laurel (Kalmia latifolia).
Lowland sites where pitch pine predominates have a variety of shrubs. Common ones include sheeplaurel (Kalmia angustifolia), leatherleaf (Chamaedaphne calyculata), staggerbush (Lyonia mariana), inkberry (Rex glabra), dangleberry, highbush blueberry (Vaccinium corymbosum), and swamp-honeysuckle (Rhododendron viscosum) (9).
Diseases and Parasites
Several fungi that attack pitch pine but usually do not cause serious or extensive damage are stem rusts such as Cronartium comptoniae, C. quercuum, C. quercuum f. sp. fusiforme, and C. comandrae (1); several needle rusts and blights such as nine species of Coleosporium, Ploioderma lethale, and P. hedgcockii; twig cankers such as Diplodia pinea; root rots such as Heterobasidion annosum; and trunk rots, chiefly Phellinus pini. Heart rot as a result of P. pini does not become important in stands until the trees are 75 years old (3,12).
Many insects attack pitch pine (6). The most important are the tip moths (Rhyacionia frustrana and R. rigidana), the pitch pine looper (Lambdina athasaria pellucidaria), the sawflies (chiefly Neodiprion lecontei, N. pratti paradoxicus, and N. pinusrigidae), the southern pine beetle (Dendroctonus frontalis), the pine webworm (Tetralopha robustella), and the pine needleminer (Exoteleia pinifoliella). Loopers periodically cause extensive damage to pitch pine in Massachusetts; in 1954 they defoliated pines on more than 20 230 ha (50,000 acres) of Cape Cod (9).
Fire adaptations: Pitch pine has numerous fire adaptations that allow it to establish and/or regenerate on burned sites. Postfire pitch pine regeneration may be through bole and crown sprouting from epicormic buds protected by thick bark, sprouting from basal buds protected by crooks and/or soil, and/or through seedling establishment from seed in nonserotinous cones or opened serotinous cones . In a 1930s survey completed by 41 leading foresters of the time, asked to rate a list of tree species in order of increasing fire resistance, pitch pine was considered most resistant. Foresters were asked to consider trees between 40 and 80 years old .
FIRE REGIMES: Pitch pine forests have been aptly described as "fire-dependent ecosystems" by Vogl . Pitch pine forests support vegetation with various fire-adapted regeneration strategies and persist in and foster environments conducive to ignition, combustion, and fire spread. Vogl further substantiates his claim by noting that exclusion of fire or decreased fire frequency in pitch pine forests produces more "dramatic" effects than increased fire frequency .
Early anthropogenic fires: While much of the information regarding the burning done by Native Americans is speculative, there is ample evidence of widespread, frequent burning after eastern United States settlement by Europeans. Often the lack of lightning in the Northeast is considered evidence of Native American fire starts; however, while lightning is rare compared to the western United States, it does occur (see Fire season, weather, and fuels). Early written accounts, a lack of lightning, and the existence of many fire-tolerant or fire-adapted species and landscapes suggests that Native Americans burned at least portions of Virginia. Fire was likely used to maintain travel routes and to find and gather food . Fossil pollen and charcoal records from the Horse Cove bog in Macon County, North Carolina, indicate that fires have been important for the last 3,900 years. Because of the lack of lightning in the area, researchers suspect selective burning by Native Americans contributed much to the charcoal record . There is considerable evidence that early European settlers burned in the New Jersey Pine Barrens and some speculation that Native Americans did too. Lightning is rare in the Pine Barrens, especially compared to the occurrence of lightning in western US forests. Early European settlers burned clearings around cranberry (Viburnum spp.) bogs and set fires to cover up illegal timber harvests. Locomotives were another ignition source. Generally, fires in the Pine Barrens were not controlled since the pines were not valued for timber, and often flames of fires in nearby timber-valued forests were directed to the Pine Barren plains during fire fighting . For more on the use of fire by Native Americans in the Northeast, see Day .
Fire season, weather, and fuels: Climate, fuels, and terrain in pitch pine habitats are favorable to fire. Often pitch pine trees have drooping, slender branches along the lower bole, and persistent dead branches contain more resin than live branches . In the New Jersey Pine Barrens, a long growing season, high maximum temperatures, strong winds, and level to rolling terrain encourage fire ignition and spread . In dwarf pitch pine-dominated plains, vegetation regeneration provides enough fuel to carry another fire 4 to 5 years after a crown fire .
Fire records and weather data from Maine indicate that severe fire years occur about every 15 years and are associated with short periods (1-2 months) of "intense" drought [130,131]. In Pennsylvania, lightning ignitions burned large areas in drought years. From 1912 to 1917, lightning struck pitch pine trees 417 times: 139 times in July and 169 times in August. From 1960 to 1997, lightning-caused fires burned 2,279 acres (922 ha), and there were lightning fires in nearly every year . At least a few lightning ignitions occur each year in the New Jersey Pine Barrens. Most lightning comes from June to September. As many as 26 ignitions have been recorded in a year, and ignitions increase with drought and "dry lightning" events . National Forest records from northern Georgia and southern Virginia indicated that there were 6 lightning fires/year/400,000 ha between 1960 and 1971. Ninety percent of lightning fires occurred from April to August, and 40% in May. Human-caused fires were most common from March to May and October to December, when litter was driest. Human-caused fires were often more severe than lightning-caused fires .
Past/present fire-return intervals and fire behavior: Dwarf pitch pine communities burned at 5- to 15-year intervals, and tree-sized pitch pine forests and woodlands burned at 15- to 150-year intervals. Fire-return intervals of about 30 years or more increase the likelihood of associated deciduous tree reproduction, and as fire-return intervals increase, so does the chance of pitch pine replacement by hardwoods.
Fire scars on 4 eastern white pine and 2 pitch pine trees from the east ridge of Catocin Mountain Park in Thurmont, Maryland, revealed that the time between 2 successive fires ranged from 5 to 49 years. The mean fire-return interval from 1813 to 1900 was 29 years. From 1813 to 1985, the mean fire-return interval was 21.5 years. Fire occurrence increased in the early 1900s, but there were no fires in the area after Park establishment in 1936 .
Fire scar data show that western Great Smoky Mountains National Park mixed-pine forests of upper southern slopes burned on average every 12.8 years between 1856 and 1940. From 1920 to 1949, 96% of the mixed-pine forests burned. After creation of the Park in the 1930s and the active exclusion of fire, the average sizes of lighting-caused and human-caused fires were 8.4 acres (3.4 ha) and 13.3 acres (5.4 ha), respectively. With fires of this size, in would take 2,000 years to burn the 22,000-acre (9,100 ha) study area within the western part of the Park . Fire exclusion converted xeric sites occupied by open-canopy forests with rich herbaceous understories, which were common in the early 20th century, to mature, closed-canopy stands with low herbaceous cover and richness by the late 20th century. Researchers predict that these canopy structure changes will affect fire behavior. In the early 20th century, open-canopy stands had limited woody fuels but a highly flammable understory; fire ignition will likely be more difficult, spread more limited, and fire size smaller in closed-canopy than historically open-canopy stands .
Age structure and fire scars from Table Mountain pine-pitch pine stands in the southern Appalachians revealed an all-aged distribution and frequent periodic or continuous regeneration from about 1800 to 1950 in 3 northern Georgia and 2 southern South Carolina stands. Age structure suggested that stand-replacing fires were unlikely, and periodic, low-to moderate-severity surface fires were common in these stands. In another South Carolina stand and 3 stands in Tennessee, age distribution was unimodal, suggesting stand-replacing events had occurred; however, evidence of stand-replacing fire was lacking. Researchers suggested that these stands have been maintained through recurring low- to moderate-severity surface fires and likely a variety of other disturbances such as droughts, hurricanes, insect outbreaks, thunderstorms, and logging. All stands experienced 3 to 8 fires since the 1850s, but pine regeneration has been rare since the 1950s . Frost  suggests that Table Mountain pine-pitch pine forests in the southern Appalachians burned at 5- to 7-year intervals in "understory shrub fires" and less frequently, every 75 years, in "catastrophic" stand-replacing fires before European settlement.
New Jersey Pine Barrens: Researchers have extensively studied fire ecology in pitch pine communities of the New Jersey Pine Barrens and found that fire frequency decreases from the dwarf pitch pine plains to the tree-sized pitch pine barrens. From fire scars and tree dating, Lutz  proposed that fires in plains communities burned on average of every 6 years. Transition communities burned at 12-year intervals, and barrens communities burned at 16-year intervals. Fire severity was greatest on the plains, because of the low stature of the vegetation, greater air movement, and higher evaporation rates, which produced drier fuels. For brief community descriptions, see Habitat Types and Plant Communities. Others reported that fire intervals range from 6 to 8 years in the plains and 16 to 26 years in tall pitch pine forests in the Lebanon State Forest (McCormick and Gill, cited in ).
From fire tolerance rankings created from an in-depth analysis and review of the distribution and fire tolerance of New Jersey's Pine Barrens species, Windisch  estimated the fire-return interval and fire behavior for upland communities and peripheral habitats in the barrens. Reported fire frequencies and behaviors were those that would perpetuate the vegetation type .
|Fire behavior descriptions and fire-return intervals for upland and peripheral New Jersey Pine Barrens communities|
|Plant community||Dominant species||Physiognomy||Mean fire-return interval (years)||Typical fire type||Fire ecology|
|pine plains||dwarf, serotinous pitch pine; shrub oaksÂ¹||open shrubland <1 m tall||5-15||mixed crown and surface||MFRI* allows shrub oaks and pitch pine to regenerate but too frequent for tree-form oaks and pines|
|pine plains||dwarf, serotinous pitch pine; shrub oaks||closed shrubland 1-3 m tall||sporadic 15-60||mixed crown and surface||sporadic MFRI of 15-60 years reduces ground cover diversity, fire "intensity" increases as canopy and heath cover increase|
|pitch pine-shrub oak barrens||tree-form, serotinous and nonserotinous pitch pine; shrub oaks||open-canopy pitch pine; lacks tree oaksÂ²||15-25||more crown than surface||tree-form pitch pine prevails over dwarf form, MFRI too frequent for tree oaks and shortleaf pine to reproduce successfully|
|pitch pine-post oak-shrub oak woodland||tree-form, serotinous and nonserotinous pitch pine; 5-10% post oak; shrub oaks||open-canopy pitch pine; sparse post oak understory||25-30||more crown than surface||20-30 year MFRI allows post oak but not other tree oaks to reproduce|
|pitch pine-tree oak-shrub oak woodland||tree-form, serotinous and nonserotinous pitch pine; tree oaks; shrub oaks||open-canopy pitch pine-oak; 5-25% tree oak cover||30-40||more crown than surface||MFRI allows tree oaks to reproduce but frequent enough to keep canopy open and shrub oak dominance|
|mixed pine-tree oak-shrub oak woodland||tree-form, nonserotinous pitch pine; shortleaf pine; tree oaks; shrub oaks||open-canopy mixed pine-oak ; 5-25% tree oak cover||30-40||more surface than crown||less "intense" burning allows shortleaf pine to codominate|
|pine-oak forest||tree-form, nonserotinous pitch pine; shortleaf pine; tree oaks; <5% shrub oaks||closed-canopy pine-oak; 25-50% tree oak cover||40-60||mixed crown and surface||MFRI allows strong tree oak reproduction and codomination; fire frequent enough for pitch pine successful reproduction|
|oak-pine forest||tree oaks; tree-form, nonserotinous pitch pine; shortleaf pine||closed-canopy oak-pine; >50% tree oak cover||60-100||surface fire predominant; crown fire rare||MFRI allows strong tree oak reproduction and is frequent enough for pine persistence|
|oak-pine-holly (Ilex spp.) forest||tree oaks; tree-form, nonserotinous pitch pine; shortleaf pine; <5% hickory||closed-canopy oak-pine; >50% tree oak cover||100-150||surface fire predominant; crown fire rare||MFRI allows for tree oak and holly dominance, hickory present but limited by fire; pines persist but not dominant|
|Â¹shrub oaks: blackjack and bear oak; Â²tree oaks: post, scarlet, white, and/or chestnut oaks. |
*MFRI: mean fire-return interval.
Increased pitch pine associated with European settlement disturbances: In parts of Maine, New York, Massachusetts, and West Virginia, disturbances and fires associated with land clearing and cultivation by early European settlers increased the abundance and prevalence of pitch pine. Pollen and charcoal records from the Newfield Marsh of southern Maine suggest that disturbances and fire regime changes occurred with European settlement of the area. Over the past 200 years, vegetation dominance has shifted to more "fire-prone, xeric-species". Early seral pitch pine-dominated communities dominating the Waterboro Barrens were likely present in presettlement time but with distribution limited to severely or repeatedly burned sites, xeric southern or western slopes, and areas of exposed bedrock. Increases in pitch pine-dominated communities are thought to be a result of postsettlement disturbances including logging, charcoal manufacture, and blueberry (Vaccinium spp.) cultivation .
Using pollen records, historic maps, and early logging and land clearing records, researchers concluded that the range of pine barrens and dwarf pine plains in central Suffolk County, New York, expanded over the last 3 centuries due to increased human disturbances. After European settlement of the area, fire frequency increased as land was burned for cultivation and grazing. Sparks thrown from wood- and coal-burning locomotives also increased the fire frequency. Mixed oak-pitch pine and pitch pine-mixed oak forests were abundant before substantial European settlement (1640-1680), but pitch pine-oak/heath woodlands, pitch pine-scrub oak (bear and/or dwarf chestnut oaks) barrens, and dwarf pine plains probably covered less than 17,000 acres (7,000 ha) before substantial European settlement. By the late 19th century, pine barrens covered about 250,000 acres (100,000 ha) in Suffolk and eastern Nassau counties. Fire exclusion in the 20th century led to some conversion of pitch pine barrens and woodland vegetation back to pitch pine-mixed oak or mixed oak-pitch pine .
Pollen and charcoal sediment from ponds on Cape Cod revealed that vegetation changes over a 2,000-year period were most dramatic during European settlement. Ponds were in pitch pine-mixed oak forests. Although archaeological evidence indicates that Native Americans were present from the Holocene, their impacts on the vegetation were not considered substantial. The first European settlements on Cape Cod occurred in the 1630s, but most areas were unoccupied until 1700. Prior to European settlement, beech and hickory (Fagus and Carya spp.) pollen was more common than at present. With European settlement, herbs and grasses increased, suggesting forest clearing and the creation of an open landscape. Charcoal influx was significantly (P<0.05) greater in postsettlement than presettlement time. Fires were more common in the past 300 years than in the previous 1,500 years [128,129]. "Following the decline of oak and other hardwood taxa, pitch pine has become a more common feature of the modern forests" . Pitch pine especially increased during the 20th century as fields and pastures were abandoned and reforestation began [128,129].
Charcoal and pollen from sediment cores taken from Green Pond in Augusta County, West Virginia, suggest that pitch pine dominance in the uplands surrounding the pond is likely a result of increased fires after European settlement, which began about 1750 .
Decreased pitch pine associated with fire exclusion: While pitch pine has increased in some areas due to increased fire frequencies associated with European settlement, in the same and other areas, fire exclusion since the early 1900s has allowed hardwoods to replace pitch pine. Aerial photos of the central Pine Barrens of Long Island showed that in 1938, 90% of the study area was open-canopy vegetation such as dwarf pine plains, pitch pine-scrub oak woodlands, heathlands, pitch pine-heath woodland, and scrub oak shrublands. From 1938 to 1994, wildfire size decreased, annual area burned decreased, and there were no fires in over 70% of the study area. Open-canopy barrens decreased to about 45% of the study area, mostly due to conversion of pitch pine-scrub oak to pitch pine-oak forests, although some of decreases were the result of residential and industrial development. Most wildfires occurred in the spring when winds were high, humidity was low, and surface fuels and litter were dry, but deep duff was moist. Researchers indicated that preserving open-canopy barrens will require active fire management .
From historical documents and forestry reports, Arabas  calculated a mean fire-return interval of 10 years for the 1888 to 1978 time period for a 500-acre (200 ha) study area in the Nottingham Serpentine Barrens of Pennsylvania. From 1937 to 1993, dominance of the study area changed from open- to closed-canopy vegetation. Fires were not actively suppressed in the area before 1930, but with increased fire exclusion the average annual area burned decreased from 200 acres (100 ha) before 1957 to 67 acres (27 ha) after 1957. Since about 1950, the area dominated by pitch pine savannahs and woodlands has decreased, and the area dominated by hardwood forests (75-100% deciduous tree cover) has increased .
Fire records and published primary and secondary literature show that an average of 55,280 acres (22,369 ha) of the 1,400,000-acre (550,000 ha) New Jersey Pine Barrens burned annually in wildfires from 1906 to 1939. This decreased to an average of 20,050 acres (8,115 ha)/year from 1940 to 1977. In the early 1900s, fire size averaged 110 acres (45 ha), and between 1940 and 1977, average fire size decreased to 15 acres (6 ha). Extensive fires occurred when there were drought conditions and high winds, which in this area can occur in any month of the year. Area burned in wildfires decreased after 1940. This may have been the result of decreased railroad usage, effective fire suppression, and/or increased winter and spring prescribed fires .
The following table  provides fire-return intervals for plant communities where pitch pine occurs. This list may not include all the plant communities in which pitch pine occurs.
|Fire regime information on vegetation communities in which pitch pine may occur. For each community, fire regime characteristics are taken from the LANDFIRE Rapid Assessment Vegetation Models . This vegetation model was developed by local experts using available literature and expert opinion as documented in the .pdf files linked from the Potential Natural Vegetation Groups listed below. Cells are blank where information is not available in the Rapid Assessment Vegetation Model.|
|Vegetation Community (Potential Natural Vegetation Group)||Fire severity*||Fire regime characteristics|
|Percent of fires||Mean interval |
|Minimum interval |
|Maximum interval |
|Rocky outcrop pine (Northeast)||Replacement||16%||128|
|Surface or low||52%||40|
|Surface or low||65%||12|
|Oak-pine (eastern dry-xeric)||Replacement||4%||185|
|Surface or low||90%||8|
|Southern Appalachians Forested|
|Surface or low||89%||6||3||10|
|Oak (eastern dry-xeric)||Replacement||6%||128||50|
|Surface or low||78%||10||1||10|
|Southern Appalachians Woodland|
|Table Mountain-pitch pine||Replacement||5%||100|
|Surface or low||92%||5|
|*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. 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. 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 [63,82].|
Fire Management Considerations
Prescribed fire is often necessary to suspend succession to hardwood-dominated stands and to maintain pitch pine populations. Deep-burning fires are necessary to convert sprout-origin pitch pine stands to seed-origin stands in New Jersey. Guidelines on the use of prescribed fire in pitch pine and pitch pine-hardwood stands to achieve various management goals are provided by Little .
In the Warm Springs District of the George Washington and Jefferson National Forests of Virginia, researchers promoted pitch pine regeneration through spring and subsequent fall prescribed fires. Fires burned in stands that were succeeding to scarlet and chestnut oak because of about 60 years of fire exclusion. The spring fire decreased hardwood canopy dominance, and the fall fire opened the canopy, exposed mineral soil, and stimulated pitch pine regeneration . In the Nantahala National Forest, pitch pine regeneration was "good" in stands that were selectively logged and burned . In mixed oak-pitch pine forests in the Nantahala National Forest, mid-September prescribed fires following clearcutting produced temperatures of only 113 to 138 Â°F (45-59 Â°C) at 1 to 2 inches (2.5-5 cm) below the soil surface. Large woody fuel consumption was minimal and likely a reason for the low soil temperatures produced. Researchers described the fires as "high intensity, low duration". At the time of the fire, mineral soils and duff were moist, resulting in minimal duff consumption. Soil temperatures and penetration depths were measured through thermologgers and tiles with temperature-sensitive paint .
A discussion on the use of prescribed fire to regenerate pines in southern habitats is provided by Van Lear and Waldrop . Considerations of wildlife needs and management goals are included.Soil/mycorrhizae: Soil nutrients were measured periodically for up to 5 years after felling and burning in mixed oak-pine stands in the Nantahala National Forest. For study results, see Knoepp and others . Changes in ectomycorrhizal diversity and soil nutrient availability following prescribed fires in pitch pine-mixed oak communities in southern New Jersey are presented by Tuininga and Dighton . For information on fire's effect on soils within mixed oak-pitch pine forests of the New Jersey Pine Barrens, see Burns .
Broad-scale Impacts of Plant Response to Fire
Pitch pine postfire regeneration methods may be influenced by tree size, bark
thickness, time since last fire, surface soil conditions, fire
severity, and/or fire season.
The thickness of pitch pine bark affects the survival of basal and dormant buds along the
trunk and within the crown. Devet  indicates that
pitch pine's scale bark is "very heat resistant". The build up of scale bark
creates a heat-resistant periderm that covers and protects the inner bark. At
the lower portion of the trunk, bark may be 1 to 2 inches (2.5-5 cm) thick .
In the Northeast, the average maximum inner and outer bark
thicknesses of pitch pine were 0.14 inch (0.36 cm) and 0.9 inch (2.3 cm), respectively.
The sizes of sampled trees were not reported (Stickel 1936, cited in ).
In Great Smoky Mountains National Park, researchers
found that pitch pine bark thickness increased with increasing DBH.
However, when trees reached about 9.8 inches (25 cm) in DBH, bark thickness
generally decreased with increasing DBH .
When dormant basal, trunk, or crown buds survive fire, pitch pine may regenerate by
Crown or bole sprouting:
Pitch pine is capable of producing sprouts from
buds at the internodes of multinodal stems. Dormant buds may be
concealed by bark or may develop into short branches of isolated or
fascicled needles. After fire, dormant buds along the bole may grow "prolifically and profusely" .
Spring prescribed fires in barrens vegetation in Pennsylvania's Centre County
consumed all pitch pine foliage. Trees were black and appeared dead, but within
several days needle fascicles appeared along the bole and larger branches. Within several weeks
of the fire, foliage appeared on smaller crown branches as well. One
year after the fire, the only evidence of fire was the blackened pitch pine trunks .
Basal buds are protected by thick bark and/or basal crooks.
Postfire sprout production from dormant basal buds can be affected by fire
severity, fire season , and time since last fire
. Studies in the Central Pine Barrens on Long
Island after the Sunrise Fire of August 1995 led researchers to suggest that
basal buds, located in or just under the duff layer, are most likely damaged or
killed during severe, growing-season fires .
Dwarf pitch pines in the New Jersey Plains often survive frequent fire and severe,
growing-season fires through sprouting. Frequently burned sites produce minimal
humus, and basal bud survival is high. In stands unburned
for about 25 years, humus layers are thicker, and basal bud survival decreases because of smoldering
conditions. In old stands unburned for 50 years or more, fire severity and pitch
pine mortality increase. When basal buds are killed, postfire regeneration
depends on seedling establishment. However, cone production is
typically reduced in old stands (unburned for over 50 years), which may reduce the regeneration potential of pitch pine in long
unburned, then severely burned stands. As time since last fire
increased in the New Jersey plains from 12 to 20 to 47 years, pitch pine mortality
increased and the number of seedlings produced increased
from 1,667 to 2,625 to 5,561 seedlings/ha, respectively. In stands unburned for 25 or more years, the percentage of
trees without cones was significantly higher (P=0.001) than
in stands unburned for 13 years or less . For a description of basal crooks
and a discussion of sprout production as it relates to stool age, see Sprouting from dormant basal buds in crooks and stools.
Cone survival and seedling establishment:
Pitch pine seed production and establishment are successful on frequently burned sites. Seeds
are produced at a young age, and seedling establishment is best on
exposed mineral soil. Open-grown seedlings produce mature cones as early as 10
years of age, and 3- to 4-year-old sprouts produce cones .
Serotinous cones are commonly produced in areas that experience
frequent fire. Seeds within serotinous cones are insulated from high
temperatures. In controlled studies, researchers found that
seeds from serotinous cones retained viability after 3 minutes of exposure to temperatures of 210
Â°F (100 Â°C). Linear regression
models predicted that serotinous cones collected from dwarf plains in
New Jersey could be exposed to oven temperatures of 790 Â°F (421 Â°C)
before internal seed locations reached 210 Â°F (100 Â°C). When a Bunsen burner was used
to heat cones, external temperatures of 946 Â°F (508 Â°C) were required to produce internal
temperatures of 210 Â°F (100 Â°C). The researcher noted that sample sizes were small, and
the controlled conditions may not entirely equate to field conditions .
Following a "high intensity" prescribed fire in pitch pine-dominated stands
that had not burned for over 70 years in the Nantahala National Forest, pitch
pine seedling density increased by 400% to 5,500 seedlings/ha. The fire burned
in April, and comparisons were made to prefire conditions. Time since fire was
not reported . Serotinous cone production is rare in the southern Appalachians, suggesting that
seeds came from open, persistent cones in the canopy or cones that remained on the
ground. In areas burned severely by the August 1995 Sunrise Wildfire in the
central pine barrens of Long Island, pitch pine seeds and cones were consumed in
the dwarf pine plains (unpublished data reported in ).
Sprouting and seedling establishment:
Pitch pine's POSTFIRE REGENERATION STRATEGY may depend on fire severity, fire season,
time since last fire, origin of burned stems, stem size, postfire seed
predation, and/or whether or not serotinous cones are produced on the site.
Development and subsequent regeneration of burned stands is affected by the
pitch pine regeneration strategy. Sprouts may grow more slowly than seedlings.
Sprouts of repeatedly burned trees, like those in the dwarf New Jersey pine plains, grow
very slowly .
Researchers studying dwarf pitch pine stands of various ages in New Jersey's
East and West Plains found that basal sprouting was the dominant postfire
regeneration method. Less than 1% of pitch pine stems were seedlings .
Sprouting was predominant 1
year after a May wildfire in an area of the New Jersey Pine Barren Plains that
had last burned 20 years earlier. Sprout density
ranged from 173,000 stems/ha to 868,000 stems/ha and averaged 418,500 stems/ha.
An average of 65 sprouts was produced per genet.
There were no pitch pine seedlings, which was unexpected since serotinous cone production in this area
was nearly 100%. The researcher did, however, observe many eastern towhees feeding
on dropped and opened pitch pine cones. Within 1 week of the fire,
almost all seeds were removed from the soil surface . Stem production occurs
with a "burst" right after fire. Production continues at a slower rate for at least 15
years after fire. The high density of small, low-growing stems in this area
promotes increased fire severity and frequency. High surface area per unit
biomass creates conditions that may support high fire intensity, and high
densities of compact stems promote fire spread .
Pitch pine produced sprouts and seedlings after a crown fire in
Pennsylvania's Sproul State Forest. The fire burned in late April 1990 when
winds were strong in mixed oak-red maple stands that were over 80 years old. The
fire size increased by 400 ha/hour. Three to 4 years after the fire, pitch pine
averaged 17Â±17 (SE) sprouts/ha on burned sites. Pitch pine seedling density
averaged 116Â±62 seedlings/ha on burned sites. There were no pitch pine seedlings
or sprouts on unburned sites. Pitch pine sprout and seedling densities were not
significantly different (P<0.05) on burned and unburned sites , likely because of variable
After a mid-July wildfire in 70-year-old Table Mountain pine-pitch pine
forests in Shenandoah National Park, Virginia, and adjacent private lands, pitch
pine produced both seedlings and sprouts. The fire burned with high and low
severities, with severity evaluated from cumulative tree mortality, crown
consumption, and average stem char heights. Low-severity sites experienced
surface fires that reduced total basal area less than 33%. Average stem char
heights were 7 feet (2 m) or less. High-severity sites experienced surface and
crown fires that reduced total basal area by 67% or more. Average stem char
heights were 20 feet (6 m) or greater. Pitch pine sprouting frequency soon after
the high-severity fire was over 50%, but 2 years after the fire only 4% of trees
had living sprouts. Researchers indicated that the high-severity fire may have
killed or damaged pitch pine roots through the removal of the forest floor and
heating of the mineral soil. Soil depths in the area were less than 6 inches (15
cm). Associated hardwood sprouts survived better than pitch pine sprouts. Pitch
pine seedling densities in the second postfire year averaged 2,186 seedlings/ha
on the high-severity burned site, 3,723 seedlings/ha on the low-severity burned
site, and 386 seedlings/ha on an unburned site . Carbon and nitrogen levels
were significantly lower (P<0.05) on high-severity burned than
low-severity burned and unburned sites. For more information on the soil
nutrients following this fire, see Groelschl and others .
Postfire seedling establishment was more common than sprouting after fire in
tall-stature, intermediate-stature, and dwarf pitch pine stands in eastern Suffolk
County, New York. For a complete summary of this study, see Fire Case Studies.
General fire effects:
Pitch pine density increased after "hot" fires but decreased after "cool" fires in the western
part of Great Smoky Mountains National Park. Some plots were burned in 1976 or 1977
in "hot" fires that removed over 25%
of the basal area; others burned in "cool" fires that removed less than
25% of the basal area; and other plots were unburned since before 1942. Pitch pine density
increased from 22 stems/ha in 1970 to 53 stems/ha in 1995 on "hot"-burned plots.
Density decreased from 133 stems/ha in 1970 to 108 stems/ha in 1995
on "cool"-burned plots and from 73 stems/ha in 1970 to 61 stems/ha in 1995 on
unburned plots . For pooled "cool" and
"hot" fire data, there was a general trend of increased pitch pine seedling and
sapling density but decreased canopy density after fire. Postfire litter depth was lowest after summer fires
and highest after winter fires. In the fourth postfire season, pine (Pinus
spp.) seedling densities were significantly (P=0.07) negatively
correlated with postfire litter depths .
In the southern Appalachians, decreases in canopy and midstory pitch pine trees were greatest after a
spring fire, when early postfire (3 months-1 year after fire) recovery was
evaluated on fall- and spring-burned pitch pine and Table
Mountain-pitch pine sites. Burned stands occurred on the Warm Springs Ranger District of the George
Washington and Jefferson National Forest in Virginia and on the Grandfather
Ranger District of the Pisgah National Forest in North Carolina. There were
3 prescribed fires: 1 in the fall on the Warm Springs Ranger District, 1 in
the spring on the Warm Springs Ranger District, and 1 in the spring on the
Grandfather Ranger District. None of
the stands were harvested or burned since the late 1930s or early 1940s. Pitch
pine decreases in the canopy and midstory layers occurred after all fires.
However, the greatest reductions occurred after spring fire on the Grandfather
Ranger District, where pitch pine basal area and density were reduced by almost
50% from prefire levels in the canopy and midstory. The spring Grandfather fire
produced the greatest flame and scorch heights of all 3 fires. There were 15,000
pitch pine seedlings/ha after the fall fire and 8,000/ha after the spring fire
on the Grandfather Ranger District, but researchers predicted future pitch pine
seedling mortality would be high . For a more complete summary of this study, see
Early postfire response of southern Appalachian Table Mountain-pitch pine stands to prescribed fires in North Carolina and Virginia.
The frequency, density, and basal area of pitch pine were lower than prefire
levels 3 months after a fire in pitch pine- and chestnut oak-dominated ridges in
the Nantahala National Forest. Stands were unburned for at least 70 years prior
to the spring prescribed fire. The fire was stand replacing, and the understory
was consumed. Pitch pine mortality averaged 18.5%. The number of pitch pine seedlings increased by 358% in the third postfire
month, but seedling density was 35% of the prefire density a year following the fire. It was
hoped that this prescribed fire would encourage pitch pine regeneration, but the
early mortality of postfire pitch pine seedlings suggested that successful
regeneration may require another fire or canopy-opening disturbance . For a more complete summary of this study, see
Early postfire effects of a prescribed fire in the southern Appalachians of North Carolina.
Wildfires and prescribed fires in New Jersey encouraged pitch pine seedling
recruitment. Wildfire-burned and 53-year-old unburned sites were in southeastern
Atlantic County. Prescribed fires burned in the Lebanon State Forest between
Burlington and Ocean counties. Both wildfires and prescribed fires burned in the
spring, but wildfires were considered more severe than prescription fires.
Postfire pitch pine recovery was evaluated 1 to 3 years after the fires. Pitch
pine biomass was significantly lower (P value not reported) on wildfire
than prescribed fire sites. Seedling densities on
both wildfire and prescribed fire
sites increased with increased time since fire, whereas mean aboveground biomass
decreased. Because the fires burned after seed fall and no serotinous cones were
observed in the wildfire burned area, the researcher considered a large, viable
seed bank unlikely. Seed on wildfire burned sites was likely dispersed from
nearby unburned stands and from surviving pines once able to reproduce again.
The researcher indicated that prescription fires did not burn into the humus layer,
and pitch pine seeds within this layer were not killed. Seedling survival, however, was
considered more likely in the wildfire burned
stands with a less dense canopy than the prescribed fire stands. Pitch pine stem
mortality was 17.3% and 23.1% on 2- and 3-year-old stands burned by wildfires, respectively.
The researchers observed no standing dead pitch pine stems on unburned or prescribed fire sites . For
information on the nutrient levels in aboveground postfire biomass, see Boerner .
For soil nutrient levels on burned and unburned sites, see Boerner .
|Pitch pine biomass and seedling density |
on unburned, wildfire burned, and prescribed burned sites 
|Time since last fire (years)||53||2||3||1||3|
|Fire season||unburned||May||April||March||Early spring|
|Mean aboveground biomass (kg/ha)Â±SE||42,089Â±28,986||22,121Â±9,494||9,692Â±4,975||35,923Â±7,911||31,488Â±12,570|
|Seedling density (seedlings/ha)||0||289||15,825||3,350||6,675|
Fire behavior in the New Jersey Pine Plains:
Stand characteristics and stem survival were studied in the Pine
Plains after a 1,400-acre (570 ha) fire on sites that had not burned for 20, 34, and 47 years. The plains region
burns primarily in crown fires, but "low-intensity"
surface fires are important in maintaining patches of low-growing, closed-canopy
plains and taller, open, semiopen, transition vegetation. A late-July crown fire
in the East Plains burned over 90% of the area, a convective column surface fire burned 1.4%
of the area, a wind-shift surface fire burned 5% of the area, and a man-made
backing fire burned the remaining 3.6% of the area. Fireline intensities of these fires,
calculated from scorch heights, averaged over 5529 kW/m for crown fires, 48.3
kW/m for convective column surface fires, and 13.5 kW/m for wind-shift surface and
man-made backing fires. Pitch pine canopy stem survival generally decreased from man-made backing fires to
wind-shift surface fires to convective column surface fire to crown fires.
Percent canopy stem survival increased from low-growing,
closed-canopy plains vegetation (dominated by 2- to 7-foot (0.5-2 m)-tall pitch pine, bear
oak, and black jack oak) to plains vegetation (5- to 7-foot (1.5-2 m)-tall canopy but with
≥1,000 stems/ha) to transition
plains vegetation (open to semiopen canopy of 10- to 20-foot (3-6 m)-tall pitch pine and open oak
shrub canopy) .
Plant Response to Fire
Pitch pine has numerous fire adaptations that allow it to establish and/or regenerate on burned sites. Postfire pitch pine regeneration may be through bole and crown sprouting from epicormic sprouts protected by thick bark, basal sprouting from the root crown, which is protected by crooks and/or soil, and/or through establishment from seed in nonserotinous or opened serotinous cones . As damage from fire increases, the predominant form of postfire regeneration changes from bole and crown sprouting to basal sprouting to seedling establishment .
Â© Tom Palmer, Friends of the Blue Hills. Photos taken 10 weeks after a late April fire
Broad-scale Impacts of Fire
Pitch pine survival may be affected by tree size, bark thickness, time since last fire, surface soil conditions, fire severity, and/or fire season. Fires first produce damage to pitch pine foliage and well-developed buds; additional heat is required to damage or kill the cambium. If dormant buds along the trunk and within the crown survive, pitch pine survives through crown regrowth and/or epicormic sprouting. Fires that kill dormant buds in the crown or along the trunk may still not produce sufficient temperatures to damage basal buds and kill the tree. Large trees are often less susceptible to fire damage than small trees because of thicker bark and higher crowns. However, old trees with low "vigor" are more likely to be fire killed than younger, more vigorous trees. Fire severity also affects survival and postfire regeneration. "Large head fires" killed 68% of 5- to 8-inch (13-20 cm) DBH pitch pines. "Slow-burning side fires" rarely killed trees of that size class. Fire season may also affect pitch pine survival and regeneration method. Fire damage is typically less when air temperatures are low than when temperatures are high and fuels are dry .
In pitch pine-dominated stands of Burlington County, New Jersey, a "light" prescribed fire killed pitch pine trees less than 6 inches (15 cm) in DBH, but a severe fire killed trees in the 11- to 15-inch (28-38 cm) DBH size class in open-canopy, upland sites . Fourteen months after an April 1933 surface wildfire in mixed-oak stands in Ulster County, New York, 6 live "butt-scorched" trees found in August were still alive. The surface fire killed the continuous mountain-laurel understory .A cross section from a pitch pine that was 12 inches (30 cm) at stump height in Monroe County, Pennsylvania, revealed that the tree survived and recorded 9 fires over its nearly 120-year lifespan. Researchers noted that the growth rate slowed with successive fires , but it seems that tree age may have also reduced growth rate.
Immediate Effect of Fire
More info for the terms: adventitious, root crown, shrub, tree
POSTFIRE REGENERATION STRATEGY :
Tree without adventitious buds and without a sprouting root crown
Tall shrub, adventitious buds and/or a sprouting root crown
Crown residual colonizer (on site, initial community)
Secondary colonizer (on-site or off-site seed sources)
More info for the terms: association, climax, density, hardwood, mesic, natural, seed tree, succession, tree, xeric
Commonly pitch pine is an early seral species that is replaced by hardwoods in the absence of fires that expose mineral soil and allow light to the reach the forest floor. However, in some very harsh habitats, pitch pine may represent a climax forest species. Clements  characterized pitch pine-dominated forests as fire subclimaxes. In New England and upstate New York, for example, pitch pine is considered a subclimax species that is maintained through fire [96,171]. On Mt Desert Island, the pitch pine vegetation association is a pioneer type that in the absence of disturbance is eventually replaced by red and/or white spruce (Picea rubens, P. glauca) . Ledig and Fryer  indicate that successional replacement of pitch pine communities is rapid when fire is excluded.
In xeric rock outcrops in the Shawangunk Mountains, pitch pine forests were characterized as a "physiographic climax" following a study of stand age, tree growth rates, and climate data. Pitch pine trees as old as 320 years occurred in the area. While small amounts of black tupelo (Nyssa sylvatica) and chestnut oak were present, site conditions were thought too severe for these trees to dominate. Researchers noted a lack of pitch pine recruitment since the 1970s, but noted that pitch pine turnover likely occurred through tree-by-tree replacement . In the Pitch Pine Ecological Preserve of Haut-Saint-Lauret, pitch pine may be a physiographic climax type in the outcrop habitats .
Shade: Pitch pine is generally considered shade intolerant . Results of a 1950s questionnaire showed that 35% of foresters surveyed rated pitch pine as intermediate in shade tolerance, 45% rated it as intolerant, and 20% rated it very shade intolerant .
Old field succession: Invasion by pitch pine occurs early after the abandonment of agricultural or pasture lands. Hotchkiss and Stewart  considered pitch pine a "pioneer" tree after studying secondary succession in abandoned fields in what is now the Patuxent Research Refuge in Maryland. In the southwestern Piedmont of Virginia, dendrochronological studies indicate that the oldest pitch pine in the area established in 1904. Fields were abandoned in the early 1900s. At time of study (2002), pitch pine was successionally "over mature" and no longer dominant due to increases in scarlet and chestnut oak .
On Martha's Vineyard, eastern redcedar (Juniperus virginiana) is often the first tree to establish after field abandonment. Pitch pine seedlings typically appear 15 to 40 years after field abandonment. Once pitch pine trees reach 50 to 100 years old, oak prominence increases. Sassafras (Sassafras albidum), beech (Fagus spp.), sweetgum (Liquidambar styraciflua), and pignut hickory (Carya glabra) appear when oaks are 125 to 300 years old .
The rapidity of old field invasions on the Burlington-Colchester-Essex sand plains of Vermont was associated with seed tree distance. In fields abandoned for about 10 years, pitch pine density was 1 tree/25 mÂ² when the nearest seed tree was 890 feet (270 m) away. When the nearest seed tree distance was 560 feet (170 m), pitch pine density was 3 trees/25 mÂ². When the nearest seed tree distance was 250 feet (75 m), pitch pine density was 7 trees/25 mÂ². Pitch pine density decreased to about 1 tree/25 mÂ² after fields had been abandoned for about 60 years. The researcher noted that pitch pine appeared to pave the way for the establishment of eastern white pine, which commonly occurs beneath the shade of pitch pine .
Pitch pine dominance decreased as old field succession progressed in southern New Jersey. Researchers monitored succession for 18 years in a shortleaf pine-pitch pine stand that established on an old field, abandoned in 1932, in what is now the Green Bank State Forest. In 1953, when the study was initiated, there were few hardwoods over 0.55 inch (1.4 cm) DBH. Periodic winter burning (March 1954, 1957, and 1962) was conducted on some sites. By 1971, hardwood densities increased greatly while the number of pines decreased greatly, regardless of winter burning. Researchers suggested that this successional pattern is natural for areas that support an oak-hickory climax .
Forest succession: In most cases, pitch pine is an early seral species that is replaced by hardwoods, spruces, or other pines in the absence of severe disturbance. In the Southeast, pitch pine is a temporary type that is replaced by hardwoods or shortleaf pine on mesic and xeric sites, respectively. The deep shortleaf pine taproot is able to penetrate rock crevices beneath shallow mountain soils, so shortleaf pine has better stability than pitch pine, which produces shallower roots .
On Mt Everett, pitch pine is a dominant species but its replacement or continued dominance of the harsh sites is unclear. Pitch pine recruitment has been continuous since the 1860s, although surveys of the area suggested no "significant" fires burned in the 20th century. Researchers suggest that winter storms, harsh climatic conditions, poor soils, high winds, droughts, and slow growth rates may maintain pitch pine's dominance. However, the importance of red maple and oaks increased over the 20th century, and these species grow faster than pitch pine, suggesting a possible hardwood conversion .
Insect outbreaks: Southern pine beetles can act as a successional agent in pitch pine forests. "Overmature" and/or stressed pitch pine trees are preferred by southern pine beetles, which often contribute to the turnover of forests to pine and hardwood seedlings and saplings . In pitch pine-dominated pine-oak forests in the Coweeta Basin, a drought-induced southern pine beetle attack accelerated the loss of pitch pine and succession to mixed oak-hardwood stands. Pitch pine regeneration was lacking in the study plots, but there was some regeneration in disturbed sites (roadsides, areas of fallen trees) outside of the study area. Pitch pine was lost at a rate of up to 10 ha/year when southern pine beetle populations reached epidemic proportions. The researcher noted that without fire, southern pine beetle outbreaks may accelerate the conversion of pitch pine-dominated forests to mixed oak-hardwood woodlands .
Fire: Pitch pine is well adapted to postfire regeneration through asexual and sexual means. Most of the information regarding pitch pine regeneration and succession following fire is provided in Fire Effects.
Seed production: Farrar  reports that "good" crops of pitch pine cones are produced every 4 to 9 years. Seed predation can be high and affect seed survival and success. Pitch pine trees as young as 3 years old may produce cones. Researchers observed seed cones on trees less than 3 feet (1 m) tall in Maine's Acadia National Park .
|In 1996 and 1997, pitch pine in Acadia National Park averaged 60 full-sized seeds/cone, and the range was 12 to 84 full-sized seeds/cone. The average number of viable seeds/fertile scale was 0.4. Researchers reported that the theoretical maximum number of viable seeds/fertile scale was 2.0, assuming no disease, predation, fertilization problems, or abortion . |
Seed predation: Squirrels, eastern towhees, and likely other wildlife feed on pitch pine seeds. High levels of seed predation and rapid seed removal can occur. Studies in Acadia National Park revealed 85% cone survival in 1996 and a decrease by 6 times in 1997, due largely to squirrel predation . Helm and others  noted that squirrel predation can affect collections of pitch pine cones from coastal locations in Massachusetts, New York, and/or New Jersey. Eastern towhees fed on pitch pine seed released from serotinous cones and picked seeds from opened cones 1 year following a wildfire in the New Jersey Pine Barrens. Within 1 week of burning, there were almost no pitch pine seeds on the soil surface .
Â© Tom Palmer, Friends of the Blue Hills. Photo taken 38 days after an early May fire.
Serotinous and nonserotinous cone production: Studies have shown that the highest levels of serotinous cone production are concentrated in or near the Coastal Plain region. Researchers indicate that serotinous cone production is highest in those areas that burn at high frequencies. While serotinous cone production is greatest in the Coastal Plain, low levels of serotiny occurred on Crozier Island, Canada . A low level of serotinous cone production outside of the most severely and frequently burned sites may offer pitch pine a regeneration advantage when sites eventually burn.
Serotinous cones were rare except from populations on or near the Coastal Plain when cones were collected throughout the pitch pine range (Quebec south to Georgia and west to Kentucky and central Ohio). Cones came from 509 trees in 79 stands. In the plains area of the New Jersey Pine Barrens, serotinous cone production was nearly 100%. Away from the Coastal Plain, just 9 trees produced serotinous cones. Six of these trees were in Clinton County, Pennsylvania, and regenerated after an "extremely" severe fire. Researchers concluded that serotinous cone production was advantageous on sites that burned severely and frequently but disadvantageous on sites that did not experience frequent, severe fires. They suggested that the distribution pattern of serotinous cone producing populations was dictated by the combined effects of fire occurrence, fire severity, and gene flow through pollen and seed dispersal .
Givnish  studied pitch pine in the New Jersey Pine Barrens and suggested that fire history was more important than gene flow in determining serotinous cone production at a scale beyond a few kilometers. Within the plains, serotiny averaged 98.8% in upland sites and 84% in lowland sites. Generally, serotinous cone production decreased with increasing distance from the plains; however, sites 2 to 12 miles (3-19 km) to the northeast, southeast, and west of plains averaged 90% to 98% serotinous cone production. These sites were downwind and in the likely direction of fire advancement from the center of the plains .
Some studies suggest that levels of serotinous cone production may decrease on sites burned less frequently or less severely than in the past. In the Central Pine Barrens of Long Island, New York, serotiny levels were about 90% on undisturbed dwarf pine plains but decreased to around 50% in cleared areas where taller, single-stemmed pitch pines were establishing. Researchers suggested that the lack of fire to open serotinous cones led to the invasion and successful establishment of seeds from surrounding pitch pine woodlands (unpublished data reported in ).
Seed dispersal: Studies of pitch pine seed dispersal were lacking as of the writing of this review (2007). However, several studies indicate that seeds from nearby stands can be important to the regeneration of burned stands when on-site seeds are consumed  or when there is a lack of fire to open serotinous cones . Wildlife likely aid in the dispersal of pitch pine seed [17,57,68]. In a review, Fowells  reports that although pitch pine seed is winged, wind does not disperse seed long distances.
Germination: Based on field studies, pitch pine seed germination is best on exposed mineral soil sites that are protected from predation. In the barrens of central Pennsylvania, overall germination was better when seeds were covered with litter and protected from predation. Seedlings on unprotected sites rarely survived 2 growing seasons. Poor germination occurred on unmanipulated seed beds in aspen (Populus spp.), scrub oak (bear and chestnut oak), and grass (poverty oatgrass (Danthonia spicata) and bluestem (Andropogon spp.)) communities . Studies conducted in Orono, Maine, revealed that pitch pine germination decreased in seed beds with low moisture-holding capacities under relatively high temperatures [33,57].
In controlled experiments, pitch pine germination was best at 77 Â°F (25 Â°C), and final germination percentages decreased with decreasing soil moisture . Pitch pine cones collected from coastal locations in Massachusetts, New York, and New Jersey averaged 45.5 germinants/cone when grown under controlled conditions . Seeds in serotinous cones are protected from high temperatures. For more on this, see Cone survival and seedling establishment.
Temperature, pH, light levels: Controlled experiments were conducted on pitch pine seed collected from fall-harvested cones from the barrens of Centre County, Pennsylvania. The optimum germination temperature was 77 Â°F (25 Â°C). At 77 Â°F (25 Â°C), the germination percentage averaged 82.7% and ranged from 52% to 100% for 24 replicates. No seeds germinated at 40 Â°F (5 Â°C) or 50 Â°F (10 Â°C) after 60 days, and germination averaged just 16% at 59 Â°F (15 Â°C). As soil moisture of Hagerstown silt loams decreased, germination was delayed and final germination percentages were significantly reduced (F=1%). Pitch pine seeds germinated better in dark than light conditions. At pH levels of 4.5 to 8.3 there were no significant (F=1%) differences in germination percentages .
Seedling establishment/growth: A combination of field and controlled studies suggests that pitch pine seedling establishment is most successful on thick mineral soils with high light levels. However, on rapidly draining soils decreased light levels may improve seedling establishment and/or growth [33,57].
Pitch pine seedlings grow slowly. One-year-old seedlings may reach maximum heights of only 0.4 inch (1 cm) on moist sites and 0.2 inch (0.5 cm) on dry sites. Growth rates increase substantially once seedlings reach a foot (0.3 m) tall. Under favorable conditions, growth rates may reach 2 feet (0.6 m)/year .
Sun and substrate effects: Field studies in Acadia National Park showed that pitch pine seedlings and saplings occurred in depressions where soil depth averaged 4.1 inches (10.5 cm) and duff layer thickness averaged 0.6 inch (1.5 cm). Researchers evaluated the conditions where 100 young pitch pine trees (1-13 years old) covered almost 1 acre (0.6 ha). Researchers concluded that thick mineral soil is favorable to pitch pine establishment .
Pitch pine seedlings grown from seed collected in Acadia National Park were monitored under controlled conditions. Seedling dry mass production was best in peat substrates exposed to full sun. Seedlings grown in sand exposed to full sun accumulated the least mass, and seedlings in sand or peat under low light (60% interception) conditions grew better than seedlings in sand and full sun. Temperatures were 7 to 9 Â°F (4-5 Â°C) cooler under shade cloth, and dry mass was evaluated after the first year of growth [33,57].
Seed origin effects: Site conditions affected seedling growth more than seed origin in a reciprocal transplant study of dwarf and normal-stature pitch pine seed collected from populations on Long Island. Regardless of seed origin, seedlings at the dwarf population site grew more slowly, experienced more mortality, and developed multiple stems more often than seedlings at the normal-stature site. However, there were differences with respect to seed origin and site related to reproductive age. Seedlings grown from seed collected in the dwarf population reproduced earlier than those from normal-stature sites, but dwarf-population seedlings reproduced earliest at normal-stature sites. Researchers concluded that plastic phenotypic environmental responses affected growth and survival more than genetic differences . In a provenance study, pitch pine trees grown from seed collected from the southern part of the Atlantic Coast Plain grew larger than those from the seed collected from the northern Atlantic Coast Plain .
Growth: Pitch pine growth can be affected by tree age, site conditions, and/or climate. Ledig and Fryer  reported that the pitch pine growth rate decreases at an "early" age. In southeastern New York, growth of pitch pine trees from 40 to 314 years old on the ridges of the Ramapo and Shawangunk mountains was monitored. For all sites, the average radial growth rate was 1.09 mm/year. Trees less than 51 years old had higher growth rates than trees over 99 years old. Growth rates ranged from a high of 2 to 3 mm/year in young trees during "favorable" years to a low of 0.25 mm/year in older trees during drought conditions . The growth of pitch pine on Mt Everett in the Taconic Mountains was very slow. The radial growth rate for the area averaged 0.47 mm/year, and some trees grew as slowly as 0.08 mm/year .
Pitch pine trees in North Carolina's Thompson Gorge grew most in the spring and fall. The average range of increases in circumference was 0.133 to 0.535 inch/year and averaged 0.24 inch/year for 11 sites monitored for 3 years . Pitch pine tree-ring growth was significantly correlated (P<0.05, r=0.53) with annual precipitation and temperature in a dry rock outcrop in the Shawangunk Mountains. Drought conditions produced decreased growth. Over the last 120 years in the area, pitch pine tree-ring growth was slow and averaged 0.33 mm/year .
On Nantucket Island, the growth of young, 3- to 8-year-old pitch pines was better within clumps of northern bayberry. Average annual growth rates of 20 young (3-8 years) and 20 older (11-25 years) pitch pine trees growing within and outside of clumps of northern bayberry were compared. In both age classes pitch pine grew more within northern bayberry clumps, but growth differences were only significant (P=0.01) for young pitch pines .
Vegetative regeneration: Pitch pine regenerates vegetatively through basal sprouting and epicormic branching [44,86], and layering may occur . On Long Island's Napeague Beach, low pitch pine branches buried by sand grew roots, and plants spread . However, on Mt Everett, prostrate-growing pitch pine did not reproduce by layering, but epicormic branching and basal sprouting occurred .
Sprouting from dormant basal buds in crooks and stools: Buds that give rise to basal or root crown sprouts form in the axils of primary needles just above the seedling cotyledons. Basal buds produce stem tissue, and sprouts are not adventitious. Buds are closely spaced, appearing clustered or whorled. Buds may occur under ground because of soil movement or litter accumulation and may form small clusters of fascicled needles. Sprouts arising from basal buds also form basal buds. Strong lateral branches can form from these buds even in the absence of stem injury .
Basal buds are protected by thick bark and/or basal crooks. Crooks form as seedling stems bend and grow horizontally before turning upright. Open-grown seedlings may form a crook at the root crown in the first year of growth. Shade-grown seedlings may not form well-developed crooks until 3 to 9 years old. In a stand of "spindly" pitch pine trees suppressed by bear oak growth, approximately 50% had poorly developed crooks, which researchers suspected would be susceptible to fire. After a prescribed fire burned pitch pine seedlings (<0.5 inch (1.3 cm) diameter), only seedlings with well-developed crooks had over 55% postfire sprouting. Researchers observed basal sprout production in pitch pine trees as old as 79 years. In New Jersey's West Plains region, sprouts came from stools that were 40 to 83 years old. The lifespan of dormant buds and stubby basal branches is estimated at 40 to 55 years. Older stools may produce many more sprouts than seedling crooks. In the plains region, as many as 249 one-year-old sprouts were produced by a single stool. Researchers suggested that stool age may affect sprout growth, and that dwarf pitch pines result from very slow-growing sprouts from old stools . In southern New Jersey, the largest trees producing basal sprouts were 8 inches (20 cm) in diameter .
The theory that old stools produce slow-growing pitch pine sprouts was substantiated in a study in the Pine Plains of New Jersey, where researchers attempted to release stems through removal of competing vegetation with fire or herbicides. Dominant stems in the plains were about 11 feet tall (3.4 m) and 27 years old. Stems were produced when stools were 40 to 60 years old. Release attempts did not encourage the production of vigorous stems likely to reach tree size. Researchers concluded that the many-decade-old plains stools were only capable of producing slow-growing, scrubby sprouts, and that the only way to convert the Pine Plains to tree-sized pitch pine forests was through seedling establishment . When pitch pine seedlings were planted in dwarf pitch pine-dominated areas of southern New Jersey, planted seedlings reached the height of 50-year-old plains sprouts in 17 years, suggesting that sprouts from stools grew more slowly than seedlings .
The number and height of sprouts from cut pitch pine trees near Mt Misery in Burlington County, New Jersey, generally increased with tree age until trees reached 34 years of age. A total of 25 pitch pine trees, thought to be of seed origin, were cut and monitored for 2 years. Sprout number and height generally increased with tree age but gradually decreased in trees over 34 years old. A 95-year-old pitch pine produced 59 sprouts within 2 months of cutting, but the entire tree died within 2 years of cutting. Average and maximum sprout heights were greatest in 4- to 11-year-old trees. Two trees, 30 and 32 years old, produced 303 and 330 sprouts 2 months after cutting, but no sprouts survived due, in part, to severe sprout browsing .
Epicormic or stem sprouting: Stem sprouts come from buds at the internodes of multinodal stems. These latent buds may be hidden under the bark or may develop into short branches with isolated or few fascicles (see photo above). Severe injury such as fire can trigger prolific, profuse dormant bud growth along the stem . For more on epicormic sprouting after fire, see Crown or bole sprouting.
Sprout vs. seedling growth: Pitch pine's predominant regeneration strategy may depend on site conditions, stem origin, and/or disturbance regimes. Vegetative regeneration was rare in the Pitch Pine Ecological Preserve of Haut-Saint-Lauret, Quebec. Regeneration was abundant and continuous since the last fire in 1957, but was not through vegetative means. Researchers suggested that pitch pine may be a physiographic climax type in the outcrop habitats . For more on pitch pine sprout and seed production after fire, see Sprouting and seedling establishment.
Growth Form (according to Raunkiær Life-form classification)
More info for the term: phanerophyte
RAUNKIAER  LIFE FORM:
Reaction to Competition
In view of its relatively low tolerance and its requirement of mineral soil for germination, pitch pine can best be maintained in stands by even-aged management with seedbed preparation and control of competing hardwoods.
Fire has been largely responsible for maintaining the pitch pine type and also has been responsible for the sprout origin, comparatively slow growth, and poor form that characterize this species. One severe fire may eliminate non-sprouting associates such as white pine (Pinus strobus); repeated severe fires may eliminate such species as shortleaf pine (P. echinata) and white oak (Quercus alba) which do not produce seed at as early an age as pitch pine and bear oak.
Possibly because pitch pine roots so deeply, it is relatively windfirm. In Maryland, Virginia pine proved much more susceptible to windthrow than pitch pine (9).
Life History and Behavior
Pitch pine cones open for pollination in April and May throughout its range [38,39,135,177]. Seeds are shed in August, September, and October [135,148].
Dormant buds capable of active growth when properly stimulated are the key to this recovery. Also, the thick bark gives a relatively high degree of protection to the dormant buds and to the cambium. Both pitch and shortleaf pines have these buds along the bole to an age of 60 years or more, but at such ages only in pitch pine do the buds at the base retain the potential for growth. In seedlings that have not yet developed thick bark, the lowermost buds may be protected by characteristic basal crooks in the stem that bring them into or against mineral soil on upland sites. Such buds often survive fires and produce new shoots (22,29,30).
Pitch pine seedlings that cannot sprout after fires are those occasional seedlings that never develop a basal crook, and around which insufficient soil accumulates to protect the buds; those that started on sphagnum or on the deep humus layer of poorly drained sites, and around which fire burns deeper than the surface where they became established; and those too young to have well-developed basal crooks. Though some open-grown seedlings may develop such crooks in their first year, shade-grown seedlings may take 9 to 10 years (20,22).
The sprouting vigor of older pitch pines varies with their life history. When single stems more than 40 years old are cut, some sprouts start but most die within 2 years. In contrast are the multi-stemmed stools that characterize some southern New Jersey localities with a history of frequent wildfires; these stools may be 60 to 90 years old and commonly have produced several generations of sprouts. The survival of new sprouts on such old stools may be associated with partial rejuvenation of the root systems (9).
Although pitch pine's sprouting ability is an asset in enabling trees to survive fire or other injuries, it is also a liability from the commercial point of view. Apparently the form and growth rate of sprouts decrease markedly with increased age of the root crown after crown age reaches about 20 years. Where wildfires have occurred at frequent intervals, often stands are composed largely of slow-growing sprouts from old stools. In many other stands the stems have been deformed by past fires and manifest holes with many small branches that have developed to replace killed crowns, boles with one or more crooks or forks where terminal shoots have been killed, or trees with flat tops where no leader has developed after the last one was killed.
Some pitch pine seeds may remain viable in the forest floor for 1 year, but there is no evidence that they can lie over for longer periods. In one instance, after heat from a July wildfire had opened many closed cones, most of the seeds germinated the following spring, though a few lay dormant until August and germinated after rains had broken a severe dry period. In another instance, when 2,400 seed spots were sown to pitch pine in late March 1955, delayed germination in the spring of 1956 provided as many as 1.4 seedlings per spot in some treatments (9). Germination of pitch pine seeds is epigeal (27).
Thick litter is unsuitable as a seedbed, even on poorly drained sites. In one study few seedlings were found in July on the thick litter of unburned sites. On similar areas treated with a severe September fire before seedfall, 16,600 to 56,300 seedlings per hectare (6,700 to 22,800/acre) were tallied on very poorly to imperfectly drained sites, and 2,200/ha (900/acre) on upland sites (21).
Droughts kill many pitch pine seedlings, but those less than 2 years old are most susceptible. A summer drought in 1957 killed 81 percent of the seedlings from a 1956 direct seeding in certain plots, and on comparable sites most of the seedlings started in 1955 survived (9).
At the end of the first year, shaded seedlings on upland sites usually have a height of about 2.5 cm (1 in), and a taproot 8 to 10 cm (3 to 4 in) long with a few laterals. In contrast, vigorous open-grown 1-year-old seedlings on upland sites may have stems 5 to 10 cm (2 to 4 in) high with a maximum height of 13 cm (5 in) and correspondingly greater root systems. On the moister, poorly drained sites, open-grown first year stems are usually 8 to 15 cm (3 to 6 in) high with a maximum height of 20 cm (8 in).
Pitch pine seedlings grow slowly for the first 3 to 5 years and then more rapidly. Some planted stands in Pennsylvania maintained an average height growth of 36 to 48 cm (14 to 19 in) between ages 6 and 17. After a seed-tree cutting in a New Jersey stand, the average height growth of dominant seedlings among the natural pitch pine reproduction was 0.5 and 0.7 m (1.5 and 2.2 ft) during the third and fourth growing seasons after the cutting, respectively (9).
Deer browsing and hardwood competition both reduce pine growth rates. In one study, young pines uninjured by deer grew 0.6 to 1.2 m (2 to 4 ft) more during a 5-year period than those that had their leaders browsed two or more times. In another study, cutting back hardwood sprouts twice resulted, after 6 years, in a 1.2 m (4 ft) increase in the height growth of the largest pines (9).
Seed Production and Dissemination
Although pitch pine is reported to bear good crops of cones at approximately 3-year intervals, production may be irregular. In southern New Jersey, good to excellent crops have occurred at intervals of 4 to 9 years. Occasionally, poor crops are borne in two successive years, although usually a poor crop is followed by fair to excellent crops for 1 to 3 years (9).
Pitch pine seeds are three-angled and 4 to 5 mm (0.16 to 0.20 in) long, although with the wings they are 15 to 21 mm (0.6 to 0.8 in) long. Because they differ in size, the number of seeds per unit of weight varies widely- from 97,700 to 181,200/kg (42,500 to 82,200/lb). In nursery practice fresh seeds need no stratification before sowing, and seeds are merely pressed in the soil at rates that produce 320 to 380 seedlings/m² (30 to 35/ft²) (27). While northern nurseries usually leave the seedlings in the seed bed for 2 years, southern nurseries lift year-old seedlings for planting.
Seed dissemination is variable, depending on the length of time that cones remain closed after maturity. On some trees, the cones open soon after maturity; at the other extreme, some cones remain closed for many years, until the heat of a fire opens them or until the trees are cut. Trees of the latter type are characteristic of the areas with a long history of wildfire.
Cone behavior is thought to be an inherited characteristic, but in southern New Jersey, groups of trees with different cone behavior are not widely separated geographically.
When cones open soon after maturity, seed dispersal begins about November 1 and ends in April in southern New Jersey (18). The pattern of dispersal seems similar to the pattern for shortleaf pine (Pinus echinata); in one study this species dropped 69 percent of its seed the first month, and 90 percent during the first 2 months. In New Jersey, probably about 90 percent of the seeds dispersed from a pitch pine source fall on the east side because the prevailing fair-weather winds are from the west (9).
On trees showing cone behavior between the two extremes, the cones open erratically within a few years after maturity. Apparently there is no fixed pattern of when, what, or how many cones open.
Although equipped with large wings, pitch pine seeds usually are not carried very far by wind. On the leeward of one stand, all natural reproduction in an abandoned field was within 90 m (300 ft) (9).
Flowering and Fruiting
Cones reach full size, 4 to 7 cm (1.5 to 2.8 in) long, and mature at the end of the second summer. Mature cones are 2.5 to 3.5 cm (1.0 to 1.4 in) wide when closed, 4.5 to 5.5 cm (1.8 to 2.2 in) wide when open. Closed cones are narrowly ovoid; open cones are ovoid and flattened at the base.
Growth and Yield
On the better sites in Pennsylvania, pitch pine maintains an average annual height growth of 0.3 m (1 ft) or more until the trees are 50 to 60 years old. The rate of height growth then starts to decline, and the trees add little to their height after they are 90 to 100 years old. On the best sites, diameter growth is 2.5 cm (1 in) in 5 years at 20 years of age, and falls to 2.5 cm (1 in) in 8 years at 90 years (9).
Total volume in cubic meters is maximum in Pennsylvania at 90 years, when fully stocked even-aged stands yield 210 to 350 m³/ha (15,000 to 25,000 fbm/acre). However, mean annual growth reaches its maximum at about 30 years- 3.0 to 5.8 m³/ha (43 to 83 ft³/acre), depending on the site (9).
In closed stands of seedling origin undamaged by fire, pitch pine self-prunes about as well as shortleaf pine, but in understocked stands it tends to produce somewhat larger and more persistent branches than shortleaf. Open-grown trees typically develop large spreading branches, which contribute to the rough appearance that many people associate with the species. Typical. pitch pine stands have been burned repeatedly, are understocked, and have suffered fire injury; consequently trees have either retained branches or have developed them from dormant buds along the boles.
Even without the stimulus of fire, pitch pines suddenly released by heavy cutting in a stand may develop branches along the bole. Pruning of living branches also may stimulate the development of new branches from buds or short shoots (9).
Molecular Biology and Genetics
Provenance/progeny tests of 156 trees in 17 natural stands of pitch pine distributed over the Atlantic coastal plain from Cape Cod, Massachusetts, to Cape May, New Jersey, were established in central New Jersey. Trees from southern seed sources grew faster than those from northern seed sources but adaptation of all sources decreased with increasing distance from the seed source. Variation among families within these 17 provenances was negligible (14).
There is an apparent contradiction between growth in provenance/progeny trials and isozyme analysis in different populations of pitch pine in central New Jersey (10). The isozyme studies seem to indicate that variation is due to differences between individuals, while the provenance/progeny trials suggest that variation in this species is due to seed source and not differences among families within a provenance. Whether to select provenances or individuals within provenances for tree improvement programs therefore is still an open question.
The differing cone-opening characteristics discussed earlier seem to be inherited, and trees at each extreme perhaps should be considered as separate races or ecotypes.
When pitch and shortleaf pines grow together, natural crossing may occasionally occur. Trees with intermediate characteristics have been seen in southern New Jersey, and similar trees have been reported in southern Pennsylvania (9).
At Placerville, CA, the Pacific Southwest Forest and Range Experiment Station crossed pitch pine with shortleaf, pond, Table Mountain, and loblolly pines. Pitch x loblolly hybrids (P. x rigitaeda) are produced in large quantities in South Korea for commercial plantings. Early field trials in Illinois, Maryland, and New Jersey showed only slight promise (9). With more careful selection of parent trees and extensive screening trials, the Northeastern Forest Experiment Station produced hybrids with exceptionally fast growth, good form, and winter hardiness for much of the natural range of pitch pine (20,21).
Barcode data: Pinus rigida
Statistics of barcoding coverage: Pinus rigida
Public Records: 5
Specimens with Barcodes: 16
Species With Barcodes: 1
IUCN Red List Assessment
Red List Category
Red List Criteria
- 1998Lower Risk/least concern (LR/lc)
National NatureServe Conservation Status
Rounded National Status Rank: N2 - Imperiled
Rounded National Status Rank: N5 - Secure
NatureServe Conservation Status
Rounded Global Status Rank: G5 - Secure
Relevance to Humans and Ecosystems
Other uses and values
A review of the ethnobotanical uses of pitch pine indicates that the Iroquois, Shinnecock, and Cherokee all utilized pitch pine. The Iroquois used the pitch to treat rheumatism, burns, cuts, and boils. Pitch also worked as a laxative. A pitch pine poultice was used by both the Iroquois and the Shinnecock to open boils and to treat abscesses. The Cherokee used pitch pine wood in canoe construction and for decorative carvings .
Value for rehabilitation of disturbed sites
Pitch pine has survived planting on and/or naturally colonized coal mines, anthracite mines, and landfill sites.
Coal mines: On coal mine spoils in Pennsylvania, pitch pine survival was 36% ten years after planting. The researcher noted pitch pine's usefulness in the revegetation of dry sites with acidic soils . Plantings were more successful than seedings on areas strip mined for coal in Ohio. Pitch pine survival ranged from 82% to 95% on 1- to 8-year-old plantings. When pitch pine was seeded, germination rates ranged from 4% to 22%, and first-year survival was 22% to 60% .
On black waste from anthracite mining in Pennsylvania, natural colonization by pitch pine was variable. Pitch pine was confined to steep north slopes with sparse abundance, although there was a nearby seed source .
Landfill: On the closed Fresh Kills Landfill on Staten Island, New York, surviving planted pitch pines averaged 4.86 feet (1.48 m) tall after 1 to 1.5 years .
Importance to Livestock and Wildlife
Pitch pine provides habitat and food for many wildlife species. Trees are used as cover, nesting, and foraging habitat. Deer browse seedlings and new sprouts, and small mammals and birds consume seeds.
Deer: Deer browse pitch pine seedlings and young pitch pine spouts. On Fire Island, where white-tailed deer populations doubled over a 5-year-period, pitch pine needles were "fairly common" in summer diets . On upland sites in the New Jersey Pine Barrens, young pitch pine is important winter deer browse . In a 1937 study by Little (cited in ), pitch pine was browsed most in an area of New Jersey where pitch pine, Virginia pine, and shortleaf pine grew together. Sixty-five percent of pitch pine seedlings or new sprouts were browsed.
Small mammals: Squirrels commonly feed on pitch pine seeds. In the Barrens Grouse Habitat management Study Area in central Pennsylvania, 512 red squirrel feeding sites were located. Feeding sites contained small piles of pitch pine cone fragments, but no intact cached cones were found. Primary feeding sites tended to be near pitch pine trees with cones . Additional information on squirrel predation of pitch pine seed is presented in Seed predation.
Birds: Numerous bird species utilize pitch pine habitats for roosting, nesting, mating, and foraging. Eastern towhees feed on pitch pine seeds. In the plains of the New Jersey Pine Barrens, the researcher observed eastern towhee feeding on pitch pine seeds released from serotinous cones and taking seed from opened cones. Within 1 week of a wildfire, there were almost no pitch pine seeds on the soil surface .
Pine warblers "favor" pitch pine forests with tall trees. Pine warbler populations in Massachusetts have declined since the 1940s and 1950s due to forest succession to oak in the absence of fire . Wild turkeys in the Nathaniel Mountain Refuge of West Virginia utilized Virginia pine and pitch pine thickets on southern aspects as winter roosting habitat . The red-cockaded woodpecker also uses pitch pine and pitch pine habitats. Red-cockaded woodpecker cavities have been found in pitch pine. Good foraging habitats for the woodpecker are southern pine or pine-hardwood stands with pine trees ≥9 inches (20 cm) DBH. Pure hardwood stands "are of little value" . Pitch pine provided 13% of the red-cockaded woodpecker cavity trees found in the London, Somerset, and Stearns ranger districts of the Daniel Boone National Forest. Average DBH of the 6 pitch pine cavity trees was 15.7 inches (39.9 cm) .
In the Barrens Grouse Habitat management Study Area, downy woodpeckers and black-capped chickadees occurred more in pitch pine in the winter than expected based on availability of pitch pine habitats. Winter avifauna foraged in the rough pitch pine bark. In the spring, great-crested flycatchers, blue jays, black-capped chickadees, black-and-white warblers, Nashville warblers, and chestnut-sided warblers were observed in pitch pine more than expected . In the Lebanon State Forest, eastern towhees and Carolina chickadees foraged less in oaks than in pines as the breeding season progressed from May to July. Pine warblers foraged mainly in pitch pine, and used oak trees less than expected based on foliage density from May to July in both oak- or pitch pine-dominated forests . In southeastern Massachusetts pine barrens with an open pitch pine canopy and a dense bear oak understory, eastern towhees, common yellowthroats, and prairie warblers made up 40% to 70% of the total breeding bird density within the study area. Prairie warblers foraged more often in pitch pine than common yellowthroats. When foraging above ground, eastern towhees preferred pitch pine and deciduous trees. Widely spaced pitch pine were important male prairie warbler song posts .
In pitch pine-scrub oak habitats southeastern Plymouth County, Massachusetts, researchers found that the majority of rare and declining bird species within a 72,731-acre (29,433 ha) study area were positively associated with early seral, open-canopy habitats. However, the majority of the study area was dominated by late-seral, closed-canopy habitats. Rare bird species hotspots made up just 2% of the total study area, suggesting active Habitat management may be necessary for the area's declining bird species .
Snakes: Pinesnakes utilized pitch pine habitats near the Toms River in Ocean County, New Jersey. Female pinesnakes were often located near pitch pine. For located males, 73% of the time the nearest vegetation was blueberry (Vaccinium spp.) or pitch pine. Males were found under logs or bark more often than females .
Insects: Diverse tiger beetles, plant hoppers, moths, and butterflies occur in pitch pine habitats. In bear oak, blackjack oak, and pitch pine pygmy forests of New Jersey's Burlington and Ocean counties, pitfall trapping of 2 tiger beetles (Cicindelidae unipuncata and Megacephala virginica) was more successful than expected, suggesting these species are more common than previously realized . Five species of plant hoppers were collected from pitch pine-bear oak barrens from New Jersey. The planthoppers were considered characteristic of northeastern pitch pine-bear oak communities .
Many moths and butterflies that utilize pitch pine habitats are of conservation concern. The endangered Karner blue butterfly is common in the pitch pine barrens of the Albany Pine Bush. In southern New England and southeastern New York, 56 Lepidoptera species of conservation concern are associated with pitch pine-bear oak barrens, ridgetop pitch pine-scrub oak barrens, scrub oak shrublands, heathlands, and maritime shrublands. Pitch pine-scrub oak communities and heathlands are the most important habitats for approximately 41% of state-listed rare and/or endangered Lepidoptera in Massachusetts and 23% of those listed in Connecticut. An additional 11 rare Lepidoptera species feed on pitch pine and are restricted to or reach their greatest abundance in pitch pine-scrub oak barrens in southern New England and southeastern New York. The sandy substrates in pitch pine-scrub oak communities are important nesting and/or foraging habitat for many arthropods. The greatest threats to these habitats are destruction and fragmentation, especially in New York, Connecticut, Massachusetts, and New Hampshire, where pitch pine-scrub oak vegetation has decreased "substantially" from historic extents. The lack of fire that maintains early seral communities is another threat .
In pitch pine-scrub oak habitats southeastern Plymouth County, Massachusetts, researchers found that the majority of rare and declining bird species within a 72,731-acre (29,433 ha) study area were positively associated with early seral, open-canopy habitats. However, the majority of the study area was dominated by late-seral, closed-canopy habitats. Rare moth species hotspots made up just 3% of the total study area, suggesting active Habitat management may be necessary for the area's declining moth species .
Nutritional value: The nutrient concentrations in 1-year-old pitch pine leaves collected for 18 months in the Brookhaven Forest, New York, are provided by Woodwell . Pitch pine leaves had much lower nutrient concentrations than associated vegetation in oak-pine forests. A database compiled by Pardo and others  also provides nutrient values for pitch pine. Values are from data presented in northeastern US publications and can be sorted by stand age, sample period, and/or region.
Cover value: Information on use of pitch pine as cover has been integrated into Importance to Livestock and Wildlife.
Pitch pine also serves as a food source for wildlife. Cones of pitch pine often remain on the trees unopened for several years or until the heat from a forest fire opens them. Seeds shed in mid-winter are an important source of food for squirrels, quail, and small birds such as the pine warbler, pine grosbeak, and black-capped chickadee. White-tailed deer and rabbits also browse young sprouts and seedlings (5,7).
Pinus rigida, the pitch pine, is a small-to-medium-sized (6–30 m (20–98 ft)) pine, native to eastern North America. This species occasionally hybridizes with other pine species such as loblolly pine (Pinus taeda), shortleaf pine (Pinus echinata), and pond pine (Pinus serotina); the last is treated as a subspecies of pitch pine by some botanists.
Pitch pine is found mainly in the southern areas of the northeastern United States, from coastal Maine and Ohio to Kentucky and northern Georgia. A few stands occur in southern Quebec and Ontario. This pine occupies a variety of habitats from dry acidic sandy uplands to swampy lowlands, and can survive in very poor conditions; it is the primary tree of the New Jersey Pine Barrens.
The needles are in fascicles of three, about 6–13 cm (2.4–5.1 in) in length, and are stout (over 1 mm (0.039 in) broad) and often slightly twisted. The cones are 4–7 cm (1.6–2.8 in) long and oval with prickles on the scales. Unusual for pines, if the main trunk is cut or damaged by fire it can re-sprout using epicormic shoots. This is one of its many adaptations to fire, which also includes a thick bark to protect the sensitive cambium layer from heat. Burnt trees often form stunted, twisted trees with multiple trunks as a result of the resprouting. This characteristic also makes it a popular species for bonsai.
Pitch pine is not a major timber tree, due to the frequency of multiple or crooked trunks; nor is it as fast-growing as other eastern American pines. However, it grows well on unfavourable sites. In the past, it was a major source of pitch and timber for ship building, mine timbers, and railroad ties because the wood's high resin content preserves it from decay. Pitch pine wood was also used for building radio towers in Germany as at Muehlacker and at Ismaning.
Pitch pine is currently used mainly for rough construction, pulp, crating, and fuel. However, due to its uneven growth, quantities of high quality can be very sought after, and large lengths of pitch pine can be very costly.
- Farjon, A. (2013). "Pinus rigida". IUCN Red List of Threatened Species. Version 2013.1. International Union for Conservation of Nature. Retrieved 17 July 2013.
- Moore, Gerry; Kershner, Bruce et al. (2008). National Wildlife Federation Field Guide to Trees of North America. New York: Sterling. p. 756. ISBN 1-4027-3875-7.
Pinus rigida, pitch pine, is a small-to-medium sized (6–30 m (20–98 ft)) pine, native to eastern North America. This species occasionally hybridizes with other pine species such as loblolly pine (Pinus taeda), shortleaf pine (Pinus echinata), and pond pine (Pinus serotina); the last is treated as a subspecies of pitch pine by some botanists.
Pitch pine is found mainly in the northeastern United States, from Maine and Ohio to Kentucky and northern Georgia. A few stands occur in southern Quebec and Ontario. This pine occupies a variety of habitats from dry, acidic sandy uplands to swampy lowlands, and can survive in very poor conditions; it is the primary tree of the New Jersey Pine Barrens.
The needles are in fascicles of three, about 6–13 cm (2.4–5.1 in) in length, and are stout (over 1 mm (0.039 in) broad) and often slightly twisted. The cones are 4–7 cm (1.6–2.8 in) long and oval with prickles on the scales. Unusual for pines, if the main trunk is cut or damaged by fire it can re-sprout using epicormic shoots. This is one of its many adaptations to fire, which also includes a thick bark to protect the sensitive cambium layer from heat. Burnt trees often form stunted, twisted trees with multiple trunks as a result of the resprouting. This characteristic also makes it a popular species for bonsai.
Pitch pine is not a major timber tree, due to the frequency of multiple or crooked trunks; nor is it as fast-growing as other eastern American pines. However, it grows well on unfavourable sites. In the past, it was a major source of pitch and timber for ship building, mine timbers, and railroad ties because the wood's high resin content preserves it from decay. Pitch pine wood was also used for building radio towers in Germany as at Muehlacker and at Ismaning. Pitch pine is currently used mainly for rough construction, pulp, crating, and fuel. However, due to its uneven growth, quantities of high quality can be very sought after, and large lengths of pitch pine can be very costly.
Gallery[edit source | edit]
View north from a fire tower on Apple Pie Hill in the New Jersey Pine Barrens. The vast pine forest is almost entirely made up of Pinus rigida.
References[edit source | edit]
- Conifer Specialist Group (1998). Pinus rigida. 2006. IUCN Red List of Threatened Species. IUCN 2006. www.iucnredlist.org. Retrieved on 12 May 2006.
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Names and Taxonomy