Pinus echinata (shortleaf pine) description

Conservation Status

Pinus echinata

Miller (1768)

Common names

Shortleaf pine, shortstraw pine, southern yellow pine; choyyihissi kobayli [Alabamian]; 萌芽松 [Chinese]; エキナタマツ [Japanese].

Taxonomic notes

This species belongs to subgenus Pinus, subsection Australes Loudon, clade Taeda (Cruz-Nicolás et al. 2024). This clade is comprised of species found in the SE US, and most of the pines that co-occur with this species in mixed stands (including P. elliottii, P. glabra, P. palustris, P. serotina, and P. taeda) are in the same subsection. P. echinata hybridizes naturally with P. taeda; the hybrids tend to closely resemble P. echinata rather than P. taeda (Edwards-Burke et al. 1997). In eastern Kentucky (and perhaps elsewhere, where sympatric), P. echinata introgresses with Pinus rigida, producing trees with very small, P. rigida-shaped seed cones, and larger seed cones that are intermediate between the normal shape for each species (R. Clark email 2009.12.15).

Description

Monoecious evergreen trees to 40 m tall and 160 cm dbh, with a straight single trunk and a rounded to conic crown of spreading-ascending branches. Bark red-brown, scaly-plated, the plates bearing resin pockets <1 mm diameter. Twigs <5 mm thick, purplish green, often glaucous, aging red-brown to gray, roughened and cracking below leafy portion. Buds ovoid to cylindric, red-brown, 5-10 mm long, resinous. Leaves 2-3 per fascicle, spreading, persistent 3-5 years, (5-)7-11(-15) cm × ca. 1 mm, straight, slightly twisted, gray- to yellow-green, all surfaces with fine stomatal lines, margins finely serrulate, apex abruptly acute; sheath 5-10(-15) mm, persistent (saplings in shaded sites can have needles more than twice as long as mature dominant trees). Pollen cones clustered, cylindric, 15-20 mm long, yellow- to pale purple-green. Seed cones maturing in 2 years, semipersistent, solitary or clustered, spreading, symmetric, lanceoloid or narrowly ovoid before opening, ovoid-conic when open, 4-6(-7) cm, red-brown, aging gray, nearly sessile or on stalks to 1 cm, scales lacking contrasting dark border on adaxial surfaces distally; umbo central, with elongate to short, stout, sharp prickle. Seeds ellipsoid; body ca. 6 mm, gray to nearly black; wing 12-16 mm. 2n=24 (Kral 1993, Farjon 2010). See García Esteban et al. (2004) for a detailed characterization of the wood anatomy.

Also see the Key to the Pines of the Southeastern United States.

There are few similar species. The small resin pits in the bark are unique among pines of the eastern U.S. The short needles and small cones are also distinctive; the most similar pine that occurs with it is P. virginiana, which differs in cone shape (spiny, conical), with shorter buds (6-10 mm) and twisted, shorter needles (4-8 cm). Shortleaf pine can often be identified at a distance by the dense profusion of seed cones retained in the upper crown, but P. virginiana sometimes does that, too.

Life History

Vegetative reproduction is uncommon, although some fire-killed trees will resprout from the base. Nearly all trees originate from seed. Seedlings are stratified during winter cold and germinate in early spring. There is no seed bank, so all seedlings are from the prior year's seed crop. Germination is best on exposed mineral soil, such as after a fire. Herbaceous and shrub competition is the greatest threat to successful establishment. Growth is slow for a year or two as the root system develops; thereafter annual height growth is about 0.3-0.9 m/year, depending on site conditions. Trees begin to bear cones with viable seed after about 20 years and achieve full maturity, at which time seed production peaks, after about 40 years. Reproduction begins with appearance of pollen and fertile seed cones in mid-March to late April, depending on local climate. Fertilized seed cones develop through two growing seasons and mature in late summer to early fall of the second season. Seedfall begins in late October, with 90% of viable seed dispersed within 2 months. The seeds are small (10.2 mg) but, as is usual in wind-dispersed pines, the vast majority of seeds fall much less than 100 m from the tree. Large cone crops occur at intervals of 3-10 years, and are less common in the colder portions of the species' range. Most mature trees in habitat are likely to be harvested, but if protected from that fate, trees can commonly survive over 100 years, or until they are removed by succession or a stand-destroying disturbance (Lawson 1990 and citations therein).

Distribution and Ecology

USA: New York, Pennsylvania, Ohio, Illinois, Arkansas, Missouri, Kentucky, Tennessee, West Virginia, Maryland, Delaware, Virginia, North Carolina, South Carolina, Georgia, Florida, Alabama, Mississippi, Louisiana, Oklahoma and Texas. It is the most widely-distributed hard pine in eastern North America; presettlement abundance is estimated at 305,000 km² (Anderson et al. 2016). Currently, though, it dominates on about 30,000 km², and the current population trend is a declining one (Moser et al. 2007, Guyette et al. 2007). The species occurs at (3-)45-460(-910) m elevation on a wide range of soils, including silicate and carbonate parent materials, and in most of these soils there is substantial subsurface clay; growth is best on deep, well-drained, acidic sand and silt loams. Compared to its common associate Pinus taeda, it performs better on dry, nutrient-poor sites (Lawson 1990 and citations therein).

Shortleaf pine's native range has a warm-temperate, summer-humid climate. Annual precipitation is 1000-1500 mm, with the driest occurrences in the west (Oklahoma) and the wettest in the south (Mississippi to Florida). In the core range (SE OK, AR, and SE MO), precipitation is near the middle of this range. Average annual temperatures historically varied from 9°C in New Jersey to 21°C in Texas (Lawson 1990). Climate change forecasts for the latter 21st century indicate that habitat is likely to remain suitable in the core range, but that suitable conditions will be much more widespread in W TN, W KY, WV, and PA, although assisted migration will be needed if the species is to colonize those areas (USFS 2025).

Distribution data from USGS (1999).

This is generally a species of uplands and low mountains, tending to be less competitive on wet sites and probably elevation-limited by low temperatures. It is a major component of two Society of American Foresters forest types: shortleaf pine and shortleaf pine-oak; together, these account for 60% of all native shortleaf pines (Moser et al. 2007). However, due to its wide environmental tolerances, shortleaf pine is also a minor component of many other forest types, including those dominated by Pinus palustris, P. pungens, P. rigida, P. strobus, P. taeda, P. virginiana, and Juniperus virginiana. Given the species' large range, it may be associated with a wide variety of shrubs and flowering trees. Example trees include scarlet oak (Quercus coccinea), southern red oak (Q. falcata), water oak (Q. nigra), willow oak (Q. phellos), blackgum (Nyssa sylvatica), sweetgum (Liquidambar styraciflua), mockernut and pignut hickories (Carya tomentosa and C. glabra), winged elm (Ulmus alata), sourwood (Oxydendrum arboreum), red maple (Acer rubrum), American beech (Fagus grandifolia), and Carolina ash (Fraxinus caroliniana). Example shrubs include mountain laurel (Kalmia latifolia), flowering dogwood (Cornus florida), redbud (Cercis canadensis), persimmon (Diospyros virginiana), blueberries (Vaccinium spp.), huckleberries (Gaylussacia spp.), Japanese honeysuckle (Lonicera japonica), greenbriers (Smilax spp.), Virginia creeper (Parthenocissus quinquefolia), poison ivy (Toxicodendron radicans), and grape (Vitis spp.) (Lawson 1990 and citations therein).

Shortleaf pine is basically a shade-intolerant fire-dependent species, with most stands being even-aged and originating in the aftermath of stand-destroying fire (Lawson 1990). Shortleaf pine lacks the strong dependence on frequent fire seen in many other pines of the southeastern U.S. such as Pinus clausa, P. elliottii, or P. palustris. Rather, it is adapted to variable severity fire with variable return intervals, and has adaptations to everything from frequent low-severity fire to stand-destroying crown fires. For frequent low-severity fire, adaptations include thick bark on the lower trunk, and resprouting from the root collar of seedlings and young trees (Little and Somes 1956, McCune 1988). However, competing pines with good low-intensity fire adaptation may outcompete it on such sites. For moderate-severity fires, shortleaf is more tolerant than P. taeda and less so than P. rigida; as in many pines, mortality tends to be proportional to the extent of crown scorch. Shortleaf pine has a high, open crown so fires tend not to crown out, it can endure moderate defoliation by fire, and trees up to about 30 years old can basally resprout after being top-killed by fire. In the event of high-severity fire, mortality is typically very high, especially if in summer; winter fires cause lower mortality. Successful regeneration commonly occurs because of the large number of seeds dispersed from high in the crown. Under all scenarios, fire produces the mineral soil substrate necessary for successful regeneration, and on most sites discourages competition from woody angiosperms, although shortleaf may not successfully dominate on sites where many of the woody angiosperms can resprout from the rootstock (Carey 1992 and citations therein).

In general, fire is a requirement for shortleaf pine to persist in a stand. Fire-history studies in Missouri suggest an optimum fire return interval of 8-15 years (Stambaugh et al. 2007). Low- and moderate-severity fire tends to eliminate or reduce competition from woody angiosperms, and prescribed fire has been successful in this. On sites with fire return intervals of less than 10 years P. echinata tends to prevail over P. taeda because of its resprouting abilities, while longer return intervals favor P. taeda, presumably due to its more rapid growth potential. Fire also increases vulnerability to insect attack, particularly by southern pine beetle. However, as the time since fire increases, hardwoods first dominate the understory, then enter the overstory, and come to dominate the forest; after 150 to 200 years, only a few relict pines may remain (Carey 1992 and citations therein).

Euroamerican settlement has also provided habitat for Pinus echinata. Especially in the latter 19th century, vast areas of farmland were abandoned within the range of shortleaf pine. The species recolonized these areas extensively, and much of this area remains under shortleaf pine cover--many of these areas, today, have become commercial forest plantations (Carey 1992 and citations therein).

A few pests and pathogens are significant within the range of P. echinata, and some others are less important or are prominent mainly in commercial plantations. The most serious pathogen is Phytophthora cinnamomi, which causes littleleaf disease. This is most common on oldfield sites with low soil nutrients and poor drainage; it causes defoliation, which is eventually fatal. Afflicted trees respond positively to large doses of nitrogen. Shortleaf pine is also affected by fungal diseases such as root rot and red heart rot, but these are basically a consequence of age and will commonly result in eventual mortality of a tree that avoids all the more common hazards (wind, fire, chainsaws, hardwood competition, etc.). Among insects, the greatest threat is the southern pine beetle Dendroctonus frontalis, which can cause extensive mortality in stressed stands of similarly-sized trees and is also the leading cause of death in old-growth trees. A 1999-2001 outbreak of the pine beetle killed over 4000 km² of pines (Nowak et al. 2008). Overall, though, shortleaf pine pathogens and pests tend to not be a problem in natural pine and pine-hardwood stands (Brinkman and Smith 1969).

Shortleaf pine provides important wildlife habitat. The seeds are an important food source for birds and small mammals, while young stands provide cover for bobwhite quail and wild turkey (Walker and Wiant 1966, Lawson 1990). Prior to logging and fire suppression, red-cockaded woodpecker (Picoides borealis, a federally-listed threatened species), pine warbler (Dendroica pinus), and Bachman’s sparrow (Aimophila aestivalis) were common in shortleaf pine forests in Missouri; however, recent surveys find that no pine-obligate species remain, with birds all representing species associated with hardwood forest (Eddleman et al. 2007). Elsewhere in its range, old-growth shortleaf pine still provides habitat for cavity dwellers, including the red-cockaded woodpecker (Conner et al. 1991). Moreover, management efforts to restore the shortleaf pine ecosystem by thinning and prescribed fire have had some success in re-establishing pineland birds (Clawson et al. 2007).

Although P. echinata is assessed as of "Least Concern" for conservation due to its very wide distribution, the same cannot be said of the shortleaf pine ecosystem, which has declined substantially due to fire suppression, conversion of natural forests to commercial plantations, increased silviculture focused on P. taeda, and habitat loss due to land conversions (Anderson et al. 2016). The same issues once affected P. palustris, the longleaf pine, and widespread recognition of this problem led to an equally widespread rehabilitation of the longleaf pine ecosystem beginning in the 1980s; as of 2025, restored longleaf forests can be found throughout most of its historic range. About 25 years later, the same kind of transformation began to affect shortleaf pine, through the establishment of cooperatives such as the Shortleaf Pine Initiative. As of 2025 there have been 6 biennial conferences to discuss the shortleaf pine ecosystem and its restoration, and an overarching restoration plan (Anderson et al. 2016) has been released. Restoration efforts and assessments are now occurring throughout the species' range. This, coupled with cultural changes such as widespread acceptance of the importance of prescribed fire, indicates a promising future for the shortleaf pine forest. However, like longleaf, most restored shortleaf forests are likely to remain "islands" within a vast sea of profitable, privately-owned loblolly pine plantations.

Remarkable Specimens

The current largest tree on record is 124 cm dbh, 32.9 m tall, crown spread 23.8 m, last measured in 2018, located in Smith County, Texas (American Forests 2023). An earlier record tree was 108 cm dbh, height 42 m, crown spread 23 m, located in Myrtle, Mississippi (American Forests 1996). The tallest known one is in the Abrams Creek watershed in Tennessee; it is 45.5 m tall (Blozan 2011). Another tree, nearby in Great Smoky Mountains National Park, was 43.16 m tall in 2004 (Robert Van Pelt e-mail 2004.02.17).

In 2007, the field class of the Dendroecological Fieldweek established a crossdated age of 324 years for tree GMX307, in the Great Smoky Mountains National Park, Tennessee (Pederson 2008). Tree LAW38 had a crossdated age of 314 years, and was alive when collected in 1980 at Lake Winona Natural Area, Arkansas by D. Stahle and G. Hawks (NCDC 2006). It would be interesting to know if this tree survives.

Ethnobotany

Aboriginal use of Pinus echinata is recorded for the Cherokee people and mentions the use of its wood for carving, construction, and building canoes. The Choctaw made a cold infusion of the buds, which they drank as a treatment for worms, and the Nanticoke used "pellets of tar" (dried resin?) as an analgesic and a cathartic. The Rappahannock used a decoction of the foliage to wash swellings, and used the grated dried bark to induce vomiting in both humans and dogs (Native American Ethnobotany Database 2025). However, this seems a very incomplete listing. Dozens of different tribes lived within the vast range of shortleaf pine, and their people likely exploited the resource widely, putting it to the same general uses as other southern pines. These include (Native American Ethnobotany Database 2025):

Wood for fuel, soot to make a black body paint, saplings used to construct pens and similar structures, bark used as a floor covering (as in P. palustris).
Basketry, furniture, adhesives, torches, decoctions used to treat sores and pains (as in P. elliottii).
Infusions and resins ingested as an emetic or laxative (as in P. rigida).
Although not recorded in the range of shortleaf pine, the pitch of various pines around the world was used for waterproofing and as a sealant, e.g. to make baskets watertight.
People also managed the pine forests, primarily through the use of fire to achieve desired conditions such as an open understory.

When settlers arrived, shortleaf pine quickly found use as a timber tree. From the colonial period into the early 1800s it was intensively exploited for construction in the Mid-Atlantic region, used in framing, flooring, finish work, and furniture. It was also used in dockyards at port cities, and for boat building in coastal areas. Some lumber was exported to Britain and the West Indies from the earliest days of colonization, as was lumber from other hard pines within its range, mainly P. palustris and P. taeda. Overharvesting largely eliminated the resource along the Atlantic seaboard by the mid-1800s, but after harvest levels declined and regeneration increased on abandoned farm fields, shortleaf pine acreage dramatically increased in the east, so much that a 1915 assessment found “shortleaf pine is the only commercial conifer on more than 100,000 square miles of upland region between Virginia and northern Alabama and Mississippi.” At that time it was used for construction, furniture, and in early automobile construction. Exploitation of shortleaf pine followed a somewhat different trajectory in the western interior of its range, such as in Arkansas, Missouri, and Oklahoma. In these areas, shortleaf was the predominant timber species from the late 1800s, with peak harvest in 1899 and virtual elimination of the original forests by about 1920, except in the Ouachita Mountains of Oklahoma, where the largest patch of old-growth shortleaf persists to this day (Sutter c.2020 and sources cited therein). However, modern techniques of silviculture had started to take root by 1905, with parties such as the U.S. Forest Service and Yale School of Forestry advocating for science-based forestry to recover and replace the destroyed southern pine forests. Paradoxically, the process was speeded up by the Great Depression, thanks to creation of the Civilian Conservation Corps in 1933. This created armies of young men entering the forests at government expense to plant trees, fence cutover lands from feral pigs, and control wildfire. Ten new national forests were created on cutover land, and others were expanded. Replanting of these lands focused on loblolly pine, which usually exceeds other southern pines in terms of growth rates, but throughout much of its northern and western range shortleaf can outperform loblolly and was planted instead. The current distribution of shortleaf, then, is a legacy of these patterns of exploitation and subsequent practical silviculture.

Today, shortleaf is one of the most economically important southern pines. The strong, dense, straight-grained wood is primarily used for lumber, plywood, other structural materials, and pulpwood; even the taproots are used for pulpwood (Larson 1990). The denser and stronger grades of wood are used in construction of bridges, docks, factories, trestles, homes, and warehouses, while lower density wood is used for interior finish, subflooring, joists, sheathing, boxes, pallets, and crates. When treated, shortleaf is also used for railroad crossties, piles, poles, and mine timbers. Shortleaf pine is particularly good for poles, preferred to both longleaf and loblolly pine due to a smaller knot size and excellent taper (Anderson et al. 2016 and sources therein). Like longleaf pine, shortleaf grows more slowly than loblolly but is a better choice for landowners interested in growing longer rotations and realizing nontimber values such as wildlife habitat, fish and game, conservation, ecosystem diversity and resilience, or recreation; and it complements longleaf because it reaches best development in a substantially cooler climate (Anderson et al. 2016 and sources therein).

Besides the uses named above, shortleaf pine is also used as an ornamental species, mainly within its native range; it grows relatively quickly and when open-grown makes a fine landscape tree. It is cold-hardy to Zone 6 (cold hardiness limit between -23.2°C and -17.8°C) (Bannister and Neuner 2001). Due to its relative drought tolerance and greater tolerance of rocky, nutrient-poor soils compared to other southern pines, it is also a popular choice for rehabilitating eroded areas and mine sites (Lawson 1990). Shortleaf pine does best on mine sites when planted as a pure stand or mixed with other pines. It also grows well with European alder (Alnus glutinosa), a nitrogen-fixing species (Vogel 1981).

Shortleaf pine has also been used in a number of dendrochronological studies, such as a fire history study in Arkansas that identified periods of differing anthropogenic fire frequencies (Guyette et al. 2006).

Observations

The best examples of shortleaf pine and shortleaf pine-oak ecosystems I have seen have been in Arkansas, particularly at Hot Springs National Park and north of there in the Ouachita National Forest. The largest surviving tract of relatively pristine shortleaf pine forest is in the McCurtain County Game Reserve in Oklahoma, but the area seems to be difficult to get into and there are some access restrictions. Some exceptionally large and tall trees occur along the Abrams Creek Trail in Great Smokies National Park (Will Blozan pers. comm. 2024.03.08). For a more expansive inventory of locations, see the map at iNaturalist.

Remarks

The epithet echinata means "spiny", likely in reference to the mature seed cones.

Citations

American Forests 1996. The 1996-1997 National Register of Big Trees. Washington, DC: American Forests.

American Forests 2023. 2021 National Register of Champion Trees, accessed 2023.02.22.

Anderson, M., Hayes, L., Keyser, P. D., et al. 2016. Shortleaf Pine Restoration Plan. Shortleaf Pine Initiative. https://www.chjv.org/wp-content/uploads/Shortleaf_Pine_Restoration_Plan_Single-Page-Layout.pdf, accessed 2025.03.23.

Blozan, Will. 2011.05.14. Abrams Creek, TN- new shortleaf pine and paw-paw record. www.ents-bbs.org/viewtopic.php?f=74&t=2487, accessed 2011.05.15.

Brinkman, K. A., and R. C. Smith. 1969. Managing shortleaf pine in Missouri. Station Bulletin 875. Agricultural Experiment Station, University of Missouri, Columbia, MO. 35 p.

Carey, Jennifer H. 1992. Pinus echinata. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: https://www.fs.usda.gov/database/feis/plants/tree/pinech/all.html, accessed 2025.03.15.

Clawson, R., C. Steen, K. Houf, and T. Thompson. 2007. Avian response to pine restoration at Peck Ranch Conservation Area. Pp. 176-178 in Kabrick, J. M., Dey, D. C., and Gwaze, D. (eds.), Shortleaf Pine Restoration and Ecology in the Ozarks: Proceedings of a Symposium. General Technical Report NRS-P-15, USFS Northern Research Station, Newtown Square, PA.

Conner, R. N., D. C. Rudolph, D. L. Kulhavy, and A. E. Snow. 1991. Causes of mortality of red-cockaded woodpecker cavity trees. Journal of Wildlife Management 55(3):531-537.

Cruz-Nicolás, Jorge, Juan Pablo Jaramillo-Correa, and David S. Gernandt. 2024. Stochastic processes and changes in evolutionary rate are associated with diversification in a lineage of tropical hard pines (Pinus). Molecular Phylogenetics and Evolution 192:108011, https://doi.org/10.1016/j.ympev.2024.108011.

Eddleman, W. R., R. L. Clawson, and J. Eberly. 2007. Birds of shortleaf pine forests in Missouri: an historical and contemporary perspective. Pp. 168-175 in Kabrick, J. M., Dey, D. C., and Gwaze, D. (eds.), Shortleaf Pine Restoration and Ecology in the Ozarks: Proceedings of a Symposium. General Technical Report NRS-P-15, USFS Northern Research Station, Newtown Square, PA.

Edwards-Burke, M. A., J. L. Hamrick, and R. A. Price. 1997. Frequency and direction of hybridization in sympatric populations of Pinus taeda and P. echinata (Pinaceae). American Journal of Botany 84(8):879-886.

Guyette, R. P., Spetich, M. A., and Stambaugh, M. C. 2006. Historic fire regime dynamics and forcing factors in the Boston Mountains, Arkansas, USA. Forest Ecology and Management 234(1–3):293–304. https://doi.org/10.1016/j.foreco.2006.07.016.

Guyette, R. P., R.-M. Muzika, and S. L. Voelker. 2007. the historical ecology of fire, climate, and the decline of shortleaf pine in the Missouri Ozarks. Pp. 19-27 in Kabrick, J. M., Dey, D. C., and Gwaze, D. (eds.), Shortleaf Pine Restoration and Ecology in the Ozarks: Proceedings of a Symposium. General Technical Report NRS-P-15, USFS Northern Research Station, Newtown Square, PA.

Lawson, Edwin R. 1990. Pinus echinata Mill. shortleaf pine. Pp. 316-326 in R. M. Burns and B. H. Honkala (technical coordinators), Silvics of North America. Volume 1. Conifers. Agric. Handb. 654. Washington, DC: U.S. Forest Service. Available: https://www.srs.fs.usda.gov/pubs/misc/ag_654/volume_1/pinus/echinata.htm, accessed 2025.03.23.

Little, S., and H. A. Somes. 1956. Buds enable pinch and shortleaf pines to recover from injury. Station Paper No. 81. USFS Northeastern Forest Experiment Station, Upper Darby, PA. 14 p.

McCune, Bruce. 1988. Ecological diversity in North American pines. American Journal of Botany 75(3): 353-368.

Miller, P. 1768. The Gardener's Dictionary, ed. 8. London. Pinus no. 12. Available: botanicus.org/title/b12066618, accessed 2011.05.20.

Moser, W. K., Hansen, M., McWilliams, W. H., and Sheffield, R. M. 2007. Shortleaf pine composition and structure in the United States. Pp. 19-27 in Kabrick, J. M., Dey, D. C., and Gwaze, D. (eds.), Shortleaf Pine Restoration and Ecology in the Ozarks: Proceedings of a Symposium. General Technical Report NRS-P-15, USFS Northern Research Station, Newtown Square, PA.

[NCDC 2006] Data accessed at the National Climatic Data Center World Data Center for Paleoclimatology Tree-Ring Data Search page. http://hurricane.ncdc.noaa.gov/pls/paleo/fm_createpages.treering, accessed 2006.09.08, now defunct.

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Nowak, J., Asaro, C., Klepzig, K., and Billings, R. 2008. The southern pine beetle prevention initiative: working for healthier forests. Journal of Forestry 106:261–267.

Pederson, N. 2008. Eastern OlDLIST: A database of maximum tree ages for Eastern North America. http://people.eku.edu/pedersonn/oldlisteast/, accessed 2008.12.07, now defunct.

Stambaugh, M. C., R. P. Guyette, and D. C. Dey. 2007. What fire frequency is appropriate for shortleaf pine regeneration and survival? Pp. 121-128 in Kabrick, J. M., Dey, D. C., and Gwaze, D. (eds.), Shortleaf Pine Restoration and Ecology in the Ozarks: Proceedings of a Symposium. General Technical Report NRS-P-15, USFS Northern Research Station, Newtown Square, PA.

Sutter, Robert. c.2020. Ecological and social history of shortleaf pine. https://shortleafpine.org/why-shortleaf/history, accessed 2025.03.24.

[USFS] United States Department of Agriculture, Forest Service. 2025. Climate Change Atlas, shortleaf pine (Pinus echinata). (Forecasts for RCP 4.5 and RCP 8.5 emissions scenario). https://www.fs.usda.gov/nrs/atlas/tree/110, accessed 2025.03.235.

Vogel, Willis G. 1981. A guide for revegetating coal minespoils in the eastern United States. Gen. Tech. Rep. NE-68. USFS Northeastern Forest Experiment Station, Broomall, PA. 190 p.

Walker, Laurence C., and Harry V. Wiant Jr. 1966. Silviculture of shortleaf pine. Bull. No. 9. : Stephen F. Austin State College, School of Forestry, Nacogdoches, TX. 59 p.