Shortleaf Pine Community Description

Authored By: R. E. Masters

Ecosystem Distribution and Extent

Shortleaf pine (Pinus echinata P. Mill.) has the widest geographic range of the southern pines (Mattoon 1915, Lawson and Kitchens 1983). East of the 100^th meridian it is second only to eastern white pine in extent of distribution (Little 1971).

Figure 1

See Figure 1 for range map (after Mattoon 1915, Haney 1962, Silker 1968, Little 1981, Lawson 1990). Following the latest period of glaciation shortleaf pine reached its current northwest limit in Missouri some 4,000 years ago and southern pines in general reached their current distributional limits in the east, a relatively recent 2,000 years ago (Buckner 1989, Delcourt and Delcourt 1991). While Mattoon (1915) listed shortleaf as occurring in 24 states and encompassing 1.14 million km²(440,000 mi²) more recent authors cite its historical and current geographical distribution as encompassing 22 states and the District of Columbia. This includes AL, AR, DC, DE, FL, GA, IL, KY, LA, MD, MO, MS, NC, NJ, NY, OH, OK, PA, SC, TN, TX, VA, and WV (Haney 1962, Lawson and Kitchens 1983, Lawson 1990, NatureServe 2006; USDA Plants Database 2006). Some evidence indicates it may have once occurred in Michigan (Fowells 1965). This species occurs from extreme southeastern New York west, sporadically through parts of Pennsylvania and the south central part of southern Ohio, then in southwest Illinois and southern Missouri south and west through eastern Oklahoma and east Texas, east through interior states and the Gulf states, then north through the Atlantic states to Delaware and New Jersey (Mattoon 1915, Sargent 1965, Sternitzke and Nelson 1970, White 1980, Williston and Balmer 1980, Lawson 1990). Shortleaf pine is currently listed as endangered in the state of Illinois (http://dnr.state.il.us/espb/datelist.htm). Its current conservation status is S1: Critically Imperiled, in Illinois, Pennsylvania and New York and S3: Vulnerable in Delaware (NatureServe 2006).

Shortleaf pine occurs across a number of physiographic regions including the eastern and western Gulf Coastal Plain, Atlantic Coastal Plain, Piedmont, Southern Appalachians and the Interior Highlands of the Ozark Plateaus and Ouachita Mountains (Mattoon 1915, Harlow and Harrar 1969, Nelson and Zillgett 1969, White 1980, Guldin 1986). Shortleaf does not occur in the Mississippi Valley (White 1980) which separates eastern and western populations by a distance varying from 50 to 225 km (31 to 140 mi). The highest concentrations of this species are found west of the Mississippi River in the Interior Highlands of Arkansas and eastern Oklahoma and also the upper Coastal Plain of southern Arkansas, northern Louisiana and east Texas (Mattoon 1915, Sargent 1965, Sternitzke and Nelson 1970, Guldin 1986). Concentrated populations also occur in the Piedmont and upper Coastal Plain in Virginia, North Carolina and South Carolina (White 1980, Guldin 1986, Nelson and Zillgitt 1969, Lawson 1990).

Marked declines, from 20-40 percent of the original range, in the extent of shortleaf pine as a relatively pure type were noted as early as 1915 as a result of extensive logging and land use change (Mattoon 1915). Substantial declines occurred from the 1900s to the 1950s with the onset of logging particularly in the Ouachita Highlands of Arkansas (Smith 1986). Although many of these logged areas initially came back with some presence of shortleaf, unless converted to other land use, the next major decline in this area occurred in the 1970s because of large industrial forest acquisitions and conversion to loblolly pine (J. Guldin, USFS, personal communication). Further declines have been noted in extent and frequency of occurrence in the south and east (White 1980). The wide-spread planting of loblolly pine north of its native range and in industrial forest operations has, in part, been responsible for some declines (Guldin 1986) as conversion to loblolly plantations have been made based on short-term growth and yield characteristics (Williston and Balmer 1980). Also, in the eastern part of the range, loblolly has been favored over shortleaf on sites where littleleaf disease (associated with Phytophthora cinnamomi) was common as loblolly is less susceptible (Campbell et al. 1953). The industrial timber company perspective that focuses on short-rotation loblolly pine plantations also limits the use of fire in managed stands (Masters et al. 2005).

The wide spread planting of loblolly north of its range also has implications for maintaining genetic integrity of shortleaf in the heart of its range as these two species readily hybridize west of the Mississippi and a high proportion of hybrid individuals have been reported from central Arkansas (Lawson 1990; Edwards and Hamrick 1995; Raja et al. 1998; Chen et al. 2003, 2004). Shortleaf also has been reported to naturally hybridize with pond pine (Pinus serotina Michx.), spruce pine (Pinus glabra Walt.), and pitch pine (Pinus rigida P. Mill.) (Dorman 1976, Little 1979). Fire suppression across its range has allowed midstory hardwoods to supplant this species as the dominant species in the canopy in many areas and prevent regeneration by this relatively shade intolerant species.

Environment

Climate

As expected, given its wide geographical distribution, shortleaf pine occurs throughout a wide range of climatic conditions, including subtropical humid and temperate continental climates (Trewartha 1968). Average annual temperatures range from 21ºC (70ºF) in southeast Texas to 9ºC (48ºF) in New Jersey (Mattoon 1915, Lawson 1990). Potential temperature extremes across its range from northeastern winter lows to southwest summer highs span 57ºC (134ºF) (Mattoon 1915). The northern limit of distribution corresponds with the 10ºC (50ºF) isotherm (Guldin 1986). The northern and western limits of shortleaf roughly coincide with the average annual rainfall isohyets of 114 cm (45 in) and 102 cm (40 in) respectively (Mattoon 1915). Average annual rainfall extremes range from less than 94 cm (37 in) in southeast Oklahoma to over 203 cm (80 in) in the southern Appalachians (Nelson and Zillgitt 1969, Guldin 1986). The rainfall zone averaging about 127 cm (50 in) in the southern Piedmont, south central Arkansas and northern Louisiana is where individuals of this species show their best development (Mattoon 1915, Guldin 1986, Lawson 1990). The species also tolerates a range of seasonal precipitation patterns throughout the year (Guldin 1986), from relatively well distributed rainfall across all months in the northeast and southeast to near annual drought conditions in late summer in the western portion of its range. To the west in east-central Oklahoma and in east-central Texas several small islands of shortleaf extend westward and are disjunct from the main body of its western distribution, 32 and 145 km respectively (20 and 90 miles), and extending in Texas to the average rainfall zone of 89 cm (35 in) (Silker 1968). These pine islands are associated with pockets of a podzolic soil type similar to that found in the main body of its range in east Texas and eastern Oklahoma and in other parts of its range (Billings 1938, Silker 1968, Nelson and Zillgitt 1969). Evapotranspiration on these sites is offset by the moisture storage and retention capacity (moisture availability) of the soils thus allowing shortleaf to persist on these islands (Silker 1968). At the northwestern limit in Missouri, the species appears to be limited by winter precipitation at the 43 cm (17 inches) winter precipitation isohyet. Shortleaf is also physiological limited by winter moisture as water uptake by roots decreases with decreasing temperatures (Fletcher and McDermott 1957).

Climate also affects growth rates of established shortleaf pine. A number of studies have documented how precipitation, temperature and soil moisture influence the growth response of shortleaf across various parts of its distribution (Schulman 1942, Fletcher and McDermott 1957, Friend and Hafley 1989, Grissino-Mayer and Butler 1993, Stambaugh and Guyette 2003). Shortleaf pine is sensitive to spring temperatures and initiates earlier cambial growth in response to early springs. However, cambial growth is limited in late spring and summer by water stress (Friend and Hafley 1989). Late summer moisture conditions also strongly influence the next years cambial growth in the winter part of shortleafs range (Schulman 1942, Friend and Hafley 1989, Stambaugh and Guyette 2003). Similar to work by Schulman (1942) in Arkansas, research in northern Georgia found that shortleaf responded with increased growth with greater than average rainfall and negatively to above average temperatures during the current growing season (Grissino-Mayer and Butler 1993). Schulman (1942) also noted that higher late-summer temperatures are strongly correlated with lower rainfall. In Missouri, extreme winter temperatures, in addition to the influence of limited moisture availability shows a decided negative relationship (Stambaugh and Guyette 2003).

Soils

The wide distribution of shortleaf pine is also related to its tolerance of a variety of sites, soils, soil moisture conditions and topography. It performs best on deep, well-drained upland soils such as on the upper Coastal Plain and Piedmont (Mattoon 1915, Harlow and Harrar 1969, Guldin 1986), but typically occurs on a range of well drained sites with from slightly coarse to loamy textured soils and within a soil moisture regime ranging from very dry to moist and where soils are moderately low in fertility and neither strongly acidic or alkaline (Fowells 1965, Harlow and Harrar 1969, Wright and Bailey 1982, McCune 1988, Duryea and Dougherty 1990, Masters et al. 2003). Generally shortleaf does not occur on deep, coarse sands which might be termed excessively drained or on poorly drained, fine textured soils with high clay content (Mattoon 1915, Nelson and Zillgitt 1969, Guldin 1986, McCune 1988, Lawson 1990, Masters et al. 2003). Fletcher and McDermott (1957) also stated that shortleaf pine did not occur on fine silt-loam windblown deposits (loess soils). In Illinois, this species appears to be associated with St. Peter sandstone parent material (Fletcher and McDermott 1957).

Shortleaf occurs broadly on Hapludults in the Coastal Plain and Piedmont to Paleudults in the Interior Highlands and western Gulf Coastal Plain, within the Ultisol order, characterized by clay accumulations in subsurface horizons, typically with some slope and moderate to moderately low mineral content respectively. In the southern Appalachians it occurs on Dystrochrepts in the Inceptisols order, characterized by weakly differentiated horizons and low calcium in subsurface horizons and these sites are usually moist (Nelson and Zillgitt 1969). Its southern distribution into peninsular Florida appears to be limited by soil type, which tends to be deep sands that are excessively drained.

Shortleaf pine occurs on a range of topographic sites from near sea level to elevations of 914 m (3,000 ft) (Mattoon 1915, Harlow and Harrar 1969). However, its ecological importance as a dominant forest type increases on sites where soils are thin and low in nutrient status and on exposed topographic sites prone to disturbances such as drought, at times of the year, and periodic fire and in areas where climatic conditions tend to be somewhat harsh (White 1980, Guldin 1986). On nutrient poor sites, shortleaf apparently has a better developed root system than competitors (McQuilken 1935). It is most prevalent on southern and western exposures in the Interior Highlands of Arkansas and Oklahoma but may occupy all types of sites at all elevations except steep north slopes (Palmer 1924, Johnson 1986, Foti and Glenn 1991, Kreiter 1994). Shortleaf also is a common early seral species establishing within 3-5 years on abandoned agricultural fields where an adjacent seed source is present (Billings 1938).

Ecosystem Description

Old-growth Condition and Forest Structure

Typical old-growth conditions are difficult to characterize because of the variety of sites on which the species could potentially occur and also the variety of species that could potentially co-mingle. In areas and in systems where all natural disturbance processes, including fire, are allowed to freely operate, old-growth stands may be characterized by open canopy pure or nearly pure pine stands with limited midstory and a bluestem dominated understory (See Vogl 1972, Komarek 1974, Fryar 1991, Sparks and Masters 1996, Batek et al 1999, Masters et al 1995)(Figure 9).

Figure 9

Hardwoods may be present to varying degrees depending on site characteristics (Vogl 1972, Fryar 1991, Kreiter 1994, Masters et al. 1995).

As shortleaf ages it becomes less tolerant of shade and neighboring crowns. By age 50 the crowns of trees develop a distinctly irregular shape and the canopy may be punctuated with numerous gaps (Mattoon 1915). However, where limited advanced regeneration is present in mixed stands, shortleaf often fails to colonize these gaps and is out-competed by hardwoods (Stambaugh 2001)(Figure 10). The structure of presettlement stands was variable (Bragg 2002). Stand structure was determined by the range of disturbance that initiated the stand and periodic disturbance events that continued throughout the life of the stand (see Turner 1935, Bragg 2002, Masters et al. 2005).

Figure 10

Stands that initiated following catastrophic disturbance typically develop as even-aged stands (Turner 1935) but this also applies to small old-field stands. However, in old growth stands excluded from anthropogenic disturbance, canopy dominant old-growth pines undergo senescence and are prone to bark beetle infestation and lightning strikes which predispose them and their neighbors to bark beetle depredations (Figure 11). Thus the pines begin dropping out of the stand and long-lived midstory hardwoods eventually take their place in a relatively short period of time (Kreiter 1994, Masters et al. 1995, Cain and Shelton 1996)(Figure 12).

Figure 11

The prevalence of even-aged old-field stands of shortleaf or in a mix with loblolly across the south, the even-aged condition of many second-growth shortleaf stands in the Ouachita Mountains, and the relative shade intolerance of shortleaf has led many silviculturalists to conclude that even-aged structure in shortleaf was the norm and that this was the usual mode of stand initiation and structural development (e.g., Lawson and Kitchens 1983, Guldin 1986). However, early foresters and ecologist reported the normal stand structure in old-growth shortleaf as decidedly uneven-aged (e.g., Turner 1935, Bragg 2002). Stand structure undoubtedly varied along a continuum of even-aged to uneven-aged conditions and varied in density according to the frequency and nature of the disturbance pulse (Turner 1935, Bragg 2002). One common characteristic irrespective of the age-distribution was the open nature of these forests and the occurrence of numerous canopy gaps depending on site conditions and fire regime (Little 1946, Fryar 1991, Murphy and Nowacki 1997, White and Lloyd 1998, Bragg 2002, Stambaugh et al. 2002). Within this context regeneration may occur under conditions as varied as even-aged cohorts in stand replacement disturbance events, even-aged patches under large canopy gaps within stands, or as individuals or small groups of distinct age-cohorts of different size classes (Bragg 2002, Stambaugh et al. 2002, Cassidy 2004).

Figure 12

Major Species

Because shortleaf is so widely distributed, it is found in many different community types and grows in association with a high number of tree species from pines to hardwoods. The relative density and distribution of shortleaf pine in these various mixtures is dependent on the disturbance regime.

Associated species on drier sites often include post oak (Quercus stellata Wangenh.), blackjack oak (Q. marilandica (L.) Muenchh.), black oak (Q. velutina Lam.), mockernut hickory (Carya tomentosa (L.) Nutt. ex Ell.), and in the western part of its range, black hickory (C. texana Buckl.). In uplands of the northwestern part of its range shortleaf pine, xeric oaks (Quercus spp.) and some hickories (Carya spp.) dominate the overstory with a high percentage of oak on steep north slopes and on post oak flats (Johnson 1986). This pine is often emergent on upper slopes. Stand density increases with available moisture. In the southern part of its range, loblolly pine is the most common pine associate. But shortleaf is also associated with longleaf and to a lesser extent spruce, pond and slash pine. In the northeast it is most commonly associated with pitch pine and Virginia pine (Pinus virginiana P. Mill.) and to a lesser extent Table Mountain pine (P. pungens Lamb.) and eastern white pine (P. strobus L.) (Lawson and Kitchens 1983). These associations follow an environmental gradient of moisture availability, soil type and fire frequency. On drier sites with frequent fire, shortleaf will assume canopy dominance. Under a fire regime of less than 3-year intervals and on coarser textured soils, longleaf pine with assume dominance where their ranges overlap (Masters et al. 2005). In longleaf-dominated stands associated with shortleaf where fire frequency is lengthened or excluded, the stands will gradually succeed to shortleaf (See Walker 1991, Pyne et al. 1996).

Other species associated with shortleaf in various part of its natural range include bear oak (Q. ilicifolia Wangenh.), black oak (Q. velutina Lam.), chestnut oak (Q. prinus L.), white oak (Q. alba L.), northern red oak (Q. rubra L.), scarlet oak (Q. coccinea Muenchh.), southern red oak (Q. falcata Michx. ), water oak (Q. nigra L.), willow oak (Q. phellos L.), blackgum (Nyssa sylvatica Marsh.), sweetgum (Liquidambar styraciflua L.), yellow poplar [tuliptree] (Liriodendron tulipifera L.), eastern red cedar (Juniperus virginiana L.), winged elm (Ulmus alata Michx.), red maple (Acer rubrum L.), white ash (Fraxinus Americana L.), and persimmon (Diospyros virginiana L.) (Palmer 1924, Marks and Harcombe 1975, Eyre 1980, Lawson 1990, Masters 1991a, White and Lloyd 1998, Murphy and Nowacki 1997, Smith et al. 1997, Rideout and Oswald 2002, Stambaugh et al. 2002).

Small trees, shrubs, and vines that frequently occur in shortleaf stands include serviceberry (Amelanchier arborea (Michx. f.) Fern.), rusty blackhaw (Viburnum rufidulum Raf.), flowering dogwood (Cornus florida L.), hawthorns (Crataegus spp.), American beautyberry (Callicarpa Americana L.), deerberries (Vaccinium spp), greenbriars (Smilax spp.), grape (Vitis spp.), Virginia creeper (Parthenocissus quinquefolia (L.) Planch.), eastern poison-ivy (Toxicodendron radicans (L.) Kuntze), smooth sumac (Rhus glabra L.), winged sumac (Rhus copallina L.), fragrant sumac (Rhus aromatica Ait.), various St. Johnswort (Hypericum spp.), and in the eastern part of its range, mountain laurel (Kalmia latifolia L.) (Palmer 1924, Buell and Cantlon 1953, Johnson 1986, Lawson 1990, Masters 1991a, Murphy and Nowacki 1997, White and Lloyd 1998, Rideout and Oswald 2002, Stambaugh et al. 2002).

In the understory, various bluestems (Andropogon spp. and Schizachyrium scoparium (Michx.) Nash), panicums (Panicum spp., Dichanthelium spp.), nutsedges (Scleria spp.), sedges (Carex spp.), and legumes (Lespedeza, Desmodium and others) are conspicuous across its range where fire is an integral part of stand history (Masters 1991a, Buell and Cantlon 1953, Sparks 1996, Sparks et al. 1998). Other important forbs include numerous asters (Asteraceae), tick-seed (Coreopsis spp.), pussytoes (Antenaria spp.), gayfeathers (Liatris spp.), sunflowers (Helianthus spp.), and wild petunia (Ruella spp.) (Masters 1991a, Sparks 1996, Smith et al. 1997, Crandall 2003).

Associated Ecosystems

Shortleaf Pine occurs in some 18 Society of American Foresters cover types (Eyre 1980). In three of these shortleaf is listed as a stand dominant; Shortleaf pine, Loblolly-shortleaf pine, and Shortleaf pine-oak. Often it occurs as a mixed type with various species of oaks and hickories (Braun 1950). The oak-pine forest is currently the largest forest type in the eastern United States (Lotan et al. 1978). In spite of its prevalence and importance, little research exists on the management of the oak-shortleaf pine type (Komarek 1981, Murphy and Farrar 1985). Frequent fire can shift forest community composition in the Ouachita Mountains (Little and Olmstead 1931) and in the Ozarks (Guyette and Dey 1997) from an oak-pine mixture to pine dominance. The oak-pine forest is a fire subclimax association and will succeed to an oak-hickory (Carya spp.) climax in the absence of fire (Bruner 1931, Little and Olmstead 1931, Braun 1950, Oosting 1956). Although fire is considered a major determinant in shaping the oak-pine ecosystem (Garren 1943, Oosting 1956), little is known about the influence of fire in this cover type in the Piedmont and Cumberland Plateau in the east (Lotan et al. 1978).

Subsections found in Shortleaf Pine Community Description

Stand Establishment

Click to view citations... Literature Cited

Batek, M. J., et. al. 1999. Reconstruction of early nineteenth-century vegetation and fire regimes in the Missouri Ozarks. Journal of Biogeography. 26: 397-412.
Billings, W. D. 1938. The structure and development of old field shortleaf pine stands and certain associated physical properties of the soil. Ecological Monographs. 8(3): 437-500.
Bragg, D. C. 2002. Reference conditions for old-growth pine forests in the Upper West Gulf Coastal Plain. Journal of the Torrey Botanical Society. 129(4): 261-288.
Braun, E. L. 1950. Deciduous forests of east North America. Philadelphia, PA: Blakiston Co. 596 p.
Bruner, W. E. 1931. The vegetation of Oklahoma. Ecological Monographs. 1: 99-188.
Buckner, E. 1989. Evolution of forest types in the Southeast. In: T.A. Waldrop. Pine-hardwood mixture: A symposium on management and ecology of the type. Asheville, NC: U.S. Department of Agriculture, Forest Service, Southeastern Forest Experiment Station: 17-33.
Buell,Murray F.;Cantlon,John E. 1953. Effects of prescribed burning on ground cover in the New Jersey pine region. Ecology. 34(3): 520-528.
Cain, M. D., and Shelton, M. G. 1996. The R. R. Reynolds research natural areas in Southeastern Arkansas: a 56-year case study in pine-hardwood overstory sustainability. Journal of Sustainable Forestry. 3(4): 59-73.
Campbell, W. A. (et al). 1953. Managing shortleaf pine in littleleaf disease areas. Asheville, NC: U. S. Department of Agriculture, Forest Service. Station Paper No. 25. 12 p.
Cassidy, P. D. 2004. Dynamics and development of shortleaf pine in east Tennessee. Knoxville, TN: University of Tennessee. 99 p. Ph.D.
Chen, J., C. G. Yauer and Y. Huang. 2003. Observations on mitochondrial DNA inheritance and variation among three Pinus species. Forest Genetics. 10: 271-276.
Chen, J., et. al. 2004. Bidirectional introgression between Pinus taeda and Pinus echinata: evidence from morphological and molecular data. Canadian Journal of Forest Research. 34: 2508-2516.
Crandall, R. M. 2003. Vegetation of the Pushmataha Wildlife Management Area. Pushmataha County, Stillwater,Oklahoma: Oklahoma State University. 114 p. M.S.
Delcourt, H.R.;Delcourt, P.A. 1981. Vegetation maps for North America. Geobotany. 11: 123-165.
Dorman, K.W. 1976. The genetics of breeding southern pines. Agric. Handb. 471. Washington, DC: U.S.DepartmentofAgriculture,ForestService. [Numberofpagesunknown] p.
Edwards, M. A., and J. L. Hamrick. 1995. Genetic variation in shortleaf pine, Pinus echinata Milll. (Pinaceae). Forest Genetics. 2(1): 21-28.
Eyre, F.H. 1980. Southern forest region. Forest cover types of the United States and Canada. Washington, DC: Society of American Foresters: 51-77.
Fletcher, P. W. and R. E. McDermott. 1957. Influence of geologic parent material and climate on distribution of shortleaf pine in Missouri. Research Bulletin 625. Columbia, MO: University of Missouri, College of Agriculture Agriculture Experiment Station. 43 p.
Foti, T.L.;Glenn, S.M. 1991. The Ouachita Mountains landscape at the time of settlement. In: Henderson, D.; Hedrick, L.D., eds. Restoration of old-growth forests in the interior highlands of Arkansas and Oklahoma. OuachitaNationalForestandWinrockInternationalInstituteforAgriculturalDevelopment: [Numberofpagesunknown].
Fowells, H. A. 1965. Silvics of Forest Trees of the United States. Washington, D. C.
Friend, A. L. and W. L. Hafley. 1989. Climatic limitations to growth in loblolly and shortleaf pine (Pinus taeda and P. echinata): a dendroclimatological approach. Forest Ecology and Management. 26: 113-122.
Fryar, R. D. 1991. Old growth stands of the Ouachita National Forest. In: D. Henderson and L. D. Hedrick , eds. Resotration of Old Growth Forest in the Interior Highlands of Arkansas and Oklahoma. Morrilton, Ark: Winrock International: 105-113.
Garren, K. H. 1943. Effect of fire on vegetation of the southeastern United States. Botanical Review. 9: 617-654.
Grissino-Mayer, H. D. and D. R. Butler. 1993. Effects of climate on growth of shortleaf pine (Pinus echinata Mill.) in northern Georgia: a dendroclimatic study. Southeastern Geographer. 33(1): 65-81.
Guldin, J. M. 1986. Ecology of shortleaf pine. In: P. A. Murphy , eds. Symposium on the shortleaf pine ecosystem. Monticello, Ark: Arkansas Cooperative Extension Service: 25-40.
Guyette, R.P.; Dey, D.C. 2000. Humans, topography, and wildland fire: the ingredients for long-term patterns in ecosystems. In: Yaussy, D.A., comp. , eds. Proceedings: workshop on fire, people, and the central hardwoods landscape, March 12-14, 2000, Richmond, Kentucky. Gen. Tech. Rep. NE-274. Newtown Square, PA: U.S. Department of Agriculture, Forest Service, Northeastern Research Station: 28-35.
Haney, G. P. 1962. A revised shortleaf pine bibliography. Asheville, NC: USDA Forest Service Southeastern Forest Experiment Station. 155. 74 p.
Harlow, W. M., and E. S. Harrar. 1969. Textbook of Dendrology Fifth edition. New York: McGraw Hill. 512 p.
Johnson, A. F., and W. G. Abrahamson. 1990. A Note on the Fire Responses of Species in Rosemary Scrubs on the Southern Lake Wales Ridge. Florida Scientist. 53: 138-143.
Johnson, F. L. 1986. Woody vegetation of southeastern LeFlore County, Oklahoma, in relation to topography. Oklahoma Academy of Science. 66: 1-6.
Johnson, Paul S.; Dale, Charles D.; Davidson, Kenneth R.; Law, Jay R. 1986. Planting northern red oak in the Missouri Ozarks: A prescription. Northern Journal of Applied Forestry. 3: 66-68.
Komarek, Edwin V. 1974. Effects of fire on temperate forests and related ecosystems: Southeastern United States. In: Kozlowski, Theodore T.; Ahlgren, Charles E. Fire and Ecosystems. New York: Academic Press: 251-277.
Kreiter, S. D. 1994. Dynamics and Spatial Patterns of a Virgin Old-growth Hardwood-pine Forest in the Ouachita Mountains, Oklahoma, from 1896-1994. Stillwater, Oklahoma: Oklahoma State University. 141 p. M.S.
Lawson, E. R., and R. N. Kitchens. 1983. Shortleaf pine. Washington, D. C.: U. S. Department of Agriculture, Forest Service. Agriculture Handbook No. 445. 157-161 p.
Lawson, E.R. 1990. Shortleaf pine, Pinus echinata Mill. In: Burns, Russell M.; Honkala, Barbara H., tech. comps. Silvics of North America: conifers. Agric. Handb. 654. Washington, DC: U.S. Department of Agriculture, Forest Service: 316-326 Vol. 1.
Little, E. L., Jr. 1971. Atlas of United States Trees. Washington, D. C.: U. S. Department of Agriculture, Forest Service. 1146. unpaged p.
Little, E. L., Jr. 1981. Forest trees of Oklahoma. Oklahoma City, Oklahoma: State Division of Agriculture. Pub. No. 1 Re. ed. no 12:240p p.
Little, E. L., Jr. and E. E. Olmstead. 1931. An ecological study of the southeastern Oklahoma protective unit. Oklahoma City, Oklahoma: Oklahoma Forest Service. 53 p.
Little, E.L., Jr. 1979. Checklist of United States trees (native and naturalized). U.S. Department of Agriculture Handbook. Washington, D.C.: U.S. Government Printing Office. 375 p.
Little, S. 1946. The effects of forest fires on the stand history of New Jersey's pine region. Forest Management Paper. No 2. Upper Darby, PA: U.S. Department of Agriculture, Forest Service, Northwestern Forest Experiment Station: 43 p.
Lotan, J. E., et. al. 1978. Effects of fire on flora - A state-of-knowledge review. WO-16. Washington, DC: U. S. Department of Agriculture Forest Service. GTR-16. 71 p.
Marks, P.L. and P.A. Harcombe. 1981. Forest vegetation of the Big Thicket, southeast Texas. Ecological Monographs. 51: 287-305.
Masters, R. E. 1991a. Effects of timber harvest and prescribed fire on wildlife habitat and use in the Ouachita Mountains of eastern Oklahoma. Stillwater. OK: Oklahoma State University. 351 p. Ph.D.
Masters, R. E. et al. 2003. Red Hills Forest Stewardship Guide. Tall Timbers Research Station Misc. Publ. 12.
Masters, R. E. et al. 2005. The missing ingredient for natural regeneration and management of souther pine. In: Proceedings of the Joint Conference, Society of American Foresters and Canadian Institute of Forestry. Edmonton, Alberta, Canada: Canadian Institute of Forestry: CD-ROM.
Masters,R.E.;Skeen,J.E.;Whitehead,J. 1995. Preliminary fire history of McCurtain County Wilderness Area and implicationsfor red-cockaded woodpecker management. In: Kulhavy,D.L.;Hooper,R.G.;Costa,R. Red-cockaded woodpecker:recovery,ecology and management. Nacogdoches,TX: Stephen F. Austin State University,School of Forestry,Center for Applied Studies: 290-302.
Mattoon, Wilbur R. 1915. Life History of Shortleaf Pine. Agric. Bull. 244. Washington, DC: U.S. Department of Agriculture. 58 p.
McCune, B. 1988. Ecological diversity in North American pines. American Journal of Botany. 75(3): 353-368.
McQuilken, W. E. 1935. Root development of pitch pine with some comparative observations on shortleaf pine. Journal of Agricultural Research. 51(11): 983-1016.
Murphy, P. A. and G. J. Nowacki. 1997. An old-growth definition for xeric pine and pine-oak woodlands. Asheville, NC: U. S. Department of Agriculture Forest Service Southern Research Station. 7 p.
Murphy, P. A., and R. M. Farrar, Jr. 1985. Growth and yield of eneven-aged shortleaf pine stands in the Interior Highlands. SO-218. U. S. Department of Agriculture Forest Service. SO-218. 11 p.
Nelson, T., and W. M. Zillgett. 1969. A forest atlas of the south. Asheville, NC: U. S. Department of Agriculture Forest Service Southern Forest Experiment Station and Southeastern Forest Experiment Station. 27 p.
Oosting, H.J. 1956. The study of plant communities. San Francisco: FreemanPublishing,Inc. 440p p.
Palmer, E. J. 1924. the Ligneious Flora of rich Mountain, Arkansas and Oklahoma. Journal of the Arnold Arboretum. 5: 108-135.
Pyne, S.J.;Andrews, P.L.;Laven, R.D. 1996. Introduction to wildland fire. New York: John Wiley. 769p p.
Raja, R. G. et al. 1998. Regeneration methods affect genetic variation and structure in shortlead pine (Pinue echinata Mill.). Forest Genetics. 5(3): 171-178.
Rideout, S., and B. P. Oswald. 2002. Effects of prescribed burning on vegetation and fuel loading in three east Texas state parks. Texas Journal of Science. 54: 211-226.
Rideout, S.; Oswald, B.; Legg, M. 2003. Ecological, political, and social challenges of prescribed fire in east Texas pineywoods ecosystems: a case study. Forestry. 76(2): 261-269.
Sargent, C. S. 1965. Manual of the trees of North America. New York: Dover Publications, Inc. Vol 1 p.
Schulman, E. 1942. Dendrochronology in pines of Arkansas. Ecology. 23(3): 309-318.
Silker, T. H. 1968. Bio-economic assay of conditions related to pine management on tension-zone sites. In: The Ecology of Southern Forest. 17. Baton Rouge, LA: Louisiana State University: 3-19.
Smith, D.M.; Larson, B.C.; Kelly, M.J.; and Ashton, P.M.S. 1997. The practice of silviculture: Applied forest ecology. New York: John Wiley and Sons. 537 p.
Smith, K.L. 1986. Sawmill: the story of cutting the last great virgin forest east of the Rockies. Fayetteville, AR: The University of Arkansas Press. 246 p p.
Sparks, J. C. 1996. Growing-season and dormant-season fire behavior and effects on vegetation in the Ouachita Mountains, Arkansas. Stillwater, OK: Oklahoma State University. 186 p. M.S.
Sparks, J. C. and R. E. Masters. 1996. Fire seasonality effects on vegetation in mid-tall-, and southeasten pine-grassland communities: a review. Trans. North American Wildlife and Natural Resources conference. 61: 230-239.
Sparks, J. C. et al. 1998. Effect of late growing-season and late dormant-season prescribed fire on herbaceous vegetation in restored pine-grassland communities. Journal of Vegetation Science. 9: 133-142.
Stambaugh, J. R., R. Muzika, and R. P. Guyette. 2002. Disturbance characteristics and overstory composition of an old-growth shortleaf pine (Pinus echinata Mill.) forest in the Ozark Highlands, Missouri, USA. Natural Areas Journal. 22: 108-119.
Stambaugh. M. C. and R. P. Guyette. 2003. Long-term growth and climate response of shortleaf pine at the Missouri Ozark forest ecosystem project. In: Proceedings of the 14th Central Hardwood Forest Congerence. Northeast Experiment Station GTR-NE-316: U. S. Department of Agriculture Forest Service: 448-458.
Sternitzke, H.S.;Nelson, T.C. 1970. The southern pines of the United States. Economic Botany. 24: 142-150.
Trewartha, G.T. 1968. An Introduction to Climate, 4th edition. New York, NY: McGraw Hill.
Turner, L. M. 1935. Catastrophes and pure stands of southern shortleaf pine. Ecology. 16(2): 213-215.
Vogl, R. J. 1972. Fire in the southeastern grasslands. Proceedings Tall Timber Fire Ecology Conference. 12: 175-198.
Walker, L. C. 1991. the southern forest: a chronicle. Austin, TX: University of Texas Press. 322 p.
White, D. L.; Lloyd, F.T. 1997. In: An Old-Growth Definition for Dry and Dry-Mesic Oak-Pine Forests. Asheville, North Carolina: USDA Forest Service, Southern Research Station.
White, David L.; F. Thomas Lloyd. 1998. An Old-Growth Definition for Dry and Dry-Mesic Oak-Pine Forests. In: General Technical Report SRS-23. Asheville, NC: Southern Research Station.
White, F. 1980. Forest cover types of the United States and Canada. In: F. H. Eyre. Shortleaf Pine. Washington, DC: Society of American Foresters: 53-54.
Wright, H.A.;Bailey, A.W. 1982. Fire ecology. New York: John Wiley. [Not paged] p.

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