“Now the Pine Forests Opened to View

A stand of 200-year-old longleaf pine growing in the red hills area north of Tallahassee, Florida.

Bartram (Van Doren 1928) described a typical longleaf

dominated stand as he traveled through the lower coastal plain in 1773:

“Now the pine forests opened to view. We left the magniﬁcent savanna and its delightful groves, passing through a level, open, airy pine forest, the stately trees scatteringly planted by nature, arising straight and erect from the green carpet, embellished with various grasses and flowering plants; … we joyfully entered the borders of the level pine forest and savannas which con-

42 tinued for many miles, never out of sight of little lakes or ponds, environed with illumined meadows, the clear waters sparkling through the tall pines.”

SPRING 2000

NATIVENATIVE PLANTS JOURNALJOURNAL THE Longleaf Pine ECOSYSTEM OF THE SOUTH

Kenneth W Outcalt

Abstract ongleaf pine (Pinus palustris longleaf pine area and techniques P. Mill. [Pinaceae]) and incentives to do so are discussed. Longleaf pine (Pinus palustris P. Mill. L is synonymous with [Pinaceae]) was once the most Southern forests. It once dominated Habitat prevalent pine type in the southern 24.3 million ha (60 million ac) in Longleaf pine was native to a wide US. Stands of longleaf were also the southeast (Figure 1), stretching range of ecosystems. Along the habitat for a vast array of plant from southeastern Virginia, south to Atlantic and Gulf coastal plain, it species. Decades of timber harvest central Florida, and west into eastern was the dominant tree on wet followed by conversion to agricul- Texas (Stout and Marion 1993). It flatwoods and savannas. It also ture, urban development, or to other was a co-dominant along with dominated higher, droughty sand pine species, have reduced longleaf loblolly (Pinus taeda L. [Pinaceae]) deposits such as the fall line sand- dominated areas to less than 5% of and shortleaf (Pinus echinata P. Mill. hills, where the Piedmont meets the its original range. My paper discusses [Pinaceae]) pine and hardwoods on upper coastal plain; the central ridge the habitat and history of this once an additional 14.6 million ha (36 of Florida and rolling sandhills of vast resource, outlining its key role million ac) in a band along the lower Mississippi; as well as central as an integral part of native plant northern portion of its range (Frost Louisiana and east Texas (Stout and communities. I also focus on the 1993). Marion 1993). Longleaf pine even more recent recognition of the My objective is to give a better extended onto the mountain slopes ecological importance of longleaf appreciation for longleaf pine and its and ridges of Alabama and north- pine ecosystems. This appreciation, role as a native plant in many of the west Georgia, where it was found along with advances in technology communities of the South. This is growing at elevations up to 600 m and additional information, are done by highlighting the importance (1970 ft)(Boyer 1990). combining to reverse the long-term of the longleaf pine, which was the The flatwoods sites of native trend and should help ensure that key tree species in a complex of fire- longleaf are primarily sandy deposits longleaf communities remain as a dependent ecosystems long native to that formed in shallow seas (Stout viable and valuable part of the the southeastern US (Figure 2). It and Marion 1993). Because of South’s heritage. was found on a variety of sites from abundant rainfall and slow surface wet coastal flatwoods to dry moun- drainage, flatwoods sites are KEYWORDS: restoration, wiregrass, tain ridges where it formed the seasonally wet with water at or above fire, bluestem

canopy over highly diverse ground the surface at least part of the year; 43

SPRING 2000 2000 SPRING SPRING NOMENCLATURE: (plants and cover communities. An understand- usually during the dormant season, animals) ITIS (1998); (fungi) ing of the decline of longleaf systems from December to February. Tainter and Baker (1996) is conveyed through a brief history However, during the spring and of the use of this once vast resource. early summer dry period, these The recent interest in increasing sandy sites can become quite

VOLUME 1 • NUMBER 1 Figure 1 • The natural range of longleaf pine (modiﬁed from Little 1971).

droughty. Longleaf pine is able to longleaf sites, producing the best 1920 when the great forests of tolerate such fluctuations and once growth and largest trees. longleaf pine were cleared from the formed pure stands across most of Carolinas to Georgia and Florida, the coastal plain (Ware and others History across Alabama and Mississippi, into 1993). Much of the former longleaf The history of longleaf pine is Louisiana and finally Texas. In 1896, occurred on sandhills where deposits intertwined with the history of logging yielded 8.6 million m3 (3.7 of coarse sands with hilly topogra- European settlement of the South. billion board feet) of longleaf pine phy were quite droughty. Mountain Naval stores, which began with the timber (Mohr 1896). Production longleaf forests also occupied first settlements in Virginia (Frost reached its peak in 1907, when 30.6 droughty, but rocky sites. The 1993), were so called because the million m3 (13 billion board feet) species also tended to be more pine tar produced was first used were cut (Wahlenberg 1946). By prevalent on the drier south– and extensively for wooden sailing ships 1930, nearly all old-growth longleaf west–facing slopes. to seal cracks and preserve ropes and had been harvested. Although able to dominate poor sails. From Virginia, the naval stores It seemed that longleaf pine, a sandy and rocky sites, longleaf also industry moved into North Carolina tree so well adapted to the southern occurred on better soils. Embedded then expanded south and west, region and covering such vast areas in the coastal plain, especially on the eventually to the limits of the would surely replace itself on inner and higher terraces, are longleaf range in east Texas. Many harvested areas, but it usually did deposits of finer-sized clay and silt acres of longleaf were destroyed not. Mohr (1888) concluded that that developed soils that are less because wildfires would readily the prospect of maintaining longleaf 44 droughty and more fertile. Hills with ignite the pitch soaked boles left forests seemed hopeless. Although clay parent material, such as the Red after a naval stores operation. smaller and poor quality trees that Hills area of southern Georgia, also Lumbering also began with the first were capable of producing seed were had extensive longleaf forests. Other European settlements; trees were cut left on most areas (Wahlenberg upland sites with better soils were with axes to build log structures 1946), many sites did not regenerate found from North Carolina to Texas. (Croker 1987). The boom years of

SPRING 2000 These were the most productive southern lumbering were 1880 to see Longleaf Pine on page 47

NATIVE PLANTS JOURNAL

45 SPRING 2000 SPRING

VOLUME 1 • NUMBER 1 46

SPRING 2000

NATIVE PLANTS JOURNAL Longleaf Pine from page 44

to longleaf pine. Along the Atlantic coastal plain of the Carolinas, the more competitive loblolly pine captured many sites following harvest of longleaf pine. On other flatwoods areas further south, slash pine (Pinus elliottii Engelm. [Pin- aceae]) replaced longleaf, while hardwoods and shortleaf pine became dominant on many of the upland areas of the interior. The longleaf pine’s irregular seed production—it has good seed years only every 5 y or more (Boyer 1990)—made re-establishment uncertain. In some areas a good seed crop may occur only once every 20 y. Also, because seeds are large, a number of insects, rodents, and birds eat them (Wahlenberg 1946). With such long intervals between seed crops and seed predation, it is not surprising that other more prolific seed producing pines became dominant on many sites. Even where pre-existing longleaf seedlings survived logging opera- tions, they often did not survive heavy feeding by feral hogs (Schwarz 1907). The settler’s free-ranging and feral hogs not only consumed the large longleaf seeds, but, even worse, they would root out young seedlings to get at the starch filled longleaf roots. Many acres of potential longleaf pine forest were thereby lost. The destructive potential was demonstrated by early fencing studies, which showed that only 20 seedlings per ha (8 per ac) survived in unfenced areas, while fenced sites contained over 14,826 seedlings per ha (6000 per ac) (Mattoon 1922). Harvest of second-growth stands began in earnest in the late 1940s. Because planted longleaf pine seedlings had very poor survival and perceived slow early growth because of the grass stage, it was seldom 47 selected for reforestation. Therefore, 2000 SPRING many second-growth longleaf stands were clear-cut, mechanically site prepared, and converted to planta- tions of loblolly or slash pine. On some private lands, owners made no

VOLUME 1 • NUMBER 1 effort to reforest harvested sites. Many such sites were captured by loblolly or slash pine also, from seedlings in place or seed dispersed from adjoining uncut trees. Old fields also were planted with loblolly or slash pine, or were colonized by these more aggressive pines following abandonment. As a result of cumulative impacts wrought by 3 centuries of land use, longleaf pine, once one of the most prevalent forest types in the South, has declined dramatically. By 1900, logging, naval stores, and agriculture had reduced the area dominated by longleaf pine by more than half (Frost 1993)(Figure 3). By 1935, the Figure 2 • Longleaf pine and other flora after a spring burn in the Apalachicola National Forest original longleaf forest had been cut; in Florida. only scattered fragments remained. Second-growth longleaf stands moisture content in the spreading hardwoods because it is able to occupied one-third of the species’ needle arrangement of grass stage tolerate frequent surface fires. Both original range (Wahlenberg 1946). seedlings tends to cool flames, loblolly and slash pine are capable of Conversion to other species and loss furnishing protection for the central growing on many sites historically to urban expansion continued to bud during fast-moving surface fires. dominated by longleaf and produce reduce the longleaf area through After a fire, seedlings can quickly more seeds on a more regular basis. 1985 (Kelly and Bechtold 1990). grow new needles from the un- Thus, they can out compete longleaf Over the next decade, longleaf pine harmed central meristem. Because by flooding areas with faster growing was reduced to less than 5% of its there is no stem, there is very little seedlings. However, loblolly and original area (Outcalt and Sheffield exposure of cambium that would be slash pine seedlings are thin-barked 1996). susceptible to damage in a thin and quite susceptible to fire-caused barked young seedling. When mortality. Longleaf pine usually Fire and Community sufficient root reserves have accumu- dominated sites where fire was Composition lated, seedlings achieve a height of 1 frequent. With longer fire-return Key to the success of longleaf pine is to 2 m (3.2 to 6.6 ft) in a few years. intervals, slash and loblolly pine its adaptation to fire. Longleaf pine Such rapid growth puts the terminal seedlings can grow large enough to ecosystems are fire-shaped and fire- bud beyond the reach of most survive surface fire. Therefore these maintained. Before landscape surface fires. species tended to be found on wetter fragmentation, natural fire occurred Adaptation to fire is evident areas where fires occurred at intervals every 2 to 8 y across the species’ throughout the life of longleaf pine. of 10 y or more. Slash pine was range (Christensen 1981; Abraham- The bark is relatively thick and typically found along the margins of son and Hartnett 1990; Ware and protects the cambium from lethal cypress strands and ponds (Abraha- others 1993). Lightning is a frequent heating by surface fires (Wahlenberg mson and Hartnett 1990), while occurrence across the South and 1946). It tends to naturally prune, loblolly most often occupied wetter historically was the ignition source thus providing a clear bole between riverside sites (Mohr 1896). for fires that shaped the vegetation of the crown and surface fuels. This Variation in weather and topogra- the region (Komarek 1964; Robins prevents fires from traveling easily phy and thus fire return, resulted in and Myers 1992). Native Americans into crowns where damage would be a transition zone of mixed pine augmented those effects by using fire more severe. Longleaf also favors species, especially along the northern to manage vegetation. regeneration in open areas, rather fringe of the longleaf range (Frost 48 Longleaf pine dominated much than under parent trees (Brockway 1993). In this transition zone, relief of the South because it was better and Outcalt 1998); keeping ladder is greater and thus aspect becomes able to tolerate fire than its competi- fuels away from crowns of adult more important. Pure stands of tors. Longleaf has evolved the trees. longleaf tended to occupy drier seedling grass stage, where root Longleaf pine has a competitive south facing slopes and ridges. More growth is favored and the top advantage over other southern pines mesic sites were often a mixture of

SPRING 2000 remains a tuft of needles. A higher like slash and loblolly, as well as longleaf, loblolly, and shortleaf pine

NATIVE PLANTS JOURNAL along with hardwoods such as oak. Fire frequency was likely less in this more dissected northern portion of the longleaf range, which would favor mixed stands (Frost 1993). Young hardwoods are also quite susceptible to top kill by fire; and frequent fires kept hardwood sprouts at low stature in longleaf stands (Komarek 1977; Landers and others 1990). Occasionally, random Hectares in thousands variation in fires or protective micro- site conditions allowed a hardwood stem to survive a few fires and become large enough to resist future surface fires (Rebertus and others 1993). Scattered hardwood trees occurred then, in the canopy or subcanopy of longleaf-dominated stands (Greenberg and Simons 1999). In the absence of fire, hardwoods were able to quickly emerge from the understory and form a dense midstory that could 60 27 20 12 7 5 4 3 shade out herbaceous species and 1700 1900 19 3 5 195 5 1965 1975 1985 19 95 longleaf seedlings. Some have argued Year that longleaf not only needs fire for site domination, but that it actually Figure 3 • Change in forest area dominated by longleaf pine since European settlement. Based on data perpetuates frequent surface fire from Wahlenberg (1946), Kelly and Bechtold (1990), Frost (1993), Stout and Marion (1993), and Outcalt and Sheffield (1996). Multiply ha by 2.5 to obtain acres. through the production of long flammable needles, that as litterfall, promote the spread of frequent sampled a number of relatively (5495 to 6993 lb/ac) in 3 to 4 y surface fire (Landers 1991). undisturbed dry sandhills sites where (Parrott 1967). Lightning-caused Another important component of wiregrass was a co-dominant with fires can spread quickly through this the fuel matrix in longleaf stands piney woods dropseed (Sporobolus fine-fuel matrix (Abrahamson and were the grasses. While longleaf pine junceus (Beauv.) Kunth [Poaceae]). I Hartnett 1990). Wiregrass therefore dominated the overstory of southern also have sampled flatwoods sites tended to shorten the fire-return ecosystems, grasses dominated the where Curtis’ dropseed (Sporobolus interval, which favored longleaf pine understory. In the eastern portion of curtissii (Vasey ex Beal) Small ex domination. the longleaf range, wiregrass Scribn. [Poaceae]), a bunch grass Grass- and pine needle-fueled (Aristida stricta Michx. [Poaceae] and with a growth form similar to fires also control brown spot needle Aristida beyrichiana Trinius & wiregrass, dominated the understory. blight (Mycospaerella dearnessii Barr Ruprecht [Poaceae]) was the major The leaves of these grasses [Loculoascomycetes]), which can understory species (Figure 4). typically arch outward from the severely limit growth and survival of However, because it was not the center of the bunch and overlap with longleaf seedlings (Boyer 1975). In dominant grass in all longleaf pine adjoining individuals. Wiregrass addition, most biomass is below communities, wiregrass is not an leaves are short-lived; 85% die ground, 60% to 80% for wiregrass essential component of the system. within 12 mo of formation (Parrott (Parrott 1967), where high root In the Gulf coastal region, little 1967). Dead wiregrass leaves remain turnover adds significant amounts of bluestem (Schizachyrium scoparium attached to the plant (Landers 1991) organic matter to the soil. Thus, a (Michx.) Nash [Poaceae]) was the and decay quite slowly (Christensen wiregrass understory maintains a 49 dominant native grass (Frost 1993). 1993). Living and dead wiregrass better soil environment by improv- 2000 SPRING Even within its major range, wire- leaves intercept the shed needles of ing soil structure and water- and grass was not always the dominant overstory pines, causing an accumu- nutrient-holding capacity (Snedaker grass on all sites. A number of other lation of dead biomass in a very and Lugo 1972). grass species native to the area must flammable configuration. These fuels Although longleaf dominated the have dominated some sites. I have reach a peak of 6160 to 7840 kg/ha overstory and grasses the understory,

VOLUME 1 • NUMBER 1 these communities contained many and several members of the Aster- listed at least 191 taxa of vascular other plant species. Common woody aceae: deer tongue (Carphephorus plants that are endemic to or exist associates in the flatwoods were saw Cass. spp.), dog fennel (Eupatorium largely in longleaf communities. palmetto (Serenoa repens (Bartr.) L. spp.), grass-leaved gold aster Walker (1993) reported 96 plant Small [Arecaceae]), gallberry (Ilex (Pityopsis graminifolia (Michx.) species as local endemics associated glabra (L.) Gray [Aquifoliaceae]), Nutt.), gold aster (Chrysopsis with longleaf pine ecosystems. wax myrtle (Morella cerifera (L.) gossypina (Michx.) Ell.), asters (Aster Not surprisingly, there are a Small [Myricaceae]), and runner L. spp.), and blazing stars (Liatris number of animal species that oaks (Quercus minima (Sarg.) Small Gaertn. ex Schreb. spp.). Woody depend on the longleaf ecosystem & Q. pumila Walt. [Fagaceae]) (Peet associates in mountain longleaf for much of their habitat, including and Allard 1993). Composites were communities included blackjack oak the red-cockaded woodpecker often the dominant forbs. Other and shortleaf pine (Harper 1905). (Picoides borealis Vieillot) and the common herbaceous species Herbaceous species were mostly the gopher tortoise (Gopherus polyphe- included meadow beauty (Rhexia L. same as occurred on dry sandhills mus Daudin). The tortoise is spp. [Melastomataceae]), beakrush sites, including grass-leaved gold especially critical because its burrows (Rhynchospora Vahl spp. [Cyper- aster, sweet goldenrod (Solidago provide homes for many secondary aceae]), and yellow-eyed grass (Xyris odora Ait. [Asteraceae]), bracken fern users from snakes to insects (Speake L. spp. [Xyridaceae]). Wet savannas (Pteridium aquilinum (L.) Kuhn 1981; Jackson and Milstrey 1989). contained a mix of herbaceous [Dennstaedtiaceae]), and sunflowers. The longleaf-grass systems also are species, including insectivorous In the mountain longleaf stands in vital to the maintenance of a number pitcherplants (Sarracenia L. spp. northern Alabama, Mohr (1901) of embedded ecosystems that occur [Sarraceniaceae]) and sundews noted a number of legume species across the South (Landers and others (Drosera L. spp. [Droseraceae]), including beggar lice, bush clover 1990). Many of these communities numerous orchids, and showy, (Lespedeza Michx. spp. [Fabaceae]), require periodic fire to maintain flowered composites like sunflowers and goat rue (Tephrosia virginiana structure and health (Kirkman and (Helianthus L. spp. [Asteraceae]). (L.) Pers. [Fabaceae]). others 1998). Fire begins in the Upland longleaf stands had a very Although some herbaceous longleaf-grass type and then spreads lush understory that contained many species were found on most sites of into adjoining habitats such as herbaceous species, including a similar nature across the longleaf seepage slopes, canebrakes, treeless number of composites and legumes range, there was a lot of variation in savannas, and sand pine scrub. (Peet and Allard 1993). Common the understory community. Thus, Without periodic fire, these commu- woody species were blackjack oak even though it is commonly referred nities, like the longleaf systems, (Quercus marilandica Muenchh. to as the longleaf ecosystem, it is not change in ways that make them less [Fagaceae]), sand-post oak (Quercus uniform. Peet and Allard (1993) suitable to the plants and animals margarettae Ashe [Fagaceae]), and divided the longleaf ecosystem into 4 that have evolved with fire. persimmon (Diospyros virginiana L. major units (xeric, subxeric, mesic, [Ebenaceae]). On sandhills sites, and seasonally wet) based on soil Longleaf Importance turkey oak (Quercus laevis Walt. moisture. They further divided these Because of continued losses, habitat [Fagaceae]) is such a common classes into 23 communities on the reduction became a cause for woody associate of longleaf pine that basis of geographic location and concern among natural resource and many have referred to it as the physiographic province. Each conservation organizations in the longleaf-turkey oak association extremely diverse community has its South. Longleaf pine communities (Stout and Marion 1993). Other own characteristic plant species. The are extraordinarily diverse and common woody associates include diversity of ground cover plants per because they once covered such a bluejack oak (Quercus incana Bartr. unit area makes longleaf pine large area, many plants and animals [Fagaceae]), persimmon, dwarf ecosystems among the most species- adapted to the habitat they provided. huckleberry (Gaylussacia dumosa rich plant communities outside the There is also growing evidence that (Andr.) Torr. & Gray [Ericaceae]), Tropics. Peet and Allard (1993) ranges of certain amphibians and and blueberries (Vaccinium L. spp. reported finding as many as 140 reptiles coincide with the longleaf [Ericaceae]). The understory on species of vascular plants in a ecosystem (Guyer and Bailey 1993). 50 these dry sites often can appear to be 1000 m2 (10,772 ft2) area and in About 10% of all arthropods found a sea of grass, especially where many longleaf communities, equally in longleaf ecosystems are endemics wiregrass is dominant. However, impressive counts of more than 40 (Folkerts and others 1993). It has many other species grow among the species in a m2 (10.8 ft2). Many of become apparent that if the decline grass clumps in these communities. these species are restricted to or are continues there will be many more Common herbs include beggar lice found principally in longleaf endangered and threatened species

SPRING 2000 (Desmodium Desv. spp. [Fabaceae]) habitats. Hardin and White (1989) needing special management. A

NATIVE PLANTS JOURNAL growing realization of the importance of the longleaf forest type has fostered committed efforts to restore and manage these ecosystems (see Kush 1998, 1999). Habitat protection is only 1 reason for the growing interest in longleaf pine. Early on, some industrial owners recognized the potential of longleaf pine (Croker 1987). Its many advantages have been well documented by Landers and others (1995), including the ability to produce as much or more fiber than other southern pines on dry, sandy sites (Outcalt 1993). Longleaf pine is less susceptible than other southern pines to damage and mortality from insects, like bark beetles, and diseases such as fusiform rust (Cronartium quercuum (Berk.) Miyabe: Shirai f. sp. Fusiforme Burds. et Snow [Melampsoraceae]). It also is very versatile, producing Figure 4 • Longleaf pine with a flowering wiregrass understory on a sandhills site in the Ocala high-value products like poles and National Forest in Florida. peeler logs. Poles are the highest value wood product taken from southern timberlands, and longleaf the payment the landowner gets per ate longleaf following harvest, stands can yield from 30% to 80% acre, is higher for longleaf pine to thereby conserving intact longleaf poles (Boyer and White 1990). offset the higher cost of seedlings. habitat where it does exist. This Longleaf forests also produce pine Recent changes in the US Depart- would reverse a long-term trend of straw, which has both wholesale and ment of Agriculture (USDA) converting to other species following retail landscaping markets (Williston Conservation Reserve Program also harvest. The flow of information and others 1990). encourage landowners to plant from organizations like the Longleaf Even some private landowners longleaf on areas enrolled in the Alliance (for example, Franklin who hold only small parcels of land program by designating longleaf 1997) is also increasing awareness are considering longleaf pine when plantings as priority areas. Areas that and helping bolster the visibility of choosing a species for their sites. would be planted to longleaf are longleaf pine, which may encourage Improved techniques for growing, given preference for enrollment. The landowners to choose longleaf pine handling, and planting longleaf Natural Resource and Conservation for their sites. seedlings are not just raising survival Service, a branch of USDA that Because the habitat is good for rates to levels comparable to other handles enrollment, recently white-tailed deer (Odocoileus pines; they also are decreasing time announced that over 41,310 ha virginianus Zimmermann) and seedlings are in the grass stage, (102,000 ac) of cropland in the bobwhite quail (Colinus virginianus thereby increasing growth rates South will be planted with longleaf L.), longleaf pine forests can also (Barnett and others 1990). Longleaf pine. produce income from hunting leases. also can be managed using the Planting longleaf pine will not Larger, private landowners have long shelterwood (Boyer and White ensure re-establishment of the recognized and valued such opportu- 1990) or the all-aged selection longleaf ecosystem, because the nities. Many hectares of the longleaf systems of harvest and regeneration diversity exists in the understory type exist because of the hunting (Farrar and Boyer 1991) as well as community. However, even old field provided in addition to timber 51 the even-aged clear-cut system. plantings furnish some of the production. This is only one aspect 2000 SPRING Assistance programs, which pay a functions of a complete system. of the longleaf tradition in southern portion of the cost of planting trees, Also, such sites serve as areas culture. Many longtime southerners are available that encourage land- demonstrating the benefits of remember longleaf pine from an owners to reforest sites. In North longleaf pine. This should encourage earlier era. Even though much of the Carolina, the cost share rate, that is others to plant or naturally regener- region was converted to other pines,

VOLUME 1 • NUMBER 1 Farrar RM, Jr., Boyer WD. 1991. Managing home sites and cities still contain References longleaf pine trees from the original longleaf pine under the selection system – Abrahamson WG, Hartnett DC. 1990. Pine promises and problems. In: Coleman SS, or second growth forest. Old flatwoods and dry prairies. In: Myers RL, Neary DG, editors. Proceedings of sixth longleaf can be seen in neighbor- Ewel JJ, editors. Ecosystems of Florida. biennial southern silvicultural research hoods from Wilmington, North Orlando (FL): University of Central Florida conference; 1990 Oct 30–Nov 1. Carolina, to Valdosta, Georgia, to Press. p 103–149. Memphis, TN. New Orleans (LA): USDA Barnett JP, Lauer DK, Brissette JC. 1990. Gulfport, Mississippi, and in most Forest Service, Southeastern Forest Regenerating longleaf pine with artificial Experiment Station. General Technical other southern cities and towns as methods. In: Farrar RM, editor. Proceed- Report SE-70. p 357–368. well as around older rural home ings of the symposium on the management Folkerts GW, Deyrup MA, Sisson DC. 1993. sites. of longleaf pine; 1989 Apr 4–6; Long Arthropods associated with xeric longleaf Beach, MS. New Orleans (LA): USDA pine habitats in the southeastern United Summary Forest Service, Southern Forest Experiment States: a brief overview. In: Hermann SH, Station. General Technical Report SO-75. editor. Proceedings of the 18th Tall Healthy longleaf pine ecosystems are p 72–-93. Timbers fire ecology conference. The aesthetically pleasing. The tall pines Boyer WD. 1975. Development of brown-spot longleaf pine ecosystem: ecology, with clear boles for half or more of infection in longleaf pine seedlings stands. restoration and management; 1991 May the total tree height are topped with New Orleans (LA): USDA Forest Service, 30–Jun 2; Tallahassee, FL. Tallahassee Southern Forest Experiment Station. a spreading crown of long needles, (FL): Tall Timbers Research Station Research Paper SO-108. 10 p. 18:159–192. making the trees stately and pictur- Boyer WD. 1990. Longleaf pine. In: Burns Franklin RM. 1997. Stewardship of longleaf esque. The profusion of flowers after RM, Honkala BH, technical coordinators. pine forests: a guide for landowners. growing season burns, followed by a Silvics of North America. Volume 1, Andalusia (AL): Auburn University, Solon waving sea-like cover of grasses, is Conifers. Washington (DC): USDA Forest Dixon Forestry Education Center. Longleaf Service. Agriculture Handbook 654. p also visually appealing. Managing Alliance Report Number 2. 44 p. 405–412. Frost CC. 1993. Four centuries of changing and restoring longleaf pine systems is Boyer WD, White JB. 1990. Natural landscape patterns in the longleaf pine now recognized as appropriate for regeneration of longleaf pine. In: Farrar ecosystem. In: Hermann SH, editor. conserving species, generating RM, editor. Proceedings of the symposium Proceedings of the 18th Tall Timbers fire economic returns, and creating on the management of longleaf; 1989 ecology conference. The longleaf pine Apr 4–6; Long Beach, MS. New Orleans pleasing visual landscapes. This ecosystem: ecology, restoration and (LA): USDA Forest Service, Southern Forest management; 1991 May 30–Jun 2; realization, coupled with cultural ties Experiment Station. General Technical Tallahassee, FL. Tallahassee (FL): Tall and improved knowledge gained Report SO-75. p 94–113. Timbers Research Station 18:17–44. from research and adaptive manage- Brockway DG, Outcalt KW. 1998. Gap-phase Greenberg CH, Simons RW. 1999. Age, ment, along with better dissemina- regeneration in longleaf pine wiregrass composition, and stand structure of old- ecosystems. Forest Ecology and tion of information are leading to a growth oak sites in the Florida high pine Management 106:125–139. landscape: implications for ecosystem concerted effort to maintain and Christensen NL. 1981. Fire regimes in management and restoration. Natural restore longleaf pine communities. It southeastern ecosystems. In: Mooney HA, Areas Journal 19:30–40. is hoped this will reverse the long- Bonnicksen TM, Christensen NL, Lotan JE, Guyer C, Bailey MA. 1993. Amphibians and term loss of the longleaf pine type. Reiners WA, technical coordinators. reptiles on longleaf pine communities. In: Proceedings, fire regimes and ecosystem Thus, although longleaf pine will Hermann SH, editor. Proceedings of the properties; 1978 Dec 11–15; Honolulu, 18th Tall Timbers fire ecology conference. not be re-established on most of the HI. Washington (DC): USDA Forest The longleaf pine ecosystem: ecology, areas it once occupied, it should Service, Washington Office. 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USDA Forest Service, System. 1998. Biological names. Version 320 Green Street Southern Region. Forestry Report R8-FR7. 4.0 (on-line database). URL: http:// Athens, GA 30602 37 p. www.itis.usda.gov/plantproj/itis/ 52 Outcalt_Kenneth/srs_athens itis_query.html (updated 15 December @fs.fed.us 1998). Jackson DR, Milstrey ER. 1989. The fauna of gopher tortoise burrows. In: Diemer JE, editor. Proceedings of the gopher tortoise relocation symposium. Tallahassee (FL): Florida Game and Fresh Water Fish Commission. p 86–98.

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