Remnant OldGrowth Longleaf Pine (Pinus palustris Mill.) Savannas and Forests of the Southeastern USA: Status and Threats J. Morgan Varner, III
CONSERVATION ISSUES ABSTRACT: Old-growth savannas and forests dominated by longleaf pine (Pinus palustris Mill.) are of great conservation and research interest. Comprehensive inventories of old-growth communities,' however, are lacking for most of longleaf pine's natural range. We searched the literature, interviewed regional experts, queried email discussion lists and newsletters, and made on-site inspections to com pile a revised list of old-growth longleaf pine stands, their acreage and status. We found 15 old-growth longleaf pine stands covering 5095 ha in only four of the nine states in the natural range of longleaf • pine. Our area estimate represents 0.004% of extant acreage and 0.00014% of the presettlement extent. Further, some community types (e.g., Piedmont, West Gulf Coastal Plain provinces) have no extant old-growth longleaf pine reference sites. Old-growth stand ownerships are diverse, with large tracts (> Remnant Old 400 hal in both public and private ownerships. Half of the remnants face serious management threats, most due to difficulty in managing fire in the contemporary southeastern landscape and tenuous land tenure. In light of the shrinking nature of this resource, statewide searches, restoration, and conservation Growth Longleaf should continue in earnest.
Pine (Pinus palustris Index terms: conservation, fire exclusion, old-growth forests, Pinus palustris, restoration Mill.) Savannas INTRODUCTION Hermann 1993, Landers and Boyer 1999, and Forests of the Varner et al. 2003b). The structure of Ecosystems dominated by longleaf pine old-growth pinelands is typified locally Southeastern USA: (Pinus palustris Mill.) blanketed a wide by patchiness and, at larger scales, by an range of landscapes within the Atlantic inverse exponential diameter distribution and Gulf Coastal Plains, Piedmont, Ridge (Figure 2; Reed 1905, Schwarz 1907, Status and Threats and Valley, Cumberland Plateau, and Blue Forbes 1930, Wahlenberg 1946, Platt et Ridge Physiographic Provinces of the al. 1988, Varner et al. 2003b). Old-growth southeastern USA, perhaps covering as pinelands contain many age classes, with J. Morgan Varner, 1111 much as 37 million ha prior to European peaks of individuals exceeding 200 y (Platt School of NaturalResources settlement (Frost 1993). Pristine longleaf et al. 1988, Platt and Rathbun 1993, Varner & Environment savannas and forests (hereafter "pine et al. 2003b). University of Florida lands") were dominated by an open to Gainesville, FL 32611 USA closed (basal area ranged from 12 to 35 Old-growth characteristics provide re m2 ha-1; stand density ranged from 130 searchers and managers with: 1) bench John S. Kush to 400 trees ha-1), patchily distributed, marks for natural area management (Noel uneven-aged longleaf pine canopy (Figure et al. 1998); 2) valuable data to reconstruct School of Forestry and Wildlife Sciences 1; Reed 1905, Schwarz 1907, Chapman climatic and disturbance events (West et Auburn University, AL 36849 USA 1909, Forbes 1930, Wahlenberg 1946). The al. 1993, Meldahl et al. 1999, Varner et overstory of virgin longleaf pinelands con- . al. 2003b); 3) among the highest qual sisted of individual pines with diameters up ity habitats for native flora and fauna • to 100 cm, and heights up to 40 m (Reed in existence (Engstrom 1993, Engstrom 1905, Chapman 1909). Large snags aver and Sanders 1997, Varner et al. 2003a, aged 15 ha- 1 (Schwarz 1907), and downed Hermann et al. In press); and 4) valued large coarse woody debris averaged 5.7 ha-1 aesthetic and cultural resources. To the de (Hermann 1993). These conditions were gree researchers and managers understand maintained by low-intensity, low-severity these characteristics, we are limited in our surface fires occurring every one to five management, research, and conservation of years (Chapman 1932). all pinelands landscapes (sensu Swanson and Franklin 1992). Old-growth longleaf pinelands have sev eral characteristics distinguishing them While many imperIled ecosystems have 1 Corresponding author: Box 118526, University of Florida, from mature pinelands. Old-growth stands been inventoried (Davis 1996), old-growth Gainesville, FL 32611-8526 USA; contain many large pines (> 50 cm dbh) longleaf pine acreage, until recently, has 352-392-1891; fax: 352-392-3993; with only a few primary branches and been unknown. Means (1996) conducted [email protected] substantial percentages of woody biomass a survey of managers, scientists, and con in heartwood (Wahlenberg 1946). Stands servationists and found 14 stands, covering Natural Areas lournaI24:141-149 consist of persistent large snags and large ca. 3902 ha. For all longleaf pinelands, downed woody debris (Schwarz 1907, approximately 97% is lost (Outcalt and
Volume 24 (2),2004 Natural Areas Journal 141 Sheffield 1996) and 99.99% is no longer old-growth. These results, added to al ready dismal reports of pinelands decline, prompted Noss and others (1995) to list longleaf pine ecosystems as the second most endangered ecosystem in the United States. This information helped promote efforts in both conservation and investi gation of the extant old-growth longleaf pine stands.
Longleaf pinelands are significant and increasingly rare elements of southeastern USA biodiversity. Pineland floral species richness is unparalleled outside of the trop ics, and includes many rare, endemic, and carnivorous species (Folkerts 1982, Hardin and White 1989, Peet and Allard 1993). Pinelands fauna includes a high diversity of reptiles and amphibians (Guyer and Bailey 1993), avian species (Engstrom 1993), and greater than 5000 invertebrate species (Folkerts et al. 1993). These facts, and their role in propagating fire across many fire-dependent plant communities, Figure 1. Typical historical photograph of a park-like virgin stand of longleaf pine on Great Southern Lumber Company land near Lake Ponchartrain, Louisiana, ca. 1920. Pinelands like these were intensively have lead many to consider longleaf pine utilized for lumber, naval stores, grazing, and agriculture following European settlement. Photograph lands as a regional landscape keystone with permission from the George Jason Streator Papers, Department of Special Collections and Archives, (Noss 1989, Means 1996). The decline Kent State University Libraries. and subsequent rarity of longleaf pinelands began with 18th century turpentining, log ging, burning cessation, development, and 50 ~------~ site conversion (Croker 1987, Frost 1993). Far from diminishing, longleaf pinelands decline was> 27% over the most recent 40 decade (1985 to 1995) of federal inventory (Outcalt and Sheffield 1996). ~ (tI t) a> 30 Rationale I.... a> In 1997, we conducted informal visits to c.. the old-growth longleaf pine stands listed en a> 20 in Means (1996). Several sites listed by a> .... Means were either in serious management I-- predicaments, not old, or had since been 10 logged. Because the list compiled by Means was brief and informal, we were encour aged to gather more information from regional managers and scientists and visit known and potential old-growth stands. 5 10 15 20 25 30 35 40 45 50 55 60 65 70 75 80 85 We initiated this project to address these Diameter Class (em) shortcomings and to assess management dilemmas in all extant old-growth longleaf pinelands. Figure 2. Diameter distributions (+1- 95% confidence intervals) of 14 old-growth longleaf pine stands taken from published data in Schwarz 1907, Forbes 1930, Varner et al. 1999, and Varner et al. 2003b. Old-growth longleaf pinelands contain many large pines (> 50 cm DBH) and many age classes, with peaks of individuals exceeding 200 years old.
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METHODS For each old-growth stand, we listed its fires often result from these restoration location .( county, state), tract size, owner fires, killing large percentages of remnant We focused our survey in three areas: ship (public, private entity), management pines (Kush et al. 1999b, Gordon and personal contacts, library research, and regime (harvesting, burning regime), and Varner 2002). These so-called "duff fires" field visits. We contacted land managers, threats to site integrity. Biological and are problems in other North American conservationists, landowners, and scientists ecological characteristics were also listed coniferous forests (e.g., Swezy and Agee via telephone, email, the Longleaf Alliance for each stand, including significant species 1991, Ryan and Frandsen 1991, Haase Forum
Table 1. Remnant old-growth longleaf pine acreage in 2004. See text for discussion of each site, its threats, and significance.
Site Name, State Size Ecosystem Ownership Intact Intact Protected Fire (ha) Type(s)" Flora Fauna LandscaEe R'eglme b Flomaton Natural Area, AL 23 CLH Private/ MOD R Mountain LongleafNWR, AL 45 MT PubliC/NWR X XC M,R Big Pine Tract, FL 170 SH&CLH Public X R Eglin Air Force Base, FL 3650 SH&FW Public X X X M,R Goethe State Forest, FL 75 FW Public X X X M Platt Branch, FL 160 FW Public X X X M Venus Flatwoods, FL 40 FW TNC X M Greenwood Plantation, GA 200 CLH&SV TNC X X X 'M Moody Tract, GA 120 CLH&SH TNC X X R Wade Tract, GA 83 CLH Private (Easement) X X X M Red Hills Plantations, GA 400 CLH Private (Easements) X X X M,R Bonnie Doone, NC 65 CLH Public ? Boyd Tract, NC 24 CLH&SH Public R Camp Lejeune, NC 20 SH Public X X X M Croatan Sandhill, NC 20 SH Public X X X M Total old-growth longleaf Eine acreage 5095 ha
a Ecosystem types are as follows, CLH= clayhills & loarnhills; SH = sandhills; FW = flatwoods; MT = mountains; SV = savannas. b Fire regime values are as follows, R = restoration phase (reintroduction underway, severity irregular); M = maintenance phase (frequent regime, severity low). CSee text for discussion ofthe precarious protection status of Mountain Longleaf National Wildlife Refuge's wildlands.
Volume 24 (2), 2004 Natural Areas Journal 143 " "" ."
Figure 3. Locations of remnant (2004) old-growth longleaf pine stands in the southeastern USA. Shaded areas represent the historic range of longleaf pine, with significant physiographic provinces labeled. Map numbers represent approximate stand locations, corresponding to descriptions in the text.
Mountain Longleaf National Wildlife pany) prescribed fire and hardwood harvesting be Refuge, Alabama). Ecosystem 'TYpe: Clay-Loamhill gan in 1994. Active education, restoration, and science programs are in place (Zutter The Flomaton Natural Area is a classic et al. 2003). The current ownership protects Extant old-growth longleaf pine example of a fire-excluded pineland. The the stand, and management is guided by stands 23 ha site contains many large, old pines a Memorandum of Understanding (MOU) 1. Flomation Natural Area (Esc ambia (maximum age> 300 y) with heavy forest among several public and private partners. County, Alabama) floor accumulations (Varner et al. 2000a). Current threats include non-native species, Owner: Private (International Paper Com- An aggressive restoration program utilizing duff fire danger, Wildland-Urban Interface
144 Natural Areas Journal Volume 24 (2), 2004 (WUI), and uncertain ownership future degraded from decades of fire exclusion. Air Force) (Figure 4). An active restoration program is underway, Ecosystem types: Sandhill and Flat including hardwood harvesting and bum woods 2. Mountain Longleaf National Wildlife ing. Many longleaf pinelands-associated Refuge (Calhoun County, Alabama) species are present, including gopher tor Eglin's tracts represent the largest remain Owner: Public (U.S. Fish and Wildlife toises and numerous herbaceous species. ing old-growth landscape at ca. 3650 ha, Service/Joint Powers Authority) Big Pine Tract is one of the few extant with many high-quality pineland commu Ecosystem Type: Mountain stands within a large protected wildland, nities and intact focal animal assodates. heightening its conservation value and Noteworthy tracts at Eglin include the The recently dedicated (June 2003) Moun integrity. CUfl'ent threats include duff fire Patterson Natural Area (375 to 2031 ha tain Longleaf National Wildlife Refuge was danger and non-native species invasions. area), White Point (58 ha), Brandt Pond created from a portion of Fort McClellan, (155 ha), Field One (507 ha), Sandy Mount a U.S. Army garrison and formerly active 4. Eglin Air Force Base Tracts (Okaloosa (200 ha), and several others (pel's. comm., military installation. Military training and Walton Counties, Florida) K. Hiers, Niceville, Fla.). Estimates of the with flammable flares and explosives Owner: Public (Department of Defense, total old-growth acreage at Eglin range helped maintain several large patches of frequently burned mountain longleaf pinelands. The Refuge contains the only known old-growth longleaf pine forest in the Blue Ridge Physiographic Province. Twelve old-growth stands cover ca. 45 ha, ranging in size from 1 to > 15 ha, in the mountainous portions of the Refuge and its boundaries (Varner et aI. 2000b). Four noteworthy tracts include stands at Red tail Ridge (4 ha), Caffey Hill (8 ha), Mt. Tylo (10 ha); and the border stand along Skeleton Mountain (8 ha). These tracts have undergone varied burning regimes, from annual to extremely fire-excluded (> 40 y since fire), and contain the only known frequently bumed old-growth stands in the Mountain Province. Stands contain species-rich groundcovers (Vamer et al. 2003a) and patchy old (trees> 230 y) pine canopies (Vamer et al. 2003b). The pres ent boundaries of the Refuge omit three old-growth longleaf pine stands: Skeleton Mountains (under consideration), Ford Hill (under consideration), and Reynolds Hill (unlikely for inclusion); this uncer- . tain ownership future is among the acute threats to these mountain pinelands. Other CUfl'ent threats include duff fire danger and WUI issues.
3. Big Pine Tract (Hernando County, Florida) Owner: Public (Florida Fish and Wildlife Conservation Commission) Ecosystem type: Sandhill and Loamhill
The Big Pine Tract is a large (170 ha), Figure 4. Extant old-growth longleaf pineland at the Flomaton Natural Area, near Flomaton, Alabama. infrequently bumed site on the Brooksville The Flomaton tract contains many of the most common threats to remnant old-growth stands: high fuel loads resulting from decades of fire exclusion, degraded groundcover species composition, presence Ridge. Many trees are young (ca. 160 y), of invasive species, highly fragmented, and located near smoke-sensitive and ever-urbanizing suburban and the groundcover communities are areas.
Volume 24 (2),2004 Natural Areas Journal 145 from our conservative 3650 ha figure (fit wildland is embedded in a large, unpro many old trees (> 250 y). Most floral and ting data gathered by a Florida Natural tected wildland complex with scattered old faunal elements remain, including abundant Areas Inventory survey; pers. comm., growth candidate stands throughout. Platt wiregrass, gopher tortoises (Gopherus C. Kindell, Tallahassee, Fla.) to areas as Branch stands are old, frequently burned, polyphemus), and a few red-cockaded large as 21,000 ha (pers. comm., K. Hiers, and contain characteristic pinelands flora woodpeckers. The large size of the tract, Niceville, Fla.), highlighting the need for and fauna, including red-cockaded wood its inclusion in a large protected landscape more formal surveys of Eglin's old-growth peckers. Acreage additions should be (ca. 1650 ha), and its location in the lower pinelands. Eglin's old-growth stands have sought from adjacent areas to maintain site Atlantic Coastal Plain make it unique in this experienced various burning regimes, from integrity. No acute threats to Platt Branch list. The Moody Tract is a recent purchase annual fire to extremely fire-excluded. were found. by The Nature Conservancy of Georgia. Intense management attention has been Duff fire dangers are the greatest threat focused on Eglin's remnant old-growth 7. Venus Flatwoods (Glades County, to Moody's stands, with substantial pine pinelands (McWhite et a1. 1999), a fact Florida) mortality in recent restoration prescribed encouraging to many conservationists Owner: Private (The Nature Conservancy bums (J.M. Varner, unpublished data). concerned for the status of southeastern of Florida) old-growth longleaf pine stands. Eglin's Ecosystem type: Flatwoods 10. Wade Tract (Thomas County, Geor stands contain very old trees; it is common gia) to encounter 400 year-old longleaf pines. Venus Flatwoods is a 40 ha, frequently Owner: Private (Tall Timbers Research Owing to its history of erratic burning, burned flatwoods site with intact pinelands Station Conservation Easement) several stands have recently experienced vegetation. The stand contains many old Ecosystem type: Clayhills mortality resulting from re-introduction pines (> 200 y) and has abundant regenera of fire. Eglin's stands face the least risk tion. Its small size, location within a dis The 83 ha Wade Tract represents the ar from urban areas, and represent the most turbed landscape, and isolation from other chetypal old-growth longleaf pine savanna. ecologically viable old-growth longleaf pinelands communities are the most acute The Wade Tract is frequently burned, con pine landscape in existence. Current threats threats to the Venus Flatwoods' stand. tains an intact pinelands fauna and flora, to Eglin's stands include duff fire danger and is the best remnant pineland landscape. and isolated WUI issues. 8. Greenwood Plantation (Thomas County, Studies from the Wade Tract frame the con Georgia) temporary view of pinelands ecology and 5. Goethe State Forest (Levy County, Owner: Private (The Nature Conservancy stand dynamics (Platt et al. 1988, Hermann Florida) of Georgia) 1993, Platt and Rathbun 1993, Grace and Owner: Public (Florida Division of For Ecosystem type: Clayhills and Savanna Platt 1995, Engstrom and Sanders 1997, estry) Hermann et a1. In press). The Wade Tract Ecosystem type: Flatwoods Often called "The Big Woods", this 200 ha faces limited threats from the WUI. site may best represent how many virgin Goethe State Forest is a large north Florida longleaf pine stands appeared at settlement. 11. Red Hills Plantations (Thomas County, pinelands landscape containing two old The site is frequently burned, contains many Georgia) growth longleaf pine flatwoods stands old trees, a diversity of plant communities, Owner: Private (Tall Timbers Research totaling 75 ha. Goethe's two tracts contain and native flora and fauna characteristic of Station and The Nature Conservancy of red-cockaded woodpecker (Picoides borea pristine pinelands. The Plantation has been Georgia Conservation Easements) lis) clusters and intact, frequently burned carefully managed for timber, bobwhite Ecosystem type: Clayhills groundcover communities. The overstory quail (Colinus virginianus), and aesthetics contains multiple age classes above 150 y, under the Stoddard-Neel Selection System The Red Hills Plantations are a collection with individuals exceeding 225 y. Isolated for 50 y (Neel 1971). Small groundcover of privately owned stands covering ca. 400 old (> 200 y) longleaf pines are common disturbance patches are scattered across ha (L. Neel, pers. comm., Thomasville, over much of Goethe's landscape. No acute the site, characteristic of quail-managed Ga.) in various states of conservation and threats to Goethe's old-growth pinelands lands. Greenwood faces limited threats management. Most sites are frequently were found. from the WUI. burned and managed for bobwhite quail and timber (Neel 1971). In combination 6. Platt Branch (Glades County, Florida) 9. Moody Tract (Appling County, Geor with Greenwood Plantation and the Wade Owner: Public (Florida Fish and Wildlife gia) Tract, the Red Hills comprise a large, high Conservation Commission) Owner: Private (The Nature Conservancy quality pinelands landscape. Conservation Ecosystem type: Flatwoods of Georgia) efforts are prioritized here to preserve this Ecosystem type: Clayhills and Sandhills magnificent, biologically significant land Platt Branch is a large south Florida scape. Dominant threats to the Red Hills pinelands landscape containing 160 ha of The Moody Tract contains an estimated Plantations are their uncertain ownership old-growth longleaf pine flatwoods. This 120 ha of old-growth pinelands, with future and limited WUI threats.
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12. Bonnie Doone Tract (Cumberland intact pinelands flora and is within a large Japanese climbing fern (Lygodium ja County, North Carolina) protected landscape. More field surveys ponicum [Thunb.] Sw.), Chinese tallow Owner: Public (City of Fayetteville, North should be made. Another old longleaf pine (Sapium sebiferum [L.] Roxb.), Chinese Carolina) stand (Catfish Lake Bay Rims) is nearby, wisteria (Wisteria sinensis [Sims] Sweet), Ecosystem type: Clayhills and may be included (pers. comm., Mike and several others. Non-natives reduce Schafale, Raleigh, NC). The only acute native species and rapidly degrade pine Bonnie Doone is a 65 ha rare stand of threat to this site is reckless Off-Road- lands. Often these stands' losses are due unturpentined (so called "round timber") Vehicle (ORV) use. ' to their proximity to the WUI, but exotics pineland in the s,andhills region of North like cogongrass (Imperata cylindrica [L.] Carolina. The site is fire-excluded and in P. Beauv.) invade large interior protected DISCUSSION need of a careful ecological restoration areas. Feral and introduced animals (do program. Threats include duff fire danger mesticated cats and dogs, wild hogs, and Summary of threats to extant old and WUI issues. nine-banded armadillos (Dasypus novem growth longleaf pinelands cinctus)) also threaten pineland plant and 13. Boyd Tract (Moore County, North Few remnant old-growth longleaf pine animal biodiversity. Carolina) lands could be described as pristine or Owner: Public (North Carolina Division approximating presettlement conditions Two sites have uncertain futures because of Parks & Recreation) (Figure 4). Acute threats to remnants of uncertain preservation and conservation. Ecosystem types: Sandhills and Clayhills include: urban interface issues (60% of The recent transfer of Fort McClellan to the extant stands), duff fire danger (47%), Mountain Longleaf National Wildlife Ref The Boyd Tract is an old (trees> 350 y non-native species invasions (20%), and uge omitted three old-growth longleaf pine old) 24 ha stand that underwent 80 y of uncertain management future (13%). Other stands and a large portion of its surrounding fire-exclusion (Gilliam et al. 1993) prior major threats include fragmentation and landscape and smokeshed from protection. to recent restoration efforts. Fire and fire groundcover degradation, which to some The privately-owned Red Hills Plantations surrogates (duff raking) have been under degree threatens all remaining sites. ' face strong development pressures from way for several years, with some areas of Tallahassee and Thomasville exurbaniza high pine mortality resulting. Turpentining Nine sites face present threats from com tion. Efforts should be heightened to protect scars are present on most trees, providing a plications associated with the Wildland these tracts with more stable conservation unique aspect and cultural history. Current Urban Interface (WUI). Dominant WUI measures (easements, purchase, further threats include duff fire danger, WUI issues, threats include smoke restrictions, burning Congressional protection). and non-native species invasions. fears, feral animal problems, exotic plant invasions, and direct development pres All sites, excepting one or two stands, have 14. Camp Lejeune Tract (Onslow County, sures. Most sites will face this threat in a history of groundcover degradation (e.g., North Carolina) the future without large landscapes being wildlife food plots, fire lines, fire exclu Owner: Public (Department of Defense, preserved. sion). Groundcover disturbances create Marine Corps) entry points for exotics, are difficult to Ecosystem type: Sandhills Seven sites have either experienced restore, and blur perspectives on pinelands problems from duff fires (most notably ecology. "Old-growth groundcover com The 20 ha Camp Lejeune Tract is a poorly Flomaton Natural Area and Eglin Air munities" are likely as important (or more known, frequently burned sand ridge site. Force Base) or face threats when fire is so) as old pine canopies. All sites should The authors have not visited the area and re-introduced. Duff fires smolder for hours work to maintain any intact groundcover no detailed reports on its status, threats, to days post-ignition, consuming forest and restore their degraded ones. cor quality were found. floor pine roots and potentially girdling trees. Fire-suppression's legacy of heavy Potential additions to old-growth 15. Croatan Ridge (Craven County, North fuel accumulations will be one of the most longleaf pine acreage Carolina) pressing issues in longleaf pine restoration Owner: Public (U.S. Forest Service, Cro management (Kush et al. 1999b). Current The ,lack of geographical and ecosystem atan National Forest) research is underway (Gordon and Varner representation is alarming. The states of Ecosystem type: Sandhills and Savanna 2002) to better understand duff fires and Mississippi and Virginia may have no their management to minimize old-growth old-growth remaining (and potentially This site, also known as the Pringle Road longleaf pine mortality. Louisiana, South Carolina, and Texas). The Tract, is a 20 ha frequently burned sand lack of Piedmont, Ridge and Valley, and hill ridge in the Croatan National Forest. Three sites are contaminated with non Cumberland Plateau Province stands is also The site strings a long, narrow pond pine native plant species, principally Chinese troubling. These points highlight the fact (Pinus serotina Michx.) pocosin, so our privet (Ligustrum sinense Lour.), Japanese that our understanding of pineland dynam acreage estimate is just that. The site has an honeysuckle (Lonicera japonica Thunb.), ics and benchmark conditions is based on
Volume 24 (2), 2004 Natural Areas Journal 147 a few isolated clayhill and sandhill stands several of these stands. This represents longleaf pine stand dynamics. that may not be generalizable to the varied another void in longleaf pine research and sites represented by pinelands. This fact conservation. This, and more detailed sur LITERATURE CITED should encourage us to search harder to veys and projects, should give researchers find these missing links. a better understanding of past southeastern Chapman, H.H. 1909. An experiment in logging climates, disturbance factors, harvesting, longleaf pine. Forest Quarterly 7:385-395. At least six sites have reports of old-growth and effects of ecological restoration on the Chapman, H.H. 1932. Is the longleaf type a longleaf pine stands but have not been pinelands landscape. The fact that so few climax? Ecology 13:328-334. visited and warrant investigation: old-growth stands remain should encourage Croker, T.C. 1987. Longleaf pine: a history of quicker action to conserve these remnants man and a forest. For. Rep. R8-FR7. U.S. AI. Kaul Tract (Coosa County, Alabama) - in the form of careful restoration, so Department of Agriculture, Forest Service, Atlanta, Ga. 36 pp. - A large mountain longleaf pineland lidifying land tenure, and land purchases reported to have been logged; site visits and/or easements. Davis, M.B. 1996. Eastern Old-Growth Forests: Prospects for Rediscovery and Recovery. are warranted. Island Press. Washington, D.C. 383 pp. ACKNOWLEDGMENTS Engstrom, R.T. 1993. Characteristic mammals A2. Number One Island (Ware County, and birds of longleaf pine forests. Pro Georgia) - A large (120 ha) frequently We thank the managers and owners of these ceedings of the Tall Timbers Fire Ecology burned flatwoods stand within the Okefe stands for their far-sighted conservation Conference 18:127-138. nokee National Wildlife Refuge. Wildfires and management of these resources and Engstrom, R.T., and FJ. Sanders. 1997. Red during 2002 may have impacted this site; their assistance with this project, spe cockaded woodpecker foraging ecology in field visits are warranted. cifically - International Paper (Flomaton an old-growth longleaf pine forest. Wilson Natural Area, Alabama), many with U.S. Bulletin 109:203-217. A3. Ca1casieu Tract (Rapides Parish, Department of Defense (Fort McClellan, Folkerts, G. W 1982. The gulf coast pitcher plant Louisiana) - A large tract (ca. 100-250 Ala.; Eglin Air Force Base, Fla.; Camp bogs. American Scientist 70:260-267. ha) within the Calcasieu District of the Lejeune, N.C.), Florida Fish and Wildlife Folkerts, G.W, M.A. Deyrup, and D.C. Sis Kisatchie National Forest. If this site is Conservation Commission (ChinsegutNa son.1993. Arthropods associated with xeric old-growth, it would represent the only ture Center, Fla.; Platt Branch Mitigation longleaf pine habitats in the southeastern old-growth longleaf pine stand in Louisiana Park, Fla.), Florida Division of Forestry United States: a brief overview. Proceedings of the Tall Timbers File Ecology Conference and potentially the largest west of the Mis (Goethe State Forest), Leon Neel (Red Hills 18:159-192. sissippi River. Field visits are needed. Plantations), The Nature Conservancy (Red Forbes, R.D. 1930. Timber Growing and Hills Plantations; Moody Tract, Ga.; Venus Logging and Turpentining Practices in the A4. Cape Fear Stand (Cumberland? Flatwoods, Fla.), Tall Timbers Research Southern Pine Region. U.S. Department of County, North Carolina) - A privately Station (Red Hills Plantations), U.S. Forest Agriculture, Forest Service, Technical Bul owned stand rumored to contain 20 ha of Service (Croatan Ridge, N.C.; Calcasieu letin No. 204. Washington, D.C. 115 pp. old-growth longleaf pineland; site visits Stand, La.), and North Carolina Division of Frost, C.C. 1993. Four centuries of changing needed. Parks and Recreation (Boyd Tract, N.C.). landscape patterns in the longleaf pine This compilation would not have been ecosystem. Proceedings of the Tall Timbers AS. Boykin Spring Longleaf (Angelina possible without the encouragement and Fire Ecology Conference 18:17-43. County, Texas) - A potential 40 ha stand knowledge of Leon Neel, Bruce Means, Gilliam, ES., B.M. Yurish, and L.M. Goodwin. in the Angelina National Forest that con Julie Moore, Bill Boyer, Janisse Ray, 1993. Community composition of an old tains red-cockaded woodpeckers and a Allison McGee, Mike Schafale, Carolyn growth longleaf pine forest: relationship to soil texture. Bulletin of the Torrey Botanical frequently-burned groundcover. Site visits Kindell, Bill Platt and many others. Club 120:287-294. and data are needed. Goebel, P.e., B.l. Palik, L.K. Kirkman, M.B. Drew, L. West, and D.C. Pederson. 2001. A6. Lufkin Tract (Angelina County, Texas) Morgan Varner is a Graduate ResearchAs Forest ecosystems of a Lower Gulf Coastal - A very large (1200 ha) frequently-burned, sistant in the School of Natural Resources Plain landscape: multifactor classification biologically intact stand owned by Temple & Environment at the University ofFlorida. and analysis. 10urnal of the Torrey Botanical Inland Corporation. Reports of this stand His research interests are fire ecology, Society 128:47-75. are impressive and warrant visits. ecological restoration, silviculture, and Gordon, D.R., and 1. M. Varner. 2002. Old natural history of the southeastern United Growth Longleaf Pine Restoration and States. Management. Final Report to Eglin Air SUMMARY Force Base 1ackson Guard, Niceville, Fla. 33 pp. However preliminary, this list represents John S. Kush is a Research Associate in the the best and most complete information School of Forestry & Wildlife Sciences at Grace, S.L., and WI. Platt. 1995. Effects of adult tree density and fire on the demogra known on old-growth longleaf pine stands. Auburn University. His research interests phy of pregrass stage juvenile longleaf pine As is obvious, very little is known about are restoration ecology, fire ecology, and (Pinus palustris Mill.). Journal of Ecology
148 Natural Areas Journal Volume 24 (2), 2004 83:75-86. on longleaf pine growth. Pp. 265-285 in R. of Pacific Northwest forests. Ecological Guyer, c., and M.A. Bailey. 1993. Amphibians Wimmer and R.E. Vetter, eds., Tree Ring Applications 2:262-274. and reptiles of longleaf pine communities. Analysis: Biological, Methodological, and Swezy, D.M., and J.K Agee. 1991. Prescribed Proceedings of the Timbers Fire Ecology EnvironmentalAspects. CAB International, fire effects on fine-root and tree mortality Conference 18:139-158. Wallingford, UK in old-growth ponderosa pine. Canadian Haase, S.M., and S.S. Sackett. 1998. Effects Neel, L. 1971. Some observations and com Journal of Forest Research 21:626-634. of prescribed fire in giant sequoia-mixed ments on the red-cockaded woodpecker in Varner, J.M., J.S. Kush, andR.S. Meldah1.1999. conifer stands in Sequoia and Kings Can the Thomasville-Tallahassee Game Preserve Old-growth montane longleaf pine stand yon National Parks. Proceedings of the Region. Pp. 137-139 in R.L.Thompson; ed., structure: a preliminary assessment. Pages Tall Timbers Fire Ecology Conference The Ecology and Management of the Red 606-608 in J.D. Haywood, ed., Proceedings 20:236-243. Cockaded Woodpecker. U.S. Department of of the 10th Southern Silvicultural Research the Interior, Bureau of Sport Fisheries and Harcombe, P.A., J.S. Glitzenstein, R.G. Knox, Conference. Gen. Tech. Rep. GTR- SRS- Wildlife and Tall Timbers Research Station, S.L. Orzell, and E.L. Briges. 1993. Vegeta 30. U.S. Department of Agriculture, For Tallahassee, Fla. tion of the longleaf pine region of the west est Service, Southern Research Station, Gulf Coastal Plain. Proceedings of the Tall Noel, J.M., w.J. Platt, and E.B. Moser. 1998. Asheville, N.C. Timbers Fire Ecology Conference18:83- Structural characteristics of old- and sec Varner, J.M., J.S. Kush, and R.S. Meldahl. 104. ond-growth stands of longleaf pine (Pinus .2000a. Ecological restoration of an old palustris) in the Gulf Coastal Region of the Hardin, E.D., and D.L. White. 1989. Rare vas growth longleaf pine stand utilizing pre USA. Conservation Biology 12:533-548. cular plant taxa associated with wiregrass scribed fire. Proceedings of the Tall Timbers (Aristida stricta) in the southeastern United Noss, R.E1989. Longleaf pine and wiregrass: Fire Ecology Conference 21:216-219. States. Natural Areas Journal 9:234-45. keystone components of an endangered eco Varner, J.M., J.S. Kush, and R.S. Meldahl. system. Natural Areas Journal 9:234-235. Hermann, S.M. 1993. Small-scale disturbances 2000b. The Mountain Longleaf Pine in longleaf pine forests. Proceedings of Noss, R.E, E.T. LaRoe, and 1M. Scott. 1995. Resources of Fort McClellan, Alabama: the Tall Timbers Fire Ecology Conference Endangered ecosystems of the United Final Report on Their Status, Ecology, 18:265-274. States: a preliminary assessment of loss and Management Needs. Final Report to and degradation. Biological Report 28. U.S. Directorate of Environment, Fort McClel Hermann, S.M., 1M. Kane, and L.C. Anderson. Department ofInterior, National Biological lan, Ala. 83 pp. In press. A vascular flora of the Wade Tract Service, Washington, D.C. 58 pp. Preserve, an old-growth longleaf pine-wire Varner, J.M., J.S. Kush, and R.S. Meldahl. grass stand in southern Georgia. Castanea. Outcalt, KW., and R.M. Sheffield. 1996. The 2003a. Vegetation of frequently burned longleaf pine forest: trends and current old-growth longleaf pine (Pinus palustris Kush, J. S., J.M. Varner, and R.S. Meldahl. conditions. Resource Bulletin SRS-9, U.S. Mill.) savannas on Choccolocco Mountain, 1999a. Old-growth longleaf pine forests in Department of Agriculture, Forest Service, Alabama, USA. Natural Areas Journal Alabama: what have we done, what can we Southern Research Station, Asheville, N.C. 23:43-52. do? Fifth Eastern Old-growth Forest Confer 23 pp. ence. Duluth, Minn. [unpaged] Varner, J.M., J.S. Kush, and R.S. Meldahl. Peet, R.K, and D.J. Allard. 1993. Longleaf 2003b. Structure of old-growth longleaf pine Kush, J.S., J.M. Varner, and R.S. Meldahl. pine vegetation of the southern Atlantic and (Pinus palustris Mill.) forests in the moun 1999b. Slow down, don't burn too fast... eastern Gulf Coast Regions: a preliminary tains of Alabama. Castanea 68:211-221. got to make that old-growth last. Pp.109- classification. Proceedings of the Tall Tim 111 in Proceedings of the 4th Longleaf Wahlenberg, w.G. 1946. Longleaf Pine: Its bers Fire Ecology Conference 18:45-82. Alliance Conference. Longleaf Alliance Use, Ecology, Regeneration, Protection, Report No.4. Platt, w.J., G.w. Evans, and S.L. Rathbun. and Management. Charles Lathrop Pack 1988. The population dynamics of a long Forestry Foundation and U.S. Department Landers, 1L., and W.D. Boyer. 1999. An old lived conifer (Pinus palustris). American of Agriculture, Forest Service. 429 pp. growth definition for upland longleaf and Naturalist 131:491-525. south Florida slash pine forests, woodlands, West, D:C., T.W. Doyle, M.L. Tharpe, J.J. and savannas. General Technical Report Platt, w.J., andS.L. Rathbun. 1993. Dynamics Beauchamp, w.i. Platt, and D.J. Downing. SRS-29. U.S. Department of Agriculture, of an old-growth population of longleaf 1993. Recent growth increases in old-growth Forest Service, Southern Research Station, pine. Proceedings of the Tall Timbers Fire longleaf pine. Canadian Journal of Forest Asheville, N.C. 15 pp. Ecology Conference 18:275-297. Research 23:846-853. McWhite, R.w., J. Furman, C.J. Petrick, and Reed, EW. 1905. A Working Plan for Forest Zutter, B.R., C.K McMahon, D.E. Whitehouse, S.M. Seiber. 1999. Integrated Natural Re Lands in Central Alabama. U.S. Department D.D. Wade, and 1S. Kush. 2003. The Flo sources Transitional Plan, Eglin Air Force of Agriculture, Forest Service Bulletin 68. maton Natural Area (FNA): demonstrating Base, 1998-2001. EgIinAFB Jackson Guard, Washington, D.C. 71 pp. the benefits of fuel management and risks Niceville, Fla. 229 pp. Ryan, KC., and W.H. Frandsen. 1991. Basal of fire exclusion in an .old-growth longleaf pine ecosystem. Pp. 164-165 in Proceedings Means, D.E. 1996. The longleaf ecosystem, injury from smoldering fires in mature Pinus of the 4th Longleaf Alliance Conference. going, going ... Pp. 210-219 in M.B. Davis, ponderosa Laws. International Journal of Longleaf Alliance Report No.6. ed., Eastern Old-Growth Forests: Prospects Wildland Fire 1:107-118. for Rediscovery an:d Recovery. Island Press. Schwarz, G.E 1907. The Longleaf Pine in Washington, D.C. Virgin Forest: a Silvical Study. 1 Wiley, Meldahl, R.S., N. Pederson, J.S. Kush, and 1M. New York, 135 pp. Varner. 1999. Dendrochronological investi Swanson, EJ., and J.E Franklin. 1992. New gations of climate and competitive effects forestry principles from ecosystem analysis
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