J. Morgan Varner III

RESEARCH ARTICLE ABSTRACT: Fire-maintained longleaf pine (Pinus palustris Mill.) savannas are imperiled throughout their range in the United States, reduced in extent by 97% since European settlement. Only half of extant stands show evidence of fire; fire suppression has led to dramatic changes in composition an.d declines iIl plaIlt diversity. Examples of pristine vegetation are kIlown for most of 'I()ngleaf pine's distinct regions-a notable exception being mountainous communities in Alabama and Georgia, USA, collec­ tively termed the mountain longleaf pine savannas. We sampled over- and understory vegetation in two • frequently burned old-growth mountain longleaf pine stands at Fort McClellan, a U.S. Army garrison in the Blue Ridge Physiographic Province of Alabama. Over the spring, summer, and fall 1999 study period, 82 native plant species were encountered in plots, representing 60 genera and 35 families. Vegetation of Overstory composition was dominated by longleaf pine, with blackjack oak (Quercus marilandica Muenchh.) and sand hickory (Cwya pallida [Ashe] Eng!. & Graeb.) as co-dominants in both stands. Understory communities were species-rich, dominated by grasses (principally Andropogon ternarius Frequently Burned Michx.), asters (Coreopsis major Walt., Chlysopsis graminifolia [Michx.] Elliott, Helianthus micro­ cephallus Torrey & Gray, Solidago odora Ait., and others), and many legumes. Non-native species were Old-Growth not encountered in sampling plots. Evidence suggests that historic fires in mountain longleaf pine . savannas were frequent (1-to 5-y return interval) and frequent fire is needed to maintain this ecosystem Longleaf Pine (Pinus in the contemporary southeastern landscape. Mill.) Vegetaci6n en Savanas Maduras de Pino de Hoja Larga (Pinus palustris Mill.) palustris Frecuentemente Quemadas en Choccolocco Mountain, Alabama, USA Savannas on RESUMEN: Las savanas de pino de hoja larga (Pinus palustris Mil!.) mantenidas con fuego estan en peligro en toda su distribucion en Estados Unidos, y fueron reducidas en el 97% de su extension desde el asentamiento de los europeos. Solamente la mitad de los lotes muestran evidencia de fuego; la Choccolocco supresion del fuego ha llevado acambios dramaticos en la composicion y la disminucion en la diversidad de plantas. Se conocen ejemplos de vegetacion prfstina para la mayorfa de las distintas Mountain, Alabama, regiones de pino de hoja larga - una notable excepcion son la comunidades montanosas en Alabama y Georgia, USA, colectivamente denominadas las savanas montanosas de pino de hoja larga. Muestrea­ mos la vegetacion del dosel y del soto bosque en dos lotes frecuentemente quemados de pino de hoja USA larga maduros en Fort McClellan, una guarnicion del ejercito en la provincia fIsiografica de Blue Ridge en Alabama. Durante el perfodo de estudio de primavera, verano y otono de 1999, 82 plantas nativas fueron encontradas en los lotes, representando 60 generos y 35 farnilias. La composicion del dosel estuvo dominada por el pino de hoja larga, con el roble blacIgack (Quercus marilandica Muenchh.) y J. Morgan Varner II J1 el nogal arena (Carya pallida [Ashe] Eng!. & Graeb.) como codominantes en ambos lotes. Las John S. Kush comunidades del sotobosque fueron ricas en especies, dominadas por pastos (principa1rnente Andro­ pogon ternarius Michx.), asters (Coreopsis major Walt., Chrysopsis graminifolia [Michx.J Elliott, Ralph S. Meldahl Helianthus microcephallus Torrey & Gray, Solidago odora Ait., y otras), y muchas leguminosas. No encontramos especies exoticas e~ los lotes de muestreo. La evidencia sugiere que los fuegos historicos en las savanas montanosas de pino de hoja larga fueron frecuentes (intervalos 1 a 5 anos) y el fuego Longleaf Pine Stand frecuente es necesario para mantener este ecosistema en el paisaje contemporaneo del sudeste. Dynamics Laboratory School of Forestry Index terms: fire regimes, mountain longleaf pine savanna, Pinus palustris, presettlement conditions, and Wildlife Sciences species composition Auburn University Auburn, AL 36849 USA

• INTRODUCTION were ignited by lightning and, in the re­ cent past, Native Arnericans (Bartram 1791, Savannas and forests dominated by Komarek 1964, Robbins and Myers 1992). lbngleaf pine- (Pinus·pahtsttis P.Miller) Longleaf pinesavahhas-' doniinated-poi'~ and a diverse bunchgrass-dominated her­ tions of the Coastal Plain, Piedmont, Cum­ I Corresponding author current address and e-mail: College of Natural Resources & baceous layer covered an estimated 37 berland Plateau, Ridge and Valley, and Blue Environment, Box 1/8526, University of million ha of the southeastern United States Ridge physiographic provinces from coast­ Florida, Gainesville, FL 32611-8526 prior to European settlement (Frost 1993). al Virginia through eastern Texas (Figure USA; [email protected] Their open structure and diverse plant spe­ 1). Because fIres in longleaf pine savannas cies assemblages (Peet and Allard 1993) often CatTY into adjacent communities (e.g., Natural Areas Joumal 23:43-52 were maintained by frequent (once every pitcher plant [Sarracenia L. spp.] bogs, 1 to 10 y), low-intensity sUlface fIres that shrub bogs, cypress strands), these savan-

Volume 23 (1),2003 Natural Areas Journal 43 nas have been referred to as the keystone suffered. A recent evaluation lists longleaf (Figure 1), less than 40,000 ha of moun­ community of the southeastern landscape pine savannas and forests as the second tain longleaf pine ecosystems remained by (Noss 1989, Myers 1990, Means 1996). most endangered ecosystems in the Unit­ 1995 (Outcalt and Sheffield 1996, Varner ed States, second only to the Florida Ever­ 2000). Of this total, only 20 ha were iden­ Since European settlement, land conver­ glades (Noss et al. 1995). tified as old growth (Varner 2000). sion has resulted in a reduction of longleaf pine acreage to less than 3% of its former Mountain longleaf pine savannas are a Fire is noticeably absent or much reduced range, covering fewer than 1.2 million ha loosely defined community type in north­ over most of the southeastern mountain (Outcalt and Sheffield 1996). The remain­ ern Alabama and Georgia. No rigorous landscape. In the absence offrequent, low­ ing acreage contains few pristine stands or delineation of a "mountain" (often termed intensity (every 1 to 5 y) frres, Coastal landscapes (Ware et al. 1993, Means 1996); "montane") type exists, though most ob­ Plain longleaf pine savannas succeed to a indeed only about half of the remnants servers agree that Blue Ridge, Ridge and closed-canopy, mixed hardwood forest, show evidence of frre (Outcalt 2000). With Valley, and Cumberland Plateau sites are termed the Southern Mixed Hardwood the precipitous decline of longleaf pine, within the mountain region. If the Pied­ Forest (SMHF) (Ware et al. 1993). In the the associated flora and fauna have also mont Physiographic Province is included mountain region, the same successional trajectory prevails, with communities shift­ ing toward either the SMHF (Ware et al. 1993) or Oak-Hickory-Pine (OHP) forest (Kuchler 1964, Skeen et al. 1993). Typical successional overs tory species in the moun­ tains include red maple (Acer rub rum L.), several oaks (particularly Quercus veluti­ na Lam. and Q. prinus L.), other pines (Pinus taeda L., P. echinata Mill., and P. virginiana Mill.), sourwood (Oxydendrum arboreum [L.] DC), and blackgum (Nyssa sylvatica Marsh.) (Golden 1979, Maceina 1997, Varner 2000). Understory succes­ sional species include many shrubs, green­ brier (Smilax rotundifolia L.), and low­ bush blueberry (Vaccinium pallidum Chapman), the latter often forming dense, monospecific stands in infrequently burned stands (Golden 1979, Varner 2000). Land use and contemporary forest management in the mountain longleaf pine region have created isolated patches of erratically burned vegetation within a larger fire-sup­ pressed forested mosaic (Alabama Natu­ ral Heritage Program 1994, Varner 2000), making approximations of pristine or pre­ settlement conditions difficult.

Although mountain pinelands represent one of the most unique longleaf pine eco­ systems-they occur at high elevations (up to 600 m above msl), commonly experi­ ence ice and snow storms, are located at the maximal distance from tropical storms, occupy extremely dissected topography, and are in the most climatically distinct region of longleaf pine's natural range (Craul 1965)-they have undergone very Figure 1. Native range of longleaf pine in Alabama and Georgia, USA (light and dark shaded areas). little study. Surveys of longleaf pine veg­ Dark shaded region denotes the range of mountain longleaf pine savannas and forests. Physiographic provinces represented by this region are in white type. The white triangle represents the approximate etation have been undertaken in the west­ location of Fort McClellan in Alabama. ern Gulf Coastal Plain (Marks and Har-

44 Natural Areas Journal Volume 23 (1),2003 combe 1981, SchafaleandHarcombe 1983, burned old-growth longleaf pine commu­ sitioning to clay-loams in subsoils (Harlin Bridges and Orze1l1989, Harcombe et al. nities. These benchmark data and their et al. 1961). Slopes averaged 40% to 60% 1993), eastern Gulf Coastal Plain (Pen­ information could then be utilized for iden­ at Caffey Hill and 30% to 45% at Red-tail found and Watkins 1937, Clewell 1986, tiftcation, conservation, and restoration of Ridge. Caffey Hill had a SSE aspect, while Peet and Allard 1993, Drew et al. 1998), this rare ecosystem. An additional objec­ Red-tail Ridge's aspect was WSw. Atlantic Coastal Plain and Sandhills (Harp­ tive was to establish permanent plots for er 1906, Walker and Peet 1983, Frost and long-term studies on this disappearing Data Collection Musselman 1987, Gilliam et al. 1993, Peet community type. and Allard 1993), Upper Coastal Plain Understory Species Composition (Beckett and Golden 1982), and Piedmont METHODS (Golden 1979). In contrast, mountain Within each intensive study area, we in­ longleaf pine ecosystems in the Ridge and stalled plot centers and vegetation sam­ Study Areas Valley, Blue Ridge, and Cumberland Pla­ pUng quadrats (Caffey Hill-ftve plots; teau have undergone very little study (but Fort McClellan is a 7,300-ha U.S. Depart­ Red-tail Ridge-four plots). From each see Maceina et al. 2000). ment of Defense Army garrison in eastern plot center, four 1.1-m X 1.1-m quadrats Calhoun County, northeastern Alabama were placed in cardinal directions, each 5 Previous research on longleaf pine com­ (33 0 42' N, 85 0 45' W). Fort McClellan con­ m from plot center, for a total of 20 quad­ munities in the mountain region have been tains a large portion of Choccolocco Moun­ rats at Caffey Hill and 16 at Red-tail Ridge. based on degraded stands and were gener­ tain and its spur ridges, all capped with We sampled all quadrats once during spring ally limited in scope. Golden (1979) ana­ Weisner quartzite. Climate is warm and (April), summer (June), and fall (Septem­ lyzed longleaf pine communities in the humid, with a mean annual temperature of ber) 1999. Within each quadrat, all plant lower Piedmont Province, providing quan­ 17°C. Winters are short and mild; sum­ species were identifted (nomenclature ac­ titative data, albeit based on stands with mers are long, humid, and warm. Precipi­ cording to Kartesz 1994) and assigned an infrequent ftre and dubious management tation is evenly distributed throughout the approximate cover value (1 % to 100%) histories. Using research based at Fort year, with small peaks associated with based on visual estimation. Since sam­ McClellan, Alabama, USA, Maceina summer thunderstorm events, and troughs pling took place over three seasons, all (1997) described the overstory species occurring during October. Annual precip­ three (spring, summer, and fall) cover val­ composition and successional trajectory itation averages 125 cm, with small ues were averaged for calculations. Rela­ of a second-growth mountain longleaf pine amounts falling as ice and snow (Harlin et tive frequency (RF) and relative cover (RC) stand. Maceina's tract had an erratic burn­ al. 1961, Craul 1965). were calculated for both sites. Importance ing history and was subject to mechanical values (IV) were calculated for individual scraping several years prior to her survey At Fort McClellan, we selected two stands species as the sum of relative frequency (Maceina1997). In the same tract, Mace­ with histories of frequent burning (1- to 3- (RF) and relative cover (RC), for a poten­ ina et al. (2000) surveyed the mountain y cycles) and an old-growth longleaf pine tial importance value equal to 200. longleaf pine flora utilizing meander sur­ overstory (oldest age classes preceding veys, a method that does not easily allow European settlement, ca. 1850). The two. To describe and compare these communi­ for comparisons or quantiftcation. Simi­ stands, hereafter Caffey Hill and Red-tail ties, we assigned each species a form class. larly, Whetstone and others (1996) ob­ Ridge, were located at ca. 450 and 350 m Tree seedlings (T) were classifted as woody served 871 plant species in their survey of above msl, respectively. Caffey Hill is an species capable of reaching the canopy. Fort McClellan's landscape. However, they upper slope stand covering 1.5 ha abutting Shrubs (S) were classifted as woody spe­ did not attribute species to speciftc com­ a downslope 250-ha matrix of younger, cies normally incapable of reaching the munity types, nor did they recognize moun­ frequently burned stands. Red-tail Ridge canopy. The other two classes were forbs, tain longleaf pine communities in their spans a mid- to upper slope position cov­ including woody vines (F), and grasses survey. From these reports emerge an in­ ering 1.8 ha abutting a frequently burned, (G). complete description of the vegetation and somewhat disturbed 20-ha tract below on a rudimentary understanding of mountain side slopes. At both sites, soils were Overstory Species Composition longleaf pine community composition, mapped as Rough Stony Land-Sandstone, -confounded by a lack· of data or-under-­ a miscellaneous Typic -Udultwith many All longleaf pine stems (100% sample at standing from pristine sites .. outcrops of quartzite and sandstone bed­ each site) > 2.5 cm dbh (where dbh = rock and loose rock fragments, represen­ diameter at breast height [1.37 m]) within We initiated a study in 1998 to address the tative of many mountain longleaf pine the two intensive study areas were mapped lack of data and understanding of moun­ ecosystems (Craul 1965). These sites are (azimuth and horizontal distance to plot tain longleaf pine communities. The pri­ located upslope of, and often intergrade centers) and measured. All other overs tory mary objective of this study was to survey with, the Anniston soil series. Anniston species> 2.5 cm dbh were identified, count­ and quantify overs tory and understory plant soils are clayey, kaolinitic, thermic Rhod­ ed, and measured (dbh only). From these species composition of two frequently ic Paleudults. Smface soils are loams, tran- measurements, average and relative stem

Volume 23 (1),2003 Natural Areas Journal 45 1 density (trees ha- ; RD) and average and Table 1. Understory plant species (stems < 2.5 cm dbh) list from two frequently burned old· relative basal area (m2 ha- 1; RBA) were growth mountain longleaf pine stands located at Fort McClellan, Alabama, USA. Values listed calculated. Importance values were calcu­ are Importance Values, where IV = Relative Frequency + Relative Cover. Asterisks in columns lated as the sum of RD and RBA, for a denote IVs < 1. Both frequency and cover values were averaged over the three seasonal samples potential importance value of 200. for each quadrat. Form refers to plant form, either tree (T), shrnb (S), forb (F), or grass (G). Nomenclature follows Kartesz 1994. Long-term fIre histories of the two stands are, as with all but a handful of sites in the Southeast, unavailable beyond the recent FAMILY Importance Values forest management era (1960 to present). Species Form Caffey Hill Red·tail Ridge Assessment of longer term fue history can be approximated based on overs tory spe­ AMARYLLIDACEAE cies composition. While understory com­ Hypoxis hirsuta (L.) Coville F munities are unlikely to contain species that are relicts of fire suppression, overs to­ ANACARDIACEAE ry species have a much higher probability Rhus copallina L. S 12 9 of remaining, even after long-term resump­ Rhus radicans L. F 0 tion of active fire management (Boyer * Rhus toxicodendron L. F 3 0 1990b). To detect relicts and overstory species stability, we used under-represen­ tation techniques (Golden 1979). Under­ ANNONACEAE representation, as defined by Golden Asimina parviflora (Michx.) Dunal S 3 * (1979), is the condition where there are fewer individuals in two consecutive size APIACEAE classes than in the subsequent larger size Angelica venenosa (Greenway) Fernald F 0 class. This technique identifies species that colonized at a point in time, but at present ASCLEPIADACEAE are failing to regenerate; that is they are Asclepias amplexicaulis Smith F 0 relicts of past favorable ecological condi­ tions. Species that either (1) were under­ ASTERACEAE represented at both sites, or 2) were under­ Aster dumosus L. F I 0 represented at one site and absent at the Aster patens Aiton F 2 8 other, were classified as relict species. Aster undulatus L. F 2 6 Chrysopsis graminifolia (Michx.) Elliott F 9 6 RESULTS Coreopsis major Walter F 13 3 Elephantopus tomentosus L. F 2 0 Understory Species Composition Helianthus microcephalus Torrey & Gray F 6 13 Helianthus mollis Lamarck F Seventy-two understory plant species were * encountered over the three sampling peri­ Krigia biflora (Walter) Blake F 2 * ods. Though most species were represent­ Liatris graminifolia Willdenow F 1 ed at both sites (42), Red-tail Ridge had Silphium terebenthinaceum Jacquin F 0 3 more species and higher cover values than Solidago erecta Pursh F 5 * Caffey Hill (Table 1). Red-tail Ridge had Solidago odora Aiton F 10 4 more forb (47 versus 34) and legume (10 versus 7) species than Caffey Hill. Num­ CAMPANULACEAE bers of grass (Poaceae), aster (Asterace­ Specula ria peifoliata L. A.DC F ae), shrub, and tree species were similar between sites. Both sites were character­ COMMELINACEAE ized by a diverse groundcover layer, dom­ Tradescantia hirsuticaulis Small F 0 inated by Andropogon ternarius Michx., * Chrysopsis graminifolia (Michx.) Elloitt, Coreopsis major Walt., Euphorbia corol­ CONVULVULACEAE lata L., Helianthus microcephalus Torrey Ipomoea lacunosa L. F 0 3 & Gray, Panicum commutatum Schult., continued

46 Natural Areas Journal Volume 23 (1),2003 Pteridium aquilinum L., Rhus copallina Table 1, continued L., and Solidago odora Ait. Asters, grass­ es, and legumes were the best represented FAMILY Importance Values .. families, together comprising 50% and 56% importance at Caffey Hill and Red­ Species Form Caffey Hill Red-tail Ridge tail Ridge, respectively. Herbaceous spe­ cies, in general, were most important at EBENACEAE both sites, accounting for 71.5% of total Diospyros virginiana L. S 8 3 understory importance at Caffey Hill, and 85% at Red-tail Ridge. Seedlings. of tree species were rare, together comprising only ERICACEAE 5% and 4% of total importance at Caffey Vaccinium arboreum Marshall S 4 6 Hill and Red-tail Ridge. Shrubs were also rare, accounting for 23.5% and 11 % total EUPHORBIACEAE importance at Caffey Hill and Red-tail Euphorbia corollata L. F 9 9 Ridge. Non-native species were not en­ countered in any of the 36 sampling quad­ FABACEAE rats in any season, or at either of the two Clitoria mariana L. F 3 6 sites outside of sampling quadrats. Desmodium viridiflorum (L.) De Candolle F o Galactia volubilis (L.) Britton F 4 Individual sample species richness maxi­ Lespedeza intermedia (Watson) Britton F 2 ma were 22 species per quadrat at Red-tail Ridge and 19 per quadrat at Caffey Hill. Lespedeza procumbens Michx. F * The lowest individual samples at each site Lespedeza stuevei Nutt. F o 1 were 8 and 10 species per quadrat. Cover Lespedeza virginica (L.) Britton F o 3 values per quadrat averaged 82% (± 6.4%) Rhynchosia tomentosa (L.) Hook. & Am. F 5 5 at Red-tail Ridge, 68% (± 5.3%) at Caffey Senna marilandica (L.) Link F 2 2 Hill. Individual quadrat cover maxima / Tephrosia virginiana (L.) Pers. F 5 6 minima were 109% / 50% at Red-tail Ridge, and 83%/45% at Caffey Hill. FAGACEAE Quercus marilandica Muenchh. T 6 3 Overstory Species Composition Quercus prinus L. T o *

Eighteen overstory (dbh >2.5 em) woody HYPERICACEAE plant species were recorded at the two Hypericum gentianoides (L.) B. S. P. F study sites; Red-tail Ridge had 16 species, * Hypericum hypericoides (L.) Crantz. F o Caffey Hill 9 species (Table 2). Longleaf * pine was the most frequent and important Hypericum punctatum Bush F * * species at both sites, representing 63.3% and 94.5% of basal area at Red-tail Ridge IRIDACEAE and Caffey Hill, respectively. Total basal Iris verna L. F o 3 area at Red-tail Ridge and Caffey Hill were Sisyrinchium angustifolium Mill. F * 2 1 2 1 20.6 m ha- and 8.8 m ha- , respectively. Overs tory density at Red-tail ridge and JUGLANDACEAE 1 Caffey Hill was 721 and 535 trees ha- , Carya pallida (Ashe) Engler & Graebner T 8 5 with longleaf pine representing 39.2% and 55.6% of all trees at each site. The most LAMIACEAE I I importanLspecies~ at .both _sites were Salvia urticifolia (L.) _ F 2 3 • longleaf pine, blackjack oak, sand hicko­ ry, and to a lesser extent, blackgum and LAURACEAE rock chestnut oak (Quercus prinus L.). Sassafras albidum (Nutt.) Nees. S 20 0

I Only three of the eighteen species were NYSSACEAE (1) present at both sites, and (2) not signif­ Nyssa sylvatica Marshall T 0 icantly underrepresented «25 % of size classes underrepresented): longleaf pine, continued !

I Volume 23 (1),2003 Natural Areas Journal 47

I blackjack oak, and sand hickory (Table 3). In addition to the previous three species, Table 1, continued blackgum, post oak (Q. stellata Wagenh.), sassafras (Sassafras albidum (Nutt.) Nees.), FAMILY Importance Values tree sparkleberry (Vaccinium arboreum Species Form Caffey Hill Red-tail Ridge Marsh.), red maple, sourwood, flowering dogwood (Cornus florida L.), and sweet- ONAGRACEAE gum (Liquidambar styraciflua L.) were Circaea lutetiana Aschers. & Magnus F 0 well represented in our two stands. Relict Gaura longiflora Spach F 2 species included rock chestnut oak, short- leaf pine (Pinus echinata Mill.), black cher- OXALIDACEAE ry (Prunus serotina Ehrh.), black oak (Q. velutina), Virginia pine, Sonderegger pine Oxalis dillenii Jacquin F 1 4 (P. X sondereggeri H. H. Chapm.), and Oxalis violacea L. F 0 1 loblolly pine. PINACEAE Pinus echinata Mill. T 0 1 DISCUSSION Pinus palustris Mill. T 0 Understory Species Composition POACEAE Fire-maintained understory communities Andropogon ternarius Michx. G 16 20 in longleaf pine savannas are renowned Panicum commutatum Schult. G 9 5 for being extremely species-rich (Walker Panicum virgatum L. G 3 and Peet 1983, Peet and Allard 1993). * Panicum spp. G 3 0 These two stands at Fort McClellan were no exception. First, even with limited sam- Poa spp. G 0 * pling, 72 native understory plant species Piptochaetium avenacea L. G * 8 were encountered. Single sample maxima of 22 and 19 species per 1.21 m2 quadrat POLEMONIACEAE are comparable to those found at many Phlox amoena Sims F 0 sites listed in Peet and Allard (1993). Our findings are significant, in light of the fact POLYPODIACEAE that data from mountain longleaf pine com- Asplenium platyneuron (L.) Oakes F . 0 1 munities were not included in Peet and Pteridium aquilinum L. F 10 9 Allard (1993) or any other works. RUBIACEAE Very few tree seedlings and saplings were Galium pilosum Ait. F 0 4 found in sampling the two study stands. Longleaf pine, which overwhelmingly dominated the overstory of both sites (Ta- SAXIFRAGACEAE ble 2), was only encountered as a seedling Hydrangea quercifolia Bartram S 0 4 once at Caffey Hill, and none were ob- served at Red-tail Ridge. As with the SCROPHULARIACEAE longleaf pine overs tory, seedlings and sap- Agalinus purpurea (L.) Pennell F 0 1 lings are concentrated in even-aged patch- Aureolaria pedicularia (L.) Raf. F 4 0 es under gaps in the canopy (Wahlenberg Pentstemon pallidus Small F 0 2 1946, Brockway and Outcalt 1998). There- Seymeria cassioides (Walt.) Blake F * 0 fore, although seedling stocking at Caffey Hill was excellent in areas (pers. observa- SMILACACEAE tion, 1.M. Varner), our sampling revealed Smilax glauca Walt. F 4 5 very few seedlings. At Red-tail Ridge, due to its proximity to several military firing ranges, nearly annual burning had taken VIOLACEAE place for 11 y prior to sampling, so seed- Viola palmata L. F 0 2 lings were unable to grow to a size (ca. 0.8 Viola pedata L. F 0 cm ground-line diameter) where they could

48 Natural Areas Journal Volume 23 (1),2003 Table 2. Overstory' (stems> 2.5 cm dbh) species of two frequently burned (1-3 year return . the Piedmont of Alabama. However, he interval) old-growth mountain longleaf pine stands at Fort McClellan, Alabama, USA. Impor­ also found several Coastal Plain species tance values are sums of relative density (trees ha-1) and relative basal area (m2 ha-1), for a (Symplocos tinctoria [L.] L'Heritier, potential sum of 200. BA=basal area, IV=importance value. Gelsemium sempervirens [L.] Ait., and others) and co-dominance by southern red Caffey Hill Red-tail Ridge oak (Q.falcata Michx.), which typically is not found in mountain savannas. In fact, Trees BA Trees BA the composition reported by Golden was 1 2 1 1 1 Species (18) ha- (m ha· ) IV ha- (m2ha- ) IV more similar to Upper Coastal Plain longleaf pine communities (Beckett and P. palustris 282.82 13.01 297.64 102.5 8.29 150.2 Golden 1982, Peet and Allard 1993). Fi­ Q. marilandica 163.23 1.44 29.6 174.35 0.32 36.2 nally, in contrast to other mountain longleaf Carya pallida 110.09 2.30 26.4 20.34 0.04 4.3 pine community investigations (Golden Nyssa sylvatica 64.96 0.95 13.6 4.36 0.02 1.0 1979, Whetstone et al. 1996, Maceina et Q. prinus 20.94 0.98 7.7 21.07 0.07 4.7 al. 2000), we found no non-native species P. echinata 13.41 0.57 4.6 in any of our 36 sampling quadrats. This Quercus stellata 18.79 0.34 4.3 1.45 <0.01 0.3 finding suggests that prescribed fIre is an Primus sera tina 17.18 0.32 3.9 effective management tool for maintenance Quercus velutina 6.44 0.25 2.1 5.09 0.01 1.1 of longleaf pine community integrity Sassafras albidum 9.45 0.01 1.9 (Drew et al. 1998). P. virginiana 3.75 0.23 1.6 Vaccinium arboreum 8.59 0.03 1.3 Overstory Species Composition 4.83 Acer rub rum 0.06 1.0 Longleaf pine dominated both study sites, Oxydendrum arboreum 2.68 0.04 0.6 1.45 0.01 0.4 with importance values similar to those in Comus florida 2.15 om 0.3 other mountain longleaf pine co~unities P. X sondereggeri 0.54 0.02 0.2 (Golden 1979, Maceina 1997). Overstory P. taeda 0.54 0.01 0.1 co-dominants (blackjack oak and sand hick­ Liquidambar styraciflua 0.54 <0.01 0.1 ory) were typical of other sites in the Pied­ mont (Golden 1979), Upper Coastal Plain Totals 721.48 20.56 535.20 8.76 (Beckett and Golden 1982, Peet and Allard 1993), Fall Line Sandhills (Gilliam et al. 1993), and other Blue Ridge sites (Maceina 1997). Differences between these stands, while most pronounced in the herbaceous and shrub layers, can also be observed in survive even cool surface fires (Boyer ates, and are all somewhat fIre-adapted. tree species presence data. Co-dominants in 1990a). Longleaf pine saplings « 10 cm Furthermore, all three of these species were the lower Piedmont (Golden 1979) and dbh) were abundant at Caffey Hill, and observed as small individuals, mostly less Upper Coastal Plain (Beckett and Golden less common at Red-tail Ridge (Varner than 0.5 m tall (J.M. Varner, pers.observ.). 1982, Peet and Allard 1993) included south­ 2000), again the latter due to the aggres­ Low shrub importance is' a characteristic ern red oak, flowering dogwood, post oak, sive military burning regimes. Other tree of frequently burned longleaf pine savan­ loblolly pine, shortleaf pine, and sweetgum. species seedlings were also rare (Table 1), nas throughout their range (Wahlenberg Black oak and turkey oak (Q. laevis Walt.) resulting in under-representation of these 1946, Boyer 1990a, Peet and Allard 1993, were co-dominants in a fire-suppressed normally fIre-tolerant species (Table 3). Landers and Boyer 1999). North Carolina Sandhills stand (Gilliam et al. 1993). Turkey oak, while known to occur Shrub and small tree species were rare at Previous studies (Golden 1979, Maceina in mountain longleaf pine stands at Fort both sites, except for sassafras, ~ winged ~ -1997) have described mountain longleaf ~ McClellan (ANHP 1994, Varner 2000); is a sumac (Rhus copallina L.), and persim­ pine ecosystems with similar composition, rare mountain congener. Black oak is sus­ mon (Diospyros virginiana L.) (Table 1). but often with obvious differences or signs ceptible to high mortality where frequent Sassafras and persimmon, while both ca­ of degradation. Golden (1979) found sev­ fires occur, is a poor sprouter (Sander 1990), pable of reaching the canopy, are common eral common mountain associates (Pte rid­ and therefore is an indicator of extended as understory species in frequently burned hun aquilinul11, Tephrosia virginiana [L.] fire-free intervals. The aspect dominance­ longleaf pine savannas throughout the Pel's., Vaccinium arboreum, Coreopsis open canopy of longleaf pine with a patchy Southeast (Peet and Allard 1993). All three major, Asimina parviflora [Michx]. Du­ midstory of blackjack oak and sand hicko­ species are common longleaf pine associ- nal, and others) in longleaf pine stands in ry-common to both sites, and to frequently

Volume 23 (1),2003 Natural Areas Journal 49 burned communities in the mountain region, Co-dominant species were of limited im­ intolerant species, as is the case with sev­ is strikingly similar to Coastal Plain longleaf portance in these mountain longleaf pine eral oak species in Coastal Plain land­ pine sites. As with Coastal Plain communi­ stands (most with IVs < 10 and/ or under­ scapes (e.g., Quercus geminata domes, ties, deviations from this aspect suggest some represented; Tables 2 and 3). Maceina Myers 1990). Similarly, an extended fIre­ history of fire exclusion or alteration. (1997), in a stand about 1 krn from this free interval (a long enough period to al­ Caffey Hill site, found similar trends in low for establishment and growth to a stage importance values for most of the same where fIre-susceptibility was low, ca. 15 y Table 3. Diameter class under-representa­ species, along with blackgum. At Red-tail or more) may have led to the establish­ tion of overstory tree species in two fre­ Ridge and Caffey Hill, overstory impor­ ment and survival of these normally fIre­ quently burned old-growth mountain longleaf pine stands at Fort McClellan, tance Was d_ominated by longleaf pine, killed species. For the latter explanation, Alabama, USA. Bold type indicates relict blackjack oak, and sand hickory; all fIre­ these atypical species will be killed by species in our analysis. Nomenclature fol­ adapted species. Longleaf pine's adapta­ continued burning and lost without replace­ lows Kartesz (1994). * denotes the absence tions to fIre are well known (e.g., Chap­ ment in such a hostile seedling environ­ of that species at that particular site, or man 1932), and both blackjack oak and ment. Whether or not this fIre-free interval better conceptualized as total, or 100% under-representation. sand hickory are common associates of was within some historical range of vari­ frequently burned longleaf pine savannas ability or was an extreme event cannot be in Fall Line communities, and in the case stated with assurance. Further study should Percentage of Size Classesa with Under-representationb of blackjack oak, throughout much of examine the spatial distribution of these longleaf pine's range (Peet and Allard fire-intolerant species, and their status Caffey Red-tail 1993). These three species were the only should be monitored with continued burn­ Species Hill Ridge species well represented in all size classes ing applications. (Table 3). Underrepresented species or Pinus palustris 10 0 atypical assemblages of establishing spe­ Only a handful of longleaf pine communi­ Quercus marilandica 0 0 cies indicate a new community trajectory, ties in pristine condition can be found at Carya paUida 0 0 largely due to altered fire regimes (Golden Fort McClellan and elsewhere in the moun­ Nyssa sylvatica 50 0 1979, Maceina 1997). Blackgum and oth­ tain region of Alabama and Georgia. Sam­ Quercus prinus 17 17 ers (most notably red maple, sweetgum, pling of other frequently burned stands, Pinus echinata * 25 and sourwood) were not signifIcantly un­ particularly in these peripheral regions, Quercus steUata 0 29 derrepresented in the two study stands, but will surely prove valuable and contribute Prunus serotina * 17 since they were absent or underrepresent­ to an approximation of presettlement land­ Quercus velutina 20 40 ed at one site, the hypothesis can be made scape conditions in the southeastern Unit­ Sassafras albidum 0 that with continued sampling of more ed States. The next step that must be un­ * stands, these species would be classifIed dertaken is preservation and management Pinus virginiana 44 * as relicts. of remnant mountain longleaf pine com­ Vaccinium arboreum 0 * munity types. However, this step will only Acer rubrum 0 * The higher than expected species richness be possible when the role of fIre as a dom­ Oxydendrum arboreum 25 0 (18 species) in the overstory was surpris­ inant environmental and evolutionary force Comus florida * 0 ing for these frequently burned stands. in mountain longleaf pine savannas is ac­ Pinus X sondereggeri * 50 Longleaf pine savannas are normally as­ knowledged. Pinus taeda * 33 sociated with remarkably low tree species Liquidambar styraciflua 0 diversity (Bartram 1791, Reed 1905, * ACKNOWLEDGMENTS Schwarz 1907, Wahlenberg 1946, Boyer 1990a, Landers and Boyer 1999). Indeed, R. Sampson, C. Avery, S. Harrison, and D. a 5-cm size classes were used longleaf pine rarely co-occurs with other Spaulding provided fIeld and laboratory b Under-representation, as defined in Golden canopy trees (Landers and Boyer 1999). assistance for this study. Discussions with (1979), is the condition where there are Its life history traits from seed to seedling D. Folkerts, M. MacKenzie, and W. Boyer fewer individuals in two consecutive size classes than in the subsequent larger size to sapling to adult all require frequent fIre, and the comments of S. Orzell and two class. Augmenting his definition with the which few species can withstand (Wahlen­ anonymous reviewers greatly enhanced the condition that if a species is absent from a berg 1946). Even though most species were quality of this paper. The assistance pro­ site, then it is also underrepresented. There­ of low importance (most with IV < 15; vided by Directorate of Environment and fore, if a species was underrepresented at Table 2) or were under-represented (Table U.S. Fish and Wildlife Service personnel one site and absent from the other, we clas­ sify it as a relict species. Percentage of size 3), their curious occurrence may be a prod­ at Fort McClellan, notably R. Smith, G. classes with under-representation was cal­ uct of a fIre intensity mosaic or a relict of Horsley, and B. Garland, was invaluable. culated as: % = # of size classes with con­ a past fire-free interval. Erratic fire behav­ Funding was provided by a U.S. Depart­ secutive under-representation / # of total ior in these undulating, steep landscapes ment of Defense Legacy Fund grant be­ size classes. may aid in creation of patches of fire- tween the U.S. Army Directorate of Envi-

50 Natural Areas Journal Volume 23 (1),2003 ronment, U.S. Forest Service Southern climax? Ecology 13:328-334. and savannas. General Technical Report Research Station, and the Auburn Univer­ Clewell, A.F. 1986. Natural setting and vege­ SRS-29, U.S. Department of Agriculture, sity School of Forestry and Wildlife Sci­ tation of the Florida panhandle. Report Forest Service, Southern Research Station, Asheville, N.C. 15pp. ences. COESAMlPDEI-86/001, U.S. Army Corps of Engineers Mobile, Ala. 773 pp. Maceina, E.C. 1997. Characterization ofa Craul, ?J. 1965. Longleaf pine site zones. montane longleaf pine community type on Morgan Varner is Graduate Research As­ Unpubl. final report on file with the U.S. Fort McClellan, Alabama: community struc­ sistant in the College ofNatural Resources Department of Agriculture, Forest Service, ture within pine-hardwood forest type. M.S. at the University of Florida. His research Southern Research Station, Auburn, Ala. thesis, Auburn University, Auburn, Ala. 180 interests are fire ecology, ecological resto­ 58 pp. pp. ration, silviculture, and natural history of Drew, M.B., L.K Kirkman, and A.K Ghol­ Maceina, E.C., J.S. Kush, and R.S. Meldahl. 2000. Vegetational survey of a montane the southeastern United States. son, Jr. 1998. The vascular flora of Ichau­ way, Baker .County, Georgia: a remnant longleaf pine community at Fort McClel­ lan, Alabama. Castanea 65:147-154. John S. Kush is a Research Associate in longleaf pine/wiregrass ecosystem. Casta­ nea 63:1-24. Marks, P.L., and P.A. Harcombe. 1981. Forest the School of Forestry and Wildlife Scienc­ vegetation of the Big Thicket, Southeast es at Auburn University. His research in­ Frost, C.C. 1993. Four centuries of changing landscape patterns in the longleaf pine eco­ Texas. Ecological Monographs 51:287-305. terests are restoration ecology, fire ecolo­ system. Proceedings of the Tall Timbers Means, D.B. 1996. The longleaf ecosystem, gy, and longleaf pine stand dynamics. Fire Ecology Conference 18: 17 -43. going, going. .. Pp. 210-219 in M.B. Frost, C.C., and LJ. Musselman. 1987. Histo­ Davis, ed., Eastern Old-growth Forests: Ralph S. Meldahl is an Associate Profes­ ry and vegetation of the Blackwater River Prospects for Rediscovery and Recovery. sor in the School of Forestry and Wildlife Ecologic Preserve. Castanea 52: 16-46. Island Press, Washington, D.C. Sciences atAuburn University. His research Gilliam, F.S., B.M. Ymish, and L.M. Good­ Myers, RL. 1990. Scrub and high pine. Pp. interests focus on the quantitative aspects win. 1993. Community composition of an 150-293 in R.L. Myers and J.J. Ewel, eds., of longleaf pine stand dynamics. old-growth longleaf pine forest: relation­ Ecosystems of Florida. University of Cen­ ship to soil texture. Bulletin of the Torrey tral Florida Press, Orlando. Botanical Club 120:287-294. Noss, RF. 1989. Longleaf pine and wiregrass: LITERATURE CITED Golden, M.S. 1979. Forest vegetation of the keystone components of an endangered ec­ osystem. Natural Areas Journal 9:234-235. Alabama Natural Heritage Program. 1994. lower Alabama Piedmont. Ecology 60: 770- Natural heritage inventory of Fort McClel­ 782. Noss, RF., E.T. LaRoe, and J.M. Scott. 1995. lan, Main Post. Alabama Natural Heritage Harcombe, P.A., J.S. Glitzenstein, RG. Knox, Endangered ecosystems of the United States: Program, Montgomery. 191 pp. S.L. Orzell, and E.L. Bridges. 1993. Vege­ A preliminary ass.essment of loss and deg­ radation. Biological Report 28, U.S. De­ Bartram, W. 1791. Travels through North & tation of the longleaf pine region of the partment of Interior, National Biological South Carolina, East & West Florida. James West Gulf Coastal Plain. Proceedings of the Service, Washington, D.C. 58 pp. and Johnson, Philadelphia, Penn. 441 pp. Tall Timbers Fire Ecology Conference 18:83-104. Outcalt, KW. 2000. 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A phytogeographical sketch Southern Research Station, Asheville, N.C. ture Handbook 654, U.S. Department of of the Altamaha Grit Region of the coastal 23 pp. Agriculture, Forest Service, Washington, plain of Georgia. New York Academy of D.C. Science 7:1-415. Peet, R.K, and D.J. Allard. 1993. Longleaf pine vegetation of the southern Atlantic and Boyer, W.D. 1990b. Growing season burns for Kartesz, J.T. 1994. Synonymized checklist of eastern Gulf Coast Regions: a preliminary control of hardwoods in longleaf pine stands. the vascular flora of the United States, Can­ classification. Proceedings of the Tall Tim­ Research Paper SO-256, U.S. Department ada, and Greenland. 2nd Ed. Timber Press, bers Fire Ecology Conference 18:45-82. of Agriculture, Forest Service, Southern Portland, Ore. 1438 pp. Research Station, New Orleans, La. 7 pp. Komarek, E.V. 1964. The natural history of Penfound, W.T., and A.J. Watkins. 1937. Phy­ tosociological studies in thepinelandsof i3rockway,~b:G.,al1dK.W. 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Volume 23 (1),2003 Natural Areas Journal 51 review. Miscellaneous Publication No.8, Varner, J.M., III. 2000. Species composition, Ware, S., C.C. Frost, and P.D. Doerr. 1993. Tall Timbers Research Station, Tallahas­ structure, and dynamics of old-growth Southern mixed hardwood forest: the former see, Fla. 96 pp. mountain longleaf pine forests of Fort Mc­ longleaf pine forest. Pp. 447-493 in W.H. Sander,l.L. 1990. Quercus velutina Lam. black Clellan, Alabama. M.S. thesis, Auburn Martin, S.G. Boyce, A.C. Echternacht, eds., oak. Pp. 744-750 in R.M. Burns and B.H. University, Auburn, Ala. 128 pp. Biodiversity of the Southeastern United Honka1a, tech. coords., Silvics of North Wahlenberg, W.G. 1946. Longleaf pine: its States: Lowland Terrestrial Communities. America, Vol. 2: Hardwoods. Agriculture use, ecology, regeneration, protection, and Wiley,_ New York. Handbook 654, U.S. Department of Agri­ management. Charles Lathrop Pack Forest­ Whetstone, R.D., J.M. Ballard, L.M. Hodge, culture, Forest Service, Washington, D.C. ry Foundation and U.S. Department of and D.D. Spaulding. 1996. Vascular flora Schafa1e, M.P., and P.A. Harcombe. 1983. Agriculture, Forest Service, Washington, of Fort McClellan, Calhoun County, Ala­ Presettlement vegetation of Hardin County, D.C. 429 pp. bama. Whetstone Consulting, Inc., Annis­ Texas. American Midland Naturalist Walker, J., and R.K. Peet. 1983. Composition ton, Ala. 153 pp. 109:355-366. and species diversity of pine-wiregrass sa­ Schwarz, G.F. 1907. The Longleaf Pine in vannas of the Green Swamp, North Caroli­ Virgin Forest: A Silvical Study. Wiley, New na. Vegetatio 55:163-179. York. 135 pp. Skeen, J.N., P.D. Doerr, and D.H. Van Lear. 1993. Oak-hickory-pine forest. Pp. 1-33 in W.H. Martin, S.G. Boyce, A.C. Echternacht, eds., Biodiversity of the Southeastern Unit­ ed States: Upland Terrestrial Communities. Wiley, New York.

52 Natural Areas Journal Volume 23 (1),2003