sign in sign up
<<
Home , Carya pallida, Quercus nigra

Prior Prevalence of Shortleaf Pine-- Woodlands in the Tallahassee Red Hills Author(s): Andre F. Clewell Source: Castanea, 78(4):266-276. 2013. Published By: Southern Appalachian Botanical Society DOI: http://dx.doi.org/10.2179/13-022 URL: http://www.bioone.org/doi/full/10.2179/13-022

BioOne (www.bioone.org) is a nonprofit, online aggregation of core research in the biological, ecological, and environmental sciences. BioOne provides a sustainable online platform for over 170 journals and books published by nonprofit societies, associations, museums, institutions, and presses. Your use of this PDF, the BioOne Web site, and all posted and associated content indicates your acceptance of BioOne’s Terms of Use, available at www.bioone.org/page/ terms_of_use. Usage of BioOne content is strictly limited to personal, educational, and non-commercial use. Commercial inquiries or rights and permissions requests should be directed to the individual publisher as copyright holder.

BioOne sees sustainable scholarly publishing as an inherently collaborative enterprise connecting authors, nonprofit publishers, academic institutions, research libraries, and research funders in the common goal of maximizing access to critical research. CASTANEA 78(4): 266–276. DECEMBER Copyright 2013 Southern Appalachian Botanical Society

Prior Prevalence of Shortleaf Pine-Oak-Hickory Woodlands in the Tallahassee Red Hills

Andre F. Clewell* Tall Timbers Research Station, 13093 Henry Beadle Drive, Tallahassee, Florida 32312

ABSTRACT Shortleaf pine-oak-hickory woodlands provided the principal vegetation cover in the Tallahassee Red Hills prior to land clearing for plantation agriculture in the 19th century. Ample historical documentation and extant remnants of that community, including old-growth, support this conclusion. This woodland was maintained by surface fires and consisted principally of open stands of shortleaf pine, post oak, Spanish oak, black oak, mockernut hickory, and dogwood. The species- diverse and predominantly herbaceous ground cover was dominated by grasses, legumes, and composites. Coppice sprouting of and shrubs after fires was common. In the absence of fire, shortleaf pine-oak-hickory woodlands convert to oak-hickory forest with similar species composition and loss of herbaceous species. Within the past 130 years, nearly all stands of shortleaf pine-oak-hickory woodlands and oak-hickory forests have been extirpated or compromised beyond recognition by intrusions of offsite tree species that are typical of moist soils at less elevated landscape positions. Key words: Apalachee, climax, ecotone, , magnolia-beech, mesophication, Tallahassee Red Hills, Tall Timbers.

INTRODUCTION The Tallahassee Red other regions in their scarcity or absence of Hills region was mapped by Roland Harper saw-palmetto and wire-grass among the (1914) as an 880 km2 physiographic unit of undergrowth. Both short- and long-leaf northern Leon County, Florida, lying north of the pine forests are subject to occasional fires. Cody scarp and extending a short distance into On some of the hillsides and richer uplands adjacent Grady and Thomas counties, Georgia dense hardwood forests with considerable (Figure 1). Harper characterized the Tallahassee humus can be seen, and the branches and Red Hills (TRH) by their deep deposits of red creeks are bordered by wet or sandy clay and rather hilly topography in which swamps. (p. 271) streams were scarce and lakes and ponds common. Harper (1914) described upland vege- Species listed by Harper (1914, with updated tation in the TRH: nomenclature) that occur in ‘‘uplands’’ or ‘‘dry woods’’ included P. echinata (Chapm. ex En- The drier uplands seem to have been gelm.) Vasey ex Sarg. (shortleaf pine), Cornus covered originally with comparatively open florida L. (flowering dogwood), forests of short-leaf pine (Pinus echinata), Michx. (Spanish or southern red oak), Carya red oak, hickory, dogwood, etc. Consider- tomentosa Nutt. (mockernut hickory), Carya able areas of this forest still remain, though glabra (Mill.) Sweet (pignut hickory), Q. stellata a good deal of it may be second growth. On Wangenh. (post oak), Q. velutina Lam. (black sandier soils near the center of the region oak), Nyssa sylvatica Marshall (blackgum), and there are limited areas (perhaps several Oxydendrum arboreum (L.) DC. (sourwood). In hundred acres) of long-leaf pine forest, this paper, I call this vegetation the shortleaf differing from the typical piney woods of pine-oak-hickory (SPOH) woodland community. Shortleaf pine-oak-hickory woodland is a pyro- *email address: [email protected] Received May 24, 2013; Accepted September 6, 2013. genic community, which, in the absence of fire, DOI: 10.2179/13-022 will convert to a closed canopy oak-hickory 266 2013 CLEWELL: SHORTLEAF PINE-OAK-HICKORY WOODLANDS 267

vegetation that occur elsewhere in the South- east, principally inland towards the continental interior. Beginning in the late 19th century in response to land usage, offsite tree species from less elevated positions in the landscape invaded the remaining original open woods described by Harper (1914) and eventually compromised the species composition of nearly all of them beyond recognition. Most offsite species were typical of flood plains and other sites that approach wetland status. They included Acer rubrum L. (red maple), Liquidambar styraciflua L. (sweet- gum), Pinus taeda L. (loblolly pine), Michx. (diamond-leaf oak), Q. nigra L. (water oak), and Q. virginiana Mill. (live oak). Other offsite species were typical of magnolia-beech forest (Fagus grandifolia Ehrh., Figure 1. The Tallahassee Red Hills according to Harper L.), particularly Prunus (1914) superimposed on a map by Griffith et al. (2001) showing the western end of Level IV ecoregion 65o. The serotina Ehrh. (black cherry) and Magnolia location of Tall Timbers Research Station is indicated by grandiflora. I call this spontaneously occurring ‘‘TT.’’ secondary forest the water oak-sweetgum-lob- lolly pine community, after three of its most forest. The inclusion of Oxydendrum arboreum abundant and characteristic species. Several in the SPOH woodland community, as intimated upland species contributed to this community by Harper (1914), is disputable; sourwood on upland sites, particularly P. echinata, C. occurs on relatively steep gradients leading from florida, and Quercus hemisphaerica W. Bartram SPOH woodlands to the heads of springs and ex Willd. (laurel oak), which either persisted seeps, and it is better considered an ecotonal after offsite colonization began or established species at the upland border with forested seedlings concurrently with offsite species. wetlands. Quercus hemisphaerica is becoming increasing- The TRH region lies within the western one- ly abundant in the TRH on unburned sites with third of the Tallahassee Hills/Valdosta Limesink well drained soils. Level IV ecoregion (Griffith et al. 2001, mapping Trees that contributed to the water oak- unit 65o). The TRH are commonly conceived to sweetgum-loblolly pine community were spared extend northward at least to Thomasville and from land clearing in the 19th century, because eastward to the Aucilla River, but soils in these the land where they grew was too moist for outlying areas are not as consistently clayey. The agricultural purposes or was protected from TRH form part of a larger physiographic unit lethal fires. Their establishment on abandoned called the Tallahassee Hills Northern Highlands agricultural land was facilitated by the absence consisting of the dissected sedimentary remains of competing trees (Clewell 2011). Upland tree of Neogene fluvial, and delta plain, deposits species were at a disadvantage in colonizing (Schmidt 1997). Upland soils belong to the abandoned agricultural fields, because land Norfolk-Ruston-Orangeburg association (Beck- clearing for agriculture had substantially re- enbach and Hammett 1962). A preponderance of duced the cover of SPOH woodlands that could broadleaved hardwood tree growth distinguishes serve as seed sources. Once established on the TRH from vegetation in surrounding regions, upland sites, water oak-sweetgum-loblolly pine which are, or were formerly typified by, longleaf forests persist indefinitely (Van Lear 2004) by pine-wiregrass savannas (Pinus palustris Mill., means of mesophication (Nowacki and Abrams Aristida stricta Michx.) (Harper 1914, Clewell 2008). Mesophication occurs when fire-main- 1986). Broad regions covered by these savannas tained open SPOH woodlands become closed- isolate the TRH and its predominant SPOH canopied in the absence of fire, which favors vegetation from lands with similar soils and colonization by shade-tolerant and fire-sensitive 268 CASTANEA VOL.78 species that produce a less flammable fuel Southeast, collectively called Seminoles, assem- bed. This process promotes rapid compositional bled in the area in small numbers but were and structural changes without ecological ante- driven out by General Andrew Jackson in 1818. cedent. By the latter half of the 20th century, Spain ceded Florida to the in 1821 Harper’s descriptions of original vegetation were (Tebeau 1971). forgotten, and ecological opinion accepted for- The TRH were essentially unoccupied from ests dominated by offsite species as natural 1704 until the U.S. Territory of Florida was stages of succession leading towards a climax established in 1824 with Tallahassee as its state in which SPOH species were of minor capital city. Abandoned Apalachee old fields importance relative to offsite species. spontaneously became forested. Without human This paper resurrects Harper’s conception of residents, the incidence of fire was likely much SPOH woodland, documents it with evidence reduced, and disturbed soils would have further from 18th and 19th century literature, discerns diminished the spread of fires. Beginning in offsite tree species, and rejects the concept of a 1824, planters and numerous African slaves rigidly predictable climatic climax community. quickly established 71 large plantations between Knowledge of the history of land use in the TRH Tallahassee and Thomasville. Forests that colo- is essential for understanding how current nized Apalachee old fields following Moore’s vegetation replaced SPOH woodlands and its raid were cleared to grow cotton, corns, and associated oak-hickory forests. sweet potatoes. The plantation era abruptly came to a close after 1875, when cotton prices Land Use Considerations fell precipitously in an economic depression The region was occupied by hunters and gathers (Paisley 1968). Forests spontaneously colonized until maize was introduced sometime after 1100 abandoned farmlands, except for small patches A.D. (Hudson 1997, p. 124). Apalachee Indians of land that were farmed largely for subsistence grew maize, also beans, pumpkins and sunflow- by tenant farmers. Most former plantations were ers extensively in the TRH eastward to the purchased by northern industrialists who be- Aucilla River. Regions now comprising Grady came winter residents and hunted bobwhite and Thomas counties north of the TRH were not quail. Large tracts of former plantations continue farmed, and that landscape was used for hunting to be held as private, fire-maintained hunting and as a protective buffer from bellicose Lower preserves. Creek Indians. The Apalachees lacked metal implements. Tree removal, land clearing, and Ecological Considerations cultivation was accomplished with tools made The first theoretical ecological paper to be from conch shells, bone, and (Brown published for the TRH was that of Laura Gano 1994). (1917) who proposed that magnolia-beech forest In 1539–1540, the expeditionary force of comprised the climax vegetation in the Red Hills. Hernando de Soto, consisting of approximately Her characterization conformed to the recently 1,000 soldiers and other personnel and their conceived theory of climatic climax, which livestock, encamped at present-day Tallahassee ignored fire as positive influence on vegetation and lived off the largess of Apalachee agricul- development. Chapman (1932) argued to the tural produce for five months. The large size of contrary, that the longleaf pine cover type was this occupation attests to the extent of Apala- climax in southeast coastal plain regions, be- chee agriculture. The size of the Apalachee cause it ‘‘has reached a stage of permanent population at its apotheosis has been the subject equilibrium’’ under the constant factor of fire (p. of wide speculation and likely exceeded 10,000 333). inhabitants—perhaps several times that number. Herman Kurz (1944) accepted magnolia-beech Several Spanish missions relied on Apalachee as climax in the TRH, and he proposed that oak- agriculture in the 17th century; however, the hickory forest was subclimax, based on descrip- Apalachee population dwindled from cultural tions of study sites near Tallahassee. He did not disruption caused by warfare and European- consider the longleaf pine cover type in his borne diseases. A raid in 1704 led by Colonel analysis. Previously, Kurz (1938) noted that Thomas Moore from South Carolina drove the magnolia-beech forest occupied the lowest Spanish and Apalachees from the region. Refu- position on river bluffs, relative to other forest gees from culturally disrupted tribes in the communities. His contention that magnolia- 2013 CLEWELL: SHORTLEAF PINE-OAK-HICKORY WOODLANDS 269 beech was ‘‘climax’’ apparently assumed the Rediscovery of SPOH Woodlands eventual dominance of that community at more My introduction to SPOH woodland came in elevated landscape positions. Schwartz (1994) 1967, after I was well acquainted with secondary suggested that ‘‘It was later ecological studies, water oak-sweetgum-lobolly pine forests on perhaps influenced by Clementsian views of abandoned agricultural land. I examined an plant communities and succession, that seem upland site near Anders Branch on property to have obfuscated the importance of pine in the owned by the Tall Timbers Research Station vegetation of northern Florida’’ (p. 703). (Figure 1). The vegetation consisted of open The magnolia-beech notion of climax did not woodland of P. echinata, Quercus stellata, Q. go unchallenged. Herbert Stoddard (1931) and falcata, Q. velutina, Q. alba, C. tomentosa, Edwin V. Komarek, Sr. (1966), among others, Nyssa sylvatica, and C. florida (Clewell 1986). demonstrated the primary role of fire and the The grass-dominated ground cover was dense mostly neglected importance of longleaf pine- and diverse. The site had been annually winter- wiregrass savannas in the TRH region. The burned for at least the past 20 years. Annual ring widespread influence of Stoddard and Komarek counts from increment borings revealed an age arose not so much from their publications as of 120 years for P. echinata, 99 years for Q. through demonstrations of controlled burns and stellata, and in excess of 120 years for heart- their ecological effects for the benefit of large rotted Q. velutina. Although I had seen occa- numbers of natural resource professionals who sional individuals of these trees growing with the attended conferences at the Tall Timbers Re- typical loblolly pine-sweetgum-water oak forests search Station in the 1960s. These conferences on abandoned plantation land, I had not previ- were instrumental in changing national policy to ously witnessed a woodland community consist- accept fire as a positive ecological driver. ing only of these species in the TRH. This In spite of inroads on ecological thought by description matched that of Harper (1914) for Stoddard and Komarek, the notion of a hard- SPOH woodlands. The age of the trees suggested wood forest climax persisted, principally due to that the land had not been cultivated during the an influential paper by Elsie Quarterman and plantation era and that the vegetation was Catherine Keever (1962) who described the ‘‘original.’’ southern mixed hardwood forest type. This I discovered several more small woodlands of community was conceived as occurring through- Harper’s description in the TRH (Clewell 1986). I out coastal plain regions of the Southeast. Tree searched the archives of the Florida Department species composition for this type was locally of State in Tallahassee and read the diaries of variable. American beech and southern magnolia five teams of surveyors who conducted the were among its potential dominant species. initial public lands survey of the TRH in 1824, Quarterman and Keever (1962) rejected magno- available at the Florida Division of Archives, lia-beech forest as ‘‘climax,’’ because the mag- History, and Records Management. These sur- nolia-beech community was restricted to moist veyors listed the kinds of trees at turning points soils of river bluffs, ravines and lake margins. and other locations along section lines. Tree lists Other potentially dominant species of the south- for some locations were detailed, with canopy- ern mixed hardwood climax community includ- forming trees listed separately from under- ed the same tree species that were noted above growth trees. The species of pines were rarely for the water oak-sweetgum-loblolly pine com- distinguished, and diaries for two centrally munity. located sections of land had been lost. Nonethe- By the 1980s, ecologists were abandoning the less, I was able to make a vegetation map from climax concept, at least as classically conceived. the data (Clewell 1980), and the SPOH cover Instead, fire and hurricane-force storm events type occurred at 239 locations. The Anders (Batista and Platt 2003) were recognized as Branch region was entirely mapped as SPOH. principal determinants of vegetation. The demise Schwartz (1994) examined diaries of public land of climax theory left a void in the ecological surveyors for a broader area of northern Florida conception of TRH ecosystems for lack of and confirmed my interpretations. consensus as to what constituted the original Earlier, I suggested that the original upland vegetation prior to 19th century plantation vegetation had been longleaf pine-wiregrass agriculture. savanna and that SPOH was an artifact of 270 CASTANEA VOL.78 agriculture on abandoned fields of the Apalachee composed of sucker saplings [coppice] of the Indians (Clewell 1980). I have since abandoned oak and hickory; this description of land is that explanation in favor of one proposed by generally disposed on the exterior edges of the Kevin Robertson of Tall Timbers Research high hammocks [oak-hickory and/or magnolia- Station (pers. comm. 2010) which recognizes beech forest], and separate them from the the shortleaf pine-oak-hickory community as [longleaf] pine lands’’ (p. 89). This observation originally occurring over large areas on sandy provides succinct evidence that SPOH wood- loam soils. Longleaf pine-wiregrass savannas, lands were subject to burning, more so than wherever they occurred, occupied a more other forest communities. elevated position on the tops of hills and ridges The fourth account was that of Colonel John capped by internally well-drained sands. Lee Williams (1837), who was commissioned to Several accounts corroborated the original help select the territorial capital of Florida. His prevalence of SPOH in the TRH. The earliest book, which was probably written to attract consisted of an annotated map by the English potential land owners, contained lengthy de- topographic engineer, Joseph Purcell, who in scriptions of vegetation. Williams described how 1778 was engaged in mapping the road from the Tallahassee region was covered with ‘‘excel- Pensacola to St. Augustine, Florida (Boyd 1938). lent brown soil, and crowned with wide spread- The road passed through northern Leon County. ing and tall . ...This kind of land, Purcell indicated four cover types with symbols in some places, extends into Georgia; in other for ‘‘pine lands,’’ ‘‘oak lands,’’ ‘‘old fields,’’ and parts the pine barrens [longleaf pine savannas] ‘‘swamp lands.’’ Nearly all of the TRH was make large indentations from various directions’’ covered by oak lands including the location of (1837, p. 88). Tall Timbers Research Station (Figure 1). He The final account was written by University of wrote ‘‘Appalatchi [sic] Old Fields’’ across a large Alabama Professor Eugene A. Smith (1884), who central area of the TRH. In his caption he thoroughly described upland vegetation in the described the growth of oak lands as consisting Red Hills as part of an assessment of the cotton of ‘‘Oaks, Hickaries and Mulberries,’’ and that lands of Florida for the 10th census of the United ‘‘Appalatchi old fields . . . are chiefly grown up States. Smith described brown loam lands, with with Oaks, Hickaries, Mulberries, Pines and oak, hickory, and shortleaf pine occurring as a small Canes, and in places hilly’’ (Boyd 1938, continuous body of upland covering >3,000 km2 Plate 5). Mulberries (presumably Morus rubra L.) are minor trees in magnolia-beech forests along the Georgia state line in Gadsden, Leon, today but may have been widely planted by Jefferson, and northeastern Madison counties, Apalachees. Florida. Smith also wrote: ‘‘The timber compris- The next account was that of Captain Hugh es the usual varieties of upland oaks, such as Young (1818), who was attached to General post, red [probably Q. velutina], and Spanish . . Andrew Jackson’s army as a topographic engi- and hickory, short-leaf pine’’ and that these lands neer. Young’s comprehensive geographic report in Florida ‘‘are usually deemed the best farming contained annotated maps and detailed descrip- areas in the state’’ (1884, p. 18). tions of the vegetation and lists of plant species None of these 18th and 19th century accounts for specific locations in the TRH. Young (1818) by Purcell, Young, Vignoles, Williams, and Smith stated that ‘‘oak and hickory’’ occupied the best recognized water oak-sweetgum-loblolly pine potential farmland in the region. He recognized forest that currently prevails in regional uplands that some ‘‘oak and hickory’’ lands consisted of (Clewell 1986, 2011). Until the 19th century, P. open grassy woodland on account of frequent taeda, Quercus nigra, L. styraciflua, and Q. fires and other parcels that were densely virginiana were restricted to lowland habitats. forested where fires were generally absent. Pinus taeda is currently the principal pine The third account is that of Charles Vignoles species on ex-arable uplands. This tree was (1823) who was also a topographic engineer who originally confined to ravines and flood plains visited the TRH and said that ‘‘The oak and (Williams 1837, p. 39) and only colonized hickory lands produce almost exclusively those abandoned farmland relatively recently (Chap- two kinds of forest trees, with occasionally man 1942). Williams (1837, p. 53) listed Q. nigra gigantic pines: the underbrush is generally as a swamp species, and it was probably 2013 CLEWELL: SHORTLEAF PINE-OAK-HICKORY WOODLANDS 271 confined there on account of its susceptibility to dispersed than the generally more fecund and fire (Fowells 1965). smaller fruited tree species typical of magnolia- Liquidambar styraciflua was not mentioned beech and flood plain forests. These qualities by Young (1818). Vignoles did not mention it help explain why most SPOH species lacked either, except possibly in an appendix using the abundance in secondary upland forests. name ‘‘myrrh’’ among the trees that grew with Fagus grandifolia (Vignoles 1823, p. 9). Williams Community Characterization (1837) did not include L. styraciflua in his ample Table 1 lists some of the more common and lists of species for the several forest types he characteristic species of trees, shrubs, grasses, described, and his references to ‘‘gum’’ in the and forbs that were typical of SPOH woodlands text (e.g., p. 88) probably referred to species of as shown in Figure 2. These species were Nyssa (Schwartz 1994). Liquidambar styraci- selected from those occurring in several SPOH flua is a large, distinctive, and useful tree that woodlands described by Clewell (1986) or noted would have warranted notice if it were common in Clewell (1985). Table 1 emphasizes species of in the early 19th century. Authors of that period SPOH woodlands that help distinguish them did not mention L. styraciflua in their descrip- from other types of TRH vegetation. Detailed tions of swamps and forested flood plains, where quantitative studies of species composition of it is presently common and sometimes domi- this vegetation type in relation to soil type and nant. Smith (1884) mentioned that it grew on other factors are currently being conducted by slopes. Schwartz (1994) concluded from Public Kevin Robertson and colleagues at Tall Timbers Land Survey records that L. styraciflua was a Research Station. relatively rare tree in the early 19th century and Principal trees are P. echinata, Q. stellata, Q. did not become common and ‘‘weedy’’ until falcata, Q. velutina, C. tomentosa, and C. nearer the end of that century. florida (Table 1). Nyssa sylvatica occurs fre- Quercus virginiana is ubiquitously distribut- quently but with less abundance. Quercus alba ed throughout the TRH and is the most obvious, and sometimes C. glabra occur with moderate if not the most abundant, tree species in frequency at lower positions along elevation Tallahassee and its suburbs. The prevalence of gradients. Quercus hemisphaerica sometimes Q. virginiana is evidently a recent phenomenon. extends into SPOH woodlands from its preferred Young (1818) mentioned Q. virginiana several habitat on steep ravines and upper slopes of times, always in respect to a narrow zone of river bluffs where it is a dominant species (Kurz forests which he called ‘‘live oak swamps’’ that 1938, Clewell 1986, Kwit et al. 1998, Gibson bordered Gulf-coastal tidal marshes and the 1992). Carya pallida (Ashe) Engl. & Graibn., lower reaches of rivers. Hawkins (1848) reported which grows commonly with Q. hemisphaerica Q. virginiana only as trees encircling ponds, on river bluffs, may also extend into adjacent which is a landscape position that the species SPOH woodlands. Two short-lived and fire- occupies today at the boundary between wet- intolerant trees may also be associated with lands and uplands. Williams (1837) wrote of SPOH woodland when intervals between fires ‘‘wide spreading oaks’’ in Tallahassee, which are protracted, Malus angustifolius (Aiton) would seem to indicate Q. virginiana, but none Michx. (Harper 1914) and Sassafras albidum of his detailed species lists included that species. (Nutt.) Nees. The obvious conclusion is that water oak- Herbaceous ground cover in SPOH woodlands sweetgum-loblolly pine forest did not occur consists primarily of grasses (Poaceae) in which originally on uplands and that this community numerous forbs are interspersed. Legumes (Fa- on upland sites is an artifact of land use. Harper baceae) and composites (Asteraceae) are well (1914) and Kurz (1944) described forests con- represented among forbs. The abundance and taining mixtures of SPOH and offsite species exceedingly rich diversity of legumes suggests shortly after the abandonment of plantation the possibility that Apalachee Indians were agriculture in 1875. With the exceptions of P. enriching the abundance of native legume echinata, Q. hemisphaerica, and C. florida, species in SPOH woodlands by their manage- most trees of the SPOH community do not ment practices. Native Americans were known produce seeds in abundance annually, and their to harvest native legume seeds as a food relatively large fruits would be less effectively resource (Anderson 2005). 272 CASTANEA VOL.78

Table 1. Some characteristic species of shortleaf Table 1. Continued pine-oak-hickory woodlands Composites Trees Chrysopsis mariana (L.) Elliott (Mill.) Sweet Elephantopus elatus Bertol. Carya pallida (Ashe) Engl. & Graebn. Fleischmannia incarnata (Walter) R.M.King & H.Rob. Carya tomentosa Nutt. Helianthus hirsutus Raf. Cornus florida L. Liatris graminifolia Willd. Malus angustifolius (Aiton) Michx. Melanthera nivea (L.) Small Nyssa sylvatica Marshall Polymnia uvedalia (L.) L. Oxydendrum arboreum (L.) DC. Rudbeckia hirta L. Pinus echinata (Chapm. ex Engelm.) Vasey ex Sarg. Sericocarpus tortifolius Nees Quercus alba L. Solidago arguta Aiton Quercus falcata Michx. Symphyotrichum urophyllum (Lindl.) G.L.Nesom Quercus hemisphaerica W.Bartram ex Willd. Verbesina aristata A.Heller Quercus margaretta Ashe ex Small Vernonia altissima Nutt. Munchh.¨ Other Forbs Quercus stellata Wangenh. Aureolaria flava (L.) Farw. Lam. Cyperus retrofractus (L.) Torr. Sassafras albidum (Nutt.) Nees Dyschoriste oblongifolia Kuntze Shrubs and Woody Vines Euphorbia pubentissima Michx. Castanea alnifolia Nutt. Galium pilosum Aiton Ceanothus americanus L. Gentiana villosa L. Celtis tenuifolia Nutt. Hexalectris spicata (Walter) Barnhart Collinsonia serotina Walter Ipomoea pandurata G.Mey. Crataegus pulcherrima Ashe Lobelia puberula Nutt. Crataegus uniflora Munchh.¨ Onosmodium virginianum A.DC. Quercus pumila Walter Pteridium aquilinum (L.) Kuhn Rhus toxicarium Salisb. Rhynchospora globularis (Chapm.) Small Smilax smallii Morong Ruellia carolinensis Steud. Vaccinium arboreum Marshall Salvia azurea Michx. Vaccinium stamineum L. Scleria pauciflora Muhl. ex Willd. Viburnum rufidulum Raf. Smilax lasioneuron Hook. Grasses Aristida lanosa Muhl. ex Elliott Andropogon gerardii Vitman Andropogon gyrans Ashe Andropogon virginicus L. Figure 3 is a photo of a young oak-hickory Dichanthelium commutatum (Schult.) Gould forest that is developing near a small stream at Gymnopogon ambiguus (Michx.) Britton, Sterns the Wade Tract on Arcadia Plantation in Thomas & Poggenb. County (3084504800 N, 8480105000 W), managed by Saccharum alopecuroides (L.) Nutt. Schizachyrium tenerum Nees the Tall Timbers Research Station. Tree species Sorghastrum elliottii (Mohr) Nash composition is similar to that shown in Figure 2. Sorghastrum nutans (L.) Nash The presence of P. echinata suggest that this Sporobolus junceus (Michx.) Kunth was formerly a SPOH woodland, or an ecotone Tridens flavus (L.) Hitchc. Legumes with longleaf pine-wiregrass savanna, and these Amphicarpaea bracteata (L.) Fernald pines will eventually succumb and likely not be Centrosema virginianum (L.) Benth. replaced, owing to the inability of their seedlings Clitoria mariana L. to establish beneath dense forest cover. Hard- Dalea carnea Poir. Desmodium floridanum Chapm. wood leaf litter replaces the grass and forb cover Desmodium nudiflorum Muhl. ex Willd. seen in Figure 2. The species composition of the Desmodium paniculatum (L.) DC. forest shown in Figure 3 is similar to that Erythrina herbacea L. described by Williams (1837) for the Tallahassee Galactia volubilis (L.) Britton Lespedeza hirta Hornem. region, quoted earlier. Under original conditions, Lespedeza intermedia (S.Watson) Britton I presume that SPOH woodlands burned at Lespedeza procumbens Michx. various fire return intervals and intensities, some Lespedeza virginica (L.) Britton as frequently as longleaf pine-wiregrass savan- Phaseolus polystachios (L.) Britton, Sterns & Poggenb. Rhynchosia tomentosa Hook. & Arn. nas and others irregularly and infrequently. The Schrankia microphylla J.F.Macbr. photos in Figures 2 and 3 show the extremes, Strophostyles umbellata (Muhl. ex Willd.) Britton and most any intermediate state may have Tephrosia virginiana (L.) Pers. occurred. 2013 CLEWELL: SHORTLEAF PINE-OAK-HICKORY WOODLANDS 273

Figure 2. Frequently burned shortleaf pine-oak-hickory woodland on an upland site near Anders Branch at Tall Timbers Research Station and currently burned on a two-year fire return interval. protruding above the grasses are coppice sprouts of hardwood trees and shrubs. Figure 3. Unburned, immature oak-hickory forest at the Wade Tract.

Ecotones with Longleaf Pine-Wiregrass by long-leaf pine and black-jack, there being all Savannas graduations between pure oak and hickory Smith (1884) described broad ecotones between uplands and the genuine pine woods’’ (p. 18). SPOH woodlands and longleaf pine-wiregrass Smith repeated this observation for regions of savannas: ‘‘As this class of soil [brown loam] Florida west of the Apalachicola River; he grades into the sandier varieties, so the short- observed that with ‘‘gradual improvement to leaf pine and upland oaks are gradually replaced the soil [in longleaf pine savannas], black-jack, 274 CASTANEA VOL.78 post, and other species of oak, with short-leaf eliminating the once abundant herbaceous pine, [form] a gradual transition into oak ground cover. The site is converting to oak- uplands’’ (p. 19). I have observed this transition hickory forest and will serve as welcome legacy in several places north of the Red Hills (Clewell representing the forest that Williams (1837) 1986) within ecoregion 65o (Figure 1), where described for Tallahassee, but it is no longer a longleaf pine-wiregrass savannas prevail. Short- SPOH woodland community. The Florida Natu- leaf pine-oak-hickory woodlands occur at lower ral Areas Inventory (FNAI) recognizes SPOH elevations along topographic gradients with woodlands, which it designates as ‘‘upland broad transition zones separating them from mixed woodland’’ (FNAI 2010) and recognizes longleaf pine-wiregrass savannas. Quercus mar- as a rare natural community in Florida (G2S2). ilandica grows as a midstory tree in this ecotone The Florida Natural Areas Inventory records where the subsoil is clayey. Quercus margaretta known locations of ‘‘upland mixed woodland’’ in is a common midstory tree on deep, sandy soils its database, and these listing are considered in in these ecotones (Godfrey 1988). evaluations of properties proposed for state In the TRH, longleaf pine savannas were acquisition under the Florida Forever Program. notably less extensive than in surrounding This effort to search for and protect SPOH regions, and wiregrass was curiously rare. woodlands is encouraging because of the com- Harper (1914) suggested that the scarcity of pelling evidence for the prevalence of these longleaf pine in Florida may have been related to woodlands prior to the 19th century. Doubt the intensity of Apalachee agriculture, and remains as to the nature of oak-hickory forest if Clewell (1989) mentioned that shallow-rooted fire is removed. Will it maintain itself as ‘‘pure’’ Aristida stricta could easily be removed with primitive farming implements. The Apalachees oak-hickory, or will it eventually be overtaken by likely grew crops in ecotones between pine species typical of contiguous magnolia-beech savannas and SPOH woodlands and perhaps on forest? Perhaps continued search will reveal an longleaf pine savannas. After abandonment, as yet undiscovered forest that will provide SPOH woodlands may have colonized these answers. A long-term and well-conceived exper- fields, as Purcell’s map from 1778 indicated iment in ecological restoration could also gen- (Boyd 1938). erate an answer to this question. Conservation ACKNOWLEDGMENTS I am grateful to Shortleaf pine-oak-hickory woodland ranks Ann Johnson, Kevin Robertson, and John Tobe among the most endangered ecosystems in whose reviews of earlier drafts contributed Florida. Any existing stands deserve immediate appreciably to the final preparation of this protection. The SPOH woodland near Anders paper. Branch at Tall Timbers Research Station is currently well managed and may be afforded LITERATURE CITED protection indefinitely. Several other relatively Anderson, M.K. 2005. Tending the wild: Native undisturbed stands of SPOH woodland occur on American knowledge and the management of private lands, most of them in Georgia north of California’s natural resources. University of the TRH, and some enjoy protection in conser- California Press, Berkeley, California. vation easements. Some stands of SPOH woodlands that I Batista, W.B. and W.J. Platt. 2003. Tree popula- examined earlier (Clewell 1986) have suc- tion responses to hurricane disturbance: syn- cumbed to development. One woodland at dromes in a south-eastern USA old-growth Welaunee Plantation northeast of Tallahassee forest. J. Ecol. 91:197–212. (308300N, 848090W) came into public ownership, Beckenbach, J.R. and J.W. Hammett. 1962. and an 18 ha tract was designated as the North General soil map of Florida. Florida Agricul- Florida Red Oak Woods within the Miccosukee tural Experiment Stations and Soil Conserva- Canopy Road Greenway, managed by the Leon tion Service. County Division of Parks and Recreation. The tract has not been burned recently, and the Boyd, M.F. 1938. A map of the road from shade and leaf litter produced by young hard- Pensacola to St. Augustine. Florida Historical wood trees growing from coppice sprouts are Quarterly 17:15–23. 2013 CLEWELL: SHORTLEAF PINE-OAK-HICKORY WOODLANDS 275

Brown, R.C. 1994. Florida’s first people: 12,000 Godfrey, R.K. 1988. Trees, shrubs, and woody years of human history, rev. ed. Pineapple vines of Northern Florida and adjacent Geor- Press, Sarasota, Florida. gia and Alabama. University of Georgia Press, Chapman, H.H. 1932. Is the longleaf type a Athens, Georgia. climax? Ecology 13:328–334. Griffith, G.E., J.M. Omernik, J.A. Comstock, S. Lawrence, G. Martin, A. Goddard, V.J. Hulcher, Chapman, H.H. 1942. Management of loblolly and T. Foster. 2001. Ecoregions of Alabama pine in the pine-hardwood region in and Georgia [color poster with map, descrip- and in west of the River. tive text, summary tables, and photographs]. Yale University School of Forestry Bulletin Reston, Virginia, U.S. Geological Survey (map No. 49. New Haven, Connecticut. scale 1:1,700,000). Clewell, A.F. 1980. The vegetation of Leon Harper, R.M. 1914. Geography and vegetation of County, Florida. p. 386–441. In: Tesar, L.D. northern Florida. p. 266–279. In: Florida (ed.). The Leon County bicentennial survey Geological Survey, 6th Annual Report. Talla- project: An archaeological survey of selected hassee, Florida. portions of Leon County, Florida. Florida Division of Archives, History and Records Hawkins, B. 1848. A sketch of the Creek Country Management, Bureau of Historic Properties, in the years 1798 and 1799. Reprint edition by Misc. Project Report. Series No. 49, Tallahas- the Georgia Historical Society, Savannah, see, Florida. Georgia, 1982. Clewell, A.F. 1985. Guide to the vascular plants Hudson, C. 1997. Knights of Spain, warriors of of the Florida Panhandle. Florida State Uni- the sun: Hernando de Soto and the south’s versity Press/University Presses of Florida, ancient chiefdoms. University of Georgia Tallahassee, Florida. Press, Athens, Georgia. Clewell, A.F. 1986. Natural setting and vegeta- Komarek, E.V., Sr. 1966. The meteorological tion of the Florida Panhandle: An account of basis for fire ecology. Proc. Annual Tall the environments and plant communities of Timbers Fire Ecol. Conf. 5:85–125. northern Florida, west of the Suwannee River. Kurz, H. 1938. A physiographic study of the tree COESAM/PDEI-86/001 U.S. Army Corps of associations of the Apalachicola River. Proc. Engineers, Mobile, Alabama District. Florida Acad. Sci. 3:78–90. Clewell, A.F. 1989. Natural history of wiregrass Kurz, H. 1944. Secondary forest succession in the (Aristida stricta Michx., Gramineae). Nat. Tallahassee Red Hills. Proc. Florida Acad. Sci. Areas J. 9(4):223–233. 7:3–42. Clewell, A.F. 2011. Forest succession after 43 Kwit, C., M.W. Schwartz, W.J. Platt, and J.P. years without disturbance on ex-arable land, Geaghan. 1998. The distribution of tree species northern Florida. Castanea 76:386–394. in steepheads of the Apalachicola River bluffs, Florida Natural Areas Inventory (FNAI). 2010. Florida. J. Torrey Bot. Soc. 125:309–318. Guide to the natural communities of Florida, Nowacki, G.J. and M.D. Abrams. 2008. The 2010 edition (http://www.fnai.org). FNAI, Tal- demise of fire and ‘‘mesophication’’ of forests lahassee, Florida. in the eastern United States. BioScience 58: Fowells, H.A. 1965. Silvics of forest trees of the 123–138. United States. Agricultural Handbook 71. Paisley, C. 1968. From cotton to quail. University USDA, Forest Service, Washington, DC. of Florida Press, Gainesville, Florida. Gano, L. 1917. A study in physiographic ecology Quarterman, E. and C. Keever. 1962. Southern in northern Florida. Botanical Gazette 63:337– mixed hardwood: climax in the southeastern 372. coastal plain, USA. Ecol. Monogr. 32:164–187. Gibson, D.J. 1992. Vegetation–environment rela- Schmidt, W. 1997. Geomorphology and physiog- tionships in a southern mixed hardwood raphy of Florida. p. 1–12. In: A.R. Randazzo forest. Castanea 57:174–189. and D.S. Jones (eds.). The geology of Florida. 276 CASTANEA VOL.78

University Presses of Florida, Gainesville, vice, Southern Research Station, Asheville, Florida. North Carolina. Schwartz, M.W. 1994. Natural distribution and Vignoles, C. 1823. Observations upon the Flori- abundance of forest species and communities das. E. Bliss & E. White, New York. Reprinted in northern Florida. Ecology 75:687–705. in 1977 by The University of Florida Press/ University Presses of Florida, Gainesville, Smith, E.A. 1884. Report on the cotton produc- Florida. tion of the State of Florida with an account of the general features or the State. U.S. Williams, J.L. 1837. The territory of Florida. A.T. Bureau of Census, 10th Census of the U.S. 6: Goodrich, New York. Reprinted in 1962 by The 175–257. University of Florida Press, Gainesville, Flor- ida. Stoddard, H.L., Sr. 1931. The bobwhite quail, its habits, preservation and increase. Charles Young, H. 1818. A topographical memoir on East Scribner’s Sons, New York, New York. and West Florida with itineraries of General Jackson’s army, 1818. p. 292–336. In: Records Tebeau, C.W. 1971. A history of Florida. Univer- of Reports, July 3, 1812–October 4, 1823, US sity of Miami Press, Coral Gables, Florida. Army Corps of Engineers, Washington, DC. Van Lear, D.H. 2004. Upland oak ecology and Reprinted serially in The Florida Historical management. p. 65–71. In: M.A. Spetich (ed.). Society Quarterly Vol. 13(1):16–50 (1934); Vol. Upland oak ecology symposium: History, 13(2):82–104 (1934); and Vol. 13(3):129–164 current conditions, and sustainability. General (1935). Introduction and annotations by Mark Technical Report SRS-73, USDA, Forest Ser- F. Boyd and Gerald M. Ponton.