Influence of Historic Upland Silviculture on the Composition of Ravine Forests along the Apalachicola River, , USA

RESEARCH ARTICLE ABSTRACT: The ravines along the Apalachicola River are recognized for their species-rich mixed hardwood forests. Although steep slopes may have limited logging, much of the surrounding sandhill and scrub uplands were extensively modified for silviculture in the late 1950's. We examined how contrasts in the intensity of historic upland silviculture have influenced present-day woody species composition in ravine slope forests. To maximize ravine differences due to upland silvicultural intensi­ ties while concurrently minimizing the potentially confounding influences of aspect and slope angle, • ravine sampling areas were delineated using a combination of historical air photos, field reconnaissance, and a digital elevation model. Ordination (principal coordinates analysis) and permutation-based tests Influence of Historic of group differences (multiple response permutation procedures) confirmed that where uplands were selectively-cut longleaf pinelands, composition on neighboring ravine slopes was dominated by three species: Quercus hemisphaerica Bartr., Pinus glabra Walt., and Quercus alba L. By contrast, where Upland Silviculture the uplands had undergone intensive mechanical site preparation for slash cultivation, only one species, Quercus hemisphaerica, was dominant. Intensive upland silviculture was also correlated with a decreased importance of Fagus grandi/alia, a species identified as a major constituent of presettiement on the Com position ravine forests. We discuss how fire suppression, geomorphic disturbance, and logging along upper ravine of Ravine slopes may explain these compositional differences. Index terms: Apalachicola, disturbance, fire suppression, land use legacies, Quercus hemisphaerica, Forests along the ravines Apalachicola River, INTRODUCTION elliottii Engelm.) silviculture expanded across much of the uplands surrounding the Florida, USA The species-rich mixed hardwood forests ravines. Intensive site preparation practices in the Apalachicola River ravines of the accompanying slash pine silviculture have Florida panhandle have been studied for been often suggested as a direct and indirect J. Anthony Stallins 1 well over a century (Chapman 1885, influence upon present-day ravine slope Florida State University Harper 1914, Kurz 1938, Kwit et al. forest composition (Schwartz 1990, Platt Department of Geography 1998). Much of this work has focused on and Schwartz 1990, Kwit et al. 1998), but Room 323 Bellamy Building the endemic and disjunct popula­ this hypothesis has not been examined in Tallahassee, FL 32306 USA tions of the ravines, which include two detail. We describe the variability in ravine globally endangered , the Florida woody tree and shrub species composition yew (Taxus fioridana Nutt.) and Torreya as it relates to upland land use history. Jennifer Griggs2 taxifolia Arnott (Schwartz et al. 1995). Clarifying the changes that may have been Undergraduate Honors Program These plant species, and others typical of initiated by past land use provides a foun­ Florida State University cool climates, were stranded within the dation for directing future research needs, Department of Geography mild microclimates in the river valley and setting conservation priorities, and focus­ tributary ravines following the last pulse of ing ecological restoration efforts (Black et • Pleistocene glaciation (James 1961). al. 1998, Kettle et al. 2000) . The protected, relict quality of ravine BACKGROUND vegetation reflects, in part, a unique eco­ system that has evolved independently The Apalachicola River begins at the of the surrounding uplands. However, borders of Alabama, Florida, and Georgia current ideas in disturbance theory and at the confluence of the Flint and Chat­ landscape ecology suggest a potential tahoochee Rivers. Along a 15-km stretch I Corresponding author: interaction between the ravines and their of the eastern shore of the Apalachicola in [email protected]; (850) 644-8385; uplands (Swan~on et al. 1988, Parker and Liberty County, Florida, fluvially-eroded fax: (850) 644-5913 Bendix 1996, Reiners and Driese 2001). ravines, or 'steepheads' , form a network of 2 Current address: Disturbance effects may propagate along steep-walled tributary valleys. These land­ Graduate Program in the School of Forest Resources & Environmental Science, gravitational gradients, linking vegetation forms are the result of spring seepage in the Michigan Tech, 1400 Townsend Drive, changes at lower elevations to upland land floor of the ravines, which undercuts over­ Houghton, MI 49931-1295 USA use (Nakamura et al. 2000). Past land use lying sands and causes tributary streams to practices may continue to influence com­ migrate inland by erosion (Rupert 1991). positional patterns and disturbance through This unique topography provides a refuge Natural Areas Journal 24:242-250 ecological legacies (Foster et al. 2003). within the surrounding xeric uplands for During the late 1950's, slash pine (Pinus plant species common to moister soils and

242 Natural Areas Journal Volume 24 (3),2004 cooler temperatures. Because of increasing longleaf pine (Pinus palustris Mill.) in in the 1800's. development pressures in the Florida pan­ the southeastern U.S. (Means 1996, Smith handle, ravine communities are classified et al. 2000), slash pine silviculture grew METHODS as imperiled by the Florida Natural Areas in economic importance and areal extent. Inventory (FNAI 1990). These early slash pine silvicultural methods Study Area were often associated with fire suppression Quercus hemisphaerica, Carya spp., Fagus and intensive mechanical site preparation Our study area was purchased in 1982 by grandifolia Ehrh., Magnolia grandiflora (Abrahamson and Hartnett 1990). Fire The Nature Conservancy and is known L., Quercus alba, and Pinus glabra are the suppression could be expected to aug­ as the Apalachicola Bluffs and Ravines dominant overstory species in the present­ ment the cover of fire-intolerant species Preserve (ABRP) (Figure 1). Current land day ravine forests. Important understory in the uplands, some of which may be management in the ABRP has focused on tree species include Osmanthus arnericanus able to germinate in the protected ravines restoring the uplands from rows of planted (L.) Benth. & Hook., Kalmia latifolia L., (Kwit et. al. 1998). Moreover, in sandy slash pine to longleaf pine forest. Restora­ Vaccinium arboreum Marsh., Ilex opaca unconsolidated substrates of the humid tion efforts have included prescribed fire, Ait., Ostrya virginiana (Mill.) K. Koch, subtropical Gulf Coastal Plain, increased physical removal of old plow lines and Oxydendron arboreum (L.) DC., Illicium erosion, downslope sediment transport, and successional hardwoods, and groundcover floridanum Ellis., and Ilex coriacea (Pursh) vegetation change are well-documented restoration. Chapm. (Kwit et al. 1998). Slope position responses to agricultural alteration of up­ is an important control on the distribution land ground cover (Magilligan and Stamp Logging of longleaf was persistent in the on these woody species (Kwit et al. 1998). 1997, Edwards, 2001). Platt and Schwartz uplands throughout the first half of this Lower slopes are typically dominated by (1990) speculate along these lines that century. Aerial photographs from 1940 in­ more mesic species, such as Ilex coriacea logging-enhanced erosion initiated in the dicated that the uplands surrounding ABRP and Illicium floridanum. The more xeric­ Apalachicola uplands is one explanation were open and sandy with well-spaced adapted species Quercus hemisphaerica for the difference between present-day longleaf pine. However, by 1957, most and V. arboreum increase in abundance ravine vegetation and descriptions recorded of the longleaf pine had been timbered. at upslope positions. Midslopes may be characterized by a higher cover of Quer­ cus alba, F. grandifolia, and Osmanthus americanus. Aspect has an overall weaker effect on species distributions (Kwit et al. 1998).

Public lands survey descriptions of ravine slope forests from the early 1800's note a paucity of oaks and hickories when compared to the present-day vegetation. Magnolia grandiflora and F. grandifolia were more important (Delcourt and Del­ court 1977). Timbering within the ravines is one of several hypotheses to explain this change. Even though the present-day ravine forests may have been selectively cut in the 1930's, it is generally acknowledged that their steep slopes have limited the extent of intensive timbering (Schwartz 1990). A complementary hypothesis is that upland land use changes may have cascaded downslope -into the ravines, -a scenario strengthened by the recognition that sedi­ ments, plant propagules, and disturbance can propagate across some landscapes, often asynchronously to the initiating event (Swanson et al. 1988, Parker and Bendix 1996). Figure 1. Our study area along the Apalachicola River. Windrowed uplands correspond to ABRP land management units 25, 26, and 27. Native uplands correspond toABRP units 5 and 21. Rectangles in land management units indicate areas in which transects were randomly located. Lowermost ravine system is Following the widespread decline of Kelly Branch. Little Sweetwater Creek is above it. The northernmost ravine is Beaverdam Creek.

Volume 24 (3), 2004 Natural Areas Journal 243 In 1958, the uplands were converted to between 0° ± 20° and south was similarly flected in p values is robust or an artifact of slash pine monoculture. Aerial photographs restricted to 180° ± 20°. Slope angles were within-group variability no different from from 1959 show several different slash initially confined to those between 20-30°. random expectation. This statistic typically pine silvicultural practices in the uplands, In the areas within land management units ranges from 0-1. Values converging upon including the emplacement of the elevated delimited by these topographic parameters, zero indicate that the heterogeneity within plow-line berms that are still visible in three transects were randomly established. groups equals expectation by chance. the present-day landscape. These plow Each transect was aligned perpendicular to Values approaching one indicate that all lines, or windrows, were bulldozed piles the true (steepest) dip of the ravine slope. items within groups are identical and the of woody debris and groundcover placed A total of 24 transects were sampled, 12 hetero geneity within groups is greater than in rows during the preparation process for each in native and windrowed ravines random expectation. Statistical significance planting pine. Windrows have been found (Figure 1). The transect starting point was in conjunction with high A values is in­ to increase overland erosion, especially positioned to correspond to the topographic dicative of stronger group differences. In where upslope contributing areas are large break at the bottom of each ravine slope. In community ecology, an A value of 0.3 is (Prosser and Abernethy 1999). In the some cases the exact upper slope terminus fairly high. All MRPP significance tests majority of the ABRP, woody debris was point was difficult to perceive because the were conducted at the 0.05 significance either pushed into the ravines or bulldozed boundary between the ravines and the up­ level using PC-Ord Version 4.04 (McCune into windrows just upslope from the ravine lands was indistinct. End points for these and Mefford 1999). Euclidean distances edge (Schwartz 1990). transects were arbitrarily located. Transects were employed as the distance metric ranged from 30-40 m in length. (Zimmerman et al. 1985). A few areas in the Apalachicola Bluffs and Ravines Preserve were not windrowed or Three 100 m2 circular plots, each with Principle coordinates analysis (PCO) was converted to slash pine silviculture. These a radius of 5.64 m, were systematically used to visualize compositional trends tracts oflongleaf pine, although selectively distributed along the centerline of each tested with MRPP (MVSP software, Ko­ cut in the past and left to regenerate, are transect. These quadrats corresponded vatch Computing Service 1999). PCO is the only remaining areas of native uplands to lower, middle, and upper ravine slope similar to principal components analysis in in the preserve. They are characterized by positions. Within each quadrat, diameter at that it maximizes the variance in the data an overstory of longleaf pine, midstory breast height (dbh) was recorded for every set along the first axis. However, PCO is hardwoods such as turkey oak (Quercus stem 2.0 cm dbh and greater. If stems less a distance-based ordination method and is laevis Walt.) , and a groundcover dominated than 2.0 cm dbh were present, they were better suited to process the large number by wiregrass (Aristida stricta Michx.). counted and recorded as recruits, with an of zero values that typify ecological data We defined two ravine types based upon estimated 1.0 cm dbh. All woody seedlings sets (Legendre and Legendre 1998). Dele­ their differences in past upland land use. and saplings were tallied irrespective of tion of infrequently encountered species is 'Windrowed' ravines were those ravine height. Raw dbh data for each species was generally not necessary. Bray-Curtis was slope forests where upland windrows and converted to basal area (cm2/m2), density employed as the distance metric. slash pine plantings extended to the edge (stems/m2) and importance. Importance of the ravines, utilizing the most land was defined as the average of relative RESULTS possible for silviculture. 'Native' ravines percent dominance (basal area in cm2/m2) were those ravine slope forests where non­ and relative percent density. All sampling A total of 49 woody tree and shrub species mechanized selective cutting of longleaf occurred along Kelley Branch and Little were identified from a total of 4131 stems. pine, rather than windrowed slash pine, Sweetwater creeks, just north of Bristol, Species richness was slightly greater in na­ characterized historic upland land use. Florida, in May through August 2002. tive ravines (44 vs. 38 spp.). Total woody Field reconnaissance, aerial photos, and species basal area was greatest on native consultation withABRP staff facilitated the slopes (40.8 vs. 28.9 cm2/m2). Total stem Data analysis selection of the specific land management density was slightly higher in windrowed units to sample. To test for compositional differences ravines (0.6 vs. 0.5 stems/m2). Native between ravine slopes based upon their slopes (30.7 ± 7'so) had a higher mean slope Variability in slope angles and aspects upland land use type, we employed multi­ angle than windrowed slopes (24.2 ± 3.7°). were expected to confound comparisons response permutation procedures (MRPP). did not occur along any of species composition among these two MRPP non-parametrically measures the of our transects. Only three individuals of ravine types. To address this limitation, a extent to which two or more multivariate Taxus fioridana were encountered, all on U.S. Geological Survey (1:24,000) digital data sets differ based on comparisons of the the ravine slopes of native uplands. elevation model with 30-m resolution was observed and permutated average within­ used to select a subset of ravine slopes group distance among samples (Biondini Quercus hemisphaerica was the most having a uniformity in these descriptors, et a1. 1991, McCune and Grace 2002). important species in both upland land use which were then subsequently field veri­ MRPP employs an effect size statistic (A) types. The largest importance value for fied. North was constrained to those slopes to gauge whether statistical significance re- this species (36.3%) was observed in the

244 Natural Areas Journal Volume 24 (3), 2004 ravines downslope of windrowed uplands native sites was highest for Pinus glabra its relative contribution to stem density on (Figure 2). Windrowed ravines had a (22.1 %), followed by Quercus hemi­ windrowed sites. Lower relative densities greater importance value for Ilex coriacea sphaerica, Quercus alba, R grandifolia, were observed for Ilex coriacea, Illicium (9.9%) and an overall greater occurrence of andM. graridiflora. Onwiildr6Wed sites, floridanum,· V. arboreum; andIlex glabra xeric sandhills species among those with Quercus hemisphaerica had the largest (L.) Gray. importance values> 1.0 [Pinus palustris, percentage relative dominance, followed CornusfloridaL., V. arboreum and Quercus by Pinus palustris, M. grbndifolia, Carya Compositional differences were also ob­ laevis]. Ravines adjoining native uplands tomentosa (Poir. in Lam.) Nutt., and Pinus served among slope positions (Figure 3). had a higher importance of Pinus glabra, glabra. The relative percent density of the For upper and middle slopes, importance Quercus alba, R grandiflora and the woody shrubs S. tinctoria, Illicium floridanum, values for Quercus hemisphaerica were shrubs Symplocos tinctoria (L.) L'Her. and K. latifolia, and Hamamelis virginiana much higher on windrowed slopes, ranging K. latifolia. L. exceeded Quercus hemisphaerica on as high as 50-57%. Windrowed transects native sites. By contrast, Quercus hemi­ also had greater positional importance val­ Species relative percent dominance on sphaerica surpassed all other species in ues for Ilex coriacea (lower slopes), Pinus

Native uplands· All slope positions

45 40 • Importance 35 D Relative density 30 +'c CI) 25· II Relative dominance u ...CI) 20 Q. 15 -- 10 5 0

Windrowed uplands· All slope positions

45 40 35 C 30 CI) 25

~CI) 20 -- Q. 15 -- - 10 -- 5 -- IIIDII o l=i ,~}-{t .JIII JI-1I .• Jmmll ~ -=-II • III J 1

Figure 2. Importance values for tree and woody shrub species by upland land-use category. Only species with importance values> 1.0 are shown. Taxonomic nomenclature follows Clewell (1985).

Volume 24 (3), 2004 Natural Areas Journal 245 palustris (upper slopes), and Pinus elliottii the individual land management units for positional differences than aspect. Upper (upper slopes). On native ravine slopes, each upland land use type were non-sig­ and middle slope species dominance and high importance values were observed nificant [native units (A = 0.04, p = 0.22) densities were significantly different with for Pinus glabra (all slope positions), S. windrowed units (A = 0.04, p = 0.28)]. relatively high A values. tinctoria (upper slopes), and Quercus alba Comparisons of land use types by slope (middle slopes). Overall, importance values position indicated that middle slopes (A PCO results corroborated the composi­ for the species accompanying Quercus = 0.28, P < 0.01) had significantly differ­ tional differences observed with MRPP. hemisphaerica on windrowed transects ent species importances. Upper and lower Ordination of species importance data for were skewed toward relatively lower slopes were significantly different, but all 24-ravine slope transects indicated a values (Figure 3). By contrast, on middle lower A values suggested that these dif­ clear separation of upland land use types and upper slopes of native sites where ferences were not as robust. Differences along the first axis (Figure 4). The spe­ Quercus hemisphaerica was abundant, its in species importance between north and cies importance values having significant importance did not greatly exceed other south aspects and for aspects by slope correlations (Spearman's non-parametric species. position were similarly weak. Thus, the rank order coefficient, rs) with the first strongest differences in species importance, axis include Ilex coriacea (rs = -0.75, > MRPP tests indicated significant differ­ as indicated by statistical significance and 0.01), Pinus glabra (rs = 0.75, P < 0.01), ences in species importance between land high A values, can be attributed to land Quercus alba (rs = 0.60, p < 0.01), and use categories (A = 0.22, P < 0.01; Table use, particularly on middle slopes. MRPP Quercus hemisphaerica (rs = -0.64, p < 1). This result was not a consequence of comparisons of relative percent dominance 0.01). Axis 2 had an eigenvalue < 1 and within land use group variability, as the and relative percent density corroborated is not reported. differences in species importance among that land use accounted for greater COffi-

Quercus hemisphaerica Pinus glabra Fagus grandfflolia Magnolia grandfflora 60 ------~- 20 ------20 ------20 50 16 16 16 40 12 12 12 30 8 8 8 20 10 4 4 4 O~--~~--~=-~ o -I--'---'--~-L-L-._"__--'__I O~=-~-L-L--~~~ o ~::l--.,.-.L...J.--~~~ U M L U M L U M L U M L

Hamamelis virginiana Symplocos tinctoria Illicium f10ridanum lIex coriacea 20 20 20 20 16 16 16 16 12 12 12 12 8 8 8 8 4 4 4 4 [ 0 0 0 0 --=-- U M L U M L U M L U M

Carya tomentosa Quercus alba Osmanthus americana Oxydendron arboreum

20 ~-~-~-- ~~l 20 20 20 16 16 16 16 12 12 12 12 8 8 8 8 4 4 4 4 0 0 0 0 U M L U M L U M L U M L

Pinus elliottii Pinus palustris Vaccinium arboreum Quercus laevis 20 ---_.. _--_..• _---- 20 20 20 ------"1 16 16 16 16 12 12 12 12 8 8 8 8 4 4 4 4 0 0 0 0 U M U M L U M L U M L

'igure 3. Tree and woody shrub species by upland land use and slope position. Unshaded bars represent ravine slopes adjacent to selectively-cut longleaf pine plands. Shaded bares represent ravine slopes adjacent to mechanically windrowed slash pine uplands.

.46 Natural Areas Journal Volume 24 (3),2004 ------_._._------.------

DISCUSSION and Pinus glabra). They sampled the same influenced ravine tree distribution, a result two ravines as our study, but included we additionally confirmed. The results of earlier studies conducted in an adjacent ravine system to the north these ravines confirmed that our sampling (Beaverdam Creek) and a wider range of Strong differences in ravine slope forest adequately characterized ravine slope aspects and slope angles. Schwartz (1990) composition and structure were observed vegetation. Kwit et al.'s (1998) list of the sampled this same northern ravine along among our two upland land use history six most important ravine slope tree spe­ with one of the two examined in our study categories. Quercus hemisphaerica was cies in the ABRP were replicated in our (Little Sweetwater Creek). His values for the most abundant species in both land use investigation in the very same rank order species relative density and relative domi­ types, but its importance was much higher (Quercus hemisphaerica, Carya spp., F. nance were also similar to ours. Kwit et in the ravines of windrowed uplands. Total grandi/alia, M. grandiflara, Quercus alba, al. (1998) noted that aspect only weakly stem density for Quercus hemisphaerica

Table 1. MRPP comparisons of species composition by land use category (native vs windrow), aspect (north vs. south), and slope position (lower, middle, upper). MRPP tests for land use and aspect compared two groups of twelve transects. MRPP tests for slope position by land use and aspect compared two groups of twelve quadrats.

ImEortance Relative Dominance Relative Density Groups A p A p A p Land use (native vs. windrow) 0.22 < 0.01 0.09 0.01 0.18 < 0.01 Upper slopes 0.11 < 0.01 0.03 0.13 0.24 < 0.01 Mid slopes 0.28 < 0.01 0.22 < 0.01 0.19 < 0.01 Lower slopes 0.07 0.01 0.03 0.13 0.07 < 0.01 Aspect (north vs. south) 0.06 0.04 0.02 0.20 0.04 0.05 Upper slopes 0.03 0.13 0.04 0.09 0.01 0.28 Mid slopes 0.03 0.14 -0.01 0.76 0.06 0.03 Lower slopes 0.05 0.02 0.04 0.09 0.04 0.04

o 00 o o

Ravine slope forest type AXIS 1 !J:, Native uplands and northern aspects D. Native uplands and southern aspects ® Windrowed uplands and northern aspects o Windrowed uplands and southern aspects

Figure 4. Principal coordinates scatterplot of transect-level data (n = 24). Axis 1 extracted 33% of the underlying variance in the data set.

Volume 24 (3), 2004 Natural Areas Journal 247 was four times greater in ravines adjacent of canopy disturbance is low (Batista et commonly occurs in sandy xeric uplands to windrowed uplands (0.17 stems/m2 al. 1998). as an early successional species follow­ vs. 0.043 stems/m2). These ravine slopes ing fire, or may re-invade a site after fire also had a greater frequency of sandhills The high stem density and overall greater exclusion. It is a generalist species that species, particularly on upper and middle importance of Quercus hemisphaerica in responds to increased inputs of moisture, slopes. By contrast, Pinus glabra, a windrowed ravines is likely due to land often becoming abundant in mesic habitats more typical of rich and use history. Quercus hemisphaerica is fire (Veno 1976, Vaitkus and McLeod 1995). hammocks, was more important in native intolerant and frequently weedy in north These conditions were likely made avail­ ravines. Accompanying Pinus glabra were Florida. In addition to open woods, it occurs able by the suppression of fire in the higher basal areas of Quercus alba and F in disturbed areas, cut-over forests and old managed uplands and by the proximity of grandifolia, particularly at mid and lower fields (Hall 1999). One possible scenario is mesic ravine edge habitats. slope positions. The relative abundance that the silvicultural modification of these of F grandiflora has been proposed to uplands facilitated the downslope expan­ CONCLUSION decrease during periods of frequent canopy sion of Quercus hemisphaerica into the disturbance and to increase when the rate adjacent ravines. Quercus hemisphaerica Although historic land use has often been

Native uplands - Middle slope position

45 40 • Importance - 35 r- Relative density 30 o - JIIII Relative dominance ~ 25 r- CI) 20 Q. 15 r 10 • 5 l r- • • I--J Ll-il-lrI.L.. • .'l. o • t-t .- .- •-

Windrowed uplands - Middle slope positions

70 60 50 I- c: 40

~CI) 30 Q. 20 10 .1. 111m ...... n ...... ______....., .....--J o ---

Figure 5. Ravine mid-slope woody species importance values by upland land use type. Only species with importance values> 2.0 are shown.

248 Natural Areas Journal Volume 24 (3), 2004 suggested as an important influence on Lower slope angles were observed in of the information needed to address this Apalachicola ravine forests, our paper ravines adjacent to windrowed uplands, management issue. documents this relationship. We observed suggesting that slumping processes may be strong differences in ravine vegetation more active at these locations. Moreover, ACKNOWLEDGMENTS among contrasting upland lands use increased retreat of ravine slopes into types, particularly at mid-slope positions. upland habitats and the gravity-induced We thank Darroc Goolsby for assistance We viewed the composition of mid-slope transport of soil and seed beds downslope with field data collection, as well as Greg ravine forests in windrowed uplands as may in part explain the increased impor­ Seamon, Leigh Brooks, and the rest of the representative of the broad vegetation tance of sandhills species in windrowed staff at the Nature Conservancy's Apala­ changes initiated by slash pine silviculture ravines. This scenario would have also chicola Bluffs and Ravines Preserve for (Figure 5). Overstory tree composition increased gap establishment opportunities generous logistical support and advice. shifted from three tree species (Quercus in the ravines for the generalist Quercus Local botanists Angus Gholson, Wilson hemisphaerica, Pinus glabra and Quercus hemisphaerica. Alternatively, selective log­ Baker, Gil Nelson, and Loran Anderson alba) to one (Quercus hemisphaerica). ging on the upper slopes of the windrowed (FSU Department of Biology) provided Fagus grandifolia, a dominant species ravines could have also led to more xeric taxonomic assistance. Mark Schwartz recorded in the Public Land Survey of the conditions and the increased abundance and our anonymous reviewers provided presettlement ravines in the 1820's (Del­ of Quercus hemisphaerica. Quercus hemi­ insightful comments on an earlier version court and Delcourt 1977), was abundant sphaerica, as a natural component of the of the manuscript. where the uplands had not been windrowed. transition zone between the ravines and This lends more support to the idea that the uplands (Schwartz 1990), may have upland silvicultural practices have allowed expanded its range downslope with the 1. Anthony Stallins is an Assistant Professor oaks, particularly Quercus hemisphaerica, removal of ravine slope hardwoods. Tree in the Department of Geography, Florida to dominate in the ravines. ring chronologies, soil profile descriptions, State University, where he studies vegeta­ and quantification of geomorphic processes tion dynamics and biogeomorphology. Fire suppression accompanying slash on ravine slopes are needed to examine pine cultivation appears to have initiated these mutually compatible hypotheses in Jennifer Griggs is presently a doctoral this shift by creating more opportunities more detail. candidate in the School ofForest Resources for the encroachment of the fire-intoler­ and Environmental Science at Michigan ant, generalist species such as Quercus A larger geographical study is needed to Technological University. hemisphaerica (Quarterman and Keever ascertain the generality of our findings 1962). However, it is likely that fires were to other ravine systems in the Florida LITERATURE CITED suppressed not only in the windrowed panhandle (see Phillips 2001, Swetnam uplands, but also in the uplands with et al. 1999). For example, the extensive Abrahamson, w.G., and D.C. Hartnett. 1990. longleaf pine (see Outcalt 2000), as these ravine system in Torreya State Park, just Pine flatwoods and dry prairies. Pp. 103- land management units were embedded north of our study area, differs in both 149 in R.L. Myers and J.J. Ewel, eds., Ecosystems of Florida. University of Central within the fire-suppressed silvicultured physiography and upland land use history Florida Press, Orlando. uplands. In this case, a greater abundance (Platt and Schwartz 1990). However, our Batista, W.B., W.J. Platt, and R.E. Macchiavelli. of Quercus hemisphaerica should have present study underscores how the lega­ 1998. Demography of a shade-tolerant tree been observed in native ravines. In other cies of land use activities may continue to (Fagus grandifolia) in a hurricane-disturbed words, if fire suppression was the only influence species abundances and manage­ forest. Ecology 79:38-53. factor initiating vegetation change, there ment strategies long after their occurrence Biondini, M.E., P.w. Mielke, and E.F. Redente. should have been fewer compositional (Foster et al. 2003, Suding et al. 2003). 1991. Permutation techniques based on Eu­ differences between ravine types. Two We are uncertain how upland restoration clidean analysis spaces: a new and powerful hypotheses may explain why ravine slopes of native longleaf pine groundcover in the statistical method for ecological research. were not more similar. ABRP may influence ravine slope forest Pp. 221-240 in E. Feoli, L. Orl6ci, eds., composition and structure. Reinstatement Computer Assisted Vegetation Analysis. Kluwer Academic Publishers, Dordrecht. Geomorphic disturbance (Swanson et of fire frequencies that existed before al. 1988, Parker and Bendix 1996) may intensive-silviculture may not necessarily BlackA.E., E. Strand, R.G. Wright, J.M. Scott, P. Morgan, and C.Watson. 1998. Land use have been an additional component in the revert slope ravines back to their previous history at multiple scales: implications for sequence of events leading to vegetation state (Vale 1982, Wallin et al. 1994). Other conservation planning. Landscape and Ur­ change in windrowed ravines. Platt and factors, including climate variability and ban Planning 43:49-63. Schwartz (1990) suggested that removal of forest pathogens (see Schwartz et al. 1995), Chapman, A.w. 1885. Torreya taxifolia Ar­ native vegetation in the uplands may have may have also influenced the changes in nott. A reminiscence. Botanical Gazette accelerated rates of groundwater seepage ravine vegetation initiated by fire suppres­ 10:251-254. into the steepheads, thereby increasing the sion, geomorphic disturbance, and selec­ Clewell, A.F. 1985. Guide to the Vascular frequency of slumping and disturbance. tive logging. Our study is one component of Florida. University of Florida Press,

Volume 24 (3), 2004 Natural Areas Journal 249 Tallahassee. Means, D.B. 1996. Longleaf pine forest, going, Schwartz, M.W. 1990. Conserving forest di­ Delcourt H.R, and P.A. Delcourt. 1977. Pre­ going, ... Pp. 210-219 in M.B. Davis, ed., versity in northern Florida: from landscapes settlement magnolia-beech climax of the Eastern Old-Growth Forests: Prospects for to popUlations. Ph.D. diss., Florida State Gulf coastal plain: quantitative evidence Rediscovery and Recovery. Island Press, University, Tallahassee. from the Apalachicola River bluffs, north­ Washington D.C. Schwartz, M.W, S.M. Hennann, and C.S. Vogel. central Florida. Ecology 58:1085-1093. Magilligan, EJ., and M.L. Stamp. 1997. His­ 1995. The catastrophic loss of Torreya taxi­ Edwards, L.A. 2001. Vegetation dynamics torical land-cover changes and hydrogeo­ folia: assessing the environmental induction and biogeomorphic influences in a highly morphic adjustment in a small Georgia of disease hypotheses. Ecological Applica­ eroded landscape. Ph.D. diss., University watershed. Annals of the Association of tions 5:501-516. of Georgia, Athens. American Geographers 87:614-635. Smith, G.C., M.W Patterson, and H.R. Tren­ Florida Natural Areas Inventory 1990. Guide Nakamura, E, EJ. Swanson, and S.M. Wond­ dell. 2000. The demise of the longleaf­ to the natural communities of Florida. zell. 2000. Disturbance regimes of stream pine ecosystem. Southeastern Geographer Florida Department of Natural Resources, and riparian systems: a disturbance-cas­ 40:75-92. Tallahassee. cade perspective. Hydrological Processes Suding, KN., KL. Gross, and G.R House­ 14:2849-2860. Foster D., E Swanson, J. Aber, J. Burke, N. man 2003. Alternative states and positive Brokaw, D. Tilman, A. Knapp. 2003. The Outcalt, KW. 2000. Occurrence of fire in feedbacks in restoration ecology. Trends in importance of land-use legacies to ecology longleaf pine stands in the southeast United Ecology and Evolution 19(1):46-53. and conservation. Bioscience 53:77-88. States. Proceedings of the Tall Timbers Fire Swanson, EJ., T.K. Kratz, N. Caine, and R.G. Ecology Conference 21:178-181. Hall, D. 1999. Laurel oak or swamp laurel oak? Woodmansee. 1988. Landform effects on Circular ENH-134. Department of Environ­ Parker, KC., and J. Bendix. 1996. Landscape­ ecosystem patterns and processes. Biosci­ mental Horticulture, Florida Cooperative scale geomorphic influences on vegetation ence 38:92-98. Extension Service, Institute of Food and patterns in four environments. Physical Swetnam, T.W, C.D.Allen, andJ.L. Betancourt. Agricultural Sciences, University of Florida, Geography 17:113-141. 1999. Applied historical ecology: Using the Gainesville. Phillips, J.D. 2001. Human impacts on the en­ past to manage for the future. Ecological Harper, RM. 1914. Geography and vegetation vironment: Unpredictability and the primacy Applications 9: 1189-1206. of northern Florida, 6th Annual Report. of place. Physical Geography 22:321-332. Vaitkus, M.R, and KW McLeod. 1995. Pho­ Florida Geological Survey, Tallahassee. Platt, WJ. and M.W Schwartz. 1990. Temper­ tosynthesis and water-use efficiency of 2 James, C.W 1961. Endemism in Florida. Brit­ ate hardwoods forests. Pp. 194-229 in RL. sandhill oaks following additions of water tonia 13:225-244. Myers and J.J. Ewels, eds., Ecosystems and nutrients. Bulletin of the Torrey Botani­ of Florida. University of Central Florida cal Club 122:30-39. Kettle, WD., P.M. Rich, K Kindscher, G.L. Press, Orlando. Pittman, P. Fu. 2000. Land-use history in Vale, T.R. 1982. Plants and People: Vegetation ecosystem restoration: a 40-year study in the Prosser, J.P., and B. Abernethy. 1999. Increased Change in North America. Association of prairie-forest ecotone. Restoration Ecology erosion hazard resulting from log-row con­ American Geographers, Washington, D.C. 8:307-317. struction during conversion to plantation Veno, P.A. 1976. Successional relationships forest. Forest Ecology and Management Kovatch Computing Services. 1999. MVSP: of 5 Florida plant communities. Ecology 123:145-155. Multivariate Statistical Package, Version 57:498-508. 3.1, Wales. Reiners, WA., and KL. Driese. 2001. The Wallin, D.O., E J. Swanson, and B. Marks. propagation of ecological influences through Kurz, H. 1938. A physiographic study of the 1994. Landscape pattern response to changes heterogeneous environmental space. Biosci­ tree associations of the Apalachicola River. in pattern generation rules: Land-use ence 51:939-950. Proceedings of the Florida Academy of Sci­ legacies in forestry. EcologicalApplications ences 3:78-90. Rupert, ER, 1991. The geomorphology and 4:569-580. geology of Liberty County, Florida. Open Kwit, C., M.W Schwartz, W.J. Platt, and J.P. Zimmennan, G.M., H. Goetz, and P.W Mielke File Report no. 23. Florida Geological 1985. Use of an improved statistical method Geaghan. 1998. The distribution of tree Survey, Tallahassee. species in steepheads of the Apalachicola for group comparisons to study effects of River Bluffs, Florida. Journal of the Torrey Quartennan, E., and C. Keever. 1962. South­ prairie fire. Ecology 66:606-611. Botanical Society. 125:309-318. ern mixed hardwood forest: climax in the southeastern Coastal Plain, USA. Ecological Legendre, P., and L. Legendre. 1998. Numerical Monographs 32:167-185. Ecology. Elsevier Science, Amsterdam. McCune, B., and J.B. Grace. 2002. Analysis of Ecological Communities. MjM Software Design, Gleneden Beach, Ore. McCune, B., and M.J. Mefford. 1999. PC-ORD. Multivariate Analysis of Ecological Data, Version 4. MjM Software Design, Gleneden Beach, Ore.

250 Natural Areas Journal Volume 24 (3), 2004