Jnurnul of the Torrey Botcmicd Socirty 129(4), 2002, pp. 261-288

Reference conditions for old-growth pine forests in theUpper West Gulf Coastal Plain

Don C. Bragg’U.S. Department of Agriculture, Forest Service, Southern Research Station.

P.0. Box 3516 UAM, Monticello, AR 71656

DON C. BKAGG (U.S. Department of Agriculture, Forest Service, Southern Research Station). Reference con-ditions for old-growth pine forests in the Upper West Gulf Coastal Plain. J. Torrcy. Bot. Sot. 129: 261-288.2002.-Ecosystem restoration has become an important component of forest management, especially on publiclands. Howeber, determination of manageable reference conditions has lagged behind the interest. This paperoresents a case studv from Dine-dominated forests in the Upper West Gulf Coastal Plain NJWGCP), with special. .emphasis on southern Arkansas. Decades of forest management, fire exclusion, exotic species invasion, and otherecological changes have converted the small remnants of mature shortleaf (Pirius echirm/cl Mill.) and loblollypine (Pinus trteda L.) stands into ineffectual models for restoring presettlement-like conditions. However, suf-ficient information can be gathered from available references to more reliably describe the boundaries of thedesired reference environment. Early explorer accounts, maps, survey records, historical trade and technicalpublications, and modern scientific journals were consulted to reconstruct presettlement (pre- 1900) forest con-ditions for pine-dominated landscapes of the UWGCP On average, virgin UWGCP pine forests had considerablymore shortleaf pine (especially in the uplands) than contemporary natural stands, with relatively low basal areaand standing volume concentrated in large trees. Presettlement pine timber also had less uniform structural andspatial patterns than modern examples of mature pine. Assuming most of the critical processes ilre still present,it appears possible to recreate the compositional and structural attributes of virgin pine forests.

Key words: Pinus, loblolly pine, shortleaf pine, presettlement vegetation, virgin forest.

Interest in old-growth forests has increased inrecent decades as issues of endangered species,wilderness, and biological and social legacieshave been raised on public lands. Although mil-lions of hectares of old-growth forests remain inthe western United States, the status of easternold-growth is more precarious. Of the nearly 154million hectares of forestland in the eastern Unit-ed States, only 798,000 hectares (approximatelyone-half of one percent) are primary forest (Davis1996), with most of this concentrated in a fewlarge tracts on public lands. Restoration of old-growth has been advocated as a means to supple-ment dwindling mature forests, even if the endproduct is not exactly equivalent to virgin timber.

Reconstructing an approximation of old-growth is not easy, however, in the highly al-tered ecosystems of modern North America. Inaddition to the lack of representative old-growthexamples, new land use patterns, modified nat-ural disturbance regimes, climate change, pol-lution, exotic species, extinction or extirpationof native species (or overabundance of others),and landscape fragmentation have affected theinnate capacity of the environment to return to

’ The comments and insights of O.H. (“Doogie”)Darling, John Gray, Tom Foti, Mike Shelton, BeverlyCollins, and several anonymous reviewers consider-nbly improved this manuscript.

Received for publication January I 1, 2002. and inrevised form May IO, 2002.

conditions similar to those prior to Euroameri-can settlement. Furthermore, some sensitive old-growth-dependent species (e.g., red-cockadedwoodpecker (Picodes boredis Vieillot)) maynot survive under current forest conditions longenough to benefit from natural rates of systemrenewal (Bukenhofer et al. 1994). Efforts are Lm-derway to restore presettlement ecological com-munities using silviculturdl treatments to accel-erate the development of desirable stand features(e.g., Bukenhofer et al. 1994; Gaines et al. 1997;Huffman and Werner 2000). Even though theseefforts cannot replace current unmanaged old-growth stands (Tyrrell 1996), managing for old-growth characteristics may permit a balance be-tween ecologically and socially desirable con-ditions and some commodity production (Len-nartz 198X; Guldin 1991).

Specific targets for prcsettlement conditionsshould be developed before attempting to usesilvicultural manipulation to achieve old-growth-like characteristics (Trombulak 1996;Clewell and Rieger 1997; Clewell et al. 2000).Limited descriptions of old-growth forests ineastern North America have been provided fromexisting examples (e.g., Walker 1963; Jones etal. 198 1; Cain and Shelton 1994; Harms 1996;Greenberg et al. 1997; Murphy and Nowacki1997; Tyrrell et al. 1998; Landers and Boyer1999). Some contemporary old-growth com-munities differ little from presetllement times.

261

262 JOURNAL OF THE TORREY BOTANICAL SOCIETY [VOL. 129

The remaining old-growth northern hardwoodstands in north-central North America, for in-stance, are comparable to similar presettlementforests because of their remoteness and intactnatural disturbance regime (wind-dominated,rather than fire). Unfortunately, most old-growthdescriptions for the southeastern United Statesprovide only limited information based on high-ly altered contemporary examples (White andLloyd 1995). Thus, those engaging in ecosystemrestoration have to consider other options whendefining their reference conditions.

Researchers have used early land surveys toprovide at least a qualitative description of pre-settlement vegetation (e.g., Stearns 1949; Bour-do 1956; Delcourt 1976; Schafale and Harcombe1983; Foti and Glenn 1991; White and Mlad-enoff 1994; Black and Abrams 200 1). Inferencesalso can be made by examining period photo-graphs, paintings, sketches, or written accountsof early travelers (e.g., Nelson 1957; I-lough1965; Delcourt 1976; White 1984; Foti andGlenn 199 1; Hammett 1992; Strausberg andHough 1997). Other information sources includeearly technical publications, stand inventories,and current research papers (e.g., Olmsted 1902;Chapman 1912: Dickson 1991). Even old tradejournals (e.g., American Lumberman) or pro-motional publications produced by railroads,timber companies, land speculators, or localgovernments can contribute to restoration ef-forts. For example, many large lumber compa-nies in the southern United States were featuredin trade magazines that, while emphasizing themilling, financing, and staffing of the operation,often provided photographs of virgin timber-lands or individual big trees (e.g., Anonymous1904a,b; 1905; 1906; 1909).

The U.S. Department of Agriculture, ForestService is in the process of establishing a re-search project on the treatment of upland pine(i%1Ll.T spp.) forests for old-growth characteristicson the Crossett Experimental Forest in AshleyCounty, Arkansas. Most of the natural divisionsof Arkansas do not have representative examplesof contemporary old-growth to emulate (Pell1981), making it necessary to find other meansto identify and describe the desired ecological at-tributes. This work details the acquisition of ref-erence conditions for virgin pine forests of UpperWest Gulf Coastal Plain (UWGCP) using histor-ical literature, photographs, and other relevant ac-counts (with special emphasis placed on southernArkansas) to restore mature pint forests consis-tent with presettlement patterns.

Materials and Methods. STUDY AREA

DESCRIPTION. The Gulf Coastal Plain is subdi-vided by the Mississippi River into East andWest provinces composed of similar parent ma-terials and geological development. The WestGulf Coastal Plain can be split further into “Up-per” and “Lower” subregions based on subtledifferences in elevation, parent materials, andkey overstory species. Schultz ( 1997) distin-guished the Lower West Gulf Coastal Plain(LWGCP) as level to gently rolling, fairly sandyplains below 30 m in elevation; the UWGCPincluded hills and plains above this level. Pre-settlement forests of the LWGCP were predom-inantly longleaf pine (Pinus pulustris Mill.); theUWGCP was primarily shortleaf pine (Pinusechinutu Mill.); and loblolly (PinLL.s tczedcr L.)was common to both subregions. The UWGCPextends west of the Mississippi River Delta fromnorth-central Louisiana and central Arkansas(south of the Ouachita Mountains) to southeast-ern Oklahoma and northeastern Texas.

Although minor Holocene-era alluvial bottom-lands are widespread, the UWGCP is primarilycomposed of marine sediments deposited duringthe Cretaceous and early Tertiary periods, withsome areas of Pleistocene river terraces. Consid-erable variation in internal soil drainage can befound across the UWGCP, ranging from some-what excessively well drained to very poorlydrained, with an abundance of somewhat poorlydrained sites. Soils also tend to be deep and me-dium textured, with relatively low nutrients andorganic content (Pell 1983; Walker and Oswald2000). Precipitation on the UWGCP averagesfrom < 100 cm annually in Oklahoma and Texasto > 13.5 cm in southeastern Arkansas and centralLouisiana, and the frost-free growing seasonlength varies from 200 to 250 days (Skiles 1981;Walker and Oswald 2000). The nearby Gulf ofMexico provides moist, unstable air that may trig-ger extreme weather events like thunderstorms,tornadoes, hurricanes, and ice storms. Droughtsare not unusual in this region (Stahle et al. 1985),and when particularly severe, widespread firesmay occur. A long history of human occupationhas also influenced the vegetation and disturbancepatterns of the UWGCF?

HISTORICAL COVIZ TYIXS. The expansive nat-ural distributions of loblolly and shortleaf pineresult in considerable geographic overlap be-tween these species, although they are found lo-cally in distinct habitats. The virgin shortlealand loblolly pine forests that once covered mil-

2002] BRAGG: OLD-GROWTH PINE REFERENCE CONDITIONS 263

lions of hectares have been reduced greatly bytimber harvest, settlement, and altered distur-bance regimes. Tellingly, several recent publi-cations (e.g., Nowacki and Trianosky 1993;Gaines et al. 1997; Tyrrell et al. 199X) identi-fying old-growth covertypes in the eastern Unit-ed States do not even list a loblolly-shortleafpine type comparable to what once existed, pos-sibly because there are so few examples. Theseforests were rarely pure pine, even in presettle-ment times (Harvey 1883; Mattoon 19 15). Someof the most homogeneous virgin pine standswere found on fire- or overflow-prone sites (e.g.,shortleaf stands in northwestern Louisiana andsouthwestern Arkansas (Mattoon 19 IS) or lob-lolly flatwoods in Arkansas and Texas (Mohr1897; Forbes and Stuart 1930)).

It is also important to recognize the impactthat Native Americans had on presettlement veg-etation. These first inhabitants used fire, landclearing, and hunting to both directly and indi-rectly alter vegetation patterns for millennia be-fore Euroamerican exploration and settlement(Forbes and Stuart 1930; Dclcourt 1976; Ham-mett 1992; Strausberg and Hough 1997; Hameland Buckner 1998; Key 2000). Native Americanuse of these landscapes helped structure naturalcommunities, but the true extent of their influ-ence on presettlement vegetation prior to European exploration will never be adequately doc-umented. Their decimation from disease and re-lated upheavals starting in the 1500s fundamen-tally changed the dynamics of the UWGCPcenturies before any chroniclers could reporttheir impacts (Hamel and Buckner 199X; Carrollet al. 2002). The lapse of many decades betweenhistorical Native American cultures and thosetribes eventually removed in the early IgOOs,coupled with considerable cuitural changes innative populations, also affected vegetationcomposition, structure, and dynamics.

M~~~;.IIN VEGETATION PATTEIINS. Pine, hardwood, and mixed pine-hardwood forests domi-nate the current natural upland communities ofthe UWGCP with loblolly and shortleaf pine,oak (Quercus spp.), gum (N~ssu spp. and Liq-uiclun?hctr sp.), and hickory (CCIY~YI spp.) of no-

table importance (Foti et al. 1994; Rosson et al.1995). Contemporary mature pine and pine-hardwood upland forests typically have a don-inant pine overstory with various hardwoods,shrubs, vines, and forbs beneath them. Large rc-gions of the UWGCP are intensively managedloblolly pine stands of both natural and planted

origin. Competition control is frequently used toimprove pine growth, but most managed standsstill have abundant understories of oak, gum,elm (IJIIIIL~S spp.), maple (Acer spp.), greenbrier(SwiIux spp.), honeysuckle (Lonicem spp.),American beautyberry (Crrliicar~x~ urnericanuL.), and many other species. Most of the uplandforested areas that were converted to agricultureor pastureland beginning in the middle of the1800s have long since reverted back to even-aged pine-, oak-, and gum-dominated forests(Reynolds 1980). Very few terrace prairies andopen, grassy woodlands originally found in theUWGCP remain; most were converted to riceand cotton farms or commercial forestland.

Current forest stand composition, density, andstructure depend largely upon silvicultural prac-tices. Loblolly pine and certain red oak taxa arepreferred timber species, and shortleaf pine andother hardwood species are often cut to favor themore rapidly growing commodities. Stand den-sities are typically maintained at much higherlevels than historical records suggest. Few treesare allowed to grow larger than 50 cm DBH oncommercial timberlands in the UWGCP, regard-less of species.

SAMPLING. Reconstruction of historical con-ditions depends upon the discovery and inter-pretation of reliable information. Scores ofsources were examined for their appropriateness.Available references included accounts of earlytravelers and residents, original General LandOffice (GLO) survey notes, historical photo-graphs and sketches, promotional brochures,early research and technical reports, and con-temporary scientific publications. Not surpris-ingly, most information was qualitative, but anyinsights that could be used in management toachieve the desired restoration goals were notedand placed in the context of other availableknowledge. Many of the presettlcment and con-

temporary pine stands cited in this work areidentilied in Figure I.

Most definitions of “presettlement” and “old-growth” are at best imprecise, and at worst ar-bitrary assignments. Presettlement, for example,has been variottsly used to describe conditionsbefore any human settlement, or the arrival ofChristopher Columbus in 1492, or at the time ofNative American removal and Euroamericansettlement, or before widespread commercial ex-ploitation (e.g., Hamel and Buckner 1998). Sim-ilar uncertainty is found in old-growth delinea-tion (Hunter and White 1997; Helms 1998).

264 JOURNAL OF THE TORREY BOTANICAL SOCIETY [vol.. I29

Reed (1905)AA

Alabama\

* Watson (I 957)

Significant location

A Historical old-growth

A Contemporary old-growth

,_.._.. -.”

“LO

P

__... -Warr

d,-

A-A I““0 I “Monticello

700 miles---~~~‘~~~‘~“~‘~~‘~~~~.. . . . . . . . .

I Levi Wilcoxon1 5 0 k m Demonstration Forest

(The Timber Company)Forest (USFS)

50 miles

7 5 k m

Fig. I, Map of the Midsouth. includin, c~7 sic?nificnnt locations used in this study.

Thus, it may be most beneficial to use the era found. Old-growth consists of relatively undis-with the most reliable information that still re- turbed stands for which the dominant trees cx-tains the ecological integrity of early landscapes. wed 100 years old.For this study, the period from 1850 to 1900A.D. was chosen to represent virgin forest con- Results and Discussion. IDENTII-‘YING P~xstrr-ditions heca~tse reasonably good records can be ~rt.twltm’r RI;I:EIWNCI: CO N D I T I O N S . White and

2002 ] BRAGG: OLD-GROWTH PINE REFERENCE CONDITIONS 265

h

= 202 Adapted from Reed (1905)5 16 (central Alabama)

3 12&8

Pine lands, “Block 1”

" 4

00 10 20 30 40 50 60 70 80 90 100 110

Diameter class (cm)

(b)I

2.4: = Loblolly pine

p 2.1 : 0 Shortleaf pineLongleaf pine

3i4

0.9

m 0.60 . 30 . 0

0 10 20 30 40 50 60 70 80 90 100 110

Diameter class (cm)

Fig. 2. Frequency ((I) and basal area (hi distribu-tion by pine species for a tract within the “pine lands”area of Bibb County, in central Alabama (adapted fromReed ( 1905)).

Lloyd ( 1995) cautioned that present-day exam-ples of old-growth may not reflect the dynamicnature of virgin forests, and thus serve as poormodels for restoration (see also Bourdo 1956).Since old-growth forests consist of more thanjust big or aged trees, efforts were made to yuan-tify as many attr ibutes of undisturbed old-growth from as close to the presettlement periodas feasible. These include: species composition,size and age structure, growth performance, treeform, overstory spatial pattern, understory andforest floor conditions, disturbance regimes, de-gree of heart rot in live trees, and large woodydebris.

Loblolly rarely occurred in pure stands, ex-cept in the flatwoods in Texas and southern Ar-kansas (Mohr 1897; Forbes and Stuart 1930).Pure shortleaf pine stands were encountered onfrequently burned sites in the UWGCP (Foster19 I2), although Mattoon (19 15, p. 4) stated “[illis doubtful whether shortleaf is now found inpure type on more than from 20 to 40 per centof its former range.” A mixture of loblolly andshortleaf pine was more typical for the UWGCPMattoon (19 15, p. 4-5) mentioned “especiallyheavy” stands of “complementary” shortleaf-loblolly pine in Arkansas and Louisiana, withshortleaf dominating “drier and lighter” soilsand loblolly predominant on “heavier, moistsoils.” Zon (1905) found decidedly more lob-lolly than shortleaf pine in several differentstands in eastern Texas (Table l), but Heptingand Chapman (1938) described the opposite:some small (< 5 ha) old-growth remnants inTexas were predominantly shortleaf, with lessthan 8% of their stocking in loblolly.

Species Cornpo.sifiorz. Pine composition var- Harvey (1883) noted the fraction of pine in-ied considerably in the prcsettlement forests of creased as one went south in Arkansas. Mohrthe southeast, but was usually prominent. For (1897, p. 119) stated that “. . it can be safelyinstance, some pinelands in Georgia were esti- assumed that about one-half of the lumber cutmated to have been 89 to 99% pine (Plummel and shipped as ‘Yellow Pine’ to Northern mar-1975). In a central Alabama mixed pine stand, kets from southwestern Arkansas is LoblollyReed ( 1905) reported that more than 43% of the Pine, the other half being Shortleaf.” Mohr’strees were lobiolly pine: shortleaf pine com- maps of loblolly and shortleaf pine distributionprised almost 39% of stems; and longleaf pine showed these species to have roughly the samecontributed 18%, (Fig. 2). Reynolds et al. (1984) stocking across much of southern Arkansas.

stated that the virgin upland forests of southernArkansas and northern Louisiana were about50% loblolly pine and 25% shortleaf pine, withthe rest in hardwoods.

Loblolly is currently the dominant pine spe-cies in the UWGCP (Rosson et al. 1995; Schultz1997). However, many early accounts and pho-tographs (e.g., Fig. 3) suggest that shortleaf pinewas the predominant conifer in upland virginpine forests of southern Arkansas, northern Lou-isiana, and northeastern Texas (Foster 19 12;Harper 1914). Loblolly pine was historicallyconsidered more of a bottomland or old fieldspecies, with shortleaf pine dominating drier orfire-prone upland sites (Mohr 1897; Reed 1905;Record 1907; Foster 1912; Chapman 1913; Mat-toon 191.5; Westveld 1935). Recent gains of lob-1011~ pine at the expense of shortleaf can be par-tially attributed to fire exclusion, managementdiscrimination against shortleaf, loblolly’s natu-

ral colonization of old fields and clearcuts, andthe widespread planting loblolly pine (White1984; Schultz 1997).

266 JOURNAL OF THE TORREY BOTANICAL SOCIETY (VOL. 129

F i g . 3 . The dominance of shortleaf pine in the virgin forests of UWGCP can be seen in many early pho-tograp 1s of the region. including this picture taken near Hamburg, Arkansas circa 1937. (Picture courtesy of theCrossett City Library.)

Near Pine Bluff, Arkansas, Olmsted (1902) re-ported the following abundances for shortleafand loblolly pine for “pine ridge,” “pine flat,”and “hardwood bottom,” respectively: 38% ver-sus 16%; 20% versus 34%; and 2% versus 3%.Even though loblolly pine proved more abun-dant in two out of the three forest categories, thepine ridge type covered 80% of the 769 hectaresincluded in the survey, compared to 12% for thepine llat and 8% for the hardwood bottom.Though not as quantitative as other estimates,Chapman (1913, p. 4) reported that the uplandtimber on an almost 1 1,000 hectare tract southof- Crossett, Arkansas consisted of “. shortleafand loblolly pine in almost equal mix-ture. . [and]. forin[eci] almost pure stands onall the higher lands. .”

Estimates of the stocking of non-pine speciesare far less reliable. Since most of these specieswere not considered valuable (Reynolds 19X0),they were often excluded from early inventories(e.g., Mohr 1897; Chapman 19 13). Others, whileproviding more information on non-pine taxa.

aggregated them into broad groupings like“hardwoods” or “gum” or “white oaks” (e.g.,Olmsted 1902; Reed 1905; Zen 1905; Walker1963; Reynolds et al. 1984). Non-pine speciesmay have constituted as little as < 1% of uplandforests (Chapman 1913) to IS to 40% of somemesic stands (Morbeck 1915) to almost everytree in some bottomland sites (Ohnsted 1902;Reed 1905). Delcourt (1976) estimated fromland survey records that dogwoods (Cormaspp.), red oaks (mostly Quc~rcu.s ,frrlcatu Michx.a n d Q 4 4 ~ r c ~ 4 . s pa~r~ckr Raf.), and post oak (Quer-CLIS .stdltrtrr Wang.) comprised about one-quarterof the presettlement upland pine communities innorthern Louisiana. The hardwoods on ArkansasLumber Company pine lands in Bradley Countywere estimated to be 35% oak, 30% hickory,25% gum, and 10% baldcypress (Tuxodium dis-tich~4m (L.) Rich.) (Anonymous 1906). Morbeck( 19 15) reported that white oak (Querc~~s u/ha L.)was the most common hardwood in many pine-dominated virgin stands near Fordyce, Arkansas,with only a limited stocking of blackgum (Ny.s.su

2002] BRAGG: OLD-GROWTH PINE REFERENCE CONDITIONS 267

Table I. Descriptions of loblolly and shortleaf pine-dominated stands from historical literature and somecurrent examples.

AhllIKl~lllCC Basal arez~ M&Y.Cicncral \Ullld p i n elocittim AELI TOt:ll Shortleai I.ol~lolly Total Pine DBH

SlaIltl n a m e (1x1) (mx\/lXI) (%)” (%)h (m’nla) (%>)‘I (Cnlp Notes’

HISTORICAI. REI:EIU~N~~.SGurdon, southwestern Arkansas (Mohr 1897)

Gurdon #I 0.4 54.4 100.0 1 0 . 4 100.0 >63.0 1Gurdon #2 0.4 7 4 . 1 < 100.0 16.X 100.0 122.0 I

Ashley County, Arkansas (Chapman 19 13)Ashley #I 1 6 . 2 50.0 -50.0~ -50.0’ 1 3 . 8 100.0 > 106.7 IAshley #2 1 6 . 2 64.9 -50.0~ -so.oa 1 0 . 9 100.0 >101.6 I

Pine Bluff, Arkansas (Olmsted 1902)Pine lands 706.2 203.x 35.7 1 x . 9 1 4 . 2 65.5 91.4 2

Enstern Texas (Zen 1905)Stand #I 1 .h 307.0 2.4 78.5 31.5 94.3 83.8 3Stand #2 4.9 410.5 6 . I 41.2 25.4 73.2 8X.9 3

Eastern Texas (Carver and Miller 1933, Hepting and Chapman 1938)Stand #I 2.6 108.7 -92.58 -7.se 78.7 4Stand #2 4.9 318.8 --92.Y ---7.5a 7 1 . 1 4

Bibb County, central Alabama (Reed 1905) (pine lands only)Block #I 45.3 149.6 38.6 43.4 1 0 . 4 61 .s 76.2 3

CURRENT EXAMPLES

Levi Wilcoxon Demonstration Forest, Ashley County, Arkansas (this paper)Main stand 6 . 1 387.5 5.0 1 3 . 2 31.8 57.2 92.5 5

R.R. Reynolds Research Natural Area, Ashley County, Arkansas (this paper)Units 41&42 32.4 414.0 1 . 6 1 7 . 2 34.4 54.5 91.9 5

,I Total number of trees per hectare, including all species (if reported).I’ Fraction of pine stems from total number of stems tallied in the stand.‘ Total stand basal area, including all species (if reported).(’ Fraction of pine (shortleaf + loblolly) basal area from total stand basal area.c Maximum shortleaf or loblolly pine diameter (other species not included).’ Notes on data: 1 = values are for pines larger than 30 cm DBH, 2 = values are for a11 trees larger than 5

cm DBH. 3 = values are for all trees larger than 2.5 cm DBH, 4 = values are for all pines larger than 10 cmDBH, 5 = values are for all trees larger than 9 cm DBH.

I: Composite species proportion estimates.

sylvuticu Marsl~.), swcctgum (Liquidcrmhar styr-UC$LKI L.), hickory, and ash (Frcrxirzus spp.). Inhis assessment of lands south of Crossett, Chap-man (1913, p. 5) ~OUII~ “. the only hardwoodgrowth is a few very stunted and deformedo a k s . ” w i t h “1 h]etter hardwoods, includingwhite and black oaks and some sweet gum andhickory. near streams where the soil is fairlywell drained, moist and deep.”

Contrast these historical descriptions to thoseof the few existing examples of old-growth pine-hardwood stands in southern Arkansas. On theproposed R.R. Reynolds RNA, loblolly pinecomprised 65% of the total pine stems and 77%of total pine basal area in 1993 (Cain and Shel-ton 1996). Hardwoods and pine switched abun-dances fi-om 1937 to 199.7, with pines dropping

from 80% to 20% of merchantable sterns, andconcurrent increases in hardwood frequency (es-pecially during the last decade). Interestingly,the proportion of pine basal area has changedlittle over the 60+ year observation period ofthe stand, although it is anticipated that hard-woods will increase in importance in the future(Cain and Shelton 1996). The gradual replace-ment of intolerant pines to more shade toleranthardwoods has been noted in other old-growthloblolly-shortleaf stands (e.g., Watson 1957;Lipps and de Selm 1969; Jones 1971; StalterI97 1; Jones et al. 198 1; Fail 199 I; Harringtonet al. 2000).

268 JOURNAL OF THE TORREY BOTANICAL SOCIETY IVOI.. 129

(a)-136

3 32f

2

28

24e 202s 16 12

i$8" 4

0 10 20 30 40 50 60 70 80 90 100 110

Diameter class (cm)

(W3 . 33 . 02 . 7

7 2 . 43 2.15 1 . 8I 1 . 5$I

1 . 20 . 9

d 0. 60 . 30 . 0

0 10 20 30 40 50 60 70 80 90 100 110

Diameter class (cm)

Fig. 4. Frequency (rr) and basal area (hj distribu-tions by species and diatneter class fix a > 60 yearsold loblolly pine stand in eastern Texas (adapted fromZen ( 1005)).

can also be reconstructed from historical data.Although open stands were more common, pine-dominated old-growth (Table 1) in the UWGCPcould produce relatively high stocking and standdensities (Westveld 1935; Walker 1963). Someeast Texas pine stands reported by Zen (1905)had > 300 trees/ha and stand densities > 25 m’lha, of which small hardwoods comprised muchof the stocking and basal area. The pines inZen’s inventory were smaller (on average, < 40cm DBH) and they showed a distinctly modaldiameter distribution (Fig. 4). The stands inven-toried by Zen ( 1905) and Reed ( 1905), thoughdominated by small diameter trees, still con-tained scattered large (> 70 cm DBH) individ-uals. Hepting and Chapman (1938) describedtwo Texas old-growth shortleaf pine stands thataveraged 109 trees/ha and 319 trees/ha greaterthan IO cm DBH.

Mohr (1897, p. 96 and 1 19) provided sum-maries of an “average” acre of both shortleafand loblolly pine near Gurdon, Arkansas (Table1, Fig. 5). IJsing diameter class means for thesestands, shortleaf pine stand had 54.4 trees great-cr than 30 cm DBH (10.4 m’ of basal area) per

hectare and loblolly pine stand averaged 74.1trees/ha and 16.X m*/ha. The loblolly acre con-tained more stems < 45 cm DBH than largetrees, while the shortleaf acre was dominated bylarge (> 4.5 cm DBH) pines. Chapman (1913)reported on uncut pine stands south of Crossett,Arkansas (Table 1). The tract in Figure 6a has arelatively even distribution of trees from 30 to90 cm DBH, gradually tapering off by 110 cmDBH (a pattern consistent with uneven-agedold-growth stands (Smith 1986)). Althoughmore evenly distributed than Mohr’s stands,stocking was still low, with an average of SOtrees/ha and 13.8 m*/ha in basal area (concen-trated in the 55 to 85 cm DBH class range, Fig.6a). Chapman’s (19 13) second stand consistedof young and mature pine with better stockingin the smaller size classes (Fig. 6b), althoughsome very large trees were present (1.2 trees/ha> 90 cm DBH). Pines < 60 cm DBH were moreabundant, but stocking (65 trees/ha) and basalarea (10.9 m?/ha) remained low.

In a stand near Pine Bluff, Arkansas, Olmsted( 1902) differentiated between loblolly and short-leaf pine and tallied stems down to 5 cm DBH.Shortleaf pine dominated most size classes (Fig.7), with its stocking in some diameter classestriple that of loblolly pine. The distribution oftrees > 30 cm was similar to Chapman’s inven-tory, except Olmsted (1902) reported no stems> 95 cm DBH. Diameter class basal area peakedat 55 cm and tapered off rapidly, with littlefound in trees > 85 cm DBH. Inclusion of thesmaller (< 30 cm) size classes yielded 203.8trees/ha, or about four times the stocking ofMohr’s (1897) and Chapman’s (1913) stands(Table 1). However, additional stocking in thesmallest diameter classes did not result in higherstand density as the parcel averaged only 14.2m?/ha of basal area, with approximately one-quarter of the total stand basal area in trees <30 cm DBH.

Contemporary old-growth stands almost al-ways have greater tree density than virgin for-ests. A photograph taken (circa 1948) of a signat the entrance to the Levi Wilcoxon Demon-stration Forest lists some of its attributes, in-cluding an average stocking of about 193 trees/ha greater than 15 cm DBH (Johnson et al.1994). This stand now has almost 390 stems/hagreater than 9 cm DBH, most of which are smallhardwoods. The proposed R.R. Reynolds Re-search Natural Area on the Crossett Experimen-tal Forest has an average stocking of 414 trees/ha and a density of 34.4 m’/ha of basal area

2002] BRAGG: OLD-GROWTH PINE REFERENCE CONDITIONS 269

4o : 0 Frequency (trees/ha) i

3

35 : m Basal area (m*/ha) i

k 3 0 j: I

itv) 2 5

I

“Average” shortleaf

z pine standz 2 0

30 38 45-36 -44 -51 -64 -38 -58 -76 -91 -122

Diameter class (cm)Fig. 5. Freuuencv and basal area distribution for “average” shortleaf and “average” loblolly pine stands

near%urdon, A;kansas (Mohr 1897).

(Table I), with a gradually increasing represen-tation of hardwoods (Cain and Shelton 1996;Shelton and Cain 1999).

Muximum Tree Dimensions. While neverreaching the maximum dimensions of westernyellow pines like ponderosa (Pinus pondero.saDougl. ex Laws.), both loblolly and shortleafpine can grow to impressive size in the UWGCP(Table 2). The biggest pines tended to grow asscattered individuals on moist, fertile bottom-land sites (Record 1907). Individual loblollypines can exceed 5.5 m in height (Table 2), withheights of 30 to 40 m probably typical of canopytrees in most virgin stands. Mattoon (1915) be-lieved that 40 m was the maximum height forshortleaf pine, although canopy trees in old-growth shortleaf stands on poorer sites rarely ex-ceed 25 m (Mattoon 1915; Fountain and Swee-ney 1985; Fountain 1991). Loblolly pine also“rows to larger diameters than shortleaf pineb(Table 2). Pines exceeding 100 cm DBH in pre-settlement old-growth forests of the UWGCPwere not uncommon (Chapman 1942; Reynoldset al. 1984). A review of the CL0 survey notesfor Ashley County, Arkansas found examples ofpine (species were not distinguished) up to 183

cm in diameter, although most were < 120 cm(see also White 1984). Buckner (1979, p. 8) re-ported an interview with A.C. Moncrief, Sr. (along-time employee of Crossett Lumber Com-pany) who said the townsite of Crossett wasoriginally in the midst of virgin pines “. three,four, and five feet in diameter. . ” (90 to 150cm DBH). The Morris Pine (near Hamburg, Ar-kansas) was - 137 cm DBH when an early ar-ticle about this loblolly pine was published(Anonymous 1950), and currently has a diamc-ter of 142 cm.

With the possible exception of baldcypressand some select oak species, very little attentionwas given to the size of non-pine taxa. Eventhough most other taxa do not grow as tall asthe pines, many are capable of reaching heightsof 30 to 45 m. Early surveyor’s records of Ash-ley County provide numerous examples of bald-cypress, oak, and sweetgum > 125 cm DBH,especially near the bottoms of the Saline andOuachita Rivers and Bayou Bartholomew (seealso Chapman 19 13). Reynolds et al. ( 1984) not-ed that hardwoods > 60 cm DBH were commonin the virgin pine forests of southern Arkansasand northern Louisiana. Early trade journals of-

Adapted from Mohr (1897)

(both stands nearGurdon, Arkansas)

“Average” loblollypine stand

30 38 58 76 91 >I22

2 7 0 JOURNAL OF THE TORREY BOTANICAL SOCIETY [VOL. 129

(al40 3 . 336 Evenly mixed 3 . 0

3 32 shortleaf 8 loblolly pine 2 . 7f 28 II Frequency 2 . 4 3I - Basal area

a 242.1 3

L Ec 20 1. 82

Adapted from Chapman(1913)

5 16(mature and old pines) 1 . 5

1 . 2;

2 12F8

0.9 -ii0 . 8 $

" 4 0. 30 0 . 0

0 30 40 50 60 70 80 90 100 110

Diameter class (cm)

(W40 3 . 336 Evenly mixed 3 . 0

;ii 32 shortleaf 8 loblolly pine 2 . 7E 26 = Frequency 2 . 4 3! 24 - Basal area 2.1 3L 1.8 &.= 202

Adapted from Chapman (1913)(young and mature pine) 1.5 L

E 16 1. 2 k3 12E8

0.9 f0. 6 pl;

" 4 0 . 30 0 . 00 30 40 50 60 70 80 90 100 110

Diameter class (cm)

Fig. 6. Frequency and basal area distribution formixed shortleaf and loblolly pine stands south of Cros-sett, Arkansas (Chapman 1913). Chapman did notsample trees < 30 cm DBH.

ten published pictures of “trophy” hardwoodsand baldcypress, many of which exceeded 120cm DBH (e.g., Anonymous 1909). Isolated syc-amore (Platunus occidentalis L.), sweetgum,

I Lobiolly pine

Adapted from Olmsted (1902)(near Pine Bluff. Arkansas)

!II; ?ri.,,

20 30 40 50 60 70 80 90 100 110

Diameter class (cm)

04

rmm~l Shortleaf pine111 Loblolly pine

: 0.9$ 0.6

0 . 30 . 0

0 IO 20 30 40 50 80 70 80 90 100 110

Diameter class (cm)

Fig. 7. Frequency (ai and basal area (h) distribu-tions of loblolly and shortleaf pine for stands near PineBluff. Arkansas (adapted from Olmsted (I 902)).

post oak, and white oak > 100 cm DBH canstill be found across the UWGCP.

Growth und Yield. Recovering growth ratesfor presettlement pines was particularly difficult

Table 2. Maximum loblolly and shortleaf pine dimensions reported for UWGCP virgin forests.

Hciaht Diameter-Source Location Specie\ (in In) (in cm) NOLCS’

American Forests (2000) Warren, Arkansas loblolly 4s 152 1Myrtle, Mississippi shortleaf 27 1 1 2 1

White (10X4) Urania, Louisiana loblolly 54 2Urania, Louisiana loblolly 56 2

Chapman (19 13) Crossett, Arkansas loblolly/sl~ortleai~ - - > 100 3Chapman (1942) Urania, Louisiana loblolly 50 132 4

Urania, Louisiana loblolly - I37GLO survey notes Ashley County, Arkansas loblolly (‘!) 1 8 3 5

Ashley County, Arkansas loblolly/shortleaf - > 100 3

Mohr ( 1897) Gurdon, Arkansas loblolly 1 2 2This study Hamburg, Arkansas loblolly 142 6

Hamburg, Arkansas shortleaf 43 90 7Ashley County, Arkansas shortlenl 1 2 7 8

d Notes on data: 1 = current national champions, 2 = “Senlinel Pines,” 3 = did not distinguish loblolly fromshortleaf pine, 4 = “BuLlard Pine,” 5 = judging from pine’s location, this is probably a loblolly, 6 = “MorrisPine” near the Levi Wilcoxon Demonstration Forest, 7 = new state champion shortleaf pine near the LeviWilcoxon Demonstration Forest, 8 = see Figure X.

2002 ] BRAGG: OLD-GROWTH PINE REFERENCE CONDITIONS 271

Tdle 3. Average growth and yield of even-agedstands of pine “under ordinary conditions” in AshleyCounty, Arkansas (adapted from Chapman ( 19 12, p.469) and Chapman (I 9 13, p. 8)).

Ag:e ol‘ .stitntl Yield <hWlh( y e a r s ) (m’/ha). (ill ‘h/y I-)

50 29.3 0.5960 3x.5 0.6470 47.2 0.6X80 55.9 0 . 7 190 64.2 0 . 7 1

I 00 72.2 0.73110 79.3 0.73I20 X6.4 0 . 7 II30 92.0 0.7 I140 96.X ( ) . 6 X150 100.3 0.66160 102.7 0.64170 102.7 0.6 1I 80 101 .s 0.57190 9 X . 6 0.52200 94.4 0.472 10 X X . 7 0.45220 80.5 0.38230 7 I .o 0.3 I

, ’ 1,000 board feet per acre (I Mbf/ac) = 5.X3 m’ perhectare.

because early observers were interested in yield,not increment. Virgin loblolly and shortleafpines were several times more likely to producehigh annual ring density boards (> 8 rings pet2.54 cm) than those second-growth (Davis1931). The height and volume growth rates ofunmanaged loblolly and shortleaf pine declinedat 30 to 50 years of age (Mlodziansky 1896;Mohr 1897), although diameter increment can

remain appreciable for several more decades.Loblolly pine generally grew faster than short-leaf pine (Mohr 1X97; Olmsted 1902; Record1910; Mattoon 1915). For example, Mohr(1 X97) estimated that an “average” loblollywould reach 29.9 m in height and 49.5 cm DBHat 100 yr, while a comparable shortleaf wouldbe 24.7 m tall and 43.2 cm DBH.

Typically, average stand production rates inold-growth pine stands are low. Bond (1939) re-ported an average annual growth of 1.4 m’/ha (1m? = 423.7 board feet, 1 ha = 2.47 acres) foran old-growth pine forest in the UWGCP Chap-man ( 19 12; 19 13) estimated the annual volumeincrement of virgin pine forests in southeasternArkansas (Table 3) peaked at 100 years (0.73m’/ha/yr) and declined in productivity to at leastage 230 (0.3 1 m’/ha/yr). Second-growth naturalpine stands under uneven-aged management inthis same area on similar sites can grow 1.9 to3.1 mi/ha/yr, depending on initial stocking (Wil-liston 1978; Baker and Murphy 1982).

Most aboveground live volume in virgin for-ests was concentrated in a few large individuals(Fig. 8). Although individual virgin pines typi-cally yielded less than 2.1 rn?, some grew sub-stantially bigger (Morbeck 1915; Mattoon 1926;Chapman 1942; White 1984). Louis L. Morris(for whom the Morris Pine was named) statedthat when he first started to work for the CrossettLumber Company in 1907 “. there were vir-gin trees with as much as 7,500 board feet [ 17.7mi] in them. ” and “. a single log was aheavy load for an ox wagon. .” (Anonymous

Fig. 8. Only a hantll’ul of large pines could contrihutc the majority of a stand’s voltmx. This I90 year oldshortleaf pint was cut in the Crossett, Arkansas arca around 1943. The tree produced a merchantable bole I X.3m long. 127 cm diameter on the butt log, and 8 I cm diameter at the top of the top log (notice the man in thebackground). The bottom log alone producccl 3.2 m’ (I ,350 board i’eet) of sawtimber. (Picture courtesy of theCrossett City Library.)

272 JOURNAL OF THE TORREY BOTANICAL SOCIETY [VOL. 129

Table 4. Sawtimber yields reported for virgin pine forests in the Upper West Gulf Coastal Plain. Few author-sreported the log scalin, a rule they used, so yields were converted as provided (I 000 board feet per x1-e (I Mb</ac) = 5.83 m3 per hectare).

Yield (m’/ha).rmye [ averitge /

Anonymous ( 1890) Ashley County, Arkansas na” [46.7] average over X55 km” “pinecordon” area

south ArkansasPine Bluff‘, Arkansas

Pine Bluff‘, Arkansas

Arkansas

Chapman (19 12) Ashley County, ArkansasChapman (19 13) Ashley County, ArkansasMorbeck ( 15) IS) south-central Arkansas‘Garver and Miller (I 933) east Texas flatwoodsCruikshank ( 1938) northwest Louisiana

northwest LouisianaMaxwell and Martin (I 970) eastern TexasReynolds (I 9X0) Cross&, Arkansas

34.9-46.7 [na] 50% shortleaf, 50% loblollynil 134.71 pine ricige type, 64% short-

leaf, 36% loblollyna 13X.71 pine flat type, 68% lohlolly,

.32% shortleaf29.3-87.6 [40.X] pine-dominated rolling low-

lands“ordinary” pint stands50% shortleaf, 50% loblollyX0% pint, 20% hardwoods92.5%) shortleaf, 7.5% loblollypure pine typeshortleaf-loblolly-hardwoods

71.0-102.7 [na]na 147.4340.8-5X.3 [mlna [hl.XJna 174.01n3 [S I .‘,I30.0-37.0 [ml]up to 146 ]41.0]

Mohr (1897)Olmsted i I’)O2)

Record ( 1907)

,I na = not available (no value was presented in the original citation)

1950, p. 3-4). A photograph taken in 1910 inBradley County, Arkansas shows a man standingnext to a felled pine with the number “7226”scrawled on the cut face (Eagle Democrat 199 1).Presumably, as was the custom of this era, thisnumber represented the board foot tally of thetree (equal to 17.1 ml). Record (1910) reporteda single loblolly pine cut in Pike County, Ar-kansas that yielded seven 3.7 m long logs scal-ing 4.08, 3.43, 3.12, 2.83, 2.69, 2.29, and 2.19m’ (a grand total of 20.63 ml). The largest pinecut from the Fordyce Lumber Company’s landsin south-central Arkansas tallied 21 .O rn? (Mor-beck 1915) and Chapman (1942) described alobloly pine near Urania, Louisiana that scaled25.5 In3 (both are greater than some stand av-erages).

The impressive size of some pines did notusually translate into high stand yields, as manylarge-scale estimates of pine volume were sur-prising low (Table 4). For example, Harvey(1883) estimated the 51.800 km’ of pine landsin Arkansas (including areas outside of theUWGCP) averaged 18.2 to 22.9 m?/ha in treesgreater than 38 cm in diameter. Harvey’s es t i -mate predates most of the land clearing associ-ated with logging and agricultural operations, solow yields arose largely from the openness olvirgin pine stands. Spatial heterogeneity ofstocking also produced noticeable yield varia-tion. Westveld (1935) provided a figure of 58m’/h;l for a “typical” virgin loblolly/shortleaf

pine stand, with some locations approaching 175mi/ha.

Table 4 summarizes other stand-level yield re-ports for the UWGCP. Most of these estimatesplace average volume yields of presettlementold-growth pine forests in this region at 30 to70 m?/ha. Even though none of the authors notedmore than 150 ml/ha in the pine component assuggested by Westveld (1935), some limited ar-eas likely approached this volume. Few observ-ers reported non-pine yields from virgin pineforests of the UWGCP, but only scattered mer-chantable hardwoods were probably cncoun-tered. Of the pine-dominated forests of south-central Arkansas, Morbeck (1915) placed thehardwood contribution to average merchantableyield at 20%, primarily from white oak. Mor-beck’s non-pine fraction is similar to those pro-vided by Baker and Bishop (1986), who esti-mated the yield of the proposed Reynolds RNAto include 47.7 m’/ha of pine and 13.0 m’/ha inhardwoods.

Age Srruct~we. Advanced age is an impor-tant attribute of old-growth. Gaines et al. (1997)set the minimum age for old-growth consider-ation (beginning at 100 to 140 years) at one-bullthe species longevity. Recent surveys of old-growth loblolly pine stands in the southeasternUnited States have found numerous individualsLIP to 200 years old (Jones 1971; Stalter 1971;Pcderson et al. 1997). White (1984) believed

1 P,liA(;G: OI.Ib(;ROW 1’11 I‘INF I~lil~lXFiNC‘E C‘ONI~I’I’IONS 273

that loblolly pine co~tld rc‘ach 400 years old, al-though a peak age of I SO years was n~orc’ COIII-

tnon. Mattoon ( 19 IS) rcporled ;I similat. ntaxi-t n u t n a g e (400 years) I’ot- short leaf pine, andcalled ~XYX in the 200 IO 300 year ranpc “con-nion.” Cliapinaii ( I9 13, p. 6) staled that the pint(both lohlolly and shortIeal”) he inventor-ids o u t h of Crossett “. rarely cxceecicct I soyears. although occasional vei-y old trees niuyreach 200 years.” The Morris Pint was esritnat-ed to he 250 years old ii i 1050 (Aiio~iyn~ot~s19X)), and this WOLIICI place its age at -- 300years today. The proposed R.R. Reynolds Re-search Natural Area on the Crossett Iixpet-inten-tal Forest has no pints older than IS0 years(Shelton and Cain IOc)c)), but this is ;I lirrtitzttionimposed by high-gradin, 0 during lhe original hiir-vest circa 1915.

Forbes and StLtar~ ( 1930) hclieveci that rnos(preseltlcnient pine forests in the Soulh wcrc kin-even-aged. The irrcgulnr diaincter ciistribuCi0i-r inmany of the virgin pine stands reporlecl in thisstudy suggest lhal nirtltiple age classer were iti-deed present. Howevet; the l‘reclttency of fire,win&brow, and other large-scale catastrophicdisturbances WOLII~ have crts~~reci that large areasof’ the UWGCP were occupied by even-aged for-ests. As an exan~ple, TLI~IIW ( 1935) attributccithree virgin even-aged shortld‘ pine stands it1southwestern At-kansas to tornadoes.

Trw I;07777. Individtt:rl tree form i s an int-portant yet intangible property of olcl-growth.Tree character develops from cnvironmcntalconditions, genetics. aiid ape, and cott~t-ihutessubstantially to the itnpressioti of old-growth.The lack or Iowa- branches. cleacl tops, hollowstuns. fire scars, snroorh bat-k, I-eclucecl bolt (a-per, distorted crown shnpes. and low crown vig-or has of&ii been associated with old lrees (c.g.,Jones 1900; Morbeck I9 IS; 13ruttet- IYX); Chapman 1042: J o n e s I97 I ; Stnhlc and <‘haney1094), and may be as noticeable as any old-growth attribute. For inslance, Figure 3 &XII-lyshows the lla~tcneci tops and cle;tr boles of‘ theshot-tlcal’ p i n e cnnopy i n a vir”iti ,st;itiil neareHamburg, Arkansas.

The openneax oi‘ many presettlenient standsproducccl rob~is(, sprcading crown\ suppot-ted bymany large branches and STOLID holes. legaciesthat can stil l be i’o~~nd (Shelton and Murphy19%); M:rrks and Garclescu 200 I ). f3ole taper incanopy dominants wits oiien very slight, pro-d u c i n g almost colutnn~tr stems i n titc biggestpines. ~‘h~ipni;iii ( I94.Z) rcportecl ii I~7blolly pine

near Urattia, Louisiana [hat was 137 cm a t DRHand IO2 cm in diameter ;II ;t height of’ almost 30m ahovc the stump. Shelton and Murphy ( I990,p. 62 1 ) dcscribcci old shortleal’ pines in theOuachita Mountains oi‘ Arkansas a s “ . a p p a r -

ent i’rotn their slick bark, flat or clecurved branch~lll~les. and 11at “l’lY2’ cr0w114.-

l~oblolly 3ncl shortleaf pines growing in old-crrowtlt stands i i i s o u t h e a s t e r n Alabatna andenctrthertt <ieorpin had sttbstnntially clearer wood(T~thle 5) that second-growth stands in thosesane regions (Spillers l939a,h). A study of lttm-her grade recovery f1-om sawlogs (Davis 193 I)taken l’rotrt old-growth pine stands in Arkansas,Louisiana, a11c1 Mississippi and second-orowthpine fi-oni Georgia and South Carolina Gtnd ahigher proporlion of qttali~y lumber in the old-“rowth. t)vcn though boards I’rom these standscwere tnore likely to have narttral defects like de-c a y . w o r m h o l e s . shake, a n d pitch streaks (Table5).

Sl~rlictl Prrttr771. Well-stocked, managed pineii,rcsts tend 10 have their sterns evenly distributed,while relatively tindisttirbed old-growth forestsare tnore inconsistent in their spatial struclttre(Smith 1086: Tut-chin et al. 1999). Unfortunate-ly, descriptions of Ihc spatial pattern of preset-tlctiictil pine I)resls at-e decidedly less yuanti-tative than cliameter or age distributions. Heptingand C’hapman ( 1938, p. I 194) noted one of thed&growth sitortleaf pine stands they satnpled inTexas a s j*_consisting chiefly 01‘ large, oldshortleal‘ pines, with :I l.ew small pines and ascattering 01. small, poor quality hardwoods.”Historical photographs from the late 19” andearly 20”’ centttry of the IJWGCP (e.g., Fig. 9)

olien showed open pine Ibrests dominated bylarge shot-tleal‘ atxi loblolly pine. with few othetspccics apparettt (Anonymotts 190s ; I 906;Ruckner 1979: Reynolds 1980; White 1084).P’t-eqttent tires at-e ttsunlly attributed as the t-cumli)r- the open stancls and sparse tinderstories (e.g.,Olms~ctl I902), :~ldio~~gh unfavorable site con-ditions also contt-ibtttecl to die presence oi scat-tcretl prairies and woodlards i n Ashley andDrew Vounties (Wackerman 1929). Mattoon( 1915, p, 34) ntentioned that older stands ofshortlcal‘ pine a r c “_irrcgtilar in density, withiiiaiiy sirtall openings. _” Clial~iiiaii (1913, p .

6) d e s c r i b e d IIILICII o f h i s \tLtcly area s o u t h 01

C’rosaett iis

~tl’l”-oximately even-aped, but seldomcontinttotis over very large seas. It is morelikely to be broken up into diKerent age

274 JOURNAI. OF THE TORREY BOTANICAL SOCIETY [VOL. I29

Table 5. Bole quality (percent of volume) of sawtimber-si/cd shortleaf and lobloliy pines in old-growth versussecond-growth stands by various grading methods.

Spillers (I 939a,b)southeast Alabama

northern Georgia

Davis (I 93 1)”Arkansas Louisiana,I >

and MississippiGeorgia and South

Carolina

old-growth 94second-growth 31old-growth 84second-growth so

BLUld

be t t e r

old-growth

second-growth

IS

3

6 051 1XI6 04s 5

No. 1Common

and c

26

19

No. 2 No. 3Common Common

44 15

60 18

(1 Trees with 2 6. t m of clear bole and at least 50% 01‘their usable length virtually free of limbs and indicationsof knots.

h Trees with at least 3.7 m of clear bole and 30 to 49% of their usable length ii-ee of limbs and indications ofknots.

c Trees with less than 3.7 m of clear bole, or less than 30% of their usable length free of limbs and indicationsof knots.

(1 Graded using American Lumber Standards from -- 1930, with superior boards receiving a B grade or higher,and the lowest quality boards receiving a grade of Number 3 Common.

classes, clumps of large, overmature treesbeing interspersed among groups of’ youngtimber, small poles, or seedlings.

Similar accounts of patchy forested landscapesin the UWGCP has been provided by others.Olmsted (1902, p. 19) described the pines in hisridge type as “. . occur[ingj either in verysmall groups or scattered about by single trees;more commonly the latter.” He found the hard-w o o d s t o “. occur [as] single trees, quiteevenly distributed.” Pine seedlings were“ . exceedingly scarce in this type of for-est. . ,” probably due to frequent fire. Preset-tlement southern pine forests are ol’ten thoughtof as multi-aged, although this age structureprobably occurred on a greater scale than un-even-aged stands dominated by more shade tol-erant species. When developing a means to es-timate yield for inconsistent uneven-aged stands,Chapman (19 12) introduced a process thatmapped “veterans,” “mature,” and “young

merchantable” classes identified by trainedcrews. The Ashley County stand maps redrawnin Figure 10 represent the same 8 ha stand fromthe perspective of three different crews. Thoughdiscrepancies in crew interpretation make de-tailed comparison of the results difficult, thepatchy nature of these stands is apparent. Most

of the area was covered with by small- to mod-erate-sized trees, with the veterans occurring asscattered clusters or individuals. Because of thecloser correlation between tree size and age, amulti-tiered size structure in stands dominatedby shade intolerant species suggests an uneven-aged forest.

Understory and Forest Floor Conditions.Very little information on the understory andforest floor of the virgin forests exists. Standscomparatively free of undergrowth were com-monly described for presettlement pine forestsin the southeastern United States. Reed (1905,p. 13) remarked that forests in central Alabamahad “. ground cover consist[ing] of’ a thin andstraggling growth of grass and other herbaceousplants. particularly on steep rocky slopes oron the tops of the high ridges. ,” however,overgrazing may have contributed to this con-dition. Maxwell and Martin (1970, p. 2) reportedthe original east Texas pinery as

great pine stands. largely free of un-dergrowth and travelers remarked on thepark-like appearance of the forest floor.One observer pictured the forest as “in itsvirgin state there was little or no under-growth save along the watercourses, but the

2002j BRAGG: OLD-GROWTII PINE REFERENCE CONDITIONS 275

Fig. 9. The openness 01‘ upland pine s(ands in the Upper West Gulf Coastal Plain is frequently observed inhistorical photographs. This picture. taken in 1934 near Crossett, Arkansas, clearly shows the scattered largepint intermixed with small hardwoods. Note the large burn scars on the pine and open understory, suggestingthe role of fire in prescttlemcnt forests. (Picture from the U.S. Department of Agriculture, Forest Service’sCrossctt Experimental Forest archives.)

trees rose in stately grandeur t-ram a litxu-rimt carpet of the finest green.”

It was not unusual for the General Lmci Of-fice (GLO) surveyors to repot-t undergrowth asranging frotn dense to virtually absent (Delcourt1976; Foti and Glenn I99 I ). Briars and cant(Ar-urztlir~crriu gigtr~rrcr (Walt.) Muhl.) were oftenmentioned in the CL0 sut-vcys of‘ southern AI--kansns, as was the occasional pint or hardwoodsprout. Record (I 907, p. 208) I;~tnd the loblollypine flatlands of‘ Arkansas dominated by‘. aground cover var[ yingj l‘rom weeds and“‘hgrass to dense thickets of wax-myrtle [( Msr-ic,tr~Y>,-@JU L.)], br~tnil~lcs, sumac I(/Zlz~s spp.)]. andhardwood sprouts . ” and tlic shortleaf pine-dominated ridges were usually occupied by waxmyrtle and huckleberry (Vrrc~~ir7irrru spp.). Oln-stcd ( 1902, p. 19) described the undergrowth 01pint ridge areas near Pine Bluff‘. Arkansas as.L found both in large and small groups andxatterecl openly and irregularly, while over

large areas it is entirely absent, leaving theground clear and bare under mature trees.”

Mot-beck (I 9 IS) also reported open understo-ries in upland pine stands near Fordyce, Arkan-sas, with good oak and pine regeneration inmany places (see Figs. 2 and 9). Pine regener-ation in places was so succcssf~tl that Mohr(I X97, p. 1 OX) repeated a quaint proverb that inupland southern forests of’ the late 19”’ century,,. the pine is crowding out the hard-woodt i m b e r . .” Pine establishment is also aided bythe exposure of tnineral soil and the limited ac-cumulation of’ litter. Oltnsted (I 902, p. IX) haddescribed the humus as

almost entirely absent, and the groundcover consists of a thin and scattered layetof’ needles and leaves, together with grass,weeds, and f’erns. On the most open placesand it-regular patches throughout the forestare tnorc or less dense growths of’ Huckle-berry, Laurel, Swamp Bay, and briers.

276 JOURNAI, OF THE TORREY BOTANICAL SOCIETY [VOL. 120

. . . . . I. . 1. . . . . q. .. . . . . .. . . . . . . .

Lq

Veteran timber-- large(> 30 cm DBH) trees“past prime” and“decadent”

Mature/youngmerchantable timber-vigorous treesz+ 30 cm DBH

Immature timber-trees < 30 cm DBH(includes seedlingsand saplings)

Fig. IO. A~~~~roximatio~ls 01‘ stand structure developed by one of’ H.H. Chapman’s field clilsses for 8. I ha ofvirgin pint south of Crossctt. Arkansas (acl;lptcd f’rom Chapmnn (I 9 12)). Each map is a different group’s inter-pretation of structure for the same stand. While the subjectivity inherent to using multiple groups to create thesemaps is un&niable, it provides a t‘itrc glimpse of the spatial pattern of loblolly/sltortleaf pine stands early in the20” century. Note the patchy distribution of “vetcran ‘. timber in a matrix of mature, young merchantable, andimmature arc3h.

Olmsted ( 1902) attributed the lack of SLII-face or-

gnnic matter to frequent fires. As for the pineflats, Oltnsted (p. 22) noted the following:

Over consideruhle areas on these flats, andespcci:llly in the open spaces, there is adense and olicn quite high growth of grass,

and the usual ground cover ol‘ leaves,weeds, ferns, and huckleberries is commonthroughout. As on the [pine] ridges, the hu-I~US layer is exceedingly thin or entirelyabsent.

Vines of grape (Viris spp.), rattan (Berchenlicl

2002 I BRAGG: OLU-GROWTH PINE REFERENCF: CONDITIONS 277

scun&rz.r (Hill) K. Koch), honeysuckle, poisonivy (Toxicockndron rudicans (L.) Kuntze), andgreenbrier were frequently encountered by theGLO surveyors, although these species are rare-ly seen in historical photographs of UWGCPpine stands (Fig. 9 and Anonymous (1904a,b;1905; 1909)). Since climbing vines arc verycommon in modern forests, their absence in his-torical photographs suggests that their abun-dance (or at least their vertical distribution) haschanged notably in recent decades. This mayprove an artifact of an altered fire disturbanceregime, as exposed, thin-barked vines are easilykilled and even severed by light surf&e fires.Disturbance from agricultural and silviculturalpractices may have also significantly improvedestablishment conditions for many vine species,thus contributing to their increased success.

Not surprisingly, the undergrowth for currentexamples of old-growth in the UWGCP appearsquite different from presettlement stands. Chap-man (1942) cited the successful exclusion of fireas primarily responsible for the accumulation ofpine-inhibiting litter and proliferation of hard-wood and brush thickets. Dense overstories,whether pine or hardwood, effectively eliminatethe high-light conditions needed to ensure goodpine reproduction (Jackson and Harper 1955;Stalter 197 1). These patterns are similar to thosenoted for other old-growth southeastern pinestands (Lipps and de Selm 1969). The understo-ry of the proposed Reynolds RNA is dominatedby woody shrubs and hardwood seedlings andsaplings, with sparse cover of graminoids andforbs (Cain and Shelton 1994). Poison ivy,grape, honeysuckle, rattan, and greenbrier vinescan be found extending into the mid- and over-story tree canopies of both the Reynolds RNAand the nearby Levi Wilcoxon DemonstrationForest, as well as many other mature forests ofthe region.

Historical Dixturbtrrlce Patterm. Recon-struction of presettlement disturbance patternsfor the UWGCP is a necessary part of any res-toration effort intended to be self-maintainingbecause the observable features of these historiclandscapes arose, in part, from the events thatperturbed them. Old-growth is a product of adynamic environment that helps to both organizeand disassemble communities and landscapes,and when decoupled from this system, primaryforests deviated from presettlement patterns. Un-derstanding the range of variation and unique-ness of virgin forests affected by perturbation

should allow for improved restoration efforts,because it is virtually impossible to maintain anexact, unchanging ecosystem to meet statutoryor regulatory interests (Noss 1985).

Changes from presettlemcnt natural distur-bance regimes are often some of the most no-ticeable differences in old-growth stands, past topresent (Chapman 1947; Dickson 1991). Out-understanding of the patterns and processes ofpresettlement disturbance is sketchy, and spec-ulation on the dynamics of these events domi-nates our knowledge. Unique presettlcment vcg-etative states arose from differences in distur-bance type, intensity, or periodicity (e.g., Frelichand Lorimer 199 1; Shinneman and Baker 1997),and even the same type of disturbance (e.g.,wind or fire) could yield significantly differentoutcomes, especially when impacting stands ofdifferent ages. Disturbances are also inconsistentin their degree of “harmfulness.” Even thoughperturbations killed or injured at least some ofthe standing timber, this damage is beneficial toother organisms. For example, ice and windstorms often breaks branches from the crownsof the dominant pines. These wounded trees arevulnerable to heartrot, which in turn produces adecay column in the still-living pine that can becolonized as nesting habitat.

Fire. The sparsely stocked pine-dominatedforests of the southern United States have beenthought the product of frequent burning, whethernatural or human caused (e.g., Olmsted 1902;Reed 1905; Chapman 1942; 1947; Dickson199 1 ; Key 2000). Naturally- and anthropogeni-tally-ignited fires anually affected a large por-tion of the presettlement UWGCP (Hamel andBuckner 199X; Carroll et al. 2002), although itis virtually impossible to reliably quantify theextent burned. Not surprisingly, the reckless andindiscriminate use of fire and vulnerability olcutover lands spurred many calls for fire sup-pression, especially when only negative com-mercial effects like the limitation of forest pro-ductivity and the destruction of commoditieswere considered (Olmsted 1902; Reed 1905;Rothkugcl 1907; Record 1907; Bruner 1930).However, even with the tire-related problems ofthe UWGCP, some early observers recognizedtire’s usefulness to control hardwood and brushcompetition (Rothkugel 1907; Chapman 1942;Bruce 1947; Harrington and Stephenson 1955).

Shortleaf and loblolly pine. though not as tire-tolerant as longleaf‘ pine, can often withstand re-peated burning because both species become re-

sistant to fire at an early age (Olmsted 1902;McNab 1977). Young shortleaf also have a sub-stantial capacity to resprout from rootstock it’ thefire has not been too intense (Olmsted 1902;Mattoon I9 IS). Fire scar frequency increaseswith tree size (and, presumably, age): both Bru-ner (I 930) and Forbes and Stuart (I 930) report-cd research that found up to 30 to 50% ol‘ large(> 46 cm) stems had visible fire scars. Garvet-and Miller (1933) and Hepting and Chapman( 19.38) found that 9 to 13% of‘ the pines in someold-growth shortleaf stands in Texas had firewounds, while less than 2% of second-growthpines in southern Arkansas showed such dam-age. However, injuries from fire were commonin second-growth pine stands when exclusionwas inef‘fective (Garren 1941). Davis (1931) at-tributed the relatively high occurrence of’ pitchand pitch streaks in old-growth pine of Arkan-sas, Louisiana, and Mississippi to their frequentexposure to fire.

In pat-t, fires were considered limiting to pro-ductivity because they destroyed advanced re-generation (Oltnstcd 1902; Reed 1905; Chap-man 191.3; Forbes and Stuart 1930; McNab1977). Bruner (I 930, p. 23) noted that “typical”old-growth pine forests of east-central Arkansasw e r e “. growing trees 31 only 53 per cent oftheir capacity. . ,” and that “. one-fourth ofthe area was producing only at 30 per cent. .”Many hardwoods have a greater sproutilig C;I-pacity than the pines, but repeated severe tiresoften resulted in stunted and decayed survivorsand, possibly, death (Morbeck I9 IS; Bruner1930; Westveld 1935; Harrington and Stephen-son 19%).

Fire damage often interacted with other typesof’ disturbance. Chapman (1942) mentioned thatrepeated fires weakened standing trees and madethem more susceptible lo toppling. Oltnstcd( 1902) recounted the early shingle making prac-tice of’ cutting into standing live trees to checkfor their quality. Individuals that did not meet

standards wcrc left, and highly flammable pitchseeped out of the injuries. The pitch co~11d thenharden into masses and become a pathway for afire to burn into the trunk 01‘ the tree. makingthem mot-e susceptible to windthrow. Olmsted( 1902, p. 9) reported that “[al large part of the‘down’ timber on the tract has been lhrown inthis way. ,” although hc did IIO~ cluanti<y thisstatcmcnt litrthcr.

Wirltltlrrobt~. Windthrow was a major causeol’ mortality in the prescttlement pine t’orcsts 01

the UWGCI? Early surveyors commonly report-ed wind damage in their notes, with scores ofareas identified in southeastern Arkansas alone.Shallow rooting depths and high soil moisturecontributed to windthrow (Chapman I9 IX), al-though Westveld (19X) and Fredericksen et al.(I 993) downplayed the risk to lobloily andshortleaf pine because of their strong taproots.However, the UWGCP is susceptible to severewind disturbances (primarily tornadoes, frontalsystetns. and hurricanes) that can produce con-siderable damage regardless of rooting habit.

Zen ( 1905) described extensive windthrow(probably hurricane-related) in 1865, I X73, and1900 for loblolly pine-dominated stands in east-ern Texas. Tornadoes were occasional visitors tothe UWGC’P (Cole 1927). Interviews with long-time Crossett, Arkansas residents suggested tot--nadoes s t r u c k t h e area i n 1875, 1893, 1915,1919, and 1938, resulting in the loss of tnillionsof board feet of‘ standing titnber (Ashley CountyGenealogical Society 1995). Turner (I 935) re-marked that the young, altnost pure stands ofpine that arose in blowdown areas (dubbed“hurricane Ibrcsts”, although most were causedby tornadoes) tnay reflect a primary mechanismin which even-aged stands of pine were formed.

I(~(> Shower. Severe ice stortns have pro-duced widespread damage to UWGCP pine for-ests (e.g., Mattoon 1915; McKellar 1942; Muntz1947; Watts I95 I). For instance, a single icestorm in Texas and Louisiana covered at least3.2 million hectares and inflicted severe damageover one-quarter of this area (White 1944). TheUWGCP usually experiences between one andIi)ur damaging ice storms pet- decndc (Bennett1959; Cool et al. 1970; Guo 1999). Most eventsare relatively tninor, rarely exceeding I cm ofice accumulation, but some ice storms have pro-duccd mot-e than 5 cm of glaze (Bennett 1959).Mattom (I 9 IS, p. 39) reported tt glazing eventin southwestern Arkansas in December of‘ 1898that “. broke down so many trees that it com-pletely blocked road traffic over all of the tim-bered roads for nearly one week, .”

Clam slorms are particularly injurious to cx-posed. spindly, decayed. or asymtnetrical trees(McKellar 1942; Nelson I95 I; Shepard 197X).Open-grown trees are thought to be less vulner-able to ice damage than those li)und in closedstands. allhough dense stands may prove resis-tant if the spacing is such that individuals treescan support each other (Cool et al. 197 I ; Schultz1997). Kccently exposed young pines often ex-

20021 BRAGG: OLD-GROWTH PINE REFERENCE CONDITIONS 279

perience the heaviest damage (Brender and Ro-mancier 1960; Burton and Gwinner 1960; Shep-ard 1978), although severe ice storms can evendamage or kill large trees. Lipps and de Selm(1969) reported the loss of approximately one-quarter of the big pines in the Marshall Forestof Georgia from the 1960 ice storm.

In the open virgin pine forests of the UWGCP,glaze storms probably helped thin the smallestsize classes while having little impact on thelargest trees. Ice storms may also prove ecolog-ically important over large spatiotemporalscales, for they can shift the composition ofstands via differential response of species to iceloading. For example, Burton and Gwinner(1960) reported more damage in young loblollypine than comparable shortleaf pine after an icestorm struck the southern Appalachians andCumberland Plateau. The longer needles, thickerfoliage, and more rapid growth of loblolly pinemay cause it to be more susceptible to breakageor uprooting, thus putting it at a disadvantage toshortleaf pine.

Drought, Flood, cud Lightning. The occur-rence of droughts, floods, and lightning are in-dependent of the developmental stage of the for-est, but the impacts of these disturbances on theecosystem are not. Vegetative cover and age, rel-ative tolerance of extreme moisture conditionsby species, and tree size and robustness all in-teract to influence the response of forests tothese perturbations. Additionally, these factorscan contribute to the severity and extent of otherdisturbances like fire, insect outbreaks, and dis-ease by weakening live trees and providing fa-vorable habitats.

The UWCCP is periodically affected by ex-tremes in moisture (Chapman 1942; Stahle et al.1985), whether they manifest themselves as se-vere droughts or flooding. Excessively low soilmoisture can limit forest productivity, especiallyfor a region where growing season demands formoisture are substantially higher than prccipita-tion inputs (Reynolds 1958). Hardwood under-stories significantly increased the depletion ofsoil water by pine forests in southern Arkansas(Zahncr 1958), and well-stocked stands provedmore consumptive than thinned ones (Moyle and

Zahner 1954). Thus, it may be inferred that vir-gin pine forests, with their lower stocking and

sparse hardwoods, may have been less suscep-tible to drought. Floods are uncommon acrossmost of UWGCP except in the bottomlands. Ex-ccssivc water is more likely to affect loblolly

pine because this species is more abundant inlow or wet areas than shortleaf pine.

Lightning kills a considerable number ofpines every year in the UWGCP (Baker andLangdon 1990). Reynolds (1940) attributed 70%of the volume lost to natural causes over a two-year period on the Crossett Experimental Forestdirectly or indirectly (via post-strike insect mor-tality to struck and adjacent trees) to lightning.Since isolated big trees have a higher probabilityof being struck, lightning mortality could be ex-tensive in mature timber during a particularlysevere storm (Reynolds 1940). Therefore, onemay expect that old-growth forests would sufferdisproportionately higher losses than younger,even-aged stands. Lightning is also responsiblefor many of the fires in Arkansas, both past andpresent (Bruner 1930).

Insects N~LI Other Animals. Even thoughsome early writers dismissed insect damage asminor (Reed 1905), others felt it was one of themost important causes of timber loss (Chapman19 13). Curry ( 1953) described insect problemsprior to 1940 as insignificant, but that a sawfly(probably Neodiprion rue&e linmris Ross) out-break beginning in 1940-I 941 in southern Ar-kansas was of great concern. Chapman (1913)felt that up to 5% of mature pines in southernAshley County were killed over a few growingseasons by insect outbreaks (especially bark bee-tles (Dendroctonu.s spp.)). Indeed, bark beetleinfestation has caused widespread mortality tothe pine ovcrstory on the proposed ReynoldsRNA (Cain and Shelton 1996) and the MurderCreek RNA in central Georgia (Harrington et al.2000).

Insect outbreaks are often confounded withthe occurrence of other types of disturbance.Logging, severe weather, or fire can woundstanding timber, thus providing an attractant forinsects, especially bark beetles (Reynolds 1940;Garren 194 1 ; Cool et al. 197 1 ; Ku et al. 1980).For- instance, Jones (1900) decried the loss ofresidual pine timber to insects and disease fol-lowing summer harvesting in Texas. However,the thinning of mature loblolly pine stands ap-pears to help minimize the spread of southernpine beetle (Dendroctonus frontulis Zimm.)(Tut-chin et al. 1999). The lower density of vir-gin pine forests probably helped limit the sever-ity of beetle outbreaks, although much of theuniyuc spatial pat tern of presettlement old-growth (open stands interspersed with solitarydominants or small patches of large trees) re-

280 .IOC!IZNAI. Of; ‘1‘111; ‘I‘OlZIZliY I3OI‘ANI(‘Al. SOC’lft’l Y 1 VOL. I29

sulted from earlier beetle kills. Insect clanragcpatterns have also changeci tiuc to the loss olold-growth habitat of woodpcckera, one of theimost imporrant nalural predatoi-s.

Native and introduced n~an~n~als can alsocause tree mortality. although rarely XC theydamaging to large trees. While presettlcmei~t es-timates of deer densities arc uncertain (Schoenet al. 198 1 ), white-lailed deer (I~c/o~~~~i/c~r.r i~ir-gir7ianms Zimm.) herds in eastern North Americahave grown rapidly in recent dccadcs. I .eopoldet al. ( 1947) mapped much of the I JWGCP ashaving few to no deer, although this probablyreflects an incomplete recovery of deer popula-t ions I’dlowing their ne;lr extirrclion earlicxr inthe century. White-tailed deer (as well as anyother overabundant herbivore) can alter forestsuccession through lethal browsing of seedlingsand saplings. Mortality to yo~mp tree can alsoarise from the polishing of‘ deer antlers on treetrunks, but this is not a major problem in rltostareas. Other large native u~~gulates like hison(Ri.wr7 hison L.) and elk (Co-i1lf.s ~~/LI/~/IIIS I , .)were probably of little consccluence in the pineli)rests of the UWCCP before they were cxlir-pated.

Introduced livestock, however. did post a sc-rious threat to I‘oresls, at least by the early 3(Ycentury. Feral hogs GILISC localized problems byconsuming the mast of’ many spccics, damagingunderstory vegetation, and uprooting pine seed-lings (Maxwell and Martin 1970; Wood andLynn 1977). Interestingly, Olmsted ( 1902) de--

lied conventional forestry wisdom by promolinghog ranging on land near Pine Hluff. Arkansasbecause the hogs cons~~mcd mostly acorns andapparently did little damage to die prcfcrred pineseedlings, while Ihcir li)ruging improved dieseedbed. Much of‘ the Inn&cape was convertedto pasture following the removal of the virginforest, with some lumber companies importingexotic breeds of cattle in an i~nsucccs~f~~l tit-tempt to encourage pustoralism (and, lheorctially, increase the value of their cutover property) ( M e t y 1952: Kcynoltls 1980). IFirc wascommonly used to suppress the pines, ha&woods, vines, and briars that could rapidly re-colonize pastures, but when cnrelcssly applied.these fires dnmagcd uncut timber (Rod~kugcl1907; StraLisberp and Hough 1097). Hunts alsofreclucntly scl lirc 10 southern li)ratc Lo iiiipuveconditions for game, alien with little regard tothe consequences to tii1ihcr and far-ms (I;os,lei1912; Key 2000).

The changes wrought by settlemcn~. land dc-

veiopin”nl, and gai-tte ll~anagcnrcnl h a v e Lln-cloubtably infltiencetl animal communities(Schocn ct al. I98 1 ; Dickson I99 1 ; Hamcl andBucknet- 199X) and thus alterctl the potential tocompletely rcslorc historical forest conditions.The red-co~katlecl woodpecker, for instance,seeks out nta(urc live pints infected with redheart in open wootlla~ids. Large pines infectedwith red hcnrt arc c0111n~on in olcl-growth (e.g.,Nelson 193 I ; Jones 197 I) and the virgin forestsalien hurl sparse. forh- and grass-dominated un-derstorics chnraclerislic 01‘ frccluently b u r n e decosystems (Bukenho~cr et al. 1994; Carroll etal. 2002). ‘l‘hr eliininalion 01‘ this habitat acrossmuch of the southern linited States has pushedthe red-cockaded woodl~cker to the brink of ex-tinction (Steirly I953: James and Burnside 1979;Lennart/ IOXX). The systematic management ol-‘l’oresls primarily f01 game species has also un-clouh~edly affected die dynamics OF the system(Wood and Idynn 1977; Key 2000). especially byfavoring mast species and early successionalco\~el- rypW

Ncorr Rot turd Orho- Trw Drcqy. The decayol‘ live lrees i,s an allrihurc that olten manifeslsitself most nolably in d&growth stands. Healthytrees are usually vipororis enough to repel moslinvasions, but with time and the accumulationof damage, pathways for infection become moreprominent. Youiig shortleaf and lohlolly pine areusually resistant to the principal heart rot of theregh, “red heart” or “red rot” (primarily li-omthe ll~ngus Phc/li7rr/.s piui Ames). Slow growthand ;I high proportion 01‘ heartwood are impor-kinI factors in the tlevelopnient of heart rot inmany soulhern pines (Lightle anti Starr 1 X7),which typically enta.s throuph branch stubs ortire scars (Hepling and Chapman 193X; GarrenI94 I ). Al‘feltranger ( IO7 I ) reported the tdlow-ing factors plnyecl an important role in red heartpresence: ~lancl age (especially for ~inevcn-agedstands); Iarge, pcrsistcnl branches; increasingproportion 01‘ pine: ;intl excessively drained,shallow, or soils with higli nitrogen content.

It is not unusual to spot red heart cankers onold pines in historical photographs (e.g., Anon-yll1olls 1004h; I906; c’hupnKln IO 1.3). Roth zon( I 005. for loblolly pine) and Marloon ( 19 15, I'mshorllcaf pine) cle~cribcci heart rot its rare in treesless than ;i century old, bcir increasing beyondthis age. This trcntl is supported by nuIncrousother authors (Table h), who noted increasingcull proportions from decay in oltlcr stands. Forcxamplc. a m i l l Iun1her recovery sample con-

posed of a mixture 01‘ shortleal‘ and lob lo l ly pinefrom virgin stands in Arkansas, l,ouisianu, and

Miss iss ipp i had tnany t i m e s grcatcr occu r rence

of’ shake , wortn h o l e s . d e c a y , p i t c h , and p i t c h

s t r e a k s t h a n one laken l‘t-om seconcl-growthstands in Gcorgin a n d S o u t h C‘;trolin;r (I>avis19.11).

Lcrrgr Wootly Orhris (I- WI)). The ~CCLII~LI-

lation of‘ L W D has b e e n recogni/etf iis an itn-

portant indicator of’ old-prowrh conditions (Ma-

tin I99 1 ; Dcvall and Ramp 1992: Spe(ich ct al.1999), but Ihe hutnitl clitnate und ahuntlat~t de-

t r i t i v o r e a n d deco~ripocitio~i~rl conitnunities 01

the UWGCP con t r i bu te f o tqid LWD loss ( see

Cain 19%). LWD longevity depends on species,

wood density, Ihe manner of tree death, the local

tnicroclitnate, and other losses due to salvaging,

lire, o r cielritivores (Mohr I897; L o n g 19 1 7 ;

Harmon e t al. I986; Van Lear 10%; Spelich et

al. 1999). Decay resistant heat-twood makes up

a larger fraclion oi’ hole vol i in ie with increasinga g e , w i t h o l d pints olicn possess ing tw ice rhc

prol~ot-(ion of‘ heartwood 21s younger s;tcnis (Da-

vis I93 I ; J>>emnion 19.36). While pil~/i-s;t(ttr;t1c’cff

hear twood in o ld . s low-growing t rees resists b i -

o t i c decotnposirion, th is “ r i ch p ine ” i s h igh l yflammable and therel’ore more sttsceplible to lit-e.

Reconstt-uclion 01‘ prescttlcnxxtt LWD loads isdi f l iculc heca~rse of‘ tlte lack of historical reporrs.L igon ( 197 I ) believed die occtfl-rcnce 01‘ srancl-

ing dead timber in southern l’oi-ests wits uitcon-tnon in l~rcscltlemcnt rimes because of its coti-sitniption by fire (see also Westvelcl 1935). Eveirthough he did not spec~ilate on lhe C;~LISC of themortality, Zen ( IOOS) noted the presence of” cie:tdlohlolly p ine in Iii\: 3>ua5 pint \tairtls (coillpi-i\-

-

slalld age I’wccnt

, ycx\) c u l l rolnmenlal-y

70-x0 /I reel heart it3 loblotly“01d” 3-10 red heart and butt rot60 65 2 hcwirot i n d~orrleai

170 I7 dixascci shortleaf“vit-pin” 1 l -14 utl5ound logs at sawmills40 -90 5 red heat-t in lohlollyoo-- 140 I9 red heart in lohlolly

140 100 ho red heart in loblollyl90~-230 72 reel heat-t in lohlolly“7”“ p,jwQ,” 1 loh lo l l y and shot-tlcitl‘~oltl-gr-owth” 7 iltortlcal65 21 \Iow growing lohlolly and

,shorllcnt’90 6 ol&lieltl Iohlolly

“vit-gill” I4~--

inp from 2 to 13% of a11 s;tetns). Studies ot‘co11-t e m p o r a r y olci-growth have J’ountl I*WD v o l -umes in the range of 30 I0 >. 300 ni’/hn in east-ern li)rests (c.g.. Harmon et a l . I986; McMinn

and I-lx& 19%; Greenberg et a l . 1997; S p e t i c h

e t al. IYN; Zhang 2000). Cain and Shelton( 1996) encottnteretl 8 to 56 snags per hectare int h e R e y n o l d s R N A , prinxrrily la rge p ines k i l led

by bark beetles. Decadent (i.e., hollow, spike-topped. or otherwise dnniagccl) live 1rccs lend tobe m o t - c common i n untnanaged o l d - g r o w t h

(I)avis 1031: Greenberg c l al. 1907).

Numtrrr /r?//~rr/~c~~.c. The l ong h i s to ry o f hu -

man seltlerncnt in die southeastern United Statescotripl icnles the understanding of presettletnentli)rest cond i t ions . Changes in agr i cu l tu ra l and

settlenienl p rac t i ces . popu la t i on dens i t y , and

comntcrcial interests resulted in a patchwork of

successional stages and sland structures. NativeAmer icans c leared land fix agr i cu l tu re , then

aha~ncioned their f i e lds when the p r o d u c t i v i t y

dropl~~I. t he Itrtners did, or the t r ibes werei‘orcecl 01‘1‘ o f their lands. E a r l y Euroamerican

seltlers ol‘tcii i i~l lowetl similar pat terns 01‘ landUSC: Wltite and I, loyd (I 095) described a stand01‘ .a old-growth” in Sottth Carolina (the 200 yearold John de I;t Ilowe Tract) [hut arose followingtlevelopti ient in the iii& 1700s.

Native Anrericmns occupied the UWGCP forttt i l lcnnia bel’ore Euroantet-icnn development he-gan in the seventeenth mtl eighteenth centuries.I<veli thottgl t Native American populdon esti-m:t(cs XC ttttcertain for the UWGCP at the timeof‘ the lit-St contact with Europeans, they proha-b l y IXII into the ntany thousands un t i l d i sease

:intl strice recli.icetl them 10 perhaps 5 IO 10% ot

282 JOURNAL OF THE TORREY BOTANICAI, SOCIETY [VOL. 129

Fig. 11. Intensive logging begun during the late 19”’ and early 20”’ centuries continues to the present day,although very few pines this size remain in the Upper West Gulf Coastal Plain. (Photo courtesy of the CrossettCity L.ibrary.)

their historical levels (Schambach and Newell1990; Carroll et al. 2002). Alter this reduction,the landscapes of the UWGCP were sparselypopulated for many decades until the beginningof the 20”’ century, during which they were pri-marily used for subsistence agriculture, grazing,hunting, and trapping. Conflicts over the use olthe land after Euroamerican exploration has-tened the anthropogenic perturbation of theUWGCP (Key 2000) and the removal of NativeAmericans and subsequent development broughtthis region starkly different disturbance regimes.

Small-scale timber removals began in the1830s and 1840s in southern Arkansas (Ether-idge 1959), but due to the primitive sawing tech-niques, lack of local markets and labor, the Civil

War and Reconstruction, and difficulty in trans-porting materials, extensive exploitation waiteduntil late in the 19”’ century (Curry 1953; 1960).Commercial harvesting of the virgin timberlandsof the UWGCP began in earnest by the 1880swhen railroads were extended to most areas(Anonymous 1904; Balogh 1985) and continuedrelatively unabated for the next half-century(Fig. 11). Many lumber companies acquiredtheir land holdings l’rom local farmers, who wereonly too glad to get rid of their “worthless” tim-berlands (Morbeck 1915; Curry 1953; Buckner1979; Reynolds 1980). Only the largest trees(those > 30 cm in stump diameter) in the virginforest were initially considered desirable (Mor-beck 1915; Mety 1952; Reynolds 1980). Fires

were common in the slash and litrle effort wastaken to protect the remaining forest.

Scientifically-based forestry was virtually un-known at the ~LI~II of the 20”’ century in America,so lumber companies cxploitcd the forests untilexhausted, then tnoved on or closed permanently(Curry 19%; Reynolds 19X0; Shipley 19X7). Forexample, the Crossett Lumber Company had nointention of engaging in forest management untilthe late 1920s when it became apparent that theirattetnpts to promote their cut-over lands as ag-

ricultural and pastoral properties wo~ild not SLIC-wed (Reynolds 1980). By 1922 only abo~~t

16,200 hectares of the company’s original101 ,000 hectares contained virgin titnber, bolt itwas 192X before cutting was reduced to prolongthe harvest of the remaining timber (Curry1953). The recognition of titnber shortages and

expansion of professional forestry eventually in-duced some of the remaining timber companiesto tnanage for sustained production. Effectivefire suppression was implemented in most of‘UWGCP by the mid-1930s (Mcty 1952; Reyn-olds 1980). Fire exclusion further altered thenatural dynatnics of the region’s pine-dominatedforests, contributing noticeably to rueI accumu-lation, expansion of hardwoods, brush, andvines, and altered pine regeneration success(Chapman 1942; 1952). In the intervening de-cades, tnany natural upland stands have beenconverted to plantations, and landscapes havebeen continually fragmented.

Considerations and Cautions of Using His-torical Documentation. Signilicant ecologicalchange during the past two centuries of settle-ment, fire exclusion, forest t~ia~xige~~ictit, and

landscape development suggests that currentfragments of old-growth stands are inadeyuateexamples of presettlement conditions. Addition-ally, contetnporary reference sites at-c often toosmall to provide the appropriate sP”tiotetnpor~tlcontext, or tnay be the product of’ an historic butt-are event atypical of the desired ecological slate(White and Walker 1997). Judging ft-mm this rc-view on old-growth pine li)rests in Ihc UWGCP,constructive information li)r ccosystcm rcstora-(ion can be gained from the usscmblage of his-torical ~OCLIIII~~IS, photographs, and inventories.Even though not all areas have such ;I wealth ofsuch docLtiiientatioti on early forest conditions,the presettlement conditions li)r marty locationscan be bcttcr dcscrihed through this approachIhan from the examination of remnant standa.

Those involved in t-cstot-ing prcseftlement

conditions should first spend as much time aspossible reviewing historical sources beforemodicying existing comtnunities. This is trueeven if the information is limited to qualitativedescriptions or period photographs, as theseSOLII-ccs can at least define reasonable boundaries(Egan and Howell 2001 ). However, the use ofhistorical references is not without challenges.Sources of yuantitative data, such as titnber ill-ventories, may date back to target periods, butcan he short on detail regarding minor or non-commercial taxa. Historical research reports orinventories rarely provide more than basic sum-maries, but are still highly relevant sources ifavailable.

Bc careful not to over-analyze historical datathat were not collected specifically for ecologi-cal purposes, for this tnay lead to improper as-sumptions on reference conditions. The earlyCL0 surveys, though an important window topresettletncnt vegetation patterns, have oftenbeen inappropriately interpreted (Whitney andDeCant 2001 ). For example, surveyors were in-structed to select witness or tnarker trees thatthey felt had the best chance of long-term sur-vival (Stewart 19.15; Bureau of Land Manage-tnent 1947), and thus may not reflect either thesize class or compositional distribution of thestand in which they were chosen (Bourdo 1956).

Inaccurate descriptions can often be found incolloquial accounts of vegetation, as many earlyobservers WOLII~ comment on species withwhich they were unfamiliar, or described con-ditions with unscientific zeal. Some early trade@trnal articles consistently refer to the pine for-ests in their photographs as shortleaf, eventhough loblolly was obviously present (e.g.,Anonymous 1904n,b; see also Mot-beck 19 IS).This usage of “shortlcaf” was probably tneantto differentiate PI’I~Ks ra&r and P. eclzincm frotnP. pu/u.stris (true longleal‘ pine). Blatantly incor-rect material also occasionally appears in sci-entific or technical accounts. For example, anearly paper by Professor FL. Harvey reportedthat 1176: botanist Thomas Nuttall had identifiedtwo pine species (P. I+~~L/o Mill. and P. ir7op.sAit.) in Arkansas (Harvey 1X83), neither ofwhich naturally occurs anywhere near the state.

Conclusions. Once the ob,jectivcs of the res-toration have been identified, developing refer-cnce conditions for the area to be reconstructedis ;I critical tirst step in designing an effectiverestoration program. To do this. one must be-come familiar with the history and documenta-

284 JOURNAL OF THE TORREY BOTANICAL SOCIETY [VOL. I29

tion of the area of interest (Egan and Howell2001). Records from the pioneering efforts ofGLO surveyors and those seeking to bring sci-entific forestry to the South also proved invalu-able, and even the photographs and personal ob-servations of early settlers have contributed tothis effort. Although many sources of informa-tion were strictly qualitative, enough yuantita-tive data co~tld also be gleaned to produce a listof specific management targets. For instance,data on species, size class, and tree density dis-tributions were readily available for virgin pineforests of the UWGCF! More valuable informa-tion was thus gathered than could be gainedfrom the structural analysis of any existing re-search natural :lrecl or demonstration forest.

This effort allows those interested in the re-construction of historical conditions to envisionwhat virgin pine forests of the UWGCP werelike before Euroamerican settlement. Usingthese historical sources as a guide, the virginpine forests often appeared as open stands, withextensive grass and forb understories only oc-casionally interrupted by clumps of shrubs andtree saplings. Overstories in most upland pinestands were dominated by irregularly scatteredshortleaf and loblolly pines. Nearby locationswith better site quality have increasing levels ofmature oak, hickory, and gum, with scattered su-percanopy pine. Frequently, these isolated over-story pines were very large and centuries old,frequently fire-scarred, with twisted crowns,smooth bar-k, and abundant heart rot. Largetwisted piles of downed trees covered in vinesand briars provided further evidence of the nat-ural catastrophes that periodically swept downupon the virgin pine forest of’ the UWGCP. Pineregeneration, when not destroyed by frequentsurface fires, was thick in exposed mineral soil.Even-aged patches of young and maturing pinearose from the gaps formed by windthrow, bee-tle outbreaks, tires, or ice storms. The juxtnpos-itioning of young, mature, and veteran timberover much of the landscape left the impressionof an uneven-aged landscape, with multiple tiersof crowns. Red-cockaded woodpeckers wcrccommon visitors to the decadent old pines, andelk and perhaps even bison joined the white-tailed deer to grake the available forage.

While it may not prove possible to extensive-ly reconstruct the virgin pine forests of theIJWGCP on ;L large scale, understanding therange of variation, dynamics, and uniqueness otvirgin forests should improve restoration ei’forts

like those being attempted on the Gulf CoastalPlain of southern Arkansas.

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