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The vegetation of Pulliam Prairie, Chickasaw County, Mississippi: a significant remnant of pre-Columbian landscape in the Black Belt.

INTRODUCTION

The Black Belt of Mississippi and Alabama is a distinctive calcareous section of the Upper East Gulf Coastal Plain in the southeastern U.S.A. It is largely underlain by Upper Cretaceous sediments that are mostly "chalk and marly chalk containing fewer impurities than underlying and overlying formations" (Moore 1985). There has been a substantial recent increase in research on the native vegetation of this region during recent years (Peacock and Schauwecker 2003, Barone 2005, Barone and Hill 2007, Schotz and Barbour 2009, Campbell and Seymour ....a). Much attention has focused on remnants of the characteristic native grasslands or "prairies" of this region, but it has been difficult to describe the full pattern of varied grassland and woodland types that existed on the landscape before first European contact. Because of general agricultural conversion, individual study-sites rarely allow insight to the broader patterns. This paper describes vegetation at the Pulliam Prairie in eastern Chickasaw County, Mississippi, a tract of ca. 250 acres [100 ha] with much evidence of the original gradients in composition related to topography and soils.

Most good remnants of Black Belt prairie are only 5-20 acres [3-12 ha] in size, but a few cover up to 500 acres [200 ha] if adjacent thickets and open woodlands are included (Wiygul et al. 2003, Barone and Hill 2007, Schotz and Barbour 2009). In Mississippi, the Tombigbee National Forest has begun to restore significant remnants within ca. 500 acres [200 ha] of Pontotoc County (Wiygul et al. 2003), a few miles north of Pulliam Prairie, and there is potential for a much broader effort in this neighborhood, given the several nearby pastures and roadsides with variously degraded remnants of the original grassland (our pers. obs.). The Osborn Prairie includes ca. 178 acres [72 ha] of grassland in Oktibbeha County ("Sixteenth Section Prairie"), and is currently leased and managed by Friends of the Black Belt in cooperation with Mississippi State University (Wiygul et al. 2003, Hill and Selzer 2007). A smaller remnant--the Morton Hill Prairie--occurs at Noxubee National Wildlife Refuge (Noxubee Co.), where restoration is also proceeding (Hill 2004).

In recent years, there been increasing focus on conservation in the Black Bel involving several governmental, academic and non-profit partners (Wiygul et al. 2003; Schotz and Barbour 2009; D. Coggin and J. Hill, pers. comm.). In addition to supporting healthy populations of deer and turkey, marginal farmlands and old fields in the Black Belt form a prioritized region for recovery of quail in the Mississippi. These efforts will clearly benefit from more detailed understanding of what the native vegetation was, where the best remnants are, and how they can be restored under current conditions.

The Pulliam Prairie offers special insight to the original landscape since it includes varied soils and native vegetation types. Also, there are several nearby woods and fields with some degree of natural quality, covering 100s of additional acres for extended study in the future. In addition to extensive grassland on the more calcareous slopes and adjacent uplands, there are strips of wooded riparian zones and patches of woods on the broader ridges. Although much of the area has been farmed to some extent, there has been little persistent cropping on the uplands, and diverse native species have recovered in many areas. According to local history, the Pulliam family purchased land in this neighborhood from the Chickasaw tribe during the 1820s. In recent years, they have consolidated some of the tracts. They have used the areas with native grassland primarily for hunting, and have applied fire to maintain the open character.

SITE CONTEXT: TOPOGRAPHY AND SOILS. The Pulliam Prairie lies within the western side of the Black Belt region (Fig. 1), mostly on relatively gentle topography (Fig. 2). It drains into Sakatonchee [or Chuquatonchee] Creek, a major tributary of the Tombigbee River. The broad ridge west of these slopes is connected to the southern terminus of the Pontotoc Ridge (Lowe 1921, Harper 1943), which is an extension of the "Northern Hilly Gulf Coastal Plain" that has been mapped as an ecological region here and across much of north-central Mississippi (Chapman et al. 2004). Elevation around the Pulliam Prairie ranges from about 250 feet [76 m] asl along Sakatonchee Creek to 400 feet asl [122 m] on ridges west of the study area. The Pulliam tract itself is mostly at 260 to 310 feet asl [85-95 m], and the mile-long corridor along the access road from the closest paved road extends up to about 340 feet asl [105 m].

Campbell and Seymour (...a; their Fig. 1) have outlined the overall patterns of variation among soil series of the Black Belt. Soils of this region (USDA 2010a), and at the Pulliam Prairie or nearby, can be arranged conceptually along two largely independent gradients (or "catenas"). There is an elevation-related gradient, from alluvial lowlands, to the chalky soils on gentle side slopes, to the overlying clays and sands on broader ridges. And there is a drainage-related gradient, from poorly drained flats with generally deeper soils, to better drained slopes with generally shallower soils. Most soil series on the chalk itself are classed as vertisols--with expansive "shrink-swell" clays, and "self-mulching" of organic matter into deep A horizons. More local soils include entisols (especially on more recent colluvium/alluvium), inceptisols (especially on deep/damp colluvium/alluvium), mollisols (presumably with more stable grass, cane or cedar cover), and alfisols (on more weathered loamy uplands with more woodland history just above the chalk). In contrast, loamy ultisols (with generally less base saturation) predominate above the chalky soils, on more sandy uplands or on high terraces with a history of woodland.

Soil series mapped by USDA (2010b) at the Pulliam Prairie cover the full range of catenas illustrated for the region by Campbell soils on chalk (Binnsville and Sumter series) and the ultisols on more sandy ridges are not mapped here. Also, several intervening soil series along the catenas are not mapped here, though they may be present to a limited extent. Within three miles of our study area, a more complete range of soils is represented, including sandy transitions to the Pontotoc Ridge.

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METHODS

We made six collecting trips to the Pulliam Prairie in 2009, covering 14 days: April 20-21, June 6-8, July 10-12, August 22-24, September 24-25 and October 28. During each trip, except the last, we systematically covered the study area, using an electric "Stealth Predator XR" vehicle for transport and collection platform for herbarium specimens (Campbell and Seymour ...c). On each trip we visited most or all sections of the study area, defined as 15 grid units of 1000 x 1000 feet [305 x 305 m].

We assigned initial codes for habitats at multiple sites within each grid unit, in order to map vegetation and to provide label data for our collected species. These codes were as follows: W1a = Cedar Woods; W1b = Post Oak Woods; W2a = Oak Hickory Woods; W2b = Riparian Mixed Woods; W3a = Green Ash Woods; W3b = Willow Oak/Water Oak Woods; S1 = Dry Scrub; S2 = Seasonally Wet Scrub; S3 = Riparian Scrub; G1 = Grassland on eroded/dry upland soils; G2 = Grassland on moderately dry soils; G3 = Grassland on deeper/damper soils. We eventually converted this system into the vegetation types outlined below. In the Appendix to our accompanying floristic paper (Campbell and Seymour ....c), we used a related series of codes to indicate the typical position of each species along ecological gradients.

In addition to our field work, we used recent aerial photographs (from USDA Farm Service Agency), topographic mapping (USGS 1987: 7.5 min topographic quadrangle "Buena Vista"), and soil mapping of NRC (http://websoilsurvey.nrcs.usda.gov/app/ WebSoilSurvey.aspx) to interpolate our observations over the site. Also, we used our broad regional review of soils (USDA 2010a) and vegetation in order to refine our definitions of habitat types (Campbell and Seymour ....a). We incorporated these observations into a map of vegetation types at the site, and into a diagram of the major gradients in vegetation. Botanical nomenclature primarily follows the list of vascular flora in Mississippi that is being developed at the Pullen Herbarium in Oxford (McCook and Kartesz 2010), based initially on Kartesz (1999). For a few taxa, we use significantly different names, which we detail and explain in the accompanying floristic paper (Campbell and Seymour ....c).

RESULTS

The following notes outline composition in each vegetation type at the Pulliam Prairie. This general classification matches the regional outline presented already, based on review of the literature, with the same lettered codes from "a" to "n" (Campbell and Seymour ....a). Fig. 3 presents a map of vegetation over the site. The mapping units correspond to the vegetation types outlined below, as much as possible, but several mapping units include transitions and mixtures, as explained in the key. There is, of course, much intergradation between types. The lists of characteristic species provided below are best estimates, offered here for continual revision with further field work on the whole flora. For linkage with our accompanying floristic paper, the ecological codes given to characteristic species in that paper are added in brackets after each vegetation title below [with "C&S ....a"]. Fig. 4 presents the scheme of ecological relationships among our interpreted vegetation types.

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Uplands with Acid Soils Overlying Calcareous Sediments: including damp flats

(a) Oak woods on ridges and knolls [see species A1/A2 and B1/B2 in C&S ....c]. There are virtually no areas of such woods at the Pulliam Prairie itself, but they are scattered near the paved road a few hundred meters to the west. In older woods, post oak (Quercus stellata) is common, with most canopy trees 2-4 dm, plus a few 5-6 dm or more. Other trees include southern red oak (Q. falcata), blackjack oak (Q. marilandica), hickories (Carya glabra,, C. ovata and C. tomentosa) and winged elm (Ulmus alata). A distinctive shrub is Vaccinium arboreum. On the ground, there are patches of Vitis rotundifolia, scattered graminoids (e.g. Dichanthelium dichotomum, Eleocharis cf. tenuis) and legumes (e.g. Desmodium laevigatum, Lespedeza intermedia).

(b) Open oak woodlands and thickets [see species A3 and B3 in C&S ....c]. On uplands within the Pulliam Prairie, there are a few patches of thin woods dominated by post oak or blackjack oak, plus scattered winged elm. These woods have been burned, browsed or otherwise disturbed for at least several decades, and probably much longer. In one area at the head of an ephemeral stream, there are several larger water oaks (Q. nigra) in a transition from post oak to riparian woods (see j below). Hickories are curiously absent. Loblolly pine (Pinus taeda) has been planted on similar soils near some of these woods, but it is probably not native to the site. In more open areas or at woodland edges, there also some distinctive shrubby species, but mostly in small numbers: Vaccinium arboreum (on more acid soils), Rhus copallina (locally abundant in transitions to grassland), Toxicodendron pubescens, Ceanothus americanus, Crataegus engelmannii, Hypericum hypericoides, Rosa carolina and Rubus spp. (with R. trivialis locally common). The alien Japanese honeysuckle (Lonicera japonica) is frequent in some areas. On the ground, frequent graminoids include Carex spp. (leavenworthii, muhlenbergii, cf. umbellata), Danthonia spicata, Dichanthelium spp. (dichotomum, linearifolium, cf. meridionale, sphaerocarpon), Panicum anceps var. rhizomatum, and Scleria spp. (especially triglomerata). Locally abundant herbs include Helianthus spp. (especially hirsutus and pauciflorus) and Verbesina helianthoides. Other characteristic species include Desmodium spp. (glabellum, laevigatum, marilandicum), Lespedeza spp. (intermedia, procumbens, violacea), Phlox spp. (carolina, pilosa), Ruellia cf. ciliosa, Salvia lyrata, Stylosanthes biflora, Symphyotrichum patens,, Tephrosia virginiana and Tragia urticifolia.

(c1) Relatively well-drained grassland on ridges [see species A4/A5 and B4/B5 in C&S ....c]. Most of the ridges with more acid clayey substrate above the chalk have been farmed, especially on deeper soils. However, the older fields have a diverse mix of native species, and there are some less disturbed areas, especially on old roadsides, in swales and other transitions down to the chalk grassland (see d, e and f below). Locally abundant grasses include broomsedges (especially Andropogon virginicus), switch grass (Panicum virgatum), Indian grass (Sorghastrum nutans), and little bluestem (Schizachyrium scoparium). Other graminoids are characteristic, including Aristida spp. (longespica, oligantha), Carex spp. (especially bushii and hirsutella), Gymnopogon ambiguus, Muhlenbergia capillaris and Scleria spp. (including ciliata). Locally frequent herbs include Agalinis spp. (especially purpurea/fasciculata), Ambrosia psilostachya, Brickelia eupatorioides, Buchnera americana, Cirsium discolor, Desmodium spp. (canescens, glabellum, sessilifolium), Eryngium yuccifolium, Eupatorium spp. (hyssopifolium, pubescens, serotinum), Helenium nudiflorum, Helianthus spp. (especially mollis and pauciflorus), Liatris spp. (spicata, squarrosa, squarrulosa), Linum medium, Solidago spp. (including nitida), and Symphoricarpus spp. (ericoides, dumosus, patens).

(c2) More poorly-drained grassland on ridges; somewhat xerohydric [see species A4/B4 as well as H/HX in C&S ....c]. On broader flats, there are seasonally wet conditions in some areas, but generally less than an acre in extent. There are a few small ephemeral ponds that appear to be at least partly artificial. Composition is highly varied, depending on hydrology and disturbance history. Several graminoids appear to be characteristic: Andropogon tenuispatheus, Cyperus spp. (echinatus, pseudovegetus), Dichanthelium scoparium, Eleocharis spp. (compressa, erythropoda), Juncus spp. (acuminatus, biflorus, effusus, scirpoides), Saccharum spp. (contortus, giganteus), and Tridens strictus. Also, Tripsacum dactyloides has filled a small brushy pond. There are few characteristic broad-leaved herbs, but the rather weedy Eupatorium serotinum is relatively frequent. Uncommon species of interest include Mecardonia acuminata and Spiranthes vernalis. One of the ephemeral ponds has abundant Potamogeton diversifolius, and is lined by Trachelospemum difforme.

Uplands with Calcareous Soils: mostly on slopes. Note that the grassland types (d), (e), (f) and (g) outlined below are usually intermixed at small scales. In areas larger than 0.1-1 acres [0.04-0.4 ha], it is often impossible to assign a predominant type.

(d) Upland grassland on shallow, partly bare soils; mostly xeric-tending or xerohydric [see species C4/C5 as well as X/XH in C&S ....c]. A more detailed analysis of microsites would separate further variants here related to degrees of soil formation versus erosion. In general, the most common graminoids include three-awns (especially Aristida longespica), dropseeds (Sporobolus vaginiflorus, S. clandestinus, S. compositus var. drummondii), little bluestem (Schizachyrium scoparium), glade panic (Panicum flexile), switch grass (P. virgatum), sedges (Carex crawei, C. meadii) and glade irisette (Sisyrinchium albidum). Locally frequent herbs include Agalinis spp. (gattingeri, oligophylla), Asclepias spp. (especially viridiflora), Coreopsis lanceolata, Dalea spp. (candicans, purpurea), Erigeron strigosus (with some var. calcicola), Eurybia hemispherica, Houstonia spp. (lanceolata, nigricans), Hypericum sphaerocarpon, Liatris spp. (especially squarrosa var. alabamense), Linum sulcatum, Lobelia spicata var. leptostachya, Ruellia cf. humilis, Spiranthes magnicamporum (especially in disturbed, puddled areas), Solidago spp. (especially nemoralis) and Symphyotricum spp. (especially laevis var. purpuratus and patens var. nov.). Alien species are usually rare or absent.

(e) More disturbed variants, often transitional from shallow to deeper soils [see, especially, species C5 in C&S ....c]. Composition appears related to disturbance history and gradients in soil. There is much intermixing among species characteristic of drier and damper types (from c to g), especially where the ground is plowed, gullied, compacted or stripped of organic matter. More weedy grasses tend to predominate, including several annuals in Aristida, Eragostis and Panicum. Typical perennial grasses include broomsedges (Andropogon virginicus and its allies) and locally Muhlenbergia capillaris. Typical herbs include Ambrosia spp. (especially the perennial, A. psilostachya), Asclepias spp. (tuberosa, viridis), Croton spp. (especially monanthogynus), Erigeron strigosus, Neptunia lutea, Plantago virginica and Symphyotrichum spp. (especially dumosum var. subulifolium). Several aliens are usually present: white sweet clover (Melilotus alba) is locally frequent; large plots of sericea lespedeza (L. cuneata) have been established in some areas; and patches of Johnson grass (Sorgum halepense) occur on deeper soils.

(f) Upland grassland on average soils; typical subxeric slopes [see species C4 in C&S ....c]. The most common graminoids are little bluestem (Schizachyrium scoparium), Indian grass (Sorghastrum nutans) and switch grass (Panicum virgatum). Sedges are often present, but most typical of transitions to swales and low thickets (see d, g and h). Frequent herbs include Asclepias spp. (tuberosa, viridis), Agalinis spp. (especially oligophylla?), Allium canadense var. mobilense, Arnoglossum plantagineum, Blephilia ciliata, Buchnera americana, Dalea spp. (especially purpurea), Desmanthus illinoensis, Eryngium yuccifolium, Eupatorium altissimum, Eurybia hemispherica, Liatris spp. (especially aspera), Physostegia angustifolia, Polytaenia nuttalii, Prenanthes aspera, Ratibida pinnata, Silphium spp. (laciniatum, terebinthinaceum), Solidago spp. (especially nitida and rigida), Symphyotrichum spp. (especially dumosum), and Valerianella radiata. Additional legumes are locally frequent, especially at upper levels: Chamaecrista fasciculata, Galactia regularis, Desmodium spp. (especially ciliare), Lespedeza spp. (especially virginica). Invasive aliens are mostly excluded, except in formerly plowed or planted areas.

(g1) Variants on deeper or damper soils; relatively mesic to slightly hydric [see species D4 in C&S ....c]. In areas with relatively little agricultural disturbance, remnants of taller grassland are usually dominated by big bluestem (Andropogon gerardii); other common prairie grasses are also frequent (switch grass, Indian grass, little bluestem) plus, especially in low brushy transitions (see h2), wild rye (Elymus glabriflorus). In more disturbed areas, especially on damper soils, the broomsedge Andropogon tenuispatheus is often dominant; also, Saccharum giganteum and Paspalum floridanum var. glabrum are locally common. Frequent herbs include Agalinis spp. (especially auriculata and purpurea), Allium canadense, Apocynum cannabinum, Asclepias tuberosa, Desmanthus illinoensis, Desmodium spp. (especially paniculatum and sessilifolium), Dracopsis amplexicaulis, Gaura longiflora, Helenium flexuosum, Helianthus resinosus, Liatris spp. (especially aspera and spicata), Monarda fistulosa, Ratibida pinnata, Physostegia angustifolia, Silphium spp. (especially terebinthinaceum), Solidago spp. (altissima, gigantea, rigida), Symphyotrichum spp. (especially novae-angliae) and Valerianella radiata. Aliens occur in some areas, especially where there has been agricultural disturbance. In old fields, Johnson grass (Sorghum halepense) is locally dominant and fescue (Festuca arundinacea) is also frequent.

(g2) Variants on the wettest soils; somewhat hydric or hydroxeric [see species D4 as well as H/HX in C&S ....c]. This vegetation of swales or slope bases is rather heterogeneous; it generally matches the "seepy inclusions" of NatureServe (2010; under CEGL 4664). These sites tend to be in rather small patches or narrow zones, and are often fragmented or disturbed by farming of various kinds. It is likely that gamma grass (Tripsacum dactyloides) was formerly common, but there are only scattered patches now. Other tall native grasses include scattered Saccharum giganteum and Paspalum spp. Instead, shorter graminoids are locally common, including Scirpus pendulus, Carex spp. (especially granularis), Cyperus spp. (especially echinatus), and Juncus spp. (especially biflorus and torreyi). Lythrum alatum var. lanceolatum is often dominant in mid-summer. Other characteristic herbs may include Agalinis spp. (heterophylla, purpurea), Eupatorium spp. (coelestinum, fistulosum, serotinum), Ptilimnium nuttalii, Solidago spp. (altissima, gigantea) and Symphyotrichum praealtum. Aliens are infrequent; none are abundant.

(h) Upland thickets, edges and associated transitions from grassland to woodland.

These are varied types of transition, including isolated shrubby patches within the grassland, broader shrubby zones at upper or lower levels, narrower edges along woodland borders, and relatively thin woods that have been burned or otherwise disturbed. The most abundant shrubby species on more calcareous soils is usually roughleaf dogwood (Cornus drummondii). Also common are chickasaw plum (Prunus angustifolia), which is locally dominant, especially at upper levels; redbud (Cercis canadensis), especially at lower levels; and supplejack (Berchemia scandens), which often takes hold in viney tangles, especially at lower levels. Other locally frequent woody species include osage-orange (Maclura pomifera), honey-locust (Gleditsia triacanthos), sugarberry (Celtis laevigata), ashes (Fraxinus americana and, on wetter sites, F. pennsylvanica), sumacs (Rhus glabra at lower levels, R. copallina at upper levels), blackberries (especially Rubus trivialis at upper levels). Bully-buckthorn (Bumelia lycioides) and hawthorns (Crataegus crus-galli and, at upper levels, C. engelmanii) are infrequent, but may have been much more common in the pre-Columbian landscape (Campbell and Seymour ....a). Red-cedar (Juniperus virginiana) is generally absent from recently burned, mowed or plowed areas, but tends to increase where such disturbance has waned. In ground vegetation, as well as climbing high, vines are locally frequent: greenbriars (especially Smilax bona-box on drier sites), grape vines (several species), peppervine (Ampelopsis arborea on damper sites), poisonivy (Toxicodendron radicans var. pubens) and trumpet-creeper (Campsis radicans). The Japanese honeysuckle (Lonicera japonica) is also locally abundant, being virtually the only alien woody species in this type.

Herbaceous composition is highly varied and differs greatly between upper levels and lower levels, as follows.

(h1) Transitions at upper levels [see species B2 and B3 in C&S ....c]. Graminoids are locally abundant, including Carex spp. (leavenworthii, texensis, cf. umbellata), Danthonia spicata, Dichanthelium spp. (especially acuminatum and allies), Panicum anceps var. rhizomatum, Scleria spp. (especially triglomerata). The more distinctive herbs include Desmodium spp. (especially glabellum), Helianthus spp. (especially hirsutus), Lespedeza spp. (especially frutescens), Ruellia cf. ciliosa, Salvia lyrata and Verbesina helianthoides.

(h2) Transitions at lower levels [see species D2 and D3 in C&S ....c]. Graminoids are locally abundant, including Carex spp. (blanda, cherokeensis, oxylepis), Chasmanthium latifolium, Elymus spp. (virginicus in shade, glabriflorus in open), Festuca spp. (subverticillata in shade, paradoxa in open), Sphenopholis spp. (intermedia in shade, obtusata in open). The more distinctive perennial herbs include Dasystoma macrophylla, Desmodium spp. (especially paniculatum), Erigeron philadelphicus, Helianthus resinosus, Rudbeckia sp. nov., Ruellia strepens, Packera spp. (obovata, glabella), Verbesina virginica, Vernonia gigantea and Viola cf. missouriensis. There are also a few winter annuals in the shade, including Chaerophyllum tainturei, Galium aparine and Myosotis macrosperma.

(h3) Wet thickets. Young thickets dominated by green ash are noted below under swampy woods (section n). With more analysis, it might be reasonable to provide separate descriptions.

(i) Red cedar woods, on drier uplands (i1) and on damper soils (i2). Apart from the dominance of red-cedar itself, there may be little general difference in associated species between these young woods and the thickets or "transitions" outlined in preceding paragraphs. But the ground cover below canopy of red cedar is sparse, except for abundant mosses (especially Thuidium delicatum (Hedw.) B.S.G.). Spleenwort (Asplenium platyneuron) is one of the few vascular species typical of this habitat at the Pulliam Prairie. A curious feature is the frequent persistence of the alien, sericea lespedeza (L. cuneata), into more open stands of red cedar, where it has perhaps invaded old fields sown with sericea some decades ago. The vegetation varies from uplands to lowlands. Some relatively infrequent species appear characteristic of more open stands on drier ground (i1): Agalinis gattingeri, Crotolaria sagittalis, Dichanthelium laxiflorum, Potentilla simplex, Spiranthes ovalis, Symphyotrichum laeve var. laeve. Lower woods (i2) are dense and extensive along some gullies, down into adjacent lowlands, and associated species are similar to the lower thickets outlined above (h2).

(j) Submesic mixed hardwoods. Woods of submesic streamheads, toe-slopes or terraces (j3) are treated here provisionally under "Lowlands" (type m) but in a broader regional analysis it might be reasonable to segregate some of these woods from true lowlands. More mesic woods below bluffs locally with Acer floridanum (j2), were outlined for the region by Campbell and Seymour (....a) but do not occur at the Pulliam Prairie.

Lowlands with Alluvial Soils. As noted by Campbell and Seymour (....a), distinction of lowland and uplands in the Black Belt is somewhat arbitrary.

(k) Lowland grassland. This is not clearly recognized at the Pulliam Prairie, but the wettest grassland variant outlined above (type h2) may have graded into true lowland prairie during pre-Columbian times. Gamagrass (Tripsacum dactyloides) is present locally in that variant, and used to be extensive on the adjacent lowlands before conversion to agriculture, when it was known as "chicken corn" by early settlers (S.D. Pulliam, pers. comm.). The species remains locally abundant along roadsides on lowlands within a few miles of the site.

(l) Lowland canebrakes and associated thickets. Cane (Arundinaria gigantea) was not found at the Pulliam Prairie, but it occurs on lowlands within a mile or two. Some areas of current soybean fields directly below the site might well have supported cane in the original vegetation.

(m) Riparian hardwoods; generally submesic [see species D1/D2 and E1/E2 in C&S ....c]. These woods occur at the Pulliam Prairie in narrow strips and small patches within riparian zones, excluding the more poorly drained (hydric) areas outlined below (type n). They have varied soils, hydrology and disturbance history, and only a provisional generalization is provided here. Larger trees are mostly 2-4 dm in diameter. The most common trees include sugarberry (Celtis laevigata), osage-orange (Maclura pomifera), chinquapin oak (Quercus muhlenbergii), and elms (especially Ulmus americana). Also scattered in some areas are ashes (Fraxinus americana, F. pennsylvanica), honey locusts, (Gleditsia triacanthos), black walnut (Juglans nigra), and pecans (Carya myristicaeformis, C. illinoenensis). Distinctive shrubby species are relatively infrequent: Amorpha croceolanata, Asimina triloba, Ilex decidua, Symphoricarpos orbiculatus, and Viburnum rufidulum. Elderberry (Sambucus canadensis) is remarkably absent here, and it appears uncommon to rare elsewhere in the northern Black Belt (USDA 2010b). However, there are several frequent, distinctive vines: Ampelopsis spp. (especially cordata), Berchemia scandens, Campsis radicans, Toxidodendron radicans var. pubens, Smilax spp. (especially hispida), and Vitis spp. (especially cinerea and vulpina). The alien privet (Ligustrum sinense) and Japanese honeysuckle (Lonicera japonica) are local problems. The low thickets and other transitions described above (types h2, h3, i2) intergrade with the more mature woods outlined here, and many of the characteristic herbs are shared. Some grasses are locally abundant, especially wild-rye (Elymus virginicus) and wild-oats (Chasmanthium latifolium). Other graminoids include Carex spp. (blanda, cherokeensis, oxylepis), Festuca subverticillata, Leersia virginica, Muhlenbergia spp. (sylvatica, cf. frondosa), and Poa autumnalis. Locally frequent herbs in deeper shade include Cryptotaenia canadensis, Galium spp. (aparine, circaezans), Packera obovata, Sanicula odorata, and Viola cf. missouriensis.

(n) Swampy woods with green ash (n1) or oaks (n2); subhydric to hydric [for n1 see species E1/E2 as well as H in C&S ....c]. The study area does not extend into the adjacent bottomlands of Sakatonchee Creek, but it does include small patches of swampy woods, rarely more than an acre in extent, on more poorly drained bottoms along ephemeral streams. There is much intermixing with the more widespread submesic riparian woods (type m) and thickets (types h2 and h3). Most woods are young but some larger trees are 3-5 dm in diameter. The most common tree is green ash (Fraxinus pennsylvanica); other common trees include sugarberry (Celtis laevigata) and elm (Ulmus americana). Sweetgum (Liquidambar styraciflua) is infrequent. There are also small groups of oaks (Quercus phellos, Q. nigra) or pecans (Carya illinoensis) associated with seasonally drier soils, and a segregate characterized by Q. nigra might be recognized along ephemeral streamheads (n2). Locally frequent graminoids include Carex spp. (aureolensis, cf. normalis, cf. socialis), Cinna arundinacea, Elymus virginicus, and Glyceria striata. Characteristic herbs include Trepocarpus aethusae, usually in more open thickets.

DISCUSSION

Vegetation of Pulliam Prairie: pattern, process and prospects for restoration. Our initial ecological overview has emphasized the general gradient from lowlands to uplands, which is relatively stable, and a more complex gradient from deeper shade to more open grassland (Fig. 4). The latter gradient must involve several dynamic processes and will deserve more detailed analysis. To some extent the most calcareous soils on slopes between flatter ridges and bottomlands may be inhospitable to woody plants and favorable for grassland. In addition to the shallow rooting depth of some calcareous soils, much is classed as vertisol, characterized by seasonal swelling and shrinking along with associated extremes of wetness and dryness. Even without fires, there would probably be some degree of grassy opening on the chalk in this region's woodland.

Various disturbance regimes have been overlayed on the Black Belt's edaphic patterns during recent centuries and millenia, probably lacking clear, consistent correlations with soil types (Campbell and Seymour ....a). Before settlement, it is likely that larger herbivores congregated around streamheads, seeps and exposures of bedrock, in order to drink, lick, play and bask. Fires, due to lightning or people, may have spread out from ignitions in more flammable vegetation on the chalk, especially where dense short grass was mixed with red cedars. With increased human ignitions during the past few thousand years, it is possible that fires often spread up from the lowlands with gamagrass, cane and corn-fields. Also, there may have been regular ignitions along nearby upland trails through blackjack oak, post oak and shortleaf pine. After settlement, burning was replaced by cropping on flatter ground, especially lowlands. Livestock eventually increased on less productive soils, especially after much marginal cropped land was abandoned ca. 1930-1950.

Although we have begun to document the flora and vegetation at this site, a deeper ecological analysis should pay more attention to historical details of land uses here. We have only initial general impressions from inspecting the site and from talking to a few people familiar with the locality. Some of the flatter parts appear to have been used for row crops about 50-100 years ago. There are a few strips of slightly raised or shelved ground that suggest limited terracing for plowed plots. But cattle and hogs may have become the main focus of farming here during ca. 1930-1960. That scenario would be consistent with regional trends (Gibson 1941).

During the past 20 years or so, the site has been used mostly for hunting, especially deer. The landowners have conducted regular burning over much of the site, often down into riparian zones. They have also plowed some plots for temporary wildlife-food crops, and ryegrass (Lolium perenne) has been sown in one upland field within the past few years. It is remarkable that native species often recover dominance in these plowed plots within a few years, except where there is dense sericea lespedeza (L. cuneata) or Johnson grass (Sorghum hapenense).

Those two alien plants now present serious problems. Johnson grass was widely promoted in the Black Belt for hay and forage ca. 1870-1900, when it became one of the most abundant grasses in the region (Harper 1913). It is likely that sericea lespedeza was sown in some drier fields and eroding parts of the site when the farming was largely abandoned here after the 1960s. The Johnson grass could be effectively reduced to a much lower level in a few years, by using grass-specific herbicides from a wick-applicator, with treatments in June-July when most native grasses are still relatively short. The sericea lespedeza can be eliminated using broadleaf herbicides, but most other forbs would also be removed. Fortunately, this alien is concentrated in some relatively degraded areas that total only about 10-15 acres [4-6 ha]. After treatment, these areas could be reseeded with forb species using sources on the site.

In addition to delving into local history of the land, it would be useful to develop a dynamic model of the vegetation that can be linked with experiments in restoration. Deeper study of the Pulliam Prairie, with integration of similar efforts across the region, could lead to predictive models that can guide management, especially for restoring degraded areas. About a third of the site can be considered significantly degraded, due to insufficient disturbance and resultant invasion of red cedar (especially on drier ground), green ash (on wetter ground) and other woody plants, or due to establishment of locally abundant aliens, especially sericea lespedeza and Johnson grass. Monitoring of changes at the site, with or without various management practices, will be invaluable. The Pulliam Prairie could also supply seed to initiate substantial efforts in restoration at other sites. The populations of dominant grasses and composites, in particular, could be easily harvested for seed.

Regional Significance of Pulliam Prairie. This paper and the accompanying floristic survey (Campbell and Seymour ....c) provide evidence that the Pulliam Prairie is among the most significant known remnants of native grassland in the Black Belt, at least within Mississippi. But a more comprehensive review of ecological data from the region would be useful for comparing sites. Such review could be developed as part of ecoregional planning that is focused on the Black Belt, or perhaps nested within a broader review of the whole Upper East Gulf Coastal Plain. In addition to incorporating historical information on orginal vegetation across the region, there should be more analysis of the potential for extending and linking sites like Pulliam Prairie with other potentially conserved areas in their neighborhoods.

Ideally, some larger conserved areas should combine lowlands, intervening slopes and upland ridges, with a wide range of geology, topography and soils. Our initial reconnaissance of the land around Pulliam Prairie indicates much potential for a broader long-term plan. Including some old pastures and hayfields with lower quality, it is likely that at least 1000 acres [400 ha] of native grassland remnants can be identified within 10-20+ square miles [25-50+ [km.sup.2]] along or near the low ridge that extends north from the Pulliam Prairie. This ridge extends into the eastern edge of Tombigbee National Forest, where prescribed burning has already begun in their best remnant. Whether all these areas have sufficient quality and feasibility for restoration remains to be seen.

The comprehensive survey of prairie remnants in Alabama's Black Belt by Schotz and Barbour (2009) deserves to be extended into Mississippi. They identified 15,509 acres [6,276 ha] of prairie, mostly in the "average" subxeric type outlined above (f). Their most extensive and significant sites--or site complexes--were as follows: (1) Bragg-Ridgefield, with ca. 1430 acres [580 ha] clustered in ca. 32 square miles [83 [km.sup.2]]; (2) Tilden-Carlowville, with 690+ acres [281 ha] in ca. 16+ square miles [40 [km.sup.2]]; (3) Old Bluffport, with ca. 602 acres [244 ha] in ca. 8 square miles [20 [km.sup.2]]; (4) Elm Bluff, with ca. 284 acres [115 ha] in ca. 3 square miles [8 [km.sup.2]]; (5) Jones Bluff-House Bluff, with ca. 281 acres [114 ha] in ca. 3 square miles [8 [km.sup.2]]; and (6) Pleasant Ridge, with ca. 210 acres [85 ha] in ca. 10 square miles [25 [km.sup.2]]. Other identified sites in Alabama had no more than ca. 75-125 acres [30-50 ha], clustered or scattered over ca. 1-10 square miles.

The Pulliam Prairie is comparable with some of these better sites in Alabama, based on its area, ecological range, and species diversity. Although degraded in some sections, the Pulliam Prairie extends onto relatively deep, damp soils with taller grassland dominated by big bluestem and associates. Schotz and Barbour (2009) noted big bluestem at only one of their sites (Pleasant Ridge), and did not list gamagrass at any site. While most of the sites in Alabama tend to be on relatively dry ridges and bluffs, remnants of native grassland within the northern arc of the Black Belt in Mississippi tend to cover more gentle topography, often with sites grading into old pastures and cropped fields on toe-slopes and lowlands. There should be more recognition of the former extension by native grassland onto these landtypes (Campbell and Seymour ....a).

The USDA Farm Service Agency has in place a $300 million "Conservation Reserve Enhancement Program" to help reverse the loss of the Longleaf Pine Ecosystem, which lies mostly on sandy soils south of the Black Belt (USDA FSA 2006). The Black Belt Ecosystem has suffered even more than the Longleaf Pine, and it has become virtually extirpated as a type of natural landscape. Yet much can be done, even in the short-term, to improve management for wildlife (such as deer and quail), enhance remnants of native vegetation, reseed native grassland, and restore riparian buffers for water-quality. The Black Belt is a vital component of natural heritage in Mississippi and Alabama, and there would be much ecological, social and historical benefit from a more concerted program to establishment a network of conserved areas.

ACKNOWLEDGEMENTS

This study would not have been possible without permission of the landowner, Dr. Joe Pulliam, and his father, Sam D. Pulliam. We are deeply grateful for the opportunity to explore the "lost world" of their Black Belt heritage. Also invaluable was the introduction to this site, and continual guidance, by Daniel Coggin of Wildlife Mississippi, as well as the fundamental support of Roundstone Native Seed, Inc. Several people have provided significant further assistance. Jeff Levy and Tres Seymour helped make the GIS maps. Other people provided information and advice in various matters, especially John Barone, Steve Brewer, Richard Brown, Charles Bryson, John Gruchy, JoVonn Hill, Brad Lieb, John Pascarella, Al Schotz, Jennifer Seltzer, and some anonymous reviewers. Above all, our wives (Saundra Campbell and Donna Seymour) put up with our frequent absences from domestic duties, and tried to believe that we were not just having fun.

LITERATURE CITED

Barone, J.A. 2005. Historical presence and distribution of prairies in the Black Belt of Mississippi and Alabama. Castanea 70: 170-183.

Barone, J.A., and J.G. Hill. 2007. Herbaceous flora of Blackland Prairie remnants in Mississippi and western Alabama. Castanea 72: 226-234.

Brown, R.L. 2003. Paleoenvironment and biogeography of the Mississippi Black Belt: evidence from insects. Pages 1-26 in Peacock and Shauwecker (see below).

Chapman, S.S., G.E. Griffith, J.M. Omernik, J.A. Comstock, M. C. Beiser and D. Johnson. 2004. Ecoregions of Mississippi (color poster with map, descriptive text, summary tables, and photographs). U.S. Geological Survey (map scale 1:1,000,000). Reston, Virginia.

Gibson, J.S. 1941. The Alabama Black Belt: its Geographic Status. Econ. Geogr. 17: 1-23.

Harper, R.M. 1913. A botanical cross-section of northern Mississippi, with notes on the influence of soil on vegetation. Bull. Torrey Bot. Club 40: 377-399.

Hill, J.G. 2004. A note on a continuing Black Belt prairie restoration effort at Noxubee National Wildlife Refuge. J. Mississippi Acad. Sci. 49: 225-226.

Hill, J.G., and J.L. Selzer. 2007. A note on additional plants found at Sixteenth Section (Osborn) Prairie. J. Mississippi Acad. Sci. 52: 295-297.

Kartesz, J.T. 1999. A synonymized checklist and atlas with biological attributes for the vascular flora of the United States, Canada, and Greenland. First Edition. In: Kartesz, J.T., and C.A. Meacham. Synthesis of the North American flora, version 1.0. North Carolina Botanical Garden, Chapel Hill, NC.

MARIS [Mississippi Automated Resource Information System]. 2009. County 10-meter digital elevation model [available at http://www.maris.state.ms.us].

McCook, L.M., and J. Kartesz. 2010. A preliminary checklist of the plants of Mississippi [first posted in 2000 at website: http://herbarium.olemiss.edu/ checklist.html]. Department of Biology, University of Mississippi, Oxford.

Moore, W.H. (ed.) 1985. Geologic Map of Mississippi. Originally compiled in 1969 by A.R. Bicker, and revised with data submitted by Dr. E. E. Russell. Mercury Maps Inc., Jackson, Mississippi.

NaturalVue. 2000. North America--South [saved at resolution of 150 m]. Derived from GeoCover Ortho Program of NASA; available from http://www.gaf.de/content/naturalvue 2000.

NatureServe. 2010. National Vegetation Classification. Details included as "Ecological Communities and Systems" within website of NatureServe (2010a). [Explanation of full system and partnership included at: http:// www.natureserve.org/library/ vol1.pdf; see also, http://biology.usgs.gov/npsveg/ nvcs.html].

Peacock, E., and T. Schauwecker (eds). 2003. Blackland Prairies of the Gulf Coastal Plain: Nature, Culture, and Sustainability. University of Alabama Press, Tuscaloosa. 348 pp.

Schotz, A., and M. Barbour. 2009. Ecological Assessment and Terrestrial Vertebrate Surveys for Black Belt Prairies in Alabama. Alabama Department of Conservation and Natural Resources, Montgomery, Alabama.

U.S.D.A. Farm Service Agency [FSA]. 2006 [October]. Fact Sheet. Conservation Reserve Program. Longleaf Pine Initiative. 10 pages [http://www.fsa.usda.gov/ Internet/ FSAFile/crplongleaf06.pdf; this also became an "Enhancement Program" with more diffuse documentation.]

U.S.D.A. Natural Resources Conservation Service [NRCS]. 2010a. Official Soil Series Descriptions (OSD) with series mapping capabilities [http:// soils.usda.gov/technical/ classification/osd/index.html].

U.S.D.A. Natural Resources Conservation Service [NRCS]. 2010b. Soil Surveys for Counties of the U.S.A. [http:// websoilsurvey.nrcs.usda.gov/app/ WebSoilSurvey.aspx].

U.S.D.A. Natural Resources Conservation Service [NRCS]. 2010c. PLANTS Database [http://plants.usda.gov]. National Plant Data Center, Baton Rouge, Louisiana.

Wiygul, S., K. Krans, R. Brown, and V. Maddox. 2003. Restoration of a prairie remnant in the Black Belt of Mississippi. Pages 254-261 in Peacock and Schauwecker (see above).

J.J.N. Campbell (1) * and W.R. Seymour, Jr. (2)

(1) Bluegrass Woodland Restoration Center, 3525 Willowood Road, Lexington, Kentucky 40517

(2) Roundstone Native Seed, 9764 Raider Hollow Road, Upton, Kentucky 42784-9216

Corresponding Author: julian.campbell@insightbb.com
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Author:Campbell, J.J.N.; Seymour, W.R., Jr.
Publication:Journal of the Mississippi Academy of Sciences
Article Type:Report
Geographic Code:1U6MS
Date:Oct 1, 2011
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