ORIGINS OF THE FLORA

The Ketona Glades and their environs provide suitable habitat for more than 60 vascular or nonvascular plant taxa of conservation concern. This is an extraordinary number for a habitat that totals roughly only 100 hectares, marking these areas as one of the most significant reservoirs of botanical diversity in the eastern United States. While all of the factors responsible for the development of the unique Ketona Glade ecosystem may never be fully understood, especially the full phylogenetic histories of the strict endemics, enough is known to suggest some hypotheses.

It seems certain that edaphic factors, especially soil chemistry, have played a major role in the development of the characteristic Ketona Glade communities. In addition to the four samples of topsoil indicated previously as having been submitted to the state Soil Testing Laboratory of Georgia, we gathered samples from several other rock outcrop communities and submitted them also, for comparison. The results are summarized in Table 3. When compared to other outcrop soils, that of Ketona Glades is seen to be extremely high in the element magnesium. The Ketona Formation is, as stated earlier, an exceptionally pure dolomite, comparatively free of the usual siliceous impurities that serve to dilute the levels of magnesium and calcium in a mineral otherwise composed almost entirely of CaCO₃ and MgCO₃, in a ratio of about 55:45 (Rheams 1992).

Of the soils sampled, only those derived from serpentine and diabase have comparable levels of magnesium. They differ critically in being acidic, while those formed from Ketona Dolomite have a mildly alkaline pH. The former also differ in containing much lower levels of calcium and much higher levels of heavy metals. Outcrop soils that have a pH greater than 7, in common with Ketona Dolomite soil, have lower levels of magnesium. In short, the Ketona Glades have a distinctive soil chemistry, in particular a combination of very high magnesium levels and mildly basic soil reaction. In addition, the shallowness of soil and high insolation values make the glades drought-prone in the extreme. The peculiar soil chemistry and low organic matter content of the Ketona Glades, in combination with extremely low soil moisture levels in summer, provide conditions that are hostile to ordinarily adapted plants. Such a habitat strongly selects for specialized adaptations, promoting endemism or, at the very least, ecotypic differentiation. And because the Ketona Formation is restricted to central Alabama, and outcrops of other rocks as high in magnesium, such as serpentine, are unknown in Alabama, at least as pavements (Adams et al. 1926), this community is isolated both geographically and edaphically. Our choice of the term "Ketona Glades" to refer to these places is due to our conviction that the chemical composition of the bedrock is chiefly responsible for the unusual plant assemblage.

Earlier, under "Biological Communities," we listed a number of characteristic plants of limestone glades that are absent or essentially so from the Ketona Glades. Some of these taxa may be unable to tolerate the presumably higher summer temperatures of glades at the latitude of Bibb County, while the propagules of others may simply have failed to reach the Ketona Glades. However, the distributions of several characteristic species of limestone glades (e.g., Heliotropium tenellum, Hypericum sphaerocarpum, Ratibida pinnata, and Verbena simplex) appear to skip over the Ketona Glades only to recur south of them, on chalk glades of the Black Belt region. This lends support to the theory that the absence of some limestone glade taxa from the Ketona Glades may be due to an inability to thrive in soil with such a high concentration of magnesium.

It is conceivable that some of the Ketona Glade endemics, such as the new Dalea or Liatris, may once have been more widespread, with a refugium in that portion of the Ridge and Valley that would likely have experienced the mildest climate during the Pleistocene. If so, it would seem that they were unable to compete within the new communities that developed over most of their former range as the climate subsequently moderated, and thus were unable to reclaim their former territory. Some elements of the flora seem clearly to represent relictual biota, such as Paronychia virginica, Silene regia, and Solanum pumilum, while others, such as Astrolepis integerrima and possibly Liatris cylindracea, appear to represent long-distance immigration.

Perhaps Ketona Glade communities once occurred outside of Bibb County, or possibly still exist and remain to be discovered. The Ketona Formation is not restricted to Bibb County, but is mapped (Szabo et al. 1988) as extending into Blount, Chilton, Shelby and, most extensively of all, Jefferson County. The formation takes its name from a place in the latter county, and Ketona is home to several quarries in the formation. The exceptional purity of the formation has made it valuable as a "flux" stone in the steelmaking process (Rheams 1992), and its abundance, combined with ready access to water and to deposits of iron ore and coal, is responsible for making Birmingham, the county seat of Jefferson County, a major steelmaking center. Our efforts to find any glades within the extensive areas mapped in Jefferson County as Ketona Dolomite were, with one exception,⁽¹⁰⁾ unsuccessful, for much of the area has been quarried, and most of the rest is occupied by commercial or residential development. Their comparative remoteness from Birmingham is probably all that has saved the Bibb County glades from a similar fate.

In summary, the existence of the extraordinary assemblage of plant taxa on the Ketona Glades of Bibb County, Alabama is best explained as resulting from a combination of strong selection for edaphic specialization, geographic isolation, freedom for many millennia from drastic climate change (fostering relictualism), chance long-range dispersal events, and placement within a rural setting where human impacts to the landscape have been comparatively moderate.

CONSERVATION STATUS

Over the several years since the discovery of the Ketona Glades, The Nature Conservancy of Alabama, U.S. Alliance Coosa Pines Corporation, and the Bibb County Commission have worked together cooperatively to permanently protect about 120 hectares of glades and surrounding hardwood and longleaf pine forest. Currently, many of the largest and highest quality glades are protected within the Conservancy's Bibb County Glades Preserve. The preserve also protects more than a mile of riverine and riparian habitat along the Little Cahaba River, which harbors many imperiled and narrowly distributed aquatic species, including rare snails, mussels, fishes, caddisflies and the spectacular shoals spider lily, Hymenocallis coronaria (Chris Oberholster, The Nature Conservancy of Alabama, pers. comm. 2000).

CATALOG OF VASCULAR FLORA OF KETONA GLADES AND THEIR ENVIRONS

The following is an annotated list of taxa that we recorded from one or more Ketona Glades or their immediate vicinity. Except where indicated, nomenclature is that accepted at the time of writing by the PLANTS Database (USDA 2000). The names of taxa designated (at the time of writing or at any time during the study) to be of conservation concern by the Alabama Natural Heritage Program are preceded by an asterisk (*); names of taxa considered to be exogenous additions to the flora are indicated by a plus sign (+). For the most part, only uncommon or phytogeographically noteworthy taxa were recorded from surrounding areas, beyond glade-forest ecotones; these names are placed within curved brackets, { }. Synonyms that have been used in recent decades are provided within square brackets, [ ]. Finally, for those taxa of which we made specimens, we append the collection number of a voucher deposited (except where a different acronym is provided) at the herbarium of the University of Alabama (UNA).

PTERIDOPHYTES

ASPLENIACEAE

Asplenium platyneuron (L.) B.S.P.

Asplenium resiliens Kunze

DENNSTAEDTIACEAE

Pteridium aquilinum (L.) Kuhn

ISOËTACEAE

*Isoëtes butleri Engelm., A. and S. 7552

OPHIOGLOSSACEAE

*Ophioglossum engelmannii Prantl, A. and S. 7646

POLYPODIACEAE

Pleopeltis polypodioides (L.) Andrews & Windham var. michauxiana (Weatherby) Andrews & Windham [Polypodium polypodioides (L.) Watt var. michauxianum Weatherby]

PTERIDACEAE

{Adiantum capillus-veneris L.}

*Astrolepis integerrima (Hook.) Benham & Windham [Notholaena integerrima (Hook.) Hevly, Cheilanthes integerrima (Hook.) Mickel], A. 6695

*Cheilanthes alabamensis (Buckl.) Kunze, A. 12574

Cheilanthes lanosa (Michx.) D. C. Eat.

Pellaea atropurpurea (L.) Link

GYMNOSPERMS

CUPRESSACEAE

Juniperus virginiana L. var. virginiana

PINACEAE

Pinus echinata P. Mill.

Pinus palustris P. Mill.

Pinus taeda L.

ANGIOSPERMS

ACANTHACEAE

Ruellia humilis Nutt., A. and S. 7787

Ruellia strepens L.

ACERACEAE

Acer leucoderme Small

AGAVACEAE

Manfreda virginica (L.) Salisb. ex Rose [Agave virginica L., Polianthes virginica (L.) Shinners]

Yucca filamentosa L.

ANACARDIACEAE

Rhus aromatica Ait.

Rhus copallinum L.

Rhus glabra L.

Toxicodendron pubescens P. Mill. [T. toxicarium Gillis, Rhus toxicodendron L.]

Toxicodendron radicans (L.) Kuntze [Rhus radicans L.]

APIACEAE

{Eryngium integrifolium Walt.} (single locality), A. and S. 8112

Eryngium yuccifolium Michx. var. yuccifolium

Thaspium barbinode (Michx.) Nutt. var. chapmanii Coult. & Rose (variety not accepted in PLANTS [USDA 2000]), A. and S. 7558

Thaspium trifoliatum (L.) Gray var. aureum (L.) Britt.

Zizia aptera (Gray) Fern.

APOCYNACEAE

Amsonia ciliata Walt. var. tenuifolia (Raf.) Woods. [A. ciliata Walt. var. filifolia Woods.], A. 10566

Amsonia tabernaemontana Walt.

Apocynum cannabinum L.

AQUIFOLIACEAE

{Ilex ambigua (Michx.) Torr.}

ARALIACEAE

*{Panax quinquefolius L.}

ARECACEAE

*{Rhapidophyllum hystrix (Pursh) H. Wendl. & Drude ex Drude}, J. Allison and C. Oberholster 7057 (VDB)

Sabal minor (Jacq.) Pers.

ASCLEPIADACEAE

Asclepias tuberosa L.

{Asclepias variegata L.}

Asclepias verticillata L.

Asclepias viridiflora Raf.

Asclepias viridis Walt.

Matelea gonocarpos (Walt.) Shinners, A. 10898

ASTERACEAE

Acmella oppositifolia (Lam.) R. K. Jansen var. repens (Walt.) R. K. Jansen [Spilanthes americana (Mutis) Hieron.] (single locality)

Ambrosia artemisiifolia L.

Brickellia eupatorioides (L.) Shinners [Kuhnia eupatorioides L.]

{Carphephorus odoratissimus (J. F. Gmel.) Herbert} [Trilisa odoratissima (J. F. Gmel.) Cass.], A. and S. 7276 (VDB)

Chrysopsis mariana (L.) Ell. [Heterotheca mariana (L.) Shinners], A. 12575

{Coreopsis auriculata L.}

{Coreopsis grandiflora Hogg ex Sweet var. grandiflora} A. 11330

*Coreopsis grandiflora Hogg ex Sweet var. inclinata J. Allison, A. and S. 7223

Coreopsis grandiflora Hogg ex Sweet var. inclinata J. Allison V C. pubescens Ell., A. and S. 6719

Coreopsis major Walt.

{Coreopsis pubescens Ell.} (seemingly only introgressants in vicinity of glades), A. 11933

*Echinacea purpurea (L.) Moench, A. 6782

Erigeron annuus (L.) Pers.

Erigeron pulchellus Michx., A. and S. 7551

Erigeron strigosus Muhl. ex Willd. var. beyrichii (Fisch. & C. A. Mey.) Torr. & Gray ex Gray, A. 12089

*Erigeron strigosus Muhl. ex Willd. var. dolomiticola J. Allison, A. and S. 7947

Eupatorium album L. var. album, A. and S. 7881

Eupatorium rotundifolium L., A. 12446

Fleischmannia incarnata (Walt.) King & H. E. Robins. [Eupatorium incarnatum Walt.], A. 12573

Gaillardia aestivalis (Walt.) H. Rock var. aestivalis, A. 12568

Helenium autumnale L.

Helianthus divaricatus L.

Helianthus hirsutus Raf., A. et al. 7839

Helianthus microcephalus Torr. & Gray, A. and S. 8011 (VDB)

Helianthus resinosus Small [H. tomentosus auct. non Michx.]

*Helianthus smithii Heiser, A. and S. 8478

Heliopsis helianthoides (L.) Sweet var. gracilis (Nutt.) Gandhi & Thomas [H. gracilis Nutt.], A. and S. 7116

Ionactis linariifolius (L.) Greene [Aster linariifolius L.]

*{Jamesianthus alabamensis Blake & Sherff}, A. and S. 7128

*Liatris cylindracea Michx., A. and S. 6877

Liatris Vfreemaniana J. Allison [L. cylindracea Michx. V L. oligocephala J. Allison], A. and S. 7237

Liatris Vmacdanieliana J. Allison [L. cylindracea Michx. V L. squarrosa (L.) Michx.], A. and S. 8533

*Liatris oligocephala J. Allison, A. and S. 7119

Liatris pilosa (Ait.) Willd. [L. graminifolia Willd.]

Liatris squarrosa (L.) Michx. var. squarrosa, A. and S. 6892

*Marshallia mohrii Beadle & F. E. Boynt., A. and S. 7720

*{Marshallia trinervia (Walt.) Trel.}, A. 6656

Oligoneuron rigidum (L.) Small [Solidago rigida L.]

Packera anonyma (Wood) W. A. Weber & A. Löve [Senecio anonymus Wood, S. smallii Britt.]

Pseudognaphalium obtusifolium (L.) Hilliard & Burtt [Gnaphalium obtusifolium L.]

Rudbeckia fulgida Ait. var. fulgida, A. and S. 7030

{Rudbeckia fulgida Ait. var. umbrosa (C. L. Boynt. & Beadle) Cronq.}

*Rudbeckia triloba L. var. pinnatiloba Torr. & Gray, A. 11023

Sericocarpus tortifolius (Michx.) Nees [Aster tortifolius Michx., A. bifoliatus (Walt.) Ahles], A. and S. 7965 (VDB)

Silphium asteriscus L. var. angustatum Gray

{Silphium asteriscus L. var. angustatum Gray V S. trifoliatum L. var. latifolium Gray}

Silphium asteriscus L. var. asteriscus V S. glutinosum J. Allison, A. and S. 8474

Silphium compositum Michx., A. and S. 6712

Silphium compositum Michx. V S. laciniatum L. (two localities, one a glade-ecotone, the other a logging road near a glade), A. 12508

*Silphium glutinosum J. Allison, A. and S. 8107

Silphium glutinosum J. Allison V S. trifoliatum L. var. latifolium Gray, A. and S. 7301

Silphium laciniatum L., A. and S. 6714

*Silphium trifoliatum L. var. latifolium Gray

Smallanthus uvedalius (L.) Mackenzie ex Small [Polymnia uvedalia (L.) L.]

Solidago nemoralis Ait.

Solidago odora Ait.

Solidago rugosa P. Mill.

Solidago ulmifolia Muhl. ex Willd.

Symphyotrichum concolor (L.) Nesom [Aster concolor L.], A. 12565

*Symphyotrichum georgianum (Alexander) Nesom [Aster georgianus Alexander, A. patens Ait. var. georgianus (Alexander) Cronq.], A. and S. 7318

*Symphyotrichum laeve (L.) A. & D. Löve var. concinnum (Willd.) Nesom [Aster concinnus Willd], A. 6757

{Symphyotrichum. oblongifolium (Nutt.) Nesom} [Aster oblongifolius Nutt.], A. 12580

Symphyotrichum patens (Ait.) Nesom [Aster patens Ait.], A. 12566

Symphyotrichum pilosum (Willd.) Nesom [Aster pilosus Willd.], A. and S. 7316

Symphyotrichum shortii (Lindl.) Nesom [Aster shortii Lindl.]

Tetragonotheca helianthoides L.

Verbesina occidentalis (L.) Walt.

Verbesina virginica L.

BETULACEAE

Ostrya virginiana (P. Mill.) K. Koch

BIGNONIACEAE

Bignonia capreolata L. [Anisostichus capreolata (L.) Bureau]

BORAGINACEAE

Lithospermum canescens (Michx.) Lehm., A. 10464

Lithospermum tuberosum Rugel ex DC.

*Onosmodium decipiens J. Allison, A. and S. 8235

Onosmodium decipiens J. Allison V O. virginianum (L.) A. DC. (single locality), A. and S. 8231

Onosmodium virginianum (L.) A. DC., A. and S. 8232

BRASSICACEAE

+Arabidopsis thaliana (L.) Heynh.

Arabis canadensis L.

*Arabis georgiana Harper, A. et al. 6681

+Cardamine hirsuta L.

*Leavenworthia exigua Rollins var. lutea Rollins, A. and S. 7509

*Leavenworthia uniflora (Michx.) Britt., A. and S. 8156

BUDDLEJACEAE

Polypremum procumbens L.

CACTACEAE

Opuntia humifusa Raf. var. humifusa [O. compressa auct. non J. F. Macbr.]

CAMPANULACEAE

Campanulastrum americanum (L.) Small [Campanula americana L.]

Lobelia spicata Lam.

CAPRIFOLIACEAE

+Lonicera japonica Thunb.

Lonicera sempervirens L.

{Triosteum aurantiacum Bickn.} (single locality)

Viburnum rufidulum Raf.

CARYOPHYLLACEAE

{Arenaria lanuginosa (Michx.) Rohrb.}

Minuartia patula (Michx.) Mattf. [Arenaria patula Michx.]

*Paronychia virginica Spreng., A. and S. 6891

*{Silene caroliniana Walt. ssp. wherryi (Small) Clausen}, A. and S. 7591

*Silene regia Sims, A. and S. 6886

CELASTRACEAE

*{Euonymus atropurpureus Jacq.}, A. et al. 9109

CISTACEAE

Lechea mucronata Raf. [L. villosa Ell.], A. 12088

CLUSIACEAE

Hypericum densiflorum Pursh, A. et al. 9463

Hypericum frondosum Michx.

Hypericum hypericoides (L.) Crantz ssp. hypericoides [Ascyrum hypericoides L. var. hypericoides]

*{Hypericum nudiflorum Michx.}

{Hypericum punctatum Lam.}

COMMELINACEAE

Commelina erecta L.

{Tradescantia hirsuticaulis Small}

CONVOLVULACEAE

Calystegia catesbiana, s. str., A. and S. 7562 (identification tentative)

Dichondra carolinensis Michx. [D. repens J. R. Forst. var. carolinensis (Michx.) Choisy]

Ipomoea pandurata (L.) G. F. W. Mey.

Stylisma humistrata (Walt.) Chapman, A. and S. 6722

CORNACEAE

Cornus florida L.

CRASSULACEAE

*{Sedum nevii Gray}

CYPERACEAE

Carex cherokeensis Schwein.

Carex crawei Dewey, A. and S. 7576

*Carex eburnea Boott, A. and S. 6717

Carex laxiflora Lam., A. and S. 7582

Carex striatula Michx., A. and S. 7577

Fimbristylis puberula (Michx.) Vahl, A. 6747

*Rhynchospora capillacea Torr. (single locality), A. et al. 6676

*Rhynchospora colorata (L.) H. Pfeiffer [Dichromena colorata (L.) A. S. Hitchc.], A. 6691

Rhynchospora divergens Chapman ex M. A. Curtis, A. 6793

Rhynchospora globularis (Chapman) Small var. globularis, A. and S. 7732

Rhynchospora globularis (Chapman) Small var. pinetorum (Britt. & Small ex Small) Gale, A. and S. 7147 (VDB)

*Rhynchospora thornei Kral, A. et al. 6754

Scleria oligantha Michx.

Scleria verticillata Muhl. ex Willd. (single locality), A. and S. 7964

EBENACEAE

Diospyros virginiana L.

EUPHORBIACEAE

Cnidoscolus stimulosus (Michx.) Engelm. & Gray

*Croton alabamensis E. A. Sm. ex Chapman var. alabamensis, A. 6777

Croton capitatus Michx.

Croton monanthogynus Michx.

Euphorbia commutata Engelm.

Euphorbia corollata L., A. and S. 7976

{Euphorbia pubentissima Michx.}

*Leptopus phyllanthoides (Nutt.) G. L. Webster [Andrachne phyllanthoides (Nutt.) Coult.], A. 6657

{Tragia cordata Michx.}

Tragia urticifolia Michx.

FABACEAE

Amorpha fruticosa L., A. et al. 6869

*{Baptisia australis (L.) R. Br. ex Ait. f. var. australis}, A. and S. 7724

Centrosema virginianum (L.) Benth.

Cercis canadensis L.

Chamaecrista fasciculata (Michx.) Greene [Cassia fasciculata Michx.]

Crotalaria sagittalis L.

*Dalea cahaba J. Allison, A. and S. 6876

Desmanthus illinoënsis (Michx.) MacM. ex B. L. Robins. & Fern.

Galactia volubilis (L.) Britt.

Orbexilum pedunculatum (P. Mill.) Rydb. var. pedunculatum [Psoralea psoralioides (Walt.) Cory var. eglandulosa (Ell.) Freeman]

*Pediomelum subacaule (Torr. & Gray) Rydb. [Psoralea subacaulis Torr. & Gray] (single locality), A. et al. 8177

{Rhynchosia reniformis DC.}

Rhynchosia tomentosa (L.) Hook. & Arn.

Stylosanthes biflora (L.) B.S.P.

Tephrosia spicata (Walt.) Torr. & Gray

Tephrosia virginiana (L.) Pers.

+Trifolium campestre Schreb.

FAGACEAE

{Quercus alba L.}

Quercus austrina Small, A. and S. 6735

Quercus coccinea Muenchh.

Quercus hemisphaerica Bartr. ex Willd.

Quercus incana Bartr., A. and S. 7586

Quercus margarettiae Ashe ex Small, A. and S. 7063

Quercus marilandica Muenchh.

Quercus muehlenbergii Engelm.

Quercus nigra L.

Quercus shumardii Buckl.

Quercus stellata Wangenh.

GENTIANACEAE

*{Gentiana saponaria L.}

*{Gentiana villosa L.}, A. and S. 7496

{Obolaria virginica L.}

Sabatia angularis (L.) Pursh

HYDRANGEACEAE

Hydrangea quercifolia Bartr.

IRIDACEAE

Hypoxis hirsuta (L.) Coville, A. et al. 6753

Iris verna L.

Sisyrinchium atlanticum Bickn., A. and S. 7589

JUGLANDACEAE

Carya glabra (P. Mill.) Sweet

Carya pallida (Ashe) Engl. & Graebn., A. and S. 7550

JUNCACEAE

Juncus effusus L.

Juncus filipendulus Buckl., A. and S. 6709

Luzula bulbosa (Wood) Smyth & Smyth

LAMIACEAE

Blephilia ciliata (L.) Benth.

{Collinsonia tuberosa Michx.}

Isanthus brachiatus (L.) B.S.P. [Trichostema brachiatum L.]

+Lamium amplexicaule L.

{Monarda fistulosa L}

{Monarda punctata L.}

(+?)Prunella vulgaris L.

Pycnanthemum pycnanthemoides (Leavenworth) Fern.

Salvia azurea Michx. ex Lam.

Salvia lyrata L.

Salvia urticifolia L.

*Scutellaria alabamensis Alexander, A. and S. 7737

Scutellaria incana Biehler var. punctata (Chapman) C. Mohr, A. 12505

Scutellaria parvula Michx. var. parvula, A. and S. 7648

LAURACEAE

Persea palustris (Raf.) Sarg.

LILIACEAE

Aletris farinosa L.

Allium canadense L. var. canadense

Allium canadense L. var. mobilense (Regel) Ownbey [A. mutabile Michx.], A. and S. 7564

{Camassia scilloides (Raf.) Cory}

*{Melanthium latifolium Desr.} [M. hybridum Walt.], A. and S. 7794 (VDB)

*{Melanthium woodii (J. W. Robbins ex Wood) Bodkin} [Veratrum woodii J. W. Robbins ex Wood] (single locality), A. and S. 7854

{Melanthium virginicum L.} (single locality)

Nothoscordum bivalve (L.) Britt. [Allium bivalve (L.) Kuntze]

{Polygonatum biflorum (Walt.) Ell.}

*Schoenolirion croceum (Michx.) Wood, A. and S. 7588

Trillium cuneatum Raf., A. et al. 8775

Trillium stamineum Harbison, A. et al. 8776

{Uvularia perfoliata L.}

LINACEAE

Linum sulcatum Riddell var. sulcatum, A. et al. 6865

LOGANIACEAE

Gelsemium sempervirens St.-Hil.

Mitreola sessilifolia (J. F. Gmel.) G. Don [Cynoctonum sessilifolium J. F. Gmel.]

*Spigelia gentianoides Chapman in A. DC. var. alabamensis K. Gould, A. 6688

Spigelia marilandica (L.) L., A. 6763

MALVACEAE

*Callirhoë alcaeoides (Michx.) Gray, A. 11025

*Sida elliottii Torr. & Gray, A. 8555

MENISPERMACEAE

Cocculus carolinus (L.) DC.

MYRICACEAE

Morella cerifera (L.) Small [Myrica cerifera L.]

NYCTAGINACEAE

*Mirabilis albida (Walt.) Heimerl, A. 6759

OLEACEAE

Chionanthus virginicus L.

Forestiera ligustrina (Michx.) Poir.

Fraxinus americana L.

Fraxinus quadrangulata Michx., A. and S. 7734

+Ligustrum sinense Lour.

ONAGRACEAE

Gaura filipes Spach, A. 12397 (OKL)

Ludwigia microcarpa Michx.

Oenothera biennis L.

Oenothera fruticosa L. ssp. glauca (Michx.) Straley [O. tetragona Roth]

+Oenothera speciosa Nutt. (adventive from farther west)

ORCHIDACEAE

Hexalectris spicata (Walt.) Barnh., A. and S. 7413

*Ponthieva racemosa (Walt.) C. Mohr, A. and S. 7960 (VDB)

Spiranthes lacera (Raf.) Raf. var. gracilis (Bigelow) Luer [S. gracilis (Bigelow) Beck var. gracilis]

*{Spiranthes lucida (H. H. Eat.) Ames} (single locality), A. and S. 8224-p

*Spiranthes magnicamporum Sheviak, A. and S. 7437

OROBANCHACEAE

*Orobanche uniflora L. (single locality)

OXALIDACEAE

Oxalis priceae Small ssp. priceae, A. et al. 6672

Oxalis violacea L.

PASSIFLORACEAE

Passiflora lutea L.

PLANTAGINACEAE

*{Plantago cordata Lam.}

+Plantago lanceolata L.

Plantago virginica L.

POACEAE

Andropogon gerardii Vitman, A. et al. 7837

Andropogon glomeratus (Walt.) B.S.P.

Andropogon gyrans Ashe var. gyrans [A. elliottii Chapman], A. et al. 8763

Andropogon virginicus L.

Anthaenantia villosa (Michx.) Beauv., A. et al. 9459

Aristida lanosa Muhl. ex Ell., A. and S. 7413

Aristida longespica Poir.

Aristida purpurascens Poir., J. Allison and C. Oberholster 7049

{Arundinaria gigantea (Walt.) Muhl.}

Bouteloua curtipendula (Michx.) Torr.

{Brachyelytrum erectum (Schreb. ex Spreng.) Beauv.}, J. Allison and T. McQuilkin 11014

Bromus pubescens Muhl. ex Willd. [B. purgans auct. non L.]

Chasmanthium latifolium (Michx.) Yates [Uniola latifolia Michx.], A. 12171

Chasmanthium sessiliflorum (Poir.) Yates [Uniola sessiliflora Poir.], A. 12173

Dichanthelium (A. S. Hitchc. & Chase) Gould spp. [Panicum L. spp.]

Eragrostis spectabilis (Pursh) Steud.

Gymnopogon ambiguus (Michx.) B.S.P.

Melica mutica Walt., A. 8174

Muhlenbergia capillaris (Lam.) Trin.

{Muhlenbergia sylvatica Torr. ex Gray}, A. 12581

Panicum anceps Michx.

Panicum capillare L.

Panicum flexile (Gattinger) Scribn.

Panicum virgatum L.

Paspalum floridanum Michx.

Piptochaetium avenaceum (L.) Parodi [Stipa avenacea L.]

+Poa annua L.

Schizachyrium scoparium (Michx.) Nash [Andropogon scoparius Michx.], A. and S. 8140

Sorghastrum nutans (L.) Nash

Sphenopholis filiformis (Chapman) Scribn.

{Sphenopholis obtusata (Michx.) Scribn.}, A. 8173

Sporobolus clandestinus (Biehler) A. S. Hitchc., A. and S. 7104

Sporobolus junceus (Beauv.) Kunth

Sporobolus vaginiflorus (Torr. ex Gray) Wood

Tridens flavus (L.) A. S. Hitchc. var. chapmanii (Small) Shinners, A. 12569

Tridens flavus (L.) A. S. Hitchc. var. flavus

POLEMONIACEAE

Ipomopsis rubra (L.) Wherry, A. et al. 6665

Phlox amoena Sims, A. 10563

{Phlox glaberrima L.}

{Phlox pilosa L.}

*{Phlox pulchra (Wherry) Wherry}, A. et al. 8343

POLYGALACEAE

*Polygala boykinii Nutt.

Polygala grandiflora Walt.

PRIMULACEAE

Dodecatheon meadia L.

Lysimachia ciliata L.

Lysimachia quadriflora Sims, A. 6783

RANUNCULACEAE

Anemone berlandieri Pritz. [A. heterophylla Nutt. ex Torr. & Gray], A. and S. 7502

{Aquilegia canadensis L.}

Clematis glaucophylla Small, A. 6692

Clematis viorna L., A. et al. 8345

Delphinium carolinianum Walt. ssp. carolinianum

*Enemion biternatum Raf. [Isopyrum biternatum (Raf.) Torr. & Gray], A. 6344

Thalictrum thalictroides (L.) Eames & Boivin [Anemonella thalictroides (L.) Spach]

Trautvetteria caroliniensis (Walt.) Vail (two localities, one a wet glade-ecotone, the other along a spring-fed stream near a glade)

RHAMNACEAE

Berchemia scandens (Hill) K. Koch

Ceanothus americanus L.

Frangula caroliniana (Walt.) Gray [Rhamnus caroliniana Walt.]

ROSACEAE

+Aphanes microcarpa (Boiss. & Reut.) Rothm. [Alchemilla microcarpa Boiss. & Reut.]

Crataegus crus-galli L., A. 11022

Crataegus uniflora Muenchh., A. et al. 9452

*{Neviusia alabamensis Gray} (single locality), A. and S. 8481

Physocarpus opulifolius (L.) Maxim., A. and S. 6711

Rubus L. spp.

RUBIACEAE

Diodia teres Walt.

Hedyotis nigricans (Lam.) Fosb. [Houstonia nigricans (Lam.) Fern.], A. 6798

Houstonia caerulea L. [Hedyotis caerulea (L.) Hook.]

Houstonia purpurea L. var. calycosa Gray [Hedyotis purpurea (L.) Torr. & Gray var. calycosa (Gray) Fosberg], A. and S. 7587

Houstonia pusilla Schoepf [Hedyotis crassifolia Raf.]

RUTACEAE

Ptelea trifoliata L.

*{Zanthoxylum americanum P. Mill.}, A. and S. 7727

SALICACEAE

Salix caroliniana Michx., A. 11931

SAPOTACEAE

Sideroxylon lycioides L. [Bumelia lycioides (L.) Pers.]

SAXIFRAGACEAE

*{Parnassia grandifolia DC.} (single locality), A. and S. 7436

SCHISANDRACEAE

*{Schisandra glabra (Brickell) Rehder} [S. coccinea Michx] (Brickell's basionym misattributed to Bicknell in PLANTS [USDA 2000]), J. Allison and C. Oberholster 7044

SCROPHULARIACEAE

Agalinis purpurea (L.) Pennell, A. and S. 7230

Agalinis tenuifolia (Vahl) Raf., A. and S. 7029

Aureolaria flava (L.) Farw.

Aureolaria pectinata (Nutt.) Pennell

Aureolaria virginica (L.) Pennell

Buchnera americana L., s. str.

*Castilleja kraliana J. Allison, A. and S. 7575

Leucospora multifida (Michx.) Nutt., A. et al. 8344

Mecardonia acuminata (Walt.) Small var. acuminata

Pedicularis canadensis L.

*Penstemon tenuiflorus Pennell, A. 10561

Seymeria cassioides (J. F. Gmel.) Blake

Seymeria pectinata Pursh

+Veronica arvensis L.

*Veronicastrum virginicum (L.) Farw. (single locality), A. 12504

SMILACACEAE

Smilax bona-nox L.

Smilax glauca Walt.

Smilax smallii Morong, A. 12577

SOLANACEAE

Physalis heterophylla Nees, A. and S. 7580

Solanum carolinense L. var. carolinense A. and S. 7590

*Solanum pumilum Dunal [S. carolinense L. var. hirsutum (Nutt.) Gray, accepted by PLANTS (USDA 2000)], A. and S. 7644

STYRACACEAE

{Styrax grandifolius Ait.}

ULMACEAE

Celtis tenuifolia Nutt. [C. georgiana Small], A. and S. 7647

Ulmus alata Michx.

{Ulmus rubra Muhl.}

VALERIANACEAE

Valerianella radiata (L.) Dufr.

VERBENACEAE

Callicarpa americana L.

Stylodon carneus (Medik.) Moldenke [Verbena carnea Medik.], A. and S. 7657 (VDB)

Verbena simplex Lehm. (single locality), A. 12395

VIOLACEAE

{Hybanthus concolor (T. F. Forst.) Spreng.}, A. 8175

Viola bicolor Pursh [V. rafinesquii Greene]

Viola pedata L.

Viola walteri House, A. and S. 7414

VISCACEAE

Phoradendron leucarpum (Raf.) Reveal & M.C. Johnston [P. flavescens Nutt. ex Engelm., P. serotinum (Raf.) M.C. Johnston]

XYRIDACEAE

*Xyris tennesseensis Kral, A. and S. 7067  

ACKNOWLEDGMENTS

Partial funding of this project was provided by the United States Fish and Wildlife Service (Grant No. 1448-0004-93-926). Particular thanks go to Vicky Holifield for preparing all of the line drawings. Special thanks go also to James Matthews of the University of North Carolina at Charlotte for making chromosome counts for us of Silphium, and to Richard LaFleur of the University of Georgia for critiquing the Latin diagnoses. We are also grateful to the curators and other staff and researchers at herbaria consulted during this study, including the late John Freeman at AUA; Wilbur Duncan and the late Michael Moore at GA; Emily Wood at GH; David Whetstone and Dan Spaulding at JSU; the late William D'Arcy at MO; Pat Holmgren, Noel Holmgren, and the late Rupert Barneby at NY; Ernest Schuyler at PH; Robert Haynes and Steve Ginzbarg at UNA; David Lellinger at US; and Robert Kral at VDB. Thanks to them for valuable discussions as well as to W. Edward Osborne and Alan Weakley, and onsite (and elsewhere) with James Affolter, Angus Gholson, the late Robert Godfrey, Scott Gunn (who found Jamesianthus and Plantago cordata), Greg Krakow, Robert Kral, John MacDonald, Sidney McDaniel, Mincy Moffett, Cary Norquist, Chris Oberholster, Tom Patrick, Milo Pyne (thanks also for directions to the population of Onosmodium molle ssp. hispidissimum closest to Alabama, in Marion County, Tennessee), Jim Rodgers, Debi Rodgers, Al Schotz, Paul Somers, Dan Spaulding, and David Whetstone. Additional thanks go to Mr. Gholson for personal guidance to occurrences of Spigelia gentianoides var. gentianoides. Mr. Krakow was also very helpful with various word processing challenges, especially in layout of the tables. The manuscript was improved by suggestions from Zack Murrell, Dan Spaulding, and, especially, Tom Patrick. Finally, much thanks to Charles Williams of Limestone Park, outfitter for our many canoeing expeditions, for logistical help and for sharing his knowledge of the people and places of Bibb County.

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