State Records
Next to the newly described taxa, the most significant discoveries of the present study were seven species never before reported from Alabama. Two of these are characteristic elements of the Ketona Glades, a Paronychia found in open, exposed situations, and a Solanum, usually found in partial shade. The remaining five species are either too uncommon on the glades to be considered characteristic, or are plants of habitats peripheral to Ketona Glades.
Astrolepis integerrima (Hook.) Benham & Windham
On June 1, 1992, after the rest of the members of the original canoeing expedition had returned home, Allison visited by car and foot two glades that had been spotted from the canoes but not visited then due to time constraints. The prime new find there was of a fern, at first mistaken, due to its brown, inrolled leaves in drought response, for Cheilanthes tomentosa Link. Upon bending closer to make a collection (A. 6695, UNA, VDB), it was clear that it was not that species at all, but seemingly a smaller version of a fern Allison had found in 1980, disjunct in Meriwether County, Georgia, Astrolepis sinuata (Lag.) Benham & Windham.
Upon returning home, Allison determined the fern to be Astrolepis integerrima (Hook.) Benham & Windham, better known at the time as Notholaena integerrima (Hook.) Hevly. This fern is disjunct on the order of 1000 km to the east of populations in Texas. On the Ketona Glades it is a scarce plant of scanty soil accumulations on rock ledges and boulders. We were only able to find two small populations of this species, but possibly other populations occur on one or more of the comparatively inaccessible rock bluffs along the Little Cahaba River.
Baptisia australis (L.) R. Br. ex Ait. f. var. australis
During our several canoe trips on the Little Cahaba River in the summer of 1992, we noted a rather glabrous Baptisia on gravelly bars and islands and on the river banks. As it was past flowering before our first expedition, it could not be conclusively identified. The following spring, after he took part in a canoe trip sponsored by The Nature Conservancy of Alabama, Chris Oberholster reported to us (pers. comm. 1993) that the flowers were blue, confirming our expectation that the plant was Baptisia australis (L.) R. Br. ex Ait. f. var. australis, known from the adjoining states of Georgia and Tennessee, but not from Alabama. We obtained a voucher on June 13, 1993 (A. and S. 7724, AUA, UNA).
The two other varieties of Baptisia australis, var. minor (Lehm.) Fern., of states west of the Mississippi River, and var. aberrans (Larisey) M. Mendenhall, have a different habitat, namely dry, rocky, calcareous glades and bluffs, usually far from water. Although B. australis var. aberrans is often a component of calcareous glades and barrens in the adjoining states of Georgia and Tennessee, it is absent from the Ketona Glades and, so far as known, from Alabama as a whole.
Only in a very few cases where the Ketona Glade habitat extends all the way down to the right bank of the Little Cahaba River is Baptisia australis var. australis found in association with some of the characteristic elements of the glade flora. It is not, therefore, to be considered a species of the Ketona Glades, strictly speaking.
Paronychia virginica Spreng.
Early in our explorations of the Ketona Glades we found that an unfamiliar, presumptive Paronychia was a characteristic perennial species of open, xeric habitats on the more centrally located outcrops. Determination to species had to await anthesis, and on July 19, 1992 we were rewarded with the first flowers, confirming our suspicion that the plant was Paronychia virginica Spreng. (A. and S. 6891, UNA).
The newfound Alabama populations have considerable phytogeographic significance, as they bridge an enormous gap between Virginia and Arkansas. The fact that Paronychia virginica is considered rare in these and the other states where it has been found (Missouri, North Carolina, Oklahoma, Texas, West Virginia) and exhibits such wide gaps in its distribution, suggests that it is a relictual species. The local abundance of such relicts (e.g., Silene regia), in combination with the richness of endemics for so small a geographic area, is evidence that the Ketona Glades have supported natural communities similar to those of the present day for a very long time.
Rhynchospora capillacea Torr.
One of the first finds on the initial, canoe-based expedition of May 30, 1992, after Marshallia mohrii and Spigelia gentianoides, was of a wiry-leaved sedge in crevices at the very base of a strongly sloping glade above the Little Cahaba River. The plant proved to be Rhynchospora capillacea Torr. (A. et al. 6676, UNA, VDB), a range extension of over 400 km to the southwest from the nearest previous collection site in northeastern Tennessee (Campbell County). The species has not been seen in Tennessee in many years and is possibly extirpated (Robert Kral, pers. comm. 1992). The Bibb County population would be by far the southernmost known, but for the relatively recent discovery of a population in Kerr County, Texas (Jones and Jones 1990).
Rhynchospora capillacea has yet to be found anywhere in Alabama except the few m2 of nearly bare rock just above the Little Cahaba River where we collected it. Because it has been found only in the glade-river ecotone of a single site, R. capillacea, like Baptisia australis, cannot be considered part of the characteristic Ketona Glade flora.
On June 21, 1992 we had the pleasure to show some of the glades we had discovered up to that time to Angus Gholson, Robert Godfrey and Robert Kral. One of the many high points of the day was the collection (A. et al. 6754, NY, UNA, US, VDB) of Rhynchospora thornei Kral, both a state record and the first collection of the species outside of the Coastal Plain.
This is among the most diminutive species of the genus, very similar to Rhynchospora divergens Chapman ex M.A. Curtis, a common but easily overlooked species primarily of the Coastal Plain. Rhynchospora thornei differs chiefly by having achenes with perianth bristles (Kral 1977). These bristles seem somewhat vestigial in nature, usually very short, unpigmented (translucent), and easily detached. Plants collected (A. and S. 7014, 7015, VDB) from one Ketona Glade population differed from all previous collections in that the bristles are somewhat longer and more pigmented, but were determined by Robert Kral to be R. thornei.
Prior to the discovery of this species in Alabama, in the Ridge and Valley Province, this former federal Candidate (C2) species was known from only a half-dozen collections, all from the Coastal Plain and from three states: Florida, Georgia, and North Carolina (Richard LeBlond, N.C. Natural Heritage Program, pers. comm. 1995). It was found subsequently at Coastal Plain localities in Alabama, beginning later in 1992 with a chalk glade in Dallas County (A. and S. 6921, VDB). Rhynchospora thornei has since been found at additional Ridge and Valley sites in Cherokee County, Alabama (J. Allison and A. Schotz 9519, UNA, VDB) and Floyd County, Georgia (A. 7776, GA, VDB), suggesting that the geographic range of Rhynchospora thornei is not fully known.
On the Ketona Glades and elsewhere, Rhynchospora thornei is a plant of microhabitats that are usually moist during winter and spring, irrigated by seepage, but which may become quite dry in summer and fall. Another Bibb County habitat for the species, and one unlike any in which it had previously been found, is crevices and shallow soil accumulations in a rocky creek bed (A. 6764, AUA, VDB). There, R. thornei is exposed during low water but inundated for much of the winter and after heavy rains. Perhaps this moisture regime is not very different in effect from the few seepage areas on Ketona Glades where R. thornei grows.
Solanum pumilum Dunal
Thomas Nuttall (1834) described Solanum hirsutum Nutt., from material collected by Samuel Boykin (Boykin s.n., PH) from the vicinity of Milledgeville, Georgia. Boykin collected the Solanum again in 1837 (specimen at NY, cited in D'Arcy 1974), from near Columbus, Georgia.
Two decades previous to Nuttall's publication, Michel Felix Dunal (1813) had described a different Solanum hirsutum (and Roxburgh yet another, soon thereafter). Nuttall's name was therefore (doubly!) a later homonym and must be rejected. Dunal (1852) rectified the situation by renaming Boykin's plant S. pumilum Dunal.
Asa Gray (1878) reduced the taxon to a variety of Solanum carolinense L., resurrecting Nuttall's epithet at varietal rank, a disposition followed by William D'Arcy (1974). As this was a taxon known only from three old specimens, the conservative treatments of Gray and D'Arcy were entirely defensible.
As of 1992, the two collections by Boykin, three sheets more than a century and a half old, were still the only known material of this taxon, and the plant was regarded as probably extinct (Shortia Torr. & Gray, after all, was lost to science for less than 90 years). On April 26, 1993 we discovered small populations of this Solanum (Figure 14) on two outcrops of Ketona Dolomite near the Little Cahaba River (A. and S. 7557, NY; A. and S. 7563, US). Subsequent exploration revealed that this taxon occurs about the margins of many of these glades and on at least one small glade of a different dolomitic formation (A. and S. 7651, VDB).
On May 1, 1994 we visited some amphibolite outcrops approximately 57 km to the southeast of the Ketona Glades. West of the Coosa River, in Chilton County, Alabama, we found (A. and S. 8239, MO, NY, VDB) one moderate-sized population of Solanum pumilum, and east of the river, in Coosa County, one rather small population (A. and S. 8241, UNA).
Other populations exist on Bibb County glades, but we saw many of these only in vegetative condition and they were not vouchered. After observing about 20 populations of this taxon, we have concluded that, though clearly related to Solanum carolinense, it should be recognized at the species level. Michael Nee, of the New York Botanical Garden, to whom we sent material of S. pumilum in 1998, concurred, writing (pers. comm. 1998), "its similarity to S. hieronymii and other spp. from Bolivia/Paraguay is striking, but obviously a good, distinct species."
There are multiple characters by which Solanum pumilum can be distinguished from S. carolinense, whereas the two varieties of S. carolinense, var. carolinense and var. floridanum (Shuttlw. ex Dunal) Chapman, reportedly can be distinguished only by leaf shape and, moreover, intergrade freely (D'Arcy 1974). Solanum pumilum differs from S. carolinense by (1) its smaller stature, less than 2 dm, vs. usually 3-10 dm; (2) stems without spines; (3) leaves rounded to obtuse at the apex, entire to shallowly sinuate, their prickles absent or few, weaker, and confined to the midvein [vs. mid- and late-season leaves acute, coarsely dentate (or deeply lobed in var. floridanum), prominently spiny on midvein and often also on secondary veins]; (4) corollas always pure white (usually lavender in S. carolinense, although white morphs–—f. albiflorum O. Kuntze–—are occasional); and (5) flowers sweetly fragrant, vs. odorless (pers. obs. of Allison over southeastern states from Arkansas to North Carolina) in S. carolinense.
Solanum carolinense is a common, native weed in the southern states (and beyond) and is found on a few Ketona Glades where there has been some disturbance, e.g. from road construction or lumbering activities. These plants are usually stunted by the extreme conditions prevailing on the glades, and thus resemble S. pumilum in stature. In all other respects, most noticeably in having cauline prickles and upper leaves coarsely dentate, they are easily recognizable as S. carolinense.
Spiranthes lucida (H. H. Eat.) Ames
Like Rhynchospora capillacea, Spiranthes lucida is very rare as far south as Tennessee, its previous southern limit, and is thus far known in Alabama from a (different) single rocky place on the right bank of the Little Cahaba River. Allison discovered this, the latest of the state records, while canoeing the Little Cahaba River with Jim Affolter and David Handlay, on April 29, 1994. We could find only ten or so individuals, and the population was vouchered by photographic slides and prints the authors took the next day, A. and S. 8224-p (AUA, JSU, UNA, VDB). We revisited the site a month later, hoping to collect seed, but, perhaps due to a very dry May, the infructescences had senesced, apparently without perfecting any seed.
The single locality known is a ledge of dolomite just above the river, where a glade extends down to the water's edge and where Spiranthes lucida is associated with Rhynchospora colorata and R. thornei. It is probably slightly inundated at times in winter and after heavy rains. Like Baptisia australis var. australis and Rhynchospora capillacea, Spiranthes lucida is considered a peripheral element of the Ketona Glade flora.
Other Rarities
Noteworthy additional rare taxa found to date on one or more Ketona Glades include county records for another 16 taxa tracked by the Alabama Natural Heritage Program (ALNHP). Most notable were federally Endangered Xyris tennesseensis Kral (A. and S. 7004, VDB), previously known in Alabama only from a small population about 160 km to the northwest, and a current and a former Federal Candidate (C2) Species, Symphyotrichum georgianum (Alexander) Nesom (A. and S. 7306, VDB), and Leavenworthia exigua var. lutea (A. and S. 7506, VDB). The 13 other county record rarities were Echinacea purpurea, Helianthus smithii Heiser, Isoëtes butleri, Leavenworthia uniflora, Mirabilis albida, Ophioglossum engelmannii, Orobanche uniflora L., Pediomelum subacaule, Ponthieva racemosa (Walt.) C. Mohr, Schoenolirion croceum, Spiranthes magnicamporum, Symphyotrichum laeve var. concinnum, and Veronicastrum virginicum (L.) Farw. The identification of the Spiranthes is tentative: some or all of these plants may be referable to S. cernua (L.) L. C. Rich., "contaminated" with genes from S. magnicamporum, as reported from other states by Sheviak (1991).
We also found new, Ketona Glade locations, for another 18 rare taxa that others had collected previously somewhere in Bibb County. These taxa included federally Threatened Marshallia mohrii, which achieves its greatest abundance on these glades; Silene regia, a former federal Candidate (C2) species that had been feared extinct in Alabama; one current and two other former C2 species, Arabis georgiana, Croton alabamensis var. alabamensis, and Rudbeckia triloba var. pinnatiloba; and 13 additional taxa tracked by the ALNHP: Callirhoë alcaeoides, Carex eburnea, Cheilanthes alabamensis, Enemion biternatum Raf., Gentiana villosa L., Leptopus phyllanthoides, Liatris cylindracea, Penstemon tenuiflorus, Polygala boykinii, Rhynchospora colorata, Scutellaria alabamensis, Sida elliottii, and Silphium trifoliatum var. latifolium.
Davison and Schotz recently (1998) reported finding, on one of the large glades, two rare liverworts: Plagiochasma crenulatum Gott, new to the eastern United States, and Cheilolejeunea clausa (Nees & Mont.) Schust. It seems certain that still other botanical (and zoological?) rarities occur on these glades and await future detection.
Within about 100 meters of one or more Ketona Glades we recorded at least 16 other rarities, including Melanthium woodii (J. W. Robbins ex Wood) Bodkin, known previously in Alabama from a single population more than 240 km to the southeast, and three more former C2 species: Jamesianthus alabamensis Blake & Sherff, a species thought at the time to be endemic to a tiny area in Alabama more than 175 km to the northwest, and two plants named by the same botanist for their same discoverer, Sedum nevii Gray and Neviusia alabamensis Gray, the latter reported, at the time, from only 15 other populations. The remaining rarities found near one or more Ketona Glades were Gentiana saponaria L., Hypericum nudiflorum Michx., Marshallia trinervia (Walt.) Trel., Melanthium latifolium Desr., Panax quinquefolius L., Parnassia grandifolia DC., Plantago cordata Lam., Phlox pulchra (Wherry) Wherry, Rhapidophyllum hystrix (Pursh) H. Wendl. & Drude ex Drude, Schisandra glabra (Brickell) Rehder, Silene caroliniana Walt. ssp. wherryi (Small) Clausen, and Zanthoxylum americanum P. Mill.
Finally, Bibb County's trove of botanical rarities is by no means limited to plants found in the vicinity of Ketona Glades. Additional rarities that we encountered during our Bibb County explorations, but at distance from any of the Ketona Glades, included Carex impressinervia Bryson, Kral & Manhart, Cladrastis kentukea (Dum.-Cours.) Rudd, Corallorrhiza wisteriana Conrad, Croomia pauciflora (Nutt.) Torr., Cystopteris bulbifera (L.) Bernh., Euonymus atropurpureus Jacq., Hymenocallis coronaria (Le Conte) Kunth, Pachysandra procumbens Michx., Phacelia dubia (L.) Trel. var. dubia, and Trillium decumbens Harbison.
The ALNHP Tracking List is revised at intervals, and a few of the taxa named in these paragraphs have subsequently been removed from it, due at least in part to additional occurrences discovered in Bibb County.