Supplement to a paper published in Castanea, Vol. 66(1-2):154–205, March/June 2001
Musical selection: Gavotte et Doubles from Nouvelles Suites de Pičces de Clavecin by Jean-Philippe Rameau (1683-1764)
Vascular Flora of Ketona Dolomite Outcrops in Bibb County, Alabama
Liatris oligocephala is the only known Liatris lacking tiny bristles on the ribs of its achenes. It is also the only Liatris that produces mostly one to three heads or at most a simple cyme. Liatris cymosa (Ness.) K. Schum. and L. ohlingerae (Blake) B. Rob. often have the heads arranged in a simple cyme, but vigorous individuals of both these species bear compound cymes. Despite its unique features, the Ketona Glade plant is clearly a Liatris because of its stocky, often globose "corm," lack of any bracts between the florets, and the fact that it hybridizes with L. cylindracea.
At right is the color version of Figure 9 from the Castanea
relates to hybridization in Ketona Glade species of Liatris.
photographs taken 13 July 1993 at the type locality of
cylindracea X L.
showing differences in head arrangement: A.
cylindracea. B. L. Xfreemaniana.
Middle row, pressed heads of same taxa, from
collections [UNA] made 18 July 1993, showing differences in outer
shape and in degree of ciliation: D.
Allison & Stevens 7800.
E. Allison & Stevens
7801. F. Allison
& Stevens 7815.
Bottom row, photographs taken 19 August 1999 at the type locality of
(L. cylindracea X L.
squarrosa), showing differences
in phyllary shape and orientation:
H. L. Xmacdanieliana.
A 2005 paper by T. M. ("Mike') Hardig, James R. Allison and Edward E. Schilling was focused on Liatris oligocephala, L. cylindracea and putative hybrids (L. Xfreemaniana). Titled "Molecular Evidence of Hybridization between Liatris oligocephala (Asteraceae) and More-Widespread Congener: A Preliminary Assessment of the Potential for Extinction" (Castanea 70:246-254, available at http://www.bioone.org/doi/pdf/10.2179/0008-7475%282005%29070%5B0246%3AMEOHBL%5D2.0.CO%3B2), its abstract is quoted here in entirety:
Hybridization may increase the risk of extinction for some rare and endangered plant species, either by increasing the rate of random genetic drift affecting populations or by introgression-mediated genetic assimilation. The former may occur when hybrids are less fit than non-hybrids and backcrossing would be less likely to occur, the latter may occur when hybrids are as fit, or fitter, than parents, and backcrossing is fairly common. To assess the risk of extinction by hybridization one must assess whether hybridization is actually occurring, and, if so, whether the hybrids are sterile, or are backcrossing. We analyzed one population each of the rare Liatris oligocephala and L. cylindracea and seven field-identified hybrids for ribosomal DNA and chloroplast DNA markers, and morphological characteristics to make a preliminary determination of the potential for L. oligocephala extinction via hybridization. We found unequivocal evidence that the two species are hybridizing, but no unequivocal evidence of backcrossing.
Further study will be necessary to determine whether Liatris oligocephala is threatened at all by genetic "swamping," which could result from backcrossing from F1 hybrids with L. cylindracea. Greater impact to L. oligocephala populations may well come from competition with F1 hybrids (L. Xfreemaniana) in microsites that are transitional from the severe microhabitats to which L. oligocephala seems better adapted to those microhabitats more favorable to L. cylindracea, with deeper soil and/or more shade.
That same year (2005), Guy L. Nesom published a paper titled "Infrageneric classification of Liatris (Asteraceae: Eupatorieae)" [Sida 21:1305-1321 (available for download at http://www.guynesom.com/LiatrisInfragClass.pdf)]. In the Castanea paper (p. 178) I had written that "This new Liatris appears to be sufficiently distinctive to merit the establishment of a new series, perhaps even a new section, to accommodate it, but we will leave that decision to a future monographer." In his 2005 paper, Nesom made L. oligocephala the type-species of his newly created Section Vorago. I was surprised to find a second species included there with it, the Texas endemic L. tenuis Shinners, a plant quite dissimilar to L. oligocephala in gross appearance. I suspect that future molecular studies will not support this disposition; this would apparently come as no surprise to Nesom, either, as he was explicit that the association of these two species was tentative:
Liatris oligocephala is similar to L. tenuis in features of the involucre, corolla, and pappus, but because no synapomorphy is evident, it seems likely that these two species originated independently from the ancestral stock of sect. Liatris. Thus, sect. Vorago, as delimited here, may not be monophyletic.... The name of the section (“vorago,” Latin, gulf) alludes to the range of the two species on the Gulf Coastal Plain as well as to the considerable morphological “gulf” between them."
Nesom's choice of the epithet Vorago was rather unfortunate, as the type species, Liatris oligocephala is not found on the Gulf Coastal Plain, but instead in the Ridge and Valley Province (its obligate association with a Cambrian-age sedimentary rock formation makes this unequivocal). Nesom may have gotten this misconception from certain literature containing the same error, perhaps owing to the fact that the majority of Bibb County, Alabama does in fact lie within the Gulf Coastal Plain.
Guy L Nesom also contributed the FNA treatment of Liatris (Vol. 21, 2006). His treatment of L. oligocephala is available online at http://www.efloras.org/florataxon.aspx?flora_id=1&taxon_id=250067110. Unlike full species, hybrids are of course not treated individually. Those that have received names as nothospecies, including L. Xfreemaniana and L. Xmacdanieliana, are listed in FNA with their putative (sometime speculative) parents (as can be viewed at http://www.efloras.org/florataxon.aspx?flora_id=1&taxon_id=242416766).
In stature and flower color, Onosmodium decipiens more closely resembles O. virginianum. The former has longer hairs on its stem (> 2.5 mm vs. <2 mm), but the critical differences are in the flowers. The photo at left shows dissected corollas rehydrated from pressed specimens of O. virginianum (left half) and O. decipiens (right half). The consistently shorter filaments of O. virginianum cause the anther apices to be positioned distinctly below the sinuses (notches) of the corolla, while the consistently longer filaments of O. decipiens result in anther apices that reach—and often slightly exceed—the bases of the sinuses. A further difference is seen in the shape of the corolla lobes, more narrowly and sharply triangular in O. virginianum than in O. decipiens.
The corollas of Onosmodium decipiens are more like those of O. molle ssp. hispidissimum, except that they are pale yellow (subspecies of O. molle have white corollas with greenish-tinged lobes). The pictures above, right (O. decipiens) and at left (O. molle ssp. hispidissimum) show plants early in anthesis. The marked differences between these two species in habit (ascending vs. erect) and in average stature (most plants less than 6.5 dm in O.decipiens, with a maximum of 8.4 dm, while plants a meter or more tall are frequent in subspecies of O. molle) are apparent. Also evident is the tendency for O. decipiens (like O. virginianum), to produce several stems, while taxon hispidissimum (like other subspecies of O. molle), produces one or very few. While the long, stiff pubescence of both taxa appears similar, a look at the foliage with a hand lens reveals another distinction. Between the veins, taxon hispidissimum has an under-pubescence of shorter, more appressed hairs beneath the long, spreading, conspicuous ones (another common feature of the subspecies of O. molle), while in O. decipiens (and O. virginianum) such hairs are confined to the veins. A further difference between these two is found in the nutlets, which are smaller and gradually tapered to the truncate base in O. decipiens (left half of image at right, above), while nutlets with distinctly "collared" bases have always been considered a hallmark of taxon hispidissimum (right half of image at right, above).
The major differences among
these taxa of Onosmodium
the following table: Morphological
molle ssp. hispidissimum O. decipiens O. virginianum stem
length to ca. 12 dm to ca. 6.5 (-8.4) dm to ca. 5 dm stems
per plant few few to several stem
hair length > 2.5 mm > 2.5 mm < 2.0 mm leaf
indument double simple simple corolla
color white w/ greenish lobes light yellow light yellow corolla
lobes nearly deltoid nearly deltoid acuminate anther
apices at corolla sinuses at sinuses below sinuses nutlet
base constricted unconstricted unconstricted
of three Onosmodium taxa
O. molle ssp. hispidissimum
to ca. 12 dm
to ca. 6.5 (-8.4) dm
to ca. 5 dm
stems per plant
few to several
stem hair length
> 2.5 mm
> 2.5 mm
< 2.0 mm
white w/ greenish lobes
at corolla sinuses
The tenth annual (2008) Spring Symposium at Eastern Oregon University (La Grande) included a poster presentation by Kali Miller, titled "Comparing Pyrrolizidine Alkaloids in Onosmodium decipiens and O. virginianum." I am not aware of any publication of results of this study. The abstract (found at www.eou.edu/sprsymp/documents/program.pdf) is as follows:
Pyrrolizidine alkaloids (PA’s), potentially toxic natural secondary products, are typically found in the plant family Boraginaceae. As a member of the family, the genus Onosmodium can be expected to contain PA’s. Onosmodium virginianum is found throughout the southeastern United States, while the recently described closely related O. decipiens is prevalent only in Bibb County, Alabama. Previous work on O. decipiens established the presence of the PA’s uplandicine and lithospermine. A preliminary GC/MS investigation of O. virginianum has discovered the main PA to be a novel saturated analog of the PA echimidine with lesser amounts of the PA’s uplandicine, intermidine, echimidine, and supinine. The exact structure of the alkaloids found in the O. virginianum will be elucidated using NMR experiments. A chemosystematics comparison between the two Onosmodium species will be presented.
The combination of gradually reduced leaves, and heads usually with 13 rays align Silphium glutinosum with the S. asteriscus complex (e.g., S. asteriscus L., S. trifoliatum L. and their varieties), but none of these has glandular stems, leaves, or involucres, and their leaves average narrower than those of S. glutinosum.
After showing Robert Kral Silphium glutinosum in the field in Bibb County, prior to anthesis, he called attention to densely glandular specimens labeled S. integrifolium he had collected from the Black Belt province of Alabama, in Dallas County (Kral 48820, VDB) and Perry County (Kral 47891, VDB). Tim Stevens and I visited both of these populations, and "windshield surveying" along highways and back roads has resulted to date in the discovery of well over a dozen additional populations of a densely glandular Silphium in those two counties. After comparison of living populations and preserved material of these plants with S. integrifolium and with S. glutinosum, we determined that a suite of characters exists that distinguish the glandular Silphium of Dallas and Perry Counties from either of those species.
Silphium integrifolium differs from S. glutinosum in many respects. The former is, under favorable conditions in the wild and in the garden (e.g., garden of Allison), a taller plant with nodes slightly closer together and therefore more numerous. Those leaves that persist until anthesis in S. integrifolium are normally sessile and, like the stem, eglandular. It also has a narrower inflorescence, on average with fewer but larger heads, these with consistently more numerous rays and phyllaries (the latter never with the revolute margins seen in S. glutinosum) and with acute receptacular bracts (pales or chaff).
Plants of the Dallas and Perry County Silphium are intermediate between S. glutinosum and S. integrifolium for some characters and for others resemble one or the other of those species, as indicated in the following table:
mostly 8-15 dm
mostly 15-20 dm
often >15 dm
nodes below inflorescence
mostly 6 or 7
often 10 or more
stem and leaf glandular hairs
lowest petiole length at anthesis
mostly 6-15 cm
mostly 5-8 cm
mostly 0 cm
open or ± congested
obtuse to acutish
number of rays per head
(8) 12-14 (16)
(17) 19-23 (33)
(16) 21-34 (35)
At left is the color version of Figure 12 from the Castanea paper. Top row, Silphium glutinosum, Bibb County, Alabama: A. Habit, including open inflorescence; "Desmond's Glade," 9 September 1993. B. Head, frontal view, with the normal 13 rays; "Goat Glade South," 14 June 1992. C. Head, lateral view, showing outer phyllaries the longest and with recurving margins; "Browne's Dam North Glade West," 3 July 1992. Middle row, S. perplexum: D. Habit; Perry County, Alabama, Co. Rd. 6, 19 August 1999. E. Heads, frontal view, showing maximal no. of rays per head; same date and location as in D. F. Head, three-quarters view showing the species' comparatively "ordinary" involucre; Dallas County, Alabama, type locality, 18 August 1999. Bottom row, S. integrifolium: G. Habit, including more numerous nodes, Lowndes County, Mississippi, Alt. U.S. Hwy. 45, 9 August 1992. H. Heads, frontal view, with numerous rays; Sumter County, Alabama, Ala. Hwy. 17, 19 August 1999. I. Comparatively congested inflorescence, same date and location as in G.
The image at lower left shows the chaff (pales, receptacular bracts) from a rehydrated and dissected head of Silphium glutinosum. The chaff apices are consistently obtuse. The chaff of S. integrifolium (right) is dramatically different—strongly acute. While intermediate between these two species in many respects, S. perplexum tends to have chaff apices more similar to those of S. glutinosum.
The treatment of Silphium in FNA (Vol. 21, 2006) was contributed by Jennifer A. Clevinger, of James Madison University. Her treatment of S. glutinosum is available online at http://www.efloras.org/florataxon.aspx?flora_id=1&taxon_id=250067527. As stated on page 187 of the Castanea article, "Given the frequency of hybridization in human-perturbed habitats, it appears that, among related taxa of Silphium, barriers to gene flow are chiefly ecological. Alteration of the landscape by humans appears to have disrupted these barriers, just as in Coreopsis." It follows that any taxonomic treatment of Silphium faces similar impediments to a thorough understanding to those confounding a treatment of Coreopsis, as discussed above under that genus. Except that the mostly tall Silphium spp. are even more subject to be represented by "top-snatched" material in herbaria! The mostly herbarium-based Silphium treatment in FNA reflects this unfortunate state of affairs. In particular the elements of the S. asteriscus complex, perhaps necessarily "lumped" in large part as a practicality, are desperately in need of studies that include a strong field component that focuses on habitats not created or transformed by humans, a quest that becomes increasingly difficult, if not impossible, every day.
I would feel better about the failure to accept Silphium perplexum in FNA if what was written about it there were more accurate. According to Clevinger, "James Allison (in protologue) suggested that Silphium perplexum is a hybrid between S. glutinosum and S. integrifolium. It has glandular hairs on its stems, leaves, and phyllaries as in S. glutinosum; it is also found on chalk, has taller stems, larger leaf blades, and deeper yellow corollas. Further study is needed to determine its status." Most unfortunately, the most salient difference between S. glutinosum and S. perplexum received no mention, namely the consistently greater number of ray flowers in the latter (or ovaries, as can be determined by dissection of a head once some of the ray flowers have abscised). It is unusual for the number of rays/ovaries in a head of S. glutinosum to number other than 13, unheard of for S. perplexum to have so few. I interpreted this and other features of their comparative morphology as well as their geographic ranges as suggestive of an origin of S. perplexum in prehistoric hybridization of S. glutinosum with the [today] wholly allopatric S. integrifolium. This is not at all the same thing as calling Silphium perplexum "a hybrid between S. glutinosum and S. integrifolium" as misstated by Clevinger, which would imply that these two allopatric species are in genetic contact today or even in the recent past. All three species are in fact allopatric.
I am much less concerned about the eventual vindication of Silphium perplexum as distinctive as I am about the edaphically less specialized taxa regarded as varieties of S. asteriscus. While introgression of S. perplexum with other species is indeed occurring along roadsides—a hybrid swarm within the city limits of Selma (west side of Bell Rd, ca. 0.2 rd. mi. S of jct. U.S. Hwy. 80) is especially memorable)—a sizeable amount of its chalk prairie habitat around the type locality of Old Cahaba (Dallas County) is now protected (another 3,020 acres as recently as September 2003), and hybrids are unlikely to have greater fitness there than S. perplexum, except along roadsides.
That Silphium perplexum is as about as close morphologically to S. integrifolium as it is to S. glutinosum (and thus can satisfactorily be made a mere variety of neither) is borne out by Clevinger's 1999 annotation of the only 19th century collection known of S. perplexum (at GH, shown at right) as S. integrifolium (likewise determined by all other annotators).
An additional factual error in the FNA treatment of Silphium begs correcting: the habitat of S. glutinosum is given there as "Limestone glades, along rocky streams." While a very few occurrences along rocky streams are over limestone (e.g., near Pratt's Ferry, as mentioned above, immediately preceding the section titled "Biological Communities), the glades on which S. glutinosum grows are always underlain by dolomite. While limestone and dolomite are both calcareous sedimentary rocks, considering the terms interchangeable (as also in Lawless et al., 2006) seems analogous to regarding the words "daisies" and "goldenrod" equivalent because they belong to the same botanical family!
In discussing Spigelia gentianoides Chapm. in A. DC., the Castanea paper indicated that
Many, if not most, Florida populations of Spigelia gentianoides have been lost or are in the process of disappearing as a result of fire suppression. By contrast, the Ketona Glade habitat is probably maintained more by rockiness and edaphic conditions. However, occasional wildfire in the ecotones would have served to limit encroachment from the glade margins. Such a role in the past for wildfire in maintaining the Ketona Glade ecosystem is indicated by the persistence, in the forest matrix surrounding some of the glades, of longleaf pine and such wiregrass-country associates as Sericocarpus tortifolius (Michx.) Nees and, less commonly, Carphephorus odoratissimus (J. F. Gmel.) Herbert.
A 2007 paper examined the effects of prescribed fire on rare plants of Ketona dolomite glade communities. Titled "The Effect of Fire Reintroduction on Endemic and Rare Plants of a Southeastern Glade Ecosystem" (Restoration Ecology16:39-49), this paper by R. Scot Duncan, Corinna B. Anderson, Heather N. Sellers, and Erin E. Robbins, is available from http://www3.interscience.wiley.com/cgi-bin/fulltext/119413438/PDFSTART.
The abstract is quoted in full as follows:
Open habitats dominated by herbaceous plants on thin, rocky soils occur within the forests of eastern North America. Although these habitats vary in origin, structure, geology, and species composition, all contribute greatly to regional biodiversity by harboring endemic and/or rare plants. Little is known about how disturbances affect plant populations in these ecosystems. Fire once was a frequent natural disturbance in the Ketona dolomite glades of Alabama, an ecosystem harboring eight endemic taxa and numerous other species of conservation concern. We designed an experiment to determine how the reintroduction of fire into the glades and surrounding longleaf pine forests affects populations of rare glade plant species. Experimental and control plots were established within the glades. Experimental plots were burned in April 2004, and all plots were surveyed during two subsequent growing seasons (2004 and 2005). Populations of three of 14 species of conservation concern declined significantly after the initial fire but recovered the next year. Among other herbaceous species, only five and two differed in population size in 2004 and 2005, respectively. In 2004, more species were more abundant in control than burned plots, but this difference was not detected in 2005. Multivariate community-level analyses of species presence-absence suggested that the effects of fire were negligible by the 2005 survey. Populations of young trees that had invaded the glades declined dramatically as a result of treatment fires. These results suggest that the reintroduction of fire will not harm glade species and may help prevent encroachment of the surrounding forest.
Which is good, since The Nature Conservancy of Alabama and its partners have conducted a number of controlled burns to date at the Kathy Stiles Freeland Bibb County Glades Preserve (photo at right; note blackened lower trunks of pines). After seeing the results, I have no doubts that these prescribed fires have enhanced the vigor and diversity of the herbaceous component in glade-forest ecotones, and served to reverse the slow encroachment upon these glades of adjacent woodland communities.
Considerable recent information about Spigelia gentianoides can be found in "Spigelia gentianoides, Gentian pinkroot, 5-Year Review: Summary and Evaluation" (2009), [available at http://www.fws.gov/southeast/5yearReviews/5yearreviews/20090121GentianPinkroot.pdf] by the U.S. Fish and Wildlife Service, Panama City, Florida Field Office (Vivian Negrón-Ortiz, Lead Recovery Botanist). Among the findings reported: there are five locations where the endangered variety gentianoides is known to be extant, including two sizeable occurrences on state-owned land in Jackson County, Florida. At Apalachee Wildlife Management Area (where discovered in 2007), more than1,700 individuals are known, divided into two subpopulations, and Three Rivers State Park is estimated to support more than1000 individuals. A third Jackson County population, on private land, was thought to consist of only four plants. The fourth Florida population, of about 100 plants, is known from Calhoun County, on a preserve owned by The Nature Conservancy. The Castanea paper (p. 190) reported a personal communication from John MacDonald that he had discovered this variety in adjacent Alabama. This occurrence, in the Geneva State Forest (Geneva County) was estimated in 2007 to contain about 400 individuals.
Regarding variety alabamensis K. Gould, Negrón-Ortiz et al., using belt transects, estimated there were a total of 3,653 individuals on 4 of the 17 Ketona dolomite glades where it had previously been found.
The recommendations of the USFWS report include the suggestion that "a taxonomic study using a multi-data approach (e.g., morphology, molecular studies) is encouraged for discerning whether the two varieties represent distinct species. If the results suggest that the varieties represent distinct species, then both should be considered for federal protection." However, the only information available since the Castanea paper and relevant to this issue that was mentioned in the USFWS report was the following:
Electrophoretic studies indicated that the genetic identity between the S. gentianoides varieties is high. The Florida sample used in the study, presently growing at the Historic Bok Sanctuary (Sanctuary), from seeds collected at Calhoun County, included a relatively narrow subset of the genetic variation found in the alabamensis populations (Affolter 2005*). The majority of allozyme diversity in var. alabamensis was found to be distributed within rather than among glades, therefore the author suggested that nearly all the genetic diversity will be captured by preserving the largest glades.
* Affolter, J. M. 2005. Conservation biology of Spigelia gentianoides and S. marilandica: genetic variation, reproduction biology, and propagation. Final project to the Georgia Cooperative Fish and Wildlife Research Unit. 33 pp.
At the top of page 195 of the Castanea article I wrote, "It seems certain that still other botanical (and zoological?) rarities occur on these glades and await future detection." Some of the insects, at least, of the Ketona Glades have received attention from zoologists. In 2006 JoVonn G. Hill, of the Mississippi Entomological Museum at Mississippi State University (Starkville) reported "The First Record of the Green Stinkbug Chlorochroa persimilis Horvath (Hemiptera: Pentatomidae) from Alabama" [Marginalia Insecta 1(2):1-2]. The next year Hill reported on "The Grasshoppers (Orthoptera: Acrididae) of the Bibb County Glades, Alabama" [Marginalia Insecta 2(1):1-3. 2007]. According to Hill, "The Mississippi Entomological Museum (MEM) visited these glades three times (1-2 May, 4-6 June, and 6-7 August 2003) for the annual William H. Cross Collecting Expedition," when nine species of grasshoppers were found. Hill returned to the glades twice in 2005, on August 13th and on September 8th and 9th. "These trips yielded 10 additional species of grasshoppers resulting in a total of 19 species for the site." Hill's 2007 paper continues:
No new state records or new species were found in these habitats, but there were a few interesting finds. Both Spharagemon cristatum (Scudder) and S. marmorata picta (Scudder) are species that are typically associated with sandy habitats (Dakin and Hayes, 1970 and Otte 1984). Here, they were found dwelling on the rocky dolomite outcrops on the most extensive glade on a bluff overlooking the Little Cahaba River. (Figure 1*). Although S. cristatum is known to occupy sandy areas near rivers (there were some sandy margins on the river below), there were significant numbers of this species occupying the rocky outcrops. Both Spharagemon species were camouflaged quite well when resting on the exposed dolomite. Another interesting find was that of Melanoplus impudicus Scudder, which can be found in dry areas of sparse vegetation on rocky or sandy areas (Gurney, 1941 and Dakin and Hayes, 1970). Gurney (1941) figured a shale barren habitat at Short Mt. near Mt. Jackson, Massanutten Mts. Virginia where M. impudicus was found and this shale barren is similar in appearance to the Bibb County Glades.
* Reproduced at right, with a Pinus palustris that I recognize as the leftmost in a grouping of that species in a photo featured elsewhere on this web site (http://www.jimbotany.com/pinuspal.jpg), and marking the site to me as the one I named "South Goat Glade."
Hill's 2006 paper is available at http://mississippientomologicalmuseum.org.msstate.edu//MEM.Pubs/Marginalia.Insecta/2006.papers/Marginalia.Insecta.Vol.1.2.pdf, the 2007 paper at http://mississippientomologicalmuseum.org.msstate.edu/MEM.Pubs/Marginalia.Insecta/2007.papers/Marginalia.insecta.Vol.2.1.pdf.
Page 157, first paragraph: replace "Psora rubiformis (Ach.) Hooker vel aff." with "Psora pseudorussellii Timdal" (I have re-identified my collection, which is apparently yet another range extension (according to the Consortium of North American Lichen Herbaria, previously recorded from no closer than Arkansas; see http://symbiota.org/nalichens/index.php).
Pages 163, 168, 178 (twice), 182, 186, 196: correct the spelling of "Brown's" Dam to "Browne's" Dam, as this historic site was named for William P. Browne, according to Bibb County, Alabama: The First Hundred Years, by Rhoda C. Ellison (The University of Alabama Press, 1984).
Page 169: first line of third paragraph: delete "a decumbent habit,"
Page 191, end of third paragraph: delete ", UNA"
Page 195, line 11: delete "(Muhl. ex Willd.)"
Page 199, under ASTERACEAE, in text for Coreopsis grandiflora var. inclinata X C. pubescens, replace "A. & S. 6719" with "A. 11927"
Page 200, under CONVOLVULACEAE: the correct spelling of the specific epithet of the Calystegia is apparently "catesbeiana"
Page 200, under CYPERACEAE, in text for Rhynchospora globularis var. pinetorum: replace "(Britt. & Small ex Small) Gale" with "(Britt. & Small) Gale"
Page 201, under FAGACEAE, correct "Quercus margarettiae" to "Quercus margarettae" [an error copied from USDA 2000 (PLANTS Database, since corrected there)].
Page 201, line 11: add "(AUA)" after "7724"
Page 202, line 43: replace "A. and S. 7413" with "A. and S. 7430"
Page 202, line 45: replace "J. Allison and C. Oberholster 7049" with "A. and S. 7413b"
These are mostly additions to the flora or are taxonomic changes (made subsequent to going to press, or then somewhat controversial but since gaining wider acceptance by systematists). I cannot claim to have kept up with all of these; most of the credit must go to Alan Weakley, via his "Flora").
Page 154: the first author's current address for correspondence: 2242 Highway 212 SW; Conyers, Georgia 30094.
Page 159, end of second paragraph: the URL for the referenced web site has changed to http://www.jimbotany.com/.
Page 199, before AGAVACEAE add "ADOXACEAE" and list under it "Viburnum rufidulum Raf." (transfer this from CAPRIFOLIACEAE. page 200).
Page 199, under APIACEAE, Thaspium barbinode var. chapmanii: add "[T. chapmanii (Coult. & Rose) Small]" as this taxon may reasonably be treated as a full species (e.g., 2010 edition of "Weakley's Flora").
Page 199: transfer taxa listed under ASCLEPIADACEAE to APOCYNACEAE.
Page 199, under APOCYNACEAE (ASCLEPIADACEAE): the correct name for the "Matelea" is now regarded to be Gonolobus suberosus (L.) R. Brown.
Page 199, under ASTERACEAE, Acmella oppositifolia var. repens: add "[A. repens (Walt.) L.C. Rich. in Pers.], as this taxon may reasonably be treated as a full species (e.g., 2010 edition of "Weakley's Flora").
Page 199, under ASTERACEAE, after "Coreopsis major Walt." add the clarifying "var. major"
Page 199, under ASTERACEAE, Silphium astericus var. angustatum, add "[S. asteriscus var. dentatum (Ell.) Chapm.]" as the former taxon may be considered synonymous with the latter (e.g., Flora of North America North of Mexico, Vol. 19).
Page 199, under ASTERACEAE, Silphium trifoliatum var. latifolium, add "[Silphium asteriscus L. var. latifolium (A. Gray) J.A. Clevinger]" as this taxon is so treated in Flora of North America North of Mexico, Vol. 19.
Page 200, under BRASSICACEAE, Arabis canadensis: add "[Boechera canadensis (L.) Al-Shehbaz], as there is now strong evidence for placing most southeastern "Arabis" spp. in Boechera Löve & Löve.
Page 200, after BUDDLEJACEAE, add "[TETRACHONDRACEAE]", as Buddleja seems to belong to the Scrophulariaceae (so no Buddlejaceae), and while the placement of Polypremum procumbens remains controversial, it is perhaps best placed with Tetrachondra Petrie [Oxelman, B., M. Backlund, and B. Bremer. 1999. Relationships of the Buddlejaceae s.l. investigated using parsimony jackknife and branch support analysis of chloroplast ndhF and rbcL sequence data. Syst. Bot. 24: 164-182].
Page 200, under CAMPANULACEAE: add "Triodanis perfoliata (L.) Nieuw."
Page 200, under CARYOPHYLLACEAE: at end of column 1 add "[S. caroliniana Walt. var. wherryi (Small) Fern.]".
Page 200, CLUSIACEAE: see first note below for page 201.
Page 200, under CONVOLVULACEAE: delete "(identification tentative)" from the Calystegia.
Page 200, under CYPERACEAE, Rhynchospora globularis var. pinetorum: after "Gale" insert "[R. pinetorum Britton & Small]".
Page 200, under CYPERACEAE: Scleria verticillata, delete "(single locality)".
Page 200, under EUPHORBIACEAE, transfer the Leptopus to PHYLLANTHACEAE after adding this family name before PLANTAGINACEAE (page 202).
Page 201, before IRIDACEAE, add "HYPERICACEAE" and transfer the Hypericum spp. from page 200 under CLUSIACEAE (delete the latter family). After Hypericum densiflorum text add "[referable to the narrow-leaved plants named as H. interior Small, probably a good subspecies but the combination apparently not yet made]".
Page 201, before IRIDACEAE, add "HYPOXIDACEAE" and transfer the Hypoxis from the former family to the latter.
Page 201, under LILIACEAE: Aletris is now regarded as belonging in NARTHECIACEAE (insert this family before NYCTAGINACEAE, page 202, Allium and Nothoscordum in the ALLIACEAE (insert before ANACARDIACEAE, page 198), Camassia and Schoenolirion in the AGAVACEAE (page 198). Polygonatum has been placed in the RUSCACEAE (insert before RUTACEAE, page 203), Trillium in the TRILLIACEAE (insert before ULMACEAE, page 203), and Uvularia in the COLCHICACEAE (insert before COMMELINACEAE, page 200). The three species listed as Melanthium have been transferred to Veratrum, in the MELANTHIACEAE (insert this family after MALVACEAE, page 201. In the entry for M. latifolium, after "Walt." insert ", Veratrum latifolium (Desr.) Zomlefer" and in the entry for M. virginicum, after "L.}" insert "[Veratrum virginicum (L.) Ait.]".
Page 201, under LOGANIACEAE: Gelsemium has been made the type-genus of its own family; insert "GELSEMIACEAE" before GENTIANACEAE and transfer G. sempervirens.
Page 202, under POACEAE: after Sporobolus vaginiflorus insert "Tridens chapmanii (Small) Chase" and place the combination "Tridens flavus var. chapmanii" (with the corresponding authorities) in brackets as a synonym.
Page 202, under PRIMULACEAE: Dodecatheon L. is now regarded as insufficiently distinct from Primula L.; insert the new combination "Primula meadia (L.) A.R. Mast & Reveal" and place Dodecatheon meadia L. in brackets as a synonym.
Page 202, under PRIMULACEAE: Lysimachia is now regarded as belonging to the MYRSINACEAE; insert this name following MYRICACEAE (page 201) and transfer the Lysimachia spp. there.
Page 202, under RANUNCULACEAE: add "Thalictrum revolutum DC."
Page 203, under ROSACEAE: add "Prunus angustifolia Marsh. (single locality)".
Page 203, under RUBIACEAE: add "Houstonia canadensis Willd."
Page 203, under SAXIFRAGACEAE: the only genus listed, Parnassia, is now (again) regarded as the type genus of its own family; insert "PARNASSIACEAE" before PASSIFLORACEAE, transfer P. grandifolia to the latter, and delete "SAXIFRAGACEAE."
Page 203, under SCROPHULARIACEAE: the genera Agalinis, Aureolaria, Buchnera, Castilleja, Pedicularis, and Seymeria are now regarded as members of the Orobanchaceae (page 202); make the necessary transfers. Leucospora, Mecardonia, Penstemon, Veronica, and Veronicastrum are now placed in the Plantaginaceae (page 202); make these transfers as well, and delete "SCROPHULARIACEAE."
Page 203, under ULMACEAE: Celtis L. is probably not a member of the Ulmaceae, but I defer placing it in another family (e.g. Celtidaceae, Cannabaceae) pending clarification from additional studies.
Page 203, under VERBENACEAE: Callicarpa L. may better be regarded as a member of the Lamiaceae, but I defer making this transfer pending further studies (which may support a "Callicarpaceae").
Page 203, under XYRIDACEAE: after "Xyris tennesseensis Kral" insert "[X. spathifolia Kral & Moffett]" (described in J. Bot. Res. Inst. Texas 3:469–478. 2009 and available at this writing for download at http://www.brit.org/fileadmin/Publications/JBotResInstTexas_3_2/469-478_KralMoffett_Xyris_JBRIT3_2__01.pdf). Unlike the other Ketona Glade endemics, this entity is known to date from a solitary occurrence, where it grows in association with the closely related X. tennesseensis. Until an additional natural occurrence is discovered, it seems best regarded as a mutant derived in situ from its intimate associate. Although multiple distinguishing features are cited, these are of a kind to suggest pleiotropy (a single developmental gene affecting several distinct and seemingly unrelated phenotypic characteristics). Sequencing the two xyrids would be helpful in testing this hypothesis. In the meantime, I hope that the publication results in site protection and conservation management, as well as other ex situ cultivation efforts in addition to those mentioned in the paper, so that the mystery of the evolution of "Xyris spathifolia" can be unraveled before this interesting phenotype becomes extinct.
Page 205, bibliographic entry beginning "USDA": the PLANTS Database has since changed its URL to http://plants.usda.gov/.
As more is learned about the flora of the Ketona Dolomite Glades (or additional errata detected in the Castanea paper) this page will be updated.
Most recent update: September 30, 2012