Gentians at their Geographic Fringe

Panhandle naturalists may have a lopsided view of the Gentians, and for good reason. Our local flora features fewer than twenty herbaceous species, the most common being twelve lovely Sabatia we encounter most often. Four rarely-seen but strikingly handsome Gentiana and three slight and evanescent Bartonia round out native representation of Gentians here along the Apalachicola River.

Sabatia decandra

In addition to the genera in our immediate area, the Florida flora includes three others: Obolaria (one species reported from Jefferson County), Eustoma (one species reported from coastal Peninsular counties (and Escambia County), and Voyria, the Ghostplant, Florida’s only representative of this genus of achlorophyllous (lacking chlorophyll) plants native to Tropical America.  You’ll not see Voyria in the Panhandle; it’s reported solely from the most Southern areas, Miami-Dade and the Keys. Regionally, the Gentians are a nice, approachable plant group, easy to get your mind around.

In the greater schema, however, the Gentian family isn’t so straightforward, including over 100 genera and more than 1600 species.  The type genus, Gentiana, so unassuming locally, encompasses nearly 350 species across its natural range, spanning the entire Northern Hemisphere and temperate zones in South America. Like our native plants, most family members are herbaceous though botanists in Central and South America may be more familiar with Gentianaceae through the common and attractive woody shrub Lisianthius (not Lisianthus.)

Keys to Gentians in Flora of Florida begin with a couplet that segregates plants with reduced, scale-like leaves from those with normal green leaves. That pulls out Voyria, Bartonia, and Obolaria, of which Bartonia is the only plant you’ll likely encounter in the Apalachicola Flora, leaving Gentiana and Sabatia as the “normal” seeming plants to identify and study locally.

When in bud or flower, distinguishing Gentiana from Sabatia is simple. Gentiana produces a conspicuously cylindrical, somewhat inflated flower with petal lobes shorter than the tube – a form fairly easy to identify. This contrasts with Sabatia, in which petals appear totally separate, though they are united basally as a short, practically imperceptible tube.

Gentiana pennelliana (above) flowers (as with other Gentiana, develop a cylindrical corolla tube with lobes no longer than the tube. Sabatia stellaris is typical of that genus, with a short, barely perceptible tube with conspicuous lobes.

There are taxonomically useful similarities.  The plants are all herbaceous, and are typically glabrous (no conspicuous hairs except Gentiana villosa). Sabatia and Gentiana have simple, opposite cauline leaves (leaves born along the stem) that lack petioles and stipules (leaves are neither petiolate nor stipulate.) None bear huge leaves; descriptions give 12 cm (about 5”) as the longest you might encounter. Bartonia, which has very small, scale-like leaves is the sole genus in which leaves might be alternately arranged.  Sabatia and Bartonia stems are upright, while Gentiana stems may be lax or sprawling. 

Stamens number the same as sepals and petals (petal lobes), alternating with petals in attachment.  Sepals, petals, and stamens are four (4-merous) in Bartonia, five (5-merous) in Gentiana, as well as five for most species of Sabatia, though a few have a variable number, from seven to fourteen parts in each whorl (7-14-merous).  Flower colors range from white to yellowish for Bartonia, white into the blues for Gentiana, and white to deep pink for Sabatia.  None bear strong yellow, orange, or red flowers, though many European Gentiana as well as the tropical Lisianthius are noted for bright yellow colors.  Corollas are rotate, i.e. radially symmetrical (also termed actinomorphic). All have a simple, superior pistil that is 2-carpellate and typically uniloculate (one chamber made of two carpels) with parietal placentation.  Consistently, the pistil bears a single style with two characteristic stigmatic lobes (very conspicuous in Sabatia.)

Compared to other families in their Order (Gentianales), Gentians lack milky sap, which we find in Apocynaceae and some Rubiaceae.  Our Gentians are not vining, nor woody, nor yellow-flowered like the Yellow Jessamines, Gelsemium (Gelsemiaceae). And the Gentians of the Apalachicola region do not bear flowers in curled inflorescences called scorpoid cymes as do the local Loganiaceae (Mitreola and Spigelia). Moreover, Gentians should not be easily confused with Rubiaceous plants which have conspicuous stipules as well as inferior fruit. 

Apart from other members of the Gentianales, one could imagine similarity to the herbaceous Hypericum gentianoides and H. mutilum, which are readily distinguished in having yellow petals as well as free stamens. Of our Gentians, Sabatia flowers are the most normal-seeming, to the point of challenging the novice field naturalist to determine the correct plant family. Various Caryophyllaceae (herbs that also have opposite leaves) can seem very similar, but Caryophylls will often have swollen nodes, produce flowers in tight clusters, bear separate (free), deeply bifid “petals” (which morphologically are considered petal-like anther), and produce free functional stamens. Many Caryophylls are pubescent whereas Sabatias are typically glabrous.  Regardless, you’ll not be alone in mistaking Gentians for Caryophyllaceae; Linnaeus included the plant we know as Bartonia virginica in the European Caryophyllaceous genus Sagina.

A colony of Sabatia decandra

The birth of Gentianaceae as a recognized family dates to 1789, with Antoine Laurent de Jussieu’s Genera Plantarum, in which he describes “Ordo XIII, Gentinae, Les Gentianes,”  including thirteen allied genera.  Jussieu’s publication culminated emerging thoughts among France’s botanists that plants evidence “natural” similarities that should be reflected in the manner in which they are organized, in botanical terms, how they are “systematized.”  Today, this concept feels normal, but in the 18th century, it was novel.

Jussieu’s treatment of the Gentians (in Latin), 1789

In fact, just a few decades earlier, Linnaeus, who dominated plant taxonomy for much of the 18th century, had devised a very different system of organization based on sexual structures, with the numbers and arrangement of stamens and pistils generating a matrix of convenient pigeonholes for genera. Plants with four anthers would be in one class, Tetrandria (four males); those with five anthers in the next, Pentandria (five males). In the first edition of Species Plantarum (1753), Linnaeus included most plants Jussieu would later treat as Gentians, but relying on numbers of anthers meant he organized them differently.  Exacum is found on page 112, with plants showing 4 stamens and 1 pistil.  Bartonia (classified in his genus Sagina) was categorized as having four stamens and four carpels.  Spigelia, Ophiorhiza, and Chironia (Linnaeus’s name for Sabatia) fall out together on pages 149, 150, and 189, as having 5 stamens and 1 pistil.  Linnaeus lists the other plants Jussieu recognized as Gentianaceae, Swertia and Gentiana, on pages 226 and 229, among plants with 5 stamens and 2-carpellate pistils (showing two distinct styles).  His system, though clearly objective, suffered from using a simplistic formula to negotiate fixed assets and was soon abandoned. Linnaeus’s truly potent legacy has been the rigor of applying binomials (the paired genus and specific name) to every plant (as well as each animal), his 1753 application constituting the basis for plant names we use today.

Frasera (formerly Swertia) in Missouri, a 4-merous flower

Imagining a more natural arrangement did not mean abandoning ancient practice altogether. Researchers in France remained faithful to the basic philosophical assumption that plants appeared at one moment, on the third day of Creation. That commanded plant kinds to be static in form and of a finite number, gifted to humans to be named and used for our benefit.  Enlightenment and the idea of a clockwork universe meant that organisms didn’t materialize helter skelter. There was a plan under which plants were created, a plan we would understand at some future time when all plants have been named and cataloged. Systematists accepted a pious aspect to plant study. Their challenge, to reconcile on-going discovery of new kinds of plants, touched on the quest for devine purpose. 

Linnaeus saw the plan through a matrix of sexual parts in flowers; the French saw different kinds of plants as being links in a chain of biodiversity, a “Chain” of life. A certain number of links, kinds or species, would be a Genus.  Some wondered whether there was a perfect number that might constitute that leg of chain.  With a large enough shackle of chain you’d connect several genera as a Family (or Order), a “natural” order divined through examining plant characters link one element to the next.  Jussieu listed thirteen genera in his Order Gentianae (in the sequence shown), most of which remain part of the family today:

  • Gentiana
  • Vohiria – Voyria
  • Coutoubea
  • Swertia
  • Chlora – Blackstonia
  • Exacum
  • Lisianthus – Lisianthius
  • Tachia
  • Chironia – (embraced plants we call Sabatia today)
  • Nigrina – Chloranthus (Chloranthaceae)
  • Spigelia – Loganiaceae
  • Ophiorrhiza – Rubiaceae
  • Potalia 

In the next century, as plant discoveries and descriptions burgeoned, especially given intermediate sorts and extinct fossilized forms, it became apparent the chain of life needed branches and even looped back on itself (reticulated).  For Jussieu, however, the ruling dictum for all beings remained: Natura non facit saltus, that is, “the body, or chain of life must be linear and complete,”  Linear thinking, characterizing the spirit of the times, was at its most insistent. In the 19th century, that model would yield to a more contemporary construct – the tree of life, culminating in Darwinism.

Jussieu’s System: Unearthing rules governing the chain of life promised revelation of botany’s equivalent to the theory of relativity.  Jussieu’s system included a stab at this with a method of higher classification, the most resilient fallout of which was segregating Monocots from Dicots.  Still and yet, a glimpse at his system is worth a moment’s consideration.  His Class I consists of plants with no Cotyledons, the Fungi, Algae, Mosses, Ferns, etc..  Classes II, III, and IV comprise the Monocotyledons, with various staminal structures.  Dicotyledonous plants were considered as Apetalous (Classes V, VI, and VII), Monopetalous (those with tubular corollas) constituting Classes VIII, IX, X, XI), and Polypetalous (plants with clearly separate petals) being Classes XII, XIII, and XIV.  Class XV was a true catchall, grouping plants with irregular flowers and fruit, ranging from Euphorbs, to Cucurbits, Nettles, Aments, and the Conifers.  The important, lasting contribution of Genera plantarum is the natural approach to grouping Genera as Families (which he called Orders), one of many propositions that set the stage for 19th Century discussion leading to evolutionary theory and 20th Century research that would explain the genetic basis of life. 

Jussieu wasn’t the first in France to publicize these concepts.  His Uncle, Bernard de Jussieu had been active in discussing a natural system three decades earlier, though it was something he never published.  Eager to give his Uncle credit, Antoine de Jussieu includes a summary of Bernard’s 1859 system in Genera, perhaps most pointedly to establish precedent over the work of Michel Adanson, whose Familles des plantes, published a quarter century earlier, in 1763, walked the same path and was prescient in many regards. 

Adanson’s Familles was an enormous undertaking, a tour de force of typesetting necessary to chart his thoughts about relationships of the world’s plants. In one modest segment, Adanson makes it clear he was aware of relationships at the core of the Gentians.  He had  aggregated five of the same genera treated 25 years later as Gentianaceae by Jussieu, along with three new genera (including Sabatia), to form a meaningful grouping which he explained was to be tucked into his Family 23, the Apocynaceae.  As mentioned earlier, Apocynaceae is still regarded as Gentianales, so Adanson was onto something. But he was also a bit maniacal, derided by noted botanist Julius von Sachs.  It’s easy to see how Sachs considers Familles a train wreck, clearly animated by Adanson’s desire to gain credit for moving beyond Linnaeus. But in the frenzy of Familles there are solid constructs.

Knowledge of many plants we include in the family, of course, precedes the work of Adanson and Jussieu.  The generic name Gentiana is credited technically to Tournefort, as the source accepted and published in Linnaeus’s Species plantarum.  Examining Tournefort’s  Institutiones rei herbariae (1719) we see he lists 20 different polysyllabic names for these plants, drawn almost exclusively from Caspar Bauhin’s Pinax, who noted that Gentians were known to Dioscorides, a famed physician believed to have authored de Materia Medica around 70 AD.  Gentians were named well before Tournefort therefore. Plant knowledge which had been limited, become instantly more available with the rise of printing in Europe (i.e. 1455, Gutenberg). Movable type gave the first herbalists opportunity to publish renditions of Dioscorides, bringing ancient knowledge to rapidly expanding audiences. Fuchs and others interpreted Gentians as highly valued medicinals, tracing their use and name to Gentius, said to be a tribal king of early Indo-European Illyrian peoples who treated people with these herbs.

Illustration of a European Gentiana lutea, from Fuchs, 1542. Note drawings in herbals often depict roots, which were believed to be important sources of medicines. (Univ. of Cambridge Royal Library digital library)
Fuch’s treatment of the yellow-flowered European Gentiana lutea, 1542

Globally, our simplest understanding regarding the number of plant species on Earth has expanded from 8500 in the 18th century to over 300,000. During the same spell of time, knowledge of the Gentian family burgeoned from less than 100 named species to over 1600 today.  The fascinating reality is that with so much published information and so many names of regional plants well-established, there is still more to be learned about the diversity, life histories, and ecological interactions of Apalachicola’s Gentians – plenty of room for lay naturalists to contribute to that body of knowledge. And our Gentians are a delight to study..

And, of course, much has been written about plants in our flora since 1800.  Many post-Linnaean specialists interested in North American plants, i.e. Michaux, Muhlenberg, Willdenow, Rafinesque, Nuttall, Pursh, Eaton, and Persoon, as well as more recent botanists (Chapman, Gray, Britton, Pennell, Fernald, Wilbur, and Reveal) had input and disagreements regarding circumscription and naming of genera and species we recognize today.  Of genera now included in Gentianaceae, three are to be encountered in the Apalachicola flora – Gentiana, Sabatia, and Bartonia. For anyone who would like to study these plants in the field, their histories might prove of interest or value.

Gentians of Apalachicola: As mentioned earlier, Gentiana encompasses over 300 species, four of which are considered part of our flora, none of which are commonly encountered. Three, Gentiana saponaria, G. villosa, and G. catesbaei (catesbyi) are basically blue-flowered and show overlapping ranges throughout the Southeast, reaching down into the Florida Panhandle (with catesbaei reported from the Peninsula), while the fourth, Gentiana pennelliana is white-flowered and endemic to the Central Panhandle.

Distribution map for Gentiana pennelliana from Weakley, Flora of the Southeastern US

The most common of blue-flowered species in our region seems to be Gentiana catesbaei (altered to catesbyi in Flora of the Southeastern US.) The plant is a modest perennial herb found (rarely) in shady, nearly boggy soils, producing a tight cluster of one to a few flowers, all of which are held erect (as is typical for the genus).

Single flower bud of Gentiana catesbyi
Open flower of Gentiana catesbyi. Note the petal lobes extending much further than the fringed alternatig appendages

It was first described by Thomas Walter in his 1788 Flora, as modestly distinguished from Linnaeus’s Gentiana saponaria. Flora of Florida separates the two species based on catesbaei having leaves more ovate (rather than elliptical) and more pronounced petal lobes.

Walter’s 1788 characterization of Gentiana saponaria and publication of his Gentiana catesbaei (catesbyi)

Apalachicola Flora Endemic: Tredding through wintry pine flatwoods in the Central Panhandle, you will occasionally encounter a large-flowered, white Gentian, nestled face-up amid understory grasses and forbs. That flower, fully two inches across is our Wiregrass Gentian.

Gentiana pennelliana flower, showing expanded stigma and post-mature (empty) anthers. Flowers are protandrous. At anthesis (flowering lasts several days), anthers are loaded with pollen and the stigma has not elongated.

Based on a short description published in 1940 by M.L. Fernald, we recognize Wiregrass Gentian as Gentiana pennelliana, a name commemorating study by Francis Pennell.

A single “clump” of Gentiana pennelliana in pine flatwoods, Wakulla County, Florida. In this plant, ten stems rose from the base, each of which produced a single terminal flower. Examining the base, there were abandoned stems from a previous flowering. By early Spring, shortly after fruiting, the plant was already sending out new stems, each of which would terminate with one (sometimes two) large flowers.

As taxonomic stories go, this one seems tame. There are only a few synonyms, seen here in Kew’s Plants of the World OnLine (POWO):

A quick summary proves otherwise. The currently accepted name is Gentiana pennelliana, but in previous years people considered other genera, breaking American Gentians away from European species. Based on a plant Croom referred to as Gentiana alba (not validly published), the eccentric, enthusiastic, and arrogant Rafinesque described Diploma tenuifolia (1837), a move botanists ignored at the time.

His contemporary, August Grisebach, retained Gentiana as an inclusive group, lumping Wiregrass Gentian in his concept of Gentiana angustifolia var. floridana. Grisebach’s treatment accepted Michaux’s G. angustifolia as valid (now treated as a synonym to Gentiana autumnalis), a standard usage adopted by Chapman in all editions of his Southern Flora.

Chapman’s Gentiana angustifolia (a Michaux name now considered synonymous with G. autumnalis) describes a variation “corolla green without and white within” – perfectly describing Gentiana pennelliana

Griseback soon moved on, abandoning Michaux’s angustifolia he came to realize that epithet belongs to an unrelated Spanish plant. Having studied a Thomas Drummond specimen with white flowers collected in Florida, Grisebach updated his concept, designating Drummond’s Wiregrass Gentian as Gentiana frigida var. drummondii. That was an even worse decision; G. frigida is very different plant native to Middle Europe.

In 1903 J. K. Small published his Flora of the Southeastern US, splitting US Gentians off as Adanson’s Dasystephana (along with Sabatia, one of several genera Adanson introduced in 1763.) Having spent most of his time in Eastern and Peninsular Florida, Small probably never saw Wiregrass Gentian in habitat and thus regarded the plant as natural variation of a wildflower he designated Dasystephana porphyrio (now also relegated to Gentiana autumnalis). It was Pennell, in 1919, who discussed Wiregrass Gention in his series Plants of the Southern United States, arguing for Rafinesque’s treatment of the plant at species status, the result being his combination Dasystephana tenuifolia.

Throughout Small’s epoch, historically conservative Harvard botanists (home of Gray’s Manual) retained the broader Gentiana. Rejecting New York’s splitters while accepting Pennell’s arguments, M. L. Fernald published the binomial Gentiana tenuifolia in 1939. Learning the epithet tenuifolia was no longer available, Fernald coined the new epithet pennelliana (see below.)

More recently, Czeck botanist Josef Halda took exception with the multiplicity of taxa and, in 1995, recognized Linnaeus’s Gentiana autumnalis as including Coastal Plain plants with lax stems and linear leaves, relegating Wiregrass Gentian to varietal status as Gentiana autumnalis var. pennelliana. Halda’s treatment hasn’t been accepted by US botanists, but recognition the Pinebarren Gentian and our Wiregrass Gention are closely related does not seem so far off-base. Growth habits of G. autumnalis and G. pennelliana are nearly identical, however petal color of pennelliana is consistently white as contrasted with the deep blue of autumnalis, and pennelliana blossoms in December, rather than September for autumnalis.

Gentiana autumnalis (above) from dissertation of Ryan Rebozo (2015). Gentiana pennelliana (right)
Fernald’s brief description suggests relationship of Wiregrass Gentian to G. stoneana, which he had recently named.

Verschlimmbesserung, a German term suggesting attempts at improvement that inherently reflect failure, has meaning in Wiregrass Gentian taxonomic history. Had Croom’s observation been accepted and published in 1832, the plant would have held the name Gentiana alba today, and might have been recognized from the very start as a unique endemic to regional pine flatwoods. But Croom was the first, and for nearly a century, the only taxonomist to realize the unique nature of Wiregrass Gentian. Considering A.W. Chapman’s knowledge of the local flora, it is surprising he didn’t describe Wiregrass Gentian as a separate species, relegating the plant most common in Apalachicola’s flora as an un-named variant of G. angustifolia (now G. autumnalis).

Gentiana pennelliana lives parsimoniously, producing a few pairs of linear leaves (71-10 nodes) along willowy stems seldom 24 cm long. Leaning regularly against other plants in the grassy pine understory throughout the region, the foliate stems camouflage growth, which eventually yields a russet-colored terminal flower bud that inflates slowly, opening eventually as a creamy white vase for December’s peak flowering. Stems broken or those nipped off early in the year will branch to send out another flowering axis, but branching otherwise is uncommon.

Because of their dark green, narrow foliage and slight stems, Wiregrass Gentian plants are difficult to spot threading their way through the undergrowth.

Flowers persist several days, often showing mechanical damage or loss due to herbivory. It’s common to site one or just a few scattered blossoms in an area, though David Roddenberry has documented a population of hundreds in a recently-burned acre of open pine canopy.

A Dozen Rose Pinks: In the Apalachicola flora people might encounter any of a dozen species of Sabatia populating flatwoods and wet prairies. Except for Sabatia quadrangula, (published by Wilbur in 1955) these plants have been studied under stable names for a couple of centuries or more. At least four species were already known by the time Linnaeus published Species Plantarum in 1753 (Sabatia angularis, S. campanulata, S. dodecandra, and S. difformis) assigned by Linnaeus to the genera Chironia and Swertia. The genus Sabatia had not been defined – yet.

Examining a Sabatia flower bud you see the distinct nature of the calyx lobes but not the short corolla tube. Note the slightly convolute (twisted) character of petals as they form. Termed “convolute aestivation” this characteristic is shared with other members of the Gentianales, most specifically the Apocynaceae.
In side-view, as the flower opens the short corolla tube becomes obvious. Moreover, one can observe the tubular union of a calyx tube and the basal corolla tube. The character and relative proportions of the tube and calyx become useful features distinguishing species.

A decade later, the remarkable and botanically-driven Michel Adanson (after whom the Baobab, Adansonia, was named) managed to publish the spreadsheet that had been materializing in his minds-eye, a grand multi-dimensional chart in which every plant genus was colocated with others he believed most similar, those groups then matrixed as explication of diversity he observed in the world of plants – an outgrowth of his work with the Jussieu family in Paris. Adanson’s chart included brief and cryptic notes to establish new genera as necessary, one of which was Sabatia.

The two panels show the legend for Adanson’s charting (above) of Sabatia, with delineation of the genus in the lower panel. The catagories move sequentially from Leaves, Flower, Calyx, Corolla, Stamens, Stigma, to Seed. Note his plant showed a 12-merous flower (12, calyx lobes, 12 petals, and 12 stamens), but the stigma of the specimen he examined hadn’t opened, thus he marked it as cylindrical.

Reading the chart above, for his Sabatia we see Adanson examined a specimen with 12-merous flowers (“merous” indicating the number of parts for each flower whorl, 12 sepals, 12 petals, 12 stamens). Adanson indicates the plant forming his foundation for Sabatia was a Gentian Gronovius had published based on Clayton’s collection 120 in Flora Virginica. That would be the entry Linnaeus cited as Chironia dodecandra, eventually designated as the type for Sabatia.

The entry for Gronovius’s Gentiana floribus duodecim petalis…. in Flora Virginica

See the voucher of Clayton’s specimen #120 below

The Clayton specimen 120, held in the Herbarium at London’s Natural History Museum, is the type for Sabatia dodecandra, which is the type species for the genus. You can access images of specimens by clicking on the Data Portal label below.

Though Linnaeus had first catalogued the Clayton Gentian along with African plants as Chironia (in Species Plantarum, misprinting the page as 27, but citing Clayton’s specimen #120), Adanson, who had lived and collected in Africa rejected both Chironia for American plants, especially a plant that differs so greatly in being 12-merous.

We haven’t come to total resolution, however. We are indebted to Frederick Pursh who recognized the significance of Adanson’s cryptic entry and assembled fragments of Sabbatia (his spelling). Though excellent in pulling together species of Sabatia, Pursh operated under less-stringent rules and was free to ignore priorities and complications imbedded in previous works; thus Pursh treated the Rose Gentian as Sabbatia chloroides, utilizing an epithet Michaux had assigned in 1803. It’s a tangled web; here’s a bulleted chronology of the history that lead to the name Sabatia dodecandra, which we have used since 1888:

  • 1734-1739 – Clayton collected a specimen, his #120, the base for publication of a Gentiana in Flora Virginica. Clayton first sent specimens and notes to Mark Catesby, with whom he had a working friendship since meeting Catesby in Virginia. Occupied with his own publication, Catesby forward the material and correspondence to Gronovius, establishing a direct exchange between Clayton and Gronovius. That created a linkage with Linnaeus, who was responsible for taxonomic decisions published by Gronovius in the Flora.
  • 1739 – Gentiana floribus duodecim petalis… Citing Clayton’s notes for his Centaurium minus floribus carneis in undecim vel duodecim segment divisis, Gronovius publishes this treatment in Flora Virginica, utilizing Clayton’s notes and specimens and working with Linnaeus. In the same Flora, Gronovius treated Sabatia difformis as Swertia difformis.
  • 1753 – Chironia dodecandra was Linnaeus’s designation in Species Plantarum, 1753. Linnaeus also included Sabatia angularis and Sabatia campanulata in Chironia
  • 1763 – Adanson established two relevant genera, Chlora and Sabatia, basing his Chlora on Linnaeus’s Gentiana perfoliata (which was based on a 1611 illustration by Paul Renaulme and basing Sabatia on Gronovius’s publication of the Clayton specimen.
  • 1767 – Chlora dodecandra is the binomial assigned for Sabatia dodecandra in Linnaeus’s 1767 Systema Naturae Tomas II (the last edition Linnaeus himself edited, the following final edition being edited by Johann Gmelin). Because the Linnaean system considered numbers and arrangement of anthers as paramount, this treatment ignored Adanson’s establishment of Sabatia and aligned Rose Gentian with yellow-flowered 6-8-merous Eurasian gentians in Adanson’s Chlora, which are classified as Blackstonia today.
  • 1788 – Chironia dodecandra is the listing for Rose Gentian in Walter’s 1788 Flora caroliniana.
  • 1803 – Chironia chloroides – Michaux’s treatment in Flora boreali-Americana reasserts Chironia as the appropriate genus for Rose Gentian, assigning a name that refers to the genus Chlora, where the plant was assigned in the 12th edition of Systema naturae.
  • 1814 – Sabbatia chloroides – In his Flora Americae Septentrionalis (i.e. septentrionales = “northern” as adapted from the Septentrionales, Latin for the seven brightest stars of Ursa Major), Pursh fleshed out Adanson’s genus Sabatia, encompassing five species: S. gracilis, S. stellaris (newly described in his Flora), S. angularis, S. calycosa, S. chloroides (adhering to Michaux’s logic), and S. paniculata (Sabatia difformis)
  • 1888 – Sabbatia dodecandra – Based on priority of publication, Britton, et al made the combination restoring the earliest (and current) specific epithet for Rose Gentian. Rules established after the date of this publication allow for orthographic changes to names (endings affecting proper Latinization) but do not support spelling corrections, thus the reversion to Adanson’s single-b Sabatia.
Sabatia “decandra” – early stage of flowering, with intact anthers and the unopened stigma which appears cylindrical before the two lobes spread apart
Later stage flower, the anthers having fallen from filaments and the stigma fully opened and turned out

That, of course is not the end for the beautiful Rose Gentian, for this plant has a twin, and in our region, that twin, Sabatia decandra, is the more common of the two. This was explained by Thomas Walter in his 1788 Flora, which described a plant (noting he assigned it to the genus Chironia) differing in developing ten petals and producing more linear, succulent leaves. Botanists have accepted the distinction. See below:

The last two entries, no. 5 and no. 6, are simple diagnoses of the distinction Walter draws between Sabatia dodecandra and Sabatia decandra. In the field, Sabatia decandra is quite erect with linear leaves, and (I’m told) a somewhat smaller flower as compared to S. dodecandra.

Our most commonly-encountered Sabatia, however, is the more delicate Sabatia stellaris, found in pine flatwoods but evident in disturbed edges of roadways.

Sabatia stellaris, described by Pursh in 1814, is found in pine flatwoods natively, but this also is the roadside Sabatia, seeding in and thriving along verges in the southern tier of counties. Note the variability in coloration and petal shape.

Closing out discussion of Sabatia, we need to explain Adanson’s source of the genus. In Sabatia , he commemorates Roman botanist Liberato Sabbatini, lettered with a single “b”. A second genus with spelling Sabbatia was published by Conrad Moench in 1794 for a plant in the Lamiaceae, now synonymized to Micromeria.

Sabbatini (1714-1779)managed the botanical garden in Rome, aggregating specimens in herbaria (books) he shared with others. Sabbatini was a major early contributer to Hortus Romanus, a luxurious multi-volume florilegium produced from 1772 to 1784. featuring native and exotic plants.

Bartonia – A Comedy of Errors

Adanson, Michel ( 7 April 1727 – 3 August 1806) – French.   1763 Familles des Plantes. In his Addendum, Adanson listed Menyanthes, Centaurion (Centaurium), Dasystefana, Cicendia, Gentiana, Chironia, Chlora, Sabatia, Tretorrhiza, Exacon (Exacum), Swertia, Ciminalis  to be added as a group to the Apocynaceae.  It would be hard to deny Adanson appreciated this as a natural grouping. Cryptically incorporated were three newly published genera: Sabatia, Cicendia, and Ciminalis.  You’ll see, in the treatment of Sabatia, that many 19th century authors “corrected” spelling of this genus to Sabbatia, properly reflecting Liberato Sabbati’s name.  Moench, however, published (1794) a genus of mints (now considered synonymous to Micromeria) with this second spelling.  

Aublet, Jean Baptiste Christophore Fusée (4 November 1720 – 6 May 1778) – French.  As Aublet’s long-term mentor, Jussieu was quite familiar with Histoire des plantes de la Guiane Françoise, published in 1775. That work culminated Aublet’s productive but unsettled life as a pharmacist and botanist, with stints in Grenada, then in Mauritius, and finally (at 45), three years in French Guiana. The descriptions and drawings in Histoire, brought Aublet lasting botanical fame, documenting over 1200 species in 576 genera, a third of which were new to science. His work contributed significantly to Jussieu’s treatment of the Gentianaceae; four of ten genera in the family were tropical plants described by Aublet: Tachia, Potalia, Voyria (Jussieu mssp. was Vohiria), and Coutoubea.

Britton, Nathaniel Lord, E. E. Sterns, & Justus F. Poggenburg, with Addison Brown, Thomas C. Porter, & Arthur Hollick. A Committee of the Torrey Botanical Club,1888. Preliminary Catalogue of the Anthophyta and Pteridophyta Reported as Growing Spontaneously within One Hundred Miles of New York City, New Yor, April 25th, 1888. Cited variously,

Eaton, Amos (17 May 1776 – 10 May 1842) – New York, USA.  Beginning in 1817, Eaton published his Manual of Botany, which he saw through several editions and title alterations (the 3rd in 1822, 5th in 1829, 6th in 1833, 7th edition in 1836. ) The first edition had been published by students at Williams College, thus the 1818 edition becomes the first commercially-set and distributed text.

Eaton’s biography, related in Wikipedia, tells us so very much about family relationships of earlier years, particularly addressing survival chances of women: “Following the death of his first wife, Polly Thomas, Eaton was remarried to Sally Cady in 1803, who bore him five sons. After her death he was remarried to Anna Bradley in 1816, by whom he had three sons and two daughters. She having died, Eaton married Alice Johnson in 1827, who bore him one son, and survived him about four years. Three of his children showed a preference for natural philosophy. Hezekiah Hulbert Eaton (1809-1832) became a chemist at Transylvania University but died at the age of twenty-three.[17] Major General Amos Beebe Eaton (1806-1877) was a U.S. Army officer interested in natural philosophy. Sara Cady Eaton (1818-1881) taught natural sciences and modern languages in a young woman’s seminary at Monticello, Illinois. Eaton’s grandson, Daniel Cady Eaton (1834-1895), professed botany at Yale College in the 1860s.

Fernald, Merritt Lyndon (5 October 1873 – 22 September 1950) – Maine, USA.  A driving force in Harvard Botany, Fernald published many papers and edited the 7th edition of Gray’s Manual,  Fernald must have been firm in his convictions. My major prof at Vanderbilt, Ben Channell, referencing Fernald’s initials, referred to him as “My Lord Fernald.”   Weatherby, Charles Alfred (1875 – 1949)- Connecticut & Colorado, US.  Curator of Gray Herbarium Harvard.  see: Fernald, M. L. and C. A. Weatherby. 1932. Bartonia; a comedy of errors. Rhodora 34: 164—167. You should look up this paper and take in its smug critique.  

Gillett, John Montague (26 November 1918 – 27 December 2014) – Ottawa, Canada.  His Ph.D. thesis at Washington University (St. Louis) was a monograph of Gentianella. In addition to work on Gentians and a love of floristic field work, Gillett specialized in Trifolium.  Relevant to our flora is his 1959 paper, A Revision of Bartonia and Obolaria (Gentianaceae), Rhodora, 61: 43-62 .

Gilg, Ernest Friedrich (12 January 1867 – 11 Octob434 1933) – German.  Gilg was a curator at Berlin’s Botanical Museum, working with Engler.  Among many diligencies, Gilg specialized in Grasses., with 42 taxa to his credit.  In 1895, his treatment of Gentianaceae  was published in Engler & Prantl, Die natürlichen Pflanzenfamilien 4(2): 50-108.

Hill, John (1716 – 1775) English. In 1756, Hill issued The British Herbal: an History of Plants and Trees, Natives of Britain, Cultivated for use, or Raised for Beauty. Alongside his description of Gentians, he resurrected Centaurian, a genus used for Gentians by earlier writers, but not adopted by Linnaeus. The fascinating aspect of this book is the more natural approach Hill takes in classification. But for the fact that he was totally despised by many educated people might explain the reason he was never credited with his simple scheme.

de Jussieu, Antoine Laurent – (12 April 1748 – 17 September 1836) – French. 1789.  Genera Plantarum.  Jussieu’s book, which by code is the starting point for authority of plant family names, covers his system for natural classification of the world’s known plants, with accepted genera parsed into 100 Orders (Families in contemporary terminology) treated in 453 pages of text.  The Gentianaceae, page 141, is “Order XIII: Gentianae, Les Gentianes, Class VIII: PLANTAE DICOTYLEDONES MONOPETALAE – Corolla Hypogyna.”  Genera included: Gentiana, Vohiria, Coutoubea, Swertia, Chlora, Exacum, Lisianthus Tachia, Chironia Nigrina, Spigelia, Ophiorrhiza, and (with affinity) Potalia.

Linnaeus, Carl – (23 May 1707 – 10 January 1776)  Though authority for binomials in plant taxonomy begins with Linnaeus’s first edition of Species Plantarum (1753), checking Plants of the World On Line (POWO) you’ll note the cited authority gives credit to various earlier authors.  In the case of Gentiana, it was clear Linnaeus adapted his treatment from Tournefort, thus POWO cites authority for Gentiana as Tournefort ex Linnaeus, which means it was a Tournefort name first published validly by Linnaeus. Most American publications omit this credit, and continue to utilize the traditional citation, the “L.”

Michaux, André  (8 March 1746 – 11 October 1802) – French.  Noted for his 1803 Flora Boreali-Ameriana, in which he published the genus Centaurella (Volume 1: 97, 98 with illustrations in t. 12), a taxon considered synonymous to Muhenberg’s Bartonia, which had been published by Willdenow in 1801.  Fernald & Weatherby (1932) discuss the confusion surrounding application of these names.  

Muhlenberg, Gotthilf Heinrich Ernst (Henry)  (17 November 1753 – 23 May 1815) Pennsylvania, USA.  Muhlenberg moved to Germany for his college education, after which he returned to Pennsylvania as a Lutheran Minister, later earning his Doctorate in Divinity from Princeton. Retiring from ministry in 1779. Muhlenberg dedicated himself to botany and was elected to the American Philosophical Society in 1785.  His best-known works are the 1813 Catalogus Plantarum Americae Septentrionalis (Catalog of the Plants of North America) and his 1817 Descriptio Uberiaor Graminum et Plantarum Calamariarum Americae Septentrionalis Indiginarium et Cicurum. (A More Detailed Description of the Grasses and Reeds of North America, Native and Naturalized)  (Sidenote:  the translation programs will interpret Calamariarum as Squid).  In his 1813 Catalog, Muhlenberg classifies Bartonia as Tetrandria Monogynia, noting Michaux’s Centaurella as a synonym and listing two species: B. verna and B. paniculata. Benjamin Smith Barton, however, citing a Rafinesque manuscript, had made the combination Boltonia verna just the year before in his 1812 edited version of Gronovius’s Flora Virginica.  Until recently, citations for Bartonia credited this recognition to Muhlenberg, but more recent publications have taken note. Not surprising, Barton of course would have been keen to keep the record straight on plants in the genus named for him.

Nuttall, Thomas (6 January 1786 – 10 September 1859) – English.  Nuttall, whose first collecting expedition to North America was cut short by the 1812 War, returned to the US in 1815, where , in 1818, he published Genera of North American Plants.  Holding positions at Harvard Botanical Gardens and eventually at Philadelphia’s Natural Academy of Sciences, Nuttall published his 1841 North American Sylva – Trees of North America not described by F. A. Michaux on the eve of his departure to England, where he retired on an inheritance as a country gentlemen.

Pursh, Frederick Traugott  (4 February 1774 – 11 July 1820)  Employed initially by William Hamilton as botanist at Hamilton’s Philadelphia estate, ‘The Woodlands’ (1803-1805,) Pursh began workng for Benjamin Smith Barton on Barton’s dream project, a flora of North America, which gave him access to specimens from the Lewis and Clark Expedition. The flora was never completed, but Pursh, who returned to England in 1813 , published Flora americae septentrionalis(1814); or A Systematic Arrangement and Description of The Plants of North America, a treatment ridiculed by Rafinesque (1818).   Pursh returned to North America, where he collected extensively in Canada, Those collections were destroyed in a fire, and Pursh’s life took a nosedive.  He died in Montreal in 1820.

In the 1814 Flora, Pursh used the Linnaean system, including Sabbatia, genus 171, on pg 137, as Pentandria monogynia and Gentiana, genus 227 on pg 185-187 in Pentandria Digynia. He recognizes Centaurella Michaux on pages 99-100, subsuming Bartonia Muhlenberg.

Persoon, Christian Hendrik – (1 February 1761 – 16 November 1836)  South African who relocated  permanently to Paris in 1802.  A colorful individual, his life and works have been praised:  “Persoon was the first successful mycological systematist”  Journal of Wild Mushrooming.  But he also published, in 1805, his Synopsis plantarum,seu Enchiridium botanicum, complectens enumerationem systematicam specierum hucusque cognitarum, which translates: “Synopsis of plants, or Enchiridium botanicum, comprising a systematic enumeration of the species hitherto known.”  An “Enchiridium” is a handbook, or small manual, Though a bit arcane, Persoon’s Synopsis was also well-known, and early in ratifying publication of Bartonia tenella as the founding taxon of that genus.  

Persoon, though, remained a fascinating character. From Chester R. Benjamin’s ‘Biographical notes on Persoon’, Proceedings Iowa Academy of Science, V60 Article 11:  “He loved with a botanist’s passion both living and dried plants, though his awkward hand tore them in the gardens or broke them in herbariums. Surprised in flagrant offense at a public garden, he was warned several times by the gardener to no avail and he l?st much in the public estimate. However, he possessed an out- standing reputation as author of the first reasonable classification of the fungi, .of the first synopsis treating these organisms, as well as of a manual published in Paris, in which he described more than 22,000 plants and of which the diagnoses are regarded as a model of conciseness and exactitude. A mycology of Europe, a treatment of the edible fungi, and a number of memoirs on various scientific subjects established his reputation on a firm foundation. He was regarded as one of the foremost botanists of Europe and considered, particularly by the Germans, as the “prince of mycologists”.”

Persoon lived his early life in South Africa, and was never priveleged or even well-compensated.  Benjamin relates a description: “According to Fee, Persoon was large, lean, bony, with long legs and long arms attached to an uncomely body; his head large, supported by a small neck and tilted over awkward shoulders; his skin red, scaly, and covered with pale blemishes; his scanty hair grizzled, waving over a forehead wrinkled and bunched; his enormous mouth containing a small number of shaky teeth and always full of saliva, much of which escaped in jets with each word he spoke; his eyes grey and watery, half-opened and almost lost behind prominent, puffy cheeks; his ears enormous and directed forward; and his face wrinkled and without expression. He was coarsely clad. His clothes were bizarre of form and strange of color, worn through by long service. He often lacked many of the most common necessities.” Persoon’ s archive is at Leiden University, but Hunt Institute maintains a copy of the materials.

Rafinesque-Schmaltz, Constantine Samuel (22 October 1783 – 18 September 1840) – French, born near Constantinople.  When discussing history of the North American flora, you’re bound to encounter Rafinesque, on of the botany’s brilliant bad boys.  Fernald and Weatherby (1932) cite comments by Rafinesque in the 1818 American Monthly Magazine and Critical Review (which they abbreviate as Am. Mo. Mag.)  His article 2, January, pg 170, is an extensive  (nearly 6 full, double-column pages), critical — yes, excoriating review of Pursh’s 1814 Flora Americae Septentrionalis, 

Robinson, B. L. (8 November 1864 – 27 July 1935) - Illinois, USA. In 1908, Robinson published ’Notes on the Vascular Plants of the Northeastern United States’ (Rhodora 10: 29-35.) On page 35, he makes the combination Bartonia paniculata, not aware that Muhlenberg had completed the task in 1813.

Sims, John (13 October 1749 – 26 February 1831) – English. One of many Quaker botanists, Sims was the first editor to succeed Curtis at the Botanical Magazine. His contribution to discussion of Gentianaceae revolves around misinformation common to the horticultural trade. The plant he illustrates as a Bartonia in Bot Mag, is an unrelated Mentzellia. Because Sims misinterpreted Bartonia, based seemingly on information from Nuttall and Pursh, he continued to accept the genus Centaurella for the Gentianaceous plants.

From Sims’s treatment in Botanical Magazine, 1812

Sprengel, Kurt Polycarp Joachim ( 3 August 1766 – 15 March 1833) – German. International Plant Names Index (IPNI) lists 5205 names he published. Relevant to Gentianaceae, Sprengle edited the 16th edition of Linnaeus’s Systema Vegetabilium, introducing the genus Andrewsia as a substitute for Bartonia.

de Tournefort, Joseph Pitton (5 June 1656 – 25 Deember 1708) – French.  1719  Josephi Pitton Tournefort Aquisextiensis, doctoris medici Parisiensis … Institutiones rei herbariae. Though bearing a title beginning with his name and qualifications, it’s common to reference this work as  Institutiones rei herbariae. It’s a significant, luxurious 3-volume set, two volumes of which are fine illustrations by Claude Aubriet, showcasing the genera described in Vol 1.  Tournefort describes Gentiana, as a component of “Classis Prima, de Herbis et Suffruticibus, Flore monopetalo, Campaniformi,” further segregated in “Sectio III.  De Herbis flore monopetalo, campaniformi, cujus pistillum abit in fructum siccum seu testam unicapsularem in nonnullis generibus, in aliis verò multicdapsularem.”  Translated, that description tells us: “Of Herbs with a monopetalous (petals united), bell-shaped flower, the pistil of which turns into a dry fruit or a unicapsular shell in some species, but in others multicapsular. ” Other genera Tournefort includes in this section are:  Cerinthe (Boranginaceae), Hydrophyllon (Hydrophyllum, Boraginaceae), Soldanella (Primulaceae), Convolvulus (Convolvulaceae), Tithymalus (Euphorbiaceae), and Oxys (Oxalis, Oxalidaceae).  His treatment, therefore, resembles a modern artificial key, in which plants are grouped based on simple, somewhat easily-detected physical properties of the flowers, with no supposition as to natural affinity.

Weakley, Alan Stuart (b. 1957 contemporary) – Maryland, USA.  Weakley directs the Herbarium at University of North Carolina, and holds a professorship at the Chapel Hill campus.  His evolving Flora of the Southeastern US is available on-line.  2023, Flora of the Southeastern United States Gentianaceae:  Gentiana:  Sabatia:   Bartonia

Willdenow, Carl Ludwig (22 August 1765 – 10 July 1812) – German.  Willdenow died at the age of 46, having been curator of the Berlin Herbarium for several years.  In 1803, he published XXVI Zehn Neue Gattungen von gewâchsen beschrieben von Herrn Professor C. L. Willdenow (Ten new genera of plants described by Professor C. L. Willdenow), in a somewhat obscure journal: Der gesellschaft naturforschender freunde zu Berlin neue schriften  (New Writings of the Society of Natural rRsearch Friends in Berlin).  

Willdenow begins treatment of Bartonia with the following:  “VI.  Bartonia.  Herr Prediger Mühlenberg, der diese Gattung entdeckte, hat sie zum Andenken des Herrn D. Benjamin Smith Barton in Philadelphia benannt.  Sie gehürt in die erste Ordnung der vierten Klasse (Tetrandria Monogynin) nach Scoparia.  Der Bau der Blüthentheile ist folgender.” which translated (Google) states;  “Mr. Preacher Mühlenberg, who discovered this genus, named it in memory of Mr. D. Benjamin Smith Barton of Philadelphia. It belongs to the first order of the fourth class (Tetrandria Monogynin) after Scoparia. The structure of the flower parts is as follows” .

Link to this Page: https://botanyincontext.com/gentianaceae/