Bartonia – A Comedy of Errors

An unusually tight cluster of Bartonia verna

I’m not the first to use this title for a short essay.  Fernald and Weatherby, in a brief 1932 paper published in Rhodora, selected this wording to signal the complicated taxonomic history of a small and distinctive genus first described as Bartonia in 1801.  We’ll get to that history, but first, something about the plant itself.

In the Apalachicola Flora, one might encounter three different species of Bartonia, though I’ve only met the more conspicuous B. verna.  All are very slight annuals.  By slight, I mean wispy or thin and grassy, though the term frail doesn’t apply, at least to plants of B. verna.  It’s curiously robust for such a modest plant.  

My first encounter was in late January, investigating boggy pine flatwoods of Apalachicola National Forest with David Roddenberry. Scattered through the groundcover of grasses we spotted starry white flowers, a half-inch across, floating 5-10 inches above the ground as though generated by the grasses themselves.  My first inclination was to follow the narrow stems to their bases, imagining I’d find some rosette of leaves, because the stem and flowers had the gestalt of a sundew.  But the flower, with 4 petals and 4 stamens, wasn’t right (though there is a sundew native to Australia that has 4-merous flowers); neither was the inflorescence.  And there were no leaves at the base. 

With just a handlens, you can make out a few bractlike leaves along the reddish stem, but it seemed improbable that this little foliage could support a plant with such relatively-large flowers.  There had to be a base in the soil, some evidence of foliage gone dormant. But no, there’s neither a sturdy underground root nor shriveled rosette.  

The only book we had at the moment was the 2011 Wunderlin & Hansen Guide.  Gathering my thoughts as to who this plant might be, I was contemplating the dreaded walk through family keys.  Here was an erect herb, with scale-like leaves, and simple flowers with 4 sepals, 4 white petals, 4 stamens, and a single pistil that might be 2-locular (based on what appeared to be a 2-parted stigma).  Thankfully, David recalled this would be Bartonia, a plant he had encountered years before.  Checking descriptions, and later investigating possibilities with the ISB website (Florida Plant Atlas) and the full treatment in Flora of Florida (Vol 5, Wunderlin, Hansen, & Franck, 2018), identification as Bartonia verna was straightforward. 

Of course, having the right name is the handle connecting a plant to the world of published information.  This is an annual, which is somewhat amazing.  It’s not a saprophyte.  The stem and leaves are all it has, photosynthetically. There’s very little means of support.  If it weren’t for the tangle of grasses through which it weaves, I’m not certain there’s enough substance in the stem to keep a plant upright.  And how does it get around (disperse itself)?  The plant is common locally; we found it throughout the area.  Each year there has to be enough seed crop from tiny drying capsules for the plant to salt and pepper its way through the landscape.  Something of a delightful miracle.

So what about the other two plants.  The keys tell me to expect Bartonia virginica and B. paniculata also, segregating my plant, B. verna, as having the larger petals, from 5 to 10 mm long, while the other two have shorter petals.  Moreover, Bartonia verna blooms in winter through spring, while the other two flower in warm weather.  So expect the other two plants to show up later in the year, but to be less easily spotted. 

Studying the key to species, distinguishing Bartonia virginica from B. paniculata requires comparing inflorescences (narrow versus spreading), leaf placement, and anther shape.  Bartonia virginica anthers form mucronate tips (suggesting a small bump or tit at the tip of the anther).  Summer is ahead, so I’m hoping to have a chance to see these plants and characters for myself.

Bartonia verna, showing the two stigmatic lobes and 4 anthers

As to Bartonia verna, which is flowering right now (February), though it is small and simple, that should be interpreted as wand-like, almost wirey.  I captured two for closer examination with a dissection scope (and yes, I do have a collecting permit issued by officials of Apalachicola National Forest, for which I am very grateful).  Setting the whole plants, with their very modest root base, in a deep vase of water (actually a recycled soda bottle), I discovered they will stay perky for days.  Individual flowers last for more than a week, and existing buds continue to develop, opening for the full several days also.   

Searching literature on the internet (outside the paywalls), I don’t find much about life history or autecology.  No wonder.  These are tiny plants that go through their life cycles with little public notice.  Just examine images of herbarium specimens in the ISB website – an Emily Dickinson kind of presence, resolute and unresolved.

Taxonomically, it’s a more complex story.  A lot of taxonomists have taken notice of these plants, almost all descriptions and decisions based completely on nearly-leafless specimens like the ones you can view in ISB.  The simplest way to explain the convoluted history is through a timeline:  Below are critical events in nomenclature of Bartonia; complete citations and additional notes are clustered at the end of the Gentianaceae family page.

1753 – Carl Linnaeus, citing Gronovius’s manuscript for  Flora Virginica, included Sagina virginica as the first valid publication of what is, today, a Bartonia.  The genus Sagina, typified by Sagina procumbens, remains valid, and is classified as Caryophyllaceae, thus our first reckoning of Bartonia has it indexed by Linnaeus in his class and order Tetrandria Tretragynia (4 stamens, 4 carpels), in a Caryophyllaceous genus.

1756 – Hill published, validly, the genus Centaurium, resurrecting a name many pre-Linnaean authors used for Gentians. This name is one of many that has been applied to Bartonia. It’s worth noting that Hill is one of few authors who ignored Linnaeus’s sexual schema for classifying plants.

1801 – Carl Willdenow, curator of the Berlin Herbarium, published a list of 10 genera described by his correspondent Henry Muhlenberg, a native-born American and retired clergyman working in Pennsylvania.  Among Muhlenberg’s new genera was Bartonia, named for then famous American botanist Benjamin Smith Barton.  The species name Bartonia tenella is part of that description.  

Description of Bartonia from Muhlenberg ex Willdenow

1803 – André Michaux, in his Flora Boreali-Americana, published the genus Centaurella, describing two species, C. verna and C. paniculata.  He illustrated both, classifying them (using the Linnaean system) as Tetrandria Monogynia.  These appear to be the first published illustrations of Bartonia.

Tab 12, Centaurella, from the Michaux Flora

1805 – Christian Persoon, a South African newly arrived in Paris, must have gained access to a wide scope of literature and specimens.  Though famous for his specialization in fungi, in this first year in Europe, Persoon published Synopsis plantarum, covering the plant kingdom.  Not only did he ratify the Muhlenberg ex Willdenow Bartonia (his genus # 290), he also accepted Michaux’s species of Centaurella.  Aware, perhaps, of the segregate genus Centaurium that Hill created in 1756, and perhaps also not approving the word Michaux selected, Persoon adopted the genus Centaurium, thus creating the combinations Centaurium vernum and C. autumnale (renaming Centaurella paniculata).  Recall that at this moment, it’s likely only three botanists had seen plants in nature: Clayton (whose specimens Gronovius co-opted), Muhlenberg (who lived and collected in the Eastern US), and MIchaux (who explored and studied plants in the US for over a decade, from 1785 to 1796).

1812 – John Sims, the first editor (after Curtis) of Botanical Magazine, introduced a beautiful drawing of a Mentzelia (Loasaceae), describing it as a new, segregate genus Bartonia, with one species, Bartonia decapetala. The drawing and name were based on horticultural plants said to have been introduced by Nuttall, supported with information conveyed by Pursh.  (We’ll run into Pursh and Rafinesque’s opinion of his work a bit later).  Having incorrectly determined Sims’s use of Bartonia to be valid, Nuttall retained Michaux’s Centaurella in his 1818 Genera of North American Plants as the proper name for the small herbs we know as Bartonia today. Learning this, I finally understood the reason one ends up in Mentzelia when querying Flora North America for Bartonia.

From Curtis’s Botanical Magazine, 1812. The plant Sims named Bartonia

1812 – Benjamin Smith Barton published his update of Gronovius’s Flora Virginica, in which he cited a Rafinesque manuscript that established the combination Bartonia verna.

1813 – Henry Muhlenberg published his Catalog plantarum…, including the combinations Bartonia verna and B. paniculata.  This publication was accepted as citation for B. verna, only recently superseded by the 1812 publication in Barton. Oddly, subsequent workers did not uniformly note the associated transfer for Bartonia paniculata.

1814 – Pursh also retained Centaurella in his Flora Americae-Septentrionalis, utilizing the Persoon specific epithet autumnalis without citation.

1818 – Constantine Rafinesque published his 5 page excoriation of Pursh’s Flora, insisting correctly that the genus Bartonia should be used for these plants.  In the same article, he casually introduced Nuttallia (Nuttallae sic) as a proposed replacement for Sims’s plant. The Rafinesque diatribe merits a read. One takeaway is that Rafinesque, a devoted Linnaean, had by that time abandoned Linnaeus’s system of classification, and was questioning authors who continued to group plants based on such simple reasoning: “These plants are arranged according to the sexual system of Linneus, with some trifling alterations: strange as it may seem, nothwithstanding the superstitious veneration which the disciples of Linneus entertain for the most trivial part of the labours of that great man, almost every one of them endeavour to alter or mend that falling system: they may be compared to masons endeavouring to sustain, by patch-work, an old building erected by an able architect with bad materials, and now falling to ruins… Let us hope that the labours of Jussieu, Decandolle, Brown, and Rafinesque, will soon supersede those wretched attempts.”

1822 – Amos Eaton, in a series of editions of his Manual, recognized Bartonia as the proper name for the genus.  From the Fernald & Weatherby text, Eaton is quoted as writing: “It is thought best to retain this name, until the fancies of our verbifacient botanists shall become so nearly stationary, that one or two changes more may settle upon this little plant a permanent name.“  A few years later, we learn also from Fernald & Weatherby, Eaton rechristened the Sims version of Bartonia, authoring a fourth genus by the name Torreya.

1825 – Kurt Sprengel, in editing the 16th edition of Linnaeus’s Systema Vegetabilium, described the genus Andrewsia, with two species, A. verna and A. autumnale.  Perhaps he was responding to the Rafinesque complaint that neither Michaux’s Centurella, nor Persoon’s (i.e. Hill’s) Centaurium would suit Linnaeus, as the rules he established for generic names prohibit basing the name of one genus on that of another.  Or, having seen the chaos, he decided to craft a new name, which indeed must have been his go-to solution.  As you’ll see in the notes, Sprengel is credited with having published 5,205 plant names.

1888 – Working as a committee of the Torrey Botanical Club to generate a catalog of plants growing spontaneously within 100 miles of New York City, Nathaniel Lord Britton, E. E. Sterns. Justus Poggenburg, with. Addison Brown, Thomas Porter. and Arthur Hollick published the combination Bartonia virginica, thus bringing the plant credited to Linnaeus, Sagina virginica, into the genus, and setting the original Bartonia tenella into synonymy.

1903 – John Kunkel Small, in his Flora of the Southeastern United States, treated three Bartonia species, including his new Bartonia lanceolata, while not citing B, paniculata. Just a few years later (1908), Benjamin Robinson published the combination Bartonia paniculata (based on Michaux’s Centaurella paniculata), as synonymous to Small’s new species, and bearing priority.  Fernald and Weatherby (1932) note the much earlier publication of the Bartonia paniculata combination by Muhlenberg was commonly missed, perhaps because it failed to make Index Kewensis. Though taking us off the track for a moment, I’m making note of the fact that Small was a fixture at New York Botanic Garden, while Robinson was (at the time) Curator of Harvard’s Gray Herbarium. This was not the only time the Harvard and NYBG teams sparred, or simply corrected one another. Fernald and Weatherby, of course were Harvard. Britton and Brown were NYBG. 

1932 – M. L. Fernald and Charles Weatherby published the article from which this essay takes its name:  ‘Bartonia: A Comedy of Errors’.  Both taxonomists were keen on detail, but even they missed the 1812 publication by Benjamin Smith Barton, reporting Rafinesque’s transfer of Centaurella to synonymy with Bartonia.

We are, after these many notes, back at the beginning.  There are, currently, three species of Bartonia reported from the Panhandle, Bartonia verna, B. virginica, and B. paniculata.  

For specialists, the trail of evidence documented here is immortalized in nearly cryptic annotations, such as the author citation and heavily-italicized complex synonymies with their abbreviated sources.  I know that in simply attempting to identify and learn about a plant, I usually blur over those notes.  But in this Flora I’m taking time to examine the histories, hoping that by documenting the stories behind plants we encounter I can construct a context for the manner in which those plants are circumscribed and understood.

Reflecting on this sequence, it’s clear that any taxonomist would benefit from a photographic memory.  The descriptive information is challenging on its own, but overlay that with the archival aspects of nomenclature, the tracking down and comparing what others have published, and it becomes clear plant nomenclature is a massive, detail-oriented challenge that burgeons over the decades.  That of course explains the need for specialists – someone has to be familiar with the details, as well as generalists – someone has to have a grasp on the overall picture, at a floristic and ecological level.

In the case of Bartonia, there is yet plenty to learn, at least for me.  I still hope to encounter and present the other representatives (two similar species) that should be in our area.  And I’ve seen very little that would tell me more about how these plants are pollinated, mature and disperse seed, or germinate and establish a new population each year.  I’ve not discovered much information in writing, but someone who reads this might know of a not-readily indexed publication, or even an unpublished thesis that fills in these gaps.   Let me know.

Best regards, Jim.