Platanthera – On the Fringes

Warning: Details of floral structure in orchids can become complex, tedious, and convoluted. For that reason, I’ll simplify names at moments in which using the binomial or any of the common names would be cumbersome, creating references for each Platanthera recorded for the Apalachicola flora by utilizing specific epithets as nouns: Ciliaris substitutes for Platanthera ciliaris, Blephariglottis for P. blephariglottis var. conspicua, Cristata for P. cristata, Chapmanii for P. chapmanii, Integra for P. integra, Nivea for P. nivea, Flava for P. flava, and Clavellata for P. clavellata.

Platanthera blephariglottis
Platanthera chapmanii

The Main Story

Our native fringed and fringeless orchids are masters of camouflage, a subterfuge made easier since several months of each year they live totally underground.  And over the two months during which plants send up young stems and foliage, they are incredibly difficult to distinguish among grasses and other herbs. For a search image, imagine stiffly-upright terrestrial herbs, softly vegetative, with glabrous and modestly glaucous stems, and sheathing strap-like leaves we associate with monocots.

Platanthera cristata inflorescences showing color and a single open flower

With experience, one can pick out an emerging stem, but they are difficult to spot until the buds show color and the flowers begin to open. Then, for 2-3+ weeks, individuals of most species are hard to miss, and reasonably easy to identify glowing in yellow and orange, gleaming in green, or shining in white. With flowers faded, the fruiting stems blend back into surrounding vegetation.  At maturity they become simply more dry, brown stems, shattering with others. 

No surprise that any key to identification of Platanthera demands access to flowers and understanding the structure of those flowers. Borne in terminal spikes, we are reminded that technically the flowers are sessile; what appears to be a pedicel is actually the inferior ovary.  Platantheras become more easily recognizable as the knobby buds develop, each showing an emerging nectary parallel to the ovary, eventually revealing hints of color.  

Describing fully-developed flowers for the entire group demands interpolating flower structure for the distinct Snowy Orchid (Platanthera nivea), which produces “nonresupinate” flowers; that is, their flowers develop with the labellum uppermost.  Developmentally, all of the Platanthera flowers begin like P. nivea (with the labellum uppermost), but those that are not Nivea, each flower develops a twist in the ovary that positions the lip (the labellum) lowermost, offering the labellum as a landing pad for pollinators (except for hovering moths).   Most orchids achieve this repositioning ( “resupination”) in one way or another,

Platanthera cristata flower – the two vertical lines are the pollinial chambers. The structures that look like a pair of “eyes” are the auricles

Each Platanthera flower has 3 somewhat similar sepals and 3 petals. The two lateral petals mirror each other while the third forms the lip. For the resupinate species, the “dorsal” sepal is uppermost, flanked by the lateral petals and forming a hood over the column while the two lateral sepals fold back, out of the way. In the Fringed Orchids the labellum is largest and most conspicuous segment, triangular to strap-shaped, with various degrees of fringing. (see Platanthera ciliaris flower, below)

Platanthera ciliaris, showing spread between viscidia

The other four species (Nivea, Integra, Flava, and Clavellata) have triangular to strap-shaped, entire labellae. In every species, the labellum generates a basal spur (nectary) projecting back along the ovary.  As is generally true for orchids, the column faces the labellum.  (You may hear a botanist say the column is opposite the lip, but “opposite” has a specific, very different technical meaning in flowers.) 

In describing orchids, the term “column” has to be explained. It isn’t so easily understood unless a person has had the opportunity to inspect and dissect a lot of flowers.  Since blossoms of the larger, epiphytic tropical orchids (such as Cattleya, Cymbidium, or even Epidendrum) are more common in horticulture, many people will have examined the column in one or more of these plants.  It’s a succulent (cartilaginous in texture) structure formed seamlessly as a union of a single anther (yes, a stamen) with style and stigma.  The anther generates masses of pollen in miniature specialized structures that, as a whole, are called pollinaria.  The Platanthera column, however, is quite different.  The name Platanthera helps – this is the flat anther club.  The column we encounter in these terrestrials (and their relatives in the Orchis subfamily) is not similar to the nose-like column peeking out from a Cattleya corsage.  In Platanthera, the two anther chambers (locules) are paired, distinct, each chamber projecting as a separate planar structure. In form, the two chambers have the shape of  pair of stubby wings, the connecting body housing the stigmatic surface.

Platanthera ciliaris, with pollinium teased out, but remaining attached at the viscidium

Each anther locule sequesters an exquisitely structured pollinium (this term applies when the anther locales are separated and two distinct pollen-bearing structures form).  Each Platanthera pollinium is a club-like structure with a sticky pad (viscidium) formed at the base of a stalk (stipe) that enlarges as a mass of mealy pollen.  The viscidium adheres to any smooth surface, a  needle, a straw, or  more naturally, the proboscis or compound eye of a pollinating insect. 

Pollinium of Platanthera ciliaris, self-adhering to a penny

If you have a host of Ciliaris or Cristata flowers available, touch a probe to the minuscule shiny pad at the tip of an anther wing (I also use the word stirrup) of a fresh flower, and you will extract a pollinium.  Then touch the pollen mass to the stigmatic surface and you will have helped with self-pollination (Linnaeus would have used more suggestive wording).  Orchids generally are self-fertile, so you may have just parented a couple of thousand seed.

Of course the insect visitor is drawn and guided to a reward, which for Platantheras is nectar produced in the spur.  Structural features of the flower direct insect behavior, ensuring pollinia can be properly attached to the visitor so as to be positioned for transfer and contact with the stigma of another flower.  The structures and sequence are called the “pollination mechanism” and are directly relevant to species barriers. Plants of an interbreeding population share the same pollination mechanism, and (theoretically) plants of each species, though scattered in sites across the range, retain the potential to interbreed through that mechanism.

For orchids most particularly, this suggests that understanding how floral structure impacts pollination will provide relevant characters tied directly to the reproductive biology, and thus the identity of a species.  Orchid flowers have stories to tell, Platantheras being no exception.

So what are the characters we use to distinguish the eight Platantheras recorded for the Apalachicola flora?  To me, these plants are readily identified in three groupings:

  • Flowers yellow or white with the labellum lowermost and the spur curving down from the base of the lip , variously fringed – Platanthera ciliaris, P. blephariglottis, P. chapmanii, and P. cristata.   (Outside the Panhandle, additional species have been described.)  Because it is yellow, Platanthera integra keys out with the Fringed Orchids, but it is distinct, by many authors aligned with Platanthera nivea, often in another genus.
  • Flowers snow white, small, with the lip uppermost (non-resupinate), the spur therefore arising above the ovary. – Platanthera nivea, which also is classified varyingly, often with the the two green-flowered species
  • Flowers green to brownish-green, the labellum lowermost and entire (with no fringing) – Platanthera clavellata and P. flava  (Taxonomists have expressed different opinions as to the inclusion of these plants in the same genus as the “fringed” orchids.)

Platanthera ciliaris, P. blephariglottis, P. chapmanii, & P. cristata

The “Fringed Orchids” constitute a definable group, which in the Apalachicola flora consists of four species and occasional hybrids.  These plants are interfertile and known to cross-pollinate, so we encounter intermediates in mixed populations.  But the vast majority of plants can be identified to species, if you appreciate the structure. The easiest to determine is Platanthera blephariglottis (sensu lato), a plant with large white flowers, each of which has a long spur careening toward and well past the stemSpecimens reported from our region have been determined as variety conspicua.  Some populations of White Fringed Orchids have barely any fringing along the lip, and have been segregated as Platanthera integrilabia (or P. blephariglottis var. integrilabia), but those aren’t reported in the regional flora.

Platanthera blephariglottis flowers

It’s tempting to think of P. blephariglottis as a white form of the better known Platanthera ciliaris, but the flower structure is noticeably different, having a greater spread separating the viscidia of the two pollinaria, a longer ovary/pedicel, and a longer spur. 

These are important distinctions, since White Fringed orchids are predominately moth-pollinated, something I personally observed as a younger person while studying the plants in my late 20s, in a South Carolina population with scores of plants.  Camped out at the population late into the evening, I was prepared with camera, flashlights, insect nets, and a body wash of insect repellent, wondering how it would be possible to observe pollination in the dark, in the shrubby edges of a bog.  With the sun gone from sight, in the gloaming, as my eyes dark adapted and lost access to color, the last distinguishable elements in the landscape were the torch-like P. blepahriglottis white-flowered, growingly fragrant spikes.  I recall the magical moment when suddenly an inflorescence began to shake, then another.  Flashlights and camera were worthless, but I had a net with a long handle.  Hurling myself through the population, I made a frantic sweep at a bobbing head, decapitating the inflorescence but scoring a sphingid moth bearing pollinia adhering to its compound eyes.  It was a sloppy bit of science, but amazingly satisfying.  Minimally, I can confirm that the broader spread of the column (separation between the two viscid) and the longer spur have real meaning in relationship to pollinators.  Most importantly, the flower must open completely enough, with petals sufficiently spread so as to provide hovering space for a feeding moth.

The other large-flowered Fringed Orchid is Platanthera ciliaris, the only plant of this group known to earlier botanists whose work was summarized by Linnaeus (note he also documented Platanthera flava), and perhaps the most well-known of the Fringed Orchids.  It’s difficult to ignore, an extravagant spike of striking yellow-orange flowers embellished with long yellow fringing and nectaries that can exceed 3 cm in length.  What’s not to love.

Platanthera ciliaris inflorescence

Platanthera ciliaris was the first orchid I studied in the field, having encountered a population near my hometown of Eufaula, AL as a botany student at Auburn (I learned the plant as Habenaria ciliaris).  And it was the first orchid in which I studied pollination, learning that large Papilios flock to the inflorescences, studiously probing flowers to the point that numerous pollinia will attach to their compound eyes – a bit of a detriment it seems.  Observing these plants one observes there’s a relationship between the spacing of the viscidia, the length of the spur, and the positioning of fringe on the labellum that meets the anatomy and behavior of these large butterflies.  Unlike pollination of P. blephariglottis, for which the lip and fringe might impede probing by a hovering moth, the fringe provides scaffolding to which the visiting butterfly clings while feeding.  The spur must be long enough to force a butterfly to jam its head into the column, and the spread between viscidia must match the dimensions between the butterfly’s compound eyes.  Finally, once attached to the butterfly and extracted, the pollinia have to collapse forward, so as to contact the stigma of a subsequent flower.  The pollinial stipe, in fact, must be long enough to assure that contact.  To appear as helter-skelter as one might imagine, based on the seemingly faltering, irradic movement of butterflies, the mechanism is one of micro-precision.  If youi examine the head of the butterfly, you’ll see a yellow speck which is one of several pollinia adhered to the compound eyes. It is a wonder.

Pollination of Platanthera ciliaris

At the other end of this spectrum is the much smaller, somewhat more common Platanthera cristata.  This orchid is more difficult to circumscribe than its larger cousin, being variable in aspect, from very short plants with a few flowers on the scapes to taller plants with long, densely-flowered spikes.  The flowers range more widely in color, from a paler yellow to an orange equivalent to that of P. ciliaris.  Flowers are small, with short, reasonably straight nectaries that typically are no longer than the ovary.  The dorsal sepal cups more tightly over the column and the two winglike pollinial chambers are closed up, like a butterfly with closed wings.  The viscidia point straight, down toward the lip and not out. 

Platanthera cristata flower, note the straight line of the paired pollinial chambers, and the positioning of the viscidia (which in the flower are darkened)

Clearly this orchid is structured to attach pollinia to smaller objects, most likely the base of a short proboscis.  But having spent scores of hours in populations of Platanthera cristata, and having seen many visiting insects, I was never convinced I’d seen the effective pollinators.  I imagine it could be a small butterfly, or fly, or most likely a bee, but have no solid clues to offer.

The image above shows Ciliaris to the left, Cristata to the right, and Chapmanii positioned in the middle. Between the large, inviting blossom of P. ciliaris and the small, more enclosed flower of P. cristata, we find intermediates.  J. K. Small noticed particular specimens in material collected by Dr. Chapman, describing them as Blephariglottis chapmanii in his monumental 1903 Flora of the Southeast.  In his 1910 monograph, Oakes Ames re-amalgamated the genus Habenaria, treating Small’s taxon as a hybrid – Habenaria ×chapmanii.  Luer accepted that concept in his 1972 treatment of Florida orchids, publishing the binomial Platanthera ×chapmanii.  That was the situation when I came into this story.

As a graduate student at Vanderbilt, working with Dr. R. B. (Ben) Channell, I entered the world of biosystematics (a term that has been erased from the systematics vocabulary).  Naturally occurring hybrids were interesting, with many studies documenting routes of speciation, from introgression to stabilization through tetraploidy.   Ben had encountered a mixed population of what he called Habenarias, and had noted documentation about the Chapman hybrid, suggesting I contact Donovan Correll, who had attended Duke with Ben, had studied with Oakes Ames, and, in 1950, the year I was born, had published his Native Orchids of North America.  Correll responded, and I even visited with him at Fairchild Tropical Gardens.  I borrowed specimens of Habenaria (Platanthera) ×chapmanii, as well as the putative parents, Platanthera ciliaris and P. cristata, read through the literature, and began field collections in the spring of 1975. 

Ben and I traveled to the Tennessee mixed population where he had seen both putative parents, and I was able to find my first bona fide hybrid.  This seemed propitious.  Back in the lab, studying specimens, I knew there was a broad geographic range to explore, but the fascinating aspect of specimens was the number of “hybrids” collected in the Florida Panhandle.  Still, I fanned out across the Southeast, South Carolina, Georgia, Alabama, inspecting historical collecting sites, but intermediates were very rare. Importantly I also made a beeline to the Panhandle.  Contacting Norris Williams (who was at Florida State in Tallahassee at that time), he agreed to meet me in Apalachicola National Forest when he was conducting a field trip.  There he would show me Chapman’s Orchid.  On a warm Saturday morning, along with his students, I encountered a population of scores of plants, all meeting the description of the “putative hybrid.”

Back in Nashville, pouring over the literature and specimens, I realized I’d failed to make the salient connection that Dr. Chapman worked out of Apalachicola, and Small’s plant was described from that area, from Chapman’s collections.  There were, I suddenly understood, two plants, similar, related, yet distinct.  

In mixed populations, not routinely, but occasionally, one might encounter spontaneous hybrids between Platanthera ciliaris and P. cristata. (which I would later name Platanthera ×channellii.) In fact, in mixed populations of any Fringed Orchids, you may encounter a few hybrids.  But then, in the Panhandle I encountered what seemed to be the hybrid form, in pure.  Amazingly, later I would examine similar populations in the Texas Big Thicket, north of Beaumont – several hundred miles away. 

The realization that Chapman’s Orchid presented itself as species, existing in pure populations, altered the plan.  Suddenly my putative hybrid wanted to be a stabilized species of hybrid origin – exciting!  Might I be encountering another instance of examples documented in research in which a hybrid form became tetraploid – the golden ticket to instant speciation.  That meant I’d need to examine chromosomes.  And of course I’d have to conduct controlled crosses, simply to confirm the production of viable seed from cross-pollination.  There was a lot to be done, all of which centered on Apalachicola.  With permission from the Forestry service, I camped out at Wright Lake for several weeks over three summers, working most intensely in one study population.  

Summer of 1976, counting chromosomes by twilight

Cutting to the chase, I was able to establish there was no tetraploidy.  Platanthera chapmanii had the same number of chromosomes as its putative parents.  I confirmed that cross pollination produced viable seed, though I never mastered a sterile medium that would support production of mature plants.  Most importantly, I was able, on multiple occasions, to observe pollination in Chapmanii populations.  To me, that was the proved to be the most important connection. 

Platanthera chapmanii is pollinated by the large Papilios, just as I had observed in Platanthera ciliaris.  However, with a much shorter spur (about the length of the ovary), butterflies large Papilios visiting Chapmanii never contact the column with their eyes.  Instead, pollinia adhere to the proboscis, halfway along its length.  This doesn’t provide for absolute isolation between Chapmanii and Ciliaris, because a butterfly with Chapmanii pollinia on its proboscis would likely deposit pollen on the stigma of any Ciliaris flower visited.  It does, however, ensure successful pollination of Chapmanii, which exists in many populations where there are no plants of Ciliaris (there’s a curious lack of Platanthera ciliaris populations in the Apalachicola area.)

Platanthera chapmanii pollination

The pollination mechanism for Chapmanii also explains the most taxonomically useful morphological feature of these plants.  If you examine spontaneous hybrids in mixed populations of Ciliaris and Cristata, you’ll see that the column shape is intermediate between the two parents.  Chapman’s orchid, however, shows a distinct adaptation for successful pollination.  The column in flowers of these plants develops a bend at the end where the viscidia are presented, positioning the viscidia close together and turned down toward the entry to the spur at the base of the labellum.  That seems to improve their odds in attaching pollinia to a narrow proboscis. The image below, taken from my Vandy thesis, shows profile drawings of columns from spontaneous hybrids, Platanthera ×channellii (above), and drawings of Platanthera chapmanii columns (below).

Of interest, Platanthera chapmanii and Platanthera cristata do occur together frequently in the Panhandle.  And, I have seen Papilios visiting Cristata, not many times, but on rare occasions.  I believe it would be possible for a butterfly to pick up pollinia of Cristata, but less likely that Chapmanii pollinia could pollinate a Cristata flower, which has a very short spur.  Moreover, I have no evidence that Papilios are pollinators for Cristata. 

For all of the work of many people (including myself), there remain mysteries.  The pollination story is not complete.  Observing pollination takes presence and patience, in many case an extreme, even unattainable quantity.  The segregate Texas population hasn’t been explained, though I know that was being investigated by C. R. Bumbly.  The life histories of these plants are not well-documented.  I followed individuals in a population for many years, but they were all extirpated at some point, and the population never recovered.  But I’ve watched another population increase from just a handful of individuals in 1975 to thousands this past year, all seemingly benefitting from distribution by highway mowing equipment.  Populations that were rich in numbers in open areas nearly fifty years ago, retain just a few flowering specimens along edges of closed forest today.  There’s plenty for more Grad students to explore.

Platanthera nivea

Platanthera nivea

Shifting gears to the non-fringed orchids, I’ve already suggested that I prefer the Rafinesque and Rydberg segregation of the Fringed orchids as Blephariglottis, but I also understand recognition for the very different Snowy Orchid, currently classified as Platanthera nivea.  

Nuttall described it as Orchis nivea.  Lindley, of course, classified it as Platanthera nivea.  Sprengel consolidated it as Habenaria nivea, which was supported by Ames and later by Correll.  In 1898, the Orchid world’s unpredictable Kraenzlin, (who later would be the first to gain access to Reichenbach’s herbarium), decided it should be classified in one of Blume’s SE Asian genera, as Peristylus niveus,. Rydberg, in 1901, segregated it as Gymnadeniopsis nivea (a genus he erected to include three species reported in Florida).  I’m fond of the concept of segregating Nivea, but that is not the current consensus, though the column profile argues for a relationship to other Platantheras.  Let’s examine the flowers; you can make your own decision in this matter.

Up front I have to admit there are difficulties in viewing Nivea flowers.  Every component (but for the pollen) is white, small & compact, a bit succulent, and tightly compressed.  You have to examine fresh flowers using a dissection scope to appreciate the floral structure.  And yes, the basic flower plan is in line with that of Fringed Orchids.  The column shows two separate chambers, each housing a single pollinium that is basically club-shaped, but with very short stipes and a large foot-like viscidium – at least as compared to the Fringed Orchids. The most convincing similarity is presence of “auricles” – these are textured structures, one on either side of each anther locule.  In Chapmanii and Ciliaris, auricles are curious cellular knobs positioned toward the base of the column, but in Nivea, they are comparatively much larger, almost as large as the pollen masses, and (due to the compact nature of the column) one sits directly to the outside of each pollen mass.  In cleared samples (preserved in fixative that makes the structure somewhat transparent), the auricles appear as though they are sterile pollinia.  In that instance, the flower almost appears to produce four pollen masses, giving pause to wonder more deeply about their structural origins (remember, every basis for Orchid interpretation considers this to be a single anther, not two anthers).  If anything ties Nivea strongly to the Fringed Orchids, it would be that anomalous structure, the auricle.

Similarities have landed Nivea in Platanthera, currently with about 150 species.  But differences from the Fringed Orchids (and perhaps other Platantheras) are real.  The flower is non-resupinate, positioning the labellum uppermost.  The strap-like lateral petals, practically as large as the labellum, fold back (as do the sepals), whereas, in the Fringed Orchids, those two lateral petals are relatively small, standing upright alongside the dorsal sepal, like mere decorations. The labellum curves upward like a tail of smoke, in some flowers rolling backward, like a windsock.  At its base the labellum forms a nectary, a 14-18 mm long, very slender capillary tube.  Hovering over the nectary orifice is the crystalline white, compact column, enclosing pale yellow pollinia.  The viscidia of the pollinia converge, hovering at the nectary orifice, perfectly located to attach to anything probing the nectary.  Employing the most slender bristle I could control, I’ve been able to slip a filament into the nectary various times, on each occasion extracting one of the pollinia. 

Platanthera nivea flower. Note the lip is the attenuated segment curling out and up from the flower center.

Adaptations that orient the viscidia into the top of the nectary also generate distinct departure from the structure of Fringed Orchid flowers. Nivea maintains the paired projecting column structure which I’ve described as a stirrup, seeming to represent the projecting horns or stirrups of the Fringed Orchid column, but the Nivea stirrup turns a bit down, arching around the nectary orifice. Most significantly, the stipe and viscidium of each pollinium in Nivea is not imbedded in the stirrup (as it is with Fringed Orchids). The structure is present, but the function is different. In the Fringed Orchids, this process projects outward or downward, precisely positioning the viscidia. In Nivea, the visicidia develop in the cavity between, just above the stigma.

Platanthera nivea, sketch, oriented upside down

In frontal view, the Nivea column is very different from that of Ciliaris and the other fringed orchids. However, in profile, the similarities are more obvious. My interpretation of the structure I see under magnification (remember we are talking about a structure less than 2 mm from top to bottom) is that Nivea actually produces similar processes to those of the Fringed Orchids, but with a real twist. In the Fringed Orchids, the projecting stirrups support the viscidia, positioned at the tips. In Nivea, the viscidia develop between the stirrups, hovering at the nectary orifice, the processes (stirrup) then framing the lower half of the opening. What I have not been able to determine is the exact structure of the stigmatic surface, which seems to be an area just below the viscidia, between and underneath the paired appendages that create a stirrup or collar partially surrounding the nectary orifice.  Logically the stirrup assists in aligning a proboscis with the viscidia, but I know of no studies in this regard. 

Sketch of Platanthera nivea column, about 2 mm along the “face” – oriented so as to compare to other Platantheras, but it’s upside down compared to its normal position

This “apparatus” is remarkably detailed as well as miniature, either pollinium barely exceeding 1 mm in length.  Once past flowering or dried, the architecture becomes inscrutable.  To interpret and explain this structure, we need an excellent draftsperson with patience, steady hands, and access to a high quality video dissection scope.  As with so many orchids, I can’t imagine understanding or interpreting this structure having only studied dried plant material.  

The Remainder:  Platanthera integra, P. flava, and P. clavellata.

You’ll have to stay tuned.  I can’t speak from personal experience.  Though I do have images and a video of Platanthera integra, I’ve failed to take the opportunity to examine a fresh flower – which is crucial to my capacity to explain its structure.  As to Platanthera flare and P. clavellata, I’ve never encountered living material, but hope to have an opportunity this year (though P. flava may have already flowered.  I’d truly appreciate information from anyone reading this who can point me to living, flowering plants.

The Bigger Story – Platantheras,  Floral Structure, & Taxonomic History

When botanists talk about flowers, there’’s seldom a perfect example available.  Tulips are useful because the parts are easily seen, the ovary is superior (you can show students the entire pistil inside the flower), but their sepals and petals are so similar we call them “tepals” – making a convoluted tale in describing “calyx” for students.  Roses are available, but so many cultivated roses sacrifice stamens to double up with petals; even in single roses, the pistils are a mess, tucked down inside the rose hip.  Peonies, if single ones are available, are wonderful, but a rare site at many times.  We have Magnolias here in the Panhandle, enormous flowers that are easily seen, but where do sepals end and petals begin?  And what do you make of that cone-like spiraling of pistils? 

Magnolia grandiflora

Take some advice and don’t start a discussion with Hibiscus (below).

Hibiscus sinensis hybrid

Orchids, however, take the prize in deviating from textbook sepal-petal-anther-pistil diagrams.  Like Tulips, the sepals and lateral petals are similar, defying illustrations that show a distinct calyx.  One of the petals, the labellum (the “lip”) is normally shaped and even colored very different from the others, often functioning as a landing or positioning platform for pollinators.  The ovary is not obvious, being “inferior” – embedded in the flower stem (the pedicel) rather than positioned atop the petals.  And the single stamen (only the Cypripediums and earlier orchids have more than one stamen) is united with style and stigma tissue, forming a structure called the “column”.  

Cattleya complex hybrid, showing the column

As small as an orchid anther is, the structural intricacies of the column are highly detailed, even intricate and precise.  Short of the wondrous gynostegium we see in Asclepiads, no other plant group has evolved a mechanism so elaborate.  Moreover, the particularities can be minutely differentiated, showing combinations of features (presence, number, and nature of pollinia, stipes, & viscid) that coincide closely to the major evolutionary clades, and often to key differences between genera and species.  Though we appreciate this today; none of these aspects of orchid flowers were obvious to Renaissance scholars.  

We recall it wasn’t until Camerarius (1694) and Vaillant (1718) explained the role of pollen in sexual reproduction that flower structure was seen as biologically significant.  Back in the day (that is, before 1700), roots were given great attention.  Herbalists included illustrations and descriptions of plant roots, envisioned to some extent as the “mouth” of a plant, consuming water and nutrients, as well as harboring  much of the plant’s curative possibilities.  Most orchids first known to Western science (that is, to European herbalists and early taxonomists) were terrestrial herbs, of interest for putative medicinal properties.  The paired, swollen testicular roots were, of course, interpreted as as a sign the plants might have aphrodisiac qualities, and suggested the plant name, ‘orchis”, which had been adapted from the Greek for testicle. 

Linnaeus had, of course, given attention to the flowers.  He knew that Orchis flowers had what appeared to be two anthers, which today we interpret as a single anther with widely-separated chambers.  And he appreciated the structure of the lip, with its long nectary (spur).  Indeed, Linnaeus separated Ophrys based on the remarkable wasp-like lip of those plants.

The first Fringed Orchid known to Western scientist fit the fleshy-rooted model understood for Orchis.  Morison (1799), Ray, Gronovius (1739), and Linnaeus considered this North American plant an Orchis, and Linnaeus conferred the epithet ciliaris in 1753.  Orchis ciliaris was one of 62 orchid species (in 8 genera) Linnaeus included in Species Plantarum, those plants constituting slightly over 1% of the nearly 6,000 species he presented.  

A lot has changed.  Today orchid species number nearly 30,000, and constitute as much as 10% of flowering plant species, and Linnaeus’s eight genera have given way to hundreds.  Additional Fringed Orchids were added by Michaux, who described Orchis cristata, as well as the Green Wood Orchid, Orchis clavellata in 1803.  Willdenow, assuming authorship for the 4th edition of Species Plantarum, published the genus Habenaria in 1805, segregating two Jamaican terrestrials with paired labellar appendages, the first Rein Orchids.  In the same publication, he described the White Fringed Orchid, Orchis blephariglottis.  Along comes Robert Brown, in 1813, who recognized our native species as distinct from Orchis, reclassifying them as species of Willdenow’s Habenaria.

Later, in 1817, Louis-Claude Richard established the genus Platanthera to segregate a European plant from the traditional concept of Orchis.  The following year (1818), Thomas Nuttall, still clinging to Orchis for the the American Fringed Orchid complex, added Orchis integra, the Fringeless member, as well as Orchis nivea, the snowy outcast (which Lindley and other botanists have exiled to other genera) 

In 1835, the great orchidist John Lindley pulled the Fringed Orchids into Richard’s genus Platanthera, generating several binomials we recognize in the ISB Florida Atlas today: Platanthera blephariglottis, P. ciliaris, P. cristata, and even Platanthera flava.  This move segregated the more tropical Habenarias (with their bi-lobed lips) from the temperate-zone Platantheras that have triangular or strap-shaped lips.  In the 1848 second edition of Louis C. Beck’s Botany of the Northern and Middle States (retitled Botany of the United States, North of Virginia), Asa Gray made the combination adding Platanthera integra to the roles.

The taxonomy was not settled, however.  The creative Constantine Rafinesque constructed a new genus, Blephariglottis, in Flora Telluriana (1836), segregating the white and yellow-flowered Fringed Orchids (with a new name for ciliaris that has “fallen into synonymy”.)  In establishing this genus, Rafinesque penned a memorable justification:

Whoever deems the numbers of Stamina unimportant in Grasses, Lilies, etc., must regret that this Genus and many other Orchids are based on their double number, altho’ a most essential character: 1 and 2, 3 and 6, 5 and 10 stamina, always distinguish very distinct Genera, and whoever unites Azalea to Rhododendron sins against Linneus and Nature! meantime notwithstanding the learned labors of Swartz, Brown, Richard, Lindley… on Orchides, they have left Habenaria, Orchis, Epidendron, and other Genera in utter perplexity.  The last char. of Habenaria is merely a ‘Cor. ringens, labellum calcaratum, antheris nudis, binis’ – while other Genera have elaborate definite characters of 50 0r 60 words.  Habenaria thus includes many blended Genera and requires complete reform.  I shall now begin it, and give a new essential character of Habenaria Raf. ‘Cor. ringens, lab. ad basi calcorato, glandulis nullis, col. simplex, antheris 2 divisie nudis anticis vertical.  Caulescens. fl. racemosis’ – Types the Amer. Sp. having those char. H. fimbirata, O. habenaria, and akin Sp.

P. A. (Per Axel) Rydberg, writing the Orchid treatment for Britton’s 1901 Manual….,) accepted and amplified Rafinesque’s segregate genus to include the more Northern B. lacera, B. leucophaea, B. grandiflora, B. psychodes, and B. peromoena (not reported as part of the Florida flora).  In accepting Willdenow’s epithet ‘blephariglottis’  for the White Fringed Orchid, Rydberg created the sparkling tautonym Blephariglottis blephariglottis (a double name against current rules for plants.)  John Kunkell Small, who worked with Britton and Rydberg at New York Botanical Garden, followed Rydberg’s lead, describing Chapman’s Orchid, Blephariglottis chapmanii, in the remarkable 1903 Flora of the Southeastern US. Both treatments also determined that the other four species reported in Florida and currently classified as Platantheras (the non-fringed ones) should be split into two genera, Platanthera flava was treated as Perularia while Plathanthera nivea, P. clavellata, and P. integra were classified as Gymnodeniopsis.  

These segregate genera (which make some sense to me) did not reign a full decade.  The situation changed radically in 1910 with Oakes Ames’s publication of a complete bibliography and citing of specimens for Habenaria sensu lato, basically restoring and expanding Willdenow’s genus and corroborating Robert Brown’s work.  Though monumental, this was no monograph.  Curious how Ames, who in the treatment never described a single plant, patronizingly critiqued Rydberg:

“Dr. Rydberg has gone further than any other author of modern times in the splitting up of the genus, and has given us several new segregate genera.  He is not at all in sympathy with the conservatism of Bentham and Hooker, Torrey, Gray, Cogniaux and others, and is much opposed to the maintenance of large groups which in any way may be divided.’  He concluded: “…the single characters assigned have all proved either so variable from species to species, or so little in accordance with any other distinction, that I feel compelled to reunite the proposed genera after the example of A. Gray and some other recent bothanitst, although I cannot go so far as to agree with Grenier and Gordon in unitizing the whole genus with Orchis…”   As America’s great Orchidist in the first half of the 20th Century, Oakes Ames had great influence.  Wealthy, erudite, focused, and Harvard solid, he was the man.  Everybody folded and came under this umbrella and we were taught to treat these orchids as Habenaria, again.  (see Appendix for more extensive statements from Ames)

Only with Carlyle Luer’s 1972 book, The Native Orchids of Florida, did taxonomists re-adopt Lindley’s treatment.  In that publication, Luer made transfers to create Platanthera clavellata, Platanthera nivea, and Platanthera ×chapmanii.  The Luer book remains the basis for classification you see in the ISB Plant Atlas today.  However, examining recent work by other researchers, we see Blephariglottis resurrected, with combinations published that update Fringed Orchids in a separate genus.

The Side Story

As was said earlier, floral structure, on paper being sepals, petals, anthers, and pistils – is not so simple.  Indeed, this is an ultimate rabbit hole, given the possible variations.  I believe Linnaeus realized this when he organized nearly 6,000 plant species in Classes based on their number of stamens, one stamen would be MONANDRIA, two DIANDRIA, three TRIANDRIA.., running out of steam with POLYANDRIA.  

And because stamens are not so regular from one kind of flower to another, what about flowers with unequal stamen lengths? OK, Linnaeus devised DIDYNAMIA to accommodate the Scrophs and some of the Mints, any with two short and two long stamens.  But the Mustards have 2 short and 4 long stamens, so Linné had to erect a Class called TETRADYDINAMIA.  Some kinds of flowers show fusion of stamens into clusters; enter the catch-all Classes MONADELPHIA, DIADELPHIA, and POLYADELPHIA.  Hibiscus and Geranium, with stamens united in a brotherhood, would be categorized in MONADELPHIA.  And those Composites (as well as Lobelias), with their fused anthers, were accommodated in the SYNGENESIA

When confronted with the curious floral structure of Orchids and Passifloras, Linnaeus classified them as GYNANDRIA, the term itself explaining that these are plants in which the anthers (male = andro) and the pistils (female = gyno) form a single structure.  The Orchids, he determined, have two stamens, as contrasted with Passifloras with five, thus Orchids had their own Order, DIANDRAE.

But Orchids didn’t require major effort for Linnaeus.  He had studied collections, even published what is considered the first treatment of the tribe, his ‘Species orchidum et affinium plantarum’, in 1740, including 38 species in 10 genera.  In 1753, his Species Plantarum included 8 genera with 62 species, about 1% of the plants treated, as compared to current understanding that estimates 10% of flowering plant species are Orchids.  

Until the 19th Century, when tropical plant collecting and “stoves” (the early English term for greenhouses) changed the cultural impact of the word “orchid”, botanists thought of orchids in terms of early European herbalists.  Linnaeus’s Species Plantarum (1753), always the beginning of the plant name database, listed 8 genera and 62 species in his GYNADRIA DIANDRIA, the Class of plants that had stamen united , mostly being those plants known to Europeans for hundreds of years.  At first, those genera absorbed orchids showing up from the Americas and the Far East. Below, I’ve included Linnaeus’s entire schema, so that you may see how he classified differing plants we encounter here in the Panhandle. These are genera he knew and included in Species Plantarum, though their generic limits have been altered radically over the 370 years of discovery and practice since the 1753 publication. Notice that any family with distinctive stamens (of the same number) falls together in a Class. But you’ll see the Grasses show up in separate locations, as do Azalea and Rhododendron.

  • Classis 1. Monandria: 1 stamen, such as Canna
  • Classis 2. Diandria: 2 stamens, i.e. Jasminum, Chionanthus, Salvia
  • Classis 3. Triandria: 3 stamens: Crocus, Gladiolus, Iris, Xyris, Cyperus, Panicum, Aristida…., Eriocaulon,
  • Classis 4. Tetrandria: 4 stamens: Diodea, Houstonia, Callicarpa, Polypremum, Cornus, Ludwigia, Ilex,
  • Classis 5. Pentandria: 5 stamens, Lithospermum, Lysimachia, Spigelia, Azalea, Phlox, Ipomoea, Physalis, Solanum, Rhamnus, Itea, Vitis, Asclepias, Hydrocotyle, Rhus, Parnassia, Linum, Crassula
  • Classis 6. Hexandria: 6 stamens, Tillandsia, Burmannia, Crinum, Amaryllis, Allium, Lilium, Tulipa, Aletris, Yucca, Orontium, Juncus, Rumex, Alisma
  • Classis 7. Heptandria: 7 stamens, Aesculus
  • Classis 8. Octandria: 8 stamens: Osbeckia, Rhexia, Gaura, Vaccinium, Polygonum
  • Classis 9. Enneandria: 9 stamens: Laurus
  • Classis 10. Decandria: 10 stamens: Cassia, Guaiacum, Tribulus, Melastoma, Kalmia, Rhododendron, Clethra, Silene, Sedum, Oxalis, Agrostemma, Phytolacca
  • Classis 11. Dodecandria: 12 stamens: Asarum, Styrax, Lithrum, Euphorbia 
  • Classis 12. Icosandria: flowers with 20 (or more) stamens: Cactus, Myrtus, Prunus, Rosa, Rubus
  • Classis 13. Polyandria: flowers with many stamens: Sanguinaria, Podophyllum, Papaver, Sarracenia, Nymphaea, Mimosa, Delphinium, Aquilegia, Magnolia, Annona, Clematis
  • Classis 14. Didynamia: flowers with 4 stamens, 2 long and 2 short: Teucrium, Lamium, Origanum, Thymus, Ocimum, Scutellaria, Gerardia, Gesneria Antirrhinum, Scrophularia, Bignonia, Lantana, Buchnera, Linnaea, Orobanche, Ruellia
  • Classis 15. Tetradynamia: flowers with 6 stamens, 4 long and 2 short – Draba, Lepidium, Thalapsis, Alyssum, Dentaria, Arabis, Brassica, Cleome
  • Classis 16. Monadelphia; flowers with the anthers separate, but the filaments united at the base – Geranium, Side, Hibiscus, Stewartia, Camellia, 
  • Classis 17. Diadelphia; flowers with the stamens united in two groups – Fumaria, Polygala, Erythrina, Genista, Amorpha, Crotalaria, Lupinus, Robinia, Phaseolus, Pisum Lathyrus, Vicia, Indogofera, Clitoria, Trifolium, Lotus, Medicago, 
  • Classis 18. Polyadelphia; flowers with the stamens united in several groups: Theobroma, Citrus, Hypericum
  • Classis 19. Syngenesia; flowers with 5 stamens having anthers united at the edges – Tragopogon, Sonchus, Lactuca, Crepis, Elephantopus, Echinops, Carduus, Eupatorium, Ageratum, Artemisia, Baccharis, Erigeron, Senecio, Aster, Solidago, Chrysanthemum, Helianthus, Silphium, but also Verbena, Lobelia, Viola, Impatiens
  • Classis 20. Gynandria; flowers having stamens united to the pistils: Orchis, Ophrys, Satyrium, Serapias, Limodorum, Arethusa, Cypripedium, Epidendrum, Sisyrinchium, Nepenthes, Passiflora, Aristolochia, Grewia, Arum, Calla
  • Classis 21. Monoecia: monoecious plants: Callitriche, Lemna, Typha, Zea, Carex, Phyllanthus, Betula, Buxus, Urtica, Morus, Ambrosia, Amaranthus, Zizania, Ceratophyllum, Sagittaria, Quercus, Juglans, Fagus, Platanus, Liquidambar, Pinus, Cupressus, Acalypha, Croton, Jatropha, Cucurbita 
  • Classis 22. Dioecia: dioecious plants: Najas, Salix, Visnum, Myrica, Pistachia, Cannabis, Smilax, Dioscorea, Juniperus, Taxus
  • Classis 23. Polygamia: polygamodioecious plants: Musa, Veratrum, Andropogon, Cenchrus, Atriplex, Acer, Begonia, Gleditsia, Fraxinus, Diospyros, Nyssa, Panax, Ficus
  • Classis 24. Cryptogamia: organisms that resemble plants but don’t have flowers, which included fungi, algae, ferns, and bryophytes: Equisetum, Ophioglossum Osmunda, Asplenium, Polypodium, Adiantum, Trichomones, Isoetes, Marsilea, Lycopodium, Sphagnum, Polytrichum, Mnium, Bryum, Hypnum, Jungermannia, Marchantia, Anthoceros, Lichen, Fucus, Ulva, Spongia, Agaricus, Bolletus, Phallus
  • Palmae: Chamaerops, Cycas, Phoenix, Areca, Caryota 


I found it interesting to read more concerning Oakes Ames’s justification regarding Habenaria. You might also. From his preface:

“In the Evolution of the Orchidaceae,’ in the Orchid Review (February, 1910), R.A. Rolfe treats the Habenaria group under two sub tribes, namely, Gymnadenieae and Habenarieae, the former having one and the latter two distinct stigmas.  he says that Platanthera and Gymnadenia are sometimes united with Habenaria, an arrangement that makes Habenaria one great chaotic aggregate whose characters cannot be defined with any degree of precision. This statement leads to the conclusion that the characters on which the three genera depend for purposes of classification are not sufficiently clear to allow subgeneric characterization under Habenaria.  This conclusion is palpably incorrect, as the basis on which systematists have relied who have upheld the disjunction of Habenaria is the distinctness of these characters.  This basis hardly leads to chaos whether adopted or rejected, although it does not simplify classification, as I attempted to show.”  

“The genus Habenaria has undergone a series of changes at the hands of those botanists who have given it special attention until at the present time it is an arduous task to comprehend its rational limits.  No two systematists agree in their revisions; and while some have considered the treatment in Bentham and Hooker’s Genera Platinum too broad, including as it doe stat recognized groups of many authors under one general heading, namely, Habenaria, others have not been content to reestablish as genera such subgeneric groups as Gymnadenia, Platanthera, Peristyles and Coeloglossum, but have made segregates from several of them.  Although there may be excellent reasons both for and against the maintenance of Habenaria in the sense in which it was understood by the authors of the Genera Plantarum, the weight of authority seems to uphold the broader view.  So eminent a student of the Orchidaceae as Lindley expressed grave doubts as to the validity f the characters on which he admitted as distinct groups Gymnadenia and Peristyles win his Genera and Species of Orhidaceous Plants.  In fact it is usually with apologies, doubts, or detailed explanations that authors accept the smaller groups which have been at one time or another put into or removed from Habenaria.”

…Under Platanthera he (Lindley) wrote, in reference to the recognition by Robert Brown of a section based on the anthers: ‘The genus has to depend upon another distinction which I can discover no exception, namely, to the absence of the fleshy processes of the lower lip of the stigma. Otherwise Platanthera is the same as Habenaria.’  Yet in his discussion of Gymnadenia we find the following, which in conjunction with his remarks on Platanthera is of more than casual interest:’It is scarcely possible to find any very precise limits between this genus and Platanthera..’”

“We are indebted for the most recent revision of a part of the genus Habenaria as represented in North America north of Mexico to Dr. P. A. Rydberg of the New York Botanical Garden.  Dr. Rydberg has gone further than any other author of modern times in the splitting up of the genus, and has given us several new segregate genera.  He is not at all in sympathy with the conservatism of Bentham and Hooker, Torrey, Gray, Cogniaux and others, and is much opposed to the maintenance of large groups which in any way may be divided.  An illustration of what I mean may be obtained from Britton’s Manual of the Flora of the Northern States and Canada, for which Dr. Rydberg prepared a revision of the Orchidaceae.  In this work eighteen species are admitted which were formerly included by Gray and Button and Brown, in the genus Habenaria.  Dr. Rydberg has distributed these species among severn genera, of which three by him are newly established.”

“…the single characters assigned have all proved either so variable from species to species, or so little in accordance with any other distinction, that I feel compelled to reunite the proposed genera after the example of A. Gray and some other recent bothanitst, although I cannot go so far as to agree with Grenier and Gordon in unitizing the whole genus with Orchis…”

Synonymous Names


  • Platanthera blephariglottis (Willd.)Lindl. var. conspicua (Nash)Luer Native Orch. Fl. 140 (1972)
  • Synonyms of Platanthera blephariglottis (Willd.)Lindl. Genera Sp. Orchid. Pl (1835):
  • Orchis blephariglottis Willd. in Sp. Pl., ed. 4. 4: 9 (1805)
  • Habenaria blephariglottis Hooker
  • Habenaria blephariglottis var holopetala (Lindl.) A. Gray
  • Habenaria conspicua Nash in Bull Torr Bot Cl23:100 (1896)
  • Habenaria blephariglottis var. conspicua (Nash)Ames Rhodora 10:70 (1908)
  • Blephariglottis blephariglottis (Hooker)Rydb. in Britton, Manual, 296(1901)
  • Blephariglottis blephariglottis var. holopetala (Lindl.)Rydb.
  • Blephariglottis alba House in Muhlenbergia 1: 127 (1906)
  • Blephariglottis alba var. conspicua (Nash)W.J.Schrenk
  • Blephariglottis conspicua (Nash)Small
  • Blephariglottis albiflora
  • Blephariglottis albiflora f. holopetala (Lindl.)Baumback & P.M.Br.
  • Platanthera ciliaris var. blephariglottis (Willd.) Chapm. in Fl. South. U.S.: 460 (1860)
  • Platanthera blephariglottis var. holopetala (Lindl.)Torr.
  • Kew lists 2 Accepted Infraspecifics
  • Platanthera blephariglottis var. blephariglottis
  • Platanthera blephariglottis var. conspicua (Nash) L
  • Platanthera chapmanii (Small)Luer emend Folsom
  • Blephariglottis chapmanii Small in Fl. S.E. U.S.: 314 (1903)
  • Habenaria chapmanii (Small) Ames in Orchidaceae 4: 155 (1910)
  • Platanthera ciliaris (L.)Lindl. Genera Sp. Orch. Pl. 292 (1835)
  • Orchis ciliaris L. in Sp. Pl.: 939 (1753)
  • Blephariglottis ciliaris (L.) Rydb. in N.L.Britton, Man. Fl. N. States: 296 (1901)
  • Habenaria ciliaris (L.) R.Br. in W.T.Aiton, Hortus Kew. 5: 194 (1813)
  • Heterotypic Synonyms
  • Blephariglottis flaviflora Raf. in Fl. Tellur. 2: 39 (1837)
  • Platanthera clavellata (Michx.)Luer
  • Homotypic Synonyms
  • Denslovia clavellata (Michx.) Rydb. in Brittonia 1: 85 (1931)
  • Gymnadeniopsis clavellata (Michx.) Rydb. in N.L.Britton, Man. Fl. N. States: 293 (1901)
  • Habenaria clavellata (Michx.) Spreng. in Syst. Veg., ed. 16. 3: 689 (1826)
  • Orchis clavellata Michx. in Fl. Bor.-Amer. 2: 155 (1803)
  • Peristylus clavellatus (Michx.) Kraenzl. in Orchid. Gen. Sp. 1: 521 (1898)
  • Heterotypic Synonyms
  • Gymnadenia tridentata (Muhl.) Lindl. in Gen. Sp. Orchid. Pl.: 277 (1835)
  • Gymnadenia tridentata var. clavellata Alph.Wood in Class-book Bot.: 683 (1861)
  • Gymnadeniopsis clavellata var. ophioglossoides (Fernald) W.J.Schrenk in Orchidee (Hamburg) 28: 69 (1977)
  • Gymnadeniopsis clavellata f. wrightii (L.S.Olive) P.M.Br. in Wild Orchids Canad. Marit. & N. Gr. Lakes: 284 (2006)
  • Habenaria clavellata var. ophioglossoides Fernald in Rhodora 48: 161 (1946)
  • Habenaria clavellata var. wrightii L.S.Olive in Bull. Torrey Bot. Club 78: 291 (1951)
  • Habenaria maritima Raf. in Herb. Raf.: 74 (1833)
  • Habenaria tridentata (Muhl.) Hook. in Exot. Fl. 1: t. 81 (1823)
  • Orchis clavellata var. tridentata Muhl. in Cat. Pl. Amer. Sept.: 80 (1813)
  • Orchis tridentata Muhl. ex Willd. in Sp. Pl., ed. 4. 4: 41 (1805), nom. illeg.
  • Platanthera clavellata var. ophioglossoides (Fernald) P.M.Br. in Wild Fl. Notes 3: 21 (1988)
  • Platanthera clavellata f. slaughteri P.M.Br. in N. Amer. Native Orchid J. 1: 200 (1995)
  • Platanthera cristata (Michx.)Lindl.
  • Blephariglottis cristata (Michx.) Raf. in Fl. Tellur. 2: 39 (1837)
  • Habenaria cristata (Michx.) R.Br. in W.T.Aiton, Hortus Kew. 5: 194 (1813)
  • Orchis cristata Michx. in Fl. Bor.-Amer. 2: 156 (1803)
  • Heterotypic Synonyms
  • Blephariglottis cristata f. straminea (P.M.Br.) Baumbach & P.M.Br. in N. Amer. Native Orchid J. 15: 74 (2009)
  • Platanthera cristata f. straminea P.M.Br. in N. Amer. Native Orchid J. 1: 12 (1995)
  • Platanthera flava (L.)Lindl.
  • Habenaria flava (L.) R.Br. ex Spreng. in Syst. Veg., ed. 16. 3: 691 (1826)
  • Orchis flava L. in Sp. Pl.: 842 (1753)
  • Perularia flava (L.) Rydb. in N.L.Britton, Man. Fl. N. States: 292 (1901)
  • Tulotis flava (L.) Senghas in Orchidee (Hamburg) 24: 97 (1973)
  • Includes 2 Accepted Infraspecifics
  • Platanthera flava var. flava
  • Platanthera flava var. herbiola (R.Br.) Luer
  • Platanthera integra (Nutt.)A.Gray ex L.C.Beck
  • Gymnadeniopsis integra (Nutt.) Rydb. in N.L.Britton, Man. Fl. N. States: 293 (1901)
  • Habenaria integra (Nutt.) Spreng. in Syst. Veg., 3: 689 (1826)
  • Orchis integra Nutt. in Gen. N. Amer. Pl. 2: 188 (1818)
  • Heterotypic Synonyms
  • Gymnadenia flava Lindl. in Gen. Sp. Orchid. Pl.: 279 (1835)
  • Habenaria crocea (Raf.) Raf. in Atlantic J. 1: 119 (1832)
  • Habenaria elliottii L.C.Beck in Bot. North. Middle States: 348 (1833)
  • Orchis crocea Raf. in Atlantic J. 1: 119 (1832)
  • Perularia flava (Lindl.) Schltr. in Repert. Spec. Nov. Regni Veg. 16: 286 (1919)
  • Platanthera nivea (Nutt.) Luer
  • Gymnadenia nivea (Nutt.) Lindl. in Gen. Sp. Orchid. Pl.: 280 (1835)
  • Gymnadeniopsis nivea (Nutt.) Rydb. in N.L.Britton, Man. Fl. N. States: 293 (1901)
  • Habenaria nivea (Nutt.) Spreng. in Syst. Veg., ed. 16. 3: 689 (1826)
  • Orchis nivea Nutt. in Gen. N. Amer. Pl. 2: 188 (1818)
  • Peristylus niveus (Nutt.) Kraenzl. in Orchid. Gen. Sp. 1: 520 (1898)
  • Heterotypic Synonyms
  • Gymnadenia conica Lindl. in Gen. Sp. Orchid. Pl.: 280 (1835)


Willdenow (Linnaeus), 1805.  Species plantarum : exhibentes plantas rite cognitas ad genera relatas, cum differentiis specificis, nominibus trivialibus, synonymis selectis, locis natalibus, secundum systema sexuale digestas, Tomus 4 Pars 1

Morison, RobertPlantarum historiae universalis Oxiniensis

Holding Institution: Biblioteca Digital del Real Jardín Botánico de Madrid

Tomas primus, 1715.  Permalink:

Pars secunda, 1680.  (part 2 was the first fully published volume)

Pars tertia, 1699  Permalink:  (Orchis palmata elegans…)

Richard, Louis-Claude, 1817.  De Orchideis Europaeis annotationes, praesertim ad genera dilucidanda spectantes, BHL Holding Institution: NYBG

Mandelbrote, Scott, 2015.  ‘The Publication and Illustration of Robert Morison’s Plantarum historiae universalis Oxoniensis’, Huntington Library Quarterly , Vol. 78, No. 2 (Summer 2015), pp. 349-379 

Stable URL:

Mandelbrote, Scott, 2013.  OUPblog:  ‘Robert Morison’s Plantarum historiae universalis Oxoniensis’.

Linnaeus, C.  1753.  Species plantarum : exhibentes plantas rite cognitas ad genera relatas, cum diferentiis specificis, nominibus trivialibus, synonymis selectis, locis natalibus, secundum systema sexuale digestas

Biodiversity Heritage Library, Holding Ins: 1908 facsimile from NYSU

tomus 1:

tomus 2:

Gronovius, Johannes F.   1739  Flora Virginica Exhibens Plantas quas V. C. Johannes Clayton in Virginia Observavit atque collegia.

PDF Available from Google Books

Biodiversity Heritage Library, Holder: Harvard University Botany Libraries:

Torrey, J & Asa Gray, 1838-1840.  A flora of North America :containing abridged descriptions of all the known indigenous and naturalized plants growing north of Mexico, arranged according to the natural system

Biodiversity Heritage Library, Holding Inst. New York Botanic Garden



vol2 Missouri BG:

Lindley, John, 1835.  The genera and species of orchidaceous plants. (pgs 291-292)

Biodiversity Heritage Library, Holding M


Nuttall, Thomas, 1818.  The genera of North American plants, and a catalogue of the species, to the year 1817.  (for Orchis, see v2, page 188)

Biodiversity Heritage Library, Holding Inst: Missouri Botanical Garden

vol 1: DOI

vol 2: DOI

Asa Gray in Louis C. Beck, 1848. Botany of the United States North of Virginia, comprising descriptions of the flowering and fern-like plants hitherto found in those states, arranged according to the natural system, with a synopsis of the genera according to the Linnaean system  a sketch of the rudiments of botany, and a glossary of terms. 2nd edition, revised and enlarged.  New York, Harper & Brothers.

Biodiversity Heritage Library, Holding Inst. Library of Congress


Vaillant, Sébastian, 1718.  Discours sur la structure des fleurs, leurs differences et l’usage de leurs parties; : Prononcé a l’ouverture du Jardin Royal de Paris … Et l’etablissement de trois nouveaux genres de plantes l’Aral…. 

Biodiversity Heritage Library, Holding: Welcome


See also:  Bernasconi, Paul & Lincoln Taiz, trans., 2002.  ‘Sebastian Vaillant’s 1717 lecture on the structure and function of flowers’, Huntia 11(2): 97-118 – You will really want to read this!!!

Camerarius, Rudolf Jacob, 1694.  De sexu plantarum epistola (Academiae Caesareo-Leopold. N.C. Hectoris II., Rudolphi Jacobi Camerarii, Professoris Tubingensis, ad Thessalum, D. Mich. Bernardum Valentini, Professorem Giessensem excellentissimum, de sexu plantarum epistola)

Source:  MDC

Description from noscemus (Univ. Innsbruck):  “This work comprising some 80 pages is styled as a letter to the author’s friend and colleague Michael Bernhard Valentini (1657–1729). For the first time in history, a number of proofs for the sexual propagation of plants are given and the difference between monoicous and dioicous plants is explained. Camerarius’ new insights were made possible by the application of the microscope to plant anatomy. The letter belongs to those texts that stimulated Linnaeus to base his novel classification of plants on the form of their sexual organs….An appendix contains a remarkable Alcaic ode (under the mysterious title A.A. Ode) in 26 stanzas summarizing the most important arguments of the letter. Moreover, two academic orations, originally given at the bestowal of baccalaureus and magister grades respectively, treat oak apples (De quercuum gallis) and the nature of semen”

Small, John Kunkel, 1903  Flora of the southeastern United States; being descriptions of the seed-plants, ferns and fern-allies growing naturally in North Carolina, South Carolin, Georgia, Florida, Tennessee, Alabama, Mississippi, Arkansas, Louisiana, and the Indian Territory and in Oklahoma and Texas East of the One-Hundredth Meridian. (Note:  BHL cites Per Axel Rydberg as co-author)

Holding Institution:  Missouri Botanical Garden


Brendel, Frederick, 1879. Historical Sketch of the Science of Botany in North America from 1645 to 1840, The American Naturalist , Dec., 1879, Vol. 13, No. 12 (Dec., 1879), pp. 754-771.

Britton, Nathaniel Lord, 1901.  Manual of the flora of the northern states and Canada, by Nathaniel Lord Britton, Director-in-Chief of the New York Botanical Garden: Emeritus Professor of Botany in Columbia University: Vice-President of the New York Academy of Sciences

Holding Institution: MBLWHOI – Wood’s Hole


Chapman, A. W, 1860. Flora of the southern United States:containing abridged descriptions of the flowering plants and ferns of Tennessee, North and South Carolina, Georgia, Alabama, Mississippi, and Florida: arranged according to the Natural System

Holding Institution:  Missouri Botanical Garden

Chapman, A.W, 1889. Flora of the southern United States: containing an abridged description of the flowering plants and ferns of Tennessee, North and South Carolina, Georgia, Alabama, Mississippi, and Florida: arranged according to the Natural System, 2nd Edition.

Holding Institution:  New York Botanical Garden


Chapman, A.W., 1897.  Flora of the southern United States: containing an abridged description of the flowering plants and ferns of Tennessee, North and South Carolina, Georgia, Alabama, Mississippi, and Florida: arranged according to the Natural System, 3rd edition – Holding Institution:  New York Botanical Garden – DOI:

Ames, Oakes, with Illustrations by Blanche Ames, 1910.  Orchidaceae: Illustrations and Studies of the Family Orchidaceae – The Genus Habenaria in North America.

Holding Institution:  New York Botanical Garden

Habenaria clavellata – pgs 29-38

Habenaria flava – pgs 41-49

Habenaria integra – pgs 49-52

Habenaria nivea – pgs 52-53

Habenaria cristata – pgs 152-155

Habenaria ×chapmanii – pgs 155-156

Habenaria ciliaris – pgs 156-163

Habenaria ×canbyi – pgs 163-164

Habenaria blephariglottis – pgs 164-171

Rafinesque, Constantine S., 1836.  Flora Telluriana, pts 1-4

Holding Institution:  New York Botanical Garden


Fringed Orchids in Pt 2: 36-39

Jarvis, Charlie & Phillip Cribb, 2009. Linnaean source and C]concepts of orchids, Annals of Botany 104:365-376



%d bloggers like this: