Chamaecrista fasciculata – More than a single Stanza

Illustration from Curtis’s Botanical Magazine, 1790 (v3, plate 107) of the plant known then as Cassia chamaecrista

The Back Story – Ancient Taxonomy

Everyone wants a piece of Patridge Pea, but as we know from other American plants, E. L. Greene was in line early, and he took his portion so as to have the last word.  But Greene wasn’t the first.  Michaux, in his 1803 Flora Boreali-Americana, appears to be the first botanist to have provided the plant we call Partridge Pea with a specific epithet that has stuck. Even that effort required two entries, designating the first from Pennsylvania, Cassia chamaecrista (in line with Linnaeus) and a second, from Virginia and South Carolina), with smaller flowers, as Cassia fasciculata.  Transfer his second entry to Chamaecrista and the story is told.

MIchaux’s 1803 Flora Boreali-Americana, showing 3 Cassia species

But as taxonomy goes, there’s a messy backstory.  And it starts with Linnaeus as well as some earlier botanists.   If you check the botanical scriptures, you’ll see that in the beginning, along with other random-seeming plants having 10 stamens and a single pistil, Linnaeus grouped several significant legumes in Class X, DECANDRIA, Order MONOGYNIASophora, Cercis, Bauhinia, Parkinsonia, Poinciana, Caesalpinia, and his 26 species of Cassia  The plants he selected to bear the name Cassia included many old-world legumes of note for their historical herbal uses.  Very different plants (Cinnamons in the Laurel family) are also historically associated with the ancient word Cassia, though botanically they are unrelated.  The linguistic conflation of these very different groups of plants remains problematic today, especially given public interest in herbal medicine.

Staying with genus Cassia (the Legume) doesn’t eliminate confusion.  Linnaeus also elected to include the plants within his Cassia umbrella that we call Senna and Chamaecrista today.  That alone is not problematic; botanists and horticulturists would automatically transpose connections, knowing that a given Chamaecrista might also be called Cassia.  The confusion comes when people do not agree as to which specific epithet is valid and applicable, and for what reasons.  

Partridge Pea, a plant we call Chamaecrista fasciculata, was commonly listed as Michaux’s Cassia fasciculata throughout the 20th century, including the entry in Clewell’s 1985 Guide to the Vascular Plants of the Florida Panhandle.  That’s cool; you just have to know that for some reason botanists have decided to “segregate” this plant and its nearest relatives from a broader concept called Cassia.  But Linnaeus didn’t refer to our Partridge Pea as C. fasciculata, rather he called it Cassia chamaecrista.  His entry #22 for the plant is given below, detailing a habitat in Jamaica, Barbados, Virginia:

Looking at the single specimen from Linnaeus’s herbarium, his sample (which is labeled Cassia chamaecrista) appears to match the plant we call Chamaecrista fasciculata, which means today’s taxonomies have accepted both plants Michaux listed as being represented by the one fasciculata epithet – and, in fact, the epithet chamaecrista has been specifically rejected – i.e. a legal move called nomen utique rejiciendum. That makes sense as long as we keep Partridge Pea in genus Chamaecrista –tautologies (using the same word for both genus and species) are not allowed in botanical nomenclature. But it isn’t clear the full tale is told; you might agree after reading further.

Specimen 528-30 from Linnaeus’s herbarium

In Species Plantarum, Linnaeus includes another plant native to Virginia, addressing it as #26, Cassia nictitans.

Linnaeus’s Cassia nictitans, which Moench selected as the basis for Chamaecrista

That taxon was singled out in 1794 by Conrad Moench as meriting its own generic standing.  Following the lead of earlier botanists whose writings Linnaeus chose to abandon, Moench elected to restore Chamaecrista to generic status, listing Linnaeus’s plant as Chamaecrista nictitans.  

But good news sometimes travels slowly. A full century would pass until the restless Edward L. Greene recovered the threads of that conversation in his 1897 article ‘The Genus Chamaecrista’.  In that article Greene makes the combination we accept today – Partridge Pea bears the botanical name Chamaecrista fasciculata (Michaux) Greene, indicating that E. L. Greene is the author we credit for transferring Michaux’s fasciculata to the genus Chamaecrista. That united the two related plants we still pair together today – the larger flowered Chamaecrista fasciculata and the smaller-flowered Chamaecrista nictitans. The curiosity here is that Michaux had distinguished his fasciculata from Linnaeus’s chamaecrista in writing it has smaller flowers. This leaves me wondering if a Michaux specimen exists, and what type specimen has been designated for our plant. I haven’t unearthed that fistful of worms yet. But the Linnaeus part is easy. You can investigate these concepts yourself, on the Web. Linnaeus’s collections are digitized and readily available through searching for “linnean-online.” Below is an image of his nictitans specimen.

Linnaeus’s specimen 528-33 of Cassia nictitans, from Linnean-online

The genus Chamaecrista is here to stay, and the type is a plant described by Linnaeus, most likely based on the specimen you just viewed. According to Greene, numerous distinctions mark Chamaecrista, any of which would suggest corralling these plants in their own genus, here reproduced from his 1897 statement:

It will thus be seen that what marks so strangely the Chamaecrista corolla is not an inversion, but something still more interesting and curious, namely a torsion of ninety degrees to the left—a character which, by itself alone, if considered in connection with the peculiar habit of the plant and its rather numerous allies, would establish the whole group as securely win the rank of a distinct genus as any genus could nee to be established.  I shall, however, here enumerate the actual characters of the genus CHAMAECRISTA as I have recently observed them in living plants, and in dried specimens of many species.  (1.) Flowers axillary or supra-axillary and solitary or few and fascicled, never terminally clustered as in Cassia. (2.) Buds slender-conical and acuminate (always subglobose or ovoid and obtuse in Cassia.)  (3.) Sepals plane, slenderly acuminate, then-membranous (in Cassia firm-herbaceous obtuse concavo-convex).  (4.) Flower on a twisted pedicel, its banner and keel petals thus made to appear lateral, and one wing enlarged and placed lowermost, the other reduced and becoming uppermost.  (5.) Pods thin, compressed, very promptly dehiscent, never subterete and indehiscent as in most or all Cassias.

You can be certain Greene had his thoughts aligned, his article being so pointed as to merit its own resurrection.  For that reason, I’ve entered the full text, including the single figure, in the Appendix.  

As with other groups we have examined in this series, the pendulum quickly rocked back to conservative treatments for next century, with the broader generic limits favored.  In contemporary science, however, given molecular and cladistic studies, systematists have acknowledged that many large genera are “polyphyletic” – which means that different evolutionary lineages have been assembled based on a far-too-limited number of shared characteristics.  The basic concept today (very different from older philosophies) is that a genus should reflect an assemblage of species, all of which we believe evolved from the same ancestor – that is, they are “monophyletic”.

Given that stance, as well as recent work on Chamaecrista nodulation genetics, (instructions that direct roots to make nodules where nitrogen fixing bacteria thrive), one wonders about the future of the very broad and singular definition of Chamaecrista fasciculata, at least compared to treatments of it’s  “sister species”, Chamaecrista nictitans.  That twinned plant, Chamaecrista nictitans, on which Moench based the genus, was described by Linnaeus as native to Virginia, but is now defined in Plants of the World OnLine (POWO) as native to  “Central & E. U.S.A. to Tropical & Subtropical America…  an annual or perennial and [that] grows primarily in the temperate biome.”  That extensive range and its different history of taxonomic treatment means POWO accepts 13 infraspecific taxa (subspecies and varieties) for Chamaecrista nitictans.  The story is radically different for the POWO description of Chamaecrista fasciculata, listed as having an overlapping native range of  “Central & E. U.S.A” – still a big footprint.  Though POWO reveals a litany of 49 published synonyms, many of which are proposed varieties, this modern, queryable Kew Index accepts no infraspecific taxa.  What factors are at work such that two similar plants, with massive and overlapping ranges, would be regarded so differently – one with 13 varieties and subspecies, the other with none?

I leave that to the specialists.  For now, until I have more information on other species of Chamaecrista in this area, it’s satisfying to have delved into the past of this one taxon.

The Main Story – Chamaecrista fasciculata

The plant I once called Cassia fasciculata, but now deem Chamaecrista fasciculata, gives every impression as one of those delightful but worrisome non-native “weeds” that have become common components of the ruderal flora.  It germinates readily in any disturbed soil in my garden, and matures rapidly.  I see it also in waysides and cleared lots.  The plant appears to thrive on disturbance. (Note: the best description I found on the Web is at plant.orghttps://bplant.org/plant/578).  

Keys to Florida plants take the reader directly to a finally choice between Cassia, Senna, and Chamaecrista, then lead to a dichotomy that allows choice based on flower size—if the flower is under 1 cm, that is Chamaecrista nictidans.  

If it’s over 2 cm, you are, indeed, looking at Chamaecrista fasciculata, which is one of those herbs that wilts readily, but recovers in a vial of water.  Left to its own devices, C. fasciculata is abundant and floriferous.  I let hundreds of seedlings develop in around our house this year, to get a better measure of their behavior.  In habit, Partridge Pea has an open architecture,.  Lacking a strong leader, seedlings quickly develop several main branches that yield what arborists would call a “deliquescent” form.  The point at which branches develop varies; buds from nodes close to the ground may take off, or the main stem could reach 10” (and over 12 nodes) by the time side branches emerge.  Regardless, the product is an arching many-stemmed herb.  Indeed, when in flower, the final effort yields what is, in effect, a large, leafy, compound inflorescence.

Though leaves are generated in various ranks about the stem, foliage splays out to make planar surfaces, each leaf being everything you’d need to define an evenly-pinnate exemplar (botanists would describe this as ‘paripinnate’, with paired leaflets terminating the blade).

They even have evident stipules, which is great for teaching – not all plants make stipules. Touching the foliage, you might note leaflets closing upwards from the base, folding at the at their stalks along the rachis (midrib) – or nothing may happen.  This tardy nyctinasty (leaf movement) fails to impress as completely as what we see in other Legumes, such as Mimosa, but it seemed memorable for E. L. Greene (you’ll have to read his article).  The leaves do close during darkness.

Photo taken late at night of a branch from Chamaecrista fasciculata showing folded leaves
Abaxial view, showing “supra-axillary” origin of the short inflorescence

Flower buds form in short, supra-axillary inflorescences (up the stem, not tucked into the axil), each stalk producing sequential flowers that open to the abaxial (lower side) of the leaf plane. That means you’ll need to squat or lie down to take in the spectacle of a line of many bright yellow corollas of several flowers.  The buds, as Greene noted, are distinctly narrowly pointed.

Note the fully-developed bud hovering above the open flower

If you visit a population early enough in the morning, you’ll enjoy the organized way in which stamens are asymmetrically deployed, but hang around for a bit and battalions of larger bees, feeding on pollen (there is no nectar reward), maraud the blossoms, leaving anthers disheveled.

Figure borrowed from Wolfe & Estes (1992).

I got my lessons in floral morphology from some delightful research published in 1992 by Wolfe and Estes who interpreted the form as well-evolved for pollinators.  The corolla is marked by a cucullus, which is the non-conforming petal that arches forward, hovering over the cluster of 9 stamens that stand apart from the style and the tenth, lone (“deflexed”) stamen. The sheltered stamens are sometimes called “fodder” anthers, with an assumption the flower offers their pollen as food reward to bees, the other stamen (i.e. anther) being the “pollinator”.  Wolfe and Estes, however demonstrated pollen to be equally viable, regardless as to source.  The telling discovery came with altered flowers, from which researchers removed the cucullus before visitation, with pollination success in altered flowers plummeting compared to the normal. 

Individual flowers from the same plant, but showing reversed orientation, with the flower to the left positioning the cucullus to the left, the reverse being true for the flower to the right in the photo.

If you read E. L. Greene’s article in the Appendix, you’ll see that he describes the cucullus as being to the left-hand side of the flower, but in our area, the floral pedicels might twist in either direction, disposing the cucullus to the left, or to the right.  That happens even on different branches of the same plant.

Greene also calls out the fruit as distinct in character from both Cassia and Senna. However, the Wunderlin & Hansen key tells us the fruit of Chamaecrista are “elastically dehiscent”, which differs from Greene’s characterization. I’ll need to sort this out as the season progresses. After all, it is written: “By their fruit, ye shall know them.”

Sequel

Something about C. fasciculata doesn’t seem so very serious.  It’s annual, having a weedy, light-weight look about it.  And, it’s an outlier. Kew’s Plants of the World OnLine currently lists Chamaecrista as a very large genus of 367 accepted species. We learn that our herbaceous representatives, on which the genus is based, are fringe elements, temperate relatives of the core diversity comprising woody plants in South America. One wonders how long that will hold, taxonomically. Is it possible, in any realm of scientific evidence and political suasion for trees native to Brasil to remain classified in a genus based on a small herbaceous plant specimen held in a British museum that had been owned by a Swedish botanist, but was collected in North America, then authored by a German and validated by a mildly-egotistical American botanist who abandoned his life as an Episcopal priest?

Despite its modest presence and minor status in a huge genus, do not be deceived; Chamaecrista fasciculata has become a biological heavyweight.  While our taxonomic heads were turned, plant ecologists and physiologists have fallen head-over-heals for Partridge Pea.  Heading down this rabbit hole doesn’t fall comfortably in a simple article introducing plants of the local flora.  But I can’t ignore the importance of this plant is various realms of cutting edge research.  Chamaecrista fasciculata research and potential has application in studies of nitrogen fixation in Legumes (genetic basis of “nodulation”), re-imagining the value of certain characteristics in plant systematics, dismantling the long-lived classification scheme that holds Caesalpinoid legumes together, and potential for development of economically-valuable crops.  In the Appendix, I’ve cited a few papers about this Future, including extracted text that (I hope) provides a sense of research direction.

Appendix

The Future

Singer, Susan R., Sonja L. Maki, Andrew D. Farmer, Dan Out, Gregory D. May, Steve B. Cannon and Jeff J. Doyle, 1009.  Venturing Beyond Beans and Peas:  What Can We Learn from Chamaecrista?  Plant Physiology 151(3): 1041-1047.  http://dx.doi.org/10.1104/pp.109.144774

As a nitrogen fixer, Chamaecrista fasciculata likely has advantages in our nutrient poor soils.  But the story of C. fasciculata nitrogen fixation has made this plant important internationally.  Here, for example, is a 2009 statement from a publication by Singer, et al:   Among the 330 species of Chamaecrista, practical considerations make C. fasciculata well suited to be a model species. It includes both annual and perennial genotypes, is small and herbaceous, making it tracta- ble in laboratory and field settings. Related Chamaecrista species have small genomes, approximately 650 Mbp.

“There is growing interest within the legume community in C. fasciculata as a complementary legume model for a number of reasons, including phylogenetic position, nodulation within a clade of limited nodulating spe- cies, nonpapilionoid floral morphology, herbaceous growth habit, and tractability in laboratory and field settings. “

WTS (Whole-transcriptome sequencing) from shoots, roots, and nodules provides the opportunity to explore the gene space of C. fasciculata and begin addressing questions about genome evolution in legumes, the origins of nodulation, and floral development in nonpapilionoid legumes. Emerging functional genomics tools, a range of C. fasciculata populations, and venues for undergraduates to participate in genomics research will leverage the value of the WTS resource. Our approach to developing C. fasciculata community resources models a culture shift where the big science of genomics provides potential opportunities to integrate research and education at big and small schools, enabled by outsourcing of increasingly affordable high-tech, high-throughput science.”

Janet I. SprentJulie ArdleyEuan K. James, 2017.  Biogeography of nodulated legumes and their nitrogen-fixing symbionts.  New Phytologist 215:40-56. First published: 17 February 2017 https://doi.org/10.1111/nph.14474

“In the last decade, analyses of both molecular and morphological characters, including nodulation, have led to major changes in our understanding of legume taxonomy. In parallel there has been an explosion in the number of genera and species of rhizobia known to nodulate legumes. No attempt has been made to link these two sets of data or to consider them in a biogeographical context.”

It is generally agreed that the earlier division of legumes into three subfamilies, Caesalpinioideae, Mimosoideae and Papilionoideae, is no longer appropriate, the Caesalpinioideae in particular being paraphyletic… Clearly, flower morphology is no longer a safe taxonomic character, whereas nodule characters are arguably (this review) more reliable.

Tucker, Shirley C., 2003.  Floral Development in Legumes,  Plant Physiol 131: 911–926

Species of flowering plants are most reliably identified by their flowers, the sexually reproductive or- gans. A flower is similar to a vegetative short shoot (lacking appreciable internodes) that bears four kinds of laterally attached organs in successive whorls: se- pals, petals, stamens, and carpels. Significant floral distinctions among plant families include symmetry , whether organs are organized in whorls or helically, number of parts per whorl, carpel position relative to the surrounding organs, fusion among organs within a whorl or between different whorls, and whether both male and female organs are present in the same flower.

A flower, like a vegetative shoot, has a terminal floral apical meristem that initiates organs laterally, usually in acropetal succession (although exceptions are common in some taxa): sepals first, followed by petals, stamens, and carpels. Because each flower lives for a very short time, the floral apical meristem is determinate, meaning that it ceases activity after a certain number of organs have initiated. Vegetative apical meristems, in contrast, are usually indeterminate, continuing to initiate new organs, such as leaves, indefinitely.

Shared assemblages of specialized characters gen- erally are viewed as evidence of close evolutionary relationships. Nevertheless, in one genus, Cassia sensu lato in the caesalpinioid tribe Cassieae, shared specializations are deceptive. Howard S. Irwin (Long Island University, Greenvale, NY) and Rupert C. Barnaby (New York Botanical Garden, Bronx, NY) (Polhill and Raven, 1981, and refs. therein) split Cas- sia into three genera: Cassia sensu stricto, Chamaec- rista, and Senna. These are superficially similar and share many specialized characters such as yellow flower color, pentamerous corolla, zygomorphy, dor- siventral heterostemony, and poricidal stamen dehis- cence. Comparison of floral ontogenies (Tucker, 1996) showed marked developmental differences in their inflorescence architecture, phyllotaxy, bracteole formation, order of organ initiation, amount of over- lap in time between whorl initiations, the basis for asymmetry, and the basis for poricidal dehiscence. These differences in development strongly suggest that the superficial similarities in the flowers have resulted from evolutionary convergence.

Bueno, Erika, Ted Ksha, Sonja L. Maki, Eric J. B. von Wettbert, and Susan Singer. 2019.  Genetic diversity oof Chamaecrista fasciculata (Fabaceae) from the USDA germplasm collection, BMC Research Notes s12:117. DOI.org/10.1186/s13104-019-4152-0

Note that we think of Chamaecrista as a genus of small herbs, though literature tells us this is a large genus of over 330 species [367 accepted species in POWO currently}, with woody trees and shrubs in Brasil being at the center of diversity.  Bueno, et al condense this a bit too much (ignoring our herbaceous species), but make a point:  “As the only temperate annual in a large tropical tree genus, a wealth of information exists on the ecology of C. fasciculata including the characterization of locally adaptive traits in response to climate change, key pollinators, and gene flow and genetic structure among naturally occurring populations”

Chamaecrista fasciculata is a widespread annual legume across Eastern North America, with potential as a restoration planting, biofuel crop, and genetic model for non-papillinoid legumes. As a non-Papilinoid, C. fasciculata, belongs to the Caesalpiniod group in which nodulation likely arose independently of the nodulation in Papilinoid and Mimosoid legumes. Thus, C. fasciculata is an attractive model system for legume evolution. In this study, we describe population structure and genetic diversity among 32 USDA germplasm accessions of C. fasciculata

Holly L. Williams and Charles B. Fenster, 1998.  Factors Contributing to the Low Frequency of Male Sterility in Camaecrista fasciculata (Fabaceae), Am. J. of Botany 85(9):  1243-1250  https://www.jstor.org/stable/2446634

Male sterility, in studies such as this, might provide more understanding as factors in the evolution of monoecy and dioecy.  “Bumble bee pollinated Chamaecrista fasciculata provides pollen as the sole reward to it pollinators.  Male sterility expressed as an absence or nearly complete absence of pollen production, occurs in low frequency in populations of C. fasciculata.  Here we describe experiments using C. fasciculata, to examine frequently cited determinants of the spread and maintenance of male sterility: compensation and the genetic basis of male sterility….”

The Past

‘The Genus Chamaecrista’ E. L. Greene, 1887.  Pittonia 3(17)

Note:  Throughout, Greene uses “fascicularis” rather than  Michaux’s “fasciculata”  

In the early years of my herborizing in these Eastern States no summer-blooming herbs had so strong a fascination for me as our annual sensitive-leaved Cassias; and after very many years of travel and residences afar beyond their habitat, I but lately renewed acquaintance with them.

In the far West and Southwest I had, in the long interval, grown familiar with other Cassias of different habit, different foliage, and different flowers, insomuch that on first beholding again a luxuriant growth of what is called C. chamaecrista in full flower, I found its floral structure not only obviously very unlike that of any typical Cassia species, but even very perplexing. In order that each one of  my readers, even those not conversant with the plan of the flower in leguminous plants, may see how I could be perplexed in this instance, I must state that in all normal and typical Caesalpineous Leguminosae the flower, though almost regular and roseous, so to speak, as to its corolla, is nevertheless just like the papilionaceous flower in its plan.  There is in the corolla of both one odd petal which is uppermost and may in either family be called the banner or vexillum.  next below the banner comes a pair of petals called wings; then a lowermost pair denominated the keel.  In shorter phrase, the ordinary caesalpineous corolla, though quite regular, is like the papilionaceous corolla, made up of banner, wing and keel petals; the banner being always an odd petal and uppermost in position.  Now the puzzling circumstance in the case of my Chamaerista corollas, as newly observed after having become familiar with those of real Cassias, lay in the obvious fact that, in the place of one lone petal at the top of the flower where the banner should be, there was a pair of petals.  Then at the base of the flower, where a closely matched pair should be, in place of the papilionaceous keel-petals, there was seen to be a solitary petal larger than any of the others.  At this point I seemed at first to have solved all difficulty by assuming that in this species the flower is inverted, turned up side down by a probable twist in its pedicel.  Such inversion of corollas is not rare in the plant world; though it never occurs in the same genus with plants having flowers in normal posture.  It is the mark more often of an entire family or natural order than even of a genus.  The Violaceae, the Lobeliaceae and the Orchidaceae are familiar examples of large families, one of the essential characters of each of which is an inverted corolla.  Logically, therefore, this new discovery would separate the Chamaecrista group, if not altogether from the family of Caesaplineae, at least from Cassia, to form a distinct genus.  Yet hitherto, as another day’s observation proved, I was in error; and my perplexity deepened as I was driven from my assumed theory that the Chamaecrista corolla is inverted.  That larger odd petal at the bottom of the flower proved to be by no means the banner, but one of the wing-petals, its mate being one of those two small ones at the top of the flower.  The real banner-petal, perfectly demonstrated as such by its position in the bud as enfolded by the wings,  is the large concave lateral petal at the left-hand side of the flower [Note – this is the cucullus, and I find it can also present in a right-sided position].  In the accompanying illustration Figure 1 represents very accurately the corolla of Cassia marylandica, b indicating the banner-petal, w w the wings, k k the keel.  Figure 2 as perfectly displays the same organ in what is called in our books Cassia chamaecrista, the same letters indicating the petals by name.  

It will thus be seen that what marks so strangely the Chamaecrista corolla is not an inversion, but something still more interesting and curious, namely a torsion of ninety degrees to the left—a character which, by itself alone, if considered in connection with the peculiar habit of the plant and its rather numerous allies, would establish the whole group as securely in the rank of a distinct genus as any genus could need to be established.  I shall, however, here enumerate the actual characters of the genus CHAMAECRISTA as I have recently observed them in living plants, and in dried specimens of many species.  (1.) Flowers axillary or supra-axillary and solitary or few and fascicled, never terminally clustered as in Cassia. (2.) Buds slender-conical and acuminate (always subglobose or ovoid and obtuse in Cassia.)  (3.) Sepals plane, slenderly acuminate, then-membranous (in Cassia firm-herbaceous obtuse concavo-convex).  (4.) Flower on a twisted pedicel, its banner and keel petals thus made to appear lateral, and one wing enlarged and placed lowermost, the other reduced and becoming uppermost.  (5.) Pods thin, compressed, very promptly dehiscent, never subterete and indehiscent as in most or all Cassias.

The genus was founded by Johann Commelin, just two centuries ago (1697), on a West Indian plant, this same becoming the type of Cassia chamaecrista, Linn.  Linnaeus suppressed the genus.  Moench restored it in 1694 [sic, its 1794], and seems to have insisted on the large size of the lower petal (the left-hand wing petal) as the most essential character; so that he certainly observed the structure of the corolla to be widely at variance with that of Cassia.  In the course of the present century a number of authors seem to have maintained it; but in so far as I have been able to read the history, the real, and superlatively numerous characters of the genus are here for the first time clearly indicated.

I do not attempt to transfer to Chamaecrista any more than a small proportion of the species which belong to it.

Large-flowered species, all the petals spreading, only moderately unequal.—Chamaecrista proper.

  1. C. PAVONIS, Cass.  Dict. viii, 78 (1817).  Cassia chamaecrista, Linn, not of recent American authors.  Chamaecrista pavonis major, Commelin, Hort. Amit. i. 53, t. 37 (1697).  Plants indigenous to the West Indies and perhaps northern South America; the type of the species, according to all the figures cited by linnaeus, having solitary supra-axillary flowers.  The Virginian plant referred here by Linnaeus is in all probability the next species  Though possibly either Cassia robusta, Pollard, or his C. mississippiensi, or C. simpsonii may prove identical with typical Linnaean C. chamaecrista, and more or less recently introduced from Jamaica or Barbados, from the island to which Linnaeus credits his type.
  2. C. FASCICULARISCassia fascicularis, Michx. Fl. i.262 (1803).  C. chamaecrista of American authors, scarcely of Linnaeus.  The very considerable fascicle of flowers, as well as the other characters assigned this species by the extremely critical and careful author of the Michaux Flora preclude my accepting the doctrine that this plant can be identical with the first species of this series.  Mr. Pollard trusts that Mr. Bentham was correct in suppressing it; but all who know Mr. Bentham’s views and methods will doubtless agree with me that he would have been likely to have reduced Linnaeus’ C. chamaecrista every one of these large-flowers species which Mr. Pollard has proposed as new.  C. fascicularis I suppose incudes all of the northern plant with the large corollas; and my own opinion is that all Mr. Pollard’s new species are perfectly distinct from this; though, as I said above I suspect that some one of them may be the true Chamaecrista pavonis.
  3. C. DEPRESSACassia depressa, Pollard, Bulll. Torr. CLub xxii. 515, 5. 251 (1895).  This Floridan species bears solitary flowers, and is thus allied to C. pavonis; but its habit is so entirely peculiar that I cannot doubt its perfect validity as distinct from the old original type of Chamaecrista, whoever that may ultimately prove to be. [ISB sinks this into C. fasciculata]
  4. C. FLAVICOMACassia flavicoma, HBK. Nov. Gen. v9 366(1823).  A most beautiful South American representative of the genus; apparently a large plant, and shrubby.
  5. C. TRISTICULACassia tristicula, HBK. l.c. 267.  Near the last, and of similar geographical range.
  6. C. PALMERICassia palmeri, Wats. Proc. Am. Acad. xx99. 408(1887).  A neat representative of the low suffrutescent type of Chamaecrista, not rare in middle and northern sections of Mexico; the flows as large as in the larger species.

**Small-flowered species; one of the petals much larger than the others and spreading, the other four minute, erect, forming a cup.—Genus NICTITELLA, Raf.

7. C. NICTITANS, Moench, Meth. 272 (1794), excluding the synonym C. chamaecrista, Linn.  Cassia niditans. Linn. Sp. 30(1753).  Of the northern U.S., with about the same general range as C. fascicularis.

8. C, ASPERA. Cassia aspera, Muhl. in Ell. Lk. i.474 (1821).  Thoroughly distinct from C. nictitans, as Elliot demonstrated, and as Mr. Pollard has reasserted.  Peculiar to the southern U.S.

9. C. MULTIPINNATACassia multipinnata, Pollard., Bull. Torr. Club. xxxii. 515, 5. 320 (1895).  A Floridan species; and Mr. Pollard’s variety Nashii looks as if it might well be specifically different; but I have not seen the living plant. [C. nictitans in ISB]

Literature

Michaux, André, 1803.  Flora boreali-americana, sistens caracteres plantarum quas in America septentrionali collegit et detexit Andreas Michaux, Source: Missouri Botanical Garden, Tomus 1: https://www.biodiversitylibrary.org/page/404743  –  Tomus 2: https://www.biodiversitylibrary.org/page/410655

Muench, Conrad, 1794.   Methodus plantas horti botanici et agri Marburgensis :a staminum situ describendi.  Source: Missouri Botanical Garden.  https://www.biodiversitylibrary.org/page/416875

Greene, Edward L., 1905.  DERIVATION OF THE NAME CHAMAECRISTA, Torreya, Vol. 5, No. 7 (July, 1905), pp. 126-128 Published by: Torrey Botanical Society

Stable URL: https://www.jstor.org/stable/40594402

Greene, Edward L.,    ‘The Genus Chamaecrista’ , Pittoniaa  series of papers relating to botany and botanists., v. 3 (17): 238-243Source: Missouri Botanical Garden https://www.biodiversitylibrary.org/page/15253790

Wolfe, Andrea D. and James R. Estes, 1882.  Pollination and the Function of Floral Part in Chamecrist fasciculata Fabaceae), Am. J. Botany 79(3): 314-317.  https://www.jstor.org/stable/2445021

Miller, Philip, 1754.  The gardeners dictionary : containing the methods of cultivating and improving all sorts of trees, plants, and flowers, for the kitchen, fruit, and pleasure gardens, as also those which are used in me…  Vol. 3 Source:  New York Botanical Garden, https://www.biodiversitylibrary.org/page/44045774 Vol 1 Source New York Botanical Garden, https://www.biodiversitylibrary.org/page/44045023

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