Varieties in S. flava: a fundamental question.

[kiwiearl]

Below is a photo of a nice S. flava var. cuprea, a "Coppertop".

I use the photo simply because it is a really nice example of the variety but the main reason for this post is to ask what I feel is relevant question as it relates to the debate on the all red flava plants in Western Florida. Specifically, I'm referring to the individuals within an otherwise broad population of fundamentally var. rubricorpora (variable as that variety inherently is) that whilst exhibiting every other aspect of that variety become fully red and are therefore accorded by some the title of var. atropurpurea as that has long referred to those all red flava found as a distinct entity in the Green Swamp, etc of the Carolinas. There are those, such as McPherson, who state that an individual var. atropurpurea can indeed be found within stands of var. rubricorpora despite their common beginning and factors beyond colour alone.

As I've done previously elsewhere, I contend that a specimen that starts out as every other var. rubricorpora to which it is clearly intimately related in the population (genetically) but develops colour over and above its more typically nominal relatives can't then have become two varieties within the same season; two things at once so to speak. And, that this highlights the potential frailty of what one might describe as the variety within varieties element within some promoted Sarracenia nomenclature.

As most are familiar with, a large percentage of S. flava that present initially as var. cuprea will fade ie, lose the copper colour to their hood and upper pitcher, often to the degree that no copper colour is evident past a certain point in the season. Often this can occur quite rapidly and disappointingly for the horticulturist.

The question I put is this: If following the rationale that a var. rubricorpora can legitimately be latter described as var. atropurpurea over the course of a season, can a var. cuprea, once faded, be just as legitimately said to have transformed into var. flava or var. ornata once the copper colour has been lost? Certainly I, for one, have never heard that stated.

[hcarlton]

I think what it starts out as should be what it stays labeled as, if we know the origins of the plant and how it looks over its lifetime. The color varieties in flava certainly are confusing and really at this point only important to horticulture at the moment, and location data seems far more stable and important. I believe at this point that color varieties can have the current names be kept, if we stick with what the original color and pattern of the plants is expressed as. A faded cuprea is still a cuprea, and often passes the trait on in hybrids, and a rubricorpora that turns solid red is just a rather fetching rubri in the end. At this point I also think that the individual plants in Fl that open fully red may need a separate name since they are so distant genetically and location-wise from the Carolina atros, as you and others have suggested on here and other forums, and due to their distinct color throughout the pitcher life from other very dark rubricorpora.

[kiwiearl]

I should make it clear that my question re. var. cuprea is entirely rhetorical. I don't buy the two things at once premise on which it is somewhat mischievously founded at all.

I recognise the logic in according variety status to those representations in S. flava that are repetitively enduring and distinct. The var. cuprea (as in my photo above) of Horry Co., South Carolina are a good example. See these Brad Wilson images of them in situ within the Lewis Ocean Bay Heritage Preserve: link and link. Note how they occur distinctly but intimately with S. flava var. flava. Under plant nomenclature, to describe cuprea as a variety in the circumstances in which they exist is a legitimate position on which to stand.

The variety rugelii is perhaps the most strong case upon which to argue the ground of variety status, occurring as it does in intimate proximity to other associated/related but different representations of S. flava and within geographical parameters. The named variety rubricorpora is one obviously associated differing representation whereby populations of like characteristics, in that case the significant infusion of red, exist alongside var. rugelii such as in the case of Liberty Co., FL. As seen in situ those red plants in question typically exist as definable populations rather than the odd red here or there. They are a body of repeated characteristics that not only include distinct colour combination presentation (as is also the case with var. rugelii) but additionally a distinct growth pattern in respect of the reduced number of pitchers produced. This led Schnell to derive the variety name rubricorpora to describe the collective phenomena.

Further however, those var. rubricorpora populations when viewed as a whole are found in terms of spectrum to be significantly variable in respect of the nominal colouration description of the variety, some having a good deal less strong representation of the yellow hood/red tube whilst some have developed a good deal more red infusion that renders them eventually entirely red. Nonetheless, and notwithstanding those entirely red individuals, they are all representations of the same expression of the populations from which Schnell recorded his description. This spectrum of variability is not only seen in situ but repeated ex situ in specimens grown from wild sourced seed. Indeed, McPherson in his earlier book, Pitcher Plants of the Americas, noted this colour diversity (reinforced with a photo line up) within var. rubricorpora.

However, within the scope of his following volume, Sarraceniaceae of North America, as a part of a sum in defining Sarracenia species into ever more varieties he proffered that any fully infused individuals within a var. rubricorpora population were in fact, or had become over time, var. atropurpurea. In my view, that was unjustified when all elements (not solely colour at any given time) of the individual specimen and its existence not only within but as part of the population are taken into account. I submit McPherson was correct the first time. And, that it remains amply sufficient to recognise that the botanical variety name rubricorpora, if utilised, is representative of what is a broad spectrum existing in specific locations and should be left at that. In all reality, if not accepting that, then a population of var.rubricorpora could conceivably be broken into myriad subjective varieties (arguably as nominated in the above work for other Sarracenia species). How far do you take it?

Of course, sub the repeated distinct representations in question are those blends between any two and the expression of downstream genetic characteristics as a result. Should they be accorded a variety name as each is encountered? Or, perhaps that is best left to Western Florida's Everyman to shoulder: var. ornata....

[Aidan]

I raised somewhat related concerns quite some time ago and without specifically addressing your examples, I have come to the (very personal) conclusion that the division of S. flava into varieties sows confusion rather than offering any clarification. This is most particularly true at the red end of the scale.

No doubt this view will be considered heretical, but the continued taxonomic hair-splitting of the Sarracenia into apparently never ending varieties and forms (and god help us, "new" species) does neither the plants nor those with an interest in them any favours.

[John Brittnacher]

On the bright side, at least Sarraceniaceae of North America didn't define the color form varieties Sarracenia purpurea var. rubricorpa, Sarracenia purpurea var. atropurpurea, and Sarracenia purpurea var. ornata. They certainly could have.

[ICPS-bob]

An excellent discussion on this topic is
Ellison, et al. 2014. Pitcher Plants (Sarracenia) Provide a 21st-Century Perspective on Infraspecific Ranks and Interspecific Hybrids: A Modest Proposal for Appropriate Recognition and Usage.
www.people.fas.harvard.edu/~ccdavis/pdfs/Ellison_et_al_Syst_Bot_2014.pdf

[kiwiearl]

Muchas gracias Bob. As you say, an excellent discussion in respect of variety status in Sarracenia. The horticulturally founded emotive according of increasing variety distinctions has become out of hand, in my view. The paper provides a worthwhile examination of this.

That said, outside of strict taxonomy, it is understandable that the colloquial usage of the nominated variety names has, for those who subjectively recognise visual differences, value. The dissemination of Sarraceniaceae of North America, for example, and its propositions via the promotion of and liberal application of variety names in its content (particularly the new delineation of S. alata - which I believe to be unjustified - following on as it does from and, in a sense, reliant on some long standing variety descriptions in S. flava) has seen them widely accepted and vigorously utilised by Sarracenia enthusiasts.

If I may quote him, John Brittnacher put this well, I believe, in a contribution on Sarracenia Forum link:
"Unfortunately real life can not always be reduced to real taxonomy in a way that makes sense because of the rules and intention of taxonomy. There are cases where I know that certain sets of plants are different and should have different names. However under the rules of taxonomy it is impossible to give them separate names because there aren't any significant taxonomic characters...to distinguish them."

Further, in the event one wishes to discuss Sarracenia with the wider Sarracenia/Carnivorous Plant community one is bound to have to use the variety element of nomenclature in order to be pertinent. I may not agree with the variety delineations, but, as sure as you like, I have to use them in discussions. Whilst I have debated the use of variety names for a good long while, I end up using them in reference, like it or lump it. Similar as the way the world is increasingly been herded toward the usage of Facebook in order to interact as significantly more and more suppliers and services enlist it as an interface toward consumers with other means becoming redundant (said by a non-Facebook user).

[Dave Evans]

Aug 7, 2014 5:33:03 GMT kiwiearl said:

The question I put is this: If following the rationale that a var. rubricorpora can legitimately be latter described as var. atropurpurea over the course of a season, can a var. cuprea, once faded, be just as legitimately said to have transformed into var. flava or var. ornata once the copper colour has been lost? Certainly I, for one, have never heard that stated.

No, this plant is S. flava cuprea regardless of any other features. Any other variety format can be "copper topped" and thus qualify as S. flava cuprea except for S. flava atropurpurea.

Also, S. flava atropurpurea is not the correct name, it is really S. flava x (S. flava x S. purpurea venosa)² and not with S. purpurea venosa burkii(!!)--that makes a different looking hybrid with different colorations. S. flava atropurpurea is likely a backcrossed variety of S. x catesbaei.

Dr. Schell observed some plants in Sumatra which are segregated into his varieties. However, I have looked at the same plants are believe they are simply old "fairy rings" where one really old clone has spread outward in all directions and center has rotted away making most of the plants present one clone, not one variety! Some of Dr. Schnells varieties are not botanically valid in my opinion. They are more accurately described as wild collected cultivars.


Also, S. flava rubricorpora and S. flava atropurpurea do not overlap! One is a hybrid with pururea venosa and there is no venosa down in Florida. S. flava rubricorpora that start to resemble S. atropurpurea cannot be considered the same. The pattern is completely different. S. flava atropururea is a solid dark purple color whereas S. flava rubricorpora has reddish purple lines that crisscross and overlap.

[ICPS-bob]

Aug 16, 2014 2:15:59 GMT Dave Evans said:

Also, S. flava atropurpurea is not the correct name, it is really S. flava x (S. flava x S. purpurea venosa)² and not with S. purpurea venosa burkii(!!)--that makes a different looking hybrid with different colorations. S. flava atropurpurea is likely a backcrossed variety of S. x catesbaei.

Dr. Schell observed some plants in Sumatra which are segregated into his varieties. However, I have looked at the same plants are believe they are simply old "fairy rings" where one really old clone has spread outward in all directions and center has rotted away making most of the plants present one clone, not one variety! Some of Dr. Schnells varieties are not botanically valid in my opinion. They are more accurately described as wild collected cultivars.

Also, S. flava rubricorpora and S. flava atropurpurea do not overlap! One is a hybrid with pururea venosa and there is no venosa down in Florida. S. flava rubricorpora that start to resemble S. atropurpurea cannot be considered the same. The pattern is completely different. S. flava atropururea is a solid dark purple color whereas S. flava rubricorpora has reddish purple lines that crisscross and overlap.

At some point in taxonomic discourse, opinions need to be published in the refereed literature and debated. And, even then, the final arbitrator is time. Over time, often decades, opinion solidifies then is overturned by a more convincing argument, which itself may eventually be replaced. There is no absolute "truth" in this business.

[kiwiearl]

Bob is spot on.

Authors have, and anybody can as time marches forward, nominated endless descriptions as they see fit as an ongoing effort to subjectively define Sarracenia more 'precisely'. Someone (and credit to them for their work) publishes their position on the delineation of the genus. Fine. But in comparative biology such propositions are never results set in stone and it is the preserve of consumers of the work to, through their own analysis, accept or reject the conclusions. The consideration of what is "valid" or "not valid" is inherently the prerogative of those an author delivers their submissions to. Indeed, the essence of authored scientific extrapolation is review with resultant questioning; authors expect challenge as part of the evolution of understanding. Naturally, a consensus may or may not develop in response.

Certainly, whatever its physical volume, it would be fraught to accept any publication as biblical despite this being a common phenomenon. By that measure, as Aidan touches on, it is not heresy to disagree. The floor of the colossal vault that contains Biology's past phylogenies, descriptions etc is neck deep in the rejected or superceded.

A legitimate consideration is to what degree the vigorous factor of horticultural thinking, translated desire perhaps, is superimposed on analysis of wild Sarracenia. In my view, in league with others, the promotion of an ever increasing delineation of Sarracenia on the variety level based on what may at root be horticultural enthusiasm is more than open to question.

Here is perhaps another way to look at it: the installation of varieties in Sarracenia based on colour alone is something we humans as, above all other senses, very visual creatures are more than prone to feel a requirement for. The question as to colour being or not being valuable represents this well. In fact, confronted visually it requires a degree of discipline to rigidly stick to the position that subspecies is the rung below which it makes no taxonomic sense to descend. A useful way to put this may be: the colloquial use of variety names based on colour allows for the identification of difference in a way that meets with human sensitivities. Although emotive, it seems to make sense to many to recognise those differences formally. Sure, not strictly taxonomic.

Its pretty obvious I have problems with McPherson's position on variety status - the variability amongst most of the red and veined S. flava and S. alata of the Gulf Coast is endless within a measured square area and, as has been argued, could ultimately be broken down into a zillion varieties if you are going to break it down at all. As John B points out, the same variety breakdown potentially but without credibility could have been applied within S. purpurea.

Then again, for the most part rugelii can seem a pretty darned hard and fast representation of the replication of colour traits that could be recognised as enduringly distinct. Trouble is, breed two rugelii or one with itself and you can get, say, a heavily veined individual expressed as offspring. Taxonomy says, no matter - they are all S. flava and the genetics of the distinct morphology of the species (colour as a determinant of variety aside) produce a range of colour patterns.

Tying in as it does with what Ellison, et al are saying, colour morphs in zoological species are not accorded variety status ie, a black jaguar is still a jaguar; there is no variety delineation beneath that fact. Same applies to all black and all white King Penguins (amazing looking things amongst the multitudes - check 'em out), black leopards, king cheetahs, melanistic pheasants etc. Canis lupus, the Gray Wolf is a good example: there is the median coat colour pattern, but there are individuals that present as entirely black, entirely white and in between within populations. Zoological taxonomy does not describe these variations on the theme as varieties. Tigers? A number of subspecies exist/existed due to geographical isolation and all that that delivered over time. The inherently variable patterns between individual tigers in each subspecies is well documented. But, varieties? No.

I agree with the authors of the paper. However, at street level I anticipate these variety names, as with any vernacular language, are here to stay (as they have been in S. flava for some time prior to McPherson's revision) and everyone will have to roll with them or not as they see fit. They are readily consumable, particularly by the enthusiastic horticulturist. And, to that end, they are notably and significantly relied on by the retailer. To many, Sarraceniaceae of North America contains THE definitive comparative analysis of the genus Sarracenia and disciples carry its message wherever they may travel. Others have assessed flaws in its propositions. No problem, that's science for you and as it should be.

[Dave Evans]

Aug 16, 2014 5:34:49 GMT ICPS-bob said:

At some point in taxonomic discourse, opinions need to be published in the refereed literature and debated. And, even then, the final arbitrator is time. Over time, often decades, opinion solidifies then is overturned by a more convincing argument, which itself may eventually be replaced. There is no absolute "truth" in this business.

But these things are measurable. When you can only reliable reproduce these kinds of "varieties" via asexual reproduction, then they aren't really varieties, just ideas. Dr. Schell says his varieties aren't taxonomically stable--when the author tells you this, you should believe it. There might be some pattern to it with some varieties being abundant in some areas and rare in others. But basically where S. flava overlaps with S. purpurea there is one set of varieties within flava and where S. flava overlaps with S. rosea there is a second set of varieties within flava.

Because of the timing of the flowering, flava and purpurea hybridize a lot; so do flava and rosea. Very colorful flava could simply be descendants of S. x catesbaei in the north; and descendants of S. x naczii in the south. See the pattern shown by S. naczii? That is where the S. flava rubricorpora comes from.

www.google.com/search?q=Sarracenia+naczii&tbm=isch&tbo=u&source=univ&sa=X&ei=TonyU86yNI75yQSvxoDQAQ&ved=0CC0QsAQ&biw=1045&bih=642