Post by waxy on Jan 16, 2014 13:15:26 GMT -5
Ok it's time this stimulating discussion had a thread of its own. It hasn't been just about red flavas in the wild for some time.
Perhaps Jonathan can/will move some of those posts here.
John, Thank you for posting the new Ellison et al paper in the science category:
A MODEST PROPOSAL FOR INFRASPECIFIC RANKS sarracenia.proboards.com/thread/1042/modest-proposal-infraspecific-ranks
Some interesting statistics and positions to digest and much fertile ground for discussion. Quick reaction - Certainly if some taxonomic ranks truly do not serve a purpose they should be abandoned, but why, after all this time have they become useless. If it is because they are no longer fashionable, for whatever reason (misunderstood, ill-defined, inconsistently applied), that is not a good reason. The problem then is with the users not the system. I will be a while studying this.
As for the question of "atropurpurea" in these infamous sites and perhaps lone individuals within var. rubricorpora stands, I still don't buy it. My views have been well laid out but put basically I don't accept the variety name based on the following while fully recognising you have visited the plants in person:
1) S.flava var. atropurpurea is a name assigned to a distinct population of a red form of plants in the Carolinas they being isolated from the ones in your photos both genetically and geographically.
2) Those Carolina var. atropurpurea share an intimate relationship with the region's S.flava var. flava as evidenced in this and other photos of wild plants (photo Don Schnell, CPN Volume 27, Number 4, December 1998, pages 116-120 )
A. var maxima, B. var atropurpurea, C. var flava, D. var ornata, E. var cuprea
3) The "atropurpurea" in your and other's photos are seen in an intimate relationship with S.flava var. rugelii. Clearly, there exists a foundational relationship between the red plants and var rugelii.
4) This relationship between red S.flava and var. rugelii occurs in a number of locations in the Florida Panhandle as you note. In the locations you highlight, the red plants are intermingled with or alongside var. rugelii. Indeed, looking at your photos I believe the hood form of the "atropurpurea" is distinctly Florida Panhandle - particularly representative of the var. rubricorpora broad, floppy morphology. By that proposition they are quite distinct from S.flava hood form in plants native to the Atlantic coastal plain.
5) The genetics in these Panhandle S.flava populations allow for the expression of the cut throat var. rugelii; the yellow lid red tube var. rubricorpora; and the all red plants wherever they are found. My position is that all three of these are related and their expression is a broad one. The expression of var.rugelii in these locations is, in fact, no less notable than that of var. rubricorpora or the all red plants other than being deemed the so by human beings.
6) S.flava var. rubricorpora has, since Schnell's 1998 paper naming the variety and beyond, been acknowledged as being highly variable with a wide spectrum. In his book, Sarraceniaceae of North America, McPherson notes this as "The overall colouration of Sarracenia flava var. rubricorpora is extremely variable" (p 249). In his earlier volume, Pitcher Plants of the Americas, McPherson highlights this theretofore accepted variability in the following photo line up of var. rubricorpora specimens. Note the fully infused red/purple specimen on the right (Photo Stewart McPherson)
7) I ask again, what of the two plants in this photo of my own Florida red wild sourced genetic stock that has remained isolated from interbreeding since germination from the same seed pod? What is the plant on the left? And, the one on the right? It is in a sense irrelevant which location these two clones are from outside of the fact they are from the Panhandle like so many others standing in the various habitats as we speak.
9) In closing, even if the submission is correct that some of them open as all red, I reiterate my contention that the "atropurpurea" in the Panhandle are sufficiently distinct from S.flava var. atropurpurea as found 700 miles east (a massive advance on 20 miles) that they should not share the name. Whilst I am happy to have the all red "atropurpurea" accepted as part of the var. rubricorpora spectrum with out the attached "atropurpurea" moniker, I am also open to those entirely red plants as evidenced in your photos to be classified with a variety name identifying them as something distinct within what are otherwise Panhandle genetics where they both are found and spring from. I guess the question then is: what name would be appropriate....?
10) As for the transplanted theory? I simply cannot accept that until evidence fixing the event comes to light. For all the reasons I have given above and elsewhere, I think the answer to these red S.flava is to be found by un-impassioned observation of the populations and, moreover, future genetic research to determine the nature of the exhibited relationship with var. rugelii.
I agree with Mike 100% on the varieties ultimately being of hybrid origin. Would that also mean the Carolina plants are derived with help from Sarracenia purpurea subsp. venosa and the Florida plants via Sarracenia rosea? Could that account for some of differences seen between the populations?
As far as what any given plant is called, according to the taxonomic rules if it matches the description, it is that name. It doesn't matter where you found it. We may not like the rules but thems the rules.
What I find bizarre is the description for Sarracenia flava var. ornata. It sounds like a failed Sarracenia flava var. atropurpurea and doesn't sound like it includes the nicely veined plants at the locations specified by Schnell. This is what he says:
Deep red to purple throat pigmentation nearly obliterated by very strong and heavy red venation throughout pitcher tube and lid.
That describes this plant:
But what is:
This is what I would like to consider "ornata" but it doesn't fit the description.
So here it is in short form without citations.
The red pigmentation in flava atropurpurea (both gulf and east coast forms) has been published as coming from purpurea and rosea by Phil Sheridan of Meadowview. Therefor, not a mutation but an expression of introgression and gene-flow.
Recent gene analyses for establishing phylogenetic relationships have strongly suggested alata and minor are sister species and put them into a clade together. That could explain the areoles. Perhaps a pre-divergence relict artifact. It might even explain the okefenokeensis populations.
The alata pollination study makes the point that the preferred or most common pollination strategies are 1) self- pollination (selfing), 2)pollination within the same species or variant, 3) pollination with different species, in that order. It puzzled me for some time why, in a genus that so easily hybridizes in cultivation, there were not more, obviously-hybrid plants in populations and how in these areas the entire field site did not blend into one big hybrid swarm but maintained separate species identities. Despite research data that shows outcrossing produces healthier more abundant seed and seedlings, and where there is oft repeated "common knowledge" that inbreeding(selfing) produces inferior, unfit plants, maintaining species identity by selfing may just be a more important advantage. Of course there may be other advantages to preferred selfing that we haven't figured out. The Ahh-Haa moment hasn't happened yet.
Frankly, while writing this, I've just come up with an aha of my own. Maintaining variability(separateness) in the face of quick and easy homogeneity would be a tremendous advantage.
As far as prey specificity, I wonder if it exists. Sarracenia seem to be opportunistic feeders, the kinds and numbers of captured prey varying with the prey populations themselves. They seem to attract all sorts of animals not just prey, from ants, flies, bugs, and wasps to spiders, moths, and hummingbirds. My personal experience adds blackbirds and our urban raccoons to the list with disasterous consequences (I now keep my plants entirely surrounded by netting).
While pigment is a distinct trait (it doesn't encode for other traits) it absolutely acts in communion with other distinct traits like nectar production and composition, combination with other chemical compounds, and even odor. Besides the odor of rotting flesh that these plants can certainly manifest, there have been various other odors that have been reported which may play a role in attraction (or even possibly repulsion) of certain potential prey. Except for possible chromosomal linkage, these traits are largely independent of each other genetically. When do they each become critical for the plants' well being or survival (known as a limiting factor) has not been studied in any depth, to my knowledge.
Pigmentation is often a protective mechanism, chlorophyll will burn out at high light levels or temperatures. Red pigment ebbs and flows in response to light levels that are not optimal but the chlorophyll is there, masked by the red, so photosynthesis is largely unaffected. Slow growth is a tendency in many clones/populations of Gulf coast flavas and doesn't seem to be related to degree of pigmentation. Large areas of unpigmented white however, definitely impact photosynthesis in a permanent way. They rarely become green once they have formed. There are other mechanisms the plants have that deal with low light.
Enjoying the interesting and thought-provoking comments and ideas here. Hope more get started like it. Thanks to all.
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Jan 15, 2014 19:42:27 GMT -8 calen said:
Thanks waxy. Such a stimulating thread.
John on the other forum mentioned something that I had been thinking about: that form, not varietal, status should be accorded in cases where a single gene accounts for the distinguishing characteristic in a taxon. This thought pretty much opens Pandora's Box, and I therefore humbly submit the following rant:
Flava varieties should be demoted to forms. If "albino" S. purpurea ssp. purpurea is forma heterophylla, why should over-expression of the gene warrant varietal status, as in red-tubed flavas? In the case of "var." rubricorpora there are entire populations exhibiting the red tube gene, which could give the superficial impression of a more significant taxon (and, surely, lay the foundations for future divergence from other populations), but is it really so? And of course even in Florida the red-tubes usually grow with rugelli, while in the Carolinas the special color "varieties" grow in sympatry as a total mixed bag. How could divergence between flava cuprea and flava flava occur under such circumstances? Seems more likely the "different" plants are just due to one gene that is getting shuffled around within the population generation to generation, while the concept that they are any more divergent than that is a misapprehension.
There are numerous other inconsistencies in the application of taxonomic categories within the genus Sarracenia, with the taxonomic double standard between S. flava and S. purpurea/rosea being the most glaring. It is interesting that no varieties analogous to those described in S. flava have been described in S. purpurea, even though the same color forms exist in both species. To wit: "veinless" purps are not var. maxima, all-red purps are not var. atropurpurea, and purps a la Belly of Blood are not var. rubricorpora, etc. The same features that are the object of so much debate in S. flava have been ignored taxonomically in S. purpurea, being blithely chalked up to the natural variability of the taxon! In my opinion this reflects a very unscientific bias towards the bigger, showier, "sexier" species S. flava. It's like a teacher only paying attention to an attractive student, while neglecting an equally interesting, but fugly, classmate!
This double standard also occurs in the case of S. rosea, aka "S. purpurea ssp. venosa var. burkii." Despite it's disjunct range and clear differences in trap and flower morphology, some regard this plant as a mere variety of purpurea. C'mon, not even a subspecies? This is certainly inconsistent with the prevailing treatment of S. flava (and, arguably, S. oreophila: I would argue that S. rosea is as distinct from purpurea as oreophila is from flava). It's as if purple pitcher plants aren't "taken seriously" while every shade of S. flava is taxonomically overestimated.
I am, in the course of this little armchair dissertation, taking issue with a portion of the work of Dr. Donald E. Schnell. I have enormous respect for Mr. Schnell, whom I have never met, and his pioneering work with the genus Sarracenia. I do, however, find many of his taxonomic arguments to be less than cogent because of the way they contradict each other.
Haha thank God this isn't a thread about S. rubra, because honestly folks I don't want to go there!
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Perhaps Jonathan can/will move some of those posts here.
John, Thank you for posting the new Ellison et al paper in the science category:
A MODEST PROPOSAL FOR INFRASPECIFIC RANKS sarracenia.proboards.com/thread/1042/modest-proposal-infraspecific-ranks
Some interesting statistics and positions to digest and much fertile ground for discussion. Quick reaction - Certainly if some taxonomic ranks truly do not serve a purpose they should be abandoned, but why, after all this time have they become useless. If it is because they are no longer fashionable, for whatever reason (misunderstood, ill-defined, inconsistently applied), that is not a good reason. The problem then is with the users not the system. I will be a while studying this.
Amazing place Mike. Must've been brilliant to visit and thanks for the posting the terrific photos and a matter worthy of debate, as you say!
As for the question of "atropurpurea" in these infamous sites and perhaps lone individuals within var. rubricorpora stands, I still don't buy it. My views have been well laid out but put basically I don't accept the variety name based on the following while fully recognising you have visited the plants in person:
1) S.flava var. atropurpurea is a name assigned to a distinct population of a red form of plants in the Carolinas they being isolated from the ones in your photos both genetically and geographically.
2) Those Carolina var. atropurpurea share an intimate relationship with the region's S.flava var. flava as evidenced in this and other photos of wild plants (photo Don Schnell, CPN Volume 27, Number 4, December 1998, pages 116-120 )
A. var maxima, B. var atropurpurea, C. var flava, D. var ornata, E. var cuprea
3) The "atropurpurea" in your and other's photos are seen in an intimate relationship with S.flava var. rugelii. Clearly, there exists a foundational relationship between the red plants and var rugelii.
4) This relationship between red S.flava and var. rugelii occurs in a number of locations in the Florida Panhandle as you note. In the locations you highlight, the red plants are intermingled with or alongside var. rugelii. Indeed, looking at your photos I believe the hood form of the "atropurpurea" is distinctly Florida Panhandle - particularly representative of the var. rubricorpora broad, floppy morphology. By that proposition they are quite distinct from S.flava hood form in plants native to the Atlantic coastal plain.
5) The genetics in these Panhandle S.flava populations allow for the expression of the cut throat var. rugelii; the yellow lid red tube var. rubricorpora; and the all red plants wherever they are found. My position is that all three of these are related and their expression is a broad one. The expression of var.rugelii in these locations is, in fact, no less notable than that of var. rubricorpora or the all red plants other than being deemed the so by human beings.
6) S.flava var. rubricorpora has, since Schnell's 1998 paper naming the variety and beyond, been acknowledged as being highly variable with a wide spectrum. In his book, Sarraceniaceae of North America, McPherson notes this as "The overall colouration of Sarracenia flava var. rubricorpora is extremely variable" (p 249). In his earlier volume, Pitcher Plants of the Americas, McPherson highlights this theretofore accepted variability in the following photo line up of var. rubricorpora specimens. Note the fully infused red/purple specimen on the right (Photo Stewart McPherson)
7) I ask again, what of the two plants in this photo of my own Florida red wild sourced genetic stock that has remained isolated from interbreeding since germination from the same seed pod? What is the plant on the left? And, the one on the right? It is in a sense irrelevant which location these two clones are from outside of the fact they are from the Panhandle like so many others standing in the various habitats as we speak.
9) In closing, even if the submission is correct that some of them open as all red, I reiterate my contention that the "atropurpurea" in the Panhandle are sufficiently distinct from S.flava var. atropurpurea as found 700 miles east (a massive advance on 20 miles) that they should not share the name. Whilst I am happy to have the all red "atropurpurea" accepted as part of the var. rubricorpora spectrum with out the attached "atropurpurea" moniker, I am also open to those entirely red plants as evidenced in your photos to be classified with a variety name identifying them as something distinct within what are otherwise Panhandle genetics where they both are found and spring from. I guess the question then is: what name would be appropriate....?
10) As for the transplanted theory? I simply cannot accept that until evidence fixing the event comes to light. For all the reasons I have given above and elsewhere, I think the answer to these red S.flava is to be found by un-impassioned observation of the populations and, moreover, future genetic research to determine the nature of the exhibited relationship with var. rugelii.
It's my belief that most, if not all of these flava variants are of hybrid origin, and we are witnessing them either early on (ie. you only see a red plant here and there) or later on (after that red plant has been inbred for many generations and is starting to stabilize).
As far as what any given plant is called, according to the taxonomic rules if it matches the description, it is that name. It doesn't matter where you found it. We may not like the rules but thems the rules.
What I find bizarre is the description for Sarracenia flava var. ornata. It sounds like a failed Sarracenia flava var. atropurpurea and doesn't sound like it includes the nicely veined plants at the locations specified by Schnell. This is what he says:
Deep red to purple throat pigmentation nearly obliterated by very strong and heavy red venation throughout pitcher tube and lid.
That describes this plant:
But what is:
This is what I would like to consider "ornata" but it doesn't fit the description.
Boggrower, I was preparing a more thorough reply to your last post with some posts to the science section of the forum but it is taking some time to organize and document and I don't want too much time to pass without a reply.
So here it is in short form without citations.
The red pigmentation in flava atropurpurea (both gulf and east coast forms) has been published as coming from purpurea and rosea by Phil Sheridan of Meadowview. Therefor, not a mutation but an expression of introgression and gene-flow.
Recent gene analyses for establishing phylogenetic relationships have strongly suggested alata and minor are sister species and put them into a clade together. That could explain the areoles. Perhaps a pre-divergence relict artifact. It might even explain the okefenokeensis populations.
The alata pollination study makes the point that the preferred or most common pollination strategies are 1) self- pollination (selfing), 2)pollination within the same species or variant, 3) pollination with different species, in that order. It puzzled me for some time why, in a genus that so easily hybridizes in cultivation, there were not more, obviously-hybrid plants in populations and how in these areas the entire field site did not blend into one big hybrid swarm but maintained separate species identities. Despite research data that shows outcrossing produces healthier more abundant seed and seedlings, and where there is oft repeated "common knowledge" that inbreeding(selfing) produces inferior, unfit plants, maintaining species identity by selfing may just be a more important advantage. Of course there may be other advantages to preferred selfing that we haven't figured out. The Ahh-Haa moment hasn't happened yet.
Frankly, while writing this, I've just come up with an aha of my own. Maintaining variability(separateness) in the face of quick and easy homogeneity would be a tremendous advantage.
As far as prey specificity, I wonder if it exists. Sarracenia seem to be opportunistic feeders, the kinds and numbers of captured prey varying with the prey populations themselves. They seem to attract all sorts of animals not just prey, from ants, flies, bugs, and wasps to spiders, moths, and hummingbirds. My personal experience adds blackbirds and our urban raccoons to the list with disasterous consequences (I now keep my plants entirely surrounded by netting).
While pigment is a distinct trait (it doesn't encode for other traits) it absolutely acts in communion with other distinct traits like nectar production and composition, combination with other chemical compounds, and even odor. Besides the odor of rotting flesh that these plants can certainly manifest, there have been various other odors that have been reported which may play a role in attraction (or even possibly repulsion) of certain potential prey. Except for possible chromosomal linkage, these traits are largely independent of each other genetically. When do they each become critical for the plants' well being or survival (known as a limiting factor) has not been studied in any depth, to my knowledge.
Pigmentation is often a protective mechanism, chlorophyll will burn out at high light levels or temperatures. Red pigment ebbs and flows in response to light levels that are not optimal but the chlorophyll is there, masked by the red, so photosynthesis is largely unaffected. Slow growth is a tendency in many clones/populations of Gulf coast flavas and doesn't seem to be related to degree of pigmentation. Large areas of unpigmented white however, definitely impact photosynthesis in a permanent way. They rarely become green once they have formed. There are other mechanisms the plants have that deal with low light.
Enjoying the interesting and thought-provoking comments and ideas here. Hope more get started like it. Thanks to all.
Jan 15, 2014 19:42:27 GMT -8 calen said:
Thanks waxy. Such a stimulating thread.
John on the other forum mentioned something that I had been thinking about: that form, not varietal, status should be accorded in cases where a single gene accounts for the distinguishing characteristic in a taxon. This thought pretty much opens Pandora's Box, and I therefore humbly submit the following rant:
Flava varieties should be demoted to forms. If "albino" S. purpurea ssp. purpurea is forma heterophylla, why should over-expression of the gene warrant varietal status, as in red-tubed flavas? In the case of "var." rubricorpora there are entire populations exhibiting the red tube gene, which could give the superficial impression of a more significant taxon (and, surely, lay the foundations for future divergence from other populations), but is it really so? And of course even in Florida the red-tubes usually grow with rugelli, while in the Carolinas the special color "varieties" grow in sympatry as a total mixed bag. How could divergence between flava cuprea and flava flava occur under such circumstances? Seems more likely the "different" plants are just due to one gene that is getting shuffled around within the population generation to generation, while the concept that they are any more divergent than that is a misapprehension.
There are numerous other inconsistencies in the application of taxonomic categories within the genus Sarracenia, with the taxonomic double standard between S. flava and S. purpurea/rosea being the most glaring. It is interesting that no varieties analogous to those described in S. flava have been described in S. purpurea, even though the same color forms exist in both species. To wit: "veinless" purps are not var. maxima, all-red purps are not var. atropurpurea, and purps a la Belly of Blood are not var. rubricorpora, etc. The same features that are the object of so much debate in S. flava have been ignored taxonomically in S. purpurea, being blithely chalked up to the natural variability of the taxon! In my opinion this reflects a very unscientific bias towards the bigger, showier, "sexier" species S. flava. It's like a teacher only paying attention to an attractive student, while neglecting an equally interesting, but fugly, classmate!
This double standard also occurs in the case of S. rosea, aka "S. purpurea ssp. venosa var. burkii." Despite it's disjunct range and clear differences in trap and flower morphology, some regard this plant as a mere variety of purpurea. C'mon, not even a subspecies? This is certainly inconsistent with the prevailing treatment of S. flava (and, arguably, S. oreophila: I would argue that S. rosea is as distinct from purpurea as oreophila is from flava). It's as if purple pitcher plants aren't "taken seriously" while every shade of S. flava is taxonomically overestimated.
I am, in the course of this little armchair dissertation, taking issue with a portion of the work of Dr. Donald E. Schnell. I have enormous respect for Mr. Schnell, whom I have never met, and his pioneering work with the genus Sarracenia. I do, however, find many of his taxonomic arguments to be less than cogent because of the way they contradict each other.
Haha thank God this isn't a thread about S. rubra, because honestly folks I don't want to go there!
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