Phrag caudatum group Taxonomy

OK, I want to start a debate regarding the group of Phragmipediums with long Petals. As some of you may know myself, Dr. Guido Braem and Sandy Ohlund wrote some articles regarding these species.

In a nutshell, Guido discovered that what we have been calling Phrag. wallisii all these years was an incorrect name. The reason is that Reichenbach who gave the name Phrag. wallisii in 1873 to that species, had named the same species twice. Twenty years earlier he had also described and named the following species (shown in the following picture, depicted in Paxton's Flower Garden (Lindley, 1850):



The name he gave to this species is Phrag. warscewiczianum. This was a species native to South America. This is clearly what we nowadays have been calling Phrag. wallisii.

see picture:



This means that if you follow the International Code, as the name Phrag. warscewiczianum was used first, this is the official name for this species, and Phrag wallisii becomes invalid.

Now later, in the trade another species showed up, that was native to Central America (Panama and Costa Rica). This is the following species:



Guido found out that this species was never officially described, but for some reason in the past it got the name Phrag. warscewiczianum associated with it. As this name was already taken by a different species we decided to officially describe it and give it the name Phrag. popowii.

Now my question for you guys is the following: It is clear there are 5 different (that we know of ) taxa within this group. We decided to give all 5 the species rank. So we came up with:

1. Phrag. caudatum
2. Phrag. lindenii
3. Phrag. warscewiczianum
4. Phrag. popowii
5. Phrag. exstaminodium

It was clear to us that Phrag. lindenii and Phrag. warscewiczianum are more closely related than to the other species, and the same is true to the two Central American species, Phrag. popowii and Phrag exstaminodium, but we decided to keep them as separate species, as we felt that they were isolated populations that grew in different areas and thus far we know no gene flow was occurring from one population to the next. I talked to a botanist (I forgot what his name is at the moment) who worked at the University in Ecuador, and he has seen Phrag. lindenii in the wild. According to him Phrag. lindenii grows in very different locations than Phrag. warscewiczianum and according to him they were 2 distinct species. Also it made it cleaner.

Cribb and Dressler on the other hand want to divide this group into 3 species. According to them lindenii is a "mutant" form of warscewiczianum, as it is bascally the peloric form of this species (It has 3 petals instead of a pouch). Aslo exstaminodium is the "mutant" form of popowii. It lacks the staminodal shield and just like Phrag. lindenii it self pollinates. However as lindenii was described before warscewiczianum and exstaminodium was described before popowii, the "mutant" forms would become the species names, and you would get the following classification (They probably would give them different variety names, but I gave them the same variety name to make it more clear):

1. Phrag. caudatum
2a. Phrag. lindenii var. lindenii
2b Phrag. lindenii var. warscewiczianum
3a Phrag. exstaminodium var. exstaminodium
3b Phrag. exstaminodium var. popowii

The third thing you can do is to call it all the same species (Phrag. caudatum) and have 5 different varieties:

1a Phrag. caudatum var. caudatum
1b Phrag. caudatum var. lindenii
1c Phrag. caudatum var. warscewiczianum
1d Phrag. caudatum var. exstaminodium
1e Phrag. caudatum var. popowii


Now my question to you guys is which classification to you prefer and why? Scenario one with 5 species; scenario two with 3 species and 2 varieties, or scenario three with one species and 5 varieties?

Robert

If it looks different, the name should different.

Your way is best. Making 5 distinct species.

Here is my opinion why...
Because they are distinct not only in appearance but also in locations. Keeping the "species" well separated with their own "simple" names eliminates confusion. Taxonomists who basically deal with only preserved specimens have no problem grouping under one specie name. But when you get into the horticultural growing side it makes more sense to have distinct names so as not to confuse the species with varieties.

I think varietal names should be reserved for individual variations that exist repeatedly within a specie population but not as separate populations on their own.

Great topic!

McCook in her Phrag thesis (1989) considered Reichenbach's description of Phrag. warscewiczianum to be a later synonym of Phrag. caudatum rather than an earlier description of the taxon now usually called Phrag. wallisii. I haven't yet examined the descriptions, so I can't say which I agree with.

McCook also noted that the Central American taxon had no valid species name of its own, but didn't describe it because she considered the Central and South American 'caudatums' to be conspecific.


I tend to believe that the name warszewiczianum should not be used in place of what is now commonly called wallisii, even if it was describing what is now commonly called Phrag. wallisii. Dressler (2005 OD) pointed out that Article 57 prohibits the use of warscewiczianum in this sense:

57.1. A name that has been widely and persistently used for a taxon or taxa not including its type is not to be used in a sense that conflicts with current usage unless and until a proposal to deal with it under Art. 14.1 or 56.1 has been submitted and rejected.

Therefore, if the 5 taxa are to be treated as species, they should be:
Phrag. caudatum
Phrag. wallisii
Phrag. lindenii
Phrag. exstaminodium
Phrag. popowii


The cloudiest issue seems to be the name of the central american very-dark taxon. So far, it changes names at least once and probably twice depending on which level it is assigned. Dressler in the 2005 OD article officially described Phrag. caudatum subsp. warszewiczii. Since article 11 of the ICBN states:

11.2. In no case does a name have priority outside the rank in which it is published (but see Art. 53.4).

both validly published names are legitimate. There may be a third legitimate name, Phrag. caudatum var. roseum, that applies to this taxon. The earliest reference I have seen about this name is the 1867 Revue Horticole. I haven't gotten a chance to read the article, so I can't say either way if it should be considered valid. Assuming that it is, this taxon now has three separate, legitimately published names:
Phrag. popowii
Phrag. caudatum subsp. warszewiczii
Phrag. caudatum var. roseum


The five taxa fall into two morphologically distinct groups: (i) caudatum, popowii, exstaminodium, and (ii) wallisii, lindenii.

I believe that (ii) wallisii and lindenii are sufficiently distinct from one another to be at the specific level.

However, I don't think there are sufficient morphological differences among group i to consider the three as distinct species. Besides the color, McCook (1989) couldn't find a consistent difference between the herbarium specimens of 'caudatum' from South America and the 'caudatum' from Central America. Phrag. exstaminodium differs from the Central American 'caudatum' by only a single morphological difference, the missing staminode, which allows the pollenia to contact the stigma during bud development and antithesis. Because of this, I consider these three taxa to be varieties of one species.

I prefer to adress the 5 taxa as:
Phrag. wallisii
Phrag. lindenii
Phrag. caudatum
Phrag. caudatum var. roseum (tentative)
Phrag. caudatum var. exstaminodium


The major issue I have with the Braem, Ohlund, & Quene article is the name chosen for the Central American species. A far, far more appropriate move would have been to validly describe the species as Phrag. humboldtii, the name Warszewicz originally intended for the taxon. Instead, the taxon was named after a friend of Braem's who is a renowned orchid smuggler.

--Stephen

Stephen very good points!

My question is why do you consider lindenii to be sufficiently distinct from wallisii. To me the morphological differences between the two are probably due to a difference in "one" or " two" genes between the two taxa. If you consider exstaminodium and popowii to be the same species; you should do the same for lindenii and wallisii.

By the way, just out of curriosity I crossed lindenii with wallisii, to see what you would get. My theory is that the primary cross will look exactly like Phrag. wallisii. If I would sib 2 of the seedlings, I am guessing that 1/4 will look like Phrag. lindenii again.....If this is true does this mean they are the same species?

I hate to agree with you on the other point regarding wallisii versus warscewiczianum, but even I prefer to keep using the name wallisii, because if I would use "warscewiczianum" for that taxon I think it creates confusion. Especially if you have a plant that has written Phrag. warscewiczianum on the label......you would start to think is this the old name or the new name.....so that is why I keep calling the South American species Phrag. wallisii, but I do call the Central American species Phrag. popowii.....now I had no part in the naming of the orchid, and have to agree with you that Phrag. humboldtii would have been more appropriate....but it has now been described as Phrag. popowii, so that is what I am naming it.

Robert

I could definitely be convinced to consider all five as species!

But for now, I consider wallisii and lindenii to be distinct enough to be considered separate species because they have more than a single consistent morphological difference. The functional third anther pressing the pollenia against the stigma was probably the first trait to evolve which separated lindenii from wallisii. Somewhere along the line, the blooms then lost their pouches. If you crossed wallisii and lindenii, I would guess that you'll get a plant that looks exactly like wallisii. If you then do an F1 sib cross, I would guess that a quarter of the offspring would be pouchless. However, I don't have a good guess as to whether or not there would be a third anther in any of the F2s, and if so, whether or not it would be formed correctly. That specific an adaptation strikes me as being too complex to be controlled by a single gene.

I don't consider exstaminodium and popowii to be separate species because they have only a single consistent morphological difference, the missing staminode. I would guess this trait is the result of a single defective recessive gene (like an albino is). Also, Dressler (2005) reported that there are intermediate forms between popowii and exstaminodium in Guatemala that have partially formed staminodes. It would be interesting to see the results of a cross between exstaminodium and popowii.

This situation is one where both arguments are convincing. I may very well go back to calling the 5 taxa separate species.

--Stephen

I have to agree with you that you can go both ways and that is why I posted this thread...my preference is still to call all 5 taxa separate species.

By the way the fact that Phrag lindenii has a 3rd pollinia in my view is linked to it having 3 petals. I have seen Paphiopedilums that sometimes have a petal instead of a pouch or have multiple pouches instead of petals, and when this is the case they sometimes have an extra pollenia. I think because it had the extra pollenia it was able to survive and become a separate entity (because it was able to selfpollinate and thus pass on it's "mutant" gene to the next generation).

We will know for sure if it is one or multiple genes if we get seggregation in the F2 generation. Say we get plants without a pouch, but with only 2 pollenia, or plants with 3 pollinia and with a pouch, but if in the F2 generation (after siibbing two F1 plants) we get 25% of the plants that look like lindenii with 3 pollenia and without a pouch, and 75% that look like wallisii that have a pouch with only 2 pollenia; it is probably only due to one gene......I can't waite to see the results.....It is good I am a patient person.

Robert

I tend to agree with Stephen that the loss of the pouch is a direct result of the evolution(well, I'd guess that it was probably more of a freak "accident" than prolonged evolution) of the third anther. Since it could self pollinate, mutations upon the labellum wouldn't make a large difference in overall fitness of the individuals. I think that is a lot more likely than a third anther and modified labellum arising at the same time as a result of the same mutation. After all, slipper parts come in threes. Three petals, three sepals, three styles(fused into one), and three stamens (1 modified to become the staminode). The addition of the 3rd anther means that lindenii would have, essentially, 4 stamens. I just don't think that mutation and the pouch loss would occur at the same time due to the same genetic mutation.

I have only one word.

Headache.

If they're in a pot and on my bench, they are called wallisii(the light one), caudatum, warscewiczianum(the dark one), lindenii, and extaminodium.

I realize taxonomists need work, and I agree that Braem is correct in his research...but I don't really care to be honest. I know that is pretty useless when it comes to having a constructive discussion, but I'm not gonna budge on this one(and I will NEVER call Laelia purpurata, Sophronitis purpurata!!). I think there needs to be a statute of limitations on some of these names. If it walks like a duck, quacks like a duck, looks like a duck, and has been called a duck for 150 years...I'm not gonna call it an owl because that was what was originally written down and lost for all that time and rename owls as hooters...though I would consider that name oke:

Jon
________
Kawasaki Zn1100B

Jon in SW Ohio said:

If they're in a pot and on my bench, they are called wallisii(the light one), caudatum, warscewiczianum(the dark one), lindenii, and extaminodium.

I realize taxonomists need work, and I agree that Braem is correct in his research...but I don't really care to be honest. I know that is pretty useless when it comes to having a constructive discussion, but I'm not gonna budge on this one(and I will NEVER call Laelia purpurata, Sophronitis purpurata!!). I think there needs to be a statute of limitations on some of these names. If it walks like a duck, quacks like a duck, looks like a duck, and has been called a duck for 150 years...I'm not gonna call it an owl because that was what was originally written down and lost for all that time and rename owls as hooters...though I would consider that name oke:

Jon

Click to expand...

I agree. These names have been used for such a long time that they should continue............switching names and concepts is too confusing and will lead to more mislabeling, misidentification and incorrectly named crosses.

In this case, the question of names isn't really an issue for the orchid grower. The 5 taxa are well known and the names don't overlap. The proper name and identity of each taxon is of interest to the taxonomists.

However, there are many cases where names are misused and confused by orchid growers to the point where you can never be sure of what plant you're buying/growing. Many orchid sellers often use the wrong name for the plants they're offering or worse still, end up selling hybrids as species.

For instance, I've seen plants of praestans, wilhelminiae, glanduliferum, gardineri, and bodegomii for sale. In reality, there are only two distinct species (and their primary hybrid) in cultivation at the moment. I bought a 'Paph. gardineri' that turned out to be a typical wilhelminiae, but it could just have easily been a praestans. I had no way of knowing until it bloomed

--Stephen

silence882 said:

For instance, I've seen plants of praestans, wilhelminiae, glanduliferum, gardineri, and bodegomii for sale. In reality, there are only two distinct species (and their primary hybrid) in cultivation at the moment. I bought a 'Paph. gardineri' that turned out to be a typical wilhelminiae, but it could just have easily been a praestans. I had no way of knowing until it bloomed

--Stephen

Click to expand...

I was recently informed by Jo Levey (and backed up by Garay) that there are no real wilhelminea in the US (they are all gardineri)??? This came about after Jo saw that I had wilhelminea on my collection list.

Even though my plant blooms on 6" leaf span growths, and the flower is a dead ringer for the insitu pic of wilhelminae in Cribbs book. Garay said it was definitely gardineri because it had twisted petals and a square staminode.

As far as the phrag debate goes:

I think its fair to give them all species status. Not only is there range and habitat separation, but separation in reproductive strategies.

Mutations are (at least one process) that make species. And I'm not aware of any rules that define how much mutation is neccesary before species status is granted from the parent species.

In this case the mutations we are seeing in the long petaled phrags seem to be producing plants with different reproductive strategies from the parent species. If this gives the new "species" a significant degree of reproductive isolation from the parent then I think that species status is justified.

Of couse the notion of "significance" will need to be debated.

The only description of gardineri is a poor line drawing from The Cruise of the Marchesa, by Guillemard (1886) (it was reproduced in the Jul/Sep 1995 OD). It's woefully inadequate to assign a taxon to the name. Cyp. gardineri should be considered a nomen nudum.

--Stephen

Rick said:

I was recently informed by Jo Levey (and backed up by Garay) that there are no real wilhelminea in the US (they are all gardineri)??? This came about after Jo saw that I had wilhelminea on my collection list.

Even though my plant blooms on 6" leaf span growths, and the flower is a dead ringer for the insitu pic of wilhelminae in Cribbs book. Garay said it was definitely gardineri because it had twisted petals and a square staminode.

Click to expand...

Eric Christenson wrote a short note in a german journal earlier this year where he stated that Selenipedium warszewiczianum was the oldest name for the Panamese taxon and made the combination to Phragmipedium. So, now it is Phrag. warszewiczianum again.
Of course, this was almost half a year ago and it really is about time to find a new name again

It may be hard to see in pressed herbarium specimens, but all the caudatum plants I have seen live and on photos I find quite distinct from the mid-american taxon, as much as from the pale wallisii. So there are caudatum, wallisii and warszewiczianum. I agree that one single different trait (missing staminode shield or lip) shouldn't make a different species, but these traits change the repoductive behaviour of the varieties lindenii and exstaminodium and together with the different geografical distribution and apparently habitat these seem to be good arguments (just to me) to treat lindenii and exstaminodium as distinct species.

Cheers, Carsten

Carsten,

Do you know when the Panamese species was described as Selenipedium warscewiczii?

If this was done after 1852 (that is the date when Rechenbach described Cyp. warscewiczianum for the South American "lighter" colored species that later became synonymous with Phrag. wallisii), we cannot use the name "warscewiczii" for the Central American species.

Anyway my point is when taxonomist's decide to combine species from different genera into a new genus, certain rules have to be followed, so if one species from Central America was described as Selenipedium warscewiczii and another species from South America as Cyperpedium warscewiczianum; when you put them together into a new genus, only one can be named Phragmipedium warscewiczianum (as the name Phrag. warscewiczii will look too similar), I am guessing that Cyp. warscewiczianum was described first which means the name "warscewiczianum" has to be applied to the South American species (later known as wallisii).

There is also a rule (that I believe) in the code that if a species has had a name for so many years even though it is not the first and valid name it can keep the name...so in this case as we have been calling the South American species Phrag. wallisii for all these years it can keep the name Phrag. wallisii. But as the name "Phrag. warscewiczianum" was already used and is now a synomym for Phrag. wallisii, I think we should abandon the name Phrag. warscewiczii for the Central American species, and thus the name Phrag. popowii would become valid again.....

I know.....those Taxonomists (I am one of them) sure make it complicated....but we are just trying to go by the rules......and I have to agree with Jon, that to me Laelia purpurescens will never be Sophronitis purpurescens....

Robert

I just emailed Dr. Guido Braem, and told him we were debating about the long petaled Phragmipedium species on this forum, and wanted his input as well, as he knows more about all the Plant Taxonomy Rules than I do, and he responded that he will contribute as well in the near future.

Robert

warscewiczianum vs. warscewiczii

yes, and it took quite some effort to type the title alright!

I only have access to a subset of the literature, my very limited knowledge of the group is based on the few articles I have:
That's Olafs article in OD, Braem et al. and Braem and Ohlund in Australian Orchid Digest (should be the english translations og the two articles in Richardiana), the article of Dressler in OD and one of Eric Christenson in the Journal für den Orchideenfreund. Also, thanks to Stephen, I have Reichenbachs Xenia Orchidea, and Hookers 1844 text.

Reading these bits left me with a few questions, some of them are probably hard to answer. Mostly nobody bothers, I'm afraid.

Taxonomy:

I think most people agree that there are 5 taxa in the caudatum group. Wether these are 5 distinct species, or three species and two subspecies is up to everyone’s personal opinion, i.e. species concept applied. Actually, we haven’t talked about the numerous varieties of caudatum yet. var. giganteum, sanderae etc. Olaf mentioned a few of those in his OD article. I don’t know wether these varieties deserve the rank variety, or if they are mere colour forms of caudatum.

Nomenclature:

Lindley described C. caudatum based on a pressed flower from the Ruiz herbarium. Hooker depicted an inflorescence 1844, coll. by Lobb in the interior of Peru, as caudatum.

Reichenbach in 1852? received C. humbodtii from v. Warsc.
Question: did he receive a life plant or pressed material? Where had v. W. been collecting this particular plant? Has it been preserved? At the Reichenbach Herbarium? I'm asking because v. Warsc. has travelled both in mid-and South America and don't know wether this particular travel led him through Peru/Ecuador only or if he reached the habitats of what's now popowii.
As for the name: I understand that Reichenbach fil published the name C. humboldtii Wzw by reducing it to synonymy with C. caudatum. Therefore that name is not available in Cypripedium anymore. The name is also not available in Phragmipedium anymore, since Attwood and Dressler published Phrag. humboldtii nom. invalid. Correct?
Rchb. treats C. humboldtii v.W. as a synonym of C. caudatum. After that he describes C. warscewiczianum based on the picture in Paxton’s flower garden(?). As far as I can tell from the picture published in the Australian Orchid Digest Article, the flower clearly resembles what was long known as wallisii.

In the Xenia O. from 1858, Rchb. fil. transferred all known species of Cypripedium from South America to Selenipedium. It reads:
1) S. caudatum. Cypripedium caudatum Lindl. Cypripedium humboldti Wzw.
2) S. warscewiczianum. Cypripedium warscewiczianum Rchb. fil.

In the Xenia O. from 1874 (1873?) Rchb. fil. describes S. wallisii:

Affine Selenipedio caudato Rchb. fil. et Warscewiczii Rchb. fil. (caudato roseo Hort.).

He continues further down: “I find it hard to determine a character despite the obviously large difference(s) to the abovementioned two species, especially after through experience I have become very careful using the staminode shape. Finally I noticed that this species lacks the hairs at the rim of the tepals which characterise the other two species.
At this occasion I may mention that I have been lucky finally to find an excellent difference between S. caudatum and S. warscewiczii. The latter has numerous small pits on the front part of the lip, which are completely absent in the former. All efforts to determine a distinguishing character from the staminode failed, with increasing numbers of plants I’ve seen.” My own humble translation.

Questions: was the transition from S. warscewiczianum to S. warscewiczii likely to be a spelling error by Rchb., or did he intend to describe a new species with this name?
Did Rchb still have access to the type of S. warsc. (the painting in Paxton’s) in 1873? Have plants fitting the type of S. warsc. been imported to Europe between 1854 and 1873? Have plants of the meso-american species been imported during that time and been available to Rchb.?
Christenson mentions a reference for C. caudatum var. roseum in Rev. Hort. 1867: 133. Anybody can provide a copy? Indications of the origin of plants and the colour, maybe a painting in there? Is this really a synonym of popowii?

I get the impression that Rchb. hasn’t seen any plant material of his S. warscewiczianum until 1873, when he described S. wallisii. Apparently, in the meantime he had received material from central America (from whom?) which for some reason he considered to be his species warscewiszii(-ianum).
Both caudatum and the mid-american species possess hairs at the rim of the petals. My popowii Fu-Manchu’s lip looks like it has been a bad case of acne when it was younger: lots of pits on a scarred lip front. On the other hand, Rchb. uses the lack of hairs on the petal rims on his wallisii as the distinguishing character.

The original description of C. warscewiczianum by Rchb was based on the staminode shape, which Rchb later on didn’t trust as a character anymore. Did he not believe in his concept of warscewiczianum anymore and silently tried to slip in caudatum roseum as warscewiczii?

Reichenbachs description of S. wallisii fits the type of S. warscewiczianum pretty well, so it leaves wallisii as a synonym of that species.
The informal description (better, comparison with related taxa) of S. warscewiczii within the text in Xenia Orch. 1873 fits what we now know as popowii, the mid-american species.

So, Christenson is right and we do have an old name (S. warscewiczii Rchb. fil.) for popowii, only if this name is valid. Validity may be questioned because of the similarity to the older name S. warscewiczianum and the doubtful intention of Rchb fil to actually newly describe a species/publish a new name S. warscewiczii. Any comments?

best wishes, Carsten

My head is spinning...

I forgot:

Christenson hasn't cited a type for Paph. warscewiczii. Did he have to, and is there one at all?
From Christenson: "The name Selenipedium warscewiczii is what we today call an avowed substitute name based on the earlier Cypripedium caudatum var. roseum Hort."

Cheers, Carsten (insane)