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Phragmipedium Section Phragmipedium The Long-Petaled Tropical American Lady’s-Slipper Orchids/By Robert L. Dressler and Franco Pupulin

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Franco Pupulin

Robert L. Dressler

The firsT european boTanisTs to see Phragmipedium caudatum were the spaniards hipólito ruiz and José pavón during their journey to peru in the late 18th century. according to their notes and diaries, they observed populations of Phrag. caudatum around Tarma, at Cuchero and Muña; there is a specimen collected at pillao in their herbarium. While the expedi-tion was in Muña in 1786, ruiz prepared a description of the plant using the name Cypripedium grandiflorum but this name was never published. The illustrator José brunete painted the first botanical illustra-tion of Phrag. caudatum while in Muña.

unfortunately, the specimens of Phrag. caudatum brought back from peru by the spanish expedition simply gathered dust until the second half of the 20th century. When John Lindley formally published Cypripedium cau-datum, he based his description on a sin-gle, somewhat dam-aged flower sent to hooker from Lima, found in 1832 in what was then in-terpreted as an “her-

barium of ruiz and pavón.” This specimen has no information on the collector, the date of collection or geographic details, though it is probable that it was collected by Juan José Tafalla, the “attaché” of the expedition, after ruiz and pavón left peru in 1788.

When Phrag. caudatum was described in 1840, the very long petals were some-thing new to the orchid world, so much so that all Phragmipedium species with long petals were called Phrag. caudatum for the next couple of decades, an error that still occasionally pops up. however, there are three distinct species with long petals, and two other forms that are self-pollinating, though each of these is closely related to one of the “normal” species. in-terestingly enough, there is a long-petaled Paphiopedilum sanderianum in borneo. it may be that all of these long-petaled species are pollinated by the same type of insects — flies of the family syrphidae,

or hover flies — in the few cases that have been observed. at least in some cases, the long petals apparently function as perfume glands. Thus, we suspect that the small “bees” that climbed up the petals of Phrag. caudatum were actually hover flies following a perfumed path to the lip (fowlie 1972).

These species make up an unusu-ally distinctive group, but they have also been one of the most confused groups in the orchid family. The first to be named, Cypripedium (now Phragmipedium) cau-

[1] The illustration of “cypripedium grandiflo-

rum” painted by by José Brunete during

the Peruvian expedition by hipólito Ruiz

and José Pavón. This is probably the

first illustration ever of Phragmipedium

caudatum.

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datum, was based, as we have seen, on a rather battered flower with no associated data. even so, there are several specimens from peru labeled as isotypes. an isotype is a duplicate specimen of the holotype, which means a part of the same plant or of the same collection (collected at the same place and the same time as the holotype). how can there possibly be any isotypes, if we don’t know who collected the holotype, or when or even exactly where?

Thus, for some years, the orchid com-munity tended to call all long-petaled tropical american lady-slippers Phrag. (or Cyp.) caudatum. Then, too, there are two self-pollinating entities, one of which, Phragmipedium lindenii, described in 1846, looks drastically different from any-thing else, but is probably a self-pollinating mutant of Phragmipedium wallisii. The other self-pollinating mutant, Phragmipe-dium humboldtii subsp. exstaminodium, really looks quite like Phrag. humboldtii subsp. humboldtii except for the lack of a staminodium.

Two species of this group were pub-lished in 1852 in a paper by h.G. reichen-bach and J. Warszewicz. Cypripedium (now Phrag.) humboldtii, from Chiriquí, was published by Warszewicz (for a review of the complex taxonomic history of Cyp. humboldtii, see pupulin and Dressler, 2011), and Cypripedium warszewiczianum was published by reichenbach. however, reichenbach (1852) insisted that Cyp. humboldtii was exactly the same thing as Cyp. caudatum of peru. Thus, no one was willing to use the name “humboldtii” at that time, because the “king of the orchids” (reichenbach) insisted that it was really Phrag. caudatum. because of this long-standing confusion, some have preferred to call the Central american species by one of its local names, “Chinela” in panama, “barbuda” in Guatemala and “Tanal de bigotes” in Mexico.

Most of the early species were first named as Cypripedium, and some of them were (nearly inexplicably) called Selenipedium. it may be less confusing if we use only Phragmipedium in the rest of this paper.

in all lady slippers, the lip (or part of it) is somewhat folded down into itself. in this group, at least, it is useful to (men-tally) divide the rim of the lip into thirds: one lower third and two upper thirds. These elements are clearly seen in most species. in Phrag. humboldtii and Phrag. wallisii, the rims are relatively uniform within each species. in Phrag. humboldtii the rim is distinctly rounded throughout, while in Phrag. wallisii the rim has a low,

narrow keel, with the lower third project-ing forward. in Phrag. caudatum, the lower third of the rim is a bit thickened and blunt, while the inner faces of the upper thirds are flattened and quite hairy.

1. Phragmipedium caudatum (Lindl.) Rolfe

This, of course, was the first species of this complex to reach europe, and is the “type species” of Phragmipedium. The flowers are white with a network of green or brownish veins, sometimes with additional brown areas on the lip or the petals. The fold

[2] The flower at the top left of the sheet is

the holotype of Cypripedium caudatum.

Andrew mathews found it in Peru in 1832,

among the specimens of an herbarium

that he supposed belonged to Ruiz and

Pavón.

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surrounding the opening of the lip is nar-rowly compressed and a bit thickened near the apex, and the fold is wider and somewhat flattened above, with an elliptic band of hairs on each side. one of the best places to find this species in nature is near Machu picchu in peru, partly because there are so many steep, inaccessible places where one may see the plant but not quite reach it. plants have been imported from bolivia, but it is not clear whether or not this species occurs in ecuador. Leslie Garay (1978) mentions a specimen “grown in puyo.” even if the plant in question is Phrag. caudatum, it is by no means certain that it had been collected in ecuador. The ecuadorian photograph of “caudatum” that we have seen (Dodson 2003) appears to represent Phrag. lindenii subsp. wallisii.

2A. Phragmipedium humboldtii (warsz.) J.T. Atwood & Dressler subsp. humboldtii

Guido braem and his colleagues in-sist that this species was not described by reichenbach, and they are right; reichen-bach would surely agree with them. The species was described, and validly by Warscewicz (pupulin and Dressler, 2011). reichenbach at first continued to insist that it was exactly the same as the peruvian Phrag. caudatum. by now, it is difficult to say whether reichenbach really thought that the panamanian “Chinela” was the same as the peruvian Phrag. caudatum, or whether he simply didn’t want Warszewicz to name a new species. anyway, between 1873 and 1881 reichenbach at least twice proposed a new epithet, “warscewiczii,” for the Chinela. his proposals were cryptic, and were not recognized for more than 100 years (Chris-tenson 2006). however, reichenbach’s de-sire to introduce a new epithet suggests that he did not want to use the name humboldtii, or, possibly he finally realized that the pana-manian Phragmipedium was not the same

[3] Longitudinal sections through the lips of

Phragmipedium species of the Phrag.

caudatum group. A. Phrag. caudatum

(Peru, morón s.n.). B. Phrag. humboldtii

(Panama, JBL-05755). c. Phrag. lindenii

subsp. wallisii (Ecuador, JBL-10030).

Scale bar = 3 cm. All vouchers at JBL.

[4] Sections through the lips (top) and

upper third of the lip rims (bottom) in

Phragmipedium sect. Phragmipedium.

a, a1. Phrag. caudatum (Peru, morón

s.n.). b, b1. Phrag. lindenii subsp. lindenii (Ecuador JBL-10030). c, c1. Phrag. humboldtii(Panama, JBL-15781). Scale bar

= 1 cm. All vouchers at JBL.

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as the peruvian Phrag. caudatum. in 1881, discussing the new garden hybrid Phragmi-pedium × grande (which occurs naturally in panama and Costa rica), reichenbach wrote of the garden hybrid, “there can be no doubt it is descended, not from genuine caudatum, but from the so-called caudatum roseum (warscewiczii rchb.f.),” or, in fact, from Phragmipedium humboldtii (Warsz.) J.T. atwood & Dressler. because C. hum-boldtii was validly published by Warscewicz, reichenbach’s replacement name, Selenipe-dium warscewiczii, was unnecessary.

While Phrag. humboldtii was validly published, it has accumulated a list of would-be synonyms: Phragmipedium exstamino-dium Castaño, hagsater & aguirre (a self-pollinated form), Phragmipedium popovii braem, ohlund & Quené, Phragmipedium warscewiczii (rchb.f.) Christenson and Phragmipedium warscewiczianum (rchb.f.) Garay.

2B. Phragmipedium humboldtii sub-sp. exstaminodium (castaño, hágsater & Aguirre) J.T. Atwood & Dressler

This self-pollinating form is known from southern Mexico and may also occur in Guatemala, but it is now rather scarce. unlike Phrag. lindenii subsp. lindenii, this population seems to suffer from genetic poverty, and is quite rare.

3A. Phragmipedium lindenii (Lindl.) Dressler & n.h. williams subsp. lindenii

This entity caused a bit of a stir, as it was described under a different generic name, Uropedium, before the publication of Phragmipedium, and we had to transfer Uropedium lindenii to Phragmipedium and treat it as a special case, to avoid transfer-ring all the species of Phragmipedium to Uropedium. in fact, Uropedium is merely a self-pollinating mutant. We believe it to be a mutant of what has been called Phrag. wal-lisii, but apparently no one has yet crossed “Phrag. wallisii” with “Phrag. lindenii” to confirm this relationship. (When “lindenii” and “wallisii” are considered two subspecies of the same species, the epithet lindenii has priority at the specific rank.)

in an interesting article by J. steyermark and the Dunstervilles (1970), one of the au-thors (probably “stalky” Dunsterville) noted that a cross between Phrag. longifolium and Phrag. lindenii “produced exactly the same result as crossing P. longifolium with the normal P. caudatum.” remember, please, that at that time, any lady slipper with long petals was “normal Phrag. caudatum.” in any case, the two hybrids in question are illustrated by Gruss (1993b), and they are, indeed, similar, though Phragmipedium (wallisii × longifolium) has somewhat darker lips than Phragmipedium (lindenii × longi-

[5] Phragmipedium caudatum from Peru.

[6] Another Peruvian specimen of Phrag.

caudatum, cultivated in guatemala.

grower: m. Dix and m. Dix.

[7] A plant of Phrag. humboldtii subsp.

humboldtii, probably from Panama, pho-

tographed in the collection of Emma de

Soto in costa Rica.

folium). it is difficult to believe that Phrag. (wallisii × longifolium) and Phrag. (lindenii × longifolium) would be so similar, unless Phrag. lindenii is a mutant of Phrag. wal-lisii, and most of the genes are the same in both hybrids, except for the gene that, when homozygous (two doses), turns the lip into a petal and eliminates the staminodium. in the above crosses, there can be only one

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Key to Phragmipedium Sect. Phragmipedium

1. flower with 3 long petals, but no lip .............................. Phrag. lindenii subsp. lindenii1. flowers with normal, cupped or saccate lips .................................................................22. fold of lip distinctly rounded throughout, not markedly thickened ..............................33. flowers without staminodium........................ Phrag. humboldtii subsp. exstaminodium3. flowers with staminodium ....................................Phrag. humboldtii subsp. humboldtii2. fold of lip markedly compressed and solid in front, not rounded .................................44. Lower margin of lip projecting forward as a narrow keel; with hairy band above on each side, but hairs small and inconspicuous ...................Phrag. lindenii subsp. wallisii4. Lower margin of lip not markedly projecting forward; with prominent hairy band on each side, hairs larger and obvious ......................................................Phrag. caudatum— Robert L. Dressler and Franco Pupulin.

[8] A detail of the flower of Phrag. humbold-

tii subsp. humboldtii photographed at

the Lankester Botanical garden.

[9] A mexican flower of Phrag. humboldtii

subsp. humboldtii.

[10] Phragmipedium humboldtii subsp.

exstaminodium from chiapas, mexico.

dose of this troublesome gene from Phrag. lindenii. it is high time that someone crosses “Phrag. wallisii” with “Phrag. lindenii.” of course, with the monumental confusion that this group has suffered, someone may have made the cross, but used the wrong names for one or both parents. We have a Phrag. lindenii subsp. wallisii in the Lankester bo-tanical Garden, but so far no Phrag. lindenii subsp. lindenii.

Phragmipedium lindenii subsp. lindenii seems to be the most widespread entity in this group. it is known from Colombia,

Venezuela, ecuador and peru. Though it is a self-pollinating form, there are quite respect-able populations, at least in ecuador.

3B. Phragmipedium lindenii subsp. wallisii (Rchb.f.) Stein.

This form of the species certainly looks very different from Phrag. lindenii subsp. lindenii. The lip is quite well developed, and there is a low, narrow keel surrounding the opening of the lip. further, the hairy bands that are so obvious in Phrag. caudatum and Phrag. humboldtii, are present, but much less obvious. The hairs are there, but they are

tiny and easily overlooked. The lip is quite pale or somewhat pinkish. This subspecies is known from ecuador and peru, though we know of no record from Venezuela. We have seen enough cultivated plants in Colombia to suggest that it is native there.

4. Phragmipedium warszewiczia-num (Rchb.f.) Schltr.

This is surely the most enigmatic name in the group. John Lindley published an illustration, as Cypripedium caudatum, in paxton’s Flower Garden in 1850. This draw-ing was mentioned by reichenbach in his de-scription of Cypripedium warszewiczianum. braem, ohlund and Quené (2004), then, interpret this drawing as what reichenbach later published as Phrag. wallisii, and thus consider Phrag, warszewiczianum as the correct name for what has long been known as Phrag. wallisii.

When Central american Phragmipe-dium reached europe, people were reluctant to use the name Phrag. humboldtii because reichenbach had insisted that Phrag. hum-boldtii was identical to the peruvian Phrag. caudatum. it was, however, unclear what Phrag. warszewiczianum might be, so the orchid community took up this name for the panamanian Chinela, which was clearly not the same thing as Phrag. caudatum. Whatever reichenbach may have intended for Phrag. warszewiczianum, the name was widely and persistently used for a taxon that does not include its type, if, indeed, there was a type (see especially Gruss [1993a], and the confusion outlined by braem and his coauthors [2004]). Thus, according to article 57.1 of the international Code of botanical nomenclature, Phrag. warszewiczianum should not be used in a sense that conflicts with current (or rather, perhaps, recent) us-age (as equal to Phrag. humboldtii) unless and until a proposal to deal with it under art. 14.1 or 56.1 has been submitted and rejected. Thus, even if reichenbach intended the epithet warszewiczianum to be used for what is better known as Phrag. wallisii, which seems very unlikely, we should continue to use Phrag. wallisii (or, in our view, Phrag. lindenii subsp. wallisii).

Referencesatwood, J.T. and r.L. Dressler. 1998. Clarifications

and new Combinations in the Phragmipedium cau-datum Complex from Central america. Selbyana 19(2):245–248.

braem, G.J., s. ohlund and r.-J. Quené. 2004. el Ver-dadero Phragmipedium warszewiczianum, una Clari-ficación del Complejo de Phragmipedium caudatum (Phragmipedium sección Phragmipedium). Orquide-ología 23:87–116.

_. 2004. The real Phragmipedium warszewiczianum, a Clarification of the Phragmipedium caudatum Complex. Orquideología 23(2):117–136.

Christenson, e.a. 2006. The former Phragmipedium cau-datum from panama. J. Orchideenfreund 13:141–143.

Dodson, C.h. 2003. Native Ecuadorian Orchids IV. Dod-son publishing, sarasota.

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fowlie, J.a. 1972. in search of Phragmipedium caudatum. Orchid Digest 36:47–48.

Garay, L.a. 1978. orchidaceae: Cypripedioideae, orchi-doideae, neottioideae. In: G. harling and b. sparre, editors. Flora of Ecuador. Opera Botanica 9:1–305.

Gruss, o. 1993a. Die Gattung Phragmipedium rolfe (Teil 1). section Phragmipedium Garay. Die Orchidee 44(3):129–136.

_. 1993b. primärhybriden — reizvolle ergänzung der natur, Teil 10: primärhybriden des Phragmipedium caudatum und der Verwandten arten. Die Orchidee 44(6):282–288.

pupulin, f. and r.L. Dressler. 2011. a nomenclatural note on Cypripedium humboldtii (orchidaceae: Cypripedioi-deae). Harvard Pap. Bot. 16(1):167–169.

reichenbach, h.G. 1852. neue orchideen der expedi-tion des herrn J. de Warszewicz. Bot. Zeit. (Berlin) 10:689–698.

solano-Gómez, r. 2011. Phragmipedium warscewiczii: a new record for the Mexican orchid flora. Revista Mexicana de Biodiversidad 82:69–75.

steyermark, J., G.C.K. Dunsterville and e. Dunsterville. 1970. finding Phragmipedium caudatum. American Orchid Society Bulletin 39(6):484–491.

Robert L. Dressler, PhD, is a research professor at the University of Costa Rica, where he works with the Lankester Botani-cal Garden, and is currently working with the genus sobralia. Author of some 225 scientific articles and author or coauthor of five books, he is a research associate of the Oakes Ames Orchid Herbarium at Harvard University and the Marie Selby Botanical Gardens, Sarasota, Florida. He is also an associate of the Angel Andreetta Research Center of the Alfredo Pérez Guerrero [Uni-versity] (e-mail kerry@bio-photo.com).

Franco Pupulin is a senior research professor at the University of Costa Rica, where he works as director of research

[11] The flowers of Phrag. lindenii subsp.

lindenii, with modified lip.

[12] Detail of a flower of Phrag. lindenii

subsp. wallisii.

[13] An Ecuadorian specimen of Phrag.

lindenii subsp. wallisii cultivated in mo-

rosolo, italy. grower: m. comunetti.

with Lankester Botanical Garden. He is especially interested in the systematics and evolution of advanced orchid groups in subtribes Oncidiinae, Pleurothallidiinae and Zygopetaliinae. Pupulin is working on several monographic and floristic projects on Neotropical orchid floras. Author of more than 150 scientific articles and seve-ral books on the orchids of the Mesoame-rican Region, he is a research associate of the Oakes Ames Orchid Herbarium at Harvard University and the Marie Selby Botanical Gardens, Sarasota, Florida. He is also the director of the Angel Andreetta Andean Orchid Research Center University Alfredo Pérez Guerrero, Ecuador.

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