Selbyana 19(2): 245-248

CLARIFICATIONS AND NEW COMBINATIONS IN THE PHRAGMIPEDIUM CAUDATUM COMPLEX FROM CENTRAL AMERICA

JOHN T. ATWOODl AND ROBERT L. DRESSLER2 IMarie Selby Botanical Gardens, 811 South Palm Ave., Sarasota, Florida 34236 2Missouri Botanical Garden, Box 299, Saint Louis, Missouri 63166-0299; Florida Museum of Natural History, University of Florida, Gainesville, Florida 32611

ABSTRACT. Members of the Phragmipedium caudatum complex (Phragmipedium section Phragmipedium) are clarified. The name Cypripedium humboldtii Warsz. ex Rchb.f. is lectotypified based on a syntype from Chiriqui, Panama and formally combined as Phragmipedium humboldtii (Warsz. ex Rchb.f.) J.T. Atwood & Dressler. The combination Phragmipedium humboldtii subsp. exstaminodium (Castaiio, Hagsater & Aguirre) J.T. Atwood and Dressler is also made.

One would think that enough has been said South American plants. McCook (1990) accepts about the attractive and widely published Phrag­ this as the same as South American P. caudatum mipedium caudatum complex or Phragmipedium on the basis of isozyme analyses as well as of Sect. Phragmipedium. Historically nearly the en­ morphological similarity, but consistent differ­ tire section was accepted as Phragmipedium ences in the flowers as well as the leaves suggest caudatum with the exception of P. lindenii to us that recognition as separate species is use­ (Lindl.) Dressler and N.Wms., an obligately au­ ful. Attempts have been made to apply the name, togamous entity ranging from Ecuador to Ven­ P. warszewiczianum (Rchb.f.) Garay3 to the ezuela with pouched lip replaced by a normal Central American entity (Garay 1979, Atwood third petal. In addition it has a third stamen with 1984), but McCook (1990) pointed out that the distinct filament. With the appearance of new type of this name is based on a South American collections into cultivation within the last 30 rather than a Central American plant. Therefore, years it has become clear that there are at least Phragmipedium warszewiczianum (Rchb.f.) three kinds of plants with normal lips corre­ Garay cannot be the same as the Central Amer­ sponding to different geographical areas. ican species, and the non-flared lip suggests that Entity one, now accepted as Phragmipedium this name must be a synonym of P. caudatum caudatum, is an enormous plant with leaves Lindl. rather than of P. wallisii. sometimes attaining 70 cm in length which ex­ Interestingly enough, there is a description ceed the inflorescence. The flower is usually preceding the original description of Cypripedi­ brownish with darker reticulation, and the orifice um warszewiczianum Rchb.f. that has been ig­ is pubescent but little flared. It hails from Peru nored by all: Cypripedium humboldti Warsz. ex and Bolivia, but doubtfully from further north. Rchb.f. A translation of the German and Latin The plant purchased in Europe and illustrated in reveals why no one has wanted to deal with this Dunsterville and Garay as P. caudatum (Vene­ name. zuelan Orchids Illustrated, v. 2, p. 265, 1961) "Cypripedium Humboldti Wszwcz. [ex has thin leaves, pendent ovaries and swellings Rchb.f.] I have a specimen and a drawing by the on each side of the orifice consistent with the author. I compare the same with specimens of garden hybrid, P. Grande registered in 1881 C. caudatum of Ruiz, [and] with the illustration casting doubt on the occurrence of P. caudatum in Hook. Ie. VII. 628 [actually this should read in Venezuela. '658']. Paxton Fl. G. 9. Also I believe in more Entity two from Ecuador (probably also Co­ species as does Mr. Von W[arszewicz]. The lombia and perhaps Venezuela) is a much small­ plant of our traveler is but exactly the same, er plant with a white lip peppered with lavender which is submitted in Hook, Ie.; the same, which and much flared at the orifice. This corresponds Ruiz collected. After a bad flower of the latter, with P. wallisii (Rchb.f.) Garay. Phragmipedium Professor Lindley erected his C. caudatum: I lindenii is probably a selfing form of this. Entity three hails from Central America. Its 3 The specific epithet is usually spelled "warscew­ leaves are the shortest and broadest in the com­ iczianum." However, the spelling in the original de­ plex; shorter than the inflorescence. The flowers scription as "warszewiczianum" corresponds with the are yellowish green with the lip darkly stained name of the collector, Joseph Ritter von Rawicz War­ around the orifice, a feature we have not seen in szewicz. 245 246 SELBYANA [Volume 19(2) know the same plant in good specimens, also mentioned in his description of Cypripedium exactly as the author himself kept the name for humboldti. the plant.-From this the plant in Fl. Gard. T. 9 The name most probably commemorates Al­ is distinguished principally by the transversely exander von Humboldt, though Reichenbach two lobed starninode. The illustration is very fails to provide the etymology. It is fitting that distinct, and Mr. Professor L. says explicitly: we finally affix a name to this grand plant from staminode transversely bilobed, setose at the Central America, but to do so requires a lecto­ apex. The illustration in Hook. Ie., on the other typification based on Warszewicz 41 and recom­ hand, evidently shows the same organ three bination with the genus Phragmipedium. Fol­ lobed, as I always find it. Although I most rarely lowing is a formal taxonomic treatment of distinguish plants after foreign illustrations and Phragmipedium humboldtii. descriptions, nevertheless it stands as self evi­ Phragmipediwn hwnboldtii (Warsz. ex Rchb.f.) dent that I trust an illustration and description J.T. Atwood and Dressler, comb. nov. Basion­ by Mr. Professor Lindley. I place the diagnoses ym: Cypripedium humboldti Warsz. ex of two [species] thus." Rchb.f., Bot. Zeitung 10:691. 1852. Lectotype Reichenbach starts a new paragraph quoting selected here: Panama, Chiriqui, Warszewicz Lindley's description of Cypripedium caudatum 41 (W 15682). Not Phragmipedium warszew­ then provides a short German and Latin diag­ iczianum (Rchb.f.) Garay. nosis for his C. humboldti citing two specimens, one from Peru and the other from Panama, fol­ Epiphytic or on steep slopes, roots 1.5-3 mm lowed by a description of C. wars;zewiczianum. in diameter, tan, velvety; stems simple. Leaves "Cypr. caudatum Lindl. Orch. 5. 31.; upper se­ several, distichous, conduplicate, 19-32 X 2.8- pal [phyllo supremo] broad, oblong, acute; syn­ 4.4 cm ligulate, obtuse or retuse. Racemes ter­ sepal [inferiori subaequali] broader; petals minal with 1-4 simultaneous flowers, peduncles [phyllis intemis] from a broader base, linear, ex­ 15-30 cm long; floral bracts 4-6.2 X 3.2-3.4 tremely long; lip [calceo] oblong, ventricose, cm ovate or obovate, obtuse; flowers yellowish with abbreviate margin, orifice velvety pilose, green with brownish reticulation; the lip heavily stained with dark red, especially near the orifice, starninode three lobate.-Hook. Ie. 1. c. C. Hum­ boldti v. Wszwcz. 'Flowers dark yellow, red the orifice with yellowish hairs; pedicel and ova­ ry velvety, 10-15 cm long; dorsal sepal 10-15 nerved, lip purple spotted.' In woods of the Pe­ X 2-2.8 cm, lanceolate, synsepal 8-12 X 3.4- ruvian Andes. Ruiz! (Herb. Berol.!) On oaks of 5.5 cm, ovate, obtuse; petals long-ciliate at the mountains, Chiriqui. Von Warszewicz!" base, 30-40 cm X ca. 18 mm, linear, twisted; Contributing to the confusion Reichenbach lip 5.5-6.2 X 3-3.7 cm, deeply saccate, hispid appears to have confused the identity of Cypri­ within, glabrous without; starninode broad, pedium caudatum, but the important point here somewhat 3-lobate or appearing 2-lobate if the is that he cites two different specimens (syn­ apex is strongly curved; capsule narrowly fusi­ types) as his C. humboldti: (1) a Peruvian spec­ form, ca. 10 X 0.8 cm. imen collected by Ruiz and the other (2) a spec­ An autogamous entity, Phragmipedium exst­ imen from Chiriqui collected by Warszewicz. aminodium Castano, Hagsater and Aguirre, was This latter specimen also seems to be the same described From Mexico on the basis of the lack that Reichenbach alludes to in the opening sen­ of starninode (Castano, Hagsater & Aguirre tence. A specimen in the Reichenbach Herbari­ 1984). However, some specimens from Guate­ um labeled C. humboldti appears to be a dupli­ mala also either lack the starninode or have ir­ cate corresponding with the Peruvian specimen regular asymmetrical starninodes showing that of Ruiz presumably destroyed at Berlin. The the two entities are not really distinct on the ba­ other is the Central American specimen of War­ sis of this feature. Furthermore, Phragmipedium szewicz along with a drawing of it on a second exstaminodium has the same short leaves and sheet. There is little doubt that the drawing and dark flower color as the allogamous entities from the specimen apply to the same plant as both Central America. We believe that this is best -contain the number "41." The specimen shows viewed as an autogamous geographic variant of the short broad leaves exceeded by the inflores­ P. humboldtii despite the recent publication of cence, features characteristic of Central Ameri­ Cox et al. (1997) with cladogram showing P. can plants, and the drawing shows the dark stain exstaminodium well separated from P. warszew­ on the rim of the lip that is less pronounced in iczianum (probably P. humboldtii). We further South American material. Unfortunately Rei­ note in their work that P. caricinum, a very dif­ chenbach failed to mention specimen number 41 ferent species with -much smaller chromosome in his text, but the specimen and drawing oth­ number, is nested within the P. caudatum group erwise correspond with the original materials despite its resemblance to P. pearcei. We sug- 1998] . ATWOOD AND DRESSLER: PHRAGMIPEDIUM HUMBOWTII 247

FIGURE 1. Engraving of Phragmipedium humboldtii published in Gardener's Chronicle «new series) 3:211. 1875) as P. cauda tum. The plant exhibits the short broad leaves of the Central American plant which are much shorter than the inflorescence. gest that these taxa may need to be resamp1ed exstaminodium agrees well with P. humboldtii owing to general confusion of identification and! but lacks a staminode, a feature whose impor­ or possible mislabeling of vials. A formal re­ tance in lady-slipper classification has some­ combination of P. exstaminodium as an autoga­ times been exaggerated. The anthers of P. exst­ mous subspecies of P. humboldtii follows. aminodium are supported on longer filaments Phragmipedium humboldtii subsp. exstami- bringing them into contact with the stigmas. The Southern range of this subspecies is un­ nodium (Castano, Hagsater & Aguirre) J.T. clear. All flowers of specimens examined from Atwood and Dressler, comb. et stat. nov. Chiriqui, Panama (N = 9 at SEL), the type "lo­ Basionym: Phragmipedium exstaminodium cality" of P. humboldtii subsp. humboldtii, have Castano, Hagsater and Aguirre, Orquidea normal staminodes. (Mex.) 9: 193. TYPE: MEXICO, Chiapas, The Phragmipedium caudatum complex in­ Tziscao, 1,700 m, Leleu s.n. (Holotype: cludes four recognized species: P. wallisii from AMO). Ecuador and Colombia, P. lindenii from Ecua­ The drawing of the type of Phragmipedium dor, P. humboldtii from Central America, and P. 248 SELBYANA [Volume 19(2) caudatum from Peru and Bolivia. Following is ACKNOWLEDGMENTS a key to species in the Phragmipedium cauda­ tum complex. We thank Hans Wiehler for checking our translation of the German, and for the reviewers' 1. Lip forming a third elongate petal, not saccate ... thoughtful comments...... P. lindenii 2. Lip saccate ...... 3 LITERATURE CITED 3. Lip white with purplish flecks, orifice flared ...... P. wallisii ATWOOD, J.T. 1984. The Relationships of the Slipper 3. Lip brown or yellow, orifice not flared .. 4 Orchids. Selbyana 7: 129-247. 4. Leaves longer than the inflorescence (Peru Cox, AV., AM. PRIDGEON, V.A ALBERT, AND M.W. and Bolivia) ...... P. caudatum CHASE. 1997. Phy10genetics of the slipper orchids 4. Leaves shorter than the inflorescence (Cypripedioideae, Orchidaceae): nuclear rDNA (Chiapas to Panama) ...... 5 ITS sequences. Pl. Syst. Evol. 208: 197-223. 5. Staminode present ...... GARAY, L.A 1979. The Genus Phragmipedium. Or­ . .. P. humboldtii subsp. humboldtii chid Digest 43: 133-148. 5. Staminode absent ...... MCCOOK, L. 1990. The genus Phragmipedium II. P. humboldtii subsp. exstaminodium Amer. Orchid Soc. Bull. 59: 153-158.