Orchid Digest Volume 60, No. 4 Oct.-Nov.-Dec., 1996 A NEW SPECIES OF TOLUMNIA RAF. FROM PUERTO RICO RUBEN P. SAULEDA AND MARVIN E. RAGAN Ruben Sauleda Ph.D. received his doctorate in biology at the University of South Florida (Tampa) with a specialty in plant taxonomy. He has been proprietor of Ruben in Orchids in Miami since 1962 and is a well-known and accomplished writer on orchids. Marvin Ragan has been an avid orchidist for thirty-one years. He is owner of MAJ Orchids in Orange Park, Florida and has been recognized as an Orchid Identification Authority by the American Orchid Society. Historically the genus Tolumnia Raf., or the miniature oncidiums as they are called in horticultural circles, have been the center of controversy and heated debates. In the sixties the debates centered around the delineation of the species, especially the white-flowered species, the classification of which was handled differently from the colored species. Later in the seventies and eighties the debate shifted to the generic level. The argument centered on the choice of the type species of the genus Oncidium Sw. One interpretation used Oncidium variegatum Sw. as the type of the genus. This interpretation would mandate that the miniature Oncidium group would be the "true" oncidiums and all of the other species would have to be transferred out of the genus. If O. variegatum was rejected as the type of the genus Oncidium, then it could be used to typify the genus Tolumnia. Proposals were made to the Botanical Congress, all with good merit, and finally the ruling was made that the genus Oncidium was not to be based on Oncidium variegatum. This opened the door for the transfer of all the species of miniature oncidiums into the genus Tolumnia. It seems that now we are returning back to the same controversy we had to endure thirty years ago. The problem in the sixties revolved around a proliferation of orchid literature generated by researchers completely ignorant of basic taxonomic and evolutionary principles. At the present time there is a proliferation of literature by researchers who although scientifically trained, either do not have a basic understanding of the definition of a species and the evolutionary principles involved in speciation, or choose to ignore these principles. Many of their interpretations are based on conjecture, personal feelings or manipulation of statistical data.

Several researchers have called the species we are naming in this paper Tolumnia variegata (Sw.) Braem. Ackerman first stated that possibly two distinct species comprised the Puerto Rico population. Now (1992) he states that the Puerto Rican species is Tolumnia variegata, but it has two phenology types, which are slightly different morphologically, but the characteristics of the two intergrades. Other researchers choose to call all of the white species T. variegata. This would be like calling all of the rock laelias one species, all the purple cattleyas one species or all the white spiranthes one species. Little regard is being given to the basic definition of a species: a population of similar individuals, which interbreed, sharing a common gene pool, but are reproductively isolated from other populations. The white tolumnias very clearly follow these rules. The reproductive isolation between most of the species is geographical. The majority of the species are endemic to their respective islands. In our estimation it is possible that at one time more than one species occurred in Puerto Rico, accounting for the high degree of variation in the population. This may be a situation similar to Tolumnia lucayana (Nash) Braem in the islands of the Bahamas. At some time in the past, two or possibly three species occurred in the Bahamas, and through natural hybridization and back-crossing a hybrid swarm has resulted. None of the original species are present. A selfing of any of the color forms which have resulted in the hybrid swarm will yield the complete range of variations. An exhaustive statistical analysis was made of the Puerto Rico population and of T. variegata and Tolumnia leiboldii (Reich. f) Braem. Measurements were taken from both herbarium specimens, live-collected material and live material, which had been cultivated for several years to eliminate differences caused by ecological factors. Computer programs, which generated scatter diagrams, percent similarity and various other statistical procedures were used. The computer-generated data from the Puerto Rico population can be interpreted in three ways: the population in Puerto Rico shares a common gene pool, can be morphologically distinguished from the other white species of tolumnias and of paramount importance is that it is geographically isolated from any other Tolumnia, meaning that it represents either a hybrid swarm or a highly variable species. After careful morphological study of both herbarium specimens and living material, making field observations and collections, and studying the statistical analysis, we have concluded that sufficient data exist to confirm that the population is distinct from the other white-flowered tolumnias and therefore must be described as a new species.

Tolumnia borinquinensis Sauleda and Ragan, sp. nov. Type: PUERTO RICO: near SW edge of Lago Tortuguero, 2.3 km NE of Manati, 17 Nov 1981, Sauleda, Sauleda, Ragan & Hansen 6835 (Holotype: NY; Isotypes: F, FTG, US, USF). Crista et isthmus labelli et fundi petalorum colore rubiginosa maculati. Crista tuberculis anticis tribus et tuberculis posticis tribus parvioribus. Plant epiphytic, rarely epilithic, rhizomatous, to 56 cm tall; root numerous, slender, velamentous; rhizome stoloniferous, to 9 cm long, repent, decumbent, wiry, remotely several sheathed, sheaths scarious, ovate-triangular, acute, to 1.2 cm long, 0.6 cm wide; stem flattened, to 0.5 cm long, 0.2 cm wide, completely enclosed by distichous leaves, to leaves; leaves coriaceous, imbricated, produced at intervals along stem conduplicate, falcate, lanceolate, acute, to 14 cm long, 1.6 cm wide, margin cartilaginous, serrulate; inflorescence lateral, to 56 cm tall, peduncle slender, erect, distantly several sheathed, racemose, rarely particulate above, to 25 flowers; floral bracts narrowly lanceolate, concave, acute to acuminate, to 0.6 cm long, 0.2 cm wide; ovary pedicellate, slender, to 1.6 cm long; sepals white, basal l/3 with reddish brown spots; dorsal sepal spatulate, basally clawed, concave, acute, occasionally apiculate, to 0.8 cm long, 0.5 cm wide; lateral sepals lanceolate, concave, acute, connate nearly to apex, to 0.8 cm long, 0.1 cm wide; petals white, basal 1/3 with reddish brown spots, spatulate, basally clawed, concave, obtuse, to 0.9 cm long, 0.4 cm wide; labellum three- lobed, white, occasionally with a pink tint, to 1.4 cm long 1.6 cm wide; lateral lobes basal, linear-elliptic to orbicular, obtuse to broadly rounded, margin entire to irregularly crenulate at apex; midlobe separated from lateral lobes by a short isthmus isthmus yellow with reddish brown spots, margin usually fringed; midlobe reniform or broadly cordate, emarginate, usually with an apicule in the sinus; callosity on isthmus yellow with reddish brown spots, with three anterior and three smaller posterior tubercles; column stout, erect, to 0.4 cm long, 0.2 cm wide, with expanded lateral wings projecting from apex, lateral wings white or pink, ovate, base obtuse, apex acute, margin irregularly lobed, to 0.2 cm long, 0.1 cm wide, anther cap white or pink; capsule pendent, ellipsoid, to 1.8 cm long, 0.8 cm wide. Etymology: Derived from the Taino Indian name for the island of Puerto Rico, Borinquin, meaning land of the haughty lord. Additional Specimens Examined: PUERTO RICO: Rio Piedras, 13 Jun 1914, Stevenson 1978 (US); west end of Laguna Tortuguero, 27 Mar 1922, Britton & Chardon 6825 (NY); 0.8 km SW of Manati, along rt.149, Sauleda et al 6840(USF); Vega Baja, 26 Mar 1906, Britton & Cowell 1432 (NY); 6.4 km SW of Dorado, 24 Nov 1981, Sauleda et al 6900 (FTG, NY, USF); Utuado, 15-20 Mar 1906, Britton & Cowell 901 (NY); 8 km N of Utuado, along rt. 10, 24 Sep 1982, Sauleda et al 7711(USF); 17.9

km S of Cieales, along rt. 149, 28 Nov 1981, Sauleda et al 6941 (USF); Maricao to Monte Alegrillo, 3 Apr 1913, Britton et al 2582 (NY); Reserva Forestal Maricao, Monte del Estado, along rt. 120, 1.9 km S of jct with rt. 366, 9 Nov 1983, Sauleda et al 8540A (USF); 1.9 km SW of Comerio, along rt. 156, 25 Nov 1981, Sauleda et al 6914 (FTG); Barrio Sonadora de Bayane, 8 Oct 1982, Sauleda et al 7944 (USF); Reserva Forestal Carite, along rt. 184, 3.2 km E of jct with rt. 152, 4 Nov 1983, Sauleda et al 8474A (USF); Reserva Forestal Guanica, 4.0 km SE of Guanica, along rt. 383, 6 Nov 1983, Sauleda et al 8521 (USF); Mount Mandios, near Jayuya, 17 Mar 1906, Britton & Cowell 903 (NY); Lares, 22 Nov 1913, Stevens & Hess 4937(NY); Playa Sucia Bay, 1 Mar 1915, Britton et al 4747 (NY, US); Abonito, 4 May 1935, Sargent 357 (US); Palo Seco, 2 Dec 1919, Stevenson 3312 (US); Aguada, near Rio Grande, 4 Mar 1886, Sintenis 5721 (US); between Morovis and Corozal, 19 Mar 1925, Britton et al 8430 (NY); in ravine, Coamo Springs, 14 June-22 Jul 1901, Underwood & Griggs s.n. (NY); Vieques Island, Calabaza to Ensenada Honda, 18 Feb 1914, Shafer 2951 (NY, US). U.S. VIRGIN ISLANDS: St. Thomas, Flag hill, Oct 1887, Eggers s.n. (US). Tolumnia borinquinensis previously was identified as Oncidium variegatum Sw. (Britton & Wilson, 1924; Cogniaux, 1910; Moir & Moir, 1970), Oncidium leiboldii Reichb. f. (Britton & Wilson, 1924; Cogniaux, 1910; Leon, 1946; Moir, 1962; Moir & Hawkes, 1967; Moir & Moir, 1970, 1973) and Oncidium sylvestre Lindl. (Cogniaux, 1910). Recent collections of living material of T. leiboldii, a Cuban endemic, and of T. variegata, an Hispaniolan endemic, have given us an opportunity to compare living specimens of these with living specimens of the Puerto Rican species. In addition, we have compared the types of T. variegata, T. leiboldii and T. sylvestre with the Puerto Rican material. Vegetatively T. variegata and T. borinquinensis are similar. Both species have broad coriaceous falcate conduplicate leaves. Tolumnia leiboldii has narrow almost terete leaves and T. sylvestre is a small plant with manyleaved elongate growths; these characteristics quickly distinguish them from the other two species and therefore they can be eliminated from any further discussion. Tolumnia variegata and T. borinquinensis occupy a wide variety of habitats. They can be found from xeric habitats at sea level to moist forests at higher elevations, which probably accounts in part for their vegetative similarities. Florally, the two species are distinct and can be easily distinguished. The isthmus of the labellum (area between the midlobe and side lobes) is usually fimbriated in T. borinquinensis, but is entire in T. variegata. The crest sculpturing on both species is somewhat similar, both have three anterior tubercles and three smaller

posterior tubercles. However, the thickness and exact position of the tubercles and the degree of development is different but consistent for each species and can be used to distinguish the species. LITERATURE CITED Ackerman, James D. (Text) and Maruja Del Castillo Mayda (Illustrations). 1992. The Orchids of Puerto Rico and the Virgin Islands. University of Puerto Rico Press. San Juan, P. R. Britton, N. L. and P. Wilson. 1924. Scientific Survey of Puerto Rico and the Virgin Islands. Orchidaceae 5: 180-217. New York Academy of Sciences. Cogruaux, C. A. 1910. Orchidaceae, In: I. Urban. Symbolae Antillanae 6: 293-696. Reprinted 1964. A. Asher & Co. Amsterdam. Garay, L. A. and H. R. Sweet. 1974. The Orchidaceae, In R. A. Howard, Flora of the Lesser Antilles. Arnold Arb., Harv. Univ., Jamaica Plain, Mass. Moir, W. W. G. and A. D. Hawkes. 1967. Studies in the Equitant Oncidiums. Phytologia 15 (1): 2-12. Moir, W. W. G. and M. A. Moir. 1970. Variegata Oncidiums. Published by the authors. Moir, W. W. G. and M. A. Moir.1973. Variegata Oncidiums, Part 1. The Florida Orchidist 16 (3): 124-131. Acknowledgments We wish to thank the directors, curators and staff members of the herbaria cited throughout the text for their cooperation and help. We acknowledge the generous field assistance given by the late Eugene Antommarchi, Myriam and Sidney Antommarchi and Julio Melendez of Puerto Rico, Patricia H. Adams of Boca Raton, Florida, Diane K. Sauleda of Miami, Florida and Bruce Hansen of the University of South Florida. Our thanks also are due to Dr. Helen B. Correll for preparing the latin description. Ruben P. Sauleda 12555 S. W. 46 Street Miami, FL 33175 Marvin E. Ragan 126 Wild Orchid Lane

Orange Park, FL 32065 32065 Orchid Digest, Oct.-Nov.-Dec. 1996