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THE AMERICAN SPECIES of THIBAUDIEAE by Albert C. Smith

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THE AMERICAN SPECIES of THIBAUDIEAE by Albert C. Smith THE AMERICAN SPECIES OF THIBAUDIEAE By Albert C. Smith INTRODUCTION The attention of the writer was first attracted to the group of plants known as Thibaudieae in 1926 and 1927, at which time he had opportunity to observe them in their natural habitat in the Eastern Cordillera of Colombia. A second trip, to Peru in 1929, furthered this interest and resulted in the addition of specimens of the group to American herbaria. These trips, both of which were under the auspices of the Smithsonian Institution and led by E. P. Killip, added so many specimens of Thibaudieae to the United States National Herbarium that to name them became highly de- sirable. In addition to these recent collections, a considerable quan- tity of other unstudied material had accumulated, owing to the efforts of several American collectors, notably Paul C. Standley, in Central America, and F. W. Pennell, H. H. Busby, E. P. Killip, and J. F. Macbride, in South America. In attempting to name these specimens it was found that the only comprehensive work on the tribe, that of R, Hoerold in 1909, was quite inadequate, having been based on the specimens in one her- barium only. In preparing the present paper, a task which has occupied much of the writer's time for two years, type material of nearly all previously described species has been seen, a privilege for which he is deeply indebted to Prof. L, Diels, director of the Botanisches Museum, Berlin, and Sir Arthur Hill, director of the Royal Botanic Gardens, Kew. At these two European institutions most of the type specimens had been deposited and were lent to the writer to facilitate his task. In addition it has been a privilege to study the specimens belonging to the Gray Herbarium of Harvard University; the Goeldi Museum of Para, Brazil; the Field Museum of Natural History; and the New York Botanical Garden. The institutions from which specimens have been studied are thus indi- cated in citations: Berlin (B), Field Museum (F), Gray Her- barium (G), Goeldi Museum (Go), Kew (K), United States Na- tional Museum (N), New York Botanical Garden (Y). To the 311 312 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM directors and curators of the above-mentioned institutions the writer wishes to acknowledge his appreciation of their cooperation. In addition the writer wishes to thank Dr. William R. Maxon and E. P. Killip for their assistance in the preparation of manu- script, Dr. J, H. Barnhart for his suggestions in certain nomen- clatorial questions, and especially Dr. H. A. Gleason for his con- stant advice and encouragement. HISTORICAL CONSIDERATION The first mention of any plant which may be placed in the tribe Thibaudieae aa it is at present understood was in 1763, when Adan- son1 published a brief description of the genus Chwpalon* His mention of the plant, in connection with Oxycoccus and Vaccinium, is so noncommittal that it can not with certainty be applied to any more recent collection. Probably a species of €mendishia was under consideration; but since no specific description is available, the name may be ignored. Among authentic genera, the first to be formally described was Ceratostema, mentioned at some length by Jussieu2 in 1789. Although no species is detailed, the specimen referred to was ob- viously Ceratostema periwianwri, subsequently published by Gmelin,8 During the following years no specimens of Thibaudieae were de- scribed, probably because collections from tropical America had not yet reached Europe in any quantity. However, the collections of Ruiz and Pavon in Peru discovered several species referable to this tribe. Unfortunately these specimens were described in the fourth volume of Flora Peruviana et Chilensis, which consists of plates only and is very rare. It is a question whether this volume was available at such an early date as 1802, but it seems best to consider this the publication date for those plates that have suitable analyses. The descriptions made use of the generic names Ceratostema and Thibattdia; the plants figured belong to the genera, as at present con- stituted, Ceratostema, Siphonandra, Psamviisia, Thibaudia, and Cavendishia. One of the plates (384, top) represents a plant best referred to the tribe Euvaccinieae and not considered here. Concerning first publication of the generic name Thibavdia, the above-mentioned plates, although adequate as specific descriptions, are not considered as establishing the genus.4 The first formal de- 1 Fam. PI. 2:164. 1763. 1 Gen. pi. 163. 1789. •Byst. Nat. 2:676. 1791. 4 On the subject of the value of plates in generic name determination, see Sprague, " OrenoaniOe vs. Villebrunea" in Kew Bull. Misc. Inf. no. 10, 1928. SMITH—AMERICAN SPECIES OP TH IB AUDIBLE 313 scription of the genus Thibaudia is in St. Hilaire.8 This descrip- tion, from the wording, probably antedates the issue of volume 4 of Ruiz and Pavon. Three species are described, obviously from the manuscript of Ruiz and Pavon, but in two cases the specific names have been slightly altered from those that appear on the plates. For the sake of uniformity, the names on the plates are in this paper considered authentic. The species first mentioned by St. Hilaire, which may be considered the type species of the genus, is T. melli- \flora (or T, mellifera according to St, Hilaire). In the subsequent 30 years a few more species were described under the generic name Tkibaiidia, mostly by Humboldt, Bonpland, and Kunth® in 1818. These species have since been referred to various genera of the tribe; the name Thibaudia as applied by authors of this period included all plants referable to the tribe. In 1836 the generic name C(uuendishia was introduced by Lindley,7 but was not understood as being the common Andean genus by sub- sequent writers, who continued to neglect it, making necessary a large number of comparatively recent correct binominals. The following year Hooker described the genus Macle<mia8 and several species. At this time species of the tribe were introduced into England as horticultural plants by commercial collectors in South America. These were described by several authors in the following years and frequently were illustrated by elaborate plates. The genus AntJiopterus was described by Hooker9 in 1839. The most elaborate and comprehensive treatment of the tribe up to this date is by Dunal.10 He describes all species known to him and adds several new descriptions, applying the names Geratostemai, Thibaudia, CavendisMa (in regard to one species only), Macleama, and Anthoplerus to American members of the tribe. Like his predecessors, Dunal used the name Thibaiudia in its broadest sense. His descriptions are brief and often inadequate, without comment as to relationships. Among the authors who described tropical American plants at this period, and among them various Thibaudieae, was Bentham, whose new species, founded on the Hartweg collection, included the genus Oreanthes.11 — - I ■ ' .—.II. ■■ . ■ ■ I . ■ I ■■■■ ■■■ * ■■■ I ' 1 1 'Expos. Fam. Nat. 362. 1805. *Nov. Gen. & Sp. 3:268-275. 3818. TBot. Beg. 21: sub pi. 1191. 1836. * Hook. Icon. PL 2: pi. 109. 1837. "Hook. Icon. PL 3: pi- 2^S. 1889. "DC. Prodr. 7:552-578. 1839. u PL Hartw. 140. 1844. * 314 CONTRIBUTIONS PROM THE NATIONAL HERBARIUM In 1851 Klotzsch12 published his " Studien iiber die naturliche Klasse Bicomes Linne," a "work whose chief value is in the strict delimitation of genera. The elaborate keys to genera published by Klotzsch are by no means natural, but his conceptions, as borne out by the species which he grouped together, were fundamentally modern. Genera of American Thibaudieae added by him include Tyria, JSatyria, Orthaea, Siphonandra, Semirarriisia, Eurygania, Polyboea, Proclesia, Tkemistoclesia, and Pscmmisia. To be sure, not all these genera &re maintained at the present time, but on the whole they represent very definite conceptions. In 1856 Planchon and Lindenia published a description using the generic name Oonocatyw, which has been neglected or placed in Oeratostemdi by subsequent authors. It is considered a distinct genus in the present treatment. In 1857 Grisebach14 published a short list of names without descriptions. These names, based on Lechler's collections from Peru, are without notation other than the collector's number; they are here considered nomdna nuda. In 1863 the Brazilian and eastern Peruvian species were considered by Meissner." Here is described the genus Biedelia, which is synony- mous with Saiyria, Another important contribution to the study of the tribe was published in 1876 by Bentham and Hooker.16 Here, of course, only the genera are discussed, but the relationships between them are more clearly expressed than in previous works. The only generic name added is Findlaya, which is synonymous with Orthaea. Pre- viously in the same year Hooker17 had published the genus Notopora. Between this time and 1909 many species were described by vari- ous authors, none of whom considered generic relationships in a way to throw light on their phylogeny. The best treatment is that of Drude,18 whose key is adequate for general use. Niedenzu19 studied in some detail the anatomy of the leaf in Ericaceae, the results being published in the form of a key that does not aid the taxonomist to a great extent. Another author whose name is found in this period is Hemsley,20 who published a summary of the Central American species, forming several new combinations in Cavendishia.. The only genus added about this time was Ru&bya Britton.21 " Linnaea 24:1-88. 1851 " Gard. Chron. 1856:152. 1856. "Lechl. Berb. Amer. Austr. 58. 1857. " Mart. Fl. Bras. *7:125-127, 171-174. 1883. "Gen. PI. 2:564-577. 1876. "Hook. Icon. PL 12:53. pi. 1159. 1876. 18 Eugl. & Prantl, Pflanzenfam. 41; 53-57.
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