THE UNIVERSITY OF ILLINOIS LIBRARY before the Return this book on or bel Latest Date stamped University of 16123 L161 H41 THE AMERICAN SPECIES OF PASSIFLORACEAE BY ELLSWORTH P. KILLIP ASSOCIATE CURATOR, DIVISION OF PLANTS UNITED STATES NATIONAL MUSEUM THE LIBRARY OF THE APR 11 1938 UNIVERSITY OF ILLINOIS BOTANICAL SERIES FIELD MUSEUM OF NATURAL HISTORY v ^ VOLUME XIX, PART I MARCH 31, 1938 p PUBLICATION 407 PRINTED IN THE UNITED STATES OF AMERICA BY FIELD MUSEUM FKESS 520,5 F6 CONTENTS - PAGE List of illustrations 5 Introduction 7 Historical survey 8 Varying concepts of classification 8 General morphology 11 Economic uses 18 General remarks 19 List of abbreviations for herbaria cited 20 Bibliography 22 Systematic treatment 23 List of exsiccatae 582 Index . 602 1 056909 LIST OF ILLUSTRATIONS TEXT FIGURES PAGE 1. a, Passiflora vitifolia; b, P. andina; c, P. foetida var. santiagana 13 2. heterohelix a, Passiflora (natural size) ; b, P. gracillima (natural size) ; c, P. nitida 15 THE AMERICAN SPECIES OF PASSIFLORACEAE ELLSWORTH P. KiLLiP 1 INTRODUCTION Passionflowers have always had great popular appeal. Early Spanish missionaries to the New World saw in the corona, the long- stalked ovary, surmounted by three styles, and the stamens a resemblance to the Crucifixion, and from this came the popular and scientific name. Drawings and descriptions of these "marvel- lous" flowers were taken back to Europe, and as early as 1610 Jacomo Boscio presented the Flos Passionis to the world as "the most wondrous example of the Croce trionfante discovered in forest or field." (See Plant lore, legends, and lyrics, by Richard Folkard, Jr., London, 1884, portions of which are reproduced in Bailey's Standard Cyclopedia of Horticulture.) The present paper had its beginning in an attempt to present a taxonomic revision of the Mexican and Central American species of Passiflora. Before that was completed, however, I had oppor- tunities for field work in parts of South America in which the genus was abundantly represented. The study of the material obtained on these expeditions and the incidental identification of other South American specimens, as well as material from the West Indies, led to an enlargement of the scope of the original project, to include all of the American Passifloraceae. Masters' monograph of the family in Flora Brasiliensis (1872) was a thorough elaboration of the American species known at that time. Since then, about 250 New World species have been proposed, but except for Harms' necessarily brief survey in the second edition of Die naturlichen Pflanzenfamilien no attempt has been made to bring together in a single treatment these recently described species, so as to establish their relationship with each other and with earlier ones. Moreover, the intensive exploration of tropical America in the last 50 years, especially in Mexico, Central America, and the Andes, has resulted in the acquisition of a large amount of herbarium material, so that there is now a far more accurate understanding of the species than was possible in Masters' time. The present survey deals with native species, and I have given little attention to the numerous hybrids developed in horticulture, 1 Published by permission of the Secretary of the Smithsonian Institution. 7 8 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIX to many of which specific names have been assigned. Masters has given a list of the species cultivated in European gardens (Journ. Hort. Soc. n. s. 4: 125, sq. 1877), and in Bailey's Standard Cyclopedia (p. 2486) there is an extensive discussion of horticultural hybrids. HISTORICAL SURVEY Because of the striking appearance of the flowers and the unusual shape of the leaves, Passifloraceae have long been a favorite subject of study for botanists. Throughout the works of the pre-Linnaean authors, as Hernandez, Pluckenet, Plumier, and Tourneforte, are to be found descriptions and illustrations of passionflowers. In an account of Passiflora by Hallman in Linnaeus' Amoenitates (1749), 22 passionflowers are treated. In the Species Plantarum (1753) Linnaeus described 24 species, a number which was increased to 35 by Lamarck (1789). The first extensive monograph of the family was published by Cavanilles in 1790 (Diss. 10), 43 species being here treated, of which 32 were figured. In 1805 Jussieu described (Ann. Mus. Hist. Nat. 6: 102-116, 388-396) 13 new species, and discussed in detail some of the generic problems connected with this group. In Rees' Cyclopedia (1819) 55 species were included. Drawings of 25 species, unaccompanied by descriptions, appeared in Velloso's Flora Fluminensis (vol. 9, 1827). There were about 145 American species in De Candolle's Prodromus (1828), and in Roemer's synopsis of the family (1846) about 225 New World species were listed as valid. The most extensive studies of the family were those made by Masters, who, until his death in 1907, was the recognized authority upon the subject. His monograph of the American species in Flora Brasiliensis (1872), a synopsis of which appeared the preceding year, contained 202 species. An account of the Colombian species by Triana and Planchon (Ann. Sci. Nat. V. Bot. 17: 121-186. 1873) discusses the collections of Lehmann, Andre", Stiibel, and Weberbauer ; and others by Masters and by Harms, a survey of the Ecuadorean tacsonias by Sodiro (1903), and numerous papers by Harms and myself, all have made known many additional species. In the present monograph 355 species of Passiflora and 10 belonging to smaller genera are recognized, of which 17 are now proposed for the first time. In a few species there are one or two varieties or forms. Passiflora foetida has 37 varieties, 13 of them now newly described. VARYING CONCEPTS OF CLASSIFICATION Passifloraceae consists of 12 genera, four of which are found in the New World. Of the strictly Old World ones, Crossostemma, AMERICAN PASSIFLORACEAE 9 Schlechterina, and Machadoa are monotypic African genera; Deidamia, with about 8 species, and Tryphostemma, with about 30, likewise are wholly African. Adenia, the largest of the Old World genera, contains approximately 80 species, most of which are restricted to Africa, though a few occur in tropical parts of Asia and on the South Pacific islands. Hollrungia and Tetrapathaea, both monotypic genera, are found in New Guinea and New Zealand, respectively. In Passi- flora, by far the most important genus of the family, less than 40 of the nearly 400 known species are endemic to the Old World; these, with the possible exception of one species in Madagascar, are known only from Asia and the South Pacific islands, and the genus is, there- fore, primarily one of the New World. Mitostemma, with three species, and Dilkea, with five, are restricted to South America. Tetrastylis contains one Central American and one Brazilian species. The families Malsherbiaceae and Caricaceae were united to Passifloraceae by early authors, but they are now considered distinct. In the following brief review of the genera and groups into which the family has at various times been divided, those genera that properly are referable to Malsherbiaceae or to Caricaceae are not included, nor are the Old World groups of Passifloraceae considered. All the passionflowers were placed by Linnaeus in Passiflora. In 1787 Medicus created the genus Cieca for the apetalous species, and revived Tourneforte's names Murucuja and Granadilla for two additional genera. In 1789 (Gen. PI. 398) Jussieu added another genus of Tourneforte's, Tacsonia, but he did not recognize Cieca. These three genera, Passiflora, Murucuja, and Tacsonia, were main- tained by Persoon (Syn. PI. 2: 219. 1807). In 1819 (Ann. G6n. Soc. Phys. Brux.) Bory St. Vincent proposed three additional segregates of Passiflora, viz., Asephananthes, Monac- tineirma, and Anthactinia. In 1822 (Me*m. Soc. Phys. Geneve 1: 434-443) De Candolle, accepting the generic concepts of Jussieu and of Persoon, subdivided Passiflora into seven sections: Astrophea, Polyanthea, Tetrapathaea, Cieca, Decaloba, Granadilla, and Dysosmia. This treatment was followed in the Prodromus, an eighth section, Tacsonioides, being added to Passiflora, and Murucuja and Tacsonia, with two and four sections, respectively, being given generic rank. In 1846 Roemer, evidently with an extremely narrow concept of a genus, raised to generic rank all of De Candolle's sections. But in placing the species Roemer unfortunately showed a lack of knowl- edge of their characters, many being assigned to certain "genera" without apparent justification. 10 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIX Bentham and Hooker adopted De Candolle's treatment, though reducing Murucuja to a section of Passiflora. In a paper in the Transactions of the Linnaean Society, issued the year before his monograph in the Flora Brasiliensis appeared, Masters recognized three genera, viz., Passiflora, Tacsonia, and Dilkea, the last being proposed at that time on the basis of recent collections from northern South America. He divided Passiflora into four subgenera: Astrophea, Plectostemma (including the sections Cieca, Dysosmia, and Decaloba), Murucuja (including the sections Eumurucuja and Psilanthus), and Granadilla. In Tacsonia he recognized two sections, Bracteogama and Eutacsonia. This early paper, in so far as it applied to New World species, was merely a synopsis of the forthcoming monograph, and many new species were mentioned by name only; in order not to burden the synonymy in the present monograph I have not cited this first use of such names. The new combinations here made date, of course, from this earlier paper. In their monograph of the Colombian Passifloraceae (1873) Triana and Planchon maintained but one genus, Passiflora, which they divided into five subgenera: Tacsonia,Granadilla, Plectostemma, Muru- cuja, and Astrophea. Within the next few years two new genera, Tetrastylis and Mito- stemma, were described from Brazil. In the first edition of Die naturlichen Pflanzenfamilien Harms recognized four New World genera: Dilkea, Mitostemma, Tetrastylis, and Passiflora.