Contents Page Introduction 25 Historical Sketch 27 Drift Fruit 34

Contents Page Introduction 25 Historical sketch 27 Drift fruit 34 Fossil species 37 Structure of the fruit 38 Relationships and evolution 41 Family Humiri&cae 44 Tribe Vantaneoideae 49 Genus Vantanea 49 Tribe Humirioideae 76 Genus Duckesia 76 Genus Endopleura 80 Genus Hylocarpa 84 Genus Humiria 87 Genus Humiriaslrum 122 Genus Schistoslemon 146 Genus Sacoglottis 161 Collections cited 187 Bibliography 206 Index 210 ill A TAXONOMIC REVISION OF THE HUMIRIACEAE Jose Cuatkkcasas Introduction My special interest in the tropical trees and shrubs of the family Humiriaceae developed many years ago while I was studying my own collections, gathered on expeditions sponsored by the regional Government of El Valle del Cauca, from the Pacific coast of Colombia. What drew my attention most were the rare fruit collected and their similarity to the fossil specimens of Sacoglottis cipaconensis presented to me some years earlier in Bogota by the geologist J. Royo G6mez. These fossils proved to belong to the genus Vantanea rather than to Sacoglottis. Notwithstanding the exceptionally good work of Urban in the "Flora Brasiliensis," the existing literature lacked information on the structure of the fruit, information indispensable to a more complete taxonomic understanding of the family. In view of the collections made since Martins' gigantic work on neo- tropical botany, some revision of the group seemed necessary. In 1951, while in Chicago, I initiated this revision with the cooperation of T. Just, who intended to prepare a section on paleobotany in the planned synopsis; however, the project was discontinued. In 1957, with the primary purpose of writing the Humiriaceae for the "Flora of Colombia," I started anew with a taxonomic revision of the entire family; the results of this study are summarized in the present publication. For this revision I have used the large collections of the U.S. National Museum, in the U.S. National Herbarium (US), which have recently been augmented by the collections formerly in the National Arboretum. I have also used those of the: New York Botanical Garden, New York (NY), Gray Herbarium, Harvard University, Cambridge, Massachusetts (GH), Arnold Arboretum, Harvard University, Cambridge, Massachusetts (A), School of Forestry, Yale University, New Haven, Connecticut (Y). Museum d'Histoire Naturelle, Paris (P), Royal Botanic Gardens, Kew, Surrey (K), British Museum (Natural History), London (BM), Naturhistoriska Museum, Stockholm (S), Botanische Staatssammlung, Munich (M), Botanical Museum and Herbarium, Utrecht (U), Museu Goeldi de Historia Natural, Belgm do Par6 (MG), Institute Agronomico do Norte, Beldm do Par& (IAN), and Institute de Ciencias Naturales, Bogota (COL). 25 26 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM The above-mentioned herbaria have the largest existing collections of members of Humiriaceae, and their examination has resulted in a considerably greater understanding of the family than we had pre- viously. These herbaria combined have a copious representation of the important historical collections made by Schomburgk, Spruce, Martius, and others, as well as of more recent collections, among them those of Ducke, Maguire, and the Bosch wezen (Forest Service) of Surinam. Nearly all types or isotypes of the described species are found among the material of the herbaria mentioned. The amount of fruiting material in the modern collections, though not abundant, is sufficient to allow us to draw new lines in the concept of the family and its genera. Especially rewarding have been the amazing Brazilian collections of Adolfo Ducke, The relative importance of the collections used can be readily seen by examining the index of collectors at the end of this paper (p. 187). The present revision is based on the classical method of comparative morphology. I utilize the fruit structure to a considerably greater extent than previous authors. The paper concludes that the Humiriaceae are to be maintained as an independent family with 8 genera, 49 species, and many subspecies, varieties, and forms. The introduction to this paper begins with a historical sketch of contributions to our knowledge of the family and a discussion of the phenomena of drift fruit and fossils. There follows, because of its taxonomic importance, a discussion of the structure of the fruit and also a discussion of taxonomic relationships and evolutionary trends. No data on the cytological structure of any Humiriaceae are available. The considerations of evolutionary trends are founded on comparative gross morphology and are merely tentative. The paleontologic data are limited to the mention of published species and to a few taxonomic suggestions. The main part of the paper is devoted to classification and taxonomic descriptions of the family and the tribes, genera, species, subspecies, varieties, and forms comprising the family. Included are plates and figures comparing and contrasting related forms. The plates are grouped following p. 84. The figures appear in the text near the species to which they refer. The citations of collections constitute an important part in a monographic work and, because of the relative scarcity of locality data, I consider it useful to publish the information given on the herbarium labels by each collector. For greater accuracy these data appear in their original language. In the citations of herbaria where specimens are deposited, I use the abbreviations of Lanjouw's "Index herbari- orum." In citing the photographic series of the Chicago Natural CUATRECASAS—HUMIRIACEAE 27 History Museum, the abbreviation "Photo F.M." has been used preceding the number. I wish to extend my thanks to the directors and curators of the herbaria who have aided by lending me collections or giving me facilities for my work, among them especially Albert C. Smith, Director of the Museum of Natural History of the U.S. National Museum; Jason R. Swallen, head curator of Botany; Lyman B. Smith, curator of phanerogams, who has checked the manuscript; Roland Brown, honorary curator of paleobotany, who has been of great help locating humiriaceous fossil collections; the artist Russell Zimmermann, who made all but four of the figures with a complete understanding of my interpretations; and Paula Gerard and Jane Roller, who made the accurate drawings of figure 1. The work for this revision has been carried out at the Department of Botany of the U.S. National Museum, with the help of a grant from the National Science Foundation. Historical Sketch 1775: Aublet publishes in his "Histoire des plantes de la Guiane franQaise" the first descriptions and drawings of Humiriaceae— Houmiri balsamijera and Vantanea guianensis—two new genera and species named after the Caribbean names "Houmiri" and "Iouantan," The first genus is characterized by small flowers with 20 stamens and the second by large flowers and many stamens. He places the genera in the class Polyandria and the order Monogynia of Linne's sexual system. 1777: Scopoli, like other botanists of the time, finding the names given by Aublet barbarous, proposes "Wernisekia" for "Houmiri" (Introd. Hist. Nat.). 1789: Schreber, for the same reason, in his eighth edition of "Linne's genera plantarum" changes "Houmiri" to "Myrodendrum" and "Vantanea" to "Lemniscia" and writes: "Aubletiano vero nomen- clature quum nimis sit barbara, aliam quin substituerem, me continere nequivi." They are included in the Polyandria Monogynia. 1789: Jussieu, trying to Latinize Aublet's name, gives it the spelling "Houmiria," in his "Genera plantarum." 1792: Lamark publishes a new species form French Guiana— Vantanea parvijlora, 1800: Willdenow, following Schreber, in his edition of Linnaeus' "Species plantarum" adopts "Lemnescia," thus correcting its spelling, and, for Vantanea guianensis, he uses "Lemnescia floribunda." 1805: St. Hilaire completes the "Latinization" initiated by Jussieu with the spelling "Humiria." He includes the plant in "Classe 28 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM Polyandrie" and "Ordre Monogynie" (Exposition des families naturelles). 1824: Necs and Martius publish the new genus Helleria, based upon H. obovata, not noticing that it is congeneric with Vantanea guianensis. The authors place the genus next to Humiria and consider its close relationship to several groups, primarily Aurantiae and Ternstroemiaceae. 1827: Martius publishes (Not. Gen. Sp. PI. Br.) detailed descriptions of Humirium crassifolium and H. floribundumt and gives a new spelling to the generic name, "Humirium" (proposed in ms. by Richard). Martius describes the genus Helleria and a new genus, SacogloitiSj based upon Sacoglottis amazonica, and distinguished by 10 stamens. He includes Sacoglottis in Linnaeus' Monadelphia- Decandria, whereas he places Humirium and Helleria in Monadelphia- Polyandria. Martius comments on the close relationship between the three genera and traces their differences with the genera considered related to Mcliaceae, Symplocaceac, and Styracaceae. 1829: Jussieu gives, in St. Hilairo's "Flora Brasiliae meridionalis," for the first time a taxonomic category (order—family) to these three genera under the name "Humiriaceae." He adds the new species Humirium montanum, H. parvifolium, and Helleria ovalifolia and lists HeUeria obovata Nees. & Mart. 18$0: Lindley (Introd. Nat. Syst. Bot.), considers the tribe Humiriaceae and their relation to the Aurantiaceae, Diosmeae, and Kutaceae. 1840: Endlicher in "Genera plan tar um" lists this family as "Ordo CCXXII-Humiriaceae," following Jussieu and recalling Martius'

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