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Flora Neotropica 123 Ghillean T. Prance Humiriaceae Flora Neotropica Volume 123 Since 1968, these monographic volumes have provided taxonomic treatments of families or other groups of plants or fungi growing in the Americas between the Tropic of Cancer and the Tropic of Capricorn. The monographs are intended to be comprehensive, so most of them also include information on economic botany, conservation, phylogenetic relationships, taxonomic history, ecology, cytology, anatomy, and phytochemistry, among other topics. Each volume is illustrated with line drawings, black and white photographs, and distribution maps. This series is the official publication of the Organization for Flora Neotropica (OFN), an organization founded in 1964 with the mission of producing a published botanical inventory of the American tropics. The current OFN Executive Director is Wm. Wayt Thomas. Flora Neotropica volumes 1–14 (1968–1974) were published by Hafner Publishing Company. Volumes 15–121 (1975 – 2019) were published by The New York Botanical Garden. All in-print volumes 1—121 are available exclusively through NYBG Shop: https://www.nybgshop.org/nybg-press/books-in-series/flora- neotropica. Volumes 122 and going forward are available exclusively from Springer. More information about this series at http://www.springer.com/series/16365 Ghillean T. Prance Humiriaceae Ghillean T. Prance Royal Botanic Gardens, Kew Richmond, Surrey, UK ISSN 0071-5794 ISSN 2330-202X (electronic) Flora Neotropica ISBN 978-3-030-82358-0 ISBN 978-3-030-82359-7 (eBook) https://doi.org/10.1007/978-3-030-82359-7 © The Editor(s) (if applicable) and The Author(s), under exclusive license to Springer Nature Switzerland AG 2021 This work is subject to copyright. All rights are solely and exclusively licensed by the Publisher, whether the whole or part of the material is concerned, specifically the rights of translation, reprinting, reuse of illustrations, recitation, broadcasting, reproduction on microfilms or in any other physical way, and transmission or information storage and retrieval, electronic adaptation, computer software, or by similar or dissimilar methodology now known or hereafter developed. The use of general descriptive names, registered names, trademarks, service marks, etc. in this publication does not imply, even in the absence of a specific statement, that such names are exempt from the relevant protective laws and regulations and therefore free for general use. The publisher, the authors, and the editors are safe to assume that the advice and information in this book are believed to be true and accurate at the date of publication. Neither the publisher nor the authors or the editors give a warranty, expressed or implied, with respect to the material contained herein or for any errors or omissions that may have been made. The publisher remains neutral with regard to jurisdictional claims in published maps and institutional affiliations. Cover illustration: Project Flora da Reserva Ducke INPA/DFID This Springer imprint is published by the registered company Springer Nature Switzerland AG The registered company address is: Gewerbestrasse 11, 6330 Cham, Switzerland This monograph is dedicated to Jose Cuatrecasas (1903–1996) pioneer in the study of the Humiriaceae Acknowledgements I thank Andrew Brown for the drawings of each genus (Fig. 10.1, 10.3, 10.5, 10.9, 10.11, 10.16, 10.18, 10.21, 10.24, and 10.26) and Michella del Rei and Matrisabel Adrianzén for Fig.10.12. I thank Rachel Crosby for much help with compiling the database for mapping. I am grateful to the herbarium of the Royal Botanic Gardens Kew for facilities to carry out this work and to Elizabeth Gjieli for the preparation of the distribution maps. I thank Thomas Zanoni and Charles Zartman for most helpful reviews of the original manuscript, and Ana Maria Ruiz for editorial assis- tance. Thanks to the IUCN Red List Unit in Cambridge, UK and to Barbara Goettsch who worked with me to determine the conservation status for each species. I thank The International Cosmos Prize for the funding that enabled this study. vii Abstract Ghillean T. Prance (Royal Botanic Gardens, Kew, Richmond, Surrey, TW9 3AB, UK). Humiriaceae. Flora Neotropica Monogr. 122. A taxonomic monograph of the Humiriaceae is presented. The family is almost exclusively neotropical with a single species in West Africa (which is included here). Humiriaceae is a member of the order Malpighiales close to the Erythroxylaceae, Ctenolophonaceae and Linaceae. Sixty-five species in 8 genera are treated as well as 14 subspecific taxa. Two new species are added, Humiriastrum purusensis and Vantanea aracaensis. Introductory material on the wood anatomy and pollen and on the systematic position of Humiriaceae is provided. Full descriptions, nomenclature, identification keys, at least one illustration of each genus, and distribution maps of all species are provided. Resumo Ghillean T. Prance (Royal Botanic Gardens, Kew, Richmond, Surrey, TW9 3AB, UK). Humiriaceae. Flora Neotropica Monogr. 122. Esta monografia presenta os oito gêneros da família Humiriaceae. Todas espécies, menos uma da África, são neo- tropicais, distribuidos de Nicaragua até o sul do Brasil. Humiriaceae pertence à Malpighiales perto à Erythroxylaceae, Ctenophonaceae e Linaceae. Há 65 espécies em oito gêneros. São descritas duas espécies novas, Humiriastrum purusensis, e Vantanea aracaensis. São incluidas descrições, nomenclatura, sinonímia, chaves para identificação, e pelo menos uma illustração de cada gênero, além mapas de distribuição de todas as espécies. ix Contents 1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 2 Taxonomic History of Humiriaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 3 Relationships and Phylogeny . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 4 Fossils . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11 5 Wood Anatomy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13 6 Pollen . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15 7 Chemistry . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17 8 Chromosomes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19 9 Morphology of Humiriaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21 9.1 Habit . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21 9.2 Leaves . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21 9.3 Vestiture . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21 9.4 Inflorescence . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22 9.5 Calyx . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22 9.6 Corolla . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22 9.7 Stamens . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22 9.8 Disc . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22 9.9 Gynoecium . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23 9.10 Fruit . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23 9.11 Seeds . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25 10 Systematic Treatment of Humiriaceae . . . . . . . . . . . . . . . . . . . . . . . . . 27 10.1 Duckesia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30 10.2 Endopleura . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 35 10.3 Hylocarpa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40 10.4 Humiria . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 44 10.5 Humiriastrum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 70 xi xii Contents 10.6 Sacoglottis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 98 10.7 Schistostemon . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 127 10.8 Vantanea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 143 Numerical List of Taxa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 181 List of Exsiccatae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 185 References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 229 Index of Local Names . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 235 Index of Scientific names . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 239