The Genus Diplusodon (Lythraceae) (Flora Neotropica, 122) 3030658740, 9783030658748

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Flora Neotropica 122

Taciana Barbosa Cavalcanti

The Genus Diplusodon (Lythraceae)

Flora Neotropica Volume 122

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

Taciana Barbosa Cavalcanti

The Genus Diplusodon (Lythraceae)

Taciana Barbosa Cavalcanti

Plant Systematics Laboratory

Herbarium CEN Embrapa Genetic Resources and Biotechnology Brasilia, Brazil

ISSN 0071-5794     ISSN 2330-202X (electronic) Flora Neotropica ISBN 978-3-030-65874-8    ISBN 978-3-030-65875-5 (eBook) https://doi.org/10.1007/978-3-030-65875-5 © 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 reserved 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. This Springer imprint is published by the registered company Springer Nature Switzerland AG The registered company address is: Gewerbestrasse 11, 6330 Cham, Switzerland

Preface

My entire career as a botanist has led to this book. My work started in the master’s program at the University of São Paulo, studying the floristics of Lythraceae in Serra do Cipó, in the Espinhaço Range, Minas Gerais state, Brazil. Here, I first encountered Diplusodon in the field. Due to the beauty of its flowers, the diversity of the genus, and taxonomic questions, my supervisor, Dr. Ana Maria Giulietti, and I realized that the genus would be an excellent subject for Ph.D. dissertation and a good candidate for the Flora Neotropica Series. During my doctoral studies, I visited seven international herbaria and was able to see the original material of almost all species of Diplusodon. I got to know the interior of Brazil during several expeditions to collect plants, and encountered a wide diversity of species of Diplusodon, native to the mountains of Goiás and Minas Gerais, with restricted distributions and not yet known to science. Visits to Brazilian herbaria also brought more and more novelties in the genus and the constant feeling that each small mountain range I visited would bring something new, leading me to continue exploring new locations in the Cerrado before finalizing the taxonomic revision of Diplusodon. In 1990, I accepted a position as a researcher at Embrapa Genetic Resources and Biotechnology (Embrapa Cenargen), in Brasília, where I still work. At Embrapa Cenargen, I took on new assignments that the work required, including projects related to collection and conservation of the Cuphea species throughout the range of the genus. In 1988, the USDA contacted Embrapa Cenargen to collaborate on a Cuphea germplasm collection expedition program in Brazil. The collaboration between the two institutions was established, and as a taxonomist, I was in charge of activities such as collecting Cuphea germplasm in Brazil, ex situ germplasm conservation, and conducting taxonomic studies in Cuphea, aiming at definition of the limits and variability of their species. I also conducted and oversaw studies related to cytogenetics, phenology, phylogeny, and population to guide sampling strategies for collecting germplasm.

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Although the work with Cuphea at Embrapa made my exclusive dedication to the taxonomic revision of Diplusodon impossible, it greatly facilitated visits to Brazilian and foreign herbaria and the exploration of new areas for the collection of Lythraceae throughout Brazil, especially in the central plateau of Brazil, and I quickly discovered that the mountain chains of this geographic region was even richer in species of Diplusodon than the mountains of Espinhaço Range in Minas Gerais and Bahia. Many new species in Lythraceae and even Diplusodon were published during this period. In 2008, Brazil affirmed its commitment to comply with the Global Strategy for Plant Conservation (GSPC) as part of the Conference of the Parties to the Convention on Biological Diversity (CBD) and initiated the “Flora do Brasil Project,” and established a “Management Committee of Flora do Brasil.” I am part of this committee and I was part of the strategies definition for formatting the “List of species of Flora of Brazil” (2010) and “Flora do Brasil” online (2020). I collaborated in the preparation of documents and articulation of strategies to orchestrate the work of 979 botanists from 224 institutions, and I am also the author of the Lythraceae from Brazil for this ambitious project. Much of my work with Lythraceae has been done in collaboration with Dr. Shirley A.  Graham (Missouri Botanical Garden) and Glocimar Pereira da Silva (Embrapa Cenargen). During this time, we have gathered rich collections of Lythraceae (deposited at the CEN Herbarium), and studied and published articles on Lythraceae from several Brazilian states, which culminated in 2020 with the work on Lythraceae from Brazil available online on the “Flora do Brasil” website. In these 30 years at Embrapa Cenargen, I have also coordinated the publication of the series Flora do Distrito Federal, Brazil, which now has 14 published volumes, and I have supervised master’s and Ph.D. students in botany at the University of Brasília, among them is Marlon Garlet Facco, a young researcher who will continue his studies of Lythraceae with me. The 40-year experience in the study of neotropical Lythraceae brings in 2021 the satisfaction of publishing this book, bringing together information from all known Diplusodon species into a single volume, providing a comprehensive overview of the genus, its range of distribution, its environmental requirements, endemism by region, along with the opportunity to highlight persistent unanswered questions and the satisfaction of making this knowledge available through the Flora Neotropica series. Taciana Barbosa Cavalcanti

Brasília, Brazil

Acknowledgments

This monograph began in 1986 as my doctoral thesis at the University of São Paulo (Herbarium SPF) and with my studies in 1989 at the Royal Botanic Gardens, Kew (Herbarium K), in London. The work was continued at the Embrapa Genetic Resources and Biotechnology (Brasília, DF, Brazil – Herbarium CEN), with studies at the New York Botanical Garden (New York, U.S.A. – Herbarium NY) and at the Missouri Botanical Garden (Saint Louis, Missouri, U.S.A. – Herbarium MO). Visits have been made to study original collections in the following herbaria: Botanic Garden and Botanical Museum Berlin-Dahlem, Berlin, Germany (B); Botanic Garden Meise, Meise, Belgium (BR); Conservatory and Botanical Gardens of Geneva, Switzerland (G); Munich Botanical Garden (Botanische Staatssammlung München), Munich, Germany (M); Muséum National d’Histoire Naturelle, Paris, France (P); Natural History Museum of Vienna, Vienna, Austria (W); Naturalis Biodiversity Center (Leiden, The Netherlands), Royal Botanic Gardens, Richmond, England (K); British Museum of Natural History, London, England (BM); University of Oxford, England (OXF); Royal Botanic Garden, Edinburgh, Scotland (E); Smithsonian Institution, Washington, U.S.A. (US). To David Cutler, Peter Gasson, Mary Gregory, for use of the facilities for anatomical studies at the Jodrell Laboratory, Kew. To Madeline Harley for orientation at the Palynology Laboratory at Kew. To researchers at the Royal Botanic Gardens, Kew, Brian Stannard, Eimear Nic Lughadha, Gwillam Lewis, and Raymond Harley, and my advisor at Kew, Simon Mayo, for their support during the working period at Herbarium K. Special thanks to Hans-Joachim Esser, curator of Herbarium M, for sending valuable information and images about the Martius’s collections and comments on the manuscript; the Herbarium OXF staff, Dennis Filer, Serena K.  Marner, and Stephen Harris (curator), for sending images and barcodes of type material; Marc Jeanson, responsible for collections of Herbarium P, for sending information on barcodes of type material; and Vladimir Dorofeyev and Larisa V.  Orlova at Herbarium LE, Komarov Botanical Institute. To Barbara M.  Thiers, Douglas Daly, Jacquelin Kallunki, Patricia K. Holmgren, Piero Delprete, Scott Mori (in memorian), Wayt Thomas, and Amy Litt for their support in using the facilities at NY, and Wayt and Piero for the rich discussions about typifications in Diplusodon. To Peter H. Raven, vii

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James C.  Solomon, John Pruski, Shirley A.  Graham, and Victoria Hollowell for their support in using the facilities at MO.  To Bruno Wallnöfer for the valuable information on the Pohl’s collections. I would like to thank all technical staff and curators at the institutions mentioned above and to the following herbaria for the contribution facilitating the loans of specimens and/or photographs and/or for permitting access to their collections, as well as providing important information on types and collector’s field books: B, BHCB, BM, BR, CEPEC, CESJ, CH, ESA, ESAL, F, G, GFJP, HB, HEPH, HPN, HRB, HRCB, HUEFS, HUFU, HTO, IAN, IBGE, ICN, INPA, K, LE, M, MBM, MG, MO, NY, OXF, OUPR, P, R, RB, SP, SPF, SPSF, TEPB, UB, UEC, UFG, UFMT, US, and W. To Tarciso Filgueiras (in memorian) for the discussions about Diplusodon, and help with the Latin diagnoses and problems with nomenclature. To Gabriel H. Rua for his participation in the studies of inflorescence typology. To Peter Inglis for his valuable participation in the studies of molecular phylogeny and Ana Y.  Ciampi, Vania Cristina Rennó Azevedo, and Zilneide Amaral for the use of the Genetics Laboratory at Embrapa/Cenargen. To Sergio Eustáquio Noronha for his dedicated and accurate work on mapping the geographic data and preparation of the maps. To my botanicals colleagues at the University of São Paulo (Herbarium SPF), especially José Rubens Pirani, Inês Cordeiro and Renato Mello Silva (in memorian), who accompanied the beginning of this work and the first field trips. To Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES, Ministry of Education) for the grant to conduct research at the Royal Botanic Gardens, Kew. To the National Council of Scientific and Technological Development (CNPq, Ministry of Science and Techonology) for the research grant (PQ no 300976/95-0). To the Brazilian Agricultural Research Corporation (EMBRAPA), CNPq, and especially Ariane Luna Peixoto for the grant to continue research at the New  York Botanical Garden and Missouri Botanical Garden. To Emiko Naruto (in memoriam) for the magnificent line drawings, and also for the other illustrators Darli Nuza, Marina de Souza Gomes, and Jeanitto Sebastião Gentilini. To Embrapa Genetic Resources and Biotechnology for providing laboratories, field supplies, and vehicles. To Mauro Carneiro for granting me extra time to work on this monograph during the period I was holding the position of the  head of R&D at Embrapa Cenargen. To the Herbarium CEN staff, Aécio Amaral dos Santos, Andrea Claudia Santana Santoro, Andrielle Câmara Amaral Lopes, Elisângela dos Santos, Gabriela Silva Riberio, João Benedito Pereira, José Geraldo Alves Vieira (in memoriam), Juarez Pereira do Amaral, Nilton Ferreira Barbosa, and Valdeci Ferreira Gomes, for their constant support in managing, sorting, and solving problems with the specimens. To Andrielle Câmara Amaral, Giselle Lopes Moreira, José Floriano Pastore, João Bernardo de Azevedo Bringel, Thaisa Sales Villar, Marlon Garlet Facco, Laiana Moraes Brauner, Moises Mendoza, and André Luiz Moreira for sharing moments and helping in the final moments of this work, and especially to Micheline Carvalho Silva, for the essential help with the data organization and encouragement, and to João Bernardo de Azevedo Bringel for helping in checking type-specimens images in virtual herbaria. To Andrielle Câmara Amaral Lopes for helping in organizing the figures and Gabriela Ribeiro for helping with specimens and database, my good friends. To my colleagues of botany and

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ecology at Embrapa/Cenargen, Anderson Castro Sevilha, Bruno Machado Teles Walter, Daniel Luiz Macias Vieira, Luciano de Bem Bianchetti, Marcelo Brilhante de Medeiros, and Marcelo Fragomeni Simon, and all my students for discussions and encouragement. A special thanks to Shirley A. Graham, a partner in science, whose research with the Brazilian Lythraceae brought us together in 1991, and who became a special friend; thank you Shirley for sharing your experience, for the stimulating discussions, and constant incentive. To Glocimar Pereira da Silva, for his friendship and valuable help with my field work in Brazil, for the large number of rich Diplusodon specimens that he found during his field work in Goiás and Tocantins, for his tireless search for specimens in the “type-localities” of Diplusodon, and for his support in administration of other projects while I was working on this monograph. To Joseph Kirkbride, Marccus Alves, and Frank Almeda for the detailed review that enriched this work. To Piero Delprete for the kind offer to read the manuscript and for the valuable information about Casaretto collections and rich suggestions to the text. To Lawrence M. Kelly, Editor-in-Chief of Flora Neotropica, for detailed reading of this manuscript and valuable suggestions. To Aldicir O. Scariot for the technical contribution and logistical support to finalize this work, and especially for the encouragement and affection. To my friend Graça Wanderley, who always encouraged me. To my beloved family, Aldicir O.  Scariot; Ionaldo Cavalcanti, my father (in memorian); Maria Zuleide, my mother, who gave all the support at home; Carlos André, my brother; Ana Elizabeth, my sister in law; and my dear friend and cousin, Ana Maria Giulietti, for accompanying and encouraging my work life in Botany. To my beloved Andrew, my son, and Mariana, my goddaughter, to whom I dedicate this work.

Abstract

Diplusodon is a monophyletic genus of shrubs and subshrubs, with showy, 6-merous, actinomorphic flowers, and floral tubes on which the sepals alternate with conspicuous epicalyx segments. The capsular fruit contains winged seeds and, uniquely for the family, is divided by a bipartite placenta with semi-lunate septa. Diplusodon is the second largest genus in the Lythraceae and occurs mostly in the Cerrado Biome, the floristically diverse savannah that covers more than 2 million km2 of the Central Brazilian Plateau, extending west into Bolivia, south to Paraguay and east to the Caatinga. A high degree of narrow endemism is evident, with many species restricted to specialized microhabitats of campos rupestres on quartzite rocks, while others are found in cerrado and more rarely, in gallery, or dry forest margins. A total of 104 species and 8 varieties are recognized in the genus, for which 47 lectotypes, 1 neotype, 1 new status and 1 new combination are designated, 9 new species are described, and 15 taxa are placed in synonymy. New information on floral and vegetative morphology, pollen, cytology, chemistry, floral biology, and habitat are provided for the genus. In addition, keys to the species are accompanied by descriptions, illustrations, distribution maps, and assignment of conservation status. Molecular phylogenetic analysis using calibrated Bayesian analysis suggests that speciation was rapid, with a mean date estimate of 3.46 Ma for the origin of Diplusodon (95% HPD = 2.46–4.57 Ma). Species with the most plesiomorphic features occur in the clades east of the São Francisco River (Espinhaço Range and Chapada Diamantina clades), while the greatest morphological diversity is found in species west of the São Francisco River (Plateau Goiano clade). With a predicted late Pliocene date for the crown node of the genus, lineages likely diversified in the campos rupestres, where most of the later Pleistocene speciation bursts also appear to have occurred. These patterns of diversification in Diplusodon appear to agree with Old Stable Landscapes theory, in which high levels of in situ speciation in islands of campo rupestre, isolated by the greater fire-susceptibility of the surrounding cerrado vegetation, resulted in accumulation of high numbers of microendemic species. However, the adaptability of the genus is evidenced by multiple dispersal events to lower elevation cerrado, and the presence, in several clades, of species found both in

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campo rupestre and cerrado habitats. The chromosome number seems to be constant for the genus (n = 15) indicating that speciation took place at the diploid level. As x = 8 is the base number for the family, the origin of the genus is most likely a reduction from n = 16.

Contents

 1 Introduction��������������������������������������������������������������������������������������������������  1   2 Historical Survey������������������������������������������������������������������������������������������  3  3 Morphology��������������������������������������������������������������������������������������������������  9 3.1 Habit and Life Forms����������������������������������������������������������������������������  9 3.2 Indumentum, Trichomes, and Colleters����������������������������������������������  11 3.2.1 Indumentum����������������������������������������������������������������������������  11 3.2.2 Trichomes�������������������������������������������������������������������������������  12 3.2.3 Colleters����������������������������������������������������������������������������������  15 3.3 Leaves ������������������������������������������������������������������������������������������������  15 3.3.1 General Aspects����������������������������������������������������������������������  15 3.4 Leaf Anatomy��������������������������������������������������������������������������������������  22 3.5 Inflorescences��������������������������������������������������������������������������������������  27 3.5.1 Typology ��������������������������������������������������������������������������������  28 3.5.2 Evolution and Diversification ������������������������������������������������  32 3.6 Bracts, Prophylls, and Peduncles��������������������������������������������������������  33  4 Flowers��������������������������������������������������������������������������������������������������������  37 4.1 Androecium����������������������������������������������������������������������������������������  41 4.2 Gynoecium������������������������������������������������������������������������������������������  42   5 Fruit, Seeds, Dispersal, and Germination������������������������������������������������  45 5.1 Fruit����������������������������������������������������������������������������������������������������  45 5.2 Seeds ��������������������������������������������������������������������������������������������������  46 5.3 Dispersal and Germination ����������������������������������������������������������������  47  6 Pollen ����������������������������������������������������������������������������������������������������������  51   7 Floral/Reproductive Biology and Phenology������������������������������������������  55  8 Chromosomes ��������������������������������������������������������������������������������������������  63  9 Chemistry����������������������������������������������������������������������������������������������������  65

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10 Phylogeny and Evolution��������������������������������������������������������������������������  67 11 Geographic Distribution and Ecology������������������������������������������������������  73 12 Conservation����������������������������������������������������������������������������������������������  87 13 Systematic Treatment��������������������������������������������������������������������������������  91 13.1 Artificial Key to the Species of Diplusodon ��������������������������������������  94 14 Insufficiently Known Species������������������������������������������������������������������  459 15 Excluded Names ��������������������������������������������������������������������������������������  465 16 Color Plates����������������������������������������������������������������������������������������������  467 Literature Cited�����������������������������������������������������������������������������������������������  473 Numerical List of Taxa������������������������������������������������������������������������������������  481 List of Exsiccatae����������������������������������������������������������������������������������������������  485 Index of Scientific Names��������������������������������������������������������������������������������  513

About the Author

Taciana B. Cavalcanti  obtained a graduate degree in biological sciences (1984), a master’s degree in biological sciences (1988), and a PhD in botany (1995) from the University of São Paulo, with an experimental part of the PhD at the Royal Botanic Garden, Kew, London, England (1989). Since 1990, she has been working as a researcher at the Embrapa Genetic Resources and Biotechnology of Brazilian Agricultural Research Corporation, and a year later became a professor and advisor in the master’s and doctorate course in botany at the University of Brasília, where she supervised several studies at master’s and doctoral level. Taciana’s contribution in the field of botany and genetic resources has an emphasis on taxonomy, biosystematics, floristics, phylogeny, biogeography, and plant germplasm collecting in areas under impact. From 2008 to 2014, Taciana served as head of research and development at Embrapa Genetic Resources and Biotechnology. From 2008 to 2012 she served as editor of the taxonomy area of ​​the Brazilian journal Acta Botanica Brasilica, and from 2008 to present is Member of the Steering Committee of “Flora do Brasil 2020”. From 1991, Taciana is curator of Herbarium CEN at Embrapa Genetic Resources and Biotechnology, and coordinator and editor of the series Flora of the Federal District, Brazil, which consists of volumes that present botanical families with descriptions, keys and illustrations, and conservation status. As specialist in the Lythraceae family, Taciana’s research spans three decades centered on the vast study of the Lythraceae of Neotropics. Her long-term monographic studies of two largely tropical American genera, Cuphea and Diplusodon, have resulted in several publications on these groups on morphological and evolutionary aspects, including the monograph Diplusodon, which was an important part of Taciana’s career. In 2001, she was a visiting researcher at The New York Botanical

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Garden, USA, in 2005, a visiting researcher at Missouri Botanical Garden, USA, and from 2019 to 2020, a visiting researcher at Wageningen University, the Netherlands, working in collaboration with researchers of the Biosystematics group on phylogeny and biogeography of Lafoensia (Lythraceae) based on Next-­ Generation Sequencing methodology and chloroplast genome.

Chapter 1

Introduction

Lythraceae (sensu lato) is composed of 27 genera and approximately 600 species. The family consists of subshrubs, shrubs, small to very tall trees, and less often herbs. They occupy diverse habitats, including swamps, dunes, humid tropical forests, and especially cerrados and dry or rocky campos. The woody genus Diplusodon Pohl is one of the ten genera of Lythraceae that is native to Brazil, and with 104 species, it is the second largest genus in the family, after Cuphea, which has approximately 240 species (Graham et al. 2013). Diplusodon is characterized by a predominantly subshrubby or shrubby habit, showy 6-merous actinomorphic flowers, floral tubes on which the sepals alternate with conspicuous epicalyx segments, a style turned to one side of the corolla, and two whorls of stamens. The capsular fruit is divided by a bipartite placenta with lunate septa and contains winged seeds. The species inhabit environments including grassy fields, cerrados of low elevation, and, less frequently, dense cerrados and margins of gallery and dry forests. However, the species are predominantly narrow-­ endemic to the campos rupestres and high-altitude rocky cerrados, where the largest diversity of Diplusodon species is found, occurring in small populations and often restricted to just one or a few adjoining mountains. The generic name Diplusodon is compound of the Greek words diplus (double) and odon (tooth), referring to the calyx and epicalyx whorls of the flowers.

© The Author(s), under exclusive license to Springer Nature Switzerland AG 2021 T. B. Cavalcanti, The Genus Diplusodon (Lythraceae), Flora Neotropica, https://doi.org/10.1007/978-3-030-65875-5_1

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Chapter 2

Historical Survey

The genus was described by Pohl (1827a) in the March issue of Flora, and later Pohl (1827b) described and illustrated 16 species in his work Plantarum Brasiliae icones et descriptiones hactenus ineditae. In both of Pohl’s 1827 publications, specimens were not cited, but in the second publication (1827b), the author cited localities. Pohl did not designate a type species for the genus. Other generic names were published with the same morphological delimitations as Diplusodon, but all were subsequent to Pohl’s March 1827 publication. Friedlandia Cham. & Schltdl., the generic name created by Martius and applied by him to diverse material in his herbarium, was published by Chamisso and Schlechtendal in June 1827. Diplodon Sprengel, created by Sprengel, was published in 1830 in Genera Plantarum. Other names for Diplusodon are found in the literature, but are not cited in the synonymy of the genus because they were not effectively published. Candolle (1828), Saint-Hilaire (1833), and Koehne (1877) accepted the name Diplusodon but noted that it was irregularly constructed and a better name would have been Diplodon. Lourteig (1989) designated Diplusodon virgatus, a species described by Pohl (1827a) when he established the genus, as a type of Diplusodon. She justified this choice by the fact that Koehne (1903) used this species to illustrate the genus in his world monograph of Lythraceae. Following the original description of Diplusodon, additional new taxa were added to the genus, along with proposals for infrageneric groups. Candolle (1828) added 20 new species and was the first author to recognize 4 informal groups based on differences in leaf venation: (1) linear, univeined leaves; (2) three-veined leaves, with one central vein and two lateral veins; (3) pinnate-veined leaves; and (4) palmately multiveined leaves. Saint-Hilaire (1833) listed ten species from Minas Gerais and Goiás in the Flora Brasiliae Meridionalis, describing two new species, D. rosmarinifolius A.St.-Hil. and D. kielmeyeroides A.St.-Hil., and a series of infraspecific taxa.

© The Author(s), under exclusive license to Springer Nature Switzerland AG 2021 T. B. Cavalcanti, The Genus Diplusodon (Lythraceae), Flora Neotropica, https://doi.org/10.1007/978-3-030-65875-5_2

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2  Historical Survey

Koehne (1877) recognized 42 species for the states of Minas Gerais, Goiás, and Bahia. In this first work leading to his hallmark revision, he presented the first key for identification of Diplusodon species and organized them into groups based, as did Candolle (1828), on leaf venation. Some years later he published a series of works on Lythraceae and Diplusodon, which focused on taxonomy, floristics, morphology, and phytogeography (Koehne 1881, 1882a, b, 1883, 1886, 1887, 1893, 1897, 1900, 1903) and resulted in his great summary for the family (Koehne 1903). The 1903 monograph of Koehne merits special recognition, because it was the most complete treatment of Diplusodon up to that time; it formed the basis of all taxonomic work on the genus that followed. It included detailed analyses of the principal morphological characteristics, with rich detail on their variations, and generated a great number of infraspecific varieties and forms. Koehne (1903) recognized 53 species, of which 52 occurred in the states of Minas Gerais and Goiás. He proposed an infrageneric classification consisting of 4 sections and 15 series based on leaf venation (Table 2.1). Following his revision (Koehne 1903), Koehne (1907) added one new species to the genus, Diplusodon ulei Koehne from Bahia, bringing to 53 the number of species that he recognized. Pilger (1937) described D. bradei Pilger and D. mononeuros Pilger, both species from Minas Gerais. Lourteig (1963) described two new species from Goiás, D. astictus Lourteig and D. caesariatus Lourteig, and in the following year (Lourteig 1964) she described D. thysanosepalus Lourteig from Goiás and D. smithii Lourteig and D. buxifolius (Cham. & Schltdl.) DC. var. naudinii Lourteig from Minas Gerais. In 1988, Cavalcanti produced a taxonomic treatment of Lythraceae for Serra do Cipó in Minas Gerais, which initiated this revision of Diplusodon. Later, Lourteig (1989) published a synopsis for Diplusodon in which most of the species recognized by Koehne were maintained and 18 new species were described, mostly from Goiás. Lourteig (1989) utilized Koehne’s sections (1877, 1903), except that she synonymized section Palmatipenninerves Koehne with section Palmatinerves Koehne, on the basis that both sections are characterized by having more than one vein arising from the base of the central rib, and correctly referred to section Subuninerves Koehne as section Diplusodon. She typified each section, selecting as types D. virgatus for section Diplusodon, D. oblongus Pohl for section Penninerves, D. rotundifolius DC. for section Palmatinerves, and D. villosissimus Pohl for the synonymized section Palmatipenninerves. The molecular phylogenetic analysis of Diplusodon performed by Inglis and Cavalcanti (2018) provides strong evidence for the rejection of the existing taxonomic classification of Koehne (1877, 1903) and Lourteig (1989). The taxonomic sections are polyphyletic, corroborating the anatomical and X-ray studies of Diplusodon. In addition, the results of Inglis and Cavalcanti (2018) do not support the taxonomic sections inferred based on flavonoid analysis (Blatt 1991; Blatt et al. 1993, 1994) and disagree with the phylogenetic hypothesis within Diplusodon proposed by the authors in which section Penninerves was assumed to be primitive in relation to sections Diplusodon and Palmatinerves based on patterns of flavone and flavonol distribution (Blatt et al. 1994; Salatino et al. 2000).

2  Historical Survey

5

Table 2.1  Koehne’s (1903) and Lourteig’s (1989) subgeneric classifications of Diplusodon. Section Subuninerves (= Diplusodon)

Series Two, based on a revolute or plane leaf margin

Characterization Leaves with a single central vein or 2–4 inconspicuous lateral veins

Koehne (1903) D. candollei, D. ciliiflorus, D. glaziovii, D. gracilis, D. hexander, D. myrsinites, D. punctatus, D. uninervius, D. virgatus

Lourteig (1989) D. astictus, D. candollei, D. ciliiflorus, D. ericoides, D. glaziovii, D. gracilis, D. hexander, D. mononeuros, D. myrsinites, D. psammophilus, D. punctatus, D. ulei, D. uninervius, D. vidalii, D. virgatus (continued)

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Table 2.1 (continued) Section Penninerves

Series Six, based on the pilosity of the plants, type of leaf base, and position of the epicalyx appendages

Characterization Leaves with 1–11 pairs of lateral veins, prominent on the abaxial surface, sunken or only slightly prominent on the adaxial surface

Koehne (1903) D. buxifolius, D. capitatus, D. epilobioides, D. helianthemifolius, D. hirsutus, D. humilis, D. incanus, D. kielmeyeroides, D. lanceolatus, D. lythroides, D. macrodon, D. microphyllus, D. nigricans, D. nitidus, D. oblongus, D. panniculatus, D. parvifolius, D. pulchellus, D. quintuplinervius, D. ramosissimus, D. rosmarinifolius, D. serpyllifolius, D. strigosus, D. subsericeus, D. thymifolius, D. velutinus, D. villosus

Lourteig (1989) D. appendiculosus, D. argenteus, D. bradei, D. buxifolius, D. caesariatus, D. capitatus, D. cordifolius, D. epilobioides, D. hatschbachii, D. helianthemifolius, D. hirsutus, D. incanus, D. irwinii, D. kielmeyeroides, D. lanceolatus, D. leucocalycinus, D. lythroides, D. macrodon, D. microphyllus, D. minasensis, D. nigricans, D. nitidus, D. oblongus, D. panniculatus, D. parvifolius, D. puberulus, D. pulchellus, D. quintuplinervius, D. ramosissimus, D. rosmarinifolius, D. serpyllifolius, D. smithii, D. strigosus, D. subsericeus, D. thymifolius, D. urceolatus, D. villosus, D. xerampelinus (continued)

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Table 2.1 (continued) Section Palmatinerves

Series Two, based on the number of stamens

Palmatipenninerves Five, based on the pilosity of the plants, position of the epicalyx appendages, the form of the floral tube and the number of stamens

Characterization Palmate-veined leaves, 2–6 veins depart at the base of the midvein; none or rarely one departs along the midvein

Koehne (1903) D. glaucescens, D. orbicularis, D. rotundifolius

Palmate-veined leaves, 2–4 veins depart from the base of the midvein; 1–3 depart along the midvein

D. burchellii, D. crulsianus, D. decussatus, D. divaricatus, D. floribundus, D. imbricatus, D. longipes, D. marginatus, D. ovatus, D. retroimbricatus, D. sessiliflorus, D. sordidus, D. speciosus, D. villosissimus

Lourteig (1989) D. adpressipilus, D. burchellii, D. crulsianus, D. decussatus, D. divaricatus, D. fastigiatus, D. floribundus, D. glaucescens, D. heringeri, D. imbricatus, D. longipes, D. marginatus, D. orbicularis, D. ovatus, D. paraisoensis, D. retroimbricatus, D. rotundifolius, D. sessiliflorus, D. sigillatus, D. sordidus, D. speciosus, D. thysanosepalus, D. villosissimus

Since 1990, new contributions on aspects of biology, distribution, conservation, and new species continue to be published from new collections made for a broad-­ scale revision of the genus (Cavalcanti 1995, 1998, 2004, 2007a, b, 2011, 2015; Cavalcanti et al. 2013; Graham and Cavalcanti 2001). A total of 104 species and 8 varieties of Diplusodon are recognized in this study, for which 47 lectotypes, 1 neotype, 1 new status, and 1 new combination are designated, 9 new species are described, and 15 taxa are placed in synonymy.

Chapter 3

Morphology

3.1  Habit and Life Forms Most species of Diplusodon are subshrubs or shrubs, with only three taxa that are small trees (Fig. 3.1E): D. virgatus Pohl var. virgatus, D. ciliatiflorus T.B.Cavalc., and D. heringeri Lourteig. The trees grow to 2–4 m tall, with a slender unbranched trunk and a well-defined, much-branched crown. These taxa grow on the margins of gallery forests and cerrado woodlands. All other species in the genus are subshrubs or shrubs that can be sparsely to much-branched, with a single main stem. In subshrubs (Figs.  3.1A–C), wood is formed from the base to the middle or distal position of the plant. The young upper branches are not woody, hence the designation of “semishrub,” which some authors, such as Eiten (1968), apply to describe this condition. A few species, for example, Diplusodon cordifolius Lourteig, D. helianthemifolius Mart. ex DC. var. pemphoides (DC.) Koehne, and D. parvifolius Mart. ex DC., are considered shrubs (Fig. 3.1D) but are very branched from the base with secondary growth reaching the upper branches. The shrubby or subshrubby species vary greatly in size, from 7 cm to 3 m tall. The subshrubby habit in Diplusodon is commonly associated with a well-­ developed, woody, subterranean, perennial xylopodium. The xylopodia vary in size and form, and they generally produce a single aerial stem with lateral branches (Fig.  3.1F–H). Following seasonal fires, many buds develop a number of stems synchronously, at the same level from the xylopodia, forming a caespitose subshrub (Fig. 3.1B). Raunkiaer (1934) classified these plants as perennial hemicryptophytes. The xylopodia provide good protection for plants against fire, which is a common occurrence in the cerrados. Xylopodia are found in many plants of the cerrados. It is common to find species of Diplusodon that have both an erect, well-­ developed subshrubby habit and a caespitose form, the latter of which flowers precociously and is strongly associated with resprouting following a fire. The occurrence of caespitose forms that are quite distinct from the common form of a species

© The Author(s), under exclusive license to Springer Nature Switzerland AG 2021 T. B. Cavalcanti, The Genus Diplusodon (Lythraceae), Flora Neotropica, https://doi.org/10.1007/978-3-030-65875-5_3

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Fig. 3.1  Habit in Diplusodon. A. Unbranched subshrub. B. Caespitose subshrub. C. Branched subshrub. D. Branched shrub. E. Small tree. F–H. Xylopodia

3.2  Indumentum, Trichomes, and Colleters

11

sometimes creates difficulties for delimitation of taxa (see the commentary for D. ciliiflorus Koehne). The stem, beginning at the region where branches are initiated, generally possesses a rhytidome that splits into thin delicate threads, making it difficult to interpret the type of indumentum and the form of the stem in this region. The stem can, at times, display fistulas, as in Diplusodon marginatus Pohl, but ants have not been observed inside of these stems. In some species of Diplusodon, such as D. buxifolius (Cham. & Schltdl.) DC., D. lanceolatus Pohl, D. leucocalycinus Lourteig, D. orbicularis Koehne, D. paraisoensis Lourteig, D. strigosus Pohl, and D. uninervius Koehne, thickened parts of the stem and lateral branches were confirmed to be galls. The occurrence of galls in Diplusodon was recorded by several authors, such as Coelho et al. (2013) for D. buxifolius, Silva et al. (2018), and Maia and Fernandes (2004) for D. virgatus, as the first record of galls in this species. Marini-Filho and Fernandes (2012) reported that stem galls drain nutrients and decrease shoot performance in D. orbicularis and that they also reduce toxic aluminum concentration in the plant tissues. The upper lateral branches are typically decussate, and the indumentum, if present, is exceedingly variable. The branches can be cylindrical, subquadrangular (when the laterals are slightly flattened), or distinctly quadrangular. Furthermore, the four angles can develop small to prominent wings up to 2  mm wide, as in Diplusodon alatus T.B.Cavalc., or wings can be absent. The upper branches also vary in coloration, for example, brownish in D. bradei, pale yellow in D. astictus, or wine-colored to red in D. glaucescens DC. and D. sordidus Koehne.

3.2  Indumentum, Trichomes, and Colleters 3.2.1  Indumentum After Koehne’s studies, little attention was given to the value of indumentum and trichome types as diagnostic characters in Lythraceae treatment. Recently, more emphasis has again been placed on the great variability in indumentum types as well as trichome morphology. Trichomes have been of considerable importance in taxonomic analyses generally in Lythraceae and especially in Lagerstroemia L. and Cuphea P. Browne (Schoemberg and Hofmeister 1986; Amarasinghe et al. 1991). With respect to Diplusodon, Lourteig (1989) distinguished groups within the sections based solely on indumentum. The great diversity of indumentum and trichome types in Cuphea, however, has not been detected in Diplusodon, and indumentum and trichome types, although helpful in delimiting some taxa, vary widely in others. Variation in the density and types of trichomes was observed within species and among populations. Of special note is the occurrence in Diplusodon leucocalycinus Lourteig of some individuals bearing two types of intermixed trichomes, simple and stellate, while other individuals only have one or the other. The variation in density

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of trichomes among populations has been the basis for recognition of varieties and forms in many species. The following types of indumentum occur in the Diplusodon species (following nomenclature of Radford et al. 1974; Johnson 1975): canescent, ciliate, floccose, hirsute, hispid, papillose, pubescent, sericeous, stellate, strigose, velutinous, and villous. The indumentum can be composed of trichomes that are erect, appressed, oblique, or acroscopic or basiscopic (Uphof 1962). Most Diplusodon species have indumentum on all parts that falls away with age. For this reason, most species are glabrescent, and some are conspicuously pubescent when young. For example, D. adpressipilus Lourteig is characterized by sericeous or silvery indumentum but can have entire branches with leaves that become green or nearly glabrous with age. Other species, such as D. ciliatiflorus, D. decussatus Gardner, D. orbicularis, D. sordidus, and D. speciosus (Kunth) DC., are pilose when seedlings but glabrous at maturity. Some species, for example, Diplusodon ciliatiflorus and D. panniculatus Koehne, have trichomes only on the flowers. When present on flowers, indumentum generally occurs on the floral tube, sepals, and epicalyx. In a few species, such as D. leucocalycinus and D. appendiculosus Lourteig, trichomes are present on the apex of the ovary and persist on the capsule. In some individuals of D. petiolatus (Koehne) T.B.Cavalc., trichomes occur on the anthers.

3.2.2  Trichomes There are five main types of trichomes in Diplusodon: Type 1: Unicellular, non-glandular trichomes with the body elongated, conical, and narrow and the base bulbous (Fig. 3.2A, B, D, G, H, J). This type occurs frequently in species with villous, velutinous, hirsute, and sericeous indumentum on all parts of the plant, for example, Diplusodon argyrophyllus T.B.Cavalc., D. bradei, and D. incanus Gardner. Type 2: Unicellular, non-glandular trichomes forming short protuberances with thick walls and bulbous bases (Fig.  3.2C, N). This type is rare in the genus and occurs in species that are apparently glabrous or puberulous, such as Diplusodon epilobioides Mart. ex DC. and D. puberulus Koehne. Type 3: Unicellular, glandular trichomes that are botuliform and narrow or obtuse at the apex (Fig. 3.2E, F). This type is frequently found on the sepals of Diplusodon hatschbachii Lourteig and D. sessiliflorus Koehne. In D. divaricatus Pohl, these trichomes occur on the sepals and also on the floral tube. Typically, these trichomes have a narrow base and a large body with a thick cell wall and wide lumen. The body of these trichomes may become elongated and slender in the direction of the apex forming a long neck, as can be seen, for example, in D. divaricatus and D. sessiliflorus. The base is enclosed externally by a layer of elongate epidermal cells, probably with a specialized function, which differ from the other epidermal cells in size and arrangement and are strongly united at the base. The basal cells accompany

Fig. 3.2  Trichomes in Diplusodon. A. Type 1, D. candollei (Pereira et al. 790, BHCB). B. Type 1, D. paraisoensis Cavalcanti et  al. 390, CEN. C. Type 2, D. epilobioides Hatschbach 41484, NY. D. Type 1, D. bradei (Zappi et al. [CFCR] 10601, SPF). E. Type 3, D. divaricatus (Rizzo 5113 & Barbosa 4362, UFG). F. Type 3, D. sessiliflorus (Philcox & Onishi 4782, UB). G. Type 1, D. strigosus (Cavalcanti et al. 1086, CEN). H. Type 1, D. adpressipilus (Cavalcanti et al. 400, SPF). I. Type 4, D. oblongus (Cavalcanti et al. 432, SPF). J. Type 1, D. villosus (Cavalcanti et al. 1265, CEN). K. Type 5, D. rosmarinifolius (Cavalcanti et al. 478, CEN). L. Type 5, D. helianthemifolius var. helianthemifolius (Cavalcanti et al. 252, CEN). M. Type 5, D. puberulus (Irwin et al. 22974, UB). N. Type 2, D. oblongus (Cavalcanti et al. 432, CEN). O. Colleter

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the trichome when it falls away. Uphof (1962) called these accessory cells and suggested that they may be responsible for the elevating the trichome, forming a “cushion.” These trichomes are filled with a deep orange substance that is apparently viscous or oily and not water soluble. Fahn (1989) described similar glandular trichomes that he called “stinging hairs,” but with the upper part narrow instead of broad (Figs. 3.2E, F). Type 4: Unicellular, papillate glandular trichomes (Fig. 3.2I). This special trichome occurs on the inner margin of the sepals and is present in all species of Diplusodon, even though they are inconspicuous in most species. Trichomes of this type can be easily seen on the sepal margins of D. appendiculosus. Structurally, they possess a rounded or short, elongated, capitate apex with thick cell walls. In herbarium material of D. appendiculosus, there are hard, orange, translucent drops, like resin, secreted by these papillae along the sepal margins that can be seen on buds prior to anthesis. Type 5: Multicellular, stellate, non-glandular trichomes that occur in tufts and are sessile or have a small stalk (Fig. 3.2K–M). In this type, a single terminal cell is replaced by a group of cells forming a stellate trichome (Uphof 1962). The apical cells are connected only at the base, and the free parts spread horizontally or, when in tufts, are ascending. In some species, a number of contiguous cells grouped around a central cell grow together over the center cell forming the pedestal of a stellate hair (Uphof 1962; Theobald et  al. 1988). Stellate trichomes are found in Diplusodon helianthemifolius var. helianthemifolius, D. helianthemifolius var. pemphoides, D. macrodon Koehne, D. puberulus, and D. rosmarinifolius, among others. In D. helianthemifolius var. helianthemifolius, the stellate trichomes possess a stalk. Lourteig (1989) referred to the trichomes of these species as “dendritic.” However, dendritic trichomes, according to a number of authors (Metcalfe and Chalk 1988; Radford et al. 1974; Uphof 1962) are elongated trichomes, in the form of a tree, with branches or extensions along their length. This is not the case in Diplusodon; therefore, this multicellular type should be called “stellate.” All the morphological trichome types analyzed in this study have high taxonomic value for distinguishing species, especially the stellate and botuliform types, which are rare and characterize just a few species. The glandular papillose trichomes on the internal margin of the sepals, not recognized prior to this study, may be important at the generic level given that Diplusodon is defined in the family by a single autapomorphy, which is the presence of semi-lunate septal walls in the ovary (Graham et  al. 1993). A more extensive survey should be made to check for the occurrence of this character in other genera of Lythraceae. Solereder (1899 apud Uphof 1962) referred to peculiar globose trichomes that occur in some Lythraceae, such as Adenaria Kunth, Pehria Sprague, and Woodfordia Salisb., but he did not mention their presence on sepals, as in Diplusodon. The surface ornamentation of trichomes in Diplusodon can vary from rugulate to striate.

3.3 Leaves

15

3.2.3  Colleters In Lythraceae, there are structures on the branches at the base of the leaves, bracts, and prophylls, which are referred to as stipulate structures or as stipules. Dahlgren and Thorne (1984) described them as characteristic of the order Myrtales and called them stipules, describing them as lateral or axillary, rudimentary, fingerlike projections, infrequently taking the form of long trichomes. They further added that in Diplusodon, the stipules are more advanced, being divided into small structures resembling axillary leaf trichomes. In Diplusodon, these stipule-like structures are on the branches in the region of the leaf insertion near the bud, and they are arranged in a line. They are a type of multicellular trichome, without vascularization, actively secretory,  that stores a dense orange to yellow substance that is possibly mucilaginous (Fig. 3.2O, 13.44I, 13.71.D). Metcalfe and Chalk (1988) recognize that stipules generally occur in pairs and can assume many forms. In general, stipules are vascularized by lateral traces or branches of lateral traces (Metcalfe and Chalk 1988). The term colleter was primarily used by Schumann (1891 apud Metcalfe and Chalk 1988) for secretory structures found on and among the stipules of Rubiaceae. Solereder (1908) described the same structures as “shaggy hairs” and applied the term colleter to trichomes that secret mucilage and are found on axillary buds. For Metcalfe and Chalk (1988), colleters are multicellular glandular structures usually composed of an elongate axis, with reduced epidermal cells that secrete mucilage, gum, or resin and are found on leaves and stipules of many eudicot families. For reasons related to their position and structure, these  stipule-like structures in Diplusodon are here interpreted  as colleters.

3.3  Leaves 3.3.1  General Aspects The phyllotaxy in Diplusodon is invariably opposite and decussate. However, some species, such as D. uninervius Koehne, D. candollei Mart. ex DC., and D. puberulus, have liner leaves that are borne on branches with very short internodes, and the leaves appear to be verticillate or form fascicles. The external leaf morphology is a source of good characters whose states are variable among species and are therefore extensively used in the taxonomy of the genus. The leaves are always entire, and in most species they are coriaceous and deciduous. Leaf form can vary from linear to elliptic, lanceolate, ovate, or orbicular (Figs. 3.3, 3.4 and 3.5). Leaf size, presence or absence of a petiole, presence or absence of domatia, a revolute or plane margin, presence or absence of indumentum, and the type of indumentum can vary widely among the species; for this reason, these features are of taxonomic importance in the genus. Most species possess subsessile leaves, that is, with a petiole 0.5–2 mm long, while others have sessile leaves, and still fewer have long-petiolate leaves

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Fig. 3.3  Patterns of venation in Diplusodon. A. Hyphodromous, D. myrsinites (Krieger 8621, SPF). B. Eucamptodromous, D. virgatus var. virgatus (Irwin et al. 17786, UB). C. Eucamptodromous, D. buxifolius (Zappi & Scatena [CFCR] 10922, SPF). D. Hyphodromous, D. hexander (Cavalcanti et  al. 250, SPF). E. Eucamptodromous, D. ciliiflorus (Cavalcanti et  al. [CFSC] 9630, SPF). F. Eucamptodromous-craspedrodromous/hyphodromous, D. alatus (Menezes 1208, SPF). G. Acrodromous-basal, D. punctatus var. punctatus (Burchell 7659, K). H. Eucamptodromous, D. virgatus var. virgatus (Badini s.n., 3727, OUPR). I. Acrodromous-basal-supranumerary, D. quintuplinervius (Irwin et al. 23070, K). J. Hyphodromous, D. uninervius (Cavalcanti et al. 189, SPF). K. Eucamptodromous-craspedrodromous/hyphodromous, D. alatus (Pirani et al. 1751, SPF). L. Hyphodromous, D. praetermissus (Cavalcanti et  al. 401, CEN). M. Hyphodromous, D. hexander (Glaziou 19175, R). N. Eucamptodromous, D. ciliiflorus (Rossi et al. [CFCR] 3078, SPF). O. Hyphodromous, D. hexander (Souza et al. [CFCR] 8786, SPF). P. Eucamptodromous, D. ciliiflorus (Cavalcanti et al. [CFCR] 9488, SPF). Q. Hyphodromous, D. hexander (Cavalcanti et al. 201, SPF). X-ray images taken at the British Museum of Natural History, London, England

3.3 Leaves

17

Fig. 3.4  Patterns of venation in Diplusodon. A. Eucamptodromous, D. ramosissimus (Hunt & Ramos 6303, UB). B, E. Eucamptodromous, D. oblongus (Ratter et  al. 4075, UB). (continued)

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3 Morphology

(e.g., D. bradei, D. lanceolatus, D. nitidus Mart. ex DC., and D. ramosissimus Pohl have petioles up to 2.5 cm long). The petioles are generally cylindrical and thin, appearing wide and compressed in a few species, such as D. leucocalycinus, D. appendiculosus, and D. paraisoensis. Domatia are found in many species of Diplusodon and provide a significant character for distinguishing some species as D. cordifolius, D. leucocalycinus, D. ramosissimus, among others. The term domatia, as defined by Metcalfe and Chalk (1988), is applied to depressions, pockets, sacks, or tufts of trichomes that are localized in the axils of the midvein on the abaxial surface of the leaves. Frequently in Diplusodon, they are located in the axils of the midvein and the lateral veins, and they may or may not be accompanied by trichomes on the leaf blade. These domatia are classified as the “pocket” type (Metcalfe and Chalk 1988). The pocket-type domatium frequently takes the form of a flattened funnel with a broad distal opening and appears in a reduced form in Diplusodon, for example, in D. villosus and D. incanus. Based on their small size, these may be acarodomatia. The leaves of Diplusodon are also notable for the presence of more than one basic pattern of venation. This feature was first noted by Candolle (1828), who described 35 species in the genus, separating them into 4 unnamed groups based on the following leaf venation characters: (1) univeined, linear leaves (Figs. 3.3D, L, M, O, Q and 3.4G); (2) leaves with 2–3 major veins departing from the base (Figs. 3.4F, I, U, X, B1, D1 and 3.5A–D, F, G, I–K, L, N); (3) pinnate-veined leaves (Figs. 3.3B, E, F, H, K, P and 3.4A, B, E, L, O, Q, R, T, V, A1, C1, G1, H1); and (4) palmately veined leaves (Fig. 3.5M). Koehne (1877, 1903) also highlighted patterns of leaf venation, which he used to establish four sections: Subuninerves, Penninerves,

Fig. 3.4 (continued) C. Eucamptodromous, D. stellatus (Menezes 1209, SPF). D. Eucamptodromous, D. incanus (Gardner 4138, K). F. Acrodromous-basal-supranumerary, D. paraisoensis (Gardner 3721, NY). G. Hyphodromous, D. puberulus (Irwin et  al. 23692, UB). H. Eucamptodromous, D. hirsutus (Glaziou 14700, MO). I. Eucamptodromous, D. argyrophyllus (Harley et  al. 27056, K). J. Hyphodromous, D. gracilis (Gardner 4139, K). K. Eucamptodromous, D. oblongus (Cavalcanti 87, SPF). L. Eucamptodromous, D. bahiensis (Harley et al 17006, SPF). M. Eucamptodromous, D. epilobioides (Martinelli et al. 11163, BHCB). N. Eucamptodromous, D. helianthemifolius var. helianthemifolius (Longhi-Wagner [CFCR] 9325, SPF). O. Eucamptodromous, D. nitidus (Mexia 5862, NY). P,Y. Eucamptodromous, D. rosmarinifolius (Ratter et al. 3133, K). Q. Eucamptodromous, D. hirsutus (Cavalcanti et  al. [CFSC] 10099, SPF). R. Eucamptodromous, D. lanceolatus (Cavalcanti et al. 421, CEN). S. Eucamptodromous, D. microphyllus (Amaral et al. [CFSC] 8403, SPF). T. Acrodromous-basal-supranumerary, D. sigillatus (Irwin et  al. 32190, NY). U. Eucamptodromous, D. helianthemifolius var. helianthemifolius (Menezes et al. [CFCR] 10677, SPF). V. Eucamptodromous, D. lanceolatus (Cavalcanti et al. 303, CEN). W. Acrodromous-basal-­ supranumerary, D. strigosus (Cavalcanti et al. 473, CEN); X. Eucamptodromous, D. parvifolius (Zappi & Kameyama [CFSC] 9636, SPF). Y. Eucamptodromous, D. hirsutus (Glaziou 14699, K). Z. Acrodromous-basal-supranumerary, D. helianthemifolius var. pemphoides (Cavalcanti et  al. [CFSC] 9636, SPF). A1. Eucamptodromous, D. leucocalycinus (Irwin et  al. 24231, UB). B1. Acrodromous-basal-supranumerary, D. villosus (Lima 58-2994, K). C1. Acrodromous-basal-­ supranumerary, D. villosus (Prance & Silva 58184, UB). D1. Acrodromous-basal-supranumerary, D. villosus (Glaziou 21419, K). E1. Eucamptodromous, D. hirsutus (Cavalcanti et  al. [CFSC] 9683, SPF). F1. Eucamptodromous, D. bradei (Zappi et al. [CFCR] 10601, SPF). X-ray images taken at the British Museum of Natural History, London, England

3.3 Leaves

19

Fig. 3.5  Patterns of venation in Diplusodon. A. Acrodromous-basal-supranumerary, D. decussatus (Burchell 8039, K). B. Acrodromous-basal-supranumerary, D. villosissimus (Warming s.n., K). C. Acrodromous-basal-supranumerary, D. panniculatus (Mori et al. 16947, CEN). D. Acrodromous-­ basal-supranumerary, D. paraisoensis (Cavalcanti et  al. 390, CEN). E. Acrodromous-basal, D. orbicularis (Cavalcanti et  al. [CFSC] 9619, SPF). F. Acrodromous-basal-supranumerary, D. speciosus (Gardner 3726, K). G. Acrodromous-basal-supranumerary, D. marginatus (Pereira 1253, IBGE). H. Acrodromous-basal, D. glaucescens (Scatena et  al. [CFCR] 10477, SPF). I. Acrodromous-basal, D. sordidus (Cavalcanti et  al. 403, CEN); J. Acrodromous-basal-­ supranumerary, D. speciosus (Cavalcanti et al. 86, SPF). K. Acrodromous-basal-supranumerary, D. sigillatus (Cavalcanti et al. 382, SPF). L. Acrodromous-basal, D. sordidus (Cavalcanti et al. 30, SPF). M. Acrodromous-basal, D. rotundifolius (Cavalcanti et al. 217, SPF). N. Acrodromous-­ basal, D. orbicularis (Cavalcanti et al. [CFSC] 10623, SPF). X-ray images taken at the British Museum of Natural History, London, England

20

3 Morphology

Palmatinerves, and Palmatipenninerves. This classification was followed until recently when Lourteig (1989) synonymized section Palmatipenninerves with Palmatinerves and correctly changed the name of section Subuninerves to section Diplusodon. In this study, I analyzed the external morphology and venation patterns of the leaves in great detail, in addition to anatomical study of the leaves. The study of the external morphology of the venation patterns, evaluated by X-ray, shows the patterns recognized by Koehne (Figs. 3.3, 3.4 and 3.5). To use venation patterns as a taxonomic character, it is necessary to have a sound understanding of Koehne’s terminology (1877, 1903), in addition to the dicotyledonous leaf architecture classification system of Hickey (1973, 1988). Hickey proposed six basic types of venation, some of which included subtypes. In Diplusodon, adapting the terminology of Hickey (1973, 1988), three basic patterns are recognized: (1) pinnate-hyphodromous, (2) pinnate-camptodromous/eucamptodromous, and (3) acrodromous. 1. Pinnate-hyphodromous (=hyphodromous) – Pattern in which all veins except the primary vein are absent or are present but rudimentary or hidden within a coriaceous or fleshy mesophyll (Hickey 1973, 1988). This pattern occurs in some species of Diplusodon, but not in all the species that Koehne and Lourteig considered members of section Subuninerves (=sect. Diplusodon). The ­pinnate-­hyphodromous pattern will be referred from this point on and in the descriptions only as hyphodromous. X-ray analysis of the leaves, corroborated by anatomical study of the cross-­ section of the mesophyll, indicates that some of the species considered hyphodromous by Koehne (1877, 1903) and Lourteig (1989), for example, Diplusodon hexander Mart. ex DC. and D. puberulus, have a prominent primary vein and secondary lateral veins in which the vascular bundles are so poorly developed that it is sometimes difficult to find them in the mesophyll. In these leaves, viewed macroscopically, the secondary veins are not seen, and the leaves appear to bear only a midvein. In other cases, as in D. candollei, the lateral veins are not seen in microscopic cross-sections of the leaf. The leaves of all such species are described here as hyphodromous (Figs. 3.3D, J, M, O and 3.4G). In other species included in the section Diplusodon by Koehne (1877, 1903) and Lourteig (1989), such as D. appendiculosus, D. astictus, and D. virgatus var. virgatus, the anatomy of the leaf mesophyll includes secondary lateral veins with well-­ developed vascular bundles, minus the sclerenchymatous xylem sheath or with the sheath poorly developed. In this cases though the have lateral are inconspicuous is considered as eucamptodromous pattern. 2. Pinnate-camptodromous/eucamptodromous (=eucamptodromous)  – Pattern in which the secondary lateral veins separate from the primary vein, diminishing apically toward the margin, and are connected to the supradjacent secondaries by a series of cross veins (Figs. 3.3B, E, H, K, M and 3.4A, B, L, N). This pattern is found in the species of section Penninerves (Koehne 1877, 1903) and will be

3.3 Leaves

21

cited here on only as eucamptodromous. In the leaves of Diplusodon bradei and D. decussatus (Figs. 3.3H and 3.4A), some of the secondary veins appear to form arcs, connecting to supradjacent veins, thus showing a pinnate-camptodromous/ brochidodromous pattern; however, the connections to the tertiary veins are weak. 3. Acrodromous – Pattern in which two or more primary veins or strongly developed secondary veins extend in convergent arches toward the apex of the leaf. The arches are not recurved at the base. This pattern is common in Diplusodon, and Koehne included taxa with this pattern in sections Palmatinerves and Palmatipenninerves (=sect. Palmatinerves of Lourteig 1989). Koehne defined section Palmatinerves as possessing conspicuous lateral veins, all of which departed from a single point at the base of the primary central vein. This subtype 1 pattern is here referred to as the acrodromous-basal type. Koehne (1877, 1903) included only Diplusodon glaucescens, D. orbicularis, and D. rotundifolius (Fig. 3.5E, H, M, N) in this section. The section Palmatipenninerves was defined by Koehne (1877, 1903) as possessing two or more pairs of veins separating from the base of the central vein and two or more additional veins diverging along the length of the central vein. This section includes the majority of species of Diplusodon. Almost all the species included in the acrodromous-basal pattern, beyond having the central veins diverging from each side at the base of the central vein, also have supradjacent veins separating along the length of the central vein (Fig. 3.5A, F, L, N). Such veins can be opposite (Fig. 3.5J, K) or alternate (e.g., Fig. 3.5A, F, L, N) and form different angles, from 20° to 60°, in relation to the primary vein. In this cases, this acrodromous pattern, with various primary veins diverging from a single point at the base of the central vein in addition to other veins diverging from the central vein at different points above the base, is considered here as subtype 2, a new pattern, the acrodromous-­ basal-­ supranumerary pattern. Diplusodon also has intersecondary veins (Fig. 3.4H1 and 3.5C), referred to by Hickey (1988) as intermediary veins, among the secondary and tertiary veins that occur most commonly between the secondary veins in pinnate leaves. In Diplusodon marginatus, a fibrous marginal vein composed of the tertiary, quaternary, and lower-­ order veins forms a single vein that runs close to the leaf margin (Fig. 3.5G). Leaf venation characters have long been used as an important source of taxonomic information in Diplusodon. To utilize such characters, however, it is necessary to determine how species fit into the patterns defined here. This requires detailed observations based not only on the external aspects of the leaves but also on their anatomy. Sometimes the leaf venation is very difficult to define, and for this reason, it is sometimes difficult to assign species to section. For example, D. punctatus Pohl in section Diplusodon (Koehne 1877, 1903; Lourteig 1989) has acrodromous leaf venation, D. astictus in section Diplusodon (Lourteig 1989) has eucamptodromous leaf venation, and D. panniculatus in section Penninerves (Koehne 1877, 1903; Lourteig 1989) has acrodromous leaf venation. The Hickey classification of dicotyledonous leaf architecture (1973, 1988) applied to Diplusodon does not lead to obvious evolutionary interpretations, nor is there any

22

3 Morphology

other reference in which the Hickey types can be considered derived from one another. As noted, some patterns described in Diplusodon are classified differently because they occur in leaves with different forms. The venation patterns parallelodromous versus campylodromous versus acrodromous can be cited as an example. The definitions are based fundamentally on two or more primary veins departing from a single point at or near the base of the leaf and all converging at a single point at or near the leaf apex. When the leaves themselves are narrow, the veins are almost parallel from the base to the apex. Similarly when the leaves are ovate with a strongly cordate base, the veins follow the shape of the blade before converging at the apex, thus forming arcs from the cordate leaf base. Therefore, variation in leaf shape sometimes can lead to complications in the interpretation of venation patterns.

3.4  Leaf Anatomy Most species in Diplusodon are heliophytes and mostly inhabit areas subject to high temperatures and water stress. Consequently, they exhibit a series of xeromorphic features such as congested foliage with reduced leaves, coriaceous blades that are frequently concave with a revolute margin, and blades that are often covered by dense indumentum. Their leaf anatomy also reflects adaptation to xeromorphic habitats in some species (Table 3.1). Table 3.1  List of anatomical characteristics for 22 species of Diplusodon Species D. bradei

Crosssection Plane

Leaf cuticle Thick, sinuous

D. candollei

Concave Thick, almost straight

D. epilobioides

Concave Slender, striate

D. glaucescens

Plane

D. helianthemifolius Plane var. helianthemifolius D. heringeri Convex

D. hexander

Slender

Slender

Thick

Concave Slender

Stomata Epidermal surface level Sunken in small cavities

Epidermal surface level Epidermal surface level Epidermal surface level Epidermal surface level Epidermal surface level

Xylem Poorly developed internally and externally Poorly developed internally and externally, few mechanical elements Poorly developed internally and externally Strongly developed Poorly developed internally and externally Strongly developed Strongly developed

Mucilaginous cells layer Poorly developed on both surfaces Strongly developed on adaxial surface

Poorly developed on both surfaces Strongly developed in both surfaces Strongly developed on adaxial surface Strongly developed on adaxial surface Strongly developed on both surfaces

3.4 Leaf Anatomy

23

Species D. leucocalycinus

Crosssection Plane

D. macrodon

Convex

D. marginatus

Plane

D. microphyllus

Plane

D. minasensis

Convex

D. myrsinites

Plane

Slender

D. oblongus

Plane

Thick, almost straight

D. paraisoensis

Plane

Thick

D. panniculatus

D. parvifolius

D. puberulus

D. rotundifolius

D. strigosus

Leaf cuticle Straight

Stomata Sunken in small cavities Striate Sunken in small cavities Slender Sunken in small cavities Slender Epidermal surface level Thick and Epidermal sinuous surface level

Epidermal surface level Epidermal surface level

Sunken in small cavities Plane Straight Epidermal surface level Concave Thick, Epidermal straight surface level Concave Thick and Epidermal sinuous surface level Plane Straight, Epidermal slender surface level Plane Straight Epidermal surface level

D. villosus

Concave Slender

D. virgatus var. virgatus

Plane

Slender

Epidermal surface level Epidermal surface level

Xylem Poorly developed

Strongly developed Strongly developed Poorly developed internally and externally Poorly developed internally and externally, few mechanical elements Poorly developed internally and externally Poorly developed internally and externally, few mechanical elements Strongly developed Strongly developed Poorly developed

Poorly developed

Strongly developed Poorly developed internally and externally, with druses Strongly developed Poorly developed

Mucilaginous cells layer Poorly developed on both surfaces Strongly developed on adaxial surface Developed on adaxial surface Developed on adaxial surface Strongly developed on adaxial surface

Strongly developed on abaxial surface Poorly developed on both surfaces

Strongly developed on adaxial surface Strongly developed on both surfaces Strongly developed on adaxial surface Strongly developed on adaxial surface Strongly developed on both surfaces Poorly developed on both surfaces Strongly developed on adaxial surface Poorly developed on both surfaces

24

3 Morphology

In cross-section, the Diplusodon leaf blade presents different outlines. They can be plane or almost plane, as in D. oblongus, D. glaucescens, D. marginatus, and D. rotundifolius (Fig. 3.6H–J, L); they can have a series of arcs, with furrows on the adaxial surface and protuberances on the abaxial surface as in D. heringeri, D. stellatus T.B.Cavalc., and D. minasensis Lourteig (Figs. 3.6F, G, K); they can be concave (Fig. 3.7B, C), as in D. candollei, D. hexander, and D. puberulus; or they can be conduplicate as in D. conduplicatus T.B.Cavalc. (Fig. 13.26D). The leaf cuticle can vary from very thick and sinuous, as in Diplusodon minasensis and D. puberulus (Fig. 3.7C), or thick and almost straight, as in D. oblongus and D. candollei (Fig. 3.7B), to thin, as in D. glaucescens (Fig. 3.6I), D. helianthemifolius var. helianthemifolius, D. marginatus, D. myrsinites Mart. ex DC., and D. epilobioides. Eberlein (1904) considered D. epilobioides to have a striate cuticular surface, which was also observed in this study. The epidermis is formed by one layer of generally isodiametric medium- to large-sized cells with thick walls and the anticlinal walls frequently curved. Interspersed among the normal epidermal cells, and more or less surrounded by them, are conspicuous mucilaginous cells usually larger than epidermal cells (Figs. 3.6E and 3.7B, C). In some cases, the mucilaginous cells are similar in size to normal epidermal cells and difficult to distinguish. Eberlein (1904) studied the foliar anatomy of ten species of Diplusodon and described the epidermis as one- or two-layered. He noted that mucilaginous epidermal cells occurred in all species examined and that, in some species, the mucilaginous cells could be recognized by their large size and their strong protrusion into the mesophyll. The presence of a second epidermal cell layer (hypodermis), referred to by Eberlein, was not observed in the species analyzed in this study. However some epidermal cells were seen to have divided periclinally with the innermost cell in the mesophyll becoming mucilaginous (Fig. 3.6E). The presence of mucilaginous cells is common in Lythraceae, and they are frequently mentioned as present in the leaves. Fahn (1989) refers to them as large, epidermal, cellular idioblasts containing mucilage. These cells have also been described in Lafoensia Vand. (Lourteig 1986), Koehneria S.Graham (1986), and Lourtella S.Graham (1987). Mucilaginous cells are most commonly found in the adaxial leaf epidermis, but they are also found in the abaxial leaf epidermis, as well as in the mesophyll or in the veins (Gregory and Baas 1989). In leaf cross-sections of some Diplusodon species, it is possible to see mucilaginous cells in the adaxial epidermis (Fig. 3.7A–C) or on both adaxial and abaxial layers and also penetrating deeply into the mesophyll (Fig. 3.6G). In Diplusodon paraisoensis, in the interior of the mesophyll, a cavity surrounded by trichomes was observed. Possibly this is the secretory mucilage cavity referred to by Gregory and Baas (1989) as a mucilage cavity or canal originating by breakdown of groups of mucilage cells or mucilage idioblasts. Mucilage cells in Diplusodon are frequently contracted, possibly the result of the xeric environment in which they grow. The idea that mucilage can serve to store water is defended by several authors (e.g., Solereder 1908; Haberlandt 1914; Gibson 1977; Gregory and Baas 1989). Others believe that the mucilage can reduce

Fig. 3.6  Leaf anatomy in Diplusodon. A. Cross section, D. myrsinites (Krieger 8621, SPF). B. Cross section, D. candollei (Pereira et  al. 932, BHCB). C, J. Cross section, D. marginatus (Pereira 1253, IBGE). D. Paradermal view, D. panniculatus (Mori et al. 16947, K). E, K. Cross section, D. minasensis (Cavalcanti et al. 220, SPF). F. Cross section, D. stellatus (Menezes 1209, SPF). G. Cross section, D. heringeri (Pedralli et al. 3366, CEN). H. Cross section, D. rotundifolius (Cavalcanti et al. 217, SPF). I. Cross section, D. glaucescens (Scatena et al. [CFCR] 10477, SPF). L. Cross section, D. oblongus (Cavalcanti et al. 87, SPF). A–C; F–L the same scale. Images made at Jodrell Laboratory, Royal Botanic Gardens, Kew, London, England

26

3 Morphology

Fig. 3.7  Leaf anatomy in Diplusodon. A. Cross section, D. stellatus (Menezes 1209, SPF). B. Cross section, D. candollei (Pereira et al. 932, BHCB). C. Cross section, D. puberulus (Irwin et al. 22974, UB). A–C the same scale. Images made at Jodrell Laboratory, Royal Botanic Gardens, Kew, London, England

transpiration by acting as a protective layer on the leaf (Esdorn and Schanze 1954; Lyshede 1977, apud Gregory and Baas 1989). The stomata in Diplusodon are most frequently seen on the abaxial epidermis; however, they also occur on the adaxial surface. They can be at the epidermal surface level, as in D. myrsinites (Fig. 3.6A), D. microphyllus Pohl, and D. virgatus (Eberlein 1904), or they can be sunken in small cavities, as in D. candollei (Fig. 3.6B), in which case they may or may not be protected by a dense covering of trichomes. The stomata pattern in Diplusodon is sufficiently variable that it does not

3.5 Inflorescences

27

constitute a good taxonomic character at the species level. The pattern can vary from paracytic, tricytic, or tetracytic to anomocytic on a single leaf. For example, the epidermis of D. panniculatus has tricytic, tetracytic, and actinocytic patterns, all within a small region of the epidermis (Fig. 3.6D). Eberlein (1904) described two, three, or rarely four subsidiary cells surrounding the stomata. The mesophyll is generally composed of one or two layers of palisade-like parenchyma on the adaxial side, spongy parenchyma restricted to the central portion of the mesophyll, and one layer of palisade-like parenchyma on the abaxial surface. It is considered isobilateral (Figs. 3.6E, G and 3.7A). Another pattern of isobilateral mesophyll is that found for Diplusodon oblongus where in cross-section the general condition is a uniform epidermal layer on both surfaces, resembling spongy parenchyma. On close examination, however, it is evident that the cells of the adaxial epidermal are vertically arranged like palisade parenchyma cells, while the cells of the abaxial surface are horizontally arranged (Fig. 3.6H, I, L). The leaves are interpreted as having an isobilateral mesophyll with a tendency toward dorsiventral arrangement. Leaves with clearly dorsiventral mesophyll are exemplified by D. candollei (Fig. 3.7C). Eberlein (1904) considered leaves in the majority of species to be dorsiventral, although the mesophyll sometimes shows a transition toward the isobilateral condition. The species of Diplusodon examined in this study indicate that the structure of the leaf vascular bundles can be diagnostic for the species. The xylem region of the midvein is rich in mechanical elements, assuming a round shape (Figs. 3.6F–I and 3.7A) or an open arc (Fig. 3.6J), and it is accompanied by poorly developed internal phloem and well-developed external phloem. Sometimes the internal phloem is surrounded by a sheath of sclerenchymatous fibers (Fig. 3.6H). In other species, such as D. candollei, D. minasensis, and D. oblongus, the xylem is poorly developed internally and externally and has few mechanical elements (Fig. 3.6K–L). As previously mentioned, the leaf venation of Diplusodon can be prominent or inconspicuous. Initially I thought that the prominence might be due to the presence of a well-developed sclerenchymatous bundle sheath. However, I found that this is the result of a proliferation of parenchymatous cells abaxial to the vascular bundles. This is demonstrated in leaves of D. minasensis, in which the veins are very prominent macroscopically, but the vascular bundle is poorly developed (Fig. 3.6K). Other features recorded from leaves of Diplusodon are mucilage cells in the parenchyma of the veins and, frequently, calcium oxalate crystals (druses) in the mesophyll.

3.5  Inflorescences Inflorescences of all Diplusodon species were dissected and observed using a standard stereoscopic microscope. Observations were made mainly on herbarium material, although living plants were also examined when available. Inflorescence descriptions and terminology are based on the typological system proposed by Troll (1964, 1969; see also Weberling 1965, 1988, 1989; Weberling et al. 1993).

28

3 Morphology

3.5.1  Typology In Diplusodon, the synflorescences are polytelic. Two types of inflorescences are found: simple synflorescences with the axis of the inflorescence unbranched (Fig. 3.8A) and compound synflorescences with the arrangement of the individual flowers of the inflorescence simple and repeated in the branching of the inflorescence (Fig. 3.8B–E). In compound synflorescences, paraclades (hereafter “Pc”) may or may not develop below the main florescence (MF). Leaves of Diplusodon are opposite, verticillate, and fasciculate, and normally paraclades arise from both axils at each node. Troll (1964) subdivided the synflorescence into the following parts: zone of enrichment, for the area that produces paraclades; zone of inhibition, for the area that precedes the zone of enrichment where the growth of paraclades is inhibited; and zone of innovation, exclusive of perennial plants, for the basal portion of the stem in which axillary buds do not develop within the same season, but produce new branches in the next season. The three zones together form the hypotagma. According to Weberling (1988), in woody plants, especially in those that grow under environmental stress, the inflorescence can be reduced, or the zonation of the flowering system can be extremely altered. In some species of Diplusodon, flowering can occur at a stage in which the vegetative part of the plant is practically reduced to an underground system, that is, plants of 3–10 cm can produce many flowers (paedomorphic flowering). Such paedomorphic flowering is the result of an earlier initiation of the inflorescence development in comparison with related species, and the extent to which the onset of the reproductive phase is initiated seems to be controlled by environmental disturbances such as fire. Good examples of this are D. ciliiflorus Koehne and D. pygmaeus T.B.Cavalc., found in the campos rupestres of Diamantina (state of Minas Gerais, Brazil) and Chapada dos Veadeiros (state of Goiás, Brazil), respectively, two regions subject to frequent fires, where populations were found in full flowering and with complete reduction in the inhibition zone. A similar behavior has been observed in other plants of the campos rupestres (rocky outcrops) and cerrados, such as species of Euphorbia (Cordeiro 1986) and Camarea (Mamede 1988). The variation in the relative sizes of the different zones of the flowering systems is great, and it is responsible for the great diversity in inflorescences. With respect to this variation in Diplusodon, Troll (1970) mentions that in D. thymifolius Mart. ex DC. and D. virgatus var. virgatus, the main florescence (terminal on the main axis) remains very small in relation to the paracladial zone, while in other species, such as D. villosissimus Pohl and D. villosus, the principal florescence predominates even though the paracladial zone can also be well developed. On the basis of the degree of branching, the synflorescences can be classified on how many times the paraclades are repeated. A double raceme, an inflorescence simply branched once with paraclades of the first order, is called a diplobotryum (Fig. 3.8B), a triple raceme with paraclades of the first and second order is a triplobotryum (Fig. 3.8C), and those with multiple branching orders (more than three) are

3.5 Inflorescences

29

Fig. 3.8 Inflorescences in Diplusodon. A. Botryum. B. Diplobotryum. C. Triplobotryum. D. Diplothyrse. E. Diplothyrsoid. F. Dichasial cyme. AcB Accessory branch, Pc 1 Paraclade of first order, Pc 2 Paraclade of second order, MF Main florescence, PF Parcial florescence, TF Terminal flower

30

3 Morphology

called, in the case of racemes, pleiobotrya. In addition to paraclades, in synflorescences the inflorescence of the main axis, or main florescence, is also distinguished. It displays the characteristic features of the inflorescence type, and it is used as the basis for defining and classifying the synflorescence as a whole. The principal florescence is situated above the paraclades, in the distal part of the inflorescence axis (Fig.  3.8B–D). When the flowers of an inflorescence are provided with two prophylls, as in the case of Diplusodon, additional flowers can appear in the axils of these prophylls, producing what Troll calls a partial inflorescence, morphologically defined and referred to as a cyme. Inflorescences of Diplusodon are generally polytelic with individual flowers organized in botrya (or racemes). In the species with synflorescences, the principal florescence is, with rare exceptions, also a botryum (Fig. 3.8B, C). All the polytelic species of Diplusodon that have synflorescences having diplo- or triplobotrya bear a main florescence, which for this reason are called heterothetic compound racemes, in contrast to the polytelic synflorescences that are composed only of lateral racemes that lack a principal florescence; the latter are called homothetic compound racemes. Homothetic compound racemes do not occur in Diplusodon. Florescences of the synflorescence, which often appear in Diplusodon, can be spike-like botrya when the flowers are nearly sessile (ca. 1 mm peduncle) or true spikes when the flowers are sessile. Florescence anthesis occurs from the base to the apex (acropetally). In many groups of plants, it is difficult to make a distinction between the region where the inflorescence begins and the vegetative region ends. There are some cases in which the flowering region is distinct from the vegetative region, and in this case, the inflorescence is said to be bracteose. The foliage can also show a gradual reduction, then forming a frondose-bracteose inflorescence, or it can lack any significant reduction in shape and/or size, then termed a frondose inflorescence (Weberling 1989). In all the species of Diplusodon that possess simple inflorescences, they are of the frondose type. Among the species with synflorescences, they are frondose to frondose-bracteose in the majority of cases or bracteose, as in D. panniculatus, D. bradei, and D. ramosissimus. In addition to the polytelic type of inflorescence, described as the basic pattern for Diplusodon, two species were found with an unusual form: D. ovatus Pohl (Fig. 15.3F) and D. panniculatus. The inflorescences of these species show that, unlike all other species of the genus and contrary to the definition of the group as polytelic (Weberling 1988), the principal axis of the inflorescence terminates in a flower, i.e., it is monotelic (Fig. 3.8E, F). This also applies to all the paraclades. In these inflorescences, the differentiation, development, and anthesis of the terminal flowers (TF) always precede those of the paraclades, whereas paraclades of the same branching order develop basipetally. Such an inflorescence is called a thyrsoid (a thyrse-like inflorescence with terminal flowers, according to Troll 1969; Briggs and Johnson 1979; Weberling 1988). The distal paraclades (“short paraclades,” cf. Troll 1965; Weberling et al. 1993) of such inflorescences are cymose-dichasial, so that the whole structure is a thyrse-like inflorescence with a terminal flower, hence a “thyrsoid” (Troll 1969; Briggs and Johnson 1979; Weberling 1988, 1989). Such cymose

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short paraclades are initially triads but are able to produce further lateral flowers in the axils of the two prophylls of each new axis (Fig. 3.8D–F). It has been proposed elsewhere (Sell 1969, 1976; Weberling 1988) that the polytelic type of inflorescence has been derived more than once from the monotelic type during the evolution of the angiosperms. It has been hypothesized to have occurred by loss of the terminal flower and specialization of the paraclades from the monotelic system into solitary lateral flowers or lateral cymes that make up the elements of the florescence in the polytelic system. The monotelic synflorescence is a character that seems to have evolved at least four times within Lythraceae (Graham et al. 2005). Secondary monotelic synflorescences are reported, representing one or more additional transitions from polytely to monotely (Cavalcanti and Rua 2008; Inglis and Cavalcanti 2018), challenging the general view of polytelic inflorescences as derived from monotelic ones (Weberling 1985, 1989). Most Lythraceae genera, including Diplusodon, have polytelic synflorescences (Weberling 1988; Graham et al. 1993), which represents the plesiomorphic condition of the family (Graham et  al. 2005). Six genera, Duabanga Buch.-Ham., Sonneratia L.f., Punica L., Lawsonia L., Lagerstroemia, and Galpinia N.E.Br., have monotelic inflorescences, a condition that seems to have evolved at least four times within the family (Graham et al. 2005). Secondary monotelic synflorescences are here reported, which represent one or more additional transitions from polytely to monotely. Such a transition has been hypothesized to be the outcome of a rather complex evolutionary process, involving the specialization of distal paraclades (homogenization) together with the onset of acropetal flowering (racemization) and the loss of the terminal flower (truncation; Sell 1969, 1976). Thus, the regain of monotely has been implicitly considered by classical morphologists to be a very improbable evolutionary event. Nevertheless, current evidence from developmental genetics suggests the transition from monotely to polytely and vice versa is mediated by a simple regulatory genetic system (Kellogg 2000; Reinhardt and Kuhlemeier 2002; Angenent et al. 2005), so that a single mutation in the proper regulatory gene could suffice to explain the multiple reversions to monotely in Lythraceae (Cavalcanti and Rua 2008). In the partial florescences of Diplusodon bradei, the very short axes produce flowers continuously, resulting in dense glomerules. In D. bradei, D. lanceolatus Pohl, and D. ulei Koehne, for example, accessory branches (AcB), sensu Weberling (1988), develop in the axils of the bracts, in the same position where the paraclades originate (Fig. 3.8D). Accessory branches were considered by Weberling (1988) to be very common in Lythraceae and were identified in this study for the first time in Diplusodon. According to Weberling (1988), the accessory branches increase the number of flowers in the inflorescence and can be from the first to the “n” order. In D. bradei and D. lanceolatus, they are up to the third order and can form solitary flowers or dichasia whose triads continue to develop flowers in the axil of their prophylls, repeating the same pattern found in the paraclades. A higher degree of this branch formation can also be seen at the base of the synflorescence and can occur within the paraclades and even in the adjacent accessory branches.

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The types of inflorescences present in Diplusodon can be organized according to the terminology of Weberling (1988, 1989), to account for the change from the monotelic to the polytelic pattern, through the loss of the terminal flower, specialization of the paraclades, and the trend toward reduction of the number of flowers. Thus, the following inflorescence types have been observed in Diplusodon: 1  – Thyrsoid: monotelic synflorescence (with terminal flower) whose branches (long paraclades) bear short cymose branches (short paraclades) or triads (Fig. 3.8E, F) 2  – Thyrse: polytelic synflorescences (without terminal flower) whose branches (long paraclades) bear short cymose branches (short paraclades) or triads (Fig. 3.8D) 3 – Pleiobotrya, triplobotrya, or diplobotrya: polytelic synflorescences (without terminal flower) whose polytelic branches (florescences) bear solitary lateral flowers (Fig. 3.8B, C) 4 – Botrya: simple inflorescences (unbranched) with only lateral flowers (Fig. 3.8A)

3.5.2  Evolution and Diversification In the phylogeny presented to Diplusodon (Cavalcanti and Rua 2008;  Inglis and Cavalcanti 2018), some hypotheses can be offered regarding inflorescence evolution and diversification within Diplusodon. The plesiomorphic inflorescence pattern, i.e., frondose or frondose-bracteose compound racemes, seems to have been highly conserved. Alternative patterns, such as thyrses and thyrsoids, are restricted to a few species. Changes involving meristem-identity genes (Coen and Nugent 1994; Angenent et al. 2005) are probably responsible for several striking features in inflorescences of some Diplusodon species, such as the switch to monotely, the production of high-­ order lateral cymes, and the proliferation, i.e., the return to vegetative growth, of the apical meristem of the racemes. On the other hand, paedomorphic flowering seems to be the outcome of a change in flowering-time genes (Coen and Nugent 1994; Angenent et al. 2005). Another set of modifications involves variations in internode elongation and in relative development of subtending leaves. At the time a shoot apical meristem (SAM) is induced to flowering, some internodes, leaf primordia, and axillary meristems are already differentiated, whereas the meristematic apex remains undifferentiated. Both preformed and neoformed (after induction) internodes can either elongate or remain short. Differences in the distribution pattern of elongated and short internodes are largely responsible for striking variations in inflorescence appearance yet without deep structural modifications. The same is true for the hypopodium/epipodium elongation pattern. The plesiomorphic condition of elongated internodes has been repeatedly lost during inflorescence evolution of Diplusodon, and reduced (strictly, non-elongated) internodes are widespread within

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the genus. In the most radical cases, highly congested inflorescences arose (Cavalcanti and Rua 2008). Development of leaf primordia can be more or less altered after flowering induction, generally leading to a gradual reduction of inflorescence leaves: the more incipient the development of a leaf primordium at induction, the less developed the resulting mature leaf. It seems to be the case of most species of Diplusodon that a gradual reduction from foliage leaves to bracts can be observed along the synflorescence main axis. The occurrence of strictly bracteose synflorescences is apomorphic within Diplusodon. In such synflorescences, the transition between foliage leaves and bracts is abrupt. Whether it results from a sudden suppression of pre-­ existing leaf primordia or from the fact that the flowering region is entirely produced from previously undifferentiated meristematic tissues is yet to be explored (Cavalcanti and Rua 2008).

3.6  Bracts, Prophylls, and Peduncles In the frondose inflorescences of Diplusodon, bracts are found that are very similar morphologically to vegetative leaves, although they are sometimes smaller. The reduction in size is greater and gradual in species with frondose-bracteose inflorescences and abruptly evident in those with bracteose inflorescences whose bracts are sharply distinct from foliage leaves. The bracts can subtend both solitary flowers, in botrya, such as those of D. imbricatus Pohl, or in dichasial cymes, as in the thyrsoids of D. panniculatus (Fig. 3.8F). The prophylls in Diplusodon are always opposite, and in the majority of cases they are distinct in form from the foliage leaves and bracts. They are variable in form and indumentum (Fig. 3.9), showing the same diversity as foliage leaves and bracts. The relationship of prophyll length to floral tube length is used as a taxonomic character in Diplusodon and represented by the following variants: prophylls reaching the apex of the floral tube (Fig. 3.9C, G, I, L); prophylls surpassing the apex of the floral tube (Fig. 3.9A, E); prophylls reaching to the middle of the floral tube (Fig. 3.9B, J); and prophylls reaching the base or less than the base of the floral tube (Fig. 3.9F, H). The pedicel is regarded in this study as composed of the epipodium, the  axis between the flower and the last node of the axis where the flower terminates; the anthopodium (of Briggs and Johnson 1979) together with the hypopodium, the proximal axis that precedes the epipodium (the peduncle of Briggs and Johnson 1979). In Diplusodon, the peduncle is variable in size and can be articulated, that is, composed of hypopodium, mesopodium, and epipodium, or it may be composed only of the hypopodium. Two prophylls always occur on each peduncle. To evaluate the specializations of the inflorescence in Diplusodon, the tendency for reduction seen at the level of the peduncles needs to be analyzed further. In the dichasial lateral cymes, peduncles can be seen whose prophyllar buds are able to further produce two flowers. Departing from this pattern, and according to

Fig. 3.9  Flowers in Diplusodon. A. D. paraisoensis (Cavalcanti et al. 390, CEN). B. D. helianthemifolius var. helianthemifolius (Longhi-Wagner et al. [CFCR] 9325, SPF). C. D. glaucescens (Scatena et al. [CFCR] 10477, SPF). D. D. speciosus (Cavalcanti et al. 86, SPF). E. D. rotundifolius (Menezes et al. [CFCR] 10600, SPF). F. D. nitidus (Hatschbach & Nicolack 53031, MBM). G. D. sessiliflorus (Philcox & Onishi 4782, UB). H. D. panniculatus (Romaniuc & Sajo 388, CEN). I. D. minasensis (Cavalcanti et al. 220, SPF). J. D. imbricatus (Pereira-Silva et al. 10648, CEN). K. D. cordifolius (Irwin et al. 24807, UB). L. D. punctatus var. punctatus (Cavalcanti et al. 1015, CEN)

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Weberling (1988), the probable sequence of evolutionary events in Myrtales inflorescences involves a reduction from dichasium to cymes of triads, leading to a simple raceme, and finally to a spike, as found in the paraclades of Diplusodon lanceolatus. In this species the flowers are sessile, totally lacking a pedicel (lacking differentiated hypopodium and epipodium). This last pattern is further reduced in simple botrya, without branching in the inflorescence other than production of floral axes. In addition to the pattern of dichasial lateral cymes, the occurrence of two axillary peduncles initiated from the same node was also observed in Diplusodon ulei and D. ciliatus (T.B.Cavalc.) T.B.Cavalc. This suggests that a reduction in the inflorescence of Diplusodon from a more complex pattern to simple botrya and spikes has taken place.

Chapter 4

Flowers

In Diplusodon the flower buds are, in the majority of cases, obconic or, more rarely, oblong. The pedicel is short in the majority of the species, but it can be long and articulated, with a distinct hypopodium and epipodium, reaching 3 cm in D. marginatus. The pilosity of the buds and peduncles is extremely variable, with some buds totally glabrous and others densely covered with long trichomes. In bud, the sepals are valvate and curved toward the center of the flower. Mayr (1969) studied the ontogeny of the flowers in some genera of Lythraceae (Rotala L., Decodon J. F. Gmel., Lythrum L., and Lagerstroemia) and other families of Myrtales. This author considers the bending of the sepals a special formation in Lythraceae and Punicaceae, which provides special protection of the bud (Fig.  4.1E). In the later stages of flower development, the petals surpass the sepals in length, taking over the function of bud protection. In Diplusodon, the flowers are dichlamydeous, hexamerous, perigynous, and actinomorphic, although the style is curved and inclines toward one side of the flower. There are nectaries or nectariferous regions on flowers of Diplusodon similar to those of Lafoensia, Koehneria, and other genera of Lythraceae, but the sepals of all species of Diplusodon bear secretory papillae on the internal margin. The secreted substance can be seen in buds just before anthesis as a drop of mucilage or resin in the slits between the still-closed sepals. The hardened drops can be seen on herbarium specimens. The flowers of Diplusodon are characterized by a floral tube that is present in all genera of Lythraceae. The floral tube is funnel-shaped in most species (Fig. 3.9). In some species, e.g., D. oblongus and D. imbricatus (Fig. 3.9J), the floral tube is oblong. The indumentum of the tube can be totally absent, sparse, or varying to very dense. The presence of trichomes on the floral tube sometimes is unrelated to the presence of trichomes on the rest of the plant. In D. puberulus, where the entire plant bears an indumentum, including the calyx and epicalyx, only the floral tube is glabrous (Fig. 13.86F–G). Normally the floral tube bears the same type of trichomes

© The Author(s), under exclusive license to Springer Nature Switzerland AG 2021 T. B. Cavalcanti, The Genus Diplusodon (Lythraceae), Flora Neotropica, https://doi.org/10.1007/978-3-030-65875-5_4

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Fig. 4.1  Flower anatomy in Diplusodon orbicularis. A. Longitudinal section of hypanthium showing region where filaments are free and ovary with basal placentation. B. Transverse section of unilocular ovary with incomplete septa. C. Longitudinal section of anatropous ovule. D. Transverse section of floral tube, showing vascular bundle divided to petals, sepals and epicalyx segments. E. Longitudinal section of perianth showing region where filaments are free and locules (A–E. Oliveira 1991 [modified]). A–B, D–E scale = 0.2 mm. C scale = 0.05 mm

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that occur on other parts of the plant. An exception is seen in D. divaricatus, which has botuliform secretory trichomes on the entire floral tube, while the branches and leaves are clothed in a simple indumentum. Based on ontogenetic studies of Lythraceae flowers, Mayr (1969) considered the floral tube of Lythraceae to consist of three parts: the hypanthium; the perianth tube; and the calyx + epicalyx + corolla at the apex of the tube. In some genera, such as Galpinia, and Physocalymma Pohl, the epicalyx whorl is lacking. Diplusodon flowers are composed of the three parts and generally develop an epicalyx, although two taxa, D. marginatus (Fig. 13.59E–F) and D. punctatus var. punctatus (Figs. 3.9L and 13.87E–F), lack this feature. In Diplusodon, the hypanthium makes up one-­ third of the length of the floral tube and is positioned in the most basal part of the tube, beginning at the apex of the epipodium and terminating a few centimeters above in the region where the filaments separate from the floral tube (Figs. 4.1A, E and 13.44G). The formation series of primordia in Lythraceae is represented by an acropetal succession of the following: calyx-epicalyx-corolla (congentially fused at the base)-stamens-ovary-petals. The perianth begins where the filaments separate from the floral tube (Fig. 4.1E). Mayr (1969) referred to this segment as the perianthal tube. For Mayr, this designation should not be used for the tubular portion between the base of the ovary and the level of stamen insertion because the stamens are not part of the perianth. Although this author did not provide a term for this region, in my study I refer to this part of the floral tube as the hypanthium. The use of hypanthium is questionable due to the fact that some traditional definitions apply the term only to inferior-ovaried flowers. According to Font Quer (1965), for example, the hypanthium is a coalescence of the sepal, petal, and stamen whorls in flowers with an inferior ovary. According to Fahn (1989), flowers with a superior ovary can also possess a hypanthium, as is the case in the perigynous flowers of Diplusodon. In Diplusodon the hypanthium is present where the calyx, corolla, and androecium whorls are united from the base of the tube; these become free at different levels (Fig. 4.1E). In this study, flower size is a significant taxonomic character. The floral measurements are from the base of the ovary to the level where the elements of the calyx, corolla, and epicalyx separate, and this region, including the perianth tube as well as the hypanthium, is referred to as the floral tube. The calyx is composed of six equal-sized sepals. They are always triangular in shape and vary from very wide to very narrow. The sepals can vary from species to species, and the variation includes size, thickness, and indumentum, which are all characters of good taxonomic value. In the majority of Diplusodon species and also in the flowers of other genera of Lythraceae, there are epicalyx appendages situated slightly below the sepal whorl, alternating with them, and of the same number (Fig. 4.2A–F). Until the discovery of the epicalyx origin, different terms were used to characterize it, and different functions were attributed to it. Koehne (1903) called them calyx appendages and considered them to be fused stipules, analogous to the external calyx of Potentilla L. Cadet (1954, apud Cheung and Sattler 1967), based on anatomical studies, considered the external calyx, i.e., the epicalyx, to be homologous to a whorl of

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Fig. 4.2 Floral diagrams in Diplusodon showing different conditions of the androecium. A. Isostemonous androecium. B. Diplostemonous androecium. C–F. Polystemonous androecium

4.1 Androecium

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hypsophylls. Mayr (1969) followed the formation of this structure in the flower of Decodon verticillatus (L.) Ell. and observed that periclinal divisions occur below the dermatogen in the areas between the calyx primordia. When the growth of the calyx is complete, the regions between the sepals begin dividing intensely and form solid, adaxially furrowed, cylindrical points and a vascular bundle system, leading to the formation of an external calyx or epicalyx. These  observations were confirmed by Oliveira (1991) in Diplusodon orbicularis, in which the division of a vascular bundle leads to the epicalyx, calyx, and corolla. The shape of the epicalyx segments can vary from short, small protuberances between the sepals, to very long and surpassing the length of the sepals. The position of these segments in relation to the axis of the flower also is variable, and they can be deflexed, spreading (perpendicular to the axis of the flower), or suberect to erect (Fig. 3.9). The indumentum on the segments varies, just as with the sepals. The position of the epicalyx segments, as well as their length in relation to the sepals, is recognized as important taxonomically in the separation of the species of Diplusodon along with other important floral characters. Mayr (1969) considered the epicalyx segments to result from simple additional growth from the areas of fusion between the sepals. The epicalyx would then be interpreted as an emergence from within the region of the fusion of the sepals (Mayr 1969). A characteristic not yet reported for the family is the presence of prominent cordiform, callose glands that are probably associated with pollination. Similar structures occur in Cuphea. The glands occur on the internal margin of the floral tube at the base of the petals and were detected in Diplusodon astictus  (Fig. 13.7N), D. appendiculosus (Fig. 13.4G), and D. leucocalycinus. The corolla is generally showy, 2–7  cm in diameter, and is composed of six obovate petals that vary in color from lilac to red, passing through various shades of pink, which is the predominant color in most species. Only one species, Diplusodon virgatus, has white petals. The petals are crinkled and compressed within the bud, curving toward the center of the flower. According to Mayr (1969), the formation of Lythraceae petals begins on the upper margin of the inner surface of the floral tube, the region in which the sepals also appear, but the development of the petal primordia appears after the other floral parts as observed in D. orbicularis (Fig. 4.1D).

4.1  Androecium The androecium in Diplusodon can be isostemonous, diplostemonous, or polystemonous, always with centrifugal development. The number of stamens varies among species and is not always fixed in a species. Nevertheless, it is an important taxonomic character in the distinction of species. The diplostemonous condition (12 stamens; Figs. 3.9B and 4.2B) is found in 53 species (51.9%); 6 species (5.8%) are isostemonous (Figs. 3.9I and 4.2A); and the rest (49 species, 48%) are polystemonous (Figs. 3.9D and 4.2C–F).

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The suppression of the epipetalous stamen whorl is exclusively found in Diplusodon candollei, D. glaziovii, D. hexander, D. minasensis, D. orbicularis, and D. verruculosus, all of which are microendemics of campos rupestres and phylogenetically related (Inglis and Cavalcanti 2018). Reduction of the two ancestral whorls of stamens is common in Rosidae families (Chase et al. 1993), and it is a condition that is never reversed and probably related to adaptations that increase pollen dispersal efficiency (Walker-Larsen and Harder 2000). Loss of stamens is also related to the development and evolution of zygomorphy (Richards 1986) and pollination specialization (Walker-Larsen and Harder 2000). The change from the basic diplostemonous condition together with corolla size reduction and enantiostyly present in all species (the style turning to the right or to the left with the stamens turning in the opposite direction) may be a significant evolutionary event in Diplusodon related to pollination specialization (Inglis and Cavalcanti 2018). Multiplication of stamens in Diplusodon occurs exclusively in the episepalous whorls (Dahlgren and Thorne 1984). In Diplusodon the basic number of stamens is 12, and from this state, there has been loss of 1 whorl in the species with 6 stamens and multiplication in the species with 13–54 stamens. The production of abundant pollen through a secondary increase of stamen number is often found in Rosidae and is thought to be due to the division of stamen primordia, rather than by re-­ initiation of suppressed stamen primordia (Walker-Larsen and Harder 2000). Stamen multiplication allows the production of more pollen and the attraction of larger pollinators (Craene 2010). The stamens can reach the apex of the sepals to be slightly exserted, or they can surpass the sepals by up to 3 cm, depending on the species. Within flowers, the stamens are all equal in length, unlike the diplostemonous condition in Cuphea P. Browne, for example, in which different whorls are of different lengths. No species were found to have included stamens, i.e., below the sepals. The filaments are cylindrical, cream-colored, lilac, or rose-magenta, and they often assume a wine-­ colored tone near the apex or sometimes over the entire length of the filament. The anthers are dorsifixed, versatile, bilocular, extrose, and with longitudinal dehiscence.

4.2  Gynoecium The pistil in Lythraceae consists two to six carpels (Graham and Graham 2014), and Diplusodon has two (Fig. 4.1B). The ovary is superior, globose, rhombic to obtrapezoidal, with apex depressed, glabrous, rarely with trichomes at the apex. Mayr (1969) stated that in all the species of Lythraceae she studied (not including Diplusodon), the primordium of the gynoecium is always formed by a circular disc from which the carpels arise. Perpendicular to the wall of the ovary and connected to it in the middle region, there are two fleshy structures that reach the apex of the carpels (Fig. 4.1B). These structures are the septa of the ovary and seen in longitudinal section appear to be complete, dividing the ovary into two locules (Fig. 4.1A, B, E). An analysis of these structures in cross-section makes clear that the septa are

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not united at maturity (Fig.  4.1B). For this reason, the literature offers different interpretations of the Diplusodon ovary. Koehne (1903) referred to the ovary as unilocular with placentas divided by two semi-lunate septa. Lourteig (1989) called the ovary falsely bilocular with “tabiques.” Graham and Graham (2014) refers to Diplusodon ovary as two-locular, appearing unilocular. The placenta has a basal and central position in the ovary and is bipartite separating the two locules by means of the septa. The septa are semi-lunate, thickened at the ovary wall, thinning from ovary wall toward the placenta, narrowing and ultimately ceasing above the placenta (Graham and Graham 2014). The placenta has the same number of ovules on each of the two parts. The ovules are anatropous (Fig. 4.1C) and vary in number from six to ca. 60. In a study of the reproductive cycle of Diplusodon orbicularis, Oliveira (1991) described the ovules as anatropous, crassinucellate, and bitegmic with the micropyle formed by both integuments. She reported that the megasporangium arises as a protuberance that projects itself vertically in the basal part of the ovary, characterizing this primordium as initially orthotropous.

Chapter 5

Fruit, Seeds, Dispersal, and Germination

5.1  Fruit In Diplusodon, the fruits are septicidal capsules. The capsules are included or exceeding the persistent floral tube at maturity. The capsules are glabrous with the exception of D. appendiculosus (Fig. 13.4M), and D. leucocalycinus (Fig. 13.56L), which bear trichomes on the apex. In the fruit, the two incomplete septa of the ovary are transformed into two thick semi-lunate septa (Fig. 13.44J, K). The fruits display two general shapes. The most frequent is the fruit whose floral tube assumes the shape of an urn with a wide mouth and a rounded base (Fig. 13.3G, H), and the capsule is globose. In the second shape, fruits also have an urn-­ shaped floral tube, but the opening is much narrower, resulting in the strong constriction of the neck (i.e., Figure 13.71J), and the capsule is usually oblong. Sometimes this type also has a narrow, prolonged base. The floral tube develops together with the fruit, and during maturation it becomes dry, rigid, brown, and sometimes thick (Fig. 13.3G, H); it is persistent on the mature fruit. Many times the floral tube is still present on old fruits from a previous season. Old floral tubes are grayish in color and with some longitudinal fissures developing as they age. The persistent fruit of Diplusodon does not have a dispersal mechanism; dispersal is accomplished by the winged seeds. Oliveira (1991) observed that after fertilization, the cells of the ovary walls, like the structure of the floral tube, are altered. These cells undergo thickening of the cell wall, reducing greatly the cellular lumen, and thickening the cuticle. The cells of the mesophyll become compressed by the loss of water as the ovary matures.

© The Author(s), under exclusive license to Springer Nature Switzerland AG 2021 T. B. Cavalcanti, The Genus Diplusodon (Lythraceae), Flora Neotropica, https://doi.org/10.1007/978-3-030-65875-5_5

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5  Fruit, Seeds, Dispersal, and Germination

5.2  Seeds The seeds provide important diagnostic characteristics for the recognition of genera within Lythraceae. Graham and Graham (2014) reported that seed morphology provides one of the most trustworthy sources of characters in the family. In Diplusodon, the seeds do not provide good characters to distinguish species, because they are homogeneous in morphology and differ only in dimensions. The seeds are round, oblong, obovate to obtriangular (Fig. 5.1A), bilaterally compressed or slightly lenticular, and invariably winged, with the wing of 0.1–0.6 mm surrounding the seed. The hilum is elliptic to round, the embryo is straight, and the cotyledons are reniform to orbicular. Oliveira (1991) studied the seed development of Diplusodon orbicularis and described the testa of this species, as well as that of the genus Pemphis, as multiplicative, which differs from the multiplicative testa (increasing in the number of the layers in the external seed coat during development) of other genera in Lythraceae, such as Lafoensia, Lagerstroemia, and Lawsonia (Corner 1976). In addition, according to Oliveira (1991), the seeds of D. orbicularis can be classified as either endotestal, for possessing the mechanical layers in the internal layer of the testa, or as exotegmic, due to a fibrous external layer of the tegmen. Anatomically, the seeds of Diplusodon have the same type of epidermal cells as seeds of Cuphea calophylla Cham. & Schltdl. ssp. mesostemon (Koehne) Lourteig (=Cuphea calophylla Cham. & Schltdl.  var. mesostemon (Koehne) S.Graham) (Schoemberg and Hofmeister 1986; Amarasinghe et al. 1991; Schoemberg 1989) except they are straight. When the seeds come in contact with water, mucilage in the epidermal cells of the testa swells, and the cells separate from the inner spongy layers, falling away as individual globose packets of clear mucilage (Graham and Graham 2014) in the form of filaments (Oliveira 1991). These filaments originate from early in seed development, when the cellular content becomes dense and forms a filament (Oliveira

Fig. 5.1  Seed and chromosomes in Diplusodon. A. Seed of D. ovatus (Nakajima et al. 876, CEN). B. Chromosomes of D. sordidus: 2n = 30 (Cavalcanti et al. 403, CEN)

5.3  Dispersal and Germination

47

1991). When the seed is exposed to water, the external wall of the epidermal cell opens like a cap and becomes turgid pushing the filament out of the cell. Stubbs and Slabas (1982) verified that the epidermal filaments of Cuphea procumbens Ortega are composed of 55% carbohydrates and 45% protein, and other species of Lythraceae (Ammannia latifolia L., Rotala verticillaris L., Nesaea triflora Kunth, and Hionanthera garciae Fernandes & Diniz) have been shown to have the same content (Panigrahi and Sarojini 1986). The chemical content in the seed filaments of Diplusodon has not been investigated. The secretion of mucilage to aid in adherence of diaspores for dispersal is known as mixospermy. According to Boesewinkel and Bouman (1984), mixospermy is common and is an important phenomenon for diaspore adherence to animals or fixation to the ground. Van der Pijl (1972) considered mixospermy as an important process in diaspore transport of plants with hydrochoric dispersal. For example, plants such as Podostemaceae that live in running water develop mucilage in their diaspores to make possible their fixation to rocks. In a similar way, Van der Pijl (1972) suggested that myxospermia is common in drier climates and cited the aspect of diaspore fixation through mixospermy in desert plants, which after a rainfall produce mucilage that temporarily fixes the diaspore to the humid ground enabling penetration of the radicle and promoting germination. The dry season in the Cerrado Biome lasts about 6 months, and most of the Diplusodon species flower and fruit during the dry season, so mixospermy may be important strategy for the genus.

5.3  Dispersal and Germination In Diplusodon, the unit of dispersal is the seed. Initially the seeds leave the capsules and are carried by the wind, which is a common feature in campos rupestres and open areas in the Cerrado. Because of their small size and the presence of small wings, the seeds can disperse short distances before they fall to the ground. With the first rains, or increased humidity caused by the dew, the mucilaginous trichomes uncoil and protrude from the seeds (Fig. 5.2), resulting in a mass around the seeds, and germination is then initiated. Silveira et  al. (2012) reported that seeds of Diplusodon hirsutus (Cham. & Schltdl.) DC. and D. orbicularis may form buried seed banks in the soil. Cesarino et  al. (1998) reported viable seeds of D. virgatus after 12  months, and they also showed that favorable growth and germination took place under long photoperiods. In the germination experiments carried out in this study, Diplusodon seeds were collected from mature fruits, when the capsules were half-open or completely open. The seeds were collected from natural populations in the states of Goiás and Minas Gerais and the Distrito Federal, then conditioned in paper bags, and stored for about 30 days. After this period, germination was initiated in glass boxes (Gearbox) using a substrate of cotton soaked in water. Fifteen seeds were placed in each box, depending on the size of the seeds. The seeds were not sterilized or scraped. The dates and

48

5  Fruit, Seeds, Dispersal, and Germination

Fig. 5.2  Phases of germination in Diplusodon virgatus (Cavalcanti et al. 571, CEN). A. Seed. B. Seed after soaking, with mucilaginous trichomes uncoiling from seed. C–D. Beginning of radicle protusion. E–H. Appearance of reniform cotyledons. (A–C, E–H frontal view. D lateral view)

5.3  Dispersal and Germination

49

time of starting were recorded, and daily observations were made between 8 and 9 a.m. All steps in the germination process were recorded (Table 5.1). Germination was considered to have occurred when the radicle protruded. All experiments were carried out under temperatures of 22–27 °C, under fluorescent light and natural light. The stages of germination are summarized in Table 5.1, and the beginning was defined as when the seed trichomes started to uncoil. It was observed that the seeds are positively photoblastic. Diplusodon alatus, D. orbicularis, D. oblongus, D. paraisoensis, D. ramosissimus, D. villosus, and D. virgatus var. virgatus showed no seed dormancy. Germination started within 5–8 days, and the process of soaking, which involves the development of the mucilaginous trichomes, was initiated for all the species from the first day after moistening of seeds. The phases involved in seed germination are illustrated for D. virgatus var. virgatus (Fig. 5.2). The highest percentages of germination for Diplusodon orbicularis (64%) were obtained at 20 °C, in light. At 35 °C, in light, the percentage of germination was of 2% (Negreiros et al. 2002). Experiments to test the regeneration niche hypothesis were carried out by Silveira et  al. (2012) comparing the germination ecology of Diplusodon orbicularis, a Brazilian endemic species, and D. hirsutus, a more widespread species, both growing in different habitats. Their results showed that there was no temperature-­ dependent microhabitat selection and that the germinability of D. hirsutus was lower than D. orbicularis. Therefore, the species studied did not provide support for the hypothesis that germinability may limit the distribution of plant species.

Table 5.1  Phases of seed germination of Diplusodon species

Species/voucher (number of seeds) D. alatus/Cavalcanti et al. 681 (18)

D. orbicularis/no voucher (16) D. paraisoensis/ Cavalcanti et al. 648 (16)

Number of days (d) to the beginning of the seed trichome uncoiling (number of seeds) 19 d (0) 20 d (2)

19 d (3)

19 d (0) 20 d (1)

Number of days (d) to the beginning of germination (number of seeds) 22 d (1) 23 d (3) 24 d (4) 25 d (4) 26 d (5) 27 d (5) 30 d (5) 26 d (2) 27 d (3) 30 d (3) 24 d (2) 25 d (2) 26 d (3) 27 d (3) 30 d (3)

Number of days (d) to the appearance of cotyledons (number of seeds) 24 d (3)

28 d (2) 29 d (3) 25 d (2) 26 d (2)

50

Species/voucher (number of seeds) D. praetermissus/ Cavalcanti et al. 660 (16)

5  Fruit, Seeds, Dispersal, and Germination Number of days (d) to the beginning of the seed trichome uncoiling (number of seeds) 19 d (1)

D. ramosissimus/ Cavalcanti et al. 586 (20)

19 d (0) 20 d (1)

D. villosus/ Cavalcanti et al. 500 (21) D. virgatus var. virgatus/Vieira et al. 469 (23)

19 d (4)

D. virgatus var. virgatus/Cavalcanti et al. 571 (17)

19 d (9)

19 d (0) 20 d (1)

Number of days (d) to the beginning of germination (number of seeds) 24 d (0) 25 d (1) 26 d (2) 27 d (2) 30 d (3) 23 d (2) 24 d (5) 25 d (5) 26 d (6) 27 d (6) 30 d (7) 22–30 d (0)

22 d (1) 23 d (6) 24 d (10) 25 d (16) 26 d (17) 27 d (17) 30 d (17) 22 d (1) 23 d (7) 24 d (9) 25 d (9) 26 d (12) 27 d (12) 30 d (12)

Number of days (d) to the appearance of cotyledons (number of seeds) 27 d (2) 28 d (2) 30 d (2)

25 d (2) 26 d (3) 27 d (5)

–

23 d (1) 24 d (6)

23 d (1) 24 d (6)

Chapter 6

Pollen

Pollen morphology is of great importance at the generic level in Lythraceae and has been widely explored (e.g., Graham and Graham 1971; Muller 1981; Patel et  al. 1984). Laboriau and Válio (1964) studied 11 Lythraceae species from the cerrado, including the genera Cuphea, Diplusodon, and Lafoensia, and highlighted pollen diversity, referring to the family as eurypalynous. Graham et al. (1985) described pollen of ten genera of Lythraceae using light microscopy and scanning and transmission electron microscopy. Their study included 12 species of Diplusodon, and the authors described the pollen of the genus as follows: pollen prolate to prolate-spheroidal, tricolporate, the colpus inconspicuous, elongated meridionally, arranged equatorially, equidistant, straight, ca. 35 μm long, relatively inconspicuous when compared to the pore which is prominent, and frequently obscured from sight by the thick, undulating sculptural elements; pore elongate, 5 × 10 μm in diameter, situated at the midpoint of the colpus, the inner margin entire, the annulus conspicuous, the outer margin lobate; wall thick (4–5 μm), verrucate, margins of the verrucae irregular and lobate, polar region with larger verrucae in some species. Data obtained in this study  generally corroborated the description of Graham et al. (1985). Under the scanning electron microscope (SEM), pollen of Diplusodon is distinguished by the absence of pseudocolpi, poorly developed or apparently absent colpi, and a deeply verrucate or rugulate exine. In D. glaucescens and D. virgatus var. virgatus, the pollen is described as more psilate near the poles (Fig. 6.1A, B, D). Transmission electron microscopy reveals a thick wall with prominent endexine in the aperture and non-aperture regions, in addition to variable columellae (Graham et al. 1985). Of the three species described by Laboriau and Válio (1964), one of them, Diplusodon virgatus var. virgatus, was also investigated palynologically by Muller (1981) and Graham et al. (1985) and for the present study. In all three studies, the pollen grains are described as prolate in form but with differences in the amount of

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Fig. 6.1  Pollen grains in Diplusodon. A–B. Prolate to subprolate, D. virgatus var. virgatus (Rawitscher s.n., SPF). C. Semi-spherical with prominent pores, D. leucocalycinus (Irwin et al. 24231, UB). D. Prolate to subprolate, D. orbicularis (Cavalcanti et  al. [CFSC] 9619, SPF). E. Prolate to subprolate, pseudocolpus absent prominent pores, D. marginatus (Pereira 1253, IBGE). F. Semi-spherical, D. parvifolius (Cavalcanti et al. 303, SPF). G. Prolate to subprolate with pseudocolpus evident, D. sordidus (Cavalcanti et al. 403, CEN). H. Area of poles, D. hirsutus (Cavalcanti et al. [CFSC] 10099, SPF). Images made at Palynology Laboratory, Royal Botanic Gardens, Kew, London, England

6 Pollen

53

sculpturing at the poles and equator. Similar variation was observed here  for D. orbicularis. Muller’s work on Lythraceae pollen was an extension of his studies with the Sonneratiaceae. He investigated the similarity between some pollen types of Lagerstroemia and an ancestral extinct type from the Sonneratiaceae. The author cited Erdtman (1960) and van Campo (1966), who had already suggested that the pollen of Lafoensia and Diplusodon from South America showed similar structure with that of Sonneratia. The pollen grains of Diplusodon with a verrucate, homogeneous wall and almost spherical shape were considered by Muller (1981) to be the most advanced among the patterns observed in Sonneratiaceae and Lythraceae. Homogeneous walls and semi-spherical shape were observed in this study in D. leucocalycinus (Fig. 6.1C). Patterns described by Graham et al. (1985) and Muller (1981) were seen in this study and are illustrated in Fig.  6.1. The prolate to subprolate form is seen in Diplusodon virgatus var. virgatus, D. marginatus, and D. sordidus (Fig. 6.1A, B, E, G), and the semi-spherical form is seen in D. leucocalycinus (Fig. 6.1C) and D. parvifolius (Fig. 6.1F). The pseudocolpus was observed in some species but not others; for example, it was very evident in D. sordidus (Fig. 6.1G), but apparently absent in D. marginatus (Fig. 6.1E). There are generally three very prominent pores (Fig. 6.1C, E). The sculpture pattern is verrucate, composed of homogeneous-sized undulations on all surfaces, especially evident in D. leucocalycinus (Fig. 6.1C) and D. marginatus (Fig. 6.1E), or less textured in the equatorial zone and more so at the poles as in D. virgatus var. virgatus (Fig. 6.1A, B) and D. orbicularis (Fig. 6.1D). From the species studied, it is clear that pollen morphology provides a potentially valuable source of taxonomic characters and should be more thoroughly explored as a possible means of distinguishing the infrageneric categories of Diplusodon.

Chapter 7

Floral/Reproductive Biology and Phenology

Among the research related to the floral biology of Diplusodon are those of Barros (1989, 1996) who studied five species from the state of Goiás and Distrito Federal, and Faria (1994) who studied the honey flora and fauna, which included D. orbicularis and D. smithii (= D. hirsutus), from an area of campo rupestre in the Serra do Cipó, a mountain chain in southeastern Brazil (Minas Gerais State). Barros (1989, 1996) reported that in the five species studied, Diplusodon crulsianus (=D. sessiliflorus), D. oblongus, D. ramosissimus, D. rosmarinifolius, and D. villosus, generally only a single flower opens at a time in the inflorescence; when two flowers open simultaneously, they occur on opposite sides in the same inflorescence. Although this is true of some species, field observations reveal that many other species, for example, D. villosus (observed in the state of Goiás), produce numerous open flowers simultaneously in an inflorescence. Barros (1989, 1996) further described the flowers of Diplusodon as lacking odor and without nectar, in addition to being xenogamic and enantiostylic (i.e., the style turning to the right or to the left, with the stamens turning in the opposite direction). Barros (1989, 1996) also reported that anthesis occurs during the day (6 AM to 4 PM), with the flowers remaining open for 1 day, and recognized three phases in anthesis. In the bud stage, the sepals begin to separate from one another, and the style emerges in an arc-­ shaped path. In the second phase, the first petal behind the style begins to open, and in the third phase, the other petals open until the last one, which covers a group of stamens, opens with the stamens and style facing in opposite directions. The process of petal dehiscence takes 4 days from anthesis. In the phenological studies of Barros (1989, 1996), four of the species studied were in flower for 3 months, and one species, Diplusodon oblongus, was in flower for 4 months. During 2 years of study, the author noted the presence of two flowering periods for different species: December to April, the wet season, when D. oblongus and D. villosus flowered, and May to August, the dry season, when D. crulsianus Koehne (=D. sessiliflorus) and D. rosmarinifolius flowered. In both seasons, the species flowered synchronously. According to Barros (1996), synchronous

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7  Floral/Reproductive Biology and Phenology

flowering suggests a floral mutualism in which all species benefit equally through attraction of a larger number of pollinators. Synchronous flowering during two different periods was considered a primary barrier to interbreeding among species of Diplusodon. In addition, Barros (1996) presented five results: (1) All five species are capable of self-pollination, and autogamy occurs in low proportion (5–17%); (2) spontaneous self-pollination is natural in all five species (3–12%); (3) natural pollination is always very high (44–71%), being higher than artificial outcrossing; (4) interspecific hybrids are produced only between species with synchronous flowering; and (5) apomixis does not occur. With respect to pollinators, Barros (1996) recorded 16 species of small- and medium-sized bees that were attracted to Diplusodon flowers. Among them, the three most frequent, Apis mellifera, Trigona spinipes, and Augochloropsis spp., were considered the most efficient pollinators. Other frequent visitors were Augochloropsis spp., Paratetrapedia xanthopoda, Tetrapedia rugulosa, and Thectochlora alaris. According to Barros, in the enantiostylic Diplusodon a bee visiting a flower with the style turned to the left receives the pollen underneath the right side of its body. The  artificial pollination experiments of Barros (1989, 1996)  indicated that Diplusodon oblongus, D. ramosissimus, and D. rosmarinifolius are self-­incompatible and that D. crulsianus (=D. sessiliflorus) and D. villosus are partially self-incompatible. Other studies examined the reproductive biology of Diplusodon species in different regions. For example, Carmo et al. (1998a) report Apis mellifera, Bombus atratus (Fervidobombus), Halictini, Megachile spp. (Acentron), Xylocopa nogueira (Stenoxylocopa), and Xylocopa sp. as the visitors of D. orbicularis in the Serra do Cipó. Carmo et al. (1998a, b) and Jacobi et al. (2000) studied the reproductive biology of D. hirsutus and D. orbicularis in the Serra do Cipó, and they observed that both species are self-compatible, but because of herkogamy, they are greatly dependent on visitors for effective pollination. These studies reported high percentages of abscised fruits in both species and attributed this to resource limitation. In D. hirsutus, drought during seed development caused fruit abscission, and in D. orbicularis, nutrient drain by shoot-galling flies caused fruit abscission (Carmo et al. 1998a, b; Jacobi et al. 2000). Faria (1994) studied the use of floral resources by Apis mellifera in the Serra do Cipó and observed that this bee species used Diplusodon orbicularis exclusively as a source of pollen, collecting it on the front legs and touching the stigma with different parts of the body, especially in the ventral region of the thorax and abdomen. Faria (1994) classified D. orbicularis and D. smithii (=D. hirsutus), in relation to floral resources, primarily as honey producers. The phenology of all Diplusodon species, obtained from herbarium material and field observations, is summarized in Table 7.1. The flowering peak for most of the species is in the rainy season, and synchronous flowering was observed. However, flowering in the dry season was confirmed for a small percentage of species.

7  Floral/Reproductive Biology and Phenology

57

Table 7.1  Estimated phenology of 108 taxa of the genus Diplusodon Pohl (Lythraceae) based on herbarium specimen and field observations. The first row for each taxon = flowering; the second row = fruiting Nº Species J 1.

D. adpressipilus Lourteig

2.

D. aggregatifolius T.B.Cavalc.

3.

D. alatus T.B.Cavalc.

4.

D. appendiculosus Lourteig

5.

D. argenteus Lourteig

6.

D. argyrophyllus T.B.Cavalc.

7.

D. astictus Lourteig

8.

D. bahiensis T.B.Cavalc.

9.

D. bolivianus T.B.Cavalc. & S.A.Graham

10. D. bradei Pilg. 11. D. burchellii Koehne 12. D. buxifolius (Cham. & Schltdl.) DC. 13. D. caesariatus Lourteig 14. D. canastrensis T.B.Cavalc. 15. D. candollei Pohl ex DC. 16. D. capitalensis T.B.Cavalc. 17. D. capitatus (A.St.-Hil.) Koehne 18. D. chapadensis T.B.Cavalc. 19. D. ciliatiflorus T.B.Cavalc.

Months of the year F M A M J J A S

O N D

58

Nº Species 20. D. ciliatus (T.B.Cavalc.) T.B.Cavalc. 21. D. ciliiflorus Koehne 22. D. conduplicatus T.B.Cavalc. 23. D. cordifolius Lourteig 24. D. cryptanthus T.B.Cavalc. 25. D. decussatus Pohl 26. D. divaricatus Pohl 27. D. epilobioides Mart. ex DC. 28. D. ericoides Lourteig 29. D. fastigiatus Lourteig 30. D. floribundus Pohl 31. D. glaucescens DC. 32. D. glaziovii Koehne 33. D. glocimarii T.B.Cavalc. 34. D. gracilis Koehne 35. D. grahamae T.B.Cavalc. 36. D. hatschbachii Lourteig 37. D. helianthemifolius Mart. ex DC. var. helianthemifolius 38. D. helianthemifolius Mart. ex DC. var. pemphoides (DC.) Koehne 39. D. heringeri Lourteig

7  Floral/Reproductive Biology and Phenology

J

Months of the year F M A M J J A S

O N D

7  Floral/Reproductive Biology and Phenology

Nº Species 40. D. hexander Mart. ex DC. 41. D. hirsutus (Cham. & Schltdl.) DC. 42. D. imbricatus Pohl 43. D. incanus Gardner 44. D. kielmeyeroides A.St.-Hil. var. kielmeyeroides 45. D. kielmeyeroides A.St.-Hil. var. macrocalyx T.B.Cavalc. 46. D. lanceolatus Pohl 47. D. leucocalycinus Lourteig 48. D. longipes Koehne 49. D. macrodon Koehne 50. D. marginatus Pohl 51. D. mattogrossensis T.B.Cavalc. 52. D. micromerus T.B.Cavalc. 53. D. microphyllus Pohl 54. D. minasensis Lourteig 55. D. mononeuros Pilg. 56. D. myrsinites Mart. ex DC. 57. D. nigricans Koehne 58. D. nitidus Mart. ex DC. 59. D. oblongus Pohl

59

J

Months of the year F M A M J J A S

O N D

60

Nº Species 60. D. obtectus T.B.Cavalc. 61. D. orbicularis Koehne 62. D. ovatus Pohl 63. D. panniculatus Koehne 64. D. paraisoensis Lourteig 65. D. parvifolius Mart. ex DC. 66. D. petiolatus (Koehne) T.B.Cavalc. 67. D. plumbeus T.B.Cavalc. 68. D. praetermissus T.B.Cavalc. 69. D. prostratus T.B.Cavalc. 70. D. psammophilus T.B.Cavalc. 71. D. puberulus Koehne 72. D. punctatus Pohl var. dentatus T.B.Cavalc. 73. D. punctatus Pohl var. punctatus 74. D. pygmaeus T.B.Cavalc. 75. D. quintuplinervius (Nees) Koehne 76. D. ramosissimus Pohl 77. D. rectinervis T.B.Cavalc. 78. D. retroimbricatus Koehne 79. D. rosmarinifolius A.St.-Hil.

7  Floral/Reproductive Biology and Phenology

J

Months of the year F M A M J J A S

O N D

7  Floral/Reproductive Biology and Phenology

Nº Species 80. D. rotatus T.B.Cavalc. 81. D. rotundifolius Mart. ex DC. 82. D. saxatilis Lourteig 83. D. sessiliflorus Koehne 84. D. sigillatus Lourteig 85. D. sordidus Koehne 86. D. speciosus (Kunth) DC. 87. D. stellatus T.B.Cavalc. 88. D. strigosus Pohl 89. D. subsericeus Casar. ex Koehne 90. D. tenuifolius T.B.Cavalc. 91. D. thymifolius Mart. ex. DC. 92. D. thysanosepalus Lourteig & Sandwith 93. D. trigintus T.B.Cavalc. 94. D. ulei Koehne 95. D. uninervius Koehne 96. D. vacillans T.B.Cavalc. 97. D. venosus T.B.Cavalc. 98. D. verruculosus T.B.Cavalc. 99. D. vidalii Lourteig

61

J

Months of the year F M A M J J A S

O N D

62

Nº

7  Floral/Reproductive Biology and Phenology

Species

100. D. villosissimus Pohl 101. D. villosus Pohl 102. D. vinaceus T.B.Cavalc. 103. D. vittatus T.B.Cavalc. 104. D. virgatus Pohl var. occidentalis T.B.Cavalc. & S.A.Graham 105. D. virgatus Pohl var. virgatus 106. D. vividus T.B.Cavalc. 107. D. xerophilus T.B.Cavalc. 108. D. xiphodon T.B.Cavalc.

J

Months of the year F M A M J J A S

O N D

Chapter 8

Chromosomes

Most of the information on chromosome numbers in Lythraceae comes from the studies of Graham (1985, 1989, 1992). In a review of chromosome data for the family, Graham and Cavalcanti (2001) presented the chromosome number of 13 species of Diplusodon, the basic chromosome number of the genus as x = 8, and the gametic number of Diplusodon as n = 15 (2n = 30, ex. D. sordidus, Fig. 5.1B). This gametic number probably arose from an ancestral polyploid with a gametic number of n = 16, followed by the loss of a chromosome. Speciation in Diplusodon might be expected to have occurred as it does in many true island endemics, via other mechanisms (Crawford and Stuessy 1997), such as genetic isolation in campos rupestres, pointed out as an important mechanism of speciation (see Phylogeny and Evolution) considering that no  changes in chromosome numbers or chromosome rearrangements were found.  Chromosome numbers for 13 species of Diplusodon were known prior to this study, based on mitotic and meiotic counts. Table 8.1 presents a summary of counts made for Diplusodon, together with those already published.

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8 Chromosomes

Table 8.1  Chromosome reports in Diplusodon, summarized from original data and the literature Taxon D. alatus

Chromosome number 2n = 30

D. bradei D. candollei D. hexander D. hirsutus D. orbicularis

2n = 30 n = 15 n = 15 n = 15 2n = 30

D. paraisoensis D. praetermissus

2n = 30 2n = 30

D. ramosissimus

2n = 30

D. sordidus (Fig. 5.1D) D. ulei

2n = 30 2n = 30

D. villosus

2n = 30 n = 15

D. virgatus var. virgatus

2n = 30 2n = 30

Voucher Cavalcanti et al. 681

References Cavalcanti (1995), Graham and Cavalcanti (2001) Cavalcanti et al. 2301 Cavalcanti (1995) Graham 985 Graham (1992) Graham 979 Graham (1992) Cavalcanti et al. 2270 Graham and Cavalcanti (2001) Cavalcanti et al. Cavalcanti (1995), Graham and (CFSC) 10635 Cavalcanti (2001) Cavalcanti et al. 648 Graham and Cavalcanti (2001) Cavalcanti et al. 660 Cavalcanti (1995), Graham and Cavalcanti (2001) Cavalcanti et al. 586 Cavalcanti (1995), Graham and Cavalcanti (2001) Cavalcanti et al. 403 Cavalcanti (1995), Graham and Cavalcanti (2001) Cavalcanti et al. 2456 Cavalcanti (1995), Graham and Cavalcanti (2001) Cavalcanti et al. 500 Cavalcanti (1995), Graham and Cavalcanti (2001) Cavalcanti et al. 500 Graham (1985), Graham and Cavalcanti (2001) Cavalcanti et al. 571 Cavalcanti (1995), Graham and Cavalcanti (2001) Vieira et al. 469 Cavalcanti (1995)

Chapter 9

Chemistry

Since the 1970s, Lythraceae has been relatively well explored chemically. Raffauf and Altschul (1968) detected alkaloids in 25 genera of the family, and small amounts of alkaloids were reported for 6 species of Diplusodon. Ellagic acid was reported for all Lythraceae taxa, as well as quinolizidine alkaloids, naphthoquinones, and essential oils (Dahlgren and Thorne 1984). Graham and Kleiman (1987) reported that in the majority of Lythraceae genera, linoleic acid is the dominant fatty acid in seeds, while palmitic or oleic acid, or a combination of these two, is the second most abundant component. They report that Diplusodon is the only genus with capric acid as the second most abundant component of the seed oil and that in D. glaucescens both linoleic and capric acids are produced, as in Cuphea, but with a much lower percentage of capric acid. In seeds of Cuphea, fatty acids are produced in large amounts, and this has resulted in a large research investment in the genus. Santos and Salatino (1998) report the content of seed oils for 12 samples from 9 species of Diplusodon. With the exception of D. virgatus, the seeds are poor sources of oil. Palmitic, linoleic, and behenic acids appeared as predominant constituents, depending on the samples analyzed. Diplusodon ciliatiflorus and D. sordidus stood out with relatively high amounts of behenic and lignoceric acids. The observed intraspecific variation of fatty acid patterns was low in D. virgatus var. virgatus, but high in D. alatus and D. strigosus. Flavonoids have been relatively well studied in the families of Myrtales, and common flavones and their O-methyl derivatives have been reported in the order. Myricetin occurs in some genera of Combretaceae, Melastomataceae, Myrtaceae, and Onagraceae, and glycoflavones are reported in Lythraceae, Combretaceae, and Myrtaceae (Dahlgren and Thorne 1984). The flavonoid profiles of 27 Diplusodon species were determined by Blatt (1991) and Blatt et al. (1993, 1994). They reported a predominance of glycosylate flavones of quercetin and myricetin, showing a complex pattern of glycolyzation with a high incidence of diglycosides formed by the association of various sugars, such as

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glucose, galactose, arabinose, xylose, rhamnose, and glucuronic acid, the latter of which is present in ca. 15% of the flavonoids. Blatt (1991) and Blatt et al. (1993, 1994) noted further that some species of sections Palmatinerves and Penninerves have only flavone glycosides (apigenin and luteolin). Diplusodon chemistry does not support the subdivision of the genus into the sections proposed by Koehne (1903) or Lourteig (1989). Blatt (1991) and Blatt et al. (1988a, 1990b, 1991) also considered the taxonomic implications of variation in the paraffin fraction of the epicuticular wax of Diplusodon and concluded that at the species level the alkane patterns are relatively fixed, making them useful as a taxonomic marker. At the sectional level, only section Palmatinerves (sensu Koehne 1903) is supported by a C 33 peak. Blatt (1991) and Blatt et al. (1988b, 1990a, 1991) concluded that most of the flavonoids and the n-alkanes of the epicuticular wax are not correlated with Koehne’s sections.

Chapter 10

Phylogeny and Evolution

In Koehne’s (1903) classification of Lythraceae, Diplusodon is included in the tribe Lythreae, subtribe Diplusondontinae, together with Galpinia, Lafoensia, Pemphis J.R.Forst. & G.Forst., and Physocalymma. These five genera share the presence of characteristics such as actinomorphic flowers, basal placenta, interrupted septal walls of the ovary or septa divided above the placenta, and seeds dorsally compressed and winged. Koehne (1886) drew an intuitive phylogeny for the family based on geographic distribution of the genera and considered the Brazilian tree genera Physocalymma and Lafoensia to have originated from the subshrub members of Diplusodon. The topology of Koehne (1886) places the herbaceous genus Nesaea in a central position as the oldest genus from which the rest of the family diversified. Diplusodon and Pemphis, subshrubs and trees respectively, were derived from Lythrum, an essentially herbaceous genus of widespread global distribution. The latter genus would have also given rise to two other herbaceous and subshrub lineages, the American genera Cuphea and Pleurophora D.  Don, and the widely distributed Peplis L. and Rotala. Graham et al. (1993) determined the position of Diplusodon in relation to the rest of the genera of the family in a phylogenetic analysis using morphological data. In this work, the authors refer to the unique bipartite placenta of Diplusodon with its two semi-lunate septal walls. The phylogenetic analysis utilized 26 characters, including anatomical, morphological, palynological, and embryological data. The resulting topology of the five most parsimonious trees was identical in showing the genera Sonneratia, Duabanga, Punica, Lagerstroemia, and Lawsonia as composing one of two clades of the family, with the remainder of the genera, including Diplusodon, forming the second clade. The relationship of Diplusodon to other genera of the Lythraceae was not resolved, and the genus appeared in different positions, but it was always among the genera Tetrataxis Hook.f., Haitia Urb., Crenea Aubl., Lythrum, Nesaea Comm. ex Juss. (= Ammannia L.), Ammannia, Rotala, and

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Peplis. In the studies of Graham et al. (1993), the members with herbaceous habit are considered highly derived, exactly opposite to the suggestion of Koehne (1903). Combined morphological and molecular data from the nuclear ribosomal internal transcribed spacer (ITS) region, along with three chloroplast regions, rbcL, trnL-F, and the psaA-ycf3 intergenic spacer, were included in a phylogenetic analysis of Lythraceae (Graham et  al. 2005). The analysis suggested a relationship of Diplusodon to Physocalymma and Lourtella, although the support for this relationship was not strong (0.55 posterior probability in a Bayesian 50% majority-rule consensus of 4000 trees; Graham et al. 2005). A phylogenetic analysis of Diplusodon based on morphological data (Cavalcanti 1995) and ITS sequences combined morphological data (Cavalcanti and Rua 2008) supported the monophyly of the genus and concluded that Koehne’s (1903) and Lourteig’s (1989) taxonomic sections based on leaf venation were polyphyletic. Recently, Inglis and Cavalcanti (2018) worked on a time-calibrated molecular phylogeny of Diplusodon, utilizing 2 rapidly evolving nuclear loci and based on 80 of the 104 species now accepted in the genus. The Bayesian analysis, using a relaxed mixed clock model, was calibrated for divergence time based on fossil evidence, where a date of 14 Ma was used for the oldest fossil of Cuphea, 40 Ma for Punica, and 81  Ma for Lythrum (Graham et  al. 2013). Twelve genera of Lythraceae and Gaura in the sister family Onagraceae (Graham et al. 2005) were chosen as outgroups. The results obtained showed that the monophyly of Diplusodon was strongly corroborated in MP, ML, and Bayesian analyses of combined ETS and ITS and Diplusodon was placed as sister to a clade containing Cuphea, Woodfordia, Pleurophora, Adenaria, Lourtella, and Physocalymma, which, with the exception of Woodfordia, are all New-World genera. Five biogeographical areas were used by the authors in the RASP (Reconstruct Ancestral State in Phylogenies) analyses on the basis of their concentration of endemic Diplusodon species and geographical distinctiveness: (1) Cadeia do Espinhaço Central Plateau (CEP), a region of mountains around the municipality of Diamantina in Minas Gerais State, to the north, and the Serra do Cipó, to the south, including a disjunct group of mountains called the Serra do Cabral, to the west, in the central part of the Espinhaço Range, in Minas Gerais State (Fig.  10.1); (2) Canastra Arc (CA), a region of mountains encompassing the Serra da Canastra located in the south-central part of Minas Gerais State, and Northwest of São Paulo State, reaching in the eastern direction the mountains of the southern Espinhaço Range to the municipalities south of Belo Horizonte, in Minas Gerais State (Fig. 10.1); (3) Chapada Diamantina (CD), a region of mountains in central Bahia State, including Rio de Contas, Serra das Almas, and Sincorá ranges, completely surrounded by Brazilian semiarid vegetation (Fig. 10.1); (4) North of Minas Gerais (NMG), a region of mountains in the northern Espinhaço Range in Minas Gerais State, extending into the south of Bahia State, with a southern limit in the municipality of Turmalina (Fig. 10.1); and (5) Plateau Goiano (PG), the upland region in the eastern half of Goiás State and the southeast of Tocantins State, with the highest altitudes especially concentrated in the Chapada dos Veadeiros, in the Brazilian Shield (Fig. 10.1).

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Fig. 10.1  Biogeographical areas used in the RASP (Reconstruct Ancestral State in Phylogenies) analyses on the basis of concentration of endemic Diplusodon species and geographical distinctiveness. Source: Brazilian Institute of Geography and Statistics IBGE (2004) and collection data of T.B. Cavalcanti at CEN. From Inglis and Cavalcanti (2018)

The phylogenetic analysis resulted in four major geographical clades (Fig. 10.2) designated as the Cadeia do Espinhaço Plateau clade (CEP), the Plateau Goiano clade (PG), the Chapada Diamantina clade (CD), and the Cerrado clade (C; Inglis and Cavalcanti 2018). The CEP clade (Fig. 10.2) constitutes a morphologically coherent group, with species endemic to higher altitudes comprising open grassy fields on white sandy-­ rocky soils and campo rupestre, occurring in the central part of the Espinhaço Range. The CD clade (Fig. 10.2) is a small clade of four endemic species of the campos rupestres and rocky cerrado of Chapada Diamantina in Bahia State that shares with the C clade (Fig.  10.2) characters considered plesiomorphic in Diplusodon and within Lythraceae (Graham et al. 1993, 2005; Cavalcanti 1995; Cavalcanti and Rua 2008), such as the shrubby habit, dense indumentum, frondose polytelic

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Fig. 10.2  Bayesian time-calibrated phylogenetic ultrametric tree of combined ETS, ETS indel, and ITS data (50% majority rule Bayesian consensus). Diplusodon species names are followed by their current sectional assignment if available (sensu Lourteig 1989): Dip  =  sect. Diplusodon; Pal = sect. Palmatinerves; Pen = sect. Penninerves. Geographical center and mountain range: PG Plateau Goiano (chain of mountains in central west and part of north of Brazil, 1000–1500 m) and CEP Cadeia do Espinhaço Plateau (chain of mountains in the Southeast and part of northeast of Brazil, 1000–1400 m); WD widely distributed, NMG northern of Minas Gerais State, CA Canastra Arc. Posterior probabilities ≥0.9 are given above branches and mean divergence dates given at nodes. Horizontal bars at nodes represent the 95% highest posterior density of the Bayesian posterior distribution. From Inglis and Cavalcanti (2018)

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71

synflorescences, and diplostemony. Many species in the C clade are perennial hemicryptophytes, associated with a well-developed, woody, subterranean, and perennial xylopodium. In addition to species with plesiomorphic characters, the C clade contains predominantly species that are widely distributed in low-elevation cerrado, in addition to two species from ecotones with caatinga (D. quintuplinervius and D. thymifolius), and species occurring in the mountains of the southernmost Espinhaço Range and from the Serra da Canastra, southwestern mountains of Minas Gerais State. The PG clade (Fig.  10.2) contains species exclusively from the mountains of Goiás and Tocantins States in the Central Brazilian Plateau. Members of this clade have many unique morphological features in the genus, which do not occur in species of the Espinhaço Range and Chapada Diamantina, such as the following: (i) monotelic synflorescences (D. ovatus, D. panniculatus); (ii) an increase in pedicel length, from 0–3 mm to 1.5–2 cm (D. cryptanthus, D. marginatus, D. longipes); (iii) an increase in corolla diameter from 2–3 cm to 4.5–7 cm (D. ciliatiflorus, D. marginatus, D. kielmeyeroides, D. marginatus); (iv) multiplication of the stamens of the episepalous whorl to 18–54 stamens (seen in 70% of species of the PG clade); (v) presence of prominent cordiform, callose glands occurring on the internal margin of the floral tube at the base of the petals (D. astictus, D. appendiculosus, and D. leucocalycinus); (vi) leaves with 2–4 inconspicuous lateral veins obscured by a thick mesophyll and wax composed of 31–33 carbon atom alkanes (Blatt et  al. 1991); (vii) presence of hairs on the top of the ovary and glands at the base of the petals, believed to attract pollinators (D. leucocalycinus and D. appendiculosus); and (viii) lack of epicalyx (D. imbricatus and D. marginatus clade). Inglis and Cavalcanti (2018) indicated a mean date estimate of 3.46 Ma (95% HPD = 2.46–4.57 Ma) for the Diplusodon crown node (Fig. 10.2), with more intense speciation occurring approximately in the last one million years. The preponderance of terminal polytomies in the Bayesian phylogeny, as well as morphological similarities, led the authors to conclude recent and closely spaced speciation events in the campos rupestres. The authors indicated that the major divergences in Diplusodon were possibly triggered by climate change associated with Middle Pleistocene glaciation cycles. Diversity patterns and endemism of Diplusodon species in the mountains of the Espinhaço Range and Plateau Goiano pointed to allopatry, probably involving altitudinal isolation in the campos rupestres with consequent genetic isolation from lowland populations as an important mechanism of speciation in Diplusodon (Inglis and Cavalcanti 2018). In Diplusodon, the CEP, CD, and C-2 clades are composed of a majority of endemic species exclusive to campo rupestre, while some other clades contain species occupying a continuum of campo rupestre and cerrado or combinations thereof, with a few species found in forest margin habitats. The biogeographic zones of the cerrados and campos rupestres are clearly interconnected, and, given the high frequency of microendemic species, especially in the PG clade and the young age of many Diplusodon clades, an Old Stable Landscapes (OSL) model (Mucina and

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Wardell-Johnson 2011) favoring high levels of in situ speciation in the campos rupestres is more evident (Inglis and Cavalcanti 2018). Subsequent radiation and long-distance dispersal were most evident in the C clade in Diplusodon. The possession of a xylopodium as a fire survival adaptation is present in many species of the C clade; however, this feature is also present in all other Diplusodon clades (Fig. 10.2). Other adaptations and specialization such as decrease in flower size and decrease in number of stamens by loss of one whorl of stamens may be related to adaptation to the windy and open environments of the campos rupestres, leading to pollination specialization (Inglis and Cavalcanti 2018).

Chapter 11

Geographic Distribution and Ecology

Diplusodon occurs mostly in the Cerrado Biome (Figs. 11.1 and 10.1), the floristically diverse savannah that covers more than two million km2 of the Central Brazilian Plateau, extending west into Bolivia, south to Paraguay, and east to the Caatinga (Eiten 1972; Loeuille et al. 2015). The genus is most diverse in the mountains of Plateau Goiano in the west central region of Brazil (west of Rio São Francisco, in the states of Goiás and Tocantins and the Distrito Federal), and this region is considered the primary center of diversity of the genus. In the Plateau Goiano, more precisely the mountain chain that forms the Serra Geral do Paranã, Serra Geral de Goiás, Chapada dos Veadeiros, Serra dos Pireneus, and Espigão Mestre, there are 63 (60.5%) species of Diplusodon (Fig. 11.2). Of the 63 species on the Plateau Goiano, 58 species are from Goiás State, with 37 endemic species; 16 species are from Tocantins State, with 2 endemic species; and 11 species are from the Distrito Federal (Fig. 11.3). The secondary center of diversity for Diplusodon is in the mountains of Minas Gerais State (Cadeia do Espinhaço), with 44 species (42%) with 29 endemic species (Fig. 11.3). The rest of the species are distributed in the states of Bahia (12 species, 6 microendemics), Mato Grosso (8 species, 1 microendemic), São Paulo (4 species), Maranhão (2 species), Mato Grosso do Sul (2 species), Piauí (2 species), and Rondônia (1 species), in addition to 2 taxa that also occur in Bolivia. Analysis of the geographic distribution of the species reveals patterns that can be compared with those already proposed for other families or genera that, like Diplusodon, are predominantly found in the mountains of Brazil. Giulietti and Pirani (1988) recognized five patterns of geographic distribution that were detected through the analysis of various families and genera that occur predominantly in the campos rupestres of Cadeia do Espinhaço in Minas Gerais and Bahia (Chapada Diamantina): (1) species widely distributed in Tropical America, especially in the campos rupestres and cerrados; (2) species distributed in the mountains of northern South America (principally Venezuela and the Guianas) and in the Cadeia do Espinhaço, in similar habitats in both regions; (3) species that occur in the restingas and in the Cadeia do Espinhaço; (4) species that occur in the Cadeia do Espinhaço

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Fig. 11.1  Geographic distribution of the genus Diplusodon (Lythraceae) in the Cerrado Biome (gray area) of Brazil and Bolivia

and the serras of Goiás (these serras are similar in edaphic aspects and climatic features); and (5) species endemic to the Cadeia do Espinhaço (with separate patterns recognized for species that occur on more than one serra and species that are restricted to a single serra). The following three patterns of geographic distribution are recognized for Diplusodon: 1. Widely distributed species, including species that extend across different habitats within the Cerrado Biome. This pattern of widespread distributions is basically in agreement with the first pattern proposed by Giulietti and Pirani (1988), but it occurs on a smaller scale. The most widely distributed species is

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Fig. 11.2  Percentage of occurrence of Diplusodon species by states. BA Bahia, DF Distrito Federal, GO Goiás, MA Maranhão, MG Minas Gerais, MS Mato Grosso do Sul, MT Mato Grosso, PI Piauí, RO Rondônia, SP São Paulo, TO Tocantins

Diplusodon virgatus, represented in Bolivia and in 7 of the 11 states of Brazil of which Diplusodon occurs. The broad distribution of this species may be related to its capacity to occupy a variety of habitats, inhabiting gallery and dry forest margins, dense cerrado, and cerrado sensu stricto of low elevation, contrasting with the characteristic highly restricted elements of rocky cerrado or campo rupestre inhabited by most Diplusodon species. In the phylogeny presented here, the position of D. virgatus as an independent branch in the deepest node of the phylogeny in the Bayesian tree (Fig. 10.2) shows an older origin for the species. Curiously, D. virgatus is the only species of the genus with a white corolla. Other species of Diplusodon that are included in the category of widely distributed species are found in three to five Brazilian states and are also species which inhabit areas of low-elevation cerrado (400–900 m) and ecotones with caatinga. These species are D. bolivianus, which occurs in Bolivia (Santa Cruz) and Brazil (Distrito Federal, Mato Grosso, and Rondônia); D. helianthemifolius var. pemphoides, which occurs in the Cadeia do Espinhaço, in Minas Gerais, in the

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Fig. 11.3  Number of endemic species of Diplusodon by Brazilian states. BA Bahia, DF Distrito Federal, GO Goiás, MA Maranhão, MG Minas Gerais, MS Mato Grosso do Sul, MT Mato Grosso, PI Piauí, RO Rondônia, SP São Paulo, TO Tocantins

Espigão Mestre, in Bahia, and in the cerrados of Goiás, near Brasília and Maranhão; D. lanceolatus, which occurs in the Distrito Federal, Goiás, Minas Gerais, and Mato Grosso; D. oblongus, which occurs in Goiás, Tocantins, and Mato Grosso; D. ovatus, which occurs in Goiás, Mato Grosso, Mato Grosso do Sul, Minas Gerais, and São Paulo; D. strigosus, which occurs in Goiás, Tocantins, and Mato Grosso; and D. thymifolius which occurs in Bahia, Minas Gerais, Piauí, and Tocantins. This distribution pattern, where one to a few species with broad geographic distribution occur among a majority of geographically restricted species, is frequent among genera found in the campos rupestres (Giulietti 1984; Harley 1988; Giulietti and Hensold 1991; Kirkbride 1976; Semir 1991; Rapini et al. 2008). 2. Species with distributions that are disjunct between the Cadeia do Espinhaço Plateau (Minas Gerais) and the Plateau Goiano (Goiás, Distrito Federal, and part of Tocantins). Diplusodon helianthemifolius var. pemphoides (Fig. 13.46) and Diplusodon lanceolatus (Fig. 13.55) have disjunct distributions between the Cadeia do Espinhaço and the Plateau Goiano, occurring east and west of Rio São Francisco, which seems to have acted as a major barrier to most

11  Geographic Distribution and Ecology

77

species from the Espinhaço Range and the Plateau Goiano. However, the uplands of the Serra da Canastra, together with the northwest of São Paulo State and southern Minas Gerais, treated by Inglis and Cavalcanti (2018) as the Canastra Arc, appear to have been an area of communication between the chain of mountains of the Espinhaço Range and the chain of mountains of the Plateau Goiano (Fig. 10.1). The Canastra Arc may therefore act as a biotic corridor, as suggested for many other plant and animal genera occupying high-­altitude areas in southeastern Brazil (Safford 2007). The pattern of floristic similarity between the Serra da Canastra and the mountains in Goiás has already been reported for other campo rupestre groups (Dutra et  al. 2014; Nakajima and Semir 2001; MelloSilva 2004; Rando and Pirani 2011; Romero and Martins 2002). Studies of phylogenetic relationships corroborate the biogeographic pattern where species in the mountains of the Espinhaço Range and Serra da Canastra and in Goiás are closely related (Loeuille et al. 2015; Ribeiro et al. 2014). 3 . Species restricted to a single chain of mountains or to a single mountain. This is the most common pattern in Diplusodon and reflects the high degree of endemism in the genus, where the species are restricted to campos rupestres on quartzite rocks. Endemism here is used as defined for plants that grow in communities that are isolated by geographical or climatic barriers, such as islands or mountains (Campbell 1989). For Diplusodon, “microendemic” applies to species that occur on just one mountain, at one locality, or in one municipal district, and “endemic” refers to species that occur in one state or several municipal districts (Table 11.1). In Diplusodon ca. 65% of Diplusodon species are predominantly endemic to the specialized campos rupestres and high-altitude rocky cerrado (Table  11.1), frequently occurring in small populations and often restricted to just one or a few adjoining mountains. However, the species can also form dense populations. In a hectare of campo rupestre in the Serra do Cipó (Minas Gerais), Faria (1994) found D. orbicularis and D. hirsutus populations with 2518 individuals and 11 individuals, respectively. Normally it is easy to detect Diplusodon in the field due to the large number of individuals that make up the populations, and because of their synchronous flowering. In the campos rupestres, species of the genus grow in the deep cracks of the rocks or on the surface of rocks, as in Diplusodon helianthemifolius var. helianthemifolius in the Diamantina region of Minas Gerais. Other species, such as D. astictus in Minaçu in northern Goiás, grow in wet and temporarily flooded soils, such as the saturated “campos brejosos” of the Chapada dos Veadeiros (Goiás). The species that grow in savanna and forest environments are differentiated by their life form: most often small- or medium-sized subshrubs, in the open campos and campos cerrados, and tall subshrubs to small trees along the margins of forests and in dense cerrado. Most species of Diplusodon have characteristics that are typical of plants that grow in cerrado vegetation. They lose part of their leaves during the reproductive stage, and they remain perennial by means of their resistant subterranean organ, the xylopodium, which is an adaptation for seasonal drought and fire.

MG GO, TO MG GO MG MG DF, GO GO GO

1. 2. 3. 4. 5. 6. 7. 8. 9.

10. 11. 12. 13. 14. 15. 16. 17. 18.

Brazilian states GO MG GO GO GO BA GO BA DF, MT, RO

D. adpressipilus Lourteig D. aggregatifolius T.B.Cavalc. D. alatus T.B.Cavalc. D. appendiculosus Lourteig D. argenteus Lourteig D. argyrophyllus T.B.Cavalc. D. astictus Lourteig D. bahiensis T.B.Cavalc. D. bolivianus T.B.Cavalc. & S. Graham D. bradei Pilg. D. burchellii Koehne D. buxifolius (Cham. & Schltdl.) DC. D. caesariatus Lourteig D. canastrensis T.B.Cavalc. D. candollei Pohl ex DC. D. capitalensis T.B.Cavalc. D. chapadensis T.B.Cavalc. D. ciliatiflorus T.B.Cavalc.

Distribution

No. Species

– – – – – – – – –

CA, CEP PG CEP PG CA CEP PG PG PG

Biogeographic Bolivia units – PG – CEP – PG – PG – PG – CD – PG CD Santa Cruz PG CR CE, CL, CR, CS CR CE, eF CR, CS CR CE CR CE, dCE, eF

Phytophysiognomy CL, CR CR, CS CL, CR, CS CR CL, CR CR CL, CR CR, CS CE, CS, dCE, eF E E E E ME E E ME ME

Occurrence in geographic distribution ME ME ME ME ME ME E ME WD

Table 11.1  Geographic distribution of Diplusodon taxa in Brazil and Bolivia with information about phytophysionomy, endemism, and rarity

O C C R O O O R O

Frequency in the area of occurrence O O O R R R O O O

78 11  Geographic Distribution and Ecology

Brazilian states BA MG GO GO GO, TO GO, TO GO, MG MG GO GO GO MG MG MG, SP TO GO GO

19. 20. 21. 22. 23. 24. 25. 26. 27. 28. 29. 30. 31. 32. 33. 34. 35.

D. ciliatus (T.B.Cavalc.) T.B.Cavalc. D. ciliiflorus Koehne D. conduplicatus T.B.Cavalc. D. cordifolius Lourteig D. cryptanthus T.B.Cavalc. D. decussatus Gardner D. divaricatus Pohl D. epilobioides Mart. ex DC. D. ericoides Lourteig D. fastigiatus Lourteig D. floribundus Pohl D. glaucescens DC. D. glaziovii Koehne D. glocimarii T.B.Cavalc. D. gracilis Koehne D. grahamae T.B.Cavalc. D. hatschbachii Lourteig

Distribution

No. Species

Bolivia – – – – – – – – – – – – – – – – –

Biogeographic units CD CEP PG PG PG PG PG CEP PG PG PG CEP CEP CA PG PG PG Phytophysiognomy CR CL, CR CR CL, CR CE CL, CR, CS CE, CS CE, CR CR CR CE CR CR CL, CR CE CR CR, CS

Occurrence in geographic distribution E E ME E E ME E E E ME ME ME ME E ME ME ME

(continued)

Frequency in the area of occurrence O C R C O R O O C R O O R C R R R

11  Geographic Distribution and Ecology 79

43. 44. 45. 46. 47. 48.

42b.

37. 38. 39. 40. 41. 42a.

36b.

36a.

Distribution

Brazilian states D. helianthemifolius Mart. ex DC. var. MG helianthemifolius D. helianthemifolius var. pemphoides BA, GO, MA, MG (DC.) Koehne D. heringeri Lourteig GO D. hexander Mart. ex DC. MG D. hirsutus (Cham. & Schltdl.) DC. MG D. imbricatus Pohl GO, TO D. incanus Gardner BA, GO D. kielmeyeroides A.St.-Hil. var. MG kielmeyeroides GO D. kielmeyeroides var. macrocalyx T.B.Cavalc. D. lanceolatus Pohl DF, GO, MG, MT D. leucocalycinus Lourteig GO D. longipes Koehne GO D. macrodon Koehne GO, TO D. marginatus Pohl DF, GO D. mattogrossensis T.B.Cavalc. MT

No. Species

Table 11.1 (continued)

CEP, PG PG PG PG PG –

PG

– – – – – – –

PG CEP CEP PG PG PG

CEP, PG

– – – – – –

–

Bolivia –

Biogeographic units CEP

CE CR, CS CE, CR, CS dCE, CE CE, CS CE

CR

dCE, CR CR, CS CR CE, CS CE, eF CE, CR

CE, CR

Phytophysiognomy CE, CR, eF

WD ME ME E E ME

ME

E E E E E ME

WD

Occurrence in geographic distribution ME

O O O O O R

R

O C C O O R

C

Frequency in the area of occurrence O

80 11  Geographic Distribution and Ecology

D. micromerus T.B.Cavalc. D. microphyllus Pohl D. minasensis Lourteig D. mononeuros Pilg. D. myrsinites Mart. ex DC. D. nigricans Koehne D. nitidus Mart. ex DC. D. oblongus Pohl

D. obtectus T.B.Cavalc. D. orbicularis Koehne D. ovatus Pohl

D. panniculatus Koehne D. paraisoensis Lourteig D. parvifolius Mart. ex DC. D. petiolatus (Koehne) T.B.Cavalc. D. plumbeus T.B.Cavalc. D. praetermissus T.B.Cavalc. D. prostratus T.B.Cavalc. D. psammophilus Lourteig

49. 50. 51. 52. 53. 54. 55. 56.

57. 58. 59.

60. 61. 62. 63. 64. 65. 66. 67.

No. Species

Brazilian states GO MG MG MG MG GO BA, MG DF, GO, MG, MT, TO MG MG GO, MT, MS, MG, SP GO GO, TO BA GO, TO GO GO GO MG

Distribution

– – – – – – – –

– – –

Bolivia – – – – – – – –

PG PG CD PG PG PG PG CEP

CEP CEP PG, CA

Biogeographic units PG CEP CEP CEP CEP PG CEP PG

CE, CR CE, CL, CR, CS CE, CR, CS CE, CS CE, CR CR CE, CR CR

CR CL, CR, CS CE

Phytophysiognomy CE, CR CR CR CE, eF CR CE, CR CE, CR CE, CR, CS

ME E ME E ME ME ME ME

ME ME WD

Occurrence in geographic distribution ME E E ME E ME E WD

C O F O O O O O

O F R

(continued)

Frequency in the area of occurrence R C R R O R O C

11  Geographic Distribution and Ecology 81

Distribution

Brazilian states 68. D. puberulus Koehne MG 69a. D. punctatus var. dentatus T.B.Cavalc. GO, TO 69b. D. punctatus Pohl var. punctatus GO 70. D. pygmaeus T.B.Cavalc. GO 71. D. quintuplinervius (Nees) Koehne BA, MG 72. D. ramosissimus Pohl DF, GO, MG 73. D. rectinervis T.B.Cavalc. MG 74. D. retroimbricatus Koehne GO 75. D. rosmarinifolius A.St.-Hil. DF, GO 76. D. rotatus T.B.Cavalc. MA, PI 77. D. rotundifolius Mart. ex DC. MG 78. D. saxatilis Lourteig MG 79. D. sessiliflorus Koehne DF, GO 80. D. sigillatus Lourteig GO 81. D. sordidus Koehne GO 82. D. speciosus (Kunth) DC. MT, TO 83. D. stellatus T.B.Cavalc. GO

No. Species

Table 11.1 (continued)

– – – – – – – – – – –

Bolivia – – – – –

Biogeographic units CEP PG PG PG CEP PG CA PG PG – CEP CEP PG PG PG PG PG Phytophysiognomy CE, CS CE, CS CE, CS CR CAA, CR, CS CE CR CE, CR CE, CL, eF CE CR CR CE CR, CS CR CE, CS CL, CR, CS

Occurrence in geographic distribution E E E ME E E ME ME E ME ME ME E ME ME WD ME

Frequency in the area of occurrence O O C R O C R R C O F R O O O O O

82 11  Geographic Distribution and Ecology

D. strigosus Pohl D. subsericeus Casar. ex Koehne D. tenuifolius T.B.Cavalc. D. thymifolius Mart. ex. DC. D. trigintus T.B.Cavalc. D. ulei Koehne D. uninervius Koehne D. vacillans T.B.Cavalc. D. venosus T.B.Cavalc. D. verruculosus T.B.Cavalc. D. vidalii Lourteig D. villosissimus Pohl D. villosus Pohl D. vinaceus T.B.Cavalc. D. virgatus Pohl var. virgatus

98b. D. virgatus var. occidentalis T.B.Cavalc. & S.Graham 99. D. vittatus T.B.Cavalc. 100. D. vividus T.B.Cavalc.

84. 85. 86. 87. 88. 89. 90. 91. 92. 93. 94. 95. 96. 97. 98a.

No. Species

– – – – – – – Santa Cruz – – –

GO MG MG MG, SP DF, GO, MG, TO GO DF, GO, MG, MT, SP, TO MT, MS

GO MG

PG CEP

PG CEP CA CA, CEP PG CA CEP, PG

Bolivia – – – – – – –

Biogeographic units PG CEP CA CD, PG PG CD CEP

Brazilian states DF, GO, MT, TO MG GO BA, MG, PI, TO TO BA MG

Distribution

CL CR

CE, M

CE, CR CR CE, CR, Ef CE, CR, CS CE, CR, EF CE CE, eF

Phytophysiognomy CE, CL, CR, CS, eF CE, CR CE, CR CAA, CE, CR, CS CE CR CR

ME ME

WD

ME ME ME E WD ME WD

Occurrence in geographic distribution WD ME ME WD ME E E

R O

O

R R C O O R O

(continued)

Frequency in the area of occurrence C R O O R O O

11  Geographic Distribution and Ecology 83

Brazilian states GO BA MG GO

101. 102. 103. 104.

Bolivia – – – –

Biogeographic units PG CD CEP PG Phytophysiognomy CE, CR CR CE CR

Occurrence in geographic distribution ME ME ME ME

Frequency in the area of occurrence O R R R

Abbreviations Brazilian states: BA Bahia, DF Distrito Federal, GO Goiás, MA Maranhão, MG Minas Gerais, MS Mato Grosso do Sul, MT Mato Grosso, PI Piauí, RO Rondônia, SP São Paulo, TO Tocantins. Biogeographic units (Inglis and Cavalcanti 2018): CA Canastra Arc, CD Chapada Diamantina, CEP Cadeia do Espinhaço Central Plateau, PG Plateau Goiano. Phytophysiognomy: CAA transition with caatinga (dry forest), CE cerrado (Brazilian savanna), CL campo limpo (grass fields), CR campo rupestre (rocky fields including rocky cerrado (ca. 350–850 m)), CS campo sujo (fields with shrub by savanna elements), dCE cerrado denso (dense arborial closed savannas), eF edge of gallery or dry forest. Occurrence in geographic distribution: E endemic, F frequent, ME microendemic, O occasional, R rare, WD widely distributed.

D. xerophilus T.B.Cavalc. D. xiphodon T.B.Cavalc. D. capitatus (A.St.-Hil.) Koehne D. thysanosepalus Lourteig & Sandwith

Distribution

No. Species

Table 11.1 (continued)

84 11  Geographic Distribution and Ecology

11  Geographic Distribution and Ecology

85

Morphological novelties previously unknown in the genus has been registered in endemic species of Diplusodon. Recently, a high number of locally unique Diplusodon species have been described to Serra Geral de Goiás, especially in the Chapada dos Veadeiros (Lourteig 1989; Cavalcanti 2004, 2015; Cavalcanti and Graham 2005). This region has elevations between 1200 and 1600  m above sea level, and it is predominantly characterized by areas of campos limpos, campos rupestres, and campos cerrados, with gallery forests and many “veredas,” which are swampy areas dominated by “buritis” (Mauritia flexuosa Mart.). It coincides with the fact that Diplusodon has undergone intense speciation in these areas (see Phylogeny). These characteristics are each shared by two or three species and have not spread very widely in the region. Harley (1988) believes that in the mountains of Brazil, under changing climates, altitudinal migrations occurred that supplied “refugia” during the adverse periods of changing environments. These changes also resulted in the division of populations, producing a mechanism of speciation called the “species pump” (Stebbins 1974), which might account for the high diversity seen in campos rupestres. Multiple, independent dispersals from campo rupestre to cerrado have occurred in several unrelated Diplusodon lineages (Inglis and Cavalcanti 2018), adding support to the notions that the campos rupestres could have acted as a “biodiversity pump” for the surrounding cerrados, not only as a response to Pleistocene climatic fluctuations influencing their boundaries (Noonan and Gaucher 2006; Rull 2011; Antonelli et al. 2010) but by acting as a source of new lineages able to resist fire regimes by exaptation (Conceição et al. 2016; Silveira et al. 2016). It is clear from the distributional analysis of Diplusodon that it is a genus with specific ecological requirements, which define the cerrados of low elevation, such as highly dystrophic soils with low fertility and high acidity and mountain vegetation of campos rupestres with soils mainly based on quartzite-sandstone rock outcrops. The adaptability of many Diplusodon species, evidenced by their survival in both campo rupestre and in cerrado, explains in some way the remarkable species richness of the genus.

Chapter 12

Conservation

In Brazil, the Cerrado Biome occupies ca. 1.8 million km2, which is 23% of the Brazilian territory (Pagotto et al. 2006). The Cerrado is best represented in the west central part of the country, where a good part of its original area is still preserved, even though it is mixed with extensive soybean plantations. The destruction of the cerrado is evident in the extensive plains and plateaus of Goiás since these areas are favored for cultivation. The rupestrian savannahs, growing between rocks, campos rupestres and cerrado vegetation on mountains, which are localized at higher elevations and have extreme topography with sandy soils intersected by rocky outcrops and slopes are areas still less affected by agricultural activities. Especially in these areas, between 1000 and 1600 m, is where most of Diplusodon species occur. A small proportion of the species inhabit the cerrados at lower elevations, from 400 to 800 m, found mainly in Mato Grosso, São Paulo, and Minas Gerais states, are the most threatened species. The species occurring in flat areas of Cerrado are especially threatened because of exploitation for agriculture and pasture. These areas are disappearing rapidly, and the area protected in established conservation units within the Cerrado Biome is small. The Cerrado Biome is known as a hotspot and should be treated as a priority region for conservation (Myers et al. 2000; Brooks et al. 2006). Other  species of Diplusodon are threatened because of restricted distributions (Table 11.1). Of the 108 taxa of Diplusodon, 43 (39.8%) have occurrence records in conservation units, of which 30 (27.7%) have records of occurrence in national parks (Table 12.1). Despite the showy flowers of Diplusodon species, none are used for horticultural or other economic purposes.

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12 Conservation

Table 12.1  Conservation status of Diplusodon taxa

1. 2. 3. 4. 5. 6. 7. 8. 9. 10. 11. 12. 13. 14. 15. 16. 17. 18. 19. 20. 21. 22. 23. 24. 25. 26. 27. 28. 29. 30. 31. 32. 33. 34. 35. 36. 37. 38. 39. 40. 41.

Species D. adpressipilus Lourteig D. aggregatifolius T.B.Cavalc. D. alatus T.B.Cavalc. D. appendiculosus Lourteig D. argenteus Lourteig D. argyrophyllus T.B.Cavalc. D. astictus Lourteig D. bahiensis T.B.Cavalc. D. bolivianus T.B.Cavalc. & S.Graham D. bradei Pilg. D. burchellii Koehne D. buxifolius (Cham. & Schltdl.) DC. D. caesariatus Lourteig D. canastrensis T.B.Cavalc. D. candollei Pohl ex DC. D. capitalensis T.B.Cavalc. D. capitatus (A.St.-Hil.) Koehne D. chapadensis T.B.Cavalc. D. ciliatiflorus T.B.Cavalc. D. ciliatus (T.B.Cavalc.) T.B.Cavalc. D. ciliiflorus Koehne D. conduplicatus T.B.Cavalc. D. cordifolius Lourteig D. cryptanthus T.B.Cavalc. D. decussatus Gardner D. divaricatus Pohl D. epilobioides Mart. ex DC. D. ericoides Lourteig D. fastigiatus Lourteig D. floribundus Pohl D. glaucescens DC. D. glaziovii Koehne D. glocimarii T.B.Cavalc. D. gracilis Koehne D. grahamae T.B.Cavalc. D. hatschbachii Lourteig D. helianthemifolius Mart. ex DC. var. helianthemifolius D. helianthemifolius var. pemphoides (DC.) Koehne D. heringeri Lourteig D. hexander Mart. ex DC. D. hirsutus (Cham. & Schltdl.) DC.

Categories of threat (IUCN) VUD2 VUD2 ENB2a VUD2 ENB2a ENB2a VUD2 VUD2 LC VUD2 VUD2 NT VUD2 VUD2 VUD2 VUD2 PEX VUD2 VUB2aiii VUD2 VUB2aiii CRB1a VUB2biii VUB2aiii ENB2a ENB2biii,iv VUB2biii VUB2biii CRB2a ENB2biii,iv VUD2 CRB2a VUB2biii CRB2biii CRB2biii VUD2 VUB2biii

Occurrence in conservation units PNCV − PNCV − − − PNCV PEMC PNNKM PNSC PNCV − − PNSC PNSCi − − − PNCV PNCD PERP RPPNST − − − PECN − − − − − − PNSC − − − −

LC

PESP, PESN, PNSCi − − −

VUB2biii VUB2biii LC

(continued)

12 Conservation

89

Table 12.1 Continued

46. 47. 48. 49. 50. 51. 52. 53. 54. 55. 56. 57. 58. 59.

Species D. imbricatus Pohl D. incanus Gardner D. kielmeyeroides A.St.-Hil. var. kielmeyeroides D. kielmeyeroides var. macrocalyx T.B.Cavalc. D. lanceolatus Pohl D. leucocalycinus Lourteig D. longipes Koehne D. macrodon Koehne D. marginatus Pohl D. mattogrossensis T.B.Cavalc. D. micromerus T.B.Cavalc. D. microphyllus Pohl D. minasensis Lourteig D. mononeuros Pilg. D. myrsinites Mart. ex DC. D. nigricans Koehne D. nitidus Mart. ex DC. D. oblongus Pohl

60. 61. 62. 63. 64. 65. 66. 67. 68. 69. 70. 71. 72. 73. 74. 75. 76. 77. 78. 79. 80. 81.

D. obtectus T.B.Cavalc. D. orbicularis Koehne D. ovatus Pohl D. panniculatus Koehne D. paraisoensis Lourteig D. parvifolius Mart. ex DC. D. petiolatus (Koehne) T.B.Cavalc. D. plumbeus T.B.Cavalc. D. praetermissus T.B.Cavalc. D. prostratus T.B.Cavalc. D. psammophilus Lourteig D. puberulus Koehne D. punctatus var. dentatus T.B.Cavalc. D. punctatus Pohl var. punctatus D. pygmaeus T.B.Cavalc. D. quintuplinervius (Nees) Koehne D. ramosissimus Pohl D. rectinervis T.B.Cavalc. D. retroimbricatus Koehne D. rosmarinifolius A.St.-Hil. D. rotatus T.B.Cavalc. D. rotundifolius Mart. ex DC.

42. 43. 44. 45.

Categories of threat (IUCN) VUB2biii VUB2biii CRB2a

Occurrence in conservation units − − −

VUD2

−

LC VUB2biii VUB2biii VUB2biii VUB2biii ENB2a CRB2a LC VUB2biii CRB2biii VUD2 ENB2a VUB2aiii LC

PERP − RPPNST − − − − EEUFMG PEB; PNSV − PEI − − PEAMP; PNB; PNCV − PNSCi PNSC − RPPNST − − − PNCV − − RECCPI − − − − RBAE; RBIBGE PNSC RPPNST JBB; PNB − −

CRB2a LC VUB2biii CRB2biii VUB2biii VUB2aiii VUB2biii CRB2a VUD2 CRB2a CRB2biii VUB2biii VUB2biii VUB2biii VUD2 VUB2aiii LC CRB2a CRB2a CRB2biii CRB2biii VUBD2

(continued)

90

12 Conservation

Table 12.1 Continued

104.

Species D. saxatilis Lourteig D. sessiliflorus Koehne D. sigillatus Lourteig D. sordidus Koehne D. speciosus (Kunth) DC. D. stellatus T.B.Cavalc. D. strigosus Pohl D. subsericeus Casar ex Koehne D. tenuifolius T.B.Cavalc. D. thymifolius Mart. ex. DC. D. thysanosepalus Lourteig & Sandwith D. trigintus T.B.Cavalc. D. ulei Koehne D. uninervius Koehne D. vacillans T.B.Cavalc. D. venosus T.B.Cavalc. D. verruculosus T.B.Cavalc. D. vidalii Lourteig D. villosissimus Pohl D. villosus Pohl D. vinaceus T.B.Cavalc. D. virgatus var. occidentalis T.B.Cavalc. & S.Graham D. virgatus Pohl var. virgatus

105. 106. 107. 108.

D. vittatus T.B.Cavalc. D. vividus T.B.Cavalc. D. xerophilus T.B.Cavalc. D. xiphodon T.B.Cavalc.

82. 83. 84. 85. 86. 87. 88. 89. 90. 91. 92. 93. 94. 95. 96. 97. 98. 99. 100. 101. 102. 103.

Categories of threat (IUCN) CRB2a VUB2biii VUB2biii VUB2biii VUB2biii VUB2aiii LC CRB2a CRB2a LC CRB2a CRB2biii VUD2 VUD2 CRB2a CRB2a CRB2a CRB2a VUD2 LC CRB2a VUD

Occurrence in conservation units − PNB − PNCV PNCV PNCV PESCN − EEUU; PNGSV − − − PESRP; PNSV − RPPNST − − PEFBJ; PNSC JBB; PNB; REAE

LC

JBB; PEAMP; PEFBJ; PESP; PNGSV; PNSC; REAE − − − −

CRB2a VUD2 VUB2biii CRB2biii

PNNKM

Occurrence in conservation units: EEUFMG Ecological Station of the Federal University of Minas Gerais (Brazil), EEUU Ecological Station of Urucuí-Uma Piauí, Brazil), JBB Brasilia Botanical Garden (Distrito Federal, Brazil), PEAMP Altamiro de Moura Pacheco State Park (Goiás, Brazil), PEB Biribiri State Park (Minas Gerais, Brazil), PEFBJ Furnas do Bom Jesus State Park (São Paulo, Brazil), PEI Ibitipoca State Park (Minas Gerais, Brazil), PEMC Morro do Chapéu State Park (Bahia, Brazil), PERP Rio Preto State Park (Minas Gerais, Brazil), PESCN Serra de Caldas Novas State Park (Goiás, Brazil), PESN State Park of Serra Nova (Minas Gerais, Brazil), PESP Serra dos Pirineus State Park (Goiás, Brazil), PNB Brasilia National Park (Distrito Federal, Brazil), PNCD Chapada Diamantina National Park (Bahia, Brazil), PNCV Chapada dos Veadeiros National Park (Goiás, Brazil), PNGSV Grande Sertão Veredas National Park (Minas Gerais, Brazil), PNNKM Noel Kempff Mercado National Park (Santa Cruz, Bolivia), PNSC Serra da Canastra National Park (Minas Gerais, Brazil), PNSCi Serra do Cipó National Park (Minas Gerais, Brazil), PNSV Sempre Vivas National Park (Minas Gerais, Brazil), RBAE Águas Emendadas Biological Reserve (Distrito Federal, Brazil), RBIBGE IBGE Biological Reserve (Distrito Federal, Brazil), RECCPI Ecological Reserve of the Itororó Hunting and Fishing Club (Minas Gerais, Brazil), RPPNST Serra do Tombador Private Reserve of Natural Patrimony (Goiás, Brazil)

Chapter 13

Systematic Treatment

Diplusodon Pohl, Flora 10: 150. 14 Mar 1827. Type: Diplusodon virgatus Pohl. Friedlandia Cham. & Schltdl., Linnaea 2: 348. 1827. Type: Friedlandia amoena Cham. & Schltdl. Diplodon Sprengel, Gen. pl. 391. 1830. Cited in error by Lourteig (1989) as Diplocon Sprengel. Diplusodon Pohl sect. Palmatinerves Koehne, Engler Pflanzenr. 17 (IV. 216). 189. 1903. Lectotype: Diplusodon rotundifolius DC. (designated by Lourteig 1989: 211). Diplusodon Pohl sect. Palmatipenninerves Koehne, Engler Pflanzenr. 17 (IV. 216). 189. 1903. Lectotype: Diplusodon villosissimus Pohl (designated by Lourteig 1989: 211). Diplusodon Pohl sect. Penninerves Koehne, Engler Pflanzenr. 17 (IV. 216). 188. 1903. Lectotype: Diplusodon oblongus Pohl (designated by Lourteig 1989: 208). Diplusodon sect. Subuninerves Koehne, Engler Pflanzenr. 17 (IV. 216). 188. 1903. Lectotype: Diplusodon virgatus Pohl (designated by Lourteig 1989: 207). (= Diplusodon Pohl sect. Diplusodon). Subshrubs, shrubs, usually with a xylopodium, rarely small trees. Stems generally with exfoliating bark, upper branches cylindrical, subquadrangular to quadrangular, with an inconspicuous to conspicuous wing or wing absent, rust, yellow, wine-colored, whitish, brown, or greenish, glabrous or with diverse types  of trichomes, often with gall associations; colleters localized on the branches at the base of the leaves, bracts, and prophylls. Leaves opposite, decussate, sessile, or petiolate, blade pilose or glabrous, the margin entire, plane or revolute, sometimes fleshy, erose, ciliate, or with protuberances; pinnate veins hyphodromous, pinnate eucamptodromous to acrodromous with perfect base to acrodromous with supranumerary

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base, prominent or inconspicuous, frequently with domatia of the pocket type. Inflorescences frondose, frondose-bracteose, or bracteose, polytelic, of the botrya to pleiobotrya or diplothyrse to pleiothyrse types, with lateral cymes in triads or dichasia, or more rarely monotelic of the thyrsoid type. Bracts distinct from the vegetative leaves or similar to them in size and shape. Flowers actinomorphic, hexamerous, corolla in various tones of rose to red, rarely white, perigynous; floral tube funnelshaped, rarely oblong; rarely with glands under the petal insertion point; petals crinkled, obovate to elliptic, the apex obtuse or rarely acute; sepals triangular with inconspicuous or prominent papillae on the margin, epicalyx composed of six segments, varying widely in shape and position, or absent to inconspicuous; androecium diplostemonous or polystemonous to rarely haplostemonous, stamens 6–55, generally exserted, anthers dorsifixed, versatile, elliptic to reniform, yellow; pollen prolate to prolate-spheroidal, tricolporate, the colpus inconspicuous, elongated meridionally, arranged equatorially, equidistant, straight, ca. 35 μm long, relatively inconspicuous when compared to the pore which is prominent, and frequently obscured from sight by the thick, undulating sculptural elements, pore elongate, 5 × 10 μm, situated at the midpoint of the colpus, the inner margin entire, the annulus conspicuous, the outer margin lobate, wall thick (4–5 μm), verrucate, margins of the verrucae irregular and lobate, polar region with larger verrucae in some species; ovary glabrous or more rarely with trichomes at the apex, bicarpellate, bilocular with incomplete septa; placentation basal, in the fruiting stage thick and with two semi-lunar septa; style filiform, generally curved, exserted; stigma capitate, papillose. Fruit a septicidal capsule, the floral tube persistent, with a broad mouth or very rarely constricted at the mouth, capsule globose to oblong,  base obtuse to rarely acute, glabrous or more rarely with trichomes at the apex; seeds flat to lens-shaped, winged, wing small; embryo straight, cotyledons flat, orbicular or reniform, radicle scarcely elongated, epidermal trichomes of the seed coat straight. Pollen generally prolate to subprolate, rarely orbicular. 2n = 30. Etymology  Name of Greek origin, originating from the fusion of the words diplus (double) and odon (tooth), which refers to the calyx and epicalyx whorls of the flowers. Notes about Diplusodon nomenclature and typifications: 1. Koehne’s work at Berlin: Koehne’s holotypes at Berlin were destroyed in 1943 and required lectotypifications (Art. 9.2, 9.11 [Turland et al. 2018]). 2. Lourteig (1989) typified many Diplusodon names. In the majority of cases what she called a holotype was not the original specimen designated the holotype by the author or a single specimen studied by the author, but was a syntype. Such citations by Lourteig are here considered to have effected lectotypification.

13.  Systematic Treatment

93

3. Species described by Candolle from Martius collections: Candolle saw the Martius collections in Munich (ex. Prodromus 3: 94a, 94b, etc.) so the holotype resides in Munich. Fragments, presumably of the types, probably brought back to Geneva by Candolle are in the herbarium at Geneva (G). 4. Pohl’s collections: Pohl specimens may bear two numbers, a field number and a number from his diary (D) referring to the same specimen. In the typifications below, Pohl’s field number is followed by Pohl’s diary number. For example, Pohl 135  =  D 945; Pohl 589  =  D 940; Pohl 773, 1391, 1745  =  D 947; Pohl 990 = D 944; Pohl 1066 = D 937; Pohl 1365 = D 945; Pohl 1988 = D 941; Pohl 2018 = D 946; Pohl 2537 = D 939; and, Pohl 3553 = D 943. 5. The Brazilian collections of Pohl were stored in Vienna in a museum called “Brasilianisches Museum.” The first sheet of a collection got the original label, and the duplicates were stored without labels. After building the “Naturhistorische Museum,” the botanical collections kept in the “Brasilianisches Museum” were incorporated in the collection of Herbarium W. Unfortunately, some of the unlabeled duplicates were mixed up (Bruno Wallnöfer, pers. comm). 6. Duplicates of Pohl’s collections went to other herbaria without complete label information from his field diary (Bruno Wallnöfer, pers. comm). The majority of these duplicates have the collection locality, but the number is missing. These locations were places where Pohl spent only once in his collection itinerary in Brazil. For this reason, there are unnumbered duplicates of Pohl’s collections (Pohl s.n.) at BR, K, G, M, and OXF, which are here considered to be duplicates of the types. 7. Martius’s numbers: Martius “Observationes” (obs) numbers are derived from his handwritten fieldbooks. The “Herb. Fl. Bras. numbers” were been published in the journal Flora (Esser, pers. comm). 8. Some of Martius’s herbarium specimens were annotated by him with names that were never formally published. These names were mentioned as synonyms of valid species by Candolle (1828) and Koehne (1877); and are cited here as “pro syn.” (ICBN rec. 50A.1).

94

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13.1  Artificial Key to the Species of Diplusodon 1. Leaves 0.5–4 mm wide, linear, oblong-linear, narrowly oblong, narrowly elliptic, narrowly lanceolate, linear-lanceolate, elliptic-linear, linear-spathulate, narrowly triangular. 2.  Leaves with indumentum on adaxial surface, sometimes minute and inconspicuous. 3.  Epicalyx segments 0.5–2 mm long, not exceeding the length of the sepals. 4. Leaf base acute to attenuate, petiole 0.5–1.5 mm long. Epicalyx segments pubescent to hirtellous to sparsely sericeous, or with minute trichomes, inconspicuous, not ciliate. 5.  Leaves with minute trichomes, sometimes stellate. 6. Leaves hyphodromous. Epicalyx segments 1.5–2 mm; floral tube glabrous or almost glabrous; stamens 12. Bahia, Minas Gerais …………………………………………..…...………… 68. D. puberulus 6. Leaves eucamptodromous. Epicalyx segments 0.5–1 mm long; floral tube covered by inconspicuous stellate trichomes; stamens 15. Distrito Federal, Goiás ……………….....................................….….. 75. D. rosmarinifolius 5. Leaves with sericeous indumentum or finely pubescent or with sparse short trichomes, sometimes hirtellous, never stellate. 7. Leaf margin plane, lateral veins absent.................................. 33. D. gracilis 7. Leaf margin subrevolute to revolute, lateral veins 1–2 of each side of the midvein. 8. Floral tube 3.5–6 mm long, hirtellous, covered by sparse trichomes or trichomes appressed. Goiás. 9. Leaf blade 4–7 mm wide, apex acute …..........… 101. D. xerophilus 9. Leaf blade 1–3.5 mm wide, apex obtuse ............... 34. D. grahamae 8. Floral tube 2–2.5 mm long, covered by minute, inconspicuous trichomes. Bahia, Minas Gerais, Piauí, Tocantins .................................................................................... 87. D. thymifolius 4. Leaf base cordate, sessile. Epicalyx segments ciliate with sparse and spreading trichomes ……………………..................…......................……. 22. D. cordifolius 3.  Epicalyx segments 3–7 mm long, exceeding the apex of the sepals 2–3 mm. 10.  Plants from Goiás and Tocantins states. 11. Prophylls 2.5–4.2. Upper branches covered by dense indumentum with short trichomes, stellate, sometimes forming tufts or covered by finely pubescent, becoming glabrous in the upper parts. 12. Upper branches covered by dense indumentum with short trichomes, stellate, sometimes forming tufts. Leaf apex acute, margin revolute. Branches of the synflorescence not thin and delicate. 13. Leaf abaxial surface canescent. Floral tube 3.5–4 mm long, covered by stellate trichomes forming tufts. Epicalyx segments 2–4 mm long. Plant canescent, usually caespitose. ……………………....…………………....…….... 83. D. stellatus

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13. Leaf abaxial surface not canescent. Floral tube 4–4.5 mm long, covered by stellate trichomes not forming tufts. Epicalyx segments 4–6.5 mm long. Plant not canescent, not caespitose …………………....………....… 46. D. macrodon 12. Upper branches covered by finely pubescent, glabrous in the upper parts. Leaf apex obtuse, margin plane. Branches of the inflorescence thin and delicate ………..…...….. 33. D. gracilis 11. Prophylls 5–10. Upper branches hirtellous to pubescent, never looking stellate or glabrous. 14. Floral tube ca. 5 mm long, prophylls ca. 5.5 mm long. Subshrubs taller than 0.4, erect ……......…….....…..… 99. D. vittatus 14. Floral tube 7.5–8 mm long, prophylls 9–9.5 mm long. Subshrubs 0.2–0.4 m tall, prostate to suberect ……............................. ......................................................................……. 17. D. chapadensis 10.  Plants from Minas Gerais State. 15. Leaves 10–42 mm long, acrodromous or eucamptodromous, secondary veins 1–4 on each side of the midvein, base acute or attenuate, rarely obtuse, surface not warty. Stamens 12–18. 16. Pedicel 4–7 mm long, prophylls 5–7 mm long. Leaf secondary veins 1–2 on each side of the midvein, domatia present ………….............. …………………………...............………...……. 26. D. epilobioides 16. Pedicel 2.5–3.5 mm long, prophylls 10–11 mm long. Leaf secondary veins 3–4 on each side of the midvein, domatia absent ……............... ...............................…........................................…... 32. D. glocimarii 15. Leaves 3–6 mm long, hyphodromous, lateral veins absent, base retuse to cordate, surface warty. Stamens 6 ........................… ........... ......................................................................................93. D. verruculosus 2. Leaves glabrous on adaxial surface or on both surfaces (when trichomes present only on the midvein – D. uninervius, D. candollei, D. conduplicatus – or with minute, inconspicuous trichomes on the margin, D. praetermissus). 17.  Stamens 6. 18. Leaves with short dense indumentum on the abaxial surface, on the midvein only; prophylls with midvein pubescent on the adaxial surface; floral tube 3.2–3.5 mm long, with sparse trichomes; epicalyx segments 1.8–1.9 mm long, with cilia …….......................….…. 15. D. candollei 18. Leaves completely glabrous; prophylls glabrous; floral tube 4–6 mm long, glabrous; epicalyx segments 2–5 mm long, not ciliate. 19. Leaves margin revolute, glaucous ………….…….......……. 38. D. hexander 19.  Leaves margin plane, not glaucous …...........................……. 31. D. glaziovii

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17.  Stamens 12–20. 20.  Leaf margin strongly revolute. 21. Leaves with midvein densely pubescent on the abaxial surface; prophylls and epicalyx segments with sparse trichomes, sparsely ciliate, prophylls with margin subrevolute to revolute. Minas Gerais. …... 90. D. uninervius 21. Leaves completely glabrous; prophylls and epicalyx segments glabrous, prophylls with margin plane. Minas Gerais ….........….. 52. D. mononeuros 20.  Leaf margin plane. 22.  Plants 0.10–0.35 m tall, stem without ramification or caespitose. Botryum. 23. Leaves folded longitudinally in the midvein, keeled, midvein yellowish, long-ciliate, base retuse, amplexicaulous, margin dense long-ciliate …...............…….......…………….… 21. D. conduplicatus 23. Leaves expanded, not folded longitudinally in the midvein, not keeled, midvein green, not ciliate, base not amplexicaulous, margin glabrous, sometimes sparse ciliate. 24. Plant erect; flowers subsessile, pedicel 0–1 mm long; prophylls 4.5–5.5 mm, epicalyx segments linear-triangular, not ciliate. Goiás ………...................….....................….….. 27. D. ericoides 24. Plant caespitose; flowers pedicellate, pedicel 1.5–2.2 mm long; prophylls 6–6.5 mm, epicalyx segments lanceolate, slightly concave, prominently short-ciliate. Minas Gerais …...................... .................................................................... 67. D. psammophilus 22.  Plants 0.40–3 m tall, much-branched. Pleiobotryum. 25. Plants dark-brown when dry; pedicel 0–0.9 mm long …….................... .................................................................……… 65. D. praetermissus 25.  Plants not dark-brown when dry; pedicel 1.5–5 mm long. 26. Prophylls 6–9 mm long, epicalyx segments 4–4.5 mm long. Upper branches flexible, pending …......................…….. 91. D. vacillans 26.  Prophylls 2.5–5.5 mm long., epicalyx segments 0.8–3.5 mm long. Upper branches not flexible, not pending. 27. Secondary veins in the leaves 1 on each side of the midvein; floral tube 4.5–5 mm long, epicalyx segments 3–3.5 mm long. Minas Gerais .....….....................….. 14. D. canastrensis 27. Secondary veins in the leaves absent; prophylls 2.5–4, floral tube 3–4 mm, epicalyx segments 0.8–3 mm long. Goiás. 28. Leaves petiolate; petioles 1–2 mm long; blades 3–8 (−12) mm wide, slightly punctate, narrowly oblong, base acute to attenuate, apex obtuse …............. 33. D. gracilis 28. Leaves sessile; blades 0.5–0.9 mm wide, not punctate, linear, base obtuse, apex acute, minutely acuminate …......................…………………… 86. D. tenuifolius 1.  Leaves wider than 4 mm, if 3–5 mm never linear or narrow (3–7 times longer than wide). 29.  Epicalyx absent or present with reduced segments of 0.1–1.5 mm long. 30.  Plants completely covered by indumentum. 31. Branches with dense short indumentum, rust-colored; floral tube golden to orange-colored, densely covered by bulbous glandular trichomes, bottle shaped ……………………....…................................…………….... 25. D. divaricatus 31. Branches hirtellous, hirsute, velutinous, hispid, not rust-colored; floral tube not golden or orange-colored, not covered by glandular trichomes, trichomes not bottle shaped.

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32. Leaves strongly discolorous, adaxial surface dark-brown to black, abaxial surface whitish, covered by stellate trichomes forming congested tufts …….…..…................……...............….……... 36. D. helianthemifolius 32.  Leaves not strongly discolorous, stellate trichomes absent. 33.  Leaves silvery, indumentum sericeous. 34. Petiole 3–7 mm long; epicalyx segments equalling the sepals in length. Bahia. ………........................….….. 6. D. argyrophyllus 34. Petiole 0–3 mm long; epicalyx segments much shorter than the sepals. Goiás, Minas Gerais. 35. Leaves petiolate; petioles 1–3 mm long; blades 10–24 mm long, base acute to slightly attenuate, apex acute, margin plane. Floral tube 6–6.5 mm long, epicalyx segments 1–1.5 mm long, stamens 12. Minas Gerais ……..................... ……………….....................……………. 85. D. subsericeus 35. Leaves sessile; blades 4–6 mm long, base cordate, obtuse, margin strongly revolute. Floral tube ca. 4–4.5 mm long, epicalyx segments 0–0.5 mm long, stamens 18. Goiás …........ ..............................................................…. 49. D. micromerus 33. Leaves not silvery, indumentum hirsute, hirtellous, villous, pubescent, glabrescent. 36. Floral tube densely covered with white trichomes, hirsute, villous to sericeous. 37.  Plants from Minas Gerais and Bahia states. 38. Synflorescence frondose, congested, accessory branches absent. Petiole 0–6 mm long, blades 4–35, secondary veins 3–5 on each side of the midvein. 39. Plants not canescent, dark-brown or dark-green when dry. Prophylls 2.5–4 mm long reaching the middle or apex of the floral tube, stamens 12–15. 40. Leaves as long as wide, usually orbicular, 4–12 mm long, petioles to 1 mm long, base obtuse to subcordate. Bahia …………................ ....................................…….. 62. D. parvifolius 40. Leaves twice as long as wide, usually elliptic, 8–35 mm long, petioles 1–6 mm long, base acute to obtuse. Minas Gerais ……..................... .......................................…….... 39. D. hirsutus 39. Plants canescent, not dark-brown when dry. Prophylls 5–5.5 mm long, reaching the apex of the floral tube or sometimes surpassing it, stamens 6 ….. ..................................................... 51. D. minasensis 38. Synflorescence bracteose, lax, accessory branches usually present. Petiole 6–25 mm long, blades 20–90 × 10–60 mm, secondary veins 5–11 on each side of the midvein ….................................... 10. D. bradei 37. Plants from Distrito Federal, Goiás, Mato Grosso, and Rondônia (Brazil states) and Bolivia. 41.  Leaf base cordate. Prophylls 4–5.5 mm long.

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13  Systematic Treatment 42. Leaf blades 4–6 mm long, margin strongly revolute. Synflorescence frondose, sepals 2–2.5 mm long, not winish, epicalyx segments 0–0.5 mm long; stamens 18 ………….....................…….. 49. D. micromerus 42. Leaf blade 15–33 mm long, margin plane. Synflorescence bracteose, sepals winish even when dry, epicalyx segments 1.5–2.2 mm long, stamens 12–16 ..................................…… 97. D. vinaceus 41.  Leaf blade obtuse to acute. Prophylls 1.5–3.8 mm long. 43. Leaf blades 4–6 mm long, secondary veins 2–3 on each side of the midvein. Floral tube 2–3.5 mm long ………...............................……..…. 41. D. incanus 43. Leaf blade 10–55 mm long, secondary veins 3–6 on each side of the midvein. Floral 4–6 mm long. 44. Stems branched just at inflorescence, often caespitose. Bracts longitudinally folded; epicalyx segments 0.5–1.5 mm long …………............…….....……... 96. D. villosus 44. Stems branched, never caespitose. Bracts when distinguishable from leaves, not longitudinally folded; epicalyx segments 1–2 mm long. 45. Synflorescence bracteose, flowers subsessile, pedicel 0.5–1 mm long, stamens 12. Secondary veins 4–6 on each side of the midvein …...…....….....…. 9. D. bolivianus 45. Synflorescence frondose, flowers pedicellate, pedicel 1.5–2.2 mm long, stamens 18. Secondary veins 3 on each side of the midvein ………… 13. D. caesariatus 36. Floral tube covered by sparse trichomes or pubescent, not hirsute or sericeous. 46.  Prophylls 1.5–3.7 mm long. Goiás. 47.  Leaves 3–10 mm long; stamens 12. 48. Epicalyx segments deflexed …...….... 41. D. incanus 48. Epicalyx segments suberect …....... 66. D. prostratus 47.  Leaves 10–45 mm long; stamens 18. 49. Leaves elliptic to ovate-elliptic, apex obtuse, 6–13 mm wide; pedicel 1.5–2.2 mm long …….......... ..........................................….….. 13. D. caesariatus 49. Leaves narrowly elliptic, apex acute, 4–7 mm wide; pedicel 0–0.5 mm long ..........….. 101. D. xerophilus 46.  Prophylls 4.5–7 mm long. Minas Gerais. 50. Plant not dark-brown when dry; leaves 10–30, apex obtuse; petiole 3–7 mm ……......…. 2. D. aggregatifolius 50. Plant dark-brown when dry; leaves 5–7, apex acute rarely obtuse, petiole 1–1.5 mm long …...... 50. D. microphyllus

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30. Plants glabrous or apparently glabrous with minute and inconspicuous trichomes, sometimes just on the flowers. 51. Inflorescences bracteose, flowering region very distinct from the vegetative region, with significant reduction in shape and/or size of the bracts compared to leaves. 52. Leaf base acute to attenuate; venation hyphodromous to eucamptodromous. Minas Gerais …................................................……….……..… 94. D. vidalii 52.  Leaf base obtuse, venation acrodromous-basal type. Goiás. 53. Synflorescence without terminal flower in the main axis and without cymes in the lateral branches ….............. 69a. D. punctatus var. punctatus 53. Synflorescence with presence of terminal flower in the main axis and cymes in the lateral branches …….....………..….….. 60. D. panniculatus 51. Inflorescences frondose-bracteose or frondose, gradual reduction in shape and/or size of the leaves compared to bracts or lacking any significant reduction in shape and/or size. 54. Pedicel 7–47 mm long. Marginal leaf vein wine-colored to purple ................... ............................................................................................... 47. D. marginatus 54. Pedicel 0.5–6.5 mm long; marginal leaf vein not conspicuously colored, green. 55. Stamens 30–32; pedicel 7–9.5 mm long. Tocantins .......... 88. D. trigintus 55.  Stamens 12–18; pedicel 2–6.5 mm long. 56. Leaves acrodromous-basal-supranumerary, base obtuse, cordate or subcordate to amplexicaul. 57. Leaves 25–65 mm long; corolla 3–4 cm diam. Goiás, Tocantins ......................................................................... 40. D. imbricatus 57. Leaves 8–20 mm long; corolla ca. 2.5 cm diam. South of Minas Gerais …………..................................….….. 12. D. buxifolius 56. Leaves hyphodromous or eucamptodromous, base acute to slightly attenuate. 58. Plant not dark-brown when dry; prophylls 2.5–3 × 0.8–1 mm, unkeeled; floral tube 4–4.5 mm long …............ 53. D. myrsinites 58. Plant dark-brown when dry; prophylls 4–6 × 2.5–3 mm, keeled; floral tube 5–6.5 mm long ….............................. 56. D. oblongus 29.  Epicalyx present, conspicuous, segments 2–11 mm long. 59. Plants completely glabrous to glabrescent or apparently glabrous with cilia at the margins of bracts, prophylls and sepals or sparse trichomes on the leaves or covered by minute and/or inconspicuous trichomes just on the branches, pedicel or floral tube. 60.  Flowers long-pedicellate, pedicel 7–47 mm long. 61.  Leaves 8-15 cm long. Flowers with 12 stamens. Minas Gerais. 62. Leaves orbicular-reniform to oval-depressed; floral tube 4–5 mm long ………..…...………..............…........…..................….. 30. D. glaucescens 62. Leaves oval-lanceolate; floral tube 5.5–6 mm long ………........................ ..................................................................................... 77. D. rotundifolius 61. Leaves 25-70 cm long. Flowers with (13–)18–33 stamens. Goiás, Minas Gerais, Tocantins. 63. Prophylls 13–25 x 4–15 mm, sometimes hiding the floral tube. Inflorescences without terminal flower, lateral cymes absent. 64. Leaves with domatia, not glaucous, elliptic-lanceolate, base acute to obtuse; epicalyx segments 6–8 mm long; corolla ca. 4.5 cm diam. Goiás, Tocantins …................................................ 23. D. cryptanthus

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13  Systematic Treatment 64. Leaves without domatia, glaucous, ovate, base retuse, sometimes slightly amplexicaul; epicalyx segments 2.5–3 mm long; corolla 5–6 cm diam. Goiás ...............................................… 45. D. longipes 63. Prophylls 2–2.8 x 1–1.8 mm, not hiding the floral tube. Inflorescences with terminal flower, lateral cymes present. 65. Prophylls, floral tube, sepals and epicalyx segments with inconspicuous pubescence. Triplothyrsoid to pleiothyrsoid. Prophylls 2–2.8 mm long, not even reaching the base of the floral tube. Floral tube 4–6 mm long. Goiás ……......................... ............................………………………….. 60. D. panniculatus 65. Prophylls, floral tube sepals and epicalyx segments glabrous. Thyrsoid or thyrse. Prophylls 5.5–8 mm long, reaching the apex of floral tube. Floral tube 6–6.5 mm long. Goiás, Mato Grosso, Mato Grosso do Sul, Minas Gerais ..….................. 59. D. ovatus 60.  Flowers sessile, subsessile, never long-pedicellate, pedicel 0–6.5 mm long. 66. Leaves eucamptodromous (veins departing along the midvein) or hyphodromous (just the midvein evident, but if veins visible, acrodromous). 67. Inflorescences bracteose, flowering region very distinct from the vegetative region, with significant reduction in shape and/or size of the leaves compared to bracts. 68.  Prophylls reaching the middle of the floral tube or below the middle. 69.  Stamens 12; leaves 20–45 mm long. 70. Petiole 5–10 mm long; pedicel 3–5 mm long, floral tube 6–6.5 mm long, epicalyx segments 3–3.5 mm long ……… …..................................................…………. 55. D. nitidus 70. Petiole 1–1.5 mm long; pedicel 0.5–1.5 mm long, floral tube 4–4.5 mm long, epicalyx segments 0.8–1.5 mm long ……... ......................................................…………. 94. D. vidalii 69.  Stamens 18–44. Leaves 40–100 mm long. 71. Leaves glaucous, base attenuate; floral tube 6–10 mm long; stamens 26–55; upper branches wine- to rust-colored …...............………42. D. kielmeyeroides 71. Leaves not glaucous, base obtuse; floral tube 4.5–5 mm long; stamens 18; upper branches not wine- or rust-colored ………….......……… 72. D. ramosissimus 68. Prophylls surpassing the middle reaching the apex of the floral tube reaching the base of sepals. 72.  Floral tube 5–7 mm long, stamens 18–24 ............ 92. D. venosus 72. Floral tube 4.8–5 mm long, stamens 12. 73. Leaves 11–35 mm long, veins and upper branches rust to wine-colored; prophylls 3–4 mm long, reaching the middle or the apex of the floral tube. Bahia. …...… 19. D. ciliatus 73. Leaves 40–95 long, veins and upper branches straw-­colored; prophylls 6–10 mm long, surpassing the apex of the floral tube. Goiás …................................................. 7. D. astictus 67. Inflorescences frondose-bracteose or frondose, gradual reduction in shape and/or size of the leaves compared to bracts or lacking any significant reduction in shape and/or size. 74.  Stamens 26–55; corolla 6.5–7 cm in diam.; floral tube fleshy.

Artificial Key to the Species of Diplusodon

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75. Leaves base attenuate; branches unwinged or inconspicuously winged. …............................................… 42. D. kielmeyeroides 75. Leaves base retuse to obtuse-subcordate; branches markedly winged, wings 1.5–2 mm wide .............................… 3. D. alatus 74.  Stamens 12–24. Corolla 1.5–4.5 cm in diam.; floral tube not fleshy. 76. Corolla white. Inflorescence frequently with accessory branches …….……………......................................….... 98. D. virgatus 76. Corolla rose, red, pink, purple, or lilac, never white. Inflorescence without accessory branches. 77.  Plants dark-brown when dry. 78. Epicalyx segments foliaceous, hyaline, 2 mm wide, obovate to lanceolate; ovary glabrous or minutely pilose on the apex …......................……. 4. D. appendiculosus 78. Epicalyx segments cylindrical to compressed, not hyaline, up to 1 mm wide, linear; ovary completely glabrous. 79. Epicalyx segments 1–2 mm long. Leaf base obtuse. South of Minas Gerais ….......…... 12. D. buxifolius 79. Epicalyx segments 2–5 mm long. Leaf base acute to attenuate. Bahia; Distrito Federal; Goiás; central, west, and north of Minas Gerais; Mato Grosso; Tocantins. 80. Flowers sessile to subsessile, pedicel 0–1 mm long; prophylls keeled. 81. Leaves elliptic, ovate to ovate-­lanceolate, petiole 2–9 mm ….…....... 56. D. oblongus 81. Leaves narrowly oblong, petiole 0.5–1.5 mm …...........65. D. praetermissus 80. Flowers pedicellate, pedicel 2–5 mm long; prophylls unkeeled. 82. Bracts 3–4 mm wide. Leaves hyphodromous, petiole 1–2 mm long ….... ............................................ 33. D. gracilis 82. Bracts 6–12 mm wide. Leaves eucamptodromous, petiole 3–10 mm long. 83. Prophylls 1–1.5 mm, spathulate, reaching the middle or the apex of the floral tube ….....….... 19. D. ciliatus 83. Prophylls ca. 0.8 mm, linearlanceolate, reaching to the base of the floral tube ….......…... 55. D. nitidus 77.  Plants not dark-brown when dry. 84.  Leaves hyphodromous, if veins visible, acrodromous.

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13  Systematic Treatment 85. Prophylls 7–13 mm long, surpassing the apex of the floral tube; floral tube 6–7 mm long; leaves oval, oval-lanceolate to elliptic-lanceolate, base obtuse to cordate. Goiás ….....….. 11. D. burchellii 85. Prophylls 2.5–3 mm long, reaching the middle of the floral tube; floral tube 4–4.5 mm long; leaves obovate, elliptic to narrowly oblong, leaves acute to slightly attenuate. Minas Gerais ………..............… .................................……………. 53. D. myrsinites 84.  Leaves eucamptodromous. 86. Petiole 5–10 mm long; prophylls 2–3 mm long, reaching to the base of the floral tube; pedicel 3–5 mm long …….........................… 55. D. nitidus 86. Petiole 0–5 mm long. Prophylls 3.5–9 mm long, reaching the middle of the floral tube to surpassing it; pedicel 0.4–3.5 mm long. 87. Prophylls 5–9 mm long; petiole up to 1 mm long. Minas Gerais ................. 20. D. ciliiflorus 87. Prophylls 3–4.5 mm long. Petiole 1–5 mm long. Bahia. 88. Leaves with ciliate margin and prominent lateral veins on abaxial surface; epicalyx segments 2.5–4 .. 19. D. ciliatus 88. Leaves with plane and not ciliated margin; lateral veins inconspicuous on abaxial surface; epicalyx segments 4–5; inflorescence without cymes of triads …... .…….................................….... 89. D. ulei 66. Leaves acrodromous-basal-typical (all veins departing from the base of the midvein) or acrodromous supranumerary (1–4 veins departing from the base of the midvein and 1–3 veins departing along the midvein), sometimes lateral veins very inconspicuous. 89.  Stamens 19–44; epicalyx segments 5–10 mm long. 90. Bracts, prophylls, sepals and epicalyx segments bearing long cilia on the margin; leaf veins rust-colored …….........….. 18. D. ciliatiflorus 90. Bracts, prophylls, sepals and epicalyx segments completely glabrous, not long-ciliate; leaf veins not rust-colored. 91. Plants dark-brown when dry; prophylls 10–12 mm long, surpassing the apex of the floral tube ca. 5 mm; epicalyx segments 8–10 mm long; leaves acrodromous-basal-typical, blade with dark punctae over the whole surface ………............... .................................................……................… 81. D. sordidus 91. Plants not dark-brown when dry; prophylls 4–9 mm long, reaching the apex of the floral tube or slightly below; epicalyx segments 1.8–6 mm long; leaves acrodromous-basalsupranumerary, blade without dark punctae. 92. Veins 4–6 on each side, domatia absent; epicalyx segments 5–6 mm long; synflorescence frondose-­bracteose …….…… ………................................................…... 82. D. speciosus

Artificial Key to the Species of Diplusodon

103

92. Veins 2–3 along the midvein, domatia present; epicalyx segments of 1.8–2 mm long; synflorescence bracteose ……. ...............................................................….. 92. D. venosus 89.  Stamens 6–18; epicalyx segments 2–5 mm long. 93. Inflorescence bracteose, flowering region very distinct from the vegetative region, with significant reduction in shape and/or size of the leaves compared to bracts. 94. Prophylls not reaching the middle of the floral tube; petiole 3–7 mm long; leaves with conspicuous domatia …..……............ ...............................................................….. 72. D. ramosissimus 94. Prophylls almost reaching to surpassing the apex of the floral tube; petiole 0–3 mm long mm; leaves without domatia or rarely with faint domatia. 95.  Epicalyx segments 3–4 mm long, stamens 12; branches of the inflorescence thin and delicate, bracts 6–8 mm long ….......................……. 69b. D. punctatus var. dentatus 95. Epicalyx segments 1.8–2 mm long, stamens 18–24; branches of the inflorescence not thin or delicate, bracts 15–30 mm long …......................................... 92. D. venosus 93. Inflorescence frondose-bracteose or frondose, gradual reduction in shape and/or size of the leaves compared to bracts, or lacking any significant reduction in shape and/or size. 96. Inflorescence with a terminal flower (caducous), cymes present; leaves 30–95 mm long …….............................…. 59. D. ovatus 96. Inflorescence without a terminal flower, cymes absent; leaves 5–30 mm long. 97. Leaves wider than longer, triangular-deltoid, oval-depressed to obcordiform. 98. Pedicel 0.5–1.5 mm long; floral tube 6.5–8 mm long, dark-brown in fruit. Goiás ……...... 24. D. decussatus 98. Pedicel 2–7 mm long; floral tube 4–5 mm long, pale beige in fruit. Minas Gerais. 99. Stamens 6; pedicel 2–5 mm long; leaves 8–22 mm long ….................................….. 58. D. orbicularis 99. Stamens 12; pedicel 5–7 mm long; leaves 5–10 mm long ….............................….. 30. D. glaucescens 97. Leaves longer than wider, ovate, oval-elliptic, ovallanceolate to elliptic-lanceolate, 100. Plants not dark-brown when dry. Leaves strongly imbricate, hiding completely the stem …..........…........ ...............................………………..... 11. D. burchellii 100. Plants dark-brown when dry; leaves not imbricate, not hiding the stem. 101. Sepals and epicalyx segments long-ciliate; epicalyx segments ca. 5.5 mm long. Goiás …..... ..................................... 104. D. thysanosepalus 101. Sepals and epicalyx segments not ciliate; epicalyx segments 3–4 mm long. Bahia, Minas Gerais …..................…. 71. D. quintuplinervius 59.  Plants with indumentum on all parts, even though minutely pubescent.

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13  Systematic Treatment 102.  Epicalyx segments 2–3.5 mm long. 103. Plants covered by pubescent to hirsute indumentum, never silvery-sericeous. 104. Plants from Distrito Federal, Goiás, Mato Grosso, Rondônia (Brazil), and Bolivia. 105.  Prophylls 8.5–10 mm long; floral tube 7.5–8 mm long. 106. Leaves 3–5 mm wide, apex acute, secondary veins 2 on each side …..............….…… 17. D. chapadensis 106. Leaves 8–45 mm wide, apex obtuse secondary veins 3–10 on each side. 107. Leaves not dark-brown when dry; domatia absent. Stamens 18–32. Leaf abaxial surface indumentum with long trichomes with a bulbous to thickened base. 108. Sepals sericeous, densely covered by white, not secretory trichomes. Leaf secondary veins 8–10 on each side of the midvein. Shrubs, stem branched ….......... ........................... 74. D. retroimbricatus 108. Sepals golden in color, densely covered by orange, secretory trichomes bottle shaped. Leaf secondary veins 4–6 on each side of the midvein. Subshrubs, single stem, sometimes caespitose …...... ...........................…… 79. D. sessiliflorus 107. Leaves dark-brown when dry; domatia present. Stamens 12. Leaf abaxial surface indumentum with short trichomes sometimes mixed with tufts of stellate trichomes ……........................ ….......................................... 54. D. nigricans 105.  Prophylls 2.2–8 mm long; floral tube 2–6 mm long. 109. Sepals golden in color, densely covered by orange, secretory trichomes bottle shaped; stamens 24–32. Prophylls 5–10 mm wide. …....................................... ...............….........................……. 79. D. sessiliflorus 109. Sepals not golden in color, not covered by orange, secretory trichomes bottle shaped; stamens 12–18. Prophylls 1–4.5 mm wide. 110. Petioles 6–15 mm long. Flowers much concentrated at the apex of the branches, sometimes forming glomerules; synflorescence broad, bracts differentiated ……....................... ................................................ 63. D. petiolatus 110. Petioles 0–6 mm long. Flowers arranged along the branches, never forming glomerules; synflorescence not broad, bracts not differentiated. 111. Leaf base acute to attenuate, eucamptodromous.

Artificial Key to the Species of Diplusodon

105 112. Prophylls 2.2–3.8 mm long, reaching the middle or slightly beyond the middle of the floral tube. 113. Leaves 4–10 mm long, secondary veins 2–3 on each side of the midvein; floral tube 2–3.5 mm long, epicalyx segments cylindrical, strongly deflexed; stamens 18. Goiás and southwestern Bahia ……..... 41. D. incanus 113. Leaves 20–40 mm long, secondary veins 4–6 on each side of the midvein, floral tube 4–6 epicalyx segments triangular, deflexed to spreading; stamens 12. Distrito Federal, Goiás, Mato Grosso, Rondônia (Brazil) and Bolivia …........................ ........………. 9. D. bolivianus 112. Prophylls 5.2–8 mm long, reaching the apex or surpassing the apex of the floral tube. 114. Leaves 8–20 mm wide, as long as wide, dark-brown when dry, apex obtuse, margin plane, secondary veins 3–6. Stamens 12, epicalyx segments 2–3 mm long. Prophylls margin hyaline …............................... .................... 54. D. nigricans 114. Leaves 1.5–6 mm wide, 4–5 times longer than wide, not dark-brown when dry, apex acute, margin revolute, secondary veins 2. Stamens 18, epicalyx segments 3–3.5 mm long. Prophylls margin not hyaline …............ ........................99. D. vittatus 111. Leaf base obtuse to cordate, acrodromous. 115. Leaves dark-brown when dry, strongly discolorous, abaxial surface white due to dense short indumentum, sometimes stellate.

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13  Systematic Treatment 116. Leaf adaxial surface slightly verrucose or minutely pilose, abaxial surface with stellate trichomes forming congested tufts, secondary veins 2–3 on each side of the midvein, not parallel. Pedicel absent, prophylls ca. 2.5 mm long, unkeeled, epicalyx segments 1.5–2 mm long ........…. 36b. D. helianthemifolius var. pemphoides 116. Leaf adaxial surface glabrous, nitid, abaxial surface with dense indumentum, not stellate forming tufts, secondary veins 4–5 on each side of the midvein, almost parallel. Pedicel 1–1.5 mm long, prophylls 4–4.5 mm long, keeled, epicalyx segments 2–2.5 mm long ….................. .................... 64. D. plumbeus 115. Leaves not dark-brown when dry, not strongly discolorous, abaxial surface without stellate trichomes. 117. Epicalyx segments longer than the sepals, sometimes the same size. 118. Epicalyx segments with long trichomes spreading. Synflorescence frondose. Leaves 6–17 × 4–9 mm, abaxial surface hirsute, trichomes long, secondary veins 3 on each side of the midvein, domatia evident ......................... .......... 37. D. heringeri

Artificial Key to the Species of Diplusodon

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118. Epicalyx segments with short trichomes, not spreading. Synflorescence bracteose. Leaves 30–50 × 15–25 mm, abaxial surface with indumentum dense, trichomes very short, secondary veins 3–5 on each side of the midvein, domatia absent ……................... ........... 80. D. sigillatus 117. Epicalyx segments shorter than the sepals. 119. Leaves 6–10 mm long. Prophylls 5–6 mm long. Synflorescence frondose, pedicel 1.5–2.5 mm long ……................................................ 28. D. fastigiatus 119. Leaves 15–55 mm long. Prophylls 2–5 mm long. Synflorescence bracteose to frondose-bracteose, pedicel 0.5–1 mm long. 120. Petiole absent, leaf blade ovate, base cordate. Flowers congested, stamens 12–16 …………..... ...............................................…97. D. vinaceus 120. Petiole 1–2.5 mm long, leaf blade ellipticlanceolate to less often ovate, base obtuse. Flowers lax, stamens 18. 121. Subshrubs branched just at inflorescence, often caespitose; bracts longitudinally folded. Prophylls 2–3 × 1–2.5 mm, epicalyx segments 0.5–1.5 mm long …..… ......................................... 96. D. villosus 121. Subshrubs branched, never caespitose; bracts not longitudinally folded. Prophylls 4–5 mm long, epicalyx segments 2–2.5 mm long............................................ ...................................... 16. D. capitalensis 104.  Plants from Minas Gerais and Bahia states. 122. Sepals golden in color, densely covered by orange, secretory trichomes bottle shaped. Stamens 24–32. Prophylls 5–10 mm wide ............…… 79. D. sessiliflorus 122. Sepals not golden in color, not covered by orange, secretory trichomes bottle shaped. Stamens 10–18. Prophylls 1–5 mm wide. 123. Leaf secondary veins 5–11 on each side of the midvein. Synflorescence bracteose, accessory branches usually present. Leaves 20–90 mm long ……… ………………................................................................….…….. 10. D. bradei 123. Leaf secondary veins 0–5 on each side of the midvein. Synflorescence frondose to frondose-bracteose, accessory branches never present. Leaves 4–30 mm long. 124.  Species from Minas Gerais.

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13  Systematic Treatment 125. Leaf abaxial surface and upper branches with stellate trichomes, or stellate trichomes mixed with simple trichomes, sometimes forming congested tufts .................…… 36. D. helianthemifolius 125. Leaf abaxial surface and upper branches without stellate trichomes. 126.  Prophylls 2–5 mm long and leaf base acute to attenuate. 127. Leaves dark-brown when dry, hirsute or with long and sparse trichomes. 128. Upper branches and leaves densely pilose to hirtellous, domatia present. Prophylls 2.5–4 mm long, floral tube 2–3 mm long ..... 39. D. hirsutus 128. Upper branches and leaves with long, sparse trichomes, domatia absent. Prophylls 4.5–6 mm long, floral tube 4–5 mm long …......................... ....................................…. 50. D. microphyllus 127.  Leaves not dark-brown when dry, pubescent. 129. Epicalyx segments 3.1–3.5 mm long, stamens 18. Petioles ca. 1 mm long, leaf secondary veins 1–2 on each side ….............… 103. D. capitatus 129. Epicalyx segments 1.5–2.5 mm long, stamens 10–12. Petioles 3–7 mm long, leaf secondary veins 2 or 3 on each side ……............................. .....................................…. 2. D. aggregatifolius 126. Prophylls 5.5–10 mm long and leaf base obtuse, retuse to cordate. 130. Leaves hyphodromous, oblong. Subshrubs white to silvery, unbranched (sometimes branched just at inflorescence) ….........…………......... 78. D. saxatilis 130. Leaves eucamptodromous to acrodromous, oval, suborbicular, rhombic, ovate to elliptic. Subshrubs never white or silvery, branched. 131. Epicalyx segments 3.5–5 mm long, cylindrical, thick, longer than the sepals. Prophylls 8–10 mm long, spathulate …....…….… 57. D. obtectus 131. Epicalyx segments 2–2.8 mm long, narrow- to wide-lanceolate, shorter to slightly longer than the sepals. Prophylls 6–8 mm long, elliptic to rhombic. 132. Secondary veins 3–4 on each side of midvein, straight, almost parallel Epicalyx segments narrowly lanceolate …....… 73. D. rectinervis 132. Secondary veins 4–5 on each side, curved, not parallel Epicalyx segments wide-­lanceolate ……........................ 100. D. vividus

Artificial Key to the Species of Diplusodon

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124.  Species from Bahia or northeastern portion of the State of Goiás. 133. Leaf base obtuse to subcordate, secondary veins 3–5. Epicalyx segments erect, spreading or slightly deflexed. 134. Leaves 10–22 mm long, petiolate, petiole 1–3 mm long, abaxial surface not white, domatia present. 135. Prophylls 6–6.5 mm long, floral tube 5–5.5 mm long, epicalyx segments lanceolate ….…...... 8. D. bahiensis 135. Prophylls ca. 3.5 mm long, floral tube ca. 3.5 mm long, epicalyx segments sword-shape ……................... ............................……….…........…. 102. D. xiphodon 134. Leaves 4–12 mm long, subsessile, petioles 0–1 mm long, abaxial surface white, domatia absent ………........................ ................…………..........................…… 62. D. parvifolius 133. Leaf base acute to attenuate, secondary veins 2–3. Epicalyx segments strongly deflexed …………………...… 41. D. incanus 103.  Plants covered by dense, silvery-sericeous indumentum. 136. Segments of epicalyx 1–2 mm long; leaves not imbricate, showing the stem, petiole 3–7 mm long. Bahia ……................................................…...... 6. D. argyrophyllus 136. Segments of epicalyx 3–5.5 mm long; leaves strongly imbricate, not showing the stem, petiole absent. Goiás. 137. Stamens 15–18; leaves 8–12 mm wide; lateral leaf veins 4–5 ……………… ……………......................................................................... 1. D. adpressipilus 137. Stamens 12; leaves 4–6 mm wide; lateral leaf veins 2–3 ……………...…….... …………...................................................................…. 5. D. argenteus 102. Epicalyx segments 4–7 mm long (if 3.5 mm long, then floral tube and sepals covered by bulbous-based secretory trichomes). 138.  Epicalyx segments longer than the sepals. 139.  Leaves covered by silvery, sericeous indumentum, erect, imbricate. 140. Leaves sessile, with base obtuse; stamens 15–18; prophylls 7–8.1 mm long, floral tube 6–7 mm long; leaves ovate to oval-lanceolate, veins 4 or 5 on each side, 2 or 3 departing from the base of the midvein. Goiás ……......................................................................... 1. D. adpressipilus 140. Leaves petiolate, with base acute; petiole 2.5–3.5 mm long; stamens 12; prophylls 10–11 mm long, floral tube 8–9 mm long; leaves elliptic to oval-elliptic, veins 3–4 on each side of the midvein, all departing from the base of the midvein. Minas Gerais, São Paulo .................................... ...................................................................................... 32. D. glocimarii 139. Leaves not covered by silvery indumentum, indumentum not sericeous, not imbricate, spreading to suberect. 141.  Plants from Goiás and Tocantins states. 142.  Prophylls 7–11 mm long; floral tube 5.5–9 mm long. 143. Floral tube 6–9 mm long, dense indumentum with long trichomes, usually whitish. Accessory branches never present. 144. Epicalyx segments cylindrical, never foliaceous; stamens 10–18. Leaf secondary veins 3–5 on each side of the midvein.

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13  Systematic Treatment 145. Plants dark-brown or dark-green when dry; leaves petiolate, petioles 2–5 mm long. Prophylls keeled. Floral tube white due to dense indumentum, highlighted from the dark-brown color of the dry plant ……................................... .........................………. 44. D. leucocalycinus 145. Plants not dark-brown or dark-green when dry; leaves sessile. Prophylls unkeeled. Floral tube not highlighted from the color of the dry plant. 146. Epicalyx segments 7–10 mm long; floral tube densely covered by long trichomes, not bulbous, not secretory; stamens 16–18; prophylls surpassing the apex of the floral tube. Leaves ovate-lanceolate, apex obtuse, sometimes acuminate …............................. ...................................... 29. D. floribundus 146. Epicalyx segments 3.5–4.5 mm long; floral tube densely covered by secretory, bulbous-based trichomes; stamens 10–15; prophylls reaching the apex of the floral tube. Leaves oblong-lanceolate, apex acute …….................…….. 35. D. hatschbachii 144. Epicalyx segments broadly triangular with subrevolute margin, sometimes foliaceous; stamens 24–32. Leaf secondary veins 4–10 on each side of the midvein ………….................…………… 61. D. paraisoensis 143. Floral tube 4–6 mm long, covered by dense pubescent indumentum with short trichomes, not whitish. Accessory branches of the first order sometimes present ….................... ................................................................. 43. D. lanceolatus 142.  Prophylls 3–5.5 mm long; floral tube 4–5.5 mm long. 147. Plants covered by stellate trichomes forming tufts; leaves narrow, 5–10 times longer than wide. Prophylls 1–1.1 mm wide; stamens 12 ……………..........…...….. 83. D. stellatus 147. Plants never covered by stellate trichomes; leaves 0.5–2 times longer than wide. Prophylls 3–1.1 mm wide; stamens 17–26. 148. Petiole 0.5–2 mm long. Bracts margin revolute. Prophylls 3–5.5 mm long. 149. Leaves canescent, covered by long, white trichomes, base obtuse, secondary veins 3 on each side of the midvein, domatia evident. Epicalyx segments with long and sparse trichomes spreading …….....…. 37. D. heringeri

Artificial Key to the Species of Diplusodon

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149. Leaves olive-green, not canescent, covered by dense pubescent indument, short trichomes, base subcordate to cordate, secondary veins 3–5 on each side of the midvein, domatia absent. Epicalyx segments densely covered by short, thick trichomes ….................…. 80. D. sigillatus 148. Petiole 3–5 mm long. Bracts margin plane. Prophylls usually 6–8 mm long ….....….....… 43. D. lanceolatus 141. Plants from Maranhão, Mato Grosso, Minas Gerais, and São Paulo states. 150. Plants looking glabrous, with inconspicuous indumentum or minutely pubescent covered with short or minute trichomes. Maranhão and Northern Minas Gerais. 151. Prophylls 8–10 mm long; pedicel 3.5–6 mm long; epicalyx segments thick .............................................. 57. D. obtectus 151. Prophylls 4–4.6 mm long, pedicel 2 mm long epicalyx segments fine. …….…..................................... 76. D. rotatus 150. Plants hairy, with dense indumentum, conspicuous villous to hirsute, dense pubescent, trichomes not minute. São Paulo, Mato Grosso, southern Minas Gerais. 152. Plants covered by white indumentum, villous. Floral tube 6.5–9 mm long. Leaves ovate to ovate-lanceolate apex obtuse. 153. Prophylls 16–17 × 7–8 mm; floral tube 8–9 mm long; epicalyx segments 4.5–6 mm long ……….................... .......................................…..… 48. D. mattogrossensis 153. Prophylls 6–9 × 3–4 mm; floral tube 6.5–7.5 mm long; epicalyx segments 8.5–9 mm long …............................ .......................................……...… 95. D. villosissimus 152. Plants covered by olive-green indumentum, dense pubescent to hirtellous. Floral tube 4.5–6 mm long. Leaves elliptic to elliptic-lanceolate, apex acute ….........… 43. D. lanceolatus 138.  Epicalyx segments equal to or shorter than sepals. 154. Small subshrubs 3–11 cm tall; stamens 12; prophylls reaching the middle or the apex of the floral tube ………......................................... 70. D. pygmaeus 154. Subshrubs taller than 15 cm; stamens 15–34; prophylls reaching or surpassing the apex of the floral tube. 155. Leaves eucamptodromous; petiole 3–5 mm long; floral tube pubescent ……....................................................................……. 43. D. lanceolatus 155. Leaves acrodromous-basal-supranumerary; petiole 0–3 mm long; floral tube densely pilose, hirsute, silver-sericeous and/or with bulbous-based trichomes or thick trichomes.

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13  Systematic Treatment 156. Prophylls as long as wide, ovate to orbicular, 4.5–10 mm wide, usually hiding the floral tube; sepals and epicalyx segments covered by bulbous-based trichomes; leaf adaxial and abaxial surface rough, with a verrucose aspect due to the bulbous base of the trichomes, trichomes sparse and equidistant ………................. ..........................................................…...…...…. 84. D. strigosus 156. Prophylls longer than wide, never ovate or orbicular, 2.5–4.5 mm wide, not hiding the floral tube; sepals and epicalyx segments covered by trichomes without bulbous-based; leaf adaxial and abaxial surface not rough, indumentum dense. 157. Leaves imbricate, sessile, blades and floral tube covered by dense silver-sericeous indumentum …................................... ..................................................……… 1. D. adpressipilus 157. Leaves not imbricate, petiolate, petiole 0.5–3 mm long, blades and floral tube not covered by silver-sericeous indumentum. 158. Leaves 25–90 mm long, veins 4–10 on each side along the midvein; prophylls 9.5–11 mm long …................... ...................................…………… 61. D. paraisoensis 158. Leaves 6–25 mm long, veins 3–4 on each side along the midvein; prophylls 3–5.5 mm long. 159. Leaves covered by dense indumentum, trichomes very short; prophylls with 3 conspicuous veins, reddish ........................…….. .........................................…..... 80. D. sigillatus 159. Leaves hirsute, covered by long trichomes; prophylls without conspicuous veins ………...... ..............................................…. 37. D. heringeri

Descriptions

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1. Diplusodon adpressipilus Lourteig, Bradea 5(19): 236. 1989. Type: Brazil. Goiás: Alto Paraíso de Goiás, Chapada dos Veadeiros, 19 Jul 1964 (fl), G.  T. Prance & N. T. Silva 58242 (holotype, US [00289079]; isotypes, K, NY, U, UB). Figs. 13.1 F–P, 13.5, 15.1 E Subshrubs to 1 m tall, erect, with a xylopodium, branched just at the inflorescence, not dark-brown when dry. Upper branches cylindrical, unwinged, sericeous, covered by white, appressed trichomes, internodes 0.4–1.2 cm long. Leaves decussate, imbricate, erect, sessile; blades 15–25 (−48) × 8–12(−20) mm, coriaceous, not discolorous, acrodromous, supranumerary at the base, ovate to ovate-lanceolate, base obtuse, apex acute, slightly acuminate, margin plane, blades covered by dense silver-sericeous indumentum, sometimes hirsute or caducous, trichomes appressed; secondary veins 4 or 5 on each side of the midvein, 2 or 3 departing from the base of the midvein, sometimes the secondary and tertiary veins prominent on the adaxial surface, less so on the abaxial surface; domatia absent. Synflorescence frondosebracteose of the diplobotryum type, congested, with short branches concentrated on the terminal portion of the paraclades; accessory branches absent, principal florescence 4–10  cm long, paraclades 6–18  cm long, generally short; bracts similar to vegetative leaves, diminishing in size toward the apex of the branch; flowers shortpedicellate, hypopodium 0.5–1.5  mm long, epipodium 0.5–1  mm long, slightly more in fruit; prophylls 7–8.1 mm long, reaching or surpassing the apex of the floral tube, oblong to obovate, keeled, the apex acute, slightly acuminate, margin plane, silver-sericeous to hirsute; floral tube 6–7 mm long, oblong, silver-sericeous; sepals 3.5–4.5 mm long, triangular, not caudate, not deflexed, in fruit with bulbous secretory trichomes becoming orange in color; epicalyx segments (3.5–)5–5.5 mm long, the same size as or slightly surpassing the sepals; narrowly triangular, dorsiventrally compressed, erect, corolla 3–3.5 cm in diam., rose, lilac, pink, sometimes reddish petals 16–17 × 8–9 mm, elliptic, apex obtuse; stamens 15–18, free portion of the filaments 7–9 mm long, included, rarely, slightly exserted; ovary ca. 2.5 × 2–2.5 mm, globose or rounded, glabrous, style 11–15  mm long, ovules 29–30. Capsule ca. 4 × 4.3–4 mm, globose; seeds 23–25, 1.8–2.2 × 1.2–2.1 mm. Distribution and Ecology (Fig. 13.5)  Diplusodon adpressipilus occurs in campos rupestres and grassy fields only on white sand, in Chapada dos Veadeiros, Goiás, at 1050–1200 m elevation. In Parque Nacional da Chapada dos Veadeiros, it occurs sympatrically with D. alatus, D. praetermissus, and D. sordidus in patches of about 10 m2, flowering synchronously. Flowers from July to February; fruits from August to March. Specimens Examined: BRAZIL.  Goiás,  Alto Paraíso de Goiás, 14°09′02″ S, 47°45′15″ W, 20 Jan 2003 (st), Cavalcanti et al. 3159 (CEN, MO); Alto Paraíso de Goiás, Parque Nacional da Chapada dos Veadeiros, estrada para Colinas, a 34 km do entroncamento da GO, 14º10′S, 47°47′29″ W, 15 Aug 1990 (fl), Cavalcanti et al. 683 (CEN, HUEFS); Alto Paraíso de Goiás, 13°46′ S, 47°30′ W, 31 Jul 1994, Munhoz & Ribeiro 186 (UB); Alto Paraíso de Goiás, córrego dos Ingleses, 14°08′20″

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S, 47°46′08″ W, 12 Sep 1996 (fl), Mendonça et al. 2788 (CEN, IBGE); Alto Paraíso de Goiás, estrada Colinas-Rodovia GO-118, km 42 (rodovia GO-327), 14°10′ S, 47°44′ W, 8 Mar 1989 (fr), Cavalcanti et al. 400 (CEN, NY, SPF); Alto Paraíso de Goiás, entroncamento GO-118-Colinas do Sul, km 26.5, 14°07′58″ S, 47°43′55″ W, 22 Aug 2001 (fl), Pereira-Silva & Pereira 5332 (CEN, MO); Alto Paraíso de Goiás, Parque Nacional da Chapada dos Veadeiros, estrada Colinas, a 34 km do entroncamento da GO, 15 Aug 1990 (fl, fr), Cavalcanti et al. 697 (CEN, HUEFS, MO, NY); Alto Paraíso de Goiás, Parque Nacional da Chapada dos Veadeiros, na margem da cerca próximo à sede (8 km), 14°12′ S, 47°52′ W, 14 Aug 1991 (fl), Cavalcanti et al. 849 (CEN, F, K, MBM); Alto Paraíso de Goiás, estrada de chão entre Alto Paraíso e São Jorge, à 26  km de Alto Paraíso, 15 Aug 1995 (fl), Mendonça et  al. 2244 (IBGE); 14 km de Alto Paraíso/São Jorge, Mulungu a 7 km à direita da estrada de terra, 14°06′54″ S, 47°38′36″ W, 14 Nov 1996 (fl, fr), Silva & Santos 3284 (IBGE); Alto Paraíso de Goiás, Chapada dos Veadeiros, 14°10′ S, 47°47′29″ W, 2 Apr 1997 (st), Cavalcanti et al. 2187 (CEN, MO [2], NY); 27 km de Alto Paraíso/Colinas, 24 Nov 1994 (fr), Silva et al. 2396 (IBGE); Alto Paraíso de Goiás, estrada Alto ParaísoColinas do Sul, km 23, 14°07′49″ S, 47°41′45″ W, 29 Aug 2004 (fl), Cavalcanti et al. 3502 (CEN, SPF). Diplusodon adpressipilus is a single-stem shrub, branched only once in the inflorescence with short branches. It is an ornamental plant due to the shiny and silvery appearance of the leaves, which are covered by a dense, appressed, and sericeous indumentum. Although this may be a strong diagnostic characteristic of the species, it is sometimes not present, disappearing completely on old leaves. In addition to the sericeous indumentum, the species sometimes has hirsute indumentum on the leaves and, in this case, not silver. The UB isotype of Diplusodon adpressipilus has silver sericeous indumentum on some branches and leaves, whereas it is totally absent on others. The NY isotype of D. adpressipilus lacks the silvery aspect and has appressed indumentum on some branches and sparse hirsute indumentum of long trichomes on others. The NY isotype also varies widely in the type of leaf venation, ranging from acrodromous through eucamptodromous. Silvery indumentum is rare in species of Diplusodon. In addition to D. adpressipilus, it occurs in D. argenteus, also from Goiás; D. argyrophyllus, from Bahia; and D. glocimarii from southern Minas Gerais and southeastern São Paulo. Diplusodon argyrophyllus and D. glocimarii differ from D. adpressipilus by their short epicalyx segments and an androecium with 12 stamens. Diplusodon argenteus differs from D. adpressipilus by its acrodromous leaves 4–6 mm wide and also by its 12 stamens. 2. Diplusodon aggregatifolius T.B.Cavalc., Novon 8(4): 337, fig. 1. 1998. Type: Brazil. Minas Gerais: Grão-Mogol, margens do córrego na saída da cidade, estrada para o rio Ventania, 25 Feb 1986 (fl), T.B.Cavalcanti, J.  Semir, N.  S. Chukr, N. L. Menezes & R. Mello-Silva (CFCR) 9596 (holotype, SPF [00042845]; isotypes, CEN, K, MBM, NY, UEC). Figs. 13.2 A–G, 13.5

Descriptions

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Fig. 13.1  Diplusodon adpressipilus and D. argenteus. A–E. D. argenteus. A. Habit. B– C. Leaves, abaxial surface. D. Prophyll, adaxial surface. E. Flower with prophylls, without petals. F–P. D. adpressipilus. F. Habit. G–K. Leaves, abaxial surface. L. Trichome. M. Floral bud. N–O. Flower with prophylls, without petals. P. Capsule. (A–E Hatschbach & Nicolack 53969, MBM; F–H, J–N Cavalcanti et al. 400, SPF; P, I Prance & Silva 58242, UB)

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Subshrubs 0.4–2 m tall, with a xylopodium, erect, branched, pubescent, leaves concentrated at branch apices, not dark-brown when dry. Branches lax, the upper portions quadrangular to subquadrangular, unwinged, red- to wine-colored, pubescent, internodes 0.5–3 cm long. Leaves decussate, not imbricate, suberect, petiolate; petioles 3–7 mm long; blades 10–30 × 5–12 mm, coriaceous, slightly discolorous, eucamptodromous, elliptic to obovate, less often orbicular, base acute to attenuate, apex obtuse, margin plane to rarely subrevolute, adaxial surface pubescent; secondary veins 2 or 3 on each side of the midvein, departing along the midvein, strongly visible; domatia absent. Synflorescence frondose to frondose-bracteose, diplobotryum to pleiobotryum type, lax; accessory branches sometimes present, principal florescence 3–15 cm long, botryum type, generally truncate, paraclades 4–10 cm long; bracts similar in shape to the vegetative leaves, only slightly smaller; flowers short-pedicellate, hypopodium 1–2 mm long, epipodium 0–0.5 mm long; prophylls 5–7  mm long, reaching the middle of the floral tube or to the apex, elliptic to obovate, unkeeled, base narrowly attenuate, margin plane, pubescent; floral tube 4–5 mm long, green, funnel-shaped, pubescent; sepals 2–2.5 mm long, wine-colored, triangular, not caudate, not deflexed, pubescent; epicalyx segments 1.5– 2.5  mm long, flat, spreading or slightly deflexed, the same size as or slightly surpassing the sepals; pubescent; corolla 2.5–3 cm in diam., rose-magenta to purple, petals 9–12 × 6–7 mm, obovate, apex obtuse; stamens (10–)12, free portion of the filaments 5.5–6.5 mm long, exserted ca. 1 mm; ovary 2–2.8 × 1.9–2 mm, globose to rhombic, glabrous, style 12–19  mm long, rose, ovules 6–14. Capsules ca. 5–5.5 × 6 mm, globose, glabrous; seeds 4–9, 3–4 × 2–3 mm. Distribution and Ecology (Fig.  13.5)  To date Diplusodon aggregatifolius is recorded as endemic to the municipality of Grão-Mogol, in northern Minas Gerais, Cadeia do Espinhaço. It inhabits campo rupestre where there is an abundance of quartzite, at elevations of 900–1200 m. It also occurs in campo sujo vegetation of cerrado in transition to campo rupestre at 830 m. Flowers from February to April; fruits from April to June. Specimens Examined: BRAZIL.  Minas Gerais,  Grão-Mogol, arredores, 23 Apr 1978 (fl, fr), Hatschbach 41409 (MBM); próximo a Grão-Mogol, 12 Apr 1981 (fl), Furlan et al. (CFCR) 741 (K, SPF); Grão-Mogol, atrás da cidade, 13 Apr 1981 (fl), Cordeiro et al. (CFCR) 776 (SPF); Grão-Mogol, em direção nordeste da cidade, 16°32′ S, 42°55′ W, 22 May 1982 (fl), Giulietti et al. (CFCR) 3423 (SPF, OUPR, RB); Grão-Mogol, Ribeirão dos Bois, estrada Grão-Mogol-­São João da Vacaria, 16°33′51″ S, 42°53′14″ W, 29 Apr 1997 (fl), Cavalcanti et  al. 2320 (CEN, MO, NY); Grão-Mogol, Ribeirão dos Bois, 16°33′31″ S, 42°53′09″ W, 30 Mar 2004 (fl), Mello-Silva et al. 2558 (CEN, SPF); Grão-Mogol, vale do Ribeirão dos Bois, 22 May 1987 (fl), Pirani & Mello-Silva (CFCR) 10811 (K, R, SPF); Grão-Mogol, na saída da cidade em sentido ao rio Ventania, córrego dos Bois, 16°34′ S, 42°54′ W, 18 Feb 1989 (fl), Cavalcanti et al. 278 (CEN, MBM, SPF); Grão-Mogol, trilha dos garimpeiros, 14 Jun 1990 (fr), Hatschbach & Nicolack 54290 (MBM, P); GrãoMogol, estrada Grão-Mogol-­Cristália, 14 Apr 1981, Pirani et al.(CFCR) 904 (SPF).

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Fig. 13.2  Diplusodon aggregatifolius. A. Branch with inflorescence. B. Leaf, adaxial surface. C. Detail of leaf abaxial surface indumentum. D. Prophyll, adaxial surface. E. Floral bud. F. Flower with prophylls, without petals. G. Pistil. (A–D, G Cavalcanti et al. [CFCR] 9596, SPF; E–F Pirani et al. [CFCR] 904, SPF)

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Diplusodon aggregatifolius is characterized by elliptic to obovate, mediumsized, petiolate (up to 3 cm long) leaves, with acute to attenuate base and sparse, simple pubescence. This species also has a broad, lax, much-branched synflorescence, sometimes with accessory branches. The upper internodes are short, resulting in strongly clustered leaves. The androecium consists of (10–) 12 stamens. 3. Diplusodon alatus T.B.Cavalc., Novon 8(4): 339, fig. 2. 1998. Type: Brazil. Goiás: Alto Paraíso de Goiás, Parque Nacional da Chapada dos Veadeiros, na margem da cerca próximo à sede do IBDF, 14 Aug 1991 (fl), T.B.Cavalcanti, B. M. T. Walter, F. R. Ferreira, H. G. P. Santos & L. B. Bianchetti 850 (holotype, CEN [00014197]; isotypes, K, MBM, NY, SPF). Figs. 13.3 A–I, 13.5, 15.1 A–C Subshrubs 0.4–1.7  m tall, erect to decumbent, unbranched, glabrous, with a xylopodium, not dark-brown when dry. Branches lax, the upper portions markedly quadrangular and winged, wing 1.5–2 mm wide, wine-colored to chestnut brown, nitid, internodes 1.5–5 cm long. Leaves decussate, not imbricate, spreading, coriaceous, sessile-amplexicaulous; blades 30–120  ×  20–70  mm, green-glaucous, not discolorous, eucamptodromous-craspedrodromous sometimes looking hyphodromous, oblong, oblong-lanceolate, ovate or ovate-oblong, base retuse to obtuse-subcordate, amplexicaulous, apex obtuse to sometimes acute, margin plane; secondary veins 15–27 on each side of the midvein, inconspicuous; domatia absent. Synflorescence frondose to frondose-bracteose, botryum to diplobotryum spiciform type, lax; accessory branches absent, principal florescence 20–22 mm long, botryum spiciform type, paraclades 10–23 mm long; bracts similar to the vegetative leaves, decreasing in size toward the stem apex; flowers subsessile, hypopodium 1–2 mm long, epipodium 0.5–1 mm long; prophylls 5–9 × 2–5 mm long, reaching the apex of the floral tube or slightly less, oblong-lanceolate, unkeeled, base retuse, apex obtuse, margin plane; floral tube 6–7 mm long, clear green, funnel-shaped, slightly fleshy, 1–2 mm thick, large and coriaceous in fruit, sepals 7–8 mm long, triangular, with thick margin, fleshy, not caudate, coriaceous, thick and turning inward in fruit; epicalyx segments 3–4  mm long, cylindrical, narrowly triangular, spreading to erect, shorter than the sepals; corolla 6.5–7  cm in diam., rose-magenta, petals 30–35 × 20–22 mm, obovate, apex obtuse; stamens 32–55, free portion of the filament ca. 10 mm long, included; ovary ca. 3 × 6 mm, obconic, glabrous, style ca. 17 mm long, ovules ca. 35. Capsules 5–6 mm × ca. 9 mm, globose, glabrous; seeds 27–49, 3.5–4 × 3–3.5 mm. 2n = 30 (Cavalcanti 1995). Distribution and Ecology (Fig. 13.5)  Diplusodon alatus occurs in Chapada dos Veadeiros, Goiás, where it has a restricted distribution. It forms small populations in the campos rupestres, campos limpos, and campos sujos on rocky soil of white sand at 800–1300 m. Flowers from April to October. Fruits from January to October. Specimens Examined: BRAZIL. Goiás,  Alto Paraíso de Goiás, 24 Jun 1995 (fl, fr), Cavalcanti et al. 1466 (CEN, MO); estrada para Alto Paraíso de Goiás-Colinas, 1 km antes de São Jorge, do lado esquerdo da estrada, 2 Jun 1999 (fl, fr), Cavalcanti

Descriptions

119

et  al. 2507 (CEN, MO); km 23, 14°07′49″ S, 47°41′45″ W, 29 Aug 2004 (fr), Cavalcanti et  al. 3506 (CEN, SPF); fazenda Mato Fundo, 16 Oct 1990 (fl), Hatschbach et al. 54666 (MBM); estrada de terra Alto Paraíso-Colinas do Sul, ca. 30 km da GO-118, 14°09′ S, 47°47′ W, 27 Jan 1997 (fr), Walter et al. 3643 (CEN, NY); Chapada dos Veadeiros, Parque Nacional da Chapada dos Veadeiros, próximo ao rio José Jacó, 14 May 1986 (fl), Romaniuc-Neto et al. 449 (CEN, SP); Parque Nacional da Chapada dos Veadeiros, 14°08′ S, 47°31′ W, 9 May 1987 (fl), Menezes 1208 (CEN, SPF); estrada para Colinas, a 34 km do entroncamento, 15 Aug 1990 (fl, fr), Cavalcanti et al. 681 (CEN, HUEFS, MO, NY); entre São Jorge e rio Preto, 19 Oct 1990 (fr), Hatschbach et al. 54781 (MBM); 2 km E de São Jorge, 14°10′ S, 47°48′ W, 29 Apr 1984 (fl), Kirkbride, Jr. 5542 (UFG, UB); 20 May 2003 (fl), Pastore & Suganuma 650 (CEN); ca. 1 km da sede do Parque Nacional da Chapada dos Veadeiros, 7 Feb 1987 (fr), Pirani et al. 1751 (K, SPF); PARNA Chapada dos Veadeiros, 14°03′59″ S, 47°38′16″ W, 1 Jun 1995 (fl, fr), Felfili et al. 332 (IBGE); Alto Paraíso-Colinas do Sul, 28 km do entroncamento da GO-118, 14°08′ S, 47°44′ W, 2 Apr 1997 (fl, fr), Cavalcanti et  al. 2186 (CEN, MO, NY); Niquelândia, Entroncamento da rodovia para Barro Alto, km 12.9, 28 Jul 1999 (fr), Pereira-Silva et  al. 4230 (CEN); Nossa Senhora da Abadia do Muquém, Serra da Meia Noite, 14°22′31″ S, 47°54′09″ W, 5 Apr 2006 (fl, fr), Cavalcanti & Pereira-Silva 3742 (CEN); Entre Brasília e Niquelândia, 10 May 1963 (fr), Pires et al. 9656 UB). Diplusodon alatus is notable for its robust habit and chestnut-colored branches that bear prominent wings on each of the four angles of the markedly quadrangular stem (Fig.  13.3B). The leaves are glaucous and large, reaching 10  cm, with an amplexicaulous base. The leaves have the unusual number of 15–27 veins on each side of the midvein, and these veins are thin and inconspicuous on both surfaces, giving the impression that there is only a single midvein. The flowers are showy and large, up to 7 cm in diameter, with a fleshy floral tube, 32–55 included stamens, spreading epicalyx segments, much shorter than the sepals and fleshy sepals. In fruit, the floral tube becomes extremely rigid and thick, and the sepals turn inward (Fig. 13.3H). The combination of vegetative characteristics and the large flowers makes this species one of the most ornamental in the genus. 4. Diplusodon appendiculosus Lourteig, Bradea 5(19): 219. 1989. Type: Brazil. Goiás: Alto Paraíso de Goiás, Chapada dos Veadeiros, ca. 35 km N of Veadeiros, 15 Mar 1969 (fl, fr), H. S. Irwin, R. R. Santos, R. Souza & S. F. Fonseca 24414 (holotype, P [01901918]; isotypes, NY, UB). Figs. 13.4 A–N, 13.5 Subshrubs 0.6–1.4 m tall, with a xylopodium, erect, branched, glabrous, darkbrown when dry. Upper branches subquadrangular to quadrangular, with narrow wings of 0.1–0.2 mm wide, internodes 1–2.5 cm long. Leaves decussate, not imbricate, spreading, coriaceous, petiolate; petioles 1–2  mm long; blades 15–50 × 6–25 mm, not glaucous, not discolorous, eucamptodromous, elliptic-lanceolate or oblong to obovate, base acute, attenuate, apex acute to obtuse, margin

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Fig. 13.3  Diplusodon alatus. A. Habit. B. Section of internode. C–D. Leaves, abaxial surface. E. Floral bud. F. Flower with prophylls, without petals. G–H. Fruit. I. Seed. (A–C, H–I Cavalcanti et al. 850, CEN; D–G Romaniuc-Neto et al. 449, SP)

Descriptions

121

thickened, plane or slightly revolute, with protuberances, the abaxial surface with sparse warts and rust-colored veins; secondary veins 3–4 on each side of the midvein, departing along the midvein; domatia absent. Synflorescence frondose to frondose-bracteose, triplobotryum spiciform type, congested; accessory branches absent, principal florescence ca. 14 cm long, paraclades of the first order 19–30 cm long, of the second order 4–7 cm long; bracts resembling the vegetative leaves in shape, scarcely diminished in size toward the apex of the branch; flowers subsessile, hypopodium 0.5–1, epipodium 0; prophylls 6–10 × 4–6 mm, surpassing the apex of the floral tube, ovate-lanceolate or oblong to spathulate, keeled, base retuse to acute, apex obtuse, margin plane; floral tube 5.5–7  mm long, green, oblong, glabrous, internally with cordiform glands below the region of petal insertion; sepals 2.5– 4.5 mm long, triangular, caudate, deflexed, with thick, papillose margin and sparse trichomes; epicalyx segments 4.5–6 × 2 mm, green, foliaceous, hyaline, obovate to lanceolate, spreading to suberect, glabrous, longer than the sepals; corolla ca. 4.5 cm in diam., rose-­magenta to red, petals 19–20 × 10–11 mm, oblong, apex obtuse; stamens 12, free portion of the filaments 7–9 mm long, exserted; ovary 2–3 × 2.l–4 mm, obconic, glabrous or the apex with very short trichomes, style 13–17  mm long, ovules 16–34. Capsules 5–5.1  ×  4–5  mm, dark-brown, oblong, sometimes short stipitate, with white, caducous indumentum on the apex; seeds 4–17, 2.8–4 mm × 1.9–3 mm. Distribution and Ecology (Fig.  13.5)  Diplusodon appendiculosus occurs in the sandy campos and rocky outcrops of Chapada dos Veadeiros at 829–1400  m. Flowers from November to March. Fruits from November to April. Specimens Examined: BRAZIL.  Goiás,  Alto Paraíso de Goiás, ca. 10  km na estrada depois da cidade em direção a Campo Belo, próximo a Cachoeira Comendadora, 28 Jan 1991 (fl, fr), Giulietti 1084 (CEN, SPF); Alto Paraíso de Goiás, cachoeira do Poço Encantado (antiga cachoeira da Comendadora), 13°52′27″ S, 47°15′38″ W, 17 Nov 2005 (fl, fr), Cavalcanti et al. 3620 (CEN); Cavalcante, Teresina de Goiás, rodovia GO-118, 5–8 km N do rio das Almas, 14 Feb 1990 (fl), Hatschbach & Nicolack 53972 (MBM, MO); Cavalcante, Teresina de Goiás, 1 Dec 1992, Hatschbach & Barbosa 58351 (BR, MBM, MO); Cavalcante, povoado Kalunga, 6 Mar 2003 (fr), Pastore & Suganuma 447 (CEN); Cavalcante, caminho Povoado Kalunga, 3 Feb 2003 (fr), Pastore et al. 841 (CEN); Teresina de Goiás, GO-118, Teresina-Alto Paraíso de Goiás, km 18, 13°51′06.5″ S, 47°17′36″ W, 9 Jan 2004 (fl), Cavalcanti et al. 3538 (CEN, SPF); Teresina de Goiás, GO-118, estrada Teresina de Goiás-Alto Paraíso de Goiás, a 20 km de Teresina, 20 Jan 2003 (fl), Cavalcanti et al. 3149 (CEN, MO). The specimens of Diplusodon appendiculosus, when dry, have a dark-brown color that, together with the shape of the leaves and oblong floral tube, immediately associates this species with D. oblongus, a species of wider distribution that also occurs in the municipality of Alto Paraíso de Goiás. Upon more detailed examination, however, D. appendiculosus is a distinguished species by a total absence of trichomes on the branches and leaves and also by the presence of prominent narrow

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Fig. 13.4  Diplusodon appendiculosus. A. Branch with inflorescence. B. Section of internode. C. Leaf, abaxial surface. D. Prophyll, adaxial surface. E. Floral bud. F. Flower with prophylls, without petals. G. Inner surface of floral tube showing glands. H. Detail of epicalyx segments. I. Ovary with indumentum at apex. J. Sepal, outer surface. K. Sepal, inner surface. L. Fruit. M. Capsule. N. Seed. (A–G, I–N Irwin et al. 24414, UB; H, Hatschbach & Nicolack 53972, MBM)

Descriptions

123

Fig. 13.5  Geographic distribution of Diplusodon adpressipilus, D. aggregatifolius, D. alatus, and D. appendiculosus

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wings on the upper branches. In addition, in this species the prophylls are ovatelanceolate or oblong to spathulate and surpass the floral tube and the entire flower; the epicalyx segments are unusually foliaceous and lanceolate; and the sepals are caudate and reflexed and have conspicuous papillae on the apex and margin. The caudate and papillate aspect of the sepals is easily observed in the floral buds as a white, spongy column between the epicalyx segments. Another feature of D. appendiculosus that is shared with D. leucocalycinus and D. astictus, species also occurring in Chapada dos Veadeiros, is the presence of cordiform glands just below the region of petal separation. Diplusodon appendiculosus also has an oblong capsule and usually a short stipitate fruit. Diplusodon leucocalycinus is distinguished from D. appendiculosus by the presence of a villous-tomentose, whitish indumentum over the whole plant, including the flowers, sepals (which are short), and its 12–14 stamens. Diplusodon astictus is distinguished from D. appendiculosus principally by the lax, bracteose synflorescence of the former. 5. Diplusodon argenteus Lourteig, Bradea 5(19): 221. 1989.Type: Brazil. Goiás: Alto Paraíso de Goiás, Chapada dos Veadeiros, sandy campo and rocky slopes with crystalline outcrops, ca. 35  km  N of Veadeiros, 11 Mar 1969 (fl), H.  S. Irwin, R. R. Santos, R. Souza & S. F. Fonseca 24318 (holotype, P [01901917]; isotypes, K [2], NY [2], UB). Figs. 13.1 A–E, 13.9 Subshrubs 0.2–0.4  m tall, with a xylopodium, erect, slightly branched or branched just at the inflorescence, silvery, not dark-brown when dry. Upper branches cylindrical, unwinged, silvery, sericeous, covered by white, appressed trichomes, internodes 0.7–1 cm long. Leaves decussate, imbricate, erect, coriaceous, sessile; blades 7–11  ×  4–6  mm, not discolorous, acrodromous with supranumerary base, ovate-lanceolate to narrowly lanceolate, boat-shaped, keeled, base obtuse, apex acute or long-acuminate, margin plane to involute, blade covered by dense, silverysericeous indumentum; secondary veins 2–3 on each side of the midvein, 2 departing from the base of the midvein, inconspicuous on both surfaces, the midvein prominent; domatia absent. Synflorescence frondose, diplobotryum type, congested; accessory branches absent, principal florescence 6–12 cm long, paraclades 2–10 mm long, generally short; bracts resembling the vegetative leaves in shape and size; flowers sessile; prophylls 4.5–6 mm long, reaching the apex of the floral tube, elliptic to ovate, keeled, apex acute or long-acuminate, sometimes apiculate, margin plane, silver-sericeous; floral tube 4.5–6  mm long, narrowly oblong, silver-sericeous; sepals 3–3.5 mm long, triangular, not caudate, not deflexed, silver-sericeous; epicalyx segments 3–3.3  mm long, narrowly triangular, delicate, erect, the same size or slightly shorter than the sepals; corolla 2.5–2.7 cm in diam., rose to darkrose, petals 13–14 × 6.5–7 mm, obovate, apex obtuse; stamens 12, free portion of the filaments 6–6.5  mm long, slightly exserted, ovary ca. 1.5  ×  2  mm, rounded, glabrous, style 10–13 mm long, ovules 10–19. Capsules 2.7–3 × 2–3.2 mm, oblong, glabrous; seeds 9–17, 1.1–1.5 × 1–1.1 mm.

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Distribution and Ecology (Fig.  13.9)  Diplusodon argenteus is endemic to Chapada dos Veadeiros, Goiás, inhabiting humid and grassy fields with rocky soils at 872–1094  m. Flowers from November to August. Fruits from November to March. Specimens Examined: BRAZIL.  Goiás,  Alto Paraíso de Goiás, ca. 10  km na estrada depois da cidade, em direção a Campos Belos, próximo a Cachoeira Comendadora, 28 Jan 1991 (fl), Giulietti 1083 (CEN, SPF); Cavalcante, caminho para o Vão do Moleque, ca. 20  km da cidade, a partir do asfalto, 13°37′31″ S, 47°29′06″ W, 21 May 2011 (fl), Bringel & Pastore 808 (HUEFS, UB); Cavalcante, Comunidade Kalunga Engenho II dentro da comunidade, 13°34′57″ S, 47°28′21″ W, 16 Jan 2012 (fl, fr), Faria-Jr. 729 (CEN, UB); margem da estrada Cavalcante Minaçú, próximo ao povoado Kalunga Engenho II, 13°39′42″ S, 47°28′34″ W, 4 Jun 2010 (fl), Faria-Jr. et  al. 862 (CEN, UB); estrada de terra Cavalcante-Kalunga, 18  km, 13°41′31″ S, 47°27′57″ W, 18 Nov 2005 (fl, fr), Cavalcanti et  al. 3627 (CEN, MO); povoado Kalunga, 6 Mar 2003 (fl, fr), Pastore & Suganuma 445 (CEN); Cavalcante, Serra do Tombador, Reserva Fundação Boticário, 6 Nov 2012 (fr), Pereira-Silva et al. 16435 (CEN, RB); ponte sobre o rio Tocantins-Cavalcante, km 98, 13°37′46.5″ S, 47°47′18.9  W, 30 Aug 2004 (fl), Cavalcanti et  al. 3525 (CEN); Teresina de Goiás, estrada Teresina de Goiás-Alto Paraíso de Goiás, Poço Encantado, 13°52′24″ S, 47°15′36″ W, 7 Jan 2005 (fl), Pastore & Suganuma 1436 (CEN); Teresina de Goiás, Rodovia GO-118, próximo ao rio das Almas, 14 Feb 1990 (fl), Hatschbach & Nicolack 53969 (MBM). Diplusodon argenteus is represented by small (up to 0.4 m) almost unbranched subshrubs with silvery leaves contrasting with the rose flowers. Other diagnostic features of the species are the erect, strongly imbricate, ovate-lanceolate to narrowly lanceolate and keeled leaves, with apex acute to long-acuminate. The flowers are also covered by silvery-sericeous indumentum with keeled prophylls reaching the apex of the floral tube, epicalyx segments erect, the same size or slightly smaller than the sepals and an androecium with 12 stamens. The other silvery-leaved species from Goiás, D. adpressipilus, differs from D. argenteus, because it has leaves 8–12 mm wide, larger flowers with the floral tube 6–7 mm long, epicalyx segments 5–5.5 mm long, and 15–18 stamens. 6. Diplusodon argyrophyllus T.B.Cavalc., Novon 8(4): 339, fig. 3. 1998.Type: Brazil. Bahia: Abaíra, distrito de Catolés, encosta da serra do Atalho, em frente ao Mendonça, ca. 13º16′S, 41º49′W, 3 Apr 1992 (fl), W.  Ganev 11 (holotype, SPF [00088768]; isotypes, CEN, HUEFS, K, NY). Figs. 13.6 A–L, 13.9 Subshrubs to 1.2  m tall, erect, branched, not dark-brown when dry. Upper branches subquadrangular, unwinged, canescent, covered by a dense sericeous indumentum, internodes 0.9–2 cm long. Leaves decussate, not imbricate, suberect, coriaceous, petiolate; petioles 3–7  mm long; blades 15–40  ×  8–20  mm, slightly discolorous, eucamptodromous, elliptic to ovate-elliptic or rarely

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elliptic-lanceolate, base acute to slightly attenuate, apex acute or shortly acuminate to more rarely obtuse, margin plane to slightly revolute, adaxial surface olive-green, canescent, with sericeous indumentum, veins sulcate, abaxial surface densely covered by a white silvery indumentum; veins prominent; secondary veins (3–)5–7 on each side, departing along the midvein; domatia absent. Synflorescence frondose to bracteose, triplobotryum to pleiobotryum type, congested; accessory branches absent, principal florescence caducous, not seen, paraclades 3–25 cm long; bracts resembling the vegetative leaves in shape, decreasing in size toward the apex of the branch, up to 10 × 5 mm; flowers subsessile, hypopodium ca. 1 mm long, epipodium ca. 0.5 mm long; prophylls ca. 5 × 2.2–2.5 mm, reaching the apex of the floral tube, spathulate, oblong to ovate-elliptic, unkeeled, apex obtuse to acute, margin plane, densely covered by a white to silvery indumentum; floral tube ca. 4.5 mm long, funnel-shaped, densely covered by white to silvery indumentum; sepals 1.5–2 mm long, triangular, not caudate, not deflexed, indumentum the same as that of the floral tube; epicalyx segments 1–2 mm long, lanceolate, equaling the sepals in length, erect to suberect, with the same indumentum as the floral tube; corolla 2.5–3 cm in diam., rose, petals 13–14 × 7.5–8.5 mm, elliptic to obovate, base narrow, apex obtuse; stamens 12, free portion of the filaments 7–7.5 mm long, slightly exserted; ovary 2–2.2 × 3–3.2 mm, obconic, glabrous, style 12–13 mm long, stigma white, ovules 14–15. Capsules ca. 5  ×  4.6  mm, oblong, glabrous; seeds ca. 14, 2.8–3 × 2.2–2.4 mm. Distribution and Ecology (Fig. 13.9)  Diplusodon argyrophyllus is endemic to the mountains of Catolés and Rio de Contas, Bahia (Chapada Diamantina), in campos rupestres at 1018–1100 m. It flowers in April to June and fruits in May to July. Specimens Examined: BRAZIL.  Bahia,  Abaíra, Catolés, Serra da Tromba, 13°17′12″ S, 41°47′32″ W, 14 May 2000 (fr), Miranda-Silva et al. 478 (HUEFS); Água Quente, Pico das Almas, vertente oeste, trilha do povoado da Sta. Rosa, 23 km ao oeste da cidade, 13°31′ S, 42º W, 1 Dec 1988 (st), Harley et al. 27056 (CEPEC, K, SPF); Rio de Contas, Arapiranga, na beira da trilha para a Serra do Porco Gordo, 13°27′ S, 41°45′ W, 18 Jun 2000 (fl, fr), Harley et  al. 53995 (HUEFS); Rio de Contas, campo do Zé Maria, 28 Jul 1999 (fr), Giulietti et al. 1528 (CEN, HUEFS). Diplusodon argyrophyllus is a branched subshrubs entirely covered by a bright silvery indumentum. The leaves are petiolate (3–7 mm long), elliptic to ovate-elliptic of 15–40 × 8–20 mm, with an acute to slightly attenuate base, eucamptodromous and with about 5–7 veins on each side of the midvein. The flowers are covered by the same silvery indumentum as the leaves, the androecium has 12 stamens, and the epicalyx segments are erect to suberect, lanceolate, equaling the sepals in length. Silver-colored indumentum is rare in Diplusodon, and this feature is shared with D. adpressipilus and D. argenteus from Chapada dos Veadeiros in the state of Goiás and with D. glocimarii from southern Minas Gerais State to northern São Paulo State. Diplusodon argyrophyllus differs from D. adpressipilus and D. argenteus mainly by the latter being smaller plants not branched or branching only in the inflorescence, by the smaller and imbricate leaves, without domatia, and by the narrowly triangular epicalyx segments. It differs from D. glocimarii for the latter have

Descriptions

127

Fig. 13.6  Diplusodon argyrophyllus. A. Branch with inflorescence. B. Section of internode. C– D. Leaves, abaxial surface. E. Detail of leaf abaxial surface indumentum. F. Prophyll, adaxial surface. G. Floral bud. H. Flower with prophylls, without petals. I. Epicalyx segment. J. Pistil. K. Longitudinal view of ovary showing one septum as a thick structures toward edge of ovary that splits ovules into two groups. L. Simple leaf trichome. (A–L Ganev 11, CEN)

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leaves with apex acute covered by pubescent to short-villous indumentum, flowers pedicellate (pedicel 2.5–3.5  mm long), prophylls 10–11 surpassing the sepals 1–2 mm, and epicalyx segments 4–4.5 mm long surpassing the sepals. 7. Diplusodon astictus Lourteig, Los Angeles County Mus. Contr. Sci. 72: 6, fig. 1, 2. 1963. Type: Brazil. Goiás: [Alto Paraíso de Goiás], region of the Chapada dos Veadeiros, 29 Apr 1956, E. Yale Dawson 14731 (holotype, US [00117356]; isotypes, LAM, P [2], R). Figs. 13.7 A–O, 13.9 Diplusodon gracilis Koehne var. ulei Koehne, Bot. Jahrb. Syst. 29: 165. 1900. Type: Brazil. Goiás: na região do Tocantins superior, Salto, Sep 1892 (fl), E. Ule 75 (2929), (lectotype, HBG, designated by Lourteig 1989: 217; isolectotypes, P [2], R, RB). Subshrubs 0.3–1.5 m tall, erect, branched just at the inflorescence, glabrescent, usually dark-brown when dry. Lower branches fistulose, the upper ones cylindrical, slightly flattened, unwinged, chestnut brown to wine-colored, with sparse trichomes or glabrous, internodes 3.5–5 cm long. Leaves decussate, not imbricate, suberect, subsessile to petiolate; petioles 1–3 mm long; blades 40–95 × 13–55 mm, chartaceous to coriaceous, not glaucous, not discolorous, eucamptodromous, elliptic to lanceolate, base obtuse, acute, or attenuate, apex acute, rarely obtuse, slightly acuminate, margin plane, sometimes ciliate, blades glabrescent, trichomes slightly bulbous at the base, sparse, common on the abaxial surface; secondary veins 4–5 on each side, departing along the midvein, chestnut brown to wine-colored, domatia absent or rarely with faint domatia. Synflorescence bracteose, triplobotryum to pleiobotryum type, lax; accessory branches absent, principal florescence botryum type, paraclades 5–44 cm long; bracts 15–35 × 5–10 mm, glabrous, elliptic; flowers short-pedicellate, hypopodium 1–2 mm long, epipodium to 0.5 mm long; prophylls 6–10 × 3–4.5 mm, surpassing the apex of the floral tube, elliptic to ovate, keeled, base acute to obtuse, apex acute, margin plane; floral tube 5–7 mm long, funnelshaped, glabrous, internally with cordiform glands below petals; sepals 3–4  mm long, triangular, thick, apex papillose, deflexed, glabrous; epicalyx segments 3–4  mm long, slightly dorsiventrally compressed, spreading to slightly deflexed, glabrous; corolla 4.5–5.5 cm in diam., rose-magenta to red, petals 20–23 × 15–18 mm, obovate, apex obtuse; stamens 12, free portion of the filaments 10–10.5 mm long, exserted; ovary 2–2.5 × (1–)2–3.5 mm, rhombic, depressed, glabrous, sulcate, style 13–17 mm long, ovules 20–40. Capsules ca. 4.5 × 4 mm, globose, glabrous; seeds 26–32, 1.9–3 × 1.5–2.5 mm. Distribution and Ecology (Fig.  13.9)  Diplusodon astictus is restricted to the region of Chapada dos Veadeiros. It occurs at 845–1400 m, forming an extensive population of individuals growing closely together in campo limpo vegetation on humid sandy soil. Flowers from August to April; fruits from September to May. Specimens Examined: BRAZIL.  Goiás,  Alto Paraíso de Goiás, 14°09′02″ S, 47°45′15″ W, 29 Jan 2003 (fl), Cavalcanti et al. 3161 (CEN, MO); Alto Paraíso de Goiás, Parque Nacional da Chapada dos Veadeiros, 14°09′02″ S, 47°46′42″ W, 11

Descriptions

129

Mar 2014 (fl), Cavalcanti et al. 3824 (CEN); Alto Paraíso de Goiás, GO-118, Alto Paraíso de Goiás-São João da Aliança, descida para o rio Piçarras (antigo Pizarrão), lado direito, 14°13′57″ S, 47°29′08″ W, 30 Jan 2003 (fl), Cavalcanti et  al. 3166 (CEN, MO); Alto Paraíso de Goiás, Chapada dos Veadeiros, Fazenda Matão, 19 Sep 1981 (fl, fr), Caldas 17 (IBGE); Alto Paraíso de Goiás, Fazenda Matão, córrego dos Ingleses, 14°08′ S, 47°45′ W, 23 Oct 1996 (fl), Fonseca et al. 1285 (CEN, IBGE, P); Alto Paraíso de Goiás, São Jorge, acesso ao Vale da Lua, 14°09′41″ S, 47°46′15″ W, 28 Feb 1999 (fl), Calago 98 (CEN, MO, NY); Alto Paraíso de Goiás, Serra da Baliza, encostas da serra, 1.3  km da rodovia Alto Paraíso de Goiás-Brasília, 14°10′26″ S, 47°30′17″ W, 2 Dec 2003 (fl, fr), Forzza et al. 2535 (CEN, CEPEC, RB, SPF); Colinas do Sul, GO-327, Colinas-GO-118, km 32, 8 Mar 1989 (fl, fr), Cavalcanti et al. 404 (CEN, G, K, MBM, NY, SPF). Diplusodon astictus is characterized by its erect habit branched just at the inflorescence with upper branches that are chestnut brown to wine-colored and by the wide and bracteose inflorescence. The leaves are large (up to 9.5 cm long), elliptic to lanceolate, and glabrescent. The bracts are small (up to 3 cm long) and elliptic to lanceolate with a prominent midvein. Diplusodon astictus shares with D. appendiculosus and D. leucocalycinus a previously unrecognized character, which is the presence of cordiform glands below the region of petal separation. Of the three species, D. appendiculosus principally differs by its frondose synflorescence, caudate deflexed sepals, and trichomes on the ovary, and D. leucocalycinus differs by its dense indumentum covering the entire plant, frondose synflorescence, erect well-developed epicalyx segments, and trichomes on the ovary. In the protologue, Lourteig (1963) designated the Diplusodon astictus holotype as Yale Dawson 14731 (US) with an isotype at P. Later, Lourteig (1989) referred to the holotype as at P and isotype at LAM.  The holotype is at US, as cited in the protologue. Diplusodon gracilis var. ulei was described based on Ule 75 (2929). This material was examined and is, without doubt, D. astictus. Lourteig (1989) synonymized it with D. astictus, retaining that epithet because D. ulei Koehne (1903) had already been applied to a different species described from Bahia. The specimen at HGB was cited by Lourteig (1989) as holotype, considered here as the lectotype (ICBN Art. 9.17, ex. 14). 8. Diplusodon bahiensis T.B.Cavalc., Phytotaxa 38: 30, fig. 1. 2011. Type: Brazil. Bahia: Morro do Chapéu, 11°35′26″ S, 41°12′26″ W, 2 Mar 1997 (fl), E.  Lughadha, F.  França, P.  E. Gasson & R.  M. Harley 5955 (holotype, CEN [00045090]; isotypes, ALCB, CEPEC, HUEFS, K, SPF). Figs. 13.8 A–D, 13.9 Shrubs 1–1.5  m tall, erect, branched, dark-brown when dry. Upper branches cylindrical to subquadrangular, unwinged, hirtellous to rarely pubescent, trichomes white, thick and sparse, concentrated on the veins, internodes 1–1.5 cm long. Leaves decussate, not imbricate, suberect, petiolate; petioles 1–3  mm long; blades

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Fig. 13.7  Diplusodon astictus. A. Vegetative branch with xylopodium. B. Section of internode (fistulous). C. Leaf, abaxial surface. D. Detail of leaf abaxial surface indumentum. E. Leaf, abaxial surface. F. Base of inflorescence. G. Distal portion of inflorescence. H. Section of internode of inflorescence. I. Bract, abaxial surface. J. Prophyll, adaxial surface. K. Floral bud. L. Flower with prophylls, without petals. M. Apex of sepal showing glandular trichomes. N. Inner surface of floral tube showing cordiform glands. O. Ovary. (A–O Cavalcanti et al. 404, CEN)

Descriptions

131

10–22 × 5–13 mm, membranaceous, not discolorous, eucamptodromous, elliptic to ovate, base obtuse, apex obtuse, margin revolute to subrevolute, slightly folded longitudinally when dry; sparse trichomes on both surfaces; secondary veins 3–5 on each side, departing along the midvein; domatia present. Synflorescence frondose, pleiobotryum type, congested; accessory branches absent, principal florescence 5–10  cm long, paraclades 4–7  cm long; bracts similar to the vegetative leaves, smaller; flowers pedicellate, hypopodium 1–2 mm long, epipodium 0–0.5 mm long; prophylls 6–6.5 × 3–4 mm, reaching the apex of the floral tube or surpassing until the base of the epicalyx segments, elliptic to narrowly elliptic, unkeeled, apex obtuse, margin plane, white and sparse trichomes concentrated on the central vein and margin; floral tube 5–5.5  mm long, light-green when fresh, funnel-shaped, sparsely covered with white trichomes; sepals 2.5–3 mm long, long-triangular, acuminate, ciliate, not caudate, not deflexed; epicalyx segments 2.5–3 mm long, lanceolate, flat, erect, shorter than the sepals, ciliate; corolla 3–3.5 cm in diam., lilac to rose-magenta, petals 12–15 × 8.5–10 mm, oblong to elliptic, apex obtuse; stamens 12, free portion of the filaments 4.5–5 mm long, not exserted, pale yellow to lilac; ovary 2–2.1 × 1.9–2 mm, rhombic, glabrous, style 12–12.5 mm long, rose, stigma green, ovules 17–26. Capsules 4–4.5  ×  3–4  mm, oblong, glabrous; seeds ca. 2, 2.5 × 2.2 mm. Distribution and Ecology (Fig.  13.9)  Diplusodon bahiensis is known from the vicinities of Morro do Chapéu and Lençóis and Umburanas, Bahia, forming small populations along the roadsides in dry sandy soils of campos rupestres and campos sujos at 1000–1151 m. Flowers from January to May; immature fruits from January to June. Specimens Examined: BRAZIL. Bahia,  Lençóis, BR-242, 3 km da entrada para Utinga, 12°33′38″ S, 41°23′29″ W, 10 Mar 1996 (fl), Lima et  al. 2223 (ALCB); Morro do Chapéu, BR-052, vizinhanças da ponte sobre o rio Ferro Doido, ca. 18 km E de Morro do Chapéu, 17 Jun 1981 (fr), Mori & Boom 14509 (K); Morro do Chapéu, proximidades da cachoeira do Ferro Doido, 11°37′31″ S, 40°59′38″ W, 20 Apr 2001 (fl, fr), Melo et  al. 3342 (CEN, HUEFS); Morro do Chapéu, 4  km de Morro do Chapéu, na estrada para Utinga, 11°34′45″ S, 41°09′56″ W, 26 Mar 1996 (fl), Stannard et al. 2561 (HUEFS); Morro do Chapéu, 8 km SE da cidade de Morro do Chapéu, para o oeste da estrada para Utinga, 11°35′ S, 41°12′ W, 2 Jun 1980 (fr), Harley 23017 (CEPEC); Morro do Chapéu, ca. 18 km E de Morro do Chapéu, 19 Feb 1971 (fl, fr), Irwin et al. 32618 (NY); Morro do Chapéu, Morro das Pedras Duas Irmãs, 11°33′25″ S, 41°17′50″ W, 2 May 1999 (fl), França et  al. 2845 (CEN, HUEFS); Morro do Chapéu, 11°33′07″ S, 41°17′17″ W, 30 Jan 2003 (fl), França et al. 4131 (HUEFS); Morro do Chapéu, estrada Morro do Chapéu-Bonito, km 12, 11°37′ S, 41°15′ W, 1 Mar 1989 (fl), Cavalcanti et al. 359 (CEN); Morro do Chapéu, Torre Telebahia, ca. 6 km S de Morro do Chapéu, 16 Jun 1981 (fr), Mori & Boom 14453 (K, NY); Morro do Chapéu, Morrão, em torno da estação retransmissora da Telebahia, a ca. 6 km W da BA-046, entrando a ca. 1 km do entroncamento com a BA-052 (Estrada do Feijão), 14 Mar 1995 (fl), Queiroz & Nascimento 4258 (CEPEC, K, HUEFS); Morro do Chapéu, 19.5  km SE da cidade de Morro do Chapéu, na

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Fig. 13.8  Diplusodon bahiensis. A. Branch with inflorescence. B. Leaf, abaxial surface. C. Prophyll, adaxial surface. D. Flower with prophyll, without petals and stamens. (A Lughadha et al. 5955, CEN; B Cavalcanti et al. 359, CEN; C, D Queiroz et al. 5197, HUEFS)

Descriptions

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estrada BA-052, 11°38′ S, 41°02′ W, 2 Mar 1977 (fl, fr), Harley 19230 (K n.v., CEPEC); Morro do Chapéu, 16 km NW de Lagoinha 4 Mar 1974 (fl, fr) Harley 16697 (CEPEC, NY n.v.); 16  km NW de Lagoinha, 5.5  km SW de Delfino, na estrada para Minas do Mimoso, 10°22′ S, 41°20′ W, 8 Mar 1974 (fl, fr), Harley et al. 17006 (CEPEC, MO, NY, SPF); Umburanas, Serra do Curral Feio (Serra da Empreitada), entrando ca. 20 km S de Delfino, 10°22′ S, 41º19′W, 9 Apr 1999 (fl, fr), Queiroz et al. 5197 (HUEFS). Diplusodon bahiensis belongs to a group of morphologically similar species which are here called the “Hirsutus complex,” involving the binomials D. bahiensis, D. bolivianus, D. hirsutus, D. microphyllus, D. serpyllifolius, D. smithii, D. vividus, and D. xiphodon, which share vegetative similar characteristics. Usually they share a dark-brown habit when dry, with hirsute to hirtellous indumentum of white trichomes in the stem, subrevolute to revolute leaf margins, conspicuous lateral veins on the abaxial surface, with white trichomes concentrated on the veins, highly floriferous inflorescences, generally with leafy racemes with flowers partially obscured by the leaves, sessile to subsessile flowers, hirtellous to hirsute floral tubes and 12–18 stamens. This complex deserves further detailed studies including the genetic variability of populations. Diplusodon bahiensis is similar to D. hirsutus; however, D. hirsutus is defined by its short (2.5–4 mm long), obovate, keeled prophylls, floral tube 2–2.5 mm long, and spreading to slightly deflexed epicalyx segments. Diplusodon bahiensis has larger flowers, with floral tube 5–5.5 mm long, long acuminate sepals (2.5–3 mm long), erect epicalyx segments, and prophylls reaching or surpassing the floral tube apex. Also, in D. bahiensis, the flowers have sparse trichomes, not hirsute as in D. hirsutus. 9. Diplusodon bolivianus T.B.Cavalc. & S.A.  Graham, Novon 6(3): 253, fig. 1. 1996. Type: Bolívia. Santa Cruz: Velasco, Parque Nacional Noel Kempff Mercado, Campamento Las Gamas, 14º49′S, 60º23′W, 28 Mar 1993, T.  Killeen, S.  Panfil & L.  Arroyo 4845 (holotype, MO [256130]; isotypes, CEN, MO [2289179]). Figs. 13.10 A–J, 13.14 Shrubs 0.2–1.5  m tall, erect, branched, usually dark-brown when dry. Upper branches subquadrangular to quadrangular, unwinged, indumentum dense, whitish, hirsute to sericeous, trichomes yellow to white, internodes 1.5–3 cm long. Leaves decussate, not imbricate, erect, petiolate; petioles 1–6  mm long; blades 20–40  ×  9–30  mm, chartaceous, not discolorous, eucamptodromous, elliptic to broadly elliptic, base acute, apex obtuse, very shortly acuminate, margin frequently subrevolute, slightly ciliate, adaxial surface sparsely hirsute, abaxial surface densely hirsute; secondary veins 4–6 on each side of the midvein, inconspicuous on the adaxial surface, prominent on the abaxial surface; domatia inconspicuous. Synflorescence bracteose, diplobotryum to triplobotryum type, congested; sometimes with development of first-­order accessory branches, principal florescence

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Fig. 13.9  Geographic distribution of Diplusodon argenteus, D. argyrophyllus, D. astictus, and D. bahiensis

Descriptions

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short, generally caducous; bracts similar to the leaves in shape, smaller toward the apex; flowers subsessile, hypopodium 0–0.5 mm long, epipodium ca. 0.5 mm long; prophylls 3.2–3.8 × 1–2.5 mm, reaching from the middle to the apex of the floral tube, oblong to obovate, unkeeled, margin plane, ciliate, densely hirsute on the abaxial surface to sericeous; floral tube 4–6  ×  ca. 4  mm, funnel-shaped, densely hirsute to sericeous; sepals 1.5–2 mm long, triangular, not caudate, not deflexed, with the same indumentum as the tube, ciliate; epicalyx segments 1–2 mm long, triangular, deflexed to spreading, less than or equal to the length of the sepals, with the same indumentum as the floral tube; corolla 1.5–2  cm in diam., rose, petals 6–8  ×  4–5  mm, elliptic, apex obtuse; stamens 12, included or slightly exserted; ovary ca. 3 × 2.8–3 mm, obovate, glabrous, style 5.5–6 mm long, ovules 13–17. Capsules 3–4 × 4–5 mm, globose, glabrous; seeds 3–11, 2.1–3 × 2–3 mm. Distribution and Ecology (Fig. 13.14)  Diplusodon bolivianus was found for the first time in Parque Nacional Noel Kempff, southeastern Bolivia, in cerrado at 1500 m. Later, it was collected in Mato Grosso, in campo cerrado, campo sujo, and cerrado at 358–580  m. The geographic distribution is here expanded to Distrito Federal and far southwestern Rondônia in cerrado vegetation with red latosol at the edge of dry forest at 1127–1200 m. Flowers from January to Mar; fruits from April to June. Specimens Examined: BRAZIL. Distrito Federal,  Brazlândia, próximo ao Poço Azul, Lago Oeste, APA da Cafuringa, 15 May 2006 (fr), Pastore & Suganuma 1508 (CEN, NY); Brazlândia, estrada Sobradinho-Brazlândia, 15°40′ S, 48°06′ W, 1 Apr 1992 (fr), Vieira et al. 1255 (CEN). Sobradinho, 15°38′59.8″ S, 47°51′5.2″ W, 10 Mar 2005 (fl), Bringel et  al. 220 (CEN). Mato Grosso, BR 364, VilhenaComodoro, 84  km  S de Vilhena, 13°21′14″ S, 59°52′15″ W, 25 Jun 1997 (fr), Cavalcanti et al. 2381 (CEN, MO, NY); Tapurah, estrada do Capixaba, ca. 20 km NE de Tapurah, 12° 37′ S, 56° 22′ W, 12 Jun 1997 (fr), Souza et al. 17863 (CEN, ESA); Estrada para projeto Ipiranga, 55 km de Tapurah, 12°22′ S, 56°11′ W, 10 Jun 1997 (fr), Souza et al. 17617 (CEN, ESA, UB). Rondônia, Vilhena, em direção a Pimenta Bueno, 12°36′27″ S, 60°21′17″ W, 19 Jan 1999 (fl), Barboza et al. 23 (UB); Vilhena, estrada de Vilhena para Mato Grosso, 13°55′06″ S, 59°45′50″ W, 20 Jan 1999 (fl), Barboza et al. 43 (UB). BOLÍVIA. Santa Cruz,  Velasco, Parque Nacional Noel Kempff Mercado, colecta general de Huanchaca 1, 13°53′41″ S, 60°48′46″ W, en la pampa cerca de la pista de Huanchaca, Soto et al. 365 (MO); Velasco, Parque Nacional Noel Kempff Mercado, colectado a 2  km de la pista, Soto et  al. 427 (MO); Velasco, Parque Nacional Noel Kempff Mercado, colectado a 2 km de la pista, Soto et al. 463 (MO); Velasco, Parque Nacional Noel Kempff Mercado, 14°48′17″ S, 60°23′14″ W, 17 Feb 1997 (fl), Guardia et al. 202 (MO, USZ n.v.); Velasco, Parque Nacional Noel Kempff Mercado, 14°31′27″ S, 60°44′40″ W, 8 Mar 1997 (fr), Jiménez & Gutiérrez 1307 (MO).

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Fig. 13.10  Diplusodon bolivianus. A. Branch with inflorescence. B. Section of internode. C. Leaf, abaxial surface. D. Detail of leaf abaxial surface indumentum. E. Flower with prophylls, without petals. F. Prophyll, adaxial surface. G. Prophyll, abaxial surface. H. Pistil. I. Fruit. J. Capsule. (A–J Killeen et al. 4845, MO)

Descriptions

137

Within the Hirsutus complex (discussed under Diplusodon bahiensis), Diplusodon hirsutus from the Cadeia do Espinhaço, Minas Gerais, share several characteristics with D. bolivianus, including eucamptodromous leaves, hirsute floral tubes, and short epicalyx segments. Diplusodon bolivianus is principally distinguished from these two species by a distinct terminal inflorescence and by the presence of first-order accessory branches. 10. Diplusodon bradei Pilg., Repert. Spec. Nov. Regni Veg. 41: 221. 1937. Type: Brazil. Minas Gerais: Diamantina, Conselheiro Mata, Jun 1924 (fl), A. C. Brade 13406 (lectotype, RB [00540664], here designated; isolectotype, RB [01901913], fragm. P). Figs. 13.11 A–I, 13.14, 15.1 D Diplusodon rupestris T.B.Cavalc., Acta Bot. Brasil. 21(4): 811, fig. 13–16. 2007. Type: Brazil. Minas Gerais: São Roque de Minas, Cachoeira da Casca D’Anta, parte de cima, 10 Jul 1996 (fl), J. Nakajima 1941 (holotype, CEN [00046854]; isotypes, HUFU, MO). Subshrubs 0.5–2  m tall, erect, branched, velutinous, sometimes dark-brown when dry. Branches lax, the upper ones cylindrical, unwinged, rust-colored, the uppermost clear yellow, tomentose to dense pubescent, internodes 3–9.5 cm long. Leaves decussate, not imbricate, suberect, petiolate; petioles 6–25 mm long; blades 20–90 × 10–60 mm, chartaceous to coriaceous, not discolorous, eucamptodromous, ovate, elliptic-lanceolate, broadly elliptic, to less often obovate, base obtuse to acute, attenuate, apex obtuse, margin plane, rare subrevolute, adaxial surface olivegreen, pubescent to velutinous, sometimes with sparse trichomes, abaxial surface canescent, velutinous; secondary veins 5–11 on each side of the midvein, sulcate on the adaxial surface, reddish and prominent on the adaxial surface; domatia absent. Synflorescence bracteose, wide, pleiothyrse type, with cymes in triads or dichasia to diplobotryum, lax, accessory branches of the first and second order, usually present, absent in some populations; accessory branches 2–13 cm long, principal florescence 4–13(−17) cm long, thyrse type, frequently deciduous; paraclades of the first order 7–50 cm long, paraclades of the second order 6–20 cm long, paraclades of the third order 6–14 cm long; bracts of the first order 10–30 × 0.7–14 mm, elliptic to ovate, those of the next order decreasing in size; bracts of the second and third order 15–25  ×  10–15  mm; flowers sessile to pedicellate, hypopodium 0–0.5  mm long, epipodium 0.5–2.5 mm long; prophylls 3–6 × 1–2.5 mm, reaching to less than half the length of the floral tube to the middle of the floral tube, rarely reaching the apex of the floral tube, oblong, sometimes keeled, base obtuse to retuse, margin plane, slightly ciliate; floral tube 4–7 mm long, funnel-shaped, sericeous to villous, with short trichomes; sepals 2.5–4 mm long, yellow, triangular, sometimes slightly caudate, not deflexed, the same indumentum of the floral tube, margin short-ciliate; epicalyx segments 1–3.5  mm long, cylindrical to lanceolate, erect to spreading, reaching the height of or shorter than the sepals, with the same indumentum of the floral tube; corolla 2–3  cm in diam., clear rose, lilac or purple, petals 10–16  ×  6–13  mm, obovate, apex obtuse; stamens 12–18, free portion of the

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filaments 8–10 mm long, exserted 2–3 mm; ovary 2–3.5 × 2.5–3.5 mm, obovate to orbicular, glabrous, style 15–18  mm long, ovules (11–)18–29. Capsule 3.5– 5.1  ×  3–4.5  mm, globose, glabrous; seeds 10–16, 2.2–2.9  ×  2–2.5  mm. 2n  =  30 (Graham and Cavalcanti 2001). Distribution and Ecology (Fig. 13.14)  Diplusodon bradei occurs in Minas Gerais, with two major known small populations, one in the central region of the Cadeia do Espinhaço, where it grows in the typical cerrado and in the cerrado with rocky outcrops, and the other population in Serra da Canastra region, southwest of Minas Gerais, growing in campos rupestres, ca. 845 m elevation. Flowers from January to June. Fruiting from January to September. Specimens Examined: BRAZIL.  Minas Gerais,  Capitólio, estrada para a pedreira, próximo à cachoeira da rodovia, 22 May 2007 (fl, fr), Nakajima et al. 4427 (CEN, HUFU); Capitólio, estrada depois do Paraíso Perdido, região de Furnas, 23 May 2007 (fl, fr), Bernardes et  al. 229 (CEN, HUFU); Delfinópolis, trilha do S. Cannyon, 20°26′04″ S, 46°38′73″ W, 16 May 2003 (fl), Romero et al. 6917 (CEN, HUFU); Diamantina, 18°15′34″ S, 43°40′28″ W, 6 Jan 2006 (fr), Proença & Harris 3133 (CEN, UB); Diamantina, estrada de terra para Conselheiro Mata, km 166, 17 Apr 1987 (fl), Zappi et al. (CFCR) 10601 (SPF); Diamantina, próximo a Conselheiro Mata, margem da estrada para cachoeira do Telésforo, 18°17′06″S, 43°57′17″W, 24 Apr 2016 (fr), Faria-Jr. 5741 (CEN, HDJF, UB); Gouveia, 30 km pela estrada SW de Gouveia, no km 60 na estrada para Curvelo, 11 Apr 1973 (fl), Anderson 8593 (NY, UB); Gouveia, BR-259, subida da Serra do Espinhaço, 23 Feb 1975 (fl), Hatschbach et  al. 36404 (MBM, MO, SPF); Gouveia, estrada Gouveia-Curvelo, descida da serra, 24 Feb 1986 (fl), Cavalcanti et  al. (CFCR) 9574 (SPF, UEC); Gouveia, BR-259, Diamantina-Curvelo, 2 km da BR na estrada para a represa da CEMIG, 18°37′ S, 43°58′ W, 26 Apr 1997 (fl, fr), Cavalcanti et  al. 2301 (CEN, MO); Gouveia, estrada de terra para a fazenda do Sr. Everaldo, 14 Apr 1987 (fl), Menezes et al. (CFCR) 10450 (K, SPF); Gouveia, BR-259, Gouveia-Curvelo, km 34, 18°37′ S, 43°50′ W, 18 Feb 1989, Cavalcanti et  al. 259 (CEN); Gouveia, 18°35′39.6″ S, 43°57′50.4″ W, 5 Apr 1998 (fr), Souza et  al. 20900 (CEN, ESA, HUEFS, MBM); Gouveia, estrada Gouveia-Curvelo, na altura da bica de água potável (lado oposto da estrada), 18°35′42.4″ S, 43°57′16.3″ W, 4 Sep 2003 (fr), Cavalcanti et  al. 3206 (CEN); Gouveia, margem da BR 259, entre Presidente Juscelino e Gouveia, 18°35′41″S, 43°57′50″W, 15 May 2016 (fl), Faria-Jr. 5860 (CEN, HDJF, UB); Sacramento, Parque Nacional Serra da Canastra, 9 May 1995 (fl), Romero et  al. 2126 (CEN, HUFU); Sacramento, Parque Nacional Serra da Canastra, próximo à guarita de Sacramento, 16 Mar 1995 (fl), Romero et al. 1866 (CEN, HUFU); São Roque de Minas, estrada São Roque-Sacramento, próximo à torre de observação, 15 May 1995 (fl), Nakajima et al. 1113 (CEN, HUFU); São Roque de Minas, estrada São Roque-Sacramento, próximo à Guarita de Sacramento, Parque Nacional da Serra da Canastra, 21 Apr 1997 (fl), Romero et al. 4218 (CEN, HUFU); São Roque de Minas, trilha para a guarita de baixo, Cachoeira da Casca D’Anta, 10 Jul 1996 (fl), Nakajima et al. 1958 (CEN, HUFU); São Roque de Minas,

Descriptions

139

Cachoeira da Casca D’Anta, parte de cima, 10 Jul 1996 (fl), J.  Nakajima 1941 (CEN, HUFU, MO); São Roque de Minas, Cachoeira Casca D’Anta, PARNA Serra da Canastra, 17 Jul 1995 (fl), Romero et al. 2509 (CEN, HUFU); Várzea da Palma, Serra do Cabral, 12 Mar 1995 (fl), Hatschbach et al. 61811 (MBM). Diplusodon bradei is easily recognized as a subshrub with rust-colored to clear yellow upper branches and a long and broadly branched inflorescence that usually bears cymes in triads or dichasia and also with accessory branches of the first order, second order, and third order that contribute to the large increase in the number of flowers on the plant, which provide an ornamental aspect to the inflorescence. The leaves are large, ovate to elliptic-lanceolate, with a pubescent to velutinous indumentum and a long petiole (1–2.5 cm), a feature that is otherwise seen only in D. nitidus and D. petiolatus. The species was known only from populations occurring in the central region of Cadeia do Espinhaço in the municipalities of Gouveia and Diamantina. Specimens collected in the region of Serra da Canastra and surroundings, southwest of Minas Gerais, have characteristics that led Cavalcanti (2007) to describe Diplusodon rupestris. Later, with the analysis of more specimens from the same region, it was verified that the populations of Serra da Canastra have characteristics that do not occur in the populations of Gouveia and Diamantina, such as smaller size, dark-brown leaves when dry, inflorescence of the diplobotryum type without accessory branches, and flowers with 18 stamens. However, these plants still have other characteristics such as indumentum, shape of the leaves, and characteristics of the prophylls and epicalyx segments that allow them to be recognized as D. bradei, expanding the amplitude of morphological variation for this species. Diplusodon bradei resembles D. petiolatus, a species of northern Goiás, due to its broad leaves with long petioles, well developed bracteose inflorescences, and accessory branches. Diplusodon petiolatus differs from D. bradei because the former has elliptical leaves with an acute apex, and inflorescences in congested racemes usually forming glomerules, not cymes or dichasia. 11. Diplusodon burchellii Koehne, Pflanzenr. (Engler) 4(216): 208. 1903. Type: Brazil. Goiás: Cavalcante, May 1865, W.  J. Burchell 7911 (holotype, K [000528521]). Figs. 13.12 A–G, 13.14, 15.1 F Subshrubs 0.3–1.5 m tall, erect, slightly branched, with a xylopodium, glabrous, not dark-brown when dry. Upper branches cylindrical to subquadrangular, unwinged, yellow, glaucous when fresh, not dark-­brown when dry, very rarely with long and sparse white trichomes on the stem and branches, internodes 0.6–2 cm long. Leaves decussate, imbricate, congested, erect, sessile; blades 8–25 × 8–26 mm, glaucous, coriaceous, not discolorous, acrodromous with supranumerary base, looking hyphodromous because lateral veins are very inconspicuous, elliptic-lanceolate to ovatelanceolate, base obtuse, sometimes cordate to amplexicaulous, apex acute, margin plane, slightly thickened and yellow, sometimes hyaline; secondary veins 3–4 on each side, with 2–3 departing from the base of the midvein and 1–2 departing along

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Fig. 13.11  Diplusodon bradei. A. Vegetative branch. B. Detail of leaf indumentum. C. Part of inflorescence. D. Detail of branch indumentum. E. Lateral cyme. F. Floral bud. G. Flower with prophyll, without petals. H. Petal. I. Trichome of leaf. (A–H Menezes et al. [CFCR] 10450, SPF; I Zappi et al. [CFCR] 10601, SPF)

Descriptions

141

the midvein, inconspicuous on both surfaces of the leaf; domatia absent. Synflorescence frondose, botryum to diplobotryum type, congested; accessory branches absent; bracts similar to the vegetative leaves in shape, decreasing in size toward the apex of the branch; flowers sessile; prophylls 7–13 × 3–3.5 mm, surpassing the apex of the floral tube by ca. 1.5 mm, glaucous, elliptic-lanceolate, unkeeled, base obtuse, apex acute, margin plane; floral tube 6–7 mm long, green, glaucous, oblong; sepals 1.5–3 mm long, triangular, not caudate, not deflexed; epicalyx segments 3.5–5 mm long, narrowly triangular to lanceolate, dorsiventrally compressed, erect, surpassing the sepals; corolla 3.5–4  cm in diam., lilac to red, petals 17–22  ×  10–12  mm, obovate, apex obtuse; stamens 12, filaments lilac, exserted 3–4  mm; ovary ca. 3  mm wide, obovate, glabrous, style 14–15  mm long, ovules 28–30. Capsule 5–6 × 3–5 mm, oblong, glabrous; seeds 8–25, 2–3 × 2–2.5 mm. Distribution and Ecology (Fig. 13.14)  Diplusodon burchellii occurs in the northern part of Chapada dos Veadeiros, in Cavalcante, with a disjunction in the lowerelevation mountains, in the municipalities of Niquelândia and Campinaçu, at 300–1450 m and also in Paranã, Tocantins. It grows in the cerrado, campo limpo, and campo sujo, in rocky or gravelly soils with high quartz content of campos rupestres. Flowers from August to May; mature fruits in January to November. Specimens Examined: BRAZIL. Goiás,  Alto Paraíso de Goiás, Parque Nacional da Chapada dos Veadeiros, Pouso Alto, BR-010, sentido Teresina de Goiás e Cavalcante, lado esquerdo da estrada, estrada de terra para as cabeceiras do Rio Preto, área intangível do PARNA, 13°58′26″S, 47°29′55″W, 26 Nov 2014 (fl), Fernandez et  al. 159 (CEN, RB); Campinaçu, Fazenda Praia Grande, estrada de acesso pela margem esquerda do rio Praia Grande, 14°02′ S, 48°34′ W, 7 Oct 1996 (fl), Cavalcanti et al. 1824 (CEN, MO, NY); Cavalcante, UTM 23 L 02117/8512562, 18 Oct 2004 (fl), Santos et  al. 1054 (CEN, HTO); Cavalcante, estrada da balsa “Porto dos Paulistas” (rio Tocantins) para Buracão, Curral de Pedra, cerca de 5.8 km do rio, 13°27′43″ S, 48°07′16″ W, 10 Nov 2000 (fl), Pereira-Silva et al. 4384 (CEN, MO, NY); 13°40′17″ S, 47°54′41″ W, 10 Nov 2000 (fl), Pereira-­Silva et al. 4414 (CEN, MO, NY); 13°40′17″ S, 47°54′41″ W, 10 Nov 2000 (fl), Pereira-Silva et al. 4416 (CEN, MO, NY); Cavalcante, estrada entre a sede da RRPN Serra do Tombador e Cavalcante, a 9 km E da sede da reserva, 13°38′09″ S, 47°40′45″W, 1 Feb 2015 (fl), Simon et al. 2568 (CEN); Cavalcante, estrada de terra saindo da balsa do “Porto dos Paulistas”(no rio Tocantins) para o Buracão, HU Curral de Pedra, à cerca de 5.8 km do rio, 13°27′43″ S, 48°07′16″ W, 8 Nov 2000 (fl, fr), Walter et al. 4564 (CEN, MO); Cavalcante, na estrada a caminho para a cachoeira da Prata, 6 Mar 2003 (fl, fr), Pastore & Suganuma 439 (CEN); Cavalcante, 12  km a leste do rio Tocantins, em direção a Cavalcante, 13º23′43″ S, 48°07′56″ W, 20 Feb 2002 (fr), Pereira-Silva et al. 5874 (CEN); Cavalcante, estrada Vila Veneno-Serra branca, km 3.6 (margem direita do rio Macacão, 13°34′25″ S, 48°04′20″ W, 20 Feb 2002 (fl, fr), Pereira-Silva et al. 5894 (CEN, MO); Cavalcante, entroncamento Vila Veneno-balsa do rio São Félix-Cavalcante, km 43, 13°40′31″S, 47°49′24″ W, 21 Feb 2002 (fl), Pereira-Silva et al. 5952 (CEN, MO); Cavalcante, ca. 4.5 km da Vila Veneno em

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Fig. 13.12  Diplusodon burchellii. A. Branch with inflorescence. B. Section of internode. C– D. Leaves, abaxial surface. E. Flower with prophylls, without petals. F. Fruit. G. Capsule. (A–G Pastore & Suganuma 439, CEN)

Descriptions

143

direção ao rio São Félix, 13°31′36″ S, 48°03′45″ W, 25 Jan 2001 (fl), Pereira-Silva et  al. 4645 (CEN); Cavalcante, ponte sobre o rio Tocantins-Cavalcante, km 67, 13°39′47″ S, 47°55′47″ W, 30 Aug 2004 (fl), Cavalcanti et al. 3518 (CEN, SPF); Cavalcante, canteiro da obra rio São Félix, área de influência indireta da futura Hidrelétrica de Cana Brava, 13°22′32″ S, 48°03′49″ W, 19 Sep 2001 (fr), PereiraSilva et al. 5426 (CEN, MO); Cavalcante, caminho para o povoado Kalunga, 3 Feb 2004 (fl), Pastore et al. 839 (CEN); Cavalcante, estrada de terra Cavalcante-Kalunga, 13°38′42″ S, 47°28′39″ W, 20  km, 18 Nov 2005 (fl, fr), Cavalcanti et  al. 3633 (CEN, MO); Cavalcante, comunidade Kalunga Engenho II, trilha para cachoeira Santa Bárbara, 13°33′51″ S, 47°28′35″ W, 20 Sep 2009 (fl), Faria-Jr. et  al. 651 (CEN, UB); Cavalcante, estrada Cavalcante-Minaçu, km 20, 13°38′42.8″ S, 47°28′39.7″ W, 31 Aug 2004 (fl), Cavalcanti et al. 3526 (CEN, SPF); Cavalcante, ponte sobre o rio Tocantins-Cavalcante, km 98, 13°37′46.5″ S, 47°47′18.9 W, 30 Aug 2004 (fr), Cavalcanti et  al. 3528 (CEN); Niquelândia, nas margens do rio Tocantinzinho, 13°59′ S, 48°18′ W, 22 Jul 1995 (fr), Cavalcanti et al. 1586 (CEN, MO). Tocantins, Paranã, estrada São Salvador-balsa do Custódio, km 43, 12°57′01″ S, 48°08′19″ W, 10 Jun 2006 (fr), Pereira-Silva et  al. 10540 (CEN); Paranã, Fazenda Alegre, 13°06′15″ S, 48°05′59″ W, 15 May 2007 (fr), Pereira-Silva et al. 11811 (CEN); Paranã, Fazenda São João, proprietário Aldair Freire, 12°56′ S, 47°36′19″ W, 8 Sep 2003 (fr), Sevilha et al. 3418 (CEN). Diplusodon burchellii characteristically has a subshrubby habit almost unbranched; leaves erect and imbricate, with lateral veins inconspicuous, appearing hyphodromous, glaucous, elliptic-lanceolate to ovate-lanceolate; flowers with an oblong floral tube with short sepals (to 3 mm long); and epicalyx segments erect and longer than the sepals (3.5–4.5 mm long). In leaf shape D. burchellii resembles D. imbricatus, but it differs in floral features. Diplusodon imbricatus is a branched subshrub, with flowers with prophylls surpassing the apex of the floral tube and very short (or absent) epicalyx segments (to 0.7 mm long). There are no duplicates of Burchell 7911, type of Diplusodon burchellii; also there appears to be no photographic evidence of a destroyed B specimen that was studied by Koehne. The specimen at K has an original label with a printed caption “Bearbeitet für das Pflanzenreich,” annotated by Koehne as “n. sp.” in May 1902, with a short description and comparison of the new species with two other species (D. decussatus and D. imbricatus), and is here taken to be the holotype. 12. Diplusodon buxifolius (Cham. & Schltdl.) DC., Prodr. 3: 94b. 1828. Friedlandia buxifolia Cham. & Schltdl., Linnaea 2: 351. 1827. Type: Brazil. Minas Gerais, Chapada, F. Sellow 2054 (lectotype, HAL [0098373], here designated; isolectotype, K[3]). Figs. 13.13 A–L, 13.14, 15.3 A Diplusodon vaccinifolius Mart. ex DC., Prodr. 3: 94b. 1828. Friedlandia vaccinifolia Mart. ex DC., Prodr. 3: 94b, 1828, pro syn. Type: Brazil. Minas Gerais:

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inter São João Del Rei et Ouro Preto, Feb, C. F. P. Martius obs. 844 (lectotype, M [0146287], designated by Lourteig 1989: 218; isolectotype, G). Shrubs 0.4–2 m tall, erect, much-branched, glabrescent, dark-brown when dry. Upper branches quadrangular, winged, distinctly four-angled, glabrous, internodes 0.5–1 cm long. Leaves decussate, not imbricate, spread to suberect, subsessile; petioles 0–1 mm long; blades 8–20 × 7–11 mm, chartaceous, not discolorous, eucamptodromous, ovate or ovate-lanceolate, base obtuse, apex obtuse, margin plane to subrevolute, with trichome protuberances, glabrous; secondary veins 3 on each side of the midvein, inconspicuous on both surfaces, domatia absent. Synflorescence frondose to frondose-bracteose, diplobotryum spiciform to pleiobotryum spiciform type, congested; accessory branches absent, principal florescence 4–7  mm long, paraclades 6–27 cm long; bracts similar to the vegetative leaves in shape, decreasing slightly in size toward the apex of the branch; flowers subsessile, hypopodium 0–0.5, epipodium 0–0.5 mm long; prophylls 2.8–4 × 1.5–2.5 mm, reaching from the middle to the apex of the floral tube, ovate or oblong to elliptic, slightly keeled, base obtuse, apex obtuse, margin plane, generally ciliate, glabrous; floral tube 4–5 mm long, oblong, glabrous, rarely pubescent; sepals 1.8–2 mm long, triangular, caudate, not deflexed, glabrous; epicalyx segments 1–2 mm long, lanceolate, dorsiventrally compressed, spreading to deflexed, shorter than the sepals, glabrous or with long trichomes; corolla 1.5–2.5  cm in diam., lilac to purple, petals 10–12  ×  6–8  mm, obovate, apex obtuse or sometimes acute; stamens 12, free portion of the filaments 6–9 mm long, red, exserted; ovary 1.2–1.5 × 2–2.5 mm, rhombic, style 10–2.8 mm long, ovules 10–16. Capsule 1.9–2  ×  2.9–3  mm, globose, glabrous; seeds 7–16, 1.8–2 × 1–1.5 mm. Distribution and Ecology (Fig. 13.14)  Diplusodon buxifolius occurs in campos rupestres of the mountains near Belo Horizonte, Ouro Preto and vicinity, south of Cadeia do Espinhaço, Minas Gerais, at 1100–1670 m. Flowers from December to August; fruits during the whole year. Specimens Examined: BRAZIL.  Minas Gerais,  Belo Horizonte, Serra da Moeda, 7 May 1988 (fl), Braga 190 (CEN, BHCB); Belo Horizonte, Serra do Curral, 15 Mar 1942 (fl), Mendes-Magalhães 1745 (UB); Belo Horizonte, Serra do Curral, 15 Mar 1960 (fl), Mendes-Magalhães 17363 (BHCB, BHMH, CEN); Belo Horizonte, Serra do Curral, Mar 1963 (fl), Roth 2220 (CESJ, SPF); Belo Horizonte, Serra do Taquaril, 19 May 1933 (fl, fr), Mello-Barreto 3380 (RB); Belo Horizonte, Serra do Taquaril, 9 Mar 1933 (fl), Mello-Barreto 11355 (NY); Brumadinho, Retiro das Pedras, 20°05′35″ S, 43°59′01″ W, Barros & Stehmann s.n. (BHCB 40775); Brumadinho, Retiro das Pedras, 20°05′35″ S, 43°59′01″ W, 12 Aug 1997, Barros & Stehmann s.n. (BHCB 40788); cacheira do Campo, Casaretto Herb. nº. 2746 (G, TO n.v.); Claussen 12a (BR [3]); Claussen 13a (BR [2]); Claussen 14a (BR [3]); Claussen 15a (BR [2]); Caetés, Alto de Nossa Senhora da Piedade, margem da estrada Caetés e a BR-381, 19°49′24″ S, 43°41′12″ W, 21 May 2011 (fl), Faria 1234 (CEN, UB); Caeté, Serra da Piedade, 19°49′ S, 43°40′ W, 28 Apr 1985 (fl), Silva et  al. 1684 (BHCB, CEN); Caeté, Serra da Piedade, Cunha et  al. 1739 (CEN, BHCB); Caeté, Serra da Piedade, Sá et al. 1701 (BHCB, CEN); Caraça, caminho

Descriptions

145

Fig. 13.13  Diplusodon buxifolius. A. Branch with inflorescence. B. Section of internode. C. Leaf, abaxial surface. D. Foliar apex. E–F. Floral buds. G–H. Prophylls, adaxial surface. I–K. Flowers with prophylls, without petals. L. Petal. (A–E, H, K Badini s.n., OUPR 22367; F–G, I Badini s.n., OUPR 4723; J Badini s.n., OUPR 23013)

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para a piscina, 23 May 1987 (fl, fr), Horta et al. 214 (BHCB, CEN, MBM); Caraça, Serra do Caraça, estrada para o Tanque Grande, 22 May 1997 (fl), Mello-Silva et al. 1329 (SPF); Caraça, Serra do Caraça, piscina do Imperador, 26 May 1987 (fl, fr), Grandi et al. s.n. (BHCB 13277); Cascatinha, Serra do Caraça, SW de Catas Altas, ca. 20°51′ S, 43°27′ W, 14 May 1990 (fl, fr), Arbo et al. 4032 (CEN, CTES n.v., MBM, SPF); Catas Altas, Serra do Caraça, ao lado da estrada que leva ao Santuário do Caraça, no mirante, 21 Apr 2006 (fl, fr), Mello-Silva et al. 2894 (CEN, SPF); Catas Altas, Serra do Caraça, no alto da Cascatinha, 20 Apr 2006 (fl, fr), Mello-Silva et al. 2879 (CEN, SPF); Conselheiro Lafaiete, Carreira, 29 Apr 1945 (fl), Badini s.n. (OUPR 4540); Itabirito, Mina do Pico, Cava do Sapecado, 20°15′21″ S, 43°52′47″ W, Arruda & Santos 88 (BHCB); Itabirito, a 10 km da cidade, em direçao a Bahia, 14 Out 1995 (fr), Souza et al. 190 (CEN, ESA); Itabirito, Pico do Itabirito, 18 May 1994 (fl), Teixeira s.n. (BHCB 25022); Itabirito, Serra do Itabirito, 10 Apr 1974 (fl), Badini s.n. (OUPR 23014); Itabirito, Serra do Itabirito, 21 Mar 1963 (fl), Pereira 7272 (M); Itabirito, Serra do Itabirito, 25 Aug 1980 (fl), Badini & Zurlo s.n. (CEN, OUPR 1782); Itabirito, Serra do Itabirito, 25 Apr 1980 (fl), Badini & Zurlo s.n. (OUPR 25404); Itabirito, Serra do Itabirito, 30 Mar 1966 (fl), Badini s.n. (OUPR 5087); Itabirito, Serra do Itabirito, 30 Mar 1966 (fl), Badini s.n. (OUPR 5093); Itabirito, Serra do Itabirito, Badini & Zurlo s.n. (OUPR 25405); Lagoa Seca, Mar 1935 (fl), Cochran s.n. (NY, US 29248); Nova Lima, 22 May 1981 (fl), Martins et al. 8470 (BHCB, CEN); Nova Lima-Morro do Chapéu, 23 Feb 1980 (fl), Grandi 185 (BHCB); Ouro Branco, estrada para Ouro Preto, a cerca de 14  km de Ouro Branco, 5 May 2012 (fl), Delfini et al. 83 (CEN, ESA, RB); Ouro Branco, Serra de Ouro Branco, entrada para a Serra no entroncamenteo com a estrada Ouro BrancoOuro Preto, 5 May 2012 (fl), Delfini et al. 100 (CEN, ESA, RB); Ouro Branco, Serra de Ouro Branco, entrada para a Serra no entroncamenteo com a estrada Ouro Branco-Ouro Preto, 5 May 2012 (fl), Delfini et  al. 107 (CEN, ESA, RB); Ouro Branco, Serra do Ouro Branco, 12 May 1990 (fl), Mello-Silva et  al. 218 (CEN, CTES n.v., SPF); Ouro Branco, Serra do Ouro Branco, 14 May 1988 (fl), Braga et al. s.n. (BHCB 15849); Ouro Branco, Serra do Ouro Branco, 17 Apr 1957 (fl), Pereira 2943 & Pabst 3779 (HB, K, RB); Ouro Branco, Serra do Ouro Branco, 20°30′14″ S, 43°38′44″ W, 28 May 1997 (fl), Mello-Silva et al. 1390 (CEN, SPF); Ouro Branco, Serra do Ouro Branco, 20°50′07″ S, 44°03′ W, 27 Feb 2000 (est), Cavalcanti et  al. 2571 (CEN, MO); Ouro Branco, Serra do Ouro Branco, 24 Jul 1987 (fl, fr), Zappi et al. (CFCR) 11187 (CEN, SPF); Ouro Preto, 12 May 1985 (fl), Ferreira s.n. (CEN, VIC 9341); Ouro Preto, 14 May 1983 (fl, fr), Moura & Vieira 385 (CEN, VIC); Ouro Preto, 5 Apr 1972 (fl), Badini s.n. (OUPR 19977); Ouro Preto, Badini s.n. (OUPR 23013); Ouro Preto, barreira da BR-3 (atual BR-040 que liga Belo Horizonte ao Rio de Janeiro), 30 Mar 1975 (fl), Badini s.n. (OUPR 22259); Ouro Preto, base da Serra do Itatiaia, 10 Apr 1974 (st), Badini s.n. (CEN, OUPR 1781); Ouro Preto, Cachoeira do Falcão, 18 May 1991 (fl), Stehmann s.n. (BHCB 21456); Ouro Preto, campo da Caveira, s.d., Badini s.n. (CEN, OUPR 4732); Ouro Preto, Chapada, 10 Apr 1974 (fl), Badini s.n. (OUPR 3743); Ouro Preto, estação Holanda, Gouveia-Santo Antônio do Leite, 5 Apr 2000 (fl, fr), Roschel 623 (CEN, OUPR); Ouro Preto, estrada de Ouro Preto para o Rio de Janeiro, barreira da BR-3,

Descriptions

147

Fig. 13.14  Geographic distribution of Diplusodon bolivianus, D. bradei, D. burchellii, and D. buxifolius

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30 Mar 1975 (fl), Badini s.n. (CEN, OUPR 4735); Ouro Preto, estrada de terra Ouro Branco-Santo Antonio do Leite, cerca de 5 km antes de Santo Antonio do Leite, 16 Jan 2003 (st), Cavalcanti et al. 3100 (CEN); Ouro Preto, Falcão, 18 Dec 1989 (fl, fr), Simão-Bianchini 171 (CEN, SPF); Ouro Preto, Falcão, 25 Apr 1966 (fl), Reis 5094 (OUPR); Ouro Preto, Lavras Novas, 17 Jun 1999 (fr), Gonçalves 1 (CEN, VIC); Ouro Preto, Moraes, 26 May 1974 (fl), without collector (OUPR 4724); Ouro Preto, Rancharia, 15  km  S de Ouro Preto, caminho a Santa Rita de Ouro Preto, 20°26′ S, 43°31′ W, 13 May 1990 (fl), Arbo et al. 4007 (CEN, CTES n.v., MBM, SPF); Ouro Preto, Rancharia, 25 Apr 1966 (fl), Reis 5095 (OUPR); Ouro Preto, Rodrigo Silva, 29 Apr 1945 (fl), Badini s.n. (OUPR 4541); Ouro Preto, Serra de Lavras Novas, 5 Apr 1972 (fl), Badini s.n. (OUPR 19978); Sabará, Serra da Soledade, 27–31 Aug 2001 (fr), Souza et al. 26936 (CEN, ESA); Santa Bárbara, Serra do Caraça, caminho para o mirante, 23 May 1987 (fl, fr), Zappi & Scatena (CFCR) 10922 (K, P n.v., SPF); Santo Antonio do Leite, 9 May 1975 (fl), Badini s.n. (OUPR 22367); s.d., Badini s.n. (CEN, OUPR 1987); São José dos Pinhais, Caraça, 19°58′ S, 43°27′ W, 23 Mar 1981 (fl), Almeida 139 (HRB, HUEFS, K); São Sebastião das Águas Claras, 15 Apr 1981, Martins s.n. (BHCB 8451, IBGE); São Sebastião das Águas Claras, 8 May 1981, Martins 8448 (CEN); Serra de Itabira do Campo, 22 Dez 1888 (fl), Schwacke 6435 (R); Tiradentes, 21°05′11″ S, 44°09′91″ W, 16 Jan 1994 (fr), Giulietti et al. (CFCR) 1369 (SPF); Tiradentes, Serra de São José, 1997, Brina s.n. (BHCB 39436); Três Moinhos, 29 Apr 1966 (fl), Badini s.n. (OUPR 5089); Venda do Campo, 10 Apr 1972 (fl), without collector (OUPR 4728); Without municipal district: Claussen 54 (BR); Claussen 111 (G); Claussen 1054 p.p. (G; W), Claussen s.n. (K); Gardner 4570 (K, OXF, W); Martius Herb. Fl. Bras. n. 865 (BR [5], G [2], K, M [4], MO, NY, OXF, W [2]); Riedel s.n. (G, LE n.v.); Weddell 1558 (G). Diplusodon buxifolius is represented by branched shrubs with prominently winged quadrangular branches and spread oval to oval-lanceolate leaves up to 2 cm long that turn dark-brown color upon drying, glabrous, with thin, inconspicuous veins. The flowers have 12 stamens, the epicalyx segments are spreading, and the prophylls reach the middle of the tube or extend to the apex. Commonly, the plants are mostly glabrous but have trichomes on some parts of the plant. This indumentum varies from cilia along the margin of the prophylls and epicalyx segments to a rarer condition in which the floral tube and sepals are pubescent (Arbo et al. 4007, Badini s.n. [OUPR 3743], Cochran s.n. [US 29248]). The type material of Diplusodon buxifolius was in the Berlin herbarium (B) and it was destroyed. Photograph of a specimen annotated as Friedlandia buxifolia by Chamisso and Schlechtendal (F neg. nº. 17919) indicates that it corresponds to Sellow 2054 and not Sellow s.n., as stated in the original work. A Sellow specimen without a collection number annotated by Schlechtendal as “Friedlandia buxifolia Nob.” is deposited at HAL and corresponds very well to the description of Friedlandia buxifolia. This specimen was chosen as lectotype. There are duplicates of Sellow s.n. at K, originally from B, but without Schlechtendal handwriting.

Descriptions

149

13. Diplusodon caesariatus Lourteig, Contr. Sci. Nat. Hist. Mus. Los Angeles County 72: 9, figs. 3, 4. 1963. Type: Brazil. Goiás: Chapada dos Veadeiros, 20 km N de São João da Aliança, 14º30′S, 47º30′W, 19 May 1956 (fl), E. Y. Dawson 14397 (holotype, P [01901911]; isotypes, RSA [2]). Figs. 13.15 A–F, 13.19 Shrubs 1.5–3 m tall, erect, branched, not dark-brown when dry. Upper branches cylindrical to subquadrangular, unwinged, hirsute to hispid, with long and white trichomes, internodes 8–16  cm long. Leaves decussate, not imbricate, suberect, petiolate; petioles 1–2 mm long; blades 10–25 × 6–13 mm, chartaceous, slightly discolorous, eucamptodromous to acrodromous with supranumerary base, elliptic to ovate-­elliptic, base obtuse to acute, apex obtuse, margin plane to subrevolute, glabrescent, adaxial surface olive-green, with sunken veins, with sparse, short and fine trichomes or almost glabrous, abaxial surface light-­green, with abundant long trichomes on the veins or with very short trichomes on the veins, almost glabrous; secondary veins 3 on each side, departing along the midvein; domatia present. Synflorescence frondose, botryum to diplobotryum type, congested; accessory branches absent, principal florescence botryum type, 4–5 cm long, often truncate, paraclades 6–8  cm long; bracts the same shape as the leaves diminished in size toward the apex of the branch; flowers pedicellate, hypopodium 1–1.2  mm long, epipodium 0.5–1 mm long; prophylls 1.5–3.7 × 0.5–1.5 mm, reaching the base to the apex of the floral tube, spathulate, unkeeled, base acute, apex obtuse, margin plane, short-ciliate, cilia thick; floral tube 4–5.5  mm long, light-green, funnelshaped, narrow, covered with sparse, long and white trichomes or with very short and inconspicuous trichomes; sepals 2–2.8  mm long, triangular, not caudate, not deflexed, not ciliate; epicalyx segments 1–1.5 mm long, cylindrical, deflexed, ciliate, shorter than the sepals; corolla 2–3  cm in diam., rose-magenta, petals 10–15 × 5–7 mm, obovate, apex obtuse; stamens 18, free portion of the filaments 5–5.2 mm long, rose, exserted ca. 5 mm; ovary 1.7–2 mm × 1.2–2 mm, rounded, glabrous, style 9–12.5 mm long, rose, ovules ca. 13. Capsule 2.8–3 × 2.8–3.5 mm, globose, glabrous; seeds ca. 11, ca. 1 × 1.5 mm. Distribution and Ecology (Fig.  13.19)  Diplusodon caesariatus is endemic and rare in Chapada dos Veadeiros, Goiás, occurring in margins of gallery forest surrounded by cerrado vegetation, at 947 m elevation. Flowers from May to July; fruits in July. Specimens Examined: BRAZIL. Goiás,  São João da Aliança, margem direita do rio Jatobá, 21 km ao N de São João da Aliança, estrada vicinal à esquerda, 14°32′03″ S, 47°30′51″ W, 3 Jul 2015 (fl), Cavalcanti & Pereira-Silva 3982 (CEN); São João da Aliança, margem direita do rio Jatobá, 21  km ao N de São João da Aliança, estrada vicinal à esquerda, 14°32′03″ S, 47°30′51″ W, 3 Jul 2015 (fl, fr), Cavalcanti & Pereira-Silva 3983 (CEN).

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Fig. 13.15  Diplusodon caesariatus. A. Branch with inflorescence. B. Section of internode. C. Leaf, abaxial surface. D. Detail of leaf indumentum, abaxial surface. E. Prophyll, adaxial surface. F. Flower with prophylls, without petals. (A–E Cavalcanti & Pereira-Silva 3982, CEN)

Descriptions

151

Diplusodon caesariatus is a branched shrub characterized by elliptic to ovateelliptic leaves and flowers that are pedicellate with short, deflexed epicalyx segments. Until recently, this species was known only from the type material, but recent collections from the type locality have expanded our understanding of variation in D. caesariatus. For example, one individual has the same characteristic pedicellate flowers of D. caesariatus, with short, deflexed epicalyx segments, but it does not have the hirsute indumentum throughout the plant. Instead, this plant has very short, inconspicuous trichomes, and in addition the flowers have 12 stamens and not 16 as in the type material. A different individual was also collected growing in full sun in a disturbed area next to the forest with smaller leaves and inflorescences with several smaller flowers but also with the typical flowers of D. caesariatus. Despite these differences, the most recent collections are included in D. caesariatus, and it is likely that variation in indumentum is due to the stage of fruiting and exposure to the sun. 14. Diplusodon canastrensis T.B.Cavalc., Bol. Bot. São Paulo, 22(1): 1, fig. 1. 2004. Type: Brazil. Minas Gerais: Sacramento, Parque Nacional da Serra da Canastra, estrada São Roque de Minas-Sacramento, a 41 km da portaria de São Roque de Minas, 7 Nov 2002 (fl, fr), R. Mello-Silva, L. R. Lima & A. F. Pontes 1973 (holotype, CEN [00046636]; isotypes, BHCB, G, HUFU, MBM, MO, NY, RB, SP, SPF). Figs. 13.16 A–C, 13.19 Shrubs 0.4–0.6 m tall, erect, much-branched, not dark-brown when dry. Upper branches quadrangular, with much reduced wings of 0.2–0.3  mm wide, slender, congested, very slightly pilose, trichomes very short, inconspicuous, internodes 3–8  mm long on the upper branches, leaves caducous on the very old branches. Leaves decussate, not imbricate, erect, sessile; blades 5–7 × 1–2 mm, coriaceous, not discolorous, hyphodromous, narrowly oblong, base attenuate, apex obtuse, margin plane, slightly hyaline, sometimes slightly ciliate by very short, sparse trichomes, blades glabrous; secondary veins 1 on each side, inconspicuous, the midvein prominent on the abaxial surface, sulcate on the adaxial surface; domatia absent. Synflorescence frondose, pleiobotryum type, congested; accessory branches absent, principal florescences not seen, paraclades 5–10 cm long; bracts resembling the vegetative leaves; flowers pedicellate, hypopodium 1.5–2 mm long, epipodium 0–1 mm long, very slightly pilose; prophylls 4–5.5 × 1.5–3 mm, reaching to surpassing the apex of the floral tube, oblong-linear or narrowly obovate to elliptic, base acute, apex obtuse, margin plane, glabrous or rarely with very short trichomes; floral tube 4.5–5 mm long, clear green, funnel-shaped, glabrous or with inconspicuous trichomes; sepals 3–3.5  mm long, yellow, triangular, apex attenuate, slightly caudate, not deflexed, margin with secretory papillae on the inner margin, slightly hyaline, short-ciliate with sparse, reduced trichomes, trichomes short, concentrated on the apex; epicalyx segments 3–3.5 mm long, dorsiventrally compressed, with a yellow midvein, erect, reaching or surpassing the apex of the sepals, margin slightly hyaline, short-ciliate with reduced sparse trichomes or totally glabrous, papillose on

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the inner surface in the region of the apex; corolla ca. 2 cm in diam., lilac, petals 9–10 × 7.5–8 mm, long-obovate, apex obtuse to emarginate; stamens 12, free portion of the filaments 6–7 mm long, exserted ca. 1 mm; ovary ca. 1.2 × ca. 2 mm, rhombic, glabrous, style 10–11 mm long, dark rose, stigma conspicuously papillose, ovules 12–17. Capsule 3.8–4  ×  2.9–3  mm, oblong, glabrous; seeds ca. 12, 2–2.5 × 1.5–1.8 mm. Distribution and Ecology (Fig.  13.19)  Diplusodon canastrensis is an endemic species of Serra da Canastra, in Minas Gerais State and inhabits campo sujo vegetation in sandstone and campo rupestre, at 900–1200 m elevation. Flowers and fruits from August to November. Specimens Examined: BRAZIL.  Minas Gerais,  São Roque de Minas, Serra Brava, próximo ao mirante, 28 Sep 1995 (fl), Romero et  al. 2980 (CEN, MO, HUFU); São Roque de Minas, 23 Nov 1995 (fl), Nakajima et al. 1591 (CEN, HUFU, MO); São Roque de Minas, estrada para o Vale dos Cândidos, Garagem de Pedras, Parque Nacional da Serra da Canastra, 22 Aug 1997 (fl, fr), Nakajima et al. 2694 (CEN, HUFU); São Roque de Minas, 24 Nov 1995 (fl, fr), Romero et  al. 3233 (CEN, HUFU, MO). Diplusodon canastrensis is a small, dense shrub, and it is part of a group of species morphologically related by sharing small and linear to narrow leaves and androecium with 12 stamens. The leaves in D. canastrensis are narrow and oblong and, at first glance, appear to be glabrous. Under magnification, the branches, margins of the leaves, prophylls, sepals, and floral tube bear minute, sparse trichomes that are almost imperceptible. With respect to the floral characteristics, this species has prophylls reaching or surpassing the floral tube. The epicalyx segments are erect and dorsiventrally compressed, reach the apex of the sepals, and on the inner apical surface are covered with yellow, translucent glands. The sepals are caudate and have secretory papillae on the inner margin, as referred to others species as D. appendiculosus from Chapada dos Veadeiros, Goiás. Diplusodon canastrensis has leaves that are similar to those of D. myrsinites, a species of the serras south of the Cadeia do Espinhaço in Minas Gerais, but D. myrsinites differs mainly by its larger leaves of 5–12  mm wide, smaller prophylls (2.5–3 mm long), and epicalyx segments of 1–2.5 mm long, shorter than sepals. Another morphological related species is Diplusodon tenuifolius from Campo Alegre, in Goiás State, by the small leaves (0.6–13 × 0.5–0.9 mm), and flowers with 12 stamens. It differs from D. canastrensis mainly by its slender habit, linear leaves, and flowers with pedicel 2.5–3.5 mm long and shorter epicalyx segments (0.8–1 mm long). 15. Diplusodon candollei Pohl ex DC., Prodr. 3: 94a. 1828. Type: Brazil. Minas Gerais In montosis sicis inter Rio Jequitinhonha & Columbis, Cap. Min. Ger. ad Itambe, Aug 1819, J. B. E. Pohl 3156, 3553 = D 943 (first-step lectotype, designated by Lourteig 1989: 218, second-step lectotype, W [0053399], here designated; isolectotypes, BR, G, K[2], W [0053400]). Figs. 13.17 A–I, 13.19, 15.1 A

Descriptions

153

Fig. 13.16  Diplusodon canastrensis. A. Habit. B. Leaf, abaxial surface. C. Flower with prophylls, without petals. (A–C Nakajima et al. 1591, HUFU)

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Shrubs 1–1.5  m tall, erect, much-branched, sometimes dark-brown when dry. Upper branches cylindrical, unwinged, pubescent, internodes 0.3–0.5  mm long. Leaves decussate, not imbricate, suberect, subsessile; petioles (when present) to 1 mm long; blades 4–6 × 0.5–0.8 mm, coriaceous, not discolorous, not glaucous, hyphodromous, linear, base acute, apex obtuse, margin strongly revolute; secondary veins absent, with short dense indumentum on the abaxial surface, adaxial surface glabrous; domatia absent. Synflorescence frondose, diplobotryum to triplobotryum type, congested; accessory branches absent, paraclades 3–12 mm long; bracts similar in shape and size to the vegetative leaves; flowers pedicellate, pedicel pubescent, hypopodium 1–1.5  mm long, epipodium 0.3–0.5  mm long; prophylls 4–4.5 × 2.5–3 mm, reaching the apex of the floral tube, ovate to ovate-elliptic, concave, not keeled, base acute, apex obtuse, margin plane, midvein pubescent on the adaxial surface; floral tube 3.2–3.5 mm long, funnel-shaped, with sparse trichomes, glabrescent; sepals ca. 2  mm long, triangular, not caudate, not deflexed, ciliate; epicalyx segments 1.8–1.9  mm long, cylindrical, erect, slightly shorter than the sepals, with evident cilia; corolla ca. 2 cm in diam., lilac, petals 6.5–8 × 4.5–5 mm, obovate, apex obtuse; stamens 6, free portion of the filaments ca. 6  mm long, exserted ca. 2 mm; ovary ca. 1.3 × 1.5 mm, obconic, glabrous, style 11–12 mm long, ovules 5–6. Capsule 3–5 × 2–3 mm, oblong, glabrous; seeds 3–6, 1.7–3 × 1.5–3 mm. n = 15 (Cavalcanti 1995). Distribution and Ecology (Fig.  13.19)  Diplusodon candollei is endemic to the central portion of Cadeia do Espinhaço, around the municipalities of Santana do Riacho, Diamantina, and disjunctly in Várzea da Palma (Serra do Cabral), in Minas Gerais State, occurring in campos rupestres between 1000 and 1300  m, Minas Gerais. Flowers from November to February; fruits observed during the whole year. Specimens Examined: BRAZIL.  Minas Gerais,  Congonhas do Norte, Serra Talhada, 9.6 km SW da estrada Congonhas do Norte-Gouveia, entrada a 3.7 km NW de Congonhas do Norte, 18°50′26″ S, 43°4′53″ W, 19 Jan 2004 (fl), Pirani et al. 5156 (CEN, NY, RB); Datas, rodovia MG-259, 10–25 km L de Datas, 21 Nov 1997 (fl), Hatschbach & Barbosa 67333 (ALCB, INPA, MBM); BR-259, DiamantinaCurvelo, estrada para a torre da Telemar, próximo ao morro dos Camelinhos, 18°36′27″ S, 43°53′53″ W, 20 Jan 2003 (fl, fr), Cavalcanti et al. 3130 (CEN, MO); Diamantina, Serra do Rio Grande, 2 May 1931 (fr), Mexia s.n. (CEN, VIC 3260); Gouveia, estrada de acesso a torre da Embratel, entrada em frente a Usina eólica, BR-259, Gouveia-Curvelo, 27 km, 18°36′33″ S, 43°53′58″ W, 10 Feb 2015 (fl, fr), Cavalcanti et al. 3940 (CEN); Gouveia, Barro Preto, Torre da Telemig, 29 Nov 1985 (fl), Hatschbach & Silva 50304 (MBM, P, UB); Gouveia, ca. 25 km de Gouveia, na estrada de acesso à torre de televisão em Barro Preto, 29 Nov 1985 (fl), Graham 985 (MO, NY, RB); Joaquim Felício, Serra do Cabral, 17 Nov 1997 (fl, fr), Haschbach & Barbosa 67187 (ESA, FUEL, MBM); Joaquim Felício, 21 Oct 1999 (fr), Hatschbach et  al. 69427 (INPA, MBM); Joaquim Felício, Serra do Cabral, ca. 8 km W de Joaquim Felício, 17°41′34″ S, 44°11′56″ W, 30 Mar 2005 (fr), Souza et  al. 1065 (CEN, HUEFS); Santana de Pirapama, Serra do Cipó, 18°55′31″ S,

Descriptions

155

Fig. 13.17  Diplusodon candollei. A. Branch with inflorescence. B. Section of internode. C. Leaf, abaxial surface. D. Simple trichome of leaf. E. Prophyll, adaxial surface. F. Floral bud. G. Flower with prophylls, without petals. H. Petal. I. Pistil. (A Hatschbach & Silva 50304, MBM; B–I Pereira et al. 790, BHCB)

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43°47′37″ W, 27 Nov 2009 (fl), Savassi-­Coutinho et al. 1319 (CEN, ESA); Santana do Riacho, Parque Nacional da Serra do Cipó, 26 Nov 1991 (fl), Pereira et al. 790 (BHCB, CEN); Santana do Riacho, 27 Jun 1991 (fr), Pereira et al. 932 (BHCB); Santana do Riacho, Mar 1892 (st.), Schwacke 9369 (RB); Várzea da Palma, Fazenda Serra do Cabral Agro-industrial, 19 Nov 1997 (fr), Hatschbach & Barbosa 67290 (MBM, RB); without reference to municipal district: nas montanhas secas entre o rio Jequitinhonha a Calumbi, Aug 1819 (fr), Pohl 3156 (W). Diplusodon candollei is a much-branched shrub distinguished by its small (4–6 × 0.5–0.8 mm), linear, revolute and glabrous leaves, with trichomes only on the midvein of the abaxial leaf surface, and also by its long, concave prophylls with a plane margin and midvein that is pubescent adaxially. The flowers are small, and the androecium has six stamens. Diplusodon candollei resembles D. uninervius in that both have linear, hyphodromous leaves, and both species also occur in the mountains of Cadeia do Espinhaço, in Minas Gerais. However, D. uninervius has prophylls with subrevolute margins that are ciliate and an androecium with 12 stamens. Candolle (1828) published Diplusodon candollei based on a specimen identified by Pohl as “Diplusodon candollei” with two separate branches and one label numbered 3156 and 3553. These numbers correspond to the number D943 in Pohl’s field diary. The specimen “Pohl 3156 and 3553” with an original label is here selected as the second-step lectotype (W 0053399). Lourteig (1989) cited Pohl 3553 as holotype (W), considered as the first-step lectotype (ICBN Art. 9.17, ex. 14), however there are two specimens of this gathering at W. The material at G consists of a specimen, Pohl s.n., but the locality is “in montosis siccis prov. Minas Geraes. Braziliae,” which corresponds to the part of the locality for Pohl 3553  =  D943. Since duplicates of Pohl’s collections went to other herbaria without the information from his field diary (Wahlnöffer, pers. comm) and most lack a number, the material at G is regarded as a duplicate of Pohl 3156 and 3553 = D943. Koehne (1877) mentions that Candolle (1828) gave an erroneous diagnosis for Diplusodon candollei of 12 rather than 6 stamens. Analysis of the type material confirms Koehne’s observation. 16. Diplusodon capitalensis T.B.Cavalc., Acta Bot. Brasil. 21(4): 807, fig. 1–3. 2007. Type: Brazil. Distrito Federal: Brasília, Fazenda Água Limpa-FAL, no campo de arnica, 25 May 2003 (fl), J.  F. B.  Pastore, A.  S. Rodrigues, F.  F. M. Santos & J. B. A. Bringel Jr. 610 (holotype, CEN [00048277]; isotypes, K, MO). Figs. 13.18 A–C, 13.19 Subshrubs, ca. 0.80  m tall, erect, branched, not dark-brown when dry. Upper branches cylindrical, unwinged, rust-colored, covered by dense, white, short trichomes of varying sizes, sometimes equal, internodes 1.5–4 cm long. Leaves decussate, not imbricate, suberect, petiolate; petioles 1–2.5  mm long; blades

Descriptions

157

20–45 × 4–11 mm, olive-green, chartaceous, not discolorous, not dark-brown when dry, acrodromous with supranumerary base to eucamptodromous, elliptic to elliptic-lanceolate, base obtuse, apex acute, margin plane to subrevolute, blades pubescent, trichomes with thick base, abaxial surface short-villous; secondary veins 2–3 on each side, inserted along the midvein; domatia present. Synflorescence bracteose to frondose-bracteose, diplobotryum spiciform to triplobotryum spiciform type, lax; accessory branches absent, principal florescence 5–10 cm long, paraclades 13–20 cm long; bracts 7–15  ×  2–7  mm, elliptic to elliptic-lanceolate; flowers subsessile, hypopodium 0.5–1 mm long, epipodium absent; prophylls 4–5 mm long, reaching the middle of the floral tube or surpassing the middle slightly, ovate to obovate, keeled, apex obtuse, margin plane, ciliate, pubescent; floral tube 4.8–5 mm long, funnel-shaped, covered by short-­villous appressed, ascending, whitish trichomes; sepals 1.5–2.5  mm long, triangular, not caudate, not deflexed, ciliate, sparsely pubescent; epicalyx segments 2–2.5 mm long, cylindrical to slightly dorsiventrally compressed, spreading, shorter than the sepals, sparsely pubescent; corolla 2.5– 3.2 cm in diam., rose to lilac, petals 11–15 × 7–7.5 mm, obovate, base acute, apex obtuse; stamens 18, free portion of the filaments 6.5–7 mm long, exserted 1.5–2 mm; ovary 1.5–2 × 2–2.5 mm, rounded, glabrous, style 12–13 mm long, ovules 17–30. Immature capsules 3–3.5  ×  3.5–3.9  mm, globose, glabrous; seeds 9–14, 2–2.4 × 1.9–2 mm. Distribution and Ecology (Fig.  13.19)  Diplusodon capitalensis is endemic to Distrito Federal environs, in cerrado sensu stricto with rocky soil at 1000 m elevation, and vicinity of Distrito Federal, in the municipality of Luziânia, Goiás. Flowers from April to June; fruits beginning in June. Specimens Examined: BRAZIL. Distrito Federal,  Brasília, Parque Rasgado, 7 Jun 2003 (fl), Chacon & Pinho 53 (CEN); Brasília, estrada para o Poço Azul, ao lado do Parque Nacional, 25 May 2003 (fl), Pastore 641 (CEN, NY); Brasília, proximidades da CAESB, QI 19, Lago Sul, 9 Jun 1985 (fl), Legre s.n. (UB); Brasília, Lago Sul, 15°50′21″ S, 47°28′ W, 9 May 2003 (fl), Silva 5514 (CEN, IBGE); Brasília, a 32 km do Cenargen, DF 170, 15°33′ S, 48°03′ W, 4 Jun 1992 (fl, fr), Dias & Vieira 153 (CEN); Brasília, fazenda Água Limpa, mirante próximo aos campos de arnica, 25 May 2003 (fl), Pastore et al. 597 (CEN); Brasília, ca. 1.5 km W das antenas da Radiobrás, 15°36′ S, 48°08′ W, 14 Apr 1982 (fl), Kirkbride Jr. 4731 (UB); Brasília, Rodovia DF-140, condomínio Parque do Mirante, 15°57′52″ S, 47°49′10″ W, 22 Jun 2000 (fl), Pereira-Silva & Abdala 4275 (CEN). Goiás, Luziânia, fazenda Légua, 16°21′19″ S, 48°12′35″ W, 11 Jun 2003 (fl), Pereira-Silva et  al. 7905 (CEN, MO); Padre Bernardo, Fazenda particular, 6 Jun 2004 (fl), Abrahim et al. 2 (UB). Although Diplusodon capitalensis occurs in Distrito Federal and surroundings, a well collected region, it is represented in herbaria by very few specimens. The most outstanding characteristic of this species is the bracteose synflorescence, with leaves and bracts that are different. The leaves are acrodromous with a supranumerary base, and the bracts are eucaptodromous. The leaves are elliptic-lanceolate, covered with a dense, short, white indumentum and also with evident domatia. The flowers

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Fig. 13.18  Diplusodon capitalensis. A. Branch with inflorescence. B. Leaf, abaxial surface. C. Flower without prophylls, without petals. (A–C Pastore et al. 610, CEN)

Descriptions

159

are covered by the same indumentum as the leaves, but it is more dense, which gives the floral tube a whitish color. Diplusodon capitalensis is morphologically similar to D. lanceolatus, but the latter differs mainly by its wider leaves (5–30 mm large), and flowers with longer epicalyx segments (4–5.5 mm long), the same size as the sepals or longer and usually androecium with 24–26 stamens. 17. Diplusodon chapadensis T.B.Cavalc., Acta Bot. Brasil. 21(4): 809, fig. 4–6. 2007. Type: Brazil. Goiás: Alto Paraíso de Goiás, Chapada dos Veadeiros, estrada Alto Paraíso de Goiás-Colinas do Sul, a 28 km do entroncamento da GO-118, 14°08′06″ S, 47°44′04″ W, 2 Apr 1997 (fl, fr), T.B.Cavalcanti, G. Pereira-Silva & S. A. Graham 2185 (holotype, CEN [00026992]; isotypes, K, MO, NY, RB, SPF). Figs. 13.20 A–C, 13.25 Subshrubs 0.2–0.4  m tall, prostate to suberect, slightly branched, sometimes caespitose, with a xylopodium, not dark-brown when dry. Upper branches cylindrical, unwinged, pubescent to hirtellous, internodes 2–3 cm long. Leaves decussate, not imbricate, erect, petiolate; petioles 1–3  mm long; blades 12–30  ×  3–5  mm, membranaceous to chartaceous, not discolorous, eucamptodromous, narrowly elliptic, base acute to attenuate, apex acute, margin subrevolute, covered by a dense pubescent and white indumentum; secondary veins 2 on each side, departing along the midvein, inconspicuous on both leaf surfaces, domatia absent. Inflorescence frondose, botryum type, lax; accessory branches absent; bracts similar to vegetative leaves in shape but smaller; flowers pedicellate, hypopodium ca. 2 mm long, epipodium ca. 1 mm long; prophylls 9–9.5 × 1.9 mm, surpassing the apex of the floral tube, elliptic, unkeeled, margin plane, dense pubescent; floral tube 7.5–8 mm long, oblong, covered by a dense pubescent, white indumentum; sepals 2.5–3 mm long, triangular, not caudate, not deflexed, with the same indumentum as the tube, not ciliate; epicalyx segments 2.6–3.1  mm long, slightly dorsiventrally compressed, spreading, not surpassing the sepals, with the same indumentum as the tube; corolla ca. 3.5 cm in diam., bright rose, petals ca. 13 mm long, elliptic, apex obtuse; stamens 18, free portion of the filaments ca. 7 mm long, slightly exserted; ovary ca. 1.8 × 2.3 mm, obovate-depressed, glabrous, style ca. 12 mm long, ovules 22–30. Capsule 2.9–4 × 3.5 mm, oblong, glabrous; seeds 13, 2.1–2.4 × 1.7–2 mm. Distribution and Ecology (Fig.  13.25)  Diplusodon chapadensis is endemic to Chapada dos Veadeiros, in Goiás State, and is found in campo limpo and campos rupestres and grows among grasses in typical cerrado with Vellozia and D. alatus, at 1150–1200 m. Flowers from February to May; fruits from March to June. Specimens Examined: BRAZIL.  Goiás,  Alto Paraíso de Goiás, São Jorge, acesso ao Vale da Lua, 14°09′41″ S, 47°46′15″ W, 28 Feb 1999 (fl), Calago 97 (CEN, MO); Alto Paraíso de Goiás, 8 Apr 2005, Chaves 265 (CEN, UB); Alto Paraíso de Goiás, estrada para São Jorge, subida para a casinha do IBAMA, alto da serra, 11 Mar 2014 (fl), Cavalcanti & Pereira-Silva 3823 (CEN); Alto Paraíso de Goiás, rodovia Niquelândia-Alto Paraíso de Goiás, a 6 km de Alto Paraíso, Portal da Chapada, 8 May 2005 (fl), Pastore et al. 1337 (CEN); Alto Paraíso de Goiás, rodo-

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Fig. 13.19  Geographic distribution of Diplusodon caesariatus, D. canastrensis, D. candollei, and D. capitalensis

Descriptions

161

Fig. 13.20  Diplusodon chapadensis. A. Habit. B. Leaf, abaxial surface. C. Flower with prophylls, without petals. (A–C Cavalcanti et al. 2185, CEN)

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via Niquelândia-Alto Paraíso de Goiás, a 6 km de Alto Paraíso, Portal da Chapada, 8 May 2005 (fl), Pastore et al. 1338 (CEN). The prostrate habit that sometimes occurs in Diplusodon chapadensis is not common in the genus, but can occur in some species when individuals are exposed to fire. However, the combination of narrowly elliptic leaves, long prophylls surpassing the apex of the floral tube, oblong floral tube, spreading epicalyx, and 18 stamens does not allow placement of this entity in any other described species. Its narrowly elliptic, eucamptodromous leaves can be confused with D. macrodon, also in Chapada dos Veadeiros, but the latter species differs from D. chapadensis by its dense indumentum, revolute leaf margins, and 12 stamens. 18. Diplusodon ciliatiflorus T.B.Cavalc., Novon 8(4): 342, fig. 4. 1998. Type: Brazil. Goiás: Cavalcante, Colinas do Sul, na estrada Colinas do Sul-Usina Hidrelétrica de Serra da Mesa, 23.6 km antes de chegar na entrada da Usina, 14°12′ S, 48°04′ W, 730 m, 8 Mar 1992 (fr), T.B.Cavalcanti, B. M. T. Walter & D. M. S. Rocha 1046 (holotype, CEN [00017897]; isotypes, HEPH, K, NY, RB, SPF, UB). Figs. 13.21 A–M, 13.25, 15.1 B Subshrubs 0.6–2 m tall, erect, branched, with a xylopodium, glabrescent, often dark-brown when dry. Upper branches subquadrangular, rust-colored, winged, glabrous, internodes 1.5–4.5 cm long. Leaves decussate, not imbricate, suberect, sessile to subsessile, sometimes amplexicaulous; petioles (when present), to 1  mm long; blades 25–65  ×  10–35  mm, slightly glaucous, coriaceous, not discolorous, rust-colored, acrodromous with supranumerary base, lanceolate, elliptic–lanceolate, the most basal ovate to oblong, base obtuse to slightly subcordate, apex acute to obtuse or sometimes acuminate, margin plane with protuberances or ciliate, blades glabrous, adaxial surface with inconspicuous veins, abaxial surface with conspicuous veins, rust-colored; veins 4–5 on each side, 2–4 departing from the base of the midvein, rust-­colored; domatia absent. Synflorescence frondose, diplobotryum to triplobotryum type, lax; accessory branches absent, principal florescence 7–15 cm long, botryum type, paraclades 5–35 mm long; bracts decreasing from leaf size toward the apex of the branch, lanceolate to elliptic-lanceolate or rarely ovate-oblong, sometimes bearing long cilia on the margin, caducous; flowers pedicellate, hypopodium 2–2.5  mm long, epipodium ca. 1  mm long; prophylls 7–12 × 3.5–4 mm, reaching the apex of the floral tube or frequently surpassing the apex of the floral tube, elliptic, unkeeled, base obtuse, apex acute to obtuse, margin plane, generally with abundant long cilia or less often cilia sparse; floral tube 7–8 mm long, funnel-shaped, glabrous or glabrescent, trichomes sometimes sparse; sepals 5–6 mm long, triangular, generally with the apex prolonged to 9 mm in fruit, not deflexed, with long, sparse cilia; epicalyx segments (4–)7–8 mm long, dorsiventrally compressed, spreading to suberect, when erect, exceeding the sepals, with long, sparse cilia; corolla 5–7 cm in diam., lilac to reddish, petals 26–28 × 14–15 mm, obovate, apex obtuse; stamens 19–22, free portion of the filaments 12–16 mm long, exserted; ovary 2.5–2.8 × 3–3.2 mm, rhombic, glabrous, sulcate, style 20–22 mm

Descriptions

163

long; ovules 34–42. Capsule 4.5–8  ×  4–9  mm, globose, glabrous; seeds 11–36, 3–5 × 4–4.5 mm. Distribution and Ecology (Fig.  13.25)  Diplusodon ciliatiflorus is endemic to Chapada dos Veadeiros, Goiás, and ranges southwest to Niquelândia, northeast to Teresina de Goiás, and north to areas around Minaçu at lower elevations. It grows in dense cerrado forests of trees and tall subshrubs and along the margins of gallery forests at 440–800 m and in more open vegetation of the campo rupestre at 996– 1513 m. Flowers from September to April; fruits from April to June. Specimens Examined: BRAZIL.  Goiás,  Alto Paraíso de Goiás, estrada Alto Paraíso-Colinas, 20 km antes de Colinas, 8 Dec 1988 (fl), Pereira-Neto 144 (IBGE, SPF); Alto Paraíso de Goiás, Parque Nacional da Chapada dos Veadeiros, trilha a oeste da sede do Parque, 14°10′27″ S, 47°49′28″ W, 11 Mar 2014 (fr), Cavalcanti et  al. 3812 (CEN); Alto Paraíso de Goiás, estrada Alto Paraíso para São Jorge, 14°06′13″ S, 47°28′50″ W, 5 Apr 2014 (fl), Faria-Jr. et al. 3986 (CEN); Alto Paraíso de Goiás, Parque Nacional da Chapada dos Veadeiros, São Jorge, Trilha dos Saltos, 14°10′03′′S, 47°49′31″W, 24 Nov 2014 (fl), Verdi et  al. 7259 (CEN); Parque Nacional da Chapada dos Veadeiros, próximo a cachoeira do rio Preto, perto do povoado São Jorge, 6 Feb 1987 (fr), Pirani et al. 1693 (SPF); Parque Nacional da Chapada dos Veadeiros, trilha para a cachoeira das Cariocas, 13°53′ S, 48°09′ W, 11 Sep 1996 (fl, fr), Mendonça et al. 2707 (IBGE); entre São Jorge e rio Preto, 19 Oct 1990 (fl, fr), Hatschbach & Silva 54783 (CTES n.v., MBM, NY); Chapada dos Veadeiros, 3 Nov 1972 (fl, fr), Rizzo 8560 (UFG); Chapada dos Veadeiros, córrego dos Ingleses, 14°08′20″ S, 47°46′08″ W, 23 Oct 1996 (fl), Mendonça et al. 2923 (CEN, IBGE); Chapada dos Veadeiros, ca. 20 km de Alto Paraíso de Goiás, 20 Mar 1971 (fr), Irwin et al. 32760 (UB); Chapada dos Veadeiros, 23 km de Alto Paraíso para Teresina de Goiás, 13°56′38″ S, 47°29′38″ W, 12 Nov 1996 (fl), Silva & Santos 3203a (CEN, IBGE); entre São Jorge e Colinas do Sul, ca. 3  km de São Jorge, 14°13′43″ S, 47°53′52″ W, 14 Nov 1996 (fl), Mendonça et al. 2963 (CEN, IBGE); Colinas do Sul, estrada Colinas do Sul-Minaçu, estrada de acesso às torres LT 500 Kv 72 e 73, 20 Oct 1999, Amaral et al. 538 (CEN); estrada UHE Serra da MesaColinas, a 20 km da barreira da UHE Serra da Mesa, 24 Jun1995 (fr), Cavalcanti et  al. 1464 (CEN); estrada Colinas do Sul-­canteiro de obras da Serra da Mesa, 13°53′ S, 48°07′ W, 20 Oct 1996 (fl), Walter et al. 3494 (CEN, MO); estrada ColinasSerra da Mesa, 13°53′26″ S, 48°09′46″ W, 30 Nov 1996 (fl), Walter et  al. 3626 (CEN, NY); vale do rio Tocantizinho, próximo à foz, reservatório em formação do AHE Serra da Mesa, segmento IV, 13°56′ S, 48°17′ W, 29 Jan 1998 (fr), Walter et al. 4071 (CEN); Minaçu, estrada Serra da Mesa-­Colinas do Sul, mais ou menos 24 km da entrada Sul do Canteiro de Obras da Usina Hidrelétrica de Serra da Mesa, 13°52′ S, 48°08′ W, 10 Dec 1991 (fl), Walter et al. 962 (CEN, MO); Nossa Senhora da Abadia do Muquém, GO-237, entroncamento Muquém-Colinas do Sul, 5 km, na descida da Serra da Meia Noite, 14°22′29″ S, 47°53′19″ W, 5 Apr 2006 (fr), Cavalcanti & Pereira-Silva 3749 (CEN); Teresina de Goiás, estrada para Alto Paraíso, a 41  km de Teresina (GO-241), Parque Nacional da Chapada dos

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Fig. 13.21  Diplusodon ciliatiflorus. A. Branch with inflorescence. B. Section of internode. C– F. Leaves, abaxial surface. G. Flower with prophylls, without petals. H. Ovary. I–J. Prophylls, adaxial surface. K. Fruit. L. Capsule. M. Seed. (A–C, G–I Hatschbach & Silva 54783, MBM; D Pirani et al. 1693, SPF; E–F, K–M Cavalcanti et al. 1046, CEN; J, Pereira-Neto 144, IBGE)

Descriptions

165

Veadeiros, 13°57′25″S, 47°28′19″W, 26 Nov 2011 (fr), Saraiva et al. 307 (CEN, RB). Diplusodon ciliatiflorus is usually represented by tall branched subshrubs, characterized by leaves with evident rust-colored veins, especially on the abaxial surface, large flowers with long, dorsiventrally compressed epicalyx segments that are spreading or suberect, and with an androecium of about 20 stamens. The sepals and epicalyx have an abundance of trichomes on the margins that form long cilia. The species is glabrescent, and herbarium specimens either have mostly glabrous leaves or are densely pilose (e.g., Walter et al. 4071, CEN). It is also possible to find specimens with sparse trichomes on the leaf blades and floral tube; these trichomes later fall away. The seedlings are sparsely or abundantly covered with long trichomes. Individual plants of Diplusodon ciliatiflorus are variable in height, from subshrubs 60 cm tall up to 2 m. Lourteig (1989) described Diplusodon thysanosepalus, and according to her illustration and diagnosis this species resembles D. ciliatiflorus. Diplusodon thysanosepalus has acrodromous leaves with a typical five-veined base; the flowers have cilia on the sepals and epicalyx. A comparison between D. ciliatiflorus and D. thysanosepalus has been based on Lourteig’s diagnosis of the latter, the illustration of the inflorescence that accompanies the protologue and the holotype (Glaziou 21548a, P). In Diplusodon thysanosepalus the inflorescence is lax and has broad, ovate bracts with a slightly revolute margin. Flowers have narrow, erect epicalyx segments, and, as the sepals, bear short, relatively sparse cilia, and the androecium has 12 stamens. Diplusodon ciliatiflorus differs from the latter by the congested inflorescence with bracts larger than the internodes, lanceolate to elliptic-lanceolate, with a plane margin. Further, the epicalyx segments are spreading and suberect and, like the sepals, covered with long abundant cilia; the androecium has 19–20 stamens. Another species, D. sordidus resembles Diplusodon ciliatiflorus because they both have ovate-lanceolate to ovate and acrodromous leaves and flowers with spreading to suberect epicalyx segments that are 8–10 mm long. In spite of the similarity between the two species, D. sordidus is totally glabrous, does not have glaucous leaves or long-ciliate flowers, and has androecium with 26–28 stamens. 19. Diplusodon ciliatus (T.B.Cavalc.) T.B.Cavalc., comb. nov. Diplusodon ulei var. ciliatus T.B.Cavalc., Novon 8(4): 349. 1998. Type: Brazil. Bahia: Lençóis, arredores da cidade, Jul 1992 (fr), T.B.Cavalcanti & C. H. S. Ayres 1260 (holotype, CEN [00019206]; isotypes, NY, SPF). Figs. 13.22 A–F, 13.25 Subshrubs 1–2  m tall, erect, branched, dark-brown when dry. Upper branches subquadrangular to quadrangular, unwinged, wine- to rust-colored, glabrous, internodes 0.8–2.0 cm long. Leaves decussate, not imbricate, suberect, petiolate; petioles 3–7 mm long; blades 11–35 × 8–23 mm, nitid, chartaceous, discolorous (fresh), eucamptodromous, glabrous, broadly elliptic to elliptic, less often obovate, base

166

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acute, slightly attenuate, rarely obtuse, apex obtuse to acute or sometimes slightly acuminate, margin plane to subrevolute, ciliate, adaxial and abaxial surfaces glabrous or sometimes trichomes on the veins, rarely on the whole blade, abaxial surface with thin and prominent veins winish, glabrous or trichomes on the veins, rarely on the whole blade; secondary veins 3–5 on each side, departing along the midvein; domatia present, usually on the two basal pairs of veins. Synflorescence bracteose to frondose-bracteose, diplobotryum to triplobotryum type, lax; accessory branches absent, principal florescence when present ca. 4 cm long, generally caducous, botryum type, paraclades 4–30  cm long, wine- to rust-colored; bracts 7–10 × 3–5 mm, similar to the vegetative leaves in shape, decreasing in size toward the apex of the branch; flowers pedicellate, hypopodium 1–1.5 mm long, epipodium 0.5–2 mm long; prophylls 3–4 × 1–1.5 mm, reaching the middle or the apex of the floral tube, spathulate, unkeeled, base acute, sometimes prolonged, apex obtuse, margin plane, wine, ciliate; floral tube 4–4.5 mm long, green to dark-wine in color, funnel-shaped, glabrous; sepals 2.5–4  mm long, purple, triangular, caudate, deflexed, ciliate; epicalyx segments 2.5–4 mm long, dorsiventrally compressed with acute attenuate apex, curved, spreading with the apex deflexed, longer or the same size than the sepals, ciliate; corolla 2.5–3 cm in diam., rose-­magenta, petals 9.5– 12 × 5.5–8 mm, obovate-elliptic, apex obtuse; stamens 12, free portion of the filaments 6–7  mm long, exserted, surpassing the sepals 2–2.5  mm; ovary 1.9–2.2 × 2–2.5 mm, rhombic, glabrous; style 10–12 mm long, rose, ovules 11–14. Capsules 3.5–4.4 × 4–4.5 mm, globose, glabrous; seeds 8–9, 2–2.5 × 1.2–1.8 mm. Distribution and Ecology (Fig. 13.25)  Diplusodon ciliatus is endemic to Chapada Diamantina, in Bahia State, known from the municipalities of Andaraí, Lençois, and Mucugê (Bahia). Flowers February to August; fruits from May to August. Specimens Examined: BRAZIL. Bahia,   Andaraí, rodovia Andaraí-Mucugê, km 30, 20 May 1989 (fl), Mattos-Silva et al. 2808 (CEN, CEPEC); Andaraí, 12°48′ S, 41°19′ W, 4 May 2002 (fl, fr), Loureiro et al. 643 (ALCB); Cascavel, borda ocidental do Parque Nacional da Chapada Diamantina, 13°17′ S, 41°21′ W, 24 Mar 2005 (fl), Funch & Funch 760 (CEN, HUEFS); Igatu, estrada para Igatu, próximo à cidade de Igatu, 8 Aug 2006 (fl, fr), Pastore & Suganuma 1523 (CEN); Lençóis, Parque Nacional da Chapada Diamantina, 12°34′6″ S, 41°24′21″ W, 16 Jun 2005 (fl, fr), Moraes et  al. 12 (CEN, HUEFS); Lençóis, Parque Nacional da Chapada Diamantina, 12°27′35″ S, 41°26′43″ W, 29 Jul 1994, Guedes et  al. 304 (ALCB, CEN); Lençóis, Parque Nacional da Chapada Diamantina, 30 Jun 1995, Guedes et al. 2049 (CEN, ALCB); BR-242, Lençóis, ca. 15 km NW de Lençóis, km 225, 10 Jun 1981 (fl, fr), Mori & Boom 14260 (CEPEC, K, NY, US); Lençóis, Mori & Boom 14289 (K, NY, US); Lençóis, ca. 5 km N de Lençóis, 13 Jun 1981 (fr), Mori & Boom 14404 (CEPEC, K [2], NY, US); Lençóis, caminho de Lençóis para Barro Branco, ca. 8 km de Lençóis, 27 Apr 1992 (fl), Queiróz et al. 2754 (ALCB, CEN, CEPEC, HUEFS); Lençóis, Morro do Pai Inácio, BR-242 W de Lençóis, km 232, 12 Jun 1981 (fl, fr), Mori & Boom 14373 (CEPEC, K [2], NY, US); Mucugê, córrego do Boiadeiro, 14°32′05″ S, 42°31′22″ W, 13 Jun 2011 (fl, fr), Hurbath 138 (ALCB); Mucugê, Pedra Redonda, entre o rio Preto e o rio Paraguaçu, 12°57′ S, 41°24′ W, 15

Descriptions

167

Fig. 13.22  Diplusodon ciliatus. A. Branch with inflorescence. B. Leaf, abaxial surface. C. Detail of leaf abaxial surface indumentum. D. Floral bud. E. Prophyll, adaxial surface. F. Flower with prophylls, without petals. (A Mattos-Silva et al. 2808, CEN; B–F Mori & Boom 14260, K)

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Jul 1996, Bautista et al. 3621 (ALCB[2], CEN, HUEFS); Mucugê, Trilha para o rio Paraguaçu, início da estatação de tratamento de água, 13°00′01″ S, 41°23′29″ W, 22 May 2010 (fl, fr), Roque et al. 2791 (ALCB); Mucugê, ca. 3 km na estrada MucugêCascavel, vale do rio Mucugê, 20 Mar 1990 (fl), Carvalho & Saunders 2946 (CEN, CEPEC n.v.; NY); Mucugê, 3 km ao sul de Mucugê, na estrada para Jussiape, 13º S, 41°24′ W, 26 Jul 1979 (fr), Mori et al. 12570 (CEN, NY, US); Mucugê, 16 km N de Mucugê na rodovia para Andaraí, 23 Nov 1985, Graham 949 (MO, NY); Mucugê, Parque Municipal de Mucugê, 13°09″ S, 41°20′23″ W, 9 Jun 2004 (fl, fr), Souza et  al. 883 (CEN, HUEFS); Mucugê, Parque Nacional da Chapada Diamantina, 500  m da saída da cidade, estrada de terra à esquerda, 100  m dentro da estrada, 12°59′51″ S, 41°21′45″ W, 15 May 2016 (fl), Brauner et  al. 58 (CEN, UB); Palmeiras, Trilha do Poço das Águas Claras, 12°32′13″ S, 41°29′05″ W, 24 May 2014 (fl, fr), Roque et al. 4240 (ALCB, RB); Rio de Contas, cachoeira do Fraga, 13°32′ S, 41°52′ W, 23 Apr 2011 (fl, fr), Guedes et al. 18851 (ALCB). Diplusodon ciliatus is represented by branched subshrubs, characterized by ovate-elliptic leaves that have an acute apex, ciliate margins, and prominent veins on the abaxial surface. The leaves turn dark-­brown color upon drying. This species also has cilia on the margin of the prophylls, sepals, and epicalyx segments, which are 3–4.5 mm long and spreading with the apex deflexed. Diplusodon ciliatus had previously been considered as a variety of Diplusodon ulei; however, the analysis of additional specimens led me to conclude that it is a different species. Diplusodon ulei has larger flowers, coriaceous leaf blades that are glabrous, with a plane, not-ciliate margin and veins that are not prominent on abaxial surface, and it also has much longer epicalyx segments (4–5 mm long). Species rank for D. ciliatus was also corroborated by ongoing molecular phylogenetic analysis in which the two varieties previously recognized under D. ulei are placed in distant subclades. 20. Diplusodon ciliiflorus Koehne, Bot. Jahrb. Syst. 23 (Beibl. 57): 35. 1897. Type: Brazil. Minas Gerais: G. Gardner 4569 (first-step lectotype, designated by Lourteig 1989: 215, second-step lectotype, W [0010988], here designated; isolectotypes, F, K [2]). Figs. 13.23 A–K, 13.24 A–F, 13.25 Subshrubs 0.10–3 m tall, erect, branched, with a xylopodium, sometimes darkbrown when dry. Upper branches subquadrangular, unwinged, yellow, glabrous or rarely with inconspicuous pubescence, internodes 6–25 cm long. Leaves decussate, not imbricate, erect to suberect, subsessile; petioles to 1.5  mm long; blades 6–24  ×  5–10  mm, coriaceous, slightly discolorous, not glaucous, eucamptodromous, oblong, elliptic to ovate, sometimes narrowly oblong, base obtuse, apex obtuse, rarely acute, margin plane, ciliate or not, adaxial surface dark-green, nitid, glabrous, abaxial surface olive-green, rarely with sparse trichomes distributed over the blade; secondary veins 1–3 on each side, departing along the midvein, inconspicuous on the adaxial surface, chestnut brown on the abaxial surface; domatia absent. Synflorescence frondose, botryum or triplobotryum type, lax; accessory

Descriptions

169

branches absent, principal florescence 5–15  cm long, paraclades 5–35  cm long; bracts similar to the vegetative leaves, decreasing in size toward the apex of the stem; flowers pedicellate, hypopodium 1–2.5 mm long, epipodium 0.5–1 mm long; prophylls 5–9  ×  2.5–4.5  mm, reaching or sometimes surpassing the apex of the floral tube, oblong to elliptic, base acute, apex acute, margin plane, frequently ciliate, glabrous or with sparse trichomes; floral tube 4–6  mm long, larger in fruit, green to wine-colored, funnel-shaped, glabrous; sepals 4–6 mm long, slightly caudate, not deflexed, triangular, with conspicuous cilia on the margin; epicalyx segments 4–6 mm long, wine-colored at the base, cylindrical, erect, the same size as or shorter than the sepals, generally prominently ciliate; corolla 3.5–4  cm in diam., rose-magenta, petals 14–20 × 12–15 mm, obovate, apex obtuse, sometimes emarginate; stamens 12, free portion of the filaments 6.5–7.5 mm long, slightly exserted; ovary 1.8–2 × 2.5–2.8 mm, obconic, glabrous, style 14–17 mm long, ovules 15–24. Capsules globose, glabrous; seeds 4–12, 2.5–2.8 × 1.9–2 mm. Distribution and Ecology (Fig.  13.25)  Diplusodon ciliiflorus is endemic to the central portion of Cadeia do Espinhaço, in the state of Minas Gerais, around the municipalities of Diamantina to São Gonçalo do Rio Preto. This species occurs in small populations of isolated individuals at 1000–1400  m elevation. The species inhabits sandy soil in the campos limpos or rupestres. Flowers and fruits from October to April. Specimens Examined: BRAZIL.  Minas Gerais,  Conselheiro Mata, 10 Jan 1987, Grandi et al. s.n. (BHCB 8094, CEN); Datas, estrada Curvelo-Diamantina, km 108, 24 Nov 1985 (fl, fr), Pirani et al. (CFCR) 8721 (NY, SPF); Datas, entroncamento Datas-­Diamantina, km 1.5, 18°22′ S, 43°41′ W, 15 Feb 1989 (fl, fr), Cavalcanti et al. 188 (CEN, RB, SPF); Diamantina, Biribiri, 25 Feb 1975 (fl, fr), Hatschbach et al. 36488 (MBM); Diamantina, Biribiri, 14 Mar 1982 (fl), Graham 879 (MBM); Diamantina, Biribiri, 22 Nov 1985 (fl), Cavalcanti et al. (CFCR) 8579 (K, NY, SPF); Diamantina, Biribiri, 28 Jan 1986 (fl), Cordeiro et al. (CFCR) 9330 (CEN, SPF); Diamantina, Biribiri, 23 Feb 1986 (fl, fr), Cavalcanti et al. (CFCR) 9488 (CEN, SPF); Diamantina, Biribiri, 14 Apr 1987 (fl, fr), Menezes et al. (CFCR) 10566 (SPF); Diamantina, BR-367, a 7 km de Conselheiro Mata, 18°20′ S, 43°45′ W, 15 Feb 1989 (fl, fr), Cavalcanti et al. 227 (CEN, SPF, K); Diamantina, estrada para Conselheiro Mata, km 192, 4 Feb 1987 (fl, fr), Mayo et  al. (CFCR) 10438 (SPF); Diamantina, estrada para Conselheiro Mata, 25 Feb 1987 (fl, fr), Zappi et al. (CFCR) 10376 (SPF); Diamantina, Área de Proteção Ambiental Pau-de-Fruta, COPASA, 18°15′01″ S, 43°39′08″ W, 13 Feb 2001 (fl), Stehmann et  al. 2896 (BHCB); Diamantina, 19 Mar 1989 (fl, fr), Salatino et al. (CFCR) 12126 (SPF); Diamantina, estrada para Conselheiro Mata, km 8.6, 18°15′ S, 43°40′ W, 15 Mar 1995 (fl), Splett 940 (CEN, UB); Diamantina, estrada para São João da Chapada, a 14 km do asfalto, 11 Apr 1982 (fl), Furlan et al. (CFCR) 3368 (SPF, K); Diamantina, São João da Chapada, 18°11′ S, 43°43′ W, 18 Feb 1989 (fr), Cavalcanti et al. 253 (CEN, SPF); Diamantina, estrada Diamantina-Biribiri, km 4, 18°12′ S, 43°37″ W, 26 Apr 1997 (fr), Cavalcanti et al. 2295 (CEN, NY); Diamantina, 10 Jan 1987 (fl), Braga s.n. (BHCB 10662); Diamantina, Água Fria, 2 Apr 1957, Pereira 2814 &

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13  Systematic Treatment

Fig. 13.23  Diplusodon ciliiflorus. A. Branch with inflorescence. B. Leaf, abaxial surface. C. Prophyll, adaxial surface. D. Prophyll, abaxial surface. E. Detail of inflorescence. F. Floral bud. G. Flower with prophylls, without petals. H. Petal. I. Fruit. J. Immature capsule. K. Seed. (A–K Cavalcanti et al. 202, CEN)

Descriptions

171

Fig. 13.24  Diplusodon ciliiflorus. A–C. Habits. D. Leaf, abaxial surface. E. Prophyll, adaxial surface. F. Flower with prophyll, without petals. (A, D, E–F Cordeiro et al. [CFCR] 9330, SPF; C Kameyama & Esteves 30, SPF; B Cavalcanti et al. [CFCR] 9488, SPF)

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Pabst 3650 (B); ca. 5 km SW de Diamantina, km 305 na BR-259, 5 Feb 1972 (fr), Anderson et al. 35457 (NY); Diamantina, estrada que passa ao longo de rio saindo da estrada Diamantina-Biribiri, 18°12′ S, 43°41′ W, 15 Feb 1989 (fl, fr), Cavalcanti et  al. 202 (CEN, SPF); Diamantina, estrada que passa ao longo de rio saindo da estrada Diamantina-Biriri, 7 km, 18°12′ S, 43°38′ W, 16 Feb 1989 (fl, fr), Cavalcanti et al. 211 (CEN); Diamantina, estrada para Biribiri, 21 Mar 1993 (fl), Kameyama & Esteves 39 (CEN, SPF); Diamantina, Diamantina-Biribiri, 18°15′ S, 43°40′ W, 13 Mar 1995 (fl), Splett 908 (CEN, UB); Diamantina, estrada Diamantina-Mendanha, km 578, 22 Mar 1995 (fl), Splett 1028 (CEN, UB); Gouveia, estrada de terra na entrada da fazenda do Sr. Everaldo, 14 Apr 1987 (fl), Scatena et al. (CFCR) 10501 (SPF); Gouveia, BR-259, Diamantina-Curvelo, torre da Telemig, 18°36′43″ S, 43°53′88″ W, 26 Apr 2004 (fl, fr), Cavalcanti et  al. 2303 (CEN [3]); Gouveia, estrada de acesso à torre da Embratel, entrada em frente à usina eólica, BR 259, Gouveia-Curvelo, 27 km, 18°36′33″ S, 43°53′58″ W, 10 Feb 2015 (fl), Cavalcanti et  al. 3941 (CEN); Estrada Congonhas do Norte-Santana do Riacho, Serra da Carapina, 18º52′S, 43º14′W, 3 Mar 1998 (fl, fr), Forzza et  al. 724 (CEN, SPF); Pedro Lessa, ca. 5 km de Trinta Réis, em direção a Pedro Lessa, km 437 da rodovia Serro-Datas, 18°33′06″ S, 43°33′14.6″ W, 13 Mar 95 (fl, fr), Souza et al. 8440 (CEN, ESA); Presidente Kubitscheck, rod. BR-259, km 433, 17 Mar 1987 (fl), Hatschbach et al. 51010 (MBM, MO); Santana do Riacho, Serra do Cipó, 19°15′ S, 43°33′ W, 23 Apr 1997, Cavalcanti et al. 2279 (CEN); Santana do Riacho, Serra do Cipó, 18 Feb 1972 (fl, fr), Anderson et al. 36221 (NY); Santana do Riacho, Serra do Cipó, rodovia Belo-Horizonte-Conceição do Mato Dentro, km 118, 14 Feb 1963 (fl, fr), Duarte 7642 (RB); Santana do Riacho, Serra do Cipó, rodovia Belo-HorizonteConceição do Mato Dentro, 5 Apr 1995 (fl), Castro 397 (CEN, HUFU); Santana do Riacho, Serra do Cipó, rodovia Belo-Horizonte-Conceição do Mato Dentro, km 115, 4 Mar 1972 (fl, fr), Joly et al. (CFSC) 777 (SP, UEC); Santana do Riacho, Serra do Cipó, rodovia Belo-Horizonte-Conceição do Mato Dentro, km 114, 26 Jan 1986 (fl, fr), Cordeiro et al. (CFSC) 9537 (SPF); Santana do Riacho, Serra do Cipó, rodovia Belo-Horizonte-Conceição do Mato Dentro, km 125, 27 Feb 1987 (fl), Zappi et  al. (CFSC) 10040 (SPF); Santana do Riacho, Serra do Cipó, rodovia BeloHorizonte-Conceição do Mato Dentro, 31 Oct 1985 (fr), Cavalcanti et al (CFSC) 9365 (SPF); Santana do Riacho, Serra do Cipó, rodovia Belo-­Horizonte-­Conceição do Mato Dentro, 22 Feb 1986 (fl, fr), Cavalcanti et al. (CFSC) 9630 (SPF); Santana do Riacho, Serra do Cipó, rodovia Belo-Horizonte-Conceição do Mato Dentro, 10 May 1987 (fr), Cavalcanti et al. (CFSC) 10159 (SPF); Santana do Riacho, Serra do Cipó, rodovia Belo-Horizonte-Conceição do Mato Dentro, 7 Sep 1987 (fr), Cavalcanti et al. (CFCS) 10615 (SPF); Santana do Riacho, Serra do Cipó, rodovia Belo-Horizonte-Conceição do Mato Dentro, km 128, 5 Mar 1972 (fl, fr), Joly et al. (CFSC) 1149 (SP); Santana do Riacho, Serra do Cipó, rodovia Belo-HorizonteConceição do Mato Dentro, km 129, 19 Apr 1950 (fl), Duarte 2638 (RB); Santana do Riacho, Serra do Cipó, caminho para a Cachoeira Capivara, 31 Mar 1988 (fl, fr), Zappi et al. (CFSC) 10927 (SPF); Santana do Riacho, Serra do Cipó, rodovia BeloHorizonte-Conceição do Mato Dentro, km 135, 21 Mar 1983 (fl, fr), Wanderley et  al. 573 (SP); Santana do Riacho, Serra do Cipó, rodovia Belo-Horizonte-

Descriptions

173

Fig. 13.25  Geographic distribution of Diplusodon chapadensis, D. ciliatiflorus, D. ciliatus, and D. ciliiflorus

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13  Systematic Treatment

Conceição do Mato Dentro, km 142, 16 Apr 1935 (fr), Mello-Barreto & Brade 1049 (RB); Santana do Riacho, Serra do Cipó, rodovia Belo-Horizonte-Conceição do Mato Dentro, ca. 18  km após Chapéu de Sol, Fazenda Cachoeira da Capivara, 19°14′59.9  S, 43°32′44.2″ W, 10 Mar 1995 (fl), Souza et  al. 8229 (CEN, ESA); Serra do Cipó, Alto do Palácio, arredores da Sede do IBAMA, 1 May 1993 (fl, fr), Souza & Sakuragui 3349 (ESA); Santo Antonio do Itambé, descida do Morro do Pico do Itambé, 18°26′ S, 43°20′ W, 6 Apr 1982 (fr), Rossi et  al. (CFCR) 3078 (CEN, SPF); Santo Antonio do Itambé, Pico do Itambé, 18°24′ S, 43°19′ W, 7 Apr 1998, Souza et al. 21113 (CEN, ESA, HUEFS); São Gonçalo do Rio Preto, Parque Estadual do Rio Preto, 18°05′28″ S, 43°20′32″ W, 7 Apr 2000 (fl), Lombardi et al. 3757 (BHCB, CEN), Serro, rodovia MG-2, 19 Jan 1972 (fl), Hatschbach et  al. 28931 (MBM). The specific epithet alludes to the principal characteristic of this species, which is the presence of cilia on the sepals and epicalyx, with the rest of the flower being glabrous. Additionally, this species can be recognized by its nitid leaves and showy flowers with 12 stamens. Diplusodon ciliiflorus is commonly found as a subshrub 0.5–1 m tall, with lax, decussate branches and expanded leaves. However, in Diamantina, in areas subject to frequent fires, for example, in the region near Conselheiro Mata, one can find sprouts of D. ciliiflorus with a very different aspect from plants that are not burned. The individuals of these burned areas, when flowering, are caespitose, ca. 15 cm tall, with narrower leaves on short branches and other much longer leafless, carbonized branches (Cavalcanti et al. (CFCR) 9488). Such plants, although possessing a reduced habit and narrower leaves, have the characteristic flower features of D. ciliiflorus. Moreover, this provides good evidence of how much this species varies with relation to habit, ramification, and leaf width. The type material of Diplusodon ciliiflorus, Gardner 4569, is represented by three specimens, two at K and one at W.  One specimen, deposited at B, was destroyed and may have been the specimen studied by Koehne. A photograph of this specimen was examined, but there were no annotations by Koehne on it (F neg. nº 17920), as was the case with the two specimens deposited at K. The W specimen was identified by Koehne in 1901 and corresponds well to the original description of D. ciliiflorus, based on fruiting material. Gardner 4569 at W (W0010988) is here designated as the second-step lectotype of D. ciliiflorus. 21. Diplusodon conduplicatus T.B.Cavalc., sp. nov. Type: Brazil. Goiás: Cavalcante, RPPN Serra do Tombador, 3  km  S da sede, área conhecida como “Campina”, 13°40′51″S, 47°49′06″W, 31 Jan 2015 (fl, fr), M. F. Simon, J. M. Mendoza, J.  B. A.  Bringel Jr., T.  K. M.  Arquelão 2529 (holotype, CEN [00092280]; isotypes, K, NY, RB, SPF, W). Figs. 13.26 A–F, 13.30 Species with unique characteristics, with leaves and prophylls completely folded lengthwise enveloping the branch, venation acrodromous-basal with almost parallel veins and margins with long and fine cilia. The prophylls exceed the length of floral tube, and the androecium has 12 stamens.

Descriptions

175

Subshrubs 0.15–0.35  m tall, erect, unbranched, with a xylopodium, not darkbrown when dry. Upper branches subquadrangular, unwinged, yellow, not darkbrown when dry, glabrous, internodes 0.3–13 cm long. Leaves decussate, imbricate, congested, erect, sessile; blades 9–14 × 1.5–3 mm (lamina folded), glaucous, coriaceous, not discolorous, acrodromous-basal, narrowly lanceolate, folded lengthwise, keeled, midvein yellowish, long-ciliate, base retuse, amplexicaulous, apex acute, sharp, margin plane, strongly ciliate with long and fine trichomes; secondary veins 4 on each side, departing from the base of the midvein, almost parallel; domatia absent. Synflorescence frondose, botryum type, congested; accessory branches absent; bracts similar to the vegetative leaves in shape, decreasing in size toward the apex of the branch; flowers pedicellate; prophylls ca. 9 × 1–1.2 mm (folded), surpassing the apex of the floral tube by ca. 1  mm, glaucous, narrowly lanceolate, folded lengthwise, keeled, base retuse, amplexicaulous, apex acute, sharp, margin plane, strongly ciliate with long and fine trichomes; floral tube ca. 7 mm long, lightgreen, narrowly oblong; sepals ca. 2 mm long, triangular, slightly caudate, deflexed, long-ciliate; epicalyx segments ca. 1.5 mm long, narrowly triangular, dorsiventrally compressed, erect, shorter than the sepals, long-ciliate; corolla ca. 2.5 cm in diam., lilac, petals 11–11.5 × 4.5–5 mm, obovate, apex obtuse; stamens 12, filaments ca. 8 mm, exserted ca. 5 mm; ovary ca. 2.2 × 1.8 mm, obovate-oblong, glabrous, style ca. 12.5 mm long, ovules ca. 24. Capsules ca. 3 × 2.5 mm, globose, glabrous; seeds ca. 5, 1.7–1.8 × ca. 1.5 mm. Distribution and Ecology (Fig. 13.30)  Diplusodon conduplicatus is endemic to Serra do Roncador, region of Chapada dos Veadeiros, occurring in campo sujo at 805 m elevation. It was collected with flowers and immature fruits in January. Specimen Examined. BRAZIL.  The type. Diplusodon conduplicatus is a remarkable new species from Chapada dos Veadeiros region in Goiás. The species is represented by small (0.15–0.35 m tall) and unbranched subshrubs readily distinguished from any other species of the genus by its unusual leaves that are completely folded lengthwise and enveloping the stem. Furthermore, the margins of leaves, bracts, sepals, and epicalyx segments are strongly ciliate with long, fine trichomes, the leaf venation is the acrodromous-basal type with veins almost parallel, the prophylls are folded lengthwise exceeding the apex of the floral tube, and the androecium has 12 stamens. An interesting aspect of this species is that only half of the leaves are seen in the habit because the blades are completely folded, so that the leaves do not appear to have a midrib. The midrib, which is long-ciliate, appears to be the ciliate margin of an asymmetric leaf. 22. Diplusodon cordifolius Lourteig, Bradea 5(19): 222. 1989. Type: Brazil. Goiás: Alto Paraíso de Goiás, Chapada dos Veadeiros, 21 Mar 1969 (fl), H. S. Irwin, R. R. Santos & S. F. Fonseca 24807 (holotype, P [01901910]; isotypes, MO, NY [2], UB). Figs. 13.27 A–G, 13.30

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Fig. 13.26  Diplusodon conduplicatus. A. Habit. B. Section of internode. C. Vegetative branch. D. Leaf (folded lengthwise), showing half of adaxial surface. E. Prophyll. F. Flower with prophylls, without petals. (A–F Simon et al. 2529, CEN)

Descriptions

177

Shrubs 0.3–0.8 m tall, sometimes prostrate, with a xylopodium, much-branched, canescent, not dark-brown when dry. Upper branches cylindrical to subquadrangular, unwinged, densely covered with simple, long trichomes, internodes 5–6  mm long. Leaves decussate, not imbricate, suberect, sessile; blades 6–10(−12) × 1.5– 3.5 mm, coriaceous, not discolorous, eucamptodromous, lanceolate to oblong, usually with waist in the middle region, base cordate, apex acute, margin revolute; secondary veins 2–3 on each side of the midvein, prominent on the abaxial surface, adaxial and abaxial surfaces hirsute, covered with simple, long trichomes, rarely the indumentum very dense and silvery; domatia evident. Synflorescence frondose, diplobotryum to pleiobotryum type, congested; accessory branches absent, principal florescence ca. 4 cm long, botryum type, sometimes caducous, paraclades 3–15 cm long, bracts similar to the vegetative leaves in form and size; flowers pedicellate, hypopodium 1–1.5  mm long, epipodium 0.5–1.5  mm long; prophylls 3–5.5 × 1.5–3 mm, reaching the apex of the floral tube or slightly less, elliptic-lanceolate, unkeeled, base obtuse, apex acute, margin plane, ciliate, covered with long, sparse trichomes; floral tube 5–5.5 mm long, greenish, funnel-shaped, covered with simple, long trichomes; sepals 3–5.5 mm long, wine-colored, triangular, sometimes with a dorso-elongated apex, not deflexed, with long, sparse trichomes, ciliate; epicalyx segments 0.5–2 mm long, dorsiventrally compressed, spreading to suberect, ciliate, shorter than the sepals; corolla ca. 2 cm in diam., lilac to bright rose, petals 8–9 × 4.5–5 mm, obovate, apex obtuse to sometimes acute; stamens 18, free portion of the filaments 6.5–9  mm long, exserted, ovary 1.9–2.1  ×  2–2.5  mm, obovatedepressed, glabrous, style 10–12.5 mm long, ovules 13–19. Capsule ca. 3.5 × 3.2 mm, globose, glabrous; seeds 10–15, 1.8–2.5 × 1.2–2 mm. Distribution and Ecology (Fig. 13.30)  Diplusodon cordifolius is endemic to the mountains of Alto Paraíso de Goiás, São João da Aliança, and Cavalcante, in Chapada dos Veadeiros, Goiás. The species occurs in campos rupestres, campos limpos, and cerrados on rocky soil at 1000–1539 m elevation. Flowers from February to September; fruits from August to April. Specimens Examined: BRAZIL.  Goiás  Alto Paraíso de Goiás, Chapada dos Veadeiros, próximo ao Grande Hotel Paraíso, 6 Sep 1995 (fl, fr), Silva et al. 2267 (IBGE); Alto Paraíso de Goiás, 2 km de Veadeiros, 18 Jul 1964 (fl), Prance & Silva 58203 (K, NY, UB); Alto Paraíso de Goiás, Chapada dos Veadeiros, 26 Jul 1985 (fl), Romaniuc Neto & Sajo 342 (CEN, SP); Alto Paraíso de Goiás, ca. 10 km de Alto Paraíso na estrada para Teresina de Goiás, entre os km 174 e 176 da GO-118, 28 May 1994 (fl), Proença & Silva 1196 (UB); Alto Paraíso de Goiás, 16 km de Alto Paraíso para Teresina de Goiás, 14°9″ S, 47°31″ W, 29 Jul 1994 (fl), Silva et al. 2137 (IBGE); Alto Paraíso de Goiás, GO-118, Alto Paraíso-Teresina de Goiás, 12 km de Alto Paraíso, 14°03′03″ S, 47°30′48″ W, 17 Nov 2005 (fr), Cavalcanti et al. 3609 (CEN, MO); Alto Paraíso de Goiás, Chapada dos Veadeiros, 24 km de Alto Paraíso/ Nova Roma, próximo à Fazenda Parida, 14°04′15″ S, 47°19′25″ W, 19 Jul 1994 (fl), Silva et al. 2163 (IBGE, P, UEC); Alto Paraíso de Goiás, estrada para Nova Roma, rio Bartolomeu, 13 Jun 1993 (fl), Hatschbach et al. 59477 (MBM, MO); Cavalcante, Fazenda Renascer, 20 Apr 2003 (fl), Pastore et al. 468 (CEN); Cavalcante, povoado Kalunga, caminho para a cachoeira Santa Bárbara, 6 Mar 2003 (fl, fr), Pastore &

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Fig. 13.27  Diplusodon cordifolius. A. Branch with inflorescence. B. Leaf, adaxial surface. C. Detail of leaf abaxial surface indumentum. D. Floral bud. E. Prophyll, adaxial surface. F. Flower with prophylls, without petals. G. Simple trichome. (A–G Irwin et al. 24807, UB)

Descriptions

179

Suganuma 446 (CEN); Cavalcante, caminho para o povoado Kalunga, 3 Feb 2004 (fl, fr), Pastore et al. 838 (CEN); Cavalcante, estrada de terra Cavalcante-Kalunga, 18 km, 13°38′42″ S, 47°28′39″ W, 18 Nov 2005 (fr), Cavalcanti et al. 3632 (CEN, MO); Cavalcante, estrada para Arai, 13°38′57.9″ S, 47°28′29.2″ W, 13 Apr 2004 (fl, fr), Fonseca et al. 5075 (CEN, IBGE); Cavalcante, estrada Cavalcante-Minaçu, km 20, 13°38′42.8″ S, 47°28′39.7″ W, 31 Aug 2004 (fl, fr), Cavalcanti et al. 3527 (CEN, NY, P, SPF); São João da Aliança, periferia da cidade, junto ao campo de aviação, 14°43′13″ S, 47°31′11″ W, 21 May 2001 (fl), Pereira-Silva et al. 5045 (CEN, MO); São João da Aliança, periferia da cidade, junto ao campo de aviação, 14°13′15″ S, 47°31′18″ W, 28 May 2002 (fl), Cavalcanti & Pereira-Silva 2720 (CEN, MO); São João da Aliança, 10 Aug 1947 (fr), Oliveira s.n. (BHCB 41712). Diplusodon cordifolius is characterized by its reduced, lanceolate leaves ca. 1 cm long, with revolute margins and a cordate base. The leaves are covered by long trichomes, and domatia are present between the veins. The flowers are ca. 2 cm in diam., and the floral tube is covered by long trichomes; the epicalyx segments are short (0.5–2 mm long), suberect, and shorter than the sepals; and the androecium has 18 stamens. 23. Diplusodon cryptanthus T.B.Cavalc., Bol. Bot. Univ. São Paulo, 22(1): 3, fig. 2. 2004. Type: Brazil. Goiás: Minaçu, 2 km ao N do antigo aeroporto do canteiro de obras do UHE Serra da Mesa, 13°28′ S, 48°23′ W, 9 Mar 1992 (fl), T.B.Cavalcanti, B. M. T. Walter, D. M. S. Rocha, H. G. P. Santos, S. P. C. Silva, N. P. V. Batista 1053 (holotype, CEN [00017898]; isotypes, RB, SPF, UFG). Figs. 13.28 A–G, 13.30 Shrubs 0.6–1(−2) m tall, erect, much-branched, with a xylopodium, glabrescent, totally glabrous when mature, corymbiform, not dark-brown when dry. Upper branches slender, quadrangular, with small wings of 0.1–0.2 mm wide, internodes 1.5–3  cm long. Leaves decussate, not imbricate, suberect, subsessile; petioles 1.5–2 mm long; blades 40–57 × 13–25 mm, not glaucous, chartaceous to membranaceous, not discolorous, acrodromous with typical base, elliptic-lanceolate, base acute to obtuse, apex acute, margin plane, glabrous, adaxial surface with small black dots; secondary veins 2 on each side, 1 pair departing from the base of the midvein, inconspicuous on the adaxial surface, prominent on the abaxial surface; domatia evident. Synflorescence frondose, triplobotryum to pleiobotryum type, lax; accessory branches absent, principal florescence 4–7 cm long, botryum type, paraclades 10–25 cm long, bracts 10–50 × 6–20 mm, elliptic to ovate-elliptic, similar to the vegetative leaves in shape; flowers pedicellate, hypopodium 15–25 mm long, epipodium 2–2.5 mm long; prophylls 13–25 × (4–) 9–12 mm, sometimes hiding the floral tube, ovate, base and apex obtuse, margin plane; floral tube 7–8 mm long, narrowly oblong, glabrous; sepals 2–3 mm long, wine-colored, triangular, not caudate, not deflexed; epicalyx segments 6–8 mm long, green, linear, slightly dorsiventrally compressed, erect, surpassing the sepals; corolla ca. 4.5 cm in diam., rose to purple, petals 16–16.5 mm long, obovate, apex obtuse; stamens 18, free portion of the filaments 7–8 mm long, exserted ca. 2 mm; ovary 2.5–3 × 3–4 mm, rhombic, glabrous, style 14–15  mm long, ovules 26–35. Capsules globose, glabrous; seeds 19–30, 1.5–2.1 × 2–3 mm.

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Distribution and Ecology (Fig.  13.30)  Diplusodon cryptanthus is limited to northern Goiás and southern Tocantins, in the depression at the margin of Rio Tocantins, in sandy soil with milky quartz and sandy clay with gravel, at 280–560 m. Flowers from January to April; fruits from February to October. Specimens Examined: BRAZIL.  Goiás,  Campinaçu, depois de Buriti, 18  km, estrada vicinal, 13°31′08″ S, 48°11′14″ W, 5 Oct 2000 (fr), Cavalcanti et al. 2661 (CEN, MO); Campinaçu, estrada de asfalto Campinaçu-Minaçu, 13°36′42″ S, 48°20′21″ W, 4 Oct 2000 (fr), Cavalcanti et  al. 2595 (CEN, MO); Campinaçu, estrada Vila Veneno-Serra Branca, km 3.6 (margem direita do rio Macacão), 13°34′25″ S, 48°04′20″ W, 20 Feb 2002 (fr), Pereira-Silva et al. 5898 (CEN, MO); Cavalcante, ca. 4  km da vila Veneno em direção ao rio São Felix, 13°31′36″ S, 48°03′45″ W, 25 Jan 2001 (fl), Pereira-Silva et al. 4655 (CEN, MO); Cavalcante, ca. 3.5 km canteiro da obra para o rio do Carmo, 13°23′58″ S, 48°07′25″ W, 23 Jan 2001 (fl), Pereira-Silva et  al. 4585 (CEN, MO); Cavalcante, ponte sobre os rios Tocantins-Cavalcante, km 1.2, 13°23′43.7″ S, 48°08′16.1″ W, 30 Aug 2004 (fr), Cavalcanti et al. 3509 (CEN, SPF); Cavalcante, 2 km à leste da ponte sobre o rio Tocantins, na direção de Cavalcanti, 13°23′43″ S, 48°07′56″ W, 20 Feb 2002 (fl), Pereira-Silva et al. 5871 (CEN, MO); Cavalcante, canteiro da obra, rio do Carmo, km 1, próximo a porteira trancada, 13°23′43″ S, 48°07′56″ W, 10 Jul 2001 (fr), Pereira-Silva et al. 5226 (CEN, MO); Cavalcante, canteiro de obras do AHE Cana Brava, próximo à estação de tratamento de água, 22 May 2001 (fr), Pereira-Silva et al. 5057 (CEN, MO, NY); Cavalcante, estrada para Limoeiro, ca. 20 km da balsa (Porto dos Paulistas) rio Tocantins, 13°26′12″ S, 48°03′44″ W, 21 Feb 2001 (fl), Pereira-Silva et al. 4718 (CEN, MO); Cavalcante, estrada para Limoeiro, ca. 28 km da balsa (Porto dos Paulistas) rio Tocantins, 13°29′34″ S, 48°03′04″ W, 21 Feb 2001 (fr), Pereira-Silva et al. 4736 (CEN, MO); Colinas do Sul, Fazenda Cigarra, próximo ao rio Tocantins, 13°38′27″ S, 48°08′31″ W, 19 Apr 2001 (fl), Pereira-Silva et al. 4991 (CEN, MO). Tocantins, Palmeirópolis, rodovia Minaçu-Palmeirópolis, km 57 após Minaçu, 13°03′34″ S, 48°23′34″ W, 21 Feb 2002 (fl), Pereira-Silva et al. 5908 (CEN, MO); Palmeirópolis, balsa do coronel Valente-Palmeirópolis, km 8, 13°01′5″ S, 48°11′25″ W, 11 Jun 2006 (fr), Pereira-Silva et  al. 10594 (CEN); Paranã, estrada São Salvador-balsa do rio Custódio, km 43, 12°57′01″ S, 48°08′19″ W, 10 Jun 2006 (fr), Pereira-Silva et al. 10536 (CEN); Paranã, estrada São Salvadorbalsa do rio Custódio, km 30, 12°52′22″ S, 48°10′49″ W, 10 Jun 2006, Pereira-Silva et al. 10533 (CEN); Paranã, margem direita do rio Tocantins, a norte da balsa do Coronel, 13°00′30″ S, 48°08′43″ W, 24 Jan 2008, Pereira-Silva & Moreira 12877 (CEN); Paranã, ca. 2 km a norte da vila Rosario, no sentido São Salvador, 12°57′ S, 48°08′19″ W, 26 Mar 2007, Pereira-Silva & Moreira 11605 (CEN); Paranã, fazenda Alegre, 13°06′15″ S, 48°05′59″ W,15 May 2007 (fr), Pereira-Silva et  al. 11813 (CEN); São Salvador do Tocantins, margem esquerda do lago da UHE São Salvador, 12°50′37″ S, 48°16′07″ W, 24 Mar 2009 (fl, fr), Pereira-Silva et al. 14144 (CEN). Diplusodon cryptanthus is represented by erect, much-branched shrubs, readily recognized by its glabrous, elliptic-lanceolate, acrodromous leaves with prominent domatia and by its long (to 2.5 cm) pedicel. Sometimes the prophylls are broad and

Descriptions

181

Fig. 13.28  Diplusodon cryptanthus. A. Branch with inflorescence. B. Section of internode. C. Leaf, abaxial surface. D. Flower without petals, hidden by prophylls. E. Flower without petals, without one prophyll. F. Petal. G. Pistil. (A–G Cavalcanti et al. 1053, CEN)

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ovate and totally hide the floral tube when the specimen is pressed. Another characteristic of this species is the presence of small black dots on the leaf surface, which are easily seen with a hand lens. Anatomically, the black dots are mucilaginous secretory glands distributed among the epidermal cells. 24. Diplusodon decussatus Gardner & Fielding, Sert. Pl., fig. 1. 1844. Type: Brazil. Tocantins: Campos Arrayas, Mar 1840 (fl, fr), G. Gardner 3722 (lectotype, OXF [00005272], designated by Lourteig 1989: 235, isolectotypes, F, fragm. F, G [3], K [3], NY [2], P [3], W [2]). Figs. 13.29 A–G, 13.30 Subshrubs 0.3–1  m tall, erect to sometimes prostate, unbranched to slightly branched, with a xylopodium, glabrous, usually dark-brown when dry. Upper branches cylindrical to subquadrangular, unwinged, internodes 1–3 cm long. Leaves decussate, not imbricate, spreading, sessile; blades 10–22  ×  17–35  mm, strongly glaucous, coriaceous, not discolorous, acrodromous-basal-supranumerary, triangular-deltoid, base retuse or subcordate to cordate, apex obtuse, sometimes slightly acuminate, margin plane, slightly thickened and yellow to slightly wine-colored, adaxial surface with inconspicuous veins, abaxial surface with veins more apparent; secondary veins 5–9 on each side, most of them departing from the base of the midvein, 1–2 departing along the midvein; domatia absent. Synflorescence frondose, botryum to diplobotryum type, congested; accessory branches absent, principal florescence 10–15 cm long, paraclades 8–12 cm long; bracts similar to the vegetative leaves in shape and size; flowers subsessile, hypopodium 0.5–1 mm long, epipodium up to 0.5 mm long; prophylls 4.5–6.5(−8) × 3–5.5 mm, reaching to the middle of the floral tube or less, elliptic or ovate to cordate, slightly keeled, margin plane; floral tube 6.5–8 mm long, glaucous, in fruit dark chestnut brown, nitid, 8–9 mm long oblong; sepals 2.5–3.5 mm long, triangular, not caudate, not deflexed, sometimes just the top deflexed; epicalyx segments 2–2.5 mm long, slightly wine-colored, cylindrical to flattened, erect to suberect, the same size as the sepals; corolla ca. 3.5 cm in diam., lilac to rose-magenta, petals ca. 20 × 12 mm, narrowly obovate, apex obtuse; stamens 12, free portion of the filaments 7–11  mm long, slightly exserted; ovary 2.5–3  ×  2.5–3  mm, yellow-green, rhombic, glabrous, style 17–18 mm long, rose-magenta, ovules 17–33. Capsule 4–5 × 3 mm, dark-brown, oblong, glabrous; seeds 5–16, 2.5–3.3 × 1.8–2 mm. Distribution and Ecology (Fig.  13.30)  Diplusodon decussatus is known from very few herbarium specimens. This species is endemic to northern Goiás and southern Tocantins, on Serra Geral do Paranã. It inhabits campos rupestres, campos sujos with quartzite sand, campos limpos and cerrados, at 750–1200 m elevation. Flowers from November to March; fruits from November to April. Specimens Examined: BRAZIL. Goiás,  Cavalcante, estrada para a Cachoeira do Prata, 6 Mar 2003 (fl, fr), Pastore & Suganuma 448 (CEN); Cavalcante, estrada de terra Cavalcante-Kalunga, 18 km, 13°39′34″ S, 47°28′32″ W, 18 Nov 2005 (fl, fr), Cavalcanti et al. 3626 (CEN, MO); Cavalcante, Comunidade Kalunga Engenho II, Caminho para a cachoeira Santa Bárbara, 13°34′57″ S, 47°28′21″ W, 10 Jan 2010,

Descriptions

183

Fig. 13.29  Diplusodon decussatus. A. Habit. B. Leaf, abaxial surface. C. Floral bud. D. Prophyll, adaxial surface, detail of leaf abaxial surface indumentum. E. Flower with prophylls, without petals. F. Fruit. G. Capsule. (A, C–D Cavalcanti et al. 2248, CEN; B, E–G Pastore & Suganuma 448, CEN)

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Faria-Jr. 736 (CEN, UB); entre Cavalcante e Conceição, Burchell 8039 (BR, K, LE n.v., P). Tocantins, Arraias, na periferia da cidade de Arraias, morro do lado Oeste, na base do morro, 12°56′17″ S, 46°56′53″ W, 9 Apr 1997 (fr), Cavalcanti et al. 2248 (CEN, MO, NY). Diplusodon decussatus is represented by unbranched to slightly branched subshrubs. The species is also characterized by its glaucous and triangular-deltoid leaves that turn dark-brown when dry. The plants are not completely erect, and the branches are somewhat prostrate or decumbent. The glabrous and glaucous aspect of the leaves is similar to that of D. imbricatus, but D. imbricatus differs by its lanceolate leaves with a cordate base and flowers with 18, rather than 12, stamens. The seedlings sometimes have trichomes on the leaves. There are fifteen duplicates of the type (Gardner 3722) distributed among at least seven herbaria. Lourteig (1989) indicated a specimen deposited at OXF as holotype, which is considered here as the first-step lectotype (ICBN Art. 9.17, ex. 14). 25. Diplusodon divaricatus Pohl, Flora 10: 130. 1827. Type: Brazil. Goiás: in montosis graminosis inter Corumbá et rio São Marcos, Dec 1818 (fl), J.  B. E. Pohl 2537 = D 939 (first-step lectotype, designated by Lourteig 1989: 241, second-step lectotype, W [0053402], here designated; isolectotypes, K, W [0069491]). Figs. 13.31 A–J, 13.35 Subshrubs 0.5–1 m tall, erect, slightly branched, with a xylopodium, not darkbrown when dry. Upper branches rust-colored, cylindrical, unwinged, with a dense short indumentum formed by bulbous glandular trichomes, internodes 1.5–3.5 cm long. Leaves decussate, not imbricate, suberect, sessile; blades 15–45 × 10–38 mm, coriaceous, not discolorous, acrodromous-basal-supranumerary, ovate to ovate-­ lanceolate, base obtuse to cordate, apex acute to obtuse, sometimes acuminate, margin plane, adaxial and abaxial nearly glabrous, very slightly pilose, trichomes short, most easily seen along the veins; secondary veins 4–6 on each side, 2–4 departing from the base of the midvein; domatia absent or rarely with faint domatia. Synflorescence frondose-bracteose to bracteose, botryum spiciform to triplobotryum type, congested; accessory branches absent, principal florescence not seen, paraclades 2–6 cm long; bracts 5–15 × 4–10 mm, ovate, ovate-lanceolate to orbicular; flowers subsessile, hypopodium ca. 0–0.5 mm long, epipodium 0–0.5 mm long; prophylls 3–3.5  ×  2–3  mm, not reaching the middle of the floral tube, ovate to orbicular, unkeeled, concave, base obtuse, apex obtuse, margin plane, densely pilose, trichomes bulbous, white; floral tube 6–7 mm long, golden in color, infundibuliform, densely covered with bulbous glandular trichomes; sepals 3.5–4  mm long, golden in color, triangular, not caudate, sometimes slightly deflexed, covered with golden botuliform glandular trichomes; epicalyx segments to 0.5  mm long, inconspicuous, cylindrical, shorter than the sepals; corolla 3.5–4.5  cm in diam., rose-magenta or lilac to red, petals 19–20 × 10–11 mm, elliptic, apex obtuse; stamens 18–23, free portion of the filaments 7–7.5 mm long, slightly exserted; ovary

Descriptions

185

Fig. 13.30  Geographic distribution of Diplusodon conduplicatus, D. cordifolius, D. cryptanthus, and D. decussatus

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2–2.5 × 3–4 mm, rounded, glabrous, style 10–14 mm long, ovules 29–50. Capsules 5–5.6 × 4–6 mm, globose, glabrous; seeds 15–44, 1.8–2 × 1.8–2 mm. Distribution and Ecology (Fig.  13.35)  Diplusodon divaricatus is distributed in the cerrados (sensu stricto) and campos sujos of the mountains around Caldas Novas, Goiás, on quartzite soils. Koehne (1903) also listed this species from Catalão, Goiás, and from Araxá and Paracatu (Saint-Hilaire s.n.), and Uberlândia, Minas Gerais. Flowers from February to August; fruits from May to November. Specimens Examined: BRAZIL. Goiás,  Caldas Novas, Pousada do Rio Quente, 17 Aug 1972 (fr), Mello-Filho 3653 (R); Caldas Novas, Alto da Serra de Caldas Novas, 25 Apr 1970, Rizzo 5113 & Barbosa 4362 (UFG); Caldas Novas, 23 May 1970 (fl), Rizzo 5206 & Barbosa 4451 (UFG); Caldas Novas, 27 Jun 1970 (fl), Rizzo 5327 & Barbosa 4576 (UFG); Caldas Novas, Parque Estadual da Serra de Caldas, 30 Jul 1993 (fl), Santos et al. 220 (CEN, MO); Caldas Novas, Parque Estadual da Serra de Caldas, próximo ao mirante, 29 Nov 2000 (fr), Amaral-Santos et al. 871 (CEN, MO); Caldas Novas, Serra de Caldas Novas, 17°46′ S, 48°39′ W, 27 Sep 1995 (fr), Cavalcanti et al. 1734 (CEN); Caldas Novas, Serra de Caldas Novas, 29 Aug 1997 (fl), Santos et al. 509 (CEN, MO); Caldas Novas, Alternativa 15-Ponto de Hidrometria, 27 Apr 1993 (fl), Cordovil-Silva et al. 276 (CEN, MO); Caldas Novas, área de empréstimo da UHE Corumbá, perto da pista do antigo aeroporto, 1 Jun 1998 (fl), Cavalcanti et al. 2398 (CEN, MO, NY); Corumbaíba, margem esquerda do rio Corumbá, 24 Jun 1993, Santos et  al. 106 (CEN); Corumbaíba, margem esquerda do rio Corumbá, 1.5 km à montante do eixo da barragem, próximo a foz do córrego Gameleira, 25 May 1993 (fl, fr), Santos et al. 2 (CEN, MO, NY); Ipameri, mirante da Fazenda Santo Antonio do Fundão, 17°43′ S, 48°10′ W, 24 Mar 1995 (fr), Cavalcanti et al. 1402 (CEN, MO, NY); Ipameri, no mirante da Fazenda Santo Antonio do Fundão, 8 Aug 1996 (fl, fr), Cavalcanti et al. 1686 (CEN, MO, NY); Morrinhos, estrada de Morrinhos para Caldas Novas, córrego Samambaia, 25 Jul 1970 (fl, fr), Rizzo 5413 & Barbosa 4662 (UFG). Minas Gerais, Paracatú, Saint Hilaire s.n. (P); Uberlândia, Reserva Ecológica do Clube de Caça e Pesca Itororó, 17 May 1994 (fl), Romero et al. 1009 (CEN, HUFU). Diplusodon divaricatus occurs in southern Goiás and southwestern Minas Gerais. The plants are subshrubs slightly branched recognized by its ovate to ovatelanceolate leaves, which although appearing to be glabrous, are covered with short trichomes. Distinguishing floral characteristics are small, short, ovate to orbicular prophylls that do not reach as far as the middle of the floral tube and the orange- to golden-colored floral tube, which is covered with an abundance of bulbous glandular trichomes secreting an orange substance. The epicalyx segments are small (to 0.5 mm long), and the androecium has 18–23 stamens. Plants of Diplusodon divaricatus are similar to those of D. sessiliflorus, in that the latter species also has bulbous, orange secretory trichomes on the flowers. Diplusodon sessiliflorus differs from D. divaricatus because it has leaves and branches with long trichomes forming a villous indumentum, broad prophylls covering the floral tube, and epicalyx segments 2–4 mm long.

Descriptions

187

Fig. 13.31  Diplusodon divaricatus. A. Habit. B. Section of internode. C. Leaf, abaxial surface. D. Detail of leaf abaxial surface indumentum. E. Glandular trichomes of floral tube. F. Floral bud. G. Prophyll, adaxial surface. H. Flower with prophylls, without petals. I. Pistil. J. Fruit. (A–I Rizzo 5113 & Barbosa 4362, UFG; J Mello-Filho 3653, R)

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13  Systematic Treatment

Diplusodon divaricatus was described by Pohl (1827a) without citation of specimens in the protologue. In another publication, Pohl (1827b) referred to a single collection of D. divaricatus collected in December in the mountains between Corumbá and Rio São Marcos, Goiás, without mentioning his collection number. Lourteig (1989) cited Pohl s.n. (holotype W; isotype K) for Diplusodon divaricatus. At W there is a specimen with this same locality, and annotated by Pohl, numbered as Pohl 2537 = D939, which is here selected as the second-step lectotype. In addition, there is a second specimen of Diplusodon divaricatus at W with a label indicating that it is Pohl 5793, collected in dry campos near Paracatu, Minas Gerais, and Serra dos Cristais, Santa Luzia [now Luziania], Goiás. Pohl 5793 is the type of D. alutaceus Pohl, a synonym of D. lanceolatus Pohl, whose label has exactly the same locality as that of the D. divaricatus specimen. This specimen is clearly a duplicate of Pohl 2537  =  D939, as already noted by Koehne on the specimen. Pohl’s Brazilian collections were stored in Vienna in a museum called “Brasilianisches Museum” The first sheet of a collection got the original label and the duplicates were stored without labels. After building the “Naturhistorische Museum” the botanical collections kept in the “Brasilianisches Museum” were incorporated in the collection of Herbarium W.  Unfortunately, some of the unlabeled duplicates were mixed up (Bruno Wallnöfer, pers. comm). In this case, it is clear that such confusion happened, and W0069491 is a duplicate of W0053402. 26. Diplusodon epilobioides Mart. ex DC., Prodr. 3: 94a. 1828. Friedlandia epilobioides Mart. ex DC., Prodr. 3: 94b, 1828, pro syn. Type: Brazil. Minas Gerais: Habitat in campis Serro Frio ad Tejuco, Jul 1820 (fl), C. F. P. Martius s.n. (firststep lectotype, designated by Lourteig 1989: 228, second-step lectotype, M [0146281], here designated; isolectotypes, M [0146282], NY, SPF, G). Figs. 13.32 A–H, 13.35 Shrubs 0.5–3  m tall, erect, much-branched, not dark-brown when dry. Upper branches lax, cylindrical to subquadrangular, unwinged, with minute, dense, inconspicuous trichomes, internodes 0.5–1  cm long. Leaves decussate, not imbricate, erect, short-petiolate; petioles 1.5–2 mm long; blades 10–22 × 1–4 mm, coriaceous, slightly canescent, sometimes slightly discolorous, eucamptodromous, narrowly elliptic to narrowly oblong, base long-attenuate, apex obtuse to acute, margin revolute, leaves covered by minute, dense, inconspicuous trichomes, giving a velvety appearance, adaxial surface with sulcate veins, abaxial surface slightly canescent, with prominent veins; secondary veins 1(−2) on each side of the midvein; domatia present. Synflorescence frondose, diplobotryum to pleiobotryum type, lax; accessory branches absent, principal florescence 4–10 cm long, botryum type, paraclades 4–20  cm long; bracts similar to the vegetative leaves in shape, slightly smaller toward the apex of the branches; flowers pedicellate, hypopodium 2–4 mm long, epipodium 2–3 mm long; prophylls 5–7 × 1–1.5 mm, usually surpassing the apex of the floral tube 1–2 mm, sometimes not reaching the apex of the floral tube, inserted in the middle of the pedicel, narrowly elliptic or spathulate to lanceolate, unkeeled,

Descriptions

189

base attenuate, apex obtuse, margin plane to involute, covered by erect, short trichomes; floral tube 3–4.5  mm long, green, funnel-shaped, covered by minute, inconspicuous trichomes, sometimes appearing glabrous; sepals 2.5–3  mm long, wine-colored, triangular, caudate, not deflexed, with the same type of indumentum as the floral tube; epicalyx segments 3–4 mm long, cylindrical, erect, surpassing the sepals, with the same type of indumentum as the floral tube; corolla 2–2.5 cm in diam., lilac or rose-magenta to red, petals 6.1–9.5  ×  5–7.5  mm, obovate, apex obtuse; stamens 12–18, free portion of the filaments 5–9 mm, exserted, surpassing the apex of the sepals 3–3.5 mm; ovary ca. 2 × 1.5–2.3 mm, rhombic, glabrous, style 10–13 mm long, ovules 10–16. Capsules 2.8–3.1 × 2–2.1 mm, globose, glabrous; seeds 5–7, 1.5–2 × 1.8–2.9 mm. Distribution and Ecology (Fig.  13.35)  Diplusodon epilobioides is endemic to northern Cadeia do Espinhaço, Minas Gerais, occurring in cerrado and campos ­rupestres between 700–1150 m, in “carrascal” to 870 m, and in sandy soils with plinthosol of hematite to 1030  m. Flowers from July to November; fruits from September to February. Specimens Examined: BRAZIL.  Minas Gerais,  Berilo, Usina Hidrelétrica Irapé, 16°45′8″ S, 42°34′14″ W, 2 Sep 2003 (fl), Cavalcanti & Pereira-Silva 3195 (CEN, MO); Berilo, 19 Jul 1991, Carvalho & STS 378 (CEN, BHCB, IBGE); Cristália, arredores, 23 Jul 1978 (fl), Hatschbach 41484 (MO, NY); Fazenda Cabral, 17 Jul 1991 (fl), Carvalho & STS 339 (CEN, BHCB); Grão-Mogol, estrada do Cardoso, NE da cidade, 9 Jul 1985 (fl), Cerati et al. 254 (CEN, SP, SPF); GrãoMogol, estrada do Cardoso, ca. 6 km NE de Grão-Mogol, 9 Jul 1985, Kral et al. 72695 (CEN, SPF); Grão-Mogol, estrada para Montes Claros, ca. 20 km de GrãoMogol, 16°33′6″S, 43°02′40″W, 13 Jul 2001 (fl), Souza et al. 25889 (ESA); GrãoMogol, a 23 km de Barrocão na estrada para Grão-Mogol, na bifurcação Grão-Mogol e Botumirim, 16°35′03″S, 43°10′40″W, 3 Feb 2015 (fr), Cavalcanti et  al. 3890 (CEN); Grão-Mogol, estrada para Virgem da Lapa, ca. 2 km de Grão-Mogol (próximo a antena de televisão), 16°33′39″S, 42°52′32″W, 13 Jul 2001 (fl), Souza et al. 25835 (CEN, ESA); Grão-Mogol, Fazenda Boa Vista, próximo à torre de TV, 6 Sep 1986 (fl, fr), Cordeiro & Mello-Silva (CFCR) 10144 (K, F, SPF); rodovia para José Nápoles, 12 Jun 1990 (fl), Hatschbach & Nicolack 54233 (MBM); Grão-Mogol, Serra do Calixto, Sep 1945 (fl), Lanstryak s.n. (R); 16 Oct 1995 (fl), Pires et al. 557 (CEN, MBM); Grão-Mogol, Vale do Rio das Mortes, a oeste da cidade, 24 Jul 1986 (fl), Mello-Silva et al. (CFCR) 9881 (CEN, K, SPF); Grão-Mogol, Aug 1960 (fl), Mendes-Magalhães 18389 (UB); Turmalina, 19 km N Lelivelândia na BR 367 de Itaobim para Diamantina, 26 Nov 1985, Graham 954 (MO, NY); Virgem da Lapa, estrada para Igigatu ou Buriti, 7–10  km de Virgem da Lapa, 19 Jul 1985 (fl), Martinelli et al. 11163 (BHCB, MO, NY, RB). Diplusodon epilobioides is represented by usually tall, much-branched, characterized by a minute and inconspicuous indumentum that covers the whole plant and gives a velvety appearance to the surfaces, in addition to its narrowly elliptic to narrowly oblong leaves with a revolute margin, and especially its long pedicel

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Fig. 13.32  Diplusodon epilobioides. A. Branch with inflorescence. B. Leaf, abaxial surface. C. Detail of leaf abaxial surface indumentum and whole plant. D. Simple trichome. E. Floral bud. F. Petal. G. Flower with prophylls, without petals. H. Pistil. (A–H Hatschbach 41484, NY)

Descriptions

191

(generally 4–7 mm long). Candolle (1828) described the species as having 12–15 stamens, but some specimens have up to 18 stamens. The habit and shape of the leaves are similar to D. rosmarinifolius, a species of Distrito Federal and Goiás. However, D. rosmarinifolius differs in having subsessile flowers with short epicalyx segments and a stellate indumentum on the whole plant. Vegetatively, Diplusodon epilobioides resembles D. thymifolius in that the latter has narrow leaves and an inconspicuous indumentum throughout the plant; however, in D. thymifolius the flowers are smaller and subsessile (with pedicels 0.9– 1.3 mm long), the epicalyx segments are shorter (1–2 mm long) and spreading to deflexed, and the prophylls are shorter (2.5–3 mm long) and do not surpass the apex of the sepals. The type material of D. epilobioides, Martius s.n., is represented at M by two sheets, both with well-preserved branches, the identification “Friedlandia epilobifolia” by Martius, and the locality, “Serro Frio ad Tejuco” (Serro near Diamantina, Minas Gerais). These specimens are a good match for D. epilobioides, but the locality data is questionable. Apart from the type material, D. epilobioides has never been collected in the Diamantina region, in spite of the many current records for northern Minas Gerais. Another Martius name is D. thymifolius Mart. ex DC., a species from Southwest Bahia, northwestern Minas Gerais, and Piauí. The type of this species also has a label indicating “Serro Frio ad Tejuco,” a region where it is not known to occur. This leads me to suspect that there may be some confusion of locality data for the types of these two species. There is not sufficient evidence in the protologue of Diplusodon epilobioides to know whether Candolle (1828) was citing one or both sheets deposited at M. One of them bears a different label that is blue in color; this specimen, originally from the Herbarium of the Flora Brasiliensis, is here designated as the second-step lectotype. Lourteig (1989) cited Martius s.n. as holotype, which is considered the firststep lectotype (ICBN Art. 9.17, ex. 14); however this gathering is represented by two specimens at M and a second-step lectotype is designated here. 27. Diplusodon ericoides Lourteig, Bradea 19(5): 213. 1989. Type: Brazil. Goiás: Serra Geral do Paranã, 3–4 km NE of São João da Aliança, 25 Mar 1973 (fl), W. R. Anderson, M. T. K. Arroyo, S. R. Hill, R. R.Santos & R. Souza 7949 (holotype P, [01901906]; isotypes, MO, NY, UB). Figs. 13.33 A–C, 13.35 Subshrubs 10–30  cm tall, delicate, erect, unbranched, sometimes slightly branched at inflorescence, sometimes caespitose, with a xylopodium, glabrous, not dark-brown when dry. Upper branches subquadrangular, unwinged, very slightly pubescent, internodes 5–6 cm long. Leaves decussate, erect, sometimes imbricate, sessile; blades 4–8.5 × 0.8–1.5 mm, slightly glaucous, coriaceous, not discolorous, hyphodromous, linear-lanceolate, base obtuse, apex acute, margin plane; secondary veins 1 on each side of midvein, inconspicuous; domatia absent. Synflorescence frondose, botryum to diplobotryum type, congested; accessory branches absent, principal florescence caducous, paraclades 2–2.5  cm long; bract similar in shape and size to the vegetative leaves; flowers subsessile, hypopodium absent, epipodium

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ca. 1 mm long; prophylls 4.5–5.5 × 1.2–1.5 mm, surpassing the apex of the floral tube, linear-lanceolate, unkeeled, base obtuse, apex acute, margin plane; floral tube 4–4.5 mm long, oblong; sepals 1.5–2 mm long, triangular, not caudate, not deflexed; epicalyx segments 2.5–2.8 mm long, linear-triangular, dorsiventrally compressed, erect, shorter than the sepals; corolla 1.5–2.5  cm in diam., rose-magenta, petals 11–11.5  ×  4.8–5  mm, obovate, apex obtuse; stamens 12, free portion of the filaments 4–5 mm long, dark rose, exserted; ovary ca. 1.3 × 1.5 mm, orbicular, glabrous, style 5–7  mm long, dark rose, stigma green, ovules 12–15. Capsule ca. 4.3 × 2 mm, globose to oblong, glabrous; seeds 4–5, 1.5–2 × 0.9–1.1 mm. Distribution and Ecology (Fig.  13.35)  Diplusodon ericoides is endemic to the south of Chapada dos Veadeiros, Goiás, occurring in cerrado, campo sujo, and campo rupestre, growing in sandstone soil among grasses, at about 1000 m. Flowers from January to April and August; fruits registered in May. Specimens Examined: BRAZIL. Goiás,  Agua Fria de Goiás, rodovia GO-118, subida para a Torre Repetidora de Roncador, 5 Aug 2000 (fl), Hatschbach et  al. 70634 (MBM); Água Fria de Goiás, estrada de acesso a repetidora de Roncador, ca. 3 km a leste da GO-118, 14°53′06″ S, 47°33′20″ W, 4 Feb 2014 (fl), Pereira-Silva et al. 16571 (CEN); 14°53′06″ S, 47°33′25″ W, 6 Apr 2006 (fl), Cavalcanti et al. 3753 (CEN); São João da Aliança, seguindo a pista de aviação, a 4 km NE a partir da cidade, 21 May 2001 (fr), Pereira-Silva et al. 5033 (CEN, MO); 28 May 2001 (fr), Cavalcanti et al. 2721 (CEN, MO); 14°14′15″ S, 47°30′08″ W, 30 Jan 2003 (fl), Cavalcanti et  al. 3110 (CEN); 14°14′15″ S, 47°30′08″ W, 30 Jan 2003 (fl), Cavalcanti et al. 3170 (CEN); Corrente, 20 Feb 2000 (fl), Hatschbach et al. 70476 (MBM); Corrente, 20 Feb 2000 (fl), Hatschbach et al. 70485 (MBM); estrada para Vãozinho, próximo ao Athos Hotel, 7 Feb 2012 (fl), Faria-Jr. et al. 769 (CEN, UB). Diplusodon ericoides is represented by small, delicate and usually unbranched subshrubs. The leaves are small (4–8 cm long) and linear-lanceolate. The flowers are small (to 2 cm in diam.), with 12 stamens and the floral tube is oblong and has erect epicalyx segments that are larger than the sepals. Populations of D. ericoides occur in a limited area that is surrounded by soybean cultivation. Because of its restricted distribution and imperiled habitat, D. ericoides can be considered threatened. 28. Diplusodon fastigiatus Lourteig, Bradea 5(19): 236. 1989. Type: Brazil. Goiás: Chapada dos Veadeiros, 21 Dec 1968 (fl, fr), A. M. Giulietti, M. José, G.M. Barroso. 588 (holotype P, [01901905]; ­isotype, UB). Figs. 13.34 A–E, 13.35 Shrubs ca. 0.4 m tall, erect, branched, not dark-brown when dry. Upper branches subquadrangular, unwinged, pubescent, internodes 0.5–1.2 cm long. Leaves decussate, not imbricate, spreading, subsessile; petioles ca. 1  mm long; blades 6–10  ×  5–7  mm, coriaceous, not discolorous, acrodromous-basal-supranumerary, ovate-lanceolate to elliptic-lanceolate, base obtuse to subcordate, apex acute,

Descriptions

193

Fig. 13.33  Diplusodon ericoides and Diplusodon saxatilis. A–C. D. ericoides. A. Habit. B. Leaf, abaxial surface. C. Flower with prophylls, without petals. D–H. D. saxatilis. D. Habit. E. Leaf, abaxial surface. F. Prophyll, adaxial surface. G. Flower with prophylls, without petals. H. Inner surface of floral tube, without pistil and filaments. (A–C Anderson et  al. 7949, UB; D–H Hatschbach et al. 28923, MBM)

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margin revolute; blades covered by fine, short indumentum; secondary veins 3 on each side of the midvein, prominent on the abaxial surface; domatia present. Synflorescence frondose, diplobotryum to triplobotryum type; accessory branches absent, principal florescence ca. 10 cm long, botryum type, paraclades 3–6 cm long, virgate; bracts ovate to oblong, 5–8 × 2.5–4 mm; flowers pedicellate, hypopodium 1–1.5 mm long, epipodium 0.5–1 mm long; prophylls 5–6 × ca. 4 mm, reaching the middle of the tube or slightly more, ovate, unkeeled, base acute, apex obtuse, margin plane, covered with fine trichomes; floral tube 5–5.5 mm long, funnel-­shaped, covered with short and fine trichomes; sepals ca. 4 mm long, triangular, not caudate, not deflexed, covered with fine trichomes; epicalyx segments ca. 2 mm long, cylindrical, spreading to slightly deflexed, shorter than the sepals; corolla ca. 2.5 cm in diam., rose, petals ca. 14 × ca. 6 mm, obovate, apex obtuse; stamens 18, free portion of the filaments ca. 6.5 mm long, slightly exserted; ovary ca. 1.5 × ca. 2 mm, oboval, glabrous, style ca. 12 mm long, ovules ca. 14. Capsules ca. 3.5 × 4 mm, globose, glabrous; seeds 2.9–3.1 × 1.5–1.7 mm. Distribution and Ecology (Fig. 13.35)  Diplusodon fastigiatus is known from two localities in Goiás, at 1131–1800 m elevation. Flowers registered in December and fruits in August and December. Specimens Examined: BRAZIL.  Goiás,  Cavalcante, estrada CavalcanteMinaçu, km 20, 13°38′42″ S, 47°28′39 W, 31 Aug 2004 (fr), Cavalcanti et al. 3532 (CEN) Diplusodon fastigiatus was described by Lourteig (1989), who cited Barroso et al. 588 as the type. This collector number belongs to Ana Maria Giulietti (checked with the collector) although Graziela Barroso also participated in the collection. The type material is in fact Giulietti et al. 588, and the label needs to be corrected in the type material. The specimen G.M. Barroso 588 is Declieuxia fruticosa [Willd. ex Roem. & Schult.] Kuntze, collected in Distrito Federal, Brazil). In her key, Lourteig (1989) separated D. fastigiatus, with epicalyx segments half the length the sepals, from D. sigillatus, with epicalyx segments slightly longer than the sepals. In addition, the epicalyx segments of D. fastigiatus are spreading to slightly deflexed (vs. suberect in D. sigillatus), and the type of D. fastigiatus has leaves smaller and more delicate, with sparser indumentum (vs. the dense and short indumentum of D. sigillatus). 29. Diplusodon floribundus Pohl in Mart., Flora 10: 151. 1827. Type: Brazil. Goiás: In montosis graminosis prope Ant. Gonzales procul Trahiras, May–Jun 1820, J. B. E. Pohl 1885 = D942 (first-step lectotype, designated by Lourteig 1989: 241, second-step lectotype, W [0053460], here designated; isolectotypes, G, K [2], W [0053403]). Figs. 13.36 A–C, 13.40 Subshrubs (0.2–)0.5–1.5 m tall, erect, slightly branched, glabrescent, not darkbrown when dry. Upper branches cylindrical, unwinged, hirsute, with a dense indumentum of long, white trichomes, sometimes pubescent, rarely glabrous, internodes

Descriptions

195

Fig. 13.34  Diplusodon fastigiatus. A. Branch with inflorescence. B. Section of internode. C– D. Leaves, abaxial surface. E. Flower with prophylls, without petals. (A–E Giulietti et al. 588, UB)

196

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Fig. 13.35  Geographic distribution of Diplusodon divaricatus, D. epilobioides, D. ericoides, and D. fastigiatus

Descriptions

197

1–2 cm long. Leaves decussate, not imbricate, spreading to suberect, sessile; blades 10–60  ×  10–32  mm, coriaceous, not discolorous, acrodromous-basal-typical to supranumerary, ovate-lanceolate, base obtuse to slightly cordate, apex obtuse or sometimes slightly acuminate, margin plane, sometimes thickened or ciliate, hirsute, glabrescent, blade canescent mostly near the region of the veins, covered by dense, long, white, abundant trichomes, sometimes completely glabrous; secondary veins 3–4 on each side, departing from the base of the midvein, 1 along the midvein, prominent on the abaxial face, less evident on the adaxial face; domatia present. Synflorescence frondose, botryum to triplobotryum type, congested; accessory branches absent, principal florescence short, generally caducous, paraclades 5–20 cm long; bracts similar to the vegetative leaves in shape, reduced in size toward the apex of the branches; flowers sessile to subsessile, hypopodium 0.5–1 mm long, epipodium absent; prophylls 8–9 × 4–5 mm, surpassing the apex of the floral tube, lanceolate, unkeeled, base obtuse, apex acute to obtuse, margin plane, with long trichomes; floral tube 7–7.5 mm long, funnel-shaped, with dense indumentum of long trichomes; sepals 5.5–7 mm long, triangular, not caudate, not deflexed, pilose; epicalyx segments 7–10 mm long, cylindrical, narrow, erect, surpassing the sepals, with spreading trichomes shorter than on the rest of the plant; corolla 4–4.5 cm in diam., rose, petals 18–20 × 9–10 mm, obovate, apex obtuse; stamens (16–)18, free portion of the filaments 7–9 mm long, exserted; ovary 1.5–2 × ca. 3 mm, rounded, glabrous, style 10–12 mm long, ovules 28–30. Capsules ca. 4 × 3 mm, globose to obovate-depressed, glabrous; seeds ca. 35, 2–2.5 × 1.8–2 mm. Distribution and Ecology (Fig. 13.40)  Diplusodon floribundus is endemic to the cerrados of Goiás, growing on laterite or sandstone soils, at 800–900 m. Flowers from March to September; fruits registered from July to August and January. Specimens Examined: BRAZIL. Goiás,  Niquelândia, estrada de terra à direita da mina de níquel, 14°23′03″ S, 48°24′57″ W, 24 Mar 1995 (fl), Oliveira et al. 265 (CEN, IBGE); Niquelândia, estrada à esquerda da mina de níquel, 14°20′12″ S, 48°23′54″ W, 20 Jun 1995 (fl), Fonseca et al. 363 (CEN, IBGE); Niquelândia, 8 Aug 1995 (fl), Marquete et al. 2222 (IBGE, RB); Niquelândia, estrada para a Codemin, estrada vivinal para a Estância Corujinha, 14°12′47.6″ S, 48°19′01″ W, 5 Jan 2006 (fr), Pastore & Suganuma 1386 (CEN); Niquelândia, 40  km de Niquelândia, Codemin, 14°11′20″ S, 48°20′40″ W, 28 May 1996 (fl), Silva et  al. 2907 (CEN, IBGE, P, UFG); Niquelândia, estrada de acesso Rosaseana, próximo à Codemin, 14°10′38″ S, 48°21′19″ W, 21 May 1997 (fl), Cordovil-Silva et al. 604 (CEN, MO, NY); Niquelândia, cerca de 1 km abaixo da Codemin, próximo ao antigo leito do rio Bagagem, 14°09′22″ S, 48°19′ W, 8 Aug 1997 (fl), Walter et al. 3769 (CEN, MO); Niquelândia, cerca de 1  km abaixo da Codemin, próximo ao antigo leito do rio Bagagem, 8 Aug 1997 (fl), Walter et  al. 3771 (CEN); Niquelândia, estrada Niquelândia-Cia. Níquel Tocantins, km 6, 14°20′ S, 48°25′ W, 28 Jul 1999 (fl, fr), Pereira-Silva et  al. 4216 (CEN); Niquelândia, reservatório de formação do AHE Serra da Mesa, 14°09′19″ S, 48°19′35″ W, 2 Jun 1998 (fl), Walter et al. 4165 (CEN); Niquelândia, área de influência do AHE Serra da Mesa, 14°10′11″S, 48°21′43″W, 4 Jun 1998 (fl), Walter et  al. 4203 (CEN, MO, NY); Niquelândia, Fazenda Santo

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Fig. 13.36  Diplusodon floribundus. A. Vegetative branch. B. Branch with inflorescence. C. Flower with prophyll, without petals. (A Cordovil-Silva et al. 604, CEN; B–C Walter et al. 4203, CEN)

Descriptions

199

Antônio da Serra Negra, 14°07′30″ S, 48°22′ W, 21 May 1996 (fl), Cordovil-Silva et al. 452 (CEN, MO, NY); Niquelândia, Macedo, 26 Jul 1952 (fl), Macêdo 3662 (MO, SP); Niquelândia, Macedo, cerca de 15  km  N de Niquelândia, 14°18′ S, 48°24′ W, 17 May 1988 (fl), Ferreira & Rizzo 301 (NY, MO); Niquelândia, ca. 18  km ao N de Niquelandia 14º18′S, 48º23′W, 3 Aug 1992 (fl, fr), Filgueiras & Lopes 2400 (IBGE); Niquelândia, Macedo, cerca de 8  km à direita da mina de níquel, 14°24′59″ S, 48°25′16″ W, 25 Jun 1997 (fl), Fonseca et  al. 1444 (CEN, IBGE); Niquelândia, 4 km de Macedo, mina de extração de níquel, 28 May 1996 (fl), Silva et al. 2944 (CEN, IBGE, P); Niquelândia, 1ª bica do asfalto, ca. 5 km de Macedo, 14°24′06″ S, 48°26′01″ W, 30 Jun 1996 (fl), Fonseca et al. 1061 (IBGE, P); Niquelândia, próximo ao povoado de Macedo, estrada de terra à esquerda da Mina de Níquel Tocantins, passando por baixo do teleférico, 14°20′12″ S, 48°23′54″ W, 20 Sep 1996 (fl), Silva & Ferreira 3160 (CEN, IBGE); Niquelândia, GO-237, Niquelândia-Nossa Senhora da Abadia de Muquém, 5 Apr 2006 (st), Cavalcanti & Pereira-Silva 3728 (CEN); Planaltina de Goiás, Condomínio Araponga, 17 Aug 2002 (fl), Costa-Vieira 21 (CEN). Diplusodon floribundus is represented by glabrescent and slightly branched, subshrubs. Commonly, it is densely pilose and canescent, but it can have different degrees of pilosity on the leaves, or even be completely glabrous (Cordovil-Silva et al. 604, Silva et al. 2907). This species is characterized by acrodromous, ovatelanceolate leaves, long prophylls that surpass the apex of the floral tube, by long, narrow, distinctly cylindrical epicalyx segments, and by the androecium with 18 stamens. Diplusodon floribundus was described by Pohl (1827a) without citation of specimens in the protologue. In another publication, Pohl (1827b) refers to a single collection of D. floribundus collected in May–June from grassy mountains in Antonio Gonçales near Traíras, Goiás, without mentioning his collection number. Two specimens were examined at W.  One specimen bears Pohl’s handwritten label, with Pohl’s collecting number (Pohl 1885), and the reference number of his field diary (D = 942). Lourteig (1989) cited this collection as the holotype, which is considered as the first-step lectotype (ICBN Art. 9.17, ex. 14). The second-step lectotype is selected here as the specimen W005360, which has the original label annotated by Pohl. It was verified that other herbaria have specimens of Diplusodon floribundus which the label indicates Pohl s.n. Since duplicates of Pohl’s collections went to other herbaria without the information from his field diary (Wahlnöffer, pers. comm) and most lack a number, the material at G, and K are regarded as likely duplicates of Pohl 1885 = D 942. 30. Diplusodon glaucescens DC., Prodr. 3: 94e. 1828. Type: Brazil. Minas Gerais: in summo montis Itambé et Serro, C. F. P. Martius obs. 1350 (first-step lectotype, designated by Lourteig 1989: 235, second-­step lectotype, M [0146280], here designated; isolectotypes, M [0146279, 0243178], G). Figs. 13.37 A–E, 13.40, 15.1 C–D

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Friedlandia glaucescens Mart. ex Koehne, Fl. Bras. (Martius) 13(2): 322. 1877, pro syn. Friedlandia nummularifolia Mart. ex Koehne, Fl. Bras. (Martius) 13(2): 322. 1877, pro syn. Subshrubs 0.2–2  m tall, erect, branched, glabrous, not dark-brown when dry. Branches lax, cylindrical, the upper ones subquadrangular, unwinged, yellow to reddish, internodes 1–2.5 cm long. Leaves decussate, not imbricate, spreading, sessile, 8–22 × 8–26 mm, glaucous, coriaceous, not discolorous, acrodromous-basaltypical, ovate-depressed to orbicular-reniform, base cordate, apex rounded, acuminate, margin plane, reddish to wine-colored; secondary veins 3–6, departing from the base of the midvein, rarely one vein departing from a higher level of the midvein; domatia absent. Synflorescence frondose, diplobotryum to pleiobotryum type, congested; accessory branches absent, principal florescence 9.5–13 cm long, botryum type, paraclades 4–20 mm long; bracts 8–12 × 7–12, ovate-lanceolate to orbicular-reniform; flowers pedicellate, hypopodium 4–6.5  mm long, 8–11  mm long in fruit, epipodium ca. 1 mm long; prophylls 5–6 × 4–5 mm, reaching the apex of the floral tube or slightly less, ovate, unkeeled, base and apex obtuse, margin plane; floral tube 4–5  mm long, in fruit, including the sepals, 11–13  ×  7–8  mm, greenish with parts wine-colored, dark-brown in fruit, slightly nitid, funnel-shaped; sepals 3.5–4 mm long, triangular, not caudate, not deflexed; epicalyx segments 3.3– 3.5 mm long, cylindrical-triangular, erect, reaching the height of the sepals; corolla ca. 3.5 cm in diam., rose-magenta, petals ca. 17 × 12 mm, obovate, apex obtuse; stamens 12, free portion of the filaments 7–8 mm long, not surpassing the sepals; ovary 1.5 × 2.2 mm, broadly obovate-depressed, glabrous, style ca. 18 mm long, ovules 19–22. Capsules 3.8–4  ×  3.9–4  mm, oblong, glabrous; seeds 11–13, 2–2.5 × 1.5–2 mm. n = 15 (Graham 1985). Distribution and Ecology (Fig. 13.40)  Diplusodon glaucescens is endemic to the central portion of the Cadeia do Espinhaço, Minas Gerais, inhabiting campos rupestres in the municipalities of Diamantina, Data, Gouveia, Presidente Kubitschek, Serro, and Serra do Cipó, at elevations of 1200–1430 m. This species also occurs disjunctly in Serra do Cabral, Minas Gerais. Flowers from September to April; fruits from January to June. Specimens Examined. BRAZIL.  Minas Gerais,  Barão, Serra do Capão, 21 May 1931 (fl, fr), Mexia 5874 (K, MO, NY, R); Barão, Serra do Capão, 15 Jan 1969 (fl), Irwin et al. 22004 (MBM, MO, NY); Barão, Serra do Capão, 10 Apr 1973 (fl), Anderson 8590, (NY, UB); Barão, Serra do Capão, 14 Apr 1987 (fl), Scatena et al. (CFCR) 10477 (K, SPF); Conselheiro Mata, entroncamento da rodovia Datas-Serro, BR-259, com Costa Sena, cerca de 1 km, 18°33′54″ S, 43°38′40″ W, 19 Jan 2003 (fl), Cavalcanti et al. 3127 (CEN, MO); Datas, Morro do Coco, 18°26′ S, 43°41′ W, 17 Jul 1987 (fr), Pirani & Mello-Silva (CFCR) 11021 (K, F, SPF); Datas, próximo à cidade, 21 Mar 1993 (fl, fr), Kameyama & Esteves 38 (CEN, SPF); Datas, ca. 1 km SW da BR 259 (Datas-Serro), nas estrada para Costa Sena, 27 Nov 1985 (fl), Graham 962 (MO, NY); Diamantina, ca. 18 km na rodovia SW de Diamantina, na

Descriptions

201

Fig. 13.37  Diplusodon glaucescens. A. Branch with inflorescence. B–C. Leaves, abaxial surface. D. Floral bud. E. Flower with prophylls, without petals. (A Anderson 8590, UB; B–E Scatena et al. [CFCR] 10477, SPF)

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estrada para Curvelo, 10 Apr 1973 (fl), Anderson 8490 (NY, UB); São Gonçalo das Pedras, (SE de Diamantina), estrada de terra para Serro, ca. 18°20′ S, 43°30′ W, 28 May 1987 (fl), Lewis & Morão 1648 (K, RB, SPF); Diamantina, estrada de terra para Barão do Guaicui, 18°22′37″ S, 43°42′12″ W, 9 Feb 2015 (fl, fr), Cavalcanti et  al. 3934 (CEN); estrada Diamantina-Conselheiro Mata, 3–6  km da estrada Diamantina-Gouveia, 14 Mar 1999 (fl), Souza et  al. 22217 (CEN [2], ESA); Gouveia, 23 Feb 1975 (fl), Hatschbach et al. 36430 (MBM); Gouveia, ca. 25 km SW de Gouveia, 29 Sep 1985 (fl), Graham 982 (NY); Gouveia, 22 May 1989 (fl), Hatschbach et al. 53082 (MO, W); Gouveia, 18°35′12″ S, 43°53′43″ W, 3 Jan 2006 (fr), Proença & Harris 3100 (CEN, UB); Joaquim Felício, Serra do Cabral, Armazém da Laje, 16 Mar 1997 (fl), Hatschbach et  al. 66307 (CEN, MBM); Presidente Kubitscheck, 18°28′ S, 43°39′ W, 15 Feb 1989 (fl) Cavalcanti et al. 180 (CEN, HUFU, SPF); Presidente Kubitscheck, rodovia Serro-Diamantina, km 34, 18°34′47″ S, 43°34′07″ W, 24 Apr 1997 (fl), Cavalcanti et  al. 2286 (CEN, MO, NY); Presidente Kubitscheck, 13 Mar 1982 (fr), Graham 874 (MO, MBM, NY 2); Serro, rodovia Serro-Diamantina, km 23, 18°34′42″ S, 43°29′97″ W, 24 Apr 1997 (fl), Cavalcanti et al. 2284 (CEN, MO, NY); Serro, 19 Jan 1972 (fl), Hatschbach et al. 28919 (MBM); Serro, estrada Serro-Milho Verde, a 19 km de Serro, 18°29′41″ S, 43°28′42″ W, 18 Jan 2003 (est), Cavalcanti et al. 3107 (CEN, MO); Milho Verde, 13 Apr 1990 (fl, fr), Lima 2 (BHCB, CEN); Without reference to municipal district: Serra do Cipó, 21 Apr 1976 (fl), Silva s.n. (OUPR 22917); 1973 (fl), Ferreira 1416 (PAMIG); Serra de Minas, 1 Apr 1957 (fl), Pereira 2760 & Pabst 3396 (B); rio das Pedras, dans le serra, Minas Gerais, Glaziou 19168 (K, LE n.v., R); Glaziou 19169a (K); Vauthier 420 (G). Based on vegetative characteristics, Diplusodon glaucescens is almost identical to D. orbicularis, which is also from Cadeia do Espinhaço. Both species are erect and branched subshrubs with glaucous, ovate-depressed to cordiform leaves with cordate base. Stamen number is the principal difference between the two species (12 in D. glaucescens vs. 6 in D. orbicularis). The vegetative structures and flowers of Diplusodon glaucescens are somewhat larger than those of D. orbicularis, which led me to suggest that D. glaucescens might be a polyploid form of D. orbicularis (Cavalcanti 1988). However, chromosome counts later indicated that the D. glaucescens number is 2n  =  30 (Cavalcanti 1995), while D. glaucescens has a haploid number of n  =  15 (Graham 1985). Diplusodon glaucescens also resembles other glabrous species with acrodromous, glaucous leaves with reddish to wine-colored margins; this species differs from D. rotundifolius, a species with a similar habit and also occurring in Cadeia do Espinhaço, by having ovate prophylls that completely cover the floral tube. The type material of D. glaucescens, Martius obs. 1350, is represented at Munich (M) by three specimens. One of these, M0146280, is here chosen as lectotype; this specimen has a label with the locality data, whereas the other two have smaller labels with annotations by Martius about some characteristics of the species and his unpublished name.

Descriptions

203

31. Diplusodon glaziovii Koehne, Bot. Jahrb. Syst. 23(Beibl. 57): 32. 1897. Diplusodon glaziovii Koehne f. subspathulatus Koehne, Bot. Jahrb. Syst. 23(Beibl. 57): 32. 1897. Diplusodon glaziovii Koehne var. subspathulatus Koehne. Type: Brazil. Minas Gerais: Diamantina, Biri-biri, A. F. M. Glaziou 19174 (lectotype, P [01901904], designated by Lourteig 1989: 214; isolectotypes, F, K, LE n.v., MPU, P [01901903], R, RB). Figs. 13.38 A–C, 13.40 Diplusodon glaziovii Koehne f. ovalifolius Koehne, Bot. Jahrb. Syst. 23(Beibl. 57): 32. 1897. Diplusodon glaziovii Koehne var. ovalifolius Koehne. Type: Brazil. Minas Gerais: Serra do Cipó, A.  F. M.  Glaziou 19174 p.p. (lectotype, P [01901903], designated by Lourteig 1989: 215; isolectotypes, K, MPU, R). Shrubs 1–1.5  m tall, erect, much-branched, glabrous, dark-brown when dry. Branches congested, the upper ones subquadrangular, unwinged, internodes ca. 1 cm long. Leaves decussate, not imbricate, erect, subsessile; petiole ca. 1 mm long; blades 6–12 × 1.5–3.5 mm, coriaceous, slightly discolorous, not glaucous, hyphodromous, narrowly obovate to narrowly elliptic, concave, base acute, apex obtuse, margin plane, lateral veins absent, adaxial surface dark-green, abaxial olive-green. Synflorescence frondose, diplobotryum to triplobotryum type; accessory branches absent, principal florescence botryum type, ca. 4  cm long, caducous, paraclades 8–17  mm long; flowers pedicellate, hypopodium 1.5–2.5  mm long, epipodium 0–0.5 mm long, prophylls 5.5–10 × 2.7–3 mm, reaching almost to the apex of the floral tube, obovate, slightly keeled, base acute, apex obtuse, margin plane; floral tube ca. 5 mm long, green, funnel-shaped; sepals 3–4 mm long, wine-colored, triangular, slightly caudate, not deflexed, not ciliate; epicalyx segments 4–5 mm long, cylindrical, slightly dorsiventrally compressed on the upper third, erect, surpassing the sepals ca. 2  mm; corolla 1.5–3  cm in diam., bright rose, petals 12–13  ×  ca. 7 mm, elliptic, apex obtuse; stamens 6, free portion of the filaments 6–7.7 mm long, slightly exserted, surpassing the sepals ca. 0.5 mm; ovary 1.8–2.2 × 2–3 mm, rhombic, glabrous, style ca. 14 cm long, ovules 8–9. Immature capsules globose, glabrous; seeds ca. 3, immature. Distribution and Ecology (Fig. 13.40)  Diplusodon glaziovii is a rare species that is endemic to Diamantina region, Cadeia do Espinhaço, specifically in areas of rocky outcrops with white sand, at ca. 1325  m elevation. Flowers and immature fruits registered in April. Specimens Examined: BRAZIL.  Minas Gerais,  Diamantina, BR 259– Conselheiro Mata, km 26, 18°18′86″ S, 43°52′44″ W, 27 Apr 1997 (fl), Cavalcanti et al. 2305 (CEN, MO, NY); Gouveia, estrada entrando paralela aos moinhos de vento do Camilinho, sítio do Mulato, área de extração do Aliance, 18°31′50″S, 43°53′45″W, 15 Apr 2016 (fl, fr), Faria-Jr. & Otoni 5661 (CEN, HDJF, UB); s.loc., s.d., Schwacke 8266 (BHCB).

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Fig. 13.38  Diplusodon glaziovii. A. Branch with inflorescence. B. Leaf, adaxial surface. C. Flower with prophyll, without petals. (A–C Cavalcanti et al. 2305, CEN)

Descriptions

205

Diplusodon glaziovii is a rare much-branched and glabrous shrub from campos rupestres of the Minas Gerais mountains. It is characterized by narrow leaves that are glaucous and slightly concave and have plane margins. The leaves turn darkbrown when dry. The flowers are pedicellate (pedicel 2–3 mm long) and have erect epicalyx segments that reach the size of the sepals and androecium with six stamens. The morphologically closest species is Diplusodon hexander due to its narrow, glabrous leaves and flowers of six stamens. However, D. hexander presents glaucous and revolute linear leaves. Koehne (1897) described Diplusodon glaziovii with the forms ovalifolius and subspathulatus (=f. glaziovii). For the form subspathulatus, Koehne cited two collections, the syntypes Glaziou 19174 (said to have both forms mounted on the single unicate specimen) and Schwacke 8266. The material studied by Koehne for f. subspathulatus (Glaziou 19174) was destroyed at B (F neg. nº 17921). A photograph of the specimen shows the left branch bearing rounded to elliptic leaves, with the notation of the author “6 stamens, form a” (=f. ovalifolius), and the other branch with narrowly elliptic leaves and the notation “forma b” (=f. subspathulatus). This same mixture was observed on the specimen at P [P01901903] and K. Lourteig (1989) accepted Koehne’s infraspecific taxa of Diplusodon glaziovii and referred to them as varieties, but did not make the required combinations. She did not cite Schwacke 8266, and selected as lectotype of f. ovalifolius, Glaziou 19174 p.p. referring to one branch with large leaves and Glaziou 19174 p.p. with narrowly elliptic leaves as f. subspathulatus. 32. Diplusodon glocimarii T.B.Cavalc., Bol. Bot. Univ. São Paulo, 22(1): 7, fig. 4. 2004. Type: Brazil. Minas Gerais: Capitólio, MG-050, lado esquerdo da estrada, do lado direito vê-se o reservatório de Furnas, 21°04′ S, 46°26′ W, 24 Feb 2000 (fl, fr), T.B.Cavalcanti, A. Amaral-Santos, G. Pereira-Silva, T. A. B. Dias 2521 (holotype, CEN [00035408]; isotypes, B, BHCB, MBM, NY, SPF, W). Figs. 13.39 A–F, 13.40 Subshrubs 0.4–1 m tall, erect, slightly branched, with a xylopodium, not darkbrown when dry. Upper branches strongly quadrangulate, rust-colored to yellow, winged, wing 0.3–0.6  mm wide, pubescent to short-villous, indumentum white. Leaves decussate, imbricate, erect, petiolate; petioles (1.5–)2.5–3.5  mm long; blades 20–42 × (2.5–4)5–18 mm, silvery to glaucous, coriaceous, not discolorous, acrodromous-­basal, elliptic to ovate-elliptic, sometimes narrowly lanceolate, base acute to rarely obtuse or attenuate, apex acute, margin plane to rarely subrevolute, blade soft to the touch, covered by a short-villous indumentum on both surfaces, abaxial surface usually silvery or white, varying to an indumentum composed of dense, short, inconspicuous trichomes, without the silvery aspect; secondary veins 3–4 on each side of the midvein, departing from the base of the midvein, 1 on each side of the midvein when leaves are narrowly lanceolate, sulcate on the abaxial surface; domatia absent. Synflorescence frondose to frondose-bracteose, diplobotryum to triplobotryum type, congested; accessory branches absent, principal florescence 20–30  cm long, paraclades 15–25  cm long; bracts 15–25  ×  5–10  mm,

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similar to the vegetative leaves but smaller in size; flowers pedicellate, hypopodium 1.5–2 mm long, epipodium 1–1.5 mm long; prophylls 10–11 × 3–3.5 mm, surpassing the sepals 1–2 mm, elliptic, unkeeled, base and apex acute, margin plane, short-­ villous or covered by indumentum of small, inconspicuous trichomes; floral tube 8–9 mm long, yellowish brown-golden in fruit, oblong, short-villous or covered by indumentum of small, inconspicuous trichomes, silvery to whitish; sepals 2.5– 3.2 mm long, triangular, not caudate, not deflexed, short-villous or with inconspicuous indumentum; epicalyx segments 4–4.5  mm long, narrowly lanceolate, dorsiventrally compressed, spreading when fresh, suberect when dry, surpassing the sepals, with short, spreading or small, inconspicuous trichomes; corolla 2.5–3 cm in diam., lilac, petals 15–25 × 7–15 mm, elliptic to obovate, apex obtuse; stamens 12, free portion of the filament 12–14  mm long, red, exserted 5–6  mm; ovary 2–2.5 × 3–3.5 mm, obovate-depressed, glabrous, style ca. 20 mm long, wine-colored, stigma green, prominently capitate, ovules (15–)33–38. Immature capsules oblong to oboval, glabrous; seeds 8–26, 2.2–2.8 × 1.8–2.1 mm. Distribution and Ecology (Fig. 13.40)  Diplusodon glocimarii is endemic to the area of Capitólio, in the southeastern part of Serra da Canastra, Minas Gerais, and from northeast of São Paulo. It was collected from a small, local population in campo limpo near a slope, growing among Vellozia sp., at 700–750 m in rocky soil and in São Paulo, in campo sujo with outcrops, at 720 m. Flowers from November to May and fruits registered in February. Specimens Examined: BRAZIL.  Minas Gerais,  Capitólio, estrada PassosPiumbi, a 45 km de Piumbi, arredores da represa de Furnas, 20°22′98″ S, 46°11′35″ W, 12 Jan 1994 (fl), Souza et al. 5049 (CEN, RB, SPF); Capitólio, Serra da Canastra, Captólio de Lago de Furnas, ca. 900 m da MG-050, estrada para Serra da Canastra, 20°38′24″ S, 46°15′26″ W, 30 Nov 2014 (fl), Flores et al. 4461 (CEN); Capitólio, Pousada do rio Turvo, trilha para a cachoeira do Fecho, 4 Nov 2008 (fl), Kinoshita et al. 42 (HUFU); Capitólio, estrada vicinal da MG 50, à esquerda após o rio Grande, cerca de 1 km após a ponte, 2.4 km da MG-050, 20°65′ S, 45°36′ W, 24 Feb 2000 (fl, fr), Cavalcanti et al. 2513 (CEN, MO, NY, RB); Delfinópolis, Estância Carmen Silvia, 21 Dec 1999 (fl), Silva 125 (RB, SPFR n.v.); Fazenda Água da Serra, Trilha Escada de Pedras, 10 Mar 2003 (fl), Pacheco et  al. 469 (HUFU); Delfinópolis, estrada para Casa Branca, Fazenda Paraíso, Sete Cidades, 10 Apr 2002 (fl), Romero et al. 6275 (HUFU); São Roque de Minas, Parque Nacional da Serra da Canastra, Cachoeira dos Rolinhos, 15 May 1999 (fl), Farinaccio et  al. 320 (CEN, HUFU, SPF); São Roque de Minas, Parque Nacional da Serra da Canastra, Cachoeira dos Rolinhos, 21 Mar 1996 (fl), Nakajima & Romero 1664 (HUFU). São Paulo, Pedregulho, Estreito, Usina de Estreito, 20°09′52″ S, 47°16′23″ W, 18 Mar 2003 (fl), Sasaki et al. 234 (SPF). Most individuals of Diplusodon glocimarii are sparsely branched subshrubs with bright silvery foliage due to the dense villous indumentum. A few populations have individuals with gray-green leaves (also due to dense villous indumentum). Other diagnostic characteristics of the species are the presence of quadrangular branches with wings and narrowly elliptic to ovate-elliptic leaves with acrodromous-basal

Descriptions

207

Fig. 13.39  Diplusodon glocimarii. A. Branch with inflorescence. B. Detail of leaf abaxial surface indumentum. C. Section of internode. D. Leaf, abaxial surface. E. Flower with prophyll, without petals. F. Ovary. (A–D, F Cavalcanti et al. 2521, CEN; E Farinaccio 320, CEN)

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Fig. 13.40  Geographic distribution of Diplusodon floribundus, D. glaucescens, D. glaziovii, and D. glocimarii

Descriptions

209

venation formed by almost parallel veins in close proximity. In the flower, the important characteristics are the long floral tube (8–9 mm long), long prophylls that surpass the apex of the sepals, and 12 stamens. The leaves in Diplusodon glocimarii are usually elliptic to ovate-elliptic. In the populations of Serra da Canastra and surroundings, individuals have leaves narrowly lanceolate, 2.5–4 mm wide, imparting a very different aspect from the type material. The specific epithet honors Glocimar Pereira da Silva (G. Pereira-Silva), geographer and expert on the Brazilian flora, who has made an inestimable contribution to our knowledge of Brazilian Lythraceae through his tireless search for specimens and seeds on collecting expeditions throughout Brazil. 33. Diplusodon gracilis Koehne, Fl. Bras. (Martius) 13(2): 323, pl. 60, fig. 7. 1877. Type: Brazil. Tocantins: Dianópolis, Missão do Duro, Sep 1839 (fl, fr), G. Gardner 3155a (lectotype, P [01901902], designated by Lourteig 1989: 214; isolectotypes, F [2], G [3], K [3], NY, OXF, P [01901900, 01901901], W [2]). Figs. 13.41 A–G, 13.46 Shrubs 0.5–3  m tall, erect, much-branched, not dark-brown when dry. Upper branches cylindrical, unwinged, delicate, finely pubescent, becoming glabrous in the upper parts, internodes 0.5–2 cm long. Leaves decussate, not imbricate, suberect, petiolate; petioles 1–2  mm long; blades 8–12(−25)  ×  3–8(−12) mm, chartaceous to membranaceous, not discolorous, hyphodromous, finely pubescent, becoming glabrous, sometimes slightly silvery, narrowly oblong, base acute to attenuate, apex obtuse, margin plane, blades slightly punctate; lateral veins absent;  domatia absent. Synflorescence frondose, of the pleiobotryum type, lax; accessory branches absent, principal florescence caducous, paraclades 4–12  cm long, bracts similar to the vegetative leaves in shape and size, glabrous or glabrescent; flowers pedicellate, hypopodium 2.5–3 mm long, epipodium 0.5–2 mm long, finely pubescent; prophylls 2.5–3.9 × 1–1.1 mm long, reaching to the middle of the floral tube or surpassing the apex, narrowly oblong to obovate, unkeeled, base acute or attenuate, apex obtuse, margin plane; floral tube 3–4  mm long, green, funnelshaped, glabrous to finely pubescent; sepals 1.5–2.5 mm long, triangular, slightly caudate, not deflexed, finely pubescent, not ciliate; epicalyx segments 2–3 mm long, narrow cylindrical, suberect, the same length as the sepals, sometimes longer, finely pubescent; corolla 1.5–2 cm in diam., rose, petals 6.7 × 5–5.5 mm, obovate, apex obtuse; stamens 12–18, free portion of the filaments 5.5–6  mm long, slightly exserted; ovary 1.5–2.2 × 1.8–2.2 mm, rounded, glabrous, style 6.5–10 mm long, ovules 7–10. Capsules 3–4 × 3–3.5 mm, oblong, slightly narrow at the base, glabrous; seeds 3–6, 2.4–3.5 × 1.9–2.1 mm. Distribution and Ecology (Fig.  13.46)  Diplusodon gracilis is a rare, microendemic species that was known prior to this study only from the type and is still poorly represented in herbaria. This species has been collected in the municipality of Dianópolis, in open cerrado with Melocactus and Syagrus acaule, and on quartz sand at 400–720 m, and it has also been collected in Taguatinga, in high and dense

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Fig. 13.41  Diplusodon gracilis. A. Branch with inflorescence. B. Section of internode. C. Leaf, adaxial surface. D. Leaf, abaxial surface. E. Prophyll, adaxial surface. F. Flower with prophyll, without petals. G. Pistil. (A–G Gardner 3155a, K)

Descriptions

211

cerrado at 850  m, Serra Geral de Goiás (Espigão Mestre). Flowers from July to September; fruits from September to January. Specimens Examined: BRAZIL.  Tocantins,  Dianópolis, 11°33′55″ S, 46°19′50″ W, Scariot et al. 783 (CEN, K, NY); Dianópolis, em encosta de morro, 11°35′55″ S, 46°39′49″ W, 28 Sep 2003 (fr), Cavalcanti et  al. 3243 (CEN, SPF, NY); Dianópolis, beira de estrada após a Lagoa Bonita, 11°38′13″ S, 46°39′47″ W, 29 Sep 2003 (fr), Cavalcanti et al. 3489 (CEN, MO); rio da Conceição, 11°25′23″ S, 46°44′27″ W, 3 Jul 2009 (fl), Oliveira et  al. 1493 (CEN, HTO, IBGE, UEC). Taguatinga, estrada Taguatinga-Luis Eduardo Magalhães, 10 km do entroncamento, 12°20′56″ S, 46°20′19″ W, 28 Jan 2003 (fr), Cavalcanti et  al. 3148 (CEN); Taguatinga, divisa Tocantins/Bahia, km 8, subida da serra, 12°21′4″ S, 47°20′54″ W, 1 Sep 2004 (fl, fr), Cavalcanti et al. 3535 (CEN); Taguatinga, na subida da serra que faz divisa com a Bahia, a 10 km do entroncamento para Dianópolis, Espigão Mestre, 12°20′73″ S, 46°20′16″ W, 9 Apr 1997, Cavalcanti et al. 2338 (CEN). Diplusodon gracilis is distinguished by its delicate upper branches that are very thin and fragile. The plant has lax, frondose inflorescences with small, hyphodromous, narrowly elliptic leaves and bracts. The prophylls are obovate to oblong and reach the middle of the floral tube or surpasses it. The epicalyx segments are cylindrical and spreading, and they are longer than the sepals. Some plants observed in the field outside the period of flowering or fruiting, and some specimens with old fruit (Cavalcanti et al. 3148), have expanded leaves that are ca. 25 × 11 mm, elliptic, with two inconspicuous veins on either side of the midvein (eucamptodromous). There are 15 duplicates of the type material (Gardner 3155a) in European herbaria. The duplicate at the herbarium in Berlin (B) where Koehne worked was destroyed (F neg. nº 017922). Lourteig (1989) cited a specimen at P seen by Koehne as the holotype, considered here as the lectotype (ICBN Art. 9.17, ex. 14). 34. Diplusodon grahamae T.B.Cavalc., Acta Bot. Brasil. 21(4): 810, fig. 7–12. 2007. Type: Brazil. Goiás: Água Fria de Goiás, estação repetidora da Telebrasília de Roncador, 12 Jun 1993 (fl, fr), G. Hatschbach & E. Barbosa 59326 (holotype, MBM). Figs. 13.42 A–G, 13.46, 15.2 E Shrubs 0.2–1 m tall, erect, branched, sometimes caespitose, with an elongated xylopodium, not dark-brown when dry. Upper branches cylindrical to subquadrangular, unwinged, canescent, pubescent to hirsute, with white trichomes; internodes 0.1–1 cm long. Leaves decussate to verticillate, not imbricate, suberect, subcoriaceous, subsessile; petioles 0.5–1 mm long; blades 5–20 × 1–3.5 mm, slightly glaucous, sometimes discolorous, eucamptodromous to hyphodromous, narrowly elliptic to linear, base acute to attenuate, apex obtuse, margin subrevolute to revolute, adaxial and abaxial canescent, with sericeous indumentum or with sparse short trichomes, sometimes inconspicuous; secondary veins 1 on each side of the midvein, prominent; domatia present sometimes inconspicuous. Synflorescence frondose, botryum to diplobotryum type; accessory branches absent, principal

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florescence 8–20 cm long, paraclades 2–15 cm long; bracts similar to the vegetative leaves; flowers subsessile, hypopodium 0.8–1 mm long, epipodium 0–0.5 mm long; prophylls 2.5–4.5 × 1.2–1.6 mm, reaching the middle to the apex of the floral tube or sometimes surpassing the floral tube, obovate to lanceolate, unkeeled, base acute, apex obtuse, margin plane to involute; floral tube 3.5–6 mm long, when fresh green at the base, winish to the apex, funnel-shaped to oblong, pubescent, covered by short trichomes appressed and ascendent; sepals 2–2.5  mm long, triangular, not caudate, not deflexed, pubescent to sparse-sericeous, not ciliate; epicalyx segments 0.8–2 mm long, cylindrical, spreading to slightly deflexed, shorter than the sepals, pubescent to hirtellous to sparsely sericeous, sometimes canescent; corolla 2.5–3 cm in diam., rose to rose-magenta, petals 12–13 × 6–8 mm, elliptic to narrowly obovate, base acute to obtuse, apex obtuse; stamens 12–18, free portion of the filaments 5–8 mm long, surpassing the sepals 2 mm; ovary 2.5–2.8 × 2–2.5 mm, obovatedepressed to obovate-cordate, glabrous, style 7–12 mm long, ovules 10–18. Capsules 3–4.5  ×  3.1–3.5  mm, obovate to oblong, glabrous; seeds 8–15, 1.9–2.1 × 1.5–1.9 mm. Distribution and Ecology (Fig. 13.46)  Diplusodon grahamae is endemic to Goiás State, occurring in northeastern Goiás, including the region of Chapada dos Veadeiros, and Serra do Roncador. It occurs in open cerrado with rocky-sandy soil and campos rupestres at 1200 m. Flowers in May and June; fruits registered in June to July and October to November. Specimens Examined: BRAZIL. Goiás,  Água Fria de Goiás, rodovia GO-118, subida para a torre repetidora de Roncador, 8 May 2000 (fl), Hatschbach et  al. 70676 (MBM); Agua Fria de Goiás, 14°53′18.8″ S, 47°33′03″ W, 3 Jul 2015 (fl, fr), Cavalcanti & Pereira-Silva 3985 (CEN); 3.2  km GO-118, torre repetidora da Telebrasília (Roncador), 14°53′35″ S, 47°33′12″ W, Werneck 840 (CEN); Cavalcante, beira da estrada Cavalcante-Minaçu, próximo ao Kalunga Engenho II, 4 Jun 2006 (fl, fr), Dias et al. 725 (CEN); Mimoso de Goiás, Serra Geral, topo da cadeia da serra, 14°52′44″ S, 48°13′58″ W, 27 May 2015 (fl), G. Pereira-Silva et al. 16684 (B, BHCB, CEN, ESA, HUFU, HTO, K, M, MBM, MO, NY, P, RB, SPF, W); Mimoso de Goiás, Serra Geral, 14°52′32″ S, 48°15′12″ W, 23 Nov 2012 (fr), Pereira-Silva et al. 16463 (CEN, HUFU, SP); Mimoso de Goiás, no topo do morro, 14°52′46″ S, 48°14′00″ W, 9 Oct 2013 (fr), Cavalcanti & Pereira-Silva 3767 (BHCB, CEN, G, HUEFS, HTO, NY, OUPR, RB, SPF, UFMT, US, W). Mimoso de Goiás, ao norte de Padre bernardo, ca. 38,7 km do trevo GO-230 e 700 m pela trilha à esquerda, 14°52′34″ S, 48°13′43″ W, 5 Jun 2015 (fl), Mendoza et al. 5021 (CEN, USZ). Diplusodon grahamae is a slightly branched shrub recognized by its verticillate to congested decussate, linear to narrowly elliptic leaves with strongly revolute margins and sericeous, canescent pubescence. In addition, the epicalyx segments are spreading to deflexed, and the flowers have 12–15 stamens. Based on vegetative features, Diplusodon grahamae resembles D. puberulus from the states of Minas Gerais and Bahia, especially because of its linear to narrowly elliptic leaves with revolute margins. Diplusodon puberulus differs from D.

Descriptions

213

Fig. 13.42  Diplusodon grahamae. A. Habit. B. Section of internode. C. Leaf, adaxial surface. D. Leaf, abaxial surface. E. Detail of leaf abaxial surface indumentum. F. Prophyll, adaxial surface. G. Flower with prophyll, without petals. (A–G Pereira-Silva et al. 16463, CEN)

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grahamae in that it has hyphodromous venation, stellate trichomes on the branches and leaves, and a glabrous floral tube. 35. Diplusodon hatschbachii Lourteig, Bradea 5(19): 226. 1989. Type: Brazil. Goiás: Alto Paraíso de Goiás, Chapada dos Veadeiros, Sep 1975 (fl, fr), G. Hatschbach 38567 (holotype, P [01901899]; isotypes, MBM, MO). Figs. 13.43 A–H, 13.46 Subshrubs 0.4–1.2 m tall, erect, unbranched to slightly branched, with a xylopodium, canescent, not dark-brown when dry. Upper branches cylindrical to subquadrangular, unwinged, canescent, with abundant, thin, long trichomes with a thick base intermixed with very short trichomes, internodes 1.5–2 cm long. Leaves decussate, not imbricate, erect, sessile; blades 10–25 × 7–12 mm, coriaceous, not discolorous, acrodromous-basal-supranumerary, oblong-lanceolate, base obtuse to cordate, apex acute, margin plane to slightly revolute, blades slightly concave, bearing sparse thin, long trichomes concentrated on the veins and margins; secondary veins 3–5 on each side, 1–2 departing near the base of the midvein, the secondary and tertiary veins prominent on the adaxial surface, slightly less on the abaxial; domatia present. Synflorescence frondose, botryum to diplobotryum type, congested; accessory branches absent, principal florescence 2–7  cm long, botryum type, paraclades 3–8 cm long; bracts similar to the vegetative leaves, diminishing in size toward the apex of the branches; flowers pedicellate, hypopodium 1.5–2 mm long, epipodium 0.5–1 mm long, longer in fruit; prophylls 7–8 mm long, reaching the apex of the floral tube, elliptic, unkeeled, densely covered by white trichomes shorter than those of the leaves; floral tube 6–9 mm long, oblong, densely covered by white secretory, bulbous-based trichomes and in fruit with a constriction near the middle third and an acute, orange base; sepals 2.5–3 mm long, triangular, not caudate, not deflexed, ciliate, with secretory, bulbous-based trichomes, sometimes turning orange, mainly on the fruit; epicalyx segments 3.5–4.5 mm long, cylindrical to slightly dorsiventrally compressed, erect, the same size as or surpassing the sepals, ciliate, with the same trichomes as the sepals; corolla ca. 3.5 cm in diam., rose to pale lilac, petals 10–13 × 12–14 mm, obovate, apex obtuse; stamens 10–15, free portion of the filaments 7–8 mm long; ovary glabrous, style 11–18 mm long, ovules 15–42. Capsules 2.5–3.5  ×  3–3.5  mm, oblong, glabrous; seeds 9–18, 1.8–2 × 1.5–2.1 mm. Distribution and Ecology (Fig. 13.46)  Diplusodon hatschbachii is known from Chapada dos Veadeiros, in Goiás. It grows in the sandy humid soil in brejo and in the sandstone soil of campo rupestre and campo sujo at 1160 m. Flowers from April to June and September; fruits from February to June and September. Specimens Examined: BRAZIL. Goiás,  Água Fria de Goiás, estação repetidora da Telebrasília de Roncador, 12 Jun 1993 (fl, fr), Hastschbach & Barbosa 59303 (MBM); Água Fria de Goiás, rodovia GO-118, subida para a torre repetidora de Roncador, 8 May 2000 (fl, fr), Hatschbach et al. 70637 (MBM); Água Fria de Goiás, na torre repetidora de Roncador, 14°53′06″ S, 47°33′25″ W, 6 Apr 2006 (fl, fr),

Descriptions

215

Fig. 13.43  Diplusodon hatschbachii. A. Habit. B. Leaf, abaxial surface. C–D. Simple trichomes. E. Prophyll adaxial surface. F. Flower with prophyll, without petals. G. Fruit. H. Capsule. (A–G Hatschbach 38567, MBM)

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Cavalcanti et al. 3757 (CEN); Água Fria de Goiás, estarada de acesso a repetidora Roncador, ca. de 3  km da GO-118, 14°53′06″ S, 47°33′20″ W, 4 Feb 2014 (fr), Pereira-Silva et al. 16574 (CEN); Alto Paraíso de Goiás, estrada Alto Paraíso de Goiás-Terezina de Goiás, 10 Oct 1979 (fr), Heringer et al. 2310 (IBGE, NY); Alto Paraíso de Goiás, comunidade Moinho, 5 Sep 2004 (fl), Martins et al. 502 (UB). Diplusodon hatschbachii is a subshrub with oblong-lanceolate leaves that are covered with an indumentum composed of long, thin trichomes that have a slightly thickened base. The leaves also have evident domatia. The floral tube is covered with bulbous-based, secretory trichomes; the trichomes are similar to those on the flowers of D. sessiliflorus and D. divaricatus, which are also from Goiás. Diplusodon hatschbachii differs from these by the white trichomes that turn orange on and near the sepals, principally when in fruit. The fruits are urceolate shaped with a constriction in the middle third and acute base, characteristics that are also seen in the fruits of D. oblongus and D. appendiculosus. 36. Diplusodon helianthemifolius Shrubs 0.4–3  m tall, erect, much-branched, with a xylopodium, dark-brown when dry. Upper branches, whitish to rust-colored, quadrangular to rarely cylindrical, unwinged or with a narrow wing, puberulous to villous, with stellate white trichomes, or stellate trichomes mixed with simple, long or stellate trichomes with long arms, internodes 0.8–1.5 cm long. Leaves decussate, not imbricate, spreading to deflexed or suberect, sessile, subsessile to petiolate; petioles (when present), 2.5–5  mm long; blades 10–30  ×  5–36  mm, coriaceous, strongly discolorous, eucamptodromous to acrodromous-basal-supranumerary, elliptic to oblong, ovate to ovate-lanceolate, base attenuate or obtuse to cordate, apex acute, slightly acuminate, margin revolute, adaxial surface brownish-green to olive-green, sometimes dark-brown, slightly verrucose or minutely pilose, or with stellate trichomes with short arms that form very prominent clumps of trichomes, abaxial surface canescent, the stellate trichomes forming congested tufts, conferring a villous aspect to the surface, sometimes with mixed with long and simple or long-armed stellate trichomes; secondary veins 2–3 along the midvein, rust-colored on the abaxial surface, barely apparent or prominent, sulcate on the adaxial surface; domatia absent. Synflorescence frondose, pleiobotryum spiciform to diplobotryum spiciform type, congested; accessory branches absent, principal florescence 4–13.5  cm long, botryum spiciform type, paraclades 4–18.5 cm long; bracts 6–12 × 1.5–3 mm, narrowly elliptic to oblong-lanceolate, with revolute margins; flowers sessile to subsessile, hypopodium absent, epipodium ca. 1  mm long; prophylls 3–4  ×  1–2.5  mm, surpassing the middle of the floral tube or reaching the apex of the floral tube, ovateoblong or ovate to obovate, unkeeled, base acute, apex acute to obtuse, margin plane, ciliate; floral tube 4–5.5 mm long, larger in fruit, measured with the sepals 6.5–7 × 5–6 mm, funnel-shaped, minutely pilose; sepals 1.5–2 mm long, wine-colored, not caudate, not deflexed, ciliate or not; epicalyx segments 1.5–2 mm long, cylindrical or subulate, spreading to deflexed, with long cilia at the apex or not

Descriptions

217

ciliate; corolla 2.4–3 cm in diam., magenta, purple to reddish, petals 7–13 × 6–8 mm, obovate or oblong-elliptic, base obtuse, apex acute; stamens 12, free portion of the filaments 6–7  mm long, exserted, surpassing the sepals ca. 1  mm; ovary 1–2 × 1.5–1.8 mm, rhombic, glabrous, style 9–15 mm long, ovules 12–14. Capsules 4–5 mm × 3.5–4.5 mm, globose, glabrous; seeds 8–14, 2.5–3.1 × 2–2.5 mm. Distribution and Ecology (Fig.  13.46)  Diplusodon helianthemifolius occurs in Minas Gerais (Cadeia do Espinhaço)-Bahia (Espigão Mestre)-Goiás (Serra dos Pireneus) and the state of Maranhão. Key to the varieties 1. Leaves eucamptodromous, petiolate, petioles 2.5–5 mm long, elliptic to oblong, base acute to attenuate. Sepals ciliate, epicalyx segments with long cilia at the apex ...............................…………. D. helianthemifolius var. helianthemifolius 1. Leaves acrodromous-basal-supranumerary, sessile to subsessile, petioles 0–1 mm long, ovate or ovate-lanceolate to narrowly lanceolate, base obtuse to cordate. Sepals rarely ciliate, when ciliate, the cilia very short, epicalyx segments not ciliate ........……….....…..…. D. helianthemifolius var. pemphoides 36a Diplusodon helianthemifolius var. helianthemifolius Mart. ex DC., Prodr. 3: 94c. 1828. Friedlandia helianthemifolia Mart. ex DC., Prodr. 3: 94c, 1828, pro syn. Type: Brazil. Minas Gerais: Diamantina, in campis editis ad Tejuco Serro Frio (Diamantina), C. F. P. Martius obs. 1338 (lectotype, M [0146277, designated by Lourteig 1989: 229; isolectotype, G). Figs. 13.44 A–M, 13.46 Diplusodon velutinus Koehne, Pflanzenr. 4 (216): 202. 1903. Type: Brazil. Minas Gerais: Aug 1840, G. Gardner 4568 (first-step lectotype, designated by Lourteig 1989: 230, second-step lectotype, K [000528544], here designated; isolectotypes, K [000528543], SPF). Shrubs 1–3 m tall; internodes 1–1.5 cm long. Leaves 10–36 mm wide, suberect, elliptic to oblong, base acute to attenuate, petioles 2.5–5 mm long. Bracts 8–12 mm long, narrowly elliptic; flowers sessile, hypopodium and absent epipodium; prophylls 1–1.5 mm wide; sepals ciliate; epicalyx segments with long cilia at the apex; ovules ca. 12. Distribution and Ecology (Fig. 13.46)  Diplusodon helianthemifolius var. helianthemifolius is endemic to Cadeia do Espinhaço, Minas Gerais, around the municipalities of Diamantina and Serro, between 43º24′S, 18º37′W and 43º47′S, 18º04′W. It inhabits cerrado and campos rupestres at 1000–1400 m, cerradão, and areas of transition between cerrado and humid forest. Flowers from November to April; fruits from November to May.

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Specimens Examined: BRAZIL. Minas Gerais,  Diamantina, 10 Apr 1973 (fl), Anderson 8496 (NY); 18°10′5″ S, 43°37′07″ W, 26 Apr 1997 (fl), Cavalcanti et al. 2296 (CEN, NY); Diamantina, 18°10′56″ S, 43°42′18″ W, 14 Jan 1998 (fl), Forzza et  al. 631 (CEN [2], SPF); Diamantina, São João da Chapada, BR 367, km 14, 18°11′ S, 43°43′ W, 18 Feb 1989 (fl), Cavalcanti et  al. 252 (CEN, K, HB); Diamantina, na subida para a Usina, 16 Jan 1963 (fl), Duarte & Barroso 7907 (HB); Diamantina, 18 km E de Diamantina, 14 Mar 1970 (fl), Irwin et al. 27462 (MO, NY, P n.v., UB); Diamantina, ca. 18 km E na estrada a SW de Diamantina na estrada para Curvelo, 10 Apr 1973 (fl, fr), Anderson 8455 (NY, P n.v., UB); Diamantina, 16 Feb 1973 (fl), Hatschbach & Ahumada 31630 (MBM n.v., NY, P n.v.); Diamantina, estrada para Diamantina-Extração, 28 Jan 1986 (fl), Longhi-­Wagner et al. (CFCR) 9325 (CEN, K, SPF); Diamantina, estrada para Mendanha, próximo ao Morro do Cruzeiro, 19 Apr 1987 (fl, fr), Menezes et al. (CFCR) 10677 (K, SPF); Diamantina, estrada para Cristais ao longo do rio Biribiri, NW de Diamantina, 28 Nov 1985 (fl, fr), Graham 978 (MO, NY); 100 km N de Diamantina na estrada para Virgem da Lapa (BR 367), 26 Nov 1985 (fr), Graham 955 (MO, NY, RB); Diamantina, Parque Estadual do Biribiri, Alto da Jacuba, 18°11′46 S, 43°35′14″ W, 28 Fev 2016 (fl, fr), Faria 5390 (CEN, HDJF, RB, UB); Diamantina, margem da estrada de Diamantina para Milho Verde, 18° 22′ 37′′S, 43° 32′ 53′′W, 20 Mar 2016 (fl), Faria-Jr. 5546 (CEN, HDJF, UB); Pedro Lessa, 13 Mar 1995 (fl), Souza et al. 8422 (CEN, ESA); Serro, estrada de terra Serro-Milho Verde, a 11  km de Serro, 18 Jan 2003 (fl), Cavalcanti et  al. 3105 (CEN, MO); base do Pico do Itambé, 5 May 1942 (fl), Magalhães 1743 (HB). Diplusodon helianthemifolius var. helianthemifolius is represented by branched shrubs recognized by its strongly discolorous, petiolate (petioles 2.5–5 mm long), elliptic to oblong leaves with an acute to attenuate base and revolute margins, with abaxial surface whitish due to dense stellate indumentum, and flowers with epicalyx segments spreading or deflexed. The whole plant gets dark-brown when dry. Another, less conspicuous, characteristic that also identifies Diplusodon helianthemifolius var. helianthemifolius is the presence of relatively long, visibly white trichomes at the apex of the sepals and less often at the apex of the epicalyx segments. These trichomes can easily be seen on the flower buds, because the sepals are united at this stage, and the hairs form a white tuft at the apex of the bud. Candolle (1828) described Diplusodon helianthemifolius and D. pemphoides as species with stellate trichomes on the abaxial leaf surface, and he characterized D. pemphoides as having sessile or subsessile, ovate-lanceolate leaves with a cordate base. Saint-Hilaire (1833), in dealing with the same taxa, synonymized D. pemphoides and D. helianthemifolius under D. alutaceus A.St.-Hil. Koehne (1877) regarded D. helianthemifolius as a valid species and reduced D. pemphoides to a variety level, creating Diplusodon helianthemifolius var. pemphoides for specimens with leaves having obtuse or cordate leaf bases, and considered D. alutaceus A.St.-Hil. a synonym of this variety. Based on field studies, extensive herbarium material, and analysis of the types, D. helianthemifolius var.

Descriptions

219

Fig. 13.44  Diplusodon helianthemifolius var. helianthemifolius. A. Branch with inflorescence. B. Leaf, abaxial surface. C. Section of internode. D. Floral bud. E. Flower with prophylls, without petals. F. Petal. G. Inner surface of floral tube showing filaments, pistil removed. H. Prophyll, adaxial surface. I. Base of prophyll, abaxial surface, showing colleters. J. Transverse view of ovary showing two septa as thick structures toward the edge of the ovary and splitting the ovules into two groups. K. Septa of the ovary. L. Longitudinal and transversal view of the ovary showing half part of the ovules, septa removed. M. Leaf stellate trichome. (A, M Cavalcanti et al. 252, CEN; B–L Longhi-Wagner et al. [CFCR] 9325, SPF)

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helianthemifolius and var. pemphoides can be distinguished from each other. In var. helianthemifolius the leaves are strongly discolorous, oblong-lanceolate to elliptic, petiolate, with an attenuate base, besides possessing rust-­colored upper branches and white, long trichomes on the margin of the epicalyx segments. Such characteristics are best seen in the field where individuals of var. helianthemifolius reach 1–3 m in height. The individuals of var. pemphoides are smaller, 0.4–1 m tall, and have shorter internodes. The material annotated by Martius at M (Martius obs. 1338) has the original label with manuscript annotations. The fragment at G was studied by Candolle. Lourteig (1989) considered Diplusodon velutinus synonymous with D. helianthemifolius var. pemphoides, without explanation. However, the type of D. velutinus, Gardner 4568, does not differ from D. helianthemifolius var. helianthemifolius in any floral characteristic and is immediately recognized as D. helianthemifolius var. helianthemifolius by its petiolate and discolorous leaves with an attenuate base. For this reason, D. velutinus is here included in the synonymy of D. helianthemifolius var. helianthemifolius. There are two duplicates of Gardner 4568 at K. The one with a label bearing a determination by Koehne dated May 1902 was considered the holotype by Lourteig (1989) 36b Diplusodon helianthemifolius Mart. ex DC. var. pemphoides (DC.) Koehne, Fl. Bras. (Martius) 13(2): 325, pl. 60, fig. 10b, 1877. Diplusodon pemphoides DC., Prodr. 3: 94c. 1828. Type: Brazil. Minas Gerais: Serro, inter Vila do Príncipe et Diamantina, C. F. P. Martius s.n. (lectotype, M [0146278], designated by Lourteig 1989: 230; isolectotype, fragm. G). Figs. 13.45 A–H, 13.46 Diplusodon alutaceus A.St.-Hil., Fl. Bras. Merid. (A.St.-Hil.) 3: 153–154. 1833, non Pohl, Flora 10: 152. 1827. Type: Brazil. Minas Gerais: Morro do Mundo Velho, A. Saint-Hilaire s.n. (P [04713362]; photos at F [neg. nº 038310]). Diplusodon helianthemifolius DC. var. pemphoides (DC.) Koehne f. striatifolius Koehne, Bot. Jahrb. Syst. 29: 165. 1900. Type: Brazil. Minas Gerais: Serra do Cipó, 26 Apr 1892 (fl), C. A. W. Schwacke 8260 (lectotype, RB [00209380], here designated). Shrubs 0.4–1 m tall; internodes 0.8–1 cm long. Leaves 5–12 mm wide, spreading to deflexed, ovate to ovate-lanceolate to narrowly ovate-lanceolate, base obtuse to cordate, sessile to subsessile, petiole (when present) to 1 mm long. Bracts 6–7 mm long, oblong-lanceolate; flowers subsessile, hypopodium absent, epipodium ca. 1 mm long; prophylls ca. 2.5 mm wide; sepals rarely ciliate, when ciliate, the cilia minute; epicalyx segments not ciliate; ovules 14. Diplusodon helianthemifolius varies in the shape of the leaves and indumentum of the flower elements, which is the basis of two varieties. Distribution and Ecology (Fig.  13.46)  Diplusodon helianthemifolius var. pemphoides occurs in Cadeia do Espinhaço, Minas Gerais; in Espigão Mestre, Bahia; in Serra dos Pireneus, Goiás; and in the cerrado sensu stricto of Maranhão. It inhabits different environments in accordance with the elevation at which it occurs. It was

Descriptions

221

found in campo rupestre on white sandy soils at 1130–1244 m, in dense ecotonal vegetation of the carrasco type between 900 and 1110  m, and in cerrado (sensu stricto) habitats between 470 and 800 m. Flowers from September to April; fruits from September to July. Specimens Examined: BRAZIL.  Bahia,  Correntina, ca. 46.3  km  W de Correntina, na estrada para Brasília, 13°31′ S, 45º W, 16 Feb 2000 (fl, fr), Queiroz et al. 6076 (HUEFS); a 100 km de Correntina, 19 Nov 1991 (st), Vieira et al. 1163 (CEN); a 130 km de Correntina, 19 Nov 1991 (st), Vieira et al. 1165 (CEN, MO, NY); Santa Maria da Vitória à Brasília, km 190, 21 Jan 1997 (fl), Hatschbach et al. 66081 (CEN, MBM); Barreiras, rio Piau, ca. 150 km SW of Barreiras, 14 Apr 1966 (fl), Irwin et al. 14853 (NY, UB); Espigão Mestre, 34 km N of Barreiras, 2 Mar 1972 (fl), Anderson et al. 36408 (NY, UB); Espigão Mestre, ca. 100 km WSW of Barreiras, 7 Mar 1972 (fl), Anderson et al. 36736 (NY, UB); Roda Velha, 12 Jan 1977 (fl), Hatschbach 39434 (MBM); estrada para Acaba Vida, 6  km da BR-020, 24 Mar 1984 (fl), Almeida & Lourdes 280 (NY); Formosa do Rio Preto, Coaceral, 28 Mar 2000, Harley et al. 53755 (HUEFS); São Desidério, 12°28′ S 45°12′ W, 11 Apr 1989 (fl), Scariot et al. 525 (CEN, IBGE, UB n.v.). Goiás Corumbá de Goiás, Serra dos Pireneus, 20 km NW de Corumbá de Goiás, 27 Jan 1968 (fl), Irwin et al. 19262 (NY, UB); Corumbá de Goiás, Pico dos Pireneus, Serra do Catingueiro, 6  km de Cocalzinho, 52º S, 30º W, 14 Jan 1981 (fl), Nogueira et al. 84 (NY, SP); Corumbá de Goiás, Serra dos Pireneus, 18 Dec 1951 (fl), Macedo 3497 (US); Corumbá de Goiás, próximo de N.S. D’Abadia, 1840, Gardner 4566 (K); Pirenópolis, Serra dos Pireneus, ca. 20  km E of Pirenópolis, 14 Jan 1972 (fl), Irwin et  al. 34042 (NY); Pirenópolis, 8 Jan 1971, Rizzo 5883 & Barbosa 5132 (UFG); 11 Feb 1971, Rizzo 5998 & Barbosa 5246 (UFG); Pirenópolis, 7 Apr 1971, Rizzo 6188 & Barbosa 5436 (UFG); Pirenópolis, Serra dos Pireneus, estrada de terra Cocalzinho-Pirenópolis, dentro do Santuário “Morro Cabeludo” 15°49′ S, 48°53′ W, 24 Apr 1994 (fr), Splett 215 (CEN, UB); Pirenópolis, Parque Estadual Serra dos Pireneus, 30 Jan 2004 (fr), Pastore et  al. 761 (CEN). Maranhão, Balsas, Projeto de Colonização Agrícola Gerais de Balsas, região de Vila Kissy, próximo ao rio Tem Medo (lote 23), 08°38′ S, 46°43′ W, 20 Mar 2000 (fl), Walter et al. 4421 (CEN); Balsas, condomínio Kissy U17, 08°37′02″ S, 46°44′5″ W, 19 Mar 1999 (fl, fr), Pereira-Silva 4098 (CEN). Minas Gerais, Abadia, May 1840 (fr), Gardner 4133 (G, K); Bocaiúva, rodovia BR-367, descida para o rio Jequitinhonha, 20 Nov 1997 (fl, fr), Hatschbach et al. 67314 (MBM); Botumirim, contrafortes orientais da Serra da Canastra, trilha do Cruzeiro a partir do posto de gasolina até o alto da serra nas nascentes do córrego São Domingos, 29 Sep 1997 (fl, fr), Mello-Silva et al. 1458 (CEN, SPF); Botumirim, estrada para Adão Colares/Barrocão, 14 km de Botumirim, 850 m, 19 Nov 1992 (fl), Mello-Silva et al. 647 (CEN, SPF); Botumirim, 14 km NW de Botumirim, caminho até Adão Colares e Barrocão, 16° 45′ S, 43° 5′ W, 20 Nov 1992 (fl, fr), Mello-Silva et al. 727 (CEN, CTES n.v., P n.v., RB, SPF); Botumirim, Serra da Canastra, São Domingos, encostas do ribeirão São Domingos, em trilha para o alto da serra, 16°51′26″ S, 43°01′36″ W, 20 Dec 2004 (fr), Mello-Silva & Forzza 2709 (CEN, RB, SPF); Chapada Gaúcha, Serra das Araras, estrada entre Januária e Chapada Gaúcha,

222

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Fig. 13.45  Diplusodon helianthemifolius var. pemphoides. A. Branch with inflorescence. B. Section of internode. C. Leaf, abaxial surface. D. Floral bud. E. Flower with prophylls, without petals. F. Fruit. G. Ovary. H. Seed. (A–H Cavalcanti et al. [CFCR] 8528, SPF)

Descriptions

223

Fig. 13.46  Geographic distribution of Diplusodon gracilis, D. grahamae, D. hatschbachii, D. helianthemifolius var. helianthemifolius, and D. helianthemifolius var. pemphoides

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15°28′11″ S, 45°26′25″ W, 31 Jan 2019 (fl, fr), Flores et al. 441 (CEN, ESA n.v.); Chapada do Norte, vale do Jequitinhonha, próximo à nascente do rio Água Suja, 3 May 2000 (fr), Roschel & Guimarães 716 (CEN, OUPR); Cristália, Serra de Cristália, Torre, 16°43′07″ S, 42°55′21″ W, 10 Sep 2003 (fr), Conceição et al. 725 (CEN, HUEFS); Cristália, Serra de Cristália, perto da base do Morro do Chapéu, 2 Sep 1988 (fl), Wanderley & Kral 1443 (SP, SPF); morro da antena de televisão, 16°43′28″ S, 42°55′42″ W, 12 Jul 2001 (fl, fr), Souza et  al. 25818 (CEN, ESA); Chapada das Galinhas, 10 Feb 1991 (fl), Hatschbach et  al. 55006 (MBM, MO); Rodovia Grão-Mogol a Cristália, próximo do rio Soberbo, 21 Jul 2002 (fl), Hatschbach et al. 73721 (CEN, MBM); Diamantina, BR 367, Diamantina-Araçuaí, km 89, 17°32′ S, 43°09′ W, 17 Feb 1989 (fl), Cavalcanti et al. 230 (CEN, K, RB, SPF); Grão-Mogol, torre da Telemig, estrada para Salinas, BR 251, 28 km a leste do entroncamento com a MG-120, 16°13′01″ S, 42°52′17″ W, 10 Jan 2006 (fl, fr), Sena et al. 1675 (CEN, HUEFS); Grão-Mogol, Plantar MG-15-fazenda Tamanduá, 9 Oct 2005 (fl), Tameirão-Neto 4017 (BHCB, CEN); Grão-Mogol, 48.5 km W da ponte sobre o rio Vacaria, Serra da Bocaina, 28.5 km E de Caveira, 31 Jan 1991 (fl), Taylor et al. 1503 (K n.v., SPF); Grão-Mogol, 12 Nov 1938 (fl, fr), Markgraf 3426 (R); Grão-Mogol, Povoado de Alegre, 24 Oct 1991 (fr), Carvalho & STS 705 (CEN); Grão-Mogol, 7 Jun 1979 (fr), Tenório & Freitas s.n. (CEN, CETEC 01077); GrãoMogol, Jambeiro, a 7 km de Grão-Mogol, 5 Sep 1985 (fl), Cavalcanti et al. (CFCR) 8528 (K, MBM, SPF, UB); Grão-Mogol, na estrada para o rio Ventania, 16°32′ S, 42°49′ W, 5 Sep 1990 (fl, fr), Esteves et  al. (CFCR) 13372 (CEN, SPF); GrãoMogol, Freire-Fierros et al. (CFCR) 12730 (CEN, SPF); Grão-Mogol, na saída da cidade, estrada para o rio Ventania, 25 Feb 1986 (fl, fr), Cavalcanti et al. (CFCR) 9644a (K, SPF, UEC); Grão-Mogol, Cavalcanti et al. (CFCR) 9650 (CEN, F, K, SPF, UEC); Grão-Mogol, próximo à antena de TV, carrascal, 6 Nov 1986 (fl, fr), Cavalcanti et al. (CFCR) 10175 (CEN, K, SPF); Grão-Mogol, estrada Grão-Mogolrio Ventania, km 7, 16°38′ S, 42°49′ W, 22 Feb 1989 (fl), Cavalcanti et  al. 282 (CEN, HUFU, SPF); Grão-Mogol, BR 251, Ribeirão da Mortes, 16°34′ S, 42°54′ W, 22 Feb 1989 (fl, fr), Cavalcanti et al. 285 (CEN, SPF), Grão-Mogol, vale do córrego Escurona, ca. 16°36′ S, 42°58′ W, 13 May 1990 (fl, fr), Zappi et al. (CFCR) 12879 (CEN, SPF); Itacambira, estrada Juramento-Itacambira, no alto da Serra de Itacambira, á direita, 30 Sep 1997 (fl), Mello-Silva et al. 1487 (NY, SP n.v., SPF); Itacambira, entroncamento BR367-Itacambira, 11 km do entroncamento, 17°15′ S, 43°03′32″ W, 5 Feb 2015 (fl, fr), Cavalcanti et al. 3899 (CEN); Joaquim Felício, Serra do Cabral, próximo ao Armazém da Lage, 18 Nov 1997 (fl), Hatschbach et al. 67228 (MBM); Mendanha, 15 Feb 1882 (fl), Glaziou 12675 (BR, LE n.v., R); Minas Novas, Buriti, 23 Mar 1993 (fl, fr), Kameyama & Esteves 41 (CEN, SPF); Minas Novas, estrada de terra para campo Buriti, na estrada Capelinha-Minas Novas, 500 m, 17°17′38″S, 43°37′54″W, 21 Jan 2003 (fl), Cavalcanti et al. 3135 (CEN, MO); Minas Novas, estrada Chapada do Norte-Minas Novas, km 10, 17°08′35″ S, 42°34′38″ W, 3 Sep 2003 (fr), Cavalcanti et al. 3199 (CEN); Rio Pardo de Minas, Parque Estadual de Serra Nova, 15°39′14″ S, 42°43′52″ W, 20 Mar 2012 (fl), Saiter 403 (BHCB); São João da Chapada, 29 Feb 1970 (fr), Irwin et al. 28627 (NY, RB,

Descriptions

225

UB); rio Vermelho, Serra do Ambrósio, Fazenda Vargem do Anjo, 18°08′32″ S, 43°03′34″ W, 25 Apr 1997 (fl), Cavalcanti et al. 2287 (CEN, MO, NY); Santana do Riacho, Parque Nacional da Serra do Cipó, 18 Apr 1992 (fl), Pereira & Lucca 930 (BHCB); Santana do Riacho, Serra do Cipó, 22 Mar 1992 (fl), Pereira & Lucca 931 (BHCB, CEN); São Gonçalo do Rio Preto, 18°07′20–40″ S, 43°20′36″ W, trilha do alojamento até o córrego das Éguas, 19 Feb 2002, Lombardi 4556 (BHCB, CEN); Senador Mourão, 1990 (fr), Teixeira & Brina s.n. (BHCB 832); caminho da base do IBAMA do rio Cipó para o Capão dos Palmitos, 25 Mar 1991 (fl), Pirani et  al. (CFCR) 12024 (CEN, SPF); Turmalina, BR-120, próximo a margem direita do rio Itamarandiba, 17°16′30″S, 42°48′50″W, 20 Jan 2003 (fr), Cavalcanti et  al. 3132 (CEN, MO); estrada Capelinha-Minas Novas, Turmalina, 1 km do entroncamento, lado direito da estrada, 17°18′16″ S, 42°40′5″ W, 21 Jan 2003 (fl), Cavalcanti et al. 3133 (CEN, MO); Turmalina, estrada Mato Grande-Turmalina, 7 Jan 1998 (fl), Carvalho 487 (CEN, VIC). The individuals of Diplusodon helianthemifolius var. pemphoides are branched shrubs characterized by its sessile, strongly discolorous, oblong-lanceolate leaves with a cordate base, revolute margins, and the abaxial surface covered with stellate trichomes and rust-colored veins and also by flowers that have deflexed epicalyx segments. The whole plant gets dark-brown when dry. There are small variations in the indumentum of the branches and leaves in populations of the different states in which the species occurs. There is a pattern in which the indumentum is composed only of prominent stellate trichomes with short arms that form tufts or clumps on both surfaces of the leaves and branches (Cavalcanti et al. [CFCR] 9644a; Pereira & Lucca 930). In another pattern, the branches and the leaves possess longer stellate and simple trichomes, and the adaxial surface of the leaves is smooth and only minutely pilose (Gardner 4133; Irwin et al. 31343; Scariot et al. 525). In a third pattern, the abaxial leaf surface and branches have more simple trichomes than stellate ones, and the adaxial surface of the leaves is totally glabrous, smooth, and shiny (Irwin et al. 28623, Zappi et al. [CFCR] 12879). Although the species is distributed in three different Brazilian states, this variation is not linked to geographic distribution, and the three patterns were found in all three regions. Lourteig (1989) considered Diplusodon helianthemifolius var. pemphoides (DC.) Koehne f. striatifolius Koehne synonymous with D. helianthemifolius var. helianthemifolius. This form, described by Koehne (1900) from Minas Gerais based on Schwacke 8260, was distinguished on the basis of its palmate-veined, plane leaves that become pinnate-veined along the midvein. Lourteig (1989) cited “Schwacke 8260 at herbarium B, destroyed.” Material of Schwacke 8260 was located in the herbarium of the Botanical Garden of Rio de Janeiro (RB), and it does not differ from Diplusodon helianthemifolius var. pemphoides. Therefore, D. helianthemifolius var. pemphoides f. striatifolius is moved from the synonymy of D. helianthemifolius var. helianthemifolius and referred to that of D. helianthemifolius var. pemphoides.

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37. Diplusodon heringeri Lourteig, Bradea 5(19): 239. 1989. Type: Brazil. Goiás: Cavalcante, rodovia GO-12, descida para Chapada dos Veadeiros, 26 May 1975 (fl), G. Hatschbach 36895 (holotype P, [01901898]; isotype, MBM). Figs. 13.47 A–H, 13.51 Shrubs 0.5–3 m tall, erect, much-branched, canescent, not dark-brown when dry. Upper branches cylindrical, unwinged, hirsute to hispid, internodes 0.5–1 cm long. Leaves subsessile; petioles 0.5–1 mm long; blades 6–10(−17) × 4–9 mm, decussate, not imbricate, suberect, coriaceous, not discolorous, acrodromous-basal-supranumerary, ovate to elliptic, base obtuse, apex obtuse, margin revolute, adaxial surface hirsute, verrucose, abaxial surface hirsute, covered by white, long trichomes; secondary veins 3 on each side along the midvein, sometimes 2 departing from the base of the midvein, prominent domatia evident. Synflorescence frondose, pleiobotryum type, congested; accessory branches absent, principal florescence truncate, paraclades 5–10 cm long; bracts similar to the vegetative leaves in shape and size, only a little smaller; flowers sessile, in congested coflorescences at the apex of the branches; prophylls 3–4.5 mm long, reaching the apex of the floral tube or slightly less, obovate to elliptic, unkeeled, base acute, apex obtuse, margin plane; floral tube 4–4.5 mm long, yellow, funnel-shaped, hirsute; sepals 2–2.5 mm long, triangular, not caudate, not deflexed, ciliate, with long trichomes; epicalyx segments 2.5–5(−6) mm long, narrowly lanceolate to cylindrical, erect, the same size as the sepals, rarely longer, with long trichomes spreading, white and sparse; corolla 2–2.5 cm in diam., bright rose, petals 9–11 × 5–6 mm, elliptic to obovate, apex obtuse; stamens (17–)18, free portion of the filaments 7–9  mm long, slightly exserted; ovary 1.5–2 × 2–2.9 mm, rounded, glabrous, style 9–12 mm long, ovules 12–14. Capsules 3.4–3.5 × 2.9–3.5 mm, globose, glabrous; seeds 5–7, ca. 2 × 1.5–2 mm. Distribution and Ecology (Fig.  13.51)  Diplusodon heringeri is endemic to the region of Chapada dos Veadeiros and Cavalcante, in the northeast of Goiás, in highelevation rocky cerrados and campos rupestres at 725–1400  m. Flowers April to September; fruits registered from May to September and January. Specimens Examined: BRAZIL.  Goiás,  Alto Paraíso de Goiás, Chapada dos Veadeiros, a 12 km da cidade, na estrada que dá acesso à Nova Roma, 13°59′ S, 47°21′ W, 21 May 1994 (fl, fr), Silva et al. 1937 (IBGE, P n.v., RB n.v., UEC n.v.); Alto Paraíso de Goiás, a 37 km de Terezina de Goiás em direção a Alto Paraíso de Goiás, 13°58′ S, 47°30′ W, 30 May 1990 (fl), Pedralli et  al. 3366 (CEN); Alto Paraíso de Goiás, Fazenda Canastra, 30 km de Alto Paraíso, 14°02′ S, 47°26′ W, 31 May 1994, Ratter et  al. 7315 (UB); Cavalcante, estrada Teresina de GoiásCavalcante, 10 km de Cavalcante, 1 Sep 2004 (fl, fr), Cavalcanti et al. 3547 (CEN, MO); Cavalcante, 10 Jul 2011 (fl, fr), Silva 665 (UB); Teresina de Goiás, km 34 da rodovia Alto Paraíso para Teresina de Goiás, 13°45′ S, 47°20″ W, 20 Jun 1995 (fl, fr), Pereira et  al. 2774 (CEN, IBGE, UB); Teresina de Goiás, rodovia GO-118, 13°38′46″ S, 47°21′33″ W, 7 Nov 2004 (fr), Pastore & Suganuma 1108 (CEN); Teresina de Goiás, GO-118, estrada Teresina de Goiás-Alto Paraíso de Goiás, a 30 km de Teresina, 13°53′45″ S, 47°21′33″ W, 29 Jan 2003 (fr), Cavalcanti et al.

Descriptions

227

Fig. 13.47  Diplusodon heringeri. A. Branch with inflorescence. B, E. Leaf, abaxial surface. C. Detail of leaf abaxial surface indumentum. D. Simple trichome. F. Section of internode. G. Floral bud. H. Flower with prophylls, without petals. (A–H Pedralli et al. 3366, CEN)

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3155 (CEN, MO); Teresina de Goiás, estrada Alto Paraíso de Goiás-Teresina de Goiás, a 59  km de Alto Paraíso de Goiás, no camping próximo à cachoeira, 13°45′72′′ S, 47°13′20′′ W, 4 Apr 1997 (fl), Cavalcanti et al. 2211 (CEN, MO, NY); Teresina de Goiás, Fazenda Hotel Ecológico Alpes Goianos, rodovia GO-118, km 202, 13°53′59.1″ S, 47°23′48.9″ W, 31 Jul 2000, Souza et al. 24735a (ESA, PAMIG, UB); Teresina de Goiás, 45  km, 13°51′06.5″S, 47°17′36″ W, 1 Sep 2004 (fr), Cavalcanti et al. 3537 (CEN); Teresina de Goiás, GO-118. Teresina-Alto Paraíso de Goiás, km 24, 1 Sep 2004, Cavalcanti et al. 3539 (CEN, SPF). Diplusodon heringeri is characterized by canescent, branched shrubs with small (6–)10[−17] × 4–9 mm, hirsute, ovate to elliptic leaf blades with revolute margins. The flowers are small and covered by long trichomes that are less sparse on the epicalyx. The epicalyx segments are the size of the sepals, and they are erect, and the androecium consists usually of 18 stamens. Vegetatively, Diplusodon heringeri resembles D. parvifolius, a species from Bahia, which also has small, ovate leaves with revolute margins. These species differ in indumentum, and the trichomes of D. parvifolius are shorter, like those of the flowers. In addition, the flowers of D. parvifolius have spreading to deflexed epicalyx segments that are very short (ca. 1 mm long), obovate prophylls that reach the middle of the floral tube or less, and an androecium with 12 stamens. 38. Diplusodon hexander Mart. ex DC., Prodr. 3: 94a. 1828. Pemphis hexandra Mart. ex Koehne, Fl. Bras. (Martius) 13(2): 324. 1877, pro syn. Friedlandia hexandra Mart. ex DC., Prodr. 3: 94a, 1828, pro syn. Type: Brazil. Minas Gerais: in campis ad Tejuco (Diamantina) et Vila do Principe, C. F. P.Martius s.n. (first-step lectotype, designated by Lourteig 1989: 214, second-step lectotype, M [146275], here designated; isolectotypes, M [0146274], fragm. G). Figs. 13.48 A–I, 13.51, 15.3 B Diplusodon angustifolius Mart. ex DC., Prodr. 3: 94a. 1828. Diplusodon angustifolium, cited in error by Lourteig (1989). Friedlandia angustifolia Mart. ex DC., Prodr. 3: 94a, 1828, pro syn. Diplusodon hexander Mart. ex DC. f. angustifolius (Mart. ex DC.) Koehne, Fl. Bras. (Martius) 13(2): 324, pl. 60, fig. 9. 1877. Diplusodon hexander var. angustifolia [as angustifolium], cited in error by Lourteig (1989). Type: Brazil. Minas Gerais: in campis ad Serro, C.  F. P.  Martius obs. 1504 (first-step lectotype, designated by Lourteig 1989: 214, second-step lectotype, M [0146276], here designated; isolectotypes, fragm. G). Subshrubs 0.4–1.7  m tall, erect, much-branched, without a xylopodium, glabrous, usually dark-brown when dry. Upper branches subquadrangular to quadrangular, chestnut brown, unwinged, internodes ca. 1  cm long. Leaves decussate to sometimes verticillate, not imbricate, erect, subsessile to petiolate; petioles 1–1.5 mm long; blades 6–15 × 1–4 mm, glaucous, coriaceous to slightly fleshy, not discolorous, hyphodromous, linear to linear-spathulate or elliptic-linear, concave, base acute to slightly attenuate, apex acute to obtuse, margin revolute to the midvein, sometimes subrevolute, adaxial surface dark-green, glaucous, midvein very

Descriptions

229

sulcate, abaxial surface olive-green, with a very prominent midvein, lateral veins absent; domatia absent. Synflorescence frondose, diplobotryum to pleiobotryum type, lax; accessory branches absent, principal florescence 4–10 cm long, botryum type, sometimes caducous, paraclades 2.5–18 cm long; bracts similar to the vegetative leaves in shape and size; flowers pedicellate, hypopodium 1.5–3 mm long, epipodium 0–2 mm long; prophylls 5–6 × 1.5–2.5 mm, reaching or exceeding the apex of floral tube ca. 1.5 mm, obovate to widely obovate, elliptic-lanceolate to spathulate, unkeeled, base and apex acute, margin plane; floral tube 4–6 mm long, greenish, funnel-shaped; sepals 1.5–2  mm long, larger in fruit, 4.5–5.5  mm long, wine-colored, triangular, not caudate, not deflexed; epicalyx segments 2–5  mm long, cylindrical to dorsiventrally compressed, erect, longer than the sepals; corolla 1.5–3 cm in diam., rose-magenta to reddish, petals 11–13 × 7–9 mm, obovate-elliptic to oblong, apex obtuse; stamens 6, free portion of the filaments 4.5–6 mm long, exserted, surpassing the sepals ca. 1 mm; ovary 1.5–1.8 × 2 mm, obconic, glabrous, style 9–14 mm long, ovules 8–18. Capsules 4–4.5 × 3.5–4 mm, globose, glabrous; seeds 8–10, 2.5–2.8 × 2.1–2.5 mm. n = 15 (Graham 1992). Distribution and Ecology (Fig.  13.51)  Diplusodon hexander is endemic to the central portion of Cadeia do Espinhaço, around Serra do Cipó and Diamantina areas, in Minas Gerais State, occurring in campos rupestres and campos sujos, 831– 1392 m elevation. Flowers and fruits from November to June. Specimens Examined: BRAZIL.  Minas Gerais,  Conceição do Mato Dentro, Solidão, Fazenda Boa Esperança, propriedade da Anglo American, 18°55′28″ S, 43°28′40″ W, 29 Jun 2016 (fr), Faria et al. 6106 (CEN, HDJF, RB, UB); Cunha Magalhães, 15  km  N de Cunha Magalhães, 20 Jan 1972 (fl), Hatschbach et  al. 29012 (MBM); Diamantina, 18°35′35″ S, 43°57′51″ W, 6 Jan 2006 (fr), Proença & Harris 3123 (CEN, UB); Diamantina, Área de Proteção Ambiental Pau de Fruta (Copasa), 18°15′14″ S, 43°38′42″ W, 14 Feb 2001 (fl, fr), Lombardi 4252 (BHCB, CEN); Diamantina, 18°12′ S, 43°36′ W, 26 Apr 1997 (fl), Cavalcanti et al. 2293 (CEN, NY); BR-367, Diamantina-Araçuaí, km 27, 18º08′S, 43º32′W, 17 Feb 1989 (fl, fr), Cavalcanti et al. 241 (CEN, K, NY, SPF); Diamantina, estrada para São João da Chapada-BR 367, km 3, 18°03′ S, 43°48′ W, 18 Feb 1989 (fl), Cavalcanti et al. 250 (CEN, CEPEC, K, MBM, NY, SPF); Diamantina, 3  km  N de São João da Chapada, 28 Mar 1970 (fl), Irwin et al. 28443 (K, NY, P, UB); Diamantina, ca. de 25 km da cidade na estrada para São João da Chapada, 9 Jun 1991 (fl, fr), MelloSilva et al. 405 (SPF); Diamantina, estrada para Biribiri, 21 Mar 1993 (fl), Kameyama & Esteves 40 (CEN, SPF); Diamantina, 23 Jan 1978 (fl), Hatschbach 40834 (SPF, MBM, MO, P); Diamantina, Hatschbach 40839 (MBM); Diamantina, 26 Mar 1892 (fl), Schwacke 8262 (OUPR, RB); Diamantina, estrada Diamantina-Biribiri, ca. de 3 km do estádio J.K., 22 Nov 1985 (fl, fr), Cavalcanti et al. (CFCR) 8552 (K, NY, SPF, UB); Diamantina, estrada Diamantina-Biribiri, km 4.5, 18°13′ S, 43°41′ W, 16 Feb 1989 (fl. fr), Cavalcanti et al. 201 (CEN, K, MBM, SPF); Diamantina, 18°15′ S, 43°40′ W, 8 Mar 1995 (fl, fr), Splett 812 (CEN, UB); Diamantina, Sopa, 21 Mar 1892 (fl, fr), Schwacke 8259 (BHCB, RB); Diamantina, a NW de Diamantina, 28 Nov 1985 (fl), Graham 979 (MO, NY); Diamantina, Morro dis Cristaes, près

230

13  Systematic Treatment

Fig. 13.48  Diplusodon hexander. A. Branch with inflorescence. B, D. Leaf, abaxial surface. C. Leaf adaxial surface. E. Prophyll, adaxial surface. F. Flower with prophylls, without petals. G. Fruit. H. Capsule. I. Seed. (A–I Cavalcanti et al. [CFCR] 8552, SPF)

Descriptions

231

Diamantina, 4 Apr 1892 (fl, fr), Glaziou 19171 (K, LE n.v., R, P); Diamantina, subida para a Usina, 16 Jan 1963 (fl), Duarte & Barroso 7910 (HB, NY, RB); Diamantina, 25 May 1955 (fl), Pereira 1547 (RB); Diamantina, ca. 12 km NE de Diamantina, estrada para Mendanha, 28 Jan 1969 (fl), Irwin et al. 22741 (COL, K, MO, NY, P, UB); Diamantina, rodovia Mendanha-Inhaí, 18 Mar 1997 (fl, fr), Hatschbach et  al. 66450 (M, ESA, NY); Diamantina, 20–26  km WSW de Diamantina, caminho a Conselheiro Mata, MG-220, ca. 18°17′ S, 43°49′ W, 10 May 1990 (fr), Arbo et al. 4371 (CEN, CTES n.v., P n.v., SPF); Diamantina, estrada Diamantina-Conselheiro Mata, 10 Jan 1988 (fl), Mello-Silva et al. (CFCR) 11810 (CEN, SPF); Diamantina, estrada Diamantina-­Conselheiro Mata, km 185, 23 Feb 1986 (fl, fr), Cavalcanti et al. (CFCR) 9496 (K, SPF, UEC); Diamantina, estrada Diamantina-Conselheiro Mata, km 184, 23 Feb 1986 (fl), Cavalcanti et al. (CFCR) 9507 (K, SPF); Diamantina, estrada Diamantina-Conselheiro Mata, km 182, 29 Jan 1986 (fl, fr), Cordeiro et al. (CFCR) 9387 (K, SPF); Diamantina, estrada DiamantinaConselheiro Mata, km 175, 25 Feb 1987 (fr), Zappi & Salatino (CFCR) 10388 (K, SPF); Diamantina, estrada Diamantina-Conselheiro Mata, km 174, Zappi et  al. (CFCR) 9449 (K, MBM, RB, SPF); Diamantina, estrada Conselheiro Mata, a 5 km da estrada Curvelo-Diamantina, 18 Dec 1985 (fl), Souza et al. (CFCR) 8786 (K, F, SPF); Diamantina, rodovia Guinda-Conselheiro Mata, 16 Mar 1987 (fl), Hastchbach et al. 51500 (P); Diamantina, rodovia Curvelo-Diamantina, 18°35′39″ S, 43°47′50″ W, Diamantina, ca. 10  km SW de Diamantina, 3 Feb 1972 (fl), Anderson et  al. 35237 (MO, NY, P); Diamantina, 18 km na estrada SW de Diamantina, na estrada para Curvelo, 10 Apr 1973 (fr), Anderson 8501 (NY); Diamantina, 18 km na rodovia SW de Diamantina, 10 Apr 1973 (fl, fr), Anderson 8503 (P); Diamantina, Biribiri, Glaziou 19174 p.p. (P); Gouveia, ca. 37 km de Gouveia para Curvelo, 5 Apr 1998 (fr), Souza et al. 20875 (CEN, ESA, MBM); Diamantina, distrito de Milho Verde, Área de Proteção Ambiental Águas Vertentes, estrada Milho Verde-Capivari, 18° 27′ 41′′S, 43° 27′ 51′′W, 12 Fev 2014 (fl, fr), Martinelli et al. 18215 (CEN, RB); Gouveia, 30 km SW de Diamantina na estrada para Gouveia, 13 Jan 1969 (fl), Irwin et  al. 21821 (MO, NY, P); Gouveia, BR-259, Diamantina-Curvelo, 18°33′39″ S, 43°50′58″ W, 20 Jan 2003 (fl), Cavalcanti et al. 3129 (CEN, MO); Gouveia, BR-529 Diamantina-Curvelo, a 100 metros da BR na estrada para a represa da CEMIG, 18°38′93″ S, 43°59′ W, 26 Apr 1997 (fl, fr), Cavalcanti et  al. 2298 (CEN, NY); Gouveia, subida da Serra do Espinhaço, 25 Feb 1975 (fl), Hatschbach et al. 36407 (BR, MBM, MO, SPF); Gouveia, 3 km SW da ponte sobre o córrego do Tigre, 12 Mar 1982 (fl, fr), Graham 865 (MBM, NY); Gouveia, estrada de terra para a fazenda do Sr. Everaldo, 14 Apr 1987 (fl, fr), Zappi et al. (CFCR) 10451 (K, SPF); Gouveia, Fazenda Contagem, 24 Feb 1988 (fl, fr), Cavalcanti et al. (CFCR) 9567 (CEN, K, F, SPF, UEC); Mendanha, morro da torre da repetidora dentro da cidade de Mendanha, 18°06′11″ S, 43°31′14″ W, 9 Mar 2014 (fr), Cavalcanti et al. 3930 (CEN); Palmital, Serra do Palmital, 29 May 1992 (fr), Mendes-Magalhães 1750 (UB); Serra do Cipó, 26 Apr 1892 (fl, fr), Schwacke 8264 p.p. (BHCB, OUPR, RB); Serro, ca. 27 km N de Serro na estrada MG-2 para Diamantina, 26 Feb 1968 (fl), Irwin et al. 20941 (NY, P, UB); Serro, estrada Milho Verde-Diamantina, a 11  km de Milho Verde, 18°24′12″ S, 43°30′51″ W, 18 Jan 2003, Cavalcanti et al. 3114 (CEN, MO); without

232

13  Systematic Treatment

reference to municipal district: Gardner 4562 (G, K, OXF, W); Varnay 1262 (OUPR); Varnay 1265 (OUPR); Riedel 1216a (LE n.v.); Riedel 1216b (LE n.v.); Riedel s.n. (OXF, P [2]). Diplusodon hexander is represented by much-branched and glabrous subshrubs characterized by narrow, glaucous leaves, which are revolute and hyphodromous, and by flowers that have six stamens. Diplusodon glaziovii is the morphologically closest species due to the narrow leaves and flowers with six stamens. However, D. glaziovii has leaves narrowly obovate to narrowly elliptic with flat margin and not glaucous. There are two Martius specimens at M annotated by Koehne as Diplusodon hexander. One specimen has the number 1277 at the foot of the label; however, the number Martius obs. 1277 corresponds to a palm specimen (Esser, pers. comm), and therefore, this specimen of D. hexander should be cited as Martius s.n. and is here designated as second-step lectotype. Lourteig (1989) cited Martius s.n. as the holotype. Koehne (1877) studied the type material of Diplusodon angustifolius DC. (Martius obs. 1504) and recognized D. hexander f. angustifolius, based on its linear leaves with revolute margins reaching the midvein. The material cited by Koehne (1903) for f. angustifolius (Martius obs. 1504 and Schwacke 8264) and other populations in the field exhibit these characteristics. Specimens from the municipality of Gouveia (Graham 865, Hatschbach 36407 and Zappi et al. [CFCR] 10451) have more narrowly linear leaves than those from the municipality of Diamantina. In the field, D. hexander exhibits great variation in the width of individual leaves ranging from 1 to 5 cm with some individuals exclusively with one extreme of variation of this characteristic and others showing width variation on a single plant. For this reason, it is not possible to recognize infraspecific taxa based on these characteristics for D. hexander. Lourteig (1989) accepted the infraspecific taxa proposed by Koehne (1877). However, she referred to them as varieties, not forms. 39. Diplusodon hirsutus (Cham. & Schltdl.) DC., Prodr. 3: 94c. 1828. Friedlandia hirsuta Cham. & Schltdl., Linnaea 2: 352. 1827. Type: Brazil. Minas Gerais: F. Sellow 2053 (first-step lectotype, designated by Lourteig 1989: 225, secondstep lectotype, HAL [0098374], here designated; isolectotypes, HAL [0053405, 0120099], K [2], P). Figs. 13.49 A–I, 13.51 Diplusodon montanus Casar. ex Koehne, Fl. Bras. (Martius) 13(2): 3271877, pro syn. Diplusodon serpyllifolius Mart. ex DC., Prodr. 3: 94c. 1828. Friedlandia serpyllifolia DC. ex Koehne, Fl. Bras. (Martius) 13(2): 334. 1877, nom. inval. Type: Brazil. Minas Gerais: Pr. Arraial de São Gonçalo, C. F. P.Martius obs. 706 (holotype M [0146260]). Diplusodon smithii Lourteig, Sellowia 16: 148, fig. 8, 1964. Type: Brazil. Minas Gerais: Jaboticatubas, Serra do Cipó, Chapéu do Sol, 1000 m, 29 Apr 1952 (fl),

Descriptions

233

L. B. Smith, W. A. Egler, S. L. Oliveira-Silva, L. Dau, Z. Lopes-Silva, W.T. Ormond, G. C. Machine 7022 (holotype, US [00169559]; isotypes, NY, P [2], R [2], RB). Friedlandia verbenifolia Mart. ex DC., Prodr. 3: 94c, 1828, pro syn. Shrubs 0.5–3  m tall, erect, much-branched, usually dark-green to dark-brown when dry. Upper branches subquadrangular to quadrangular, unwinged, generally canescent, sometimes rust-canescent, densely pilose to hirtellous, internodes 0.6– 1.5 cm long. Leaves decussate, not imbricate, spreading to suberect, petiolate; petioles 1–6  mm long; blades 8–35  ×  5–25  mm, chartaceous, not discolorous, eucamptodromous to acrodromous-basal-supranumerary, elliptic or ovate to orbicular, base acute to obtuse, apex obtuse to acute or sometimes acuminate, margin slightly revolute to subrevolute, glabrescent, adaxial surface pilose, verrucose, abaxial surface densely hirsute, canescent principally on the veins; secondary veins 3–4 on each side, departing along the midvein or departing from the base of the midvein; domatia present. Synflorescence frondose, pleiobotryum type, congested; accessory branches absent, principal florescence of the botryum spiciform type, paraclades 5–20 cm long; bracts similar to the vegetative leaves, but smaller; flowers sessile to subsessile, hypopodium 0–1  mm long, epipodium 0–0.5  mm long; prophylls 2.5–4 mm long, reaching the middle or apex of the floral tube, obovate, oblong to elliptic, keeled, base acute, apex obtuse, rarely acute, margin plane, densely covered with white trichomes; floral tube 2–3  mm long, funnel-shaped, densely covered with white trichomes to less pilose; sepals 2–3 mm long, triangular, not caudate, not deflexed, densely covered with white trichomes to less pilose; epicalyx segments 0.5–3 mm long, lanceolate, spreading to slightly deflexed, shorter than the sepals, with the same indumentum as the sepals; corolla 3–3.5 cm in diam., clear rose, lilac or purple to reddish, petals 12–15 × 7–8 mm, obovate, apex obtuse; stamens 12–15, free portion of the filaments 7–10  mm long, exserted; ovary 1.7–2 × 2–2.5 mm, rounded, glabrous, style 14–18 mm long, stigma green, ovules 13–28. Capsule 4.5–5 × 4.4.5 mm, globose, glabrous; seeds 7–25, 2–3 × 1.8–2 mm. 2n = 30 (Graham and Cavalcanti 2001). Distribution and Ecology (Fig. 13.51)  Diplusodon hirsutus is widely distributed in Cadeia do Espinhaço, in Minas Gerais State, occurring in campos rupestres with rocky outcrops, cerrado, and carrasco at elevations of 900–1400 m. Flowers from September to July, fruits from October to July. Specimens Examined: BRAZIL. Minas Gerais,  Belo Horizonte, 12 Apr 1919 (fl, fr), Gehrt 165 (CEN, SP); Belo Horizonte, estrada Belo Horizonte-Vespaziano, km 25, 12 May 1990 (fl, fr), Grandi s.n. (BHCB 18110, CEN 19048); Brumadinho, 20°06′ S, 43°59′ W, 27 May 1998 (fl, fr), Silveira 26 (BHCB, CEN); Conceição do Mato Dentro, 28 Jun 1988 (fr), Stehmann II-34 (BHCB); Cristalia, morro da antena de televisão, 16°43′28″ S, 42°55′42″ W, 12 Jul 2001 (fl), Souza et al. 25791 (ESA); Nova Lima, Retiro das Pedras, s.d., Badini s.n. (OUPR 2742); Nova Lima, 7 May 1972 (fl), Badini s.n. (OUPR 19653); Nova Lima, 7 May 1972 (fl), Badini s.n. (CEN, OUPR 5419); Nova Lima, Serra da Moeda, s.d., Badini s.n. (OUPR 19649); Nova Lima, 2 May 1968, Duarte 10853 (P); Nova Lima, 12 May 1983 (fl), Zurlo &

234

13  Systematic Treatment

Fig. 13.49  Diplusodon hirsutus. A. Branch with inflorescence. B–C. Leaf, abaxial surface. D– E. Prophylls, adaxial surface. F–G. Floral bud. H. Flower with prophylls, without petals. I. Seed. (A Joly et al. (CFSC) 52, SPF; B Smith et al. 7022 NY; C, E, F Martius 869, M; D, G–I Joly et al. (CFSC) 2307, SPF)

Descriptions

235

Silva s.n. (OUPR 26250); Nova Lima, s.d., Kassis et al. s.n. (CEN, OUPR 5357); Botumirim, Serra da Canastra, Tinoco, 14 Jun 1991 (fl, fr), Mello-Silva et al. 502 (SPF); Cachoeira do Campo, 7 Apr 1972 (fl), Badini s.n. (OUPR 19652); 50 km N de Congonhas, ao longo da BR-135 para Belo Horizonte, 28 Feb 1976 (fl), Davidse & D’Arcy 10797 (SP); Congonhas, área da Itaminas, 16 Jun 1990 (fl, fr), Silva et al. 1649 (CEN, VIC); Entre Rios de Minas, Pedra Branca, 29 Jan 1977 (fl), Krieger 14568 (CESJ); Entre Rios de Minas, Fazenda da Pedra Branca, 29 Jan 1977 (fl, fr), Coons et  al. 77–348 (CEN, VIC); Cristália, alto do Pico do Chapéu, onde fica a antena, 16°43′38″ S, 42°55′35″ W, 29 Apr 1997 (fl), Cavalcanti et al. 2322 (CEN, MO, NY); Alto do Pico do Chapéu, aonde fica a antena de televisão, 16°43′38″ S, 42°55′35″ W, Cavalcanti et al. 2323 (CEN, MO); Morro do Chapéu, 14 Jun 1991 (fl, fr), Mello-Silva et al. 487 (CEN, SPF); Grão-Mogol, campo rupestre em direção nordeste da cidade, 22 May 1982 (fl, fr), Mamede et al. (CFCR) 3465 (CEN, SPF); Grão-Mogol, vale do rio das Mortes, 24 Jul 1986 (fl), Zappi et al. (CFCR) 9931 (CEN, SPF); Grão-Mogol, vale do Riacho Ribeirão, 16°32′ S, 42°55′ W, 21 May 1987 (fl, fr), Mello-Silva & Pirani (CFCR) 10761 (CEN, SPF); Grão-Mogol, vale do Ribeirão dos Bois, 21 May 1987 (fl), Mello-Silva & Pirani (CFCR) 10763 (K, SP, SPF); Grão-Mogol, subindo a trilha da tropa, à esquerda Riacho Ribeirão, 27 May 1988 (fr), Prado et al. (CFCR) 12013 (CEN, SPF); Itabira, Pico do Itabira do Campo, 5 (G [2], P); Itabira, Pico do Itabira do Campo, s.d., Badini s.n. (OUPR 19651); Itabira, Serra de Itabira do Campo, Apr 1892 (fl), Ule 2562 (R, US); Ouro Preto, Serra do Ouro Preto, Apr 1839, Claussen 114 (Martius Herb. Fl. Bras. n. 867) (G [2], M, P, W); Serra de Ouro Preto, 7 May 1972 (fl, fr), Badini s.n. (OUPR 19654); Ouro Preto, estrada de Ouro Preto, próximo da BR-3, 24 May 1959 (fl), Occhioni 1827 (HB); Itabirito, Serra do Itabirito, 7 May 1972 (fl, fr), Badini s.n. (OUPR 19650); Jandarela, 13 Jul 1972 (fr), Emygdio et al. 3251 (NY); Morro do Gavião, 2 Nov 1985 (fl), Rossi et al. (CFCR) 5766 (K, SPF); Piedade do Paraopeba, 25 Apr 1968, Duarte 10819 (P); Piedade do Paraopeba, Serra da Piedade do Paraopeba, 28 Apr 1970, Duarte 12412 (P); rio Acima, beira da estrada na área de mineração, 28 Jun 1988 (fl), Mendonça-Filho 8 (BHCB); Santana do Riacho, 15 Apr 1985 (fl), Lopes & Andrade s.n. (BHCB 9109, CEN 19042); Santana do Riacho, Serra do Cipó, atalho entre o km 115 e a estrada da Usina, 2 May 1993 (fl), Souza & Sakuragui 3404 (ESA); Santana do Riacho, estrada da Conceição do Cerro, km 130 a 180, 28 Jun 1972 (fl), Occhioni 5151 (US); Santana do Riacho, 19°17′ S, 43°35′ W, 6 Jun 1998 (fl), Jacobi 52 (BHCB, CEN); Santana do Riacho, Serra do Cipó, Jul 1949 (fl, fr), Vidal s.n. (R); Santana do Riacho, Serra do Cipó, 26 Nov 1991 (fr), Pereira et al. 787 (BHCB); Santana do Riacho, Serra do Cipó, 27 Jun 1991 (fl, fr), Pereira et al. 933 (BHCB); Santana do Riacho, Serra do Cipó, ca. 2 km do Chapéu do Sol, 3 Jul 1996 (fl, fr), Souza et al. 11599 (CEN, ESA); Santana do Riacho, Serra do Cipó, rodovia Santana do Riacho-Conceição do Mato Dentro, Véu da Noiva, 2 May 1986 (fl), Cavalcanti et al. (CFSC) 9683 (SPF); Santana do Riacho, Serra do Cipó, rodovia Santana do Riacho-­Conceição do Mato Dentro, km 100, 19°18′87″ S, 43°36′43″ W, 23 Apr 1997 (fl), Cavalcanti et  al. 2270 (CEN, MO, NY); Santana do Riacho, Serra do Cipó, rodovia Santana do Riacho-Conceição do Mato Dentro, km 104, 8 May 1987 (fl), Cavalcanti et  al. (CFSC) 10099 (SPF);

236

13  Systematic Treatment

Santana do Riacho, Serra do Cipó, rodovia Santana do Riacho-Conceição do Mato Dentro, km 105, ístimo que leva ao Morro do Calcáreo, 8 May 1987 (fl), Cavalcanti et al. (CFSC) 10110 (SPF); Santana do Riacho, Serra do Cipó, rodovia Santana do Riacho-Conceição do Mato Dentro, km 106, 20 Apr 1985 (fl), Amaral & Kawasaki (CFSC) 9321 (SPF); Santana do Riacho, Serra do Cipó, estrada Lagoa SantaConceição do Mato Dentro, km 106, 16 May 1995 (fl), Castro 488 (CEN, HUFU n.v.); Santana do Riacho, Serra do Cipó, rodovia Santana do Riacho-Conceição do Mato Dentro, km 107, 1 Sep 1986 (fl, fr), Zappi & Kameyama (CFSC) 9853 (SPF); Santana do Riacho, Serra do Cipó, rodovia Santana do Riacho-Conceição do Mato Dentro, km 109, 27 May 1977 (fl), Menezes (CFSC) 7311 (SPF); Santana do Riacho, Serra do Cipó, rodovia Santana do Riacho-Conceição do Mato Dentro, 6 Nov 1980 (fl, fr), Pirani (CFSC) 6166 (SP); Santana do Riacho, Serra do Cipó, rodovia Santana do Riacho-Conceição do Mato Dentro, km 110, 8 Jun 1989 (fl, fr), Barbosa 403 (BHCB, UFU); Santana do Riacho, Serra do Cipó, rodovia Santana do RiachoConceição do Mato Dentro, km 112, 16 Apr 1972 (fl), Joly et  al. (CFSC) 1653 (CEN, SP); Santana do Riacho, Serra do Cipó, rodovia Santana do RiachoConceição do Mato Dentro, km 113, 14 Apr 1935 (fl), Mello-Barreto & Brade 1270 (R); Santana do Riacho, Serra do Cipó, rodovia Santana do Riacho-Conceição do Mato Dentro, km 114, 5 Jun 1970 (fl, fr), Joly et al. (CFSC) 52 (CEN, SP); Santana do Riacho, Serra do Cipó, rodovia Santana do Riacho-Conceição do Mato Dentro, km 115 (antigo 108) da rodovia MG-010, afloramento rochoso, lado direito no sentido Santana do Riacho-Morro do Pilar, 9 Jun 2002 (fl, fr), Groppo Jr. et al. 1091 (CEN, SPF); Santana do Riacho, Serra do Cipó, rodovia Santana do RiachoConceição do Mato Dentro, km 116, Duarte 8119 (HB, RB); Santana do Riacho, Serra do Cipó, rodovia Santana do Riacho-Conceição do Mato Dentro, km 118, 18 Jun 1995 (fl, fr), Castro 491 (CEN, HUFU); Santana do Riacho, Serra do Cipó, rodovia Santana do Riacho-Conceição do Mato Dentro, km 124, 29 May 1972 (fl), Joly et al. (CFSC) 2042 (SP); Santana do Riacho, Serra do Cipó, rodovia Santana do Riacho-Conceição do Mato Dentro, km 126, 30 Apr 1973 (fl), Semir et al. (CFSC) 4159 (SP); Santana do Riacho, Serra do Cipó, rodovia Santana do Riacho-Conceição do Mato Dentro, km 130, 17 Apr 1950 (fl), Duarte 2504 (MO, NY, RB); Santana do Riacho, Serra do Cipó, rodovia Santana do Riacho-Conceição do Mato Dentro, km 132, 5 Apr 1951 (fl), Mendes-Magalhães 5976 (HB); Santana do Riacho, Estrada da Usina, 16 Apr 1972 (fl), Joly et  al. (CFSC) 1766 (CEN, SP, UEC); Santana do Riacho, Estrada da Usina, 28 May 1972 (fl), Joly et al. (CFSC) 2307 (CEN, SP); Santana do Riacho, Estrada da Usina, Cavalcanti et al (CFSC) 9691 (SPF); Santana do Riacho, estrada para a usina Dr. Pacífico Mascarenhas, 3.4 km da estrada Lagoa Santa-Conceição do Mato Dentro (MG-010), 19°15′45″ S, 43°36′16″ W, 24 Apr 2006 (fl), Mello-Silva et al. 2927 (CEN, SPF); Santana do Riacho, Serra do Cipó, 6 km ao Norte do Palácio, 19°10′ S, 43°35′ W, 29 Apr 1952 (fl), Smith et al. 6880 (R, US); Santana do Riacho, Serra do Cipó, 5 Apr 1951 (fl), Black & Magalhães 51-12077 (IAN); Santana do Riacho, base da Serra do Cipó, 16 Mar 1962 (fl), Duarte 6506 (HB, P n.v.); Santana do Riacho, Serra do Cipó, 21 May 1982 (fl), Bautista 611 (HRB, SPF); Santana do Riacho, Serra do Cipó, Lapinha, 3.2 km de Lapinha em direção a Santana do Riacho, 19°06′45″ S, 43°41′52″ W, 23 Apr 2006

Descriptions

237

(fl), Mello-Silva et al. 2914 (CEN, SPF); São João Del Rei, estrada de Morro do Ferro para Passa Quatro, 8 km de Morro do Ferro, sentido Passa Quatro, Serra das Vertentes, 21°13′ S, 44°54′ W, 26 Feb 2000 (fl, fr), Cavalcanti et al. 2562 (CEN, MO); São Tomé das Letras, 5 Feb 1973 (fl), Hatschbach & Ahumada 31275 (MBM, NY, W); São Tomé das Letras, Serra das Ninfas, fazenda dos Correa, 21°45′45″ S, 45°01′37″ W, 19 Feb 2005 (fr), Mello-Silva & Ferreira 2865 (CEN, SPF); Serra de Gandarela, 25 Jun 1884 (fl), Glaziou 14699 (G, K, P, R); Serra do Belo Vale, 6 May 1969, Duarte 11526 (P); Serra do Itatiaia, s.d., Badini s.n. (OUPR 20013); Monte Serra da Caraca, Casaretto Herb. Nº. 2989 (G, TO n.v.); without reference to municipal district: Claussen 22a (BR); Claussen 23a (BR); Claussen 115 (G); Claussen 309 (P); Claussen s.n. (G, K, OXF, P, R); Glaziou 21422b (R); Martius s.n. (fragm. G); Schüch s.n. (W); Feb 1892 (fl), Sellow s.n. (P, W). Diplusodon hirsutus belongs to the Hirsutus complex (discussed under D. bahiensis). Diplusodon hirsutus has variable indumentum, and leaf shapes vary from elliptic, with a more revolute margin and acute base, to ovate or orbicular, with the margin slightly revolute to almost plane and the base acute to obtuse. In all plants, whether glabrescent or pubescent, the leaves are hirsute and verrucose to pilose on the adaxial surface. When pilose, the species displays white, abundant indumentum covering all vegetative and floral parts. The plants of D. hirsutus turn dark-brown when dry. In the flowers, the epicalyx segments are short, varying from 0.5 to 3 mm long, and are spreading to slightly deflexed. Stamen number also varies, from 12 to 15. In São Tomé das Letras, Minas Gerais, the flowers have 15 stamens (Hatschbach & Ahumada 31275), while the specimens from other localities in Minas Gerais have 12 stamens. Lourteig (1964) described Diplusodon smithii based on specimens from Serra do Cipó and emphasized its similar habit with D. subsericeus and D. hirsutus. She separated D. smithii from D. subsericeus based on the indumentum and D. smithii from D. hirsutus based on the obconic floral tube, subulate epicalyx segments, and size of the prophylls. In her key, Lourteig (1989) later separated D. hirsutus and D. smithii based on simple trichomes in the former species and enlarged trichome bases in the latter. I studied specimens from Serra do Cipó, as well as type materials of D. hirsutus, D. smithii, and D. subsericeus, in relation to the characteristics cited by Lourteig (1964). Apart from the orbicular leaves with plane margins of D. smithii, no other characteristics, especially of the flowers, are useful to separate these taxa as different species. In all specimens examined, the flowers are densely pilose, the buds are white due to the dense indumentum, the epicalyx segments are short and spreading to slightly deflexed, and the number of stamens is the same. Diplusodon smithii is therefore considered as synonyms of D. hirsutus. Diplusodon subsericeus differs from D. hirsutus due to the former having leaves with sericeous indumentum with silvery aspect, not dark-brown when dried, and flowers with prophylls of 5–6.5 mm long, reaching the middle to surpassing the apex of the floral and longer floral tube (6–6.5 mm long).

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In the southern part of its distribution, Diplusodon hirsutus varies in vegetative characteristics. In the region from Ouro Preto to Belo Horizonte, state of Minas Gerais, where D. microphyllus occurs, there are individuals with features intermediate between the two species but with smaller, less coriaceous leaves and with less dense indumentum with sparser trichomes, as in D. microphyllus. Still other individuals in this region appear to have a gradation of the characteristics of the two species with a third species of the region, D. buxifolius, which has ovate-lanceolate, almost glabrous leaves (Williams & Assis 6745). Some specimens are difficult to determine as D. hirsutus or D. microphyllus because there is a morphological gradient from one species to the other. In the phylogenetic studies using molecular and morphological data (Cavalcanti et al. 2007), the two appear to be distinct species. From Serra do Cipó to the northern part of Minas Gerais, specimens of Diplusodon hirsutus have more orbicular, plane leaves with less dense indumentum. This pattern culminates in the populations of Grão-Mogol where the plants have elliptic to orbicular leaves with up to five veins. The material cited in Linnaea in the protologue of Friedlandia hirsuta Cham. & Schltdl. (1837) was Sellow s.n. However, the specimen annotated as F. hirsuta by Chamisso & Schlechtendal is Sellow 2053 (B, destroyed; F neg. nº 017923). Lourteig (1989) lectotypified Friedlandia hirsuta, choosing as lectotype material a specimen at P, without collector, where it appears only with Lourteig’s annotation. A duplicate of Sellow s.n. annotated by Chamisso and Schlechtendal as “Friedlandia hirsuta Nob.” is deposited at HAL.  A second-step lectotype is here designated (HAL0098374). 40. Diplusodon imbricatus Pohl, Flora 10: 150. 1827. Type: Brazil. Tocantins: Serra de São Félix, in aquosis humidis, montosis circa Rio Trahiras in mont. Serro d. Felix, Aug 1819 (fl, fr), J. B. E. Pohl 1988 = D 941 (first-step lectotype, designated by Lourteig 1989: 234, second-step lectotype, W [0053859], here designated; isolectotypes, G, K [3], W [0053858]). Figs. 13.50 A–H, 13.51 Subshrubs 0.5–2 m tall, erect, branched, stem brownish to reddish, glabrous, not dark-brown when dry. Upper branches cylindrical to subquadrangular, unwinged, internodes 3–5 cm long. Leaves not imbricate, decussate, spreading to suberect, sessile to amplexicaulous; blades 25–65 × 12–45 mm, slightly glaucous, chartaceous, not discolorous, acrodromous-basal-supranumerary, with inconspicuous reticulation, lanceolate to ovate-lanceolate, base cordate or subcordate to amplexicaulous, apex acute, margin plane, marginate-yellow; secondary veins 6–10 on each side, 3–6 departing from the base and 3–4 departing along the midvein, inconspicuous; domatia absent. Synflorescence frondose-bracteose, diplobotryum type, congested; accessory branches absent, principal florescence 15–20  cm long, botryum type, paraclades 10–20 mm long; bracts 10–20 × 5–10 mm, lanceolate to ovate-lanceolate, margin plane, marginate-yellow; flowers pedicellate, hypopodium 2–4  mm long, persistent, epipodium 0–0.5 mm long; prophylls 3.5–4 × 2.8–3 mm, reaching the middle of the floral tube, rhombic to broadly elliptic, unkeeled, base acute to

Descriptions

239

obtuse, apex obtuse, margin plane; floral tube 4–5.5  mm long, oblong; sepals 2.5–3 mm long, triangular, not caudate, not deflexed; epicalyx segments absent or 0.5–1 mm long, spreading, shorter than the sepals; corolla 3–4 cm diam., lilac to rose, petals 12–15 × 6.6–8 mm, obovate, apex obtuse; stamens 12–18, free portion of the filaments 7.5–8.5 mm long, exserted, surpassing the sepals ca. 1 mm; ovary 2–2.2  ×  1.7–3.5  mm, rhombic, glabrous, style 14–16  mm long, ovules 16–20. Capsule 5–6 × 5–6.5 mm, chestnut brown when mature, globose, glabrous; seeds 15–17, 2–3 × 2–3.8 mm. Distribution and Ecology (Fig. 13.51)  Prior to this study, Diplusodon imbricatus was known only from the type material, but it has since been collected during the last decade in northern Goiás and southern Tocantins, in the depression of the Tocantins River, in cerrado and campo sujo at 300–919  m, and in Chapada dos Veadeiros, Goiás. Flowers from February to August; fruits from March to August and November. Specimens Examined: BRAZIL.  Goiás,  Cavalcante, entroncamento vila Veneno-balsa rio São Félix, km 3.6, 13°40′53″ S, 47°51′46″ W, 21 Feb 2002 (fl), Pereira-Silva et  al. 5950 (CEN); Cavalcante, estrada para Prata (Kalungas), 13°22′24.2″ S, 47°41′9.1″ W, 14 Apr 2004 (fl, fr), Fonseca et al. 5104 (CEN, IBGE); Cavalcante, ponte sobre o rio Tocantins-Cavalcante, km 84, 13°40′42.4″ S, 47°50′36″ W, 30 Aug 2004 (fr), Cavalcanti et  al. 3524 (CEN, SPF); Cavalcante, Reserva Natural Serra do Tombador (RPPN), 13°40′38.44–44.2″ S, 47°49′47.0– 48.6″ W, 20 Feb 2013 (fl), Lombardi et  al. 9912 (CEN, HRCB). Tocantins, Palmeirópolis, alto do morro da torre de rádio da Enerpeixe, 12°47′28″ S, 48°24′09″ W, 16 Jun 2006 (fr), Pereira-Silva et al. 10762 (CEN); Palmeirópolis, 12°48′16″ S, 48°16′45″ W, 10 Nov 2003 (fr), Bucci & Verano 1559 (UB); Paranã, Fazenda São João, 12°53′23″ S, 47°36′01″ W, 25 Mar 2004 (fl, fr), Sevilha et al. 3774 (CEN); Paranã, Morro do Cruzeiro, a leste do canteiro de obras do AHE São Salvador, 12°46′32″ S, 48°13′59″ W, 4 Jul 2002 (fr), Pereira-Silva et al. 6552 (CEN, MO); Paranã, estrada Palmeirópolis-Alvorada de Goiás, a 25  km de Palmeirópolis, 12°43′87″ S, 47°28′02″ W, 6 Apr 1997 (fl), Cavalcanti et al. 2225 (CEN, MO, NY); Paranã, estrada Palmeirópolis-Alvorada de Goiás, a 35  km de Palmeirópolis, 12°54′46″ S, 47°33′42″ W, 6 Apr 1997 (fl), Cavalcanti et al. 2228 (CEN, MO, NY); Paranã, canteiro de obras do UHE São Salvador, alto do Morro da Cruz, 12°48′ S, 48°13′39″ W, 9 Jun 2006 (fl, fr), Pereira-Silva et al. 10519 (CEN); Paranã, estrada São Salvador-balsa do rio Custódio, km 51, 13°0′07″ S, 48°06′39″ W, 10 Jun 2006 (fl, fr), Pereira-Silva et  al. 10552 (CEN); Paranã, estrada de acesso ao vilarejo Rozario, ca. 3 km após a entrada principal da obra, 12°47′42″ S, 48°11′58″ W, 24 Mar 2007 (fr), Pereira-Silva & Moreira 11466 (CEN); Paranã, Vila Rozario, 13°57′39″ S, 48°07′48″ W, 24 Jan 2008 (fl, fr), Pereira-Silva & Moreira 12882 (CEN); São Salvador do Tocantins, 3.4 saindo da barragem “A”, Fazenda Serrinha, área de influência da Usina Hidrelétrica Cana Brava, influência indireta, 12°47′25″ S, 48°16′13″ W, 21 Feb 2002 (fl), Pereira-Silva et al. 5936 (CEN); São Salvador do Tocantins, entroncamento fazenda Serrinha-rio Mutum (mataburro), 12°47′18″ S, 48°16′31″ W, 14 Jun 2006 (fl, fr), Pereira-Silva et al. 10648 (CEN); São Salvador

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Fig. 13.50  Diplusodon imbricatus. A. Vegetative branch. B. Leaf, abaxial surface. C. Section of internode. D. Branch with inflorescence. E. Flower with prophylls, without petals. F. Fruit. G. Capsule. H. Seed. (A, E Pereira-Silva et  al. 10648, CEN; B–D, F–H Pereira-Silva et  al. 10762, CEN)

Descriptions

241

Fig. 13.51  Geographic distribution of Diplusodon heringeri, D. hexander, D. hirsutus, and D. imbricatus

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do Tocantins, estrada de acesso ao córrego Mutum, 12°48′10″ S, 48°16′40″ W, 25 Mar 2007 (fl), Pereira-Silva & Moreira 11543 (CEN); São Salvador do Tocantins, Margem esquerda do lago UHE São Salvador, 12°49′13″ S, 48°15′53″ W, 24 Mar 2009 (fl), Pereira-Silva et al. 14133 (CEN); without reference to municipal district: Pohl s.n. (G). The plants of Diplusodon imbricatus are subshrubs with brownish to reddish branches and recognized by its lanceolate to ovate-lanceolate sessile leaves with a cordate to amplexicaulous base and acrodromous-­basal-­supranumerary venation. Besides these characteristics, the plant is totally glabrous, the leaves are glaucous with slightly yellow margins (purple when dried), and the leaves are much larger than the bracts. Epicalyx segments are absent or diminutive and cylindrical (0.5–1 mm long). Diplusodon imbricatus is similar to D. trigintus, which differs from the former mainly by its leaves lacking yellow margins and with prominent veins, smaller prophylls (1.5–3 × 1–2.5 mm) that are inserted close to the apex of the pedicel and mostly not reaching the base of the floral tube, flowers with epicalyx segments 0.5–1.5 mm long, and 24–32 stamens. Lourteig (1989) cited Pohl 1988  =  D941 (W) as the holotype of Diplusodon imbricatus, even though D. imbricatus was described by Pohl (1827a) without citation of specimens in the protologue. Pohl (1827b), in another publication, referred to a single collection of D. imbricatus collected in August in Serra de São Félix, near to Rio Traíras, Goiás, without mentioning a collection number. At W, there are two specimens of D. imbricatus from the type locality, one of which has the original label annotated by Pohl (Pohl 1988 = D941) and was chosen here as lectotype. In addition, other specimens of D. imbricatus were examined at G and K, with labels that indicate Pohl s.n. Since duplicates of Pohl’s collections were sent to other herbaria without the information from his field diary (Wahlnöffer, pers. comm), and most lack a number, the material at G and K are regarded as duplicates of Pohl 1988 = D941. 41. Diplusodon incanus Gardner, Sert. Pl., fig. 2. 1843. Type: Brazil. Goiás: Campos between São Domingos & Posse, May 1840 (fl), G. Gardner 4140 (lectotype, OXF [0005271], designated by Lourteig 1989: 223; isolectotypes, BR, F, fragm. F, G [4], K, NY, P [3], S, W [2]). Figs. 13.52 A–P, 13.55 Diplusodon incanus Gardner var. hirtellus Koehne, Fl. Bras. (Martius) 13(2): 336, pl. 63, fig. 1b. 1877. Type: Brazil. Goiás: G.  Gardner 4137 (lectotype, P [01901891], designated by Lourteig 1989: 224; isolectotypes, BR, F [2], G [3], K [3], NY, P [01901892, 01901893], OXF, US, W [2]). Diplusodon incanus Gardner var. serpyllum Koehne, Fl. Bras. (Martius) 13(2): 336, pl. 63, fig. 1c. 1877. Diplusodon virgatus Gardner ex Koehne, Fl. Bras. (Martius) 13(2): 336. 1877, pro syn. Type: Brazil. Goiás: 1841 (fl, fr), G. Gardner 4141

Descriptions

243

(lectotype, P [01901889] designated by Lourteig 1989: 223; isolectotypes, BR, fragm. F, G [2], K [3], NY [2]; OXF, P [01901888, 01901890], S, US, W [2]). Shrubs 1.2–2.5 m tall, erect, much-branched, usually slightly canescent, heterogeneous in leaf size, not dark-brown when dry, glabrescent. Upper branches cylindrical, unwinged, densely pilose or villous, internodes 0.4–0.8  cm long. Leaves decussate, not imbricate, spreading to suberect, petiolate; petioles 0.5–2 mm long; blades 4–10 × 4–9(−13) mm, coriaceous, not discolorous, eucamptodromous, elliptic to ovate-elliptic, base acute to attenuate, apex obtuse, margin subrevolute, glabrescent, sometimes canescent, with fine trichomes, sometimes slightly sericeous or almost glabrous; secondary veins 2–3 on each side of the midvein, adaxial surface with sunken, inconspicuous veins, densely covered by abundant, white trichomes on the blade and veins, slightly prominent on the abaxial surface; domatia present. Synflorescence frondose-bracteose to frondose, diplobotryum to pleiobotryum type, congested; accessory branches absent, principal florescence botryum type, 1.5–5 cm, often truncate, paraclades 2–19 cm long; bracts 5–15 × 2–7 mm, obovate to elliptic or the same shape and size as the leaves; flowers pedicellate, hypopodium 0.3–1 mm long, epipodium 0.3–0.5 mm long; prophylls 2.2–3 × 1–1.5 mm, reaching the middle of the floral tube or slightly beyond the middle of the floral tube, oblong, unkeeled, base acute, apex acute to obtuse, margin plane to involute, the trichomes becoming very short and sparse; floral tube 2–3.5  mm long, oblong or funnelshaped, covered by dense indumentum or the trichomes becoming very short and sparse or sparsely sericeous; sepals 1.8–3  mm long, triangular, not caudate, not deflexed, pilose, ciliate or not; epicalyx segments 1.5–3(−3.5) mm long, cylindrical, strongly deflexed, the same indumentum as that of the sepals, ciliate or with very short trichomes, longer than the sepals; corolla 1.8–2.5 cm in diam., rose to lilac, petals 6–9 × 4–8.5 mm, obovate, ovate to elliptic, apex obtuse; stamens 12, free portion of the filaments 6–7  mm long, exserted; ovary 1.7–1.9  mm  ×  1.8–2  mm, rounded, glabrous, style 9–12.5  mm long, ovules 8–17. Capsules 3.5–4  ×  3.5– 3.9 mm, globose, glabrous; seeds 6–15, 2–2.9 × 1.1–2 mm. Distribution and Ecology (Fig. 13.55)  Diplusodon incanus occurs in the northeastern portion of the state of Goiás and southwestern portion of the state of Bahia, Brazil. It is found in campos rupestres and campos sujos on sandy soils and in sandstone and clay at 950–1400 m, on Serra Geral do Paranã. It was collected also in Serra Geral de Goiás and Espigão Mestre, Bahia, in cerrado with a tall, slightly dense, tree stratum at 822–870 m, among rocky outcrops of the campos cerrados and along the margins of gallery forests. Flowers from March to August; fruits registered from July to August. Specimens Examined: BRAZIL.  Bahia,  Correntina, rodovia entre Posse e Correntina, ca. 25 km de Posse, 15 Aug 2006 (fl), Pastore & Suganuma 1549 (CEN, HUEFS); Correntina, 27 km do entroncamento com a BR-020 e BR-349, 13°43′ S, 45°55′17″ W, 1 Apr 1997 (fl), Harley et  al. 28587 (CEN, HUEFS); Correntina, Fazenda Jatobá, 6 Jun 1992 (fl), Silva et al. 1319 (IBGE, RB n.v., UB); Correntina, Silva et al. 1325 (IBGE); Correntina, 27 Jun 1992 (fl), Silva et al. 1435 (IBGE);

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Fig. 13.52  Diplusodon incanus. A. Branch with inflorescence. B–D. Section of internodes. E, G, I. Leaves, abaxial surface. F, H, J. Indumentum of abaxial surface of leaves. K. Simple trichome of leaves. L–M. Floral buds. N–O. Flowers with prophylls, without petals. P. Immature fruit. (A, H, L Valls et al. 13070, CEN; B, D–E, J Gardner 4140, K; C, F–G, O Gardner 4141, H; I Rizzo & Heleno 10283, UFG)

Descriptions

245

Correntina, Fazenda Jatobá, 8 Aug 1992 (fl), França et al. 763 (IBGE, P n.v., UEC n.v.); Correntina, Fazenda Jatobá-Reserva Biológica Fazenda Jatobá, 14°02′09″ S, 45°57′46.9″ W, 29 Jul 2004 (fl, fr), Fonseca et al. 5621 (CEN, IBGE). Goiás, Posse, Serra Geral do Paranã, 3  km de Posse pela BR-220, 20 May 1983 (fl), Rizzo & Heleno 10283 (UFG); Posse, 15 km N de Posse, rodovia BR-020, 12 Jul 1979 (fl), Hatschbach & Guimarães 42273 (MBM, P); Guarani de Goiás, 8.3 km ao norte do acesso à Posse, ao longo da Rodovia Brasília-Barreiras, BR-020, 15 May 1991 (fl), Valls et  al. 13070 (CEN, SPF); São Domingos, Contagem, 15  km  L de São Domingos, 15 May 2000, (fl), Hatschbach et  al. 71131 (MBM); São Domingos, rodovia GO-108, 3 km S de São Domingos, 16 May 2000 (fl), Hatschbach et al. 71184 (MBM); São Domingos, Parque Estadual de Terra Ronca, 13°22′22″ S, 46°10′26.9″ W, 26 Apr 2013 (fl), Amaral, A.G. et  al. 2978 (CEN, HEPH, UB); Teresina de Goiás, Fazenda Hotel Ecológico Alpes Goianos, rodovia GO-118, km 202, 13°53′59.1″ S, 47°23′48.9″ W, 31 Jul 2000 (fr), Souza et al. 24735b (ESA). Diplusodon incanus is represented by tall, much-branched, canescent to glabrescent shrubs and presents a wide variation in indumentum density: the stems can be puberulous, hirtellous, or slightly sericeous, and the leaves can be glabrous on the abaxial surface and hirtellous to sericeous on the abaxial surface, when sericeous, sometimes appearing canescent to silvery. The leaves are elliptic or narrowly elliptic to ovate with subrevolute margins, and the flowers are hirsute at anthesis, becoming minutely pilose to nearly glabrous with age, with epicalyx segments that are narrowly cylindrical and deflexed, and the androecium has 12–18 stamens. Gardner (1843) called attention to the leaf size variation found on the branches of a single individual as a diagnostic feature of Diplusodon incanus, which is also observed in this study. In the vegetative aspects, Diplusodon incanus is similar to D. heringeri due to characteristics such as the much-branched shrubby habit and similarly shaped and sized leaves of similar form and size with revolute to subrevolute margins and the presence of domatia. Diplusodon heringeri differs from D. incanus in that the former has sessile flowers, with a longer floral tube (4–4.5 mm long), erect epicalyx segments, and an androecium with 17–18 stamens. Koehne (1877) described Diplusodon incanus var. hirtellus and D. incanus var. serpyllum, both collected in Goiás. Lourteig (1989) placed D. incanus var. serpyllum in synonymy under D. incanus var. incanus and recognized D. incanus var. hirtellus. The type material of D. incanus var. serpyllum is almost glabrous and does not have the same canescent aspect of the type material of D. incanus; however, the habit, leaf shape, and flowers are undoubtedly those of D. incanus, so I agree with the synonymy of Lourteig (1989). The type material of Diplusodon incanus var. hirtellus has smaller leaves that are slightly sericeous and with a canescent or slightly silvery aspect, which is not different from some variants of D. incanus; therefore, D. incanus var. hirtellus is here synonymized. There are 17 duplicates of the type material of Diplusodon incanus (Gardner 4140) in European and American herbaria, and most have a short label that lacks

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complete data such as the exact locality. Lourteig (1989) cited a specimen from OXF and cited as the holotype, which is considered here as a first-step lectotype (ICBN Art. 9.17, ex. 14). There are 17 duplicates of the type material of Diplusodon incanus var. hirtellus (Gardner 4137) in European and American herbaria. Lourteig (1989) cited a specimen, Gardner 4137a, from P as the holotype, which is considered the lectotypification (ICBN Art. 9.17, ex. 14). Although there is no letter “a” along with the Gardner number in any of the three specimens at P, the specimen chosen by Lourteig (indicated because she wrote “type” on the specimen) has a more complete label and is considered as the lectotype [P01901891]. There are 20 duplicates of the type material of Diplusodon incanus Gardner var. serpyllum (Gardner 4141) in European and American herbaria. Lourteig (1989) cited a specimen from P as the holotype. There are three specimens of Gardner 4141 at P, and Lourteig’s (1989) choice stands as the designation of a lectotype (ICBN Art. 9.17, ex. 14). 42. Diplusodon kielmeyeroides Subshrubs 0.4–1.2 m tall, erect, unbranched or branched just at inflorescence, with a xylopodium, glabrous, slightly branched, sometimes caespitose, not darkbrown when dry. Upper branches cylindrical to subquadrangular, wine- to rust-colored, lax, unwinged or inconspicuously winged, internodes 1.5–10 cm long. Leaves decussate, not imbricate, suberect, petiolate; petioles 1.5–10  mm long; blades 40–100  ×  (20–)35–70  mm, glaucous, chartaceous, not discolorous, elliptic to obovate or obovate to rhombic, base attenuate, apex obtuse, margin plane, with protuberances, sometimes with cilia; eucamptodromous-­brochidodromous, secondary veins 8–24 on each side of the midvein, inconspicuous, tertiary veins forming an inconspicuous reticulum; domatia absent. Synflorescence bracteose, botryum to diplobotryum type, lax; accessory branches absent, principal florescence truncate, paraclades 18–90 cm long; bracts 10–30 mm long, 4–15 mm long, elliptic; flowers pedicellate, hypopodium 1–2  mm long, epipodium 0.5–1  mm long; prophylls 1.5–8  ×  0.5–1.8  mm, wine-colored, reaching the middle of the floral tube or the apex, oblong to lanceolate, unkeeled, margin plane; floral tube 6–10  mm long, green, funnel-shaped, sometimes nitid in living material, fleshy or not, robust or thin; sepals 5.5–6.5 mm long, triangular, not caudate, not deflexed; epicalyx segments 2–8 mm long, dark-green or light-green, lanceolate, elliptic-lanceolate, dorsiventrally compressed, or flat, spreading to suberect, shorter or longer than the sepals; corolla 5–8  cm in diam., bright rose, rose-magenta or lilac, petals 23–30  ×  13–20  mm, apex obtuse; stamens 26–55, free portion of the filaments 8–10 mm long, not exserted or slightly exserted; ovary 2.8–3.5 × 3–4.5 mm, obconic, glabrous, style 12–18 mm long, ovules 36–48. Capsules 5–11 × 4–9 mm, globose, glabrous; seeds 23–34, 3–5 × 2.8–4 mm.

Descriptions

247

Distribution and Ecology (Fig.  13.55)  Diplusodon kielmeyeroides occurs in Minas Gerais, along the border between Minas Gerais and Goiás states, and in Goiás, near Chapada dos Veadeiros in areas of campos rupestres. Key to the varieties 1. Prophylls 1.5–2 mm long. Floral tube 6–6.5 mm long, floral tube thin; epicalyx segments 2–3 mm long, shorter than the sepals …................................................ ............................................................…. D. kielmeyeroides var. kielmeyeroides 1. Prophylls ca. 8 mm long. Floral tube 9–10 mm long, floral tube robust and fleshy; epicalyx segments 7–8 mm long, longer than the sepals ….................................. .....................................…………………..… D. kielmeyeroides var. macrocalyx 42a. Diplusodon kielmeyeroides A.St.-Hil. var. kielmeyeroides A.St.-Hil., Fl. Bras. Merid. (A.St.-Hil.). 3: 151, pl. 189, fig. b. 1833. Type: Brazil. Minas Gerais: en mai sur le plateau dit Chapadao, limites des prov. de Minas & Goyas, A.Saint-Hilaire s.n. (lectotype, P [01901886], designated by Lourteig 1989: 221; isolectotype, P [01901885]). Figs. 13.53 A–D, 13.55 Subshrubs, sometimes caespitose. Upper branches cylindrical to subquadrangular, unwinged, internodes 1.5–5 cm long. Leaves with the petioles 1.5–4 mm long; blades 25–55 mm wide, elliptic to obovate, margin without cilia; secondary veins 8–21 on each side, departing along the midvein. Paraclades 18–30 cm long; bracts 12–30 × 11–15 mm; prophylls 1.5–2 × 0.5–1 mm, wine-colored, not reaching the middle of the floral tube; floral tube 6–6.5 mm long, shiny in living material; sepals 5.5–6 mm long; epicalyx segments 2–3 mm long, dark-green, lanceolate, dorsiventrally compressed, shorter than the sepals; corolla 5–6.5 cm in diam., bright rose to rose-magenta, petals 23–25 × 13–16 mm; stamens 26–44, free portion of the filaments 8–9 mm long, slightly exserted; ovary 3–3.5 × 3–3.5 mm, style 15–18 mm long, ovules 36–45. Capsules 5–8 × 4–6.5 mm; seeds 23–24, 3–3.5 × 2.8–3.5 mm. Distribution and Ecology (Fig. 13.55)  Diplusodon kielmeyeroides var. kielmeyeroides is poorly represented in herbaria. It has a very restricted distribution, occurring in the mountains of Serra da Carcaça and Serra dos Pilões, west of Minas Gerais State, near the boundary with the state of Goiás. It grows in open cerrado and cerrado with lateritic soil, 990–1098 m. Collected with flowers and immature fruits in April. Specimens Examined: BRAZIL.  Goiás,  Catalão, Serra do Facão, 5 Apr 2008 (fl), Arantes et  al. 956 (CEN, HUFU). Minas Gerais, Coranmandel, Fazenda Mangue, 20 Apr 1989 (fr), Brandão 15379 (PAMIG); Coranmandel, Serra da Carcaça, Chapadão, entroncamento Vazante-Coranmandel, MG 188, km 38, 18°07′4″ S, 47°05′ W, 22 Apr 1997, (fl), Cavalcanti et al. 2266 (CEN, MO, NY); Coranmandel, entroncamento Vazante-Coranmandel, MG-188, km 38, Serra da Carcaça, Chapadão, 18°07′04″ S, 47°5″ W, 22 Apr 1997 (fr), Cavalcanti et al. 2267 (CEN, MO, NY); Guarda Mor, Serra dos Pilões, estrada vicinal Chapadão-Pilões,

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Fig. 13.53  Diplusodon kielmeyeroides var. kielmeyeroides and Diplusodon kielmeyeroides var. macrocalyx. A–D. D. kielmeyeroides var. kielmeyeroides. A. Part of vegetative branch. B. Branch with inflorescence. C. Floral bud. D. Flower with prophylls, without petals. E–F. D. kielmeyeroides var. macrocalyx. E. Leaf, abaxial surface. F. Flower with prophyll, without petals. (A–D Cavalcanti et al. 2266, CEN; E–F Pastore & Suganuma 1907, CEN)

Descriptions

249

km 50, na descida da serra, 17°50′79″ S, 47°08′87″ W, 22 Apr 1997 (fl), Cavalcanti et al. 2269 (CEN, MO, NY). Diplusodon kielmeyeroides var. kielmeyeroides is a rare and threatened species due to its restricted endemism in an area of active agriculture and road building. It occurs on small rocky outcrops. This variety is characterized by glaucous, elliptic to obovate leaves with 8–21 inconspicuous veins and an attenuate base (similar to the leaves of the genus Kielmeyera [Clusiaceae]), bracteose inflorescences branches, and showy flowers. The flowers are large (corolla 5–6.5 cm diam.) and have short prophylls (1.5–2 mm long), which do not reach the middle of the floral tube, and 26–44 stamens. Another glabrous species from Goiás State with large leaves and flowers is Diplusodon alatus, which differs mainly by its markedly quadrangular and winged branches and leaves sessile-amplexicaulous. Lourteig (1989) cited Saint-Hilaire s.n. at P as the holotype, which is considered here as the lectotype (ICBN Art. 9.17, ex. 14). Two specimens were examined at P. 42b. Diplusodon kielmeyeroides var. macrocalyx T.B.Cavalc., Phytotaxa 38: 32, fig. 2. 2011. Type: Brazil. Goiás. Alto Paraíso de Goiás, estrada entre Distrito de São Jorge e Colinas do Sul, 14°13′16″ S, 47°53′26″ W, 14 Mar 2007 (fl), J.  F. B.  Pastore, A.  Amaral-Santos, R.  M. Harley 1851 (holotype, CEN [00078202]; isotypes, MO, SPF, W). Figs. 13.53 E–F, 13.55 Subshrubs, slightly branched. Upper branches subquadrangular, sometimes inconspicuously winged, internodes 5–10 cm long. Leaves with petioles 2–10 mm long; blades 40–70  mm wide, obovate to rhombic, margin sometimes with cilia; secondary veins 17–24 on each side of the midvein. Paraclades 20–90  cm long; bracts 10–20 × 4–13 mm; prophylls ca. 8 × 1.5–1.8 mm, reaching the apex of the floral tube; floral tube 10 mm long, fleshy; sepals 6–6.5 mm long, fleshy; epicalyx segments 7–9 mm long, light-green, elliptic-lanceolate, flat, longer than the sepals; corolla 7–8 cm in diam., lilac, petals 27–30 × 18–20 mm; stamens 45–50, free portion of the filaments 9–10 mm long, not exserted; ovary ca. 2.8 × 4.5 mm, style ca. 12  mm long, ovules ca. 48. Capsules 9–11  ×  8–9  mm, coriaceous; seeds ca. 34, 4–5 × 3.5–4 mm. Distribution and Ecology (Fig. 13.55)  Diplusodon kielmeyeroides var. macrocalyx is known from Chapada dos Veadeiros, Goiás, in dry fields with quartzitic sandy soil, at 774–900 m. Represented by a single population with flowers registered from March to April and fruits from April to July. Specimens Examined: BRAZIL.  Goiás,  Alto Paraíso de Goiás, estrada entre Distrito de São Jorge e Colinas do Sul, 14°13′10″ S, 47°53′20″ W, 9 Apr 2007 (fl, fr), Pastore & Suganuma1907 (CEN, HUEFS, MO, NY, K, SPF, W); Alto Paraíso

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de Goiás, margem da estrada entre Alto Paraíso de Goiás e São Jorge, 17 Jul 2011 (fr), Faria-Jr. et  al. 1307 (CEN, IBGE, UB); Alto Paraíso de Goiás, margem da estrada entre Alto Paraíso de Goiás e São Jorge, 14°06′13″ S, 47°28′50″ W, 5 Apr 2014 (fl, fr), Faria-Jr. et al. 3966 (UB); Alto Paraíso de Goiás, margem da estrada de São Jorge para Colinas do Sul, 14°06′55″ S, 47°30′37″ W, 5 Apr 2014 (fr), Faria-Jr. et al. 3985 (CEN, UB). This variety is a subshrub with fleshy flowers that are consistently larger than the type variety. Other important distinctions between the varieties are the longer prophylls, floral tube, sepals, and epicalyx segments of Diplusodon kielmeyeroides var. macrocalyx. In addition, this variety occurs in northeastern Goiás, whereas the typical variety occurs in the state of Minas Gerais near the border with the state of Goiás. 43. Diplusodon lanceolatus Pohl, Flora 10: 152. 1827. Type: Brazil. Goiás: In montosis graminosis ad Serra d’ Ourada, Mar 1820, J. B. E. Pohl 1526 (firststep lectotype, designated by Lourteig 1989: 227, second-step lectotype, W [0053410], here designated; isolectotypes, G, W [0053409]). Figs. 13.54 A–G, 13.55 Diplusodon alutaceus Pohl, Flora 10: 152. 1827. Diplusodon lanceolatus Pohl var. alutaceus (Pohl) Koehne f. remotus Koehne, Fl. Bras. (Martius) 13(2): 330. 1877. Type: Brazil. “In campis siccis circa Paracatu, Cap. Minas Gerais et in Serra de Christaes, S. Luzia, Cap. Goyas, Nov 1818, J. B. E. Pohl 5793 (lectotype, W [0053411], designated by Lourteig 1989: 227; isolectotype, fragm. G). Diplusodon scaber Pohl, Flora 10: 152. 1827. Diplusodon lanceolatus Pohl var. scaber (Pohl) Koehne, Fl. Bras. (Martius) 13(2): 330. 1877. Type: Brazil. Goiás: In campis graminosis ad Serra dos Cristais et Santa Luzia, circa Paracatu et Minas Gerais, Nov et Dec 1818, J.  B. E.  Pohl 801 (lectotype, W [0053412], designated by Lourteig 1989: 227; isolectotype, G). Subshrubs 0.2–1.5 m tall, erect, branched, with a xylopodium, not dark-brown when dry. Upper branches cylindrical, lax, unwinged, pubescent, internodes 3–10 cm long. Leaves decussate, not imbricate, suberect, petiolate; petioles 3–5 mm long; blades 30–60 × 25–30 mm, olive-green, chartaceous, not discolorous, eucamptodromous, elliptic to elliptic-lanceolate, base obtuse, rarely acute, apex acute, margin plane, adaxial surface glabrescent, abaxial surface rough, covered with short, generally dense indumentum, pubescent to hirtellous; secondary veins 3–6 on each side, departing along the midvein, prominent; domatia absent. Synflorescence frondose-bracteose, diplobotryum spiciform to triplobotryum spiciform type, congested, sometimes with development of accessory branches of the first order 4–8 cm long, principal florescence 7–20 cm long, paraclades of the primary order 4–25 cm long, paraclades of the secondary order 1.5–10 cm long; bracts 10–20 × 3–6 mm, elliptic, margin plane; flowers sessile to subsessile, hypopodium 1–1.5 mm long, epipodium absent; prophylls (4–)6–8 × 3–4.5 mm, surpassing the floral tube or rarely less than the tube, elliptic-lanceolate or oblong to obovate, unkeeled, base acute, apex obtuse,

Descriptions

251

Fig. 13.54  Diplusodon lanceolatus. A. Vegetative branch. B. Branch with inflorescence. C. Floral bud. D. Prophyll, adaxial surface. E. Flower with prophylls, without petals. F. Fruit. G. Seed. (A Walter et al. 5120, CEN; B, D–G Strang 87; C Cavalcanti et al. 1254, CEN)

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Fig. 13.55  Geographic distribution of Diplusodon incanus, D. kielmeyeroides var. kielmeyeroides, D. kielmeyeroides var. macrocalyx, and D. lanceolatus

Descriptions

253

margin plane, ciliate; floral tube 4.5–6 mm long, clear green, funnel-shaped, pubescent; sepals 2–4 mm long, triangular, not caudate, not deflexed, ciliate, pubescent; epicalyx segments (3–)4–5.5 mm long, cylindrical, spreading to suberect, the same size as the sepals or a little longer, pubescent; corolla 3–4.5 cm in diam., clear rose, petals 14–17 × 8–11 mm, obovate, base acute, apex obtuse; stamens 18–26, free portion of the filaments 8.5–11  mm long, surpassing the sepals 2–3  mm; ovary 0.5–2.3  ×  2.2–2.8  mm, obconic, glabrous, style 10–16  mm long, ovules 20–33. Capsules 5.5–6 × 5–5.5 mm, globose, glabrous; seeds 10–27, 2–2.5 × 2–2.8 mm. Distribution and Ecology (Fig. 13.55)  Diplusodon lanceolatus is disjunct between Cadeia do Espinhaço in Minas Gerais State and mountains of the Brazilian Central Plateau. Its habitat is cerrado vegetation (sensu stricto) at 590–1100  m. Flowers from November to June; fruits from February to August. Specimens Examined: BRAZIL. Distrito Federal,  Brasília, QI 28, Lago Sul, 15°49′ S, 47°49′ W, 10 Jun 2004 (fl, fr), Walter et al. 5198 (CEN); Brasília, Fazenda Sucupira, encosta da margem direita do alto do córrego Riacho Fundo, 15°52′ S, 48°01′ W, 19 Apr 2001 (fl), Walter et al. 4833 (CEN). Goiás, Caldas Novas, 17°58′ S, 48°35′ W, 12 Feb 1993 (fl), Dias et  al. 383 (CEN); Caldas Novas, estrada de acesso à obra, próximo à vila de funcionários da obra, 17°55′ S, 48°33′ W, 27 Feb 1996 (fl), Pereira-Silva et al. 3358 (CEN, MO, NY); Caldas Novas, área de empréstimo da UHE Corumbá, perto da pista do antigo aeroporto, 1 Jun 1998 (fr), Cavalcanti et al. 2399 (CEN); Campo Alegre de Goiás, rodovia GO-213, a 24,2 km a leste da BR-050, 17°30″28″ S, 47°38′24″ W, 16 Nov 2016, Pereira-Silva & Bringel 16877 (CEN); Catalão, ca. 27 km NE de Catalão, 23 Jan 1970 (fl), Irwin et al. 25251 (NY, P, UB); Catalão, ca. 58 km NE de Catalão, 26 Jan 1970 (fl), Irwin et al. 25445 (NY, UB); Cristalina, 18 Mar 1964 (fr), Pereira 8980 (HB, P [2]); a 24 km NW de Goiás, 16 Mar 1982 (fl, fr), Graham 890 (NY); BR-050, Cristalina-Ipameri, km 88 às margens do rio Imbiruçú, 17°25′ S, 47°48′ W, 12 Mar 1989 (fl), Cavalcanti et al. 414 (CEN, NY, SPF); Formosa, lagoa do Perta Pé, 15°59′ S, 47°11′50″ W, 26 Mar 2002 (fl, fr), Pereira-Silva et al. 6260 (CEN, MO); Imbiruçú, à 6 km de Paraopeba, 28 Feb 1959 (fl), Heringer 11264 (B); Ipameri, estrada Ipameri-Caldas Novas, km 34, 17°43′ S, 48°20′ W, 13 Mar 1989 (fl, fr), Cavalcanti et al. 421 (CEN, NY, SPF); Ipameri, Fazenda Santo Antonio Fundão, córrego afluente da margem esquerda do rio Jacuba, 17°43′ S, 48°10′ W, 24 Mar 1995 (fl, fr), Cavalcanti et al. 1395 (CEN, MO, NY); Ipameri, lado direito da estrada de Caldas Novas-Ipameri, mais ou menos a 3  km após a Ponte São Bento, 17°44′31″ S, 48°28′08″ W, 12 Mar 1997 (fl), Cavalcanti et  al. 2150 (CEN, MO, NY); Ipameri, estrada de acesso ao córrego Jacubeiro, 14 Mar 1997 (fl, fr), Cavalcanti et al. 2174 (CEN, MO, NY); 25 km SW de Monte Alegre de Goiás, 13 Mar 1973 (fl), Anderson 7002 (UB); João Pinheiro, fazenda da Manesmann, 20 Apr 1987 (fl), Coelho s.n. (VIC 10107); without reference to municipal district: rio Paranã, Glaziou 21414a (P). Mato Grosso, Diamantino, estrada Diamantino-Sumidouro (BR 364), cerca de 26 km após o trevo para São José do Rio Claro, Fazenda Nova Era, 14°09′ S, 56°45′ W, 18 May 1997 (fl), Souza et al. 16322 (CEN, ESA). Minas Gerais, Alto Parnaíba, 9 km NW de Santa Juliana, 30 Jan 1978 (fl), Krapovickas et al. 33328 (G, P); Araguari, 4 Mar

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1996, Tameirão-Neto 1977 (IBGE); Belo Horizonte, Universidade Federal de Minas Gerais, Campus ICB, Mar 1979 (fr), Pinheiro et al. 247 (BHCB); Belo Horizonte, Morro das Pedras, próximo a Belo Horizonte, 4 Mar 1945 (fl, fr), William & Assis 5996 (R); Belo Horizonte, Campus do ICB, 4 Feb 1982 (fl), Ferrari 929 (BHCB); Braslândia de Minas, Fazenda Brejão, 1 Feb 2001 (fl, fr), Pompeu & Matoso 6 (BHCB). Buenópolis, estrada para Curimataí, 21 May 1990 (fr), Mello-Silva et al. 286 (SPF); Cabeceira Grande, região da ponte sobre o rio Preto, a 29 km do entroncamento Brasília/Unaí/Palmital, na direção de Palmital, 16°02′15″ S, 47°18′34″ W, 14 Feb 2002 (fl), Walter et al. 5009 (CEN, MO); estrada Palmital-Cabeceira Grande, ca. 7 km à esquerda, ca. 4 km após a guarita, 18°04′01″ S, 47°16′20″ W, 20 Feb 2003 (fl, fr), Amaral-Santos & Pereira 1866 (CEN, MO); Fazenda São Bento, região do rio Bezerra, 14 Feb 2002 (fl), Walter et al. 5120 (CEN, MO); Conselheiro Mata, estrada de terra, a 3 km após a saída da cidade de Conselheiro Mata, 18°43′14″ S, 43°59′14″ W, 19 Jan 2003 (fl), Cavalcanti et  al. 3119 (CEN, MO); Corinto, ca. 12 km W de Corinto, 4 Mar 1970 (fl), Irwin et al. 26883 (NY, P); Corinto, Beltrão, 13 Mar 1997 (fl), Hatschbach et al. 66154 (M, MBM n.v.); Cristália, rio Mumbuca, 11 Feb 1991 (fl), Hatschbach et al. 55060 (BR, MBM n.v., P); Diamantina, 17 Feb 1973 (fl), Hatschbach & Ahumada 31716 (NY, MBM n.v., P); Curvelo, rodovia Curvelo-Diamantina, ca. 9 km de Curvelo, em direção à Sete Lagoas, 18°49′01″ S, 44°29′18.5″ W, 5 Apr 1998 (fr), Souza et al. 20865 (ESA); Francisco Sá, 31 km NE de Francisco Sá, na estrada para Salinas, 11 Feb 1969 (fl), Irwin et al. 23061 (NY, P, UB); Gouveia, encostas pedregosas de morro, 22 Nov 1971 (fl), Hatschbach & Pelanda 27794 (MBM); Gouveia, 20 Dec 1997, Lemos-Filho s.n. (BHCB 41165); Gouveia, margem da estrada para Cuiabá, região da Bucaina, 18° 25′ 08′′S, 43° 46′ 49′′W, 1 May 2016 (fl), Faria-Jr. & Otoni 5810 (CEN, HDJF, UB); GouveiaCongonhas do Norte, ca. 6 km da estrada Gouveia-Curvelo, 4 Mar 1999 (fl), Souza & Souza 22283 (ESA); Imbiruçú, 28 Feb 1959 (fl), Heringer 6808 (HB, P, UB); Indianapolis, Fazenda Bela Tanda and neighbouring fazendas, 19°03′ S, 47°57′ W, 7 Mar 1986 (fl, fr), Gottsberger & Doring 71–7386 (CEN); Inimutaba, BR-259, a 74  km de Gouveia, 18°42′ S, 44°20′ W, 18 Feb 1989 (fr), Cavalcanti et  al. 263 (CEN, NY, SPF); Jaboticatubas, São José da Serra, 1 Mar 1987 (fl), Vieira & Hermano 560 (CEN, VIC); entre Cavalcante e Conceição, May 1865 (fl), Burchell 8046 (K); Jequitaí, BR 365, entre km 70–72, 17 Feb 1991 (fl), Hatschbach et al. 55227 (MBM n.v., P); BR-365, km 66, 13 Mar 1995, Hatschbach et  al. 61829 (MBM); Joaquim Felício, Serra do Cabral, ca. 2 km N de Joaquim Felício, 5 Mar 1970 (fl, fr), Irwin et al. 26966 (K, NY, P, UB); Joaquim Felício, Serra do Cabral, 17°45′ S, 44°12′ W, 13 Feb 1988 (fl, fr), Thomas et al. 5948 (CEN [2], SPF, NY, P); Joaquim Felício, Serra do Cabral, 2–8 km NW da cidade, na estrada para a Torre de TV e a Fazenda Bocaina, 17°45′10″ S, 44°10′56″ W, 19 Mar 1994 (fl, fr), Sakuragui (CFCR) 15174 (CEN [2], ESA); Joaquim Felício, Serra do Cabral, 17°43′44.5″ S, 44°10′33.8″ W, 13 Mar 1999, Souza & Souza 22003 (ESA); Lagoa Santa, Feb 1835 (fl), Warming s.n. (P); Lagoa Santa, estrada Lagoa Santa-Grutas, 12 Jan 1965 (fl), Handro 65 (SP); Lassance, rodovia 496, 13 Apr 1996, Hatschbach et  al. 64635 (CEN, MBM n.v.); Minas Novas, estrada de terra para Campo Buriti, na estrada Capelinnha-Minas Novas, 500  m, 17°17′38″ S, 43°37′54″ W, 21 Jan 2003 (bud),

Descriptions

255

Cavalcanti et al. 3134 (CEN, MO); Minas Novas, Saint-Hilaire B1 1336 (P); entroncamento rodovia BR-367-Minas Novas, km 8, em uma estrada vicinal a leste de Campo Alegre, 17°17′59″ S, 42°37′97″ W, 27 Apr 1997 (fl), Cavalcanti et al. 2311 (CEN, MO [2], NY); Monjolo, estrada de terra Monjolos-Diamantina, 6  km de Monjol, 18°16′57″ S, 44°05′09″ W, 7 Feb 2015 (fl, fr), Cavalcanti et al. 3919 (CEN); Montes Claros, ca. 25 km O de Montes Claros, 23 Feb 1969 (fl), Irwin et al. 23713 (NY, P, UB); Montes Claros, rodovia BR-365, próximo ao km 21, 9 Mar 1998 (fr, fl), Hatschbach et al. 67539 (MBM); Nova Ponte, 30 Mar 1987, Pedralli et al. 705 (CEN); Paracatu, 16°74′72″ S, 46°48′29″ W, 21 Apr 1997 (fl), Cavalcanti et al. 2264 (CEN); Paracatu, ca. 5 km N de Paracatu, 5 Feb 1970 (fl), Irwin et al. 26115 (NY, P, UB); Paracatu, km 477 da BR-7, Posto Sombra, 22 Feb 1964 (fl), Laboriau & Valio 1212 (SP); Paracatu, Fazenda Acangaú, 17º12′S, 47º06′W, 5 Mar 1989 (fl), Walter et al. 133 (IBGE, UB, UEC n.v.); Paraopeba, Horto Florestal, 13 Mar 1954, Heringer 3354 (P); Paraopeba, Horto Florestal, 14 Jun 1954 (fr), Heringer 3456 (P, UB); Paraopeba, Horto Florestal, 3 Mar 1955 (fl, fr), Heringer 3746 (HB, P, UB); Paraopeba, Horto Florestal, 5 Apr 1965 (fl, fr), Paula 59 (NY, SP, UB); EFLEX, 2 Mar 1987 (fl, fr), Silveira 75 (CEN, VIC); Patrocínio, ca. 4 km N de Patrocínio, 31 Jan 1970 (fl), Irwin et  al. 25745 (NY, UB); Patrocínio, MG-188 (PatrocínioCoromandel), 18°48′ S, 46°55′ W, 28 Feb 1989 (fl), Mendonça et al. 1233 (IBGE, P, UB n.v.); Pedro Leopoldo, 12 Feb 1973 (fl), Hatschbach & Ahumada 31484, (NY, MBM n.v., P); Perdizes, 5 Jan 1995, Tameirão-Neto & Werneck 1021 (CEN); Santana do Riacho, 15 Apr 1983, Andrade & Lopes s.n. (BHCB 8808); Santana do Riacho, Serra do Cipó, Rodovia Belo Horizonte-Conceição do Mato Dentro, km 103, próximo ao afloramento de calcáreo, 24 Mar 1986 (fl, fr), Zappi & Kameyama (CFSC) 9636 (SPF); Serra do Cipó, entre São José do Almeida e Vacaria, Feb 1953 (fl, fr), Vidal II-6351 (P, R); Serra do Cipó, 3 Apr 1958 (fl, fr), Strang 84 (R); 3 Apr 1958 (fr), Strang 87 (R); São Gonçalo do Rio Preto, Parque Estadual do Rio Preto, trilha do alojamento até o córrego das Éguas, 18°07′40″ S, 43°20′53″ W, 19 Feb 2002 (fl), Lombardi 4549 (BHCB, CEN); Serro, Vauthier 421 (G, P); Sete Lagoas, 24 Mar 1983 (fl), Gontijo s.n. (BHCB 9220); Triângulo Mineiro, Vale do rio Araguari, estrada da Coalbia, 25 Feb 1994, Mota 2185 (HUFU); Uberaba, ca. 78 km SE de Uberlândia, 11 Mar 1982 (fl, fr), Graham 860 (NY); Uberaba, 21 Jan 1988 (fl), Hatschbach & Cordeiro 51743 (MBM n.v., P); Uberlândia, Reserva Ecológica Panga, 20 Mar 1987 (fl, fr), Araújo 173 (CEN, HUFU); Uberlândia, 15 Feb 1993 (fl), Barbosa 586 (CEN, HUFU); Unaí, MG-188, 20 km S de Unaí, na estrada para Paracatu, 16°32′ S, 46°53′ W, 11 Feb 1988 (fl, fr), Thomas et al. 5860 (NY, P, SPF); Várzea da Palma, 16 Jan 1996 (fl), Hatschbach et al. 64200 (BR, CEN, MBM n.v.); Várzea da Palma, Clube Caça e Pesca Itororó, 9 Feb 1994 (fl, fr), Arantes 144 (HUFU); Várzea da Palma, 9 Mar 1994 (fl), Arantes 159 (HUFU); Várzea da Palma, 19 Jan 1995 (fl), Romero 1820 (CEN, HUFU); without locality, Fazenda Diamante, 9 Apr 1931 (fl, fr), Mexia 5578 (K, NY, P); Claussen s.n. (G, K); Claussen s.n. ([herb. Fl. Bras. n. 119] G); 1840, Claussen 19a (BR [2]); Claussen 308 (NY, P [3], RB); Nov 1843, Weddell s.n. (G, P); Feb 1953 (fr), Vidal II-6369 (R).

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Diplusodon lanceolatus is represented by branched subshrubs characterized by the olive-green, elliptic to elliptic-lanceolate leaves and a synflorescence with the flowers united in botrya. The flowers are sessile to subsessile with long prophylls (6–8 mm long) which usually surpass the floral tube and long epicalyx segments which are cylindrical of usually 4–5.5  mm long, spreading to suberect, with the same size as the sepals or a little longer. There is some variation from one population to another in stamen number, ranging from 18 to 26 stamens, with 24 being the most common. Diplusodon lanceolatus, like D. bolivianus, D. bradei, D. virgatus, D. oblongus, and D. ulei, at times develops accessory branches. Lourteig (1989) cited Pohl 1526 (W) as the holotype; however, Diplusodon lanceolatus was described by Pohl (1827a) without citation of specimens in the protologue. Subsequently, Pohl (1827b) referred to a single collection of D. lanceolatus collected in April in Serra Dourada, in the state of Goiás, without mentioning a collection number. Two specimens were examined at W whose labels correspond to the same locality cited by Pohl (1827b), and both are numbered as Pohl 1526. Lourteig (1989) indicated Pohl 1526 as holotype, and this choice should be interpreted as a first-step lectotypification. The specimen with the original label annotated by Pohl (W0053410) is here designated as a second-step lectotype. Diplusodon alutaceus was described by Pohl (1827a) without citation of specimens in the protologue. Shortly thereafter, Pohl (1827b) referred to a single collection of D. alutaceus collected “in siccis montosis ad Corumba et rio S.  Marcos, Capitaniae Goyaz, legi mense Decembri 1818.” At W there is just one specimen identified by Pohl as D. alutaceus (Pohl 5793) whose label indicates another locality; however, Pohl’s annotation is the evidence to consider this specimen as original material. Lourteig (1989) indicated this specimen as holotype, which is here considered as the lectotype (ICBN Art. 9.17, ex. 14). Diplusodon scaber also was described by Pohl (1827a) without citation of specimens in the protologue. Subsequently, Pohl (1827b) referred to a collection of D. scaber collected “in Serra de Cristaes et ad S.  Luzia, Capitaniae Goyas; circa Paracatu do Principe, Capitaniae Minas Geraes, legi mense Novembri et Decembri 1818.” At W, there is one specimen identified as D. scaber (Pohl 801) whose label indicates the same locality, which is annotated by Pohl. Lourteig’s citation of this specimen as “holotype” constitutes lectotypification (ICBN Art. 9.17, ex. 14). 44. Diplusodon leucocalycinus Lourteig, Bradea 5(19): 232. 1989. Type: Brazil. Goiás: Alto Paraíso de Goiás, Chapada dos Veadeiros, rodovia GO-12, 20 Feb 1975 (fl), G.  Hatschbach 36374 (holotype, P [01901877]; isotypes, BR, C, CTES n.v., MBM, MO, MU, P [01901878], S, SPF, US). Figs. 13.56 A–L, 13.60 Diplusodon irwinii Lourteig, Bradea 5(19): 231. 1989. Type: Brazil. Goiás: Alto Paraíso de Goiás, Chapada dos Veadeiros, 42 km N of Alto Paraíso de Goiás, 25 Mar 1971 (fl, fr), H. S. Irwin, R. M. Harley & G. L. Smith 33160 (holotype, P [01901887]; isotypes, NY, SP, UB, W). Diplusodon urceolatus Lourteig, Bradea 5(19): 233. 1989. Type: Brazil. Goiás: Serra Geral do Paranã, ca. 20 km S of São João da Aliança, 17 Mar 1971 (fl, fr),

Descriptions

257

H. S. Irwin, R. M. Harley & G. L. Smith 32100 (holotype, P [01902004]; isotypes, NY, U, UB). Shrubs 0.4–1.5  m tall, erect, much-branched, with a xylopodium, sometimes dark-green or dark-brown when dry. Upper branches cylindrical, sometimes rustcolored, unwinged, villous-tomentose to pubescent, internodes 1–2 cm long. Leaves decussate, not imbricate, spreading to suberect; petioles 2–5  mm long; blades 12–40 × 8–30 mm, not discolorous, coriaceous, petiolate, eucamptodromous, ovate to elliptic or rarely orbicular, base acute-attenuate to obtuse, apex obtuse to acute, margin plane, yellow, villous-tomentose, hirtellous to pubescent or with equalarmed stellate indumentum mixed with villous-­tomentose or pubescent indumentum to various degrees, from only stellate, with long trichomes rare, to only villous-tomentose or pubescent, generally canescent on the abaxial surface, trichomes bulbous-­based; secondary veins 3–5 on each side, departing along the midvein; domatia present. Synflorescence frondose, diplobotryum type, congested; accessory branches absent, principal florescence generally caducous, when present ca. 18 cm long, paraclades 8–19 mm long; bracts similar to the vegetative leaves except diminishing in size toward the apex of the branch; flowers subsessile, hypopodium 0.5–1 mm long, epipodium absent; prophylls 9–11 × 3–5 mm, surpassing the apex of the floral tube, oblong to oblong-elliptic, keeled, base obtuse to retuse, apex obtuse, acuminate to apiculate, margin plane, usually hyaline, villoustomentose; floral tube 6–9 mm long, oblong, totally covered with villous-tomentose white indumentum or sometimes hirtellous to pubescent, yellow glands below the petal insertion; sepals 1–2.5 mm long, triangular, not caudate, not deflexed, villoustomentose; epicalyx segments 4.5–6  mm long, cylindrical, dorsiventrally compressed, erect, surpassing the sepals 2–3 mm long, with short, sparse indumentum, sometimes hirsute; corolla 4–4.5  cm in diam., clear rose to rose-magenta, petals 18–22 × 8–10 mm, oblong to elliptic, apex obtuse; stamens 12–15, free portion of the filaments 9–10 mm long, surpassing the sepals ca. 1 mm; ovary 2–4 × 2.5–3 mm, obconic, sulcate-angulate with a lobed, densely pilose at the apex, indumentum whitish or glabrous, persistent on the capsule, style 13–20 mm long, ovules 12–22. Capsules 4–4.5 × 3.5–4.5 mm, oblong, with white indumentum at the apex; seeds 4–11, 2.5–3.2 × 2–2.1 mm. Distribution and Ecology (Fig. 13.60)  Diplusodon leucocalycinus inhabits campos sujos and campos rupestres of Chapada dos Veadeiros, in Goiás State, and surrounding areas at 1000–1560  m. Flowers from January to May; fruits registered from February to July and September and November. Specimens Examined: BRAZIL.  Goiás,  Água Fria de Goiás, GO-118, subida para a torre repetidora do Roncador, 14°53′06″ S, 47°33′25″ W, 5 Apr 2006 (fl), Cavalcanti et al. 3754 (CEN); Água Fria de Goiás, rodovia GO-118, subida para a Torre Repetidora de Roncador, 8 May 2000 (fl, fr), Hatschbach et al. 70654 (MBM); Água Fria de Goiás, estrada de acesso à repetidora Roncador, cerca de 5 km a leste da GO-118 (BR 010), 4 Feb 2014 (fl), Pereira-­Silva 16578 (CEN); Água Fria de Goiás, estrada de acesso à repetidora Roncador, cerca de 5 km a leste da GO-118,

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Fig. 13.56  Diplusodon leucocalycinus. A. Branch with inflorescence. B. Leaf, abaxial surface. C–F. Trichomes of leaves. G. Floral bud. H. Prophyll, adaxial surface. I–J. Flowers with prophyll, without petals. K. Epicalyx segment with revolute margin. L. Ovary. (A–C, H Cavalcanti et al. 384, CEN; D, G, I, Irwin et al. 33160, UB; E–F Anderson 7188, UB)

Descriptions

259

14°53′18.1″ S, 47°33′03″ W, 3 Jul 2015 (fr), Cavalcanti & Pereira-Silva 3986 (CEN); Alto Paraíso de Goiás, 30.6 km ao NE de Alto Paraíso de Goiás, na rodovia para Teresina de Goiás, 13°56′ S, 47°26′ W, 5 Mar 1988 (fr), Ginzbarg & César 747 (UB, TEX-LL); Alto Paraíso de Goiás, estrada Alto Paraíso de Goiás-Nova Roma, a 17 km de Alto Paraíso, 13°55′12″ S, 47°25′12″ W, 4 Apr 1997 (fl), Cavalcanti et al. 2204 (CEN, MO); Alto Paraíso de Goiás, estrada para Cavalcante, 8 Feb 1987 (fl, fr), Romaniuc-Neto et al. 605 (CEN, SP); Alto Paraíso de Goiás, ca. 20 km S de Cavalcante, 10 Mar 1969 (fl, fr), Irwin et al. 24231 (NY, S, UB, US); Alto Paraíso de Goiás, 13°44′ S, 47°22′ W, GO-118, Alto Paraíso de Goiás-Teresina de Goiás, km 31, 7 Mar 1989 (fl), Cavalcanti et al. 384 (CEN, K, NY, MBM, SPF, UFG); Alto Paraíso de Goiás, GO-118, Alto Paraíso-Teresina de Goiás, 31 km de Alto Paraíso, 13°54′09″ S, 47°21′36″ W, 17 Nov 2005 (fr), Cavalcanti et al. 3615 (CEN, MO); Alto Paraíso de Goiás, estrada Alto Paraíso-Teresina de Goiás, a 37  km de Alto Paraíso, local de camping, próximo à cachoeira, hotel ecológico Alpes Goianos, 13°54′14″ S, 47°23′79″ W, 4 Apr 1997, Cavalcanti et  al. 2209 (CEN, MO); Cavalcante, 24 km na estrada S de Teresina de Goiás, 16 Mar 1973 (fl), Anderson 7188 (NY, P, UB); Cavalcante, comunidade Kalunga, caminho para a cachoeira Capivara I, 20 Apr 2003 (fr), Pastore et al. 470 (CEN); São João da Aliança a 7 km da rodovia S de São João da Aliança, 22 Mar 1973 (fl, fr), Anderson 7658 (MO, NY, UB); São João da Aliança, ca. 30  km de Alto Paraíso, Chapada dos Veadeiros, 13°59′ S, 47°21′ W, 20 May 1994 (fl, fr), Silva et al. 1924 (IBGE, P); Teresina de Goiás, GO-118, estrada Teresina de Goiás-Alto Paraíso de Goiás, a 23  km de Teresina, 13°51′40″ S, 47°19′41″ W, 29 Jan 2003 (fl), Cavalcanti et al. 3150 (CEN, MO); Teresina de Goiás, a 30 km de Teresina, 13°53′45″ S, 47°21′33″ W, 29 Jan 2003 (fl), Cavalcanti et al. 3153 (CEN, MO); Teresina de Goiás, a 30 km de Teresina, 13°53′45″ S, 47°21′33″ W, 29 Jan 2003 (fl), Cavalcanti et al. 3156 (CEN, MO); Teresina de Goiás, GO-118, road Teresina de Goiás-Alto Paraíso de Goiás, km 45, 13°56′45.5″ S, 47°27′16″ W, 1 Sep 2004 (fl, fr), Cavalcanti et al. 3542 (CEN). The plants of Diplusodon leucocalycinus are much-branched shrubs, with branches covered by dense and white indumentum giving a canescent appearance, special to the abaxial leaf surface. The same indumentum covers the entire floral tube, making it visibly white. Besides this conspicuous characteristic, the leaves when dried become dark-green to dark-brown in color. The leaves are ovate to elliptic, and the prophylls are keeled and the same size as the floral tube or slightly longer. The flowers have long (9–11 mm long), oblong to oblong-elliptic and keeled prophylls, which surpass the apex of the floral tube. The floral tube is long (6–9 mm long), with short sepals (1–2.5 mm long), and the epicalyx segments are erect, surpassing the sepals 2–3 mm long, and the androecium has 12–15 stamens. An unusual characteristic within the flower of D. leucocalycinus is the presence of a white indumentum at the apex of the ovary that persists on the apex of the capsule. In the same publication that Lourteig (1989) described Diplusodon leucocalycinus, she also described D. irwinii and D. urceolatus from the same region, the Chapada dos Veadeiros in Serra Geral do Paranã, Goiás. The characteristics separating the three species, according to Lourteig’s key (Table 13.1), such as the shape and

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length of the epicalyx segments and the ratio of the length of the epicalyx segments to the length of the sepals, were analyzed in the type material and additional herbarium specimens of the three species. The differences in Lourteig’s (1989) key are not consistently useful to distinguish the three species; I have decided that these species should be treated as synonymous. Diplusodon irwinii and D. urceolatus are synonymized here under the binomial D. leucocalycinus, whose epithet refers to the white indumentum of the floral tube. Indumentum on the ovary is uncommon in the genus, appearing only in species on Chapada dos Veadeiros. This characteristic is shared by D. appendiculosus, a species which differs from D. leucocalycinus in several ways, principally because it is otherwise totally glabrous; it has wide prophylls and caudate, deflexed sepals; and it has epicalyx segments that are obovate to lanceolate and foliaceous. There is a minor variation in some characteristics of Diplusodon leucocalycinus, especially those related to indumentum on different parts of the plant. At one extreme are dark-brown specimens that are totally covered with white, villous indumentum made of trichomes that have an enlarged base (Anderson 7658, Irwin et al. 33160, Hatschbach 36374, Irwin et al. 32100). At the other extreme are the darkbrown but non-canescent specimens, with short, equal-armed stellate trichomes forming visible tufts. In this case, the flowers are not so densely pilose, and the epicalyx segments can have a sparser indumentum of long trichomes, sometimes with short bulbous trichomes (Anderson 7188). Between these extremes are variants in which the long trichomes are mixed with stellate ones (Cavalcanti et  al. 384, Romaniuc et al. 605). Another variable characteristic is the shape of the prophylls, which varies from narrow and oblong to more elliptic; the prophyll indumentum also varies as mentioned above. The length and width of the epicalyx segments are also sometimes variable, being wider and longer in some specimens. 45. Diplusodon longipes Koehne, Bot. Jahrb. Syst. 4: 404. 1883. Type: Brazil. Goiás: Sertão do Amaroleité, Sep/Oct 1844, H. A. Weddell 2844 (lectotype, P [01901874] designated by Lourteig, 1989: 234; isolectotypes, F, P [01901875, 01901876]). Figs. 13.57 A–G, 13.60 Subshrubs 0.4–0.6 m tall, erect, branched just at inflorescence, sometimes caespitose, with a xylopodium, glabrous, glaucous, unbranched, not dark-brown when dry. Upper branches cylindrical to subquadrangular, unwinged, light-yellow, internodes 3–4  cm long. Leaves decussate, not imbricate, erect, sessile; blades 30–70  ×  20–50  mm, glaucous, chartaceous, not discolorous, acrodromous-basal, glabrous, ovate, base retuse, sometimes slightly amplexicaulous, apex obtuse, sometimes apiculate, margin plane, light-yellow; secondary veins 4–6 on each side, inconspicuous; domatia absent. Synflorescence frondose, botryum type, congested; accessory branches absent; flowers pedicellate, hypopodium 15–40 mm long, epipodium 1–2 mm long; prophylls 18–22 × 10–15 mm, elliptic to ovate, surpassing the apex of the floral tube, totally hiding the flower, base and apex obtuse, margin plane; floral tube 9–11 mm long, clear green, oblong, narrow; sepals 2–3 mm long,

Descriptions

261

Table 13.1  Comparison of morphological characteristics among D. leucocalycinus, D. irwinii, and D. urceolatus, based on the diagnoses of Lourteig (1989) Characteristics/ species D. irwinii Leaf shape Elliptic to oblong Leaf nerves Leaf size (mm) 10–40×7–28 Leaf base Attenuate to decurrent

D. leucocalycinus Elliptic to ovate-elliptic 4–7 15–38×8–24 Acute or subacute, decurrent Appressed to subappressed

Leaf indumentum

Dendritic pubescence

Leaf color Petiole (mm) Petiole (shape) Prophyll shape

Dark-brown 0.5–5 Thick Spathulate, oblong, slightly keeled The size of the tube

The size of the tube or larger

Short (ca. 1)

Prophylls/tube

D. urceolatus D. xerampelinus Elliptic to oblong Ovate-elliptic to elliptic 4–7 2–3 15–40×10–20 12–25×6–12 Attenuate Decurrent on a thick petiole Dendritic pubescence, dense on both surfaces Dark-brown 0–3.5 Thick Oblong, keeled

Short (ca. 1)

Densely appressed to subappressed pilose – 0.5–3 Thick Oblong spathulate, keeled Almost the size of the floral tube or smaller Small (1)

–

Subulate

Subulate

Subulate

Subulate

Erect

Erect

Erect

–

Dark gray 0.5 Thick Oblong, keeled

The size of the floral tube

Sepal length (mm) Epicalyx (shape) Epicalyx (position) Epicalyx segments indumentum Epicalyx segments/sepals Stamen number Indumentum of the ovary

Pubescent

Appressed pubescent

Pubescent

Pubescent

5–6x the size of the sepals 12 Densely dendritic pilose

Ovule number

ca. 12

4–5x the size of the sepals 14 Densely pilose (trichomes dendritic) 16

5–6x the size of the sepals 14 Densely pilose (trichomes dendritic) 16

The size of the sepals or shorter 10–12 Dendritic trichomes, very short 14

triangular, not caudate, slightly deflexed, glabrous; epicalyx segments 2.5–3  mm long, dorsiventrally compressed, spreading to suberect, the same size as or slightly shorter the sepals; corolla 5–6  cm in diam., purple, petals 23–28  ×  15–17  mm, obovate, apex obtuse; stamens 13–18, free portion of the filaments 9–11 mm long, exserted 2–2.5 mm; ovary ca. 3.1 × 3.1 mm, oblong, glabrous, style 16–20 mm long,

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Fig. 13.57  Diplusodon longipes. A. Vegetative branch. B. Branch with inflorescence. C. Prophyll, abaxial surface. D. Flower with prophyll, without petals and style. E. Flower without prophylls. F. Branches with fruits. G. Fruit. (A, F–G Pereira-­Silva et al. 4422, CEN; B–E Mello-Silva et al. 2248, SPF)

Descriptions

263

ovules 30–52. Capsule 6–6.5  ×  4–6  mm, globose, glabrous; seeds 8–10, 2–2.6 × 1.6–1.9 mm. Distribution and Ecology (Fig. 13.60)  Diplusodon longipes was long known only from the type material, and it is still poorly represented in herbaria. This species is presently known to occur only on Chapada dos Veadeiros, in Goiás State, with a disjunction in the mountains of Uruaçu and the vicinity of Niquelândia, at 380– 550 m. It inhabits cerrado and campo sujo on sandstone soil. Flowers from November to February, fruits from January to February and June, old fruits registered in October and November. Specimens Examined: BRAZIL. Goiás,  Cavalcante, estrada para a balsa “Porto dos Paulistas” (rio Tocantins), 13°41′07″ S, 47°51′22″ W, 10 Nov 2000 (fl, fr), Pereira-Silva et al. 4422 (CEN, MO, NY); Cavalcante, entroncamento Vila Venenobalsa do rio São Félix-Cavalcante, km 3,2 m, 13°40′53″ S, 47°51′46″ W, 21 Feb 2002 (fr), Pereira-Silva et al. 5941 (CEN, MO, SPF); Cavalcante, Serra do Tombador, Reserva Fundação Boticário, 7 Nov 2012 (fr), Pereira-Silva et  al. 16449 (CEN); Cavalcante, estrada 18  km entre RPPN Serra do Tombador e Cavalcante, ramal 200 m ao N, 13°37′41″ S, 47°42′00″ W, 1 Feb 2015 (fl), Simon et al. 2551 (CEN); Cavalcante, Reserva Natural Serra do Tombador (RPPN), 18 Nov 2013 (fl), Barbosa et al. 1092 (CEN, HRCB n.v.); Colinas do Sul, 14°12′39″ S, 47°55′45″ W, ca. 18 km de Colinas do Sul pela GO-241, sentido São Jorge, 27 Fev 2014 (fl), Mendoza et al. 4172 (CEN, USZ); Niquelândia, 25 Nov 1992 (fl), Cordovil-Silva et al. 154 (CEN, MO, NY); Niquelândia, estrada de terra entre Niquelândia e Colinas do Sul, 14°19′41.4″ S, 48°07′59.7″ W, 5 Jun 2001 (fl, fr), Pastore & Suganuma 1399 (CEN); Niquelândia, saída para Uruaçú, fazenda após a lagoa, Amaral-Santos et al. 3677 (CEN); Niquelândia, estrada Niquelândia-Colinas do Sul, 59  km da saída de Niquelândia, 14°19′27″ S, 48°08′24″ W, 1 Dec 2003 (fl), Mello-Silva et al. 2248 (CEN, NY, K, RB, SPF); Niquelândia, rodovia GO-132, às margens da rodovia, 14°  17′ 14′′S, 48°  09′ 08′′W, 26 Nov 2014 (fl), Oliveira et  al. 467 (CEN, RB); Uruaçu, estrada de terra da GO-237, para a Fazenda Ponte Alta, cerca de 7 km da GO-237, 14°35′ S, 48°57′ W, 6 Feb 1996, Walter et al. 3008 (CEN, MO, NY); without locality, 30 km a leste de Niquelândia, 21 Jan 1992 (fl), Hatschbach & Kummrow 56304 (MBM, P). Diplusodon longipes is an ornamental subshrubs, almost unbranched, with large purple flowers (corolla 5–6 cm in diam) and large (30–70 × 20–50 mm), sessile, and glaucous leaves. The most prominent characteristics of the species are the very long pedicels (1.5–4  cm long) and the large prophylls that hide the entire floral tube. These characteristics are also found in D. cryptanthus, another species from Goiás that differs from D. longipes by its corymbiform, shrubby, much-branched habit, smaller, elliptic-lanceolate leaves, and flowers 4.5 cm in diam. Diplusodon marginatus, another species of Goiás with long pedicels and broad leaves, differs from D. longipes principally by its characteristic leaves with purplish margins, its absent or very small epicalyx segments, and also its androecium of 23–28 stamens.

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Lourteig (1989) referred to a specimen of Weddell 2844 at P as the holotype, which is here considered lectotypification (ICBN Art. 9.17, ex. 14). 46. Diplusodon macrodon Koehne, Fl. Bras. (Martius) 13(2): 338, pl. 63, fig. 3. 1877. Type: Brazil. Goiás: São Bernardo, Aug 1842 (fr), G. Gardner 4135 (lectotype, P [01901873] designated by Lourteig 1989: 228; isolectotypes, BR, F, fragm. F, G [3], K [2], NY [2], OXF, P [01901871, 01901872], US [2], W. Figs. 13.58 A–F, 13.60 Shrubs 0.5–2.5 m tall, erect, much-branched, not dark-brown when dry. Upper branches subquadrangular, unwinged, covered by dense indumentum with short trichomes oriented in different directions, sometimes looking stellate trichomes, internodes 10–12 mm long. Leaves decussate, not imbricate, suberect; petioles 1–2.5 mm long; blades 8–17  ×  1.5–3 (−4–5) mm, chartaceous to coriaceous, petiolate, not discolorous, sometimes slightly glaucous, eucamptodromous, narrowly elliptic to narrowly oblong, base acute, apex acute, margin revolute, adaxial and abaxial surfaces covered by very short trichomes, in abaxial surface more concentrated on the veins; secondary veins 2 on each side, departing along the midvein, inconspicuous on the adaxial surface; domatia absent. Synflorescence frondose, pleiobotryum spiciform type, congested; accessory branches absent, principal florescence truncate, botryum spiciform type, paraclades 4–20 mm long; bracts similar to the vegetative leaves in shape and size; flowers subsessile, hypopodium 0.1–0.2 mm long, epipodium 0.3–0.5 mm long; prophylls 3.6–4.2 × 1.5–1.8 mm, surpassing the middle of the floral tube, sometimes surpassing the apex of the floral tube, elliptic, unkeeled, base acute, apex acute, sometimes reflexed, margin plane; floral tube 3.5–4 mm long, greenish, funnel-shaped, covered by very short stellate trichomes; sepals 1.8–2 mm long, wine-colored, triangular, slightly deflexed, not caudate, not ciliate; epicalyx segments 4–6.5 mm long, narrow-cylindrical, spreading, covered by very short stellate trichomes, longer than the sepals; corolla 1.5–2.5 cm in diam., lilac to rose, petals 7–8 × 4–4.5 mm, obovate, apex obtuse; stamens 18–24, free portion of the filaments 7–7.5  mm long, exserted, surpassing the sepals 2.5–3  mm; ovary ca. 1.5 × 2–2.1 mm, obovate, glabrous, style 10.5–11 mm long, ovules 13–20. Capsules ca. 3 × 3–3.5 mm, globose, glabrous; seeds 5–11, 1.8–2.5 × 1.2–2.1 mm. Distribution and Ecology (Fig. 13.60)  Diplusodon macrodon occurs on red latosol at the edge of dense cerrado in northeast of Goiás State and southeast of Tocantins State, at 395–750 m. Flowers from March to April; fruits registered in April. Specimens Examined: BRAZIL. Goiás,  Campos Belos, 13°07′23″ S, 46°44′63″ W, 9 Apr 1997 (fl), Cavalcanti et al. 2250 (CEN, MO, NY); Monte Alegre de Goiás, beira do rio Imbé, a cerca de 4 km da rodovia, próximo à ponte do rio Paranã, 22 Nov 1991 (st.), Vieira et al. 1183 (CEN); Nova Roma, estrada para São Domingos,

Descriptions

265

Fig. 13.58  Diplusodon macrodon. A. Branch with inflorescence. B. Section of internode. C. Leaf, adaxial surface. D. Leaf, abaxial surface. E. Prophyll, adaxial surface. F. Flower with prophyll, without petals. (A Fonseca et al. 2437, CEN; B–F Cavalcanti et al. 2250, CEN)

margem do rio Paranã, 13°31′42″ S, 46°49′50″ W, 24 Apr 2009 (fl, fr), Queiroz et al. 14128 (CEN, HUEFS); Nova Roma, estrada de terra de Ouro Minas para Teresina de Goiás, 13°35′37″ S, 47°00′47″ W, 1 Apr 2008 (fl, fr), Bringel et al. 419 (CEN); Posse, estrada de chão Posse–Iaciara, 14°03′56″ S, 46°35′17″ W, 7 Mar 2002 (fl), Fonseca et al. 2437 (CEN, IBGE, UFG n.v., P); May 1840, Gardner 4136 bis (BR, G (3), K, OXF, P (2), NY, R, W); without locality, Gardner 3155b (BR); Gardner 3725 (BM, G [2], K, LE n.v., NY, OXF, P, W); 1841, Gardner 4138 (K); 1842 (fr);

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Gardner s.n. (NY [2]). Tocantins, Aurora do Tocantins, 12°38′75″ S, 46°22′43″ W, 8 Apr 1997 (fr), Cavalcanti et al. 2246 (CEN, NY). Diplusodon macrodon is a much-branched shrub with branches covered by dense indumentum with short trichomes oriented in different directions. The specific epithet of D. macrodon alludes to the principal diagnostic feature of this species, the epicalyx segments 4–6.5  mm long. Other characteristics are the short trichomes covering all parts of the flowers and leaves and the narrowly elliptic to narrowly oblong leaves. Diplusodon macrodon resembles D. stellatus, a species of Chapada dos Veadeiros, Goiás, because of its narrow leaves and long epicalyx segments. However, Diplusodon stellatus has dense tufts of stellate trichomes covering the entire plant that confers a flaky aspect to the surface. Diplusodon stellatus also has flowers with shorter epicalyx segments (3–4 mm long) and an androecium of 12 stamens. The habit of D. stellatus is usually caespitose with several branches coming out from a xylopodium, and the plant is canescent. Koehne (1877) described Diplusodon macrodon and cited the syntypes Gardner 3725, Gardner 4135, and Gardner 4136 bis (quoted by the author as a duplicate number). The gathering Gardner 4135 at B had the species name and the signature of Koehne but was destroyed (F neg. nº 017926). Lourteig (1989) cited a specimen of Gardner 4135 at P as the holotype, which is here considered as the lectotype (ICBN Art. 9.17, ex. 14). There are three specimens of Gardner 4135 at P; two of these specimens have original labels and have the initials of Koehne. 47. Diplusodon marginatus Pohl, Flora 10: 150. 1827. Type: Brazil. Goiás: In montosis pascuis ad rio São Bartolomeu et in Serra de Montes Claros, Dec 1819, J. B. E. Pohl 990 = D 944 (lectotype, W [0053422], designated by Lourteig, 1989: 233; isolectotypes, BR, G, K [2], M, OXF). Figs. 13.59 A–H, 13.60 Subshrubs 0.3–1 m tall, erect, unbranched, branched just at inflorescence, with a xylopodium, glabrous, not dark-brown when dry. Upper branches rust to wine-colored, quadrangular to flattened, unwinged, internodes 2–7 mm long. Leaves decussate, sometimes imbricate, erect, amplexicaulous; blades 35–120  ×  20–75  mm, olive-green, glaucous, chartaceous, not discolorous, acrodromous-basal-supranumerary, elliptic-lanceolate or lanceolate to ovate, base retuse to amplexicaulous, apex acute or rarely obtuse, margin plane with yellowish cast when fresh, winecolored to purple when dry; secondary veins 5–10 on each side, 4–5 of them departing from the base of the midvein, wine-colored or purple; domatia absent. Synflorescence frondose-bracteose, diplobotryum type, less often simple botryum, congested; accessory branches absent, principal florescence 10–25  mm long, botryum type, paraclades 6–25 mm long; bracts 13–55 × 10–35 mm, ovate-lanceolate to lanceolate, margin plane, wine-colored to purple; flowers long-­pedicellate, hypopodium 15–45  mm long, epipodium 0.5–1.5  mm long; prophylls 9.5– 13 × 6–9 mm, reaching the apex or slightly more than to the middle of the floral

Descriptions

267

Fig. 13.59  Diplusodon marginatus. A. Vegetative branch. B. Section of internode (fistulous). C. Branch with inflorescence. D. Prophyll, adaxial surface. E. Floral bud. F. Flower with prophyll, without petals, showing lack of epicalyx. G. Floral tube outer surface showing rudiment of epicalyx segments with cilia. H. Ovary. (A, E–F Cavalcanti et al. 1000, CEN; B–D, G–H Walter et al. 789, CEN)

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tube, ovate-lanceolate to orbicular, concave, unkeeled, base and apex obtuse, margin plane, wine-colored to purple; floral tube 10–11 mm long, green to purple, funnel-shaped; sepals 4–5  mm long, triangular, not caudate, spreading to deflexed, glabrous, rarely with 1–2 cilia at the apex; epicalyx segments absent or when present reduced to mere protuberances and rounded, glabrous or rarely with long trichomes; corolla 6–7 cm in diam., purple to clear purple, petals 23–30 × 13–17 mm, elliptic, apex and base obtuse; stamens (18–)23–33, free portion of the filaments 12–14 mm long, exserted, surpassing the sepals 2–3 mm; ovary 3.5–4 × ca. 4 mm, globose, glabrous, style purple, 17–24  mm long, ovules 25–30. Capsules 9 × 7–7.5 mm, globose, glabrous; seeds 36–42, 2.5–4 × 2.5–4 mm. Distribution and Ecology (Fig. 13.60)  Diplusodon marginatus occurs in Distrito Federal to the extreme north of Goiás, in Chapada dos Veadeiros, in areas of the campo sujo and campo cerrado, on sandstone, sandy-clay, latosol with quartz, at 500–1100 m. Flowers registered from July to April; fruits registered from January to March and from July to October. Specimens Examined: BRAZIL. Distrito Federal,  Brasília, estrada vicinal para o rio Belchior, cerca de 2 km ao norte, 15°54′27″ S, 48°11′28″ W, 10 Mar 2003 (fl, fr), Pereira-Silva et  al. 7285 (CEN, MO); estrada para Santo Antônio do Descoberto, saindo da BR-060, 15°55′12″ S, 48°11′15″ W, 11 Sep 2012 (fr), Faria-Jr. & Zanatta 2766 (CEN, UB); fazenda Salvador próxima a divisa DF/GO, Núcleo Rural Boa Esperança, 15°48′32″ S, 48°14′17″ W, 10 Oct 2005 (fl), Amaral et al. 11 (CEN); Gama, na rodovia para o rio Corumbá, 16 Oct 1963 (fl, fr), Maguire et al. 57090 (NY, P [2]); Granja Samambaia, 20 Oct 1964 (fl), Heringer 9913 (UB); 120  km sul de Brasília, 15 Jan 1967, Heringer 11313 (UB); margem da rodovia Brasília-Anápolis, 8 Dec 1965 (fl), Belém 1903 (CEPEC, UB); Without reference of municipal district: estrada rio Brechó, 22 Nov 1979 (fl), Pereira-Silva & Vieira 40 (CEN, MO, NY); DF-290, km 8, 8 Feb 1996 (fr), Munhoz et al. 332 (IBGE); Divisa com o Distrito Federal, Sete Curvas-Engenho das Lages, BR 060, 21 Sep 2003 (fl), Pastore & Suganuma 700 (CEN); a 10 km da divisa com Goiás, ao lado da BR 060, 7 Jul 2004 (fl), Pastore 1044 (CEN); divisa com Goiás, rodovia para Santo Antonio do Descoberto, 30 Jan 2004 (fr), Pastore et  al. 751 (CEN). Goiás, Corumbá de Goiás, Aparecido, ca. 50 km W de Brasília, 29 Nov 1965 (fl), Irwin et al. 10724 (IAN, K, MO, NY, P, UB); Entre Anápolis e Corumbá de Goiás, 2 Apr 1958 (fl), Lima 58–3002 (IPA, K, RB); rio Corumbá, 15 Nov 1967 (fl), Heringer 11574 (NY, P [2]); Luziânia, cerca de 800 a oeste da ponte do rio alagado, direção BR-060, 16°12′21″ S, 48°10′25″ W, 6 Nov 2002, Pereira-Silva et  al. 6843 (CEN, MO); fazenda do Sr. José Rodrigues, próximo à sede, 16°19′49″ S, 48°12′49″ W, 9 Apr 2003 (bud), Pereira-Silva et al. 7493 (CEN, MO); estrada vicinal de acesso ao rio Descoberto (captação Saneago), km 2, 16°17′17″ S, 48°10′43″ W, 2 Jul 2003 (fr), Pereira-Silva et al. 7940 (CEN, MO); Minaçu, estrada nova Minaçu-Serra da Mesa, 24 km do asfalto, 13° 40′ S, 48° 12′ W, 11 Oct 1991 (fl), Cavalcanti et al. 1000 (CEN, K, NY, SPF); a 8.9 km do portão norte do canteiro de obras, 13°29′ S, 48°24′ W, 11 Mar 1992 (fr), Cavalcanti et  al. 1150 (CEN, RB, SPF); estrada de terra Minaçu-Serra da Mesa, ca. 5  km da entrada da UHE Serra da Mesa, 13°40′ S,

Descriptions

269

Fig. 13.60  Geographic distribution of Diplusodon leucocalycinus, D. longipes, D. macrodon, and D. marginatus

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48°14′ W, 20 Nov 1991 (fl), Walter et al. 789 (CEN, MO, NY); cerca de 25 km de Minaçu, 13°49′ S, 48°26′ W, 21 Mar 1996 (fl, fr), Walter et al. 3283 (CEN, MO, NY); Santo Antônio do Descoberto, 29 Nov 1965 (fl), Cobra & Sucre 308 (UB, P); 28 Oct 1966 (fl), Duarte 9971 (HB, MBM); 25 Oct 1979 (fl), Heringer et al. 2613 (IBGE, K, MO, P [2], US); 26 Feb 1980 (fl, fr), Heringer et al. 3466 (IBGE, NY); 25 Feb 1985 (fr), Pereira 1253 (IBGE, RB, SPF). Without reference of municipal district: entre Guariroba e Siriaco, 11 Jul 1895 (fl), Glaziou 21420 (BR, K, P [2], R); Pohl s.n. (G, M, OXF). Diplusodon marginatus is an ornamental, unbranched subshrub with broad (35– 120  ×  20–75  mm), glaucous leaves with a purple margin, and purple corolla of 6–7 cm in diam. In addition to these characteristics, this species has long pedicels (1.5–4.5 cm long), similar to those of D. longipes, and it lacks an epicalyx in the floral tube. In some of the collections examined, the epicalyx segments are merely small protuberances, either glabrous or with trichomes on the apex. Diplusodon longipes, the morphologically closest species, differs by its long, narrow floral tube, epicalyx segments ca. 3 mm long, and androecium with 13–19 stamens. Lourteig (1989) cited Pohl 990 = D 244 (W) as the holotype of Diplusodon marginatus, which is here treated as lectotypification (ICBN Art. 9.17, ex. 14). It was verified that other herbaria have specimens of Diplusodon marginatus which the label indicates Pohl s.n., but without locality, and some with the indication that the material came from W. Since duplicates of Pohl’s collections were sent to other herbaria without the information from his field diary (see “Notes about Diplusodon nomenclature and typifications”), and most lack a number, the specimens at BR, G, K, M, and OXF are regarded as duplicates of Pohl 990 = D 944. 48. Diplusodon mattogrossensis T.B.Cavalc., Novon 8(4): 342, fig. 5 h–l. 1998. Type: Brazil. Mato Grosso: ca. 270 km norte de Nova Xavantina, 8 May 1968 (fl), J. A. Ratter, R. R. Santos, R. Souza, A. Ferreira 1361 (holotype, UB; isotypes, K, NY [00468429]). Figs. 13.61 A–D, 13.65 Subshrubs 0.8–1.5 m tall, erect, branched just at inflorescence, with a xylopodium, not dark-brown when dry. Upper branches cylindrical to subquadrangular, unwinged, densely covered with long, white trichomes; internodes 2–6  cm long. Leaves decussate, not imbricate, subsessile; petioles 1–3  mm long; blades 40–60 × 20–45 mm, coriaceous, not discolorous, acrodromous-basal-supranumerary, ovate-­lanceolate, base cordate to subcordate to obtuse, apex obtuse, acuminate, margin plane, blades villous due to the bulbous trichome base; secondary veins 4–7 on each side, 2–4 of them departing from the base of the midvein, prominent on the abaxial surface; domatia absent. Synflorescence frondose, botryum to diplobotryum type; accessory branches absent; flowers pedicellate, hypopodium 4–4.5 mm long, epipodium 1.5–2 mm long; prophylls 16–17 × 7–8 mm, surpassing the apex of the floral tube, elliptic-lanceolate, unkeeled, base obtuse to acute, apex acute, margin plane, pilose; floral tube 8–9 mm long, funnel-shaped, densely covered with long trichomes; sepals 5.5–6  mm long, triangular, not caudate, not deflexed, densely

Descriptions

271

covered with long trichomes; epicalyx segments 8.5–9 mm long, cylindrical, erect, surpassing the sepals; corolla 5.5–6 cm in diam., petals 23–25 mm long, obovate, apex obtuse; stamens 23–32, free portion of the filaments ca. 10 mm long, included; ovary 3–3.5 × 4.5–5 mm, obovate-depressed, glabrous, style ca. 17 mm long, ovules ca. 36. Fruits not seen. Distribution and Ecology (Fig. 13.65)  Diplusodon mattogrossensis is known only from a few collections from northeastern Mato Grosso, growing at 500 m. Flowers registered in May. Specimens Examined: BRAZIL. Mato Grosso,  estrada Xavantina-Cachimbo, 200 km de Xavantina, 29 May 1966 (fl), Hunt & Ramos 5661 (K, UB); ca. 270 km N de Xavantina, Lagoa de Leo, 12°54′ S, 51°52′ W, 29 May 1968 (fl), Ratter et al. 1574 (K, MO, NY); Serra do Roncador, ca. 160 km N de Xavantina, 27 May 1966 (fl), Irwin et al. 16139 (NY). Diplusodon mattogrossensis is represented by branched subshrubs with branches densely covered with long and white trichomes. The leaves are petiolate, with large villous and ovate-lanceolate blades, of 40–60 mm long, with cordate to subcordate base. The flowers are pedicellate, with broad prophylls (16–17  ×  7–8  mm),

Fig. 13.61  Diplusodon mattogrossensis. A. Leaf, abaxial surface. B. Detail of leaf indumentum. C. Section of internode. D. Flower with prophyll, without petals. (A–D Ratter et al. 1361, UB)

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surpassing the apex of the floral tube, floral tube with long and erect epicalyx segments surpassing the sepals, and androecium with 23–32 stamens. Diplusodon mattogrossensis is similar to D. villosissimus in the vegetative aspect and by the leaf shape and the dense indumentum. Diplusodon mattogrossensis differs by its much larger flowers and 23–32 stamens. 49. Diplusodon micromerus T.B.Cavalc., Phytotaxa 38: 33, fig. 3. 2011. Type: Brazil. Goiás: Cavalcante, caminho para cachoeira Santa Bárbara, 20 Apr 2003 (fl), J.  F. B.  Pastore, A.S. Rodrigues & E.  Suganuma 466 (holotype, CEN [00053418]; isotypes, K, MO, SPF, W). Figs. 13.62 A–D, 13.65 Shrubs ca. 0.35 m tall, erect, much-branched, congested, not dark-brown when dry. Upper branches cylindrical, unwinged, dense hirsute, whitish, internodes 0.5– 0.8 cm long. Leaves decussate, not imbricate, suberect, strongly congested, sessile; blades 4–6 × 3–6 mm, canescent, coriaceous, discolorous, acrodromous supranumerary, orbicular to oblong-lanceolate, base cordate, sometimes obtuse, apex obtuse, margin strongly revolute; secondary veins 3–4 on each side, adaxial surface hirsute-sericeous, green-grayish, with deeply sulcate veins, abaxial surface hirsutesericeous, white, with prominent veins; domatia absent. Synflorescence frondose, triplobotryum to pleiobotryum type, congested; accessory branches absent, principal florescence truncate, paraclades 3–10 cm long; bracts similar to the vegetative leaves in shape and size; flowers sessile, hypopodium and epipodium absent; prophylls 5–5.5 × 3.8–3.9 mm, reaching to the apex of the floral tube, obovate, unkeeled, margin plane; floral tube 4–4.5 mm long, funnel-shaped, densely covered with silvery trichomes, turning the floral tube shiny; sepals 2–2.5 mm long, triangular, not caudate, not deflexed, densely covered with silvery trichomes; epicalyx segments 0–0.5 mm long, shorter than the sepals, cylindrical, spreading, sericeous; corolla ca. 1.5 cm in diam.; lilac, petals 12–14 × 6.5–9 mm, elliptic, apex obtuse; stamens 18, free portion of the filaments ca. 6  mm long, shortly exserted; ovary oboval, glabrous, ca. 1.9 × 1.8 mm, style ca. 11 mm long, ovules ca. 13. Fruits not seen. Distribution and Ecology (Fig.  13.65)  Diplusodon micromerus is only known from the type locality, in cerrado and campo sujo, ca. 1080 m elevation. Flowers registered from January to April, fruits from February to April. Specimens Examined: BRAZIL.  Goiás,  Cavalcante, Comunidade Kalunga Engenho II, próximo ao córrego Quebra Coco, 13°34′57″ S, 47°28′21″ W, 16 Jan 2010 (fl), Faria-Jr. et al. 731 (CEN); Cavalcante, caminho para o povoado Kalunga, 3 Feb 2004 (fl, fr), Pastore et  al. 840 (CEN); Cavalcante, estrada para Prata (Kalungas), 13°22′24.2″ S, 47°41′9.1″ W, 14 Apr 2004 (fl, fr), Fonseca et al. 5099 (CEN, HRB n.v., IBGE, RB); Cavalcante, 23.8 km após Cavalcante, em direção ao Engenho Vão do Moleque II, 2.3  km após a bifurcação sentido cachoeira Santa Bárbara, 15°36′37″ S, 47°28′27″ W, Amaral-Santos et al. 3666 (CEN). The specific epithet was chosen to emphasize the diminutive floral parts of this species. The shrubs are strongly branched and have congested, discolorous, small

Descriptions

273

Fig. 13.62  Diplusodon micromerus. A. Branch. B. Leaf, adaxial surface. C. Leaf, abaxial surface. D. Flower with prophylls, without petals. (A–D Pastore et al. 466, CEN)

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leaves with a revolute margin and white abaxial surface. The flowers are small, congested on the branches, hidden by the bracts, and the floral tube and sepals are densely white hirsute. The epicalyx segments are absent or inconspicuous, and the prophylls reach the apex of the floral tube. Diplusodon micromerus is closely allied to D. cordifolius, but the latter differs by its oblong to lanceolate leaf shape, eucamptodromous venation, with 2–3 lateral veins and evident domatia, and also by its epicalyx segments 0.5–2 mm long, which have sparse, spreading trichomes. 50. Diplusodon microphyllus Pohl, Flora 10: 151. 1827. Type: Brazil. Minas Gerais: Ouro Preto, Nov 1821–Jan 1822, J.  B. E.  Pohl 5792 (lectotype, W [0069492], designated by Lourteig, 1989: 224; isolectotype, G). Figs. 13.63 A–E, 13.65 Diplusodon buxifolius (Cham. & Schltdl.) DC. var. naudinii Lourteig, Sellowia 16: 147. 1964. Type: Brazil. Minas Gerais: 1842, Pixis 38a (holotype, P [01901912]). Friedlandia hirtella Cham. & Schltdl., Linnaea 2: 352. 1827. Diplusodon hirtellus (Cham. & Schltdl.) G.Don, Gen. Syst. 2: 719, 1832. Type: Brasilia aequinoctialis. F. Sellow s.n. (lectotype, HAL [0107570], here designated; isolectotypes, B, HAL [0098635], K, LE n.v., W [2]). Friedlandia villosula Mart. ex DC., Prodr. 3: 94c, 1828, pro syn. Subshrubs 0.4–1 m tall, dark-brown when dry. Upper branches quadrangulate, weakly winged, with long, sparse trichomes, internodes 0.5–1.2 cm long. Leaves decussate, spreading to suberect, petiolate; petioles 1–1.5  mm long; blades 5–17 × 3–11 mm, eucamptodromous, membranaceous to chartaceous, slightly discolorous, dark-brown when dry, ovate to lanceolate, base acute, apex acute or rarely obtuse, margin slightly revolute, adaxial surface glabrous, abaxial surface olivegreen, with sparse trichomes, veins scarcely prominent; secondary veins 2–3 on each side, departing along the midvein; domatia absent. Synflorescence congested, frondose, diplobotryum to pleiobotryum type, accessory branches absent, principal florescence botryum type, 10–35 mm long, paraclades 7–30 mm long, bracts similar to the vegetative leaves, slightly smaller; flowers subsessile, hypopodium 0.7– 1.5 mm long, epipodium 0.5–0.7 mm long; prophylls 4.5–6 × 1.5–3 mm, reaching the apex of the floral tube or slightly less, elliptic, unkeeled, base and apex acute, margin plane, ciliate; floral tube 4–5  mm long, funnel-shaped, sparsely pilose; sepals 2–3 mm long, triangular, not caudate, not deflexed, almost glabrous, ciliate; epicalyx segments 1.5–2  mm long, cylindrical, curved, deflexed to spreading, shorter than the sepals, pilose; corolla 2–2.5  mm long, rose to purple, petals 10–13 × 6–9.5 mm, elliptic to obovate, apex obtuse; stamens 12, free portion of the filaments 8–9 mm long, exserted; ovary 1.5–2 × 2–2.5 mm, rhombic, glabrous, style 14–15 mm long, ovules 8–18. Capsules ca. 3.5 × 3 mm, oblong, glabrous; seeds ca. 7, 2–2.2 × 1.5–1.6 mm.

Descriptions

275

Fig. 13.63  Diplusodon microphyllus. A. Vegetative branch. B. Section of internode. C. Leaf, abaxial surface with detail of indumentum on abaxial surface of leaf. D. Prophyll, adaxial surface. E. Flower with prophylls, without petals. (A, B, D, E Badini s.n. (OUPR 4792); C Irwin et al. 29628, NY)

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Distribution and Ecology (Fig. 13.65)  Diplusodon microphyllus is endemic to the south of Cadeia do Espinhaço, in Minas Gerais State, where it grows in campo rupestre and open fields. Flowers from February to June; fruits from March to June. Specimens Examined: BRAZIL.  Minas Gerais,  Alegria, Mar 1943 (fl, fr), Badini s.n. (OUPR4261); Belo Horizonte, Pampulha, 23 Mar 1963, Pereira 7285 (P); Belo Horizonte, Serra do Curral, 29 Apr 1945 (fl), Williams & Assis 6745 (F, GH, SP); Belo Horizonte, Estação Ecológica da UFMG, 5 Mar 1991 (fl), TameirãoNeto 419 (CEN, BHCB); Belo Horizonte, Estação Ecológica da UFMG, 23 May 1990 (fl), Bacariça et al. 28 (CEN, BHCB); Bento Pires, próximo a Belo Horizonte, Mar 1945 (fl), William & Assis 6127 (R); Bocaina de Minas, Serra do Itatiaia, s.d., Badini s.n. (CEN, OUPR 4737); Congonhas do Campo, Stephan s.n. (BR); Congonhas do Campo, 20 May 1989 (fl), Hatschbach & Nicolack 52956 (MBM, P); Congonhas do Norte, Retiro do Barbado, morro à esquerda do rio Preto, 22 Apr 1982 (fl), Amaral et al. (CFSC) 8403 (SPF); Gouveia, estrada de terra para a fazenda do Sr. Everaldo, 14 Apr 1987 (fl), Zappi et al. (CFCR) 10472 (SPF); Itabirito, Pico do Itabirito, 21 May 1994 (fl), Teixeira s.n. (BHCB 25023, CEN 18967); Itabirito, Serra da Capanema, s.d., Badini s.n. (CEN, OUPR 1776); Capanema, 1 Mar 1943 (fl, fr), Badini s.n. (OUPR 4250); Itatiaçu, rodovia Fernão Dias, 19 Apr 1978, Hatschbach et al. 41195 (MBM n.v., P); Lagoa Santa, Caxoeirinha, 24 Apr 1866, Glaziou s.n. (P); Mariana, Fazenda do Macaquinho, s.d., Badini s.n. (CEN, OUPR 4792); Mariana, ca. 3 km N de Mariana, 2 Feb 1971, Irwin et al. 29628 (NY, P); Mariana, Feb 1892 (fl), Schwacke 7733 (RB); Mariana, Reserva Mineral da CVRD, estrada Santa Bárbara-Mariana, 25 Apr 1998 (fl, fr), Meira & Neto 107 (CEN, VIC); Santa Rita Durão-Mariana, 20 Mar 1974 (fl, fr), Badini s.n. (OUPR 3854); Santa Rita Durão, 10 Apr 1973 (fl, fr), Badini s.n. (CEN, OUPR 4778); Santa Rita Durão, Vauthier 423 (P); Santa Rita Durão, 20 Apr 1957, Pereira 3034 (P); Santa Rita Durão, May 1943, Badini s.n. (CEN, OUPR 4767); Ouro Preto, Camarinhas, 20°22′08.7″ S, 43°30′30.5″ W, 20 Feb 2002 (fl), Souza et al. 28058 (CEN, ESA); Cachoeira do Campo, s.d., Badini s.n. (CEN, OUPR 1811); Campo Grande, May 1943 (fl, fr), Badini s.n. (CEN, OUPR 4762); Batatais, May 1942 (fl), Badini s.n. (CEN, OUPR 4771); Mariana, Santa Rita Durão, 21 May 1974 (fl), Badini s.n. (OUPR 21668); Mesquita, May 1943 (fl), Badini s.n. (CEN, OUPR 4775); Nova Lima, Serra da Mutuca, 3 Jun 1945 (fl), Williams & Assis 7241 (MO, NY, SP); Claussen 11a (BR); Claussen 21a (BR); Ackerman s.n. (BR); Ouro Preto, subida para o Morro de São Sebastião, 20°38′15″ S, 44°50′28″ W, 27 Feb 2000 (fl), Cavalcanti et al. 2581 (CEN, MO, NY); Ouro Preto, Serra de Ouro Preto, 25 Jun 1884 (fl), Glaziou 14700 (BR, LE n.v., MO, P, R); Ouro Preto, Serra do Ouro Preto, Feb 1892 (fl), Ule 2563 (R); Ouro Preto, Serra do Ouro Preto, s.d., Badini s.n. (CEN, OUPR 4759); Ouro Preto, Serra do Itacolomi, 6 Apr 1974 (fl), Badini s.n. (CEN, OUPR 4760); Ouro Preto, Serra de Antonio Pereira, s.d., Badini s.n. (CEN [2], OUPR 4765); Itacolomi, Aug 1888, Bayern s.n. (M); Fazenda dos Macaquinhos, s.d., Badini s.n. (OUPR 21649); Ouro Preto, Lund s.n. (P); Ouro Preto, estrada nova do Itacolomi, 6 Apr 1974 (fl), Lisboa s.n. (CEN 38139, OUPR 1778); Ouro Preto, Morro do Cruzeiro, 20 Mar 1970 (fl), Emygdio & Lisboa 2882 (R); Ouro Preto,

Descriptions

277

Morro do Cruzeiro de Santa Efigênia, May 1943, Badini s.n. (CEN, OUPR 4770); Ouro Preto, Morro da Forca, May 1943 (fl, fr), Badini s.n. (CEN, OUPR 4761); Ouro Preto, Cachoeira das Andorinhas, 23 May 1979, Mautone et al. 691 (P, RB); Ouro Preto, 6 Mar 1982 (fl), Grandi 904 (CEN, BHCB); Santa Bárbara, base da Serra do Caraça, s.d., Badini s.n. (CEN, OUPR 4790); Santa Bárbara, base da Serra do Caraça, 21 May 1974 (fl, fr), Badini s.n. (OUPR 21669); Santana de Pirapama, Fazenda Inhame (Serra Mineira), 21 Mar 1982 (fl), Pirani et al. (CFSC) 8014 (SPF); Serra do Cipó, 5 Apr 1958 (fl), Strang 81 (R); Serra do Cipó, 28 Apr 1892, Glaziou 19173 (K, P); Serra do Cipó, Strang 85 (P); Serra da Moeda, 5 Jun 1988 (fl, fr), Souza s.n. (BHCB 16071); without reference to municipal district and collector (SP 18801). Diplusodon microphyllus is a member of the Hirsutus complex (discussed under D. bahiensis). Diplusodon microphyllus is a canescent, much-branched subshrub with small, short-petiolate or rarely subsessile leaves that are ovate, glabrous with pilose veins on the adaxial surface which turns dark-brown when dry. In addition, the flowers have a sparsely pilose floral tube, epicalyx segments erect to spreading and androecium with 12 stamens. Lourteig (1964) established Diplusodon buxifolius var. naudinii, which differed from the typical variety by the calyx pubescence and ciliate prophylls and leaves. The holotype of the variety (Pixis 38a) was examined, as well as the cited paratype (Williams & Assis 6745), and these specimens are here considered D. microphyllus. In Williams & Assis 6745, there are densely pilose white buds and leaves with an obtuse base and prominent veins with domatia, characteristics that do not occur in D. buxifolius. From southern Minas Gerais to Belo Horizonte, D. hirsutus occurs sympatrically with D. microphyllus and D. buxifolius, sometimes making it difficult to distinguish the limits among these three species. Perhaps the species hybridize. Some specimens have trichomes on the flowers, but some do not. Therefore, it is not possible to establish an exact morphological type that characterizes var. naudinii, because various densities of indumentum exist between pilose and glabrous flowers. Further, there was no relationship between the presence and absence of trichomes and geographic distribution that would allow establishment of a local ecotype. For this reason, Diplusodon buxifolius var. naudinii has been synonymized under D. microphyllus Pohl. At the extremes of the range of variation of the leaves, Diplusodon microphyllus and D. hirsutus are well separated. Diplusodon microphyllus has smaller membranaceous leaves, with acute base and almost glabrous, while D. hirsutus has larger leaves, coriaceous, with obtuse base and hirsute. The original material of Diplusodon microphyllus was studied together with other material cited by Koehne (1877, 1903, Glaziou 14700, Ule 2562) and Lourteig (1989; Irwin et al. 29628, Riedel s.n., Sellow s.n.). The study of the original material and additional material showed that D. microphyllus can be difficult to separate from D. hirsutus. Both species have obovate to elliptic prophylls reaching the middle of the floral tube or slightly longer, and both species have spreading to deflexed epicalyx segments 3  mm long. The difference between the two species is based on a few

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characteristics. For example, Koehne (1903) distinguished Diplusodon microphyllus from D. hirsutus based on the acute leaf base of D. microphyllus, in contrast to the rounded leaf base of D. hirsutus. Lourteig (1989) separated the two species in her key based on leaf and sepal pubescence: glabrous adaxial leaf surface and glabrous or slightly pilose sepals in D. microphyllus versus pilose adaxial leaf surface and pilose sepals in D. hirsutus. Examination of specimens from southern Minas Gerais showed the two species to be very similar, with differences in the density of the indumentum. In some specimens the trichomes are long and sparse (corresponding with D. microphyllus), while in others the trichomes are very abundant and short lending a whitish cast to the plant corresponding with D. hirsutus and exemplified by some of the specimens seen by Koehne (1877, 1903) and Lourteig (1989), such as Glaziou 14677 and Hatschbach & Ahumada 31275. In addition to these observations, Diplusodon microphyllus and D. hirsutus are here maintained as distinct species based on results of molecular phylogenetic analyses (Cavalcanti et al. 2007), which suggest that the species are distinct. Diplusodon microphyllus was described by Pohl (1827a) without citation of specimens in the protologue. In a separate publication, Pohl (1827b) cited a single gathering of D. microphyllus collected in December and January near Vila Rica (today Ouro Preto), state of Minas Gerais, without mentioning a collection number. Lourteig (1989) indicated as holotype “Pohl s.n.” at W, which is considered here as the lectotype (ICBN Art. 9.17, ex. 14). However, the specimen at W with the same locality data and annotated by Pohl has a number, Pohl 5792. Although the type material of Friedlandia hirtella at B (Sellow s.n.), known today from a photograph, was destroyed, there is another Sellow s.n. at B that could have been a duplicate of the type and was probably examined by Chamisso and Schlechtendal in describing F. hirtella. On this sheet, there is an annotation as Diplusodon microphyllus in Koehne’s writing, but without the initials that he customarily used. Similarly scant information is present on the specimen at K, which was transferred from B. Another specimen of Sellow s.n., also originally from B, was found at W and was identified by Koehne as D. microphyllus, with “F. hirtella Cham. & Schltdl.” indicated in parenthesis. Identical data are available on a duplicate specimen in Saint Petersburg (LE), which I did not study directly. The specimen at HAL (0107570) has an original label annotated by Schlechtendal and is here designated as the lectotype of F. hirtella. 51. Diplusodon minasensis Lourteig, Bradea 5(19): 229. 1989. Type: Brazil. Minas Gerais: Datas, ligação BR-250 a Costa Sena, 18 Mar 1987 (fl), G. Hatschbach, M. Hatschbach, A. C. Cervi & J. Cordeiro 51067 (holotype, P [01901870]; isotypes, C, K, MBM, MEXU, MO, MU, US, S). Figs. 13.64 A–H, 13.65 Shrubs 1–2.2 m tall, erect, much-branched, with whitish indumentum, canescent, not dark-brown when dry. Upper branches cylindrical to subquadrangular, unwinged, hirsute, lateral branches erect, internodes 5–6  mm long. Leaves decussate, not imbricate, suberect, short-petiolate; petioles to 1 mm long; blades 5–11 × 2–5 mm,

Descriptions

279

coriaceous, not discolorous, eucamptodromous, narrowly elliptic, base acute, apex obtuse, margin revolute, abaxial surface hirsute, covered by whitish trichomes with bulbous base, adaxial surface verrucose due to the bulbous trichome bases; secondary veins 2–3 on each side of the midvein, prominent; domatia absent. Synflorescence frondose, diplobotryum to pleiobotryum type, congested; accessory branches absent, principal florescence 4–10 cm long, botryum type, paraclades 2–14 cm long, bracts similar to the vegetative leaves in shape and size; flowers subsessile, hypopodium 0.5–1  mm long, epipodium 0–0.5  mm long; prophylls 5–5.5  ×  ca. 3  mm, reaching the apex of the floral tube or sometimes surpassing it, obovate, unkeeled, base acute, apex obtuse, margin revolute, 3–5-lateral veined; floral tube 4–5 mm long, oblong, hirsute, with whitish trichomes with bulbous bases; sepals ca. 2 mm long, triangular, not caudate, not deflexed, covered by whitish trichomes; epicalyx segments ca. 1  mm long, shorter than the sepals, lanceolate, deflexed; corolla 2–2.5 cm in diam., clear rose to rose-magenta, petals 10–11 × ca. 6 mm, obovate, apex obtuse; stamens 6, free portion of the filaments 8–9 mm long, exserted, surpassing the sepals 4–5 mm; ovary 1.8–1.9 × 2.1–2.2 mm, obconic, glabrous, style 10–11 mm long, ovules 15–19. Capsules ca. 3 × 2.5 mm, oblong, glabrous; seeds ca. 3, ca. 2.8 × 2.5 mm. Distribution and Ecology (Fig. 13.65)  Diplusodon minasensis is endemic to the central portion of Cadeia do Espinhaço, in Minas Gerais State, and inhabits campos rupestres at 838–1333 m and rocky cerrado. Flowers from January to May; fruits registered from February to March (immature) and in July and September. Specimens Examined: BRAZIL. Minas Gerais,  Alvorada da Serra, Distrito de Itapanhoacanga, estrada para Cachoeira Campina, a 3  km de Itapanhoacanga, 18°47′54″ S, 43°26′36″ W, 24 Mai 2009 (fl), Menini-Neto et al. 714 (CEN, K, RB, SPF); Bocaiúvas, Parque Nacional das Sempre Vivas, caminho dos Campos de São Domingos e a Serra do Landi, passando pelo córrego do Lando, 17°54′07.5″ S, 43°46′22.6″ W, 29 Apr 2007 (fr), Almeida et al. 877 (BHCB); Congonhas do Norte, Serra do Cipó, Serra Talhada, fazenda Vereda do Cambota, alto da Serra do João Camilo, 6.8 km sudoeste da estrada Congonhas do Norte-Gouveia, entrada 3.7 km noroeste de Congonhas do Norte, 18°48′39″ S, 43°45′09″ W, 19 Jan 2004 (fl), Mello-Silva et al. 2382 (CEN, SPF); Congonhas do Norte, Serra Talhada, 18°48′39″ S, 43°45′09″ W, 19 Jan 2004 (fl), Lovo et al. 19 (SPF); Conselheiro Mata, 20.3 km depois do asfalto, 18°20′ S, 43°53′ W, 23 Sep 1994 (fr), Splett & Gröger 656 (UB); Diamantina, BR-367-Conselheiro Mata, 21 km, 18°18′ S, 43°53′ W, 16 Feb 1989 (fl), Cavalcanti et al. 220 (CEN, K, NY, SPF); Diamantina, 18 km E de Diamantina, 19 Mar 1970 (fl), Irwin et al. 27908 (MO, UB); Diamantina, Parque Estadual do Biribiri, Alto da Jacuba, 18°11′46″ S, 43°35′,14″ W, 27 Fev 2016, Faria 5376 (CEN, HDJF, RB, UB); Gouveia, estrada de terra para a fazenda do Sr. Everaldo, 14 Apr 1987 (fl), Menezes et al. (CFCR) 10617 (BHCB, K, NY n.v., SPF); Gouveia, estrada paralela aos moinhos de vento, entrando à direita sentido Gouveia-Curvelo, sítio do Mulato, área de extração da empresa mineradora Tracomal, 18°31′24″′ S, 43°54′,49″ W, 4 Mar 2016 (fr), Faria et al. 5410 (CEN, HDJF, RB, UB); Gouveia, Sítio do

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Fig. 13.64  Diplusodon minasensis. A. Branch with inflorescence. B. Leaf, abaxial surface. C. Leaf adaxial surface indumentum. D. Floral bud. E. Prophyll, adaxial surface. F. Flower with prophylls, without petals. G–H. Trichomes of leaves. (A Menezes et al. [CFCR] 10617, SPF; B–H Cavalcanti et al. 220, SPF)

Descriptions

281

Fig. 13.65  Geographic distribution of Diplusodon mattogrossensis, D. micromerus, D. microphyllus, and D. minasensis

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Mulato, entrando na estrada paralela aos moinhos de vento, área da mineradora Tracomal, 18°31′24″ S, 43°54′,49″ W, 16 Mar 2016 (fl, fr), Faria & Otoni 5510 (CEN, HDJF, UB); Monjolos, mina do Geraldinho, próximo à mineradora Tracomal, 18°30′ S, 43°54′29″ W, 5 Jul 2016 (fr), Faria et al. 6204 (CEN, HDJF, UB); Serro, Pedra do Cruzeiro, 18°24′49′ S, 43°16′22″ W, 28 Mar 2001 (fl), Romero & Nakajima 6014 (HUFU). Diplusodon minasensis are much-branched, tall shrubs, characterized by its congested, elliptic-narrow leaves (to 1 cm long) that are covered with abundant, slightly whitish indumentum and also with strongly revolute margins and acute base. The flowers are small (ca. 2.5 cm in diam.) with a hirsute floral tube, very short, deflexed epicalyx segments (ca. 1 mm long), and the androecium has 6 stamens. Vegetatively, D. minasensis resembles D. cordifolius, from Chapada dos Veadeiros, in Goiás State, because both have congested branches, with small leaves that have revolute margins. Diplusodon cordifolius can be distinguished because it has a cordate leaf base, suberect epicalyx segments, and an androecium with 18 stamens. In addition to D. minasensis in Minas Gerais, very few species in the genus have an androecium with 6 stamens, which is the smallest number of stamens in the genus. 52. Diplusodon mononeuros Pilg., Repert. Sp. Nov. Regni Veg. 41: 221. 1937. Type: Brazil. Minas Gerais: Diamantina, Conselheiro Mata, Jun 1934 (fr), A. C. Brade 13405 (holotype, RB [0054671]; isotype, fragm. P). Figs. 13.66 A–D, 13.70 Shrubs 1.5–3  m tall, erect, much-branched, not dark-brown when dry. Upper branches cylindrical to subquadrangular, unwinged, indumentum inconspicuous, minutely pilose, internodes 5–7 cm long. Leaves decussate to verticillate, not imbricate, spreading to suberect, subsessile; petioles 0.5–1  mm long; blades 3–12 × 0.7–3 mm, chartaceous, slightly discolorous, hyphodromous, linear-spathulate, base acute to attenuate, apex obtuse, margin revolute, blades glabrous; lateral veins absent; domatia absent. Synflorescence frondose, pleiobotryum type; accessory branches absent, paraclades 4–16 cm long; bracts the same size and shape as the leaves; flowers pedicellate, hypopodium 2.5–4  mm long, epipodium 0.5  mm long, minutely pilose; prophylls 3.5–6 mm long, reaching or slightly exceeding the middle of the floral tube, subspathulate, unkeeled, base acute, apex obtuse, margin plane, glabrous, spreading to erect, when erect, rarely somewhat ciliate, inserted in the upper third of the pedicel; floral tube 3–4.5 mm long, funnel-­shaped, glabrous; sepals 2.5–2.8 mm long, triangular, not caudate, not deflexed, not ciliate; epicalyx segments 2.6–2.8 mm long, cylindrical, spreading, shorter than the sepals, glabrous; corolla ca. 2.5 cm in diam., light rose to lilac, petals 9–10 × 6–7 mm, obovate, apex obtuse; stamens 12, free portion of the filaments 8–8.5 mm long, exserted; ovary ca. 1.5 × 2 mm, obconic, glabrous, style 13–14 mm long, ovules ca. 20. Capsules 3.5– 3.7 × 3.9–4 mm, globose, glabrous; seeds 4–6, 2.6–2.9 × 1.9–2.1 mm. Distribution and Ecology (Fig. 13.70)  Diplusodon mononeuros grows on a hillside in disturbed woods along a river margin, on sandstone soils at about 1000 m, in

Descriptions

283

Fig. 13.66  Diplusodon mononeuros. A. Branch with inflorescence. B. Leaf, abaxial surface. C. Prophyll, adaxial surface. D. Flower with prophylls, without petals. (A–D Cavalcanti et  al. 3125, CEN)

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Cadeia do Espinhaço. Flowers registered in January and October; fruits in January and June. Specimens Examined: BRAZIL. Minas Gerais,  Conselheiro Mata, estrada de terra Conselheiro Mata-Diamantina, cerca de 1 km de Conselheiro Mata, 18°17′5″ S, 43°58′26″ W, 19 Jan 2003 (fl, fr), Cavalcanti et  al. 3125 (CEN [2], MO); Conselheiro Mata, estrada para o Telésforo, morro próximo a cidade, 21 Out 2007 (fl), Rosa et al. 950 (HUFU). Diplusodon mononeuros is represented by tall and much-branched shrubs. The species is characterized by its glabrous and linear-spathulate leaves of acute base and revolute margin. The flowers are pedicellate (pedicel 3–4 mm long), with narrow and subspathulate prophylls reaching or slightly exceeding the middle of the floral tube, epicalyx segments spreading and shorter than the sepals, and androecium with 12 stamens. The habit of Diplusodon mononeuros resembles that of D. uninervius, except that the former does not become black when dry. Diplusodon uninervius differs also by its 3–5-whorled leaves with dense indumentum on the midvein on abaxial surface. 53. Diplusodon myrsinites Mart. ex DC., Prodr. 3: 94b. 1828. Friedlandia myrsinites Mart. ex DC., Prodr. 3: 94b, 1828, pro syn. Type: Brazil. Minas Gerais: in campis prope rio Sapucaí, C. F. P. Martius s.n. (lectotype, M [0146269], designated by Lourteig 1989: 215; isolectotype, fragm. G). Figs. 13.67 A–K, 13.70 Diplusodon myrsinites DC. var. cribellinus Koehne, Bot. Jahrb. Syst. 29: 165. 1900. Type: Brazil. Minas Gerais: Ouro Preto, Serra de Ouro Preto, V. M. Silveira 938 (holotype, B, destroyed; isotypes, not found). Neotype: Brazil. Minas Gerais: Serra do Pião, entre 1350–1450 m, 14 May 1970, D. Sucre 6819 (RB00209550), here designated. Diplusodon myrsinites DC. var. cribellinus Koehne f. obtusifolius Koehne, Bot. Jahrb. Syst. 29: 165. 1900. Diplusodon koehnei Schwacke, Pl. Nov. Mineir. 2: 5. 1900. Type: Brazil. Minas Gerais: in campis elevatis in Serra de Ibitipoca, 2 Aug 1896, C. A. W. Schwacke 12410 (lectotype, RB [00209541], here designated; isolectotype, BHCB). Subshrubs 0.4–1 m tall, erect, much-branched, with a xylopodium, glabrous, not dark-brown when dry. Upper branches yellow to rust-colored, quadrangular, wing 0.2–0.5  mm wide, internodes 0.5–1.5  cm long. Leaves decussate, not imbricate, suberect to spreading, subsessile; petioles ca. 1  mm long; blades 4–22  ×  (3–)5– 12 mm, coriaceous, not discolorous, slightly glaucous, hyphodromous, obovate or elliptic to rarely narrowly oblong, base acute or slightly attenuate, apex obtuse to acute, margin plane, yellow- to wine-colored; secondary veins 1–2 on each side, departing along the midvein, inconspicuous on both surfaces of the blade; domatia absent. Synflorescence frondose, diplobotryum to pleiobotryum type, lax; accessory branches absent, principal florescence generally truncate, paraclades 4–30 cm

Descriptions

285

Fig. 13.67  Diplusodon myrsinites. A. Branch with inflorescence. B. Section of internode. C–H. Leaves, abaxial surface. I. Prophyll, adaxial surface. J. Floral bud. K. Flower with prophyll, without petals. (A–B, E–K Krieger et al. s.n., CESJ 24316; C Krieger 8621, SPF; D Carvalho et al. s.n., ESA 8319, SPF)

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long; bracts similar to the vegetative leaves in shape, decreasing in size toward the apex of the branch; flowers subsessile, hypopodium 1.8–2.3 mm long, epipodium 0.5–0.8 mm long; prophylls 2.5–3 × 0.8–1 mm, reaching from the middle to the apex of the floral tube, oblong, obovate to wide-elliptic, unkeeled, base acute, apex obtuse, margin plane, wine-colored; floral tube 4–4.5  mm long, green, funnelshaped, glabrous; sepals 2–2.5 mm long, wine-colored, triangular, not caudate, not deflexed; epicalyx segments (1–)1.5–2.5 mm long, cylindrical to narrowly lanceolate, spreading to suberect, rarely slightly deflexed, shorter than sepals or the same size; corolla 2–2.5 cm in diam., lilac to purple, petals 11–13 × 7–8 mm, obovate, apex obtuse to acute; stamens 12, free portion of the filaments 6–8  mm long, exserted; ovary 1.5–2 × 2–2.5 mm, rhombic, glabrous; style 13–18 mm long, ovules 14–23. Capsules 4.5–5  ×  3.7–4  mm, globose, glabrous; seeds (9–)12–17, 1.8–2 × 1.5–2 mm. Distribution and Ecology (Fig. 13.70)  Diplusodon myrsinites occurs most often in campos rupestres of the mountains of Cadeia do Espinhaço, in Minas Gerais State. Flowers from January to October; fruits from February to December. Specimens Examined: BRAZIL. Minas Gerais,  Bom Jardim, 10 Oct 1988 (fl, fr), Krieger et  al. s.n. (CEN 18564, CESJ 24316); Carrancas, 8 Dec 1983 (fr), Sugiyama & Chiea 423 (CEN, SP); Serra das Carrancas, 21°28′34″ S, 44°39′40″ W, 1 Jul 2011 (fl), Sobral et al. 14091 (HUFSJ n.v.; RB 2); Diamantina, Jul 1840 (fr), Gardner 4565 (K); Diamantina, estrada de terra para Barão do Guaicui, 18°22′24″ S, 43°43′36″ W, 9 Feb 2015 (fl), Cavalcanti et al. 3935 (CEN); Itumirim, Morro da Janela, Serra da Bocaina, 12 Jun 1987 (fl), Carvalho et al. s.n. (ESAL 7551, SPF); Itumirim, Serra da Bocaina, 29 Jun 1987 (fl), Kinoshita-Gouveia et al. 18983 (UEC n.v., VIC); Ingaí-Itumirim, Serra da Bocaina, 21 Jul 1987 (fl), Carvalho et al. s.n. (ESAL 8319, SPF); Itutinga, BR-265, Lavras-São João Del Rei, cerca de 25 km de Lavras, margem esquerda da estrada, 21°17′53″ S, 44°42′50″ W, 15 Jan 2003 (fl), Cavalcanti et  al. 3088 (CEN, MO); Itutinga, 10 Sep 1978 (fl, fr), Kirizawa 246 (CEN, SP [3]); Lavras, de fronte a entrada do Parque do Rio Bonito, estrada LavrasLuminárias, km-8.5, 10 Apr 2001 (fl, fr), Chaddad Jr. 39 (ESA); Lavras-São João Del Rey, 10 Oct 1992 (fl, fr), Souza & Sakuragi 2005 (CEN, ESA, HUFU, MBM, RB); Lima Duarte, Conçeição de Ibitipoca, Serra da Ibitipoca, 30 Sep 1970, Sucre et al. 7255 (P, RB); arraial de Conceição de Ibitipoca, Parque Estadual do Ibitipoca e Pedra do Gavião, 19 May 2001 (fl, fr), Castro et  al. 343 (CEN, CESJ); Lima Duarte, Serra de Ibitipoca, 20 Jun 1991 (fl, fr), Salimena-Pires s.n. (CEN 18566, CESJ 25451); Lima Duarte, Arredores do Parque Estadual do Ibitipoca, 9 Feb 2001 (fl, fr), Forzza et al. 1811 (CEN, CESJ); Lima Duarte, Ibitipoca, 14 Oct 1986, Souza & Andrade 114 (CEN, UB); Lima Duarte, Ibitipoca, 30 Aug 1987, Stehmann & Oliveira s.n. (BHCB 15703, CEN 19125); Lima Duarte, Serra de Ibitipoca, 15 May 1970 (fr), Krieger 8621 (CESJ n.v., SPF); Lima Duarte, Ibitipoca, Saint Gobain, próximo ao povoado do Souza, 14 Jul 2005 (fl, fr), Tameirão-Neto 3925 (BHCB, CEN); São João del Rei, 23 Aug 1974 (fl), Badini s.n. (OUPR 4162).

Descriptions

287

Diplusodon myrsinites individuals are much-branched, glabrous subshrubs, distinguished by the presence of hyphodromous leaves with plane margins that are yellow- to wine-colored. Its flowers are pedicellate (pedicel 2–3  mm long) with prophylls reaching from the middle to the apex of the floral tube. The epicalyx segments are spreading to suberect, and the androecium has 12 stamens. Diplusodon myrsinites resembles D. ciliiflorus with respect to the elliptic, hyphodromous leaves with yellow margins. With respect to the flower, D. ciliiflorus differs especially by the longer epicalyx segments and sepals, both of which are prominently ciliate. The types of Diplusodon myrsinites var. cribellinus and D. myrsinites var. cribellinus f. obtusifolius (Silveira 938 and Schwacke 12410, respectively) were destroyed at B. For Diplusodon myrsinites var. cribellinus, a neotype was chosen based on the gathering Sucre 6819 deposited at RB.  The specimen corresponds well to the description of the variety. For D. myrsinites var. cribellinus f. obtusifolius, two specimens of Schwacke 12410 were found in Brazilian herbaria, and the specimen at RB has the original label of the Schwacke Herbarium and is annotated with the name of the variety. The specimen at RB (RB 00209541) is chosen here as lectotype. 54. Diplusodon nigricans Koehne, Bot. Jahrb. Syst. 23 (Beibl. 57): 34. 1897. Type: Brazil. Goiás: Chapadão dos Veadeiros, 9 Jan 1895 (fl), A. F. M. Glaziou 21425 (lectotype, P [01901868], here designated; isolectotypes, BR, fragm. F, K, LE., R, S). Figs. 13.68 A–J, 13.70 Diplusodon xerampelinus Lourteig, Bradea 5(19): 230. 1989. Type: Brazil. Goiás: Alto Paraíso de Goiás, Chapada dos Veadeiros, 16  km  N of Alto Paraíso de Goiás, 5 Feb 1979 (fl), B. Gates & G. F. Estabrook 140 (holotype, P [01901995]; isotypes, RB, SP, UB). Shrubs 1–1.5  m tall, erect, branched, dark-brown to black when dry. Upper branches subquadrangular to quadrangular, unwinged, with short trichomes, internodes 1–3 cm long. Leaves decussate, not imbricate, suberect, petiolate; petioles 1–4 mm long; blades 10–35 × 8–20 mm, coriaceous, not discolorous, eucamptodromous, ovate to elliptic, base acute, attenuate; apex obtuse, margin plane, indumentum hirtellous on both surfaces, short trichomes sometimes mixed with tufts of stellate trichomes; secondary veins 3–6 on each side, departing along the midvein, prominent on the abaxial surface, domatia present. Synflorescence frondose, triplobotryum to pleiobotryum type; accessory branches absent, principal florescence 5–10  cm long, paraclades 4–20  cm long; bracts similar to the vegetative leaves, diminishing in size toward the apex of the branch, flowers sessile, hypopodium and epipodium absent; prophylls 6–8  mm long, reaching the apex of the floral tube, oblong, strongly keeled, minutely pilose, margin hyaline; floral tube 5–6 mm long, funnel-shaped, minute pilose; sepals 2–2.8  mm long, not caudate, not deflexed, minute pilose; epicalyx segments 2–3 mm long, cylindrical, suberect to spreading, the same size as or shorter than the sepals, with short cilia on the apex; corolla 2.5–3 cm in diam., rose-magenta to purple, petals 14–17 × 8–10 mm, obovate, apex obtuse; stamens 12, free portion of the filaments 9–10  mm long, exserted; ovary

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Fig. 13.68  Diplusodon nigricans. A. Branch with inflorescence. B–D. Leaf, abaxial surface. E. Floral bud. F–G. Prophylls, adaxial surface. H–I. Flowers with prophylls, without petals. J. Ovary. (A–B, D–F, H, J Gates & Estabrook 140, UB; C, G, I Glaziou 21425, CEN)

Descriptions

289

2–2.5  ×  1.8–2  mm, rhombic, glabrous, style 14–15  mm long; ovules 18–22. Immature capsules ca. 3.5 × 2.5 mm, oblong, glabrous, or with white indumentum at the apex; seeds not seen. Distribution and Ecology (Fig. 13.70)  Diplusodon nigricans is endemic to northern Goiás, in the municipalities of Alto Paraíso de Goiás and Cavalcante. Flowers from December to April; fruits (immature) in September. Specimens Examined: BRAZIL. Goiás,  Alto Paraíso de Goiás, 5 km E de Alto Paraíso de Goiás, 14º S, 47º W, 24 Jan 1979 (fl), Gates & Estabrook 23 (SP); Alto Paraíso de Goiás, Chapada dos Veadeiros, 21 Dec 1968 (fl), Barroso et  al. 624 (CEN, P, UB); Alto Paraíso de Goiás, Chapada dos Veadeiros, 4 Jan 1972, Rizzo 7374 (UFG); Alto Paraíso de Goiás, km 34, 29 Jan 2003 (fl), Cavalcanti et al. 3165 (CEN); Cavalcante, rodovia GO-118, 8–10 km N de Rio das Almas, 14 Feb 1990 (fl), Hatschbach & Nicolack 53978 (BR, MBM, NY, UB); Teresina de Goiás, a 24 km de Teresina, 13°51′49″ S, 47°19′44″ W, 1 Sep 2004 (fr), Cavalcanti et al. 3540 (CEN); Cavalcante, Fazenda Renascer, trilha para a ponte de pedra, 20 Apr 2003 (fl), Pastore et al. 475 (CEN); Cavalcante, Fazenda Renascer-trilha para Ponte de Pedra, 2 Feb 2004 (fl), Pastore et  al. 789 (CEN); Chapada dos Veadeiros, ca. 35 km N of Veadeiros, 14 Mar 1969 (fl), Irwin et al. 24321 (UB); Teresina de Goiás, a 24 km de Teresina, 13°52′29″ S, 47°20′08″ W, 29 Jan 2003 (fl), Cavalcanti et al. 3151 (CEN, MO); Teresina de Goiás, a 30 km de Teresina, 13°53′45″ S, 47°21′33″ W, 29 Jan 2003 (fl), Cavalcanti et al. 3154 (CEN, MO); without reference to municipal district: Glaziou s.n. (P). Diplusodon nigricans is represented by branched shrubs characterized by elliptic to ovate leaves that turn dark-brown to black when dry. The leaves have an indumentum of sparse simple trichomes that are sometimes mixed with stellate trichomes. In addition, the sessile flowers possess oblong, keeled prophylls which reach the apex of the floral tube, spreading to suberect epicalyx segments of 2–3 mm long, and androecium with 12 stamens. The type material of Diplusodon nigricans, Glaziou 21425, studied by Koehne at Berlin (B) was destroyed (F neg. nº 017927). None of the duplicates found in the herbaria were identified by Koehne. The P specimen was chosen as the lectotype because it is a good representative of the species. 55. Diplusodon nitidus Mart. ex DC., Prodr. 3: 94d. 1828. Friedlandia nitida Mart. ex DC., Prodr. 3: 94d, 1828, pro syn. Type: Brazil. Minas Gerais: in campis ad Serro, Jul (fl), C. F. P. Martius s.n. (first-step lectotype, designated by Lourteig 1989: 220, second-step lectotype, M [0146268], here designated; isolectotypes, M [0146267], fragm. G). Figs. 13.69 A–E, 13.70 Subshrubs 1.5–3 m tall, erect, much-branched, glabrous, sometimes dark-brown when dry. Upper branches quadrangular, winged, internodes 1.5–4 cm long. Leaves decussate, not imbricate, suberect, petiolate; petioles 5–10  mm long; blades 20–45  ×  6–25  mm, coriaceous, not discolorous, eucamptodromous, elliptic, base

290

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acute or attenuate, apex obtuse, margin plane, nitid; secondary veins 3–6 on each side, departing along the midvein, not prominent; domatia absent. Synflorescence bracteose to frondose-bracteose, diplothyrse to diplobotryum type, lax; accessory branches rarely present, principal florescence 4–7  cm long, botryum type, sometimes caducous, paraclades 4–35 mm long; bracts 6–12 mm long, elliptic, linearelliptic to narrowly obovate; flowers pedicellate, hypopodium 1.5–3  mm long, epipodium 1.5–2 mm long; prophylls 2–3 × ca. 0.8 mm, reaching to the base of the floral tube, linear-lanceolate, unkeeled, base obtuse to retuse, apex acute, margin plane; floral tube 5.5–6  mm long, wine-colored, funnel-­shaped, glabrous; sepals 4–5  mm long, triangular, with prolonged apex, not deflexed; epicalyx segments 3–3.5 mm long, cylindrical, spreading to slightly deflexed, shorter than the sepals; corolla 4–4.5 cm diam., rose-magenta, petals 15–18 × 10–11 mm, obovate, apex obtuse; stamens 12, free portion of the filaments 7–8 mm long, included to slightly exserted; ovary obovate-depressed, style 13–15 mm long, ovules 13–16. Capsules 6–7 × 5.5–7 mm, globose, glabrous; seeds 11–22, 2.5–3 × 2–3 mm. Distribution and Ecology (Fig.  13.70)  Diplusodon nitidus occurs in the mountains of the east of Brazil, from the central portion of Cadeia do Espinhaço, in the surroundings of the municipality of Diamantina, in Minas Gerais State, to the center-south of Bahia State, in campo rupestre vegetation and also on rocky outcrops in cerrado vegetation, in elevations of 700–1300 m. Flowers from March to July; fruits registered from February to July and from September to November. Specimens Examined: BRAZIL.  Bahia,  Montugaba, ca. 8  km em direção a Jacarau, 14°53′07″ S, 42°28′03″ W, 16 Mar 1994 (fl), Souza et al. 5539 (K). Minas Gerais, Botumirim, Serra da canastra, acesso pela cidade, 16°51′16″ S, 43°01′40″ W, 17 May 2008 (fl), Rosa et al. 1073 (CEN, HUFU); Conselheiro Mata, estrada de terra Conselheiro Mata-Diamantina, a 11  km de Conselheiro Mata, 18°18′45″ S, 43°54′28″ W, 7 Fev 2015 (fr), Cavalcanti et al. 3928 (CEN); Botumirim, 17 May 1998 (fl), Pirani et al. 4367 (CEN, SPF); Diamantina, 18°17′ S, 43°49′ W, 19 May 1990 (fl), Arbo et al. 4358 (CEN [2], CTES n.v., MBM, SPF); Diamantina, Rod. Guinda-Conselheiro Mata, km 10, 21 May 1989 (fl, fr), Hatschbach & Nicolack 53031 (MBM, MO); Diamantina, Rod. Guinda-Conselheiro Mata, km 178, 4 Jun 1985 (fl, fr), Tamashiro et al. 17387 (ESA); Diamantina, Serra do Capão, 19 May 1931 (fl), Mexia 5862 (MO, NY [3], VIC); Gouveia, Morro da torre da Embratel, próximo à usina eólica, 18°36′26″ S, 43°53′53″ W, 4 Sep 2003 (fr), Cavalcanti et al. 3202 (CEN, MO); BR-259, 20 km SW de Gouveia, caminho para Curvelo, 18°35′ S, 43°50′ W, 20 Mar 1990 (fl, fr), Arbo et al. 4469 (CEN, CTES n.v., SPF); Gouveia, ca. 19  km  S de Gouveia, 28 Nov 1985 (fr), Graham 981 (MO, NY); Gouveia, estrada entrando paralela aos moinhos de vento do Camilinho, sítio do Mulato, área de extração do Aliance, 18° 31′ 50′′S, 43° 53′ 45′′W, 15 Apr 2016 (fl), Faria-Jr. & Otoni 5658 (CEN, UB, HDJF); Grão-Mogol, Plantar MG-15-fazenda Tamanduá, 9 Oct 2005 (fr), Tameirão-Neto 4018 (CEN, BHCB); Itacambira, Serra de Itacambira, 1° ponto, km 910, Brandão-Laca 20818 (PAMIG); Montes Claros, ca. 30 km SE de Montes Claros, estrada para Juramento, 15 May 1977 (fl, fr), Gibbs et al. 5128 (SPF, UEC n.v.); Juramento, estrada Juramento-Itacambira, 16°57′49″ S, 42°32′14″ W, 27

Descriptions

291

Fig. 13.69  Diplusodon nitidus. A. Branch with inflorescence. B. Leaf, abaxial surface. C. Floral bud. D. Prophyll, adaxial surface. E. Flower with prophylls, without petals. (A, C Gibbs et  al. 5128, SPF; B, D–E Hatschbach & Nicolack 53031, MBM)

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Fig. 13.70  Geographic distribution of Diplusodon mononeuros, D. myrsinites, D. nigricans, and D. nitidus

Descriptions

293

Apr 2009 (fl, fr), Mello-­Silva 3183 (SPF); Retiro, 11 km para Mato Verde, 15°22′41″ S, 42°41′54″ W, 17 Mar 1994 (fl), Sakuragui et al. (CFCR) 15119 (CEN [2], ESA, K n.v., MBM, RB, SPF); Santo Antônio do Retiro, 15o20′18′′S, 42o45 09′′W, 15 Mar 2017 (fl), Sevilha et al. 6354; (CEN, MO, OUPR, SP, W); São João do Paraíso, 15o08′14′′S, 42o08′38′′W, Sevilha et al. 6866 (CEN, Vargem Grande do Rio Pardo, 15o24′23′′S, 42o21′04′′W, 23 Mar 2017 (fl), Sevilha et al. 6795 (CEN). The plants of Diplusodon nitidus are much-branched subshrubs completely glabrous with nitid and elliptic leaves of acute to attenuate base and petioles up to 1 cm long. The inflorescences are diplobotryum or diplothyrse type, the unusual type for Diplusodon where we can find cymes of triads on the paraclades. The flowers have pedicels 3–5  mm long, reduced prophylls reaching only to the base of the floral tube, spreading to slightly deflexed epicalyx segments and androecium with 12 stamens. The specific epithet refers to the shiny aspect of the leaves. In the Munich herbarium (M), there are two unnumbered Martius specimens of Diplusodon nitidus. One of them has, in addition to a blue label with the locality data “in campis Serro Frio” (today Diamantina), a white label with the annotation, “Friedlandia nitida,” also with the number of stamens and the petal color. Because this is the most complete with the largest number of flowers, it is here selected as lectotype (M 0146268). 56. Diplusodon oblongus Pohl, Flora 10: 152. 1827. Type: Brazil. Goiás: Luziânia (Santa Luzia, “Megaponte” [Meiaponte, now Pirenópolis]) in campis inter frutices, circa Vila Boa [now town of Goiás], Feb–Mar, J. B. E. Pohl 5791 (firststep lectotype, designated by Lourteig 1989: 220, second-step lectotype, W [0053425], here designated; isolectotypes, BR [2], G, M, W [0053424]). Figs. 13.71 A–L, 13.75E Subshrubs 0.4–3 m tall, erect, much-branched, with a xylopodium, dark-brown when dry. Upper branches subquadrangular to cylindrical, rarely quadrangular, rustcolored, with reduced wings 0.1–0.2 mm wide, totally glabrous or inconspicuously pilose with minute trichomes, internodes 1–3.5  cm long. Leaves decussate, not imbricate, spreading to suberect, petiolate; petioles 2–9  mm long; blades 10–7.5 × 10–38 mm, not glaucous, coriaceous, not discolorous, dark-brown when dry, eucamptodromous, elliptic, ovate or ovate-lanceolate, base acute or attenuate, apex obtuse rarely acute, margin plane, rarely with some short cilia, blades glabrous; secondary veins 3–4 on each side of the midvein, prominent, yellow in nature, wine-colored when dry; domatia absent. Synflorescence frondose, triplobotryum to pleiobotryum type, congested; sometimes with accessory branches of the first order, principal florescence 2–10  cm long, paraclades of the first order 7–27 cm long, of the second order 2–10 cm long; bracts similar to the vegetative leaves in shape, decreasing in size toward the apex of the branch; flowers subsessile, hypopodium 0.5–1 mm long, epipodium 0–0.5 mm long; prophylls 4–6 × 2.5–3 mm, reaching the middle or to the apex of the floral tube, oblong, keeled, base retuse, apex acute to obtuse, margin plane, glabrous or minutely pilose; floral tube 5–6.5 mm long, wine-colored, oblong, often glabrous, rarely minutely pilose; sepals

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Fig. 13.71  Diplusodon oblongus. A. Habit. B. Section of internode. C. Leaf, abaxial surface. D. Prophyll, lateral view showing colleters. E. Prophyll, adaxial surface. F. Floral bud. G. Flower with prophylls, without petals. H. Sepal, inner surface showing glandular trichomes. I. Ovary. J. Fruit. K. Capsule. L. Simple trichome. (A Cavalcanti et al. 454, CEN; B, C, E–G Cavalcanti et al. 432, CEN; H–J Cavalcanti et al. 437, CEN)

Descriptions

295

2–2.5  mm long, wine-colored, triangular, not caudate, slightly deflexed, thick, minutely ciliate, sometimes cilia more evident on the sepal apex, with conspicuous papillose trichomes on the margin of the inner surface; epicalyx segments 1.5–5 mm long, cylindrical, spreading to erect, the same size of the sepals, often glabrous, rarely minutely pilose; corolla 4–4.5 cm in diam., lilac to clear purple or sometimes rose-magenta, petals 13–15 × 9–10 mm, elliptic, apex obtuse to acute; stamens 12, free portion of the filaments 8–9 mm long, exserted, ovary 1.5–1.8 × 1.8–2 mm, obovate-depressed, glabrous, style 13–15 mm long, ovules 22–32. Capsules 4.8– 5.5  ×  4–4.5  mm, oblong, glabrous; seeds 9–25, 2–3  ×  1.5–2.5  mm. 2n  =  30 (Cavalcanti 1995; Graham and Cavalcanti 2001). Distribution and Ecology (Fig. 13.75)  Diplusodon oblongus is the second most widely distributed species of the genus. The species occurs in central Brazil in Distrito Federal, Goiás, and Mato Grosso and less often in the states of Tocantins and west of Minas Gerais. In Mato Grosso it occurs in cerrados at 300–400 m. In Goiás, it occurs in cerrado vegetation at 700–800 m and is also found in the campos rupestres of Chapada dos Veadeiros and campos sujos of Serra dos Pireneus, at elevations of 900–1600 m. In Distrito Federal, it occurs in open cerrado and bordering gallery forests at 1000 m and in campos sujos from Chapada da Contagem to the campos alagados of Brasília, at 1100–1250 m. Flowers from October to May, predominantly from January to April; fruits from November to August. Specimens Examined: BRAZIL.  Distrito Federal,  APA da Cafuringa, 15°31′ S, 47°58′ W, 14 Mar 1991, Dias et  al. 5 (CEN, MO); Bacia do rio São Bartolomeu, 18 Jan 1980 (fl), Heringer et  al. 3122 (IBGE, SP, MO, NY, UB); Brasília, Asa Norte, SQN 208, 15°45′ S, 45°53′ W, 20 Feb 1986 (fl, fr), Nascimento & Catharina 54 (UB); Brasília, Lago Paranoá, imediatamente E do Lago Paranoá, 12 Dec 1965 (fl), Irwin et al. 11219 (NY, UB); Brasília, Fazenda Água Limpa, 18 Aug 1985 (fr), Barros 1000 (UB); Brasília, Fazenda Água Limpa, 5 Jan 1994 (fl), Munhoz 71 (HEPH); Brasília, Fazenda Água Limpa, ca. 18 km do centro de Brasília, 21 Jan 1987 (fl), Stevens 16 (UB); Brasília, Fazenda Água Limpa, ca. 18 km SE da torre de TV de Brasília, 13 Dec 1976 (st), Ratter et al. 4032 (UB); Brasília, Fazenda Agua Limpa, 25 Jan 1977 (fl, fr), Ratter et al. 4176 (MO, NY, UB); Brasília, Jardim Botânico de Brasília, 5 Jun 1990 (fr), Azevedo & Oliveira 616 (IBGE, P n.v., UEC n.v.); Brasília, Jardim Botânico de Brasília, trilha de entrada da área de visitação, 15º52′S, 47º51′W, 24 Feb 1997 (fl), Nóbrega et al. 713 (HEPH); Brasília, Jardim Botânico de Brasília, estrada que dá acesso à CAESB, 15º32′S, 47º51′W, 19 Jun 2000 (fr), Nóbrega et  al. 1173 (HEPH); Brasília, Jardim Botânico de Brasília, Jardim Botânico de Brasília, ca. 20 km SE de Brasília, 6 Feb 1986 (fl), FurtadoNeto 3 (HEPH); Brasília, Jardim Botânico de Brasília, Furtado-Neto 5072 (HEPH); Brasília, Setor Militar Urbano, em frente ao quartel das Forças Armadas, 7 Feb 1983 (fr), Oliveira 16 (UB); Brasília, Parque Rural, fim da Asa Norte, pátio do Cenargen, 21 Feb 1977 (fr), Allem 849 (CEN, RB, UB); Brasília, Reserva Ecológica do Guará, 15°48′ S, 47°58′ W, 19 Jan 1994 (fl, fr), Pereira-Silva 2200 (CEN, MO, NY); Brasília, Reserva Ecológica do Guará, 15 Feb 1994 (fl), Pereira-Silva 2278 (CEN, MO, NY); Brasília, Reserva Ecológica do IBGE, 6 Dec 1990 (fl), Camara &

296

13  Systematic Treatment

Brochado 68 (IBGE); Brasília, Reserva Ecológica do IBGE, 15°57′03″ S, 47°52′18″ W, 6 Jan 2004 (fl), Silva 5649 (CEN, IBGE); Brasília, Reserva Ecológica do IBGE, Área do Cristo Redentor, 16 Mar 1990 (fr), Azevedo & Alvarenga 547 (IBGE); Brasília, Reserva Ecológica do IBGE, picada R-7, 6 Mar 1978 (fl, fr), Heringer et al. 393 (MO, NY); Brasília, Reserva Ecológica do IBGE, Nov 1992 (fr), Souza 139 (IBGE); Brasília, Reserva Ecológica do IBGE, 14 Jul 1981 (fr), Heringer et al. 7135 (IBGE, K, MG, MO, P); Brasília, Fundação Zoobotânica, 18 Feb 1961 (fr), Heringer 7945 (UB); Brasília, Campus da Universidade de Brasília, 13 Apr 1963 (fl), Pires et al. 9089 (NY, UB); Brasília, cerrado entre Hotel Planalto e Universidade, 2 Mar 1965 (fl, fr), Clayton 4888 (K, P, SP); Brasília, Campus da Universidade de Brasília, 10 Feb 1977 (fl, fr), Classe de Taxonomia da UNB 399 (UB 32653); Brasília, Campus da Universidade de Brasília, 25 Jan 1978 (fl), Gentry 21447 (MO, UB); Brasília, Parque Nacional de Brasília, 18 Mar 1998, Cavalcanti et al. 2389 (CEN); Brasília, Parque Nacional de Brasília, 11 Feb 1992 (fl), Barros et al. 2318 (IBGE, UB); Chapada da Contagem, rodovia para Brazlândia, ca. 1 km do trevo da BR-060, 10 Feb 1988 (fl), Pirani et al. 2129 (F n.v., NY n.v., SPF); DF-100, a 93 km da entrada para Unaí, 15°33′ S, 47°25′ W, 12 May 1992 (st), Dias et al. 111 (CEN, MO); Estação Florestal Cabeça de Veado, ca. 22 Feb 1983 (fr), Alves 6 (CEN, HEPH, IBGE); Estação Florestal Cabeça de Veado, ca. 20 km SE de Brasília, 2 Feb 1983 (fl, fr), Reis 4 (CEN, IBGE); Granja da Samambaia, 13 Nov 1964 (fl, fr), Heringer 9938 (UB); Riacho Fundo, Fazenda Sucupira, 24 Jan 1997 (fl), Sampaio et al. 102 (CEN, NY); Riacho Fundo, Fazenda Sucupira, 15°55′ S, 48°01′ W, 5 May 1994 (fr), Assis et al. 12 (CEN, MO); Riacho Fundo, Fazenda Sucupira, 15°55′27″ S, 48°02′ W, 10 Jan 1997, Sampaio et al. 76 (CEN, MO); Riacho Fundo, Oliveira 1635 (HEPH); rio Corumbá, 8 Jan 1967 (fl), Heringer 11273 (NY, UB [2]); rio São Bartolomeu, Quebrada dos Neri, 1 Mar 1991 (fl, fr), Melo & França 572 (NY, UB); Rodovia 15, 30 Jan 1975 (fl, fr), Heringer 14449 (P, UB); Saia Velha, 15 Feb 1965 (fl, fr), Heringer 10080 (UB); perto da represa Saia Velha, 16°02′ S, 47°56′ W, 11 Apr 1983 (fl), Kirkbride Jr. 5203 (UB); Samambaia, Parque Boca da Mata, 15°52′ S, 48°03′ W, 9 Feb 1996, Rezende 358 (CEN, MO); São Sebastião, 4 Apr 2003 (fl), Pastore et al. 454 (CEN); Sobradinho, 26 Feb 1965 (fl. fr), Belém 497 (UB); 13 km SE de Brasília, 4 km SE da Escola Fazendária, 23 Jan 1980 (fl), Garcia-Kirkbride 1014 (UB). Goiás, Abadiânia, 16°05′38″ S, 48°46′49″ W, 12 Mar 2002 (fr), Mendonça et  al. 4635 (CEN, IBGE); Acreúna, margem esquerda da rodovia Acreúna-rio Verde, km 147, 10 Mar 1983, Rizzo 10239 & Ferreira 135 (UFG); Alexânia, cerca de 300  m a oeste da BR 060, na direção de Corumbá de Goiás, 16°08′33″ S, 48°36′12″ W, 18 Feb 2003 (fl, fr), Pereira-Silva et  al. 7157 (CEN, MO); Alto Horizonte, Fazenda do Senhor Manoel Braz, 6 Mar 2011 (fl), Faria-Jr. 1177 (CEN, UB); Alto Paraíso de Goiás, 14°08′ S, 47°27′ W, 14 Mar 1995, Cavalcanti et al. 1330 (CEN); Alto Paraíso de Goiás, 14°08′ S, 47°27′ W, entrada do Belvedere Paraíso. A cerca de 4 km N de Alto Paraíso de Goiás, 14 Mar 1995, Cavalcanti et  al. 1347 (CEN); Alto Paraíso de Goiás, Chapada dos Veadeiros, Estrada Alto Paraíso de Goiás-Colinas do Sul, 9  km do entroncamento com a GO-118, 14°09′ S, 47°35′88″ W, 2 Apr 1997 (fr), Cavalcanti et al. 2181 (CEN, MO, NY); Alto Paraíso de Goiás, a oeste do Centro da cidade, 20 Feb 1991 (fl, fr), Pereira

Descriptions

297

et al. 1456 (IBGE); Alto Paraíso de Goiás, GO-327, 15–20 km O de Alto Paraíso de Goiás, 12 Feb 1990 (fl, fr), Hatschbach & Nicolack 53883 (MBM, MO); Alto Paraíso de Goiás, rodovia GO-118, Alto Paraíso de Goiás-Terezina de Goiás, km 8, 14°03′ S, 47°32′ W, 7 Mar 1989 (fl, fr), Cavalcanti et al. 377 (CEN, F, K, MBM, NY, SPF); Alto Paraíso de Goiás, 7 km pela estrada N de Alto Paraíso de Goiás, 4 Mar 1973 (fl), Anderson 6265 (NY, UB); Alto Paraíso de Goiás, 20  km  W de Veadeiros (Alto Paraíso de Goiás), 16 Feb 1966 (fl, fr), Irwin et al. 12928 (NY); Alto Paraíso de Goiás, 1 km W de Veadeiros (Alto Paraíso de Goiás), 13 Feb 1966 (fl, fr), Irwin et al. 12743 (K, MO, NY, UB); Alto Paraíso de Goiás, Chapada dos Veadeiros, 27  km na estrada N de Alto Paraíso de Goiás, 9 Mar 1973 (fl, fr), Anderson 6722 (NY, UB); Alto Paraíso de Goiás, ca. 6 km a leste de Alto Paraíso de Goiás, 14°07′ S, 47°30′ W, 10 Feb 1979 (fl), Ferreira & Cardoso 10 (UB [3]); Alto Paraíso de Goiás, Chapada dos Veadeiros, 21 Dec 1968 (fl), Barroso et al. 590 (UB); Alto Paraíso de Goiás, Chapada dos Veadeiros, 21 Dec 1968 (fl), Harley et al. 11458 (K, NY); Alto Paraíso de Goiás, Parque Nacional da Chapada dos Veadeiros, arredores da sede do IBDF, próximo ao rio Jacó, 14 May 1986 (fl, fr), Toledo et al. 104 (SP); Alto Paraíso de Goiás, Alto Paraíso de Goiás-São Jorge, lado direito, 13 km de Alto Paraíso, 14°09′54″ S, 47°37′16″ W, 29 Jan 2003 (fl), Cavalcanti et al. 3158 (CEN, MO); Alto Paraíso de Goiás, estrada de Alto Paraíso para São Jorge, ca. 20  km da estrada de São Jorge, 1 Feb 2004 (fl), Pastore et  al. 777 (CEN); Alto Paraíso de Goiás, entre Cavalcante e Alto Paraíso de Goiás, cachoeira dos Cristais, 20 Apr 2003 (fr), Pastore et al. 476 (CEN); Anápolis, margem da rodovia BrasíliaAnápolis, 8 Dec 1965 (fl), Belém 1917 (CEPEC); Anápolis, ramal no km 8 da estrada Anápolis para Corumbá de Goiás, 2–4 km ramal adentro, 8 Feb 1986 (fl), Carvalho & Delphim 2248 (UB); Anápolis, a esquerda do Ribeirão Dourado, 29 Jan 1969 (fl), Rizzo & Barbosa 3415 (UFG); Aparecida de Goiânia, divisa com hidrolândia, próximo à BR-153, 26 Apr 2003 (fr), Pastore & Bringel 572 (CEN); Barro Alto, foot of Hill, near hilltop, 15°02.96′ S, 48°57.12′ W, 15°03.286″ S, 48°57.323″ W, 30 Jan 2005 (fl), Reeves 2864 (CEN); Cachoeira da Vargem Grande, Rio Piçarrão, Glaziou 21548 (P); Caldas Novas, 28 Feb 1970 (fl), Rizzo 4799 & Barbosa 4045 (UFG); Bom Princípio, morro da Estação Repetidora Dois Irmãos, na subida do morro, 15°14′03″ S, 48°43′06″ W, 4 Apr 2006 (fr), Cavalcanti & Pereira-Silva 3723 (CEN); Caiapônia, bacia do rio Caiapó, 17°04′32″ S, 51°34′44″ W, 1 Jan 2007 (fr), Sousa-Silva et al. 275 (CEN, IBGE); Caldas Novas, alto da Serra de Caldas Novas, 4 Feb 1971 (fl), Rizzo 5933 & Barbosa 5181 (UFG); Caldas Novas, 12 km W da cidade de Caldas Novas, 2.6  km  N do hotel Pousada do rio Quente, 17º47′S, 48º45′W, 23 Dec 1974 (fl), Heringer & Eiten 14175 (SP, MO, UB); Cavalcante, entroncamento Minaçu-Kalunga (Engenho Dois), km 2, 13°37′08.9″ S, 47°28′30.4″ W, 31 Aug 2004 (fr), Cavalcanti et al. 3533 (CEN, SPF); Cavalcante, estrada de terra Cavalcante-Kalunga, 13  km, 13°41′31″ S, 47°27′57″ W, 18 Nov 2005 (fr), Cavalcanti et al. 3624 (CEN, MO); Cavalcante, caminho para o povoado Kalunga, 3 Feb 2004 (fr), 3 Mar 2004 (fr), Pastore et al. 843 (CEN); Cocalzinho, Fazenda Buriti, 15°44′22″ S, 48°46′18″ W, 23 Mar 2002 (fr), Silva et al. 5135 (CEN, IBGE); Corumbá de Goiás, a 10 km de Corumbá de Goiás, 15°55″35″ S, 48°51′12″ W, 13 Mar 2002 (fl, fr), Mendonça et al. 4668 (CEN, IBGE); 12 km S de Corumbá de

298

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Goiás, 2 Dec 1965 (fl), Irwin et al. 10934 (NY); Cristalina, 7 Jan 1994, Hatschbach et al. 59907 (BR, MBM); Cristalina, estrada para Salto Arrojado, 10 Feb 1990 (fl, fr), Hatschbach & Nicolack 53748 (MBM, MO); Cristalina, 20 km N de Cristalina, 7 Mar 1966 (fr), Irwin et al. 13690 (LE n.v., NY); Cristalina, 12 km na estrada N de Cristalina, 3 Apr 1973 (fl), Anderson 8001 (NY, UB); Cristalina, margem esquerda do rio Arrependido, a noroeste da área de empréstimo, 16°13′14″ S, 47°20′42″ W, 6 Mar 2002 (fl, fr), Pereira-Silva et al. 6062 (CEN, MO); Édeia, Fazenda Canadá, 4 Apr 2013 (fl, fr), Faria et al. 3375 (UB); Formosa, Fazenda Indaiá, 5 May 2003 (fr), Pastore & Suganuma 586 (CEN); Goiânia, Dec 1936 (fr), Brade 15433 (MO, RB n.v.); Goiânia, km 14 da rodovia Goiânia para Inhumas, 29 Jan 1969 (fl), Rizzo & Barbosa 3478 (UFG); rodovia Goiânia-Brasília, BR-060, a 10 km E do acesso a Alexânia, 16°10′ S, 48°30′ W, (fl), Pirani et al. 2085 (CEN, K, NY n.v., SPF, UB); Goiânia, 10 km de Goiânia, à direita da GO-7 que liga Goiânia a Guapó, 10 Apr 1968 (fr), Rizzo & Barbosa 150 (UFG); Goiânia, a esquerda da rodovia GO-7, Goiânia para Guapó, 15 May 1969 (fr), Rizzo & Barbosa 730 (UFG); Goiânia, km 14 da rodovia para Inhumas, 29 Jan 1969 (fr), Rizzo & Barbosa 135 (UFG); Goianira, 21 Mar 1970 (fl, fr), Rizzo 4867 & Barbosa 4114 (UFG); Goiás, Jeroaquara, Serra de Santa Rita, 27 Mar 1971 (fr), Rizzo 6113 & Barbosa 5261 (UFG); Goiás, Jeroaquara, Serra de Santa Rita, 26 Feb 1972 (fl), Rizzo 7733 & Barbosa 6322 (UFG); Mossâmedes, alto da Serra Dourada, 18 Dec 1968 (fl), Barroso et al. 787 (UB); Mossâmedes, Serra Dourada, 1.3 km ão da junção com a estrada para Mossâmedes na rodovia GO-070 de Goiânia para Goiás, Velho, 15°57′ S, 50o2′ W, 9 Feb 1988 (fl), Thomas et  al. 5762 (CEN, NY, SPF); Mossâmedes, divisa dos municípios de Mossâmedes ao S e Goiás ao N, 5 Apr 1969 (fl), Rizzo 4041 (UFG); Mossâmedes Serra Dourada, perto da cidade de Goiás Velho, 18 Dec 1968 (fl), Harley et al. 11330 (K, NY); Mossâmedes, Serra Dourada, ca. 20 km de Goiás, Velho, 18 Jan 1966 (fl), Irwin et al. 11716 (NY); Mossâmedes, Serra Dourada, 10 Jan 1965 (fl), Heringer 10869 (NY, UB); Mossâmedes, Serra Dourada, 16°03′ S, 50o7′ W, 17 Mar 1989 (fl, fr), Cavalcanti et  al. 454 (CEN, CEPEC, NY, SPF); Ipameri, 17°44′ S, 48°28′ W, 12 Mar 1997 (fl, fr), Cavalcanti et al. 2149 (CEN, MO, NY); Jataí, estrada de Jataí para Serranópolis, a 20  km do Ribeirão Ariranhas, Campo Serrinha, Rizzo 8853 (UFG); Leopoldo Bulhões, 16°35′ S, 48°45′ W, 20 Mar 1989 (fr), Alvarenga et al. 168 (IBGE); esquerda da estrada GO-1, GoiâniaLeopoldo Bulhões, 9 km de Goiânia, 26 Jan 1971 (fl), Rizzo 6960 (UFG); Leopoldo Bulhões, 16°36′ S, 48°35′ W, 12 Jan 1989 (fl), Filgueiras et al. 1757 (IBGE, P n.v.); Luziânia, Mansões Reunidas Heringer, 25 Feb 1980 (fl, fr), Heringer 17844 (IBGE, K, MO, NY, US); Luziânia, AHE Corumbá IV, local desmatado para construção de casa de força, margem esquerda do rio Corumbá, 16°19′52″S, 48°11′26″W, 10 Dec 2001 (fl), Carvalho-Silva 170 (CEN, MO); Luziânia, próximo ao canteiro de obras, montante, margem direita, 16°20′10″ S, 48°11′47″ W, 13 Dec 2002 (fl), Rezende et  al. 843 (CEN, MO); Luziânia, área próxima ao encontro dos rios Lagoinha e Alagado, ponto 5, 16°16′13″ S, 48°11′36″ W, 5 Nov 2002 (fr), Pereira-Silva et al. 6773 (CEN, MO); Luziânia, área de pastagem próximo à Fazenda Santa Maria, 16°14′54″ S, 48°09′40″ W, 5 Nov 2002 (fl), Pereira-Silva et al. 6778 (CEN, MO); Montes Claros de Goiás, 16°02′16″ S, 51°20′54″ W, May 2007 (fr), Mendes &

Descriptions

299

Neves 174 (BHCB); Morrinhos, 17°49′22″ S, 49°03′39″ W, 6 Sep 1998 (fr), Proença et  al. 1992 (CEN [2], UB); Morrinhos, estrada Morrinhos-Caldas Novas, 11 Feb 1993 (fl, fr), Cordovil-Silva et  al. 239 (CEN, MO, NY); Morrinhos, estrada Morrinhos-Caldas Novas, 4 Jan 1971 (fl), Rizzo 5852 & Barbosa 5101 (UFG); Morrinhos, estrada Morrinhos-Caldas Novas, 4 Feb 1971 (fr), Rizzo 5912 & Barbosa 5161 (UFG); Nerópolis, Parque Estadual Altamiro de Moura Pacheco, 16°32′12.8″ S, 49°09′17.1″ W, 30 Mar 2005 (fl), Fonseca et al. 5734 (CEN); Niquelândia, 11 km em direção a Uruaçu, 22 May 1990 (fr), Pedralli et al. 3292 (CEN, NY); Niquelândia, estrada de chão direto à reserva do IBAMA, 14°29′46″ S, 48°27′ W, 1 Mar 1996 (fl, fr), Fonseca & Barros 836 (IBGE); Niquelândia, Fazenda Traíra, base do morro, 8 Aug 1995 (fr), Marquete et  al. 2213 (CEN, IBGE); Niquelândia, 8  km  S de Niquelândia, 23 Jan 1972 (fl), Irwin et al. 34860 (NY); Niquelândia, Nossa Senhora da Abadia do Muquén, Serra da Meia Noite, 14°22′34″ S, 47°54′13″ W, 5 Apr 2006 (fl, fr), Cavalcanti & Pereira-Silva 3743 (CEN); Niquelândia, Nossa Senhora da Abadia do Muquém, GO-237, entroncamento Muquém-Colinas do Sul, 5  km, 14°26′18″ S, 48°09′54″ W, 5 Apr 2006 (fl, fr), Cavalcanti & Pereira-Silva 3737 (CEN); 70 km N de Corumbá de Goiás, na estrada para Niquelândia, 13 Jan 1968 (fl), Irwin et al. 18504 (NY, UB); 70 km N de Corumbá de Goiás na estrada para Niquelândia, no vale do rio Maranhão, 20 Jan 1968 (fl, fr), Irwin et al. 18872 (IAN, K, NY, P, UB); Paraúna, Serra da Arnica, 17°02′ S, 50o27′ W, 15 Mar 1989 (fl), Cavalcanti et al. 437 (CEN, SPF); Paraúna, Serra das Galés, 17°09′ S, 50o27′ W, 14 Mar 1989 (fl, fr), Cavalcanti et  al. 430 (CEN, NY, SPF); Paraúna, estrada para Muralha de Pedra, próximo a córrego, 17°10′ S, 50o33′ W, 14 Mar 1989 (fl), Cavalcanti et  al. 432 (CEN, SPF); Pirenópolis, Serra dos Pireneus, cachoeira da Meia Lua, 14 Jan 2002 (fl), Pereira-Silva et al. 4537 (CEN, MO); Pirenópolis, ca. 10  km E de Pirenópolis, 15 Jan 1972 (fl), Irwin et  al. 34190 (MO, NY, UB); Pirenópolis, ca. 20 km E de Pirenópolis, 14 Jan 1972 (fl), Irwin et al. 34016 (MO, NY, UB); Pirenópolis, Serra dos Pireneus, 18 Mar 1989 (fl, fr), Cavalcanti et al. 474 (CEN, SPF); Pirenópolis, alto da Serra dos Pireneus, nas base dos Três Picos, 13 Mar 1971 (fr), Rizzo 6050 & Barbosa 5298 (UFG); Pirenópolis, Serra dos Pireneus, Fazenda Solar dos Pireneus, 12 Feb 2000 (fl, fr), Hatschbach et al. 70080 (MBM); Pirenópolis, estrada Cocalzinho-Pirenópolis, via Serra dos Pireneus, 20  km, 15°48′08″ S, 48°51′45″ W, 14 Nov 2005 (fr), Cavalcanti et al. 3584 (CEN); Rio Verde, Sete Quedas, 9 Feb 1974 (fr), Hatschbach 33949 (MBM n.v., MO, NY); São Domingos, próximo de São Domingos, May 1840, Gardner 4132 (BR, G, K, OXF, P, W); Santa Cruz de Goiás, 17°21′ S, 48°31′ W, 11 Jul 1996 (fr), Cavalcanti et al. 2019 (CEN, MO, NY); Santo Antonio do Descoberto, margem direita do rio Descoberto, próximo à Barra com o córrego Engenho das Lages, 16°05′39″ S, 48°16′27″ W, 17 Feb 2003 (fl), Pereira-Silva et al. 7126 (CEN, MO); São João da Aliança, estrada para São Jorge, 14°09′14″ S, 47°32′01″ W, 28 Feb 1999 (fl. fr), Calago 106 (CEN, NY); Serra dos Caiapós, 17°12′ S, 51°47′ W, ca. 60 km S de Caiapônia na estrada para Jataí, 27 Oct 1964 (fl), Irwin & Soderstrom 7441 (B, K, NY, P, SP, UB); Serra do Caiapós, a 40 km de Amorinópolis para Rio Verde, 22 Jan 1972 (fl), Rizzo 7497 & Barbosa 6227 (UFG); Serra do Caiapós, 22 Feb 1972 (fl, fr), Rizzo 7677 & Barbosa 6266 (UFG); Teresina de Goiás, entrada para o Poço Encantado, 13°52′54″ S, 47°15′54″ W, 6 Nov 2014 (fr), Pastore & Suganuma 1096

300

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(CEN); Uruaçú, 14°38′ S, 48°57′ W, 6 Feb 1996 (fl), Walter et al. 3038 (CEN, MO, NY); Uruaçú, 14°23′ S, 49º W, 7 Feb 1996 (fl, fr), Walter et al. 3083 (CEN, MO, NY); Uruaçú, 8 km do rio Maranhão, 11 Feb 1972 (fl), Rizzo 7645 (UFG); Uruaçú, 8 Mar 1972 (fl, fr), Rizzo 7861 (UFG); Vianópolis, Santa Maria do Passa Quatro, rodovia GO-139, 11 Feb 2000 (fl, fr), Hatschbach et  al. 69980 (MBM); without reference to municipal district: Burchell 6647 (BR [2], K, LE n.v.); Burchell 6700– 13 (K); Tamberlick s.n. (W). Mato Grosso, Barra do Garças, 15°51′09″ S, 52°16′26″ W, 15 Jun 1997, Cavalcanti et al. 2339 (CEN); Barra do Garças, Distrito de Indianápolis, 15°05′35″ S, 52°25′82″ W, 20 Mar 1997 (fl), Árbocz et al. 3472 (CEN, UB); Barra do Garças, Fazenda Brazil, 15°06′59″ S, 52°24′26″ W, 24 Mar 1997 (fl, fr), Árbocz et al. 3573 (CEN, UB); Canarana, 13°58′12″ S, 52°30′96″ W, 15 Feb 2010 (fl), Zanatta & Fernandes 237 (UB); Cuiabá, rodovia Cuiabá-Chapada dos Guimarães, Cachoeirinha, desvio à direita em direção à Casa de pedras, 5 km, 18 May 1986 (fl, fr), Saddi 7637 (UFMT); Cuiabá, Chapada dos Guimarães, 7 Apr 1985 (fl, fr), Silva 64 (IBGE, SP); Cuiabá, between Buriti e Água Fria, 15°23′ S, 55°47′ W, 1 Feb 1995 (fl, fr), Dubs 1818 (ESA); Cuiabá, Chapada dos Guimarães, Cachoeira Véu da Noiva, 3 Mar 1983 (fl, fr), Moraes et al. 105 (MEG n.v., UFMT); Cuiabá, Chapada dos Guimarães, 80 km acima da Cachoeira Véu de Noiva, 27 Jan 1989 (fl), Cavalcanti et al. 87 (CEN, K, SPF, UB); Cuiabá, entre Buriti e Chapada, 5 km antes da Chapada para Água Fria, 1 Jul 1984 (fl, fr), Saddi 4602 (UFMT); Cuiabá, Chapada dos Guimarães, Fazenda Morro da Lage, 15°19′ S, 55°44′ W, 25 Feb 1997 (fl), Nave et al. 1239 (ESA); Garapu, 30 Sep 1964 (fr), Irwin & Soderstrom 6458 (K, NY [2], P, UB); Garapu, ca. 1 km NE de Garapú, 1 Oct 1964 (fr), Irwin & Soderstrom 6517 (NY); Gaúcha do Norte, Gaúcha do Norte-Paranatinga, 13°37′06″ S, 53°17′30″ W, 28 Mar 1997, Dario et al. 1078 (CEN, UB); Novo São Joaquim, ponto 18, 23 Feb 1997 (fl, fr), Arbócz et  al. 3318a (ESA); Nova Xavantina, 11.5  km  N de Xavantina na estrada para Cachimbo, 1 Jan 1968 (fl), Philcox & Fereira 3816 (K, NY, P, UB); Nova Xavantina, Aldeia dos Areões, Fazenda Coração de Jesus e Santana, margem direita do rio das Mortes, 28 Jan 1969 (fl), Onishi et al. 866 (UB); Nova Xavantina, ca. 2 km NE de Xavantina, 7 Jun 1966 (fr), Irwin et al. 16711 (MO, NY); Nova Xavantina, ca. 6 km E de Xavantina, 14°44′ S, 52°20′ W, 25 Jul 1967 (fr), Ratter & Castro 163 (K, NY, UB); Nova Xavantina, ca. 1.5 km S de Xavantina, 11 Jan 1968 (fl), Philcox & Fereira 3971 (K, NY, UB); Nova Xavantina, 2 km S de Xavantina, 26 Sep 1964 (fr), Irwin & Soderstrom 6385 (NY); Nova Xavantina, ca. 6 km S de Xavantina, 11 Oct 1967 (fr), Argent et al. 6716 (K, NY, P, UB); Nova Xavantina, aproximadamente 10 km S de Xavantina, 14°44′ S, 52°20′ W, 11 Jan 1977 (fl, fr), Ratter et al. 4075 (K, MO, NY, UB); Nova Xavantina, ca. 20 km S de Xavantina, 9 Jun 1966 (fr), Irwin et al. 16800 (IAN, K, MO, NY, P, UB); 80  km  N de Barra do Garças na estrada para Xavantina, Vale dos sonhos, 15°40′ S, 52°20′ W, 26 Aug 1972 (st), Ratter et al. 2267 (UB); Nova Xavantina, Campus Universitário da UNEMAT, Estação Biológica Maric Viana, 22 Feb 1999, Viana 126 (ESA); Torixoréu, MT-0100, entre Barra do Garças e rio Diamantino, 20 Jun 1988 (fr), Saddi 9213 (UFMT); São José do Rio Claro, 13°51′ S, 56°29′ W, 14 Jun 1997, Souza et  al. 18014 (CEN, ESA, UB). Minas Gerais, Guarda Mor, 18°05′ S, 47°4″ W, 22 Apr 1997 (fr), Cavalcanti et  al. 2265 (CEN, MO, NY); Coranmandel, entroncamento Vazante-Coranmandel, MG-188, km 38, Serra da

Descriptions

301

Carcaça, Chapadão, 18°07′4′′ S, 47°5″ W, 22 Apr 1997, Cavalcanti et  al. 2268 (CEN, MO, NY); Vazante, mina de zinco da Votorantin, 17°57′34″ S, 46 o49′19″ W, 18 Jan 2008 (fl), Resende et al. 2306 (BHCB); Umuarama, 16º05′S, 46º26′W, 27 Nov 2000, Proença et al. 2307 (CEN, UB); Unaí, Cabeceira Grande, estrada que dá acesso a AHE Queimado, ca. 1 km após Palmital, 16°08′59″ S, 47°18′11″ W, 18 Feb 2003 (fl), Amaral-Santos & Pereira 1850 (CEN, MO). Tocantins, Palmas, subbacia do Ribeirão São João, UTM 0818544/8843346, 9 Mar 2006 (fl), Santos & Pereira 328 (HTO); Pium, a 10 km de Barreira da Cruz, próximo a cidade de Pium, 11 Jan 1974 (fl), Rizzo 9335 (UFG). Diplusodon oblongus is represented by much-branched subshrubs, and it can be recognized by the glabrous or covered with minute and inconspicuous trichomes, elliptic, ovate to ovate-lanceolate leaves with acute or attenuate base, which turn dark-brown when dry. The floral tube is oblong with narrow, keeled prophylls, with an almost imperceptible indumentum visible as small protuberances on all parts of the flower. This species also has papillose trichomes that are noticeable on the inner surface of the sepals. Diplusodon oblongus strongly resembles D. appendiculosus in that the whole plant turns dark-brown when dry and also because of the elliptic leaves, oblong floral tube, and fruit with a conspicuous constriction. Diplusodon appendiculosus is distinguished by its epicalyx segments, which are foliaceous, hyaline, and obovate to lanceolate, and also by its pubescent ovary. Lourteig (1989) cited Pohl s.n. (W) as the holotype of Diplusodon oblongus, and according to the code this is an error to be corrected and should be treated as the first-step lectotype (ICBN Art. 9.17, ex. 14). Pohl (1827b) indicated a single gathering of D. oblongus collected in February and March near Vila Boa [now town of Goiás], state of Goiás, without mentioning a collection number. At W there are two specimens of Pohl 5791 from the same locality, and one is annotated by Pohl, which is here designated as a second-step lectotype (W0053425). 57. Diplusodon obtectus T.B.Cavalc., sp. nov. Type: Brazil. Minas Gerais: Itacambira, estradinha para o morro da torre repetidora, 17°4′47″ S, 43°16′35″ W, 5 Feb 2015 (fl, fr), T.B.Cavalcanti, A. C. D. Vieira, L. M. Braunner, G. PereiraSilva & G. L. Moreira 3907 (holotype, CEN [00088651]; isotypes, BHCB, K, NY, SPF, W). Figs. 13.72 A–F, 13.75 This species is distinguished within the genus by the long, spathulate prophylls that extend beyond the flower and hide the floral tube and also by the thick, long epicalyx segments that are spreading and pointing downward. Because of its orbicular to broadly elliptic leaves and androecium with 12 stamens, this species is similar to Diplusodon aggregatifolius, but the latter differs by its wide, lax inflorescences, small prophylls reaching from the middle to the apex of the floral tube, and epicalyx segments that are up to 2.5 mm long. Shrubs 0.4–3  m tall, erect, much-branched, sometimes dark-brown when dry. Branches cylindrical, unwinged, the upper portions wine-colored, internodes

302

13  Systematic Treatment

Fig. 13.72  Diplusodon obtectus. A. Branch with inflorescence. B. Section of internode. C. Leaf, abaxial surface. D. Detail of leaf abaxial surface indumentum. E. Prophyll, adaxial surface. F. Flower with prophylls, without petals. (A–F Cavalcanti et al. 3907, CEN)

Descriptions

303

0.6–1 cm long. Leaves decussate, not imbricate, suberect, petiolate; petioles 1–2 mm long; blades 11–20 × 7–15 mm, coriaceous, not discolorous, sometimes dark-brown when dry, eucamptodromous, broadly elliptic to orbicular, base obtuse to acute, apex obtuse, margin plane, adaxial surface glabrous, concave when fresh, abaxial surface weakly pubescent; secondary veins 2–3 on each side, departing along the midvein, inconspicuous; domatia absent. Synflorescence frondose, diplobotryum to pleiobotryum type, congested; accessory branches absent, principal florescence truncate, paraclades 4–14 cm long; bracts similar in shape to the vegetative leaves; flowers pedicellate, hypopodium 3–5  mm long, epipodium 0.5–1  mm long; prophylls 8–10  ×  3–5  mm, surpassing the apex of the floral 1–1.5  mm, spathulate, unkeeled, base attenuate, margin plane, pubescent on the vein in adaxial surface; floral tube 6–6.5  mm long, funnel-shaped, finely and sparsely pubescent; sepals 2–2.1 mm long, triangular, not caudate, not deflexed, glabrous or with sparse fine trichomes; epicalyx segments 3.5–5  mm long, cylindrical, thick, spreading to deflexed, longer than the sepals, minute pilose; corolla 3–4  cm in diam., rosemagenta, petals 13–14 × 7–8 mm, obovate, apex obtuse; stamens 12, free portion of the filaments ca. 6 mm long, slightly exserted; ovary ca. 2.3 × 2.5 mm, globose, glabrous, style 11–12 mm long, ovules ca. 10. Capsules 3.5–5 × 4–4.2 mm, oblong, glabrous; seeds 2–9, 3–3.5 × 2.1–2.9 mm. Distribution and Ecology (Fig. 13.75)  Diplusodon obtectus occurs in the municipality of Itacambira, Cadeia do Espinhaço, northern Minas Gerais. It inhabits campo rupestre, with a population of numerous individuals growing on white sandy soil at 1220–1383 m. Flowers from January to April; fruits from February to April. Specimens Examined: Minas Gerais,  Itacambira, estrada para Montes Claros, 9 Jan 1986 (fl), Cordeiro et al. (CFCR) 9122 (CEN, SPF); Itacambira, 2 km W da cidade, na rodovia para Juramento, no alto da serra, 17°4′ S, 43°18′ W, 14 Feb 1988 (fl), Pirani et  al. 2270 (CEN, NY, SPF), Itacambira, estrada Itacambira-Montes Claros, 4 km de Itacambira, subindo a chapada, 17°4′ S, 43°19′ W, 30 Apr 1997 (fl, fr), Cavalcanti et al. 2332 (CEN, DIAM, MO, RB, SP, US). The specific epithet alludes to the main feature of this species, which are the large bracts that cover the flowers. Additionally, this species can be recognized by the broadly elliptic to orbicular hyphodromous leaves; pedicellate flowers; thick, cylindrical, spreading to deflexed epicalyx segments that are 3–5 mm long; and the androecium with 12 stamens. Vegetatively, Diplusodon obtectus resembles D. aggregatifolius, which is also from northern Minas Gerais State, because both have broadly elliptic to orbicular leaves and the whole plants are inconspicuously pubescent. However, D. aggregatifolius has broad, lax synflorescences, prophylls reaching the middle or the top of the floral tube, and smaller flowers (floral tube 4–5 mm long) with epicalyx segments 1.5–2.5 mm long. 58. Diplusodon orbicularis Koehne, Fl. Bras. (Martius) 13(2): 322, pl. 60, fig. 5, 1877. Type: Brazil. Minas Gerais: montosis altis saxosis prope Diamantina, L. Riedel s.n. (lectotype, LE [010113838], here designated; photo CEN).

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Figs. 13.73 A–M, 13.75 Diplusodon orbicularis Koehne var. brachyander Koehne, Bot. Jahrb. Syst. 29: 164, 1900. Type: Brazil. Minas Gerais: Serra do Cipó, 22 Apr 1892 (fl, fr), A.  F. M. Glaziou 19167 (first-step lectotype, designated by Lourteig 1989: 235, second-step lectotype, P [01901866], here designated; isolectotypes, BR, K, LE, P [01901867]). Subshrubs 0.15–2 m tall, erect, branched, glabrous, not dark-brown when dry. Upper branches yellow, cylindrical, unwinged, lax, internodes 0.5–1  cm long. Leaves decussate, not imbricate, spreading, subsessile; petioles 0.5–1  mm long; blades 5–10(−15) × 5–18 mm, glaucous, coriaceous, not discolorous, acrodromousbasal-typical, cordiform to oblate-cordiform, base cordate or rarely obtuse, apex obtuse, frequently acuminate, margin plane, sometime wine-colored; secondary veins 2–3 on each side, departing at the base of the midvein; domatia absent. Synflorescence frondose, diplobotryum to triplobotryum type, lax; accessory branches absent, principal florescence 8–9  cm long, often truncate, paraclades 6–25 cm long; bracts similar to vegetative leaves in shape and size; flowers pedicellate, hypopodium 1.5–4  mm long, epipodium 0.5–1  mm long; prophylls 4–5 × 3–4 mm, slightly shorter than or reaching the apex of the floral tube, ellipticlanceolate to rhomboid-ovate, unkeeled, base obtuse to rarely acute, apex obtuse to acute, margin plane; floral tube 4–5 mm long, clear green with wine-colored apex, funnel-shaped; sepals 2–3 mm long, wine-colored, triangular, slightly caudate, not deflexed; epicalyx segments 2–2.5 mm long, cylindrical to slightly dorsiventrally compressed, erect to suberect, shorter than the sepals; corolla clear rose to lilac, 3–3.5 cm in diam., petals 12–15 × 8–9 mm, obovate, apex obtuse; stamens 6, free portion of the filaments rose to wine-colored, 7–9  mm long, exserted; ovary 1–1.5 × 1.2–1.5 mm, obconic to orbicular, glabrous, style 13–15 mm long, ovules 15–25. Capsules 3–4.5 × 3.1–4, globose, glabrous; seeds 12–17, 2.3–2.5 × 2–2.5 mm. 2n = 30 (Graham and Cavalcanti 2001). Distribution and Ecology (Fig.  13.75)  Diplusodon orbicularis occurs in the region of Serra do Cipó, Cadeia do Espinhaço, Minas Gerais, where it is locally common and known by the vernacular name of “cai-cai.” The species has its northern limit in the municipality of Guanhães. It grows in campos limpos, campos rupestres, and campos sujos, associated with sandy soils and sandstones, at 900–1400 m. Flowers from January to August; fruits from February to November. Specimens Examined: BRAZIL. Minas Gerais,  Guanhães, 21 Mar 1976 (fr), Silva s.n. (OUPR 22918); Guanhães, 21 Apr 1976 (fl), Silva s.n. (OUPR 1807); Jaboticatubas, Serra do Cipó, 11 Mar 1969 (fl), Eiten & Eiten 11028 (MO); Jaboticatubas, Serra do Cipó, 10 Apr 1980 (fl), Barros 212 (CEN); Jaboticatubas, Serra do Cipó, Souza & Abbud 33239 (ESA); Jaboticatubas, Parque Nacional Serra do Cipó, Cachoeira da Farofa, Portaria do Retiro, 19° 20′ 56′′S, 43° 34′ 38′′W, 16 Out 2013 (fr), Verdi et al. 6734 (CEN, RB); Serra do Cipó, km-01, ao lado esquerdo da estrada, 2 Apr 1983 (fl, fr), Silva et  al. 496 (CEN, VIC n.v.); Presidente Kubitscheck, 6 km de Presidente Kubitscheck, em direção a Serro, 5 Jul 1996 (fl,

Descriptions

305

Fig. 13.73  Diplusodon orbicularis. A. Branch with inflorescence. B–D. Leaves, abaxial surface. E–F. Prophylls, adaxial surface. G. Detail of inflorescence. H. Floral bud. I. Flower with prophylls, without petals. J. Petal. K. Fruit. L. Capsule. M. Seed. (A, E, F, H–J Amaral et al. (CFSC) 9322, SPF; B Duarte 2494, RB; C, D Bautista 593, SPF; G, K–M Giulietti et  al. (CFSC) 7316, SPF)

306

13  Systematic Treatment

fr), Souza et al. 11810 (ESA); Santana do Riacho, 15 Apr 1985 (fl, fr), Andrade & Lopes s.n. (BHCB 8818, CEN); Santana do Riacho, 20 Oct 1990 (fl, fr), Stehmann et al. s.n. (BHCB 18870); Santana do Riacho, estrada Lagoa Santa-Conceição do Mato Dentro, km 110, 8 Jun 1989 (fl, fr), Castro s.n. (BHCB 2255, CEN); Santana do Riacho, estrada Lagoa Santa-Conceição do Mato Dentro Castro s.n. (BHCB 1097); Santana do Riacho, estrada Lagoa Santa-Conceição do Mato Dentro, Castro s.n. (BHCB 22458, CEN, HUFU); Santana do Riacho, estrada Lagoa SantaConceição do Mato Dentro, 19 May 1990 (fl), Castro s.n. (BHCB 18378); Santana do Riacho, estrada Lagoa Santa-Conceição do Mato Dentro, 3 Mar 1981 (fl, fr), Castro 26 (BHCB); Santana do Riacho, córrego Duas Pontes, MG-010, ca. 19°17′27.1″ S, 43°34′5″ W, 10 Jun 2002 (fl), Groppo, Jr. et al. 1099 (CEN, SPF); Santana do Riacho, Serra do Cipó, 19°17′ S, 43°35′ W, 15 May 1998 (fl, fr), Jacobi 51 (CEN, BHCB); Santana do Riacho, Serra do Cipó, 22 Mar 1992 (fl), Pereira & Lucca 929 (BHCB, CEN); Santana do Riacho, Serra do Cipó, 1 Oct 1989, Souza & Abbub 12331 (ESA); Santana do Riacho, Serra do Cipó, Chapéu do Sol, ca. 9 km após Chapéu do Sol, 19°17′25″ S, 43°34′02″ W, 10 Mar 1995 (fl, fr), Souza et al. 8182 (ESA); Santana do Riacho, Serra do Cipó, 19° 40′ S, 43° 57′ W, 29 Apr 1952 (fl), Smith et al. 7024 (B, K, NY, US); Santana do Riacho, Serra do Cipó, 12 May 1990 (fr), s.n. (BHCB 17991, CEN 19167); Santana do Riacho, Serra do Cipó, MG-010, 19º20′S, 43º37′W, 21 May1982 (fl, fr), Bautista 593 (SPF); Santana do Riacho, Serra do Cipó, rodovia Belo Horizonte-Conceição do Mato Dentro, km 100, 19°18′87″ S, 43°36′43″ W, 23 Apr 1997 (fl), Cavalcanti et al. 2271 (CEN, MO, NY); Santana do Riacho, Serra do Cipó, rodovia Belo Horizonte-Conceição do Mato Dentro, km 103, 25 Mar 1995 (fl), Castro 395 (HUFU); Santana do Riacho, Serra do Cipó, rodovia Belo Horizonte-Conceição do Mato Dentro, km 105, (ca. 130 km N de Belo Horizonte), 17 Feb 1968, Irwin et al. 20370 (NY); Santana do Riacho, Serra do Cipó, rodovia Belo Horizonte-Conceição do Mato Dentro, km 106, 20 Apr 1985 (fl), Amaral et al. (CFSC) 9322 (SPF); Santana do Riacho, Serra do Cipó, rodovia Belo Horizonte-Conceição do Mato Dentro, km 106, 8 May 1987 (fl), Cavalcanti et al. (CFSC) 10097 (SPF); Santana do Riacho, Serra do Cipó, rodovia Belo Horizonte-Conceição do Mato Dentro, km 109, 6 Sep 1980 (fr), Forero et  al. (CFSC) 8725 (SPF); Santana do Riacho, Serra do Cipó, rodovia Belo Horizonte-Conceição do Mato Dentro, km 109, 2 Aug 1990 (fl), Sakuragui et al. 34 (ESA); Santana do Riacho, Serra do Cipó, rodovia Belo Horizonte-Conceição do Mato Dentro, km 110, 8 Jun 1989 (fl, fr), Barbosa 392 (BHCB, CEN [2], HUFU); Santana do Riacho, Serra do Cipó, rodovia Belo Horizonte-Conceição do Mato Dentro, entre km 110 e 112, 8 Apr 1977 (fl, fr), Menezes (CFSC) 7318 (F, SPF); Santana do Riacho, Serra do Cipó, rodovia Belo Horizonte-Conceição do Mato Dentro, km 111–120, 6 Aug 1960 (fr), Maguire et  al. 49040 (NY); Santana do Riacho, Serra do Cipó, rodovia Belo Horizonte-Conceição do Mato Dentro, km 112, 11 Jul 1965 (fr), Glassman & Gomes Jr. 8074 (SP); Santana do Riacho, Serra do Cipó, rodovia Belo Horizonte-Conceição do Mato Dentro, km 112, 2 May 1986 (fl), Cavalcanti et al. (CFSC) 9658 (SPF); Santana do Riacho, Serra do Cipó, rodovia Belo Horizonte-Conceição do Mato Dentro, km 12.5, 15 Apr 1972 (fl, fr), Joly et  al. (CFSC) 1423 (CEN, SP); Santana do Riacho, Serra do Cipó, rodovia Belo

Descriptions

307

Horizonte-Conceição do Mato Dentro, km 113, 15 Apr 1972 (fl, fr), Joly et  al. (CFSC) 1080 (SP); Santana do Riacho, Serra do Cipó, rodovia Belo HorizonteConceição do Mato Dentro, km 113, 15 Apr 1972 (fl, fr), Joly et al. (CFSC) 1517 (CEN, SP); Santana do Riacho, Serra do Cipó, rodovia Belo Horizonte-Conceição do Mato Dentro, km 114, 5 Jun 1970 (fl, fr), Joly et al. (CFSC) 1 (CEN, SP); Santana do Riacho, Serra do Cipó, rodovia Belo Horizonte-Conceição do Mato Dentro, km 114, 30 Apr 1972 (fl), Semir & Sazima (CFSC) 2046 (SP); Santana do Riacho, Serra do Cipó, rodovia Belo Horizonte-Conceição do Mato Dentro, km 114, 29 Apr 1973 (fl), Semir et al. (CFSC) 4047 (SP); Santana do Riacho, Serra do Cipó, rodovia Belo Horizonte-Conceição do Mato Dentro, km 114, 31 Oct 1985 (fr), Cavalcanti et al. (CFSC) 9376 (F, SPF); Santana do Riacho, Serra do Cipó, rodovia Belo HorizonteConceição do Mato Dentro, km 114, 30 May 1988 (fl), Menezes & Oliveira (CFSC) 11075 (SPF); Santana do Riacho, Serra do Cipó, rodovia Belo Horizonte-Conceição do Mato Dentro, km 14.5, após o córrego Duas Pontinhas, 26 Jun 1995 (fr), Salatino et al. 28 (CEN, SPF); Santana do Riacho, Serra do Cipó, rodovia Belo HorizonteConceição do Mato Dentro, km 115 antigo, 11 Mar 1989 (fl), Carmo (CFSC) 11272 (SPF); Santana do Riacho, Serra do Cipó, rodovia Belo Horizonte-Conceição do Mato Dentro, km 115, 30 Mar 1990 (fl), Oliveira & Bertoluci (CFSC) 11865 (SPF); Santana do Riacho, Serra do Cipó, rodovia Belo Horizonte-Conceição do Mato Dentro, km 116, 6 Jun 1970 (fl), Joly et al. (CFSC) 121 (SP); Santana do Riacho, Serra do Cipó, rodovia Belo Horizonte-Conceição do Mato Dentro, km 116, 6 May 1970 (fr), Joly et al (CFSC) 132 (SP); Santana do Riacho, Serra do Cipó, rodovia Belo Horizonte-Conceição do Mato Dentro, km 120, 1 May 1972 (fl), Semir & Sazima (CFSC) 2056 (CEN, SP); Santana do Riacho, Serra do Cipó, rodovia Belo Horizonte-Conceição do Mato Dentro, km 120, 26 Mar 1988 (fl), Menezes (CFSC) et  al. 11052 (SPF); Santana do Riacho, Serra do Cipó, rodovia Belo Horizonte-­ Conceição do Mato Dentro, ca. km 120, (ca. 145 km N de Belo Horizonte, 14 Feb 1968 (fr), Irwin et al. 19996 (NY); Santana do Riacho, Serra do Cipó, rodovia Belo Horizonte-Conceição do Mato Dentro, km 122, 29 May 1972 (fl, fr), Joly et  al. (CFSC) 2388 (SP); Santana do Riacho, Serra do Cipó, rodovia Belo HorizonteConceição do Mato Dentro, km 125, 7 Nov 1987 (fr), Cavalcanti et  al. (CFSC) 10622 (SPF); Santana do Riacho, Serra do Cipó, rodovia Belo Horizonte-Conceição do Mato Dentro, km 125, 7 Nov 1987 (fr), Cavalcanti et al. (CFSC) 10623 (SPF); Santana do Riacho, Serra do Cipó, rodovia Belo Horizonte-Conceição do Mato Dentro, córrego Três Pontinhas, 27 Oct 1988 (fr), Menezes et  al. (CFSC) 11258 (CEN, SPF); Santana do Riacho, Serra do Cipó, rodovia Belo Horizonte-Conceição do Mato Dentro, km 127, 16 Mar 1962 (fl), Duarte 6499 (K, NY, RB); Santana do Riacho, Serra do Cipó, rodovia Belo Horizonte-Conceição do Mato Dentro, km 128, 19 Apr 1050 (fr), Duarte 2742 (RB); Santana do Riacho, Serra do Cipó, rodovia Belo Horizonte-Conceição do Mato Dentro, km 128, 5 Mar 1972 (fl, fr), Joly et al. (CFSC) 1104 (SP); Santana do Riacho, Serra do Cipó, rodovia Belo HorizonteConceição do Mato Dentro, km 128, 5 Mar 1972 (fl), Joly et al. (CFSC) 1122 (SP); Santana do Riacho, Serra do Cipó, rodovia Belo Horizonte-Conceição do Mato Dentro, km 128, 5 Mar 1972 (fl), Joly et al. (CFSC) 1153 (SP); Santana do Riacho, Serra do Cipó, rodovia Belo Horizonte-Conceição do Mato Dentro, km 131, 16 Apr

308

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1935 (fl), Mello-Barreto 1268 & Brade 14881 (RB); Santana do Riacho, Serra do Cipó, rodovia Belo Horizonte-Conceição do Mato Dentro, km 131–134, 15 Apr 1950 (fl, fr), Duarte 2494 (F, MO, RB); Brade 14406 (MO); Santana do Riacho, Serra do Cipó, Estrada da Usina, 7 Jan 1973 (fl), Semir & Joly (CFSC) 3815 (SPF, UEC); Santana do Riacho, Serra do Cipó, km 109, Cachoeira do Sr. Nicolau, 3 Mar 1981 (fl), Castro & Mayo (CFSC) 7192 (SPF, UB); Santana do Riacho, Serra do Cipó, 26 Jan 1986 (fl), Zappi et al. (CFSC) 9654 (SPF, UEC); Santana do Riacho, Serra do Cipó, 22 Feb 1986 (fl), Cavalcanti et al. (CFSC) 9619 (F, SPF), 8 May 1987 (fl), (CFSC) 10139 (SPF), 10 May 1987 (fl), (CFSC) 10163 (SPF); Santana do Riacho, Serra do Cipó, 8 May 1987 (fl), Cavalcanti et  al. (CFSC) 10139 (SPF); Santana do Riacho, Serra do Cipó, 10 May 1987 (fl), Cavalcanti et al (CFSC) 10163 (SPF); Santana do Riacho, Parque Nacional da Serra do Cipó, Reserva do IBDF, caminho para a Cachoeira da Farofa, 21 Feb 1986 (fl, fr), Cavalcanti et al. (CFSC) 9614 (SPF); Santana do Riacho, Serra do Cipó, Serra das Bandeirinhas, Costa, caminho para a Cachoeira das Flores, 9 Nov 1987 (fr), Cavalcanti et al. (CFSC) 10635 (F, SPF); Santana do Riacho, Serra do Cipó, Alto do Palácio, 28 Mar 1996 (fl), Castellois 477 (PAMIG); Santana do Riacho, Serra do Cipó, Alto do Palácio, 30 May 1981 (fl), Martins s.n. (BHCB 8490, CEN); Santana do Riacho, Serra do Cipó, estrada Santana do Riacho-Lapinha, 4.4  km de Santana do Riacho, 19°08′23″ S, 43°41′41″ W, 22 Apr 2006 (fl), Mello-Silva et  al. 2895 (CEN, SPF); Santana do Riacho, Serra do Cipó, Apr 1977 (fl, fr), Giulietti (CFSC) 7316 (SPF); Santana do Riacho, Serra do Cipó, próximo ao córrego Vitalino, 21 Mar 1989 (fl, fr), Lewinger et al. (CFSC) 11284 (CEN [2], SPF); Serra do Cipó, 17 Feb 1972 (fl), Anderson et al. 36117 (NY); Várzea da Palma, Serra do Cabral, Agropecuária Serra do Cabral, 16 Jan 1996 (fl), Hatschbach et al. 64214 (BR, CEN, FLOR, MBM, NY); without reference to municipal district: 26 Apr 1896 (fl), Schwacke 8265 (OUPR, RB); 6 Mar 1958 (fl), Occhioni 1842 (HB, MBM); 16 Jan 1976 (fr), Silva s.n. (OUPR 22708). Diplusodon orbicularis is characterized by glaucous, cordiform leaves and acrodromous-basal-type venation. These characteristics are shared also by D. glaucescens, which has slightly larger leaves with 5–6 secondary veins on each side of the midrib and an androecium with 12 stamens (whereas D. orbicularis has only 6 stamens). The seedlings sometimes have trichomes on the leaves. Koehne received Riedel’s collection at B for his studies on Lythraceae, but from photographs of type material of Diplusodon orbicularis from the Berlin herbarium (B), probably no material of D. orbicularis identified by Koehne was retained. Specimens were transferred from B to LE and had already been identified by Koehne. One specimen at LE annotated by Koehne as “Diplusodon orbicularis” and with a short description with Koehne’s handwriting is here selected as the lectotype (LE 010113838). Koehne (1900) described Diplusodon orbicularis var. brachyander and cited the syntypes Glaziou 19167 and Schwacke 8265, collected in Serra do Cipó. Koehne distinguished the new variety by its strongly cordate leaf base. I studied several populations of D. orbicularis on Serra do Cipó (Cavalcanti 1988), as well as the

Descriptions

309

syntypes. All of Koehne’s diagnostic characteristics for D. orbicularis var. brachyander, including the shape of the leaf base, ratio of stamen length to sepal length, and ratio of prophyll length to floral tube length, are continuous characteristics within the range of D. orbicularis and do not consistently distinguish the variety. Lourteig (1989) cited Glaziou 19167 at P as the holotype, which is here considered as the first-step lectotype (ICBN Art. 9.17, ex. 14). There are two specimens of Glaziou 19167 at P, and the second-step lectotypification is here performed (P01901866). 59. Diplusodon ovatus Pohl, Flora 10: 131. 1827. Type: Brazil. Minas Gerais: in campis graminosis ad Arraial de Bambuia et Nossa Senhora do Patrocínio, Oct/ Nov 1819, J. B. E. Pohl 589 = D 940 (first-step lectotype, designated by Lourteig, 1989: 234, second-step lectotype, W [0053427], here designated; isolectotypes, BR, G, K [2], M, OXF, W [0053426]). Figs. 13.74 A–J, 13.75, 15.3 F Subshrubs 0.4–0.9 m tall, erect, unbranched, sometimes caespitose, with a xylopodium, glabrous, not dark-brown when dry. Upper branches cylindrical, flattened, unwinged, internodes 3–8 cm long. Leaves decussate, rarely 3-verticillate at basal nodes, not imbricate, suberect, petiolate; petioles 1–2  mm long; blades 30–70 × 25–50 mm, coriaceous to chartaceous, not discolorous, not glaucous, acrodromous-basal to acrodromous-basal-supranumerary, ovate to ovate-lanceolate or rarely orbicular, base obtuse to acute, apex acute to obtuse, margin plane, ciliate on the older leaves; secondary veins 3–5 on each side, 3–4 of them departing at the base of the midvein, prominent on the abaxial surface; domatia absent. Synflorescence bracteose, of the thyrsoid type, sometimes thyrse, lax; accessory branches absent, paraclades short with terminal cymes; bracts 7–30  ×  2–14  mm, ovate to elliptic; flowers pedicellate, hypopodium 2.5–6 mm long (7 mm in fruit), epipodium 2–4 mm long (7  mm in fruit); prophylls 5.5–8  ×  1–2  mm, reaching from the base to the middle of the floral tube, linear, unkeeled, base retuse, apex acute, margin plane; floral tube 6–6.5, in fruit 8–10 mm, funnel-shaped; sepals 3–4 mm long, triangular, slightly caudate, sometimes the top deflexed, slightly thickened, rarely with some trichomes on the margin; epicalyx segments 2–3 mm long, cylindrical, spreading, the same size as or shorter than the sepals; corolla 2.5–3 mm in diam., rose, petals 20–23 × 12–16 mm, obovate, apex obtuse; stamens 18, free portion of the filaments 8.5–9 mm long, included to exserted; ovary 2–2.5 × 3–3.5 mm, rhombic, glabrous, style 17–18 mm long, ovules 46–59. Capsule 5–6 × 5.5–6 mm, wine-colored, globose, glabrous; seeds 10–40, 2–3 × 2–2.5 mm. Distribution and Ecology (Fig. 13.75)  Diplusodon ovatus has a wide geographic distribution in the cerrados of São Paulo, Mato Grosso, Mato Grosso do Sul, Minas Gerais, and Goiás, up to 900 m (in Goiás). Flowers from November to May; fruits from November to June. Specimens Examined: BRAZIL.  Goiás,  Campo Alegre, 28 Nov 1992, Hatschbach & Barbosa 58271 (MBM, P); Catalão, Serra do Facão, ca. 25 km de Catalão, 24 Jan 1970 (fr), Irwin et  al. 25261 (K, MO, NY, P, RB, UB); Catalão, Balsa das Gamelas, 2 Nov 1988 (fr), Brandão 14377 (PAMIG). Mato Grosso,

310

13  Systematic Treatment

Fig. 13.74  Diplusodon ovatus. A. Habit. B. Detail of branch. C–D. Leaves, abaxial surface. E. Prophyll, adaxial surface. F, H. Floral bud. G. Terminal flower. I. Flower with prophylls, without petals. J. Pistil. (A–C, E, G–J Hatschbach & Koczicki 33061, MBM; D, F Löefgren & Edwall 2178, SP)

Descriptions

311

Santana da Chapada, 1903 (fl), Robert 571b (K). Mato Grosso do Sul, Camapuã, Capão Redondo, 11 Nov 1973 (fl), Hatschbach & Koczicki 33061 (MBM, MO, NY, P). Minas Gerais, São Roque de Minas, estrada da Cachoeira dos Rolinhos, 21 Mar 1996 (fl), Nakajima & Romero 1699 (CEN HUFU); São Roque de Minas, Parque da Serra da Canastra, 16 Dec 1998 (fl, fr), Farinaccio & Campos-Filho 256 (SPF); São Roque de Minas, Parque Nacional Serra da Canastra, após a entrada para o Retiro das pedras, 11 Jan 2003 (fr), Romero et al. 1727 (CEN, HUFU); São Roque de Minas, estrada para Sacramento, após o Curral de Pedras, 10 Mar 1995 (fr), Nakajima et al. 876 (CEN, HUFU); São Roque de Minas, Parque Nacional da Serra da Canastra, morro antes do córrego dos Passageiros, 10 Jan 1998 (fl), Romero et al. 4968 (HUFU); Uberaba, Nov 1916 (fr), Brito 50 (R); Uberaba, 14 Jan 1996, Hatschbach et al. 64063 (CEN, MBM); Uberaba, 28 km de Uberaba, 15 Oct 1966, Duarte 783 (P); Uberaba, ca. 48 km SE de Uberlândia, 11 Mar 1982 (fr), Graham 861 (MO, NY); entre o rio São Francisco e Curvelo, Riedel 2596 (K, LE, P); entre Rio Grande e Tejuco (Sul de Minas Gerais), May 1865 (fl), Burchell 5722 (BR, K, P); Uberlândia, fazenda das Perobas, 250 m da BR-497, na altura do km 18, 26 Dec 2002 (fl), Oliveira 2139 (HUFU). São Paulo, Franca, 16 Jan 1983 (fl), Loefrgren & Edwall 2178 (SP); São Paulo, Regnell 55 (P). Diplusodon ovatus is a glabrous species with flattened branches and wide, ovate leaves. It has a thyrsoid inflorescence with short lateral cymes, linear prophylls, and flowers with 18 stamens. These characteristics separate it from D. speciosus, another glabrous species with similar habit and distribution, but which has botryum to diplobotryum inflorescences, ovate prophylls, and flowers with 30–38 stamens. The presence of a terminal flower on the main axis of the inflorescence is a previously unrecognized feature in the genus. In other species, the branches of the inflorescence are described as lateral cymes, but the main axis is indeterminate. Diplusodon ovatus was described by Pohl (1827a) without citation of specimens in the protologue. Shortly after, Pohl (1827b) cited a single gathering of D. ovatus collected in October–November in the grassy fields near Bambuia and Nossa Senhora do Patrocínio, Minas Gerais, without mentioning a collection number. Lourteig (1989) cited Pohl 940 as the holotype, which should be interpreted as a lectotypification (ICBN Art. 9.17, ex. 14). I examined two type materials of Diplusodon ovatus at W: one has a simple label only with the species name; the other, chosen as lectotype, has an original label, annotated by Pohl, including the locality cited in the protologue (Pohl 1827b), his collection number, Pohl 589, and the number cited in his diary (D 940). There are specimens of Diplusodon ovatus in several herbaria with labels indicating Pohl s.n., some without locality, some with part of the locality, and the specimen at M with a clear indication that the material came from W. Because duplicates of Pohl’s collections were distributed to other herbaria without the information from his field diary (see “Notes about Diplusodon nomenclature and typifications”) and most lack either the collection or diary number, the specimens at BR, G, K, M, OXF, and W are regarded as duplicates of Pohl 589 = D940.

312

13  Systematic Treatment

Fig. 13.75  Geographic distribution of Diplusodon oblongus, D. obtectus, D. orbicularis, and D. ovatus

Descriptions

313

60. Diplusodon panniculatus Koehne, Bot. Jahrb. Syst. 23(Beibl. 57): 30. 1897. Type: Brazil. Goiás: Santa Luzia [now Luziânia], inter cabeceira rio Samambaia, 19 Sep 1894 (fl), A. F. M. Glaziou 21416 (lectotype, BR [5178995], here designated; isolectotypes, K, P, R). Figs. 13.76 A–G, 13.80 Subshrubs 0.6–1 m tall, erect, branched just at inflorescence, with a xylopodium, not dark-brown when dry. Branches lax, the upper ones cylindrical to subquadrangular, unwinged or sometimes with a narrow wing, glabrous, internodes 2–5.5 cm long. Leaves decussate, not imbricate, erect, subsessile; petioles broad, to 1  mm long; blades 25–55 × 7–25 mm, coriaceous, glabrous, not discolorous, acrodromousbasal-­ supranumerary, elliptic-lanceolate to ovate-lanceolate, base obtuse, apex acute, margin plane, sometimes ciliate; secondary veins 3 on each side, 2 of them departing from the base of the midvein, tertiary veins forming a prominent reticulum on both surfaces of the blade, more prominent on the adaxial surface; domatia absent. Synflorescence bracteose, triplothyrsoid to pleiothyrsoid type, lax; accessory branches absent, paraclades 4–20  m long, glabrous, from the second order onward pubescent to glabrous; bracts of the first order 4–20 × 2–8 mm, elliptic to elliptic-lanceolate, glabrous, those of the cymes or dichasia, 2–4 × ca. 1 mm, linear, pubescent; flowers pedicellate, hypopodium 5–6  mm long, epipodium 4–5  mm long; prophylls 2–2.8 × 1–1.8 mm, not even reaching the base of the floral tube, linear, keeled, base retuse, apex acute, margin plane, pubescent; floral tube 4–6 mm long, grayish purple, funnel-shaped, pubescent; sepals 4–5 mm long, triangular, not caudate, sometimes the top deflexed, pubescent; epicalyx segments 1.5–2.2  mm long, cylindrical, spreading to deflexed, shorter than the sepals; corolla 3–4.5 cm in diam., rose-magenta, petals 15–17  ×  10–12  mm, obovate, apex obtuse; stamens 15–18, free portion of the filaments 6–8  mm long, exserted to included; ovary 2–2.5  ×  2.5–4  mm, obovate-depressed, glabrous, style 10–12  mm long, ovules 28–30. Capsules 5–5.5  ×  4–4.5  mm, globose, glabrous; seeds 18–26, 1.8–2 × 1.5–1.9 mm. Distribution and Ecology (Fig.  13.80)  Diplusodon panniculatus occurs in Luziânia, Serra dos Cristais in the municipality of Cristalina, and Campo Alegre, in Goiás. It grows in cerrado and campos rupestres at 1100–1250  m. Flowers from April to September; fruits from July to November. Specimens Examined: BRAZIL.  Goiás,  Campo Alegre, BR-050, BrasíliaCatalão, margem esquerda do rio Imbiruçu, 17°24′30.9″ S, 47°46′58.1″ W, 7 May 2016 (fl), Cavalcanti et al. 3988 (CEN). Cristalina, área ao lado da entrada para o restaurante Milho Verde, 10 Jul 2003 (fr), Cavalcanti & Pereira-Silva 3182 (CEN, MO); Cristalina, 31 Jul 1985 (fl, fr), Romaniuc Neto & Sajo 388 (CEN, NY, SP); Cristalina, BR-040, 1 km SE de Cristalina, km 107, 24 Jul 1984 (fl), Mori et al. 16947 (CEN, CEPEC, K, NY, P, UB); Cristalina, BR-040, Brasília-Cristalina, 2 km de Cristalina, 16°43′59″ S, 47°37′16″ W, 21 Nov 2005 (fr), Cavalcanti et al. 3649 (CEN, MO); margem da BR-040 ca. 5  km ao sul de Cristalina, 16°47′12″ S, 47°34′31″ W, 30 Jul 2011 (fl), Faria-Jr. et  al. 1458 (CEN, UB). Cristalina, São

314

13  Systematic Treatment

Fig. 13.76  Diplusodon panniculatus. A. Branch with inflorescence. B. Vegetative branch. C. Leaf, abaxial surface. D. Cyme. E. Prophyll, adaxial surface. F. Flower with prophyll, without petals. G. Ovary. (A, D, G Mori et al. 16947, CEN; B, E–F Romaniuc & Sajo 388, SP; C Irwin et al. 9717, NY)

Descriptions

315

Topázio, 20 km antes de Cristalina, rod. Brasília-Belo Horizonte, 25 May 1973 (fl), Rizzo 9051 (UFG); Cristalina, Rizzo 9102 (UFG); Cristalina, Serra dos Cristais, 5 km S de Cristalina, 1 Nov 1965 (fr), Irwin et al. 9717 (NY, P, UB); Cristalina, Fazenda Lopo Botelho, 7 Jul 1963 (fr), Pires & Mattos 9820 (UB); Cristalina, Serra dos Topázios, 16°45′ S, 47°40′ W, 3 Jun 1996, Proença & Oliveira 1416 (UB); Cristalina, entroncamento da BR 050 com a rodovia de acesso à BR 251, cerca de 100 m a leste da BR, 16°43′54″ S, 47°37′14″ W, 17 Jun 2003 (fl), Pereira-Silva et al. 7928 (CEN, MO); Cristalina, área próxima à Reserva Ecológica, 17 Apr 2004 (fl), Pastore et  al. 1008 (CEN); without locality, Jul–Aug 1877, Reichardt 110 (W), without locality, Reichardt 111 (W). Diplusodon panniculatus has a monotelic inflorescence that is bracteose, broad, and much-branched, with paraclades bearing cymes of triads or dichasia. This species stands out in the genus, together with D. ovatus, in having a terminal flower on the distal part of the principal florescence, referred to generally as the monotelic condition (Weberling 1965). Other characteristics of Diplusodon panniculatus are the totally glabrous vegetative branches and leaves, elliptic-lanceolate to ovate-lanceolate leaves, and the presence of short indumentum on the 2° order and succeeding paraclades. The flowers are 3–4.5 cm in diameter, with linear keeled prophylls, short, spreading to deflexed epicalyx segments (1–2.2 mm long), and an androecium with 15–18 stamens. Diplusodon panniculatus has a habit similar to that of D. punctatus with respect to the shape of the leaves, absence of indumentum on the leaves, and a bracteose synflorescence. The latter species differs from D. panniculatus principally because it has acrodromous leaves, the absence of a terminal flower, cymes in the inflorescence, the absence of an epicalyx, and the presence of indumentum on the flowers, in addition to an androecium with 12 stamens. Koehne (1897) described Diplusodon panniculatus using a specimen that was destroyed at B (F neg. nº 017928). There are four duplicates of Glaziou 21416 in Brazilian and European herbaria, and just the specimen at BR is annotated by Koehne as “Diplusodon panniculatus sp. nov.” and is chosen as lectotype. 61. Diplusodon paraisoensis Lourteig, Bradea 5(19): 240, pl. 3, figs. a–f. 1989. Type: Brazil. Goiás: Chapada dos Veadeiros, Alto Paraíso de Goiás, 21 Mar 1969 (fl), H. S. Irwin, R. R. Santos, R. Souza & S. F. Fonseca 24793 (holotype, P [01901864]; isotype, NY). Figs. 13.77 A–F, 13.80, 15.4 A–B Diplusodon lanceolatus Pohl var. alutaceus (Pohl) Koehne f. foliosus Koehne, Fl. Bras. (Martius) 13 (2): 330. 1877. Diplusodon foliosus (Koehne) T.B.Cavalc., nom. illeg., Bol. Bot. Univ. São Paulo 22(1): 5, fig. 3. 2004. Type: Brazil. Goiás: Without locality, 1841, G.  Gardner 3721 (first-step lectotype, designated by Lourteig 1989: 227, second-step lectotype, P [01901884], designated by Cavalcanti 2004: 5; isolectotypes, F, G [2], K, NY [2], OXP, P [01901882, 01901883], US, W [2]).

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Fig. 13.77  Diplusodon paraisoensis. A. Branch with inflorescence. B. Leaf, abaxial surface. C. Flower with prophyll, without petals. D. Epicalyx segment. E. Trichomes with thickened base. F. Simple trichome. (A, C–F Cavalcanti et al. 390, CEN; B Gardner 3721, K)

Descriptions

317

Subshrubs 0.4–3 m tall, stem 1–1.5 cm diam., erect, branched, with a xylopodium, not dark-brown when dry. Upper branches subquadrangular, sometimes bright gray, unwinged, strigose to dense sericeous, internodes 1–3 cm long. Leaves decussate, suberect, not imbricate, subsessile to petiolate; petioles 1–3 mm long; blades 25–90 × 10–45 mm, coriaceous, not discolorous, acrodromous-basal-supranumerary, ovate to oblong-lanceolate, base cordate to subcordate to obtuse, apex acute to obtuse or acuminate, margin plane, blade verrucose due to the bulbous base of the trichomes, velvety to the touch, adaxial surface strigose to sericeous, abaxial surface strigose to sericeous, with dense white trichomes, sometimes sufficiently abundant to turn the leaves whitish, sometimes inconspicuous; secondary veins 4–10 on each side, 3–4 of them departing from the base of the midvein; domatia absent. Synflorescence frondose-bracteose, triplobotryum to pleiobotryum type, congested; accessory branches absent, principal florescence 4–6  cm long, paraclades 6–25 cm long; bracts 10–27 × 6–10 mm, similar to the vegetative leaves in shape; flowers subsessile, hypopodium absent, epipodium 1–2 mm long; prophylls 9.5–11  ×  2.5–4.5  mm, surpassing the apex of the floral tube, elliptic-lanceolate, unkeeled, base obtuse to acute, apex acute, margin plane, pilose; floral tube 5.5–7 mm long, funnel-shaped, increasing in size on fruit, whitish, densely pilose; sepals 3–6  mm long, wine-colored, triangular, not caudate, not deflexed, pilose; epicalyx segments 4.3–7  mm long, greenish to wine-colored, broadly triangular with subrevolute margin, sometimes foliaceous, spreading to suberect, reaching or exceeding the height of the sepals, pilose; corolla 5–6 cm in diam., clear rose to lilac or sometimes bright rose, petals 18–25 × 12–15 mm, obovate, apex obtuse, sometimes emarginate; stamens 24–32, free portion of the filaments 10–14  mm long, exserted 5–6 mm, rose, surpassing the sepals ca. 1 mm; ovary 2.1–2.8 × 3–3.7 mm, obconic, glabrous, style 14–21  mm long, stigma black-violet, ovules 36–80. Capsules ca. 5.5  ×  5.5–6  mm, globose, glabrous; seeds (33–)50–72, 3.5–4 × 3–3.5 mm. 2n = 30 (Cavalcanti 1995). Distribution and Ecology (Fig.  13.80)  Diplusodon paraisoensis occurs in Chapada dos Veadeiros, Goiás, at 900–1300 m, and in Serra Geral de Goiás (Espigão Mestre), Tocantins, at 660–700. The species grows in densely settled cerrado, campo sujo, campo limpo, and campo rupestre and in cerrado (sensu stricto) with sandy soil at lower elevations. Flowers from January to July; fruits from January to November. Specimens Examined: BRAZIL.  Goiás,  Água Fria de Goiás, 15°02′53″ S, 47°36′06″ W, 27 Jan 1997 (fl), Walter et  al. 3631 (CEN); Água Fria de Goiás, Estação Repetidora da Telebrasília de Roncador, 8 Feb 1994 (fl), Hatschbach & Silva 60152 (MBM); Água Fria de Goiás, estrada de terra Água Fria-São João da Aliança, km 36, 14°54′12″ S, 47°33′01″ W, 16 Nov 2005 (fr), Cavalcanti et al. 3599 (CEN, MO); Água Fria de Goiás, na torre repetidora de Roncador, 14°54′12″ S, 47°33′01″ W, 6 Apr 2006 (fr), Cavalcanti et al. 3755 (CEN); Alto Paraíso de Goiás, 14°07′ S, 47°26′ W, 13 Mar 1995 (fl), Cavalcanti et al. 1309 (CEN); Alto Paraíso de Goiás, Fazenda Sete Irmãos, margem esquerda do rio São Bartolomeu, 14°02′26″ S, 47°20′27″ W, 1 Nov 2012 (fr), Pereira-Silva et al. 16432 (CEN); Alto Paraíso de

318

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Goiás, 13°55′12″ S, 47°25′12″ W, 4 Apr 1997 (fl), Cavalcanti et al. 2205 (CEN, MO, NY); Alto Paraíso de Goiás, 1  km  L de Alto Paraíso na estrada para Nova Roma, 5 Mar 1973 (fl), Anderson 6336 (NY, P, UB); Alto Paraíso de Goiás, 6–7 km L de Alto Paraíso na estrada para Nova Roma, 7 Mar 1973 (fl), Anderson 6506 (NY, P, UB); Arraias, estrada GO-118, Campos Belos-Arraias, a 9 km da saída de Campos Belos, 12°59′69 S, 46°50′46″ W, 5 Apr 1997 (fl, fr), Cavalcanti et al. 2217 (CEN, MO, NY); Campos Belos, 21 Mar 1993 (fl), 8  km de Campos Belos no sentido Arraias, 21 Mar 1993 (fl), Cavalcanti & Ayres 1259 (CEN, MBM, MO, R, SPF); Campos Belos, 12 km depois de Campos Belos na estrada Campos Belos/Arraias, 21 Mar 1993 (fl), Cavalcanti & Ayres 1257 (CEN); Campos Belos, estrada Campos Belos para a Chapada dos Veadeiros, 6 Apr 1972 (fl, fr), Rizzo 7978 (UFG); Cavalcante, estrada velha para Nova Roma, a 7 km do entroncamento Teresina de Goiás-Campos Belos, 13°43′ S, 47°11′ W, 7 Mar 1989 (fl), Cavalcanti et al. 390 (CEN, NY, SPF); Cavalcante, estrada Cavalcante-Minaçu, km 20, 13°38′42″ S, 47°28′39″ W, 31 Aug 2004 (fr), Cavalcanti et  al. 3531 (CEN, SPF); Cavalcante, estrada de terra Cavalcante-Kalunga, km 13, 13°41′31″ S, 47°27′57″ W, 18 Nov 2005 (fr), Cavalcanti et  al. 3625 (CEN, MO); Cavalcante, estrada de terra Cavalcante-Kalunga, 24  km, 13°36′07″ S, 47°28′29″ W, 18 Nov 2005 (fr), Cavalcanti et  al. 3640 (CEN, MO); Cavalcante, RPPN Soluar, 13°45′00.8″ S, 47°23′54.6″ W, 18 May 2004 (fl, fr), Fonseca et al. 5258 (CEN, IBGE); Cavalcante, entre Cavalcante e Conceição, Burchell 8011 (K); Niquelândia, Fazenda Serra Negra, Serra Negra, ao lado da linha de transmissão elétrica, 11 Jun 1992 (fl, fr), Walter et al. 1617 (CEN, NY); Planaltina, GO-118, km 110, 20 km S de São João da Aliança, 14 Apr 1994 (fl, fr), Cavalcanti & Pereira-Silva 1264 (CEN, SPF); São João da Aliança, 10 km N de São João da Aliança, 14°43′ S, 57°52′ W, 15 Mar 1995 (fl), Cavalcanti et al. 1385 (CEN); São João da Aliança, 18 Feb 1975 (fl), Hatschbach et al. 36257 (MBM, P); São João da Aliança, GO-3, a 72 km do entroncamento da BR-020-DF-345, 15 Aug 1990 (fr), Cavalcanti et al. 648 (CEN, K, NY, SPF); São João da Aliança, GO-118, 10–20 km S de São João da Aliança, 11 Feb 1990 (fl), Hatschbach & Nicolack 53813 (MBM, NY); São João da Aliança, 65 km S, 27 Feb 1982 (fl), Oliveira & Anderson 474 (MBM, US); Teresina de Goiás 10  km  S de Teresina, 27 May 1975, Hatschbach et  al. 36914 (MBM, P); Teresina de Goiás, estrada Alto Paraíso-Teresina de Goiás, a 59 km de Alto Paraíso, local de camping próximo à cachoeira, Hotel Ecológico Alpes Goianos, 13°51′54″ S, 47°15′55″ W, 4 Apr 1997 (fl, fr), Cavalcanti et  al. 2210 (CEN, MO, NY); without reference to municipal district: 1839 (fr), Gardner 3152 (K); 9 Nov 1991, Hatschbach et  al. 55995 (MBM, P). Tocantins, Conceição do Tocantins, rodovia TO-050, 5 km ao S de Conceição do Tocantins, Fazenda Cartao de Visita, 11 May 2000 (fl), Hatschbach et  al. 70922 (MBM); Dianópolis, TO-040, Dianópolis-Natividade, cerca de 1  km de Dianópolis, entrada à esquerda para fazenda, 11°37′06″ S, 41°50′52″ W, 28 Jan 2003 (fr), Cavalcanti et  al. 3147 (CEN, MO); Dianópolis, Dianópolis-Natividade, beira da estrada após Lagoa Bonita, 11°44′26″ S, 46°40′57″ W, 30 Sep 2003 (fr), Cavalcanti et al. 3434 (CEN, MO); Gurupi, 5 km S de Gurupi, 11°43′ S, 49°07′ W, 11 Nov 1997 (fr), Proença et al. 1809 (UB); Natividade, Serra

Descriptions

319

da Natividade, próximo à antena (topo da serra), 11°41′37″ S, 47°42′08″ W, 17 Jul 2000, Souza et al. 24079 (ESA, UB); Natividade, Serra da Natividade, a uns 20 km de Natividade, 12 Apr 1974, Rizzo 9877 (UFG); Natividade, Fazenda São João, 12°53′23″ S, 47°36′01″ W, 25 Mar 2004 (fl), Sevilha et  al. 3761 (CEN); Paranã, 12°49′58″ S, 47°37′44″ W, 31 Mar 2004 (fl), Sevilha et al. 4031 (CEN); Pindorama do Tocantins, estrada de terra para Pindorama do Tocantins, a 18 km do entroncamento com Ponte Alta, 11°26′47″ S, 47°41′40″ W, 13 Jun 2002 (fl), Cavalcanti et al. 2738 (CEN, MO); Porto Nacional, estrada Porto Nacional-Brejinho de Nazaré, próximo à Fazenda Amélia, 11°0′34″ S, 48°54′71″ W, 27 May 1999, Lira & Noleto s.n. (HTO); Santa Rosa do Tocantins, a 6 km do lixão, 11°25′01″ S, 48°03′23″ W, 7 Jul 2009 (fl), Oliveira et al. 1580 (IBGE). Diplusodon paraisoensis is a usually tall subshrub (ca. 1.5 m high) covered by dense indument on the whole plant with ovate to oblong-lanceolate leaves of 2.5–9 cm long. The trichomes sometimes turn the leaves and the floral tube whitish. The corolla is 5–6 cm in diameter; the epicalyx segments are broadly triangular, sometimes foliaceous, and spreading to suberect; and the androecium has 24–32 stamens. This species resembles D. floribundus in the vegetative respects, but the latter differs principally because it has linear epicalyx segments and an androecium with 18 stamens. Koehne (1877) described Diplusodon lanceolatus var. alutaceus f. foliosus, which he distinguished by its velutinous, congested branches and leaves, and flowers with 22–26 stamens. In addition to these features, the type of Diplusodon lanceolatus var. alutaceus f. foliosus also has robust, oval to ovate-elliptic leaves with a cordate to obtuse base, flowers 9–11 mm long, epicalyx segments with long setae, and an androecium with 24–32 stamens. Cavalcanti (2004) elevated f. foliosus to species rank and synonymized D. paraisoensis with it. Nonetheless, because a name does not have priority outside of the rank at which it was published (Turland et al. 2018: Art. 11.2), Diplusodon lanceolatus var. alutaceus f. foliosus does not have priority over D. paraisoensis, even though the former was published first. Therefore, the name Diplusodon foliosus is an illegitimate name, because it included the type of D. paraisoensis in its circumscription, and the latter name should have been adopted (Turland et al. 2018: Art. 52.1). 62. Diplusodon parvifolius Mart. ex DC., Prodr. 3: 94c. 1828. Friedlandia parvifolia Mart. ex DC., Prodr. 3: 94c, 1828, pro syn. Type: Brazil. Bahia: Habitat in campis subalpestribus ad Sincorâ, C. F. P. Martius s.n. (lectotype, M [0146264] designated by Lourteig 1989: 223; isolectotypes, fragm. G, K). Figs. 13.78 A–H, 13.80 Diplusodon revolutifolius DC. ex Koehne, Mart. Fl. Bras. (Martius) 13(2): 326. 1877, pro syn. Shrubs 0.7–2  m tall, erect, much-branched, dark-brown when dry. Upper branches cylindrical to subquadrangular, unwinged, velutinous, hirsute, trichomes

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Fig. 13.78  Diplusodon parvifolius. A. Branch with inflorescence. B. Section of internode. C, E–F. Leaves, abaxial surface. D. Leaf, adaxial surface. G. Prophyll, adaxial surface. H. Flower with prophyll, without petals. (A, B Mori & Funch 13383, CEPEC; C–E, G–H Cavalcanti et al. 314, CEN; F Harley 19944, CEN)

Descriptions

321

with bulbous base, canescent, internodes 1–2 cm long. Leaves decussate, not imbricate, spreading to suberect, subsessile; petioles to 1 mm long; blades 4–12 × 3–7 mm, canescent, coriaceous, sometimes discolor, eucamptodromous, orbicular, ovate-­ lanceolate to elliptic-lanceolate, base obtuse to subcordate, apex obtuse, margin revolute; secondary veins 3–5 on each side, adaxial surface hispid, trichomes with bulbous base, with deeply sulcate veins, abaxial surface hirsute, white, veins prominent; domatia absent. Synflorescence frondose, diplobotryum to pleiobotryum type, congested; accessory branches absent, principal florescence botryum type, 9–14 cm long, paraclades 3–22 cm long; bracts similar to the vegetative leaves in shape and size; flowers subsessile, hypopodium 0.5–1 mm long, epipodium absent; prophylls 2.5–3 × 1.8–2 mm, reaching to the middle of the floral tube, obovate, unkeeled, base acute, apex obtuse, margin plane; floral tube 4–5 mm long, funnel-shaped, densely covered with white trichomes, turning the floral tube whitish, sepals 1.8–2 mm long, triangular, not caudate, not deflexed, pilose as the floral tube; epicalyx segments 1–2  mm long, pilose, cylindrical, spreading to deflexed, shorter than the sepals; corolla ca. 2  cm diam., rose-­magenta, petals 10–11  ×  6–6.5  mm, obovate, base obtuse; stamens 12, free portion of the filaments 5–7  mm long, exserted; ovary 1.8–2  ×  2–2.2  mm, obovate-depressed, glabrous, style 12–14.5  mm long, ovules 8–10. Capsules 2.5–3.5 × 2.5–3 mm, globose, glabrous; seeds 2–3, 2–2.1 × 1.9–2 mm. Distribution and Ecology (Fig.  13.80)  Diplusodon parvifolius grows predominantly in campos rupestres, cerrado, campo sujo, campos gerais, and carrasco, on Chapada Diamantina, Bahia, at elevations of 1000–1477 m. Flowers from November to March; fruits registered from January to March, also in July and November. Specimens Examined: BRAZIL. Bahia,  Catolés, Abaíra, ladeira rochosa entre Ouro Fino e Pedra Grande, 13°14′ N, 41°54′ W, 11 Feb 1992 (fl), Queiróz et al. H51085 (CEN, CEPEC n.v., HUEFS, K, SPF); Catolés, Abaíra, campo de Ouro Fino, 13°15′ N, 41°54′ W, 6 Feb 1992 (fl), Lughadha et al. H51051 (CEN, CEPEC n.v., HUEFS, K, NY, SPF); Catolés, Abaíra, Serra do Sul do Riacho da Taquara, 13°15′ N, 41°55′ W, 10 Jan 1992 (fl, fr), Harley et al. H51248 (CEN, CEPEC n.v., HUEFS, K, NY, SPF); Catolés, Abaíra, campo do Cigano, 13°15′ N, 41°55′ W, 25 Feb 1992 (fl, fr), Harley & Sano 52310 (CEN, HUEFS); Mucugê, caminho para Abaíra, 13°0′24″ S, 41°22′28″ W, 13 Feb 1997 (fl, fr), Atkins et al. [PCD] 5582 (ALCB, CEN); Piatã, a 10 km ao N de Piatã, 3 Mar 1980 (fr), Mori & Funch 13383 (CEPEC, NY, P n.v.); Piatã, Três Morros, estrada Piatã-Inúbia, 13°15′ S, 41°53′ W, 5 Dec 1992 (fl), Ganev 1623 (CEN, HUEFS, K, SPF); Piatã, 9.3 km de Piatã, na estrada Piatã-Boninal (BA-148) e 2.1 km W na estrada vicinal para Inúbia, 13°04′10″ S, 41°48″29″ W, 28 Dec 2004 (fl), Mello-Silva & Forzza 2809 (CEN, RB, SPF); Rio de Contas, estrada para o Pico das Almas, próximo à Fazenda Brumadinho, 18 Mar 1998 (fl, fr), Hatschbach et  al. 67911 (MBM); Rio de Contas, estrada Rio de ­Contas-Pico das Almas, 7 Jan 1999 (fl, fr), Cavalcanti et al. 2438 (CEN); Rio de Contas, ca. 3 km South of small town of Mato Grosso on the road to Vila do Rio de Contas, 13°29′ S, 41°49″ W, 24 Mar 1977, Harley et  al. 19944 (CEN); Rio de

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Contas, 2 km da cidade para Marcolino Moura, 4 Mar 1994, Sano et al. (CFCR) 14850 (CEN, SPF); Rio de Contas, ca. 5 km na estrada da cidade para o Pico das Almas, 27 Dec 1997 (fl), Carvalho et al. 6340 (CEN, CEPEC, RB); Rio de Contas, ca. 5 km na estrada da cidade para o Pico das Almas, 20 Feb 1987 (fl, fr), Harley et al. 24500 (CEN, K, SPF); Rio de Contas, estrada Rio de Contas-Mato Grosso, 13°32′20″ S, 41°51′29″ W, 22 Jan 2003 (fl), Cavalcanti et al. 3141 (CEN, MO); Rio de Contas, 12–14 km N de Rio de Contas, na estrada para Mato Grosso, 13°28′ S, 41°50′ W, 17 Jan 1974 (fl), Harley 15186 (CEPEC [2], K, MO, NY, P); Rio de Contas, 6–10 km NE de Rio de Contas, na estrada para o Pico das Almas, 13°32′ S, 41°53′ W, 21 Jul 1979 (fr), Mori et al. 12451 (CEPEC, US); Rio de Contas, Pico das Almas, vertente leste, Junco-Fazenda Brumadinho, 12–16 km ao N da cidade, 10 Nov 1988 (fl, fr), Harley et al. 26083 (CEN, CEPEC, K, MO, NY, SPF); Rio de Contas, a 6 km N de Rio de Contas, na estrada para Abaíra, 13°33′ S, 41°47′ W, 16 Jan 1974 (fl), Harley 15122 (CEPEC, K, MO, NY, P); Rio de Contas, a 9 km de Rio de Contas, na estrada para Mato Grosso, próximo a riacho, 13°29′ S, 41°51′ W, 24 Feb 1989, (fl), Cavalcanti et  al. 295 (CEN); Rio de Contas, estrada para Mato Grosso, 20 Jan 1989 (fl), Hatschbach 47367 (MBM, W); Rio de Contas, estrada Mato Grosso-Rio de Contas, km 5, 13°19′ S, 41°53′ W, 24 Feb 1989 (fl), Cavalcanti et  al. 303 (CEN, K, MBM, NY, SPF); Rio de Contas, ca. 15  km NE de Rio de Contas, próximo a Junco, 22 Jan 1974 (fl), Harley et al. 15594 (CEPEC [2], K, MO, NY, P); Rio de Contas, estrada Rio de Contas-Marcolino Moura, ca. 2.8 km de Rio de Contas, 13°35′40″ S, 41°47′17″ W, 18 Nov 2000 (fl), Juchum et al. 81 (CEN, CEPEC); Rio de Contas, estrada Rio de Contas-Rio da Caixa, km 18, 13°27′ S, 41°57′ W, 25 Feb 1989 (fl), Cavalcanti et al. 314 (CEN, F, MBM, NY, SPF); Rio de Contas, aeroporto, 21 Jan 1984 (fl), Hatschbach 47420 (BR, CEPEC, MBM, MO); Rio de Contas, ca. 19 km NE ao longo da estrada de Vila do Rio de Contas para o Pico das Almas, 13°33′ S, 41°53′ W, 21 Mar 1977 (fl), Harley 19783 (CEPEC, MO, NY); Rio de Contas, Pico das Almas, 19 Feb 1987 (fl), Harley et al. 24402 (CEN, K, P, SPF); Rio de Contas, Serra das Almas, a 5 km NW de Rio de Contas, 21 Mar 1980 (fl), Mori & Benton 13515 (CEPEC, NY, P); Seabra, ca. 23 km N de Seabra, estrada para Água de Rega, 24 Feb 1971 (fl), Irwin et al. 30912 (MO, NY, UB). Diplusodon parvifolius has a shrubby habit, with many branches and a corymbose aspect. It has small, orbicular leaves that are hirsute on the abaxial surface also with a revolute margin. The flowers are small, with 12 stamens, short, spreading to deflexed epicalyx segments and obovate prophylls that reach the middle of the floral tube or less. The populations analyzed are uniform in vegetative aspect, although there is some variation in the shape of the leaves, from orbicular to ovate-lanceolate or elliptic-lanceolate. However, the different shapes are almost imperceptible owing to the reduced leaf size. Diplusodon vividus is a species from north of Minas Gerais that vegetatively resembles D. parvifolius by the habit and by the shape of its leaves, with the strongly revolute margin. However, D. vividus has larger flowers (floral tube 5.5–6 mm long), with prophylls surpassing the apex of the floral tube and longer (2.2–2.5 mm long), wide-lanceolate and ciliate epicalyx segments.

Descriptions

323

The type material of Diplusodon parvifolius, Martius s.n., is represented at M by one specimen, with the original blue label from the Herbarium of the Flora Brasiliensis, indicating the locality, “Sincora, Bahia.” Lourteig (1989) cited Martius s.n. at M as holotype, what is considered the lectotype (ICBN Art. 9.17, ex. 14). 63. Diplusodon petiolatus (Koehne) T.B.Cavalc., Bol. Bot. Univ. São Paulo 22(1): 9, fig. 5. 2004. Diplusodon lanceolatus Pohl var. lanceolatus (Pohl) Koehne f. petiolatus Koehne, Fl. Bras. (Martius) 13(2): 330, pl. 61, fig. 4b. 1877. Type: Brazil. Province de Goyas, 1841, G. Gardner 3724 (lectotype, P [01901880] designated by Lourteig 1989: 227; isolectotypes, G [3], HAL, K, OXF, NY [2], P [01901879, 01901881], W [2]). Figs. 13.79 A–F, 13.80, 15.3 D Subshrubs 0.4–1.6 m tall, erect, branched, sometimes caespitose, with a xylopodium, not dark-brown when dry. Branches cylindrical, lax, unwinged, densely puberulous, with short thick trichomes of different lengths, irregularly arranged, sometimes forming tufts, internodes 2–6 cm long. Leaves decussate, not imbricate, suberect, petiolate; petioles 6–15  mm long; blades 45–70  ×  15–40  mm, slightly glaucous, chartaceous, not discolorous, eucamptodromous, elliptic to ovate, base acute to obtuse, apex acute to attenuate, margin plane, blades pubescent; secondary veins 4–6 on each side, departing along the midvein, tertiary veins often evident; domatia absent. Synflorescence bracteose, pleiobotryum type, lax, broad and long, beginning almost from the base of the plant, branches slender and delicate, with short trichomes of varying sizes; accessory branches of the first and second orders present, short, principal florescence generally truncate, paraclades of the first order 20–30  cm long, paraclades of the second order 10–15  cm long; bracts 20–30 × 7–10 mm, elliptic; flowers much concentrated at the apex of the branches, sometimes forming glomerules, subsessile, hypopodium ca. 1 mm long, epipodium absent; prophylls 4–5  ×  1–1.2  mm, surpassing the floral tube, narrowly oblong, keeled, base acute, apex acute, margin plane; floral tube 4–5 mm long, canescent, funnel-shaped, trichomes very short and dense; sepals 1.5–2 mm long, triangular, not caudate, slightly deflexed, margins with conspicuous papillae; epicalyx segments 2.5–4  mm long, cylindrical and narrow, erect, ca. 1.5  mm longer than the sepals; corolla 1.7–2 cm in diam., bright rose, petals 10–12 × 6–8 mm, elliptic, base acute, apex obtuse; stamens 18, free portion of the filaments 6–7 mm long, slightly exserted, anthers generally with trichomes; ovary 1.2–1.8  ×  1.8–2  mm, obconic, glabrous, style 10–13 mm long, ovules 12. Capsules ca. 3 × 2.5–3.1 mm, globose to oblong, glabrous; seeds 5–12, 1.5–2 × 1.1–1.9 mm. Distribution and Ecology (Fig. 13.80)  Diplusodon petiolatus is found at the margins of dense cerrado and also in disturbed open cerrado and campo sujo; on sandy soils and red latosols, at 260–484 m, northeast of Chapada dos Veadeiros; and in the low areas of Rio Tocantins valley. Flowers from February to September; fruits registered from March to September.

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Fig. 13.79  Diplusodon petiolatus. A. Vegetative branch. B. Leaf, abaxial surface. C. Branch with inflorescence. D. Flower with prophylls, without petals. E. Flower in longitudinal section. F. Immature capsule. (A–E Cavalcanti et al. 2220, CEN; F Cavalcanti et al. 2223, CEN)

Descriptions

325

Fig. 13.80  Geographic distribution of D. panniculatus, D. paraisoensis, D. parvifolius, and D. petiolatus

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Specimens Examined: BRAZIL.  Goiás,  Campos Belos, 5  km após Campos Belos, 21 Mar 1993 (fl, fr), Cavalcanti et  al. 1254 (CEN, MO, NY, SPF); Nova Roma, estrada Nova Roma-Teresina de Goiás, Fazenda Barra e Barreiro, 13°27′ S, 47°05′ W, 31 Jul 2000 (fl, fr), Forzza et  al. 1593 (CEN). Tocantins, Arraias, estrada Arraias-Paranã, a 9 km da saída de Arraias, 12°52′93′′ S, 46°59′78′′ W, 5 Apr 1997 (fl, fr), Cavalcanti et al. 2220 (CEN, MO, NY); Arraias, 12 Feb 1994 (fl), Hatschbach et al. 60423 (MBM, P); Arraias, 15 Feb 1990 (fl), Hatschbach et al. 53996 (MBM, P); Arraias, rodovia TO-050, km 415, 6 km O de Arraias, 10 May 2000 (fl), Hatschbach et  al. 70862 (MBM); Arraias, estrada Arraias-Paranã, entrando à esquerda a 43 km da cidade de Arraias, 12°47′20″ S, 47°15′39″ W, 22 Feb 2006 (fl), Bringel & Barbosa 267 (CEN); Arraias, Mar 1840 (fl, fr), Gardner 3723 (K); 1841 (fl), Paranã, Fazenda Petrolina, 12°48′38″ S, 47°14′45″ W, 14 Sep 2003 (fl, fr), Sevilha et al. 3561 (CEN); Paranã, Fazenda Petrolina, 12°47′20″ S, 48°14′20″ W, 3 Jul 2002 (fl, fr), Pereira-Silva et  al. 6492 (CEN, MO); Paranã, Fazenda Petrolina, 12°51′53″ S, 47°34′32″ W, 30 Mar 2004 (fl), Sevilha et al. 4009 (CEN); Paranã, Fazenda São João, 12°54′ S, 47°46′17″ W, 13 Sep 2003 (fl, fr), Sevilha et al. 3541 (CEN); Paranã, estrada Arraias-Paranã, a 53 km da saída de Arraias, 12°43′87′′ S, 47°28′32′′ W, 5 Apr 1997 (fr), Cavalcanti et al. 2223 (CEN, MO, NY); Paranã, estrada Palmeirópolis-Alvorada de Goiás, a 25 km de Palmeirópolis, 12°54′46″ S, 48°33′42″ W, 6 Apr 1997 (fr), Cavalcanti et al. 2230 (CEN, MO); Paranã, 12°44′17″ S, 47°37′25″ S, 31 Mar 2004 (fl, fr), Sevilha et al. 4039 (CEN); São Salvador do Tocantins, canteiro de obras do UHE São Salvador, margem esquerda do rio Tocantins, 12°47′5″ S, 48°14′45″ W, 12 Jun 2006 (fl, fr), Pereira-Silva et al. 10599 (CEN). Koehne (1877) described Diplusodon lanceolatus var. lanceolatus f. petiolatus, based on Gardner 3724, which has larger, petiolate leaves and smaller flowers with fewer stamens than the typical form. Lourteig (1989) considered Diplusodon lanceolatus var. lanceolatus f. petiolatus a synonym of D. lanceolatus. These characteristics of Koehne’s taxon, together with other observations from the type material and additional recent collections, led Cavalcanti (2004) to elevate it to specific rank. Diplusodon petiolatus has several characteristics that are not found together in D. lanceolatus, including the following: a broad, delicate inflorescence, similar only to that in D. bradei from Minas Gerais; small, delicate, very congested flowers forming glomerules on the apical parts of the branches; flowers with 18 stamens; anthers with trichomes; and long-petiolate leaves. There are 13 duplicates of the type material (Gardner 3724) in European and American herbaria. Lourteig (1989) cited a specimen from P as the holotype and must stand as the designation of the lectotype (ICBN Art. 9.17, ex. 14). One of the three specimens at P was annotated by Koehne. 64. Diplusodon plumbeus T.B.Cavalc., Novon 8(4): 345, fig. 6. 1998. Type: Brazil. Goiás: Cristalina, a 5 km N de Cristalina, 17 Apr 1994 (fl, fr), T.B.Cavalcanti & C. H. S. Ayres 1267 (holotype, CEN [00018555]; isotypes, F, K, MBM, NY, R, RB, SPF).

Descriptions

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Figs. 13.81 A–L, 13.85 Subshrubs to shrubs 0.4–1.2 m tall, erect, branched, with a xylopodium, darkbrown when dry. Upper branches cylindrical to subquadrangular, sometime with a faint wing, covered by stellate trichomes, internodes 0.3–1 cm long. Leaves decussate, not imbricate, spreading, sessile; blades 6–22 × 3–5 (−19) mm, coriaceous, discolorous, acrodromous-basal-supranumerary, narrowly lanceolate, base subcordate, apex acute, margin revolute, adaxial surface dark-green, nitid, glabrous, with sulcate veins, ash-gray when dry; abaxial surface totally white due to dense indumentum, when dry the veins rust-colored, almost parallel, with dense white trichomes or sometimes glabrous; secondary veins 4–5 on each side, prominent; domatia absent. Synflorescence frondose, triplobotryum spiciform type, congested; flowers petiolate, hypopodium 0.5–1.5 mm long, epipodium ca. 0.5 mm long, prophylls 4–4.5 × 2.2–2.5 mm, reaching the middle of or at times surpassing the floral tube, elliptic to obovate, keeled, base acute, apex obtuse, margin plane, puberulous; floral tube 4–5 mm long, yellow, wine-colored in fruit, oblong, pilosity inconspicuous, apparently glabrous, sepals 2–2.5  mm long, triangular, not caudate, not deflexed; epicalyx segments 2–2.5 mm long, lanceolate, dorsiventrally compressed, spreading, the same size as the sepals; corolla 3–3.5 cm in diam., rose-magenta to bright rose, petals 15–17 × 10–12 mm, obovate, apex obtuse to acute; stamens 12, free portion of the filaments 7–9 mm long, exserted, ovary obconic, glabrous, ovules 15–21. Capsules 5.5–6  ×  4.5–5  mm, oblong, glabrous; seeds 19–21, 2.5–3 × 2.1–2.5 mm. Distribution and Ecology (Fig. 13.85)  Diplusodon plumbeus occurs in Serra dos Cristais, Goiás, in mixed vegetation in high-elevation cerrado and campo rupestre on quartzite outcrops at 1070–1250 m. Flowers from January to June; fruits registered from April to June and November. Specimens Examined: BRAZIL.  Goiás,  Cristalina, área próxima à reserva ecológica, 17 Jun 2004 (fl, fr), Pastore et  al. 1023 (CEN); Cristalina, BR-040, Brasília-Cristalina, 2  km de Cristalina, 16°43′59″ S, 47°37′16″ W, 21 Nov 2005 (bud), Cavalcanti et al. 3648 (CEN, MO); Cristalina, BR-040, Brasília-Cristalina, km 102, 16°48′ S, 47°37′ W, 12 Mar 1989 (fl), Cavalcanti et al. 411 (CEN, SPF); Cristalina, estrada para Salto do Arrojado, 10 Feb 1990 (fl), Hatschbach et al. 53751 (MBM); Cristalina, Serra dos Cristais, 21 Feb 1975, Hatschbach et  al. 36396 (MBM); Cristalina, Serra dos Cristais, 22 Jan 1997 (fl), Hatschbach et al. 66126 (CEN, MBM, NY); Cristalina, Serra dos Cristais, 22 Feb 2000 (fl), Hatschbach et al. 70515 (MBM); Cristalina, 3 km N de Cristalina, 2 Mar 1966 (fl), Irwin et al. 13258 (MO, NY, RB, UB); Cristalina, ca. 5 km da cidade, na estrada para Paracatu, em vegetação mista de cerrado de altitude e campo rupestre em afloramento quartzíticos, 16°46′ S, 47°37′ W, 4 Feb 1987, Pirani et al. 1512 (CEN, F, K, MBM, NY, SPF, UB); Cristalina, 17 Apr 1989, Cavalcanti & Ayres 1268 (CEN); Cristalina, 5 km S de Cristalina, 1 Nov 1965 (fr), Irwin et al. 9737 (NY); Cristalina, RPPN Linda Serra dos Topázios, coletado entre a sede e o córrego Areião, 16°45′ S, 47°40′ W, 28 Feb 1998 (fl), Proença & Oliveira 1945 (CEN, UB); Cristalina, RPPN Linda

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Fig. 13.81  Diplusodon plumbeus. A. Habit showing xylopodium. B. Section of internode. C. Leaf, adaxial surface. D. Leaf, abaxial surface. E. Detail of leaf abaxial surface indumentum. F–G. Prophylls, adaxial surface. H. Floral bud. I. Flower with prophyll, without petals. J. Detail of floral tube indumentum. K. Ovary. L. Ovules. (A, G, H, J–L Cavalcanti & Ayres 1267, CEN; B–F, I Pirani et al. 1512, SPF)

Descriptions

329

Serra dos Topázios, 16°43′18″ S, 47°41′04″ W, 13 Jun 2004 (fr), Pastore et al. 1006 (CEN). Diplusodon plumbeus can be recognized by its acrodromous leaves that are 3–6 mm wide, with 4–5 secondary veins on each side of the midvein; the secondary veins are close to together, almost parallel to each other, and are perpendicular to the midvein. The adaxial leaf surface is glabrous, dark-green in color but grayish when dry. The abaxial surface is strongly whitish, due to a dense indumentum formed by short, spreading trichomes that are located on the leaf surface but absent on the veins. The flowers have an oblong floral tube with lanceolate, spreading epicalyx segments, and the capsule is oblong. 65. Diplusodon praetermissus T.B.Cavalc. nom et stat. nov. Diplusodon oblongus Pohl var. angustifolius T.B.Cavalc. Novon 8(4): 345. 1998. Type: Brazil. Goiás: Alto Paraíso de Goiás, cerca de 5 km depois da entrada para as cachoeiras (Vila São Jorge), 14°08′ S, 47°30′ W, 8 Mar 1992 (fl, fr), T.B.Cavalcanti, B. M. T. Walter & D. M. S. Rocha 1042 (holotype, CEN [00017896]; isotypes, CEN [00017896], NY). Figs. 13.82 A–E, 13.85, 15.3 C Subshrubs 0.4–2 m tall, erect, much-branched, with a xylopodium, dark-brown when dry. Upper branches subquadrangular, rust-colored, inconspicuously pilose, minute trichomes densely arranged almost imperceptible, internodes 0.8–1.2  cm long. Leaves decussate, not imbricate, spreading to suberect, subsessile to shortpetiolate; petioles 0.5–1.5 mm long, wide; blades 7–34 × 3–5(−8–12) mm, not glaucous, coriaceous, not discolorous, eucamptodromous to hyphodromous, narrowly oblong, base acute or attenuate, apex acute, margin plane, not ciliate, thickened, blades glabrous or with minute trichomes on the margin, almost imperceptible; secondary veins 1–2 on each side, inconspicuous, reddish to rust-colored; domatia absent. Synflorescence frondose, pleiobotryum type, congested; without accessory branches, principal florescence 5–10 cm long, paraclades 3–11 cm long; bracts similar to the vegetative leaves in shape, decreasing in size toward the apex of the branch; flowers sessile to subsessile, hypopodium 0, epipodium 0.6–0.9 mm long; prophylls 6–7.1 × 1.1–1.8 mm, reaching the the base of the sepals, oblong, strongly keeled, base retuse, apex acute, margin plane, markedly lighter, with minute trichomes almost imperceptible to glabrous; floral tube 5.5–6.5 mm long, green when fresh, oblong, with minute trichomes almost imperceptible, glabrous in fruit; sepals 1–1.8 mm long, triangular, not caudate, not ciliate, with minute trichomes almost imperceptible to glabrous; epicalyx segments 4–5  mm long, cylindrical, acute, suberect, longer than the sepals, with minute trichomes almost imperceptible; corolla 3.5–4  cm in diam., lilac to rose, petals ca. 17.5  ×  8  mm, obovate, apex obtuse; stamens 12, free portion of the filaments 6.5–7 mm long, exserted, ovary 2.5–2.6 × 1.8–2 mm, obtriangular, glabrous, style 10.5–11 mm long, ovules 14–19. Capsules 4.5–5 × ca. 3.5 mm, oblong, glabrous; seeds 8–11, 2–2.2 × 2–2.1 mm. Distribution and Ecology (Fig. 13.85)  Diplusodon praetermissus is restricted to Chapada dos Veadeiros, Goiás, where it is found in campos limpos, campos sujos

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Fig. 13.82  Diplusodon praetermissus. A. Branch with inflorescence. B. Section of internode. C. Leaf, abaxial surface. D. Prophyll, adaxial surface. E. Flower with prophylls, without petals. (A Cavalcanti et al. 401, CEN; B Pastore 237, CEN; C–E Cavalcanti et al. 3813, CEN)

Descriptions

331

(fields with shrubby savanna elements), campos rupestres, and cerrado with rocky outcrops and sandstone soils at 1000–1250 m. Flowers from August to March; fruits from January to July. Specimens Examined: BRAZIL.  Goiás,  Alto Paraíso de Goiás, estrada para Colinas, a 34 km do entroncamento da estrada, 15 Aug 1990 (fl), Cavalcanti et al. 660 (CEN); Alto Paraíso de Goiás, Parque Nacional da Chapada dos Veadeiros, na margem da cerca, próximo à sede (8 km), 14°12′ S, 47°52′ W, 14 Aug 1991 (fr), Cavalcanti et  al. 851 (CEN [2]); Alto Paraíso de Goiás, Chapada dos Veadeiros, estrada Alto Paraíso-Colinas do Sul, a 28 km do entroncamento da GO-118, 14°10′ S, 47°47′ W, 2 Apr 1997 (fr), Cavalcantiet al. 2188 (CEN, MO [2], NY); Alto Paraíso de Goiás, Chapada dos Veadeiros, estrada Colinas-GO-118, km 42, GO-327, 14°10′ S, 47°44′ W, 8 Mar 1989 (fl), Cavalcanti et al. 401 (CEN, MBM, NY); Alto Paraíso de Goiás, entrada do Parque Nacional da Chapada dos Veadeiros (à esquerda), estrada Alto Paraíso-Colinas do Sul, 4 km da GO-118, 27 Jan 1997 (fl), Walter et al. 3646 (CEN, MO, NY, SPF); Alto Paraíso de Goiás, Parque Nacional da Chapada dos Veadeiros, trilha a oeste da sede do Parque, 14°10′27″ S, 47°49′28″ W, 11 Mar 2014 (fr), Cavalcanti et al. 3813 (CEN, K, MO, SPF, UFG); Alto Paraíso de Goiás, a 2 km da entrada do Parque, 13 Dec 1992 (fl), Bianchetti & Pereira-Silva 1227 (CEN); Alto Paraíso de Goiás, próximo à cachoeira do rio Preto, perto do povoado São Jorge, 6 Feb 1987 (fl, fr), Pirani et  al. 1685 (CEN, K, SPF); Alto Paraíso de Goiás, Parque Nacional da Chapada dos Veadeiros, vila São Jorge, trilha de acesso aos saltos do Rio Preto (saltos I e II), 14° 10′ 22′′S, 47° 49′ 36′′W, 24 Nov 2014 (fl), Oliveira et al. 457 (CEN, RB); Alto Paraíso de Goiás, São Jorge, caminho para mirante, 11 Nov 2004 (fl), Rodrigues & Chesini 194 (CEN); povoado de São Jorge, beira da estrada, 4 Jan 2003 (fl), Pastore 237 (CEN, HUFU, MO). Diplusodon praetermissus is characterized by branches and flowers that are densely puberulent, narrowly oblong leaves, and flowers with an oblong floral tube, suberect epicalyx segments, 4–5 mm long, and 12 stamens. The whole plant turns dark-brown when dry. This taxon had been previously considered as a variety of Diplusodon oblongus; however, the analysis of additional specimens led me to conclude that it is a different species. Diplusodon oblongus differs from D. praetermissus because it has elliptic or ovate to ovate-lanceolate leaves, with 3–4 secondary veins on each side of the midvein, a petiole 2–9 mm long, and epicalyx segments 2–3.5 mm long. The specific rank of this taxon has been corroborated by molecular phylogenetic analyses (Inglis and Cavalcanti 2018), in which the two previously recognized varieties are found in distant subclades. 66. Diplusodon prostratus T.B.Cavalc., sp. nov. Type: Brazil. Goiás: Alto Paraíso de Goiás, Chapada dos Veadeiros, na trilha para a cachoeira Almécegas II, 3 Jun 1999 (fl), T.B.Cavalcanti & R. Mello-Silva 2505 (holotype, CEN [00039856]; isotypes, ESA, F, G, HTO, K, MBM, MO, NY, P, RB, SPF, W). Figs. 13.83 A–E, 13.85

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Fig. 13.83  Diplusodon prostratus. A. Branch with inflorescence. B. Section of internode. C–D. Leaves, abaxial surface. E. Flower with prophyll, without petals. (A–C, E Cavalcanti & Mello-Silva 2505, CEN; D Cavalcanti et al. 3751, CEN)

Descriptions

333

Diplusodon prostratus is characterized by prostrate branches and reduced elliptic to oblong leaves with revolute margins. The floral tube is 3–3.5 mm long, and the epicalyx segments are suberect, 1–1.5 mm long, cylindrical, and shorter than the sepals. It is morphologically similar to D. cordifolius, which differs by its erect, narrowly elongated leaves with a cordate base, hirsute pubescence covering the plant, and caudate sepals. Shrubs 0.4–1 m tall, branched, branches prostrate, with a thick xylopodium, not dark-brown when dry. Upper branches cylindrical, unwinged, with sparse long trichomes mixed with short trichomes, or just the short trichomes; internodes 3–5 mm long. Leaves decussate, not imbricate, patent, sessile to subsessile; petioles 0.2– 0.8 mm long; blades 3–9 × 0.9–4 mm, coriaceous, discolorous, eucamptodromous, elliptic to oblong, base obtuse, sometimes acute, apex obtuse, margin revolute; adaxial surface dark-green, pubescent, glabrescent, abaxial surface olive-colored, with trichomes on the veins; secondary veins 1–2 on each side of the midvein, inconspicuous, domatia present, inconspicuous. Synflorescence frondose, diplobotryum to triplobotryum type congested; accessory branches absent, principal florescence truncate, paraclades 1.5–15 cm long, bracts similar to the vegetative leaves in form and size; flowers pedicellate, hypopodium 1–1.5  mm long, epipodium 0–0.5 mm long; prophylls 3–3.5 × 1.3–1.5 mm, reaching the middle to the apex of the floral tube, obovate to elliptic, unkeeled, base acute, apex obtuse, margin plane, with trichomes sparse, ciliate; floral tube 3–3.5  mm long, green, funnel-shaped, covered with sparse, long trichomes; sepals 2–3 mm long, wine-colored, triangular, not deflexed, not caudate, with sparse trichomes, ciliate; epicalyx segments 1–1.5 mm long, cylindrical, suberect, with trichomes sparse, ciliate, shorter than the sepals; corolla 2–2.5 cm in diam., rose-magenta, petals 11–11.5 × 6–6.5 mm, elliptic, apex obtuse; stamens 14–15, free portion of the filaments 7–8 mm long, exserted, ovary 1.5–1.9 × 2–2.3 mm, obovate, glabrous, style 11–12 mm long, ovules 15–17. Immature capsules ca. 3 × 2 mm, globose, glabrous; seeds immature. Distribution and Ecology (Fig. 13.85)  Diplusodon prostratus is known from the mountains of Alto Paraíso de Goiás, Chapada dos Veadeiros, Goiás State. It grows in cerrado and open fields with rocky outcrops on rocky soil at 1150–1436  m. Flowers from January to June; fruits in April. Specimens Examined: BRAZIL.  Goiás,  Alto Paraíso de Goiás, Chapada dos Veadeiros, estrada de terra à leste, subindo na Serra da Baliza, nas terras do Sr. Paulo, no alto do morro, perto da casinha abandonada, 14°9′35″ S, 47°28′ W, 2 Apr 1997 (fl, fr), Cavalcanti et  al. 2193 (CEN); Alto Paraíso de Goiás, Chapada dos Veadeiros, Solarium, no caminho para a cachoeira dos Arcanjos, 12 Apr 1998 (fl), Cavalcanti et  al. 2393 (CEN); Alto Paraíso de Goiás, GO-118, Alto Paraíso de Goiás-São João da Aliança, descida para o rio Piçarras (antigo Pizarão), lado direito, 14°13′57″ S, 47°29′8″ W, 30 Jan 2003 (fl), Cavalcanti et  al. 3168 (CEN); Alto Paraíso de Goiás, GO-118, Alto Paraíso de Goiás-São João da Aliança, km 15, morro do lado esquerdo, 4 Apr 2006 (fl), Cavalcanti et al. 3751 (CEN, HUFU).

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Diplusodon prostratus is characterized by its prostrate branches with small (3–9 × 0.9–4 mm), elliptic to oblong leaves with revolute margins. The flowers are small (floral tube 3–3.5 mm long), with cylindrical, suberect epicalyx segments that are 1–1.5 mm long and shorter than the sepals. Diplusodon prostratus is morphologically similar to D. cordifolius, which also has small leaves and small flowers with epicalyx segments that are shorter than the sepals. Diplusodon cordifolius is different from D. prostratus primarily because of its erect, narrowly elongated leaves with a cordate base, hirsute pubescence covering the plant and giving it a canescent appearance, and flowers that have caudate sepals. 67. Diplusodon psammophilus Lourteig, Bradea 19(5): 212. 1989. Type: Brazil. Minas Gerais: Diamantina, rodovia Guinda-Conselheiro Mata, 17 Feb 1973 (fl), G.  Hatschbach & L.  Z. Ahumada 31687 (lectotype, P [1901862], here designated; isolectotypes, C, MBM, NY, P [1901863]). Figs. 13.84 A–F, 13.85 Subshrubs 7–25  cm tall, erect, branched, caespitose, with a xylopodium, not dark-brown when dry. Upper branches subquadrangular, unwinged, glabrous or rare sparse and minute trichomes inconspicuous, internodes 3–10  cm long. Leaves decussate, not imbricate, erect, sessile; blades 5–14 × 0.5–1.5 mm, coriaceous, not discolorous, not glaucous, hyphodromous, linear to narrowly elliptic, base acute to slightly attenuate, apex acute, minutely mucronate, margin plane, sometimes sparse ciliate, glabrous; secondary veins absent; domatia absent. Synflorescence frondose, botryum type, lax; accessory branches absent; bracts similar to the vegetative leaves in size and shape; flowers pedicellate, hypopodium 1.5–2  mm long, epipodium 0–0.2 mm long; prophylls 6–6.5 × 1.5–1.7 mm, surpassing the apex of the floral tube ca. 1.5 mm, narrowly lanceolate to linear, base obtuse to acute, apex obtuse, margin plane, glabrous; floral tube 4.5–5.5  mm long, funnel-shaped, glabrous; sepals 3.5–3.8 mm long, triangular, caudate, not deflexed, with short cilia on the margin; epicalyx segments 2.2–2.9  mm long, lanceolate, slightly concave, erect, shorter than the sepals, generally prominently short-ciliate; corolla ca. 3  cm in diam., rose-magenta, petals ca. 16 × 8 mm, obovate, apex obtuse; stamens 10–12, free portion of the filaments 6.5–7  mm long, slightly exserted; ovary 1.5–2 × 1.5–2 mm, rounded, glabrous, style 11–12 mm long, ovules 10–13. Capsules globose, glabrous; seeds ca. 6, 2–2.2 × 1.5–1.8 mm. Distribution and Ecology (Fig. 13.85)  Diplusodon psammophilus is a microendemic species in the neighboring municipalities of Conselheiro Mata and Diamantina, Cadeia do Espinhaço, Minas Gerais. It occurs in small populations at 1200–1256  m. It inhabits sandy soil of campos rupestres. Flowers in February; fruits registered in February and September. Specimens Examined: BRAZIL. Minas Gerais,  Conselheiro Mata, estrada de terra Conselheiro Mata-Diamantina, a 15  km de Conselheiro Mata, 18°18′34″ S, 43°53′04″ W, 7 Feb 2015 (fl, fr), Cavalcanti et al. 3929 (CEN); Conselheiro Mata,

Descriptions

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Fig. 13.84  Diplusodon psammophilus. A. Habit. B. Leaf, adaxial surface. C. Leaf, abaxial surface. D. Prophyll, adaxial surface. E. Epicalyx segment. F. Flower with prophylls, without petals. (A–F Cavalcanti et al. 3929, CEN)

estrada de terra Conselheiro Mata-Diamantina, 20.3 km depois do asfalto, 18°20′ S, 43°53′ W, 23 Sep 1994 (fr), Splett 635 (UB); Conselheiro Mata, estrada Conselheiro Mata-Diamantina, km 184, 23 Feb 1986 (fl), Cavalcanti et al. (CFCR) 9525 (CEN, SPF); Diamantina, Rodovia Guinda-Conselheiro Mata, Hatschbach et  al. 50956 (MBM). The specific epithet alludes to the sandy soil where the species is found. The species can be recognized by its small (7–25 cm tall), caespitose habit; hyphodromous, linear leaves; 10–12 stamens; ciliate sepals and epicalyx margins; and narrowly

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Fig. 13.85  Geographic distribution of Diplusodon plumbeus, D. praetermissus, D. prostratus, and D. psammophilus

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lanceolate to linear prophylls that surpass the apex of the floral tube. Because of its small habit and linear leaves, Diplusodon psammophilus resembles D. ericoides, a microendemic species of Chapada dos Veadeiros, Goiás State. The latter species differs by its noncaespitose habit, flowers with sepals and epicalyx segments not ciliate, and linear-triangular dorsoventrally compressed epicalyx segments. Lourteig (1989) designated a P specimen as the holotype of Diplusodon psammophilus, but the gathering Hatschbach & Ahumada 31687 is represented by two specimens at P. The specimen P (1901862) is hereby designated as the lectotype. 68. Diplusodon puberulus Koehne, Fl. Bras. (Martius) 13(2): 339, pl. 63, fig. 4. 1877. Type: Brazil. Minas Gerais, 1842, G.  Gardner 4563a (first-step lectotype, designated by Lourteig 1989: 228, second-step lectotype, P [01901860], here designated; isolectotypes, G [2], K, NY, OXF [2], P [01901861], W). Figs. 13.86 A–I, 13.90 Subshrubs 0.5–1.5  m tall, erect, much-branched, not dark-brown when dry. Upper branches cylindrical to subquadrangular, unwinged, covered with erect, minute stellate trichomes, internodes 0.4–0.5 cm long. Leaves decussate to verticillate, not imbricate, suberect, subsessile; petioles to 1 mm long; blades 6–18 × 0.8–1.5 (−4) mm, coriaceous, not discolorous, hyphodromous, linear, rarely narrow elliptic, base acute, apex obtuse, margin revolute to the midvein, adaxial surface covered with very short and erect trichomes, sulcate-veined, abaxial surface covered by minute stellate trichomes, midvein very prominent, pubescent, secondary veins absent; domatia absent. Synflorescence frondose, pleiobotryum type, congested; accessory branches absent, principal florescence botryum type, paraclades of the first order 10–19 cm long, paraclades of the second order 4.5–6 cm long; bracts similar to the vegetative leaves in shape and size; flowers pedicellate, hypopodium 2–3.5  mm long, epipodium 1–1.5 mm long; prophylls 4–4.2 × 1.1–1.5 mm, reaching the apex of the floral tube, lanceolate, unkeeled, base attenuate, apex obtuse, margins plane, covered by appressed, minute trichomes; floral tube 3.5–4 mm long, oblong-triangular, glabrous; sepals 1.5–2 mm long, triangular, not caudate, not deflexed, glabrous, with cilia on the margin; epicalyx segments 1.5–2  mm long, cylindrical, spreading to suberect, not exceeding the length of the sepals, covered by erect, minute trichomes; corolla ca. 2 mm in diam., lilac to purple, petals ca. 6.5 × 4 mm, obovate, apex obtuse to acute; stamens 12, free portion of the filaments 4.5–5 mm long, exserted; ovary ca. 1 × 1.5 mm, rhombic, glabrous, style 10–13 mm long., ovules 12–16. Capsules 3–4  ×  4–4.1  mm, globose, glabrous; seeds 6–10, 3–3.1 × 2–2.5 mm. Distribution and Ecology (Fig. 13.90)  Diplusodon puberulus inhabits campo and cerrado in Cadeia do Espinhaço and the western part of the state of Minas Gerais, at 880–1200  m. In Bahia the species occurs in areas of campos rupestres. Lourteig (1989) referred to this species as occurring in Goiás; however, the single collection that she cited for this state (Heringer 10126) is instead Diplusodon rosmarinifolius. Flowers from January to April and in August; fruits registered from January to April and from July to August.

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Fig. 13.86  Diplusodon puberulus. A. Branch with inflorescence. B. Section of internode. C. Leaf, adaxial surface. D. Detail of leaf abaxial surface indumentum. E. Leaf, abaxial surface. F. Floral bud. G. Flower with prophylls, without petals. H. Pistil. I. Multicellular stellate trichomes. (A–B, G–I Irwin et al. 22974, UB; C–F Belém & Mendes 397, CEPEC)

Descriptions

339

Specimens Examined: BRAZIL.  Bahia,  Serra Geral de Caetité, ca. 15  km  S from Caetité, along road to Brejinho das Ametistas, 14°14′ S, 42º30′W, 12 Apr 1980 (fr), Harley et al. 21299 (K, SPF). Minas Gerais, Coração de Jesus, 16°46′14″ S, 44°19′53″ W, 24 Aug 2011 (fr), Proença & Harris 3852 (UB); Diamantina, Conselheiro Mata, 12 Aug 1971 (fl, fr), Hatschbach 30204 (MBM); Diamantina, Gardner 4564 (K, NY); Francisco Sá, a 30 km NE de Francisco Sá, estrada para Salinas, 10 Feb 1969 (fl), Irwin et  al. 22974 (MO, NY, SP, UB); Grão-Mogol, Barrocão, rodovia para Grão-Mogol, km 20–25, 9 Feb 1991, Hatschbach et  al. 54939 (MBM n.v., MO); Jequitaí, 91.5 km NE de Pirapora em direção a Montes Claros, 27 Jan 1991 (fl, fr), Taylor et al. 1452 (BHCB, K n.v., SPF); Joaquim Felício, Serra do Cabral, 2–8 km NW da cidade na estrada para a Torre da TV e a Fazenda Bocaina, 17°45′10″ S, 44°10′56″ W, 19 Mar 1994 (fr), Sakuragui et  al. (CFCR) 15172 (CEN, ESA, k, SPF); Joaquim Felício, Serra do Cabral, hills above Joaquim Felício, along or near road to tv tower, rochy cerrado and transition to campo rupestre, on slope, 17°45′ S, 44°12′ W, 13 Feb 1988 (fl, fr), Thomas et al. 5945 (CEN, NY, SPF); Montes Claros, a 52 km de Montes Claros para Pirapora, 30 Jan 1965 (fl), Belém & Mendes 397 (CEPEC, UB); Montes Claros, ca. 25  km  W de Montes Claros, estrada para Água Boa, 23 Feb 1969 (fl), Irwin et al. 23692 (K, MO, NY, P, UB); Patrocínio, Fazenda Grão de Ouro, 18°48′31″ S, 46°58′ W, 28 Feb 1989 (fl, fr), Mendonça et  al. 1203 (IBGE, SP n.v., UB); Patrocínio, Fazendas Daterra, Boa Vista, 14 Jul 1998 (fr), Farah & Freitas 342 (ESA); São Gonçalo do Abaeté, rodovia BR-365, 40 km S do trevo com a BR-040, 12 Apr 1996 (fr), Hatschbach et al. 64581 (MBM); Uberlândia, Clube de Caça e Pesca Itororó, 16 Feb 1992 (fr), Leenza & Barbosa 370 (CEN, HUFU n.v.); Uberlândia, Reserva Ecológica do Clube de Caça e Pesca Itororó, 19 Apr 1994 (fr), Arantes & Marra 169 (CEN, HUFU n.v.); Uberlândia, Reserva Ecológica do Clube de Caça e Pesca Itororó, s.d.., Cortes 3 (HUFU); Várzea de Palma, Serra do Cabral, 16 Jan 1996 (fl), Hatschbach et  al. 64196 (CEN, MBM); without reference to municipal district: Weddell 1860 (G). Diplusodon puberulus is characterized by narrow, strongly revolute leaves and especially by the presence of a floral tube that is glabrous, while the sepals, epicalyx, prophylls, and pedicel are covered with minute erect trichomes. Koehne (1882) considered this species synonymous with D. rosmarinifolius, perhaps because of the similarity in habit. Diplusodon rosmarinifolius differs from D. puberulus especially by its androecium with 15 stamens, pedicel 0–1 mm long, and prophylls reaching less than half the length of the floral tube. Diplusodon puberulus differs from D. macrodon, a species from Goiás with linear leaves and stellate trichomes, mainly because of the 5–7-mm-long epicalyx segments of the latter. Diplusodon puberulus is also similar to D. uninervius and D. mononeuros, both from Minas Gerais and also with linear leaves, but both of these species have leaves with a glabrous adaxial surface. Gardner 4563 is a duplicated number, to which I added the letters a and b to distinguish the two species under which this number is cited, Diplusodon puberulus (Gardner 4563a) and D. uninervius (Gardner 4563b).

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Lourteig (1989) cited as the holotype one specimen of Gardner 4563 with a label annotated by Koehne which is considered as the first-step lectotype (ICBN Art. 9.17, ex. 14). There are nine duplicates of Gardner 4563a in European and American herbaria and two at P.  The specimen at P annotated by Koehne (P 01901860) is chosen as the second-step lectotype. 69. Diplusodon punctatus Pohl. Subshrubs 0.2–1.7 m tall, erect, branched, with a xylopodium, totally glabrous when mature, not dark-brown when dry. Upper branches cylindrical to subquadrangular, unwinged, bright yellow, internodes 3–5.5  cm long. Leaves decussate, not imbricate, erect, subsessile; petioles 1–3 mm long; blades 17–75 × 16–40 mm, glaucous, slightly coriaceous, not discolorous, acrodromous-basal type, ovate-lanceolate, with black punctae, base obtuse, apex acute, margin plane, glabrous; secondary veins 2 on each side, departing from the base of the midvein, prominent; domatia absent. Synflorescence bracteose, pleiobotryum type, lax, thin, delicate, bright yellow; accessory branches absent, principal florescence botryum type, 3–5 cm long, paraclades 3–24 cm long; bracts 6–8 × 1.5–2.5 mm, elliptic to elliptic-lanceolate, margin plane, 1 vein on each side; flowers pedicellate, pedicel not persistent, hypopodium 2.5–5.5 mm long, epipodium 0–1 mm long; prophylls 4–5.5 × 1–2 mm, reaching or surpassing the apex of the floral tube, keeled, oblong, base retuse, apex acute to obtuse, margin plane; floral tube 3.2–5  mm long, clear yellow to strawcolored in fruit, funnel-shaped; sepals 2–3  mm long, triangular, not caudate, not deflexed; epicalyx segments absent or present, 3–4 mm long, spreading, when present not surpassing the sepals; corolla 2–2.5  cm in diam., lilac to purple, petals 8–10 × 4–5 mm, obovate, apex obtuse; stamens 12, exserted, free portion of the filaments 5.5–6.3 mm long, rose to white, exserted; ovary 1.2–2.1 × 1.5–2 mm, rhombic to rounded, glabrous; style 9–11  mm long, ovules 10–24. Capsule ca. 3.5 × 3.5 mm, globose; seeds 13–16, 1.8–2 × 1.2–1.5 mm. Distribution and Ecology (Fig.  13.90)  Diplusodon punctatus occurs in campos cerrados and campos sujos of Goiás and Tocantins states, at 330–840 m. Flowers from February to June and November; fruits from June to December. Diplusodon punctatus is characterized by its much-branched synflorescence with slender, delicate, bright yellow stems, in addition to small flowers, an androecium with 12 stamens, and the absence of an epicalyx. This last feature is also seen in D. marginatus, which is also from Goiás. Diplusodon marginatus differs principally in its pedicels 15–45 cm long, flowers ca. 7 cm in diam., and androecium of 23–33 stamens. Diplusodon punctatus is also similar to D. imbricatus, but the latter species has leaves with a cordate base, prophylls reaching or nearly reaching the middle of the floral tube, and flowers with an androecium of 18 stamens. Diplusodon punctatus was described by Pohl (1827a) as possessing reduced leaves, and later Koehne (1877, 1903) included this species in the section Subuninerves, characterized by leaves having a single central vein or inconspicuous lateral veins. Lourteig (1989) retained the species in the same section. Observations

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in the field show that, in reality, the leaves of D. punctatus are well developed. The specimens that Pohl, Koehne, and Lourteig described were mounted only with the distal, branched portion of the inflorescence. When this species flowers, it loses all stem leaves, and only the bracts persist, which look like reduced leaves with inconspicuous veins. It was these bracts that led the authors to include D. punctatus in section Subuninerves. The specimen Cavalcanti et al. 1015 provides an example of plants collected in the last phase of fruiting and young plants with true leaves, which are acrodromous, erect, coriaceous, 2–7.5 cm long, and with two very conspicuous veins on each side of the midvein and departing from its base. Following Koehne’s (1903) sectional taxonomy, D. punctatus should be classified in section Palmatinerves. Key to the varieties. 1 . Epicalyx segments absent …………...................…. D. punctatus var. punctatus 1. Epicalyx segments present, 3–4  mm long …….…….…….…….…….…….… ….…….…….…….…….…….…….…….…............ D. punctatus var. dentatus 69a. Diplusodon punctatus Pohl var. punctatus. Flora 10: 151, pl. 72. 1827. Type: Brazil. Tocantins: Natividade, in campis siccis ad Rio Bagagem, Sep 1821 (fl), J. B. E. Pohl 2018 = D 946 (first-step lectotype, designated by Lourteig 1989: 215, second-step lectotype, W [0053429], here designated; isolectotypes, G, K, W [0053430, 0053431]). Figs. 13.87 A–F, 13.90 Flowers pedicellate, hypopodium 4–5.5  mm long, epipodium 0.5–1  mm long; prophylls 5–5.5 × ca. 1 mm, surpassing the apex of the floral tube; epicalyx segments absent. Distribution and Ecology (Fig. 13.90)  Diplusodon punctatus var. punctatus was known prior to this study from only two collections, including the type. It has since been collected several times in Goiás at 400–840 m. It occurs in small, infrequent populations, in cerrado and campo sujo, in sandy soils with gravel. Flowers from January to October; fruits from July to December. Specimens Examined: BRAZIL. Goiás,  Barro Alto, 38 km do trevo NiquelândiaCodemin, estrada para Colinas do Sul, 14°27′15″ S, 48°10′44″ W, 22 Jul 2007 (fr), Forzza et al. 4600 (CEN, RB n.v.); Cavalcante, Burchell 7684–2 (BR); ponte sobre o rio Tocantins-Cavalcante, km 15, 13°22′40.3″ S, 48°02′58.2″ W, 30 Aug 2004 (fl, fr), Cavalcanti et  al. 3510 (CEN, SPF); Cavalcante, RPPN Serra do Roncador, 13°40′01″ S, 47°48′04″ W, Munhoz et al. 7966 (CEN, UB); Cavalcante, Estrada de acesso à Fazenda Paraíso, 13°20′16″ S, 48°08′31″ W, 16 May 2007 (fl), PereiraSilva et al. 11844 (CEN); Cavalcante, Reserva Natural Serra do Tombador (RPPN), 9 May 2014 (fl), Rissi et al. 661 (CEN, HRCB); Colinas do Sul, 15 Jun 1993 (fl), Hatschbach et al. 59555 (BR, CEN, MBM, MO, P); Colinas do Sul, 15 canteiro da obra rio São Feliz, 13°22′32″ S, 48°03′49″ W, 19 Sep 2001 (fr), Pereira-Silva et al. 5427 (CEN, MO); Colinas do Sul, 15 entroncamento Vila Veneno-balsa rio São

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Fig. 13.87  Diplusodon punctatus var. punctatus. A. Vegetative branch. B. Inflorescence. C. Bract, adaxial surface. D. Prophyll, adaxial surface. E. Floral bud showing lack of epicalyx. F. Flower with prophylls, without petals, showing lack of epicalyx. (A–F Cavalcanti et al. 1015, CEN)

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Félix-Cavalcante, km 70, 13°36′42″ S, 47°38′51″ W, 21 Feb 2002 (fl), Pereira-Silva et al. 5954 (CEN, MO); Colinas do Sul, 15 APA Pouso Alto, Serra Branca, 13°36′ S, 48°03′54″ W, 1 Sep 2005 (fr), Mendonça et al. 5999 (CEN, IBGE); Minaçu, área de influência da HE Serra da Mesa, 13°46′51″ S, 48°15′ W, 6 Aug 1997 (fl, fr), Oliveira et  al. 872 (CEN, HEPH); Minaçu, estrada Minaçu-Balsa do Rubão, 13°36′56″S, 48°11′19″W, 6 Oct 2000 (fr), Cavalcanti et  al. 2700 (CEN, MO); Minaçu, estrada Minaçu-UHE Serra da Mesa, antigo viveiro, a 7 km da UHE, 19 Jun 1995 (fl), Cavalcanti et  al. 1421 (CEN, NY); Minaçu, margem da estrada Minaçu-Serra da Mesa; a 7 km do canteiro de obra, antigo viveiro, 13º34′S, 48º10′W, 11 Oct 1991 (fl, fr), Cavalcanti et  al. 1015 (CEN [3], K, NY, RB, SPF, UFG); Niquelândia, córrego da Serra, 16 Jun 1993, Hatschbach et al. 59610 (MBM n.v., P); Niquelândia, km 35 da estrada Niquelândia-Colinas do Sul, 14°20′–14°30′ S, 48°20′–48°40′ W, 2 Jul 1996 (fl), Pereira & Alvarenga 3105 (CEN [2], IBGE, P, UB); Niquelândia, bacia de inundação UHE Serra da Mesa, rio Bagagem, 16 Dec 1997 (fr), Amaral-Santos et al. 30 (CEN, MO, NY); Niquelândia, estrada paralela ao rio Tocantins, entro o rio Tocantins e a Serra Negra, 24 Jul 1995 (fl), Cavalcanti et  al. 1649 (CEN, NY); Niquelândia, estrada vicinal à GO-237 (NiquelândiaColinas), entrada 0.6 km da ponte sobre o Rio Bagagem, a 32 km desta, bacia de inundação da Usina Hidrelétrica da Serra da Mesa, 14 Apr 1992 (fl), Walter et al. 1306 (CEN, K, NY, SPF, UFG); Niquelândia, fazenda ao lado da CODEMIN, lado direito do rio Bagagem, área de influência da Usina Hidrelétrica da Serra da Mesa, 14°10′ S 48°22′ W, 15 Apr 1992 (fl), Walter et al. 1380 (CEN, K, NY, SPF, UFG); Niquelândia, área de influência do AHE Serra da Mesa, estrada de terra NiquelândiaMuquém, ca. 21 km a leste de Niquelândia, 14°26′40′′ S, 48°13′53′′ W, 3 Jun 1998 (fl), Walter et al. 4177 (CEN, MO, NY); Niquelândia, área de influência do AHE Serra da Mesa, estrada de terra Niquelândia-Muquém, ca. 3 km antes de Muquém, 14°31′17″ S, 48°09′21″ W, 3 Jun 1998 (fl), Walter et  al. 4187 (CEN, MO, NY); Niquelândia, Macedo, km 2 da estrada para Macedo Velho, 14°21′03″ S, 48°24′52″ W, 27 Jun 1996 (fl), Azevedo et al. 985 (CEN, IBGE); Niquelândia, Macedo, km 2 da estrada para Macedo Velho, 27 Jun 1996 (fl), Fonseca et al. 1016 (CEN, IBGE); Niquelândia, Macedo, 2 km de Macedo, 14°21′03″ S, 48°24′52″ W, 9 Mar 1995, Fonseca et al. 408 (IBGE); Niquelândia, ca. 2 km da estrada para Macedo Velho, 14°21′03″ S, 48°24′52″ W, 9 Aug 1995 (fl), Fonseca et al. 428 (CEN, IBGE); without reference to municipal district: entre Goiás e Cavalcante, Burchell 7659 (K). The two varieties of Diplusodon punctatus are vegetatively similar, with flowers the same size and with the same number of stamens. However, some populations have flowers with a well-defined epicalyx with six spreading segments. I opted to distinguish the populations at level of variety, especially because they can be sympatric, without geographic barriers between them. This may be a representation of genetic variation within populations, but I have not explored this possibility and have kept the two populations as distinct varieties. Diplusodon punctatus was described by Pohl (1827a) without citation of specimens in the protologue. In a subsequent publication, Pohl (1827b) cited a single gathering of D. punctatus collected in September in dry fields near Rio Bagagem, Goiás, without mentioning a collection number.

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Lourteig (1989) cited one specimen of Pohl 2018 = D 946 as the holotype, which stand as the designation of a first-step lectotype (ICBN Art. 9.17, ex. 14). However, I examined three specimens of Pohl 2018 = D 946 at P. The specimen bearing Pohl’s original manuscript label, annotated by Pohl, containing Pohl’s collection number (Pohl 2018) and reference number of his field diary (D = 946) is chosen here as the second-step lectotype (W 0053429). Two additional specimens of Diplusodon punctatus, with labels that indicate Pohl s.n., were examined at G and K. Because duplicates of Pohl’s collections were sent to other herbaria without the information from his field diary (see “Notes about Diplusodon nomenclature and typifications”) and most lack a number, the specimens at G and K are regarded as duplicates of Pohl 2018 = D 946. 69b. Diplusodon punctatus Pohl var. dentatus T.B.Cavalc., Bol. Bot. Univ. São Paulo 22(1): 11, fig. 6. 2004. Type: Brazil. Goiás: Cavalcante, balsa do Rio Tocantins (Serra Branca), para Serra Branca km 2, solo arenoso com afloramento de quartzo leitoso, 13°34′26″ S, 48°05′48″ W, 23 May 2001 (fl), G. Pereira-Silva, J. B. Pereira, K. S. Ribeiro & V. P. Lima 5084 (holotype, CEN [00044717]; isotypes, K, NY, SPF, W). Figs. 13.88 A–D, 13.90 Flowers pedicellate, hypopodium 2.5–3.5 mm long, epipodium 0–0.5 mm long; prophylls 4–4.5 × 1.5–2 mm, reaching the apex of the floral tube or less; epicalyx segments present, 3–4 mm long, spreading. Distribution and Ecology (Fig. 13.90)  Diplusodon punctatus var. dentatus occurs in campos cerrados and campos sujos in Goiás and Tocantins states, sometimes sympatrically with D. punctatus var. punctatus at 330–760 m. Flowers from April to June and from October to November; fruits registered in June and from October to November. Specimens Examined: BRAZIL. Goiás,  Cavalcante, estrada de terra que parte do “Porto Paulista” (balsa) no rio Tocantins para o Buracão, à ca. 4.9 km do rio, 13°28′06″ S, 48°07′08″ W, 7 Nov 2000 (fl, fr), Walter et  al. 4549 (CEN, MO); Cavalcante, estrada Balsa “Porto Paulista” (rio Tocantins), à 1.8 km do rio, 13°29′13″ S, 48°08′16″ W, 7 Nov 2000 (fl, fr), Pereira-Silva et al. 4365 (CEN, MO); Cavalcante, estrada Balsa “Porto Paulista” (rio Tocantins) para HU ca. 32 km da balsa, 13°22′52″ S, 48º W, 9 Nov 2000 (fr), Pereira-Silva et al. 4402 (CEN, MO, NY); Cavalcante, estrada Minaçu-Cavalcanti, passando pela Serra Branca (Coterra), à ca. 20 km do rio Tocantins, 13°40′17″ S, 47°54′41″ W, 10 Nov 2000 (fr), Pereira-Silva et al. 4417 (CEN, MO, NY); Cavalcante, Vila Veneno-estrada São Felix, km 4, 13°33′55″ S, 48°04′34″ W, 27 Jun 2001 (fl, fr), Pereira-Silva & Carvalho-­Silva 5208 (CEN, MO); Minaçu, margem da estrada Minaçu-Serra da Mesa, a 7 km do canteiro de obras, antigo viveiro, 13°34′ S, 48°10′ W, 11 Oct 1991 (fr), Cavalcanti et al. 1014 (CEN); Minaçu, depois de Buriti, 18 km, estrada vicinal, 13°31′08″ S, 48°11′14″ W, 5 Oct 2000 (fr), Cavalcanti et al. 2662 (CEN, MO); Minaçu, estrada obra-rio do Carmo, km 10, 13°37′07″ S, 48°06′10″ W, 21 May 2002 (fl), Pereira-­Silva &

Descriptions

345

Fig. 13.88  Diplusodon punctatus var. dentatus. A. Vegetative branch. B. Leaf, abaxial surface. C. Inflorescence. D. Flower with prophylls, without petals. (A–D Pereira-Silva et al. 5084, CEN)

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Guarino 6460 (CEN, MO). Tocantins, Palmeirópolis, estrada PalmeirópolisAlvorada do Tocantins, à 35 km de Palmeirópolis, 12°57′75″ S, 48°34′41″ W, 6 Apr 1997 (fl), Cavalcanti et  al. 2227 (CEN, MO); Paranã, estrada PalmeirópolisAlvorada de Goiás, a 25 km de Palmeirópolis, 12°54′46″ S, 48°33′42″ W, 6 Apr 1997 (fl), Cavalcanti et  al. 2229 (CEN, NY); Paranã, estrada PalmeirópolisAlvorada de Goiás, a 25 km de Palmeirópolis, 12°54′46″ S, 48°33′42″ W, 6 Apr 1997 (fl), Cavalcanti et al. 2232 (CEN, MO). 70. Diplusodon pygmaeus T.B.Cavalc., Novon 8(4): 347, fig. 7. 1998. Type: Brazil. Goiás: Alto Paraíso de Goiás, estrada vicinal à esquerda, a 5 km S de Alto Paraíso de Goiás no sentido Brasília, GO-118, Serra da Baliza, no topo do morro, 14°11′ S, 47°45′ W, 15 Mar 1995 (fl), T.B.Cavalcanti, F.  R. G.  Salimena-Pires, G.  Pereira-Silva & R.  Mello-Silva 1363 (holotype, CEN [00018545]; isotypes, F, NY, SPF, K, MBM). Figs. 13.89 A–J, 13.90 Small subshrubs, 3–11 cm tall, branched, caespitose, erect, with a broad xylopodium, not dark-brown when dry. Upper branches cylindrical, wine-colored, unwinged, subsericeous, covered with white trichomes; internodes 0.5–1.5 cm long. Leaves decussate, not imbricate, suberect, petiolate; petioles 1–2 mm long; blades 10–25 × 7–15 mm, subcoriaceous, not discolorous, acrodromous, reticulate, ovate to elliptic, base obtuse to subcordate, apex obtuse or slightly acuminate, margin plane, ciliate, with sparse white trichomes; secondary veins 3–4 on each side, 1–2 of them departing from the base of the midvein, prominent, tertiary veins conspicuous; domatia absent. Synflorescence frondose, botryum type; accessory branches absent, principal florescence 1–2 cm long; bracts similar to the vegetative leaves; flowers subsessile, hypopodium 1.5–2 mm long, epipodium 0.5–1 mm long; prophylls 7–8 mm × 2–3 mm, reaching from the middle to the apex or sometimes surpassing the floral tube, elliptic, unkeeled, base acute, apex obtuse to acute, margin plane; floral tube 9–10 mm long, base green, apex wine-colored, narrowly oblongurceolate, sericeous, constricted at the middle in fruit; sepals 4–5 mm long, triangular, not caudate, not deflexed, ciliate, sericeous; epicalyx segments 4–5 mm long, wine-colored, erect to spreading, the same size as the sepals, sericeous, cylindrical; corolla ca. 3 cm in diam., rose, petals obovate, apex obtuse; stamens 12, free portion of the filaments 7–9 mm long; ovary 3.5–4 × 2.5–3 mm, obovate, glabrous; ovules 11–14. Capsules 3.5–4  ×  2.4–3.5  mm, oblong, sometimes with a narrower base, glabrous; seeds 5–7, 1.8–2 × 2–2.3 mm. Distribution and Ecology (Fig.  13.90)  Diplusodon pygmaeus occurs in grassy fields on rocky soil and campo rupestres, at 1500 m, on Serra da Baliza, Chapada dos Veadeiros, Goiás. Flowers in March and April; fruits in April. Specimens Examined: BRAZIL.  Goiás,  Alto Paraíso de Goiás, Chapada dos Veadeiros, estrada de terra a leste, subindo na Serra da Baliza, nas terras do Sr. Paulo, no alto do morro, perto da casinha abandonada, 14°09′35″ S, 47°28′83″ W, 2 Apr 1997 (fl, fr), Cavalcanti et al. 2194 (CEN, MO, NY).

Descriptions

347

Fig. 13.89  Diplusodon pygmaeus. A–B. Habit. C. Section of internode. D, F. Leaf, abaxial surface. E. Detail of leaf indumentum. G. Prophyll, adaxial surface. H. Floral bud. I. Flower with prophylls, without petals. J. Immature capsule. (A–J Cavalcanti et al. 1363, CEN)

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Fig. 13.90  Geographic distribution of Diplusodon puberulus, D. punctatus var. dentatus, D. punctatus var. punctatus, and D. pygmaeus

Descriptions

349

Diplusodon pygmaeus has a diminutive habit, short branches, and a broad xylopodium. The flowers are large and showy, and the floral tube is long and urceolate, as are the capsules. 71. Diplusodon quintuplinervius (Nees) Koehne, Fl. Bras. (Martius) 13(2): 318, pl. 58, fig. 2. 1877. Lythrum quintuplinervium Nees, Flora 4(1): 302. 1821. Type: Brazil. Bahia: Vitória da Conquista, Barra da Vareda, 1829, M. A. P. WiedNeuwied s.n. (lectotype, W, designated by Lourteig 1989: 219; isolectotypes, BR [2], LE). Figs. 13.91 A–H, 13.95 Subshrubs 0.5–1.5 m tall, erect, much-branched, glabrous, dark-brown when dry. Upper branches quadrangular with narrow wings at the angles, dark-brown to darkreddish; internodes 10–25 cm long. Leaves decussate, not imbricate, spreading to suberect, subsessile; petioles 0.5–1 mm long; blades 10–30 × 5–15 mm, dark-brown when dry, coriaceous, not discolorous, acrodromous-basal-supranumerary, ovate or ovate-elliptic to ovate-lanceolate, base obtuse to subcordate, apex acute to obtuse, margin plane, rarely slightly revolute, adaxial surface with veins slightly prominent; secondary veins 4–5 on each side of the midvein; domatia present. Synflorescence frondose-bracteose, diplobotryum to triplobotryum type, congested; accessory branches absent, principal florescence often caducous, paraclades 4–28  cm long; bracts similar to the vegetative leaves, except diminishing in size toward the apex of the branch; flowers subsessile, hypopodium absent, epipodium ca. 1 mm long; prophylls 4.5–5 × 1.8–2.1 mm, reaching the middle or slightly more of the floral tube, elliptic to obovate, sometimes slightly keeled, base acute, apex obtuse, margin plane; floral tube 5–6 mm long, oblong; sepals 2.5–4 mm long, triangular, not caudate, not deflexed, with some papillose trichomes on the margin; epicalyx segments 3–4 mm long, cylindrical to dorsiventrally compressed, spreading to suberect, the same size as the sepals; corolla 2.5–3  cm in diam., rose to violet, petals 13–15 × 6–7 mm, oblong, apex obtuse to acute; stamens 12, free portion of the filaments 7–8 mm long, exserted, ca. 1 mm long; ovary ca. 2 × 2.5 mm, rounded, glabrous, style 13–15 mm long, ovules 10–18. Capsules 3–4 × 2.5–3.5 mm, globose, glabrous; seeds 7–16, 1.9–3 × 1.5–2.2 mm. Distribution and Ecology (Fig.  13.95)  Diplusodon quintuplinervius grows in campo sujo and campo rupestre vegetation and transitional campo rupestre/caatinga north of Cadeia do Espinhaço, Minas Gerais, and in Chapada Diamantina, Bahia, at 830–1162 m. In Minas Gerais, it can also be found in wet campo limpo. Flowers from January to March; fruits registered in February to June, and November. Specimens Examined: BRAZIL. Bahia,  Caetité, estrada Brejinho das AmetistasLicínio de Almeida, a 10 km de Brejinho, 14°25′19″ S, 42°33′04″ W, 12 Mar 1994 (fl), Roque et al. (CFCR) 15001 (CEN, ESA, SPF); Jacaraci, subida para Morro do Chapéu, 14°52′51″ S, 42°31′02″ W, 23 Feb 2014 (fr), Roque et al. 4199 (ALCB); Montugatuba, 8  km de Jacaraú, 14°53′07″ S, 42°28′03″ W, 16 Mar 1994 (fl, fr), Souza et al. 5528 (CEN, ESA); ca. 17 km SW da divisa de Minas Gerais-Bahia, ao

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Fig. 13.91  Diplusodon quintuplinervius. A. Branch with inflorescence. B. Section of internode. C–E. Leaves, abaxial surface. F–G. Floral bud. H. Flower with prophylls, without petals. (A, F, H Schwacke 8261, OUPR; B–C Irwin et al. 23070, K; D–E, G Irwin et al. 23531, NY)

Descriptions

351

longo da BR-116, 30 Mar 1976 (fl), Davidse & D’Arcy 11610 (SP); 3 km S da divisa Minas Gerais-Bahia, na BR-116 na intersecção para Grafite, 20 Nov 1985 (fr), Graham 939 (MO, NY, RB). Minas Gerais, Águas vermelhas, 30 km de Curral de Dentro para Águas Vermelhas, 29 Jan 1965 (fl), Belém & Mendes 374 (P n.v., UB); Diamantina, ca. 25 km da cidade na estrada para São João da Chapada, 9 Jun 1991 (fr), Mello-Silva et al. 404 (CEN, SPF); Divisa Alegre, BR-116, km 18 ao Sul da divisa com a Bahia, 15°49′50″ S, 41°23′28″ W, 1 Aug 2003 (st), Cavalcanti et al. 3191 (CEN); Francisco Sá, estrada para Salinas, 11 Feb 1969 (fl, fr), Irwin et al. 23070 (K, NY, P, UB, W); Grão-Mogol, ca. 15 km W de Grão-Mogol, 19 Feb 1969 (fl, fr), Irwin et al. 23531 (NY); Jequitinhonha, 16°12′06″ S, 41°03′08″ W, 25 Mar 2008 (fl), Almeida 1302 (HUFU); Joaquim Felício, 12 km de Joaquim Felício, 31 Oct 1988 (fr), Kral et al. 75419 (CEN, MO, NY, SP n.v., SPF); Joaquim Felício, Serra do Cabral, próximo ao Armazém da Laje, 18 Nov 1997, Hatschbach et al. 67251 (G, MBM, W); Joaquim Felício, Serra do Cabral, 22 Oct 1999, Hatschbach et al. 69503 (ALCB, ESA, INPA, UPCB); Montes Claros, rio das Pedras (a Sudeste de Montes Claros), 28 Mar 1892 (fl), Schwacke 8261 (OUPR); Montezuma, 15° 10′ 20′′S, 42° 26′ 45′′W, 19 Mar 2017 (fl, fr), Sevilha et al. 6673 (CEN); Novorizonte, 16°02′17″ S, 42°23′33″ W, 2 Dec 2016 (fr), Sevilha et al. 6036 (CEN); Rio Pardo de Minas, 15°26′17″ S, 42°26′26″ W, 24 May 2005 (fr), Sevilha et al. 4421 (CEN); Rio Pardo de Minas, 15°26′13″ S, 42°30′36″ W, 22 May 2005 (fr), Sevilha et al. 4229 (CEN); Rio Pardo de Minas, 15°23′36″ S, 42°28′08″ W, 23 May 2005 (fr), Sevilha et  al. 4349 (CEN); without municipality: Riacho das Varas, 25 Mar 1892 (fl), Glaziou 19169b (K, R). The plants of Diplusodon quintuplinervius are totally glabrous, with narrowly winged quadrangular branches, and shiny, ovate to ovate-lanceolate leaves. The flowers have 12 stamens, an oblong floral tube, and spreading epicalyx segments. This species resembles D. buxifolius in its dark-brown habit when dry and also because of its glabrous, shiny, ovate leaves. Diplusodon buxifolius is also found in Minas Gerais, but with a more southern distribution in the state. Diplusodon buxifolius differs from D. quintuplinervius because it has leaves with only three inconspicuous veins and flowers with short epicalyx segments (1.5–2.5 mm long). Koehne (1903) referred to Glaziou 19169 with the same characteristics to Diplusodon helianthemifolius var. pemphoides f. striatifolius. The material Glaziou 19169 has glabrous, shiny leaves with a plane margin, 2–3 veins on each side of the midrib that depart from the base of the midvein, and epicalyx segments that are spreading to suberect. It is not synonymous with D. helianthemifolius var. pemphoides f. striatifolius, but rather with D. quintuplinervius. The label of the type material examined at W, Wied-Neuwied s.n., reports that Diplusodon quintuplinervius was collected in the state of Espírito Santo, in an area known as Barra da Vareda. This reference is puzzling because the species is not otherwise known from this state, where the vegetation is predominantly Atlantic forest (in which Diplusodon does not occur). No material of the species was found in the herbaria of Espírito Santo, and there is no other evidence of this species in the state, nor any reference to the local name Barra da Vareda. In Prince Neuwied’s

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account of his voyage to Brazil (Wied-Neuwied 1821), the passage about the route to Barra da Vareda refers to its position near the mouth of the stream of Vareda that is near Inhobim, in the municipality of Vitória da Conquista, Bahia. The type of the species is thus assumed to have been collected in the state of Bahia, where it is known by several collections. Lourteig (1989) cited Wied-Neuwied s.n. as the holotype, which stands as the lectotype. 72. Diplusodon ramosissimus Pohl, Flora 10, 151. 1827. Type: Brazil. Goiás: in campis graminosis circa Ponte Feio, Oct, J.  B. E.  Pohl 2533 (lectotype, W [0053433], here designated; isolectotypes, G, W [0053432]). Figs. 13.92 A–H, 13.95, 15.3 E Diplusodon ramosissimus var. decipiens Koehne, Bot. Jarb. 23(57): 30. 1897. Type: Brazil. Goiás, Paranã, 27 May 1895 (fl), A. F. M. Glaziou 21414b (lectotype, P [01901859] designated by Lourteig 1989: 221; isolectotypes, A, BR, GH, K, LE, P [01901858], R, S.). Subshrubs 0.6–1.5 m tall, erect, much-branched, with a xylopodium, not darkbrown when dry. Upper branches subquadrangular to quadrangular, unwinged, lax, internodes 1–2 cm long. Leaves decussate, not imbricate, suberect, petiolate; petioles 3–9 mm long; blades 40–95 × 12–50 mm, chartaceous, usually discolorous, eucamptodromous to acrodromous, oval-lanceolate, with black punctae, base obtuse, apex acute, rarely obtuse, margin plane, sometimes irregularly inconspicuous denticulate and with sparse, short, appressed trichomes, glabrescent; secondary veins 3–6 on each side, prominent on the abaxial surface, sulcate on the adaxial surface; domatia present. Synflorescence bracteose, triplobotryum type, lax; accessory branches absent, principal florescence 6–19 cm long, botryum type, frequently deciduous, paraclades of the first order 6–22  cm long, paraclades of the second order 2–7 cm long, bracts of the first order 15–45 × 15–23 mm, the same shape as the vegetative leaves; bracts 15–30 × 10–14 mm, the second order bracts ellipticlanceolate, bracts of the third order 7–15  ×  3–8  mm, elliptic-lanceolate; flowers subsessile, hypopodium 1–1.5 mm long, epipodium absent; prophylls 2–2.5 × 1 mm, not reaching the middle of the floral tube, oblong to oblong-lanceolate, keeled, base retuse, apex obtuse, margin plane; floral tube 4.5–5 mm long, funnel-shaped, greenish, glabrous, rarely with some trichomes; sepals 3.5–4 mm long, slightly larger in fruit, with the sepals ca. 1 mm long, triangular, slightly caudate, not deflexed, with trichomes on the apex forming a tuft in bud; epicalyx segments 2–2.5(−3.5) mm long, cylindrical, rarely with some trichomes, deflexed, shorter than the sepals; corolla 2.5–3 cm in diam., rose-magenta, petals 13–16 × 8–12 mm, obovate, base and apex obtuse; stamens 18, free portion of the filaments 7.5–8.5 mm long, surpassing the sepals ca. 1  mm; ovary ca. 3  ×  1.8  mm, obovate, glabrous; style ca. 13 mm long; ovules 22–24. Capsules 3.5–5.5 × 4–4.5 mm, globose, glabrous; seeds 9–27, 1–3 × 1.8–2 mm. 2n = 30 (Graham and Cavalcanti 2001).

Descriptions

353

Fig. 13.92  Diplusodon ramosissimus. A. Vegetative branch. B. Detail of domatia. C. Inflorescence. D. Bract, abaxial surface. E. Floral bud. F. Prophyll, adaxial surface. G. Flower with prophylls, without petals. H. Ovary. (A–H Cavalcanti et al. 586, CEN)

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Distribution and Ecology (Fig. 13.95)  Diplusodon ramosissimus inhabits cerrado areas in Distrito Federal, to northern Chapada dos Veadeiros and low areas of Rio Tocantins valley, Goiás, as well as Serra dos Cristais, at 580–1000 m, on the border between Goiás and Minas Gerais, and extreme south of Tocantins. Flowers from January to October; fruits from June to October. Specimens Examined: BRAZIL.  Distrito Federal,  Brasília, Reserva Biológica de Águas Emendadas, cerca de 40 km de Brasília, 15°38′ S, 47°37 W, 28 Jun 1982 (fl, fr), Maury 157 (CEN, HEPH, IBGE); Brasília, Reserva Ecológica do IBGE (RECOR), 9 Aug 1978 (fl), Heringer et al. 563 (IBGE, NY); Brasília, Lago Paranoá, 3 Jun 1979 (fl), Heringer 17269 (IBGE, P n.v.); Brasília, Barragem do Paranoá, 15°48′ S, 47°47′ W, 2 Jun 1992 (fl, fr), Cavalcanti et al. 1232 (CEN, SPF); Brasília, 31 May 1983 (fl), Netto 5 (UB); Brasília, rio São Bartolomeu, 1 Jan 1990 (fl), Guala & Filgueiras 1439 (IBGE); Brasília, Bacia do rio São Bartolomeu, 26 May 1980 (fl), Heringer et  al. 4894 (IBGE, P, K); Brasília, próximo ao rio São Bartolomeu, 15º57′S, 47º42′W, 26 Jun 1999 (fl), Calago 208 (CEN, MO, NY); Brasília, Reserva Biológica do IBGE, 31 May 1983 (fl), Pereira 539 (IBGE, P, US); Brasília, Reserva Biológica do IBGE, 28 Jun 1990 (fl, fr), Azevedo 709 (IBGE, UEC n.v.); Brasília, córrego Fazendinha, 15°47′ S, 47º43′W, 22 May 1984 (fl), Neri 13 (UB); Brasília, Hakme 16 (UB); Brasília, córrego Cabeça de Veado, ca. 9.5 km SE da torre de TV de Brasília, 22 Aug 1976 (fl, fr), Ratter 3472 (UB); Brasília, entre Brasília e Fercal, 30 Jun 1964 (fl), Pires 58096 (K, NY); Brasília, 7 Jul 1966 (fl, fr), Lourteig 1846 (K, NY, R); 30 km NE de Brasília, 14 May 1966 (fl), Irwin et al. 15857 (MO, NY); Brasília, bacia do rio Maranhão, 16 Jun 1999 (fl), Calago 175 (CEN, MO); Brasília, Fazenda Palmeiras, rio Maranhão, 15°31′ S, 47°43′ W, 16 Jun 1999 (fl, fr), Calago 147 (CEN, MO, NY); Brasília, bacia do córrego Taquara, área do Cristo Redentor, 27 May 1991 (fl), Silva 1134 (IBGE); Brasília, área de treinamento da aeronáutica, 7 Aug 1990 (fl, fr), Muniz & Brochado 67 (IBGE); Brasília, Fazenda Água Limpa, Campo Experimental da Universidade de Brasília, 15°58′ S, 47°55′ W, 1120 m, 2 May 1986 (fl), Barros 1023 (SPF, UB); Brasília, Fazenda Água Limpa, Campo Experimental da Universidade de Brasília, Barros 1026 (SPF, UB); Brasília, proximidades do Centro Olimpico da Universidade de Brasília, 2 Jul 1985 (fl), Gonçalves 15 (UB); Brasília, na margem do vale do córrego Capão da Onça, 15°49′ S 47°38′ W, 23 Jun 1982 (fl, fr), Kirkbride, Jr. 4854 (CEN, UB); Brasília, perto do córrego Terra Branca, 15°34′ S, 47°45′ W, 10 Jul 1980 (fl, fr), Kirkbride, Jr. & Kirkbride 3537 (NY, UB); Brasília, SW do córrego Almecegas, 15°33′ S, 48°10′ W, 16 Jun 1982 (fl), Kirkbride, Jr. 4821 (CEN, UB); Brasília, córrego Fazendinha, 15°47′ S, 47°43′ W, 22 Apr 1984 (fl), Ferreira 7 (UB); Brasília, Jardim Botânico de Brasília, 15º52′S, 47º51′W, 19 Apr 2000 (fl), Paiva et al. 73 (HEPH); Brasília, Fazenda Sucupira, 15°55′ S, 48°01′ W, 9 Jun 1999 (fl, fr), Faria et al. 191 (CEN, MO); Brasília, Fazenda Sucupira, Faria et  al. 185 (CEN, MO); DF-100, 15°53′ S, 47°23′ W, 5 May 1992 (fl), Dias et al. 66 (CEN, MO); Brasília, Fazenda Sucupira, na borda da mata do Riacho, 15°52′ S, 48º W, 4 Mar 2000 (fl), Guarino & Pereira 299 (CEN, MO); Brasília, bacia do rio Maranhão, 15°30′ S, 47°41′ W, 16 Jun 1999 (fr), Calago 152 (CEN, MO); Brasília, fazenda córrego D’Anta, após escola Divino Pai Eterno, 4 km seguindo sempre a direita pelas estradinhas, 14°26′

Descriptions

355

S, 48°58′ W, 5 Aug 1992 (fl), Walter et al. 1891 (CEN); Brazlândia, Chapada da Contagem, ca. 20  km E de Brasília, 15 Aug 1964 (fl), Irwin & Soderstron 5135 (NY); Brazlândia, próximo ao Poço Azul, Lago Oeste, APA da Cafuringa, 15 May 2006 (fl), Pastore & Suganuma 1509 (CEN); Brazlândia, APA da Cafuringa, região da Palma, 15°34′ S, 48°02′ W, Leme 21 (UB); Gama, 3 km da estrada do Gama à Goiânia, 16°02′ S, 48°06′ W, 1 Oct 1986 (fr), Nascimento 110 (UB); Saia Velha, DF 495, clube águas Correntes, 16°03′ S, 47°56′ W, 4 Jun 2004 (fr), Proença & Harris 2769 (CEN, UB); Sobradinho, ca. 20 km E de Brasília, 1000 m, 6 Jul 1966 (fl), Irwin et al. 18037 (NY); 15°34′26″ S, 47°54′19″ W, 3 Jun 2004 (fl, fr), Pastore et al. 930 (CEN); Sobradinho, Fercal, 23 Jul 1964 (fl, fr), Duarte & Mattos 8232 (RB, UB); Sobradinho, áreas da Fercal, a 6.7 km da fábrica de cimento Ciplan, na DF-250 leste, 27 Jul 1990 (fr), Bianchetti et al. 895 (CEN, MO); Sobradinho, Fercal, APA da Cafuringa, Sobradinho, Fercal, APA da Cafuringa, a 38 km do Cenargen, 910 m, 9 Aug 1980 (fl, fr), Cavalcanti et  al. 586 (CEN, SPF); Núcleo Rural Tabatinga, Fazenda Palmeirinha, lote 49, Colonia Agricola Stanislaw, acesso pela DF-15, 23 Jun 1984 (fl), Alvin 20 (UB); Planaltina, ca. 55 km E de Brasília na estrada para Planaltina, 13 Sep 1964 (fr), Irwin & Soderstron 6165 (NY); Planaltina, 3 km ao sul da ponte sobre o rio São Bartolomeu, na DF-250, entrada a 500 m desta, 15°44′ S, 47°41 W, 20 May 1992 (fl), Dias et al. 124 (CEN, MO); Taguatinga, lado direito da estrada Taguatinga-Nucleo Bandeirante, a aproximadamente 5  km do centro de Taguatinga, 3 Jun 1984 (fl), Brito 14 (UB); without reference to municipal district: 20 km NE de Brasília, 7 Jul 1966 (fl, fr), Hunt & Ramos 6303 (NY, UB); a 10 km da divisa com Goiás, ao lado da BR-060, 7 Jul 2004 (fl, fr), Pastore 1043 (CEN); Brasília, BR-020, Brasília-Formosa, 15°35′45″ S, 48°38′45″ W, 28 May 2001 (fl), Cavalcanti et al. 2715 (CEN, MO). Goiás, 60–80 km S de Brasília, entre Cristalina e Luziânia, 8 Jul 1976 (fl, fr), Davis & Sheferd D.60025 (NY); Alto Paraíso de Goiás, 14°08′54″ S, 47°43′42″ W, 4 Jul 1998 (fl), Mendonça et  al. 3499 (CEN, IBGE); Alto Paraíso de Goiás, estrada Alto Paraíso de Goiás-Colinas do Sul, caminho para o Vale da Lua, 3 Jun 1999 (fl), Cavalcanti et al. 2502 (CEN, MO); Alto Paraíso de Goiás, São Jorge, 20 May 2003 (fl), Pastore & Suganuma 649 (CEN); Campinaçu, estrada de terra Minaçu-canteiro de obras do AHE Cana Brava, cerca de 12  km do estádio de futebol, 13°24′56″ S, 48°11′27″ W, Oct 2000 (fr), Cavalcanti et al. 2646 (CEN, MO); Catalão, BR-050, Brasília-Catalão, 17°52′48.5″ S, 47°41′34″ W, 7 May 2016 (fl), Cavalcanti et al. 3993 (CEN); Cavalcante, entroncamento Vila Veneno-balsa do rio São Félix-Cavalcante, km 3.2, 13°40′53″ S, 47°51′46″ W, 21 Feb 2002 (fl), Pereira-Silva et al. 5947 (CEN); Cavalcante, estrada Teresina de Goiás-Cavalcante, 5 km antes de Cavalcante, 9 Aug 1990, Cavalcanti & Ayres 1256 (CEN); Cavalcante, ponte sobre o rio Tocantins-Cavalcante, km 61, 13°39′15.5″ S, 47°58′36.5″ W, 30 Aug 2004 (fl, fr), Cavalcanti et al. 3514 (CEN, SPF); Cavalcante, canteiro de obra rio São Félix, km 12, marcados do dique, próximo ao Curral de Pedras, 13°22′12″ S, 48°04′19″ W, 22 Aug 2002 (fl, fr), PereiraSilva et al. 5305 (CEN, MO); Cavalcante, canteiro de obras, próximo à estação de tratamento de água, 13°23′43″ S, 49°08′13″ W, 22 May 2001 (fr), Pereira-Silva et al. 5056 (CEN); Cavalcante, estrada obra-rio São Félix, km 21, 13°37′ S, 48°06′ W, 22 Apr 2002 (fl), Pereira-Silva & Guarino 6467 (CEN, MO); Colinas do Sul,

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cerrado à margem direita do rio Tocantinzinho, 13°55′32″ S, 48°15′45″ W, 22 Jun 1999 (fr), Pereira-Silva et al. 4160 (CEN); Cristalina, 16°54′ S, 47°40′ W, 22 Jun 1997, Proença & Oliveira 1775 (CEN, UB); Cristalina, mata à esquerda da ponte (margem direita), que dá acesso à guarita da saída para Palmital, 16°12′35″ S, 47°20′24″ W, 15 May 2002 (fl), Amaral-Santos et al. 1175 (CEN, MO); Cristalina, 1.5  km após a fazenda do Sr. Edileno (após a ponte sobre o rio Preto, sentido Palmital-BR-251, na bifurcação à esquerda, 14 May 2002 (fl), Amaral-Santos et al. 1126 (CEN); Cristalina, cerrado do lado esquerdo da área de empréstimo, 16°13′5″ S, 47°20′19″ W, 15 Aug 2002 (fr), Amaral-Santos et al. 1424 (CEN, MO); Cristalina, abaixo da área de empréstimo, 16°13′19″ S, 47°20′07″ W, 27 Jun 2002 (fl, fr), Amaral-Santos et al. 1316 (CEN, MO); Cristalina, 16°13′24″ S, 47°20′01″ W, 14 Aug 2002 (fr), Amaral-Santos et al. 1391 (CEN, MO); Formosa, 29 Apr 1965, Irwin et al. 15540 (UB); Formosa, Fazenda Indaiá, 5 May 2003 (bt), Pastore & Suganuma 584 (CEN); Formoso, Alto da Serra Grande, 15 Jun 1972 (fl), Rizzo 8156 (UFG); Luziânia, 15 Jun 1984 (fl), Rodovia Brasília, km 17, lotes 8 e 9, Quadra 12, Magalhães 24 (UB); Minaçu, 13°47′ S, 48°17′ W, 26 Mar 1992, Cavalcanti et al. 1458 (CEN); Minaçu, Serra da Mesa, margem do córrego Bateinha, 10 Oct 1985 (fl, fr), Silva 1874 (R); Niquelândia, 14°01′ S, 48°17′ W, 20 Jul 1995 (fr), Cavalcanti et al. 1496 (CEN, MO); Niquelândia, nas margens do rio Tocantinzinho, 13°59′00″ S, 48°18′00″ W, Cavalcanti et  al. 1575 (CEN, MO, NY); Niquelândia, estrada Niquelândia–Indianópolis, ca. 10  km a NE de Indianópolis (passando a cidade) 14°14′29.9″S, 48°32′36″W, 11 Sep 1998 (fr), Souza et  al. 21536 (CEN, ESA); Niquelândia, Fazenda Ponte Alta, 14°38′ S, 48°58′ W, 8 Jul 1992 (fl), Pereira-Silva et al. 1227 (CEN, MO); Niquelândia, 3.7 km sul da GO - 237, a 5 km da sede da Fazenda Ouro Fino, área de influência da UHE Serra da Mesa, 14°33′ S, 48°58′ W, 7 Jul 1992 (fl), Pereira-Silva et al. 1142 (CEN, MO, NY); Niquelândia, 3.7 km sul da GO - 237, a 5 km da sede da Fazenda Ouro Fino, área de influência da UHE Serra da Mesa, Pereira-Silva et al. 1143 (CEN, MO); Niquelândia, base da vertente sudeste da Serra Negra, 14°02′46″ S, 48°21′40″ W, 8 Jul 1992 (fl), Cordovil-Silva et al. 609 (CEN, MO, NY); Niquelândia, margem esquerda do rio Bagagem, 9 Sep 1997, Cordovil-­Silva et al. 671 (CEN, MO); Niquelândia, margem esquerda do rio Traíras, a 10  km da ponte sobre este rio (descendo), próximo a Indaianápolis, 14°14′ S, 48°34′ W, 9 Jun 1992 (fl, fr), Walter et  al. 1536 (CEN, MO, NY); Niquelândia, região da Serra Negra, margem esquerda do rio Bagagem, próximo à Fazenda Aroeira, após a CODEMIN, Bacia de inundação da UHE Serra da Mesa, 14°07′ S, 48°23′ W, 15 Apr 1992 (fl), Walter et al. 1358 (CEN); Niquelândia, Fazenda Serra Negra (Niqueltocantins), arredores da Serra Negra em estrada para o rio Bagagem, 14°02′ S, 48°18′ W, 8 Jun 1992 (fl), Walter et al. 1436 (CEN, RB); Posse, 17 Aug 1990, Cavalcanti et al. 815 (CEN); São João da Aliança, 21 May 2001 (fl), PereiraSilva et al. 5041 (CEN, MO); São João da Aliança, Fazenda Barra Mata Serena, 18 Sep 1990 (fl, fr), Vaz 784 (IBGE); São João da Aliança, 72 km do entroncamento da BR-020/DF-345/GO-3, 15°38′18″ S, 47°36′03″ W, 15 Aug 1990, Cavalcanti et al. 651 (CEN); São João da Aliança, entroncamento DF 345, 72 km do entroncamento da BR 020-DF 345, 15 Aug 1990, Cavalcanti et al. 657 (CEN); Uruaçu, região a mais ou menos 800 m da margem direita do rio Acaba Saco e rio Maranhão, Fazenda

Descriptions

357

córrego D’Anta, 14°26′ S, 49º W, 4 Aug 1992 (fl, fr), Walter et al. 1848 (CEN); Uruaçu, próximo a Ponte Nova, entre Uruaçu e Niquelândia, 14°32′56″ S, 49°01′30″ W, 18 Jun 1997 (fl, fr), Cordovil-Silva et al. 622 (CEN); entre Goiás e Cavalcante, Burchell 7726 (K); entre Cavalcante e Conceição (fr), Burchell 7935 (K); Burchell 7791 (K); without reference to municipal district: Riedel 2595 (LE n.v., W); Riedel s.n. (K). Minas Gerais, Unaí, arredores da cidade de Palmital, 15 May 2002 (fl), Amaral-Santos et al. 1205 (CEN, MO); Unaí, mata baixo do vertedouro, 16°13′13″ S, 47°19′29″ W, 25 Jun 2002 (fl, fr), Amaral-Santos et al. 1281 (CEN, MO); Unaí, Cabeceira Grande, 2 km sudoeste da ponte do rio Preto, divisa de Minas Gerais com Goiás, 17 May 2001 (fl, fr), Cavalcanti et  al. 2712 (CEN, MO). Tocantins, Palmeirópolis, Linha de transmissão, torre 79, 13°08′29″ S, 48°13′15″ W, 14 Jun 2008 (fl), Pereira-Silva et al. 13,470 (CEN). Diplusodon ramosissimus is one of the several species in the genus with a welldifferentiated inflorescence from the vegetative parts, i.e., a bracteose inflorescence, in which the bracts are morphologically distinct from the vegetative leaves. This characteristic also occurs in D. astictus, D. bradei, and D. panniculatus, among others. Other distinctive characteristics of D. ramosissimus include its frequently totally glabrous, well-developed leaves up to 9 cm long and evident domatia on the abaxial leaf surface and bracts. The flowers have 18 stamens, the prophylls are shorter than half the length of the floral tube, and the epicalyx segments are short (2–2.5 mm long) and deflexed. In addition, trichomes on the apex of the sepals produce an evident tuft when the sepals are united in the bud. Diplusodon ramosissimus was described by Pohl (1827a) without citation of specimens in the protologue. In a subsequent publication, Pohl (1827b) cited a single gathering of D. ramosissimus collected in January in grass fields, in the state of Goiás, without mentioning a collection number. At W, there are two specimens numbered (Pohl 2533), from the same locality and annotated by Pohl, of which one has an original label and was chosen here as lectotype [W 0053433]. A duplicate of the collection by Pohl, Pohl s.n., was also located at G. It appears to be from the same locality as the lectotype of Diplusodon ramosissimus, but the label lacks details. Given that it is well known that many duplicates of Pohl’s collections were distributed before they received a number (see “Notes about Diplusodon nomenclature and typifications”), the specimen at G is treated as an isolectotype of D. ramosissimus. Koehne (1897) described Diplusodon ramosissimus var. decipiens, which was characterized as having only a few trichomes on both surfaces of leaves of the lower branches and a totally glabrous inflorescence. In studying various populations of D. ramosissimus, it is possible to find individuals with the full range of pubescence variation that distinguishes the varieties (pubescent, glabrescent, or glabrous). Therefore, my observations support the synonymization proposed by Lourteig (1989). Lourteig (1989) indicated for Diplusodon ramosissimus var. decipiens a specimen at P (Glaziou 21,414) as holotype, what is considered as the lectotype (ICBN Art. 9.17, ex. 14).

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73. Diplusodon rectinervis T.B.Cavalcac., sp. nov. Type: Brazil. Minas Gerais: Delfinópolis, “Condomínio das Pedras”, Parque Nacional da Serra da Canastra, 20°21′64″ S, 46°44′83″ W, 28 Nov 2003 (fl), R. Romero 7000 (holotype, CEN [00051466]; isotype, BHCB, HUFU, K, NY, P, RB, SPF, W). Figs. 13.93 A–F, 13.95 This species is characterized by canescent branches covered by long trichomes, small leaves, with acrodromous-basal venation with almost parallel veins and an inconspicuous midvein. Shrubs 1 m tall, erect, branched, foliage congested, not dark-brown when dry. Upper branches subquadrangular to quadrangular, unwinged, rust-colored, hirsute, internodes 0.4–0.6 cm long. Leaves decussate, not imbricate, spreading to suberect, sessile; blades 5–10  ×  3–6  mm, coriaceous, not discolorous, acrodromous-basalsupranumerary, ovate, oblong-ovate to elliptic, base retuse, cordate to rarely obtuse, apex obtuse to acute or sometimes acuminate, margin revolute to subrevolute, adaxial surface glabrous, with rare trichomes, abaxial surface densely hirsute on the veins, long trichomes; secondary veins 3–4 on each side of midvein, 2–3 of them departing from the leaf base, secondary veins almost parallel; domatia absent. Synflorescence frondose, diplobotryum to triplobotryum type, congested; accessory branches absent, principal florescence ca. 2 cm, paraclades 3–10 cm long; bracts similar to the vegetative leaves; flowers sessile, hypopodium absent mm long, epipodium ca. 0.5 mm long; prophylls 6–8 × 2–2.6 mm, surpassing the apex of the floral tube 1.5–6 mm, elliptic, not keeled, base acute, apex obtuse, margin plane, ciliate, densely covered with long and white trichomes; floral tube 5.5–6 mm long, oblong, covered with white and long trichomes; sepals ca. 2 mm long, triangular, not caudate, not deflexed; epicalyx segments 2–2.8 mm long, narrowly lanceolate, spreading, slightly longer than the sepals, long-ciliate; corolla 1.5–2 cm in diam., clear rose-magenta, petals 8–9 × 6–7 mm, obovate, apex obtuse; stamens 12, free portion of the filaments 8.5–9 mm long, exserted; ovary ca. 2.1 × 1.5 mm, globose, glabrous, style 14–15 mm long, ovules ca. 16. Capsules not seen. Distribution and Ecology (Fig. 13.95)  Diplusodon rectinervis is known only from the type collected in the National Park of Serra da Canastra, Minas Gerais State, and in vegetation of campos rupestres with rocky sandstone outcrops at 1178 m. Flowers in November. Specimen Examined. BRAZIL.  The type. Diplusodon rectinervis is characterized by its small (5–10 × 2–6 mm) and acrodromous leaves that have almost parallel veins and an undifferentiated midrib, in contrast to the prominent midrib on the abaxial surface of other species in the genus. In addition, the branches and leaves of D. rectinervis are covered by long trichomes, the flowers have long prophylls surpassing the floral tube, the epicalyx segments are spreading and long-ciliate, and the androecium consists of 12 stamens.

Descriptions

359

Fig. 13.93  Diplusodon rectinervis. A. Branch with inflorescence. B. Section of internode. C. Leaf, adaxial surface. D. Leaf, abaxial surface. E. Prophyll, adaxial surface. F. Flower with prophyll, without petals. (A–F Romero et al. 7000, CEN)

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74. Diplusodon retroimbricatus Koehne, Bot. Jahrb. Syst. 23(Beibl. 57): 34. 1897. Type: Brazil. Goiás: nos campos de Formosa, 10 Dec 1894, A.  F. M. Glaziou 21417 (lectotype, P [01901858], designated by Lourteig 1989: 241; isolectotypes, BR, F, K). Figs. 13.94 A–E, 13.95 Shrubs ca. 0.4–1.5 m tall, erect, branched, canescent to sometimes silvery, rarely dark-brown when dry. Upper branches cylindrical, unwinged, canescent, densely velutinous, indumentum long, dense, and whitish, internodes 1.5–5 cm long. Leaves decussate, sometimes imbricate, spreading to suberect, subsessile to short-petiolate; petioles 0.5–3.5  mm long; blades 20–55  ×  15–34  mm, coriaceous, discolor, acrodromous-­basal-supranumerary, ovate to ovate-lanceolate, base obtuse to subcordate, apex obtuse, subacuminate to acuminate, margin subrevolute, blades densely hirsute to sericeous, covered by long trichomes with the base slightly thickened, whitish to silver, adaxial surface gray-green, abaxial surface white, sometimes silvery; secondary veins 8–10 on each side, 1–2 of them departing from the base of the midvein; domatia absent. Synflorescence frondose-bracteose, botryum type, congested; accessory branches absent; bracts ovate to ovate-lanceolate, 10–15 × 7–10 mm; flowers sessile, prophylls 8.5–10 × 6.5–8 mm, reaching or surpassing the apex of the floral tube, obovate to rhombic, keeled, base acute, apex obtuse, margin plane, sometimes with involute margin at the apex, sericeous; floral tube 7.5–8 mm long, whitish, funnel-shaped, increasing in size on the fruit, densely sericeous; sepals 4.5–5 mm long, triangular, slightly caudate, not deflexed, ciliate, sericeous; epicalyx segments 2–3.5 mm long, triangular, erect, reaching to half the height of the sepals, sericeous; corolla 5–6  cm in diam., clear rose, petals 18–20 × 10–12 mm, obovate, apex obtuse; stamens (18–) 24–26, free portion of the filaments ca. 9  mm long, slightly exserted; ovary 2–2.9  ×  2.9–3.5  mm, obconic, glabrous, style ca. 15 mm long, ovules 20–30. Immature capsules ca. 4 × 4 mm, globose, glabrous; seeds 25–27, 1.9–2 × 1.2–1.5 mm. Distribution and Ecology (Fig.  13.95)  Diplusodon retroimbricatus was known only from historical gatherings until new collections were made from 2002 to 2004 in Chapada dos Veadeiros, Goiás. Although it has been recollected, D. retroimbricatus remains a rare species. It grows in campo limpo and campo sujo on sandy soil at 820–919 m. Flowers from January to March; fruits registered in March. Specimens Examined: BRAZIL. Goiás,  Cavalcante, Fazenda Renascer, 20 Apr 2003, Pastore et  al. 467 (CEN); Cavalcante, Fazenda Renascer, caminho para a ponte de pedra, 3 Feb 2004 (fl, fr), Pastore et al. 791 (CEN); Cavalcante, em frente à Fazenda São Vicente, caminho para a Cachoeira do Prata, 3 Feb 2004 (fr), Pastore et al. 844 (CEN); Cavalcante, entroncamento vila Veneno-balsa do rio São Félix, km 39, 13°40′31″ S, 47°49′24″ W, 21 Feb 2002 (fl), Pereira-Silva et al. 5951 (CEN); Cavalcante, Reserva Natural Serra do Tombador (RPPN), estrada municipal não pavimentada Cavalcante-Minaçu, 13°40′34″ S, 48°47′9″ W, 21 Feb 2013 (fl, fr), Lombardi et al. 10027 (CEN, HRCB n.v.); Cavalcante, RPPN Serra do Tombador, 3  km da sede, área conhecida como campina, 13°40′51″ S, 47°49′06″ W, 31 Jan 2015 (fl), Simon et al. 2526 (CEN).

Descriptions

361

Fig. 13.94  Diplusodon retroimbricatus. A. Branch with inflorescence. B. Leaf, abaxial surface. C. Detail of branch indumentum. D. Prophyll. E. Flower with prophyll, without petals. (A–E Pastore et al. 791, CEN)

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Fig. 13.95  Geographic distribution of Diplusodon quintuplinervius, D. ramosissimus, D. rectinervis, and D. retroimbricatus

Descriptions

363

Diplusodon retroimbricatus was known only from the diagnosis of Koehne (1897) and the type material. In his diagnosis, Koehne (1897) compared D. retroimbricatus with D. villosissimus, distinguishing the two based on the retroflexed leaves and manifestly caudate sepals of D. retroimbricatus. The retroflexed leaves are now known to be an artifact resulting from pressing. Nonetheless, D. retroimbricatus can be distinguished by its densely sericeous indumentum covering the whole plant, its strongly congested inflorescences with flowers hidden by the bracts, and its short epicalyx segments (2–3.5 mm long). The diagnosis of Diplusodon retroimbricatus was made by Koehne (1897) based on material deposited at B that was destroyed (F neg. nº 017930). Lourteig (1989) cited a specimen at P as the holotype, which is here considered as lectotypification (ICBN Art. 9.17, ex. 14). 75. Diplusodon rosmarinifolius A.St.-Hil., Fl. Bras. Merid. (A.St.-Hil.) 3: 147, pl. 189, fig. a. 1833. Type: Brazil. Goiás: en Mai sur la plaine élevee qui s’étend des limites de la province des Mines au village de S Luzia prov. de Goyaz, A.Saint-Hilaire s.n. C1 644 (lectotype, P [01902028], designated by Lourteig 1989: 228; isolectotypes, OXF, P [01902029, 01902030]). Figs. 13.96 A–G, 13.99 Subshrubs 0.5–2.5 m tall, erect, much-branched, with a xylopodium, not darkbrown when dry. Upper branches cylindrical to subquadrangular, unwinged, lax, covered by stellate trichomes, the young branches rust-colored, internodes ca. 1 cm long. Leaves decussate, not imbricate, spreading, subsessile; petioles ca. 1  mm long; blades 15–20 (−35) × 2–3 (−7) mm, coriaceous, not discolorous, eucamptodromous, linear to linear-lanceolate, base acute, apex acute, margin revolute; secondary veins 1 on each side, prominent on the abaxial surface, sulcate on the adaxial surface, adaxial surface with few trichomes evident, abaxial surface canescent with dense indumentum of stellate trichomes; domatia absent. Synflorescence frondose, pleiobotryum spiciform type, lax; accessory branches absent, principal florescence 6–10  cm long, botryum spiciform type, paraclades 6–20  cm long; bracts 17–25 × 1–5 mm, linear-lanceolate to linear, margins revolute; flowers subsessile, hypopodium 0–0.5 mm long, epipodium 0–0.5 mm long, prophylls 3–3.5 × 1–1.5 mm, less than half the length of the floral tube, oblong-linear, unkeeled, base obtuse, apex obtuse, margin plane; floral tube 3–3.5 mm long, wine-colored, funnel-shaped, covered by inconspicuous stellate trichomes; sepals ca. 2 mm long, triangular, not caudate, not deflexed; epicalyx segments 0.5–1 mm long, cylindrical, spreading to slightly deflexed, not exceeding the length of the sepals; corolla 2–2.5 cm in diam., rose-magenta to clear rose, petals 9–10 × 6–7 mm, obovate, apex obtuse; stamens 15, free portion of the filaments 6–6.5 mm long, exserted, surpassing the sepals ca. 1 mm; ovary ca. 1.5 × 2 mm, rhombic, glabrous; style 7.5–8 mm long, ovules 9–14. Capsules 2.8–3 × 2.8–3 mm, globose, glabrous; seeds 9–12, ca. 2 × ca. 2.1 mm. Distribution and Ecology (Fig. 13.99)  Diplusodon rosmarinifolius is known by a few collections in the state of Goiás at about 850 m and numerous collections from Distrito Federal, on red latosols, at 1000–1200 m. It can also be found in campo

364

13  Systematic Treatment

Fig. 13.96  Diplusodon rosmarinifolius. A. Branch with inflorescence. B. Leaf, adaxial surface. C. Leaf, abaxial surface. D. Detail of leaf indumentum. E. Multicellular trichome. F. Floral bud. G. Flower with prophyll, without petals. (A–G Cavalcanti et al. 478, CEN)

Descriptions

365

limpo and campo de murundus. Flowers from March to October; fruits from March to October. The species was reported by Koehne (1903) from Minas Gerais, based on Gardner 4563; however, this gathering is here identified as D. puberulus. Specimens Examined: BRAZIL.  Distrito Federal,  Brasília, córrego do Tororó, 24 Jun 1985 (fr), Xavier 7 (UB); Brasília, estrada Brasília-Cocalzinho, km 30, 15°40′ S, 48°20′ W, 14 May 1991 (fl), Vieira & Walter 750 (CEN, MO); Brasília, 2 km NE of Brasília, 16 Jul 1964 (fl, fr), Prance & Silva 58174 (MO, UB); Brasília, a 32 km do Cenargen, DF-170, seguir 6 km, 15°33′ S, 48°03′ W, 4 Jun 1992 (fl), Dias & Vieira 152 (CEN, MO); Brasília, a 7 km do entroncamento com a DF-001, 12 Jul 1990, (fl), Cavalcanti et al. 531 (CEN, SPF); Brasília, APA da CAfuringa, Mumunhas, 15 Jun 1985, Pena s.n. (UB); Brasília, Tororó, 25 Jun 1985 (fl, fr), Rodrigues 10 (UB); Brasília, APA da Cafuringa, Chapada da Contagem, 17.5 km due north of center of Brasília, at north edge of the Chapada da Contagem, 31 May 1973 (fl), Taxonomy Class of Universidade de Brasília 112 (UB); Brasília, Área da Aeronáutica, 5 Jul 1990 (fl), Azevedo & Alvarenga 775 (IBGE, UEC n.v.); Brasília, Asa Norte-residencial, 21 Jul 1967 (fl), Anhanguera 42 (UB); Brasília, bacia do rio Maranhão, 16 Jun 1999 (fl), Calago 182 (CEN, MO, NY); Brasília, bacia do rio São Bartolomeu, 8 Jul 1983 (fl, fr), Pereira 647 (CPAC n.v., IBGE, K); Brasília, bacia do rio São Bartolomeu, 28 May 1980, Heringer et al. 4939 (IBGE, K, MO); Brasília, ca. 20 km E de Brasília, 14 Oct 1965 (fl, fr), Irwin et al. 9222 (NY, UB); Brasília, ca. 20 km E de Brasília, 15 Feb 1982 (fl), Proença 47 (UB); Brasília, Cabeça de Veado, caminho da Papuda, 2 Sep 1960 (fl, fr), Andrade 320 & Emmerich 312 (R); Brasília, Cabeceiras do rio Gama, 3 May 1896 (fl), Glaziou 21424 (BR, K, NY); Brasília, campus da UNB, 23 Aug 1975, Oldenburger & Mecenas 1600 (UB); Brasília, Cemitério Campo da Esperança, 17 Jun 1984 (fl, fr), Salles 26 (UB); Brasília, Cerrado atrás do HFA, 9 Jun 1985 (fl), Senna s.n. (UB); Brasília, Colorado, DF-001, km 11.3 do Posto Colorado, 12 Jul 1990, (fl), Cavalcanti et al. 478 (CEN, SPF, K, NY); Brasília, DF-170, 3 km do entroncamento da DF-001, 18 Feb 1999 (fl), Cavalcanti et al. 2491 (CEN, MO, NY); Brasília, condomínio Belo Horizonte, entrada para a Ermida Dom Bosco, ao lado da QL 28, 15°48′11″ S, 47°48′26″ W, 29 Sep 2001 (fl, fr), Pereira-Silva 5493 (CEN, MO); Brasília, córrego do Tororó, 24 Jun 1985 (fr), Bernardino 16 (UB); Brasília, córrego do Tororó, Araújo 9 (UB); rodovia, próximo à cachoeira do Tororó, 47°42′30″ W, 24 Jun 1985 (fl, fr), MaiaFilho 11 (UB); Brasília, córrego do Tororó, 15°55′ S, 47°48′ W, 17 Jun 1985 (fl, fr), Guimarães-Neto 14 (UB); Brasília, córrego do Tororó, 17 Jun 1985 (fl, fr), Junior 16 (UB); Brasília, entre o córrego Piripau e Formosa, 15°34′79′ S, 47°25′57″ W, 21 Jul 2003 (fr), Proença & Harris 2653 (CEN, UB); Brasília, estrada para o Poço Azul, ao lado do Parque Nacional, Jun 2003 (fl), Pastore et  al. 638 (CEN); Saia Velha, 24 Jun 1975 (fl), Heringer 14725 (IBGE); Brasília, Fazenda Água Limpa, 15°58′ S, 47°55′ W, 24 Apr 1985 (fl), Barros 974 (UB); Brasília, Fazenda Água Limpa, 2 May 1986 (fl), Barros 1013 (SPF, UB); Brasília, Fazenda Água Limpa, 2 May 1986 (fl), Barros 1014 (SPF, UB); Brasília, Fazenda Água Limpa, 2 May 1986 (fl), Barros 1015 (SPF, UB); Brasília, Fazenda Água Limpa, 9 Jun 1976 (fl), Ratter et al. 3133 (K, MO, UB); Brasília, Fazenda Água Limpa, no campo de arnica, 25

366

13  Systematic Treatment

May 2003 (fl), Pastore et al. 604 (CEN); Parque Nacional de Brasília, 14 Apr 1963, Santos 1621 (R); Brasília, Jardim Botânico de Brasília, 15°52′ S, 47°51′ W, s.d., Boaventura 167 (HEPH); Brasília, Jardim Botânico de Brasília, 26 Jun 1987 (fl, fr), Equipe do JBB 963 (HEPH); Brasília, Jardim Botânico de Brasília, 15°52′ S, 47°50′ W, 24 Jun 1995 (fl), Dantas et al. 2 (UB); Brasília, Jardim Botânico de Brasília, área atrás do Jardim Botânico, 15º32′S, 47º51′W, 10 Jul 2000 (fl), Mendes & Nóbrega 20 (HEPH); Brasília, Jardim Botânico de Brasília, trilha ecológica, 2 Jun 1998 (fl), Oliveira et al. 1142 (HEPH); Brasília, Núcleo Bandeirantes, Fazenda Sucupira, à esquerda do capril, próximo da cerca da divisa, 28 Sep 2000 (fr), Amaral-Santos et al. 737 (CEN, MO); Brasília, Núcleo Bandeirantes, Fazenda Sucupira, entre o restaurante e a sede da fazenda, 15°52′ S, 48º W, 9 May 2000 (bud), Guarino & Pereira 179 (CEN, MO); Brasília, Núcleo Bandeirantes, Fazenda Sucupira, 15°55′ S, 48°01′ W, 19 May 1999 (fl), Faria et al. 153 (CEN, MO); 9 Jun 1999 (fl, fr), Faria et  al. 255 (CEN, MO); Brasília, Parque Nacional de Brasília, 26 Mar 1971 (fr), Lucidoi 22 (HEPH); Brasília, Parque Nacional de Brasília, 5 May 1962 (fl), Heringer 8935 (UB); Brasília, Rodovia DF-140, condomínio Parque Mirante, 15°57′52″ S, 47°49′10″ W, 22 Jun 2000 (fl, fr), Pereira-Silva & Abdala 4273 (CEN); Brasília, Sobradinho, Fercal, 15°35′ S, 47°53′ W, 20 Dec 1996, Proença 1670 (UB); a cerca de 2 km a leste do balão da Escola Fazendária, 15°50′ S, 47°48′ W, 2 Jun 1992 (fl), Cavalcanti et al. 1220 (CEN, SPF); Brasília, Tororó, 24 Jun 1985 (fl), Passos s.n. (UB); Brasília, Universidade Federal de Brasília, 14 Aug 1964 (fr), Irwin & Soderstrom 5105 (NY, UB); Gama, Parque Nacional do Gama, 25 May 1965 (fl), Sucre 271 (UB); Planaltina, 47°42′30″ W, 22 Jun 1982 (fl), without collector (UB); Planaltina, CPAC-Chapada entre Sobradinho e CPAC a direita da, 22 Jun 1982, Ribeiro & Haridasan 851 (UB); Planaltina, Parque Nacional das Águas Emendadas, 17 Jul 1979 (fl), Heringer et al. 1853 (IBGE, MO); Riacho Fundo, Fazenda Sucupira, área em frente ao capril, 15°54′53″ S, 47°59′59″ W, Oct 2005 (fl, fr), Cavalcanti et al. 3562 (CEN, MO); Sobradinho, 12 Apr 1965 (fl), Heringer 10126 (NY, UB); próximo à BR-003, 25 May 1989 (fr), Barros 1229 (UB); Taguatinga, 9 Sep 1965 (fl, fr), Irwin et  al. 8122 (K, MO, NY, UB); Taguatinga, córrego Vicente Pires, 15°47′ S, 48°01′ W, 27 Mar 1984 (fl), Pereira-Neto 9 (UB); Trevo da BR-020, com a DF-345, 15°35′5″ S, 47°39′17″ W, 28 May 2001 (fl), Cavalcanti et al. 2716 (CEN, MO); Goiás, Formosa, 10 km NW de Formosa, Fazenda J. Teles, 29 Apr 1966 (fl), Irwin et al. 15440 (NY, UB); Luziânia, rodovia Brasília-BH, km 17, lote 8 e 9, 15 Jun 1984 (fl), Magalhães 18 (UB). Diplusodon rosmarinifolius is characterized by its linear-lanceolate leaves that are sparsely arranged on the branches, short epicalyx segments, and androecium of 15 stamens. In addition, there are minute stellate trichomes covering the branches, leaves, and flowers. This dense indumentum confers a flaky aspect to the surface, with tufts of trichomes close to one another. This species resembles D. macrodon in habit, shape of the leaves, and type of indumentum. Diplusodon macrodon differs from D. rosmarinifolius by its long epicalyx segments (5–6 mm long) and androecium of 12 stamens. Diplusodon puberulus has been considered synonymous with D. rosmarinifolius because of vegetative similarity between the two. However, D.

Descriptions

367

puberulus differs from D. rosmarinifolius because it has flowers with 12 stamens, an almost glabrous floral tube, pilose sepals, and epicalyx, and it lacks stellate trichomes (but has short, simple trichomes). All the populations of this species that were examined were uniform in vegetative and floral characteristics with very few exceptions. Some variation was observed in the size of the individuals, which range from 0.5 to 2.5 m in height, but despite these differences, all plants exhibit the typical characteristics of Diplusodon rosmarinifolius. Lourteig (1989) cited the specimen of A.Saint-Hilaire s.n. at P as holotype, what is considered as the lectotype (ICBN Art. 9.17, ex. 14). 76. Diplusodon rotatus T.B.Cavalc., Phytotaxa 226(2): 145, fig. 1–2. 2015. Type: Brazil. Maranhão: Balsas, Lote 19, na descida da escarpa-Agrovila Nova de Carli, 08°41′39″ S, 46°45′45″ W, 4 Aug 1998 (fl), R. C. Oliveira, G. PereiraSilva & M.  M. R.  Cunha 1245 (holotype, CEN [00071996]; isotypes, CEN [00072526], HEPH). Figs. 13.97 A–E, 13.99, 15.4 C Shrubs 1–1.5 m tall, erect, much-branched, not dark-brown when dry. Branches lax, the uppermost delicate, rust to wine-colored, cylindrical compressed, unwinged, minutely pubescent, internodes 12–16  cm long. Leaves decussate, not imbricate, suberect, petiolate; petioles 1.5–2 mm long, thick; blades 20–35 × 12–19 mm, lightgreen, chartaceous, petiolate, not discolorous, light-green when dry, acrodromousbasal-supranumerary, oblong to ovate, base retuse, apex obtuse, acuminate, margin plane to slightly subrevolute, ciliate with short trichomes, adaxial surface minutely pubescent to glabrescent, abaxial surface minutely pubescent; secondary veins 2–3 on each side, 2 of them departing near the base of the midvein, sulcate on the adaxial surface, slightly prominent on the adaxial surface; domatia absent. Synflorescence frondose, pleiobotryum type, branches rust, thin, delicate, lax; principal florescence truncate; paraclades of the first and second order 15–30 cm long; paraclades of the third order 4–7 cm long; bracts oval to elliptic, spread to suberect, base retuse to slightly subcordate, margin plane to slightly subrevolute, pubescent, ciliate with short trichomes, apex obtuse, slightly acuminate, secondary veins 2–3 on each side of the midvein, with domatia; flowers pedicellate, hypopodium ca. 1.5 mm long, epipodium ca. 0.5 mm long; prophylls 4–4.6 × 1.2–2 mm, inserted in the upper third of the pedicel, surpassing the apex of the floral tube, narrowly elliptic, unkeeled, base attenuate, apex acute, margin plane, minutely pubescent; floral tube 4.2– 4.5  mm long, delicate, funnel-shaped, with minute trichomes; sepals 2–2.5  mm long, triangular, not caudate, not deflexed, with minute trichomes; epicalyx segments 4–4.1 mm long, fine, cylindrical, spreading, longer than the sepals, with minute, spiky trichomes; corolla 2–2.5  cm in diam., rose, petals 9–10  ×  5–7  mm, obovate, apex obtuse; stamens 12, free portion of the filaments ca. 5  mm long, slightly exserted; ovary 2–2.4 × 2.2–2.9 mm, obovate, glabrous, style 7–8 mm long, ovules 10–12. Capsules 3.6–4 × 3.5–4 mm, globose to oblong, glabrous; seeds 3–4, 2.5–2.9 × 2–2.1 mm.

368

13  Systematic Treatment

Fig. 13.97  Diplusodon rotatus. A. Branch with inflorescence. B. Section of internode. C. Leaf, adaxial surface. D. Leaf, abaxial surface. E. Flower with prophylls, without petals. (A–E Oliveira et al. 1245, CEN)

Descriptions

369

Distribution and Ecology (Fig. 13.99)  Diplusodon rotatus is a rare species that occurs in Brazilian Northeast, in Maranhão and Piauí states, known from a few collections from cerrado sensu stricto with rocky soil, at 200–447  m. Flowers from June to August; fruits registered in January and October. Specimens Examined: BRAZIL. Maranhão,  Carolina, BR 010-Balneário, ca. 500 m da pista em estrada de terra, 07°01′03″ S, 47°27′27″ W, 19 Oct 2013 (fr), Cavalcanti & Pereira-Silva 3780 (CEN, MO, NY, UFMA, W); Carolina, estrada BR 010-Balneário Encontro das Águas, km 01, área de influência indireta, 07°21′03″ S, 47°27′46″ W, 19 Jan 2010 (fr), Pereira-Silva & Moreira 14965 CEN, HUEFS, SP). Piauí, Currais, 26 Jun 2001 (fl), Barros 1281 (HUFU). Diplusodon rotatus is a shrub that loses its vegetative leaves during anthesis. The vegetative leaves are quite different from the bracts, and the two are not present on the plant at the same time. The vegetative leaves of D. rotatus are oblong and lack domatia, whereas the bracts are oval and have domatia. Diplusodon rotatus is characterized by the presence of minute puberulent indumentum throughout the plant, flowers with 12 stamens, evident epicalyx segments that are strongly spreading, and prophylls positioned in the upper third of the pedicel. The inflorescence is delicate and has small flowers. 77. Diplusodon rotundifolius Mart. ex DC., Prodr. 3: 94  f. 1828. Friedlandia rotundifolia Mart. ex DC., Prodr. 3: 94f, 1828, pro syn. Type: Brazil. Minas Gerais: Habitat in campis prope Vila Rica s. Ouro Preto, C. F. P. Martius s.n. (lectotype, M [0146263], designated by Lourteig 1989: 234; isolectotypes, fragm. G, OXF). Figs. 13.98 A–E, 13.99 Subshrubs 0.5–2  m tall, erect, branched, without a xylopodium, glabrous, not dark-brown when dry. Branches lax, the upper ones cylindrical, wine-colored, winged, internodes 0.9–1.5  cm long. Leaves decussate, not imbricate, spreading, sessile; blades 15–32 × 17–25 mm, glaucous, coriaceous, not discolorous, acrodromous-basal-typical, ovate-lanceolate, base cordate to subcordate, sometimes amplexicaulous, apex obtuse, slightly acuminate, margin plane with wine-colored margin; secondary veins 3–4 on each side, departing at the base of the midvein, prominent on the abaxial surface; domatia absent. Synflorescence frondose, diplobotryum type, rarely simple botryum or pleiobotryum type; accessory branches absent, principal florescence 7–14  cm long, botryum type, paraclades 5–19  cm long; bracts 7–15 × 6–18 mm, ovate-­lanceolate, margins plane; flowers pedicellate, hypopodium 6–7(−10) mm long, epipodium 1–1.5  mm long, prophylls 6–9 × 4.5–9 mm, exceeding the apex of the floral tube, hiding the tube, ovate-lanceolate to suborbicular, unkeeled, base and apex obtuse, margin plane; floral tube 5.5–6 mm long, funnel-shaped, glabrous; sepals 4.5–5 mm long, wine-colored, triangular, not caudate, not deflexed, glabrous; epicalyx segments 3.5–4  mm long, wine-colored, linear-triangular, erect, slightly shorter than the sepals, glabrous; corolla 3–3.5  cm in diam., clear rose to rose-magenta, petals ca. 16  ×  14  mm,

370

13  Systematic Treatment

Fig. 13.98  Diplusodon rotundifolius. A. Branch with inflorescence. B. Leaf, abaxial surface. C, E. Flower with prophylls, without petals. D. Prophyll. (A–E Menezes et al. [CFCR] 10600, SPF)

Descriptions

371

obovate, apex retuse; stamens 8–12, free portion of the filaments 6.5–7 mm long, included; ovary ca. 3  ×  3  mm, rhombic, glabrous; style 15–16  mm long, ovules 14–17. Capsules 3–5 × 3–3.2 mm, globose, glabrous; seeds ca. 14. Distribution and Ecology (Fig.  13.99)  Diplusodon rotundifolius is restricted to Minas Gerais, in Cadeia do Espinhaço, and grows in campos rupestres at 923– 1207 m. Flowers from January to April; fruits from January to June. Specimens Examined: BRAZIL.  Minas Gerais,  Conselheiro Mata, estrada Conselheiro Mata-Diamantina, 1 km de Conselheiro Mata, 18°17′02″ S, 43°58′51″ W, 7 Feb 2015 (fl, fr), Cavalcanti et al. 3925 (CEN); Diamantina, Gardner 4571 (BM, NY, OXF); Diamantina, a 3 km N de São João da Chapada, 24 Mar 1970 (fl), Irwin et al. 28203 (K, MO, NY, P, UB); Diamantina, Riacho das Varas, próximo a Diamantina, 20 Mar 1893 (fl), Schwacke 8263 (OUPR, RB); Diamantina, 9 km na estrada para Biribiri, a partir da BR 367 (Diamantina-Araçuaí), 29 Jan 2000 (fl), Mello-Silva & Forzza 1748 (CEN, SPF); Diamantina, Rodovia Guinda-Conselheiro Mata, 17 Feb 1973 (fl), Hatschbach & Ahumada 31704 (F, MBM, MO, NY, W); Diamantina, Rodovia Guinda-Conselheiro Mata, 19 Mar 1997, Hatschbach et al. 66459 (M, MBM); Diamantina, a 32  km  N da estrada Diamantina-­Curvelo, na estrada para Conselheiro Mata, 14 Feb 1982 (fl), Graham 885 (MO, MBM, NY); Diamantina, estrada Conselheiro Mata-Diamantina, km 166, 23 Feb 1986 (fl), Cavalcanti et  al. (CFCR) 9529 (F, K, SPF, UEC); Diamantina, estrada para Conselheiro Mata, km 165, 25 Feb 1987 (fl), Zappi et al. (CFCR) 10381 (K, SPF); Diamantina, estrada de terra para Conselheiro Mata, km 166, 17 Apr 1987 (fl), Menezes et al. (CFCR) 10600 (K, SPF); Diamantina, BR 369-Conselheiro Mata, km 29, 15 Feb 1989 (fl), Cavalcanti et al. 217 (CEN, CESJ, F, HUEFS, IBGE, K, MBM, OUPR, SP, SPF); Diamantina, Riacho das Varas, 24 Mar 1892 (fl), Glaziou 19166 (K, LE n.v., P, R); Diamantina, margem da estrada para Conselheiro Mata, 18° 18′ 38′′S, 43° 52′ 47′′W, 15 Mar 2016 (fl), Faria-Jr. & Otoni 5472 (CEN, HDJF, UB); Conselheiro Mata, arredores, 14 Feb 1982 (fl, fr), Hatschbach 44725 (MBM); Conselheiro Mata, estrada Conselheiro Mata-Diamantina, a 500  m da cidade de Conselheiro Mata, 18°17′02″ S, 43°58′51″ W, 19 Jan 2003 (fr), Cavalcanti et al. 3122 (CEN, MO); Rio Pardo de Minas, 15°23′36″ S, 42°28′08″ W, 23 May 2005 (fr), Sevilha et al. 4328 (CEN); Santana do Riacho, Serra do Cipó, margem da rodovia entre a estátua do Juquinha e o povoado de Serra do Cipó, 19° 17′ 40′′S, 43° 35′ 45′′W, 30 Jun 2016 (fr), Faria-Jr. 6130 (CEN, HDJF, UB); Santo Antônio do Retiro, 15° 20′ 18′′S, 42° 45′ 09′′W, 15 Mar 2017 (fl, fr), Sevilha et al. 6371 (BHCB, CEN, NY, W). Diplusodon rotundifolius is easily recognized by its ovate-lanceolate, glaucous leaves with wine-colored margins and acrodromous-basal veins, and ovate-lanceolate prophylls that cover the floral tube. This species is similar to D. glaucescens, also from Cadeia do Espinhaço, but the prophylls of D. glaucescens do not hide the floral tube, and its leaves are orbicular-reniform. Lourteig (1989) indicated a specimen of Martius s.n. at M as holotype, what is considered as the lectotype (ICBN Art. 9.17, ex. 14).

372

13  Systematic Treatment

78. Diplusodon saxatilis Lourteig, Bradea 5(19): 218. 1989. Type: Brazil. Minas Gerais: Serro, rodovia MG-2, entre Serro e Datas, 19 Jan 1972 (fl), G. Hatschbach, L. B. Smith & E. S. Ayensu 28923 (holotype, P [01902026]; isotypes, C, MBM, MO, MU, P [01902027], US). Figs. 13.33 D–H, 13.99 Subshrubs ca. 0.4 m tall, erect, unbranched, sometimes branched just at inflorescence, with a xylopodium, not dark-brown when dry. Upper branches cylindrical, unwinged, hirsute, internodes 0.3–1 cm long. Leaves decussate, slightly imbricate, erect, sessile; blades 6–12 × 3–5 mm, coriaceous, not discolorous, hyphodromous, oblong, base cordate to retuse, apex obtuse, margin revolute, hirsute, blades covered by long, abundant trichomes; secondary veins not visible; domatia absent. Synflorescence frondose, diplobotryum type, congested; accessory branches absent, principal florescence truncate, paraclades 8–11 mm long; bracts similar to the vegetative leaves in shape and size; flowers sessile, concentrated at the apex of the paraclades; prophylls 5.5–6 × 3–4 mm, reaching the apex of the floral tube, elliptic to ovate, unkeeled, margins plane, base obtuse, hirsute; floral tube 5.5–6 mm long, funnel-shaped, hirsute; sepals 4.5–5 mm long, triangular, not caudate, not deflexed, hirsute; epicalyx segments 2.5–3 mm long, narrowly triangular, erect, hirsute, distinctly shorter than the sepals; corolla lilac, petals not seen; stamens 12, free portion of the filaments 8–8.5 mm long, included within the sepals; ovary ca. 1.8 × 2 mm, obconic, glabrous, style 15–16 mm long, ovules 10. Capsules not seen. Distribution and Ecology (Fig. 13.99)  Diplusodon saxatilis is known only from the type. The species grows in sandy fields in the municipality of Serro, Cadeia do Espinhaço, Minas Gerais. Flowers in January. Specimen Examined. BRAZIL.  The type. Diplusodon saxatilis is characterized by its dense indumentum of long trichomes that cover the entire plant except the mid to lower portion of the stem where there is an exfoliating bark. The leaves are oblong and retuse to cordate at the base, the epicalyx segments are shorter than the sepals (2.5–3 mm long), the prophylls are the same size as the floral tube, and the flowers have 12 stamens. 79. Diplusodon sessiliflorus Koehne, Fl. Bras. (Martius) 13(2): 333, pl. 62, fig. 1. 1877. Type: Brazil. Minas Gerais: G. Gardner 4573 (first-step lectotype, designated by Lourteig 1989: 241, second-step lectotype, P [01902025], here designated; isolectotypes, fragm. F, G [3], K [3], NY [2], P [01902024], S, SP, US, W [2]). Figs. 13.100 A–K, 13.104 Diplusodon crulsianus Koehne, Bot. Jahrb. Syst. 23(Beibl. 57): 33. 1897. Type: Brazil. Distrito Federal: entre Torto et le Rio Bananal, 23 May 1895, A.  F. M. Glaziou 21415 (lectotype, P [01902022] designated by Lourteig 1989: 241; isolectotypes, BR, G [3], K [2], LE, MO, NY, P [01902023], R [2], RB, S, US).

Descriptions

373

Fig. 13.99  Geographic distribution of Diplusodon rosmarinifolius, D. rotatus, D. rotundifolius, and D. saxatilis

374

13  Systematic Treatment

Fig. 13.100  Diplusodon sessiliflorus. A–B. Habits. C. Detail of leaf indumentum. D, K. Glandular trichome of sepals. E. Three-verticillate leaves. F. Floral bud. G. Prophyll, adaxial surface. H. Flower with prophylls, without petals. I. Detail of glandular trichomes of sepals. J. Unicellular trichomes of leaf. (A, C, D, J–K Cavalcanti et al. 1233, CEN; B, G–I Philcox & Onishi 4782, UB; E Ratter 2855, UB; F Cavalcanti et al. 527, CEN)

Descriptions

375

Subshrubs 0.3–1.5 m tall, erect, branched just at inflorescence, sometimes caespitose, with a xylopodium, not dark-brown when dry. Upper branches cylindrical, unwinged, villous, internodes 1–6 cm long. Leaves decussate, sometimes ternate, not imbricate, subsessile to petiolate; petioles 1–2  mm long; blades 20–60  ×  10–45  mm, suberect, coriaceous, not discolorous, acrodromous-basalsupranumerary, oval to ovate-lanceolate, very rarely orbicular, base cordate, apex acute to obtuse, generally acuminate to mucronate, margin plane, adaxial and abaxial surface covered by sparse, long white trichomes with a bulbous base; secondary veins 4–6 on each side, 2–3 of them departing from the base of the midvein, and 2–3 of them departing along the midvein; domatia absent. Synflorescence bracteose to frondose-bracteose, botryum spiciform to diplobotryum spiciform type, congested; accessory branches absent, principal florescence 6–20 cm long, paraclades 6–24 cm long; bracts 7–20 × 8–18 mm, orbicular to ovate, base cordate; flowers sessile; prophylls 4–9 × 5–10 mm, reaching the apex of the floral tube, orbicular, wide, keeled, base obtuse, apex obtuse, mucronate or totally flat, margin plane, floral tube 6–9 mm long, whitish, villous, the trichomes with a very enlarged base; sepals 4–6 mm long, triangular, not caudate, not deflexed, golden in color because of the color of the trichomes, covered with secretory trichomes bottle shaped; epicalyx segments 2.5–4 mm long, cylindrical, spreading to slightly deflexed, villous, shorter than the sepals; corolla 5–5.5 cm in diam., rose-magenta, petals 18–24 × 12–15 mm, obovate, apex obtuse; stamens 24–32, free portion of the filaments 7–8 mm long, generally included; ovary 2–2.5 × 3–3.5 mm, obconic, glabrous, style 14–17 mm long, ovules 40–46. Immature capsules ca. 5 × 3.5 mm, glabrous; seeds ca. 7. Distribution and Ecology (Fig. 13.104)  Diplusodon sessiliflorus occurs in Distrito Federal and the state of Goiás in the municipalities surrounding Distrito Federal. The report of this species for the state of Minas Gerais is supported only by the type material. It inhabits cerrado and rocky vegetation at 950–1200 m, and is often found occurring sympatrically with D. ramosissimus. Flowers from January to November; fruits registered from April to June. Specimens Examined: BRAZIL. Distrito Federal,  Brasília, ca. 10 km E of Brasília, 12 Sep 1965 (fl), Irwin et al. 8243 (UB); Brasília, 2 km NE do córrego Sítio Novo, 15°33′ S, 47°31′ W, 13 Jun 1983 (fl), Kirkbride 5329 (CEN); Brasília, a 32 km do Cenargen, 6 km na DF-170, 4 Jun 1992 (fl, fr), Dias & Vieira 164 (CEN, MO, NY); Brasília, APA Cafuringa, Sito Canaã, 16 Apr 1992 (fr), Melo & França 673 (UB); Brasília, APA da Cafuringa, Chapada da Contagem, 18 km N da torre de televisão de Brasília, 15°37′ S, 48°53′ W, 22 Jun 1982 (fl, fr), Croat 53630 (MO, UB); Margem da rodovia entre Santo Antônio do Descoberto e a estrutural, 15°55′19″ S, 48°14′29″ W, 11 Sep 2012 (fl), Faria & Zanatta 2771 (CEN, UB); Brasília, bacia do córrego Taquara, área do Cristo Redentor, 27 May 1991 (fl), Silva 1135 (IBGE); Brasília, bacia do rio São Bartolomeu, 28 Apr 1980 (fl), Heringer et  al. 4541 (IBGE); Brasília, bacia do rio São Bartolomeu, Heringer et  al. 4902 (IBGE, K, MO, P, US); Brasília, bacia do rio São Bartolomeu, 26 May 1980 (fl), Heringer et al 4859 (IBGE); Brasília, Barragem do Paranoá, 15°48′ S, 47°47′ W, 2 Jun 1992 (fl), Cavalcanti et al. 1233 (CEN, SPF); Brasília, Barragem do Paranoá,

376

13  Systematic Treatment

17 May 1983 (fl, fr), Thomé 81/02686 (UB); Brasília, ca. 18 km Sudoeste da torre de televisão, 26 Mar 1976 (fl), Ratter & Fonsêca 2831 (UB); Brasília, condomínio residencial Atenas, 15°57′51″ S, 47°49′09″ W, 4 Mar 2000 (fl), Pereira-Silva & Abdala 4251 (CEN); Brasília, córrego Fazendinha, 22 May 1984 (fl), Pereira-Neto 32 (UB); Brasília, DF-170, a 7  km de entroncamento com DF-001-Chapada da Contagem, 12 Jul 1990 (fl), Cavalcanti et al. 527 (CEN, NY, SPF); Brasília, Estação Biologica Experimental da Universidade de Brasília, 5 Nov 1980 fl), Barros 108 (UB); Brasília, Estação Ecológica do Jardim Botânico de Brasília, 27 Jun 1991 (fl), Ramos 397 (HEPH); Brasília, Estação Ecológica do Jardim Botânico de Brasília, 15°52′ S, 47°51′ W, 18 May 1999 (fl), Ramos et al. 1294 (HEPH); Brasília, Fazenda Água Limpa, 2 May 1986 (fl), Barros 1017 (SPF, UB); Brasília, Fazenda Água Limpa, 20 Jun 1986 (fl), Barros 1033 (UB); Brasília, Fazenda Água Limpa, 5 Jun 1986 (fl), Barros 1031 (SPF, UB); Brasília, Fazenda Água Limpa, 1 Apr 1976 (fl), Ratter & Fonsêca 2855 (K, UB); Brasília, Fazenda Água Limpa, 2 May 1986 (fl), Barros 1022 (SPF, UB); Brasília, Lago do Paranoá, entre a Universidade de Brasília e lago, 11 Apr 1968, Philcox & Onishi 4782 (K, NY, UB); Brasília, margem do Lago Paranoá, 12 Jun 1979 (fl), Heringer 17283 (HB, IBGE); Brasília, margem oeste do Lago Paranoá, 11 Mar 1966 (fl), Irwin et al. 13897 (NY); Brasília, Núcleo Bandeirante, Fazenda Sucupira, 15°52′00″ S, 48°00′00″ W, 18 Apr 2000 (fl), Guarino & Pereira 106 (CEN, MO); Brasília, Núcleo Bandeirante, Fazenda Sucupira, 3 May 2000 (fl), Guarino & Pereira 138 (CEN, MO); Brasília, Núcleo Bandeirante, Fazenda Sucupira, 15°55′ S, 48°01′ W, 19 May 1999 (fl), Faria et al. 156 (CEN, MO); Brasília, Núcleo Bandeirante, rodovia de acesso à Fazenda Sucupira, a 300  m do portão, 1 Apr 1998 (bot), Pereira-Silva & Sampaio 3884 (CEN, MO); Brasília, Núcleo Rural Lago Oeste estrada para o Poço Azul, ao lado do Parque Nacional de Brasília, Jun 2003 (fl), Pastore et al. 639 (CEN); Brasília, Parque Nacional de Brasília - DF, entrada pelo portão 6, 15 Jun 1999 (fl, fr), Faria et al. 212 (CEN, MO); Brasília, Parque Rasgado, 7 Jun 2003 (fl), Chacon & Pinho 54 (CEN); Brasília, Reserva Biológica do IBGE, 10 Jun 1983 (fl), Pereira 560 (IBGE); Brasília, Universidade de Brasília, 29–31 May 1973 (fl), Classe de Taxonomia da Universidade de Brasília 111 (UB 31696); Planaltina, Águas Emendadas, Jun 1993 (fl), Cavalcanti 1255 (CEN); Samambaia, 15°52′ S, 48°03′ W, 17 Jan 1996 (fl), Rezende 322 (CEN, MO); Sobradinho, 10 Apr 1974 (fl), Heringer 13228 (UB); Planaltina, Estação Experimental de Águas Emendadas, 27 Apr 1983 (fl), Alves 110 (HEPH); Planaltina, GO-118, km 5–10, próximo à Planaltina, 14 Oct 1990 (fl), Hatschbach & Silva 54503 (MBM); Planaltina, Reserva Biológica de Águas Emendadas, ca. 40 km a NE de Brasília, 28 Jun 1982 (fl), Maury 161 (HEPH); Rodovia DF-140, condomínio Parque do Mirante, 15°57′52″ S, 47°49′10″ W, 22 Jun 2000 (fl, fr), Pereira-Silva & Abdala 4274 (CEN); without reference of municipalto municipal district: 25 Feb 1965 (fl), Heringer 10441 (UB). Goiás, Campo Alegre, 17°24′30.9″ S, 47°46′58.1″ W, 7 May 2016 (fl), Cavalcanti et al. 3988 (CEN). Formosa, Fazenda Indaiá, 5 May 2003 (fl), Pastore & Suganuma 585 (CEN); Formosa, rio Tiquiri, 25 May 1967 (fl, fr), Heringer 11438 (NY, UB); Luziânia, Chácara Marajoara, rodovia Brasília-BH, km 17, lotes 8 e 9, Quadra 12F, 1 Apr 1984 (fl), Magalhães 9 (UB); Montes Claros de Goiás, Fazenda Funil,

Descriptions

377

15°53′26″ S, 51°53′04″ W, 29 May 2010 (fr), Zanatta et al. 518 (UB); without reference to municipal district: 18 Apr 1967 (fl), Heringer 11429 (UB); Rizzo & Barbosa 4927 (UFG). Diplusodon sessiliflorus is characterized by a floral tube with dense, whitish indumentum and golden sepals that are colored by bulbous-based trichomes that secrete an orange substance. The species resembles D. divaricatus, which also occurs in Goiás. In addition to the very similar habit, D. divaricatus has the same leaf shape and general floral aspect. On close examination, the indumentum of the leaves and branches differs between the two species: D. divaricatus has minute trichomes, and D. sessiliflorus has long, white trichomes that form a villous indumentum on the branches in addition to sparse trichomes on the leaves. Furthermore, the flowers of D. divaricatus have reduced prophylls (3–3.5 mm long) and inconspicuous epicalyx segments (to 0.5 mm long). Fifteen duplicates of Gardner 4573 are distributed in several herbaria. One of the duplicates at P was annotated by Koehne and was cited as the holotype by Lourteig (1989), which is here considered as the first-step lectotype (ICBN Art. 9.17, ex. 14). There are two specimens of Gardner 4573 at P, and the specimen P 01902025 is here chosen as the second-step lectotype. The type of Diplusodon crulsianus corresponds exactly to the morphological delimitation of D. sessiliflorus, leaving no doubt of its inclusion in synonymy with D. sessiliflorus, as done by Lourteig (1989). A photograph (F neg. nº 017931) of one sheet from the type collection, Glaziou 21415, at B shows that Koehne annotated this specimen as D. crulsianus, sp. nov. Among the duplicates studied, no others were annotated by Koehne. Lourteig (1989) cited a specimen of Glaziou 21415 at P as holotype, which is here considered as the lectotype. 80. Diplusodon sigillatus Lourteig (emend.) T.B.Cavalc., Bradea 5(19): 237, fig. 2. 1989. Type: Brazil. Goiás, Alto Paraíso de Goiás, Chapada dos Veadeiros, ca. 20 km de Alto Paraíso de Goiás, 9 Feb 1966 (fl), H. S. Irwin, J. W. Grear-Junior, R.  Souza & R.  R. Santos 12447 (holotype, P [01902021]; isotypes, BR, NY, UB, US, W). Figs. 13.101 A–J, 13.104 Subshrubs to shrubs 1–2 m tall, erect, much-branched, not dark-brown when dry. Upper branches subquadrangular to cylindrical, unwinged, dense pubescent, whitish, internodes 0.7–2 cm long. Leaves decussate, not imbricate, spreading, subsessile; petioles 1–2  mm long; blades 30–50  ×  15–25  mm, olive-green, coriaceous, sometimes discolorous, acrodromous-basal-supranumerary, ovate-lanceolate, base subcordate to cordate, apex acute, margin plane; blades covered by dense indumentum, trichomes very short, fine with bulbous base; secondary veins 3–5 on each side of the midvein, prominent on the abaxial surface; domatia absent. Synflorescence bracteose, diplobotryum spiciform to pleiobotryum spiciform type, congested;

378

13  Systematic Treatment

Fig. 13.101  Diplusodon sigillatus. A. Branch with inflorescence. B. Section of internode. C. Leaf, abaxial surface. D. Detail of leaf abaxial surface indumentum. E. Bract, abaxial surface. F. Simple trichome. G. Floral bud. H–I. Flower with prophyll, without petals. J. Ovary. (A–G Cavalcanti et al. 382, CEN)

Descriptions

379

accessory branches absent, principal florescence 3.5–10  cm long, botryum type, paraclades 4.5–15  cm long; bracts 6–25  ×  5–13  mm, orbicular to ovate, margin revolute; flowers sessile, hypopodium and epipodium absent; prophylls 4–5.5 × 3–4.5 mm, reaching or slightly less than the apex of the tube, obovate to elliptic, unkeeled, base acute, apex obtuse, margin plane, veins ca. 3, reddish, evident; floral tube 4–6 mm long, funnel-shaped, covered with short, thick trichomes; sepals 3.5–4  mm long, triangular, not caudate, not deflexed; epicalyx segments 4–4.5 mm long, cylindrical, suberect, longer than the sepals, sometimes the same size, with the same indumentum as the sepals; corolla 3.5 cm in diam., rose-magenta, petals 12–14 × 7–8.5 mm, obovate, apex obtuse; stamens 18, free portion of the filaments 7.5–8 mm long, included; ovary 1.5–2 × 2–2.5 mm, obovate-depressed, glabrous, style 10–13  mm long, ovules 14–34. Capsules ca. 3.5  ×  4  mm, globose, glabrous; seeds 2.9–3.1 × 1.5–1.7 mm. Distribution and Ecology (Fig. 13.104)  Diplusodon sigillatus was described from Chapada dos Veadeiros, Goiás, where several additional populations were subsequently found, and the species also occurs in Serra dos Pireneus, Goiás. It inhabits campos rupestres, campos sujos, and wet campos, at 1000–1800 m. Flowers from December to June; fruits from January to July. Specimens Examined: BRAZIL.  Goiás,  Alto Paraíso de Goiás, GO 118, 5–10  km  N de Alto Paraíso, 12 Feb 1990 (fl), Hatschbach & Nicolack 53933 (HUEFS, MBM); Alto Paraíso de Goiás, 2 km de Alto Paraíso na estrada para Nova Roma 1050, 20 May 1994 (fl), Proença et al. 1141 (UB); Alto Paraíso de Goiás, rodovia Alto Paraíso de Goiás-Teresina de Goiás, km 18, 7 Mar 1989 (fl), Cavalcanti et al. 382 (CEN, K, NY, SPF); Alto Paraíso de Goiás, 20 km N de Alto Paraíso de Goiás, 19 Mar 1971 (fl), Irwin et al. 32190 (K, NY, P); Alto Paraíso de Goiás, 14°04′ S, 47°30′ W, 5 Mar 1988, Ginzbarg & César 738 (UB); Alto Paraíso de Goiás, Chapada dos Veadeiros, rodovia GO-118, 10 km ao norte de Alto Paraíso de Goiás, 14°03′04″ S, 47°30′04″ W, 3 Apr 1997, Cavalcanti et al. 2198 (CEN, MO, NY); Alto Paraíso de Goiás, rodovia GO-118, a 10 km ao N de Alto Paraíso de Goiás, 14°06′09″ S, 47°28′47″ W, 3 Apr 1997 (fr), Cavalcanti et al. 2199 (CEN, MO); Alto Paraíso de Goiás, Chapada dos Veadeiros, GO-12, 20 Feb 1975 (fl), Hatschbach et al. 36357 (MBM); Alto Paraíso de Goiás, Chapada dos Veadeiros, 21 Dec 1968 (fl), Giulietti et al. 589 (Barroso et al. 589) (UB); Alto Paraíso de Goiás, Chapada dos Veadeiros, 2 Mar 1972 (fl, fr), Rizzo 7785 (UFG); Alto Paraíso de Goiás, Chapada dos Veadeiros, 6 Apr 1972 (fl, fr), Rizzo 7962 (UFG); Alto Paraíso de Goiás, Chapada dos Veadeiros, 4 May 1972 (fl), Rizzo 8061 (UFG); Alto Paraíso de Goiás, 16 km de Alto Paraíso, km 184 da rodovia GO-188, próximo ao rio Preto, 24 May 1994 (fl), Silva et al. 2005 (IBGE); Alto Paraíso de Goiás, povoado São Jorge, 20 Jun 2003 (fl), Pastore & Suganuma 642 (CEN); Alto Paraíso de Goiás, GO-239, ca. 8.5 km do trevo GO-118, sentido Nova Roma, e subindo ca. 30 m encosta sobre a estrada a direita, 14°  05′ 52′′S, 47°  28′ 18′′W, Mendoza et  al. 4237 (CEN); Cavalcante, ponte de pedra, Fazenda Renascer, 20 Apr 2003 (fr), Pastore et al. 472 (CEN); Pirenópolis, alto da serra, 15°47′51″ S, 48°49′45″ W, 14 Nov 2005 (st.), Cavalcanti et al. 3580 (CEN).

380

13  Systematic Treatment

Diplusodon sigillatus was described with leaves of 7–20  ×  5–11  mm, but all duplicates of the type material H. S. Irwin et al. 12447 have only parts of the synflorescence; it is therefore likely that the measurements of the leaves that appear in the protologue are in fact based on the bracts. The leaves of D. sigillatus are much larger (30–50 × 15–25 mm) and usually fall off at the time of flowering. Diplusodon sigillatus is mainly characterized by the ovate-lanceolate leaves with prominent veins on the abaxial surface and dense indumentum with very short and fine trichomes that have a thick base. The bracts are orbicular to ovate, sometimes discolorous, dark-green on the adaxial surface and olive-green on the abaxial surface. The flowers have 18 stamens, suberect epicalyx segments that are not too much longer than the sepals, and the prophylls reach the apex of the floral tube. The leaf shape and indumentum of Diplusodon sigillatus resemble that of D. helianthemifolius var. pemphoides. The latter taxon can be distinguished by its darkbrown color when dry and also by its shorter, deflexed to spreading epicalyx segments (1.5–2 mm long). Diplusodon sigillatus can be confused with D. paraisoensis, although the latter has larger flowers with expanded and foliaceous epicalyx segments that are longer than the sepals and the plant is usually strigose to densely sericeous in the leaves and flowers. In some cases these two species are difficult to separate. 81. Diplusodon sordidus Koehne, Bot. Jahrb. Syst. 23(Beibl. 57): 31. 1897. Type: Brazil. Goiás: Cachoeira da Vargem Grande, próximo ao rio Pizarão, 4 Jan 1895 (fl, fr), A. F. M. Glaziou 21426 (lectotype, P [01902018], designated by Lourteig 1989: 234; isolectotypes, A, BR, GH, K, R). Figs. 13.102 A–J, 13.104 Subshrubs 0.4–3 m tall, 15–20 cm diam., erect, branched, with a xylopodium, glabrous, dark-brown when dry. Upper branches quadrangular, winged, chestnut brown, internodes 1–2 cm long. Leaves decussate, not imbricate, spreading, sessile; blades 20–45 × 15–40 mm, coriaceous, not discolorous, acrodromous-basal type, ovate-lanceolate to ovate or to suborbicular, base cordate, apex obtuse, acuminate, margin plane and yellow in color with inconspicuous glands, adaxial and abaxial surface with dark punctae over the whole surface; secondary veins 3–4 on each side, yellow; domatia absent. Synflorescence frondose, botryum to diplobotryum type, lax; accessory branches absent, principal florescence 18–21 cm long, botryum type, paraclades 11–15 cm long; bracts 10–23 × 7–20 mm, ovate to suborbicular, margins plane; flowers pedicellate, hypopodium 1–1.5 mm long, epipodium 0.5–1 mm long; prophylls 10–12 × ca. 6 mm, surpassing the apex of the floral tube ca. 5 mm, lanceolate to ovate-lanceolate, unkeeled, base obtuse, apex acute, margins plane; floral tube 6–7 mm long, greenish to wine-colored, funnel-shaped, glabrous; sepals ca. 4 mm long., longer in fruit ca. 14 × 11 mm, triangular, not caudate, not deflexed; epicalyx segments 8–11 mm long, wine-colored, cylindrical to linear, spreading to suberect, surpassing the sepals, glabrous, longer than the sepals; corolla 5.5–6 cm in diam., rose-magenta to purple, petals 25–30  ×  11–17  mm, obovate, apex obtuse; stamens (26–)28, free portion of the filaments ca. 5 mm long, surpassing the sepals

Descriptions

381

Fig. 13.102  Diplusodon sordidus. A. Branch with inflorescence. B. Section of internode. C. Leaf, abaxial surface. D. Prophyll, abaxial surface. E. Floral bud. F. Flower with prophylls, without petals. G. Fruit. H. Capsule. I. Thickened septa of capsule. J. Seed. (A–B, Cavalcanti et al. 30, SPF; C–J Cavalcanti et al. 381, SPF)

382

13  Systematic Treatment

ca. 2 mm; ovary ca. 3 × 2 mm, rhombic, glabrous, style 20–23 mm long, ovules 33–38. Capsules 6.5–7 × 6–6.2 mm, globose, glabrous; seeds 17–46, 4–5 × 3–4 mm. 2n = 30 (Cavalcanti 1995; Graham and Cavalcanti 2001). Distribution and Ecology (Fig. 13.104)  Diplusodon sordidus is restricted to the campos rupestres and campos cerrados of Chapada dos Veadeiros, Goiás, at 1000– 1600 m. Flowers from October to April; fruits from August to April. Specimens Examined: BRAZIL. Goiás,  Água Fria de Goiás, cerca de 10 km em direção à Teresina de Goiás, 14°04′21″ S, 47°30′33″ W, 14 Oct 2000 (fl), Munhoz et al. 2120 (CEN, IBGE); Água Fria de Goiás, Fazenda Água Fria, cerca de 10 km em direção a Teresina de Goiás, 14°04′21″ S, 47°30′33″ W, 30 Nov 2000 (fl, fr), Munhoz et al. 2268 (CEN, IBGE); Água Fria de Goiás, Fazenda Água Fria, cerca de 10 km em direção a Teresina de Goiás, 14°04′21″ S, 47°30′33″ W, 14 Nov 2000 (fl), Munhoz et al. 2204 (CEN, IBGE); Alto Paraíso de Goiás, ca. 5 km de Alto Paraíso, entrada para sítio, 4 Mar 1988 (fl), Cavalcanti et al. 108 (CEN, SPF); Alto Paraíso de Goiás, ca. 7.5 km de Alto Paraíso, 30 Nov 1988 (fl), Cavalcanti et al. 30 (CEN, K, SPF); Alto Paraíso de Goiás, Parque Nacional da Chapada dos Veadeiros, estrada para Colinas a 34 km do entroncamento, 14°10′01″ S, 47°47′29″ W, 15 Aug 1990, Cavalcanti et al. 662 (CEN [2], K, MO, NY); Alto Paraíso de Goiás, entrada do Belvedere Paraíso. A cerca de 4  km  N de Alto Paraíso de Goiás, 14°08′00″ S, 47°27′00″ W, 14 Mar 1995 (fr), Cavalcanti et  al. 1339 (CEN); Alto Paraíso de Goiás, Chapada dos Veadeiros. Estrada Alto Paraíso-Colinas do Sul, a 9  km do entroncamento com a GO-118, 14°09′45″ S, 47°36′28″ W, 2 Apr 1997 (fl, fr), Cavalcanti et al. 2182 (CEN, MO); Alto Paraíso de Goiás, rodovia GO-118, a 10 km ao N de Alto Paraíso de Goiás, 14°03′04″ S, 47°30′04″ W, 3 Apr 1997 (fr), Cavalcanti et al. 2197 (CEN); Alto Paraíso de Goiás, entrada do Parque Nacional da Chapada dos Veadeiros (à esquerda), estrada Alto Paraiso-Colinas do Sul, 34 km da GO-118, 14°09′49″ S, 47°47′08″ W, 27 Jan 1997 (fr), Walter et al. 3647 (CEN, MO, NY); Alto Paraíso de Goiás, Jardim Japonês, 14°08′00″ S, 47°30′35″ W, 24 Jan 1997 (fr), Assis et al. 355 (CEN); Alto Paraíso de Goiás, GO-118, km 156.7, sentido BrasíliaAlto Paraíso, 15°52′52″ S, 47°50′54″ W, 11 Nov 1995 (fl, fr), Mello et al. 69 (CEN, MO); Alto Paraíso de Goiás, estrada Alto Paraíso de Goiás-Colinas do Sul, 10 km da GO-118, 14°09′45″ S, 47°36′48″ W, 29 Aug 2004 (fr), Cavalcanti et al. 3498 (CEN, SPF); Alto Paraíso de Goiás, estrada Colinas-GO-118, km 42 (GO-307), 14°10′ S 47°44′ W, 10 Mar 1989 (est.), Cavalcanti et  al. 403 (CEN, SPF); Alto Paraíso de Goiás, rodovia para Colinas, rio das Cabras, 2 Dec 1992 (fl), Hatschbach & Barbosa 58380 (MBM, MO); Alto Paraíso de Goiás, Parque Nacional da Chapada dos Veadeiros, 9 Feb 1987 (fl, fr), Toledo et  al. 294 (CEN, SP); Alto Paraíso de Goiás, Chapada dos Veadeiros, 14º S 47º W, ca. 12 km NW de Veadeiros, na estrada para Cavalcante, 21 Oct 1965 (fr), Irwin et al. 9398 (NY, UB n.v.); Alto Paraíso de Goiás, a 15 km W de Veadeiros, 12 Feb 1966 (fr), Irwin et al. 12636 (MO, NY, RB, UB); Alto Paraíso de Goiás, a 23  km de Alto Paraíso em direção a Teresina de Goiás, 13°56′39″ S, 47°29′38″ W, 12 Nov 1996 (fl, fr), Silva et al. 3203b (CEN, IBGE); Alto Paraíso de Goiás, 19 Dec 1967 (fl), Duarte 10765 (HB, SPF); Alto Paraíso de Goiás, GO-118, ca. 26 km N de Alto Paraíso de Goiás, 13°58′ S, 47°27′

Descriptions

383

W, 8 Feb 1987 (fr), Pirani et al. 1886 (CEN, K, SPF); Alto Paraíso de Goiás, a 6 km Leste de Alto Paraíso, 14°07′ S, 47°30′ W, 15 Feb 1979 (fl), Ferreira & Cardoso 37 (UB); Alto Paraíso de Goiás, a 5 km E de Alto Paraíso, 14°47′ W, 15 Feb 1979 (fl), Gates & Estabrook 191 (RB, SP); Alto Paraíso de Goiás, rodovia GO-118, Alto Paraíso-Terezina de Goiás, km 18, 13°59′ S 47°29′ W, 7 Mar 1989 (fr), Cavalcanti et al. 381 (CEN, SPF); Alto Paraíso de Goiás, 9 km S de Alto Paraíso de Goiás, na estrada Brasília-Alto Paraíso de Goiás, 21 Mar 1993 (fl, fr), Cavalcanti & Ayres 1263 (CEN); Alto Paraíso de Goiás, rodovia GO-118, 5–10 km N de Alto Paraíso, 13 Feb 1990 (fl), Hatschbach & Nicolack 53931 (MBM); Cavalcante, Fazenda Renascer, estrada para a ponte de pedra, 3 Feb 2004 (fr), Pastore et al. 787 (CEN); Teresina de Goiás, 13°53′45″ S, 47°21′33″ W, 29 Jan 2003 (fr), Cavalcanti et al. 3157 (CEN, MO). Diplusodon sordidus is characterized by showy flowers with a rose-magenta to purple corolla that is 5.5–6 cm in diam., in addition to ovate-lanceolate to ovate or suborbicular leaves ca. 4 cm long, long epicalyx segments (8–10 mm long), and an androecium with 26–28 stamens. Koehne (1897) described Diplusodon sordidus based on Glaziou 21426, a specimen that was destroyed at B (F neg. nº 017932), where Koehne conducted his research on Lythraceae. Lourteig (1989) chose a specimen of Glaziou 21426 from P and cited as holotype, which is considered as the lectotype (ICBN Art. 9.17, ex. 14). 82. Diplusodon speciosus (Kunth) DC., Prodr. 3: 94  f. 1828. Nesaea speciosa Kunth, Nov. Gen. Spec. Pl. 6: 191. 1823. Type: Brazil. [Mato Grosso], A. R. Ferreira s.n. (lectotype, P [01902017], designated by Lourteig 1989: 234; isolectotype, K). Figs. 13.103 A–B, 13.104 Diplusodon ovatus Mart. ex Koehne, Fl. Bras. (Mart.) 13(2): 319. 1877, pro syn. Type: Brazil. Mato Grosso prope Cuiabá, A.  L. P.  Silva-Manso 69 (Herb. Fl. Bras. n. 217) (BR, K, LE n.v., MO, P). Subshrubs 0.4–1 m tall, prostrate to erect, branched just at inflorescence, sometimes caespitose, with a xylopodium, glabrous, not dark-brown when dry. Upper branches cylindrical to subquadrangular, unwinged, internodes 3–8 cm long. Leaves decussate, spreading to suberect, sessile; blades 25–75 × 20–60 mm, slightly glaucous, coriaceous, not discolorous, acrodromous-basal-supranumerary, ovate or ovate-lanceolate to rarely orbicular, base subcordate to obtuse, apex obtuse or rarely acute, margins plane and yellow in color with inconspicuous glands, adaxial surface with tertiary veins evident; secondary veins 4–6 on each side, 3–4 of them departing from the base of the midvein; domatia absent. Synflorescence lax, frondose-bracteose, botryum to diplobotryum type; accessory branches absent, principal florescence 4–15  cm long, botryum type, paraclades 4–19  cm long; bracts 10–20  ×  15–20  mm, the same shape as the vegetative leaves, decreasing in size toward the apex of the branch; flowers pedicellate, hypopodium 1–1.5  mm long,

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Fig. 13.103  Diplusodon speciosus. A. Habit. B. Flower with prophyll, without petals. (A–B Cavalcanti et al. 86, SPF)

Descriptions

385

epipodium 0.5–1 mm long; prophylls 6–9 × 4–6 mm, reaching or slightly less than the apex of the floral tube, ovate to ovate-lanceolate, rarely oblong, unkeeled, base obtuse, apex obtuse, margins plane; floral tube 6–7  mm long, greenish, funnelshaped to oblong, elongated in fruit; sepals 3–5 mm long, rose, triangular, not caudate, not deflexed; epicalyx segments 5–6  mm long, filiform, dorsiventrally compressed to cylindrical, spreading to suberect, the same size as or longer than the sepals; corolla ca. 6  cm in diam., clear rose or rose-magenta to violet, petals 21–23 × 13–15 mm, elliptic, apex obtuse; stamens (26–)30–44, free portion of the filaments 8–9 mm long, slightly exserted; ovary 2–2 × 3–3.5 mm, ovate-depressed, glabrous, ovules 36–62. Capsules 4.8–5 × 4–5 mm, globose, glabrous; seeds 20–47, 2–2.5 × 1.5–2.5 mm. Distribution and Ecology (Fig.  13.104)  Diplusodon speciosus occurs in the southern part of the state of Tocantins and also in Mato Grosso, in cerrado and campo sujo vegetation at 427–960  m. In Mato Grosso, it was found growing on plinthosols. Flowers from November to March; fruits from November to June. Specimens Examined: BRAZIL.  Mato Grosso,  Água Boa, 13°54′ S, 52°05′ W, 16 Jun 1997 (fr), Cavalcanti et al. 2351 (CEN); Canarana, estrada Água BoaNova Xavantina, ca. 40 km após entrada de Canarana, 13°54′29.3″S, 52°04′39″ W, 23 Mar 1997 (fr), Souza et al.1940 (ESA); Cuiabá, Mar 1911 (fl, fr), Hoehne 4812 (R); Cuiabá, Chapada dos Guimarães, 80 m acima da Cachoeira Véu da Noiva, 27 Jan 1989 (fl, fr), Cavalcanti et al. 86 (CEN [3], K, MO, NY, SPF); Cuiabá, Geodésico, 28 Jan 1988 (fl, fr), Coelho & Meirelles s.n. (CESJ 23663); Cuiabá, Marco da Geodésia e Topografia do IBGE, 1 Mar 1983 (fr), Cunha et al. 722 (UFMT); Cuiabá, Marco da Geodésia e Topografia do IBGE, 11 Nov 1985 (fl, fr), Guarim-Neto 1140 (UFMT); Cuiabá, estrada da Chapada dos Guimarães, proximidades do Ribeirão da Mutuca, s.d., Erendorfer 2103 (UFMT); Cuiabá, rod. Cuiabá-­ Chapada dos Guimarães, km 27.7, entre o asfalto e o casarão, 15 Jan 1986 (fl, fr), Saddi 7230 (UFMT); Cuiabá, rodovia Cuiabá-Chapada dos Guimarães, imediações da Mata Fria, 2  km depois do Portão do Inferno, 1 Dec 1984 (fl), Saddi 4483 (UFMT); Cuiabá, Morro Podre, Chapada, Mar 1911 (fr), Hoehne 1025 (R); Paranapitinga, Bananalzinho, afluente de Paranatinga, Nov 1914 (fl), Kuhlmann 1076 (R); Várzea Grande, 10 km de Cuiabá, 2 Jan 1968 (fl), Saddi 36 (UFMT); Xavantina, aproximadamente 10 km S de Xavantina, 14°44′ S, 52°20′ W, 11 Jan 1977 (fl), Ratter et al. 4076 (K, MO, UB); without reference to municipal district: Ferreira 415 (K); Martius herb. Fl. Bras. n. 217 (BR, K, LE n.v., MO, P); Riedel 749 (K, LE n.v., P); Sellow 194 (P). Tocantins, Almas, Rodovia entre Almas e Natividade, estrada de terra com entrada em frente a primeira entrada para a cidade de Almas, 11°24′37″ S, 47°08′17″ W, 16 Jan 2008 (fl, fr), Pastore et al. 2460 (CEN); Arraias, Mar 1840 (fl), Gardner 3426 (K, W); Arraias, Gardner 3726 (K, W); Dianópolis, TO-040, entre Novo Jardim e Dianópolis, à 6 km de Dianópolis, 11°37′41″ S, 46°46′30″ W, 28 Jan 2003 (fl, fr), Cavalcanti et al. 3146 (CEN, MO); Dianópolis, estrada da divisa BA/ TO, para Dianóplois, Ca. 9 km da entrada da cidade de Dianópolis, 11°37′45″ S, 46°46′51″ W, 28 Jan 2014 (fl), Walter et al. 6671 (CEN, RB, UB); Dianópolis, rodovia Almas-Dianópolis, a 8 km antes de Dianópolis, 11°37′46″ S, 46°54′47″ W, 8

386

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Fig. 13.104  Geographic distribution of Diplusodon sessiliflorus, D. sigillatus, D. sordidus, and D. speciosus

Descriptions

387

Apr 1997 (fl), Cavalcanti et al. 2237 (CEN, MO); Dianópolis, estrada DianópolisTaguatinga do Tocantins, 4 km de Dianópolis, 8 Apr 1997 (fl, fr), Cavalcanti et al. 2243 (CEN, MO, NY); Goiatins, comunidade indígena Krahô, aldeia Serra Grande, 15 Apr 2001 (fl), Amaral-Santos & Nascimento 937 (CEN); Palmas, estrada vicinal partindo à direita da rodovia TO-010, sentido Lageado, próximo ao km 8, 10°07′12″ S, 48°17′54″ W, 14 Jan 1999 (fl), Árbocz 6513 (HTO); Natividade, 3 km antes de Natividade, estrada de acesso ao morro da antena da repetidora, 11°41′65″ S, 47°42′10″ W, 7 Apr 1997 (fr), Cavalcanti et  al. 2234 (CEN, MO [2], NY); Natividade, Dec 1841 (fl), Gardner 3154 (G, K, NY [2], OXF, P [3], SP); Pindorama, estrada para Pindorama, ca. 10 km do entroncamento com a BR-010. 26 Mar 2011 (fl), Bringel & Moreira 717 (UB); Taipas do Tocantins, estrada entre Dianópolis e Taipas, 12°04′07″ S, 46°54′58″ W, 15 Jan 2008, Pastore et al. 2444 (CEN); without reference to state and municipal district: Pohl 93 (P). Mature individuals of Diplusodon speciosus are totally glabrous and have sessile, ovate to ovate-lanceolate (or rarely orbicular) leaves. The seedlings sometimes have trichomes on the leaves. The flowers are large, up to 6 cm in diam., with ovate prophylls reaching the apex of the floral tube, and also with spreading, suberect epicalyx segments 4–6 cm long, and an androecium of 27–38 stamens. The species is similar vegetatively to D. ovatus, which also occurs in Goiás and Mato Grosso. Diplusodon ovatus differs from D. speciosus principally based on its inflorescences with lateral cymes, linear prophylls, and flowers with 18 stamens. Although the type material lacks the exact locality of collection, it is known that Alexandre Rodrigues Ferreira collected in Brazil from 1792 to 1798, in the states of Amazonas and Mato Grosso. For this reason, and because no species of Diplusodon occur in Amazonas, the type locality is probably in Mato Grosso. Lourteig (1989) chose a specimen of Ferreira s.n. at P and cited as holotype, which is considered as the first-step lectotype (ICBN Art. 9.17, ex. 14). 83. Diplusodon stellatus T.B.Cavalc., Phytotaxa 226(2): 147, fig. 3. 2015. Type: Brazil. Goiás: Parque Nacional da Chapada dos Veadeiros, estrada para Colinas, a 34  km do entroncamento da GO, 15 Aug 1990 (fl), T.B.Cavalcanti, G. Pereira-Silva, I. Cordeiro & L. P. Queiroz 684 (holotype, CEN [00021148]; isotypes, MO, NY, SPF). Figs. 13.105 A–I, 13.109 Subshrubs 0.4–1 m tall, erect, much-branched, usually caespitose, with a xylopodium, not dark-brown when dry. Upper branches cylindrical, lax, unwinged, covered by stellate trichomes forming tufts, internodes 0.8–3 cm long. Leaves decussate, not imbricate, suberect, petiolate; petioles 1.5–2  mm long; blades 15–35  ×  1.5– 4(−6) mm, coriaceous, slightly discolorous, eucamptodromous, narrowly lanceolate or narrowly elliptic to linear, base acute, apex acute, margin revolute, adaxial surface olive-green, covered by dense stellate trichomes forming tufts; abaxial surface canescent, covered by dense stellate trichomes in tufts; secondary veins 1–2 on each side along the midvein, prominent on the abaxial surface; domatia absent. Synflorescence frondose, diplobotryum spiciform to pleiobotryum spiciform type,

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Fig. 13.105  Diplusodon stellatus. A. Habit. B. Detail of vegetative branch. C. Section of internode. D. Leaf, adaxial surface. E. Detail of leaf adaxial surface indumentum. F. Leaf, abaxial surface. G. Detail of leaf adaxial surface indumentum. H. Flower without petals. I. Detail of floral tube indumentum. (A–I Cavalcanti et al. 684, CEN)

Descriptions

389

lax; accessory branches absent, principal florescence 3–5 mm long, botryum spiciform type, paraclades 5–17 mm long; bracts similar to the vegetative leaves, except diminishing in size toward the apex of the branch; 10–17 × 1.5–3 mm wide, linear, margin revolute; flowers subsessile, hypopodium ca. 0.5 mm long, epipodium ca. 0–0.3 mm long; prophylls 3.1–3.2 × 1–1.1 mm, slightly surpassing the middle of the floral tube, narrowly oblong, with strongly prominent midrib, base retuse, apex acute, margin plane; floral tube 4–4.5 mm long, canescent, funnel-shaped, covered by dense stellate trichomes forming tufts; sepals 1.5–1.6 mm long, rose, triangular, not deflexed, not caudate, not ciliate, covered by dense stellate trichomes forming tufts; epicalyx segments 3–4 mm long, rose, cylindrical, erect, exceeding the sepals 2–3  mm, covered by dense stellate trichomes forming tufts; corolla 3.5–4  cm in diam., rose-magenta to lilac, petals 21–22 × 6–7.5–10 mm, obovate, apex obtuse; stamens 12, free portion of the filaments ca. 5.5 mm long, rose, 2–2.1 mm exserted, surpassing the sepals ca. 1 mm; ovary ca. 1.8 × 1.9 mm, obovate-depressed, glabrous, style 10–11  mm long, rose, ovules 10. Capsules 2.5–2.8  ×  2.6–2.9  mm, oblong, glabrous; seeds ca. 5, 1.9–2 × 1.2–1.5 mm. Distribution and Ecology (Fig. 13.109)  Diplusodon stellatus occurs on sandstone soils of campos rupestres, campos sujos, campos limpos, and cerrado with rocky outcrops of Chapada dos Veadeiros, Goiás, at 880–1200 m. Flowers from April to August; fruits registered in January and from June to August. Specimens Examined: BRAZIL.  Goiás,  Alto Paraíso de Goiás, 15 May 1986 (fl), Toledo et al. 113 (IBGE); Alto Paraíso de Goiás, 14 Jun 1993 (fr), Hatschbach et al. 59523 (BR, MBM, MO); Alto Paraíso de Goiás, Parque Nacional da Chapada dos Veadeiros, estrada para Colinas, a 34 km do entroncamento da GO, 14°10′00″ S, 47°47′29″ W, 15 Aug 1990 (fr), Cavalcanti et al. 682 (CEN, SPF); Alto Paraíso de Goiás, povoado São Jorge, 20 May 2003 (fl), Pastore & Suganuma 651 (CEN); Alto Paraíso de Goiás, estrada Alto Paraíso de Goiás-São Jorge, próximo à casa do IBAMA, 29 Jan 2003 (fr), Cavalcanti et  al. 3162 (CEN, MO); Alto Paraíso de Goiás, estrada Alto Paraíso para São Jorge, 14°06′13″ S, 47°28′50″ W, 5 Apr 2014 (fl), Faria-Jr. et al. 3959 (CEN, UB n.v.); Alto Paraíso de Goiás, São Jorge, caminho para mirante, 11 Apr 2004 (fl), Rodrigues & Chesini 191 (CEN); Alto Paraíso de Goiás, estrada Alto Paraíso de Goiás-Colinas do Sul, a 1 km de São Jorge, 2 Jun 1999 (fr), Cavalcanti & Mello-Silva 2508 (CEN, MO, SPF); Alto Paraíso de Goiás, estrada Alto Paraíso-­Colinas do Sul, km 23, 14°7′49″ S, 47°41′45″ W, 29 Aug 2004, Cavalcanti et al. 3508 (CEN, SPF); Alto Paraíso de Goiás, Chapada dos Veadeiros, 28 Jul 1985 (fl, fr), Romaniuc-Neto & Sajo 380 (SP); Alto Paraíso de Goiás, Chapada dos Veadeiros, 14o8′ S, 47o31′ W, 19 May 1987 (fl), Menezes 1209 (K, SPF); Alto Paraíso de Goiás, Parque Nacional da Chapada dos Veadeiros, trilha a oeste da sede do Parque, 14o10′27″ S, 47o49′28″ W, 11 Mar 2014 (fl), Cavalcanti et  al. 3820 (CEN); Cavalcante, GO-327, Colinas do Sul-GO-118, 14o10′ S, 47o49′ W, 8 Mar 1989 (fl), Cavalcanti et al. 394 CEN, IBGE, K, NY, SP). Diplusodon stellatus is characterized by the dense tufts of stellate trichomes covering the entire plant, which confers a flaky aspect to the surface. It is characterized

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also by its narrowly lanceolate or narrowly elliptic to linear leaves with revolute margins, 12 stamens, and epicalyx segments 3–4 mm long. Diplusodon stellatus resembles D. rosmarinifolius in that both have stellate trichomes over the entire plant and both have similarly shaped leaves with revolute margins. Diplusodon rosmarinifolius differs because it has 15 stamens and spreading to deflexed epicalyx segments that are ca. 0.5 mm long. 84. Diplusodon strigosus Pohl, Flora 10: 151. 1827. Type: Brazil. Goiás: Ad Engento de S. Izidro et dos Bois versus Arnicuro, Cap. Goyaz in pascuis graminosis, Mar and Apr, J. B. E. Pohl 1365 = D 945 (first-­step lectotype, designated by Lourteig 1989: 227, second-step lectotype, W [0053436], here designated; isolectotypes, F, G, K, W [0053435, 0053437]). Figs. 13.106 A–H, 13.109, 15.4 D Subshrubs 0.1–1.5 m tall, erect, branched, often caespitose, with a xylopodium, not dark-brown when dry. Upper branches cylindrical to subquadrangular, sometimes with a narrow wing, lax, commonly associated with galls, villous, with bulbous-based trichomes, internodes 3–9  cm long. Leaves decussate, not imbricate, suberect, short-petiolate; petioles 2–3 mm long; blades 30–90 × 20–50 mm, coriaceous, not discolorous, acrodromous-basal-supranumerary, oblong, elliptic, ellipticlanceolate to ovate-lanceolate or orbicular, base obtuse to subcordate, apex obtuse to acute, generally acuminate, margin plane, adaxial and abaxial surface rough, with a verrucose aspect due to the bulbous base of the trichomes, trichomes sparse and equidistant; secondary veins 6–8 on each side, departing along the midvein, prominent; domatia present. Synflorescence bracteose to frondose, diplobotryum spiciform to botryum spiciform type, lax; accessory branches absent, principal florescence 6–28  mm long botryum spiciform type, paraclades 4–33  mm long; bracts of the first order 30–60 × 24–30 mm, of the second order 10–25 × 8–18 mm, orbicular to ovate, margins plane; flowers sessile to subsessile, hypopodium absent, epipodium 0–0.5 mm long; prophylls 6–9(−13) × 4.8–8(−10) mm, surpassing the floral tube, sometimes hiding the entire tube, ovate to orbicular, keeled, base obtuse, apex obtuse to acute, margin plane, glabrescent; floral tube 5.5–7 mm long, winecolored to greenish, funnel-shaped, covered by bulbous-based trichomes; sepals 4–5 mm long, triangular, fleshy, not caudate, not deflexed, ciliate, covered by bulbous-based trichomes, longitudinally sulcate; epicalyx segments 3.5–6  mm long, wine-colored at the base, triangular, erect, thick, covered by bulbous-based trichomes, segments the same size or slightly larger or sometimes shorter than the sepals; corolla 4–4.5 cm in diam., bright rose, petals 15–17 × 10–11 mm, obovate, apex obtuse to acute; stamens (15–)22–34, free portion of the filaments 7–9 mm long, included, ovary 2–3.5 × 3.5–4 mm, obconic, glabrous, style 11–17 mm long, ovules 36–84. Capsules 4.9–5  ×  4.5–5  mm, globose, glabrous; seeds 23–40, 1.8–2.8 × 1.5–2.1 mm. Distribution and Ecology (Fig. 13.109)  Diplusodon strigosus is found in distributed areas of cerrado in Distrito Federal and the states of Goiás, Mato Grosso, and Tocantins, at 400–1050 m. It also occurs in campos sujos with rocky soil and cam-

Descriptions

391

Fig. 13.106  Diplusodon strigosus. A. Habit. B. Detail of leaf abaxial surface indumentum. C. Floral bud. D–E. Prophylls. F. Flower without petals and prophylls. G. Detail of sepal indumentum, showing glandular trichomes. H. Simple trichome of leaves. (A–D, H Cavalcanti et al. 1086, CEN; F–G Pires et al. 9715, UB; E Burchell 7693, K)

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pos rupestres at 1000–1270 m on Serra dos Pireneus and Chapada dos Veadeiros, Goiás. In Niquelândia, Goiás, D. strigosus was found in campos sujos, campos brejosos, mesophytic forests on calcareous outcrops at 470 m and swampy areas (brejo). Flowers from January to November; fruits registered from March to October. Specimens Examined: BRAZIL.  Distrito Federal,  Brasília, bacia do rio Descoberto, 19 May 1999 (fl), Calago & Silva 142 (CEN, MO, NY); Brasília, Barragem do Paranoá, 15°48′ S, 47°47′ W, 17 May 1983 (fl, fr), Thomé 13 (UB). Goiás, Abadiânia, rodovia Anápolis-Abadiânia, entrada da Fazenda Papuazal, Fazenda Mato Comprido, 16°14′17″ S, 48°45′48″ W, 26 Mar 2002 (fl), Silva et al. 5264 (CEN, IBGE, UB); Alto Horizonte, Fazenda do Senhor Joaquim Braz, 19 Feb 2011 (fl), Faria-Jr. 1122 (CEN, UB); Alto Paraíso de Goiás, 14 Jun 1994, Walter & Bridgewater 2140 (CEN); Alto Paraíso de Goiás, Chapada dos Veadeiros, ca. 5 km SE de São Jorge, acesso para o Vale da Lua, 14°48′ S, 47°46′ W, 11 Jul 2005 (fl, fr), Rua 638 (CEN); Anápolis, 21 Jul 1952 (fl, fr), Macêdo 3586 (NY); Aparecida de Goiânia, 4 km de Aparecida de Goiânia, Chácara Santo Antonio, 5 Mar 1966 (fl), Rizzo 16 (UB); Aparecida de Goiânia, entorno da Chácara Jatobá, 28 Oct 2003 (fl, fr) Pastore 104 (CEN, MO); Aparecida de Goiânia, divisa com hidrolândia, próximo à BR-153, 26 Apr 2004 (fr), Pastore & Bringel 571 (CEN); Aparecida de Goiânia, BR-153, 10 km oeste de Anápolis, 22 May 1975 (fl, fr), Hatschbach 36677 (MBM, MO); Araguaína, próximo ao rio Lontra, Fazenda Baixa Mata, 7 Apr 1974 (fl), Rizzo 9782 (UFG); Barro Alto, 14°38 S, 49º W, 8 Feb 1998 (fl), Walter et al. 3165 (CEN, MO); Caiapônia, Serra do Caiapó, ca. 12 km S de Caiapônia, 30 Apr 1973 (fl), Anderson 9488 (UB); Caiapônia, Serra do Caiapó, Anderson 9498 (UB, NY, P); Caldas Novas, Parque Estadual da Serra de Caldas, 28 Mar 2009 (fr), Iglesias et al. 159 (CEN [3]); Caldas Novas, Parque Estadual da Serra de Caldas, 21 Mar 2008 (fl, fr), Junqueira et al. 129 (CEN); Cavalcante, estrada da balsa “Porto dos Paulistas” (rio Tocantins) para a HU, cerca de 5.8 km da balsa, 13°40′17″ S, 47°54′41″ W, 10 Nov 2000 (fl), Pereira-Silva et al. 4412 (CEN, NY); Cavalcante, ponte sobre o rio Tocantins-Cavalcante, km 61, 13°39′15.5″ S, 47°58′36.5″ W, 30 Aug 2004 (fr), Cavalcanti et al. 3517 (CEN, SPF); Cavalcante, ponte sobre o rio Tocantins-Cavalcante, km 78, 13°40′51.5″ S, 47°52′54.6″ W, 30 Aug 2004 (fl), Cavalcanti et al. 3519 (CEN, SPF); Cavalcante, Serra Branca, balsa do rio Tocantins para Serra Branca, km 3, 13°34′41″ S, 48°05′47″ W, 23 May 2001 (fl), Pereira-Silva et  al. 5086 (CEN, MO); Cavalcante, estrada Vila Veneno-Serra Branca, km 3.6, margem direita do rio Macacão, 13°34′25″ S, 48°04′20″ W, 20 Feb 2002 (fl), Pereira-Silva et  al. 5884 (CEN, MO); Cavalcante, Vila Veneno-rio São Félix, 13°32′10″ S, 48°03′25 W, 27 Jun 2001 (fr), Pereira-Silva et al. 5200 (CEN, MO); Ceres, rodovia Belém-Brasília, próximo a Ceres, 14°57′ S, 49°22′ W, Guimarães 10 (RB); Cocalzinho, estrada Cocalzinho-Pirenópolis, Serra dos Pireneus, próximo aos 3 Picos, 28 May 1998, Forzza et al. 905 (CEN, SPF); Cocalzinho de Goiás, estrada de cesso a Serra dos Pireneus, a cerca de 8 km de Cocalzinho, 28 May 1998 (fl, fr), Assis et al. 549 (CEN); Couto Magalhães, de Pequizeiro a Couto Magalhães, 30 km antes de Couto Magalhães, 15 May 1974, Rizzo 9828 (UFG); Corumbá de Goiás, Rizzo & Barbosa 1327 (UFG); Corumbá de Goiás, 16 May 1973 (fl), Anderson

Descriptions

393

10418 (NY n.v., UB); Corumbá de Goiás, 14 km S de Corumbá de Goiás, 30 Nov 1965 (fl), Irwin et al. 10760 (NY, P n.v.); Formosa, 10 km de Formosa, 29 Apr 1966, Irwin et  al. 15496 (UB); Formoso, Serra Grande, 14 Jan 1972 (fl), Rizzo 7445 (UFG); Formoso, 10 Feb 1972 (fl), Rizzo 7627 (UFG); Formoso, 13 May 1972 (fl, fr), Rizzo 8103 (UFG); Goiânia, 10  km de Goiânia, 6 Apr 1968 (fl, fr), Rizzo & Barbosa 66 (UFG); Goiânia, junto ao morro Santo Antônio, 7 Jun 1968 (fl, fr), Goiânia, à esquerda da rodovia Goiânia para Trindade, 10 Jun 1968, (fr), Rizzo & Barbosa 1443 (UFG); Goiânia, 29 Jan 1969 (fl), Rizzo & Barbosa 3499 (UFG); Goiânia, 5 Mar 1969 (fl), Rizzo & Barbosa 3923 (UFG); Goiânia, 21 Jan 1969 (fl), Rizzo & Barbosa 3589 (UFG); Goiânia, esquerda da estrada GO-1, de Goiânia para Leopoldo de Bulhões, 9  km de Goiânia, 1 Mar 1971 (fl), Rizzo 6985 (UFG); Goianira, 18 Apr 1970 (fl, fr), Rizzo 5012 & Barbosa 4261 (UFG); Jaraguá, Serra do Jaraguá, próximo à cidade de Jaraguá, 14 Apr 1984 (fr), Rizzo 10407 (UFG); Goiás, 29 Jan 1972, (fl), Rizzo 7516 & Barbosa 6246 (UFG); Mara-Rosa, Amarolândia, 14°05′33″ S, 49°16′44″ W, 25 Jun 1998 (fl), Fonseca et  al. 1956 (IBGE); Mimoso de Goiás, topo do morro, 14°52′46″ S, 48°14′ W, 9 Oct 2013 (fr), Cavalcanti & Pereira-Silva 3769 (CEN); Minaçu, a 7 km da estrada N do canteiro de obras da UHE, 13°28′ S, 48°23′ W, 10 Mar 1992 (fl), Cavalcanti et  al. 1086 (CEN [2], NY, SPF); Minaçu, próximo a área de empréstimo de argila ME 33 (Usina Hidrelétrica da Serra da Mesa), 13°28′ S, 48°23′ W, 10 Mar 1992 (fl), Cavalcanti et al. 1120 (CEN); Minaçu, estrada de terra Minaçu-Serra da Mesa, ca. 5 km da entrada N para o canteiro de obras da UHE, 20 Nov 1991 (fl), Walter et  al. 792 (CEN); Minaçu, estrada Minaçu/Serra da Mesa, cerca de 25 km de Minaçu, 13°49′ S, 48°26′ W, 21 Mar 1996 (fl), Walter et  al. 3290 (CEN, MO); Minaçu, estrada Minaçu-UHE Serra da Mesa. Antigo viveiro, a 5  km da UHE Serra da Mesa, 13°49′00″ S, 48°21′00″ W, 19 Jun 1995, Cavalcanti et  al. 1428 (CEN); Minaçu, estrada de acesso a área indígena, próximo a área de emprestimo. 500 m da barreira da UHE-Serra da Mesa, 13°47′ S, 48°17′ W, 20 Jun 1995 (fl), Cavalcanti et al. 1441 (CEN); Niquelândia, 11 km em direção a Uruaçu, 22 May 1990 (fl), Pedralli et al. 3297 (CEN, NY); Niquelândia, ponte sobre o rio Bagagem, distante 32.5  km da cidade de Niquelândia, estrada Niquelândia-Colinas do Sul, 14°22′ S, 48°12′ W, 12 Apr 1992 (fl), Walter et al. 1106 (CEN); Padre Bernardo, estrada Brazlândia-Padre Bernardo, 7 May 1991 (fl, fr), Vieira et al. 717 (CEN, MO, NY); Padre Bernardo, próximo à Jaboquinha, 15°33′00″ S, 48°15′00″ W, 9 Jul 1991 (fr), Vieira et al. 833 (CEN, MO, NY); Niquelândia, Fazenda Engenho, ca. 11  km Niquelândia-Dois Irmãos, local chamado de Lama Preta, 28 Jan 1997 (fl), Fonseca et al. 1491 (CEN, IBGE); Niquelândia, Fazenda Aranha, próximo à dolina, 14°15′43″ S, 48°42′58″ W, 23 May 1997 (fr), Cordovil-Silva et al. 618 (CEN, MO, NY); Niquelândia, estrada de terra vicinal GO-237 (Niquelândia/Colinas), entrada a 600 m da ponte sobre o Bagagem; a 2.5 km desta, 13 Apr 1992 (fl), 14°21′ S, 48°12″ W, Walter et al. 1236 (CEN, MO); Niquelândia, área de influência do AHE Serra da Mesa. Estrada de terra Niquelândia-Muquém, cerca de 3  km antes do Muquém, 14°32′17″ S, 48°09′21″ W, 3 Jun 1998 (fl, fr), 14°32′ S, 48°9″ W, Walter et al. 4194 (CEN, MO, NY); Niquelândia, Nossa Senhora da Abadia do Muquém, GO-237, entroncamento Muquém-Colinas do Sul, 5 km, Serra da Meia Noite, 14°26′18 S, 48°09′54″ W, 5

394

13  Systematic Treatment

Apr 2006 (fl, fr), Cavalcanti & Pereira-Silva 3739 (CEN); Niquelândia, Bom Princípio, ca. 8 km ao norte de Assunção de Goiás, lado direito da rodovia BR-414, 15°08′5″ S, 48°42′54″ W, 4 Apr 2006 (fl, fr), Cavalcanti & Pereira-­Silva 3726 (CEN); Pirenópolis, Cachoeira do Abade, 25 May 1968 (fl), Onishi et al. 69 (UB); Pirenópolis, Serra dos Pireneus, 4 Jun 1971 (fl, fr), Rizzo 6388 & Barbosa 5636 (UFG); Pirenópolis, Serra dos Pireneus, 9 Jul 1971 (fr), Rizzo 6517 & Barbosa 5166 (UFG); Pirenópolis, Serra dos Pireneus, 15°52′00″ S, 48°50′00″ W, 18 Mar 1989 (fl), Cavalcanti et  al. 473 (CEN, NY, SPF); Pirenópolis, estrada Cocalzinho-­ Pirenópolis, via Serra dos Pireneus, 8.1  km, 15°48′08″ S, 48°51′45″ W, 14 Nov 2005 (fr), Cavalcanti et  al. 3583 (CEN); Tupiratins, à esquerda da estrada para Tupiratins, a 6 km da cidade, 18 Mar 1972 (fl), Rizzo 7895 (UFG); Serra Dourada, ca. 16 km S de Goiás Velho, 11 May 1973 (fl), Anderson 10107 (NY [2], UB, P); without municipality: Entre Goiás e Cavalcante, Burchell 7693 (BR, K); entre Lage e Corumbá, 14 Jul 1895 (fl, fr), Glaziou 21413 (BR, K, LE n.v., MO, NY, P, R); entre Brasília e Niquelândia, 10 May 1963 (fl), Pires et al. 9715 (NY, P, UB); 6 Apr 1968 (fl, fr), Rizzo & Barbosa 123 (UFG); GO-118, a 15 km do trevo, em direção a São Gabriel, 15°28′26″ S, 47°30′38″ W, 28 May 2001 (fl, fr), Cavalcanti & PereiraSilva 2719 (CEN, MO). Mato Grosso, Água Boa, 14°24′41″ S, 52°13′41″ W, 15 Jun 1997 (fr), Cavalcanti et  al. 2347 (CEN, MO); BR-158, Água Boa-Ribeirão Cascalheira, 62 km de Água Boa, 13°46′28″ S, 51°56′26″ W, 26 Jun 1997, Cavalcanti et al. 2354 (CEN); Água Boa, Terra Indígena, Areões, 14°35′36.8″ S, 52°11′42″ W, 30 Jul 2002 (fr), lima et al. 597 (CEN); Barra do Garças, BR-158 para Xavantina, Vale do Sonho, 20 Jun 1988 (fl, fr), Saddi 9171 (HC); Barra do Garças, entre Barra do Garças e Campinápoles, 21 Feb 1997 (fl), Arbocz et al. 3292 (ESA); Canarana, Água Boa-Nova Xavantina, 13°23′15.8″ S, 51°56′52.4″ W, 23 Mar 1997, Souza et al. 15042 (CEN, ESA); 8 km NE da base do acampamento, próximo à estrada Xavantina-São Félix, 12º54′S, 51º52′W, 10 Apr 1968 (fl), Ratter et al. 965 (K, NY, P, UB); estrada entre a base do acampamento e a estrada principal, 26 Feb 1968 (fl), Philcox & Fereira 4361 (K, NY, P, RB, UB); Nova Xavantina, Campus Universitário UNEMAT, Estação Biológica, Mário Viana, trilha do Pequi, próximo ao cismógrafo, 24 Mar 2000 (fr), Viana 302R (CEN); Nova Xavantina, Parque Municipal do Bacaba, 29 Apr 2005 (fl), Santos et  al. 46 (CEN); Xavantina, ca. 86  km  N de Xavantina, Serra do Roncador, 3 Jun 1966 (fr), Irwin et al. 16524 (IAN, K, LE n.v., MO, NY, P, UB, W); Xavantina, 7 km SW de Xavantina, 14°44′ S, 52°20′ W, 3 Apr 1968 (fl), Ratter et  al. 829 (K, MO, NY, P); Xavantina, 1.5  km  S de Xavantina, 14°44′ S, 52°20′ W, 17 Aug 1967 (fr), Ratter & Ramos 374 (NY, P, UB); Xavantina, 15  km  S de Xavantina, 10 Jun 1966 (fr), Irwin et  al. 16862 (MO, NY, P, UB). Tocantins, Porto Nacional, de Porto Nacional para Ponte Alta do Norte, 20 km de Ponte Alta do Norte, 5 Apr 1974 (fl) Rizzo 9744 (UFG). Plants of Diplusodon strigosus commonly have galls that occur on the youngest branches. In addition, the branches, leaves, and especially the flowers are covered by long, bulbous-based trichomes that confer a verrucose aspect to the surfaces. The leaves are large, 3–9 cm long, the prophylls are broad and rounded, and the floral tube, calyx, and epicalyx are thick and fleshy. One interesting characteristic of the

Descriptions

395

species is the shape of the epicalyx segments, which are triangular like the sepals, so that the external calyx (epicalyx) and the internal calyx are equal in size and shape. In the field, some variation in habit correlated with the environment is noteworthy. The species can form small, delicate, caespitose plants up to ca. 15 cm tall, as seen in campos sujos and campos brejosos in northern Goiás. Larger, more robust subshrubs are seen at the same location but in areas where the cerrado vegetation becomes taller and more dense. Diplusodon strigosus was described by Pohl (1827a) without citation of specimens in the protologue. In another publication (Pohl 1827b), the author refers to a single collection of D. strigosus collected in March–April in Engenho de São Isidro e Engenho dos Bois, Goiás, without mentioning a collection number. Three specimens were examined at W.  The specimen bearing Pohl’s original manuscript label, annotated by Pohl, containing Pohl’s collecting number (Pohl 1365) and the reference number of his field diary (D = 945), is chosen here as lectotype (W0053436). Lourteig (1989) cited one specimen at W as holotype, considered as the first-step lectotype (ICBN Art. 9.17, ex. 14). Two specimens of Diplusodon strigosus were examined at G and K, whose labels indicate Pohl s.n. Since duplicates of Pohl’s collections were sent to other herbaria without the information from his field diary (see “Notes about Diplusodon nomenclature and typifications”) and most lack a number, the specimens at G and K are regarded as duplicates of Pohl 1365 = D 945. 85. Diplusodon subsericeus Casar. ex Koehne, Fl. Bras. (Martius) 13(2): 329. 1877. Type: Brazil. Minas Gerais: Cachoeira do Campo, Mar 1836 (fl), P. Claussen 113 [Casaretto herb. n. 2826 n.v.] (first-step lectotype, designated by Lourteig 1989: 223, second-step lectotype, P [01902015], [ex TO], here designated; isolectotypes, BR, F [2], fragm. F, K [3], NY, P [01902016], TO, W [C. F. P. Martius herb. Fl. Bras. n.869]; BR, K, M, NY). Figs. 13.107 A–G, 13.109 Subshrubs 0.4–1  m tall, erect, with a xylopodium, branched, young plants unbranched, slightly silvery, not dark-brown when dry. Upper branches subquadrangulate, not winged, hirsute to villous, white to silvery, internodes 1–1.8 cm long. Leaves decussate, not imbricate, suberect, petiolate; petioles 1–3 mm long; blades 10–24 × 5–12 mm, usually slightly silvery, coriaceous, not discolorous, acrodromous with supranumerary base to eucamptodromous, elliptic, rarely obtuse, base acute to slightly attenuate, rarely ovate, apex acute, sometimes slightly acuminate, margin plane, blades covered by a short-villous indumentum on both surfaces, slightly silvery; secondary veins 2–4 on each side, 1–2 of them departing from the base of the midvein, sulcate on the abaxial surface; domatia absent. Synflorescence frondose, diplobotryum to pleiobotryum type, congested; accessory branches absent, principal florescence truncate, paraclades 3–15 cm long; bracts similar to the vegetative leaves but smaller in size; flowers subsessile, hypopodium 0–1 mm long, epipodium 0–0.5  mm long; prophylls 5–6.5  ×  2.1–2.7  mm, reaching the

396

13  Systematic Treatment

Fig. 13.107  Diplusodon subsericeus. A. Branch with inflorescence. B. Section of internode. C. Leaf, abaxial surface. D. Detail of leaf abaxial surface indumentum. E. Prophyll, adaxial surface. F–G. Flower with prophylls, without petals. (A–G Cavalcanti & Pereira-Silva 3793, CEN)

Descriptions

397

middle to surpassing the apex of the floral tube, obovate to elliptic, unkeeled, base acute, apex acute to obtuse, margin plane, sericeous; floral tube 6–6.5  mm long, greenish when fresh, sometimes silvery, oblong, sericeous to villous; sepals 3–4.5 mm long, triangular, slightly caudate, not deflexed, sericeous; epicalyx segments 1–1.5 mm long, triangular to lanceolate, dorsiventrally compressed, suberect, much shorter than the sepals, with sericeous indument; corolla ca. 3 cm in diam., rose, petals 12–14 × ca. 8 mm, broadly obovate, apex obtuse; stamens 12, free portion of the filament 6.5–7 mm long, not exserted; ovary ca. 1.9–2.1 × 2.2–2.5 mm, depressed-obovate, glabrous, style 12–15 mm long, stigma capitate, ovules 26–36. Capsules ca. 4.2 × 3.9 mm, oblong, glabrous; seeds ca. 6, ca. 2.1 × 1.9 mm. Distribution and Ecology (Fig.  13.109)  Diplusodon subsericeus is presently known only from the municipality of Rio Acima, in Serra da Gandarela, Minas Gerais. It was collected from a population in cerrado with outcrops, at 1100  m. Flowers and fruits registered in March. Specimens Examined: BRAZIL. Minas Gerais,  Rio Acima, a 7 km do entroncamento de Rio Acima, 20°06′15″ S, 43°44′09″ W, 9 Mar 2014 (fl, fr), Cavalcanti & Pereira-Silva 3793 (B, BHCB, CEN, G, ESA, HUEFS, HUFU, HTO, K, LAM, MBM, M, MO, NY, OUPR, OXF, P, RB, SPF, UEC, UFMA, UFMT, US, W); Rio Acima, 20°06′15″ S, 43°44′09″ W, 7 Mar 2008 (fl), Garcia 3470 (CEN, PMSP); without reference to municipal district: Claussen s.n. (BR, OXF). Most individuals of Diplusodon subsericeus have silvery foliage due to its sericeous indumentum. Other diagnostic characteristics of the species are the elliptic to rarely ovate leaves with an acute to slightly attenuate base and 2–4 pairs of lateral veins. The flowers are subsessile with an oblong floral tube and short (1–1.5 mm long), suberect epicalyx segments that are much shorter than the sepals, and the androecium has 12 stamens. In flowers of individuals from a single population, variation was observed in the shape and length of prophylls in relation to the floral tube. Some flowers from younger, unbranched individuals had obovate prophylls reaching the middle of the floral tube, while flowers from mature, branched individuals had flowers with elliptical prophylls, reaching the apex of the floral tube or even surpassing it. This variation is unusual Diplusodon. Koehne (1877), in describing Diplusodon subsericeus, cited Casaretto 2826, Claussen 113, and Riedel s.n. The diary of Casaretto was consulted by Piero Delprete (pers. comm.), and it was found that when Casaretto traveled in Brazil, he did not collect in Minas Gerais. In Casaretto’s notes the information appears that during his stay in the country, he purchased four boxes, each with 100 specimens, from Claussen. This suggests that the materials cited by Koehne as Claussen 113 and Casaretto 2826 may be the same collection, the Casaretto number having been assigned on receipt of the specimen in Casaretto’s herbarium (TO). In the same way, Claussen 113 received the Herbarium of the Flora Braziliensis number 869 (Herb. Fl. Bras. n. 869), as seen in the notes of Martius and Koehne (1903). Duplicates were distributed to several herbaria in Europe. In Casaretto’s diary, there is a

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reference to number 2826 alongside the indication of D. subsericeus as a new species, which was named by Casaretto but never published, then later published by Koehne. I examined other duplicates of Casaretto herb. n. 2826 (except from TO), and only one has the manuscript note by Koehne of “Diplusodon subsericeus Casar,” along with his initials. The photographed specimen at B (F neg. nº 19,601) was also verified by Koehne before it was destroyed, and there was no indication that Koehne had further material of Casaretto herb. n. 2826. Lourteig (1989) chose Claussen 113, P ex TO, as the lectotype of Diplusodon subsericeus, cited as holotype, considered as the first-step lectotype (ICBN Art. 9.17, ex. 14). There are two specimens of Claussen 113 at P, and the one with the original label of Claussen was chosen as the second-step lectotype (P01902015). 86. Diplusodon tenuifolius T.B.Cavalc., sp. nov. Type: Brazil. Goiás: Campo Alegre, BR-050, Brasília-Catalão, margem esquerda do rio Imbiruçu, 17°24′30.9″ S, 47°46′58.1″ W, 7 May 2016 (fl), T.B.Cavalcanti, G.  PereiraSilva & G. L. Moreira 3989 (holotype, CEN [00101933]; isotypes, B, HUFU, K, M, MBM, MO, NY, P, RB, SPF, W). Figs. 13.108 A–D, 13.109, 15.4 E Diplusodon tenuifolius is characterized by the plant being a much-branched shrub with thin, erect branches and thin leaves. The plant is completely glabrous and presents flowers with reduced corolla of 1–1.5 cm in diameter, linear profiles that surpass the apex of the floral tube, pedicels of 2.5–3.5 mm long, and short epicalyx segments of 0.8–1 mm in length, reaching half the length of sepals. The closest species is Diplusodon canastrensis from Serra da Canastra, in Minas Gerais State, another glabrous and much-branched shrub with flowers of 12 stamens. Diplusodon canastrensis differs mainly by its narrowly oblong leaves, flowers with longer prophylls (4–5.5 mm long), and longer epicalyx segments (3–3.5 mm long), surpassing the apex of the sepals. Shrubs 0.4–1.5 m tall, erect, much-branched, not dark-brown when dry. Upper branches subquadrangular, unwinged, erect, slender, lax, glabrous, internodes 3–6  mm long on the upper branches, leaves caducous on the very old branches. Leaves decussate, not imbricate, erect, sessile; blades 0.6–13 × 0.5–0.9 mm, coriaceous, not discolorous, not glaucous, hyphodromous, linear, base obtuse, apex acute, minutely acuminate, margin plane, blades glabrous; secondary veins 0, the midvein inconspicuous on adaxial surface, visible on abaxial surface; domatia absent. Synflorescence frondose, pleiobotryum type, lax; accessory branches absent, principal florescences not seen, paraclades 4–25  cm long; bracts resembling the vegetative leaves; flowers pedicellate, hypopodium 2–2.5  mm long, epipodium 0.5–1 mm long, glabrous; prophylls 3.5–4 × 0.3–0.4 mm, reaching or slightly surpassing the apex of the floral tube, linear, base obtuse, apex acute, margin plane, glabrous; floral tube 3–3.5 mm long, winish, funnel-shaped, glabrous; sepals 1.5– 1.9 mm long, winish, triangular, not caudate, slightly mucronate, not deflexed, margin short-ciliate or glabrous; epicalyx segments 0.8–1  mm long, dorsiventrally

Descriptions

399

Fig. 13.108  Diplusodon tenuifolius. A. Branch with inflorescence. B. Leaf, adaxial surface. C. Prophyll, adaxial surface. D.  Flower with prophylls,  without petals. (A–C Cavalcanti et  al. 3989, CEN)

400

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compressed, erect, shorter than sepals, reaching half the length of sepals, glabrous, margin with sparse and short cilia or glabrous; apex acute; corolla 1–1.5  cm in diam., rose-magenta, petals 6.5–9 × 3.5–4.5 mm, oblong, apex obtuse; stamens 12, free portion of the filaments 5–6 mm long, not exserted; ovary 1.5–2 × 1.5–2.3 mm, elliptic, glabrous, style 6.5–8 mm long, winish, thick, stigma conspicuous, ovules 4–6. Fruits not seen. Distribution and Ecology (Fig.  13.109)  Diplusodon tenuifolius is known from areas of cerrado with rocky outcrops and open fields in southeast of Goiás, growing in rock crevices in white lithosol or clay-sandy soil with quartz, at 868–890 m elevation. Flowers registered from May to July and November; old fruits (without seeds) in November. Specimens Examined: BRAZIL.  Goiás,  Campo Alegre de Goiás, Serra do Facão, estrada em direção à fazenda Helena, região do córrego Imburuçu e ribeirão Ponte Alta, 12 Jul 2007 (fl), Arantes et  al. 38 (CEN, HUFU); Campo Alegre de Goiás, margens do rio Imbiruçu, BR-050, 17°24′29″ S, 47°46′35″ W, 15 Nov 2016 (fr), Pereira-Silva et al. 16863 (CEN); Campo Alegre de Goiás, rodovia GO-213, a 13,2 km a leste da BR-050, 17°32′58″ S, 47°41′58″ W, 16 Nov 2016 (fr), PereiraSilva et  al. 16867 (CEN); Campo Alegre de Goiás, GO-213, km 18 a leste da BR-050, 16 Nov 2016 (fl, fr), Pereira-Silva & Bringel 16875 (CEN, HTO, HUFU, K, NY, P, W); Campo Alegre de Goiás, ao longo da GFO-213, 13 km NE de Campo Alegre, 17°33′04″ S, 47°41′56″ W, 28 Jun 2017 (fl), Simon et al. 3080 (CEN, HTO, IBGE, RB, SP, UB). Diplusodon tenuifolius is a much-branched glabrous shrub with thin and erect branches and linear leaves of about 1 cm long, aspects which characterize the new species together with small flowers of 1–1.5 cm in diameter, linear prophylls surpassing the apex of the floral tube, pedicel of 2.5–3.5 mm long, and epicalyx segments of 0.8–1 mm long, shorter than sepals, reaching half the length of sepals. The closest related species is Diplusodon canastrensis from Serra da Canastra, in Minas Gerais State, by the small, narrowly oblong leaves and flowers with 12 stamens. The species differs from D. tenuifolius in that Diplusodon canastrensis presents longer prophylls (4–5.5 mm long) and floral tube (4.5–5 mm long) with longer epicalyx segments (3–3.5 mm long) surpassing the apex of the sepals. Besides these characteristics, the branches, margins of the leaves, prophylls, sepals, and floral tube in D. canastrensis bear minute, inconspicuous, and sparse trichomes. Diplusodon tenuifolius and D. canastrensis are geographically related species, since both occur in the Canastra Arc, an area of communication between the species of Diplusodon of the chain of mountains of Minas Gerais and of the chain of mountains of Goiás (Inglis and Cavalcanti 2018). Another morphologically similar species to Diplusodon tenuifolius is D. vacillans due to linear and glabrous leaves. Diplusodon vacillans, a species of north of Minas Gerais State, differs mainly by its peculiar plant architecture where the branches are pendulous and the flowers are facing down. In addition, the flowers in

Descriptions

401

Fig. 13.109  Geographic distribution of Diplusodon stellatus, D. strigosus, D. subsericeus, and D. tenuifolius

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13  Systematic Treatment

D. vacillans are larger (corolla of 2.5–3 mm in diam.), the prophylls surpass the floral tube, and epicalyx segments are 4–4.5 mm long. 87. Diplusodon thymifolius Mart. ex DC., Prodr. 3: 94b. 1828. Friedlandia thymifolia Mart. ex DC., Prodr. 3: 94b, 1828, pro syn. Type: Brazil. Minas Gerais: Diamantina, in dry fields of Serro, Jul, C. F. P. Martius s.n. (first-step lectotype, designated by Lourteig 1989: 224, second-step lectotype, M [0146429], here designated; isolectotypes, M [0243898, 0243899]); photos at F [neg. nº19602]. Figs. 13.110 A–F, 13.114, 15.5 A Shrubs 0.5–3  m tall, erect, much-branched, not dark-brown when dry. Upper branches cylindrical to subquadrangular, not winged, pubescent, covered by short trichomes, internodes 0.5–0.7  cm long. Leaves decussate, spreading, subsessile; petioles ca. 0.5  mm long; blades 4–11  ×  1–4  mm, coriaceous, not discolorous, eucamptodromous, oblong-lanceolate, narrowly elliptic, apex obtuse, margin subrevolute, base acute, blades covered by very short trichomes, denser and oriented on the veins; secondary veins 1 on each side, sulcate on the adaxial surface, prominent on the abaxial surface, sometimes surfaces sericeous; domatia absent. Synflorescence lax, frondose, diplobotryum to pleiobotryum type; accessory branches absent, principal florescence ca. 10  cm long, paraclades 7–30  cm long; bracts similar to the vegetative leaves in shape and size; flowers subsessile, hypopodium 0.8–1 mm, epipodium 0.1–0.3  mm; prophylls 2.5–2.6(−2.9)  ×  1–1.2  mm, oblong-lanceolate, obovate to elliptic, apex obtuse, base acute, reaching to the apex of the floral tube or slightly less, inserted in the apex of pedicel; floral tube 2–2.5  mm long, funnelshaped, covered by very short, inconspicuous trichomes; sepals 1.2–1.3(−2) mm long, triangular, not caudate, not deflexed, covered by very short, inconspicuous trichomes; epicalyx segments 1.1–3  mm long, cylindrical, spreading to suberect, with the same type of indumentum as the sepals; corolla 1.5–2 cm diam., lilac, rose to rarely white, petals 5–6.5 × 2.2–3 mm, oblong to obovate, apex obtuse; stamens 12, exserted to 1  mm, free portion of the filaments 4–5  mm long; ovary 1.2– 1.1(−1.8) × ca. 1.1(−1.9) mm, obovate, glabrous, style 9–10 mm long, ovules ca. 10. Capsules oblong, seeds ca. 6, seeds, 4–5 × 3.5–4 mm long, ca. 3 × 2.9–3 mm long. Distribution and Ecology (Fig.  13.114)  Diplusodon thymifolius is known from areas of sandy soil, in the northwest of Minas Gerais, at 720 m; in the west of Bahia State, in Espigão Mestre, in habitats of open tree cerrado and campo sujo of the cerrado at 510–750 m and also in caatinga in the municipality of Ibotirama at 830 m; and in the center of Bahia, in Chapada Diamantina, in campos rupestres at 880– 1260 m. To the north, the species grows in the southwest of Piauí State, in cerrado vegetation between 400 and 530 m, and in in the eastern portion of Tocantins, in Serra Geral, at 690–720 m. Flowers and fruits from November to July. Specimens Examined: BRAZIL. Bahia,  Abaíra, Bem Querer, Catolés de Cima, 13°17′ S, 41°53′ W, 23 Jun 1994 (fl), Ganev 3401 (CEN, HUEFS, SPF); Piatã, encosta do Morro do Santana, 9 Jun 1992 (fl), Ganev 448 (CEN, HUEFS); Baianópolis, rodovia BA-242, entre Barreiras e Baianópolis, 12°11′3″ S, 44°37′35″

Descriptions

403

Fig. 13.110  Diplusodon thymifolius. A. Branch with inflorescence. B. Leaf, adaxial surface. C. Leaf, abaxial surface. D. Detail of leaf adaxial surface indumentum. E. Prophyll, adaxial surface. F. Flower with prophylls, without petals. (A Martius s.n., M; B–F Scariot et al. 485, CEN)

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W, 15 Apr 1997 (fl), Cardoso et al. 30 (CEN, NY, UB); Barreiras, Espigão Mestre, ca. 25 km W de Barreiras, 3 Mar 1971 (fl), Irwin et al. 31343 (NY, P); Barreiras, 20 Mar 1981 (fl, fr), Silva 221 (ALCB, CEPEC); Barreiras, km 33 da estrada Roda Velha-Sítio Grande, 19 May 1984 (fl, fr), Silva & Viegas 349 (ALCB, CEPEC, HC, NY); Barreiras, estrada para o aeroporto, a ca. 15 km de Barreiras, 12°06′ S, 44°59′ W, 14 May 1997 (fl), Proença et al. 1757 (HUFU, SPF, UB); Barreiras, 30–40 km de Roda Velha, 2° Jun 1986 (fl), Hatschbach & Silva 50536 (MBM, P); Caetité, morro com torre de transmissão de TV, 14°04′ S, 42°30′ W, 25 May 1985 (fl), Noblick & Clodoaldo 3758 (CEN [2], HUEFS); Caetité, 6 km S de Caetité, caminho para Brejinho das Ametistas, 10°02′ S, 42°32′ W, 20 Nov 1992 (fl, fr), Arbo et al. 5640 (CTES, SPF); Caetité, Brejinho das Ametistas, estrada entre Brejinho das Ametistas e Caetité, 22 May 1999 (fl), Souza et al. 23096 (ESA); Caetité, ca. 5 km de Caetité em direção à Brejinho das Ametistas, 14°06′25″ S, 42°30′20″ W, 14 Jul 2002 (fl), Souza et al. 25955 (CEN, ESA); Caetité, BR 122, Brejinho das Ametistas, estrada para i lixão, 14°05′09″ S, 42°30′53″ W, 20 May 2004 (fl, fr), Pereira-Silva et al. 9066 (CEN); Caetité, ca. 3 km da cidade, estrada para Brejinho das Ametistas, 14°05′19″ S, 42°30′19″ W, 28 Apr 2003 (fl), Bothelho et  al. 8 (ALCB); Cocos, Fazenda Trijunção, área da sede Santa Luzia, 14°53′26″ S, 45°52′ W, 6 Jul 2001 (fl), Fonseca et al. 2857 (CEN, IBGE); Correntina, 10 km N de Correntina na estrada para Inhaúmas, 13º16′S, 44º39′W, 20 Apr 1980 (fl, fr), Harley 21916 (CEPEC, K, P); Correntina, estrada Correntina-Posse, 13°39′27″ S, 45°47′ W, 21 Nov 1999 (fl), Simon et  al. 273 (CEN, UB); Ibotirama, BR-242 (Ibotirama-Barreiras), km 86, 12°07′ S, 44°02′ W, 7 Jul 1983 (fr), Coradin et  al. 6644 (CEN, K, MO, P); São Desidério, 12°28′ S, 45°12′ W, 10 Apr 1989 (fl), Scariot et al. 485 (CEN, IBGE, P); São Desidério, caminho para Estivas, 12°21′ S, 44°58′ W, 30 Mar 2002 (fl), Loureiro et al. 568 (ALCB); São Desidério, assentamento Thainá, 12°36′ S, 44°46′ W, 2 Jul 2001 (fr), Alves et  al. 258 (ALCB); Érico Cardoso, Poção, estrada VeredaCurralzinho, 13°16′16″ S, 42°5′54″ W, 4 Jul 2001 (fr), Ribeiro 344 (HUEFS, RB); Rio de Contas, 16 km da cidade, na direção à Fazendola, 13°27′42″ S, 41°51′34″ W, 24 May 2001 (fl, fr), Harley & Giulietti 54422 (CEN, HUEFS); Rio de Contas, 20 km da cidade, na direção à Fazendola, 13°27′19″ S, 41°52′16″ W, 20 May 2001 (fl, fr), Harley & Giulietti 54410 (CEN, HUEFS); Rio de Contas, 13°29′46″ S, 41°51′38″ W, 22 Jan 2003 (fr), Cavalcanti et al. 3142 (CEN, MO); Vanderlei, ca. 45 km de Vanderlei ao longo da BR-242, s.d. (fl), Davidse & D’Arcy 12041 (SP). Minas Gerais, Formoso, Parque Nacional Grande Sertão Veredas, 15°9′ S, 45°47 W, 31 Jul 1989 (fr), Dias et al. 68 (CEN, IBGE, SPF, UB); Formoso, Parque Nacional Grande Sertão Veredas, 15°19′06″ S, 46°36″ W, 20 May 1998 (fl, fr), Fonseca et  al. 1876 (IBGE); Monte Azul, Parque Estadual Caminho das Gerais, 14°55′44″ S, 43°02′53″ W, 13 Jun 2009 (fl), Viana et  al. 4253 (BHCB). Piauí, Baixa Grande do Ribeiro, a 10.5 km da fazenda Cana Brava, direção W-E, limite Sul da Estação Ecológica Urucuí-Uma, 21 May 1982 (fl), Castro & Bernadete s.n. (HGB 2639); Gilbués, Serra Guariba, 20 May 1997 (fl), Félix 7725 (RB); Ribeiro Gonçalves, Estação Ecológica de Urucuí-Uma, 21 May 1981 (fl, fr), Freire s.n. (HGB 2038). Tocantins, Galheiros, Mangabeiras, chapada arenosa próximo a São Pedro, May 1840, Gardner 4139 (BR, G, K, NY, OXF, P).

Descriptions

405

Diplusodon thymifolius is characterized by small leaves (4–11 × 1–4 mm) with a subrevolute margin, covered by an abundance of very short trichomes that are primarily located on the veins. In some individuals (e.g., Fonseca et al. 2857, Gardner 4139), the leaves become slightly sericeous and silver due to the dense concentration of trichomes on the abaxial surface. The flowers are small with inconspicuous indumentum; short, spreading to suberect epicalyx segments (1–3 mm long); and 12 stamens. A group of specimens analyzed (e.g., Simon et al. 273, Scariot et al. 485) corresponds with the morphological characteristics of the type material (Martius s.n.), with delicate, lax upper branches (inflorescence), small leaves (4–6 mm long), and flowers with epicalyx segments that are 1–2  mm long and the same length as or slightly shorter than the sepals. From the same area of distribution, there is another group of specimens in which the upper branches are not lax and delicate, the leaves are longer (7–11 mm long), and the flowers are slightly larger with epicalyx segments that are 2–3 mm long and longer than the sepals. The latter morphotype led me to consider whether these plants may represent a different species. However, because Simon et al. 273 and Scariot et al. 485 also had some longer leaves as in the second morphotype, I have retained both forms within Diplusodon thymifolius. Vegetatively, Diplusodon thymifolius resembles D. epilobioides based on the narrow leaves and inconspicuous indumentum throughout the plant. Diplusodon epilobioides differs by its distinctive long pedicel (generally 4–7  mm long), its larger flowers with epicalyx segments that are erect and longer than the sepals, and also by its longer prophylls, which surpass the apex of floral tube. Koehne (1877) described two varieties from Goiás for specimens with slightly larger leaves than those described by Candolle (1828): var. β based on the collection Gardner 4136, with larger and thicker trichomes and cuneate-lanceolate or oblong prophylls, and var. γ, based on the collection Gardner 4139, with narrow prophylls and linear, strigose leaves that are univeined with one lateral inconspicuous vein. Later, Koehne (1903) cited also the collection Gardner 3723 for the variety β. Lourteig (1989) included the two varieties in the synonymy of Diplusodon thymifolius. I examined these specimens (Gardner 3723, 4136, 4139). Gardner 3723 is a poorly pressed sample of Diplusodon petiolatus (Koehne) T.B. Cavalcanti. Specimen Gardner 4136 has linear to narrowly elliptic leaves with revolute margins and epicalyx segments that are twice the length of sepals; this specimen instead belongs to D. macrodon. Gardner 4139, the type material of Diplusodon thymifolius var. γ, has narrow, small, nearly plane leaves covered by a sericeous indumentum that also occurs on the branches and flowers and belongs to the variation found within D. thymifolius. These varieties were never named. The type material of Diplusodon thymifolius, Martius s.n., is represented at M by three specimens, with the locality, “Serro Frio” (Serro, Minas Gerais). However, the species has never been collected in this region, but only in northwestern Minas Gerais, Bahia, and Piauí, which suggests the labeling could be erroneous. One of the three specimens of Martius s.n. at M was indicated by Lourteig (1989) as the holotype, which is here considered as the first-step lectotype (ICBN Art. 9.17, ex. 14). A

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specimen with a blue original label is here designated as the second-step lectotype (M0146429). 88. Diplusodon trigintus T.B.Cavalc., Novon 8(4): 337. 1998. Type: Brazil. Tocantins: Ponte Alta do Norte, de Porto Nacional para Ponte Alta do Norte, 20 km de Ponte Alta, 13 Jun 1974 (fl), J. A. Rizzo 9904 (holotype, UFG). Figs. 13.111 A–G, 13.114 Subshrubs, 0.8–1.3  m, erect, branched, glabrous, not dark-brown when dry. Upper branches cylindrical to subquadrangular, weakly winged, yellow to reddish, glabrous; internodes 2–2.4 cm long. Leaves decussate, not imbricate, erect, sessile; blades 20–30  ×  15–26  mm, glaucous, coriaceous, not discolorous, acrodromousbasal-supranumerary, ovate to ovate-lanceolate, base cordate, apex acute or acuminate, margin plane, glabrous, slightly reticulate; secondary veins (3–)4 on each side along the midvein, prominent; domatia absent. Synflorescence bracteose, diplobotryum to triplobotryum type, lax; accessory branches absent, principal florescence when present ca. 12 cm long, paraclades 5.5–18 mm long; bracts similar to the vegetative leaves in shape, base obtuse, smaller; flowers long-pedicellate, hypopodium 6–8.5  mm long, persistent, epipodium ca. 1  mm long; prophylls 1.5–3 × 1–2.5 mm, reaching from the base to the middle of floral tube glabrous, oblong to elliptic, unkeeled, margin plane, base retuse, apex obtuse; floral tube 4–5.5  mm long, campanulate, glabrous; sepals 2.5–3.5  mm long, triangular, not caudate, not deflexed; epicalyx segments 0.3–1 mm long, cylindrical to dorsiventrally compressed, spreading to slightly deflexed, shorter than the sepals; corolla 3.5–4 cm in diam., lilac, petals 14–15 × 9–10 mm, oboval, apex obtuse; stamens 24–32, free portion of the filaments 7.5–8 long, exserted, surpassing the sepals ca. 2  mm; ovary 2–2.2  ×  3–3.8  mm, rhombic-depressed, glabrous; style 12–15  mm long, ovules 16–20. Immature capsules ca. 4 × 5.5 mm, globose, glabrous; seeds 4–7, 2.1–2.5 × 2–2.2 mm. Distribution and Ecology (Fig.  13.114)  Diplusodon trigintus occurs in cerrado vegetation with sandy soil at ca. 400 m, in the state of Tocantins. Flowers from May to August; immature fruits in August. Specimen Examined: BRAZIL. Tocantins,  Almas, Fazenda Minnehaha, arredores do córrego Cachorro, ca. 70 km a nordeste da cidade de Almas, 11°06′55″ S, 47°07′46″ W, 10 Aug 2004 (fl, fr), Walter et  al. 5273 (CEN, IBGE); Miracema, estrada da UHE Lajeado, e o Ribeirão Mares, margem esquerda do rio Tocantins, 09°45′09″ S, 48°23″ W, 19 Jul 2000 (fl), Soares et al. 839 (HTO); Palmas, serra do Lajeado, altos do córrego Ubim e Cedro, 26 May 1994 (fl), Ramos & Barros 840 (HEPH, HTO). Diplusodon trigintus has ovate to ovate-lanceolate leaves, with prominent acrodromous venation and slightly conspicuous second- and third-order veins. The main characteristics are the long persistent pedicels, cordate leaf base, small prophylls, and 24–32 stamens. Diplusodon trigintus is similar to D. imbricatus, but the latter

Descriptions

407

Fig. 13.111  Diplusodon trigintus. A. Branch with inflorescence. B. Section of internode. C. Leaf, abaxial surface. D. Floral bud. E. Prophyll, adaxial surface. F. Flower with prophylls, without petals. G. Pistil. (A–G Rizzo 9904, UFG)

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has acrodromous-basal-supranumerary venation, flowers with shorter hypopodium, epicalyx segments absent or 0.5–1 mm long, and 12–18 stamens. 89. Diplusodon ulei Koehne, Bot. Jahrb. Syst. 41: 98. 1907. Type: Brazil. Bahia: Sincorá, Serra da Vendinha, Nov 1906 (fl, fr), E. Ule 7111 (lectotype, HBG n.v., designated by Lourteig 1989: 216; isolectotypes, P n.v., R). Figs. 13.112 A–G, 13.114 Subshrubs 0.50–3  m tall, erect, much-branched, sometimes dark-brown when dry. Upper branches quadrangular to subquadrangular, unwinged, yellow to winecolored, glabrous, internodes 0.8–2.0  cm long. Leaves decussate, not imbricate, suberect; petioles 3–7 mm long; blades 11–35 × 8–23 mm, nitid, chartaceous, petiolate, not discolorous, eucamptodromous, glabrous, orbicular, wide-elliptic to obovate, sometimes ovate, base acute, slightly attenuate, rarely obtuse, apex obtuse to acute or sometimes slightly acuminate, margin plane, winish, slightly thickened with protuberances, not ciliate, adaxial and abaxial surfaces glabrous, abaxial surface with not prominent veins, glabrous; secondary veins 3–5 on each side, along the midvein; domatia absent, when rarely present on the two basal vein pairs, inconspicuous. Synflorescence bracteose to frondose-bracteose, diplobotryum to triplobotryum type, lax; accessory branches absent, principal florescence when present ca. 4 cm long, generally caducous, botryum type, paraclades 4–30 cm long, winecolored; bracts 7–10 × 3–5 mm, similar to the vegetative leaves in shape, decreasing in size toward the apex of the branch; flowers pedicellate, hypopodium 2–2.5 mm long, epipodium 0.5–2 mm long; prophylls 3.6–4 × 1.5–1.8 mm, reaching the middle to the apex of the floral tube, elliptic or obovate, unkeeled, base acute, apex obtuse, margin plane, not ciliate, inserted in the upper third of the pedicel; floral tube 5–5.5 mm long, green to dark-wine, funnel-shaped, glabrous; sepals 3–3.5 mm long, reddish to dark-wine, triangular, caudate, apex deflexed and papillose, sometimes with 1–2 trichomes, not ciliate; epicalyx segments 4–5 mm long, dorsiventrally compressed with acute apex, spreading, longer than the sepals, not ciliate, sometimes with 1–2 trichomes on the apex; corolla 2.5–3 cm in diam., rose-, petals 9.5–12 × 5.5–8 mm, obovate-elliptic, apex obtuse; stamens 12, free portion of the filaments 7–8.5  mm long, exserted, surpassing the sepals 2–3  mm; ovary 1.8–2 × 2–2.2 mm, obovate, glabrous; style 11–12 mm long, rose, ovules 16–18. Capsules 4.5–4.8  ×  5–5.5  mm, globose, glabrous; seeds 4–9, 3–3.2  ×  ca. 2  mm. 2n = 30 (Graham and Cavalcanti 2001). Distribution and Ecology (Fig.  13.114)  Diplusodon ulei inhabits campos rupestres in the mountains of Chapada Diamantina (Bahia), at 1200–1700 m. Flowers from November to April; fruits registered from November to January, April and July. Specimens Examined: BRAZIL.  Bahia,  Abaíra, Salão, 9  km de Catolés na estrada para Inúbia, 13°15′ S, 41°53′ W, 28 Dec 1991 (fl), Harley et  al. H50520 (CEPEC n.v., HUEFS, K n.v., MO, NY, SPF); Abaíra, campo de Ouro Fino, 13°15′ S, 41°54′ W, 8 Jan 1992 (fl, fr), Hind & Queiróz H50025 (CEN, CEPEC n.v., HUEFS, K, NY, SPF); Abaíra, Tijuquinho, 13°16′ S, 41°54′ W, 26 Feb 1992 (fl),

Descriptions

409

Fig. 13.112  Diplusodon ulei. A. Branch with inflorescence. B–C. Leaves, abaxial surface. D. Detail of lateral cyme. E. Floral bud. F. Prophyll, adaxial surface. G. Flower with prophylls, without petals. (A, C–G Harley et al. 27845, SPF; B Harley H50666)

Sano & Laessoe H52343 (CEN, CEPEC n.v., HUEFS, K, NY, SPF); Abaíra, caminho Serrinha-Capão de Levi, 13°19′ S, 41°51′ W, 22 Nov 1992 (fl), Ganev 1154 (HUEFS, K, SPF); Barra da Estiva, ca. 10 km N of Barra da Estiva on Ibicoara road, by the rio Preto, 13°35′ S, 41°38′ W, 2 Feb 1974 (fl), Harley 15829 (CEN, K, RB); Boninal, Mar 1980 (fl), Pinto s.n. (ALCB (9232); Catolés, 13°17′ S, 41°51′ W, 20 Dec 1991 (fl, fr), Harley et  al. H50146 (CEN, CEPEC n.v., HUEFS, K, NY, SPF); Catolés, Lagoinha, 13°19′ S, 41°48″ W, 16 Dec 2003 (fl), Oliveira et al. 260 (CEN, HUEFS); Catolés, na saída da cidade em direção à Catolés de Cima, 10 Jan

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1999 (fl), Cavalcanti et  al. 2475 (CEN); Catolés de Cima, enconsta da serra do Barbado, 13°18′ S, 41°50′ W, 10 Jul 1995, Queiróz et  al. 4398 (CEN, HUEFS); Catolés, na subida para a forquilha da serra, 8 Jan 1999 (fl), Cavalcanti et al. 2452 (CEN); Catolés, vale frente à Serra do Guarda Mor, 13°17′ S, 41°52′ W, 27 Dec 1988 (fl), Harley et al. 27845 (CEN, SPF); Ibicoara, Chapada Diamantina, 7 Mar 2011 (fl), Ogasawara 95 (ALBC); Jussiape, estrada de terra Jussiape-Capão de Volta, a 18 km de Jussiape, 13°26′42″ S, 41°30′38″ W, 25 Jan 2003 (fl), Cavalcanti et al. 3144 (CEN, MO); Vassouras, caminho para Tapera, 13°17′ S, 41°51′ W, 22 Apr 1994, Ganev 3095 (HUEFS); Mucugê, 13°0′ S, 41°23′ W, Ferreira et al. 78 (CEN, HUEFS); Mucugê, caminho para Guiné, 12°56′23″ S, 41°28′31″ W, 15 Feb 1997 (fl), Atkins et al. 5680 (CEN, ALBC); Mucugê, 6–12 km toward Serra do Pina, NW of Mucugê, 23 Jul 1985 (fr), Kral et al. 72917 (CEN, SP); Mucugê, subida do Beco, 12°45′ S, 41°30′ W, 17 Apr 2005 (fl, fr), Conceição & Cardoso 1324 (CEN, HUEFS); Piatã, encima do morro cuja estrada sai do final da cidade, 9 Jan 1999 (fl), Cavalcanti et al. 2456 (CEN); Piatã, estrada para Unúbia, ca. 31 km, 15 Feb 1987, Harley et al. 24307 (CEN, K, SPF); Piatã, estrada Piatã-Abaíra, 4 km após Piatã, 13°09′ S, 41°47′ W, 7 Jan 1992 (fl, fr), Harley et al. H50666 (CEN, CEPEC n.v., HUEFS, RB, SPF); Piatã, quebrada da Serra do Atalho, 13°13′ S, 41°50′ W, 26 Dec 1991 (fl), Harley et al. H50391 (CEN, CEPEC n.v., HUEFS, K, NY, SPF); Rio de Contas, 1 km antes do distrito de Mato Grosso, 20 Dec 1997 (fl, fr), Carvalho et al. 6432 (CEN, CEPEC); Rio de Contas, Pico das Almas, vertente leste, trilha da Fazenda Silvina-Queirós, 13°32′ S, 41°36′ W, 12 Nov 1988 (fl), Harley et al. 26124 (CEN, K, MO, NY, RB, SPF); Rio de Contas, Pico das Almas, vertente leste, trilha da Fazenda Silvina-­Queirós, 13°33′ S, 41°57′ W, 17 Feb 1977 (fl), Harley 19512 (K, SPF); Rio de Contas, Pico das Almas, vertente leste, trilha da Fazenda SilvinaQueirós, 13°32′ S, 41°57′ W, 29 Nov 1988, Harley et al. 26682 (CEN, K, MO, NY, RB, SPF); Rio de Contas, Pico das Almas, vertente leste, trilha da Fazenda SilvinaQueirós, 13°32′ S, 41°56′ W, 27 Nov 1988, Harley et al. 27006 (CEN[2], K, MO, NY, SPF); Rio de Contas, Pico das Almas, vertente leste, trilha da Fazenda SilvinaQueirós, 10 Apr 1999 (fr), Forzza et al. 1149 (CEPEC n.v., NY, SPF); Rio de Contas, a 7 km da cidade para o vilarejo Bananal, 13°31′38″ S, 41°51′13″ W, 5 Mar 1994, Roque et  al. (CFCR) 14789 (SPF); without reference to municipal district: Serra Narsalina, 1913, Lutzelburg 78 (M); Serra Narsalina, Minas de Contas, 1914, Lutzelburg 122 (M). Diplusodon ulei is characterized by its glabrous habit, orbicular to broadly elliptic or obovate leaves, with an acute base, a slightly thickened margin, and inconspicuous veins on the adaxial surface. The flowers have a wine-colored or dark-wine floral tube, spreading epicalyx segments 4–5 mm long, and 12 stamens. The holotype (Ule 7111) deposited at B and annotated by Koehne as “n. sp.” was destroyed (F neg. nº 017934). There are three duplicates of the type material in European and Brazilian herbaria. Lourteig (1989) cited a specimen from HGB as the holotype because it was seen by Koehne, which is here interpreted as the lectotype (ICBN Art. 9.17, ex. 14). 90. Diplusodon uninervius Koehne, Fl. Bras. (Martius) 13(2): 340, pl. 63, fig. 5. 1877. Type: Brazil. Minas Gerais: Serra da Lapa, L. Riedel 909 (first-step lec-

Descriptions

411

Fig. 13.113  Diplusodon uninervius. A. Branch with inflorescence. B. Section of internode. C. Leaf, abaxial surface. D. Prophyll, abaxial surface. E. Floral bud. F. Flower with prophylls, without petals. (A–F Cavalcanti et al. [CFCR] 10185, SPF)

412

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totype, designated by Lourteig 1989: 217, second-­step lectotype, LE, here designated; isolectotypes, G, K; photos at CEN, F, SPF). Figs. 13.113 A–F, 13.114 Diplusodon candollii A.St.-Hil., nom. illeg., Fl. Bras. Merid. (A.St.-Hil.) 3: 146, pl. 188. 1833, non Pohl ex DC. Type: Brazil. Minas Gerais: In campis prope praedium Bom Fim et pagum Nossa Senhora da Penha, Minas Novas, A.  SaintHilaire s.n. (catal. B1 nº 1174) (lectotype, P [01902008], designated by Lourteig 1989: 217); isolectotypes, P [01902009, 05092622, 05092626]. Shrubs 0.5–4  m tall, erect, much-branched, dark-brown when dry. Upper branches subquadrangular, unwinged, pubescent, internodes 0.3–0.5  cm long. Leaves decussate to 3–5-whorled, subsessile; petioles to 1  mm long; blades 4–12  ×  0.7–1  mm, suberect, coriaceous, not discolorous, hyphodromous, linear, base acute, apex obtuse, margin revolute, blade glabrous, dense indumentum on the midvein on abaxial surface; lateral veins absent, with dense indumentum. Synflorescence congested, frondose, diplobotryum to pleiobotryum type; accessory branches absent, paraclades 3–20 cm long; bracts similar to the vegetative leaves in shape and size; flowers pedicellate, hypopodium 1–1.5  mm long, epipodium 0.5–1 mm long; prophylls 3–4 × 1.8–2 mm, lanceolate to linear-lanceolate, unkeeled, with conspicuous cilia, margin subrevolute, reaching the apex of the floral tube; floral tube 2.5–3 mm long, funnel-shaped, glabrous; sepals 2–2.5 mm long, triangular, not caudate, not deflexed; epicalyx segments 1.8–2 mm long, cylindrical, suberect to spreading, with conspicuous cilia, shorter than the sepals; corolla ca. 2 mm in diam., rose-magenta, petals 8–11 × 5–7 mm; stamens 12, exserted, free portion of the filaments 7–8  mm long; ovary ca. 1.5  ×  2  mm, obovate-depressed, glabrous, style 12–14 mm long, ovules 15–18. Capsules 4–5 × ca. 4.5 mm, globose; seeds 5–10, 2–4 × 2–3.5 mm. Distribution and Ecology (Fig.  13.114)  Diplusodon uninervius occurs in the municipalities of Serra do Cipó, Diamantina, Datas, Gouveia, and Rio Vermelho in the vicinity of Cadeia do Espinhaço, Minas Gerais. It inhabits campos rupestres on sandy soil and sandstone at 1000–1400 m. Flowers from September to May; fruits from September to July. Specimens Examined: BRAZIL. Minas Gerais,  Buenópolis, Curimatai, arredores de Buenópolis, 19 May 2001 (fl, fr), Hatschbach et al. 72217 (MBM); Cunha Magalhães, rio Jequitinhonha, 16 Nov 1971, Hatschbach & Pelanda 28030 (MBM, NY, P); Serra do Cipó, 26 Apr 1992 (fr), Schwacke 8264 p.p. (RB); Datas, rodovia BR-367, entroncamento Datas-Diamantina, km 1.5, 18°22′ S, 43°41′ W, 1 Feb 1989 (fr), Cavalcanti et al. 189 (CEN [2], UFG); Datas, estrada Datas-Serro, ca. 2 km de Datas, 24 Nov 1985 (fl), Cavalcanti et  al. (CFCR) 8702 (SPF); Diamantina, Diamantina-Biribiri, a 14  km de Diamantina, 31 Oct 1981 (fl), Giulietti et  al. (CFCR) 2481 (K, SPF); Diamantina, Biribiri, 25 Nov 1998 (fr), Nogueira et al. s.n. (CEN 46566, BHCB 48592); Diamantina, Biribiri, 18°10′ S, 43°37′ W, 8 Apr 1982

Descriptions

413

Fig. 13.114  Geographic distribution of Diplusodon thymifolius, D. trigintus, D. ulei, and D. uninervius

414

13  Systematic Treatment

(fl, fr), Hensold et  al. 3098 (MBM, NY); Diamantina, Biribiri, 15 Oct 1984 (fl), Isejima et al. (CFCR) 5537 (CEN, K, SPF); Diamantina, Biribiri, 5 Jun 1985 (fr), Cruz 78 (CEN, SP); Diamantina, Biribiri, 22 Nov 1985, (fl), Cavalcanti et  al. (CFCR) 8597 (K, NY, SPF); Diamantina, a ca. de 35 km SW de Gouveia, km 243 na MG-259, 2 Feb 1972 (fr), Anderson et al. 35140 (NY, P); Diamantina, Aug 1949, Vidal s.n. (P); Diamantina, km 14 após Diamantina, 27 Oct 1984 (fl) Brandão 10883 (PAMIG); Diamantina, 22 Nov 1964 (fl), Duarte 8511 & Pereira 9281 (P); Diamantina, 30 km NE de Diamantina, 1 km do rio Jequitinhonha, 26 Nov 1985 (fl, fr), Graham 956 (MO, NY); Diamantina, na estrada para Cristais, ao longo do rio Biribiri, 28 Nov 1985 (fl, fr), Graham 980 (MO, NY); Diamantina, estrada para Biribi, 8 Dec 1997 (fl, fr), Lima et al. 3 (CEN, SPF); Diamantina, rodovia GuindaConselheiro Mata, 16 Mar 1987, Hatschbach et al. 50966 (MBM, P); Diamantina, estrada Diamantina a Corinto, até 20 km, 1 Dec 1976 (fl, fr), Shepherd et al. 3930 (NY, RB, UEC); Diamantina, subida para o Cruzeiro, 14 Jan 1963 (fl), Duarte & Barroso 7878 (HB, NY); Diamantina, estrada para Conselheiro Mata, a 5 km da estrada Curvelo-Diamantina, 18 Dec 1985, Simão-Bianchini et al. (CFCR) 8784 (F, SPF); Diamantina, entroncamento da rodovia Datas-Serro, BR-259, com Costa Sena, cerca de 1 km, 18°33′54″ S, 43°38′40″, 19 Jan 2003 (fl, fr), Cavalcanti et al. 3126 (CEN, MO); Diamantina, estrada para Extração, 16 Mar 1970 (fl), Irwin et al. 27672 (K, MO, NY, P); Diamantina, estrada para Extração, 28 Jan 1986 (fl, fr), Zappi et al. (CFCR) 9294 (SPF); Formigas, próximo à Formigas, Gardner 4563b (W); Gouveia, estrada Gouveia-Curvelo, na altura da bica de água potável, lado oposto da estrada, 18°35′42″ S, 43°57′16″ W, 4 Sep 2003 (fr), Cavalcanti et al. 3205 (CEN, MO); Gouveia, Fazenda do Sr. Everaldo, 11 Sep 1986 (fl. fr), Cavalcanti et al. (CFCR) 10313 (SPF); Guapé, Serra do Rio Grande, 2 May 1931 (fr), Mexia 5741 (MO, NY, P [2], RB); Montes Claros, rodovia BR-365, descida da Serra, 9 Mar 1998 (fl, fr), Hatschbach et al. 67551 (MBM); Penha da França, saída de Penha da França, Serra do Espinhaço, 18º S, 43º W, 11 Mar 1995 (fl), Splett 858 (CEN, UB); Rio Vermelho, Pedra Menina, Serra do Ambrósio, Espigão do Meio, 14 Jul 1984 (fr), Giulietti et al. (CFCR) 4464 (K, SP, SPF); rio Vermelho, Pedra Menina, Serra do Ambrósio, 31 Mar 1985 (fl), Menezes et al. (CFCR) 7681 (CEN, K, SP, SPF); rio Vermelho, Pedra Menina, Serra do Ambrósio, Fazenda Vargem do Anjo, Morro do Espigão do Meio, 13 Oct 1984 (fl), Isejima et al. (CFCR) 5413 (K, MBM, NY, SPF, UB); rio Vermelho, Pedra Menina, Serra do Ambrósio, 10 Jun 1990 (fr), Mello-Silva et al. 416 (CEN, SPF); Rio Vermelho, Pedra Menina, Serra do Ambrósio, Fazenda Vargem do Anjo, 8 Sep 1986 (fl), Cavalcanti et al. (CFCR) 10185 (F, K, MO, MBM, SPF); rio Vermelho, Pedra Menina, Serra do Ambrósio, Fazenda Vargem da Angélica, Morro da Virada do Mato Virgem, 14 Oct 1984 (fl), MelloSilva et al. (CFCR) 5494 (CEN, K, SPF); Santana do Riacho, UCAT, 15 Apr 1985, Andrade & Lopes s.n. (BHCH 8812). São Gonçalo do Rio Preto, Parque Estadual do Rio Preto, 18°05′28′′S, 43°20′32′′W, Lombardi 3373 (CEN, BHCB); São Gonçalo do Rio Preto, Parque Estadual do Rio Preto, 7 Apr 2000 (fl, fr), Lombardi et al. 3754 (BHCB, CEN); Serro, Grota Seca, São Gonçalo do Rio das Pedras, 12 Jan 1998 (fr), Vasconcelos s.n. (CEN 33603, BHCB 40209); Serro, 5 km s de Pedro Lessa na BR-259, 27 Nov 1985 (fl, fr), Graham 961 (MO, NY); Serro, estrada de

Descriptions

415

terra Serro-Milho Verde, a 10 km de Serro, 18°32′19″ S, 43°26′18″ W, 18 Jan 2003 (fl, fr), Cavalcanti et al. 3102 (CEN, MO); Várzea da Palma, Serra do Cabral, 16 Jan 1996, Hatschbach et al. 64204 (CEN, MBM); without reference to municipal district: Parque Nacional das Sempre Vivas, fazenda do “seu Geraldão”, acesso por Inhaí, 17°33′14″ S, 43°31′51″ W, 2 May 2010 (fr), Mello et al. 152 (UB); Cerrado do Jequitinhonha, 29 Nov 1992 (fl), Stehmann s.n. (BHCB 21085); Glaziou 19172 (P [2]); Riedel s.n. (K, P [3], W [2]). Diplusodon uninervius is recognized by linear, hyphodromous leaves and short internodes; linear-lanceolate prophylls with subrevolute, ciliate margins; a glabrous floral tube; and 12 stamens. The populations studied were homogeneous with respect to the morphological characteristics listed above. The only evident variation was in the length of the leaves. The specimens from the municipality of Rio Vermelho had short leaves (4 mm long), which is the lower limit for the species. Saint-Hilaire (1833) recognized, but did not name, three forms of Diplusodon candollei Pohl ex DC. One of these was based on Saint-Hilaire s.n., from Bom Fim, Minas Gerais. Koehne (1877) proposed a new species, D. uninervius, to accommodate this variant citing the syntypes Riedel 909 and Saint-Hilaire s.n. One sample of the Riedel 909 collection, identified as D. uninervius, was located at K, but was not annotated by Koehne. A specimen of Riedel 909, annotated by Koehne as a new species, was found at G, and another collection of Riedel 909, at LE, also annotated by Koehne is here designated as the lectotype of D. univervius, in view of the fact that specimens annotated by Koehne were transferred from Berlin to LE. The firststep lectotype designated by Lourteig (1989) is not accepted because she failed to cite a specific Riedel collection. 91. Diplusodon vacillans T.B.Cavalc., sp. nov. Type: Brazil. Minas Gerais: Rio Pardo de Minas, Serra Nova, sítio 25, ponto 2, 15°38′3″ S, 42°42′51″ W, 23 Jun 2017 (fl), A.  C. Sevilha, G.  Pereira-Silva, J.  B. Pereira 7020 (holotype, CEN [00102140]; isotypes, B, BHCB, K, MBM, MO, NY, OUPR, P, RB, SPF, US, W). Figs. 13.119, 15.5 C Diplusodon vacillans is an erect subshrub with flexible branches that become pendulous with flowers facing down. The whole plant is glabrous, with linear leaves, pedicellate flowers, prophylls surpassing the floral tube, epicalyx segments spreading to erect, and 12–20 stamens. Diplusodon tenuifolius and D. epilobioides are morphologically similar due to linear leaves. Diplusodon tenuifolius differs by its smaller flowers, shorter prophylls, and epicalyx segments of 0.8–1 mm long, much shorter than sepals. Diplusodon epilobioides differs mainly by the dense indument of minute trichomes, peciolate, eucamptodromous leaves with revolute margins, with one prominent vein on each side of the midvein. Shrubs 1.2–1.5  m tall, pendant, much-branched, not dark-brown when dry, totally glabrous. Upper branches flexible, rusted, pendant, cylindrical, unwinged, internodes 0.4–1  cm long. Decussate, slightly imbricate, erect, sessile; blades 10–30 × 1–2 mm, coriaceous, not discolorous, hyphodromous, linear to narrowly

416

13  Systematic Treatment

oblong, base acute, apex acute, margin plane, secondary veins 0, the midvein sulcate on adaxial surface, visible on abaxial surface; domatia absent. Synflorescence frondose, pleiobotryum type, lax; accessory branches absent, paraclades 10–40 cm long; bracts similar to the vegetative leaves in shape; flowers pedicellate, hypopodium 1–3 mm long, epipodium 1.5–2.5 mm long; prophylls 6–9 × 1.5–1.9 mm, surpassing the apex of the floral tube ca. 1  mm, inserted in the upper third of the pedicel, narrowly elliptic, unkeeled, base acute, apex acute, margin plane; floral tube 4–5 mm long, green, funnel-shaped; sepals 2–4 mm long, sometimes slightly wine-colored, triangular, caudate, not deflexed; epicalyx segments 4–4.5 mm long, cylindrical, spreading to erect, surpassing the sepals; corolla 2.5–3  cm in diam., lilac to pale-rose, petals 11–13 × 6–7 mm, obovate, apex obtuse; stamens 12–20, free portion of the filaments 5–5.5  mm, exserted; ovary 1.1–1.5  ×  1.5–2.3  mm, obovate-depressed, glabrous, style 9–12  mm long, ovules 8–14. Capsules 2.8–3 × 2.5–2.8 mm, globose, glabrous; seeds ca. 7, 1.5–1.8 × 1.5–1.7 mm. Specimens Examined: BRAZIL. Minas Gerais,  Rio Pardo de Minas, Vereda Funda, barranco próximo à estrada, depois do campo de Arnica, 15 Dec 2008 (fr), Sevilha et al. 5145 (CEN, NY); Rio Pardo de Minas, sítio 16, ponto 3, 15°27′44″ S, 42°31′14″ W, 27 Mar 2017 (fl), Sevilha et al. 6985 (BHCB, CEN, K, M, NY, OUPR, RB, SP, US, W). Distribution and Ecology (Fig. 13.119)  Diplusodon vacillans is known only from the municipality of Rio Pardo de Minas. It inhabits environments of rocky cerrado with lithic soil with quartz at 813–920 m elevation. Flowers from March to June; fruits registered in December. Diplusodon vacillans is an amazing ornamental species due to the architecture of its habit unusual for the genus. The individuals are subshrubs erect; however, the branches are flexible becoming fully pendulous swinging back and forth with pink and lilac flowers arranged at the ends of the branches and facing down (Fig. 15.5 C). In addition, other characteristics define the species as the absence of indumentum, linear leaves, flowers with pedicel evident (2.5–5.5 mm long), prophylls narrowly elliptic (1.5–1.9 mm wide) surpassing the floral tube, epicalyx segments 4–4.5 mm long spreading to erect, and 12–20 stamens. The most morphologically similar species to Diplusodon vacillans are D. tenuifolius and D. epilobioides, due to linear leaves though neither species shows the peculiar architecture with pendulous branches. Diplusodon tenuifolius, a species of southeastern Goiás, differs by its small flowers with corolla of 1–1.5 cm, shorter prophylls 3.5–4 mm long, and epicalyx segments 0.8–1 mm long, much shorter than sepals. Diplusodon epilobioides so with D. vacillans is a species from the north of Minas Gerais but differs mainly by the whole plant to be covered by a dense indument of inconspicuous trichomes, giving a velvety appearance, peciolate leaves with revolute margins, eucamptodromous, with one prominent vein on each side of the midvein. 92. Diplusodon venosus T.B.Cavalc., Phytotaxa 226(2): 149, fig. 4. 2015. Type: Brazil. Goiás: Cavalcante, entroncamento Minaçú-Kalunga (Engenho Dois),

Descriptions

417

Fig. 13.115  Diplusodon venosus. A. Part of vegetative branch with elliptic leaves. B. Part of vegetative branch with narrowly elliptic leaves. C. Part of inflorescence. D. Bract. E. Section of internode. F. Prophyll, abaxial surface. G. Flower with prophylls, without petals. H. Capitate stigma. (A Pastore & Suganuma 444, CEN; B–D Pastore et al 473, CEN; E–H Fonseca et al. 5095, CEN)

418

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km 5, 13°36′06″ S, 47°28′29″ W, 31 Aug 2004 (fl, fr), T.B.Cavalcanti & G. Pereira-Silva 3534 (holotype, CEN [00061825]; isotype, SPF). Figs. 13.115 A–H, 13.119, 15.5 E Subshrubs 0.35–0.7 m tall, erect, branched, glabrescent, not dark-brown when dry. Branches slightly quadrangular and flattened, with an inconspicuous wing, the upper branches straw-colored, with minute and very inconspicuous trichomes to glabrous, internodes 3–4.5  cm long. Leaves decussate, rarely verticillate, not ­imbricate, erect, subsessile; petioles 0.5–1  mm long; blades 5.5–7  ×  0.6–2.6  cm, coriaceous, not discolorous, not glaucous, eucamptodromous to acrodromousbasal-supranumerary, narrowly elliptic, narrowly lanceolate to elliptic, base obtuse to acute, apex acute, slightly acuminate, margin plane, irregular, with minute and inconspicuous cilia, blades glabrescent to glabrous, trichomes when present slightly bulbous at the base, sparse, common on the abaxial surface, very prominent; domatia present, inconspicuous. Synflorescence bracteose, triplobotryum to pleiobotryum type, lax; accessory branches absent, principal florescence botryum type, usually truncate, paraclades 5–25 cm long; bracts 15–30 × 1–4 mm, glabrous, linear to narrowly elliptic; flowers short-pedicellate, hypopodium 0.5–1 mm long, epipodium to 0.5 mm long; prophylls 4–4.2 × 1.5–1.7 mm, surpassing the middle of the floral tube, almost reaching the apex of the floral tube, narrowly elliptic to obovate, keeled, base retuse to acute, apex acute, margin plane; floral tube 4.5–5 mm long, funnelshaped, glabrous; sepals 2–2.2 mm long, triangular, not deflexed, glabrous; epicalyx segments ca. 2 mm long, slightly dorsiventrally compressed, spreading, glabrous, shorter than the sepals; corolla 3.5–4  cm in diam., lilac to rose-magenta, petals 13–19 × 7–10 mm, obovate, apex obtuse; stamens 18–24, free portion of the filaments 6–6.5  mm long, slightly exserted; ovary 2–2.2  ×  2.2–3  mm, obovatedepressed, glabrous, style 7–10 mm long, ovules 22–42. Capsules 4.5–5 × 4–4.5 mm, globose, glabrous; seeds ca. 31, 1.5–2.2 × 1.5–2.8 mm. Distribution and Ecology (Fig. 13.119)  Diplusodon venosus is known only from the municipality of Cavalcante in the vicinity of the traditional community of Kalunga. It inhabits cerrado and fields on sandy soil and sandstone at 919–1030 m. Flowers from February to August; fruits registered in March, August, and November. Specimens Examined: BRAZIL.  Goiás,  Cavalcante, aldeia Kalunga, 6 Mar 2003 (fl), Pastore & Suganuma 444 (CEN); Cavalcante, estrada para Prata (Kalungas), 13°22′24″ S, 47°41′09.1″ W, Cavalcante, comunidade Kalunga, caminho para a cachoeira Capivara, 20 Apr 2003, (fl), Pastore et  al. 471 (CEN); Cavalcante, comunidade Kalunga, Pastore et  al. 473 (CEN); Cavalcante, estrada para povoado do Engenho, 13°40′40″ S, 47°28′22″ W, 30 Mar 2011 (fl, fr), Bringel & Moreira 753 (CEN, UB); Cavalcante, estrada para Prata (Kalungas), 13°22′24″ S, 47°41′9.1″ W, 14 Apr 2004 (fl), Fonseca et al. 5095 (CEN, HRB, IBGE); Cavalcante, RPPN Serra do Tombador, 7 Nov 2012 (fr), Pereira-Silva et  al. 16445 (CEN); Cavalcante, 10 Jul 2011 (fl, fr), Silva 662 (UB); Cavalcante estrada municipal não pavimentada Cavalcante-Minaçu, 13°34′20.0″ S, 47°31′25.1″ W, 19 Feb 2013 (fl), Lombardi et al. 9882 (CEN, HRCB).

Descriptions

419

Diplusodon venosus is an unusual species of the genus from the mountains of Goiás. The upper branches are straw-colored, and the bracteose inflorescence is long, broad, and conspicuous, with linear to narrowly elliptic bracts that are different from the vegetative leaves, especially in size. The leaves vary in width, ranging in shape from narrowly elliptic to elliptic and lanceolate, but always with strongly prominent veins, which are also present in the bracts. Based on the bracteose inflorescence, D. venosus resembles D. astictus and D. ramosissimus, which also occur in the region. Diplusodon astictus can be distinguished because it has upper branches that are wine-colored to chestnut and larger flowers with 12 stamens. Diplusodon ramosissimus can be distinguished because the leaves have conspicuous domatia and the flowers have very short prophylls that do not reach the middle of the floral tube. 93. Diplusodon verruculosus T.B.Cavalc., Phytotaxa 226(2): 151, fig. 5. 2015. Type: Brazil. Minas Gerais: Santana de Pirapama, Serra do Cipó, Capelão de São José, subida da Senhorinha, 18°57′14″ S, 43°45′31″ W, 6 Mar 2012 (fl, fr), D. C. Zappi, J. G. Rando, A. C. Araújo, V. Scalon, Y. Antonini, C. Martins & E. Cândido 2733 (holotype, CEN [00074415]; isotype, SPF, RB [2]). Figs. 13.116 A–H, 13.119 Subshrubs, erect, with a xylopodium, strongly branched from the base, caespitose, not dark-brown when dry. Upper branches cylindrical, unwinged, strigose, internodes 0.8–2 cm long. Leaves decussate, slightly imbricate, erect, coriaceous, sessile; blades 3–6 × 1–1.5 mm, not discolorous, hyphodromous, narrow triangular, base retuse to cordate, apex acute, margin revolute to the midvein, strigose, blades covered by sparse bulbous-based trichomes which give the leaf surface a verrucose aspect; lateral veins absent, hispid; domatia absent. Synflorescence frondose, diplobotryum type, congested; accessory branches absent, principal florescence truncate, paraclades 4–9 mm long; bracts similar to the vegetative leaves in shape and size; flowers sessile, concentrated at the apex of the paraclades; prophylls 5–5.5 × 2.8–3 mm, surpassing the apex of the floral tube, elliptic, slightly keeled, margins subrevolute, base acute, with sparse and fine trichomes, almost glabrous; floral tube 3.5–3.8 mm long, funnel-shaped, with sparse and fine trichomes, almost glabrous; sepals 3.8–4 mm long, triangular, caudate, not deflexed, with sparse and fine trichomes, almost glabrous, strongly ciliate; epicalyx segments 3–3.2 mm long, narrowly triangular, erect, ciliate, the same size as or a little shorter than the sepals; corolla ca. 2 cm in diam., deep rose, petals 9–10 × 4.5–5 mm; stamens 6, free portion of the filaments 6–6.5  mm long, included within the sepals; ovary ca. 2.5– 2.7 × 2–2.6 mm, obconic, glabrous, style 10–11 mm long, ovules 18–20. Capsules not seen. Distribution and Ecology (Fig. 13.119)  Diplusodon verruculosus is known only from the region of Santana de Pirapama. The species grows in campos rupestres of Cadeia do Espinhaço, Minas Gerais at 1264 m. Flowers from January to March; fruits registered in March.

420

13  Systematic Treatment

Fig. 13.116  Diplusodon verruculosus. A. Habit. B. Section of internode. C. Leaf, adaxial surface. D. Detail of leaf adaxial surface showing bulbous-based trichomes. E. Leaf, abaxial surface. F. Detail of leaf abaxial surface. G. Prophyll, adaxial surface. H. Flower with prophylls, without petals. (A–H Zappi et al. 2733, CEN)

Specimens Examined: BRAZIL. Minas Gerais,  Santana de Pirapama, Serra do Cipó, estrada S.  José da Cachoeira-Inhame, trilha da Senhorinha, 19°0′0″ S, 43°45′47″ W, 19 Feb 2007 (fl), Zappi et al. 830 (ESA, RB, SPF). Diplusodon verruculosus is characterized by small, narrowly triangular leaves, with strongly revolute margins and an indumentum of short bulbous-based

Descriptions

421

trichomes. The flowers have long sepals and epicalyx segments that are distinctly ciliate with bulbous-based trichomes, and the androecium has 6 stamens. The species name refers to the verrucose leaf surface aspect given by the bulbous base of the trichomes. Diplusodon verruculosus is vegetatively similar to D. saxatilis; however, D. saxatilis is well defined by its hirsute branches, leaves, and floral tube and its short (2.5–3 mm long) epicalyx segments that are distinctly shorter than the sepals, in addition to its androecium with 12 stamens. 94. Diplusodon vidalii Lourteig, Bradea 5(19): 216. 1989. Type: Brazil. Minas Gerais: Thermópolis, Balneário de Águas Quentes, Apr 1945, J.  Vidal I-713 (holotype, R [000043428]; isotypes, P, R [000043428a]). Figs. 13.117 A–H, 13.119 Subshrubs 0.2–1  m tall, erect, much-branched, glabrous, delicate, not darkbrown when dry. Upper branches quadrangular, yellow when young, rust-colored when older, winged, upper internodes slightly compressed, internodes 2–5 cm long. Leaves decussate, not imbricate, erect, petiolate; petioles 1–1.5  mm long; blades 3–4  ×  0.5–1.5  cm, membranaceous, not discolorous, punctate, hyphodromous, eucamptodromous, elliptic, base acute or attenuate, apex acute to obtuse, sometimes acuminate, margin plane; veins ca. 3, inconspicuous on the adaxial surface, thin on the abaxial surface, midvein prominent, the secondary veins only sulcate or slightly raised; domatia absent. Synflorescence bracteose, diplobotryum to pleiobotryum type, lax; accessory branches absent, paraclades 4–12  cm long; bracts 10–18  ×  2.5–5  mm, elliptic-linear; flowers pedicellate, hypopodium 0.5–1  mm long, epipodium 0–0.5 mm long; prophylls 2–2.5 mm long, reaching the middle or slightly below the middle of the floral tube, linear to linear-spathulate, unkeeled; floral tube 4–4.5 mm long, green, funnel-shaped; sepals 2.7–3 mm long, triangular, sometimes slightly caudate, not deflexed, apex minutely ciliate; epicalyx segments 0.8–1.5 mm long, cylindrical, deflexed to spreading, shorter than the sepals; corolla ca. 2 cm in diam., deep rose to lilac, petals 7–7.5 × 4–4.5 mm, obovate, apex obtuse; stamens 12, free portion of the filaments 7–8  mm long, exserted 4–5  mm; ovary 1.8–2 × 2–2.5 mm, oboval, glabrous, style 13–14 mm long, ovules 23–26. Capsules not seen. Distribution and Ecology (Fig.  13.119)  Diplusodon vidalii is endemic to the mountains of Termópolis and Chapadão in southern Minas Gerais. It occurs in dense cerrado and the margins of forests on the slopes of the mountains and open cerrado and campos rupestres on the mountains, on quartzite sands at 1050–1130 m. Late stages of flowers and immature fruits from April to May. Specimens Examined: BRAZIL.  Minas Gerais,  São Sebastião do Paraíso, Distrito de Termópolis, Estância Hidromineral de Termópolis, 20°55′58″ S, 46°49′57″ W, 13 Jan 2003 (st), Cavalcanti et al. 3063 (CEN, MO, SP); São Sebastião do Paraíso, Distrito de Termópolis, Estância Hidromineral Thermópolis, Serra de Termópolis, 20°55′58″ S, 46°49′57″ W, 29 Apr 2003 (fl), Cavalcanti et  al. 3171

422

13  Systematic Treatment

Fig. 13.117  Diplusodon vidalii. A. Vegetative branch. B. Leaf, abaxial surface. C. Branch with inflorescence. D. Schematic representation of inflorescence. E. Bract. F. Prophyll, adaxial surface. G. Flower with prophylls, without petals. H. Ovary. (A–H Cavalcanti et al. 3171, CEN)

Descriptions

423

(CEN); São João do Paraíso, Termópolis, 24 May 1994 (fl), Simão-­Bianchini 455 (CEN, SP). The species is characterized by its bracteose inflorescences, which are well differentiated from the vegetative part of the plant, and by its small flowers. Diplusodon vidalii is similar to D. virgatus due to elliptical, membranaceous leaves and flowers with 12 stamens. However, the flowers in D. virgatus are white, and in D. vidalii are deep rose to lilac. In southern Minas Gerais, the two species occur sympatrically in dense cerrado and on the edges of forests. On the high parts of mountains of Termópolis, there is a dense population of caespitose individuals of D. vidalii; on the lower slopes, the individuals are tall and not caespitose. Diplusodon vidalii is frequent in the mountainous and rocky area where it occurs. 95. Diplusodon villosissimus Pohl, Flora 10: 151. 1827. Diplusodon lythroides Mart. ex DC. var. villosissimus (Pohl) DC., Prodr. 3: 94e, 1828. Type: Brazil. Minas Gerais: Ouro Preto, in montosis graminosis, circa Vila Rica, G. Schüch 5966 (first-step lectotype, designated by Lourteig 1989: 238, second-step lectotype, W [0069494], here designated; isolectotype, W [0069493]). Figs. 13.118 A–G, 13.119 Friedlandia stachyoides Cham. & Schltdl., Linnaea 2: 353. 1827. Diplusodon stachyoides (Cham. & Schltdl.) DC., Prodr. 3: 94e. 1828. Type: Brazil. Brazil. Locality unknown, s.d., F.  Sellow s.n. (holotype, B, destroyed; isotypes, not found). Diplusodon lythroides Mart. ex DC., Prodr. 3: 94e. 1828. Friedlandia lythroides Mart. ex DC., Prodr. 3: 94e. 1828, pro syn. Diplusodon lythroides Mart. ex DC. var. glabrescens DC., Prodr. 3: 94e. 1828. Type: Brazil. Minas Gerais austral et São Paulo, C. F. P.Martius s.n. (lectotype, M [146273], designated by Lourteig 1989: 225; isolectotype, G). Subshrubs 0.3–0.9  m tall, erect, usually branched just at inflorescence, sometimes caespitose, with a xylopodium, not dark-brown when dry. Upper branches subquadrangular to cylindrical, unwinged, lax, densely covered by long, white trichomes, wine-colored on the younger parts, internodes 2–6 cm long. Leaves decussate, not imbricate, suberect, subsessile; petioles 1–2  mm long; blades 20–60 × 15–45 mm, coriaceous, not discolorous, acrodromous-basal-supranumerary, concave when fresh, villous, ovate or ovate-lanceolate to rarely elliptic, base obtuse to subcordate, apex obtuse, frequently acuminate, margin plane, adaxial and abaxial surface covered by long, white trichomes, sparse on the leaves; veins 4–6 on each side, 2–3 of them departing from each side of the base, 1–2 departing on each side along the midvein, yellow; domatia absent, sometimes present and inconspicuous. Synflorescence frondose-bracteose, botryum to diplobotryum type or rarely triplobotryum, congested; accessory branches absent, principal florescence 4–23 cm long, botryum type, paraclades 4–18  cm long; bracts 14–18  ×  7–9  mm, slightly shorter than the vegetative leaves, much shorter than the paraclades; flowers pedicellate, hypopodium 1–1.5  mm long, epipodium 0–0.5  mm long; prophylls

424

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Fig. 13.118  Diplusodon villosissimus. A. Habit. B. Section of internode. C. Leaf, abaxial surface. D. Detail of leaf indumentum. E. Trichome of leaf. F. Leaf, acuminate apex. G. Flower with prophyll, without petals. (A Martens 258, SPF; B–G Badini s.n., OUPR 21534)

Descriptions

425

6–9 × 3–4 mm, surpassing or the same size as the floral tube, elliptic-lanceolate, unkeeled, apex acute, margins plane, ciliate, densely pilose; floral tube 6.5–7.5 mm long, funnel-shaped, densely covered with long, white trichomes turning grayish, wine-colored under the trichomes; sepals 4–5 mm long, triangular, not caudate, not deflexed, pilose as on the floral tube; epicalyx segments 4.5–6 mm long, cylindrical, erect, pilose as on the floral tube, surpassing the sepals; corolla 4.5–5 cm in diam., lilac to purple, petals 16–22 × 8–14 mm, narrowly obovate, apex obtuse; stamens 15–19, free portion of the filaments 7.5–8.5 mm long, included; ovary 2–3 × 2.5– 3.5 mm, obovate-depressed, glabrous, style 11–13 mm long, ovules 30–56. Capsules 4.8–5 × 4–4.3 mm, globose, glabrous; seeds 16–43, 2–2.2 × 1.7–2.5 mm. Distribution and Ecology (Fig.  13.119)  Diplusodon villosissimus occurs in the states of São Paulo and southern Minas Gerais, in cerrado, campo rupestre, and rocky campo sujo vegetation, at 1000–1500 m. In Itabirito it grows sympatrically with D. buxifolius and D. virgatus. Flowers from January to May; fruits from August to December. Specimens Examined: BRAZIL.  Minas Gerais,  Aiuroca, 16 Feb 2000 (fl), Tameirão-Neto 2877 (BHCB); Alpinópolis, arredores da barragem de Furnas, 29 Nov 1969 (fl), Emygdio 2809 (R); Araxá, 1 Feb 1993 (fl), Brandão 22439 (PAMIG); Barbacena, Sítio Rancho Novo, 9 Mar 2013 (fl), Gonzaga 171 (CEN, CESJ n.v); Belo Horizonte, Lagoa Seca, 5 Feb 1900 (fl), Schwacke 18802 (SP); Serra do Taquaril, 7 Feb 1933 (fl), Mello-Barreto 3373 (CEN, RB, SPF); Serra do Curral, Mar 1953 (fl, fr), Roth 2219 (CESJ, SPF); Brumadinho, 20o8′S, 44º13′W, Serra da Moeda, Retiro das Pedras, 13 Feb 1990 (fl), Martens 258 (SPF); Cachoeira do Campo, 7 Apr 1976 (fl), Badini s.n. (OUPR 22937); Cachoeira do Campo, Badini s.n. (OUPR 23809); Caldas, Feb/Mar 1967 (fl), Regnell 116 (NY); Cristais, morro ao lado da cidade, 14 Feb 1998 (fl), Goldenberg et al. 565 (HUFU); Delfinópolis, subida da Serra da Canastra, a 14 km da ponte sobre o rio Grande, 22°12′30″ S, 47°06′16″ W, 14 Jan 2003 (fl), Cavalcanti et al. 3073 (CEN); Serra da Canastra, ca. 45 km de Delfinópolis, em direçao a Peixoto, 6 Sep 1998 (fl, fr), Souza et al. 21226 (ESA); Entre rios, Fazenda da Pedra Branca, 29 Jan 1977 (fl), Coons et al. 77–349 (CEN, VIC); Furnas, estrada Furnas-Belo Horizonte, 6 Jul 1985 (fl), Lombardi 843 (BHCB); Hargreaves, Baeta 10525 (OUPR); Itabirito, Serra do Itabirito, ca. 50 km SE de Belo Horizonte, 10 Feb 1969 (fl), Irwin et al. 19722 (NY, UB); alto do Morro do Cristo, acesso pela cidade de Itabirito, 20°25′27″ S, 44°50′28″ W, 27 Feb 2000 (fl), Cavalcanti et al. 2584 (CEN, MO, NY); Lagoa Santa, Warming s.n. (K, NY n.v., P, W); Morro do Ferro, estrada de Morro do Ferro para Passa Quatro, 8 km de Morro do Ferro, Serra das Vertentes, 21°13′ S, 44°54′ W, 26 Feb 2000 (fl, fr), Cavalcanti et al. 2561 (CEN, MO, NY); Nova Lima, Serra do Cachimbo, 14 Mar 1999 (fl), Pompeu 392 (BHCB, CEN); Nova Lima, estrada de terra para Barão de Cocais, sobre a Serra da Gandarela, a 5  km do entroncamento do rio acima, 20°06′11″ S, 43°45′10″ W, 9 Mar 2014 (fl, fr), Cavalcanti & Pereira-Silva 3787 (CEN); Ouro Branco, Serra do Ouro Branco, 28 Feb 1975 (fl), Badini s.n. (OUPR 22156); Serra de Ouro Branco, 20°29′ S, 43°39′ W, 8 Mar 1995, Souza et al.7913 (CEN, ESA); Ouro Preto, Chapada, 23 Feb 1975 (fl), Badini s.n. (OUPR 23053);

426

13  Systematic Treatment

Fig. 13.119  Geographic distribution of Diplusodon vacillans, D. venosus, D. verruculosus, D. vidalii, and D. villosissimus

Descriptions

427

Ouro Preto, Chapada, Badini s.n. (OUPR 21534); Ouro Preto, Serra da Chapada, Badini s.n. (OUPR 19670); Rodrigo Silva, 26 May 1974 (fl, fr), Badini s.n. (OUPR 21692); Rodrigo Silva, Badini s.n. (OUPR 22818); Rodrigo Silva, Badini s.n. (OUPR 26078); São Julião, 9 Mar 1891 (fl), Schwacke s.n. (R); 26 Feb 1978 (fl), Badini s.n. (OUPR 24378); Passos, estrada entre Furnas e Passos, ca. 16 km do rio Turvo, em direção a Furnas, Fazenda Paredão, Usina Açucareira Passos, 1 Jul 1996 (fl), Souza et al. 11481 (ESA); São Roque de Minas, Parque Nacional da Serra da Canastra, 10 Apr 1999 (fl), Mota 44 (BHCB); Santa Rita de Cássia, Morro da Furna, Mar 1945 (fl), Vidal I-524 (R); São Roque de Minas, Parque Nacional da Serra da Canastra, 16 Feb 2000 (fl), Mota 131 (BHCB, CEN); próximo a Casa de Pedra, 23 Feb 1994 (fl), Nakajima & Romero 197 (HUFU); morro próxima à sede administrativa, 19 Feb 1997 (fl), Romero et al. 3829 (CEN, HUFU); estrada para Sacramento, após o vale da nascente do rio São Francisco, 18 Apr 1994 (fl, fr), Romero et al. 911 (CEN, HUFU); final da estrada para a Fazenda do Fundão, 22 Feb 1997 (fl, fr), Nakajima et al. 2242 (CEN, HUFU); Cachoeira Casca D’anta, 20 Mar 1995 (fl), Nakajima et al. 930 (CEN); Serro, Martius obs. 724 (M); Serro, Martius obs. 723 (M [2]); Tres Moinhos, Badini s.n. (OUPR 22040); Tres Moinhos, 19 Mar 1986 (fl, fr), Badini s.n. (OUPR 26769); Sete Lagoas, 13 Feb 1973 (fl), Hatschbach & Ahumada 31509 (MBM, NY); Três Corações, rodovia Três Corações-São Tomé das Letras, 4 Feb 1973 (fl), Hatschbach & Ahumada 31252 (MBM); without reference to municipal district: BR-58, entre Caxambú e Pouso Alto, 29 Mar 1964 (fl), Trinta 672 & Fromm 1748 (HB, NY); base da Serra do Itatiaia, 19 May 1992 (fr), Badini s.n. (OUPR 19669); Serra dos Ilheus, 21 Jan 1889 (fl, fr), Glaziou 17617 (K, P, R); Glaziou 17619 (K); Claussen s.n. (BR, K, OXF). São Paulo, Pedregulho, Parque Estadual de Furnas do Bom Jesus, 20°14′52″ S, 47°27′137″ W, 13 Dec 2003 (fl, fr), Sasaki & Sasaki 879 (CEN, SPF); Pedregulho, 2 May 1995 (fl), Marcondes-Ferreira et al. 1109 (CEN, UEC); Pedregulho, 10 Apr 1920 (fl, fr), Gehrt s.n. (CEN, 29268, SP 4017); Parque Estadual de Furnas do Bom Jesus, 27 Jan 1993 (fl), Macedo 57 (SP); Pedregulho, Parque Estadual de Furnas do Bom Jesus, 18 Mar 1994 (fl), Marcondes-Ferreira et al. 806 (CEN, UEC); Pedregulho, estrada para Estreito, 17 Jan 1996 (fl), Marcondes-Ferreira & Belinello 1248 (SP, UEC); Pedregulho, Estreito, Fazenda Três Irmãos, 18 Jan 1996 (fl), Marcondes-Ferreira & Belinello 1270 (CEN, UEC [2]); Raposo, 20°01′79″ S, 43°76′40″ W, 5 Aug 2010 (fr), Mansur & Tameirão-Neto 91 (BHCB). Without reference to state or municipality: Martius Herb. Fl. Bras. n. 868 (K, M [2], P; W); Riedel 2597 (LE n.v.); Riedel s.n. (K, LE n.v., W). The specific epithet of Diplusodon villosissimus alludes to the great quantity of long white trichomes that cover the entire plant and confer a grayish aspect. This species is also characterized by ovate or ovate-­lanceolate to rarely elliptic leaves, flowers with elliptic-lanceolate prophylls reaching or slightly surpassing the apex of the floral tube, erect epicalyx segments, and 15–19 included stamens. Pohl (1827a) described Diplusodon villosissimus without citation of specimens in the protologue. In a later publication, Pohl (1827b) cited a single gathering of D. villosissimus collected by Schüch in November, near to Vila Rica (now Ouro Preto),

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Minas Gerais. Lourteig (1989) cited Schüch s.n. (W) as the holotype of D. villosissimus, which should be treated as a first-step lectotype (ICBN Art. 9.17, ex. 14). There are two specimens of this gathering at W: One has a label annotated by Pohl and is numbered as Schüch 5966; this specimen is here designated as the secondstep lectotype (W0069494). The other specimen is labeled as Schüch s.n. but is clearly a duplicate of Schüch 5966. The herbarium W database refers to R. Schüch instead G. Schüch; however, the collector is Guilherme Schüch, naturalist, born in Ouro Preto, Brazil, in 1824, also known as Baron de Capanema. Diplusodon lythroides was described by Candolle (1828: 94e) based on an inflorescence branch (Martius s.n.) with bracts that are smaller than the vegetative leaves. This specimen is here regarded as D. villosissimus. Lourteig (1989) recognizes D. lythroides; however, all specimens that she cited, apart from the type, are from Bahia, and these specimens are treated here as D. bahiensis. Lourteig (1989) indicates the specimen of Martius s.n. at M as holotype, which is considered as the first-step lectotype (ICBN Art. 9.17, ex. 14). 96. Diplusodon villosus Pohl, Flora 10: 151. 1827. Type: Brazil. Tocantins: Ad Ponto Alto (Ponte Alta), in pascuis graminosis, Jan 1820, L. B. E. Pohl 1066 = D 937 (lectotype, W [0053440], designated by Lourteig 1989: 225; isolectotypes, BR, K [2], M). Figs. 13.120 A–H, 13.122, 15.5 F Diplusodon humilis Koehne, Bot. Jahrb. Syst. 23(Beibl. 57): 33. 1897. Type: Brazil. Goiás: córrego do Brejo, 27 Apr 1895 (fl), A.  F. M.  Glaziou 21423 (first-step lectotype, designated by Lourteig 1989: 225, second-step lectotype, P [01902002], here designated; isolectotypes, BR, fragm. F, G, K, P [01902001], R). Diplusodon pulchellus Koehne, Bot. Jahrb. Syst. 23(Beibl. 57): 32. 1897. Type: Brazil. Goiás: Córrego do Brejo, 24 Feb 1895, A. F. M. Glaziou 21422a (firststep lectotype, designated by Lourteig 1989: 225, second-step lectotype, P [01902000], here designated; isolectotypes, F, K). Subshrubs 0.2–1.2  m tall, erect, usually branched just at inflorescence, often caespitose, with a xylopodium, not dark-brown when dry. Upper branches subquadrangular, unwinged, villous-tomentose, internodes 2–5.5  cm long. Leaves decussate, not imbricate, suberect, sessile to subsessile; petioles ca. 1 mm long; blades 20–55  ×  10–22  mm, slightly ashy gray, chartaceous, sometimes discolorous, acrodromous-­ basal-­ supranumerary, elliptic-lanceolate to less often ovate, base obtuse, apex obtuse to acute or acuminate, margin revolute, adaxial surface with long, sparse trichomes, abaxial surface with long, sparse trichomes more concentrated on the veins; secondary veins 3–4 on each side, 1–2 of them departing from the base of the midvein; domatia present. Synflorescence bracteose to frondosebracteose, diplobotryum or triplobotryum type or less often botryum type, congested; accessory branches absent, principal florescence 7.5–28 cm long, paraclades 4–20 (−30) cm long, botryum type, often truncate; bracts 6–20 × 3–8 mm, elliptic, longitudinally folded, with the same indumentum as the leaves, margins revolute;

Descriptions

429

Fig. 13.120  Diplusodon villosus. A. Habit. B–C. Leaves, abaxial surface. D. Detail of leaf veins showing domatia. E. Prophyll, adaxial surface. F. Prophyll, abaxial surface. G. Flower with prophyll, without petals. H. Trichome. (A Irwin et al. 34260, NY; B, D Almeida 924, UB; C, E, F Cavalcanti et al. 500, CEN; H Cavalcanti et al. 1265, CEN)

430

13  Systematic Treatment

flowers sessile, prophylls 2–3 × 1–2.5 mm, less than half the length of the floral tube, elliptic to orbicular, unkeeled, base retuse, apex obtuse, margin plane; floral tube 4–5 mm long, funnel-shaped, villous-tomentose; sepals 2.5–3 mm long, winecolored, triangular, not caudate, not deflexed, villous-tomentose; epicalyx segments 0.5–1.5 mm long, wine-colored, cylindrical to rounded, spreading, longitudinally ciliate, shorter than the sepals; corolla 3–4 mm in diam., rose to rose-magenta, petals 10–12(−17) × 7–9(−12) mm, obovate, apex obtuse to acute; stamens 18, free portion of the filaments 7–8 mm long, exserted 1–2 mm, rose; ovary 2.5–3 × 2.2– 2.5  mm, rhombic, glabrous, style 11–12  mm long, rose; ovules 31–38. Capsules 1.5–2.1 × 1.5–2 mm, globose, glabrous. n = 15 (Graham 1985); 2n = 30 (Cavalcanti 1995). Distribution and Ecology (Fig.  13.122)  Diplusodon villosus occurs in Distrito Federal and the states of Goiás and Minas Gerais and is known from the state Tocantins only from the type. It grows in cerrado vegetation, on red-yellow latosols, in campos rupestres, and wet fields or adjacent to gallery forests, at 600–1250 m. Flowers from November to August; fruits from November to July. Specimens Examined: BRAZIL. Distrito Federal,  Adjacências da barragem do rio Descoberto, 27 Feb 2004 (fl), Pastore et al. 867 (CEN); Águas Emendadas, 2 Jul 1971 (fl, fr), Ferreira 766 (BHCB, BHMH, CEN, HEPH); APA da Cafuringa, Fercal, 15°27′27″ S, 47°58′ 00″ W, 22 Mar 2006 (fl), Amaral et al. 27 (CEN); Barra Alta, córrego São Gonçalo, 15°48′ S, 47°31′ W, 19 Feb 1981 (fl), Kirkbride 3884 (NY); Brasília, rodovia de acesso à fazenda Sucupira, 1 Apr 1998 (fl, fr), PereiraSilva & Sampaio 3885 (CEN, MO); Brasília, fazenda Sucupira, 15°55′ S, 48°01′ W, 5 May 1994 (fl), Assis et al. 10 (CEN, MO); Brasília, fazenda Sucupira, 22 Apr 1999 (fl), Faria et al. 1 (CEN, MO); Brasília, fazenda Sucupira, 22 Apr 1999 (fl, fr), Faria et al. 2 (CEN, MO); Brasília, fazenda Sucupira, campos à nordeste da fazenda, 26 Feb 1999 (bud), Sampaio 346 (CEN, MO); Brasília, fazenda Sucupira, estrada entre o restaurante e a sede da fazenda, 15°52′ S, 48º W, 9 May 2000 (fl, fr), Guarino & Pereira 180 (CEN, MO); Reserva Biológica do Jardim Botânico de Brasília, 15°52′ S, 47°41′ W, 16 Apr 1993 (fl, fr), Proença 887 (UB); 15 km ao SE do centro de Brasília, 16 May 1984 (fl), Fonseca 6 (UB); a 17 km a SE do centro de Brasília, 16 May 1984 (fl), Neves 9 (UB); 17 km em linha reta ao SSE do centro de Brasília, 16 May 1984 (fl), Menezes 4 (UB); Jardim Botânico de Brasília, entre as divisões DEA e DECO, 15°52′ S, 47°51′ W, 21 Feb 1997 (fl), Nóbrega et al. 694 (HEPH); Trilha Ecológica, 6 Apr 1995 (fl), Martins 80 (HEPH); Jardim Botânico de Brasília, 15°52′ S, 47°51′ W, 6 Feb 1996 (fl), Boaventura & Silva 231 (HEPH); Parque Rural, Fundação Zoobotânica, 10 Jan 1967 (fl), Duarte 10146 (CH); Parque Rural, Fundação Zoobotânica, 3 Mar 1977 (fl), Monteiro s.n. (CEN 1011); Parque Rural, pátio do Cenargen, 21 Feb 1977 (fl, fr), Allem 812 (CEN); entre a Colina e Lago Norte, 1 Apr 1980 (fl), García-Kirkbride 1162 (CEN, UB); Parque Nacional de Brasília, 15 Apr 1998 (fl), Amaral-Santos&Cordovil-Silva 96 (CEN, MO); Parque Nacional de Brasília, 18 Mar 1998, Cavalcanti et al. 2391 (CEN, MO, NY); Parque Nacional de Brasília, 4 Feb 1992 (fl), Barros et al. 2243 (UB); Parque Nacional de Brasília, 5 Feb 1992 (fl, fr), Barros et al. 2282 (UB); Parque Nacional de Brasília, 5

Descriptions

431

Feb 1992 (fl, fr), Barros et al. 2281 (UB); Parque Nacional de Brasília, 13 Feb 1968 (fl), Philcox & Onishi 4306 (UB); Parque Florestal, 12 Apr 1963 (fr), Pires et al. 9005 (UB); SQN 208, 15°45′ S, 45°53′ W, 20 Feb 1986 (fl), Nascimento & Catharina 56a (UB); Cidade Universitária, 6 May 1964 (fl, fr), Jesus 7 (UB); Cidade Universitária, 16 Jan 1965 (fl), Handro 110 (SP); Cidade Universitária, 29 May 1965 (fr), Sucre 409 (UB); Cidade Universitária, perto do Centro Olímpico, 4 Feb 1977 (fl), Aires 4869 (UB); Cidade Universitária, 4 Feb 1977 (fl, fr), Aires 4860 (UB); Campus, Aug 1980 (fl), Taxonomy Class of The Universidade de Brasilia 362 (MO, UB); Campus, 27 Mar 1968 (fl, fr), Lima 140 (NY, UB); Fazenda Água Limpa (FAL), Universidade de Brasília, 22 Jun 1967 (fl), Santiago 25 (UB); Fazenda Água Limpa (FAL), Universidade de Brasília, 10 May 1980 (fl), Santos et  al. 8 (UB); Fazenda Água Limpa (FAL), Universidade de Brasília, 15 Mar 1976 (fl, fr), Ratter & Fonsêca 2763 (MO, UB); Fazenda Água Limpa (FAL), Universidade de Brasília, 22 Jan 1980 (fl), Cesar 68 (UB); Fazenda Água Limpa (FAL), Universidade de Brasília, 2 Apr 1983 (fl), Haridasan 109 (UB); Fazenda Água Limpa (FAL), Universidade de Brasília, 2 May 1986 (fl), Barros 1018 (SPF, UB); Fazenda Água Limpa (FAL), Universidade de Brasília, 2 May 1986 (fr), Barros 1021 (UB); Fazenda Água Limpa (FAL), Universidade de Brasília, 23 Apr 1984 (fl), Haridasan 454 (UB); Fazenda Agua Limpa - FAL, perto do laboratório de Ecologia, 2 Jun 1985 (fr), Barros 985 (UB); Sul de Brasília, 8 Dec 1965 (fl), Irwin et al. 11096 (MO, NY). Brazlândia, Pires 275 (CEN, UB); Fazenda José Pires, ca. 4  km de Brazlândia, 15°40′ S, 48°12″ W, 28 May 1991 (fl, fr), Vieira et al. 775 (CEN, MO); Brazlândia, próximo ao Poço Azul, Lago Oeste, APA da Cafuringa, 15 May 2006 (fl, fr), Pastore & Suganuma 1507 (CEN); Núcleo Bandeirante, APA Gama e Cabeça de Veado, ARIE do córrego do Cedro SPMW, Q 26, conjunto 3, área vizinha à Associação dos Empregados da Embrapa, 15°54′48.9″ S, 47°57′50.2″ W, 29 Apr 2003, Fonseca & Alvarenga 4621 (CEN); Q26 conjunto 12, 15°53′36.1″ S, 47°57′10.4″ W, 23 Apr 2003, Fonseca & Alvarenga 4597 (CEN); córrego Vicente Pires, 15°47′ S, 48°01′ W, Sato 3 (UB); Setor de Indústria, 15 May 1965 (fl), Heringer 10422 (UB); Fazenda Vargem Bonita, ca. 10 km S de Brasília, 4 Feb 1966 (fl), Irwin et al. 12246 (MO, NY); São Bartolomeu, 3 Mar 1980 (fl), Heringer et  al. 3604 (IBGE, MO, NY); Reserva Biológica do IBGE, 3 Mar 1978 (fl, fr), Heringer et al. 386 (IBGE); Reserva Biológica do IBGE, 22 Jan 1987 (fl, fr), Silva & Vianna Jr. 317 (IBGE); Reserva Biológica do IBGE, 5 Mar 1988 (fl, fr), Azevedo 38 (IBGE); Reserva Biológica do IBGE, 15°56′ 47″ S, 47°52′ 35″ W, 28 Apr 1988 (fr), Alvarenga 16 (IBGE); Reserva Biológica do IBGE, 27 Apr 1988 (fl), Mendonça 921 (IBGE); Reserva Biológica do IBGE, 27 Apr 1988 (fl), Mendonça 955 (IBGE, RB); Reserva Biológica do IBGE, 5 May 1988 (fr), Alvarenga 39 (IBGE, RB); Brazlândia, a 5 km Norte, 8 Mar 1982 (fl), Pires 273 (CEN); DF-3, km 17, após a 2ª ponte sobre o rio do Sal, 24 Apr 1996 (fl), Gomes 35 (CEN); DF-3, km 17, após a 2ª ponte sobre o rio do Sal, Gomes 36 (CEN. MO); Gama, Parque do Gama, 17 Mar 1982 (fl), Graham 896 (MO, NY); 16°02′24″ S, 48°03′14″ W, 1 Apr 2004 (fl), Pastore et al. 883 (CEN); Reserva Ecológica do IBGE (RECOR), 6 Feb 1979 (fl, fr), Heringer et  al. 1021 (IBGE); área da aeronáutica, córrego Taquara/Gama, 24 Jan 1990 (fl), Azevedo & Lopes 479 (IBGE, UEC n.v.); próximo à Estação Meteorológica, 15°56′ S, 47°53′

432

13  Systematic Treatment

W, 27 Apr 1993 (fl, fr), Dias 544 (UB); área da aeronáutica, próximo à Estação Meteorológica, 15°57′ S, 47°52′ W, 18 May 1992 (fr), Proença 754 (IBGE); Gama, 1 km W, 9 Apr 1971 (fl, fr), Sastre 1109 (UB); Gama, 24 May 1972 (fl), Ferreira 1481 (HEPH); Guará, 15°48′ S, 47°58′ W, 19 Jan 1994 (fl), Pereira-Silva 2215 (CEN, MO, NY); Guará, 26 Jan 1994 (fl, fr), Pereira-Silva 2225 (CEN, MO); Reserva Ecológica do Guará, 15°48′ S, 47°58′ W, 15 Feb 1994 (fl), Pereira-Silva 2280 (CEN, MO, NY); Guarazinho, 18 Feb 1983 (fl), Figueira 21 (UB); Chapada da Contagem, 31 Jan 1980 (fl), Garcia-Kirkbride 1054 (UB); Chapada da Contagem, DF-001, km 11.3 do Posto Colorado, 12 Jul 1990 (fr), Cavalcanti et al. 500 (CEN, UEC); córrego Barrocão, 15°37′ S, 48°09′ W, 2 Mar 1983 (fl, fr), Kirkbride Jr. 5273 (UB); córrego Carirú, 15°52′ S, 47°47′ W, 6 Feb 1981 (fl, fr), Kirkbride Jr. 3735a (UB); Poço Azul, 22 Jan 2003 (fl), Pastore et al. 263 (CEN); Poço Azul, 4 Apr 2003 (fl), 22 Apr 2003 (fl), Pastore et al. 281 (CEN, MO); São Sebastião, 4 Apr 2003 (fl), Pastore et al. 464 (CEN); divisa Distrito Federal-Goiás, BR-060, 8 Feb 2004 (fl), Pastore & Suganuma 845 (CEN). Goiás, Abadiânia, estrada Abadiânia-Silvânia, 16°13′52″ S, 48°38′52″ W, 26 Mar 2002 (fl), Fonseca et  al. 3370 (CEN, IBGE); Alexânia, cerca de 300  m a oeste da BR 060, na direção de Corumbá de Goiás, 16°08′33″ S, 48°36′12″ W, 18 Feb 2003 (fl), Pereira-Silva et al. 7176 (CEN, MO); Alto Paraíso de Goiás, Chapada dos Veadeiros, entrada para a estrada de terra para a Cachoeira dos Cristais, lado direito, 14°05′11″ S, 47°31′10″ W, 17 Nov 2005 (fr), Cavalcanti et al. 3603 (CEN); Alto Paraíso de Goiás, GO-118, Alto Paraíso-Teresina de Goiás, 31  km de Alto Paraíso, 13°57′03″ S, 47°27′31″ W, 17 Nov 2005 (fr), Cavalcanti et al. 3617 (CEN); Alto Paraíso de Goiás, Portal da Chapada, próximo aos chalés do Portal da Chapada, 18 Jun 2011 (fl), Faria et al. 1275 (CEN, HUEG, UB); Alto Paraíso de Goiás, GO-12, 23 May 1975 (fl), Hatschbach 36737 (MBM, MO); Alta Paraíso de Goiás, 14°02′ S, 47°26′ W, 30 May 1994 (fl), Ratter et al. 7273 (IBGE, UB); Chapada dos Veadeiros, 18 Jul 1964 (fl, fr), Prance & Silva 58184 (K, NY, UB); 12 km de Alto Paraiso/Colinas, Fazenda São Bento, Parcela VII, 14°09′55″ S, 47°35′68″ W, 20 Jul 1994 (fl), Silva et al. 2177 (IBGE); Anápolis, road, 31  km from Brasília, 22 Feb 1965 (fl), Clayton 4813 (UB); Caldas Novas, Serra de Caldas Novas, Termas do Rio Quente, 4 Jan 1979 (fl), Heringer 16664b (IBGE); Cidade Eclética, 50  km de Brasília, 10 Mar 1974 (fl), Heringer 13176 (UB); Corumbá de Goiás, topo do Pico dos Pireneus, 6 km de Cocalzinho, 51º S, 30° W, 14 Jan 1981 (fl), Nogueira et al. 139 (SP, UB); Cocalzinho, 12 Mar 2000 (fl), Serra dos Pireneus, Hatschbach et al. 70003 (MBM); 20 km de Corumbá de Goiás, próximo ao Pico dos Pireneus, 26 Jan 1968 (fl), Irwin et al. 19226 (NY, UB); ca. 15  km  N de Corumbá de Goiás, 16 May 1973 (fl, fr), Anderson 10389 (UB); Corumbá de Goiás, estrada velha da Cidade Eclética para Anápolis, 15°45′ S, 48°18 W, 13 Nov 1990 (fl), Vieira et al. 619 (CEN); Serra do Pireneus, ca. 20 km E de Pirenópolis, 16 Jan 1972 (fl), Irwin et al. 34260 (NY); entre Anápolis e Corumbá de Goiás, 2 Apr 1958 (fl), Lima 58–2994 (K); Cristalina, Serra dos Cristais-Santa Luzia, 17 Jan 1895 (fl), Glaziou 21419 (K, LE n.v., NY, P [2]); entre Formosa e Lagoa Feia, 6 Feb 1895, Glaziou 21418 (K n.v., LE n.v., NY, P); Luziânia, Santo Antonio do Descoberto, Fazenda Boa Viagem, 18 Apr 1985, (fl, fr), Silva 374 (IBGE); Luziânia, Jardim Marajoara, 22 Dec 1990 (fl), Melo & França 421 (CEN);

Descriptions

433

Jardim Marajoara, 9 Feb 1991 (fl), Melo & França 518 (CEN); estrada vicinal de acesso ao rio Descoberto (captação Saneago), km 2, 16°17′17″ S, 48°10′43″ W, 2 Jul 2003 (fr), Pereira Silva et al. 7946 (CEN, MO); ca. 9 km de Luziânia, entrada para a Fazenda São Lucas, 16°17′13″ S, 48°43″ W, 11 Jun 2002 (fl), Fonseca et al. 3440 (CEN, IBGE); Niquelândia, 6  km de Niquelândia, plantação de Eucalipto, 14°25′26″ S, 48°28′19″ W, 28 Apr 1995 (fl), Fonseca et  al. 259 (IBGE); Padre Bernardo, Fazenda Engenho Queimado, 10 km de Brazlândia, 15°43′ S, 48°12′ W, 13 Nov 1990 (fl), Vieira et  al. 594 (CEN, MO); Pirenópolis, Serra dos Pireneus, Fazenda Solar dos Pireneus, 15 Feb 2000 (fl), Hatschbach et  al. 70284 (MBM); Pirenópolis, Serra dos Pireneus, estrada Cocalzinho-Pirenópolis, via Serra dos Pireneus, 8.1 km, 15°47′51″ S, 48°49′45″ W, 14 Nov 2005 (fr), Cavalcanti et al. 3579 (CEN, MO); Planaltina de Goiás, Serra da Biboca, próximo à fazenda Quintas, 15°23′01.6″ S, 47°41′07.5″W, 20 Mar 2003 (fl), Fonseca et al. 4405 (CEN, IBGE); Planaltina, Chapada entre Sobradinho e CPAC, à direita da rodovia, 3 Feb 1983 (fl), Almeida 924 (UB); rio Piripau, SE de Planaltina, 19 Feb 1970 (fl), Irwin et al. 26402 (MO, NY, UB); APA da Cafuringa, Fazenda Palestina, 20 Jul 1992 (fr), Proença & Souza 812 (IBGE); Saia Velha, 15 Feb 1965 (fl, fr), Heringer 10082 (HTO, UB); Santo Antonio do Descoberto, BR-060, 15°55′ S, 48°12′ W, 2 Mar 1992 (fl), PereiraSilva & Pereira 953 (CEN, MO, NY); São João da Aliança, Rodovia GO-118-São João da Aliança, 14°43′ S, 47°31′ W, 6 Mar 1989 (fl), Cavalcanti et al. 373 (CEN); 21 May 2001 (fr), Pereira-Silva et al. 5038 (CEN, MO); a 3 km S de São João da Aliança, 15 Mar 1971 (fl), Irwin et al. 31890 (UB); a 3 km S de São João da Aliança, 16 Mar 1971 (fl), Irwin et al. 31900 (MO, NY); GO-118, km 130, 14°43′ S, 47°31′ W, 14 Apr 1994 (fl), Cavalcanti & Pereira-Silva 1265 (CEN, SPF); a 35 km N de Formosa na estrada para São Gabriel, 28 Mar 1966 (fr), Irwin et al. 14171 (NY, UB, W); córrego das Brancas, próximo à barra do Jacaré, 9 Feb 1994 (fl), Hatschbach et al. 60250 (MBM); Distrito de São Gabriel, GO-118, cerca de 2 km a norte de São Gabriel, 28 May 2001 (fl), Cavalcanti & Pereira-Silva 2722 (CEN, MO); Serra Dourada, 26 Jan 1968 (fl), Eunice et al. s.n. (NY, UB); Silvânia, 16°42′ S, 48°36′ W, 12 Jan 1989 (fl), Filgueiras et al. 1788 (IBGE, SPF); Estação Florestal Experimental, 21 Mar 1989 (fl), Alvarenga et  al. 206 (IBGE, UEC n.v.); without reference to municipal district: Pereira 7402 (HB n.v., M). Minas Gerais, Unaí, Fazenda em Unai, 24 Nov 1986 (fl), Souza s.n. (UB). Diplusodon villosus is characterized by the villous-tomentose indumentum of its branches and flowers and also by its concave bracts, short spreading to deflexed epicalyx segments, prophylls that are shorter than half the length of the floral tube, and 18 stamens. It is frequently confused with D. lanceolatus, which differs by its pubescent indumentum, flowers with longer epicalyx segments ([3–]4–5.5  mm long), and prophylls that surpass the floral tube. Lourteig (1989) also cited D. villosus for the state of Minas Gerais based on Hatschbach & Pelanda 27794; however, this collection is here identified as D. lanceolatus. Diplusodon villosus was described by Pohl (1827a) without citation of specimens in the protologue. In a later publication, Pohl (1827b) cited a single gathering of D. villosus collected in January in the mountains of Ponte Alta, Tocantins State, without mentioning a collection number. This collection was examined at W, and

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the original label corresponds to the locality cited by Pohl (1827b) and is annotated by the author. Also the label indicates Pohl 1066 and Pohl’s diary number D 937. Lourteig (1989) cited this specimen in error under the incorrect number Pohl 137, as the holotype at W, which is nonetheless considered lectotypification (ICBN Art. 9.17, ex. 14). It was verified that other herbaria have specimens of Diplusodon villosus, on which the labels indicate Pohl s.n., which were donated by W. Since duplicates of Pohl’s collections were distributed to other herbaria without the information from his field diary (see “Notes about Diplusodon nomenclature and typifications”) and most lack a collection number, the specimens at BR, K, and M are here regarded as duplicates of Pohl 1066 = D 937. Lourteig (1989) treated Diplusodon humilis Koehne and D. pulchellus Koehne as synonyms of D. villosus. I studied the type material of both taxa, and I agree with both synonymization, given that the material of Glaziou 21422a and Glaziou 21423 corresponds exactly to the characteristics of D. villosus. As for collection Glaziou 21422, two specimens with this number correspond to two different species, to which I added the letters a and b to distinguish them from each other. The specimens Glaziou 21422a are deposited at P and K and correspond to Diplusodon pulchellus, which is a synonym of D. villosus. Glaziou 21422b, on the other hand, deposited at R, corresponds to D. microphyllus, but it probably has a wrong label, because this species does not occur in Goiás. 97. Diplusodon vinaceus T.B.Cavalc. sp. nov. Type: Brazil. Goiás: Campo Alegre de Goiás, ao longo da GO-213, 7  km NE de Campo Alegre, 17o35′48″ S, 47o42′45″ W, 28 Jun 2017 (fl), M. F. Simon, A. Amaral-Santos, M. G. Facco & T.B.Cavalcanti 3079 (holotype, CEN [00101350]; isotype NY). Figs. 13.122, 15.5 D Diplusodon vinaceus is characterized by the bracteose and dark-wine-colored inflorescence with densely arranged flowers. The prophylls are narrow and keeled, the epicalyx segments are thin and short (1.5–2.2 mm long), and the androecium has 12–16 stamens. Due to the leaves ovate and indumentum hirsute, the species resembles D. floribundus, a species whose main diagnostic feature is the presence of the longest epicalyx segments of all species of the genus (7–10 mm long). Subshrubs 0.5–1.5 m tall, erect, with a xylopodium, branched just at inflorescence, glabrescent, not dark-brown when dry. Upper branches cylindrical, unwinged, hirsute, with a dense indumentum of long, white trichomes, trichomes usually reflexed, internodes 2–2.5 cm long. Leaves decussate, not imbricate, spreading, sessile; blades 15–33 × 12–22 mm, coriaceous, not discolorous, acrodromous-basaltypical to supranumerary, ovate, base cordate, apex obtuse, slightly acuminate, margin plane, ciliate, blades glabrescent, hirsute, covered by dense, long, white trichomes, slightly bulbous at the base; secondary veins 3–6 on each side, departing from the base of the midvein, 1–2 along the midvein, prominent on the abaxial face, less evident on the adaxial surface; domatia present. Synflorescence bracteose, botryum to diplobotryum type, strongly congested; accessory branches absent,

Descriptions

435

principal florescence 4–11 cm, paraclades 4–14 cm long; bracts 5–10 × 3–5 mm, ovate, winish, hirsute, trichomes white; bracts 7–10 × 2–6 mm, ovate to narrowly lanceolate, sometimes naviculate; flowers sessile, hypopodium and epipodium absent; prophylls 4–5 × 1–2 mm, reaching the apex of the floral tube, dark-wine in color, narrow-oblong, keeled, base obtuse, apex acute, margin plane, hirsute, ciliate; floral tube 5.5–6  mm long, funnel-shaped, winish even when dry, hirsute, with dense indumentum of long, white trichomes; sepals 4.5–5 mm long, winish, triangular, slightly caudate, hirsute, white trichomes, ciliate with stiff trichomes; epicalyx segments 1.5–2.2 mm long, dark-wine in color, cylindrical, thin, erect, shorter than the sepals, ciliate with spreading, stiff, white trichomes; corolla 2–2.5 cm in diam., rose-magenta to purple, petals 15–17 × 7–9 mm, obovate, apex obtuse; stamens 12–16, free portion of the filaments, 7–10 mm long, slightly exserted; ovary 1.8–2 × 2–2.5 mm, rounded, glabrous, style 8–9 mm long, ovules 24–28. Capsules ca. 4 × 3 mm, globose, glabrous; seeds 8–12, 1.8–2 × 1.8–1.9 mm. Distribution and Ecology (Fig. 13.122)  Diplusodon vinaceus is a rare plant in the region of Campo Alegre, Goiás, and to the present the species is microendemic of this mountainous area of the southeast of Goiás, occurring in stony field of cerrado vegetation at 830 m elevation. Specimens Examined: BRAZIL.  Goiás,  Campo Alegre de Goiás, Serra do Facão, estrada em direção à fazenda Helena, região do córrego Imburuçu e ribeirão Ponte Alta, 12 Jul 2007 (fr), Arantes et  al. 33 (CEN, HUFU); Campo Alegre de Goiás, estrada para rancharia, fazenda de porteira branca, 7 Nov 2007 (fl, fr), Arantes et al. 24 (CEN, HUFU). Diplusodon vinaceus presents the branches of the inflorescence, prophylls, floral tube, sepals, and epicalyx segments dark-wine in color (Fig. 15.5 D) even when dry. In addition, the inflorescence is bracteose with reduced bracts that lie hidden among the flowers. The flowers are densely arranged and have narrow (1–2 mm wide) and keeled prophylls, short (1.5–2.2 mm long) and thin epicalyx segments with stiff and spreading trichomes. The androecium has 12–16 stamens. The entire plant is covered by hirsute and white indumentum; however, the leaves are glabrescent and may present lower density of trichomes when older. Diplusodon vinaceus resembles D. floribundus in the vegetative aspect, due to both species present ovate and hirsute leaves. However, the flowers in D. floribundus are larger, with a floral tube 7–7.5 mm long; unkeeled prophylls of 8–9 × 4–5 mm, surpassing the floral tube; and epicalyx segments 7–10 mm long, the longest found in the genus. 98. Diplusodon virgatus Pohl. Shrubs or subshrubs to small trees, slender or robust when shrubs or subshrubs (0.4–)1.5–4 m tall, erect, much-branched, slender-stemmed, not dark-brown when dry. Upper branches subquadrangular, sometimes with a reduced wing, light-colored, fistulose, lax, glabrous to minutely pilose to pubescent, internodes 1–3  cm long. Leaves decussate, spreading, subpetiolate, rarely subsessile or sessile; petioles

436

13  Systematic Treatment

(when present) to 8 mm long; blades 1.5–7 × 0.5–3 cm, membranaceous to chartaceous or coriaceous, discolorous, eucamptodromous, elliptic, elliptic-lanceolate to oblong or ovate to ovate-lanceolate, base acute to attenuate to obtuse, apex acute or obtuse to retuse, rarely obtuse, margin plane to subrevolute, blades totally glabrous or with sparse trichomes concentrated on the veins, adaxial surface dark-green, abaxial surface clear green; secondary veins 2–3 on each side, departing along the midvein, inconspicuous on both surfaces; domatia absent. Synflorescence frondose, pleiobotryum type, lax; frequently with development of accessory branches, principal florescence 7–15 cm long, paraclades 5–30 cm long; bracts similar to the vegetative leaves in shape, only decreasing in size toward the apex of the branch; flowers delicate or robust, pedicellate, hypopodium 2–5  mm long, epipodium 0.5–1  mm long; prophylls 4.5–7 × 2.5–5 mm, reaching or sometimes surpassing the apex of the floral tube, ovate to obovate, unkeeled, apex obtuse or sometimes acute, margin plane to subrevolute, ciliate or not; floral tube 4–6 mm long, clear green, slightly fleshy or not, funnel-shaped, glabrous; sepals 4–5 mm long, triangular, caudate or not, deflexed or not, glabrous or ciliate; epicalyx segments 2.5–4.5 mm long, linear or lanceolate, dorsiventrally compressed, slightly revolute, deflexed to spreading, glabrous or generally ciliate or with cilia concentrated only on the apex; corolla 2.5–3 cm in diam., white, petals 13–14 × 7–10 mm, oblong to obovate, apex obtuse; stamens 12–14, free portion of the filaments 5.5–7  mm long, slightly exserted, white; ovary 2–2.5  ×  3–3.5  mm, obconic, glabrous, style 9–11  mm long, white, stigma yellow, ovules 38–56. Capsules 4–4.5 × 4–5.2 mm, globose, glabrous; seeds (19–)34–52, 2.5–4 × 2–3 mm. Distribution and Ecology (Fig.  13.122)  Diplusodon virgatus occurs in Distrito Federal, Goiás, Mato Grosso, Minas Gerais, São Paulo, and Tocantins, in cerrado vegetation and along the margins of gallery forests, and in Bolivia, in cerrado areas extending into Bolivia. Diplusodon virgatus is the type species of the genus, designated by Lourteig (1989) because it was used by Koehne (1903) to illustrate the principal characteristics of the genus. In spite of this, D. virgatus is the most distinctive species in the genus. It is a tree that can grow up to 4 m tall, with a well-defined trunk and crown. It occurs in gallery forests and tree cerrado, which are unusual environments for the genus. Morphologically, it has membranous leaves, rather than coriaceous leaves that are typical of Diplusodon, and it is the only species in the genus with a white corolla. The geographic distribution of the species is also unique for Diplusodon. Diplusodon virgatus is widely distributed in six Brazilian states, and it also occurs in Bolivia, whereas most other Diplusodon species are narrow endemics. Key to the varieties.

Descriptions

437

1. Prophylls 2.5–3.5 mm wide, not hiding the entire floral tube; flowers delicate, floral tube thin, 4–4.5 mm long; leaves petiolate, base attenuate; epicalyx segments 2.5–3.5 mm long..…….......................………… D. virgatus var. virgatus 1. Prophylls ca. 5 mm wide, hiding the entire floral tube; floral tube thicker, 5–6 mm long; leaves sessile, base not attenuate; epicalyx segments 3–4.5 mm long….... …….....................................................…….…….. D. virgatus var. occidentalis 98a. Diplusodon virgatus Pohl var. virgatus. Flora 10: 151. 1827. Type: Brazil. Goiás: In campis ad Rio S. Marcos, ad Caretao, Pillar, Trahiras, Arnicuns – in collibus sept. pr. Villa Rica, Minas Gerais, Mayo-­Julio, 1818–1821, J.  B. E. Pohl 773, 1391, 1745 = D 947 (lectotype, W [0053445], here designated; isolectotypes, BR, F, K, M, NY, OXF, W [0053442, 0053443, 0053444]). Fig. 13.121 A–H, 13.122, 15.5 B Friedlandia albiflora Mart. ex DC., Prodr. 3: 94b. 1828, pro syn. Friedlandia amoena Cham. & Schltdl., Linnaea 2: 350. 1827. Diplusodon amoenus (Cham. & Schltdl.) DC., Prodr. 3: 94b. 1828. Type: Brazil. Minas Gerais: na direção para Chapada e pouso alegra, F. Sellow s.n. (first-step lectotype, designated by Lourteig 1989: 217, second-step lectotype, HAL [0098372], here designated; isolectotypes, BR, P, W). Friedlandia leucantha Mart. ex Koehne, Fl. Bras. (Martius) 13(2): 314. 1877, pro syn. Friedlandia myrtifolia Mart. ex Koehne, Fl. Bras. (Martius) 13(2): 314. 1877, pro syn. Shrubs or subshrubs to small trees (0.4–)1.5–4 m tall. Leaves petiolate, rarely subsessile; petioles 3–8  mm long; blades elliptic or elliptic-lanceolate to oblong, base acute to attenuate, apex acute or rarely obtuse, membranaceous to chartaceous, glabrous or with sparse trichomes concentrated on the veins. Flowers delicate, pedicellate, hypopodium 2–5  mm long, epipodium 0.5–1  mm long; prophylls 4.5– 6.5 × 2.5–3.5 mm, reaching the apex of the floral tube, obovate; floral tube 4–4.5 mm long, not fleshy; sepals not caudate, not deflexed; epicalyx segments 2.5–3.5 mm long, spreading to deflexed, lanceolate, dorsiventrally compressed, slightly revolute; ovules 44–56. n = 15 (Graham 1985); 2n = 30 (Cavalcanti 1995). Distribution and Ecology (Fig. 13.122)  Diplusodon virgatus var. virgatus occurs in almost all the contiguous areas of cerrado in Distrito Federal and the states of Goiás, Mato Grosso, Mato Grosso do Sul, Minas Gerais, São Paulo, and Tocantins. There are reports of the species occurring in the states of Rio de Janeiro and Rondônia, although I am unable to verify this. Unlike most other species in the genus, D. virgatus var. virgatus inhabits gallery forest margins and sparse low woods in the cerrado and cerradão, at elevations of 500–1500 m. Flowers and fruits throughout the year. Specimens Examined: BRAZIL. Distrito Federal,  Entre Brasília e Fercal, 30 Jun 1964 (fr), Pires s.n. (NY, UB); Fercal, APA da Cafuringa, a 38 km do Centro

438

13  Systematic Treatment

Fig. 13.121  Diplusodon virgatus var. virgatus and Diplusodon virgatus var. occidentalis. A–H. D. virgatus var. virgatus. A. Branch with inflorescence. B. Section of internode. C. Leaf, abaxial surface. D. Detail of leaf abaxial surface indumentum. E–F. Prophylls, adaxial surface. G. Flower with prophyll, without petals. H. Seed. D. virgatus var. occidentalis. I. Leaf, abaxial surface. (A–D, F Cavalcanti et al. 1266, CEN; E, G–H Pedralli et al. 3318, CEN; I Anderson 9795, UB)

Descriptions

439

Fig. 13.122  Geographic distribution of Diplusodon villosus, D. vinaceus, D. virgatus var. virgatus, and D. virgatus var. occidentalis

440

13  Systematic Treatment

Nacional de Pesquisa de Recursos Genéticos (Embrapa/Cenargen), 9 Aug 1990 (fl, fr), Cavalcanti et al. 571 (CEN, SP); APA Cafuringa, córrego Taquari, 16 Apr 1992 (fl), Melo & França 651 (UB); Fazenda Sucupira, mata do Riacho Fundo, 15°52′ S, 48°00′ W, 3 May 2000 (fr), Guarino & Pereira 131 (CEN, MO);Fazenda Sucupira, 5 May 1994 (fr), Assis et al. 2 (CEN, MO); Brazlândia, Fazenda José Pires, ca. 4 km de Brazlândia, 28 May 1991 (fl), Vieira et al. 774 (CEN, MO, NY); Brasília, 15 May 1966 (fr), Hunt & Ramos 5496 (NY, UB); Brasília, Centro Olimpico, UnB, 2 Jul 1985 (fl, fr), Boiteux s.n. (UB); Catetinho, 15º57′S, 47º59′W, 8 Mar 1965 (fl), Smith A-51 (UB); Catetinho, 12 Apr 1963 (fl), Pires et al. 9047 (NY, UB); Catetinho, 12 Apr 1963 (fl), Santos 1648 & Sacco 1881 (HB n.v., NY); Rio Paranoá, 15º49′S, 47º42′W, 5 May 1968 (fl), Philcox & Onishi 4917 (NY, UB); adjacências da mata após o córrego Cachoeirinha, afluente da margem esquerda do Rio Paranoá, 26 May 1982 (fl, fr), Pereira 268 (IBGE, K, L n.v., MO, US); Córrego Vicente Pires, Fundação Zoobotânica, 18 Apr 1963 (fl), Pires et al. 9225 (NY, UB n.v.); Jardim Botânico de Brasília, 15º52′S, 47°51′ W, 20 May 1987 (fr), Equipe do JBB 918 (HEPH); próximo ao Rio Verde, 15º38′S, 48º21′W, 14 May 1991 (fl), Vieira & Walter 761 (CEN, MO, NY); Perto da DF-2, 15º32′S, 46º50′W, 19 Jun 1980 (fr), Kirkbride & Kirkbride Jr. 1261 (CEN, UB); Chapada da Contagem, a 7  km da BR-010, na rodovia para Brazlândia, BR-080, 15°35′ S, 47°57′ W, 16 Apr 1997 (fl), Cavalcanti & Graham 2258 (CEN, NY); Sobradinho, 15°39′26″ S, 47°54′19″ W, 3 Jun 2004 (fl, fr), Pastore et al. 933 (CEN); Área de Proteção Ambiental da bacia do rio São Bartolomeu, 20 May 1987 (fr), Mendonça 836 (IBGE, UB, UEC n.v.); bacia do rio São Bartolomeu, 15 May 1979 (fl), Heringer et al. 1409 (IBGE, MO); bacia do rio São Bartolomeu, 22 Oct 1979 (fl, fr), Heringer et al. 2547 (IBGE, K, NY, MO, US); bacia do rio São Bartolomeu, 5 Mar 1980 (st), Heringer 3779 (IBGE, P n.v.); bacia do rio São Bartolomeu, 11 Mar 1980 (fl), Heringer et al. 4234 (IBGE); córrego Cabeça de Veado, ca. 9.5 km SE da Torre de TV de Brasília, 8 Mar 1976 (fl), Ratter 2867 (MO, NY, UB); Estação Florestal Cabeça de Veado, ca. 20 km a SE de Brasília, 12 Apr 1983 (fl), Alves 83 (CEN, HEPH); Gama, mata próxima ao rio Gama, 17 Mar 1965 (fl), Smith 15071 (NY); Parque do Gama, 19 Apr 1968 (fr, fl), Fonseca 837 (UB); Parque Municipal do Gama, 16°01′ S, 48°03′ W, 21 Mar 1966 (fl), Irwin et al. 14154 (LE n.v., NY, UB); Parque Municipal do Gama, 25 Mar 1966 (fl), Lima & Cobra 6 (UB); RECOR, picada M-1, 14 Apr 1978 (fl), Heringer et al. 445 (IBGE), P n.v.); Planaltina, Águas Emendadas, 7 Jul 1971 (fr), Ferreira 756 (HEPH); Reserva Biológica de Águas Emendadas, ca. 40  km a NE de Brasília, 15°32′/15°38′ S, 47°33′/47°37′ W, 1984 (fr), Ramos et  al. 4127–1 (HEPH); Planaltina, 1 km Oeste da DF-130, próximo ao córrego Saco dos Pilões, afluente do rio São Bartolomeu, 15°44′ S, 47°41′ W, 20 May 1992 (fr), Walter et al. 1422 (CEN, MO, NY); beira do Corregozinho, à base de um morro com um cruzeiro, 6 km S de Planaltina, 15°40′ S, 47°40′ W, 9 May 1983 (fl), Kirkbride, Jr. 5289 (HPN, INPA, UB); Reserva Biológica de Águas Emendadas, entre 15°32′ – 15°38′ S e 47°33′ – 47°37′ W, 23 May 1982 (fl), Maury 149 (HEPH); Reserva Biológica de Águas Emendadas, 24 Feb 1993 (fl), Munhoz 14 (NY, UB); Reserva Ecológica do IBGE, 15°56′41″ S, 47°53′07″ W, 4 May 1994 (fl), Silva 2207 (IBGE); córrego Bananal, 15°35′ S, 47°55′ W, 19 Apr 1983 (fl), Netto 17 (UB); Estrada para Unaí, 16°02′ S,

Descriptions

441

47°18′ W, 18 Jul 1993 (fr, fl), Proença 873 (UB); DF-100, a 500 m do trevo com a estrada para Barreiras (BR-030), 15°36′ S, 47°22′ W, 5 May 1992 (fl, fr), Dias et al. 44 (CEN, MO); a 93 km da entrada para Unaí, 15°33′ S, 47°25′ W, 12 May 1992 (fr), Dias et al. 115 (CEN, MO); perto do rio Descoberto e matadouro, 5 May 1982 (fl), 15°37′ S, 48°12′ W, Kirkbride, Jr. 4771 (HPN, UB, UFG); bacia do rio Descoberto, caminho para a cidade Eclética, 19 May 1999 (fl, fr), Calago & Silva 128 (CEN, MO); córrego do Reino, beira de mata, 4 May 1972 (fl, fr), Silva 3 (HEPH). Goiás, Abadiânia, BR 060, próximo à divisa com Anápolis, 16°16′53.4″ S, 48°47′38.7″ W, 27 Mar 2003 (bud), Fonseca et al. 4488 (CEN, IBGE); Aparecida de Goiânia, condomínio Rosa dos Ventos, divisa com Hidrolândia, 5 May 2002 (fl), Pastore 26 (CEN, MO); Barro Alto, 9 Jun 1994, Brina s.n. (BHCB 35583); Caldas Novas, alternativa 15, ponto de hidrometria, 27 Apr 1993 (fl), Cordovil-Silva et al. 273 (CEN, MO, NY); Caldas Novas, margem direita do rio Corumbá, próximo ao eixo da barragem, 26 Apr 1994 (fl), Santos et al. 242 (CEN, MO); Caldas Novas, 2 km à montante do eixo da barragem, 26 May 1993 (fl), Santos et al 42 (CEN, MO); Caldas Novas, margem direita do rio Corumbá, na borda da mata, 28 Oct 1993 (fr), Vieira et al. 1777 (CEN); Cocalzinho de Goiás, Parque Estadual da Serra dos Pirineus, 10 Mar 2007 (fr), Bosquetti et al. 685 (CEN, ESA); Colinas do Sul. 2 km da estrada Serra da Mesa-Colinas do Sul, 13°50′ S, 48°17′ W, 12 Dec 1991 (fl), Walter et al. 1079 (CEN, MO, NY); região próxima à foz do rio Tocantinzinho, no rio Tocantins, 13°56′ S, 48°16′ W, 8 Aug 1997 (fr), Walter et al. 3855 (CEN, NY); Caiapônia, Serra do Caiapó, ca. 20 km S de Caiapônia, próximo à rio Claro, 29 Apr 1973 (fl), Anderson 9439 (MO, NY); ca. 40 km S de Caiapônia, 26 Jun 1966 (fr), Irwin et al. 17786 (MO, NY, UB); 50 km S de Caiapônia, na estrada para Jataí, 25 Oct 1969 (est), Irwin & Soderstrom 7331 (NY); Campinaçu, 13º46′S, 48º17′W, 9 Oct 1991 (fr), Cavalcanti et al. 886 (CEN); estrada da Jacira, entrada para Boa Nova, 20 Jun 1995 (fl, fr), Cavalcanti et al. 1432 (CEN, NY); Campinaçu, Alto da Serra Grande, 12 May 1972 (fl), Rizzo 8093 (UFG); Campinaçu, ao longo da estrada de acesso à Fazenda Aroeira, 25 Apr 1995 fl, Santos et al. 445 (CEN); Corumbá de Goiás, Serra dos Pireneus, ca. 15 km N de Corumbá de Goiás, 16 May 1973 (fl), Anderson 10465 (MO, NY); Corumbá de Goiás, Fazenda Burita dos Alves, 11 Sep 1990 (fl), Vieira et al. 469 (CEN); encosta a 5 km do rio dos Macacos, 15°45′ S, 48°18′ W, 13 Nov 1990 (fl), Vieira et al. 611 (CEN, MO); Corumbaíba, margem esquerda do rio Corumbá, 1.5 km a montante do eixo da barragem, 25 May 1993 (fl), Santos et al. 1 (CEN, MO, NY); 1 km acima da S.O. 4, margem esquerda do rio Corumbá, 22 Jun 1993 (fl, fr), Cordovil-Silva et al. 319 (CEN); Cristalina, estrada de acesso à área de empréstimo, cerca de 600 m da jazida, 16°12′38″ S, 47°20′09″ W, 7 Mar 2002 (fl), Pereira-Silva et al. 6092 (CEN); BR-050, Cristalina-Ipameri, km 31, às margens do rio Veríssimo, 12 Mar 1989 (fr), Cavalcanti et al. 417 (CEN, MO, NY, SPF); rod. BR-020, Lageado, 12 Apr 1981 (fl), Hatschbach 43828 (MBM, MO); Formosa, margem esquerda da Lagoa Perta Pé, área do exército, 15°59′ S, 47°11″50″ W, 20 Mar 2002 (fl), Pereira-Silva et  al. 6258 (CEN, MO); Serra do Morcego, córrego Estrema, ca. 35 km NE de Formosa, 22 Apr 1966 (fl), Irwin et al. 15266 (MO, NY, UB); Fazenda J.  Teles, ca. 10  km NW de Formosa, próximo à ponte do rio Paranã, 29 Apr 1966 (fl), Irwin et al. 15479 (MO, NY, UB); 35 km N

442

13  Systematic Treatment

de Brasília, rio Contagem, 8 May 1966 (fl), Irwin et  al. 15725 (NY, UB); BR 020-Fazenda Bambuzal, entrada no km 35, para o Buraco das Araras, 15°21′42″ W, 47°09′2.8″ W, 19 Mar 2003 (fl), Fonseca et  al. 4348 (CEN, IBGE); Formoso, Formoso para Campinaçu, Alto da Serra Grande, 13 Apr 1972 (fl), Rizzo 8017 (UFG); Goianápolis, Parque Estadual Altamiro de Moura Pacheco, 16°32″29.8″ S, 49°08′24.9 W, 12 Apr 2005 (fl, fr), Fonseca et al. 5810 (CEN); Goianira, margem esquerda da estrada que demanda a Fazenda Louzanira, 18 Apr 1970 (fl), Rizzo 4994 & Barbosa 4242 (OUPR); a 2 km da margem esquerda do rio Meia Ponte, na Fazenda Louzandira, 18 Apr 1970 (fl), Rizzo 4967 & Barbosa 4215 (UFG); a 2 km da margem esquerda do rio Meia Ponte, na Fazenda Louzandira, 16 May 1970 (fr), Rizzo 5153 & Barbosa 4402 (UFG); de Goianira a Leopoldo Bulhões, 18 km de Goiânia, 16 May 1968 (fl), Rizzo & Barbosa 754 (UFG); Ipameri, margem esquerda do rio Corumbá, 21 km da ponte que liga Ipameri à Cladas Novas, 27 May 1993 (fl, fr), Santos et al. 63 (CEN, MO, NY); Leopoldo de Bulhões, 16°34′ S, 48°43′ W, 20 Mar 1989 (fl), Mendonça et al. 1314 (IBGE, P n.v., UB n.v.); Luziania, sítio do Dr. Jesus Reis, 26 Feb 1975 (fl), Heringer 14429 (IBGE, P n.v.); Luziania, sítio do Dr. Jesus Reis, 12 Mar 1981 (fl, fr), Heringer 18307 (IBGE, K n.v., L n.v., MO, US n.v., MG n.v.); estrada Alexânia-Silvânia, km 37, 16°17′16″ S, 48°26′32″ W, 11 Apr 2003 (fl), Pereira-Silva et al. 7561 (CEN, MO); À direita da GO-9, para Nerópolis, 15 km de Goiânia, 17 Apr 1968 (fl), Rizzo & Barbosa 497 (UFG); Minaçu, fazenda Cigana, próximo ao rio Tocantins, 13°38′27″ S, 48°08′31″ W, 19 Apr 2002 (fl), Pereira-Silva et al. 4990 (CEN, MO). Morrinhos, estrada Morrinhos para Caldas Novas, córrego Samambaia, 25 Apr 1970 (fl), Rizzo 5089 & Barbosa 4338 (UFG); Morrinhos, 25 Apr 1970 (fl), Rizzo 5093 & Barbosa 4342 (UFG); Niquelândia, estrada de terra para Colinas, ca. 4 km de Niquelândia, 14°26′59″ S, 48°21′17″ W, 28 May 1996 (fl), Silva & Jesus 2937 (CEN, IBGE); nas margens do rio Tocantinzinho, 13°59′ S, 48°18′ W, 22 Jul 1995 (fl, fr), Cavalcanti et al. 1593 (CEN, MO, NY); estrada de terra vicinal à GO-237, (Niquelândia-Colinas), entrada 0.6 km da ponte sobre o rio Bagagem, a 7 km desta, bacia de inundação da UHE Serra da Mesa, 14 Apr 1992 (fl), Walter et al. 1209 (CEN, F, K, MBM, UB, MO); a 32 km desta, bacia de inundação da UHE Serra da Mesa, 14 Apr 1992 (fl, fr), Walter et al. 1302 (CEN, NY, SPF); estrada de terra vicinal à GO-237, Niquelândia-Colinas, entrada a 600 m da ponte sobre o rio Bagagem, a 7 km desta, 14°20′ S, 48°12′ W, 14 Apr 1992 (fl), Walter et  al. 1269 (CEN); Fazenda Engenho, ca. 11  km de Niquelândia/Dois Irmãos, 27 Jan 1997 (fl), Fonseca et  al. 1468 (CEN, IBGE); Niquelândia, Fazenda Serra Negra, (Niqueltocantins), arredores da Serra Negra na estrada para o rio Bagagem, 14º02′S, 48º18′W, 8 Jun 1992 (fl, fr), Walter et al. 1439 (CEN, RB); região da Serra Negra, margem esquerda do rio Bagagem, próximo à fazenda Aroeira, após a CODEMIN, ca. 10 km desta, 7 Oct 1995 (fr), Walter et al. 2698 (CEN, MO); rio Paranã, 35 km N de Formosa, 30 Mar 1966 (fl), Irwin et al. 14290 (LE n.v., NY, UB); São João da Aliança, GO-118, km 130, 14 Apr 1994 (fl), Cavalcanti & Pereira-Silva 1266 (CEN, F, K, MBM, NY, SPF, UB); 12 km S de São João da Aliança, 19 Mar 1982 (fl), Graham 906 (MO, NY); 15 km S de São João da Aliança, 21 Mar 1973 (fl, fr), Anderson 7551 (UB, MO, NY); Padre Bernardo, Chapada da Vargem Grande, cerca de 30 km de Brazlândia, 15º33′S, 48º15′W, 7

Descriptions

443

May 1991 (fl), Vieira & Pereira 741 (CEN, NY); Piranhas, 6 km NW de Piranhas, 24 Jun 1966 (fr), Irwin et al. 17699 (MO, NY, UB); Pirenópolis, estrada CocalzinhoPirenópolis, via Serra dos Pireneus, 13.4 km, 15°54′30″ S, 48°51′44″ W, 14 Nov 2005 (fr), Cavalcanti et al. 3587 (CEN); São João da Aliança, margem direita do Rio Jatobá, 21  km ao norte de sâo João da Aliança, estrada vicinal a esquerda, 14°32′03 S, 47°30′51″ W, 3 Jul 2015 (fr), Cavalcanti & Pereira-Silva 3984 (CEN); Serra do Facão, 3 Apr 2008 (fl), Arantes et al. 943 (HUFU); without reference to municipal district: 1841, Gardner 3727 (NY); Gardner 3729 (NY); Gardner 4134 (BM n.v., NY); entre Brasília e Niquelândia, 10 May 1963 (fl, fr), Pires et al. 9672 (UB); rodovia Anápolis-Brasília, 15 May 1965 (fl), Heringer 10429 (NY, UB). Mato Grosso, Buriti, 27 Mar 1927 (fr), Collenette 117 (NY); Cuiabá, rodovia Cuiabá-Chapada dos Guimarães, Cachoeirinha, desvio à direita em direção à Casa de Pedras, 5 km, 18 May 1986 (fl), Saddi 7646 (UFMT); Cuiabá, Martius Herb. Fl. Bras. n. 866 (M [2], MO, NY [2]), Riedel s.n. (M, MO, NY, OXF [2]); Torixoréu, proximidades de Diamantino, s.d., Macedo et al. 1062 (UFMT). Guiratinga, margem norte do arroio Piauí pouco ao norte de Guiratinga, 16°18′00′′ S 53°46′ 00′′ W, 24 Apr 1995 (fl), Valls et al. 13684 (CEN). Minas Gerais, Aiuroca, PCH Aiuroca, rio Aiuroca, 20 May 2000 (fl), Tameirão-Neto 2951 (BHCB); Baependi, próximo à Cachoeira do Gamarra, 28 Mar 2002 (fl), Forzza et  al. 2129 (CESJ); Toca dos Urubus, 3 May 2002 (fl), Ferreira & Ferreira 45 (CESJ); Barbacena, 28 Mar 1964 (fl), Trinta 574 & Fromm 1650 (HB n.v., NY); Belo Horizonte, campus da UFMG, Estação Ecológica, trilha das Flores, 30 May 1990 (fr), Tameirão-Neto 54 (BHCB, CEN); Morro das Pedras, 25 Mar 1945 (fl), Williams & Assis 6491 (GH n.v., NY); Morro das Pedras, 24 May 1945 (fl), Williams & Assis 7044 (GH n.v., MO, SP); Campus-Veterinária, 27 Mar 1981 (fl), Grandi 622 (ESA); Serra do Curral, 2 May 1953 (fl), Roth 2232 (SPF, UFJF n.v.); Botumirim, Serra da Limeira, na estrada para Itacambira, 14 Jun 1991 (fl, fr), Mello-Silva et al. 496 (CEN, SPF); Bueno Brandão, Serrinha, 22°28′28″ S, 46°22′09″ W, 27 Mar 1999 (fl, fr), Stehmann et  al. 2439 (BHCB, CEN); Caldas, Regnell 142 (NY); Congonhas do Norte, estrada para Costa Senna, 18°45′04″ S, 43°40′43″ W, 1 Mar 1998 (fl), Pirani et al. 4084 (CEN, SPF); Congonhas do Norte, 5–10 km N, 20 May 1989 (fl, fr), Hatschbach et al. 52969 (MBM n.v., MO); Carangola, Ponte Alta, Apr 2004, Leoni & Silva 5725 (GFJP, CEN); Corinto, Fazenda Diam.ante, córrego Diam.ante, 9 Apr 1931 (fl), Mexia 5574 (MO, NY); Datas, 17  km SE da interseção da estrada Diamantina-Curvelo com a estrada para Serro, 13 Mar 1982 (fl), Graham 878 (NY [2]); 5 km pela rodovia NE de rio Manso e Couto Magalhães de Minas, 13 Apr 1973 (fl), Anderson 8742 (MO, NY); Diamantina, Barão, base da Serra do Capão, 21 May 1930 (fr), Mexia 5897 (MO, NY); entre Rios de Minas, Pedra Branca, 29 Jan 1977 (fl, fr), Krieger 14546 (CESJ n.v., SPF); Formoso, Parque Nacional Grande Sertão Veredas, margens do rio Mato Grande, 15°19′6″ S, 45°59′9″ W, 29 Apr 1999 (fl), RodriguesSilva et al. 267 (CEN, IBGE); Gouveia, rod. BR-259, ponte córrego do Tigre, 20 Mar 1987 (fl), Hatschbach 51136 (MBM, MO); Guarapuava, 16°02′ S, 46°18′ W, 26 Nov 2000, Milhomens et  al. 62 (CEN, UB); Ibiá, MG-230, distante 2  km da BR-262, 2 Mar 1952 (fl), Walter et  al. 60 (IBGE, P n.v., UB n.v., UEC n.v.); Indianópolis, fazenda Bela Tanda and neighbouring fazendas, 6 km de Indianópolis,

444

13  Systematic Treatment

3 Mar 1986 (fl, fr), Gottsberger & Doring 137–3386 (CEN); Itabirito, Serra do Itabirito, 25 Apr 1980 (fl), Badini s.n. (OUPR 25529); Pico do Itabirito, 1 Feb 1995 (fl), Teixeira s.n. (CEN 18948, BHCB 26302); Ituiutaba, 10 Apr 1948 (fl), Macedo 1048 (NY); Itutinga, 9 km W de Itutinga ao longo da MG-265 para Barbacena, 27 Feb 1976 (fl), Davidse & Ramamoorthy 10767 (MO, NY); Jaboticatubas, 10 km N de Lagoa Santa, 28 Apr 1952 (fl), Smith 6954 (R, US n.v.); Jacuí, estrada de terra Jacuí-São Sebastião do Paraíso, 21°05′35″ S, 46°43′04″ W, 13 Jan 2003 (st), Cavalcanti et al. 3062 (CEN, MO); Lagoa Santa, 19º40′S, 43º55′W, 30 Apr 1952 (fl), Smith et al. 6734 (NY, R, US n.v.); Lavras, 3 Mar 1939 (fl), Heringer 191 (CEN, ESAL, SP); Serra da Bocaina/Poço Bonito, 3 Apr 1987 (fl), Carvalho et  al. s.n. (ESAL 6735, SPF); Serra de Lavras, 11 km S de Lavras, caminho a Poço Bonito, 21º 20′S, 45º W, 11 May 1990 (fr), Arbo et  al. 3883 (CEN, CTES n.v., SPF); Mariana, Santa Rita Durão, 10 Apr 1973 (fl), Badini s.n. (OUPR 22587); fazenda dos Macaquinhos, 5 Apr 1974 (fl, fr) Badini s.n. (OUPR 3727); entre Mariana e Bento Rodrigues, 9 Apr 1972 (fl), Badini s.n. (OUPR 2663); entre Mariana e Bento Rodrigues, 9 Lisboa s.n. (CEN 38140, OUPR 1817); ca. 3 km N de Mariana, 3 Feb 1971 (fl), Irwin et al. 29695 (NY); Morro Feio, 5 km N de Hidrolândia, 16°55′ S, 49°14′ W, 12 Apr 1988 (fl, fr), Brooks et al. 36 (MO, NY, UFG n.v.); Nazareno, 1990, Teixeira & Brina s.n. (BHCB 35831); Nova Lima, São Sebastião das Águas Claras, Macacos, 3 Apr 1981 (fl), Martins s.n. (CEN, 19056, BHCB 8457, ESA); Nova Lima, estrada para Barão de Cocais, sobre a Serra da Garandela, a 5 km do entroncamento do rio Acima, 20°06′11″ S, 43°45′10″ W, 9 Mar 2014 (fl), Cavalcanti & Pereira-Silva 3791 (CEN); Ouro Branco, 14 May 1988 (fl), Júlio s.n. (BHCB 12569, CEN 19117); vila do Itatiaia, 19 Feb 1977 (fl), Badini & Lisboa s.n. (OUPR 4960); Ouro Branco, estrada entre Ouro Branco e o Morro do Gabriel, ca. 9 km de Ouro Branco, 20°29′20.6″ S, 43°39′49″ W, 8 Mar 1995 (fl, fr), Souza et al. 7912 (CEN, ESA); Vila da Itaparica, 19 Feb 1977 (fl, fr), Lisboa s.n. (CEN 38145, OUPR 4763); Paraopeba, Reserva do Horto Florestal, 14 Jun 1954 (fl), Heringer 3468 (UB); Paraopeba, Reserva do Horto Florestal, 4 Apr 1965 (fl), Paula 32 (CEN, NY, SP); Paula 40 (CEN, SP); Passa Tempo, estrada de Morro do Ferro para Passa Quatro, 18 km de Morro do Ferro, Serra das Vertentes, 21°13′ S, 44°54′ W, 26 Feb 2000 (fl), Cavalcanti et  al. 2567 (CEN, MO, NY); Ouro Preto, Antonio Pereira, May 1943 (fl), Badini s.n. (OUPR 4332); base do Pico do Itacolomi, 20o24′S, 43º 40′W, 13 May 1990 (fl, fr), Arbo et al. 3992 (CEN, CTES n.v., MO, SPF); Bailão, 21 Apr 1943 (fl), Badini s.n. (OUPR 4230); Capanema, May 1943 (fl), Badini s.n. (OUPR 4331); Salto, 22 Apr 1966 (fl, fr), Badini s.n. (OUPR 5090); Porto de Santa Cruz, 22 Apr 1966 (fl, fr), Reis 5091 OUPR; Falcão, 25 Apr 1966 (fl, fr), Reis 5097 (OUPR); Patos de Minas, 40  km O da cidade, 16 May 1963 (fl, fr), MendesMagalhães 19026 (HB n.v., NY); Paraopeba, EFLEX - Paraopeba, 25 May 1994 (fl, fr), Paula 3461 (UB); Patrocínio, BR-365, Fazenda Chalé, 18°55′57″ S, 47°08′26″ W, 1 Mar 1989 (fr), Mendonça et  al. 1244 (IBGE, P n.v., SPF, UB, UEC n.v.); Patrocínio, BR-365, Fazenda Chalé, Mendonça et al. 1259 (IBGE, P n.v.); Perdões, rio Grande, 26 Apr 1997 (fl, fr), Vasconcelos s.n. (BHCB 37892, CEN 33612); Perdões, rio Grande, 29 May 1994 (fr), Tameirão-Neto & Werneck 1020 (CEN, BHCB); Pouso Alegre, 30 Apr 1927 (fl), Hoehne s.n. (CEN 29272, SP 19286); Rio

Descriptions

445

Manso, 5 km pela rodovia NE de Rio Manso e Couto Magalhães de Minas, 13 Apr 1973 (fl), Anderson 8742 (NY); Ritápolis, estrada para Ritápolis, no entroncamento para a entrada da cidade, predomínio de gramíneas, 21,0430 S, 44,3231 W, 4 May 2012 (fl), Delfini et al. 62 (CEN, ESA); Santana do Riacho, estrada para Cardeal Mota, via Melo, 9.9  km da bufurcação para a usina Coronel Américo Teixeira, 19°13′59″ S, 43°39′58″ W, 23 Apr 2006 (fl), Mello-Silva et al. 2915 (CEN, SPF); São João del Rei, 23 Apr 1888 (fl), Glaziou 12421 (P n.v.); São João del Rei, 23 Apr 1888 (fl), Glaziou s.n. (NY [2]); São João del Rei, 26 Mar 1970 (fl), Krieger 8283 (CEN, UFJF); São Roque de Minas, Serra da Canastra, Mar 1991 (fl), Faria s.n. (CEN, SPF); São Roque de Minas, estrada para Parque Nacional da Serra da Canastra, cerca de 2 km antes da portaria São Roque de Minas, 16 May 2012 (fl), Delfini et al. 497 (CEN, ESA, RB); São Sebastião do Paraíso, 20°55′58″ S, 46°49′57″ W, 29 Apr 2003 (fl, fr), Cavalcanti et  al. 3179 (CEN, MO); Tapina, Triângulo Mineiro, Aug 1988 (fr), Souza s.n. (BHCB 17131, CEN 19103); Três Corações, rodovia Três Corações-São Tomé das Letras, 4 Feb 1973 (fl), Hatschbach & Ahumada 31244 (CEN, MBM, NY); Uberlândia, posto de piscicultura, 19 Feb 1981 (fl), Araújo 2 (CEN, HUFU, NY); Reserva Ecológica Panga, 27 Mar 1987 (fl), Araújo 233 (CEN, HUFU, NY); vereda 3, 26 May 2000 (fl), Barbosa 3242 (CEN, HUFU); Reserva Ecológica Panga, 2 Jun 2000 (fr), Barbosa s.n. (CEN 38150, HUFU 23429); Clube de Caça e Pesca, 13 Aug 1993 (fr), Arantes 39 (CEN, HUFU); Reserva Ecológica Panga, 19 Apr 1994 (fl, fr), Arantes & Marra 180 (CEN, HUFU); Unaí, cerca de 1  km à montante do eixo da barragem, margem direita do rio Arrependido, 16°12′43″ S, 47°16′40″ W, 27 Mar 2002 (fl), Pereira-Silva et al. 6306 (CEN, MO); fazenda Experimental da Glória, 16 Mar 1994 (fl), Nakajima et al. 237 (CEN, HUFU); rodovia Uberlândia-­Araguari, 29 Apr 1994 (fl, fr), Kerr s.n. (CEN 38159, HUFU 5873); without reference to municipal district: Sabara et Ouro Preto, 1907, Glaziou 13818 (LE n.v., NY, P); Claussen 312 (NY [2]); 1840, Claussen s.n. (NY, OXF, Vila Rica e Sabará, Martius obs 1379 (M [3]). São Paulo, Águas da Prata, 30 Apr 1941 (fl), Viegas s.n. (CEN 25498, SP 48763); Aracariguana, Morro da Voturuna, 9 Mar 1989 (fl), Macedo s.n. (ESA); Baruerí, 29 Mar 1929 (fl), Hoehne 16 (SP); Baruerí, 30 Apr 1941 (fl), Viegas 6252 (ESA); Brotas, NW da interseção da estrada para Campo Alegre com a Brotas-Tirapina, 22º17′S, 47º56′W, 16 Jun 1961 (fr), Eiten et al. 2973 (NY, SP); Buritizal, Serra do Buritizal, 20°12′26″ S, 47°45′22″ W, 27 Jul 1994 (fr), Barreto et al. 2725 (CEN, ESA); Buritizal, Serra do Buritizal, 14 Apr 1981 (fl, fr), Leitão-­Filho et al. 14491 (CEN, UEC); Caieiras, Campo Alto, 4 May 1942 (fl, fr), Kuhlmann & Kuhn 1132 (CEN, SP); Conchal, Estação Experimental do Instituto Florestal do Estado de São Paulo, 11 May 1983 (fl), Castañeda & D’Affonseca s.n. (CEN 34035); Corumbataí, Reserva de Cerrado de Corumbataí, 18 May 1977 (fl, fr), Marinis & César 18 (CEN, SPF); Reserva de Cerrado da UNESP, 11 May 1983 (fl, fr), Saraiva 19 (CEN, HRCB); Reserva de Cerrado da UNESP, 26 Mar 1985 (fl, fr), Campos 87 (CEN, HRCB); Itapira, 12 May 1927 (fl), Hoehne s.n. (CEN 29257, SP 20258); Itirapina, 22°14′22″ S, 47°48′14″ W, 26 Apr 1994 (fl), Barreto et al. 2314 (CEN, ESA); Jaraguá, Apr 1912 (fl), Luederwaldt s.n. (CEN 29198, SP 14229); Jaraguá, 21 Apr 1921 (fl), Hoehne 5473 (SP); Pico do Jaraguá, Taipas, 30 Apr 1939 (fl), Hashimoto 57 (SP); Jundiaí, 4

446

13  Systematic Treatment

Apr 1915 (fl), Brade 5663 (CEN, SP); Jundiaí, 5 Apr 1939 (fl, fr), Viegas s.n. (SP 42029); Itu, 15 Dec?, (fl, fr), Russel 322 (CEN, SP); Magda, Fazenda CFM, 17 May 1995 (fl, fr), Bernacci et  al. 1792 (CEN, IAC n.v.); Moji-Guaçu, 25  km NW de Moji-Guaçu, Fazenda Campininha, 4 Feb 1977 (fl), Gibbs & Leitão-Filho 4353 (IBGE, NY, RB, UEC n.v.); Fazenda Campininha, Campo das Sete Lagoas, 5.3 km NW de Pádua Sales, 14 Apr 1961 (fl), Eiten & Eiten 2586 (CEN, MO, SP, UB n.v.); Fazenda Campininha, 23 May 1961 (fl, fr), Mattos 9004 (CEN, SP); Fazenda Campininha, 24 May 1965 (fl, fr), Mattos 12242 (SP); Fazenda Campininha, 24 May 1965 (fr), Mattos 12280 (CEN, SP); Fazenda Campininha, 12 Feb 1980 (fl, fr), Melo & Chiea 194 (CEN, NY, SP); Fazenda Campininha, Melo & Chiea 195 (SP); Fazenda Campininha, 9 Apr 1980 (fl), Mantovani 654 (CEN, SP); Fazenda Campininha, 27 Apr 1981 (fl, fr), Mantovani & Suagyiama 1829 (CEN, SP); Martinho Prado, Reserva Biológica da Fazenda Campininha, 27 Jan 1981 (fl, fr), Sugyiama & Mantovani 12 (SP); Martinho Prado, Reserva Biológica da Fazenda Campininha, 12 Feb 1981 (fl, fr), Mantovani 1676 (CEN, SP); Martinho Prado, Reserva Biológica da Fazenda Campininha, 26 May 1981 (fr), Mantovani 1365 (CEN, SP); Martinho Prado, Reserva Biológica da Fazenda Campininha, Mantovani 1865 (SP); Martinho Prado, Reserva Biológica da Fazenda Campininha, 27 Apr 1981 (fl, fr), Sugyiama & Mantovani 163 (CEN, NY, SP); Martinho Prado, Reserva Biológica da Fazenda Campininha, 19 Mar 1985 (fl), Cerati 158-A (SP); Mococa, na estrada de Casa Branca, Mococa, 21 May 1953 (fl, fr), Dedecca 388 (UB); MojiMirim, km 150 da estrada de rodagem, 10 Feb 1946 (fl), Viegas & Toledo 8045 (SP); Parnaíba, beira do rio Tietê, Apr 1901 (fl), Edwall 5772 (CEN, SP); Perus, 25 Mar 1907 (fl), Usteri s.n. (CEN 29196, SP 14225); Pedregulho, Parque Estadual das Furnas do Bom Jesus, 20°11′50″ S, 47°25′10″ W, 18 Apr 2003 (fl), Sasaki et al. 425 (CEN); Pirassununga, Emas, 1 Apr 1942, Rawitscher s.n. (SPF 147413); Pirassununga, cerrado de Emas, 22°02′ S, 47°30′ W, 4 May 1994 (fr), Batalha & Mantovani 103 (CEN, SP); Pirassununga, sítio Bom Retiro, 26 Mar 1947 (fl), Pickel s.n. (NY, SPSF); Pirassununga, cerrado de Emas, 13 Apr 1977 (fl, fr), Kirizawa 108 (CEN, SP); Rio Claro, 25 May 1988 (fr), Loefgren 533 (SP); São Carlos, 6 Apr 1980 (fr), Linhares 11129-A (CEN, UEC n.v.); São Paulo, via Anhanguera, km 30, 11 May 1949 (fl, fr), Hoehne s.n. (CEN 25962, SPF 12247); próximo ao Jaraguá, 10 Mar 1942 (fl, fr), Hoehne s.n. (CEN 25959, SPF 11038); a 2 km oeste de Jeriquara, Mattos & Bicalho 11495 (CEN, SP); Butantã, 10 Apr 1917 (fl, fr), Hoehne 7 (SP); Butantã, 10 Apr 1917 (fl), Hoehne s.n. (CEN 29260, SP); Butantã, 3 Mar 1938 (fl), Campos 17 (CEN, SP); a 2  km oeste de Jeriquara, 16 Mar 1964 (fl), Mattos & Bicalho 11496 (SP); without reference to municipal district: Campina Verde, 29 Jun 1943 (fr), Macedo 57 (SP). Tocantins, Xambioiá, Piraquê, próximo ao rio Lontra, na Fazenda Baixa, 15 May 1974 (fl), Rizzo 9838 (UFG); Colméia, Pequizeiro, a 9 km de Pequizeiro em direção à Conceição do Araguaia, 8º32′S, 49º01′W, 24 May 1990 (fl, fr), Pedralli et al. 3318 (CEN, NY); do entroncamento da Belém-Brasília com a Tranzamazônica, direção a Araguatins, a 80  km, 17 May 1974 (fl), Rizzo 9845 (UFG).

Descriptions

447

Diplusodon virgatus var. virgatus has a slender habit, long-attenuate petiolate leaves, and more floriferous, divergent branches than D. virgatus var. occidentalis. Diplusodon virgatus was described by Pohl (1827a) with no collection cited in the protologue. In a subsequent publication, Pohl (1827b) cited several gatherings collected from 1818 to 1821, near Vila Rica (now Ouro Preto), Minas Gerais, and five localities in Goiás State, without mentioning collection numbers. With direct examination of Pohl’s collections at W, it was observed that for D. virgatus, Pohl assigned a series of collection numbers to one large set of the same species, rather than to individual collections. Specimens of D. virgatus collected at different times and in different places (in Goiás and Minas Gerais state) have one single label with Pohl’s field annotations that provides three numbers, Pohl 773, 1391, and 1745, and the corresponding number from his diary, D947, to the same set of specimens without exact localities. The specimen “Pohl 773, 1391, 1745 = D 947” with an original label is here selected as the lectotype of this taxon (W0053445). Friedlandia amoena was described by Chamisso and Schlechtendal (Sellow s.n.). Lourteig considered the type destroyed at B, and chose a lectotype at P, considered as the first-step lectotype (ICBN Art. 9.17, ex. 14). There is a specimen of Sellow s.n. at HAL with an original label annotated by Schlechtendal. This specimen is here designated as the lectotype of F. amoena (HAL 0098372). 98b. Diplusodon virgatus var. occidentalis T.B.Cavalc. & S.Graham, Novon 15(1): 67. 2005. Type: Bolivia. Santa Cruz: Velasco, Parque Nacional Noel Kempff Mercado, Campamento Huanchaca I, 13°53′55″ S, 60o48′46″ W, 17 May 1994, L. Arroyo, B. Mostacedo, H. Gonzales, S. Cabrera & J. Surubí 684 (holotype, MO [2272371]; isotype, USZ n.v.). Figs. 13.121 I, 13.122 Robust shrubs, 0.7–2 m tall. Leaves sessile; blades coriaceous, ovate to ovatelanceolate, base obtuse to slightly acute, not attenuate, eucamptodromous, apex obtuse to retuse, glabrous. Flowers robust, pedicellate, hypopodium ca. 2 mm long, epipodium absent; prophylls 5–7 × ca. 5 mm, reaching the apex of the floral tube, hiding the floral tube, ovate; floral tube 5–6 mm long, slightly fleshy; sepals caudate, apex deflexed; epicalyx segments 3–4.5 mm long, deflexed, linear-oblong, not revolute; ovules 38. Distribution and Ecology (Fig. 13.122)  Southeastern Bolivia, also northeastern Mato Grosso and northern Mato Grosso do Sul, Brazil, in woody cerrado adjacent to mata ciliar, 500–650 m. Flowers from March to May; fruits registered from April to May. Specimens Examined: BRAZIL. Mato Grosso,  Barra do Garças, ca. 9 km NE of Barra do Garças, 6 May 1973, Anderson 9795 (MO, NY, UB); Coxim, rod. Campo Grande-Cuiabá, 16 May 1973 (fl), Hatschbach 31981 (MBM, MO); Diamantino, Fazenda Pequeno Figueiredo, cerca de 4  km em linha reta de Diamantino, 14°20′ S, 56°27′ W, 22 May 1997, Souza et al. 16844 (CEN, ESA); Juruena, Apr 1909 (fl), Hoehne 1796 (R); Juruena, Apr 1909 (fl), Hoehne 1798 (R);

448

13  Systematic Treatment

Juruena, May 1909 (fl), Hoehne 1846 (R); Nova Ubiratã, estrada para o rio Teles Pires, 13°34′60″ S, 55°16′10″ W, 1 May 1997 (fl), Nave et al. 1415 (CEN, HUEFS); Tapurah, estrada do Capixaba, cerca de 20 km de Tapurah, 12°37′ S, 56°22′ W, 12 Jun 1997, Souza et al. 17830 (CEN, ESA); Pontes e Lacerda, Serra da Santa Bárbara, 40  km em vicinal na MT-473, 28  km ao sul de Pontes e Lacerda, 15°39′30″ S, 59°21′57″ W, 24 Mar 2014 (fl), Simon et al. 2333 (CEN). Mato Grosso do Sul, Paranaíba, fazenda Nova Ponte, 19°18′16.5″ S, 51°05′21″ W, 16 Apr 2004 (fl, fr), Sciamarelii et al. 1299 (CEN, CGMS); Paranaíba, fazenda Nova Ponte, 19°18′16.5″ S, 54°21′51.4 W, 16 Apr 2004 (fl, fr), Jacques 1299 (CEN, HMS); Selúria, Fazenda de Ensino e Pesquisa da Unesp, Campus Ilha Solteira, 22 Apr 1991 (fl), Santos 370 (UB); Selúria, Fazenda de Ensino e Pesquisa da Unesp, Campus Ilha Solteira, 5 Apr 1991 (fl), Tiritan 584 (UB); Selúria, Fazenda de Ensino e Pesquisa da Unesp, Campus Ilha Solteira, 7 May 1991 (fl, fr), Santos 413 (UB). BOLIVIA.  Santa Cruz,  Velasco, Parque Nacional Noel Kempff Mercado, Campamento Huanchaca I, 13°53′55″ S, 60o48′46″ W, 17 May 1994, Arroyo et al. 684 (MO; USZ n.v.). Diplusodon virgatus var. occidentalis is a robust shrub with more coriaceous, glabrous leaves and fewer, more robust, fleshy flowers with epicalyx segments more deflexed and longer than the typical variety. The principal diagnostic characteristics of var. occidentalis are the sessile leaves with an attenuate base and inflorescences with sparse flowers on short ascending branches. One collection, morphologically and geographically intermediate between the two varieties (Anderson 9795), from southeastern Mato Grosso, near the border with Goiás, has subcoriaceous sessile leaves and widely divergent branches of the inflorescence. Diplusodon virgatus var. occidentalis is the second taxon of Diplusodon to be discovered in Bolivia. Diplusodon bolivianus is also known from the cerrados of the Parque Nacional Noel Kempff Mercado in an area to the south of D. virgatus var. occidentalis. It is easily separated from D. virgatus var. occidentalis by the rosecolored flowers in a distinct terminal inflorescence and yellow hirsute to sericeous indumentum that covers all parts of the plant. 99. Diplusodon vittatus T.B.Cavalc., Phytotaxa 226(2): 152, fig. 6. 2015. Type: Brazil. Goiás, Alto Paraíso de Goiás, povoado de São Jorge, na beira da estrada, 4 Jan 2003 (fl), J. F. B. Pastore & E. Suganuma 234 (holotype, CEN [00050979]). Figs. 13.123 A–E, 13.127 Subshrubs, erect, branched, caespitose, with a xylopodium, not dark-brown when dry. Upper branches subquadrangular, congested, unwinged, hirtellous to rarely pubescent, covered by white and sparse trichomes, internodes 0.7–1.1  cm long. Leaves decussate, not imbricate, suberect, petiolate; petioles ca. 1 mm long; blades 15–35  ×  1.5–4(−6) mm, coriaceous, not discolorous, eucamptodromous, narrowly lanceolate, narrowly elliptic, base acute, apex acute, margin revolute, adaxial surface sericeous to hirtellous covered by white and fine trichomes; abaxial surface covered by hirtellous to villous covered by white trichomes; secondary

Descriptions

449

Fig. 13.123  Diplusodon vittatus. A. Branch with inflorescence. B. Leaf, adaxial surface. C. Leaf, abaxial surface. D. Detail of leaf abaxial surface indumentum. E. Flower with prophyll, without petals. (A–E Pastore et al. 234, CEN)

450

13  Systematic Treatment

veins 2 on each side of the midvein, almost parallel, prominent on the abaxial surface; domatia present. Synflorescence frondose, diplobotryum spiciform type, lax; accessory branches absent, principal florescence 10–12  mm long, botryum spiciform type, paraclades 5–7 mm long; bracts similar to the vegetative leaves, except diminishing in size toward the apex of the branch, 15–25 × 1.5–3 mm wide, linear, margin revolute; flowers subsessile, hypopodium absent, epipodium ca. 0.5  mm long; prophylls ca. 5.5 × 1.5 mm, slightly surpassing the apex of the floral tube ca. 0.5 mm, narrowly oblong, with 2 inconspicuous veins on each side of the midvein, base acute, apex acute, margin plane; floral tube ca. 5  mm long, funnel-shaped, sparsely covered with white and fine trichomes; sepals 2.8–3 mm long, triangular, not deflexed, not caudate, ciliate, covered by sparse, short, white and trichomes; epicalyx segments 3–3.5 mm long, cylindrical, spreading, covered by sparse, short, white and sparse trichomes; corolla ca. 4 cm in diam., lilac, petals ca. 21 × 6 cm; stamens 18, free portion of the filaments ca. 8 mm long, ca. 1 mm exserted; ovary ca. 2 × 2.5 mm, obovate, glabrous, style ca. 13.5 mm long, ovules ca. 21. Fruits not seen. Distribution and Ecology (Fig. 13.127)  Diplusodon vittatus occurs only in campo limpo vegetation. Flowers in January. Specimens Examined: BRAZIL.  The type. Diplusodon vittatus is characterized by its habit with the xylopodium and ascending branches covered by sparse, white trichomes, narrow leaves with one prominent vein on each side of the midvein, epicalyx segments 3 mm long, and flowers with 18 stamens. Because of its narrowly lanceolate to narrowly elliptic leaves with the base and apex acute, it resembles D. macrodon. However, the latter species differs from D. vittatus mainly because of the presence of dense indumentum of short trichomes covering all the vegetative and floral parts and longer epicalyx segments (4–6.5 mm long) much longer than the sepals. 100. Diplusodon vividus T.B.Cavalc., sp. nov. Type: Brazil. Minas Gerais: Santo Antonio do Retiro, alto da Serra do Espinhaço, base do morro do Formoso,15o13′33″ S, 42o49′46″ W, 26 Mar 2017 (fl, fr), A.  C. Sevilha, G. Pereira-Silva, J. B. Pereira, S. E. Noronha 6876 (holotype, CEN [00101934]; isotypes, K, NY, BHCB, SPF, W). Figs. 13.124 A–F, 13.127, 15.5 G Diplusodon vividus is characterized by small leaves, with strongly revolute margins and abundant indumentum throughout the plant which create a canescent aspect. The upper branches are rust-colored, the flowers have epicalyx segments that are distinctly broadly lanceolate, and the androecium has 12 stamens. The new species is distinguished from Diplusodon parvifolius mainly by the prophylls that reach or surpass the apex of the floral tube, by epicalyx segments that are 2.2– 2.5 mm long, broadly lanceolate and long-ciliate, and by flowers with epicalyx segments that are longer (1.5–2 mm long) than those of D. vividus.

Descriptions

451

Fig. 13.124  Diplusodon vividus. A. Branch with inflorescence. B–D. Leaves, abaxial surface. E. Detail of leaf abaxial surface indumentum. F. Flower with prophyll, without petals

452

13  Systematic Treatment

Shrubs ca. 0.8  m tall, erect, much-branched, canescent, not dark-brown when dry. Upper branches subquadrangular to quadrangular, rust-colored, unwinged, tomentose to villous, canescent, internodes 0.7–1 cm long. Leaves decussate, not imbricate, congested, spreading, sessile to subsessile; petioles 0–0.5  mm long; blades 7–15  ×  5–9  mm, canescent to, coriaceous, strongly discolorous, acrodromous with supranumerary base, lanceolate, wide-lanceolate, oval, suborbicular, rhombic to rare elliptic, base cordate, subcordate to obtuse, apex obtuse, margin strongly revolute; secondary veins 4–5 on each side, adaxial surface verrucose, sparse-tomentose, glabrescent, grayish green, with deeply sulcate veins, abaxial surface tomentose to villous, white, veins prominent, rusted-colored; domatia absent. Synflorescence frondose, pleiobotryum type, congested; accessory branches absent, principal florescence botryum type, 4–10  cm long, often truncate, paraclades 4–20 cm long; bracts similar to the vegetative leaves in shape and size; flowers subsessile, hypopodium ca. 0.5 mm long, epipodium ca. 0.5 mm long; prophylls 6–7 × 3.5–4.5 mm, surpassing the apex of the floral tube, rhombic to widely rhombic, unkeeled, base acute, apex obtuse, margin plane, ciliate, involute at the apex; floral tube 5.5–6  mm long, funnel-shaped, tomentose to villous, densely covered with long and white trichomes, turning the floral tube whitish, sepals 2–2.8  mm long, triangular, not caudate, sometimes deflexed, pilose as the floral tube, longciliate; epicalyx segments 2.2–2.5 mm long, long-ciliate, wide-lanceolate, spreading to deflexed, shorter than the sepals; corolla ca. 2.5  cm diam., rose-magenta, petals 12–15 × 9–12 mm, obovate to wide-elliptic, base obtuse; stamens 12, free portion of the filaments 6–7 mm long, slightly exserted; ovary 1.9–2 × 2–2.2 mm, obovate, glabrous, style 13–14  mm long, ovules 18–20. Capsules 4.2–4.9  ×  4.5– 4.9 mm, globose, glabrous; seeds ca. 14, 2.1–3 × 1.9–2.1 mm. Distribution and Ecology (Fig.  13.127)  Diplusodon vividus occurs with occasional individuals at the type locality, on white sandy soil of campos rupestres and rocky cerrado, at elevations of 1311–1334 m. Flowers and fruits in March. Specimens Examined: BRAZIL.  The type. Diplusodon vividus belongs to the Hirsutus complex (discussed under D. bahiensis). Diplusodon vividus is a dense, much-branched, and canescent shrub, with rustcolored branches and small (7–15  ×  5–9  mm), densely congested, discolorous leaves with revolute margins and dense indumentum. The flowers bear petals of intense and bright rose-magenta color that are almost as broad as long; a tomentose to villous floral tube, with broadly lanceolate, long-ciliate, and spreading to deflexed epicalyx segments; and an androecium with 12 stamens. The most similar species is Diplusodon parvifolius, from Bahia, which resembles D. vividus based on small leaves (4–12 × 3–7 mm), with revolute margins, and 12 stamens. The species differ mainly because D. parvifolius has a dark-brown color when dry, eucamptodromous leaves, small prophylls reaching to the middle of the floral tube, and smaller epicalyx segments (1–2 mm long) that are cylindrical and not ciliate.

Descriptions

453

Fig. 13.125  Diplusodon xerophilus. A. Branch with inflorescence. B. Section of internode. C. Leaf, abaxial surface. D. Detail of leaf abaxial surface indumentum and domatia. E. Flower with prophyll, without petals. F. Prophyll, adaxial surface. (A Faria et  al. 3983, CEN; B–F Cavalcanti et al. 2192a, CEN)

454

13  Systematic Treatment

101. Diplusodon xerophilus T.B.Cavalc., sp. nov. Type: Brazil. Goiás: Chapada dos Veadeiros, estrada Alto Paraíso de Goiás-Colinas do Sul, quase em frente ao povoado São Jorge, 14°10′85″ S, 47°49′08″ W, 2 Apr 1997 (fl, fr), T.B.Cavalcanti, G. Pereira-Silva & S. A. T. Graham 2192a (holotype, CEN [00026999]; isotypes, MO, NY). Figs. 13.125 A–F, 13.127 Diplusodon xerophilus is characterized by narrowly elliptic leaves with an acute apex and prominent veins and domatia on the abaxial surface; flowers with epicalyx segments that are shorter than the sepals and deflexed; and 18 stamens. The leaf shape of D. xerophilus is similar to that of D. vittatus, but the latter species differs from the new species by its longer prophylls (5.5 mm long) and epicalyx segments (3–3.5 mm long). Subshrubs 0.4–1 m tall, erect, much-branched, with a xylopodium, sometimes prostrate, not dark-brown when dry. Upper branches cylindrical, unwinged, hirtellous to hirsute, long trichomes mixed with very short ones, internodes 1.5–2.4 cm long. Leaves decussate to fasciculate, not imbricate, suberect to spreading, petiolate; petioles 1–1.5 mm long; blades 15–45 × 4–7 mm, chartaceous, not discolorous, slightly canescent, not dark-brown when dry, eucamptodromous, narrowly elliptic, base acute, apex acute, margin subrevolute, hirtellous, with fine and long sparse trichomes, glabrescent; secondary veins 1–2 on each side, departing along the midvein, on the abaxial surface the basal vein inconspicuous or absent, the median vein prominent, adaxial surface with sunken veins, both surfaces covered by fine and long trichomes; domatia present, prominent. Synflorescence frondose-bracteose to frondose, diplobotryum to pleiobotryum type, lax; accessory branches absent, principal florescence botryum type, ca. 7 cm long, often truncate, paraclades 5–15 cm long; bracts 10–15  ×  2–4  mm, the same shape as the leaves diminished in size toward the apex of the branch; flowers subsessile, hypopodium absent, epipodium ca. 0.5 mm long; prophylls 2.5–2.6 × 0.8–1 mm, reaching the middle of the floral tube, oblong, midvein prominent, base acute, apex obtuse, usually reflexed, margin plane, ciliate, the trichomes fine and long; floral tube 3.5–4 mm long, funnel-shaped, hirtellous, covered by sparse and long trichomes; sepals 2–2.5 mm long, triangular, not caudate, not deflexed, hirtellous; epicalyx segments 1–1.3 mm long, cylindrical, slightly compressed, deflexed to spreading, the same indumentum as that of the sepals, ciliate, shorter than the sepals; corolla ca. 2.5 cm in diam., rose to lilac, petals 9–10 × 5–6 mm, broadly obovate, apex obtuse; stamens 18, free portion of the filaments ca. 4.5 mm long, slightly exserted; ovary 1.7–1.8 × 2–2.1 mm, obovaldepressed, glabrous, style 6.8–7 mm long, ovules 18–24. Capsules 4–4.5 × 4.5–5 mm, globose, glabrous; seeds 13–24, 2–2.8 × 2.1–2.8 mm. Distribution and Ecology (Fig.  13.127)  Diplusodon xerophilus is known only from the municipalities of Teresina de Goiás and Alto Paraíso de Goiás, Chapada dos Veadeiros, in the northern part of the state of Goiás. It grows in campo rupestre, campo limpo, campo sujo, and cerrado. Flowers and fruits from February to July.

Descriptions

455

Specimens Examined: BRAZIL.  Goiás,  Alto Paraíso de Goiás, Chapada dos Veadeiros, estrada ligando São Jorge à Colinas do Sul, ao lado da entrada para Raizama, 1 Feb 2004 (fl, fr), Pastore et al. 784 (CEN); Alto Paraíso de Goiás, margem da estrada de São Jorge para Colinas do Sul, 14°07′16″ S, 47°30′45″ W, 5 Apr 2014 (fl, fr), Faria-Jr. et al. 3983 (CEN, UB n.v.); Alto Paraíso de Goiás, rodovia Colinas do Sul, 7 km antes de São Jorge, sentido Alto Paraíso, Amaral-Santos et al. 3678 (CEN); Teresina de Goiás, rodovia GO-118, km 205, ca. 50 km de Alto Paraíso de Goiás, 13°54′ S, 47°21′ W, 31 Jul 2000 (fl, fr), Forzza et al. 1607 (CEN, SPF). Diplusodon xerophilus is characterized by the narrowly elliptic leaves with an acute apex and prominent veins and domatia on the abaxial surface. The flowers have epicalyx segments that are shorter than the sepals and usually deflexed (sometimes spreading) and an androecium with 18 stamens. The specific epithet refers to the dry habitat where the specimens were collected. Diplusodon xerophilus is similar to D. grahamae which also has canescent leaves, flowers with 18 stamens, and deflexed epicalyx segments. However, the leaves of D. grahamae are linear and strongly revolute, the flowers are larger, and the prophylls are longer (sometimes exceeding the floral tube), and they have involute margin. Furthermore, the leaves and flower indumentum are sericeous in D. grahamae. Diplusodon xerophilus is also similar to D. vittatus based on leaf shape, but the flowers in D. vittatus are longer (floral tube ca. 5 mm long), as are the epicalyx segments (3–3.5 mm long). 102. Diplusodon xiphodon T.B.Cavalc., Phytotaxa 226(2): 155, fig. 7. 2015. Type: Brazil. Bahia: Mucugê, Vale do Medronho, 13°1′ S, 41°21′ W, 27 Mar 2009 (fl), R. M. Harley & E. Ribeiro Jr. 55896 (holotype, HUEFS). Figs. 13.126 A–F, 13.127 Shrubs to 1 m tall, erect, branched, dark-brown when dry. Upper branches cylindrical, unwinged, hirtellous to hirsute, trichomes white, thick and sparse, internodes 1.5–4.5 cm long. Leaves decussate, not imbricate, patent to suberect, petiolate; petioles 1–2 mm long; blades 10–20 × 7–15 mm, coriaceous, slightly discolorous, acrodromous with supranumerary base to eucamptodromous, orbicular, ovate to broadly elliptic, base obtuse to retuse, apex obtuse, margin revolute, hirtellous on both surfaces, adaxial surface gray-colored, abaxial surface rust-colored; secondary veins (3–)4 on each side, departing along the midvein; domatia present, inconspicuous. Synflorescence bracteose to frondose-bracteose, diplobotryum to triplobotryum type, lax; accessory branches absent, principal florescence ca. 9 cm long or truncate, paraclades 4–9 cm long, delicate; bracts 4–12 × 3–10 mm, orbicular; flowers subsessile, hypopodium ca. 1  mm long, epipodium 0–0.5  mm long; prophylls ca. 3.5  ×  1.8–2  mm, reaching the middle of the floral tube, clavate to spathulate, unkeeled, apex obtuse, margin plane, with sparse, white and thick trichomes, ciliate with thick trichomes; floral tube ca. 3.5 mm long, funnel-shaped with an attenuate base, sparsely covered with white thick trichomes; sepals ca. 2 mm long, triangular, ciliate, slightly caudate, apex deflexed; epicalyx segments ca. 2  mm long,

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Fig. 13.126  Diplusodon xiphodon. A. Branch with inflorescence. B. Leaf, abaxial surface. C. Prophyll, adaxial surface. D. Flower with prophyll, without petals. E. Epicalyx segment. F. Capsule. (A–F Harley & Ribeiro Jr. 55869, HUEFS)

Descriptions

457

Fig. 13.127  Geographic distribution of Diplusodon vittatus, D. vividus, D. xerophilus, and D. xiphodon

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13  Systematic Treatment

sword-shape, flat, spreading, slightly shorter than the sepals, glabrous but strongly ciliate with thick trichomes; corolla ca. 1.5 cm in diam., dark rose, petals ca. 6 x 4 mm, elliptic, apex obtuse; stamens 12, free portion of the filaments ca. 7 mm long, exserted ca. 2 mm; ovary ca. 2 × 2 mm, slightly rhomboid, glabrous, style 10–18 mm long, ovules ca. 7. Capsules 4.5–5 × 3–3.5 mm, elliptic-fusiform, slightly stipitate, glabrous; seeds 5–6, 2.8–3 × 1.8–2 mm. Distribution and Ecology (Fig.  13.127)  Diplusodon xiphodon is known from Mucugê and Rio de Contas, Bahia (Chapada Diamantina), growing in rocky-sandy soil of campos rupestres. Flowers registered in November and March; fruits registered in November. Specimens Examined: BRAZIL. Bahia,  Rio de Contas, vertente leste, Campo do Queiróz, ao lado leste, 13°32′ S, 41°57′ W, 29 Nov 1988 (fl, fr), Harley & Stannard 26681 (CEN [2], CEPEC, K, SPF). Diplusodon xiphodon is part of the Hirsutus complex (discussed under D. bahiensis). Diplusodon xiphodon is characterized by orbicular to broadly elliptic leaves and lax inflorescences with small flowers and bracts. Diplusodon xiphodon is similar to D. bahiensis in that both have dark-brown leaves when dry, hirtellous indumentum on the leaves and flowers, and leaves with 3–5 secondary veins. Diplusodon bahiensis differs because it has congested, frondose inflorescences, larger flowers (floral tube 5–5.5 mm long), and longer prophylls (6–6.5 mm long) that reach the apex of the floral tube or surpass it arriving to the base of the epicalyx segments.

Chapter 14

Insufficiently Known Species

103. Diplusodon capitatus (A.St.-Hil.) Koehne, Fl. Bras. (Martius) 13(2): 337. 1877. Diplusodon microphyllus Pohl f. capitata A.St.-Hil., Fl. Bras. Merid. (A.St.-Hil.) 3: 149, 1833. Type: Brazil. Minas Gerais: dans les campos près Chapada “Minan Nova” (Minas Novas), A. Saint-Hilaire s.n. (catal. B1 n° 1596) (lectotype, P [00506524], designated by Lourteig 1989: 224; isolectotypes, P [00506526, 00506525]). Figs. 14.1 and 14.3. Subshrubs, erect, branched. Upper branches cylindrical, young branches flattened, pubescent to hirtellous, rust-colored, not dark-brown when dry. Leaves decussate, not imbricate, suberect, short-petiolate; blades ca. 12  ×  8  mm, coriaceous, not discolorous, eucamptodromous, obovate to lanceolate, base attenuate, apex obtuse, margin subrevolute, short pubescent-villous, adaxial surface punctate; secondary veins 1–2 on each side, departing along the midvein. Synflorescence triplobotryum type; accessory branches absent; flowers subsessile, compact and capitate at the branch apices; prophylls ca. 5.2 × 2 mm, reaching or nearly reaching the apex of the floral tube, spathulate, apex obtuse, margin plane, ciliate, hirtellous; floral tube ca. 4.5 mm long, funnel-shaped, hirtellous; sepals ca. 3.5 mm long, triangular, not caudate, not deflexed, hirtellous; epicalyx segments 3.1–3.5  mm long, slightly exceeding the sepals, narrow and dorsiventrally compressed with spreading trichomes, erect; stamens 18, free portion of the filaments 8–9 mm long, exserted; ovary ca. 2 × 3 mm, rounded, glabrous, style ca. 13 mm long, ovules ca. 22. Fruits not seen. Distribution and Ecology (Fig. 14.3)  Diplusodon capitatus is only known from the type material collected in northern Minas Gerais. Specimen Examined. Brazil  The type.

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Fig. 14.1  Diplusodon capitatus (Saint-Hilaire s.n., P)

The diagnosis of Saint-Hilaire (1833) for D. microphyllus f. capitatus was  expanded when Koehne (1877) elevated this form to a species based on the examination of the one flower present on the type material. There were several unsuccessful attempts to recollect the species in the region of Minas Novas, Minas

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14  Insufficiently Known Species

Fig. 14.3  Geographic distribution of Diplusodon capitatus and D. thysanosepalus

Gerais, which is the type locality cited by Saint-Hilaire (1833). The natural vegetation of the region has been almost completely obliterated by agricultural activities, remaining narrow areas of native cerrado. The habit and bracts of Diplusodon capitatus are similar to the type of D. serpyllifolius (=D. microphyllus). However, the epicalyx segments are long and narrow, unlike those of D. microphyllus. Because only the type material was examined, making it difficult to take a position on this species, I prefer to tentatively maintain D. capitatus in this study. Lourteig (1989) indicated a specimen of Saint-Hilaire s.n. at P as holotype, which is here considered as first-step lectotype (ICBN Art. 9.17, ex. 14). There are three specimens of Saint-Hilaire s.n. at P linked with the name Diplusodon capitatus, and a second-step lectotype is proceeded (P00506524). 104. Diplusodon thysanosepalus Lourteig & Sandwith, Sellowia 16: 145, fig. 6. 1964. Type: Brazil. Goiás: Rio Pizarão, 5 Jun 1895 (fl), A. F. M. Glaziou 21548a (holotype, P [00506523]; isotype, K). Figs. 14.2 and 14.3. Subshrubs ca. 0.4  m tall, erect, branched. Upper branches quadrangular?, winged?, glabrous, dark-brown when dry. Leaves decussate, not imbricate, sessile; blades 11–17  ×  7.5–12  mm, coriaceous, not discolorous, acrodromous-perfect-­ basal, ovate to oblong-ovate, base obtuse to subcordate, apex acuminate, margin

462

Fig. 14.2  Diplusodon thysanosepalus (Glaziou 21548a, P)

14  Insufficiently Known Species

14  Insufficiently Known Species

463

ciliate at the apex of young leaves; secondary veins two on each side of the midvein departing from the base; domatia absent. Synflorescence frondose, diplobotryum type, congested, florescence of the botryum type; accessory branches absent; flowers pedicellate, hypopodium ca. 1.5 mm, longer in fruit; prophylls surpassing the apex of the floral tube, obovate, unkeeled, apex subacute; floral tube oblong; sepals ca. 5 × 2.5 mm, triangular, slightly caudate, not deflexed, sparsely ciliate; epicalyx segments ca. 5.5 mm long (in the diagnosis ca. 0.75 mm long), linear, acute, sparsely ciliate, slightly longer than the sepals; corolla ca. 18 × 14 mm, rose, petals obovate; stamens 12; ovary round, glabrous, style very long, twice the length of the calyx, ovules 22. Fruits not seen. Distribution and Ecology (Fig.  14.3)  Known only from the type material collected in Goiás. Collected with flowers in June. Specimen Examined. Brazil  The type. Lourteig (1989) described D. thysanosepalus from the state of Goiás. According to the illustration and diagnosis, it has acrodromous 5-veined leaves, and the flowers have cilia on the sepals and epicalyx and an androecium of 12 stamens. I examined the type and was unable to clarify important details of the flowers, and no additional collections were located in other herbaria, so it appears that this species has not been recollected. The collection locality for Glaziou 21548 is listed in Glaziou (1909: 302) as, “rio Pizarão, dans le campo, Goyaz, no 21548. Arbuste, fl. roses. Janvierfévier. C.” ‘Pizarão’. There is Rio Piçarrão, 47°37′ W, 14°20′ S, in Goiás, and that is probably what Glaziou attempted to write. I prefer to maintain this species in this study despite several unsuccessful attempts to relocate the species at the type locality and nearby areas at different times of the year as recently as 2015.

Chapter 15

Excluded Names

Diplusodon arboreus Poepp. & Endl. Nov. Gen. et Sp. 2: 66. 1817. = Physocalymma scaberrimum Pohl Diplusodon ginorioides Griseb., Cat. Pl. Cub. 106. 1866. = Ginoria ginorioides (Griseb.) Britton.

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Chapter 16

Color Plates

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Fig. 16.1  A–C. Diplusodon alatus. D. D. bradei. D. burchellii. E. D. adpressipilus. F. D. burchellii. Photos A–D, T. B. Cavalcanti; photo F, M.F. Simon

16  Color Plates

469

Fig. 16.2  A. Diplusodon candollei. B. D. ciliatiflorus. C–D D. glaucescens. E. D. grahamae. Photos A–D, T. B. Cavalcanti; photo E, G. Pereira-Silva

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16  Color Plates

Fig. 16.3  A. D. buxifolius. B. D. hexander. C. D. praetermissus. D. D. petiolatus. E. D. ramosissimus. F. D. ovatus. Photos A, C–D, T.  B. Cavalcanti; photo B, C.E.B.  Proença; photo E, M.F. Simon; photo F, G. Pereira-Silva

16  Color Plates

471

Fig. 16.4  A– B. D. paraisoensis. C. D. rotatus. D. D. strigosus. E. D. tenuifolius. Photos A–C, T. B. Cavalcanti; photos D–E, M.F. Simon

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Fig. 16.5  A. D. thymifolius. B. D. virgatus var. virgatus. C. D. vacillans. D. D. vinaceus. E. D. venosus. F. D. villosus. G. D. vividus. Photos A–B, E–F, T.  B. Cavalcanti; photo C, G. Pereira-Silva; photo D, M.F. Simon; photo G, A. C. Sevilha

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Numerical List of Taxa

  1.   2.   3.   4.   5.   6.   7.   8.   9. 10. 11. 12. 13. 14. 15. 16. 17. 18. 19. 20. 21. 22. 23. 24. 25. 26. 27. 28. 29.

Diplusodon adpressipilus Lourteig Diplusodon aggregatifolius T.B.Cavalc. Diplusodon alatus T.B.Cavalc. Diplusodon appendiculosus Lourteig Diplusodon argenteus Lourteig Diplusodon argyrophyllus T.B.Cavalc. Diplusodon astictus Lourteig Diplusodon bahiensis T.B.Cavalc. Diplusodon bolivianus T.B.Cavalc. & S.Graham Diplusodon bradei Pilg. Diplusodon burchellii Koehne Diplusodon buxifolius (Cham. & Schltdl.) DC. Diplusodon caesariatus Lourteig Diplusodon canastrensis T.B.Cavalc. Diplusodon candollei Pohl ex DC. Diplusodon capitalensis T.B.Cavalc. Diplusodon capitatus (A.St.-Hil.) Koehne Diplusodon chapadensis T.B.Cavalc. Diplusodon ciliatiflorus T.B.Cavalc. Diplusodon ciliatus (T.B.Cavalc.) T.B.Cavalc. Diplusodon ciliiflorus Koehne Diplusodon conduplicatus T.B.Cavalc. Diplusodon cordifolius Lourteig Diplusodon cryptanthus T.B.Cavalc. Diplusodon decussatus Gardner Diplusodon divaricatus Pohl Diplusodon epilobioides Mart. ex DC. Diplusodon ericoides Lourteig Diplusodon fastigiatus Lourteig

© The Author(s), under exclusive license to Springer Nature Switzerland AG 2021 T. B. Cavalcanti, The Genus Diplusodon (Lythraceae), Flora Neotropica, https://doi.org/10.1007/978-3-030-65875-5

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482

30. 31. 32. 33. 34. 35. 36. 37. 38. 39. 40. 41. 42. 43. 44. 45. 46. 47. 48. 49. 50. 51. 52. 53. 54. 55. 56. 57. 58. 59. 60. 61. 62. 63. 64. 65. 66. 67. 68.

Numerical List of Taxa

Diplusodon floribundus Pohl Diplusodon glaucescens DC. Diplusodon glaziovii Koehne Diplusodon glocimarii T.B.Cavalc. Diplusodon gracilis Koehne Diplusodon grahamae T.B.Cavalc. Diplusodon hatschbachii Lourteig Diplusodon helianthemifolius Mart. ex DC. a. var. helianthemifolius b. var. pemphoides (DC.) Koehne Diplusodon heringeri Lourteig Diplusodon hexander Mart. ex DC. Diplusodon hirsutus (Cham. & Schltdl.) DC. Diplusodon imbricatus Pohl Diplusodon incanus Gardner Diplusodon kielmeyeroides A.St.-Hil. a. var. kielmeyeroides b. var. macrocalyx T.B.Cavalc. Diplusodon lanceolatus Pohl Diplusodon leucocalycinus Lourteig Diplusodon longipes Koehne Diplusodon macrodon Koehne Diplusodon marginatus Pohl Diplusodon mattogrossensis T.B.Cavalc. Diplusodon micromerus T.B.Cavalc. Diplusodon microphyllus Pohl Diplusodon minasensis Lourteig Diplusodon mononeuros Pilg. Diplusodon myrsinites Mart. ex DC. Diplusodon nigricans Koehne Diplusodon nitidus Mart. ex DC. Diplusodon oblongus Pohl Diplusodon obtectus T.B.Cavalc. Diplusodon orbicularis Koehne Diplusodon ovatus Pohl Diplusodon panniculatus Koehne Diplusodon paraisoensis Lourteig Diplusodon parvifolius Mart. ex DC. Diplusodon petiolatus (Koehne) T.B.Cavalc. Diplusodon plumbeus T.B.Cavalc. Diplusodon praetermissus T.B.Cavalc. Dipludoson prostratus T.B.Cavalc. Diplusodon psammophilus Lourteig

Numerical List of Taxa

  69. Diplusodon puberulus Koehne   70. Diplusodon punctatus Pohl a. var. punctatus b. var. dentatus T.B.Cavalc.   71. Diplusodon pygmaeus T.B.Cavalc.   72. Diplusodon quintuplinervius (Nees) Koehne   73. Diplusodon ramosissimus Pohl   74. Diplusodon rectinervis T.B.Cavalc.   75. Diplusodon retroimbricatus Koehne   76. Diplusodon rosmarinifolius A. St.-Hil.   77. Diplusodon rotatus T.B.Cavalc.   78. Diplusodon rotundifolius Mart. ex DC.   79. Diplusodon saxatilis Lourteig   80. Diplusodon sessiliflorus Koehne   81. Diplusodon sigillatus Lourteig   82. Diplusodon sordidus Koehne   83. Diplusodon speciosus (Kunth) DC.   84. Diplusodon stellatus T.B.Cavalc.   85. Diplusodon strigosus Pohl   86. Diplusodon subsericeus Casar. ex Koehne   87. Diplusodon tenuifolius T.B.Cavalc.   88. Diplusodon thymifolius Mart. ex. DC.   89. Diplusodon thysanosepalus Lourteig & Sandwith   90. Diplusodon trigintus T.B.Cavalc.   90. Diplusodon ulei Koehne   92. Diplusodon uninervius Koehne   93. Diplusodon vacillans T.B.Cavalc.   94. Diplusodon venosus T.B.Cavalc.   95. Diplusodon verruculosus T.B.Cavalc.  96. Diplusodon vidalii Lourteig  97. Diplusodon villosissimus Pohl  98. Diplusodon villosus Pohl  99. Diplusodon vinaceus T.B.Cavalc. 100. Diplusodon virgatus Pohl a. var. virgatus b. var. occidentalis T.B.Cavalc. & S.Graham 101. Diplusodon vittatus T.B.Cavalc. 102. Diplusodon vividus T.B.Cavalc. 103. Diplusodon xerophilus T.B.Cavalc. 104. Diplusodon xiphodon T.B.Cavalc.

483

List of Exsiccatae

Abrahim, M. A. et al., 02 (16). Ackerman, J. D., s.n. (50). Aires, J., 4860 (96); 4869 (96). Alkimim, W., 25 (75). Allem, A. C., 812 (96); 849 (56). Almeida, E. F. & Lourdes, M., 280 (36b). Almeida, E. P., 139 (12). Almeida, S. P., 924 (96). Almeida, T. E., 1302 (71). Almeida, T. E. et al., 877 (51). Alvarenga, D., 16 (96); 39 (96). Alvarenga, D. et al., 168 (56); 206 (96). Alves, L. J. et al., 258 (87). Alves, M. A., 6 (56); 83 (98a); 110 (79). Alves, M.C. et al., 8 (43). Alvin, L. C., 20 (72). Amaral, A. C. & Moreira, G. A., 44 (69a). Amaral, A. C. & Santos, A. A., 2 (79). Amaral, A. C. et al., 11 (47); 27 (96). Amaral, A. G. & Eugênio, C. U. O., 1541 (56); 1637A (79). Amaral, A. G. et al., 813 (96); 978 (96); 1130 (96); 1797 (79); 2096 (79); 2428 (75); 2978 (41). Amaral, M. C. E. & Kawasaki, M. L., CFSC 9321 (39). Amaral, M. C. E. et al., CFSC 8403 (50); CFSC 9322 (58); 97/86 (95). Amaral-Santos, A. & Cordovil-Silva, S. P. 96 (96). Amaral-Santos, A. & Nascimento, A. R. T., 937 (82). Amaral-Santos, A. & Pereira, J. B., 1850 (56); 1866 (43).

© The Author(s), under exclusive license to Springer Nature Switzerland AG 2021 T. B. Cavalcanti, The Genus Diplusodon (Lythraceae), Flora Neotropica, https://doi.org/10.1007/978-3-030-65875-5

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List of Exsiccatae

Amaral-Santos, A. et  al., 30 (69a); 538 (18); 539 (18); 569 (18); 615 (45); 616 (69a); 737 (75); 871 (25); 996 (96); 998 (96); 999 (96); 1008 (96); 1015 (96); 1126 (72); 1175 (72); 1205 (72); 1243 (75); 1281 (72); 1316 (72); 1391 (72); 1424 (72); 3131 (61); 3665 (61); 3666 (49); 3671 (56); 3677 (45); 3678 (101). Amorim, E. H. et al., 458 (43), 477 (43), 601 (43), 710 (43), 760 (43), 817 (43), 833 (43). Anderson, W.R., 6265 (56); 6336 (61); 6506 (61); 6722 (56); 7002 (43); 7188 (44); 7265 (5); 7551 (98a); 7658 (44); 8001 (56); 8455 (36a); 8490 (30); 8496 (36a); 8501 (38); 8503 (38); 8590 (30); 8593 (10); 8742 (98a); 9439 (98a); 9488 (84); 9498 (84); 9795 (98b); 10107 (84); 10389 (96); 10418 (84);10465 (98a); 35457 (20). Anderson, W. R. et al., 7949 (27); 35140 (92); 35237 (38); 35457 (18); 36117 (58); 36221 (20); 36408 (36b); 36736 (36b). Andrade, A. G. & Emmerich, M., 320 (75). Andrade, P. M. & Lopes, M. A., 1038 (53); s.n. BHCB 8808 (43); s.n. BHCH 8812 (90); s.n. BHCB 8818 (58). Anhanguera, U. L., 42 (75). Anjos, B. A., s.n. (61869) (19). Aragaki, S. & Batalha, M., 330 (98a). Arantes, A. A., 39 (98a); 144 (43); 159 (43). Arantes, A. A. & Gonçalves, D. J. P., 1779 (68). Arantes, A. A. & Marra, K., 169 (68); 180 (98a). Arantes, A. A. et al., SF24 (97); 33 (97); 38 (86); 700 (43); 943 (98a); 956 (42a); 1637 (32); 1724 (32). Araujo, A. O. et al., 101 (36b); 119 (36b). Araújo, G., 2 (98a); 173 (43); 233 (98a). Araújo, L. H., 9 (75). Arbo, M. M. et al., 3883 (98a); 3992 (98a); 4007 (12); 4032 (12); 4358 (55); 4371 (38); 4469 (55); 5640 (87). Árbocz, G. F., 6513 (82). Árbocz, G. F. et al., 2075 (50); 3292 (84); 3318a (56); 3472 (56); 3573 (56). Argent, G. C. G. et al. 6716 (56). Arroyo, L. et al., 684 (98b). Arruda, L. J. & Santos, L. M., 88 (12). Arruda, L. et al., 303 (43) Assis, L. C. S. et al., 708 (95); 715 (98a). Assis, M. C. et al., 2 (98a); 10 (96); 12 (56); 355 (81); 549 (84). Atkins, S. et al., 5680 (89); [PCD] 5582 (62). Azevedo, I. N. et al., 318 (56); 375 (98a); 428 (79). Azevedo, M. L. M., 38 (96); 709 (72). Azevedo, M. L. M. & Alvarenga D., 547 (56); 775 (75). Azevedo, M. L. M. & Lopes, E. C., 479 (96). Azevedo, M. L. M. & Oliveira, F. C. A., 616 (56). Azevedo, M. L. M. et al., 180 (72); 985 (69a).

List of Exsiccatae

487

Bacariça, E. M., 28 (50). Badini, J., s.n. OUPR 1776 (50); OUPR 1781 (12); OUPR 1811 (50); OUPR 1987 (12); OUPR 2742 (39); OUPR 3727 (98a); OUPR 3743 (12); OUPR 3854 (50); OUPR 4162 (53); OUPR 4230 (98a); OUPR 4250 (50); OUPR 4261 (50); OUPR 4331 (98a); OUPR 4332 (98a); OUPR 4540 (12); OUPR 4541 (12); OUPR 4732 (12); OUPR 4735 (12); OUPR 4737 (50); OUPR 4759 (50) OUPR 4760 (50); OUPR 4761 (50); OUPR 4762 (50); OUPR 4765 (50); OUPR 4767 (50); OUPR 4770 (50); OUPR 4771 (50); OUPR 4775 (50); OUPR 4778 (50); OUPR 4790 (50); OUPR 4792 (50). OUPR 4792 (50); OUPR 5087 (12); OUPR 5089 (12); OUPR 5090 (98a); OUPR 5093 (12); OUPR 5419 (39); OUPR 19649 (39); OUPR 19650 (39); OUPR 19651 (39); OUPR 19652 (39); OUPR 19653 (39); OUPR 19654 (39); OUPR 19669 (95); OUPR 19670 (95); OUPR 19977 (12); OUPR 19978 (12); OUPR 20013 (39); OUPR 21534 (95); OUPR 21649 (50); OUPR 21668 (50); OUPR 21669 (50); OUPR 21692 (95); OUPR 22040 (95); OUPR 22156 (95); OUPR 22259 (12); OUPR 22367 (12); OUPR 22587 (98a); OUPR 22818 (95); OUPR 22937 (95); OUPR 23013 (12); OUPR 23014 (12); OUPR 23053 (95); OUPR 23809 (95); OUPR 24378 (95); OUPR 25529 (98a); OUPR 26078 (95); OUPR 26230 (95); OUPR 2663 (98a); OUPR 26769 (95). Badini, J. & Lisboa, C., s.n. OUPR 4960 (98a). Badini, J. & Zurlo, M.  A., s.n. OUPR 1782 (12); OUPR 25404 (12); OUPR 25405 (12). Baeta, A., 10525 (95). Barbosa, A. A. A., 392 (58); 403 (39); 586 (43); 3242 (98a); s.n. CEN 38150, HUFU 23429 (98a). Barbosa, E. G. et al.1092 (45); 1093 (11). Barboza, M. A. et al., 23 (9); 43 (9). Barcelos, L., s.n. HUFU 59165 (98a); HUFU 72352 (98a). Barreto, K. D. et al., 2314 (98a); 2725 (98a). Barros, A. & Stehmann, J. R., 1 (12); s.n. BHCB 40775 (12); s.n. BHCB 40788 (12). Barros, F., 212 (58); 2681 (98a). Barros, M. G. A., 108 (79); 974 (75); 985 (96); 1000 (56); 1013 (75); 1014 (75); 1015 (75); 1017 (79); 1018 (96); 1021 (96); 1022 (79); 1023 (72); 1026 (72); 1031 (79); 1033 (79); 1229 (75). Barros, M. G. A. et al., 2281 (96); 2282 (96); 2318 (56); 2243 (96). Barros, R., 1281 (77). Barroso, G. et al., 590 (56); 624 (54); 787 (56). Batalha, M. & Mantovani, W., 47 (98a); 103 (98a). Batista, J. A. N. & Martins C. A. N., 2065 (12). Bautista, H. P., 593 (58); 611 (39). Bautista, H. P. et al., 3621 (19). Bayern, s.n. (50). Belém, R. P., 497 (56); 1903 (47); 1917 (56). Belém, R. P. & Mendes, J. M., 374 (71); 397 (68). Bernacci, L. C. et al., 1792 (98a). Bernardes, P. H. N. et al., 121 (32); 181 (32); 229 (10); 307 (32).

488

List of Exsiccatae

Bernardino, J., 16 (75). Bianchetti, L. B. & Pereira-Silva, G., 1227 (65). Bianchetti, L. B. et al., 895 (72). Bianchini, R. S., 471 (98a). Black, G. A. & Magalhães, M., 51-12077 (39). Boaventura, M., 167 (75). Boaventura, M. & Silva, J. V., 231 (96). Boiteux, M. S., s.n. (98a). Bordon, N. & Réu-Júnior, W. s.n., (25). Bosquetti, L.B. et al., 685 (98a). Bothelho, R. B. et al., 8 (87). Brade, A. C., 5663 (98a); 13405 (52); 13406 (10); 14406 (58); 15433 (56). Brade, A. C. & Apparicio 20495 (98a). Brade, A. C. & Barbosa, A. 17703 (55) Braga, M. M. N., 190 (12); s.n. BHCB 10662 (20). Braga, M. M. N. & Chaves, A. L. F., 412 (38); Braga, M. M. N. et al., s.n. BHCB 15849 (12). Brandão, M., 10883 (90); 14377 (59); 15379 (42a); 22439 (95). Brandão, M. & Laca-Buendia, J. P., 20818 (55). Brauner, L.M. et al., 58 (19). Brina, s.n. BHCB 35583 (98a); s.n. BHCB 39436 (12). Bringel, J. B. A. & Barbosa, N. F., 267 (63). Bringel, J. B. A. & Chacon, R. G., 222 (96). Bringel, J. B. A. & Ferreira, V. V., 913 (20). Bringel, J. B. A. & Moreira, H. J. C., 717 (82); 752 (22); 753 (92); 839 (61); 932 (3); 948 (69a); 961 (11); 974 (56). Bringel, J. B. A. & Pastore, J. F. B., 752 (22); 775 (61); 802 (61); 804 (22); 808 (5). Bringel, J. B. A. et al., 220 (9); 419 (46); 549 (96); 567 (36b); 1063 (18); 1266 (47); 1282 (56); 1301 (72). Brito, L., 50 (59). Brito, S. M., 14 (72). Brooks, R. R. et al., 36 (98a). Bucci, F. & Verano, C., 1559 (40). Burchell, W. J., 5722 (59); 6647 (56); 6700-13 (56); 7659 (69a); 7684-2 (69a); 7693 (84); 7726 (72); 7791 (72); 7911 (11); 7935 (72); 8011 (61); 8039 (24); 8046 (43). Calago, K. 97 (17); 98 (7); 106 (56); 147 (72); 152 (72); 175 (72); 182 (75); 208 (72). Calago, K & Silva, S., 128 (98a); 142 (84). Caldas, T. S. P., 17 (7). Camacho, G. P. et al., 4, (98a). Camara, P. S. & Brochado, A. L., 68 (56). Campos, J., 17 (98a). Campos, M. J. O., 87 (98a). Campos-Rocha, A & Moura, K. A., 446 (98a), 869 (61); 896 (98a) Cardoso, D. & Santos, R. M., 2012 (8). Cardoso, C. F. R. et al., 30 (87).

List of Exsiccatae

489

Carmo, R., CFSC 11272 (58). Carvalho, A. F., 487 (36b). Carvalho, A. M. & Delphim, C. F. M., 2248 (56). Carvalho, A. M. & Saunders, J., 2946 (19). Carvalho, A. M. et al., 6340 (62); 6432 (89). Carvalho, D.  A. et  al., s.n. ESAL 8319 (53); s.n. ESAL 6735 (98a); s.n. ESAL 7551 (53). Carvalho, F. J. et al., 154 (72); 292 (47); 399 (96). Carvalho, M. G. & STS, 339 (26); 378 (26); 705 (36b). Carvalho-Silva, M., 170 (56). Carvalho-Silva, M. et al., 240 (72); 1390 (50). Casaretto, J., 2746 (12); 2826 (85); 2989 (39). Castañeda, L. A. & D’Affonseca, S.L.C., s.n. CEN 34035 (98a). Castellois, I., 477 (58). Castro, N. M., 26 (58); 395 (58); 397 (20); 488 (39); 491 (39); s.n. HUFU 12396 (56); s.n. BHCB 1097 (58); s.n. BHCB 2255 (58); s.n. BHCB 18378 (58); s.n. BHCB 22458 (58). Castro, N. M. & Bernadete, s.n. HGB 2639 (87). Castro, N. M.& Mayo, S. CFSC 7192 (58). Castro, R. M. et al., 343 (53). Castro, V. G. P., s.n. BHCB 18378 (58). Cavalcanti, T. B., 1255 (79). Cavalcanti, T. B. & Ayres, C. H. S., 1256 (72); 1257 (61); 1259 (61); 1260 (19); 1263 (81); 1267 (64); 1268 (64). Cavalcanti, T. B. & Graham, S. A., 2258 (98a). Cavalcanti, T. B. & Mello-Silva, R. M. 2505 (66); 2508 (83). Cavalcanti, T. B. & Pereira-Silva, G., 1264 (61); 1265 (96); 1266 (98a); 2719 (84); 2720b (22); 2722 (96); 3182 (60); 3195 (26); 3723 (56); 3726 (84); 3728 (29); 3737 (56); 3739 (84); 3742 (3); 3743 (56); 3749 (18); 3767 (34); 3769 (84); 3780 (76); 3787 (95); 3791 (98a); 3793 (85); 3794 (12); 3982 (13); 3983 (13); 3984 (98a); 3985 (34); 3986 (44). Cavalcanti, T. B. et al., 30 (81); 86 (82); 87 (56); 108 (81); 180 (30); 188 (20); 189 (90); 201 (38); 202 (20); 211 (20); 217 (77); 220 (51); 227 (20); 230 (36b); 241 (38); 250 (38); 252 (36a); 253 (20); 259 (10); 263 (43); 278 (2); 282 (36b); 285 (36b); 295 (62); 303 (62); 314 (62); 359 (8); 373 (96); 377 (56); 381 (81); 382 (80); 384 (44); 390 (61); 394 (83); 400 (1); 401 (65); 403 (81); 404 (7); 411 (64); 414 (43); 417 (98a); 421 (43); 430 (56); 432 (56); 437 (56); 454 (56); 473 (84); 474 (56); 478 (75); 500 (96); 527 (79); 530 (16); 531 (75); 571 (98a); 586 (72); 648 (61); 651 (72); 657 (72); 660 (65); 662 (81); 681 (3); 682 (83); 683 (1); 684 (83); 697 (1); 815 (72); 818 (98a); 849 (1); 850 (3); 851 (65); 886 (98a); 1000 (47); 1014 (69b); 1015 (69a); 1042 (65) 1046 (18); 1053 (23); 1086 (84); 1120 (84); 1150 (47); 1220 (75); 1232 (72); 1233 (79); 1254 (63); 1309 (61); 1330 (56); 1339 (81); 1347 (56); 1363 (70); 1385 (61); 1395 (43); 1402 (25); 1421 (69a); 1428 (84); 1432 (98a); 1441 (84); 1458 (72); 1464 (18); 1466 (3); 1496 (72); 1575 (72); 1586 (11); 1593 (98a); 1649 (69a); 1686 (25); 1734 (25); 1824

490

List of Exsiccatae

(11); 1936 (98a); 2019 (56); 2149 (56); 2150 (43); 2174 (43); 2181 (56); 2182 (81); 2185 (17); 2186 (3); 2187 (1); 2188 (65); 2192a (101); 2193 (66); 2194 (70); 2197 (81); 2198 (80); 2199 (80); 2204 (44); 2205 (61); 2209 (44); 2210 (61); 2211 (37); 2217 (61); 2220 (63); 2223 (63); 2225 (40); 2227 (69b); 2228 (40); 2229 (69b); 2230 (63); 2232 (69b); 2234 (82); 2237 (82); 2243 (82); 2246 (46); 2248 (24); 2149 (56); 2250 (46); 2264 (43); 2266 (42a); 2265 (56); 2267 (42a); 2268 (56); 2269 (42a); 2270 (39); 2271 (58); 2279 (20); 2181 (56); 2284 (30); 2286 (30); 2287 (36b); 2293 (38); 2295 (20); 2296 (36a); 2298 (38); 2301 (10); 2303 (20) 2305 (31); 2311 (43); 2320 (2); 2322 (39); 2323 (39); 2332 (57); 2338 (33); 2339 (56); 2347 (84); 2351 (82); 2354 (84); 2381 (9); 2389 (56); 2391 (96); 2393 (66); 2398 (25); 2399 (43); 2438 (62); 2452 (89); 2456 (89); 2475 (89); 2491 (75); 2496 (96); 2502 (72); 2507 (3); 2513 (32); 2521 (32); 2558 (98a); 2561 (95); 2562 (39); 2567 (98a); 2570 (95); 2571 (12); 2581 (50); 2584 (95); 2586 (98a); 2595 (23);2646 (72);2661 (23); 2662 (69b); 2700 (69a); 2712 (72); 2715 (72); 2716 (75); 2721 (27); 2738 (61);3062 (98a); 3063 (94); 3073 (95); 3088 (53); 3100 (12); 3102 (90); 3105 (36a); 3107 (30); 3110 (27); 3114 (38); 3119 (43); 3122 (77); 3125 (52); 3126 (90); 3127 (30); 3129 (38); 3130 (15); 3132 (36b); 3133 (36b); 3134 (43); 3135 (36b); 3141 (62); 3142 (87); 3144 (89); 3146 (82); 3147 (61); 3148 (33); 3149 (4); 3150 (44); 3151 (54); 3153 (44); 3154 (54); 3155 (37); 3156 (44); 3157 (81);3158 (56); 3159 (1); 3161 (7); 3162 (83); 3164 (101); 3165 (54); 3166 (7); 3168 (66); 3170 (27);3171 (94); 3179 (98a); 3191 (71); 3199 (36b); 3200 (43); 3201 (20); 3202 (55); 3205 (90); 3206 (10); 3207 (60); 3243 (33); 3434 (61); 3489 (33); 3498 (81); 3502(1); 3506 (3); 3508 (83); 3509 (23);3510 (69a); 3514 (72); 3517 (84); 3518 (11); 3519 (84); 3524 (40); 3525 (5); 3526 (11); 3527 (22); 3528 (11); 3531 (61); 3532 (28); 3533 (56); 3534 (92); 3535 (33); 3536 (96); 3537 (37); 3538 (4);3539 (37);3540 (54); 3542 (44); 3543 (96); 3547 (37); 3562 (75); 3579 (96); 3580 (80); 3583 (84); 3584 (56); 3587 (98a); 3596 (101); 3599 (61); 3603 (96); 3608 (81); 3609 (22); 3615 (44); 3616 (101); 3617 (96); 3618 (101); 3620 (4); 3624 (56); 3625 (61);3626 (24); 3627 (5); 3632 (22); 3633 (11); 3640 (61);3648 (64); 3649 (60); 3751 (66); 3753 (27); 3755 (61); 3757 (35); 3754 (44); 3800 (12); 3807 (12); 3811 (3); 3812 (18); 3813 (65); 3820 (83); 3823 (17); 3824 (7); 3890 (26); 3897 (36b); 3899 (36b); 3907 (57); 3915 (43); 3919 (43); 3925 (77); 3928 (55); 3929 (67); 3930 (38); 3934 (30); 3935 (53); 3936 (38); 3940 (15); 3941 (20); 3975 (61); 3980 (81); 3988 (60); 3989 (86); 3990 (97); 3993 (72). Cavalcanti, T. B. et al., CFCR 5494 (78); 8528 (36b); 8552 (38); 8579 (20); 8597 (90); 8702 (90); 9488 (20); 9496 (38); 9507 (38); 9525 (67); 9529 (77); 9567 (38); 9574 (10); 9596 (2); 9644a (36b); 9650 (36b); 10175 (36b);10185 (90); 10313 (90). Cavalcanti, T. B. et al., CFSC 9365 (20); 9614 (58); 9630 (20); 9376 (58); 9614 (58); 9619 (58); 9630 (20); 9658 (58); 9683 (39); 9691 (39); 10097 (58); 10099 (39); 10110 (39); 10139 (58); 10159 (20); 10163 (58); 10615 (20); 10622 (58); 10623 (58); 10635 (58). Cerati, T. M., 158-A (98a). Cerati, T. M. et al., 254 (26).

List of Exsiccatae

491

Cesar, H. L., 68 (96). Chacon, R. G. & Pinho, J.C., 53 (16); 54 (79). Chacon, R. G. et al., 38 (79), 445 (75); 757 (72). Chaddad Jr., J., 39 (53); 57 (53). Chaves, E., 265 (17). Classe de Taxonomia da Universidade de Brasília, 111 (79); 112 (75); 362 (96); 399 (56). Claussen, P., 5 (39); 11a (50); 12a (12); 13a (12); 14a (12); 15a (12); 19a (43); 21a (50); 22a (39); 23a (39); 54 (12); 66 (39); 111 (12); 113 (85); 114 (39); 115 (39); 308 (43); 309 (39); 312 (98a); 1054 p.p. (12); s.n. (12); s.n. (85); s.n. (39). s.n. (43), s.n. ([herb. Fl. Bras. n. 119]) (43); s.n. (95); s.n. (98a). Clayton, W. D., 4813 (96); 4888 (56). Cobra, L. Q. & Sucre, D., 308 (47). Cochran, D., s.n. US 29248 (12). Coelho, B. B. S. & Meirelles, F.M., s.n. CESJ 23663 (82). Coelho, J. P., s.n. VIC 10107 (43). Collenette, C. L., 117 (98a). Colletta, G. D. et al., 69 (50). Conceição, A. A. & Cardoso, D., 1324 (89). Conceição, M. C. L. & Silva, J.C.S., 60 (79). Conceição, A. S. et al., 725 (36b). Coons, M. P. et al., 77-348 (39); 77-349 (95). Coradin, L., 4227 (79). Coradin, L. et al., 6644 (87). Cordeiro, I. & Mello-Silva, R. et al., CFCR 10144 (26). Cordeiro, I. et al.,CFCR 776 (2); 9122 (57); 9330 (20); 9387 (38). Cordeiro, I. et al., CFSC 9537 (20). Cordeiro, J. et al., 4951 (5). Cordeiro, L., 32 (98a). Cordovil-Silva, P. et al., 154 (45); 239 (56); 273 (98a); 276 (25); 319 (98a); 452 (29); 604 (29); 609 (72); 618 (84); 621 (56); 622 (72) ; 671 (72). Cortes, R. C. 3 (68). Costa, J. M. et al., 1088 (36b). Costa, L.V. s.n., (BHCB 5454). Costa, L.V. & Amado, J. C., 221 (36b). Costa-Vieira, R., 21 (29). Croat, T. B., 53630 (79). Cruz, M. A. V., 78 (90). Cunha, C. N. et al., 722 (82). Cunha, L. et al., 1739 (12). Cunha, M. M. R., 3 (29). Dantas, J. N. et al., 2 (75). Dario, F. R. et al., 1078 (56). Davidse, G. & D’Arcy, W. G., 10797 (39); 11610 (71); 12041 (87). Davidse, G. & Ramamoorthy, T. P., 10767 (98a).

492

List of Exsiccatae

Davis, P. H. & Sheferd, G. J. D., 60025 (72). Dawson, E. Y., 14397 (13). Dedecca, D. M., 388 (98a). Delfini, C. et al., 62 (98a); 83 (12); 100 (12); 107 (12); 497 (98a). Delprete, P. G. et al., 9610 (56). Deus, F.F., s.n. HUFU 13040 (43). Dias, B. J. et al., 66 (72); 68 (87). Dias, E. B. A. & Nogueira, L. M. G., 187 (56). Dias, E. B. A. et al., 149 (75); 223 (56); 259 (56); 359 (56); 367 (96); 448 (79); 558 (96); 615 (56); 724 (61); 725 (34). Dias, T. A., 544 (96). Dias, T. A. B. & Vieira, R.F., 152 (75); 153 (16); 164 (79). Dias, T. A. B. et al., 5 (56); 44 (98a); 111 (56); 115 (98a); 124 (72); 125 (79); 217 (79); 219 (75); 233 (56); 259 (56); 383 (43); 680 (98a). Duarte, A. P., 2494 (58); 2504 (39); 2638 (20); 2742 (58); 6499 (58); 6506 (39); 7642 (20); 8119 (39); 9971 (47); 10146 (96); 10765 (81); 10819 (39); 10853 (39); 11526 (39); 12412 (39). Duarte, A. P. & Barroso, G., 7878 (90); 7907 (36a); 7910 (38). Duarte, A. P. & Mattos, A., 8232 (72). Duarte, A. P. & Pereira, E., 8511 (90). Duarte, L., 783 (59). Dubs, B., 1818 (56). Duval, F. G. et al., 114 (96). Edwall, G., 5772 (98a). Eiten, G. & Eiten, L. T., 2586 (98a); 11028 (58). Eiten, G. et al., 2973 (98a). Emygdio, L., 2809 (95). Emygdio, L. & Lisboa, 2882 (50). Emygdio, L. et al., 3251 (39). Equipe do Jardim Botânico de Brasília, 918 (98a); 963 (75). Erendorfer, F., 2103 (82). Esteves, G. L. et al., CFCR 13372 (36b). Eunice et al., s.n. (96). Farah, F. T. & Freitas, C. A., 342 (68). Faria, C. A. et al., 146 (72); 206 (79); 216 (75). Faria, G. M., s.n. (98a). Faria, J. G. et al., 1 (96); 2 (96); 100 (79); 153 (75); 155 (16); 156 (79); 185 (72); 191 (72); 212 (79); 216 (75); 229 (98a); 255 (75); 263 (96). Faria-Jr., J. E. Q., 729 (5); 736 (24); 1122 (84); 1177 (56); 1234 (12); 3823 (72); 5376 (51); 5390 (36a); 5514 (20); 5531 (20); 5546 (36a); 5611 (90); 5741 (10); 5767 (77); 5860 (10), 6130 (77); 6134 (39). Faria-Jr., J. E. Q. & Moreira, H. J. C., 2032 (15). Faria-Jr., J. E. Q. & Otoni, T.J.O. 5472 (77); 5510 (51); 5658 (55); 5661 (31); 5792 (43); 5810 (43). Faria-Jr., J. E. Q. & Zanatta, M. R. V., 2766 (47); 2771 (79).

List of Exsiccatae

493

Faria-Jr., J. E. Q. et al., 405 (56); 651 (11); 731 (49); 769 (27); 862 (5); 1177 (56); 1275 (96); 1307 (42b); 1365 (58); 1458 (60); 3171 (56); 3298 (56); 3375 (56); 3775 (98a); 3959 (83); 3966 (42b); 3983 (101); 3985 (42b); 3986 (18); 5410 (51); 6106 (38); 6204 (51). Farinaccio, M. A. & Campos-Filho, E. M., 256 (59). Farinaccio, M. A. et al., 320 (32). Felfili, J. M. et al., 332 (3). Félix, L. P. 7725 (87). Fernandes, M. H. et al., 168 (98a). Fernandez, E. P. et al., 159 (11). Ferrari, J. M., 438 (12); 929 (43). Ferreira, s.n. (82). Ferreira, A. A. A., s.n. VIC 9341 (12). Ferreira, A. R., 415 (82). Ferreira, F. M., 45 (98a); 449 (98a). Ferreira, H. D. & Rizzo, J. A., 301 (29). Ferreira, M. B., 756 (98a); 766 (96); 1416 (30); 1481 (96). Ferreira, M. P., 7 (72). Ferreira, M. S. G. & Cardoso, F., 10 (56); 37 (81). Ferreira, R. M. & Ferreira, C. F., 45 (98a). Ferreira, V. S., 78 (89). Figueira, M. L. F. R., 21 (96). Filgueiras, T. S. & Lopes C. R. D., 2400 (29). Filgueiras, T. S. et al., 1757 (56); 1788 (96). Figueredo, S. et al., 92 (75). Flores, J. M. M., 4461 (32). Flores, T. B. et al., 441 (36b). Fonseca, M. L. & Alvarenga, D., 4621 (96); 4597 (96). Fonseca, M. L. & Barros, B. S., 836 (56). Fonseca, M. L. et al., 259 (96); 363 (29); 408 (69a); 428 (69a); 1016 (69a); 1061 (29); 1285 (7); 1444 (29); 1468 (98a); 1491 (84); 1876 (87); 1956 (84); 2437 (46); 2857 (87); 3370 (96); 3440 (96); 4348 (98a); 4405 (96); 4488 (98a); 5075 (22); 5095 (92); 5099 (49); 5104 (40); 5258 (61); 5380 (98a); 5621 (41); 5734 (56); 5810 (98a); 5832 (98a). Fonseca, N., 6 (96). Fonseca, S. G., 837 (98a). Fontelas, J. C. et al., 143 (23); 98 (90). . Fontes, C. G. & Barbosa, N. F., 207 (72). Fontes, C. G. et al., 109 (56); 195 (79). Fontella, G. 1522 (96). Forero et al., CFSC 8725 (58). Foster, W. et al., 913 (16). Forzza, R. C. et al., 631 (36a); 724 (20); 905 (84); 1149 (89); 1593 (63); 1607 (101); 1811 (53); 2129 (98a); 2535 (7); 4600 (69a); 6316 (98a); 6351 (50); 6403 (12). França, F. et al., 763 (41); 2845 (8); 4131 (8).

494

List of Exsiccatae

Freire, s.n. HGB 2038 (87). Freire, F. M. T. & Lopes, A. S., s.n. HUFU 13103 (87). Freire-Fierro, A. et al., CFCR 12730 (36b). Freitas, F. S. & Albuquerque, L.S., 445 (43). Funch, R. & Funch, L. S., 760 (19). Furlan, A. et al., CFCR 741 (2); 3368 (20). Furtado-Neto, A., 3 (56); 5072 (56). Ganev, W., 11 (6); 448 (87); 1154 (89); 1623 (62); 3095 (89); 3401 (87). Garcia, A., s.n. HUFU 59183 (98a). Garcia, R. J. F., 3470 (85). Garcia-Kirkbride, M. C., 1014 (56); 1054 (96); 1105 (56); 1162 (96). Gardner, G., 3152 (61); 3153 (33); 3154 (82); 3155a (33); 3155b (46); 3426 (82); 3721 (61); 3722 (24); 3723 (63); 3724 (63); 3725 (46); 3726 (81); 3727 (98a); 3729 (98a); 4132 (56); 4133 (36b); 4134 (98a); 4135 (46); 4136 (46); 4136 bis (46); 4137 (41); 4138 (46); 4139 (87); 4140 (41); 4141 (41); 4562 (38); 4563a (68); 4563b (90); 4564 (68); 4565 (53); 4566 (36b); 4568 (36a); 4569 (20); 4570 (12);4571 (77); 4573 (79); s.n. (46). Gates, B. & Estabrook, G. F., 10 (56); 23 (54); 140 (54); 191 (81). Gehrt, A., 165 (39); s.n. CEN 29268 (95). Gentry, A., 21447 (56). Gibbs, P. E. & Leitão-Filho, H. F., 4353 (98a). Gibbs, P. E. et al., 5128 (55). Giordano, L. C. & Toscano 572 (58). Ginzbarg, S. & César, P., 738 (80); 747 (44). Giroldo, A.B. et al., 233 (96). Giulietti, A. M., 1083 (5); 1084 (4). Giulietti A. M. et al., 588 (28); 589 (80); 1528 (6). Giulietti, A. M. et al., CFCR 1369 (12); 2481 (90); 3423 (2); 4464 (90). Giulietti A. M., CFSC 7316 (58). Glassman, S. F. & Gomes Jr., J. C. 8074 (58). Glaziou, A. F. M., 12421 (98a); 12675 (36b); 13818 (98a); 14699 (39); 14700 (50); 17617 (95); 17619 (95); 19166 (77); 19167 (58); 19168 (30); 19169a (30); 19169b (71); 19171 (38); 19172 (90); 19173 (50); 19174 p.p. (31); 19174 p.p. (38); 21413 (84); 21414a (43);21414b (72); 21415 (79); 21416 (60); 21417 (74); 21418 (96); 21419 (96); 21420 (47); 21422a (96); 21422b (50); 21423 (96); 21424 (75); 21425 (54); 21426 (81); 21548 (56); 21548a (104); s.n. (50); s.n. (54); s.n. (98a). Goldenberg, R. et al., 565 (95). Gomes, B. M., et al., 41 (98a). Gomes, S. M., 35 (96); 36 (96). Gonçalves, L. A., 1 (12). Gonçalves, M., 15 (72). Gontijo, T. A., s.n. BHCB 9220 (43). Gonzaga, D. R., 171 (95). Gottsberger, G. K. & Doring, 71-7386 (43); 111-25771 (41); 137-3386 (98a).

List of Exsiccatae

495

Graham, S. A., 860 (43); 861 (59); 865 (38); 874 (30); 878 (98a); 879 (20); 885 (77); 890 (43); 896 (96); 906 (98a); 939 (71); 949 (19); 954 (26); 955 (36a); 956 (90); 961 (90); 962 (30); 978 (36a); 980 (90); 981 (55); 982 (30); 985 (15). Grandi, T. S. M., 185 (12); 622 (98a); 904 (50); 1413 (58); s.n. BHCB 17991, CEN 19167 (58); s.n. BHCB 18110, CEN 19048 (39). Grandi, T. S. M. et al., s.n. BHCB 8094 (20); s.n. BHCB 12510 (12); s.n. BHCB 13277 (12). Groppo, M. Jr. et al., 1091 (39); 1099 (58). Guala, G. F. & Filgueiras, T. S., 1439 (72). Guardia, C. et al., 202 (9). Guarim-Neto, A. 1140 (82). Guarino, E. S. G. & Pereira, J. B., 106 (79); 131 (98a); 138 (79); 179 (75); 180 (96); 299 (72); 638 (96). Guarino, E. S. G. & Salles, P. A., 615 (56). Guedes, M.  L. et  al., 304 (19); 2049 (19); 12388 (89); 13320 (19); 17136 (19); 18851 (19). Guimarães, J. G., 10 (84). Guimarães-Neto, A. B., 14 (75) Haas, J. H. & Belém, 361 (5). Haas, J. H. et al., 312 (5). Haidar, R., 478 (98a). Hakme, N. A., 16 (72). Handro, W., 65 (43); 110 (96). Haridasan, M., 109 (96); 454 (96). Harley, R. M., 15122 (62); 15186 (62); 16697 (8); 19230 (8); 19512 (89); 19783 (62); 19944 (62); 21916 (87); 23017 (8). Harley, R. M. & Giulietti, A. M., 54410 (87); 54422 (87). Harley R. M. & Ribeiro Jr., E. 55896 (102). Harley, R. M. & Sano, P. T., 52310 (62). Harley, R. M. & Stannard, B., 26681 (102). Harley, R. M. et al., 11330 (56); 11458 (56); 15594 (62); 15743 (89); 15829 (89); 17006 (8); 24307 (89); 24402 (62); 24500 (62); 26083 (62); 26124 (89); 26682 (89); 27006 (89); 27056 (6); 27845 (89); 28587 (41); H50146 (89); H50391 (89); H50520 (89); H50666 (89); H51248 (62); 53755 (36b); 53995 (6); 54410 (87). Hashimoto, G., 57 (98a). Hastchbach, G., 27840 (38); 30204 (68); 31981 (98b); 33949 (56); 36374 (44); 36677 (84); 36737 (96); 36883 (101); 36895 (37); 38567 (35); 39434 (36b); 40834 (38); 40839 (38); 41409 (2); 41484 (26); 43828 (98a); 44725 (77); 47367 (62); 47420 (62); 51136 (98a). Hastchbach, G. & Ahumada, L. Z., 31244 (98a); 31252 (95); 31275 (39); 31484 (43); 31509 (95); 31630 (36a); 31687 (67); 31704 (77); 31716 (43); 33949 (56). Hastchbach, G. & Barbosa, E., 58271 (59); 58351 (4); 58380 (81); 59303 (35); 59326 (34); 67187 (15); 67290 (15); 67333 (15). Hastchbach, G. & Cordeiro, J., 51743 (43). Hastchbach, G. & Guimarães, 42273 (41).

496

List of Exsiccatae

Hastchbach, G. & Koczicki, C., 33061 (59). Hastchbach, G. & Kummrow, R., 56304 (45). Hastchbach, G. & Nicolack, V., 52956 (50); 53031 (55); 53748 (56); 53813 (61); 53883 (56); 53931 (81); 53933 (80); 53969 (5); 53972 (4); 53978 (54); 54233 (26); 54290 (2). Hastchbach, G. & Pelanda, P., 27794 (43); 28030 (90). Hastchbach, G. & Silva, J. M., 50304 (15); 50536 (87); 54503 (79); 54783 (18); 60152 (61); 64200 (43). Hastchbach, G. et al., 28919 (30); 28923 (78); 28931 (20); 29012 (38); 36257 (61); 36357 (80); 36396 (64); 36404 (10); 36407 (38); 36430 (30); 36488 (20); 36914 (61); 41195 (50); 50956 (67); 50966 (90); 51010 (20); 51067 (51); 51500 (38); 52969 (98a); 53082 (30); 53751 (64); 53996 (63); 54666 (3); 54781 (3); 54939 (68); 55006 (36b); 55060 (43); 55227 (43); 59477 (22); 55995 (61); 59523 (83); 59555 (69a); 59610 (69a); 59907 (56); 60250 (96); 60423 (63); 61811 (10); 61829 (43); 64063 (59); 64196 (68); 64204 (90); 64214 (58); 64581 (68); 64635 (43); 64794 (43); 66081 (36b); 66126 (64); 66154 (43); 66307 (30); 66450 (38); 66459 (77); 67228 (36b); 67251 (71); 67314 (36b);67539 (43); 67551 (90); 67911 (62); 69427 (15); 69503 (71); 69980 (56); 70003 (96); 70080 (56); 70284 (96); 70476 (27); 70485 (27); 70515 (64); 70634 (27); 70637 (35); 70654 (44); 70676 (34); 70862 (63); 70922 (61); 71131 (41); 71184 (41); 72217 (90); 73721 (36b). Hattori, E. K. O. et al., 305 (43); 484 (32). Hensold, N. et al., 3098 (90). Heringer E. P., 191 (98a); 1150 (98a); 3122 (56); 3354 (43); 3456 (43); 3468 (98a); 3746 (43); 3779 (98a); 4541 (79); 4859 (79); 4902 (79); 6808 (43); 7945 (56); 8935 (75); 9913 (47); 9938 (56); 10080 (56); 10082 (96); 10126 (75); 10422 (96); 10429 (98a); 10441 (79); 14449 (56); 10869 (56); 11264 (43); 11273 (56); 11313 (47); 11429 (79); 11438 (79); 11574 (47); 13176 (96); 13228 (79); 14429 (98a); 14449 (56); 14725 (75); 16664b (96); 17269 (72); 17283 (79); 17844 (56); 18307 (98a). Heringer, E. P. & Eiten, G., 14175 (56). Heringer, E. P. et al., 386 (96); 393 (56); 445 (98a); 563 (72); 1021 (96); 1409 (98a); 1853 (75); 2310 (35); 2547 (98a); 2613 (47); 3122 (56); 3466 (47); 3604 (96); 3779 (98a); 4234 (98a); 4541 (79); 4939 (75); 4859 (71); 4894 (72); 4902 (79); 7135 (56). Hind, D. J. N. & Queiróz, R. F., H50025 (89). Hoehne, F. C., 7 (98a); 16 (98a); 1025 (82); 1796 (98b); 1798 (98b); 1846 (98b); 4812 (82); 5473 (98a); s.n. CEN 25959, SPF 12247 (98a); s.n. CEN 25962, SPF 11038 (98a); s.n. CEN 29257, SP 20258 (98a); s.n. CEN 29272, SP 19286 (98a); s.n. CEN 29260 (98a). Honório, L. C. & Mendes, T. M., 9 (56). Horta, M. B. et al., 214 (12). Hunt, D. R. & Ramos, T.F., 5496 (98a); 5661 (48); 6303 (72). Hurbath, F., 138 (19).

List of Exsiccatae

497

Irwin, H. S. & Soderstrom, T. R., 5105 (75); 5135 (72); 6165 (72); 6385 (56); 6458 (56); 6517 (56); 7331 (98a); 7441 (56). Irwin, H. S. et al., 8122 (75); 8243 (79); 9222 (75); 9398 (81); 9717 (60); 9737 (64); 10724 (47); 10760 (84); 10934 (56); 11096 (96); 11219 (56); 11716 (56); 12246 (96); 12447 (80); 12636 (81); 12743 (56); 12928 (56); 13258 (64); 13690 (56); 13897 (79); 14154 (98a); 14171 (96); 14290 (98a); 14853 (36b); 15266 (98a); 15440 (75); 15479 (98a); 15496 (84); 15540 (72); 15725 (98a); 15726 (98a); 15857 (72); 16139 (48); 16524 (84); 16711 (56); 16800 (56); 16862 (84); 17699 (98a); 17786 (98a); 18037 (72);18504 (56); 18872 (56); 19226 (96); 19262 (36b); 19722 (95); 19996 (58); 20370 (58); 20941 (38); 21821 (38); 22004 (30); 22741 (38); 22974 (68); 23061 (43); 23070 (71); 23531 (71); 23692 (68); 23713 (43); 24231 (44); 24318 (5); 24321 (54); 24414 (4); 24793 (61); 24807 (22); 25251 (43); 25261 (59); 25445 (43); 25745 (43); 26115 (43); 26402 (96); 26883 (43); 26966 (43); 27462 (36a); 27672 (90); 27908 (51); 28203 (77); 28443 (38); 28623 (36b); 28627 (36b); 29628 (50); 29695 (98a); 30912 (62); 31343 (87); 31890 (96); 31900 (96); 32100 (44); 32190 (80); 32618 (8); 32760 (18); 33160 (44); 34016 (56); 34042 (36b); 34190 (56); 34260 (96); 34860 (56). Iglesias, J. O. V. et al., 149 (84). Isejima, E. M. et al., CFCR 5413 (90); 5537 (90). Jacques, E., 1299 (98b). Jacobi, C., 51 (58); 52 (39). Jesus, J. O., 7 (96). Jiménez, S. & Gutiérrez, E., 1307 (9) Joly, A. B. et al., CFSC 1 (58); 52 (39); 121 (58); 132 (58); 777 (20); 1080 (58); 1104 (58); 1122 (58); 1149 (20); 1153 (58); 1423 (58); 1517 (58); 1653 (39); 1766 (39); 2042 (39); 2307 (39); 2388 (58). Juchum, F. et al., 81 (62). Júlio, C. I., s.n. BHCB 12569, CEN 19117 (98a). Junior, F., 16 (75). Junqueira, D. I. et al., 129 (84); 152 (56). Kameyama, C. & Esteves, G. L., 30 (20); 38 (30); 39 (20); 40 (38); 41 (36b). Kassis, A. et al., s.n. OUPR 5357 (39). Kinoshita-Gouvêa, L. S. et al., 18983 (53). Kerr, W. E., s.n. CEN 38159, HUFU 5873 (98a). Killeen, T. et al., 4845 (9). Kirizawa, M., 108 (98a); 246 (53). Kirkbride Jr., J. H., 3735a (96); 3884 (96); 4731 (16); 4771 (98a); 4821 (72); 4854 (72); 5203 (56); 5273 (96); 5289 (98a); 5329 (79); 5542 (3); Kirkbride, M. C. G. & Kirkbride Jr., J.H. 1261 (98a); 3537 (72). Kral, R. et al., 72695 (26); 72917 (89); 75419 (71). Krapovickas, A. et al., 33328 (43). Krieger, L., 8283 (98a); 8621 (53); 14546 (98a); 14568 (39). Krieger, L. et al., s.n. CEN 18564 (53), CESJ 24316 (53). Kuhlmann, J. G., 1076 (82). Kuhlmann, M. & Kuhn, 1132 (98a).

498

List of Exsiccatae

Laboriau, L. & Valio, I. F. M., 1212 (43). Lanstryak, L., s.n. (26). Leenza, E. O. & Barbosa, A. A. A., 311 (68); 312 (68); 370 (68). Legre, M. G., s.n. (16). Leitão-Filho, H. F. et al., 14491(98a). Leme, R. O., 21 (72). Lemos-Filho, J. P., s.n. BHCB 29092 (30); BHCB 41165 (43). Leoni, L. S. & Silva, E., 5725 (98a). Lewinger, M. E. et al., CFSC 11284 (58). Lewis, G. P. & Morão, L. M. M., 1648 (30). Lima, A., 58-3002 (47); 58-2994 (96). Lima, A. A. A., 2 (30). Lima, A. & Cobra, L., 6 (98a). Lima, C. R., s.n. BHCB 16974 (58). Lima, E. S. et al., 597 (84). Lima, L. R. et al., 3 (90); 239 (90). Lima, R. et al., 2223 (8). Lima, N., 140 (96). Linhares, A. X., 11129-A (98a). Lira, R. G. & Noleto, W., s.n. (61). Lisboa, C. E., s.n. (98a); s.n. CEN 38145, OUPR 4763 (98a); Lisboa, M.  A., s.n. CEN 38139, OUPR 1778 (50); s.n. CEN 38140, OUPR 1817 (98a). Löefgren, A., 533 (98a). Löefgren, A. & Edwall, G., 2178 (59). Lombardi, J. A. A., 843 (95); 3373 (90); 4252 (38); 4303 (77); 4549 (43); 4556 (36b). Lombardi, J. A. A. et al., 3754 (90); 3757 (20); 9882 (92); 9912 (40); 10027 (74). Longhi-Wagner, H. M. et al., CFCR 9325 (36a). Lopes, M. A. & Andrade, P. M., s.n. BHCB 9109, CEN 19042 (39). Lorenzi, H., 4237 (36b); 5359 (98a); 5367 (56). Loureiro, D. M. et al., 568 (87); 643 (19). Lourteig, A., 1846 (72). Lovo, J. et al.19 (51); 326 (36a). Lucca, M., 44 (36b). Lucidoi, 22 (75). Luederwaldt s.n., CEN 29198, SP 14229 (98a). Lughadha, E. et al., 5955 (8). Lughadha, E. N. et al., H51051 (62). Lund, P. W., s.n. (50). Lutzelburg, P., 78 (89); 122 (89); 16046 (87). Macêdo, A., 57 (98a); 1048 (98a); 3497 (36b); 3586 (84); 3662 (29). Macedo, E. E., 57 (95). Macedo, J. C. R., s.n. (98a). Macedo, M. et al., 1062 (98a). Machado, A. I. M. R. 85 (20).

List of Exsiccatae

499

Machado, F. A., 29 (90). Magalhães, 1743 (36a). Magalhães, B., 9 (79). Magalhães, F. B., 18 (75); 24 (72). Maguire, B. et al., 49040 (58); 57090 (47). Maia-Filho, P., 11 (75). Mamede, M. C. H. et al., CFCR 3465 (39). Manso, s.n. herb. Fl. Bras. n. 217 (81); Mansur, T. & Tameirão-Neto, 91 (95). Mantovani, W., 654 (98a); 1365 (98a); 1676 (98a); 1865 (98a). Mantovani, W. &Sugyiama, M., 1829 (98a). Marcondes-Ferreira, W. & Berinello, R., 1248 (95); 1270 (95). Marcondes-Ferreira, W. et al., 806 (95); 1109 (95). Marinis, G., 1393 (98a). Marinis, G. & César, O., 18 (98a). Markgraf, F., 3426 (36b). Marques, D. et al. 70 (43); 358 (90). Marquete, R. et al., 2213 (56); 2222 (29); 3059 (53). Martens, L., 258 (95). Martinelli, G. 861 (58); 913 (58). Martinelli, G. et al., 11163 (26); 16565 (44); 16439 (65); 18215 (38); 18274 (77). Martins, C. R., 1227 (98a); 1336 (96); 3344 (98a); 1654a (96); 1654b (56); 2392 (56); 2651 (75). Martins, R. C., 80 (96); 190 (98a). Martins, R. C. et al., 502 (35); 596 (72). Martins, R.  P., 8448 (12); s.n. BHCB 8451 (12); s.n. CEN 19056, BHCB 8457 (98a); s.n. (BHCB 8490) (58). Martins, R. P. et al., 8442 (98a); 8457 (98a); 8470 (12). Martius, C. F. P. Herb. Fl. Bras. n., 217 (82); 865 (12); 866 (98a); 867 (39); 868 (95); 869 (85). Martius, C. F. P. obs. 424 (95); 706 (39); 707 (98a); 723 (95); 724 (95); 844 (12); 1338 (36a); 1350 (30); 1439 (98a); 1504 (38); Martius, C. F. P., s.n. (26); s.n. (36b); s.n. (38); s.n. (39); s.n. (53); s.n. (55); s.n. (62); s.n. (77); s.n. (87); s.n. (95); s.n. (98a). Mattos, A. et al. 9820 (60). Mattos, E. C. et al., s.n. BHCB 100306 (98a). Mattos, J. & Bicalho, H., 11495 (98a); 11496 (98a). Mattos, J., 9004 (98a); 12242 (98a); 12280 (98a). Mattos-Silva, L. A. et al., 2808 (19). Maury, C. M., 149 (98a); 157 (72); 161 (79). Mautone, L. et al. 691 (50). Mayo et al., CFCR 10438 (20). Mazine, F. F. et al., 913 (98a); 929 (98a). Meira, R. M. S. A. & Neto, J. A. A. M., 107 (50). Meira, R. M. S. A. et al., 31 (12).

500

List of Exsiccatae

Mellazo, A. F. O., 23 (68). Mello, C., 69 (81). Mello, F. N. A. et al., 18 (38); 263 (90); 470 (90). Mello-Barreto, H.  L., 3373 (95); 3380 (12); 8694 (58); 9366 (38); 9427 (90); 11355 (12). Mello-Barreto, H. L. & Brade, A.C., 1049 (20); 1052 (58); 1268 (58); 1270 (39). Mello-Filho, 3653 (25); Mello-Silva, R. & Ferreira, F. M., 2865 (39) Mello-Silva, R. & Forzza, R. C., 1748 (77); 2709 (36b); 2809 (62). Mello-Silva, R. & Pirani, J. R., CFCR 10761 (39); 10763 (39). Mello-Silva, R. et al., 218 (12); 286 (43); 404 (71); 405 (38); 416 (90); 487 (39); 496 (98a); 502 (39); 647 (36b); 727 (36b); 1329 (12); 1390 (12); 1458 (36b); 1487 (36b); 1973 (14);2248 (45); 2251 (18); 2382 (51); 2485 (38); 2558 (2); 2879 (12); 2894 (12); 2895 (58); 2914 (39); 2915 (98a); 2927 (39); 2930 (53); 3019 (36b); 3271 (53); 3183 (55). Mello-Silva, R. et al., CFCR 5494 (90); CFCR 9881 (26); 11810 (38). Mello, T. R. B. et al. 152 (90). Melo, E., 4807 (8). Melo, E. & França, F., 421 (96); 518 (96); 572 (56); 595 (56); 651 (98a); 673 (79). Melo, E. et al., 3342 (8). Melo, M. R. F. & Chiea, S.A.C., 194 (98a); 195 (98a). Mendes, M. S. & Neves, G. S., 174 (56). Mendes, L. C. & Nóbrega, M. G., 20 (75). Mendes-Magalhães, G., 1745 (12); 1750 (38); 4506 (98a); 4519 (39); 5444 (98a); 5976 (39); 17363 (12); 18389 (26); 19026 (98a); s.n. BHCB 68501 (12). Mendonça, R. C., 836 (98a); 921 (96); 955 (96). Mendonça, R. C. et al., 1203 (68); 1233 (43); 1244 (98a); 1259 (98a); 1314 (98a); 2244 (1); 2707 (18); 2788 (1); 2923 (18); 2963 (18); 3499 (72); 4635 (56); 4668 (56); 5999 (69a). Mendonça-Filho, C. V., 8 (39). Mendoza, J.M. et al. 4237 (80); 4267 (84); 4269 (11); 4294a (79); 4294b (96); 4296 (98a); 4310 (75); 4312 (96); 4322 (96); 4329 (96); 4749 (56); 4800 (47); 4871 (3); 4889 (56); 4913 (36b); 4908 (96); 5017 (44); 5021 (34); 5193 (3) Meneguzzo, T. E. C. et al., 106 (96); 110 (79). Menezes, C. S. L., 4 (96). Menezes, N. L., 1208 (3); 1209 (83). Menezes, N. L. & Oliveira, CFSC 11075 (58). Menezes, N. L. et al., CFCR 7681 (90); 10450 (10); 10566 (20); 10600 (77); 10617 (51); 10677 (36a). Menezes, N. L. et al., CFSC 7311 (39); 7318 (58); 11052 (58); 11258 (58). Menini-Neto et al., 714 (51). Messias, M. C. T. B. et al., 1395 (50); 1741 (50); 1791 (50). Mexia, Y., 5574 (98a); 5578 (43); 5741 (90); 5862 (55); 5874 (30); 5897 (98a); s.n. VIC 3260 (15).

List of Exsiccatae

501

Milhomens, L. C. et al., 62 (98a); 314 (72). Miranda-Silva, E. B. et al., 478 (6). Monteiro, P. L., s.n. CEN 1011 (96). Monteiro, R. F. et al., 225 (15) Moraes, A. et al., 12 (19) Moraes, E. C. C. et al., 105 (56). Moraes, L. I., s.n. CEN 76508 (25). Moraes, L. P. R., et al., s.n. CEN 74252 (25); s.n. CEN 76505 (56); s.n. CEN 76507 (25); 79026 (84); s.n. ESA 102752 (36b). Mori, S. & Benton, 13515 (62). Mori, S. & Boom, B., 14260 (19); 14289 (19); 14373 (19); 14404 (19); 14453 (8); 14509 (8). Mori, S. & Funch, R., 13383 (62). Mori, S. A. et al., 12451 (62); 12570 (19); 16947 (60). Mota, A. L. P. 2185 (43). Mota, R. C., 44 (95); 131 (95). Mota, R. C. & Viana, L., 585 (50). Moura, L. S. & Vieira, M. F., 385 (12). Moura, T. M. et al., 35 (25); 47 (56); 75 (25); 146 (98a); 153 (25); 389 (25). Munhoz, C. B. R., 14 (98a); 71 (56). Munhoz, C. B. R. & Ribeiro, J. F., 186 (1). Munhoz, C. B. R. et al., 332 (47); 492 (22); 1302 (98a); 1446 (79); 1538 (79); 2120 (81); 2204 (81); 2268 (81); 2748 (72); 2820 (72); 3749 (96); 3837 (56); 3973 (96); 4211 (56); 7185 (90); 7834 (1); 7965 (5); 7967 (72); 7966 (69a). Muniz, C. H. & Brochado, A. L., 67 (72). Nakajima, J. N. & Romero, R., 197 (95); 1664 (32); 1699 (59). Nakajima, J. N. et al., 237 (98a); 876 (59); 930 (95); 1113 (10); 1591 (14); 1941 (10); 1958 (10); 2242 (95); 2694 (14); 3281 (32); 3541 (32); 4005 (32); 4016 (95); 4091 (32); 4427 (10); 4636 (38); 4649 (38); 4863 (30). Nascimento, E. A., 110 (72). Nascimento, E. A. & Catharina, B., 54 (56); 56a (96). Nave, A. G., 1239 (56). Nave, A. G. et al., 1415 (98b). Neri, C. F., 13 (72). Netto, D. A. M., 5 (72); 17 (98a). Neves, J. L. G., 9 (96). Noblick, L. R. & Clodoaldo, 3758 (87). Nóbrega, M. G. et al., 694 (96); 713 (56); 1003 (56); 1173 (56); 1407 (56); 1507 (56). Nogueira, E. et al., 84 (36b); 139 (96). Nogueira, M. M. et al., s.n. BHCB 48592, CEN 46566 (90). Occhioni, P., 1827 (39); 1842 (58); 5151 (39). Ogasawara, H. A., 95 (89). Oldenburger , F. H. F. & Mecenas, V.V., 1600 (75). Oliveira, A. A. et al., 260 (89). Oliveira, F. C. A. et al., 265 (29); 1493 (33); 1580 (61).

502

List of Exsiccatae

Oliveira, G. C., 2139 (59). Oliveira, J.A. et al. 457 (65); 467 (45); Oliveira, J. E., s.n. BHCB 41712 (22). Oliveira, J. S. et al., 53 (56). Oliveira, O. L. C., 16 (56). Oliveira, P. A. M., 1635 (56). Oliveira, P. E. 1635 (56). Oliveira, P. I. & Anderson, W. R., 474 (61). Oliveira, R. C., 2889 (79). Oliveira, R. C. & Bertoluci, J., CFSC 11865 (58). Oliveira, R. C. & Silva, G. P., 585 (36b). Oliveira, R. C. et al., 872 (69a); 1142 (75); 1245 (76); 2690 (18). Onishi, E. et al., 69 (84); 866 (56). Pacheco, R. A. et al. 92 (32); 371 (14); 469 (32). Paiva, V. F. et al., 73 (72); 204 (56), 499 (75). Passos, S. R. D., s.n. (75). Pastore, J. P. B., 26 (98a); 104 (84); 237 (65); 1033 (98a); 1043 (72); 1044 (47); 1496 (96); 1497 (96); 1504 (98a); 1505 (79). Pastore, J. F. B. & Bringel Jr., J.B.A., 571 (84); 572 (56); 2553 (96); 2566 (79). Pastore, J. F. B. & Suganuma, E., 439 (11); 444 (92); 445 (5); 446 (22); 447 (4); 448 (24); 463 (56); 464 (96); 584 (72); 585 (79); 586 (56); 642 (80); 649 (72); 650 (3); 651 (83); 700 (47); 845 (96); 1096 (56); 1108 (37); 1386 (29); 1399 (45); 1436 (5); 1507 (96); 1508 (9); 1509 (72); 1523 (19); 1549 (41); 1907 (42b). Pastore, J. F. B. et al., 234 (99); 263 (96); 281 (96); 454 (56); 464 (96); 466 (49); 467 (74); 468 (22); 469 (56); 470 (44); 471 (92); 472 (80); 473 (92); 474 (61); 475 (54); 476 (56); 597 (16); 604 (75); 610 (16); 638 (75); 639 (79); 641 (16); 751 (47); 761 (36b); 777 (56); 784 (101); 787 (81); 789 (54); 791 (74); 838 (22); 839 (11); 840 (49); 841 (4); 843 (56); 844 (74); 867 (96); 883 (96); 930 (72); 933 (98a); 1006 (64); 1008 (60); 1023 (64); 1337 (17); 1338 (17); 1340 (56); 1851 (42b); 2080 (8); 2444 (82); 2460 (82); 2474 (81); 2485 (81); 2608 (8); 2612 (8). Paula, J. A. & Velten, S. B., s.n. BHCB 17857 (39). Paula, J. E., 32 (98a); 40 (98a); 59 (43); 3461 (98a). Paula-Souza, J. et  al., 4191 (36b); 4229 (36b); 4362 (81); 8443 (56); 8446 (56); 8840 (24). Pedralli, G. et al., 705 (43); 3292 (56); 3297 (84); 3318 (98a); 3366 (37). Pena, C. R., s.n. (75). Pena, E. C., s.n. (75). Pereira, B. A. S., 268 (98a); 539 (72); 560 (79); 1253 (47). Pereira, B. A. S. & Alvarenga D., 3105 (69a). Pereira, B. A. S. et al., 1456 (56); 2774 (37). Pereira, E., 647 (75); 1547 (38); 1580 (36a); 1594 (20); 7272 (12); 7402 (96); 8980 (43). Pereira, E. & Pabst, G., 2760 (30); 2814 (20); 2943 (12). Pereira, G. A. et al., 84 (79).

List of Exsiccatae

503

Pereira, J. B. & Moreira, G. A., 29 (40); 80 (23). Pereira, M., 3034 (50); 7285 (50). Pereira, M. & Lucca, M., 929 (58); 930 (36b); 931 (36b). Pereira, M. et al., 786 (36b); 787 (39); 790 (15); 932 (15); 933 (39). Pereira-Neto, A. B., 9 (75); 32 (79). Pereira-Neto, M., 144 (18). Pereira-Silva, G., 1817 (96); 2200 (56); 2215 (96); 2225 (96); 2278 (56); 2280 (96); 4098 (36b); 5493 (75); 7979 (79). Pereira-Silva, G. & Abdala, G.  C., 4251 (79); 4252 (96); 4273 (75); 4274 (79); 4275 (16). Pereira-Silva, G. & Bringel, J. B. A. 16875 (86); 16877 (43). Pereira-Silva, G. & Carvalho-Silva, M., 5208 (69b). Pereira-Silva, G. & Guarino, E., 6460 (69b); 6467 (72). Pereira-Silva, G. & Moreira, G.  A., 11170 (11); 11466 (40); 11493 (63); 11543 (40); 11600 (23); 11605 (23); 12847 (81); 12877 (23); 12882 (40); 14965 (76). Pereira-Silva, G. & Pereira, J. B., 953 (96); 5332 (1). Pereira-Silva, G. & Sampaio, A. B., 3884 (79); 3885 (96). Pereira-Silva, G. & Vieira, J. G. A., 40 (47). Pereira-Silva, G. et al., 1142 (72); 1143 (72); 1227 (72); 3358 (43); 4160 (72); 4181 (18); 4216 (29); 4230 (3); 4365 (69b); 4384 (11); 4402 (69b); 4412 (84); 4414 (11); 4416 (11); 4417 (69b); 4422 (45); 4537 (56); 4585 (23); 4645 (11); 4655 (23); 4718 (23); 4736 (23); 4990 (98a); 4991 (23); 5033 (27); 5038 (96); 5041 (72); 5045 (22); 5056 (72); 5057 (23); 5084 (69b); 5086 (84); 5200 (84); 5226 (23); 5305 (72); 5426 (11); 5427 (69a); 5871 (23); 5874 (11); 5884 (84); 5894 (11); 5898 (23); 5908 (23); 5936 (40); 5941 (45); 5947 (72); 5950 (40); 5951 (74); 5952 (11); 5954 (69a); 6062 (56); 6092 (98a); 6258 (98a); 6260 (43); 6306 (98a); 6492 (63); 6552 (40); 6773 (56); 6778 (56); 6843 (47); 7126 (56); 7157 (56); 7176 (96); 7285 (47); 7493 (47); 7561 (98a); 7905 (16); 7928 (60); 7940 (47); 7946 (96); 9066 (87); 10435 (63); 10519 (40); 10533 (23); 10536 (23); 10540 (11); 10552 (40); 10599 (63); 10594 (23); 10648 (40); 10762 (40); 11811 (11); 11813 (23); 11827 (23); 11828 (72); 11844 (69a); 11854 (72); 11887 (72); 12308 (63); 13467 (72); 13470 (72); 14133 (40); 14144 (23); 14910 (81); 15230 (23); 16432 (61); 16435 (5); 16445 (92); 16447 (84); 16449 (45); 16457 (92); 16463 (34); 16568 (61); 16569 (61); 16571 (27); 16574 (35); 16578 (44); 16684 (34); 16818 (8); 16863 (86); 16867 (86). Philcox, D. & Fereira, A., 3816 (56); 3971 (56); 4361 (84). Philcox, D. & Onishi, E., 4306 (96); 4782 (79); 4917 (98a). Pickel, D. B., s.n. (98a). Pinna, G. F. A. M. et al. 18 (26). Pinheiro, J. M. et al., 247 (43). Pinto, G., s.n. (ALCB (9232) (89). Pirani, J. R. & Devecchi, M.F. 6568 (43) Pirani, J. R. & Mello-Silva, R., CFCR 10811 (2); 11021 (30).

504

List of Exsiccatae

Pirani, J. R. et al., 1512 (64); 1685 (65); 1751 (3); 1886 (81); 1693 (18); 2085 (56); 2129 (56); 2255 (19); 2270 (57); 4084 (98a); 4367 (55); 5156 (15); 5233 (38); 9737 (60). Pirani, J. R. et al., CFCR 904 (2); 8721 (20); 12024 (36b). Pirani, J. R. et al., CFSC 6166 (39); 8014 (50). Pires, A., 3 (67); 76 (75); 230 (96); 327 (96). Pires, F. R. S. et al., 557 (26) Pires, M. J. P., 273 (96); 275 (96). Pires, M. J. & Mattos, A., 9820 (60). Pires, M. J. et al., 9005 (96); 9047 (98a); 9089 (56); 9225 (98a); 9656 (3); 9672 (98a); 9715 (84); 58096 (72); s.n. (98a). Pixis, 438a (50). Pohl, J. B. E., 93 (82); 589=D940 (59); 773 (98a); 801 (43); 990=D944 (47); 947 (98a); 1066=D937 (96); 1365=D945 (84); 1391 (98a); 1526 (43); 1745=947 (98a); 1885=D942 (29); 1988=D941 (40); 2018=D946 (69a); 2533 (72); 2537=D939 (25); 3156 (15); 3553=D943 (15); 5791 (56); 5792 (50); 5793 (43); s.n. (40); s.n. (47); s.n. (56); s.n. (72). Pompeu, M., 392 (95). Pompeu & Matoso 6 (43) Prado, J. et al., CFCR 12013 (39). Prance, G. T. & Silva, N. T., 58174 (75); 58184 (96); 58203 (22); 58242 (1). Proença, C. E. B., 47 (75); 754 (96); 873 (98a); 887 (96); 1670 (75). Proença, C. E. B. & Harris, S. A., 1217 (1); 2653 (75); 2769 (72); 3100 (30); 3123 (38); 3133 (10); 3380 (1); 3388 (1); 3389 (72); 3533 (84); 3852 (68). Proença, C. E. B. & Oliveira, R. S., 1416 (60); 1775 (72); 1945 (64). Proença, C. E. B. & Souza, M.L., 812 (96). Proença, C. E. B. et al., 1141 (80); 1676 (79); 1757 (87); 1809 (61); 1992 (56); 2151 (96); 2182ª (75); 2307 (56); 3777 (72). Proença, P. & Silva, M. A., 1196 (22). Queiroz, L. P. & Nascimento, 4258 (8). Queiroz, L. P. et al., 2754 (19); 4398 (89); 5197 (8); 6076 (36b); 14128 (46); 14358 (98a); H51085 (62). Ramos, A. E., 397 (79). Ramos, A. E. & Barros, G. V., 840 (88). Ramos, A. E. et al., 511 (98a); 1294 (79); 2236 (72); 4127-1 (98a). Ratter, J. A., 2867 (98a); 3472 (72); Ratter, J. A. & Castro, R. A., 163 (56). Ratter, J. A. & Fonsêca, S. G., 2763 (96); 2831 (79); 2855 (79). Ratter, J. A. & Ramos, J., 374 (84). Ratter, J. A. et al., 829 (84); 965 (84); 1361 (48); 1574 (48); 2267 (56); 3133 (75); 4032 (56); 4075 (56); 4076 (82); 4176 (56); 7273 (96); 7315 (37). Rawitscher, F. s.n. SPF 147413 (98a). Reeves, R. D., 2864 (56); 2927 (29); 2946 (29); 2969 (29); 3031 (61). Regnell, A. F., 55 (59); 116 (95); 142 (98a). Reichardt, C. V., 110 (60); 111 (60).

List of Exsiccatae

505

Reis, G., 4 (56). Reis, M. A., 5091 (98a); 5094 (12); 5095 (12); 5097 (98a). Resende, S. G. et al., 2306 (56). Rezende, A. R., 339 (98a). Rezende, J. M., 322 (79); 358 (56). Rezende, J. M. et al., 843 (56). Ribeiro, J. F & Haridasan, M., 851 (75). Ribeiro, T., 344 (87). Rissi, M. N. et al. 623 (61); 661 (69a). Riedel, L., 749 (82); 909 (90); 1216a (38); 1216b (38); 2595 (72); 2596 (59); 2597 (95); s.n. (12); s.n. (38); s.n. (58); s.n. (72); s.n. (90); s.n. (95); s.n. (98a). Rizzo, J. A., 16 (84); 4041 (56); 6960 (56); 6985 (84); 7374 (54); 7445 (84); 7627 (84); 7645 (56); 7785 (80); 7861 (56); 7895 (84); 7962 (80); 7978 (61); 8017 (98a); 8061 (80); 8093 (98a); 8103 (84); 8156 (72); 8560 (18); 8853 (56); 9051 (60); 9102 (60); 9335 (56); 9744 (84); 9782 (84); 9828 (84); 9838 (98a); 9845 (98a); 9877 (61); 9904 (88); 10407 (84); s.n. (98a). Rizzo, J. A. & Barbosa, A., 66 (84); 123 (84); 135 (56); 150 (56); 497 (98a); 730 (56); 754 (98a); 1327 (84); 1443 (84); 3415 (56); 3478 (56); 3499 (84); 3589 (84); 3923 (84); 4242 (83); 4799 (56); 4867 (56); 4927 (79); 4967 (98a); 4994 (98a); 5012 (84); 5089 (98a); 5093 (98a); 5113 (25); 5153 (98a); 5206 (25); 5327 (25); 5413 (25); 5852 (56); 5883 (36b); 5912 (56); 5933 (56); 5998 (36b); 6050 (56); 6113 (56); 6188 (36b); 6388 (84); 6517 (84); 7497 (56); 7516 (84); 7677 (56); 7733 (56). Rizzo, J. A. & Ferreira, H. D., 10239 (56);10283 (41) Robert, A., 571b (59). Rocha, D. M. S., S2P1 s.n. (98a). Rodrigues, A. C. M., 10 (75). Rodrigues, A. S. & Chesini, F. L., 191 (83); 194 (65). Rodrigues-Silva, R. et al., 267 (98a). Romaniuc-Neto, S. & Sajo, M. G., 342 (22); 380 (83); 388 (60); Romaniuc-Neto, S. et al., 449 (3); 605 (44). Romão, G. O., et al., 2505 (90). Romero, R., 1820 (43). Romero, R. & Nakajima, J. N., 6014 (51). Romero, R. et al., 911 (95); 1009 (25); 1727 (59); 1866 (10); 2126 (10); 2509 (10); 2980 (14); 3108 (30); 3233 (14); 3829 (95); 3994 (98a); 4218 (10); 6275 (32); 6606 (14); 6872 (32); 6917 (10); 7000 (73); 7033 (32); 7394 (32); 7628 (32); 7852 (32); 8347 (90). . Roque, N. et al., CFCR 14789 (89); 1552 (19); 1957 (19); 1995 (19); 2791 (19); 4199 (71); 4240 (19); 4677 (14); 4968 (59); 6275 (32); 15001 (71). Rosa, P. O., 1297 (25). Rosa, P. O. et al., 900 (12); 950 (52); 1073 (55); 1081 (36b). Roschel, M. B., 623 (12). Roschel, M. B. & Guimarães, R., 716 (36b). Rossi, L. et al., CFCR 3078 (20); 5766 (39).

506

List of Exsiccatae

Roth, L., 2219 (95); 2220 (12); 2232 (98a). Roveratti, J., 426 (79); 753 (96). Roveratti, J. & Faria, C. A., 365 (75). Roveratti, J. et al., 174 (96); 282 (96); 304 (96); 671 (56); 692 (96); 722 (56); 746 (96); 777 (96); 800 (96); 836 (98a); 857 (96). Rua, G. H., 638 (84); 901 (98a). Rua, G. H. & Oliveira, R. C., 629 (69a); 639 (72). Russel, A., 322 (98a). Sá, K. et al., 1701 (12). Saavedra, M. M. et al., 391 (60); 471 (84); 501 (61). Saddi, N., 36 (82); 4483 (82); 4602 (56); 7230 (82); 7637 (56); 7646 (98a); 9171 (84); 9213 (56). Saint-Hilaire, A., s.n. catal. B1 n° 1174 (90); B1 1250 (98a); B1 1336 (43); s.n. (25); s.n. (36b); s.n. (42a); s.n. (75); s.n. (90); s.n. (103). Saiter, F. Z. 403 (36b); Sakuragui, C. M., 34 (58). Sakuragui, C. M. et al., CFCR 15119 (55); 15172 (68); 15174 (43). Salatino, A. et al., 28 (58). Salatino, M. L. et al., CFCR 12126 (20). Salimena-Pires, F. R., s.n. CEN 18566, CESJ 25451 (53). Salles, P. S. B. A., 26 (75). Sampaio, A. B., 346 (96). Sampaio, A. B. et al., 76 (56); 102 (56); 214 (65). Salino, A. & Mota, R. C., 6993 (98a). Sano, P. et al., 928 (98a). Sano, P. et al., CFCR 14850 (62). Sano, P. & Laessoe, T., H52343 (89). Santiago, A. F., 25 (96). Santos, A. A., 96 (96); 1008 (96); 1015 (96); 1243 (75); 1281 (72). Santos, A. A. & Pereira, J. B., 2575 (56). Santos, A. O. R. & Bartelli, B. F., s.n. HUFU 69909 (98a). Santos, E., 1621 (75). Santos, E. & Sacco, J., 1648 (98a). Santos, E. R. & Pereira, C. B., 328 (56). Santos, E. R. et al., 1054 (11). Santos, H. G. P. et al., 1 (98a); 2 (25); 8 (96); 42 (98a); 63 (98a); 106 (25); 220 (25); 242 (98a); 445 (98a); 509 (25). Santos, J. R. & Moreira G. A., 134 (56); 174 (96); 183 (79). Santos, J. R. et al., 1 (75). Santos, R. H., 33 (98a). Santos, T. R. R. et al., 46 (84). Santos, 370 (98b); 413 (98b). Saraiva, D. P. et al., 249 (98a); 307 (18). Saraiva, L. C., 19 (98a). Sartin, R. D., 36 (98a).

List of Exsiccatae

507

Sasaki, D. & Sasaki, C., 879 (95). Sasaki, D. et al., 234 (32); 306 (95); 425 (98a). Sastre, C., 1109 (96). Sato, M. N., 3 (96). Savassi-Coutinho, A. P. et al., 1052 (38); 1319 (15). Scariot, A. O. et al., 485 (87); 525 (36b); 783 (33). Scatena, V. L. et al., CFCR 10477 (30); 10501 (20). Schechtmann, D. F., s.n. (98a). Schiavini, I., s.n. (HUFU 13065). Schüch, G., 5966 (95); s.n. (39). Schwacke, C. A. W., 6435 (12); 7733 (50); 8259 (38); 8260 (36b); 8261 (71); 8262 (38); 8263 (77); 8264 p.p. (38); 8264 p.p. (90); 8265 (58); 8266 (31); 9369 (15); 18802 (95); 12410 (53); s.n. (95). Sciamarelli, J. et al., 1299 (98b). Sellow, F., 194 (82); 2053 (39); 2054 (12); s.n. (12); s.n. (39); s.n. (50); s.n. (95); s.n. (98a). Semir, J. & Joly, A. B., CFSC 3815 (58). Semir, J. & Sazima, M., CFSC 2046 (58); 2056 (58). Semir, J. et al., CFSC 4047 (58); 4159 (39). Sena, T. S. N. et al., 1675 (36b). Senna, L., s.n. (75). Sevilha, A. C. et al., 2130 (43); 3418 (11); 3541 (63); 3561 (63); 3761 (61); 3774 (40); 4009 (63); 4031 (61); 4039 (63); 4229 (71); 4328 (77); 4349 (71); 4421 (71); 4646 (71); 5145 (91); 5944 (71); 5979 (26); 6036 (71); 6180 (71); 6354 (55); 6362 (71); 6371 (77), 6470 (71); 6523 (55); 6673 (71); 6795 (55); 6823 (71); 6833 (55); 6866 (55), 6876 (100); 6969 (98a); 6974 (71); 6985 (91); 7020 (91); 7066 (98a); 7096 (71). Shepherd, G. J. et al., 3930 (78). Silva, A. C. B. 125 (32) Silva, A. F. et al., 451 (20); 496 (58); 1649 (39); 2536 (98a). Silva, C. I., s.n. HUFU 50009 (43). Silva, E. A, et al., 9 (43). Silva, E. H. & Mendonça, C.V., 38 (20). Silva, J. G., 1874 (72). Silva, J. S., 662 (92); 665 (37). Silva, M. A., 64 (56); 221 (87); 374 (96); 1134 (72); 1135 (79); 1874 (72); 2207 (98a); 4701 (96); 4815 (96); 4962 (96); 5514 (16); 5649 (56); 7580 (96). Silva, M. A. & Jesus, G. N., 2937 (98a). Silva, M. A. & Ferreira, C. C. S., 3160 (29). Silva, M. A. & Pinheiro, F. C., 7024 (96); 7585 (96). Silva, M. A. & Santos, A. J. V., 3203a (18); 3203b (81); 3284 (1). Silva, M. A. & Vianna, R. L. Jr., 317 (96). Silva, M. A. et al., 1319 (41); 1325 (41); 1435 (41); 1924 (44); 1937 (37); 1948 (65); 1957 (3); 2005 (80); 2137 (22); 2163 (22); 2177 (96); 2267 (22); 2396 (1);

508

List of Exsiccatae

2907 (29); 2944 (29); 3160 (29); 5049 (75); 5135 (56); 5264 (84); 5459 (96); 5461 (56); 5692 (56); 5694 (96); 5742 (75); 6150 (29); 7306 (84); 7689 (90). Silva, M. P., 3 (98a). Silva, R. et al., 1684 (12). Silva, S. B. & Viegas, D. X., 349 (87). Silva, W. M., s.n. OUPR1807 (58); OUPR 22708 (58); s.n. OUPR 22917 (30); s.n. OUPR 22918 (58). Silva-Manso, A. L. P. 69 (82). Silveira, F. A., 75 (43). Silveira, V. M., 26 (39); 938 (53). Simão-Bianchini, R. et al., 171 (12); CFCR 8784 (90); 455 (94). Simon, M. F. et al., 273 (87); 1941 (95); 2333 (98b); 2526 (71); 2529 (21); 2551 (45); 2568 (11); 3079 (97); 3080 (86). Smith, L. B., A-51 (98a); 6954 (98a); 7022 (39); 15071 (98a). Smith, L. B. et al., 6880 (39); 6734 (98a); 7024 (58); 7912 (98a). Siniscalchi, C.M. et al., 489 (61). Soares, E. A. et al., 839 (88). Sobral, M. & Alves, T. M. A.13046 (98a). Sobral, M. et al., 13080 (12); 13860 (12); 14050 (98a); 14091 (53); 15609 (53). Soto, A. et al., 365 (9); 427 (9); 463 (9). Sousa, A., s.n. BHCB 16071 (50). Sousa, G., 406 (98a). Sousa, H. C. et al., s.n. BHCB 16197 (58); s.n. BHCB 16220 (58). Sousa-Silva, S. et al., 275 (56). Souza, E. B. et al., 883 (19); 1065 (15). Souza, H. C., s.n. BHCB 17131, CEN 19103 (98a). Souza, H. C. & Andrade, P. M., 114 (53). Souza, J. P. et al., 190 (12); 4229 (36b); 4362 (81); 4497 (81); 4487 (17); 5288 (62); 5617 (71). Souza, M. A. B., s.n. (96). Souza, M. R. et al., 1 (43). Souza, M. F. L., 139 (56). Souza, V. C. & Abbud, P. V., 12331 (58); 33239 (58). Souza, V. C. & Sakuragui, C. M., 2005 (53); 3349 (20); 3404 (39). Souza, V. C. & Souza, J. P., 22003 (43); 22078 (43); 22283 (43); Souza, V. C. et al., 190 (12); 1489 (50); 1940 (82); 5049 (32); 5528 (71); 5539 (55); 7912 (98a); 7913 (95); 8182 (58); 8229 (20); 8422 (36a); 8440 (20); CFCR 8786 (38); 11481 (95); 11599 (39); 11810 (58); 15042 (84); 16322 (43); 16844 (98b); 17617 (9); 17830 (98b); 17863 (9); 18014 (56); 20865 (43); 20875 (38); 20900 (10); 21226 (95); 21536 (72); 21113 (20); 22217 (30); 23096 (87); 23943 (84); 24079 (61); 24735a (37); 24735b (41); 25224 (20); 25349 (38); 25791 (39); 27791 (39); 25818 (36b); 25835 (26); 25889 (26); 25955 (87); 26936 (50); 28058 (50); 28225 (55); 28386 (20); 29654 (90); 30119 (90); 32764 (90); 32953 (71). Splett, S., 215 (36b); 635 (67); 812 (38); 858 (90); 908 (20); 940 (20); 1028 (20).

List of Exsiccatae

509

Splett, S. & Gröger, 656 (51). Stannard, B. et al., s.n. (PCD 4914) (89); 2561 (8). Stehmann, J.  R., II-34 (39); s.n. BHCB 20931 (90); s.n. BHCB 21085 (90); s.n. BHCB 21456 (12). Stehmann, J. R. & Oliveira, A. R., s.n. BHCB 15703, CEN 19125 (53). Stehmann, J. R. et al., 2439 (98a); 2896 (20); s.n. BHCB 18870 (58). Stephan, E., s.n. (50). Stevens, A. D., 16 (56). Strang, H., 81 (50); 84 (43); 85 (50); 87 (43). Sucre, D., 271 (75); 409 (96); 458 (98a). Sucre, D. et al., 7255 (53). Sugyiama, M. & Chiea, S. C., 423 (53). Sugyiama, M. & Mantovani, W., 12 (98a); 163 (98a). Tamashiro, J. Y. et al., 17387 (55). Tamberlick, E., s.n. (56). Tameirão-Neto, E., 54 (98a); 419 (50); 1977 (43); 2335 (43); 2877 (95); 2951 (98a); 3925 (53); 4017 (36b); 4018 (55); 4457 (36b). Tameirão-Neto, E. & Werneck, M.S., 1020 (98a); 1021 (43). Taxonomy Class of the Universidade de Brasília, 112 (75); 362 (96); 4635 (79). Taylor, N. P. et al., 1452 (68); 1503 (36b). Teixeira, W. A., s.n. BHCB 25022 (12); s.n. BHCB 25023 CEN 18967 (50); s.n. BHCB 26302 CEN 18948 (98a). Teixeira, E. M. & Brina, A. E., s.n. BHCB 832 (36b); s.n. BHCB 35831 (98a). Tenório, E. C. & Freitas, L. C., s.n. CETEC 01077 (36b). Thomas, W. W. et al., 5762 (56); 5860 (43); 5945 (68); 5948 (43). Thomé, M. L. M., 13 (84); 81/02686 (79). Tiritan, O., 584 (98b). Toledo, C. B. et al., 104 (56); 113 (83); 294 (81). Trinta, Z. A. & Fromm, E., 574 (98a); 672 (95). Tsuji, R. et al., 1232 (71). Ule, E., 75[2929] (7); 2562 (39); 2563 (50); 7111 (89). Uliana, V. L. C. R. et al., 695 (37). Usteri, A., s.n. CEN 29196, SP 14225 (98a). Vale, G. D. et al., 19 (75); 395 (56); 389 (96). Valente, G. E. et al., 1226 (50). Valls, J. F. M. et al., 13070 (41); 13684 (98a). Varnay, L. et al., 1262 (38); 1265 (38). Vasconcelos, M. F., s.n. BHCB 40209, CEN 33603 (90); s.n. BHCB 37892, CEN 33612 (98a). Vauthier, M., 420 (30); 421 (43); 423 (50). Vaz, A. F., 784 (72); 882 (5). Verboonen, S. M. et al., 10 (84); 24 (56); 45 (56); 71 (56); 97 (98a); 163 (56). Verdi, M. et al., 6734 (58); 6808 (38); 6897 (38); 6834 (20); 6897 (38); 7007 (77); 7017 (38); 7029 (38); 7073 (39); 7236 (18); 7259 (18); 7282 (18); s.n. HUFU 13186 (58). .

510

List of Exsiccatae

Viana, P. L. et al., 4253 (87). Viana, R. H. O., 126 (56); 302R (84). Vidal, J., I-524 (95); I-713 (94); II-6351 (43); II-6369 (43); s.n. (39); s.n. (90). Vidal, J. R. B. et al., 64 (75). Viegas, A. P., 6252 (98a); s.n. SP 42029 (98a); s.n. CEN 25498 SP 48763 (98a). Viegas, A. P. & Toledo, J. F., 8045 (98a). Vieira, M. F. & Hermano, 560 (43). Vieira, M. F. & Reis, H. S., 560 (43). Vieira, R. F. & Pereira, J. B., 741 (98a); 775 (96). Vieira, R. F. & Silva, G., 1163 (36b); 1165 (36b). Vieira, R. F. & Walter, B. M. T., 750 (75); 761 (98a). Vieira, R. F. et al., 469 (98a); 594 (96); 611 (98a); 619 (96); 717 (84); 750 (75); 774 (98a); 775 (96); 833 (84); 1183 (46); 1205 (96); 1206 (56); 1255 (9); 1777 (98a). Volpi, R. L., et al. 382 (32), 665 (32). Walter, B. M. T. & Bridgewater, S., 2140 (84). Walter, B. M. T. et al., 60 (98a); 133 (43); 565 (98a); 789 (47); 792 (84); 962 (18); 1079 (98a); 1106 (84); 1209 (98a); 1236 (84); 1269 (98a); 1302 (98a); 1306 (69a); 1358 (72); 1380 (69a); 1422 (98a); 1436 (72); 1439 (98a); 1536 (72); 1617 (61); 1848 (72); 1891 (72); 2698 (98a); 3008 (45); 3038 (56); 3083 (56); 3165 (84); 3283 (47); 3290 (84); 3494 (18); 3626 (18); 3631 (61); 3643 (3); 3646 (65); 3647 (81); 3769 (29); 3771 (29); 3855 (98a); 4071 (18); 4165 (29); 4177 (69a); 4187 (69a); 4194 (84); 4203 (29); 4421 (36b); 4549 (69b); 4564 (11); 4791 (84); 4833 (43); 5009 (43); 5120 (43); 5198 (43); 5273 (88); 5862 (75); 6130 (96); 6220 (96); 6346 (96); 6489 (98a); 6671 (82); 6768 (56). Wanderley, M. G. L. & R. Kral, 1443 (36b). Wanderley, M. G. L. et al., 573 (20). Warming, E., s.n. (95); s.n. (43). Weddell, H. A., s.n. (43); 1558 (12); 1860 (68); 2844 (45). Werneck, W. L., 840 (34). Wied-Neuwied, M. A. P., s.n. (71). Williams, L.  O. & Assis, V., 5996 (43); 6127 (50); 6491 (98a); 6745 (50); 7044 (98a); 7241 (50). Without collector, OUPR 4724 (12); OUPR 4728 (12); SP 18801(50). Xavier, H. V., 7 (75). Xavier-Vieira, E. M., 5 (79). Yale Dawson, E., 14731 (7). Zanatta, M. R. V. & Araujo, R. C., 193 (72). Zanatta, M. R. V. & Carvalho, A. M., 1188 (56); 1195 (96). Zanatta, M. R. V. & Faria, J. E. Q., 1350 (75). Zanatta, M. R. V. & Fernandes, J. B., 237 (56). Zanatta, M. R. V. & Silva, J. S., 964 (75); 1238 (96); 1381 (79). Zanatta, M. R. V. et al., 518 (79); 685 (75); 812 (56); 833 (98a). Zappi, D. C. & Kameyama, C., CFSC 9636 (43); 9654 (58); 9853 (39). Zappi, D. C. & Salatino, A., CFCR 10388 (38).

List of Exsiccatae

511

Zappi, D. C. & Scatena, V. L., CFCR 10922 (12). Zappi, D. C. et al., 158 (96); 704 (58); 830 (93); 1721 (43); 2295 (15); 2733 (93); 2759 (43); 2794 (15). Zappi, D. C. et al., CFCR 9294 (90); 9449 (38); 9931 (39); 10040 (20); 10376 (20); 10381 (77); 10451 (38); 10472 (50); 10601 (10); 10927 (20); 11187 (12); 12879 (36b). Zurlo, M. A. & Silva, J. L., s.n. OUPR 26250 (39).

Index1

D Diplodon, 3, 91 Diplusodon adpressipilus, 7, 12, 13*, 57, 78, 88, 109, 112–114, 115*, 123*, 125, 126, 468, 481 aggregatifolius, 57, 78, 88, 98, 108, 114, 116, 117*, 118, 123*, 301, 303, 481 alatus, 11, 16*, 49, 57, 64, 65, 78, 88, 101, 113, 118, 119, 120*, 123*, 159, 249, 468*, 481 alutaceus, 188, 218, 220, 250, 256 amoenus, 437 angustifolius, 228, 232 LT appendiculosus, 6, 12, 14, 18, 20, 41, 45, 57, 71, 78, 88, 101, 119, 121, 122*, 123*, 124, 129, 152, 216, 260, 301, 481 argenteus, 6, 7, 57, 78, 88, 109, 114, 115*, 124–126, 134*, 481 argyrophyllus, 12, 18*, 57, 78, 88, 97, 109, 114, 125, 126, 127*, 134*, 481

 New names and combinations are in boldface; synonyms are in italics. Page numbers in boldface indicate primary page references; those with an asterisk (*) indicate pages with illustrations or maps. New lectotypfications are indicated by boldface (LT) after the page number. New neotypifications are indicated by boldface (NT). 1

astictus, 4, 5, 11, 20, 21, 41, 57, 71, 77, 78, 88, 100, 124, 128, 129, 130*, 134*, 357, 419, 481 bahiensis, 18*, 57, 78, 88, 109, 129, 131, 132*, 133, 134*, 137, 237, 277, 428, 452, 458, 481 bolivianus, 57, 75, 78, 88, 98, 105, 133, 135, 136*, 137, 147*, 256, 448, 474, 481 bradei, 4, 6, 11, 12, 13*, 18, 21, 22, 30, 31, 57, 64, 78, 88, 97, 107, 137–139, 140*, 147*, 256, 326, 357, 468*, 481 LT burchellii, 7, 57, 78, 88, 102, 103, 139, 141, 142*, 143, 147*, 468*, 481 buxifolius, 6, 11, 16*, 57, 78, 88, 99, 101, 143, 144, 145*, 147*, 148, 238, 351, 425, 470*, 481 LT var. naudinii, 4, 274, 277 caesariatus, 4, 6, 57, 78, 88, 98, 149, 150*, 151, 160*, 481 canastrensis, 57, 78, 88, 96, 151, 152, 153*, 160*, 398, 400, 481 candollei, 5, 13*, 15, 20, 22, 24, 25*, 26*, 27, 42, 57, 64, 78, 88, 95, 152, 154, 155*, 156, 160*, 415, 469*, 481 LT candollii, 412 capitalensis, 57, 78, 88, 107, 156, 157, 158*, 159, 160*, 481 capitatus, 6, 57, 84, 88, 108, 459, 460*, 461*, 481 chapadensis, 57, 78, 88, 95, 104, 159, 161*, 162, 173*, 481

© The Author(s), under exclusive license to Springer Nature Switzerland AG 2021 T. B. Cavalcanti, The Genus Diplusodon (Lythraceae), Flora Neotropica, https://doi.org/10.1007/978-3-030-65875-5

513

514 Diplusodon (cont.) ciliatiflorus, 9, 12, 57, 65, 71, 78, 88, 102, 162, 163, 164*, 165, 173*, 469*, 481 ciliatus, 35, 58, 79, 88, 100–102, 165, 166, 167*, 168, 173*, 481 ciliiflorus, 5, 11, 16*, 28, 58, 79, 88, 102, 168, 169, 170*, 171*, 173*, 174, 287, 481 LT conduplicatus, 24, 58, 79, 88, 95, 96, 174, 175, 176*, 185*, 481 cordifolius, 6, 9, 18, 34*, 58, 79, 88, 94, 175, 177, 178*, 179, 185*, 274, 282, 333, 334, 481 crulsianus, 7, 55, 56, 372, 377 cryptanthus, 58, 71, 79, 88, 99, 179, 180, 181*, 185*, 263, 481 decussatus, 7, 12, 19*, 21, 58, 79, 88, 103, 143, 182, 183*, 184, 185*, 481 divaricatus, 7, 12, 13*, 39, 58, 79, 88, 96, 184, 186, 187*, 188, 196*, 216, 377, 481 LT epilobioides, 6, 12, 13*, 18*, 22, 24, 58, 79, 88, 95, 188, 189, 190*, 191, 196*, 405, 415, 416, 481 LT ericoides, 5, 58, 79, 88, 96, 191, 192, 193*, 196*, 337, 481 fastigiatus, 7, 58, 79, 88, 107, 192, 194, 195*, 196*, 481 floribundus, 7, 58, 79, 88, 110, 194, 197, 198*, 199, 208*, 319, 434, 435, 482 LT foliosus, 315, 319 glaucescens, 7, 11, 19*, 21, 22, 24, 25*, 34*, 51, 58, 65, 79, 88, 99, 103, 199, 200, 201*, 202, 208*, 308, 371, 469*, 482 LT glaziovii, 5, 42, 58, 79, 88, 95, 203, 204*, 205, 208*, 232, 482 glocimarii, 58, 79, 88, 95, 109, 114, 126, 205, 206, 207*, 208*, 209, 482 gracilis, 5, 18*, 58, 79, 88, 94–96, 101, 129, 209, 212*, 211, 223*, 482 var. ulei, 128, 129 grahamae, 58, 79, 88, 94, 211, 212, 213*, 214, 223*, 455, 469*, 482 hatschbachii, 6, 12, 58, 79, 88, 110, 214, 215*, 216, 223*, 482 helianthemifolius, 6, 13*, 14, 18*, 22, 24, 34*, 58, 77, 80, 88, 97, 108, 216–218, 219*, 220, 223*, 225, 482 f. striatifolius, 220, 225, 351 LT var. pemphoides, 9, 14, 18*, 58, 75, 76, 80, 88, 106, 217, 218, 220, 222*, 223*, 225, 351, 380, 482

Index heringeri, 7, 9, 22, 24, 25*, 58, 80, 88, 106, 110, 112, 226, 227*, 228, 241*, 245, 482 LT hexander, 5, 16*, 20, 22, 24, 42, 59, 64, 80, 88, 95, 205, 228, 229, 230*, 232, 241*, 470*, 482 LT f. angustifolius, 228, 232 hirsutus, 6, 18*, 47, 49, 52*, 55, 56, 59, 64, 77, 80, 88, 97, 108, 133, 137, 232, 233, 234*, 237, 238, 241*, 277, 278, 474, 482 LT hirtellus, 274 humilis, 6, 428, 434 LT imbricatus, 7, 33, 34*, 37, 59, 71, 80, 89, 99, 143, 184, 238, 239, 240*, 241*, 242, 340, 406, 482 LT incanus, 6, 12, 18*, 59, 80, 89, 98, 105, 109, 242, 243, 244*, 245, 252*, 482 var. hirtellus, 242, 245, 246 var. serpyllum, 242, 245, 246 irwinii, 6, 256, 259–261 kielmeyeroides, 3, 6, 59, 71, 80, 89, 100, 101, 246, 247, 248*, 249, 252*, 482 var. macrocalyx, 59, 80, 89, 247, 248*, 249, 250, 252*, 482 koehnei, 284 lanceolatus, 6, 11, 18*, 31, 35, 59, 76, 80, 89, 110, 111, 159, 188, 250, 251*, 252*, 253, 256, 326, 433, 482 LT f. foliosus, 315, 319 f. petiolatus, 323, 326 f. remotus, 250 var. alutaceus, 250, 315, 319 var. lanceolatus, 323, 326 var. scaber, 250 leucocalycinus, 6, 11, 12, 18*, 23, 41, 45, 52*, 53, 59, 71, 80, 89, 110, 124, 129, 256, 257, 258*, 259–261, 269*, 482 longipes, 7, 59, 71, 80, 89, 100, 260, 262*, 263, 269*, 270, 482 lythroides, 6, 423, 428 var. glabrescens, 423 var. villosissimus, 423 macrodon, 6, 14, 23, 59, 80, 89, 95, 162, 264, 265*, 266, 269*, 339, 366, 405, 450, 482 marginatus, 7, 11, 19*, 21, 23, 24, 25*, 37, 39, 52*, 53, 59, 71, 80, 89, 99, 263, 266, 267*, 268, 269*, 270, 340, 482 mattogrossensis, 59, 80, 89, 111, 270, 271*, 272, 281*, 482 micromerus, 59, 81, 89, 97, 98, 272, 273*, 274, 281*, 482

Index microphyllus, 6, 18*, 23, 26, 59, 81, 89, 98, 108, 133, 238, 274, 275*, 276–278, 281*, 434, 461, 482 f. capitata, 459 minasensis, 6, 23, 24, 25*, 27, 34*, 42, 59, 81, 89, 97. 278, 279, 280*, 281*, 282, 482 mononeuros, 4, 5, 59, 81, 89, 96, 282, 283*, 284, 292*, 339, 482 montanus, 232 myrsinites, 5, 16*, 23, 24, 25*, 26*, 59, 81, 89, 99, 102, 152, 284, 285*, 286, 287, 292*, 482 f. obtusifolius, 284, 287 LT var. cribellinus, 284, 287 NT nigricans, 6, 59, 81, 89, 104, 105, 287, 288*, 289, 292*, 482 LT nitidus, 6, 18*, 34*, 59, 81, 89, 100–102, 139, 289, 290, 291*, 292*, 293, 482 LT oblongus, 4, 6, 13*, 17*, 18*, 23, 24, 25*, 27, 37, 49, 55, 56, 59, 76, 81, 89, 91, 99, 101, 121, 216, 256, 293, 294*, 295, 301, 312*, 331, 482 LT var. angustifolius, 329 obtectus, 60, 81, 89, 108, 111, 301, 302*, 303, 312*, 482 orbicularis, 7, 11, 12, 19*, 21, 38*, 41–43, 46, 47, 49, 52, 53, 55, 56, 60, 64, 77, 81, 89, 103, 202, 303, 304, 305*, 308, 309, 312*, 474, 477, 478, 482 LT var. brachyander, 304, 308, 309 LT ovatus, 7, 30, 46*, 60, 71, 76, 81, 89, 100, 103, 309, 310*, 311, 312*, 315, 387, 470, 482 LT ovatus, 383 panniculatus, 6, 12, 19*, 21, 23, 25*, 27, 30, 33, 34*, 60, 71, 81, 89, 99, 100, 313, 314*, 315, 325*, 357, 482 LT paraisoensis, 7, 11, 13*, 18*, 19*, 23, 24, 34*, 49, 60, 64, 81, 89, 110, 112, 315, 316*, 317, 319, 325*, 380, 471*, 482 parvifolius, 6, 9, 18*, 23, 52*, 53, 60, 81, 89, 97, 109, 228, 319, 320*, 321–323, 325*, 450, 452, 482 pemphoides, 218, 220 petiolatus, 12, 60, 81, 89, 104, 139, 323, 324*, 325*, 326, 405, 470*, 482 plumbeus, 60, 81, 89, 106, 326, 327, 328*, 329, 336*, 482 praetermissus, 16*, 50, 60, 64, 81, 89, 95, 96, 101, 113, 329, 330*, 331, 336*, 470*, 482

515 prostratus, 60, 81, 89, 98, 331, 332*, 333, 334, 336*, 482 psammophilus, 5, 60, 81, 89, 96, 334, 335*, 336*, 337, 482 LT puberulus, 6, 12, 13*, 14, 15, 18*, 20, 23, 24, 26*, 37, 60, 82, 89, 94, 212, 337, 338*, 339, 348*, 365–367, 483 LT pulchellus, 6, 428, 434 LT punctatus, 5, 16*, 21, 34*, 39, 60, 82, 89, 99, 315, 340, 341, 342*, 343, 344, 348*, 483 var. dentatus, 60, 82, 89, 103, 341, 344, 345*, 348*, 483 LT pygmaeus, 28, 60, 82, 89, 111, 346, 347*, 348*, 349, 483 quintuplinervius, 6, 16*, 60, 71, 82, 89, 103, 349, 350*, 351, 362*, 483 ramosissimus, 6, 17*, 18, 30, 49, 50, 55, 56, 60, 64, 82, 89, 100, 103, 352, 353*, 354, 357, 362*, 375, 419, 470*, 483 var. decipiens, 352, 357 rectinervis, 60, 82, 89, 108, 358, 359*, 362*, 483 retroimbricatus, 7, 60, 82, 89, 104, 360, 361*, 362*, 363, 483 revolutifolius, 319 rosmarinifolius, 3, 6, 13*, 14, 18*, 55, 56, 60, 82, 89, 94, 191, 337, 339, 363, 364*, 366, 367, 373*, 390, 483 rotatus, 61, 82, 89, 111, 367, 368*, 369, 373*, 471*, 483 rotundifolius, 4, 7, 19*, 21, 23, 24, 25*, 34*, 61, 82, 89, 91, 99, 202, 369, 370*, 371, 373*, 483 rupestris, 137, 139 saxatilis, 61, 82, 90, 108, 193*, 372, 373*, 421, 483 scaber, 250, 256 sect. Palmatinerves, 4, 7, 20, 21, 66, 70, 91, 341 sect. Palmatipenninerves, 4, 7, 20, 21, 91 sect. Penninerves, 4, 6, 18, 20, 21, 66, 70, 91 sect. Subuninerves, 4, 5, 18, 20, 91, 340, 341 sessiliflorus, 7, 12, 13*, 34*, 55, 56, 61, 82, 90, 104, 107, 186, 216, 372, 374*, 375, 377, 386*, 483 LT sigillatus, 7, 18*, 19*, 61, 82, 90, 107, 111, 112, 194, 377, 378*, 379, 380, 386*, 483 smithii, 4, 6, 55, 56, 133, 232, 237

Index

516 Diplusodon (cont.) sordidus, 7, 11, 12, 19*, 46*, 52*, 53, 61, 63–65, 82, 90, 102, 113, 165, 380, 381*, 382, 383, 386*, 483 stellatus, 18*, 24, 25*, 26*, 61, 82, 90, 94, 110, 266, 387, 388*, 389, 390, 401*, 483 strigosus, 6, 11, 13*, 18*, 23, 61, 65, 76, 83, 90, 112, 390, 391*, 392, 394, 395, 401*, 471*, 483 LT subsericeus, 6, 61, 83, 90, 97, 237, 395, 396*, 397, 398, 401*, 483 LT tenuifolius, 61, 83, 90, 96, 152, 398, 399*, 400, 401*, 415, 416, 471*, 483 thymifolius, 6, 28, 61, 71, 76, 83, 90, 94, 191, 402, 403*, 405, 413*, 472*, 483 LT thysanosepalus, 4, 7, 61, 84, 90, 103, 165, 461*, 462*, 463, 483 trigintus, 61, 83, 90, 99, 242, 406, 407*, 413*, 483 ulei, 4, 5, 31, 35, 61, 64, 83, 90, 102, 129, 165, 168, 256, 408, 409*, 410, 413*, 483 uninervius, 5, 11, 15, 16*, 61, 83, 90, 95, 96, 156, 284, 339, 410, 411*, 412, 413*, 415, 483 LT urceolatus, 6, 256, 259–261 vacillans, 61, 83, 90, 96, 400, 402, 415, 416, 426*, 472*, 483 vaccinifolius, 143 velutinus, 6, 217, 220 LT venosus, 61, 83, 90, 100, 103, 416, 417*, 418, 419, 426*, 472*, 483 verruculosus, 42, 61, 83, 90, 95, 419, 420*, 421, 426*, 483 vidalii, 5, 61, 83, 90, 99, 100, 421, 422*, 423, 426*, 483 villosissimus, 4, 7, 19*, 28, 62, 83, 90, 91, 111, 272, 363, 423, 424*, 425, 426*, 427, 428, 483 LT villosus, 6, 13*, 18*, 23, 28, 49, 50, 55, 56, 62, 64, 83, 90, 98, 107, 428, 429*, 430, 433, 434, 439*, 472*, 483 vinaceus, 62, 83, 90, 98, 107, 434, 435, 472*, 483 virgatus, 3–5, 9, 11, 16*, 20, 23, 26, 28, 41, 47, 48*, 49–51, 52*, 53, 62, 64, 65, 75, 83, 90, 91, 101, 242, 256, 423, 425, 435–437, 438*, 439*, 447, 472*, 474, 483 LT var. occidentalis, 62, 83, 90, 437, 438*, 439*, 447, 448, 483

vittatus, 62, 83, 90, 95, 105, 448, 449*, 450, 454, 455, 457*, 483 vividus, 62, 83, 90, 108, 133, 322, 450, 451*, 452, 457*, 472*, 483 xerampelinus, 6, 261, 287 xerophilus, 62, 84, 90, 94, 98, 453*, 454, 455, 457*, 483 xiphodon, 62, 84, 90, 109, 133, 455, 456*, 457*, 458, 483 F Friedlandia, 3 albiflora, 437 amoena, 91, 437, 447 LT angustifolia, 228 buxifolia, 143, 148 epilobioides, 188 glaucescens, 200 helianthemifolia, 217 hexandra, 228 hirsuta, 232, 238 hirtella, 274, 278 LT leucantha, 437 lythroides, 423 myrsinites, 284 myrtifolia, 437 nitida, 289, 293 nummularifolia, 200 parvifolia, 319 rotundifolia, 369 serpyllifolia, 232 stachyoides, 423 thymifolia, 402 vaccinifolia, 143 verbenifolia, 233 villosula, 274 L Lythrum quintuplinervium, 349 N Nesaea speciosa, 383 P Pemphis hexandra, 228