Organization for Flora Neotropica Cyphomandra (Solanaceae) Author(s): Lynn Bohs Source: Flora Neotropica, Vol. 63, Cyphomandra (Solanaceae) (Jul. 5, 1994), pp. 1-175 Published by: New York Botanical Garden Press on behalf of Organization for Flora Neotropica Stable URL: http://www.jstor.org/stable/4393848 Accessed: 02-03-2016 18:53 UTC Your use of the JSTOR archive indicates your acceptance of the Terms & Conditions of Use, available at http://www.jstor.org/page/ info/about/policies/terms.jsp JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship. For more information about JSTOR, please contact [email protected]. New York Botanical Garden Press and Organization for Flora Neotropica are collaborating with JSTOR to digitize, preserve and extend access to Flora Neotropica. http://www.jstor.org This content downloaded from 169.237.45.23 on Wed, 02 Mar 2016 18:53:32 UTC All use subject to JSTOR Terms and Conditions FLORA NEOTROPICA MONOGRAPH 63 CYPHOMANDRA (SOLANACEAE) by Lynn Bohs - _ V . _? TROPIC OP CANCER FLORA NEOTROPICA TROPIC OF CAPRICORN Publishedfor Organization for Flora Neotropica by The New York Botanical Garden New York Issued 5 July 1994 This content downloaded from 169.237.45.23 on Wed, 02 Mar 2016 18:53:32 UTC All use subject to JSTOR Terms and Conditions NYBG Copyright ? 1994 The New York Botanical Garden Published by The New York Botanical Garden Bronx, New York 10458 Composition by Carol Johnson International Standard Serial Number 0071-5794 The paper used in this publication meets the minimum requirements of American National Standard for Information Sciences-Permanence of Paper for Publications and Documents in Libraries and Archives ANSI/NISO (Z39.48-1992. Printed in the United States on recycled paper Library of Congress Cataloging in Publication Data Flora neotropica. - Monograph no. 1 - New York: Published for Organization for Flora Neotropica by the New York Botanical Garden, 1968- v.: ill.; 26 cm. Irregular. Each issue has distinctive title. Separately cataloged and classified in LC before monograph no. 40. ISSN 0071-5794 = Flora neotropica. 1. Botany-Latin America-Classification-Collected works. 2. Botany- Tropics- Classification-Collected works. 3. Botany-Classification-Collected works. I. Organization for Flora Neotropica. II. New York Botanical Garden. QK205.F58 581.98'012-dcl9 85-647083 AACR 2 MARC-S Library of Congress [8508] ISBN 0-89327-385-6 This content downloaded from 169.237.45.23 on Wed, 02 Mar 2016 18:53:32 UTC All use subject to JSTOR Terms and Conditions CYPHOMANDRA (SOLANACEAE) LYNN BOHS TABLE OF CONTENTS Abstract ....................... ............... 2 R esum en . 3 Introduction ........... ................................... 3 Generic Delimitation and Relationships . ................................ 4 Taxonomic History .............. ..... .. .... 5 M orphology ... ... ............................................. 7 Habit, Architecture, and Stems .... .... ................... 7 Trichomes ...................................... ............. 9 L eaves . .. 10 Inflorescences . .... .. 11 Flow ers . .. .... .. .. 11 Fruits . ... 12 Seeds . .................................... ....... 13 Pollen .13 .. .. .. .. .. .... ... 13 Chromosomes .............................. ........ 21 Reproductive Biology .... ..... .. .. .................. .. .22 Species Groups and Character Evolution ........... ................ .. 25 Ecology .................... ..................... 29 Habitats ................... .. ........ .. ........... 29 Pollination . .... ..................... .. .............30 Dispersal . .............................. 32 Herbivory ......3.......2............ .......... 32 Fossil and Archaeological Record ..... ................................. 35 Distribution and Biogeography ....... .............. ................. 35 Uses ... .............................................. 41 Systematic Treatment ............................. ... ........ 42 Generic Description . ... ... .... ... ..... 42 Species Concept . .. .. ... .. .. .. 43 Typification . .. 43 Notes on the Systematic Treatment . ... .... ...... .. 43 Dichotomous Key to the Species of Cyphomandra ............. .. 44 Synoptic List of Cyphomandra Characters .. .. ...... ......... 46 Cyphomandra acuminata .. .... ....... ...................... 47 Cyphomandra benensis ...................................... .. 49 Cyphomandra betacea . 50 Cyphomandra cajanumensis ............ ................. 57 Cyphomandra calycina . .. .... ... ... .. .... .... 60 Cyphomandra corymbiflora . 63 subspecies corymbiflora . 66 subspecies mortoniana . .. 68 Cyphomandra diploconos ...................... .... ... 69 Cyphomandra divaricata ..................................... 74 Cyphomandra diversifolia ...... ... .. .. .................... 77 subspecies diversifolia . 79 subspecies chlorantha . 82 Cyphomandra dolichocarpa ............. ...... ..... ... 83 This content downloaded from 169.237.45.23 on Wed, 02 Mar 2016 18:53:32 UTC All use subject to JSTOR Terms and Conditions 2 Flora Neotropica Cyphomandra endopogon ......................................... 85 subspecies endopogon ......................................... 87 subspecies guianensis ......................................... 90 Cyphomandra foetida .............. ................... ...... 90 Cyphomandra fragilis ........... .. ................ ........ 93 Cyphomandra hartwegii ................................. 96 subspecies hartwegii .......................................... 99 subspecies ramosa ............................................ 104 Cyphomandra heterophylla ........... ......................... 105 Cyphomandra hypomalaca ..................................... 107 Cyphomandra obliqua ............... ................... ...... 108 Cyphomandra oblongifolia ........................................ 112 Cyphomandra ovum-fringillae ...................................... 114 Cyphomandra pendula .......................................... 116 Cyphomandra pilosa ........................................... 119 Cyphomandra pinetorum ......................................... 123 Cyphomandra premnifolia ........................................ 126 Cyphomandra rojasiana ...................................... .... 128 Cyphomandra sciadostylis ......................................... 130 Cyphomandra sibundoyensis ........................................ 133 Cyphomandra stellata ............. ....................... ..... 134 Cyphomandra sycocarpa ......... .............. ... ... ..138 Cyphomandra tegore ........................................... 140 Cyphomandra tenuisetosa ...................................... ... 142 Cyphomandra tobagensis ...................................... ... 145 Cyphomandra uniloba ............... ...................... .147 Insufficiently Known Taxa ........... .. ................... 150 Doubtful Names ............................... .. ....... 151 Excluded Taxa ..................................... 155 Acknowledgments ...................... ................. 158 Literature Cited . .. 159 Numerical List of Taxa . ... 163 List of Exsiccatae . 164 Index of Local Names . ................ .. 171 Index of Scientific Names ................. .................. ..... 172 ABSTRACT Bohs, Lynn (Department of Biology, University of Utah, Salt Lake City, UT 84112, U.S.A.) Cyphomandra (Solanaceae) Flora Neotropica 63: 1-176, 1994. The genus Cyphomandra (Solanaceae) includes about 35 species of shrubs and small trees native to the Neotropics. Cyphomandra ranges from Mexico to northern Argentina and east to southeastern Brazil. Most species are small trees that exploit light gaps in the primary forest. Several species are grown for their edible fruits and one, Cyphomandra betacea, the tree tomato or tamarillo, is cultivated worldwide in tropical and subtropical areas. This species is currently known only from cultivation. This monograph covers aspects of the taxonomic history, morphology, ecology, reproduc- tive biology, infrageneric relationships, biogeography, and uses of Cyphomandra. Thirty-two species are recognized in this treatment, and two insufficiently known taxa are discussed. Five informal species groups are described. All issues of nomenclature and typification pertaining to the genus are comprehensively examined. The genus is defined on the basis of the enlarged anther connective. In addition, the unusually large chromosomes of Cyphomandra are useful in generic delimitation. This content downloaded from 169.237.45.23 on Wed, 02 Mar 2016 18:53:32 UTC All use subject to JSTOR Terms and Conditions Introduction 3 Unusual ecological aspects of Cyphomandra include its pollination syndrome and herbi- vore relationships. Male euglossine bees may be important pollinators of Cyphomandra flowers, and the primary attractants may not be pollen, but odor substances secreted by the anther connectives. Specialized herbivores known for Cyphomandra are larvae of neotropical butterflies of the nymphalid subfamily Ithomiinae. Further examination of these plant/insect relationships may elucidate the functional significance of the anther connective in pollination and provide insight into the comparative chemistry of Cyphomandra alkaloids. Key words: Cyphomandra, Solanum, Solanaceae, tree tomato, tamarillo, anther connective, euglossine bees, Ithomiinae. RESUMEN El genero Cyphomandra