Solanum Section Geminata (Solanaceae) (Flora Neotropica Monograph No. 84)

Organization for Flora Neotropica Solanum Section Geminata (Solanaceae) Author(s): Sandra Knapp Source: Flora Neotropic...

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Organization for Flora Neotropica

Solanum Section Geminata (Solanaceae) Author(s): Sandra Knapp Source: Flora Neotropica, Vol. 84, Solanum Section Geminata (Solanaceae) (Jul. 30, 2002), pp. 1404 Published by: New York Botanical Garden Press on behalf of Organization for Flora Neotropica Stable URL: http://www.jstor.org/stable/4393908 Accessed: 21/10/2008 12:36 Your use of the JSTOR archive indicates your acceptance of JSTOR's Terms and Conditions of Use, available at http://www.jstor.org/page/info/about/policies/terms.jsp. JSTOR's Terms and Conditions of Use provides, in part, that unless you have obtained prior permission, you may not download an entire issue of a journal or multiple copies of articles, and you may use content in the JSTOR archive only for your personal, non-commercial use. Please contact the publisher regarding any further use of this work. Publisher contact information may be obtained at http://www.jstor.org/action/showPublisher?publisherCode=nybg. Each copy of any part of a JSTOR transmission must contain the same copyright notice that appears on the screen or printed page of such transmission. JSTOR is a not-for-profit organization founded in 1995 to build trusted digital archives for scholarship. We work with the scholarly community to preserve their work and the materials they rely upon, and to build a common research platform that promotes the discovery and use of these resources. For more information about JSTOR, please contact [email protected].

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FLORA NEOTROPICA MONOGRAPH 84

SOLANUMSECTION GEMINATA(SOLANACEAE) SANDRA KNAPP

FLOR........... NEOTROPICA> 'toic

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N AP/C/WON

Publishedfor Organization for Flora Neotropica by The New York Botanical Garden Bronx, New York

Issued 30 July 2002

? 2002 by The New YorkBotanical Garden All rights reserved. Published by The New YorkBotanical Garden Bronx, NY 10458

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Library of Congress Cataloging-in-Publication Data Flora neotropica.- Monographno. 1 - New York:Published for Organizationfor FloraNeotropica by The New York Botanical Garden, 1968v.: ill.; 26 cm.

Irregular. Eachissue has distinctivetitle. Separatelycataloguedand classified in LC before monographno. 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. Organizationfor FloraNeotropica. II. New YorkBotanical Garden. QK205.F58 Libraryof Congress ISBN 0-89327-441-0

581.98'012-dcl9

85-647083

[8508]

02 03 04 05 06 07 08 09 10/98765432

1

SOLANUMSECTION GEMINATA (SOLANACEAE) SANDRA KNAPP

CONTENTS 2 Abstract/Resumen ........................................................................................................ Introduction....................................................................................................................... 3 4 History of Classification of Solanum................................................ Nomenclaturaland TaxonomicHistory of Section Geminata ........................................... 5 Morphology................................................. 8 Habit .............................................................................................................................. 8 Stems .................................................................................................. 8 . Leaves .......................................................................................................................... 10 Inflorescences.............................................................................................................. 10 Flowers ........................................................................................................................ 11 Fruits ............................................................................................................................ 12 Seeds ............................................................................................................................ 15 Trichomes .................................................................................................................... 15 Chromosomes .............................................................................................................. 22 Ecology andNaturalHistory ............................................................................................ 22 Habitatsand Distribution......................................................................................... 22 Pollination ..................................................................................................................... 25 FruitDispersal .............................................................................................................. 28 Sex Expression............................................................................................................. 29 fHerbivores.................................................................................................................... 31 Species Groupsand Species Concepts............................................................................. 32 Species Groups ............................................................................................................ 32 Species Concepts..........................................................................................................32 Taxonomic Treatment ......................................................... 38 Artificial Key to Woody, Similar Groupsof Non-Spiny Solanums ............................ 39 Artificial Key to the Species Groups............................................................................ 40 Synoptic Key to the Species ......................................................................................... 42 l. Solainumoblongifoliumspecies group ......................................................................43 II. Solanumlpseudocapsicum species group ................................................................54 1II.Solanumn ntidumspecies group...............................................................................69 IV.Solanum leucocarpon species group .................................................................... 136 V. Solanum nutans species group...............................................................152 VI. Solanum amblophyllumspecies group ................................................................ 165 VII. Solanumdeflexiflorumspecies group ................................................................. 179 VIII. Solanum arborelum species group .................................................................... 194 IX. Solanumunifoliatumspecies group ..................................................................... 223 X . Solanumrob stifrons species group ............................................. ........................ 235 XI. Solanumnarcoticosmnum species group .............................................................. 247 XII. Solanum nigricans species group....................................................................... 254 X III. Solanum arenarium species group ................................................................... 273 XIV. Solanumsessile species group........................................................................... 282 XV. Solaniumconfine species group ........................................................................... 311

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FLORANEOTROPICA XV I. Solanum dolosum species group....................................................................... 350 IncertaeSedis ........................................................................................................... . . . 356 New Species Added in Proof ..................................................................................... 361 DoubtfulNames ............................................................................................................. 363 Excluded Species ............................................................................................................ 364 Acknowledgments .......................................................................................................... 368 Literature Cited ...............................................................................................................36 9 Numerical List of Taxa ................................................................................................... 375 Exsiccatae ....................................................................................................................... 377 Index of Scientific Names .............................................................................................. 399 Appendices .....................................................................................................................403

ABSTRACT Knapp, Sandra (Departmentof Botany,The NaturalHistoryMuseum,CromwellRoad, London SW7 5BD, United Kingdom).Solanumsection Geminatas.l. [sections Geminata, Pseudocapsicum, Holophylla pro parte, and Indubitaria pro parte] (Solanaceae). Flora Neotropica Monograph84: i-iv + 1-404. 2002. -Solanum section Geminatas.l. (Solanaceae) includes about 126 species of shrubsand small trees native to the Neotropics. Membersof the sectionrangefromnorthernMexico andthe WestIndiesthroughoutSouthAmerica to UruguayandnorthernArgentina.A single species is nativeto the Paleotropics,fromChina to tropicalAustralia,and is also unusualin the group in being polyploid. Most species are shrubsor small treelets found in light gaps or along streamsin primaryforest understory.A few species arecommonin second growthforestsin manyforesttypes. The sectionas treated here containsspecies with both simple andbranchedtrichomes.It is clearfromthis studythat the trichomecharacterstraditionallyused to differentiatesectionsofSolanumaretoo variable to be of use in sect. Geminata. Some species traditionallytreated as members of sects. Pseudocapsicum,Indubitaria,andHolophyllaare includedin the monograph,as those sections as traditionallyconstitutedareartificialgroupingsof apparentlyunrelatedtaxa. The monographtreatstaxonomic history,morphology,ecology, and naturalhistory of Solanumsect. Geminatas.l. One hundredtwenty-six species arerecognizedin this treatment and fourinsufficientlyknowntaxa arediscussedin contextwith theirputativerelatives.The section has been dividedinto 16 species groupsto facilitateidentificationandfuturein-depth phylogenetic study. Species groups are defined using largely charactersof: 1) sympodial structure,2) leaf shape,3) trichomemorphology,4) inflorescencemorphology,5) pedicel scar type, 6) fruitingpedicel morphology,and 7) seed morphology.A fundamentaldifferencein seed morphologyappearsto separatethe section into two broadgroups:those species groups with flattenedreniform seeds typical of those found elsewhere in the family, and species groups with unusual ovoid reniformseeds found only in sect. Geminata.All typification andnomenclaturalissues pertainingto the section andits componentspecies arecomprehensively examined. Section Geminatas.l. may not be monophyletic,but knowledge aboutrelated groups is scanty and a definitivejudgment cannotbe made at present. Extensiveecologicaldataareincludedon pollinationandherbivory.PollinationinSolanunm sect. Geminatais similarto thatobservedthroughoutthe genus. Femalebees of severalfamilies vibratethe poricidalanthersandextractpollen. Specializedherbivoresof membersof the section arebeetles of the family Chrysomelidaeandnymphalidbutterfliesof the subfamily Ithomiinae.Larvaeof relativelyphylogenetically"advanced"butterfliesfeed on species in the section, but specific host/herbivoretrackingis not apparent.Futurecladistic studies of both hosts and herbivoresmay reveal patternsin these relationships.

RESUMEN Knapp, Sandra (Departmentof Botany,The NaturalHistoryMuseum,CromwellRoad, London SW7 5BD, United Kingdom).Solanumsection Geminatas.l. [sections Geminata, Pseudocapsicum, Holophyllaproparte,andIndubitaria proparte](Solanaceae).FloraNeotropica

INTRODUCTION

3

Monograph84: i-iv + 1-404. 2002. -Solanum section Geminatas.l. (Solanaceae)incluye alrededorde 126 especies de arbustosy pequefiosarbolesnativos del Neotr6pico.Miembros de la secci6n se distribuyendesde Mexico y las Antillas, a trav6sde Sudam6rica,hastaUruguay y el nortede Argentina.Una sola especie es nativadel Paleotr6pico,desde Chinahasta Australia,siendoinusualen el grupopor serademaspoliploide.La mayoriade las especies son arbolespequefiosque crecen en claros o a lo largo de arroyos,en el sotosbosquede bosques primarios.Pocasespeciesson comunesen bosquessecundariosde diversostipos.La secci6n,en la formaaquitratada,contieneespeciescon ambostiposde tricomas,simplesy ramificados.De esteestudioresultaclaroquelos caracteres deltricoma,tradicionalmente utilizadosparadiferenciar secciones de Solanum,son demasiadovariablesparaser empleadosen la secci6n Geminata. tratadascomo miembrosde las seccionesPseudocapsicum, Algunasespecies,tradicionalmente Indubitariay Holophylla, son incluidas en la monografia,ya que estas secciones estaban tradicionalmenteconstituidasen agrupacionesartificialesde taxasin relaci6naparentente. Lamonografiatratala historiataxon6mica,morfologia,ecologiae historianaturaldel genero Solanumsecci6n Geminatas.l. Cientoveintiseis especies son reconocidasen el tratamientoy cuatrotaxones, insuficientementeconocidos, son discutidos en contexto con sus parientes putativos.Lasecci6nha sidodivididaen 16gruposde especies,amodo de facilitarsuidentificaci6n y un futuroestudio filogen6tico profundo.Los gruposde especies son definidos utilizando fundamentalmente caracteresde: 1) estructurasimpodial,2) formade la hoja,3) morfologiade los tricomas,4) morfologiade la inflorescencia,5) tipo de cicatrizdel pedicelo, 6) morfologia del pedicelofrutal,y 7) morfologiade la semilla.Unadiferenciafundamentalen la formade las semillas parecesepararla secci6n en dos grandesgrupos:un grupode especies con semillas reniformesy aplanadas, en la familia;y otrogrupode especiesconsemillas tipicamenteencontradas ovoide-reniformes,inusuales,encontradass6lo en la secci6n Geminata.Son tratadosin extenso todaslas cuestionesnomenclaturalesy de tipificaci6nrelacionadascon la secci6n y sus especies componentes.La secci6nGeminatapodriano sermonofiletica,peroel conocimiento sobregruposafines es todaviaescaso y por el momentono puede haberunjuicio definitivo. Se incluyenextensosdatosecol6gicos sobrepolinizaciony herbivorismo.Lapolinizaci6n en Solanumsecci6n Geminataes similar a lo observado en el genero. Abejas hembrasde variasfamiliashacenvibrarlasanterasporicidas,extrayendoasiel polen.Herbivorosespecializados de miembros de la secci6n son escarabajospertenecientes a la familia Chrysomelidaey mariposas de la subfamilia Ithomiinae. Larvas de mariposas, relativamente avanzadas se alimentande especiesde la secci6n,peroaparentemente no hayunarelaci6n filogeneticamente, especifica herbivoro/huesped.Futurosestudios cladisticos de ambos, herbivoroy hu6sped, podrianrevelarrelaciones de este tipo.

INTRODUCTION SolanumsectionGeminata(G. Don) Walp.,with 126 species as treatedhere, is one of the largestsections of the extremelylargeplantgenusSolanum.Solanumhas not been comprehensivelymonographedsince the De CandolleProdromus(Dunal, 1852). In his treatmentof the group,Dunal(1852) included37 species. Latercontributionsby Van Heurck& Miiller-Argovius(1870), Morton(1944), and especially by the prolific German botanistGeorgBitter(1913, 1916, 1919a, 1920a, 1920b, 1922) greatlyincreasedthis number.Manynew species now placed in sect. Geminatas.l. were described,but no attemptwas madeto determinerelationshipsor synonymies in the section.The sectionhas been considered difficult,andspecies arevery similarmorphologically; uniformon the theyhavebeendescribedas "remarkably herbariumsheet" (D'Arcy, 1973). The authorsof two CentralAmericanfloristic treatments(D'Arcy, 1973; Gentry& Standley,1974)triedto sortoutnomenclature

and synonymyin the section, but since the greatdiversity in sect. Geminataoccurs on the easternslopes of the Andes and in the Amazon basin, their attempts, thoughlaudable,treatonly a small fractionof the species in the group. The goals of this study were twofold: first to provide a monographof the group, solving nomenclaturaldifficulties and clarifying species identities,and second to breakthis large andunwieldy group into smaller, monophyletic subgroupssuitable forfuturein-depthstudyandphylogeneticanalysis.The techniquesI have used have been for the most partthe traditionalones of systematicbotanists,examinationand analysis of the comparativemorphologyof herbarium specimens, but I have placed much emphasis on the observationof the species in theirnative habitats.This has been very importantin my eventualunderstanding of relationshipsin the group,and is crucialto the solving of problems in this complex and extremely large section of Solanum.

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HISTORY OF CLASSIFICATION SOLANUM

OF

Solanumwas one of Linnaeus' largergenera,with 23 species described in the first edition of Species Plantarum(1753). The species describedin thatwork arelargelyEuropeanorAfrican;manyof theexotic ones were known only fromcultivation.The late eighteenth andearlynineteenthcenturiesweretimesof exploration, in theAmericas(Steam,1968;Steele, 1964), particularly andplants collected on expeditions flooded European herbaria.In 1813,Michel-FelixDunal'sthesis,a worldwide monographof thegenus,included235 species.The synopsis (Dunal, 1816) of his never completed second edition included 321 species, of which many of the additionalspecies were collected by Alexandervon Humboldtand Aimee Bonpland in tropicalAmerica. GeorgeDon (1837) enumeratedanddescribed406 species of Solainum.G. G. Walpers(1844) followed Don's general system and listed 435 species and an additional 72 for which he had incomplete information. The last worldwide treatmentof Solanumwas that of Dunal(1852) forAlphonsede Candolle'sProdromus andby thattime the numberof species hadrisento 900. This increase largely reflected the enormousincrease in collections fromtropicalAmerica. Since 1852,manynew specieshavebeendiscovered anddescribed,bringingthe numberof namesin the genus to between 3000 and 5000. The numberof"good" species this representsis difficultto assess, as the genus has grown to proportionsunmanageablefor a single Withprobably1500to 2000 validspecies, monographer. Solanumis one of the largestgeneraof floweringplants. Since there exist so many species of Solanum, comprehensivemonographictreatmentof the entiregenus is problematicif not impossible, andtaxonomistshave concentratedon smaller groupswithin the genus. Linneaus(1753) dividedthe genus into two groups, the Iliermia, or unarmedspecies, and the Spinosa, or armedspecies. Dunal (1813, 1816) divided the genus similarly. His categories were Inermia, or unarmed solanums, and Aculeata, or armed solanums. In the Prodromus(1852) he divided Solanum into two sections ("sectio") which were basically the same as his earlier divisions. He named these Pachystemonum, species with stout, cylindricalanthersandno prickles, and Leptostemonum,species with attenuateanthers, trichomes,andoftenwithprickles(see TableI forDunal's characterizationof the groups). Generaltreatmentsof the entire genus since Dunal's 1852 work have been relativelyfew andlimitedto arrangements ofinfrageneric taxa. Subsequentworkershave divided the genus into subgenera and called the groupings at the next rank

below sections.Fora review of sectionalnomenclature in Solanum, see D'Arcy (1972) and Knapp (1983). Seithe (1962), building upon the voluminous work of GeorgBitter(see bibliographyin Weber,1928) andher own studiesof hairmorphologyin the genus, proposed a classificationfor Solanumbased largelyon trichome types.Thetwo majordivisionsin herclassificationwere the stellate-haired solanums (chorus subgenerum inpartto Dunal'sAculeata), Stellatipilum,corresponding and the simple- and branched-hairedsolanums (chorus subgenerumSolanum).Danert(1967, 1970), using his own workon the developmentof branchingpatterns in the genus and family,proposeda subgenericclassificationtakingintoaccountthesecharacters.Gilli (1970) provided a key to the groupingsof Seithe (1962), but did not propose any new re-arrangementsof taxa. D'Arcy (1972) lectotypifiedall of the existing subdivisions of Solanum,andproposeda provisionalconspectus of the genus. This treatmentis now thatmost commonly used by presentSolanumworkers.He divided the genus into seven subgenera(see Table II) and 52 sections, based on charactershe felt were more realistic than those used in the past. Of these systems of classification, only Seithe's (1962) specified the species to be includedin eachof the subgroupings.D'Arcy (1972) used and cited only the type or lectotype species of each section, andplacing a non-type species in his systemis difficult,if not impossible.Whalen(1984) provideda conspectus for the spiny solanums(subgenus Leptostemonum),in which he divided them into speciesgroupsandlistedthe componentspecies in each group. D'Arcy (1991) assessed revisionary work in Solainumandpresentedan identicalsubgenericscheme, the only differencebeing the replacementof the name subgenus Brevantherumwith subgenus Minon. Nee (1999) has divided the genus (treatingonly the New Worldtaxa) into three subgenerausing charactersof andinflorescencemorphology pubescence,architecture, (see TableII). Componentneotropicalspecies in each section or subdivisionare listed, makingNee's (1999) treatmentan extremelyuseful startingpoint for future work in Solanum. A comparisonof D'Arcy's (1972) and Nee's (1999) systems is presentedin Table II. Relationshipsof subgenericgroupswithinSolanum are currentlybeing investigated using the techniques of molecular systematics (Olmstead& Palmer, 1991, 1997; Spooneret al., 1993; Bruneauet al., 1995;Bohs & Olmstead, 1997, 1999). The use of these tools to identify monophyletic groups within Solanumnwill greatly facilitate futuremonographicwork. Although very few (currentlyonly about30 of some 1000-2000 species) have been analyzed,interestingpatternsof relationship are beginning to emerge. Clearly mono-

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HISTORY OF CLASSIFICATION OF SOLANUM

Table I Classification of Solanum abbreviated from Dunal (1852). Only the two major sectional groupings and the portion of the classification pertaining to section Geminata are included. Symbols are those of Dunal (1852) and are indicative of rank within his system Sectio I. Pachystemonum "Antheraebreviores, crassiores, cylindrico-vel ovato-ellipticae, nec elongatae, apice attenuatae;poris terminalibus, plerumque anticis, loculos diametro aequantibus, saepe primum poris terminalibus, dein rimis lateralibus, rariusporis terminalibus minutis sursum spectantibus, dehiscentes. -Sp. omnes inermes." Subsectio IV. Micranthes 1?. Anthoresis 2?. Anthopleuris ? 1. Oppositifolia * Indubitaria ** Lepidota *** Leiodendra ? 2. Pseudocapsicum ? 3. Bassovioides Sectio II. Leptostemonum "Antherae elongatae, apice attenuatae, poris terminalibus minutis retrorsisposticis vel sursum spectanibus. -Species nonnunquam inermes, plerumque aculatae."

phyletic groupsare identifiable,in partcorresponding to the traditionaldivisions in the genus, but some of the relationshipsrevealedby these techniquesare surprising. Based on analysesof moleculardatasets, Spooner et al. (1993) have shown that the tomatoes (formerly segregated as the genus Lycopersicon) are the sister group of the potatoes and should be considered as membersof Solanum subgenusPotatoe, Bohs (1995) has transferredall the species formerly segregatedas the genus Cyphomandra(Bohs, 1994) into Solanum, and it is clear that Seithe's (1962) chorus subgenerum Stellatipilumis polyphyletic. Since Dunal's monograph(1852), taxonomicwork in Solanumhas proceededat the sectional level and in regional floristic treatments.Recent sectional monographs in Solanum include the following: sect. Tuberarium(Dunal) Bitter(sects. Petota Dumort.and Basarthrum Bitter, pro parte, Correll, 1962); sect. Brevantherum Seithe (Roe, 1967b, 1972); sect. Acanthophora Dunal (Nee, 1979); sect. Androceras (Nutt.)Marzell(Whalen,1978, 1979);sect.Lasiocarpa D'Arcy (Whalen et al., 1981); the S. nitidumspecies group(sect.Holophyllaproparte)(Knapp,1989);sects. Lepidotum(Dunal)SeitheandCernuumCarv.(Carvalho & Shepherd, 1991; Carvalho, 1996); sect. Allophylla (Child) Bohs (Bohs, 1990); sect. Pachyphylla Dunal (as Cyphomandra, Bohs, 1994; see above); sect. Pteroidea Dunal (Knapp& Helgason, 1997); andcontinuingwork on various subgroupsof the complicated sect. Solanum L. (Maurella Dunal: Edmonds, 1972,

1977, 1978; Heiser, 1955, 1963; Heiser et al.. 1979; Henderson,1974; Schilling, 1981; Schilling & Heiser, 1976, 1979; Edmonds& Chweya, 1997). A list of regional floras treatingSolanim in some detail is provided in TableIII.

NOMENCLATURAL AND TAXONOMIC HISTORY OF SECTION GEMINATA Solanum section Geminatahad long been known as sect. Leiodendra Dunal, but the former name has priority under the rules of the InternationalCode of Botanical Nomenclature (Greuter et al., 1994; see Knapp,1983, 1984). The variousnamesthathave been appliedto the groupsof species includingthe lectotype (D'Arcy, 1972: S. nudum Dunal) of sect. Geminata are listed in the TaxonomicTreatment. Two species of sect. Geminataas treatedherewere known to Linnaeus(1753): SolanumdiphyllutmL., an ornamentalshrubprobablyfirstbroughtto Europefrom Mexico in the early seventeenth century (see discussion underS. diphyllum),and S. pseudocapsicum L., whichLinnaeusknewfromplantscollectedon Madeira, but almost certainlybroughtthereby early Portuguese traders(see Press & Short, 1994). In his 1813 thesis on the naturalhistory,taxonomy,and medical uses of Solanum,DunalincludedS. diphyllumandeleven other species in his group "c. Foliis ovatis, oblongis aut lanceolatis,glabrisautpubescentis."In a footnoteto the

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Table II Subgenericclassification of Solanum (according to D'Arcy 1972, 1991, and Nee, 1999). Characters defining the subgenera Inflorescence intra-axillary (leaf-opposed or terminal or various); anthers oblong; plants never armed; hairs simple, dendritic, or stellate Inflorescence axillary or intraaxillary in bi- or tri-furcation of stem; plants never armed; anthers oblong or tapered; hairs simple or dendritic

Characters defining the subgenera Stout anthers; simple hairs; no spines

Nee, 1999

Subgenus Bassovia

Stout anthers; simple hairs; pinnate leaves; axillary inflorescences; pointed fruits

Subgenus Bassovia

Subgenus Brevantherum (as subgenus Minon in D'Arcy, 1991) SubgenusPotatoe

Stout anthers;entire leaves; dendriticor stellate hairs

Included in subg. Solanum

Scandentspecies;pinnate leaves with interstitialleaflets; lateralpendulousinflorescences;articulatedpedicels Stout anthers; aneuploid chromosome numbers; Australia Stout anthers; South Africa

Included in subg. Solanum

D'Arcy 1972, 1991 Subgenus Solanum

Subgenus Archaeosolanum Subgenus Lyciosolanum Subgenus Leptostemonum

Tapering anthers; stellate hairs, often with spines

Subgenus Solanum

Included in subg. Solanum

Not treated Subgenus Leptostemonum

Inflorescence intra-axillary: anthers tapering; plants armed; hairs stellate, never dendritic

descriptive section of the thesis he commented, "S. raceme insteadof a cyme, see Morphology).Solanum acuminatum,gnaphalioides, fetidum diphyllum, ob- pseudocapsicum,by virtueof its possessionofangulateThismar- repandleaves, was placed by Dunal in a groupnamed longum;an sectionemnaturalemconstituunt?" ginal commentis the firstindicationof the naturalunity "Pseudo-Capsica" which also had as members S. of the group. Also included with S. diphyllumin this scabrum and S. aethiopicum, both spiny solanums. group were the following species which are still con- Otherspecies I have included in sect. Geminatawere sideredpartof sect. Geminatas.l: S. triste,S. oppositi- placed by Dunal(1816) in a groupcharacterizedby its folium(as S. urceolatum),S. acuminatum,S. oblongum, terminalinflorescences(S. sessile, S. oblongifolium,S. was assigned anonaefolium(= S. nudum))which was composed of andS.foetidum.Solanumpseudocapsicum to a hodgepodge of a grouppossessing sinuate leaves species froma numberof currentlyrecognizedsections. G. Don (1837) andWalpers(1844), followingDon's which also includedS. dulcamaraand several species of spiny solanums (Dunal, 1813). By 1816 Dunal had system almost exactly, named the group containing assigned these species and several others to a group SolanumdiphyllumandS. nudumas Geminata(Don at (butgrad.ambig.,see Knapp,1983)named"Leiodendra," the subsectionalrankandWalpersat the sectionalrank) it fromtherestofSolanumby,"Leaves possessingthe followingcharacters:"Arboress. frutices anddifferentiated and sometimes solitaryon the same raro in axillis foliis twin, entire, quite glabri, saepe geminis glabris, nervorumsubtuspilosis; racemissimplicibussymosis branch,rarelyaggregate.Calyx5-parted.Stamensequal." aut cymoso-umbellatis oppositifoliis; corollis sub-5- (G. Don, 1837:418). The speciesincludedin Don's sect. partitis."The charactersthathe consideredimportant Geminataare quite diverse;many are spiny solanums in the delimitationof the groupareclearlythe glabrous and the group as a whole is not as naturalas Dunal's natureof the plants and the leaf-opposed, simple in- muchsmallergroup"Leiodendra"(1816). NeitherDon florescence (which was misinterpretedby Dunal as a norWalperswas anexpertin Solanumorthe Solanaceae,

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HISTORYOF SECTIONGEMINATA Table III Regional floras with complete treatmentsof Solanum. COUNTRYOR REGION

AUTHOR

Brazil Costa Rica Venezuela Peru Panama Guatemala Veracruz, Mexico

Sendtner, 1846 Morton & Standley, 1938 Pittier et al., 1947 (list) Macbride, 1964 D'Arcy, 1974 [1973] Gentry & Standley, 1974 Nee, 1993

as was Dunal, so this is perhapsnot surprising. Sendtner,in his treatmentof Solanumfor Martius' Flora Brasiliensis (1846), for the most partfollowed Dunal's(1816) groupings,butconfinedhisLeiodendra Dunal to glabrous shrubs with geminate leaves and "pseudo-lateral"inflorescences.His treatmentof these species is excellent. He not only describesin detailthe Brazilianspecies, but many species fromPeruandthe Caribbeanas well. All of the species that Sendtnerincluded in his group Leiodendra are here retained as members of sect. Geminata. In his lastandmost extensivetreatmentofSolanum, Dunal(1852) definedhis groupLeiodendravery much as had Sendtner(1846). He includedLeiodendraas a subgroup of his larger group Oppositifolia (grad. ambig.),andcharacterizedit in the sameway as in 1816, "Arbores,fruticessuffruticesque;foliis saepe geminis glabris,raroin axillis nervorumsubtuspilosis; racemis simplicibus,cymosis autcymoso-umbellatis,oppositiHabitrange foliis; corollis albis subquinquepartitus." was extendedto subshrubs,andflowerswere described as white; otherwisethis descriptionis exactly the same as the one in theSolanorumSynopsis(1816), although manymore species were included(12 in 1816 vs. 37 in 1852). Dunal (1852) clearly distinguishedthose taxa with tomentoseleaf undersides(Indubitaria)andthose species with lepidote scales ("leproso-squamulosa," Lepidota) from his groupLeiodendra(see Table I for detailsof Dunal's 1852 classification).Delimitationof the groupnow known as sect. Geminatahas remained very similarto thatproposedby Dunal,but opinionsas to the relationshipsof the groupwith othersections of Solanumhave differedgreatly.Taxaincludedin Dunal's Oppositifolia but excluded from sect. Geminata as treatedhere are detailedin Excluded Species. Dunal (1852) considered those taxa with leaf-opposed inflorescences to be most closely related. He groupedLeiodendra,Indubitaria,andLepidotatogether in the largergroup Oppositifolia (see Table I). Seithe (1962, 1979), workingwith trichomemorphology,con-

cluded that the fundamentaldivision in Solanumwas taxaandthenon-stellatethatbetweenthe stellate-haired hairedtaxa.In herclassificationthe species inLepidota, possessing lepidotescales andstellatetrichomes,were placedwith otherstellate-hairedgroups.The othertwo groups from Dunal's Oppositifoliawere still consideredto be closelyrelated.SectionOppositifolium (Dlual) Seitheconsistedof threesubsections:1) Oppositifolium (Dunal)Seithe (largelyDunal'sLeiodendra),2) Silicosolanum(consistingof a singlespecies,S. trachytrichium Bitter,with unusualtrichomemorphology),and3)Indubitaria(those species with branchedtrichomes).She specified at least some of the species to be includedin each subsection,but in many cases the characterof trichomebranchingdoes notworkwhenmorethana single specimen per species is examined (see Morphology). Danert(1970, 1975),thoughconcentratingon inflorescence and sympodial structure,basically followed Seithe's (1962) system, with minor changes, none of which affect the placementof sect. Geminata.D'Arcy (1972) divided Solanum into seven subgenera (see TableII),clearlydifferentiating the simpleandbranched hairedgroupsas belongingto distinctphylogeneticlineages. His reasonsfor this arenot explicit, but fromhis conspectus,the intentis clear.In D'Arcy's (1972) system, sect. Geminata(as LeiodendraDunal)is placedin subgenusSolanum,while Indubitariais a subsection of sect.Holophylla(G. Don) Walp.in subgenusBrevantherum.Section PseudocapsicumBitteris includedas a subgroupof Indubitariain both D'Arcy's (1972) and Nee's (1999) systems. Hypothesesaboutrelationships between sections arenot presentedin D'Arcy's (1972) treatment,but from his placement of sect. Geminata, its relationshipsappearto be with sect. SolanumL. To date only three species of sect. Geminata as treatedhere have been used in molecular systematic analyses: Solanum pseudocapsicum, S. arboreum (Bohs & Olmstead, 1997; S. pseudocapsicum only in Bohs & Olmstead, 1999), and S. aphyodendron (Olmstead & Palmer, 1997). In analyses using ndhF

8

sequences (Bohs & Olmstead, 1997) and cpDNA restriction site variation (Olmstead & Palmer, 1997), a sister group relationship for S. pseudocapsicum and either S. arboreumor S. aphyodendronwas strongly supported. It is clear that past segregation of the branched-trichome taxa (cf. Dunal, 1852; D'Arcy, 1972) is not justified. All three of these taxa of sect. Geminatahave not been used in a single analysis, so molecularrelationshipswithin the groupremainto be resolved.It is also clearthatmorerobustsamplingwill greatlyimprovephylogeneticinference(see Perssonet al., 1994; Knapp et al., 1997; Chase et al., in press). However, on-going studies (Bohs, pers. comm.) indicate that members of sect. Holophylla s.str. (i.e., S. Bitter& Lillo, see Excluded Species for a argentinurm complete list) may also be partof the clade containing membersof sect. Geminata. My morphologicalworkwith sect.Geminataclearly showed that trichome morphology and inflorescence structurewere both extremely variableattributes(see discussions of these charactersin Morphology).Variation occurswithin species, populations- andeven individuals- in the degreeof trichomebranching,which traditionallyhas been one of the most importantcharacters used in Solanumtaxonomy. Section Geminata as delimitedhereincludesspecieswithsimpletrichomes and species with branched trichomes, a conclusion strongly supportedby recent molecular analyses (see above).Insteadof delimitingformalsubsectionalorother groupings at this point, I have brokensect. Geminata s.i., which is largeandunwieldy,into a set ofputatively monophyleticspecies groups.The monophylyof these smaller groups can be tested (see Knapp, 1991b), and fromthis a stableandtestableclassificationconstructed. In this monographI have treatedthe section in its broadestsense and have assembled groups of species I hypothesize to be monophyletic.This has meantthat I have includedtaxapreviouslyconsideredto be members of various other sections of non-spiny solanums: most notably Pseudocapsicum, Holophylla, and Indubitaria. This to a certain extent means that sect. Geminiatas.l. is probably not a monophyletic group, while its component species groups almost certainly are. The section as treated here is most probably a paraphyleticgradewhose relationshipsto otherclearly monophyletic groups of solanums are not at all resolved. Species groups that I think will be partof the wider set of monophyletic groups related to the taxa treated in this monographare sect. Holophylla s.str. (species list in Excluded Taxa), sect. Pachyphylla (Bohs, 1994), sect. Cyphomandropsis(Bohs, 1994), sect.Allophylla(Bohs, 1990),theSolanumthelopodium species group (Knapp, 2000), and possibly the S.

FLORA NEOTROPICA

nitidumspecies group(Knapp,1989). SectionsLepidotum,Cernuum,andExtensumarealso possibly related to these groups,althoughthis has not been traditionally considered due to their possession of lepidote scales, chaffy hairs derived from lepidote scales, and stellate hairsrespectively.Generalarchitecture andinflorescence morphology is very similar however in all of these groups. A brief key distinguishing these groups and differentiatingthem fromsect. Geminatas.l. as treated here is providedin the TaxonomicTreatment,andspecies lists areprovidedeitherin the referencescited, in the Appendices, or in Excluded Species.

MORPHOLOGY HABIT Speciesof sectionGeminataareunusualin Solanum in being plants of primaryforest understory,whereas the genus as a whole is mainly one of open, disturbed areasandlightgaps. Speciesof the sectionrangein habit fromsmallshrubsto trees 10-15 m talland20 cm diameter,butarealwayswoody. Some species alwayshave a well-definedtrunk(e.g.,S. acuminatum, S. trichonelron), while othersaremany-branchedfromthe base. Variation in habit is common within species and is often correlatedwith habitat. STEMS The young erect stem is monopodialwith alternate leaves arrangedin a 2/5 phyllotaxic spiral. When the reproductivephasebegins, stem growthbecomes sympodial(see Danert,1958, 1967, 1970;Child,1979;Child & Lester, 1991; Bell & Dines, 1995, for diagramsand explanations of terminology concerning sympodial branchingin Solanum).Eachinflorescenceis developmentally terminal,and stem growthproceeds from an axillarylateralshoot. This shoot is againterminatedby an inflorescence, the stem continuationis initiated in the axil of the leaf subtendingthe inflorescence,andso on (Danert,1967). Thusall membersof the Solanaceae are sympodialwith determinatemodules (Bell, 1991; Bell & Dines, 1995). The numberof leaves in a sympodialunitandtheirarrangementareimportantcharacters in sect. Geminata.Sympodiain sect. Geminataare either1) plurifoliate,2) difoliate,3) difoliate-geminate, 4) unifoliate,or 5) "fasciculate"(Danert,1967)(see Fig. 1 for diagramsof these sympodial types). Plurifoliate sympodial units are considered to be primitive in Solanumby Danert(1967, 1970)andChild(1979),while the othertypes arespecializationswithinthe genus.The plurifoliatespecies in sect. Geminataareprobablysecondarily so; many of the species possessing this con-

MORPHOLOGY

9

B

E

F

C

D

G H

FIc. 1. Sympodial types found in section Geminiata. The colors (black & white) represent different shoot generations. Patterns of concaulescence can be easily seen. Diagrams modified from Danert (1967). A. plurifoliate sympodial units. B. trifoliate sympodial units. C. difoliate sympodial units. D. difoliate sympodial units with geminate leaves, inflorescence internodal. E. difoliate sympodial units with geminate leaves, inflorescence leaf-opposed. F. difoliate sympodial units with geminate leaves, leaf pair markedly anisophyllous, inflorescence leafopposed. G. unifoliate sympodial units, inflorescence leaf-opposed. H. fasciculate sympodial units, inflorescence overtopping the new shoot.

dition are shrubsof river beds subject to periodic inundation,andarespecializedin otherfeatures.Thegeminateconditionso common in sect. Geminata(Fig. 1E) is caused by suppressionof growth in various partsof the stem axis, andthe two leaves of a geminate cluster are from differentshoot generations. Architecturalmodels have been applied to various

genera in the Solanaceae (Bohs, 1994; Bell & Dines, 1995), andbranchingpatternsarecommonly very precise andpredictableat the species level. In the original architecturalscheme of Halle et al. (1978) membersof the Solanaceaecan be classified in bothmonoaxialand polyaxialmodels,with threedifferenttypes ofpolyaxial models being present (Bell & Dines, 1995). Models

10

representedin the genus Solanum are Prevost's (species of Cyphomandra,see Bohs, 1994;Solanummorii, see below), Leeuwenberg's (sects. Brevantherum, (sect. Allophyllum,Holophyllaproparte),Chamberlain's Pteroidea), and Mangenot's (sect. Solanum) (Bell & Dines, 1995). Membersof sect. Geminataseem to most with closely ressembleeitherthe modelof Chamberlain, monchasialbranchinganda sympodialtrunk,or the intermediateandmixed model describedby Bell & Dines (1995) from Lycianthes sanctae-clarae (Greenm.) D'Arcy. This mixed model involves a primarymodule terminatedby a flower and one dichasial branch,this producingtwo indefinite monochasia. The bud in the axil of the final leaf also producesa monochasium,resultingin a tierofplagiotropicbranches(Bell & Dines, 1995). Members of all the species groups in sect. Geminatahave architecturesimilarto thatof some species of Lycianthes, but detailed studies remain to be carriedout. Species in the S. confine and S. dolosum species groups conform more to the model of Chamberlain, but with the addition of tiers of plagiotropic branchesas describedforLycianthes.Specimenlabels on sheets in MPU indicate that S. leucocarpon conforms to Troll'smodel (Halle 2199) with long, plagiotropic branches,or to Pr6vost'smodel (Prevost 177). Sheets of S. oppositifolium(Halle 2241) statethatthe plants conform to Troll's model and those of S. morii (Prevost 1613, Halle 2377) to Prevost'smodel. These collections date from the 1970s and undoubtedlyrepresent field assessments, but it is clear thatmore work needs to be done with members of this group. Architecturemay provide useful charactersfor the recognition of monophyletic groups within sect. Geminata. The sympodialgrowthpatternof reproductivestems often gives them a zigzag appearance,particularlyin the Solanum confine and S. dolosum species groups. Branchorientationvariesbetween species and species groups,but is difficult to assess fromherbariummaterial. Forthose species I have seen in the field, this characteris discussedin the species descriptions.Branches are generally held upright,or at an approximately45? angle. In the S. confine andS. dolosum species groups however, the branchesare held parallelto the ground, giving the shrubsa pagoda-likeaspect.This may be an advantagefor light gatheringin the darkforest understory,the common habitatfor these species. The barkof these woody plants is quite variablein color and textureon younger stems, rangingfrom reddishandexfoliatinginSolanumtuerckheimii, grayishand smoothinS. arboreum,to brightgreenandstronglywinged in S. chlamydogynum andS. oetens. Inthosespeciesthat attainlarge diameters(15-20 cm dbh), the barkof the older stems and trunksis usually smooth and grayish-

FLORA NEOTROPICA

however,herbariumlabels tinged.InS. chlamydogynum mentionthatwings on the stems arepersistenton stems up to 10 cm in diameter.Woodof species attainingsuch diametersandheights has a distinctsmell of rawpotatoes when cut and is quite soft. Detailed descriptions of the wood anatomyof threespecies of sect. Genminata (S. oblongifolium,S. trichoneuron,andS. triste)areprovided by Carlquist(1992). Solanumoblongifoliumhas no growthrings in the wood, perhapsdue to habitator relativelycontinuousgrowth(Carlquist,1992). Rhomboidal crystals and crystal sand occur in the wood of both S. triste andS. trichoneuron(Carlquist,1992). LEAVES Leaves in section Geminataare simple andusually entire,and at first glance appearquite uniformamong species.Amongthe leaf charactersusefulin distinguishing between species are leaf size, leaf shape, margin characteristics,venation, vestiture,and apex andbase shape. If present,trichomesareusually denseror only found on the undersurface.The trichomedistribution patternis an extremely useful characterfor some species, but in otherspecies can be extremelyvariableand of little use taxonomically.In general,membersof the Solanumsessile and S. oblongifoliumspecies groups have the largest leaves, while membersof the S. colnfine species group have the smallest leaves. Several species (S. monadelphum,S. amnicola,andS. imberbe) havenarrowlylinearleaves.Thesespeciesgrowin stream beds andaresubjectto periodicseverewaterloggingand swift currents.Theirpeculiarleaf shape(forSolanum) maybe an adaptationto theirextremehabitatandis typical for rheophytes(van Steenis, 1981). As in therestofthe family,isolatedidioblastsof"crystal sand" (tiny calcium oxalate crystals) occur in the parenchymalayerof the leaflaminae(Solereder,1908; Carlquist,1992).Thesize andnumberof theseidioblasts appearto differ between species, but furtheranatomical work is neededbefore conclusions can be drawnas to theirusefulness as differentiatingcharacters. INFLORESCENCES Inflorescences of section Geminataare variously modified helicoid cymes. They are usually bore opposite a geminateleaf cluster,or single leaf in the case of those species with unifoliate sympodia.They occasionallyappearto be intemodalor extra-axillary,due to adnationof the stem and inflorescence axes (Danert, 1967). This characteristicoften varieswithina species, and even within an individual plant. In the Solanum sessile andS. oblongifoliumspeciesgroups,lateralshoot growthoften lags behindinflorescence expansionand development, causing the inflorescence to appear

11

MORPHOLOGY

branch-terminal.The branch-terminalinflorescence occurs in sect. Holophylla (G. Don) Walp.and in sect. BrevantherumSeithe, but in these groups no further branchelongation occurs (see Knapp, 1989), and the inflorescenceterminatesa shoot. This superficialsimilarityhowever,has resultedin severalof the species of the S. sessile complex being previously placed in sect. Holophylla.The degree to which the lateralshoot lags behindtheinflorescenceis variable,bothbetweenplants in a populationandbetweenbrancheson an individual. Branchescollected in different stages thus appearto have completely differenttypes of inflorescences and often were classified as different species in different parts of the genus (see discussion of S. sessile and Knapp, 1991b). See Fig. 1 for diagrams of inflorescence positions commonly found in sect. Geminata. The inflorescence in species of sect. Geminata is simpleto manytimes branched,stoutto threadlike,and with few to many flowers. The arrangementof flowers on the inflorescencecan be deducedfromthe scars left by the flowerandfruitpedicels when they fall. This is a useful characterand has been largely overlooked (but see D'Arcy, 1973).Pedicelscarsareclosely spaced(oftenoverlapping)in some speciesgroups,andwidely and irregularlyspacedin others.The scarsareflushwith the inflorescence axis,notleavingsmallplatforms(see Knapp, 1989), pegs (see Bohs, 1994), or sleeves (see Knapp, 1989) as in otherpossible relatedgroups (see above). In general inflorescences are simple, but in the Solanum sessile and S. oblongifoliumspecies groups and in some species of the S. amblophyllumspecies grouptheyaremanytimesbranchedandoftenverylarge. Diagramsof the differenttypes of inflorescencesoccurringin sect. Geminataarepresentedin Fig. 2.

A

^ B

C FIG. 2. Inflorescencetypes in section Geminata.A. stout with closely spaced pedicel scars. B. filiform with widely and unevenly spaced pedicel scars. C. branched.

lobes) have distinctive and peculiar calyx lobe morphol-

ogy; these variations are discussed in the individual species treatments. Species of section Geminata have amongst them some of the smallest flowers in Solanum.Averagecorolla diameteris about 8 mm, but some species have corollas as small as 4 mm in diameter.The corolla is usually white or greenish-white, but occasionally the FLOWERS edges of the lobes are tinged purple, particularilyin As in most othersolanums,flowers of sect. Gemin- plants growing in full sun. The corolla is deeply lobed ata arepentamerous,actinomorphic,andgamopetalous. and stelliformto shallowly lobed andmorepentagonal Floweringpedicelsareusuallydeflexed andquiteslen- in outline. The lobes are usually rathernarrowand ofder.Constrictionsat the base of the calyx tube are use- ten the corollatube is minute.A few species have shalful charactersin severalspecies andspecies groups.The lowly lobed corollas with a broadlyopen corolla tube, calyx is 5-parted,with variously developed deltate to and the flowers are thus more pentagonal in outline. lanceolatelobes. In some species the lobes are minute, The degreeto which corollalobes arereflexedat anthemereprojectionsfromthe rim of the calyx tube, while sis is an importantcharacter.This characteris easily in others,such as Solanumtanysepalum,S. irregulare, seen in live plants, but with more difficulty in pressed andmembersof theS. pseudocapsicumspecies group, specimens.Onlya few species(Solanumtrachytrichium, the calyx lobes are long-triangularand occasionally S. campaniforme,andS. validinervium)have campanualmost equal to the corolla lobes in length. The calyx latecorollas;these arealso those species with shallowly lobesarenotusuallyreflexed,butreflexedcorollalobes lobed, pentagonalcorollas. In many species the lobes at anthesisare characteristicof several species. A few are held planar (i.e., perpendicularto the pedicel or species (notablyS. abitaguense with its closed calyx spreading) at anthesis, while in others the lobes are envelope, S. platycypellumwith its flattened, almost slightly reflexed. The orientationof the corolla lobes lobeless calyx, andS. pastillum with its globose calyx does not generallychange over the reproductivelife of

FLORA NEOTROPICA

12

A

B

D

C

FIG. 3. Corolla types in section Geminata. A. campanulate. B. lobes planar. C. lobes strongly reflexed. D. with unequal anthers.

a flower, which is usually five days to a week. Many membersof the S. nudumspecies groupare characterizedby theirslightlyreflexedcorollalobes. Specieswith stronglyreflexedcorollalobesarefoundin theS. confine, S. arboreum,andS. unifoliatumspecies groups.In species with this flower type the lobes are held parallelto the pedicel, and only begin to close aroundthe anthers when flower wilting begins. Interestingly,these arethe smallestflowersin the section.The stronglyreflexedcorollalobes may be a strongsignalto pollinatorsin these otherwise inconspicuous flowers. See Fig. 3 for diagramsof the majorcorollatypes in sect. Geminata. The anthers of species of section Geminata are small, oblong, and strongly to weakly connivent. The anthersare usually brightor pale yellow, but those of SolanumleucocarponandS.pseudocapsicumarebright orange-yellow.The color is not the color of the pollen, butof the antherwalls, andmay serve to attractpollinators.Theanthersof some membersoftheS. leucocarpon species group (notably S. leucocarpon) are strongly conniventandsuperficiallysimilarto theanthersof some species of sect. Pachyphylla(the genus Cyphomandra, see Bohs, 1994). They lack, however, the strongly developed antherconnective of that section. As in all species of Solanum,the anthersareporicidalat the tips. In sect. Geminata,however,as the anthersdrythe pores split open towardthe base of the antherandlengthento slits (see Bonner& Dickinson, 1989, for a detailedanatomicaldescriptionof a similarsituationinLycopersicon esculentumMill. [SolanumlycopersicumL.]). At dehiscence, the brightwhite lines along the lines of opening indicatetheprescenceof calciumoxalate,whichhas been implicatedin the dehiscence process itself and is commonin the Solanaceae(Bonner& Dickinson,1989, 1990; D'Arcy et al., 1996). The filaments are much shorterthanthe anthers,areinsertedin the corollathroat, andarebasallyconnate.Unequalanthersizes arefound

in two species, S. pseudoquina and S. reitzii. In these species, two of the anthersandtheirfilamentsare long, and the otherthree are short (Fig. 3D). The anthersin thesetwo species areless tightlyconniventthanthose in otherspecies, arecurvedin longitudinalsection,andthe pores do not split open with age or upon drying. Pollen grainsof section Geminataspecies, as in all other species of Solanum, are tricolporate,pouting at the equator,and have a minutely granularexine (see Anderson & Gensel, 1976; Punt & Monna-Brands, 1980; Edmonds, 1984; Bohs, 1994). Pollen grains of membersof sect. Geminataare on the orderof 12 gm in diameter.No notabledifferencesin pollensize ormorphology have been found between species or species groups in sect. Geminata(see Fig. 4). The ovaries in most species of sect. Geminataare glabrous, but some members of the Solanum sessile, S. nigricans, S. arenarium, and S. nutans species groupshave denselypubescentovariesandstyle bases. The trichomes are persistent on the maturefruit; furtherdetails are discussed with otherfruitcharacters. FRUITS Fully developed ovaries of species of section Geminataare globose or depressed-ovoidandusually glabrous.Occasionallythe fruitsaresomewhatpointed at the apex (e.g., Solanum nigricans, S. stipulatum). The fruitis a bicarpellate,usually bilocularberrywith axile placentation.They are usually few-seeded (1050 seeds), an unusual condition in Solanum. In some groups, particularlythose with flattened seeds (see below), berriesaremore typical forSolanumandmay contain 100-200 seeds. Berries are usually green or yellowish-green,andrelativelyhardat maturity.Many species have fruitwhich becomes softer and yellowgreenvery rapidly(overnight,or in a single day). These fruits are taken by bats or birds almost immediately

13

MORPHOLOGY

_______20_gm FIG. 4. Pollen grains in Solanum section Geminata. All polar views. A. S. arboreum, Costa Rica (Bohs 3010). B. S. rovirosanum, Costa Rica (Khan et al. 1319). C. S. incomptum,Costa Rica, (Tonduz13666). D. S. leucocarpon, FrenchGuiana (Sagot 454). E. S. leptopodum,Colombia (Philipson et al. 2184). F. S. nudum,Belize (Whitefoord9065).

14

FLORA NEOTROPICA

- ::;1: 1

A ...

''-4iy

;

..

0**

..

FIG. 5. Fruiting pedicel types in section Geminata. A. erect (Solanum arboreum). B. slightly deflexed (S. nudum). C. strongly deflexed and expanded at the apex (S. tenuiflagellatum).

(Knapp,1986a;Dinerstein,pers.comm.).The fruitflesh is usually green or pale greenish-white and hard,becoming waterywhen the fruitripens.Even in ripe fruit the flesh is not sweet. Members of the Solanum pseudocapsicum species groupandS. spirale of the S. nudumspecies grouphave more typically solanaceous brightorange or red-colored, soft, juicy fruits. The majorityof species in sect. Geminatahave glabrous fruit, but those taxa with pubescent berries fall intotwo generalcategories.Firstarethose species with simple uniseriateor dendritictrichomeson the mature berries.Solanumgnaphalocarpum,S. oblongum,and some members of the S. sessile, S. nigricans, and S. nutansspecies groupshave this type of pubescentfruit. The trichomespersistthroughovary developmentand are presenton the matureberry.The trichomes on the maturefruit are usually similar to those found on the restof theplant.Importantdifferencesaredetailedin the species accounts. The berry of S. gnaphalocarpon is extremein this regardandis denselycoveredwith elongate, echinoidtrichomessuch thatthe pericarpis never visible. Solanumabitaguense,S. cucullatum,andsome membersof the S. arboreumspecies group also have pubescent berries, but of a completely different and probablyindependentlyderivedtype. In these species, the ovary is glabrous at anthesis, but during fruit de-

velopmentacquiresa coatingof grayishorpinkishpapillate trichomes.Thesepersiston the maturefruit,butare inconspicuousunless viewed at high magnification. The orientationanddegreeof elongationand/orexpansion of pedicels in fruitare importantcharactersin sect. Geminata.The pedicels elongatesomewhatin the Solanum nudumspecies group, but are not markedly differentin shapeor orientationfromtheirconditionat anthesis.Severalgroupsof species (S.pseudocapsicum, S. arboreum,some speciesoftheS. nudumandS. sessile species groups)arecharacterizedby erect,woody, fruiting pedicels. These pedicels are not much elongated, but aregreatlyexpandedin diameterandat fruitmaturityarethickandwoody alongtheirentirelength.Fruits in these species areheld above the leaves. Anotherextreme is seen in those species with elongate pedicels and pendantfruits (S. confine and S. dolosum species groups).Here the fruitingpedicel is two to threetimes as long as the floweringpedicel andis usuallyexpanded andrathercorkyjust below the calyx tube. The berries of these species hang below the leaves. The effective pedicel length is often increased even furtherby the elongate inflorescence, which in some species can be up to 20 cm long (S. tenuiflagellatum and S. leptorhachis). Generalized fruitingpedicel types are shown in Fig. 5.

15

MORPHOLOGY Table IV Seed type distribution in the species groups of section Geminata. FLATTENED-RENIFORM

SEEDS

Solanum oblongifolium species group Solanum pseudocapsicum species group Solanum nudum species group Solanum leucocarpon species group Solanum nutans species group Solanum amblophyllum species group

SEEDS Seeds of Solanumspecies areusuallyflattened,reniform,andhavesomewhatincrassate(thickened)margins. In sect. Geminata,six species groupshave this type of seed (hereafterreferredto as flattened-reniformseeds, see Table IV). Flattened-reniformseeds are usually small, 1-2 mm in length,yellow or darkbrownin color, and the surfaces are minutely pitted (Fig. 6A). The incrassatemargin is paler than the body of the seed. The remainderof species groups in the section (the majority,see TableIV) have large,roundedseeds with very thin seed coats, throughwhich the spiralembryo characteristicof Solanumcan be clearly seen (hereafterreferredto as ovoid-reniformseeds).Ovoid-reniform seeds are larger,usually 2-4 mm in length, pale tan or brightgreenin color,andaregenerallyfew in any given fruit (Fig. 6B,C). They dry out easily due to the thin seed coat, and are difficult to germinateafterdesiccation. These seeds are unusualin the genus. Two of the few othersectionsin whichtheyoccuraresect.Pteroidea Dunal(Knapp& Helgason,1997)andtheS.thelopodium species group (Knapp,2000), but in those groups the seed coat is not thin and the testa walls are thickened (see below). The generalappearanceof the ovoid-reniform seeds in sect. Geminatais so aberrantthat they have been misinterpretedas not belonging to Solanum at all. For instance, Bitter, in a note on the holotype of S. tenuiflagellatum, commented that the fruit in the packet probably did not belong to the specimen and probablywas not even a Solanum(see discussion of S. tenuiflagellatumfor text of his comment). The seeds of S. tenuifagellatunm aretypicalof the ovoid-reniform type found in sect. Geminata. Removalof the outerseed coatwall revealspatterns of lateralcell wall thickeningthatdiffer between taxa (Soueges, 1907; Edmonds, 1983; Lester & Durrands, 1984; Knapp& Helgason, 1997; Whalen & Caruso,

OVOID-RENIFORM

SEEDS

Solanum deflexiforum species group Solanum arboreum species group Solanum unifoliatum species group Solanum robustifrons species group Solanum narcoticosmum species group Solanum nigricans species group Solanum arenarium species group Solanum sessile species group Solanum confine species group Solanum dolosum species group

unpubl.).Throughoutthe familySolanaceaelateralseed coat walls arecomposed of thickenedpyramidalbases toppedby elongate,lignifiedprojections(Sou6ges,1907; Mohan& Singh,1968;Edmonds,1983;Hoare& Knapp, 1997). The projectionsareconnectedby thinner"membranous"sections of material(see Figs. 6D, 7A,C,D). Flattened-reniform seedsin sect.Geminata(see TableIV) conformto thispattern.Ovoid-reniformseeds,however, have a very differentlateralwall structure.Lateralcell walls in these seeds are thin, and do not possess elongate finger-like projections (see Knapp, 1991b; Fig. 7B). Seed shapeandfine structurearecorrelatedin sect. Geminata,andserveto dividethe sectionintotwo major groups: Groups I-VI have flattened-reniformseeds, GroupsVII-XVI have ovoid-reniformseeds. TRICHOMES Trichome morphology has been used extensively in Solanum taxonomy (Edmonds, 1982; Roe, 1971; Seithe, 1962, 1979;Seithe& Anderson,1982;Whalen, 1978, 1979;Whalenet al., 1981).Eightbasic hairclasses occurringin Solanumaccordingto Seithe(1962, 1979) arelisted in the left-handcolumn of TableV, using her terminology.Standardizedtermsaregenerallyused in discussions of more complex hair classes (i.e; stellate hairs), but the terminology available for simple and branchedtrichomes does not adequatelyreflect their structuraldiversity.Roe (1971), in his attemptto present a standardizednomenclatureforSolanumhairs,went a long way toward solving this problem. His terminology is presentedin the right-handcolumn of TableV. Seithe(1962, 1979) consideredsimpletrichomes("finger hairs" in her terminology) to be primitive in Solanum. She believed them to representa good case of Haeckel's law of ontogeny recapitulatingphylogeny (Haeckel, 1883). Glandular"finger hairs" occur on the cotyledons and seedling leaves of all of the spe-

FLORA NEOTROPICA

16

:w,~, ~

_O~CSE~ !;":_It.

""

FIG. 6. Seed types in section Geminata A. flattened-reniform seed 50x (Solanum acuminatum, Knapp et al. 6343). B. ovoid-reniform seed 30x (S. sessile, Knapp & Mallet 6442). C. ovoid-reniform seed 30x (S. abitaguense, Knapp & Mallet 6180). D. enzymatically digested flattened-reniform seed showing the lateral cell wall thickenings 50x (S. nudum, Nee et al. 30019).

FIG. 7. Lateral seed coat wall thickenings in section Geminata. A. from flattened-reniform seeds 500x (Solanum leucocarpon,

Knapp & Mallet

6284).

B. from ovoid-reniform

seeds

500x (S. sessile,

Knapp & Mallet

6442).

C. flattened-reniform seeds with rectangular or square cells lOOOx(S. nudum, Nee et al. 30019). D. flattenedreniform seeds with sinuate-walled seeds lOOOx(S. campaniforme, Steyermark et al 88179)

17

MORPHOLOGY Table V in Solanum to trichomes (for definitions of these terms the reader Terminology applied is referredto the original publications). SEITHE(1962,1979) finger hairs branchlet hairs stellate hairs gland-tipped finger hairs gland-tipped stellate hairs

ROE(1971) simple trichomes uniseriate multiseriate stellate trichomes porrect stellate peltate multangulate multiradiate echinoid

multicellular glands prickles

dendritictrichomes furcate dendritic

bristles some modified trichome types glandular dendritic-echinoid

cies studied by Seithe, and are the least structurally complex of all of the hairclasses. Two distinctphyletic lines or transformationseries were hypothesized by Seithe: 1) gland-tippedfinger (simple) hairsto stellate hairs, and 2) gland-tipped finger (simple) hairs to branchlet(branched)hairs. Section Geminatahadtraditionallybeen characterized as those species that are glabrous or possessing only simple trichomes (D'Arcy, 1973; Gentry & Standley,1974; Seithe, 1962). Subsectionlndubitaria (see NomenclaturalandTaxonomicHistory)was characterizedby the possession of only branchedtrichomes (Seithe, 1962). Seithe (1962) recognized the close relationship of these two groups based on habit, flower and fruit morphology, and inflorescence and sympodial structure, and placed them in a single section: Oppositifolium.D'Arcy (1972), however, placed the two groups in different subgenera.While examining specimens in the course of this study, I found that in this group these two hairtypes differ only by degree, and variationandpolymorphismexist at all taxonomic levels. One memberof a pairof closely relatedspecies may possess branchedtrichomes while the other has only simple trichomes(Solanumchlamydogynumand S. obovalifolium); different populations of a single species may vary in their possession of simple or branched trichomes (S. leucocarpon, S. lindenii, S. confertiseriatum,S.pseudocapsicum);individualsin a

populationmay vary in their possession of branched or simple trichomes(S. confertiseratum);andfinally,a single individual may have simple trichomes on one leaf and branched trichomes on another (S. confertiseriatum,S. pseudocapsicum).It is clearthen that the characterof simple versus branchedtrichomes is not as simple as was previously considered in sect. Geminata.SectionGeminataas delimitedhereincludes both simple andbranchedtrichomespecies. Three basic types of unbranchedtrichomes occur in sect. Geminata.The most widespreadis the simple uniseriatetrichome,consisting of a single row of cells (Figs. 8A, 9A). The basal cell or cells surroundingthe basal cell are occasionally somewhatmodified (Figs. 8C, 9B), but generally the cells of the hair are of approximatelyequal size and shape (save, of course, the tip, which is pointed).The cell walls areoftenquitethin, and, if so, the cells collapse when dry.In the Solanum sessile species group however, the cell walls of uniseriatehairsarerelativelystiff andretaintheirshape even on herbariumspecimens. Simple uniseriatetrichomes arefoundto some degreein all species groups. The second type of unbranchedtrichome,the papillate trichome, is characteristicof the S. arboreumand S. robustifronsspecies groupsbut is also found to varying degrees in most of the other groups. Papillatetrichomes are technically a subset of simple uniseriate trichomes,but are so differentin appearanceandtaxo-

FLORA NEOTROPICA

18

A

C

B

D

FIG. 8. Unbranched trichome types in section Geminata. A. simple uniseriate. B. papillate. C. uniserate with multiseriate base. D. unicellular. All drawn approximately 10x life size.

19

MORPHOLOGY

d_f _ f.~?:;?"?~c~;c~flF~,~t~l~''~~ Cll~q~-~?~g~S~i~i'f c~-. ' ..

...

FIG. 9. Scanning electron micrographs of unbranched trichome types in section Geminata. A. simple uniseriate trichomes from the pedicel of Solanum oblongum, 500x, (Knapp & Mallet 6343). B. papillate trichome from the new growth of S. arboreum, lOOOx,(BH 84:140). C. unicellular trichome with a broad multicellular base from S. trachytrichium, 480x, (Cabrera 69). D. uniseriate trichome with multiseriate base from the new growth of S. validinervium, 204x, (Knapp & Mallet 6782).

nomic distributionamong species that I preferto treat them as a separatehairtype. They are uniseriate,or in some cases uni-cellular,with thin, crumplingwalls in dry or preservedmaterial.The distal cell of the hair is rounded(Fig. 9B). The fine structureof papillate trichomes is difficult to assess with driedmaterial,as the trichomescollapse andthe numberof cells in each hair is nearlyimpossible to determine.Papillatetrichomes are usually found on the young leaves and stems, and on the inflorescences in those species where they occur.In dry specimens they are often reddish in color. The last and least common type ofunbranchedtrichome is the multiseriatehair,foundin only a few species, e.g., SolanumvacciniiflorumandS. validinervium

(Figs. 8C, 9D). Simple multiseriatetrichomes consist of severaladjacentrows of cells. The structureis similar to that of the simple uniseriate trichomes. In S. vacciniiflorum,simplemultiseriatetrichomesarefound on the upperportionsof the stems in conjunctionwith simple uniseriatetrichomes. Fourbasictypesof branchedtrichomesoccurin sect. Geminata:echinoid/dendritic-echinoid (Christmas-treelike),arachnoid,dendritic,andfurcate(Fig. 10).Themost complex structurallyare the echinoid and tree-liketrichomes. These types of hairs occur in some species of the Solanum nutans species group and S. gnaphalocarponof the S. arenariumspecies group.Thetrichome axis or stalk is either uniseriate or multiseriatebut is

FLORA NEOTROPICA

20

B

A

C

D

FIG. 10. Branchedtrichome types in section Gemninata (see text). A. echinoid. B. arachnoid.C. dendritic. D. furcate.

oftenobscuredby the numerousbranches.Therays(side branches)alwaysprojectat all anglesalongthe axis and are very closely packed and occasionally branched themselves (Fig. O1A).Two main types of echinoid trichomesoccur in sect. Geminata:1) short,sessile trichomes with numerous branchesobscuring the stalk aretermedechinoid;and2) elongate densely branched trichomes (Roe's dendritic-echinoid)are termedhere "Christmas-tree-like," following Seithe (1962, as see Knapp, 1989). "tannenbaumartig," Manyspecies previouslyconsideredto be members of subsectionIndubitaria(includingthe lectotypespecies SolanumbrachystachysDunal= S. nigricans)have arachnoidtrichomes(Fig. 10B). Arachnoidtrichomes

are complex multicellular hairs that appear felt-like when seen with the nakedeye or the dissecting microscope. The cells are usually quite small, thin walled, and the hair itself is an irregularlybranched,fragile structure.Arachnoidhairsaresimilarin appearanceand probably structurallyrelated to papillate trichomes. These trichomesare often deciduousand so areabsent from older leaves. Like papillate trichomes, they are densest and most commonly seen on the new growth and the inflorescences of the species in which they occur. As a leaf matures, it often loses its coating of arachnoidtrichomes (e.g., S. maturecalvans). The third and most common type of branchedtrichome is the dendritic trichome (Asthaare of Seithe,

21

MORPHOLOGY

for leaf

underside

of

. nudum

500x

(from

seed

of Nee

et al. 30019,

BH

84:177).

C. glandular

trichomes

along

a vein on the abaxial leaf surface of S. oblongum, 500x, (Knapp & Mallet 6343). D. glandular trichome of S. Mallet 6343). 2000x, (Knapp oblongum, 6343). (Knapp & Mallet oblongum, 2000x,

1962), consistingof a uniseriatemain stem with few to several divergent branches seemingly distributedat random(Figs. 10C, 11B). The numberand length of the branchesvaries considerably,but any given branch is usually only one or two cells long. The branchesare not as densely packed as those of the echinoid or treelike trichomesandthe stalkis always visible. Dendritic hairsare found in several species groupsand are often foundin conjunctionwith simple, uniseriatetrichomes (see also Knapp, 199 la). The fourthtrichometype occurringin theseSolanum species is the furcate hair, technically a type of dendritichair.Furcatetrichomesarehairsthatare divided into two branches(Figs. 10D, 11A). Usually one cell is branched,and appearsbud-like. Roe (1971) com-

mentsthat"truefurcatehairsareprobablynot common in Solanum."They arevery commonin sect. Geminata. These small trichomes often appear simple and uniseriateunderthe dissecting microscope, but when viewed at high magnificationstheirstructurebecomes clear (Fig. 11A). The style base and fruittrichomesof many of the species of the S. sessile species groupare of this type. All of the largertrichome types in sect. Geminata are non-glandular.On mature foliage glandular trichomes occuron the veins andnew growthbutthe hairs themselves are always tiny (Fig. 11C,D). I have found at least some of these minute, glandulartrichomes on almost all the species of sect. Geminata. The distribution of trichomes on the plant, their

FLORA NEOTROPICA

22

color in driedmaterial,andtheir structureare all good charactersfor identificationat the species level, especially when used in conjunctionwith floral and inflorescence characters.Some species groups are characterizedby the presenceof a particulartrichometype or types:thick-walled,golden,simpleuniseriateor furcate trichomes in the Solanum sessile species group (see Knapp,1991a); reddishpapillatetrichomeson the new growth of the S. arboreumandS. robustifronsspecies groups; golden dendritictrichomes in the S. pseudocapsicum species group; densely brancheddendritic trichomesin theS. arenariumspecies group.Untilthese trichome types are betterunderstoodstructurallyand developmentally it seems most reasonablenot to use them in phylogenetic analyses. Trichome morphology has been a mainstay in Solanum taxonomy (Seithe, 1962, 1979), but careful examinationshows thatit cannotbe relied upon to the extent that it is now used, particularilyin this group. Trichome morphology of the stellate-haired groups (subgenusLeptostemonum)has been relatively better studied, and a few studies exist of non-stellate-haired groups (Edmonds, 1982; Seithe & Anderson, 1982; Bohs, 1994). It is quite clear however that much remains to be done with trichomemorphology and evolution in Solanum, particularlywith groups of nonspiny species without stellate hairs.

tetraploidthroughoutits range.Materialreportedas S. superficiens (a synonym of S. spirale) from Bogor, Java,Indonesia(see TableVI), is actuallyS. diphyllum, a nativeof Mexico commonlycultivatedas anornamental shrubthroughoutthe tropics and subtropics. Chromosome size has been used to support the monophyly of some groups within Solanum (Bohs, 1994). Cyphomandra(Solanumsect.Pachyphylla)and its sistergroup,Solanumsect. Cyphomandropsis,both have uniformly large (3-14 gfn with an average of 8 pam)chromosomeswith correspondinglylargeamounts ofDNA (Bemardello & Anderson, 1990; Bohs, 1994; Pringle& Murray,1992;Moscone, unpubl.).Members of sect. Geminatahave chromosomes of a more typical Solanummorphology:very small (less than4 tm) andindistinct.However,an intriguingexceptionto this arethe chromosomesofS. tanysepalum,which arequite large and look more like those of a typical Cyphomandra.Detailed measurementsare lackinghowever, andthese may prove to be of greatutility in the assessment of chromosomesize as an informativecharacter in sect. Geminata.

CHROMOSOMES

Species of section Geminataoccur throughoutthe Neotropics,fromca. 20? N latitudeto nearly32?S latitude (Fig. 12). Species of the section occurin all countries of continentalSouthAmericaandthroughoutthe WestIndies.A single species,Solanumn spirale,is widespreadin the Old WorldAsian andAustraliantropics, rangingfrom Chinato Australia.The greatestspecies diversity in sect. Genminatais found in the western Amazonbasin,on the easternslopesof the Andes.Other areas of high species diversity are montane coastal VenezuelaandtherelictAtlanticmoistforestsof eastern Brazil.In SouthAmericamost species arefoundin relatively low elevationforests,andmany species occurin the Amazonbasin, but in CentralAmericathe groupis largelymontane.This may reflectthe geological youth of CentralAmerica, andthe possible derived statusof the species occurringthere.Most species of the section, in both South and CentralAmerica, grow in areas of high rainfall (2500 to 12000 mm per year), but some species are only found in dry habitats.Refer to Table VII for the distributionsof membersof the section in differenthabitats(life zones) in the Neotropics. Speciesof sectionGeminataareunusualin Solanum in beingtreesandshrubsof forestinteriors.Many,if not most,otherSolanumspeciesareplantsof disturbedareas andareoftencharacterized as noxiousweeds. Roadsides

Polyploidy has been importantin the evolution of some groupsof Solanum,e.g., sect.Petota (see Correll, 1962; Magoon et al., 1962; Hawkes & Hjerting,1969; Hawkes,1979;Hawkes& Hjerting,1989;sect.Solanum, see Edmonds,1972, 1979).Thebase chromosomenumber in Solanum is n = 12. Three chromosome counts representingtwo species of sect. Geminatahave been reportedin previousliterature(D'Arcy, 1969; Randall & Symon, 1976). Aceto- and lacto-propionicorcein squashesof pollen mothercells fromplantsgrowingat the Bailey HortoriumConservatoryandcollectedin the field in modified Camoy's solution (4:3:1 chloroform: ethanol:acetic acid) have raisedthe numberof counts to 58 andthe species countedto 40. Documentedchromosome counts for species of sect. Geminataarelisted in TableVI. Voucherspecimens aredocumentedin the individualspeciesdescriptions,andaredepositedin BH, BM, or MO. All species, except S. spirale, are consistently diploid with n = 12. Thus speciation and evolution in sect. Geminatahas largelyoccurredatthe diploid level. Solanumspirale is tetraploid,with n = 24. Only populationsfrom Australiahave been counted, andS. spirale occurs from Australia through Indochina to India. Authentic material from Indonesia, India, or Indochinais needed to determinewhetherS. spirale is

ECOLOGY AND NATURAL HISTORY HABITATS AND DISTRIBUTION

23

ECOLOGY AND NATURAL HISTORY

Table VI Chromosome numbersin sectionGeminata.Collectionsby Knappandco-collectorsaredesignatedby K followedby number:details canbe foundin specimenscited.BH vouchersreferto plantsgrownin the greenhousesat the L.H.BaileyHortorium, Ithaca,NY. TAXON Solanum abitaguense S. amblophyllum S. aphyodendron S. arboreum

S. campaniforme S. confertiseriatum S. confine S. cucullatum S. diphylluni S. dissimile S. erosomarginatum S. foetens S. gratum S. hypaleeurotrichum S. hypocalycosarcum S. leptopodum S. leptorhachis S. leucocarpon

S. maturecalvans S. nudum

S. oblongifoilun S. oblongum S. ombrophilum S. oppositifolium S. pastillum S. psetudocapsicum S. ramonense S. ripense S. robustifrons S. rovirosanum S. sessile

S. sieberi S. spirale S. tanysepalum S. tenuiflagellatum S. triste S. tuerckheimii S. turgidum S. vaccinilflorum S. yanamonense

NUMBER n = 12 n = 12 n = 12 n = 12 n = 12 n = 12 n = 12 n = 12 n = 12 n = 12 n = 12 n = 12 n = 12 n = 12 n = 12 n 12 n = 12 n = 12 n = 12 n = 12 n = 12 n = 12 n = 12 n = 12 n 12 n = 12 n = 12 n = 12 n = 12 n = 12 n = 12 n 12 n = 12 n = 12 n = 12 n= 12 n= 12 n = 12 n = 12 (2 n = 24) n = 12 n 12 n = 12 n = 12 n = 12 n = 12 n = 12 n = 12 n = 12 n = 12 n = 24 n = 24 n = 12 n = 12 n = 12 n = 12 n = 12 n = 12 n = 12

SOURCE Ecuador. Tungurahua.Bafios-Mera. (K 6177) Peru. Lima. Huarochiri.Rio Macachala. (K 6309) Panama. Chiriqui. Fortuna. (K 4136) Ecuador. Tungurahua.Rio Topo. (K 6181) Costa Rica. Heredia. Finca La Selva. (K 802) Panama. Canal Area. Pipeline Rd. (K 4862) Venezuela. Tfchira. Las Dantas. (K 6835) Venezuela. Bolivar. Gran Sabana. (K 6734) Ecuador.Los Rios. Rio Palenque.(K 6220, 6226, 6228) Peru. Pasco. Rio Palcazu. (K 6649) Ecuador. El Oro. nr. Las Balsas. (K 6266) Florida. Alachua. Gainesville (D'Arcy, 1969) Florida. Tampa (BH 85:234) Venezuela.Merida.Chorreralas Gonzalez.(K 6817) Venezuela. Trujillo. Bocon6-Trujillo. (K 6777) illo.llo. Venezuela. Trujillo. Bocon (K 6788) Venezuela. Merida. Chachopo. (K 6799) Venezuela. D.F. P.N. El Avila. (K 6872) Ecuador. Loja. JardinBotanico. (K 9085, 9094) Ecuador. El Oro. nr. Las Balsas. (K 6265) Peru. Loreto. Yanamono. (K 6610) Ecuador. Pichincha. Tinalandia. (K 6160) Colombia. Cundinamarca.(K 889, BH 79:376) Venezuela. Tfchira. Las Dantas. (K 6836) Ecuador. Morona-Santiago.Gualaquiza. (K 6241) Peru. Cuzco. Ollantaytambu.(K 6351) Costa Rica. Puntarenas.P.N. Corcovado. (K 887) Panama. Veraguas. Rio Sta. Maria. (K 4325) Venezuela. Merida. La Mucuy. (K 6791) Ecuador. Napo. Rio Latas. (K 6200) Peru. San Martin. Cufiumbuque.(K 6480) Peru. San Martin. Tarapoto-Yurimaguas.(K 6515) Ecuador. Loja. Loja-Zamora.(K 6247) Peru. Pasco. Rio Cacazu. (K 6628, 6636) Venezuela. Aragua.P.N. Henri Pittier.(K 6842, 6847, 6857) Peru. Loreto. Yanamono. (K 6618) Peru. Pasco. Laguna. (K 6656) Costa Rica. Puntarenas. Monteverde. (K s. n., 6063) Cultivated. Gerasimenko & Reznikova, 1968; Randall & Symon, 1976 Panama. Chiriqui. Pate Macho. (K 4270) Venezuela. Merida. La Carbonera.(K 6813) Peru. San Martin.Cufumbuque. (K 6474) Costa Rica. Heredia. Finca La Selva. (K 804) CostaRica.Puntarenas. Monteverde.(K 844, 848, 861) Ecuador. Napo. Cascada San Rafael. (K 6207) Peru. Pasco. Oxapampa. (K 6316) Peru. Cuzco. Quincemil. (K 6391) Peru. Amazonas. Rio Chiriaco (Imaza). (K 6569) Venezuela. Falc6n. Chichiriviche. (K 6704, 6707) Australia. (BH 81:139) Australia. Queensland. Toonumbar.(Randall & Symon, 1976) Venezuela. Aragua. P.N. Henri Pittier. (K 6856) Venezuela. D.F. above Carayaca.(K 6859) Venezuela. Sucre. Cumanacoa.(K 6750) Costa Rica. Puntarenas.Monteverde.(K 610, 872) Venezuela. Sucre. Manacal. (K 6765) Costa Rica. San Jose. Cerrode la Muerte.(K 790) Peru. Loreto. Yanamono. (K 6613)

FLORANEOTROPICA

24

I

WI

-

r -L77

J.

---Jl-

* I3

-- .

!3

g!Je

I

-r

,

?.

g

I

l

l

CT

I-

-

-

i Z

-

- I

.

_^

--

w

--

-

-

....: ........ ....... :.........

FIG. 12. Worldwide distribution of Solanum section Geminata.

Table VII Distribution of species of section Geminata in habitats in the Neotropics. Forest types modified from Holdridge (1967) and Heuck & Seibert (1972). For habitats and specifics of individual species, refer to the species descriptions. The total in this table exceeds the number of species accounts in this monograph because many taxa occur in more than one habitat. HABITAT

NUMBERS OF SPECIES

above tree line (incl. paramo) montane rain forest premontane rain forest lowland tropical rain forest tropical and subtropical deciduous forest savannah& savannahedges (incl.camposlimpios& cerrados) Araucaria angustifolia forests

10 species 30 species 45 species 49 species 14 species 6 species 3 species

intropicalAmericaareprimehabitatsformanysolanums, particularlythe spiny species (see Nee, 1979;Whalen, 1979;Whalenet al., 1981). Those species now so common along roadsidesprobablywere rarebefore widespreadhabitatdestructionby manso alteredandincreased theirhabitat.Most species of sect. Geminataareprimary forest treelets and shrubs, often growing along small

streamsor in old light gaps. They rarelygrow in areas of high light intensity,or where the soil is likely to dry out for long periods.Light gaps createdby tree falls or by small streams in the forest understoryare natural andimportantpartsof the tropicalforesthabitat,andare essential in the maintenanceof the system (Hartshorn, 1978, 1980, 1989, 1990; Denslow, 1987). Many sec-

25

ECOLOGY AND NATURAL HISTORY

Table VIII Hymenopteraknown to vibrateSolanum flowers (classification mod ified from Michener, 1979; for referencessee text . HYMENOPTERA COLLETIDAE COLLETINAE:Caupolicana, Colletes, Ptiloglossa HYLAEINAE: Hvlaeus STENOTRITIDAE:Stenotritus HALICTIDAE HALICTINAE:4 ugochlora, Augochlorella, Caenaugochlora, Augochloropsis, Halictus, Lasioglossum, Neocorynura, Pseldaugochloropsis, Evvlaeus NOMIINAE: Nomia OXAEIDAE: Mesoxaea, Nofoxaea, Protoxaea, Oxaea ANDRENIDAE PANURGINAE: Protandrena, Psaenythia ANTHOPHORIDAE ANTHOPHORINAE:Anthophora, Centris, Epicharis, Exomalopsis, Melissodes, Amegilla XYLOCOPINAE:Xy,locopa APIDAE BOMBINAE: Bombiis, Eulaema, Euglossa MELIPONINAE:Melipona, Trigona

ondaryspecies germinateandgrow in these light gaps, but do not persistwhen the forestcanopy closes. Some species of sect. Geminatagerminateandgrow in these areas,but do persistas forestplants.Membersof theS. confine species group are all plants of shady forest understory,as are many other species in the section. The understoryspecies, both those startingin light gaps andthose fromdarkinteriors,can be classified as sparse(Rabinowitz,1981). Sparsespecies arethosethat arenot common locally, but may grow in few to many habitats,andover a large or narrowgeographicrange. Species of sect. Geminataare often both rareand endemic (sensu Gaston, 1994) and are also likely to be vulnerable or threatened(Davis et al., 1986), but the degree to which this is the case remains to be tested. Particularforestspeciesof sect.Geminataareoftenconfined to a single forest type, e.g., cloud forest, varzea forest, etc. Where a particularspecies is found, individuals are often ratherscarce and difficult to locate. Statementsaboutthe rangeof habitatsor the geographical rangeof species are difficult to make at present,as so few collections exist for any but the most common roadside species. Some of the forest species of sect. Geminataareknown from only a few collections, perhapsdue to theirprimaryforest habitat,but also due to their small populationsizes where they do occur. Extreme examples of these sorts of species areSolanum dasl,neuronfromthe borderof GuatemalaandMexico, S. vanamonense fromeaster Peru,andS. tenuiflagellatum from montanecoastal Venezuela. Not all species of sectionGeminataareprimaryforest plants.Mostmembersof theSolanumnudumspecies

group are shrubsand small trees of secondaryforests androadsides.Thesespeciesoftengrowin densemonospecificstandsandareusuallyrelativelycommonwhere they occur(with the possible exceptionofS. reitziiand S. intermediumof southeasternBrazil).Theserelatively weedy species grow in recenttreefalls in the forestbut do not persist in these habitatsonce the canopy closes andthe sitebecomesshady.Thesecondaryforestspecies of sect. Geminataoften attainmuchgreatersizes (when left uncut)thando the primaryforest species.Solanum acuminatum,a memberof the S. nudumspecies group, occasionally grows to be a tree 15 m tall and 20 cm in diameterin the ChanchamayoValleyin centralPeru. POLLINATION Many, if not all, solanums are buzz-pollinatedby bees (Bowers, 1975;Buchmann,1983;Buchmannet al., 1977;Linsley, 1962;Linsley& Cazier,1963, 1972;Liu et al., 1975; Michener, 1962). Vibratilepollination is relativelywidespreadin angiosperms,occurringin about 200 genera(Buchmann,1983). This phenomenonwas first documentedin Solanlumin 1962, but was known from other genera much earlier(Michener, 1962; see discussion in Buchmann, 1983). Bees of many families possess the ability to vibrateSolanumanthers(see TableVIII andreferencesabove). Interestingly,honey bees (Apis mellifera L.) are unable to vibrate the anthers,and arethus not good solanumpollinators.Bees extractpollen from the poricidal anthersby vibrating theirindirectflightmuscles locatedin the thorax,while they firmly graspthe anthercone and often "milk"the antherswiththeirmandiblesandlegs (Buchmann,198:3;

FLORA NEOTROPICA

26

..

A

B

FIG. 13. Bombus worker buzzing a Solanum pastillum flower. A. buzzing. B. cleaning and packing pollen in the scopa.

Macior, 1964; Michener et al., 1978) (see Fig. 13A). Cloudsof pollen arereleasedanddepositedon the sternum of the bee. The pollen is cleaned from the accessiblepartsof thebee andplacedin the scopaeitherwhile the bee is still hangingby a tarsalclaw fromthe flower (Fig. 13B), after the bee lands on a nearby leaf, or in flight (see Micheneret al., 1978, for mechanics). Pollination in Solanum thereforeis sterotribic. In some large-floweredspecies of Solanumonly largebees, such as Centris,Bombus, andXylocopa, are able to vibrate theentireanthercone andthuseffectpollination.Smaller bees often vibrateone antherat a time, such that their bodies do not make contact with the stigma. In these cases, small bees are ineffective as pollinatorsand are moreproperlyconsideredpollen thieves (see Eickwort, 1967; Whalen & Costich, 1986). Small bees and bees unableto vibrateanthersoften gleanpollen scatteredon thecorollalobesby buzzingbees (Linsley& Cazier,1963; Wille, 1963;Knapp, 1986a;Whalen& Costich, 1986). Pollen is collected for larval food and is the most importantproteinsourcefor all bees (Buchmann,1983; Michener, 1974). The only floral reward offered by solanums to their pollinators and/or floral visitors is

pollen. Nectar is never found in these flowers (see Buchmannet al., 1977; Knapp, 1986a). The anthersof most species ofSolanumarerelativelythick-walledand tough, as one might expect when bees manipulateand presumablycoulddamagethe anthers.Antherson older flowers of membersof sect. Geminataoften have damage dueto smallgleaningbees orto thosetryingto chew intothe anthersto "illicitly"removepollen(see Fig. 14). The antherwalls are usually brightyellow or orange, which is not due to pollen content of the anther.The pollen itself is pale cream, and the bright color of the antherwalls may serve to attractbees to the flower, or to deceive bees as to the contents of the anther(Vogel, 1978). Bees are apparentlyable to assess the pollen contentof anthersbefore buzzing them, as I have seen Bombusworkersapproachflowers,hovernearthemand then fly away. Invariablythese were flowers thatwere severaldays old, or thathadbeen visited by manybees previously. Pollen may have some odor, as in certain Myrtaceae(van Wyk,pers. comm. to D. Charlesworth; Kevan& Lack,1985),Dodecatheon Porsch, (Primulaceae; 1954), and some solanums(Buchmann,1983; D'Arcy et al., 1990), or perhapsthe bee perceives a change in

27

ECOLOGY AND NATURAL HISTORY

~~~~i. i 2

i...............................................

~~~~: ~.~... ~,~~ ,

...

~,

*.

;.

,

..........

ti'?

. 'E'; . ' 'a'j.

f.

~.

...

'7'

:...:~ ~i~ii:iiJi!!ii!i: :! ... . ..... .....

;i~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~

tSt !011;,''SES

iffiii:;,

'.'

.........i.:. ,,,,..

!!..:: . i,~~~~~~~~~~~~~~~~ :, .........................

'

;

t

i;8;ti

S

..:

FIG. 14. Damage to anthers of older flowers of Solanum sessile (Knapp & Mallet 6625: Peru) from pollencollecting bees.

the staticelectricity of the antherbroughtaboutby repeatedbuzzing(Buchmann,pers.comm.).SomeSolanum species have ultra-violetabsorbingtissue in the corolla nearthebaseof theanthercone (Utech& Kawano,1975; Buchmannet al., 1977). The anthersthemselves are stronglyultra-violetabsorbingin S. nigrum(Utech & Kawano, 1975). No members of sect. Geminatahave been tested for ultravioletabsorptionor reflectance. Flowers of species of section Geminata are quite small as solanums go, and thus even very small bees can function as effective pollinators. In a field study conducted in Monteverde, Costa Rica (see Knapp, 1986a, for details of the site andthe study),workersof Bombusephippiatuswere the most important(numerically) floral visitors of the forest species. Workersof thehighlyeusocialstinglessbeeMeliponafasciatawere most importantat the secondarygrowth species. Bagging studiesshowed thatthese bees also effected most of thepollinationin the species studied(Knapp,1986a). Other smaller bees also visited the flowers of these species(see TableIX), butwere not as importantin pollinationandoften only gleanedpollen fromthe corolla lobes (Knapp, 1986a). In a low elevation area in Ven-

ezuela, many small bees were foundbuzzingSolanum sieberi. The lack of large bees in this sample may be due to small samplesize andshortcollectingtime at the flowers. See Table IX for details of bees collected on members of sect. Geminata. In Monteverde,CostaRica, the second growthspecies of section Geminata were visited primarily by Meliponafasciata, a highly eusocial bee that recruits to food resources(Michener,1974;Hubbell& Johnson, 1978; Knapp, 1986a;Roubik, 1980). Much movement of bees is between flowers of the same plant,but since the plants often grow in dense monospecific stands, inter-plantmovement is common as well. In the forest species, on the otherhand,individualBombusworkers appearedto go fromone individualto anotherof thesame species. This was particularly obvious in Solanum pastillum,where visits were common enoughto be assessed. These bumblebees, markedwith dots of paint, werenot observedvisitingothersolanumsor otherflowers in bloom in the area. In the area, many other individuals of Bombus visited the flowers of a species of Hoffmannia(Rubiaceae),andnevervisited the flowers of S. pastillum in thatparticularforagingtrip.Vibratile

28

FLORA NEOTROPICA

Table IX Bees collected on species of section Geminata. MV-Monteverde, Puntarenas,Costa Rica; CH-Parque Nacional Morrocoy, Chichiriviche, Falc6n, Venezuela: all insects in the Cornell University Insect Collection (Lot no. 1146). Determinations by J. Labougle (Bombus), G.C. Eickwort (Halictidae, Anthophoridae), and D. Roubik (Apidae). TAXON

FLOWER VISITOR

Solanum aphyodendron MV

Meliponafasciata costaricensisCockerell(Apidae) MeliponamelanopleuraCockerell(Apidae) Neocorynura(Neocorynura)sp. (Halictidae) LasioglossumkatyaeMcGinley (Halictidae) BombusephippiatusSay (Apidae) Caenaugochlora(Ctenaugochlora)sp. (Halictidae) BombusephippiatusSay (Apidae) Caenaugochlora(Ctenaugochlora)sp. (Halictidae) Meliponafasciata costaricensisCockerell(Apidae) Exomalopsissp. (Anthophoridae) BombusvolucelloidesGribodo(Apidae) BombusephippiatusSay (Apidae) Augochloropsisvesta (Fabr.)(Halictidae) Augochloropsissp. 1 (Halictidae) Augochloropsissp. 2 (Halictidae) Exomalopsissp. (Anthophoridae) BombusephippiatusSay (Apidae)

S. pastillum MV S. pertenue MV S. rovirosan um MV

S. sieberi CH

S. tuerckheimii MV

pollination is a learned technique in eusocial bees (Buchmannet al., 1977), and the behaviorofBombus workerson flowers of sect. Geminatain Monteverde may be an example of the behavioralspecializationof individualworkers(see Heinrich, 1976, 1979; Knapp, 1986a). The primaryforest species visited byBombus occurin rare,widely scatteredpatches,andconstitutea good resource for a bee exhibiting flower constancy (Grant, 1950). Polylectic bees visiting Solanum in a desert site in the southwesternUnited States were remarkablyconstantto those species (Linsley & Cazier, 1963). Flower constancy by tropical bees has been suggested as a mechanismallowing manytropicalbee pollinatedplantsto survive andreproduceat low densities (Janzen, 1971). Foragingbehaviorof individual bees on flowers of low elevation species of sect. Geminatais not known. Bombusis not numerousoutside of the cool cloud forests in the tropical zone, but euglossine bees are common in low elevation tropical forests. Female euglossine bees have been shown to exhibit great flower constancy in lowland Costa Rica (cf. Janzen, 1971), but theirvisits to membersof sect. Geminatahave never been studied.Studiesof pollination in these sparse forest species requirea great deal of time andpatience,since bees do not visit the flowers very often (see Knapp, 1986a). Self-compatibility is widespread in Solanum (see Whalen& Anderson,1981,fora reviewofincompatibil-

ity in the genus). The potatoes and their relatives are exceptions to this rule (de Nettancourt, 1977), as are the wild species of sect. Pachyphylla (Cyphonandra sensu Bohs, 1994: all species now transferred to Solanurm,see Bohs, 1995). The few membersof sect. GeminatathatI have testedhave been self-compatible, butnot autogamous(S. diphyllum,S. aphyodendron,S. pertenue, S. pseudocapsicum,S. spirale). Very few of the species of sect. Geminataas treatedherehave been tested,however,so generalconclusionsarepremature. FRUIT DISPERSAL No rigorous studies of seed or fruit dispersal in Solanumin the Neotropicshave been done, despitethe greatdiversityin fruitsize, color, andgeneralmorphology in the genus (but see Symon, 1979b, for Australian solanums).Berriesof species of sect. Geminataare usuallyrelativelysmall(ca. 1 cm in diameter)andgreen, turningyellow-green when they areripe. Ripe berries are difficult to find, suggesting thatthey ripenand are takenby frugivoresratherquickly.Many species have berrieswhich,whenripe,easily detachfromthepedicel. No speciesthatI have seen have fruitsthatabsciseat the base of the pedicel, releasing the fruitandpedicel as a unit, as occursin some othersections of the genus (see Nee, 1979;Whalen, 1979). If an inflorescencecontains severalberries,theseripensequentiallyandratherslowly. InMonteverde,CostaRica,fruitsof all membersof sect.

29

ECOLOGY AND NATURAL HISTORY

A

B

FIG. 15. Long and short-styled flower morphs in Solanum section Geminata.

Geminataaretakenby both birdsandbats. The berries of S. aphyodendron(as S. nudumin Dinerstein, 1986) are an importantcomponent of the diet of Sturnira ludovici, a small frugivorous bat (Dinerstein, 1983, 1986). These fruits have very high CHO values and aretakenpreferentiallyduringlactationof female bats (Dinerstein,1986). Emeraldtoucanets(Aulacorynchus prasinus) have been reportedas commonly feeding on the fruitsofS. aphyodendron(as S. nudum)andmasked tityras(Tityrasemifasciata)as occasionally feeding on the same species (Wheelwrightet al., 1984). I also observed band-tailedpigeons (Columbafasciata) feeding on the fruitsof this species, which is commonin the second growthin montaneCosta Rica. Seeds of all the forestspeciesof sect. Geminatagrowingat Monteverde were also found in bat feces (Dinerstein, 1983; pers. comm.). Bats use trailsand forest openings as flyways and often defecate in flight. The distributionof these solanumsin light gaps and along trailsmay reflect this behavior.Seeds of several species of sect. Geminata germinateafterpassagethroughbatguts,indicatingthese animals are probably good seed dispersers (but see Wheelwright& Orians, 1982, for a discussion of what constitutesa "good"seed disperser).Some species of sect. Geminata,notably those of the S. arboreumspecies group, have berries smelling strongly of wintergreen when they are ripe (also true in sect. Pteroidea, another forest understory group, see Knapp & Helgason, 1997). Whatanimalrespondsto this smell is not known. Fruitsof these species are often found on thegroundbeneaththe plants,andareperhapsdispersed by small rodents living on the forest floor. The fruits and seeds of the species with brightly colored berries (S. pseudocapsicum species group and S. spirale) are almostcertainlytakenanddispersedby birds.The fruit of S. pseudocapsicum growing in its native habitat

(Paraguayand Argentina)is much dullerin color than thatof cultivatedplants.Muchmorework is needed on both frugivoryand seed dispersal in sect. Geminata. SEX EXPRESSION Derived sexual systems in Solanum have become of interest recently (Anderson, 1979; Anderson & Levine, 1982; Whalen & Costich, 1986; Anderson & Symon, 1989;Zavada& Anderson,1997;Knappet al., 1998), particularlyin light of field studiesandtheoretical work on the evolution of these systems in angiosperms (Bawa, 1980; Bawa & Beach, 1981; Charlesworth, 1984; Renner& Ricklefs, 1995). Many species of Solanumbear only hermaphroditicflowers, all capable of functioningboth in pollen donation and fruit set.Differentformsof flowerswererecognizedby Dunal (1813, 1816, 1852), when he noticedthe differencesin style length in flowers of the same specimen. Many insubgenusLeptostemonum, Solanumspecies,particularly are andromonoecious,bearinghermaphroditicand female-sterileflowers on the same plant(for a classification of sexual systems in angiosperms see Bawa & Beach, 1981). The two floral morphsare easily distinguished: the female-sterile flowers have the style included in the anthercone (Fig. 15B), while in the hermaphroditicflowers,the styleprojectsbeyondtheanther cone (Fig. 15A). The morphsare generallyreferredto as short-styled(SS) flowers and long-styled(LS) flowers. For a review of the literaturedescribingstylarheteromorphisminSolanumsee Whalen& Costich(1986). Varioushypothesesrelatingto the evolutionof these derivedsexualsystemsinSolanumhavebeenadvanced. Symon (1979a) suggested thatsince pollen is the only rewardoffered to pollinatorsby Solanumflowers, the production of excess male flowers should increase pollinatorvisitationratesandthereforeincreaserepro-

30

ductive success. The enhancementof pollination efficiency by productionof excess pollen (i.e., male flowers) has been suggested for the andromonoeciousspecies S. marginatum L. f. (Dulberger et al., 1971). Coleman& Coleman(1982), workingin BrazilwithS. palinacanthumDunal, speculatedthat andromonoecy in thatspeciespromotedoutcrossing.Whalen& Costich (1986) review severalhypothesesconcerningthe evolutionof andromonoecyin Solanumand suggest thatit is unlikely that andromonoecypromotes outcrossing, protects ovaries from predators,or enhances pollen depositionon stigmas. They concludethatthe production of excess staminateflowers aids in pollinatorattraction,and thatthe suppressionof gynoecia in those flowers providesa flexible mechanismfor controlling fruit set. Diggle (1991, 1993) has shown that both phenotypicandgenetic plasticitycontributeto the proportionof female-sterileflowers producedin response to both pollination and fruitset. Selection for increasingcross-fertilizationhas generallybeen consideredthe most importantdrivingforce in the evolution of andromonoecy (Heithaus et al., 1974). Problems with this explanation are many (Primack& Lloyd, 1980;Bawa & Beach, 1981;Bertin, 1982;Renner& Ricklefs, 1995), andit has been shown that in a self-compatible species, andromonoecywill actually always restrict and not enhance outcrossing (Whalen & Costich, 1986). Andromonoecy may be maintainedin Solanumby selection againstthe loss of stamens, as the stamens provide the sole reward for potentialpollinators,and areessential in pollinatorattraction. Bertin (1982) has suggested that the role of pollen as a food rewardpreventsthe evolution ofmonoecism from andromonoecism.No strictly monoecious Solanum species are known. Truedioecy has been reportedin several unrelated sections of Solanum in both Australia and Central America(Anderson, 1979; Anderson& Levine, 1982; Anderson & Symon, 1989; Knapp et al., 1998). The Australian species were originally thought to be androdioecious(Symon, 1970, 1979), with male flowers on one plantandhermaphroditic flowerson another, as the female flowers appearto have functional,pollenfilled anthers.It has been shown however thatthe pollen produced in the anthers of female flowers is inaperturateanddoes not germinate(Anderson, 1979; Anderson & Symon, 1989). Dioecious species in Solanumare an interestingcase: they occur in several unrelatedgroups and appearto have evolved directly from andromonoecious or hermaphroditicancestors withoutpassingthroughthe intermediatestages generally assumed(Bawa, 1980;Bawa & Beach, 1981;Ross, 1982). A model for testing the evolution of dioecy in

FLORA NEOTROPICA

Solanumwas proposedby Knappet al. (1998), and it is clear that Solanum is an ideal group with which to test hypotheses about the evolution of derived sexual systems in angiosperms. Derived sexual systems have very rarelybeen describedfor most membersof sect. Geminata,save for comments contained in the species descriptions of Dunal (1813, 1816, 1851) and Sendtner(1846) (but see Knapp et al., 1998, for a complete descriptionof themorphologyof dioecyinSolanumconfertiseriatum). Most work on this subject in Solanumhas been done in subgenusLeptostemonum. Inthe courseof the preparationof this monographI found that most species of sect. Geminatahave short-styledandlong-styledflowers in the same inflorescences. These species often set few fruit per inflorescence, thus strengthening the circumstantialevidence that they are indeed andromonoecious.I have neverseen short-styledflowerssetting fruitin the field or in the greenhouse.The evidence for andromonoecyin sect. Geminata,though strong, remainsto be tested experimentallywith pollinations in the field or greenhouse in most of the species. The distributionof short-styledflowers on a plant and in an inflorescence in species of sect. Geminatais not as obvious as in many members of subgenus wherethebasalflowersareoftenlongLeptostemonum, styled and the rest are short-styled.In sect. Geminata a few generalizationscan be made. The first inflorescencesproducedby a reproductiveshootareoftencomprised only of short-styledflowers. Subsequentinflorescences will have both floral morphs. Within an inflorescence, short-styledflowers are seemingly randomlyproduced,sometimesat the base andsometimes nearthe tip. Theproductionof short-styledflowersmay be under some environmentalcontrol as it is in other solanums(Wakhloo, 1975a, 1975b, 1975c; Whalen& Costich, 1986) or some genotypic plasticity may be involved(Diggle, 1993;Knappet al., 1998). Species of sect. Geminatagrowingin the Bailey Hortoriumgreenhouses seem to have higherproportionsof short-styled flowersthanplantsof the same species I have collected in the field. Whetherthis distributionis age-relatedor environmentallyinducedis not known. In the Solanum sessile species group, the entire range of sex expression is found: from plants with all flowers(S.turgidum)to dioeciousplants hermaphroditic (S. confertiseriatum) (Knapp,1991a). InS. rovirosanum in Costa Rica, young plantsproduceonly short-styled flowers. Largerolder plantsbearonly hermaphroditic flowers,with an occasionalshort-styledfloweron some inflorescences. Solanum sessile in eastern Peru and Ecuadorbears both long and short-styled flowers on the same inflorescence, and appears to be andro-

ECOLOGY AND NATURAL HISTORY

31

~' $rWY'4i~:~,? ',' .........

:'LI~i(~i!'~ 1~ I; ""'"' ......I":'"'"' ..... *..'.'....X

_

S 11 S

l

l

.'i ..

:il.33'i'i::i.......

S|-|S>>a<..

..............ffi

ra

...........:,.

FIG. 16. Maecolapsis sp. (Coleoptera: Eumolpinae) on leaves of Solanum pertenue (Monteverde, Costa Rica).

Table X Chryosomelid beetles collected feeding on species of section Geminata. All insects are in Cornell University Insect Collection (Lot number 1146), and were determined by C.R. Hoebeke. Parts fed on by the beetle: L-leaves, F-flowers, B-berries. HOSTPLANT

SPECIES

Solanum arboreum S. pastillum

Coptocycla cf annulus Fabr. (Cassidinae) Diabrotica sp. (Galerucinae) genus nr. Labidomera (Chrysomelinae) Aspidomorpha sp. (Cassidinae) Maecolapsis sp. (Eumolpinae) unknown genus (Eumolpinae) Maecolapsis sp. (Eumolpinae) Diabrotica sp. (Galerucinae) Aspidomorpha sp. (Cassidinae)

S. pertenue

S. ramonense S. rovirosanum

monoecious.In otherspecies groupsin the section,both long- and short-styledflower morphsoccur.

STAGE

adult (L) adult (F) adult (F) adult, larvae (L) adult (L) adult (L) adult (L) adult (F, B) adult (L).

1983; Hsiao, 1978, 1982, 1985; Hsiao & Fraenkel, 1968a, 1968b, 1968c), and ithomiine butterflies (Nymphalidae: Ithomiinae:Gilbert, 1969; Brown & HERBIVORES d'Almeida, 1970; Young, 1972, 1973, 1974a, 1974b, Theinteractionof speciesof the Solanaceaeandtheir 1974c, 1977, 1978, 1979; Rathke & Poole, 1975; host-specificherbivoresis of interestas a possible case Drummond,1976,1981; Brown, 1985; Drummond& of insect/plant biochemical coevolution (Ehrlich & Brown, 1987; Haber, 1979; Beccaloni, 1995). Much Raven, 1964; Gilbert, 1975). The best studiedof these work has also been done with the insect trappingglanherbivoresarechrysomelidbeetles (includingthe Colo- dulartrichomesof potatospecies (Gibson, 1974, 1976a, radopotatobeetleLeptinotarsadecemlineataSay:Hare, 1976b, 1978, 1979; Gibson & Turner,1977; Gibson

32

& Valencia, 1978; Tingey & Gibson, 1978), but will not be discussedhere,as no membersof sect. Geminata possess these types of trichomes. Membersof the beetle family Chrysomelidaefeed on the leaves of Solanumspecies both as larvaeand as adults(Hsiao, 1985;pers.observ.).Most workhas been done with Leptinotarsa decemlineata (the Colorado potatobeetle) and its relatives (see referencesabove), but little is knownaboutthe feeding habitatsof neotropical solanum-feedingchrysomelids.I have observed several species of chrysomelidbeetles feeding both as adults and as larvae on various species of sect. Geminata(see TableX). Membersof the subfamilies Cassidinae and Eumolpinae(Fig. 16) are often found on the leaves ofsolanaceous plants,but directevidence for feeding is difficult to obtain.These beetles may do most of theirfeeding at night, when most botanistsare inactive.Onlyrecordsof beetles actuallyfoundfeeding on Solanumspecies are includedin TableX. Adults of Diabrotica sp. were observed burrowinginto the unripe and ripe fruits of S. rovirosanumin Costa Rica, and were feeding on the seeds. The fruitsof S. rovirosanum exude a sugary solution which attractswasps and bees. This might help to deter seed predatorslike Diabrotica, but seemed to be completely ineffective in this species in Monteverde,Costa Rica (pers. obs.). Ithomiinebutterflies(Nymphalidae:Ithomiinae)are mimetic, warningly colored, commonly encountered membersof the Neotropical fauna(Fig. 17). Theirbiology has been the subject of fascination since Bates (1862) collected in the Amazon and subsequentlydescribedtheirstrikinganddiverse color patterns.These butterfliesaredistastefulto birds,andwere long thought to have acquiredthis distastefulnessas larvaefromtheir solanaceous food plants (Brower & Brower, 1964; Ehrlich & Raven, 1964; Drummond, 1976, 1981). Brown (1986) has recently shown however that the defensive compounds found in the adults are not derived from Solanaceae at all. Instead, they are dihydropyrollizidinemonoesters, obtained by adults fromvarioussources,particularlyflower nectar(Senecio spp., Asteraceae;Tournefortiaspp., Boraginaceae). These compoundsare also used by males to attractfemales, using specialized scent scales on the hindwings (Haber,1979). Species of sect. Geminataare fed upon by what are considered to be advanced groups of ithomiines(Mielke & Brown, 1979;Brown, 1986):the DircenniiniandGodyridini.Many of the generareared frommembersof sect. Geminatahave also been reared from species of Cestrum,perhapsdue to their overall vegetative similarity (entire simple leaves, forest understorytrees and shrubs).Rearingdataarepresented in TableXI. Literaturedatafor otherhost specific lepidopterousherbivoresis presentedin TableXII.

FLORA NEOTROPICA

With so few records, broad general conclusions cannotbe drawnwith any confidence, but some interestingpatternsareemerging.Ceratiniatutia(Haensch) larvae(Fig. 18) appearto feed exclusively on members of the Solanumnudumspecies group.Membersof the S. confine species group are fed upon largely by species of Pteronymia.Drummond& Brown (1987) proposed a new tribe of ithomiines including the genera Ceratinia, Ceratiscada, Prittwitzia, Episcada and Pteronymia, all of which feed on members of sect. Geminata.Of the rest of the tribe, only Godyris and Mcclungia of the Godyriniinifeed on species of sect. Geminata(see Drummond& Brown, 1987). The generic phylogeny of the ithomiine butterflies is being actively investigated(Beccaloni, pers. comm.), but at presentis not at all well known.The datapresentedhere arebutthe tip of an iceberg.Muchmore informationis needed beforewe drawconclusions aboutcoevolution or possible phylogeneticpatternsconcernedwith host relationshipsin these groups.

SPECIES GROUPS AND SPECIES CONCEPTS SPECIES GROUPS I have dividedthe 126 species of section Geminata into 16 species groupsusing largelythe following morphologicalcriteria:1) generalizedsympodialstructure, 2) leaf shape,3) trichomemorphology,4) inflorescence size and type, 5) spacing of pedicel scars within the inflorescence,6) fruitingpedicel orientationandstructure,and7) seedmorphology.I havebeenunableto place threespecies (Solanumdelitescens,S. lasiocladumand S. mapiricum)in any species group, due to their aberrantmorphologyor to the small numberof collections available to me for study.A key to the species groups in the taxonomictreatment,andbrief characterizations of the species groups,areprovided(see also TableXII). The threeisolatedspecies arealso described.The order of the groupsis not to be interpretedas a phylogenetic scheme.The speciesof sect. Geminataareall very similarto theuninitiatedobserver,so thebestway to approach the keys is to first read the descriptionsof the groups to get an idea of the variationpresentin each one, then attemptto use the key to identify the plant in hand. SPECIES CONCEPTS My views on delimitationof species basically follow what is knownas the "morphologicalcluster"species concept(Mallet, 1995), i.e., "assemblagesof individuals with morphological features in common and separate from other such assemblages by correlated morphologicaldiscontinuitiesin a numberof features"

SPECIES GROUPS AND SPECIES CONCEPTS

C

33

J7D

F

E ,

1

FIG. 17. Butterflies of the subfamily Ithomiinae (Lepidoptera: Nymphalidae). All specimens held in the entomological collections of The Natural History Museum, London. A. Pteronymia artenna. B. Ceratinia tutia. C. Episcada salvinia. D. Mcclungia galita. E. Scada karschina. F. Ceratiscada canaria. G. Godyris zavelata caesiopicta.

FLORA NEOTROPICA

34

Table XI Records of ithomiine(Lepidoptera:Nymphalidae:Ithomiinae)butterfliesfeeding on species of section Geminata(butterfly identificationsby GL- GerardoLamasM., JM-JamesMallet,PV- PhilipJ. deVries,FFY- FranciscoFemandez-Yepez).Butterfly specimensare in the privatecollectionof JamesMallet or in the collectionscited in the references.No unconfirmedrecords,or recordsforwhichI havenotseena plantvoucherspecimen,areincluded.Butterfliesrearedoffunidentifiedorotherwiseunconfirmed speciesof sectionGeminataaregroupedatthe endof the tableas Solanumspp.Knappcollectionsaredesignatedby K numberafter the butterflyname.Plantspecimensare depositedin the herbariacited in the text. HOSTPLANT

LOCALITY

BUTTERFLY

Solanum aphyodendron

Costa Rica. Monteverde Costa Rica. Monteverde Costa Rica. San Vito Costa Rica.

Episcada salvinia (Bates) (Haber, 1979)

Costa Rica. Ecuador. Tandapi S. arboreum

Costa Rica. La Selva

S. caavurana

Brazil. Espirito Santo Brazil. Rio de Janeiro Brazil. Sao Paulo Brazil. Sao Paulo Brazil. Sao Paulo

S. campaniforme

Venezuela. Kavanayen Brazil. Espirito Santo Brazil. Espirito Santo Brazil. Brazil. Brazil.

S. leptorhachis

Ecuador. Tinalandia

S. lucens

Venezuela. Tachira

S. nudum

Costa Rica. Turrialba Costa Rica. Corcovado Costa Rica. Costa Rica. Costa Rica.

Pteronymia artenna (Hew.) (K 834: PV) Episcada salvinia (Bates) (Haber, 1979) Pteronymia cotytto (Drummond& Brown, 1987) Episcada salvinia opleri (Drummond & Brown, 1987) Episcada cf. apia (Felder) (K 6272: GL) Scada zibia (Hew.) (Haber, 1979; in Drummond& Brown, 1987, as Scada zibia xanthina (Bates)) Ceratiscada canaria (Brown & d'Almeida) (Brown & d'Almeida, 1970) Episcada clausina striposis Haensch (Drummond& Brown, 1987; Brown, 1992) Pteronymia carlia Schaus (Drummond& Brown, 1987) Pteronymia sylvo (Drummond& Brown, 1987) Prittwitzia hymenaea hymenaea (Prittwitz) (Drummond& Brown, 1987; Brown, 1992) Mcclungia galita (Hew.) (K 6718: JM) Pteronymia euritea (Cramer) (Drummond& Brown, 1987) Ceratiscada canaria (Brown & d'Almeida) (Drummond& Brown, 1987; Brown & Freitas, 1994) Prittwitzia hymenaea hymenaea (Prittwitz) (Brown, 1992) Episcada carcinia (Brown, 1992) Episcada philoclea (Brown, 1992) Pteronymia glauca (Haensch) (K 6260: GL) Pteronvmia latilla (Hew.) (K 6830: JM) Ceratinia tutia (Hew.) (Haber 1979) Ceratinia tutia (Hew.) (K 887: JM) Ceratinia tutia dorilla (Bates) (Drummond& Brown, 1987) Pteronymiafulvimargo (Butl. & Druce) (Drummond& Brown, 1987) Episcada salvinia opleri (Drummond& Brown, 1987)

35

SPECIES GROUPS AND SPECIES CONCEPTS HOSTPLANT

LOCALITY

BUTTERFLY

S. nudum (cont'd.)

Ecuador. Limoncocha

Ceratiniapoeciliapoecilia (Bates) (Drummond,1976) Ceratinia tutia radiosa (Haensch) (K 6188: GL) Ceratinia tutia poecilia (Beccaloni, 1995) Ceratinia tutia (Hew.) (K 6489: JM) Ceratinia tutia (Hew.) new ssp. (K7707: GL) Ceratinia tutia (Hew.) (K 6772: JM) Pteronymia artenna (Hew.) (K 830: PV) Pteronymia artenna (Hew.) (K 828: PV) Prittwitzia hymenaea hymenaea (Prittwitz) (Drummond& Brown, 1987) Pteronymia carlia Schaus (Drummond & Brown, 1987) Pteronymia sylvo (Drummond& Brown, 1987, fide Beccaloni, pers. comm. =?carlia) Pteronymia hemixanthe (Feld. & Feld.) (Drummond& Brown, 1987) Pteronymia euritea (Drummond& Brown, 1987) Episcadaphiloclea (Brown & Freitas, 1994) Pteronymia sylvo (Brown & Freitas, 1994) Episcada polita striposis (Brown & Freitas, 1994) Scada karschina karschina (Freitas pers. comm. to Beccaloni) Pteronymia latilla Hewitson (Drummond& Brown, 1987) Godyris kedema kedema (Hew.) (Drummond & Brown, 1987, as nr. ripense) Hypomenitis andromica andromica (Drummond& Brown, 1987, as nr. ripense) Oleria denuda ssp. (Riley) (K 6370: GL) Pteronymia notilla (Butler & Druce) (Gilbert, 1969) Pteronymia agalla (Godman & Salvin) (Gilbert, 1969) Godyris zavaleta caesiopicta (Niepelt) (Drummond& Brown, 1987, as G. gonussa caesiopicta) Hypothyris lycaste linosa (Sourakov, 1995) Pteronymia simplex simplex (Drummond & Brown, 1987) Pteronymia notilla notilla (Drummond& Brown, 1987) Episcada sylpha (Haensch) (K 6700: FFY) Pteronymia alida (Hew.) (K 6859: JM)

Ecuador. Rio Latas Ecuador. Jatun Sacha Peru. Tarapoto Peru. Sarameriza Venezuela. Manacal S. pastillum S. pertenue S. pseudocapsicum

Costa Rica. Monteverde Costa Rica. Monteverde Brazil. Sao Paulo Brazil. Sao Paulo Brazil. Sao Paulo

S. pseudoquina

Brazil. Espirito Santo Brazil. Espirito Santo Brazil. Minas Gerais Brazil. Sao Paulo Brazil. Sao Paulo Brazil.

S. ripense

Venezuela. Venezuela. Venezuela.

S. robustifrons

Peru. Quincemil

S. rovirosanum

Costa Rica. San Vito Costa Rica. Osa Costa Rica. Costa Rica. Costa Rica. Monteverde Costa Rica.

S. sieberi

Venezuela. Morrocoy

S. tenuflagellatum

Venezuela. Col. Tovar

FLORA NEOTROPICA

36 HOSTPLANT

LOCALITY

BUTTERFLY

S. triste

Venezuela. Cumanacoa

Solanum spp. (sectionGeminata)

Costa Rica.

Pteronymia latilla (Hew.) (K 6750: JM) Pteronymia simplex simplex (Haberin Drummond& Brown,1987,on variousunidentified species) Pteronymia notilla notilla (Drummond& Brown, 1987, on various unidentifiedspecies) Hypoleria cassotis (Bates) (Young in Drummond& Brown, 1987) Episcada salvinia (Drummond & Brown, 1987) Ceratinia tutia tutia (Drummond & Brown, 1987) Pteronymia artena beebei (Drummond & Brown, 1987) Ceratiscada hymen (Drummond& Brown, 1987) Episcada carcinia (Brown, 1992) Episcada clausina striposis (Brown, 1992) Godyriszavaleta (Drummond& Brown, 1987) Prittwitzia hymenaeahymenaea (Prittwitz) (Drummond& Brown, 1987) Pteronymia carlia Schaus (Drummond& Brown, 1987)

Costa Rica. Costa Rica. Colombia. Meta Venezuela. Venezuela. Ecuador. Limoncocha Brazil. Sao Paulo. Japi Brazil. Sao Paulo. Japi Brazil. Acre Brazil. Rio de Janeiro Brazil. Sao Paulo. Japi

Table XII

Records of other Lepidopterafeeding on leaves of members of section Geminata(from databaseconstructed and maintained by Dr. G. Beccaloni, Department of Entomology, BM (NH)) HOSTPLANT

LOCALITY

BUTTERFLY

S. pseudocapsicum

Argentina.

Lycaenidae Arawacus ellida (Hewistson) (Robbins, pers. comm. to Beccaloni)

Brazil.

Lycaenidae Rekoa palegon (Cramer) (Robbins, pers. comm. to Beccaloni)

(Davis & Heywood 1963). Biological (Mayr, 1982), phylogenetic (Cracraft, 1981), and the host of other finely defined species concepts (see Mallet, 1995) are almost impossible to apply in practice when dealing with large, complex groups such as sect. Geminata, andarethereforeof littleutility in a practicalsense. It is importanthowever to clearly state the criteriafor the delimitationof species, ratherthandogmaticallyfollow particularideological lines (see Luckow, 1995; Davis, 1997). In this revision I have triedto emphasize similarities between populations instead of differences, which so often reflect incomplete collecting or local variation. I have not recognized subspecies or variet-

ies, as I do not feel these are useful categories, in either a taxonomic or evolutionary sense. The variation is better described and documented, ratherthan formalizedwith a name which then encumbersthe literature.I have been conservativein my approach,recognizing as distinct entities those populationsystems (sets of specimens) that differ in several morphological characteristics.Minor differences in morphology, distribution, habitat, and ecology are important in some groups such as the Solanum confine species group,where the common groundplanfor the species is very similar. On the other hand, I have recognized several extremely widespread,polymorphic species,

SPECIES GROUPS AND SPECIES CONCEPTS

37

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FLORA NEOTROPICA

38

e.g., S. nudum, S. leucocarpon, S. sessile. In these species variation exists in certain characters,but the patternof variation is such that no reliable units can be extracted.In these cases the variability within and between populations seems more importantthan the variations of the extremes others have recognized as distinct. In these cases I have describedthe variation, realizing thatothertaxonomistsmay wish to interpret it differently.

TAXONOMIC TREATMENT

species: SolanumnudumDunal (D'Arcy, 1972). Pheliandra Werderm.,Notizbl. Bot. Gard.BerlinDahlem 15: 54. 1940. Type species:Pheliandra herthaeWerderm.A. T. Hunziker,Kurtziana4: 136. 1967. Solanum subsect. TriplinerviumC. V. Morton, Contr. U. S. Natl. Herb. 29: 48. 1947. Type species: Solanum triplinervium C. V. Morton. Solanum sect. Oppositifolium (Dunal) Seithe, Bot.

Jahrb.Syst. 81: 289. 1962. Pro parte,Lectotype species: SolanumnudumDunal (Seithe, 1962). Solanum subsect. Oppositifolium (Dunal) Seithe,

Bot. Jahrb.Syst. 81: 289. 1962. Lectotypespecies: SolanumnudumDunal (Seithe, 1962).

Solanumsect. Geminata(G. Don) Walp.,Repert.Bot. Syst. 3: 58. 1844. Solanum subsect. GeminataG. Treesor shrubs,occasionallyrhizomatous,1-20 m Don, Gen. syst. 4: 418. 1838. Lectotype species: tall; pubescence of papillate or uniseriate simple or Solanum nudum Dunal (D'Arcy 1972). Danert, branched trichomes, the trichomes occasionally Kulturpflanze18: 280. 1970; D'Arcy, Ann. Mis- multiseriateat the base or echinoid. Sympodial units souriBot. Gard.59: 268,274. 1972; Knapp,Taxon most often difoliate, in some species unifoliate or 32:636. 1983. plurifoliate.Leaves geminateor not, linearto obovate, PseudocapsicaMoench,Methodus476. 1794. Type variablein size withina species,usuallyglabrousabove, species: Pseudocapsica undatifoliumMoench glabrous or pubescent with uniseriate trichomes be(=Solanum pseudocapsicum L: D'Arcy, 1972). neath,the trichomessimple or branched,in a few speSolanum subgenus Minon Raf., Autik. Bot. 108. 1840. cies echinoid; minor leaves, if present,either equal in Lectotype species: Solanum pseudocapsicum L. size and shape to the major ones or strongly aniso(D'Arcy, 1972). Solanum grad. ambig. Leiodendra Dunal, Solan. phyllous. Inflorescences usually opposite the leaves, Syn. 20. 1816; Dunal in DC., Prodr.13(1): 29, occasionally somewhatintemodalor pseudoterminal, 137. 1852. Lectotype species: Solanumnudum filiform to stout, simple to many-times furcate,papilDunal (D'Arcy, 1972). late or pubescentwith usuallyuniseriatetrichomeslike Solanumgrad. ambig.Pseudocapsica(Moench)G. those of the leaves, the trichomessimple or branched, Don, Gen. Syst. 4: 410. 1838. Lectotypespecies: usually small;pedicels at anthesis white or greenishSolanum pseudocapsicum L. (D'Arcy, 1972). white, usually deflexed; buds globose to elliptic, variSolanum subsect. Micranthes Dunal, in DC., Prodr. ously pubescent;calyx tube urceolateto broadlyconi13(1); 28, 95. 1852. Pro parte, Lectotype species: Solanum micranthum Willd. ex Roemer & cal, the lobes minutely deltoid or long-triangularand Schultes (D'Arcy, 1972). D'Arcy, Ann. Missouri acuminate;corolla white, greenish-white,occasionally Bot. Gard. 59: 271, 274. 1972. pink or tinged with purple, 0.4-3.5 cm in diameter, Solanumgrad. ambig.AnthopleurisDunal in DC., fleshy to membranous,lobed from 1/2 of the way to Prodr. 13(1): 29, 123. 1852. Pro parte, Lectotype the base to nearly to the base, the lobes campanulate, species: Solanum nudum Dunal (D'Arcy, 1972). planar,or deflexed at anthesis,the tips and marginsof Solanum grad. ambig. Oppositifolium Dunal in DC., the lobes usually densely papillate;anthers elliptic to Prodr. 13(1): 29, 123. 1852. Pro parte, Lectotype or orangishyellow, not terminallyattenuovoid, yellow species: Solanum nudum Dunal (Seithe, 1962). all anthers poricidalat the tips, the poresbecoming Solanumsect. Leiodendron (Dunal)Bitter,Bot. Jahrb. ate, slit-like upondryingandteardropshapedin all buttwo Syst. 54: 500. 1917; D'Arcy, Ann. Missouri Bot. Gard. 59: 274. 1972 (as Leiodendra). Lectotype species with dimorphicanthers(Solanumpseudoquina species: Solanum nudum Dunal (D'Arcy, 1972).

Solanumsect. Pseudocapsicum (Moench)Bitter,Bot.

and S. reitzii); free portion of thefilaments more or less

equalto the filamenttube;ovaryglabrousor pubescent with papillate or uniseriate trichomes;style straight, Solanumpseudocapsicumr L. (Bitter, 1917). glabrous or minutely puberulentat the base, usually Solanum subsect. Silicosolanum Bitter, Repert. Spec. to the length of the corolla lobes in long-styled Nov. Regni Veg. 16: 10. 1919. Type species: equal includedin theanthercone in short-styledflowflowers, Solanum trachytrichiumBitter. Seithe, Bot. Jahrb. ers; stigma capitate or more often restricted to the Syst. 81: 289. 1962; D'Arcy, Ann. Missouri Bot. unexpandedtip of the style. Berries globose or someGard. 59: 273, 274. 1972. Solanum sect. Anthopleuris (Dunal) Bitter, Repert. what ellipsoid and apically pointed, bilocular,usually Spec. Nov. Regni Veg. 16: 10. 1920. Lectotype green or yellowish green at maturity,in some species Jahrb. Syst. 54: 497. 1917. Lectotype species:

39

TAXONOMIC TREATMENT

(i.e., S. pseudocapsicum species group) brightly colored orange or reddish;fruiting pedicels erect to deflexed andgreatlyelongate,usuallythickerat the apex; seeds flattened-reniformwith incrassatemargins and lignified lateralseed coat walls or ovoid-reniformwith non-thickenedlateralseed coat walls, the surfacesminutely pitted. Chromosomenumber:n = 12 (in all but S. spirale from the Old Worldtropics, where n = 24). The section is treatedhere in the broad sense, includingall specieswith simpleandbranchedtrichomes. Furtherdiscussion relating to the utility of trichome charactersin the taxonomyof this groupwill be found in Morphology. Species are here keyed to species groups and within each group to species. As many of the species of sect. Geminataareremarkablysimilarin morphology,I have provideda synoptic, multi-access key to all the taxa.The synoptickey will be most useful for those specimensin only flower or fruit,andvegetative material should be identifiable to broad groups using the synoptic key. I have attemptedto correctlytypify all the species treatedin thismonograph.I havestrictlyfollowedguidelines laid down in theInternationalCodefor Botanical Nomenclature(Greuteret al., 1994) for the interpretationof types. I have only regardeda specimenas a holotype if it was specifically and unambiguouslydesignated as such by the original author, or if a single specimenfroma singleherbariumwas citedin the originaldescription.If a single specimenwas not designated, I have designated a lectotype from among the extant duplicates or syntypes. If a lectotype was previously designated,I have cited its place of publication.Notes on lectotypifications can be found in the individual

species accounts. Both M.-F. Dunal at MPU and later J. F. MacBride at F received small fragmentsof type specimens for inclusion in theirrespective herbaria.I have cited these fragmentsamong the isotypes where they occur and have indicated them as fragmentary (frag.).Wherethe fragmentis morethanjust a leaf and a flower and could be classed as an entire specimen, it is cited as an isotype. Species groups are listed in an arbitraryorder,but those groups with flattened-reniform seeds (the plesiomorphicstate in Solanum)are groups I through VI, and those with ovoid-reniformseeds (the derived state)areVII throughXVI (see NumericalListof Taxa). Within each species groupthe species are listed in alphabeticalorder.Species are numberedconsecutively throughoutthe monograph.All specimens cited in the monographhave been seen by the authorexcept when designated "n.v.,"including types. Not all specimens examinedduringthe courseof thistreatment(morethan 7000 collectionsof approximately2000 collectors)have been citedin the text, butall arelistedin the Exsiccatae. For widespreadspecies particularly,only representative specimens are cited in the species treatments.For rareror less well-known taxa, however, all specimens examinedarecitedin full. Severalof the species of sect. Geminataarewidely cultivatedin both the tropicsand subtropics and I have not cited obviously cultivated specimens, unless they were the only record from a particularneotropicalcountry.Lists aregiven of cultivated ranges for those commonly cultivated taxa (S. diphyllum,S. pseudocapsicum). Maps were preparedusing the Flora Neotropica Base MapNo. 1, where each symbolin a degreesquare indicatesthatthe species occurs in thatdegree square.

Artificial key to woody, similar groups of non-spiny solanums 1. Inflorescence borne in branch axils or laterally; sympodial units on reproductive branches with 3-4 leaves or plurifoliate; anthers often with enlarged or otherwise modified anther connective, tightly connivent at anthesis. 2. Anther connective thickened and prominent abaxially, sharply differentiated from the thecae; sympodial units 3-4-leaves .................. Solanum sect. Pachyphylla (Cyphomandra), (see Bohs, 1994; Appendix III for species 2. Anther connective not expanded abaxially, continuous with the thecae; sympodial units usually multi-leaved. 3. Anthers of equal size..... Solanum sect. Cyphomandropsis (see Bohs, 1994; Appendix IV for species ........................................ Solanum sect. Allophyllum (see Bohs, 1990; Appendix V for species 4. Anthers with the pores lengthening into slits, oblong; plants glabrous or with scattered lepidote scales ............... Solanum acropterum species group (see Excluded Species for species 4. Anthers with the pores not lengthening into slits, obovoid; plants glabrous, or the new growth papillate ............. Solanum havanense species group (see Excluded Species for species 3. Anthers of different sizes, two long, three short ........................................ Solanum thelopodium species group (see Excluded Species for species 1. Inflorescence terminal, lateral or leaf-opposed; sympodial units various; anthers not with a conspicuous thickening of the connective.

list)

list) list) list) list) list)

FLORA NEOTROPICA

40

5. Pedicels inserted on the axis in "sleeves" or platforms, not flush with the surface; sympodial units always plurifoliate; branching sometimes dichasial. 6. Pedicels inserted in "sleeves" ................................... Solanurmnitidum species group (see Knapp, 1989; Appendix VI for species list) 6. Pedicels inserted on platforms in congested groups, inflorescences terminal .................................................... Solanum sect. Holophylla s.str. (see Excluded Species for species list) 5. Pedicels flush with the inflorescence axis, not leaving a sleeve or platform; sympodial units only rarely plurifoliate; branching never dichasial, always monochasial. 7. Pubescence of simple or dendritic trichomes, or the plants glabrous . s.l. (this monograph) ............................................................................................... Solanum sect Geminata 7. Pubescence of stellate or lepidote trichomes (these sometimes scale-like), if trichomes simple, derived from the central midpoint of a stellate hair (i.e., single-celled). 8. Pubescence of stellate trichomes (in a single undescribed species simple as above); calyx lobes enlarged and petaloid ........... Solanum sect. Extensum (see Excluded Species for species list) 8. Pubescence of lepidote scales or other chaffy trichomes; calyx lobes not petaloid ...................................... Solanum sect. Lepidotum (see Carvalho, 1996; Appendix II for species list) ........................................ Solanum sect. Cernuum (see Carvalho, 1996; Appendix I for species list) Solanum punctulatum of the S. acropterum species group (see above) ........................................

Artificial key to the species groups 1. Sympodial units unifoliate (one leaf opposite each inflorescence); leaves not geminate. 2. Flowers large, greater than 1 cm in diameter, the lobes planar at anthesis. 3. Leaves glabrous or with simple, uniseriate trichomes ......................................... Group XI (Solanumnarcoticosmum species group, S. foetens: p. 251) 3. Leaves densely pubescent with dendritic or echinoid trichomes ...................................................................... Group V (Solanum nutans species group, S. nutans: p. 157) 2. Flowers smaller, less than 1 cm in diameter, the lobes reflexed at anthesis. 4. Pedicel scars closely and evenly spaced; inflorescence stout, not filiform; fruiting pedicels erect, usually woody ........................................ GroupIX (Solanumunifoliatum species group: p. 223) 4. Pedicel scars widely and irregularly spaced; inflorescence usually filiform (if not, greatly reduced and few-flowered); fruiting pedicels deflexed, greatly expanded at the apex. 5. Leaf margins undulate, occasionally erose, the leaf apices not ciliate ............................................................................... GroupXV (Solanumconfine species group: p. 311) 5. Leaf margins not erose, plane, the leaf apices ciliate, often minutely so .......................................................... Group XVI (Solanum dolosum species group: p. 350) 1. Sympodial units di- or plurifoliate; leaves often geminate. 6. Seeds flattened-reniform, with incrassate margins, usually many (more than 50) per fruit; lateral seed coat walls thickened and suberized. 7. Berries brightly colored when mature, usually red or orange. 8. Trichomes simple; calyx lobes deltoid; fruiting pedicel deflexed. Old World tropics .................................................... Group III (Solanum nudum species group: S. spirale: p. 122) 8. Trichomes branched (occasionally simple or some cultivated plants glabrous); calyx lobes long-triangular; fruiting pedicels erect. New World tropics ..................................................................... Group II (Solanum pseudocapsicum species group: p. 54) 7. Berries green or yellowish green when mature, not brightly colored orange or red. 9. Anthers tightly connivent, often bright orange; fruiting pedicel with a distal swelling ........................................................................ Group IV (Solanum leucocarpon species group: p. 136) 9. Anthers not orange, not tightly connivent; fruiting pedicels without a conspicuous swelling. 10. Inflorescences lateral, large and with complex branching; flowers fleshy or waxy. Plants of high elevations. 11. Trichomes of leaves dendritic; leaves to 30 cm, repand ............................................................. Group I (Solanum oblongifolium species group: p. 43) 11. Trichomes of leaves simple, only occasionally dendritic, or the plant glabrous; leaves not exceptionally large, less than 30 cm, or repand ............................................ Group VI (Solanum amblophyllum species group: p. 165)

TAXONOMIC TREATMENT

41

10. Inflorescences opposite the leaves or occasionally (only rarely) intemodal, simple, if branched only furcate; flowers various. 12. Leaf and stem trichomes all complex and branched, often echinoid ...................................................... Group V (Solanum nutans species group: p. 152) 12. Leaf trichomes simple, a few occasionally branched. 13. Trichomes abundant on the leaf undersides, often covering the surface. Usually plants of high elevations........... Group VI (Solanum amblophyllum species group: p. 165) 13. Trichomes usually in tufts in the vein axils, not on the lamina. Plants of low to middle elevations .................................. Group III (Solanumnudum species group: p. 69) 6. Seeds ovoid-reniform, usually few (less than 50) per fruit, the lateral seed coat walls not thickened. 14. Sympodial units plurifoliate; leaves not geminate. 15. Flowers large, 1-1.5 cm in diameter; the corolla lobes planar at anthesis ........................................ GroupXIV (Solanumsessile species group: S. monadelphumn:p. 287) 15. Flowers smaller, 0.8-1 cm in diameter; the corolla lobes reflexed at anthesis ........................................ GroupVIII (Solanumarboreum species group: S. amnicola: p. 197) 14. Sympodial units difoliate; leaves usually geminate. 16. Young stems and leaves glabrous; leaves drying black; fruiting calyx accrescent. Mountains of Central America ................................. GroupXI (Solanumnarcoticosmum species group: p. 247) 16. Young stems and leaves densely papillate, the trichomes usually reddish or gray in color in dry material or golden pubescent; leaves not drying black. Various habitats in Central and South America. 17. Mature leaves glabrous; trichomes on the stem, if present, uniseriate. 18. Leaves with strong parallel primary venation; new growth densely red papillose; berries usually papillose ...................... GroupX (Solanumrobustifrons species group: p. 235) 18. Leaves with venation not strongly parallel; new growth red papillose or appearing glabrous; berries glabrous or pubescent, not papillose ........................................................... GroupVIII (Solanumarboreum species group: p. 194) 17. Mature leaves pubescent; trichomes simple or branched. 19. Leaf and stem trichomes branched, arachnoid, or dendritic. 20. Trichomesarachnoid,fine structureobscure (occasionallyvery sparse on older leaves), the new growth felty pubescent.............Group XII (Solanumnigricans species group:p. 254) 20. Trichomes dendritic. 21. Inflorescences opposite the leaves, simple, few-flowered; leaves 7-12 cm long, elliptic; trichomes sometimes very sparse in vein axils ( Solanum smithii in S Ecuador)................................... Group XIII (S. arenarium species group: p. 273) 21. Inflorescencesappearingterminalor lateral, branched,many-flowered;leaves 10-30cm long, elliptic or more commonly obovate;trichomeson the veins and lamina......................................................... Group XIV (Solanum sessile species group: S. chlamydogynum, S. confertiseriatum: pp. 283, 285) 19. Leaf and stem trichomes simple or at most furcate. 22. Fruiting pedicels longer than at anthesis, expanded at the apex, strongly deflexed; leaves membranous, with undulate (ruffled) margins............................................. GroupXV (Solanumconfine species group: p. 311) 22. Fruiting pedicels not greatly expanded at the apex nor longer than at anthesis; leaves not as above. 23. Inflorescence large and many times furcate (occasionally simple); buds fleshy; pubescence of minute golden uniseriate trichomes; berries often pubescent; pedicel scars usually closely spaced; corolla lobes planar at anthesis ........................................ GroupXIV (Solanumsessile species group: p. 282) 23. Inflorescence not as above, if furcate, then slender; buds small, not fleshy, long-elliptic; pubescence, if present, of long, uniseriate trichomes; berries glabrous; pedicel scars ca. 1 mm or more apart; corolla lobes reflexed at anthesis .............................. GroupVII (Solanumdeflexiflorum species group: p. 179)

FLORA NEOTROPICA

42

Synoptic key to the species This synopsis is intended to simplify the task of identifying a member of this large and at first glance veryuniformgroup.On anygiven specimenof Solanum sect. Geminata,flowers or fruits may not be present, thus making the use of the dichotomouskey difficult. This list includes characterswhich are distinctive, sometimes at the species grouplevel, or othertimes at the species level. I have includedmany leaf andwhole plant charactersso that sterile plants can be in many cases identifiedto a choice of a few taxa.Once this step has been done, the user can read the descriptionsand matchdistributionsandothercharactersnot used in this list to the plant in hand. In the list the characterstates are followed by the numberscorrespondingto species in the taxonomictreatment.Numbersin parenthesesindicatethatthe stateis relativelyuncommonin thatspecies. Species thatvary in a particularcharacter,for example in leaf trichome distribution,will be listed for each alternativepresentin thatspecies. Not all alterative charactersarelisted:for example,I have listed fruit color otherthangreen,but not fruitsgreen. In this way diagnostic states can be easily seen. Plantsof CentralAmerica and Mexico: 5, 7, 8, 11, 19, 32, 36, 45, 48, 49, 54, 61, 63, 77, 79, 83, 96,97, 104, 113, 114, 116, 118 Plants of SE Brazil: 6, 8, 9, 12, 13, 14, 16, 17, 20, 117 21, 22, 23, 27, 29,30,65,87,88, Plantsof Caribbeanislands (including Trinidad& Tobago): 5, 8, 19, 51, 54, 99, 101 Shrubsof river courses, rheophytes (usually with very narrowleaves): (9?), 48, 52, 93, 96 Plantsgrowing in weedy thickets at roadsides: 10, 11, 12, 19,28,51,97 orPrevost's (Chamberlain's Pagoda-likearchitecture model with tiers ofplagiotropicbranches):101, 102, 103?, 104, 105, 106,108,109,110?,111, 114, 115, 117, 118?, 119 Stems markedlywinged: 3, 56, 78, 91, 123 Bark of older stems exfoliating: 45, 49, 116, 117 Sympodial units unifoliate: 37, 65, 66, 67, 68, 69, 70, 71, 78, 84, 101, 102, 104, 109, 111, 114, 115, 118, 119, 120, 121,122 Plantscompletelyglabrous,anytrichomesnotvisible to the naked eye: 5, 8 (cultivated), 29, 30, 126 New growth resinous: 33, 40 Trichomes of new growth reddish when dry: 4, (37), 43, 45, 53, 54, 58, 65, (72), (73), 75, 76 Leaf margins markedly undulate (ruffled): 101, 102, 103, 104, 105, 106, 108, 109, 110, 111, 112, 113, 114, 115, 118, 119 Leaf pubescence felty (arachnoid trichomes): 2, 58?, 65, 81, 82, 83, 84, 85

Leaves bullate or markedlyrugose: 39, 40, 45, 82 Leaves dryingblack: 12, 13, (18), (25), (27), 30, 77, 78, 79, 126 Leaf venation triplinerved with markedly parallel lateralveins (like Melastomataceae):67?, 70. Leaves of a geminate pair markedly unequal and differing in shape: 32, 35, 50, 53, 59, 106, 107, 108, 112, 117 Leaf andstem trichomesechinoidor Christmas-treelike: 2, 37, 38, 88 Trichomesdendriticorantler-like:1,3, 6, 8,9, 32, 34, 37, 39, 76, 80, (83), 87, 88, 89, 90,91,92, 99, 106 Leaftrichomessimple, long anduniseriate:7,26, 31, 44, 49, 50, 55, 68, 77, 78, 86, (92), 100, 101, 102, 106, 111,118, 124 Leaftrichomes with enlargedbases: 14, 27 Tufts oftrichomes in the vein axils beneath: 10, 11, 12, 13, 18, 19, 20, 22?, 23?, 24, 25, 26, 28, 32, 41, 43, 58?, 81?, 89, 124 Pubescence uniform and dense on leaf undersides: 2, (3), 4, 7, 8, 21, (31), (32), 34, 35, 36, 37, 38, 39, 44, 68, 76, 80, 84, 87, 88, 90, 91, (92), 106, 118, 123 Inflorescencebranchedmany times: 1, 2, 3, 4, (10), (18), (21), 30, (35), 39, 40, 42, (75), (89), 91, 92, 93, 95, 97, 98, 99, (100), (112), (116), 123, 126 Flowers sessile, no inflorescence axis present (if so very small): 64, 103, 114, 120 Inflorescence filiform, longer than 3 cm: 109, 111, 115, 121 Corollacampanulate:13,46 Corollalobes stronglyreflexed at anthesis:5,47, 51, 66, (69?), 101, 103, 106, 108, 110, 112, 114, (118?), 119, 120, (124?) Corollalobes tomentose(not merelypapilloseon the tips and margins)abaxially:1, 2, 4, 6, 35, 36, 37, (38), 76, 80, 81, (86), 87, 88, 91, (99), 123, (124) Anthersof unequallengths (3 long, 2 short):20, 21 Anthers strongly connivent: 30, 31, 32, 33, (34) Anthers obovoid in shape, the base narrow: 123 Corolla green: 38, 119 8,9, 12,29,50 Calyxlobespetaloidor long-triangular: Calyx lobes orbicular(live plants): 113 Fruitingpedicels erect: 5, 6, 8, 9, 52, 53, 54, 58, 59, 61, 62, 63, 66, 67, 69, 70, 71, 75, 100 Berriespubescent:35, 72, 73, 80, 81, 88, 91,92, 99, 100 Berries > 2 cm in diameter:72, 73, 76 Berry red or orange at maturity:6, 8, 9, 24 Seeds flattened-reniform,with incrassatemargins: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, 38,39,40,41,42,43,44,45,46 Seeds ovoid-reniform, the embryo clearly visible: 47,48,49,50,51,52,53, 54, 55, 56, 57, 58, 59,

TAXONOMIC TREATMENT43

60, 61, 62, 63, 64, 65, 66, 67, 68, 69, 70, 71, 72, 73, 74, 75, 76, 77, 78, 79, 80, 81, 82, 83, 84, 85, 86, 87, 88, 89, 90, 91, 92, 93, 94, 95, 96, 97, 98, 99, 100, 101, 102, 103, 104, 105, 106, 107, 108, 109, 110, 111, 112, 113, 114, 115, 116 117, 118, 119, 120, 121, 122

I. Solanum oblongifolium species group (S. clivorum, S. hypaleurotrichum,S. oblongifolium, S. venosum). Fig. 19A-C

opposed except occasionally in S. clivorum;pedicel scars closely spaced and overlapping.Buds globose, the calyx thick and appearingwoody when dry.Flowers white, large and usually waxy, the petals planarat anthesis. Fruit green and hard at maturity;fruiting pedicels erect to deflexed from the weight of the fruit. Seeds flattened-reniform,pale yellowish-brown, very large, with prominentlyincrassatemargins.

Distribution. Andean,Venezuelato Peru.Members of the Solanum oblongifolium species group are superficiallyreminiscentof species belongingto Solanum Treelets or small trees, young stems and leaves sect. Brevantherumin their large, complex infloresdenselydendritic pubescent.Sympodiausuallyplurifoliate, cences. However, membersof sect. Brevantherumalin Solanumclivorumdifoliateandthe leaves geminate. ways have stellate or echinoid pubescence (see Roe, Leaves large(to 30 cm) andrepand,the apex acute,the 1971) and the inflorescences arise from a dichasial base occasionallysessile andsomewhatauriculate,leaf branchfork (see Knapp, 1989) insteadof being lateral pubescence always dendritic,the trichomes transpar- as they are in the S. oblongifolium species group. In ent to beige or reddish-brown.Inflorescenceslargeand this species groupin the Andes of Ecuadorhybridizabranched,usuallyterminal,occasionallylateral,not leaf- tion appearsto be relatively common (see below).

Key to the species of Solanum oblongifolium species group 1. Trichomes of stems dendritic, transparent. 2. Flowers waxy, 1.2-2 cm diam.; fruit to 2 cm diam., green and hard at maturity; sympodial units plurifoliate .............................................................................................................................. 3. S. oblong f liu m 2. Flowers membraneous, 1-1.1 cm diam.; fruit 1-1.2 cm diam., green and soft at maturity; 1. S. clivorum sympodial units difoliate; the leaves usually geminate .................................................... I. Trichomes of stems Christmas-tree-like or dendritic, reddish or beige, never transparent. 3. Trichomes reddish, those of leaf undersides dendritic; leaves with 10-20 pairs of main lateral v ein s .................................................................................................................................................. S. venosum 3. Trichomes beige, those of leaf undersides Christmas-tree-like;leaves with 6-8 pairs of main lateral veins ...................................................................................................................................... 2. S. hypaleurotrichum

1. Solanum clivorum S. Knapp,Novon 2: 341. 1992. Type.Peru.La Libertad:BetweenHuamachucoand Cajabamba,3000-3200 m, 16 Mar 1948,Ferreyra 3059 (holotype, USM; isotypes, IBE, MO, US). Fig. 20 Bushy shrubs, 2-3 m tall; young stems and leaves densely pubescentwith dendritictrichomesca. 0.5 mm long; bark of older stems reddish or green, glabrous. Sympodial units difoliate, geminate, or appearing plurifoliatedue to rapidinflorescence growth.Leaves elliptic,commonlywith severalaxillaryshoots,glabrous (occasionallywith scatteredsimpleuniseriatetrichomes on the lamina)andwith a few dendritictrichomesalong the main veins adaxially,densely pubescentwith dendritictrichomesabaxially;majorleaves 13-21 (-30) cm long, 6-9(-17) cm wide, with 7-9 pairsof main lateral veins, these dryingyellowish, the apex acute, the base acute;petiole 2-3 cm long; minorleaves differingfrom the majorsonly in size, 7-10 cm long, 3-4 cm wide, the apexacute,thebase acute;petiole 1-2 cm long.Inflorescences oppositethe leaves or appearingshoot-terminal, 3-7 cm long, many times branched,20-40 flowered, denselypubescentwithdendritictrichomes;pedicelscars

unevenly spaced 1-2 mm apart.Buds globose, the corollaonly halfwayexsertedfromthe calyxtube.Pedicels at anthesis erect, tapering,6-8 mm long, ca. 0.5 mm diam.at the base, densely pubescentwith soft dendritic trichomes like those of the rest of the inflorescence. Flowerswith thecalyxtubeconical, 1-1.5 mm long, the lobes deltate, 0.5-1 mm long, densely pubescentwith dendritictrichomes;corollawhite, 1-1.1(-2) cm diam., lobed 1/2 way to the base, the lobes planarat anthesis, denselypubescenton abaxiallobe surfacewith minute, dendritictrichomesor occasionally glabrous;anthers 1.5-2 mm long, 1-1.5 mm wide, poricidal at the tips, the pores not opening to slits; free portion of thefilamentsca. 0.5 mmlong,thefilamenttubeca. 0.5 mm long, with 5 triangularprojectionsca. 0.5 mm long alternating withtheanthersandclosely investingthestyle;ovary glabrous;style 3-4 mm long, glabrous;stigmacapitate, the surfaceminutely papillose. Fruit a globose, green berry,1-1.2 cm diam.;fruitingpedicels woody, erect, 1.5-1.7 cm long, ca. 1.5 mm diam. at the base. Seeds darkreddish-brown,flattened-reniform with thickened 3mm 2-2.5 mm the surfaces wide, margins, long, minutely pitted. Chromosomenumbernot known.

44

FLORA NEOTROPICA

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TAXONOMIC TREATMENT

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Distribution (Fig. 21). N Peru in the province of Cajamarca, in montane forest at 2000-3200 m. Specimens examined. PERU. CAJAMARCA: Prov. Cutervo, Laguiac, rd. to Cochabamba, 2000 m, 26 Feb 1985, Llatas Quiroz 1178 (F); Cajabamba, Finca Zil, 2500 m, 22 Aug 1985, Mostacero L. & Guerra L. 63 (HUT, MO, NY); Cajamarca, between Matara and Namora, 2600 m, 16 Aug 1973, Sdnchez Vega et al. 1217 (NY); prov. Cutervo, distrito San Andr6s, above the caves, 2500 m, 13 Oct 1987, Sdnchez Vega 4510 (BM, F); prov. Cutervo, La Pucarilla, between Socotra and San

Andr6s,2500 m, 3 Nov 1991,SdnchezVegaet al. 5960 2600 m, 16 Aug 1973,Sagdstegui (F); Namora-Matara, A. 7755 (HUT, MO, NY); Huamachuco-Cajabamba, 2800 m, 16 Nov 1983,SagdsteguiA. 11144(HUT,NY). Local names. Peru.Cajamarca:shirac Solanumclivorumis probablymost closely related to S. oblongifoliumand may in fact be a geographical derivative of that species. It differs from S. oblongifoliumin its usuallydifoliate,geminatesympodial units, smaller, more membraneousflowers, and

FLORA NEOTROPICA

46

2. Solanum hypaleurotrichum Bitter, Repert. Spec. Nov. Regni Veg. 16: 100. 1916. Type. Colombia. Narifio:Pasto, Triana2243 (lectotype,W,heredesignated;isolectotypes, B [destroyed:F neg. 2664], NY). Figs. 19C, 22

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Shrubsor small trees, 2-8 m tall; young stems and leaves densely pubescentwith fawn (light beige) congested dendritictrichomesca. 1 mm long, the branches of the trichomesso closely packedthatthe structureis unclear;barkof older stems darkbrown,not glabrate. Sympodia plurifoliate. Leaves elliptic, widest at the middle,glabrousandshiningadaxially,densely pubescentabaxiallywithbeige congestedtrichomeslike those of the stems, these denser along the veins, often matted, leaf blades 3.5-15 cm long, 1.5-6 cm wide, with 6-8 pairsof main lateralveins, these impressedadaxially, the apex acute, the base acute;petiole 1.5-2.5 cm long. Inflorescencesterminal,laterlateral,2-7 cm long, 2-5 times branched,with 5-15 flowers, densely pubescent with beige congested trichomes (Christmastree-like)likethoseof thestemsandleaves;pedicelscars unevenly spaced 1-2 mm apart.Buds globose ca. 1/2 way included in the calyx tube. Pedicels at anthesis thick, 0.5-1 mm long, ca. 1 mm diam., densely pubescent with tree-liketrichomeslike those of the restof the inflorescence.Flowers with the calyx tube campanulate,constrictedatthebase,2-4 mmlong,thelobesdeltate, 1.5-3 mm long, with scatteredtree-liketrichomeslike those of the rest of the inflorescence; corolla white, fleshy, 1.6-2.4 cm diam.,lobed ca. 1/2 way to the base, the lobes planarat anthesis,the abaxial surfaces,margins and tips of the lobes densely papillose; anthers obovate, 3.5- mm wide, 2-2.5 mm wide at the apex, narroweratthebase,poricidalatthetips,theporeslengtheningto slits with age; freeportionof thefilaments1.52 mm long, the filamenttubeca. 0.5 mm long, glabrous; ovaryglabrous;style glabrous,heteromorphic,in shortstyled flowers ca. 4 mm long and included within the anthercone, in long-styled flowers 5-6 mm long, exsertedfromthe anthercone;stigma clavate,the surface minutelypapillose.Fruit a globose (wartyfide Cazalet & Pennington5511) greenberry,ca. 1 cm diam.,occasionallyapicallypointed;fruitingpedicelserect,woody, 1-2 cm long, 1-2 mm diam. at the base.Seeds reddishwithprominentincrassate brown(?), flattened-reniform margins,ca. 3 mm long, ca. 3 mm wide, the body of the seed flat, the margins minutely pitted. Chromosome number:n = 12 (voucher Knapp et al. 9085, 9094).

smaller berries on more elongate deflexed fruiting pedicels.The leaf andstemtrichomesofS. clivorumare generallysmallerandwith less elongatebranchesthan those of S. oblongifolium.In bothspecies, however,the The five projectionsfromthe trichomesaretransparent. filamenttubethatclosely investthe style in S. clivorum are occasionally present in S. oblongifolium. In S. oblongifolium,however, they are not as long and are not consistentwithin a given population.These projections are foundin all known collections ofS. clivorum. Two recent collections from Cutervo province (Sanchez Vega 4510, Stinchez Vega et al. 5960) are Distribution (Fig. 21). In dwarf cloud forest and unusualin theirvery large leaves, larger,waxier flowers, and glabrous corolla lobes. They may represent along paramoedges, at 2000-3500 m, Ecuador. merely a geographicalvariant,but could also be a difSelected specimens examined. ECUADOR. AZUAY: ferent species. No fruiting specimens have been col- Hacienda Yanasacha, 3000-3400 m, 24 Jul 1978, Boeke & Jaramillo 2530 (K, NY); Cuenca-Gir6n rd., Nudo lected from amongstthese populations.

47

TAXONOMIC TREATMENT

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48 de Portete 9 km from Cuenca, side rd. to Hacienda, 7 km from main rd., 2700 m, 20 Aug 1987, Zak 2511 (NY). BOLIVAR: Santiago-San Vicente-Alizo "San Agustin" rd., Rio Tataguazo, 2500 m, 24 Feb 1987, Zak 1742 (NY). CARCHi: Tulcan-Tufifio-Maldonado, La Pradera, 2360-2640 m, 11 Oct 1986, Zak 1384 (NY). CHIMBORAZO: Riobamba-Penipe-Bayushig-Llurayacu rd., 3400-3600 m, 18 Feb 1987, Zak 1629 (AAU). IMBABURA: Lago San Marcos, Cayambe, 11200 ft, 4 Dec 1961, Cazalet & Pennington 5511 (BM, K, NY, US); San Jose de Minas-Otovalo rd., side rd. to television antenna, NE of Peias Blancas, 19 Jan 1980, Jaramillo & Proano 1997 (NY, QCNE); Parroquia San Luis de Qichinche, San Alberto, 2700-2900 m, 0?12'S, 78?22'W, 19-21 Jul 1989, Moran et al. 57 (BM, QCNE). LOJA: Parque Universitaria, Loja, s.d., Bolivar s.n. (LOJA); Parque Educacional Recreacional Universidad de Loja, vic. of Loja, 1200-2300 m, 4?02'02"S, 79011'87"W,28 Oct 1994, Knapp et al. 9085, 9094 (LOJA, QCNE). NAPO: Julio Andrade-San Francisco, near Cocha Seca, 3100-3300 Olmedom, 30 Dec 1986, Zak 1586 (NY). PICHINCHA: Laguna San Marcos rd., 3500 m, 0?07'N, 78?00'W, 21 May 1980, Holm-Nielsen & Balslev 23714 (K, NY, U); Bosque Protector Pasochoa, 30 km SE of Quito, 28503900 m, 0? 27'S, 78?28'W, 15 Nov 1986, Zak & G6mez 1436 (AAU); old Machachi-Volcan Cotopaxi rd., ca. 2600 m, 23 May 1988, Zak 3684 (AAU); slopes of Volcan El Coraz6n, 3300 m, 0?30'S, 78?25'W, 23 May 1988, Zak & Jaramillo 3684 (BM, MO, QCNE). Possible hybrid specimens with some characters of Solanum oblongifolium, usually with the leaf trichomes arachnoid and matted. ECUADOR. AZUAY:Cuenca, Hacienda Yanasacha, 3000-3400 m, 24 Jul 1978, Boeke & Jaramillo A. 2530 (NY); PartideroLlantera-ChiquintadRepresa de la Empresa Electrica-Labrado rd., km 33, 7 Aug 1986, Jaramillo et al. 8910 (NY). BOLIVAR: Simiatug, Hacienda Talahua, 3200 m, 3 May 1939, Penland & Summers 625, 626 (COLO(Nee neg.), GH). CANAR: Cafiar-Azoguez detour to Moloboc-GrandeMolon Ventana rd., between Moloboc & Molon Ventana, 3200--3450 m, 14 Aug 1987, Zak 2443 (NY). CHIMBORAZO: Rio Bamba-Penipe-Bayashig-Llurayacu rd., near Bayushig, 3400-3600 m, 18 Feb 1987, Zak 1627 (NY); Rio Bamba-Ambato rd., 2400-2650 m, 19 Feb 1987, Zak 1659 (NY). COTOPAXI:LatacungaAngamarca rd., near Angamarca, 3100-3600 m, 17 Feb Cotacachi-Cuicocha1987, Zak 1606 (NY). IMBABURA: Apuela rd., 2800-3000 m, 5 Dec 1986, Zak 1454 (NY). NAPO: N side of Cerro Sumaco, 3100-3150 m, 0?34'S, 77?43'W, 28 Apr 1979, Holm-Nielsen et al. 17400 (NY). PICHINCHA:Quito-Guantopugro-Yanacocha, 34003500 m, 22 Mar 1987, Zak 1847 (NY). Local names. Ecuador. Imbabura: furafango. Solanum hypaleurotrichum can be difficult to distinguish from S. oblongifolium and S. venosum at a superficial glance. It differs from those species in its long-petiolate, usually somewhat smaller leaves, smaller fruits on elongate fruiting pedicels, and in its trichome

FLORA NEOTROPICA

morphology. The branches of the inflorescence in S. areusuallylongerandtherearefewer hypaleurotrichum of them than in S. oblongifolium.The trichomes of S. hypaleurotrichumare distinctive and are somewhat intermediatebetweendendriticandarachnoidin structure.They aregenerallybeige coloredandtheirdensity variesconsiderably.In some specimensofS. hypaleurotrichumthe trichomes of the abaxial leaf surfaces are densely mattedand arachnoid(see above). This probablyrepresentsan extremein intraspecificvariation,but may be indicative of hybridizationwith S. oblongifolium. For this reason I have listed these specimens separatelyso that they can be studied in detail in the future.Furtherfield studyofS. hypaleurotrichum where it is sympatricwith S. oblongifoliumis clearly needed to solve this problem. 3. Solanum oblongifolium Dunal, Solan. Syn. 14. 1816. Type.Colombia.Caldas:Quindio,Humboldt & Bonpland s.n. (holotype, P-Bonpl.; isotypes, P [Mortonneg. 8269], frag.F). Figs. 19A,B, 23 Solanum pteropodum Dunal, Solan. Syn. 14. 1816.

Type.Ecuador.Pichincha:nearQuito,Humboldt & Bonplands.n. (holotype, P-Bonpl.). Solanum jamesoni Bitter, Repert. Spec. Nov. Regni Veg. 16: 98. 1916. Type. Ecuador. Pichincha: Andes of Quito, Jameson 627 (lectotype, BM,

here designated). Solanum hypomalacophyllum Bitter ex Pittier, Man. PI. Usual. Venez. 137. 1926. Type. Venezuela. Merida: woods above Tafayes, 2600 m, 2 Sep 1921, Jahn 533 (lectotype, VEN, here designated; isolectotype, US). Solanum oblongifolium var. soukupii J. F. Macbr.,

Publ. Field Mus. Nat. Hist., Bot. Ser. 13(5B): 211. 1962. Type. Peru. Piura:Ayavaca,Soukup 4336 (holotype, F; isotype, US).

Shrubsor small trees,2-20 m tall;young stems and leaves variouslypubescentwith loose, translucentdendritictrichomesca. 1 mm long; the stems thick, erect, oftenwingedfromthe decurrentleaf bases;barkof older stems pale greenish-yellow.Sympodiaplurifoliate,often with much axillary branching,giving the stems a bushyappearance.Leavesellipticto obovate,oftenvery large, especially on juvenile plants, widest at or just above the middle, the laminaglabrousadaxiallyor occasionallysparselypubescentwith dendritictrichomes, the pubescence dense along the veins, abaxial surface densely to sparsely pubescent (a few collections glabrous) with translucentdendritictrichomes 1-2 mm long, these denseralong the veins; blades 9-30x 4-15 cm, with 15-20 pairs of main lateralveins, these drying yellowish abaxially,the apex acute to acuminate, the base acute to sessile with the basal portion of the

TAXONOMIC TREATMENT

49

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1

FIG. 23. Solanum oblongifolium Dunal. A. holotype in P-Bonpl. B. Ecuador. Mexia 7664 (BM).

leaf winged onto the stem and crotaloid;petiole 0-1.5 cm long, if leaves sessile, alwayswingedontothe stems. Inflorescenceterminal,laterlateral,5-25 cm long,many times branched,to 15 cm across,20-200-flowered, the pubescence like that of the stems;pedicel scars unevenly spaced 1-2 mm apart.Buds globose, later obovoid, strongly exserted from the calyx tube.Pedicels at anthesisstout,erect,0.2-1 cm long, 1-1.5 cm diam., denselyor sparselypubescentwithtranslucentdendritic trichomes.Flowers with the calyx tube broadly conical, 1.5-2.5 mm long, the lobes broadlydeltatewith a prominentknob ca. 0.3 mm from the apex, 0.3-1 mm long, densely or sparselypubescentlike the rest of the inflorescence, the apex always possessing a tuft of simpletranslucenttrichomes;corollawhite, fleshy, 1.22 cm diam., lobed nearly to the base, the lobes with large thin margins,planarat anthesis,the tips and distal marginsof the lobes densely papillate;anthers3-4 mm long, 1-2 mm wide, poricidalat the tips, the pores teardropshaped;freeportionof thefilaments0-0.5 mm long, the filamenttube0-0.5 mm long, glabrous;ovary glabrous;style stout,barelysurpassingthe anthercone, ca. 5 mm long; stigma capitate, the surface minutely papillose. Fruit a globose, green or yellowish-green berry,the pericarphardandwoody, 1.5-2.5 cm diam.;

fruitingpedicels woody, erect, 0.8-1.1 cm long, ca. 5 mm in diameter.Seedspale tan,flattened-reniform with a prominentincrassatemargin,3.5-5 mm long, 3-3.5 mm wide, the body of the seed flat, the margins minutely pitted. Chromosomenumberunknown. Distribution (Fig. 24). Common in secondary growthat high elevationsin the Andes fromVenezuela to northernPeru, 2000-3500 m. Selected specimens examined. VENEZUELA. MtRIDA: San Rafael, 3185 m, 4 Apr 1930, Gehriger1

(F,NY); SanRafael,3150m, 22 Jan1922,Jahn766 (US); ParqueNacionalSierraNevadade SantoDomingo,km 86 Merida-Barinas, betweenAlto de MucubajiandSanto Domingo, L6pez Figueiras 8713 (US); San Rafael de Mucuchies, 6 Feb 1928, Pittier 12915 (NY, US); El Chorrer6n-Caiadadel Ahorcado,rd. from Palmirato paramode Palmira,2560-2750 m, 30 May 1975,RuizTerdn& Dugarte 12488 (K); Merida to paramode la Culatard., 2900 m, 13 Mar 1980, Sobel & Strudwick 2180 (BH, NY); "LaEme"or Caiiadadel Cedro,above Bailadores,2300 m, 20 Sep 1942, Tamayo2391 (US); SantaCruz-ElMolinard., 2030 m, 6 Jun 1967, Trujillo 8172 (MY, NY). TACHIRA: ParamoBatall6n, toward Pregonero,2800 m, Sep 1956,Aristeguieta2529 (NY); La Grita,paramodel Rosal, 2800-3200 m, 8 Oct 1965,

50

FLORA NEOTROPICA

Cocuy, Valle de la Uvita, El Hatico, 2900 m, 7 Sep 1938, Cuatrecasas 1162 (US); Valle de Cocuy, SW slopes, 3100-3750 m, 8 Sep 1938, Cuatrecasas 1285 (F, US); paramo between Duitama and Charala, Cordillera Oriental, ca. 3050 m, 7 May 1944, Ewan 15650 (NY); Sierra Nevada del Cocuy, path from Guicanto to Hacienda Rita Cuba, ca. 3300 m, 19 Aug 1957, Grubb et al. 413 (K). CUNDINAMARCA: Boquer6n de Bogota, 2890 m, 26 Jan 1876, Andre 1289 (K); near Salto de Tequendama, 1917, 00 Ariste-Joseph All9 (US); Salto de Tequendama, Jul 1917, Ariste-Joseph A139 (US); region of Bogota, 1919, Ariste-Joseph s.n. (US); NW of Zipaquira on rd. to Pacho, near km 21, 2900 m, 26 Feb 1974, Barclay et al. 3748 (US); Salto de Tequendama, La Rinconada, 22502300 m, 2 Oct 1938, Cuatrecasas 240 (F, US); Bogota Massif, Quebrada de Chic6, 2650-2750 m, 1 Jun 1939, Cuatrecasas 5237 (F, US); Bogota Massif, Quebrada de ,.-~ 0o* #' las Delicias, 2650-2700 m, 11 Jun 1939, Cuatrecasas 5467 (F, US); San Miguel, extreme W of Sabana de Bogota, 2800 m, 15 Aug 1939, Cuatrecasas 6700 (F, US); extreme SW of Sabana de Bogota, San Miguel, 2800-3000 m, 10 Sep 1941, Cuatrecasas & Jaramillo 12006 (F, US); Bogota, Ciudad Universitaria, 2620 m, 20 Mar-20 Apr 1946, Duque-Jaramillo 2897 (NY); Laguna de Pedro Palo, 3 km N of Tena, 2080 m, 19 May 1952, Ferndndez & Mora 1436 (US); Quebrada El Chic6, extreme N Bogota, 3200 m, 3?40'N, 74?04'W, 11 Jul 1943, Fosberg & Villareal 20583 (NY, US); Bogota, Ciudad Universitaria, Sabana de Bogota, 2600 m, 10 Apr 1945, Garcia-Barriga 11621 (US); W slope, Cordillera Oriental, between San Miguel and La Aguadita, 1980-2360 m, Garcia-Barriga 12104 (US); 0FIG. 24. Distribution of Solanum oblongifoium. Anolaima rd. above La Florida, 2740 m, 3 Nov 1941, Gutierrez V 114 (GH); W slope, Cordillera Oriental, FIG. 24. Distribution of Solanum oblongifolium. paramo de Cruz-Verde, 2840 m, 21 Apr 1942, Gutierrez V 250 (GH); Guadalupe Hill, Bogota, ca. 3000 m, 23 Mar 1947, Haught 5590 (US); 8 km SW of Guasca, ca. 2900 m, 25 Jun 1947, Haught 5894 (US); Pacho-San Bernardi 10927 (K); headwaters of Rio Quinimari along Cayetano, 2500 m, 7 Aug 1947, Haught 6036 (US); Quebrada Agua Negra, trail to paramo de Judio (Apure Tequendama, ca. 2400 m, 11 Sep 1949, Haught 6604 border), 5 km S of San Vicente de la Revancha, 15 km S (US); Mt. Guadalupe, above Bogota, Sep 1917, Idinael of Providencia, SE of Santa Ana, 2100-2400 m, 7?25'N, 8 (NY); near Facatativa, 2700 m, Jan 1883, Lehmann 72?25'W, 23 Oct 1984, Knapp & Mallet 6822 (BH, K, 2594 (US); SE of Bogota on trail to Ubaque, 9600 ft, 7 MY, NY, US, VEN); below paramo de Tama, Colombian Jan 1945, Little & Little 9236 (GH, US), Little & Little border, 2475-2550 m, 18-20 May 1967, Steyermark et 9237 (NY, US); W slopes Cordillera Oriental, Bogotaal. 98413 (VEN, US); W of paramo de La Negra, be- Villaviencio rd., km 18, 3100 m, 7 Jan 1976, Luteyn et tween Sabana Grande and paramo de La Negra, 2800al. 4744 (MO, NY); Bogota plateau, San Crist6bal, 28 3000 m, 29 Aug 1966, Steyermark& Rabe 96867 (VEN, Dec 1926, Niemeyer 155 (US); Rio San Francisco above US); headwaters of Rio Quinimari, near Las Copas, base Bogota, 2700-2800 m, Pennell 1941 (GH, NY); of Pefia del Pato de Judio below paramo de Judio, 15 km Tequendama, 2400 m, 15 Sep 1917, Pennell 1962 (GH, S of San Vicente de la Revancha, 30 km S of Alquitrana, NY); Cerro Monserrate, 11000 ft, 12 Jan 1969, PlowSW of Santa Ana, 2400 m, 10-11 Jan 1968, Steyermark man 2205 (GH, K, US); 3 km S of Usme on rd. to et al. 100734 (VEN, US); Quebrada Agua Azul, S of El Chisaca, 2975 m, 4 May 1972, Plowman 3199 (F, GH, Reposo, 14 km SE of Las Delicias, 2150-2300 m, 22-23 US); L bank of Rio San Crist6bal, 25 Apr 1951, Romero Jul 1979, Steyermark& Liesner 118496 (VEN, MO, NY). Castaneda 2458 (F); Hacienda Chic6 near Usaquen, 11 COLOMBIA. Niebli, 17 Jun 1876, Andre K645 (K); Feb 1945, Scheifer 443 (GH); Suba Hill, near Bogota, 7 s.loc., Apolinar Maria 31 (F); s.loc., 1760-1808, Mutis Sep 1945, Scheifer 904 (GH, US); Bogota, mtns. back 2011 (US). ANTIOQUIA:Paramo de Sons6n, 2700-2850 of Calle 75, 11 Oct 1945, Scheifer 929 (GH); paramo m, 26 Jan 1945, Daniel 3415 (US); Cerro de la Vieja, de Chipaque, 24 Feb 1951, Schultes 11615 (GH, NY); 2600 m, Daniel 1697 (US). BOYACA:Between Soata and above Usme, 17 Mar 1970, Schultes & Cabrera 26030

800

i

TAXONOMIC TREATMENT (GH); Boquer6n de Bogota, Jan 1925, Schultze 105 (US); below paramo de Chisaca, ca. 3000 m, 15 Aug 1963, Soejarto 424 (US); Bogota, 2700 m, 1851-1857, Triana s.n. (BM, NY, US). NORTE DE SANTANDER: Paramo de

Fontibon, Pamplona, 2400 m, 19 Feb 1939, Alston 7144 (U, US); Cordillera Oriental, Sarare, Quebrada de Rio Chitaga between Fontib6n and La Acbuya, 2200 m, 24 Oct 1941, Cuatrecasas et al. 12596 (F, GH, U, US); Pamplona, near Escuela Nacional, 15 Feb 1945, Garganta 936 (F); Culaga valley, near Tapata, N of Toledo, 1500-2100 m, 3-8 Mar 1927, Killip & Smith 20154 (A, GH, NY, US). SANTANDER: Edge of paramo de las Vegas, 3300-3700 m, 20-21 Dec 1926, Killip & Smith 15762 (A); mtns. E of La Vegas, 3000-3300 m, 20-21 Dec 1926, Killip & Smith 15850 (A, GH, NY, US); vic. California, 3000 m, 11-27 Jan 1927, Killip & Smith 16947 (A, GH, NY, US); vic. California, 2500 m, 11-27 Jan 1927, Killip & Smith 16983 (US); vic. La Baja, 2700-3500 m, 14-31 Jan 1927, Killip & Smith 17156 (A, GH, NY, US); vic. Vetas, 3100-3250 m, 1620 Jan 1927, Killip & Smith 17302 (A, US); vic. Vetas, 3100-3200 m, 16 Jan 1927, Killip & Smith 17351 (A, GH, NY, US); vic. Vetas, 3100-3200 m, 16 Jan 1927, Killip & Smith 17389 (A, GH, NY, US); W slope paramo de Rico, 3200 m, 15-19 Jan 1927, Killip & Smith 17863 (US); vic. Vetas, 3100-3250 m, 16-20 Jan 1927, Killip & Smith 17887 (A, GH, NY, US); S slope of paramo de Romeral, 3800 m, 29-31 Jan 1927, Killip & Smith 18711 (A, GH, NY, US); vic. Charta, 2000-2600 m, 111 Feb 1927, Killip & Smith 19249 (A, GH, NY, US). ECUADOR. AZUAY: Along Rio Matadero, vic. Cuenca, 8200-8900 ft, 16 Apr 1945, Camp E-2704 (NY); vic. Cuenca, 17-24 Sep 1918, Rose et al. 22912 (GH); km 12-15 of Gualeceo-Macas rd., 2700-2900 m, 6 Aug 1986, Jaramillo et al. 8873 (QCA, NY); Cuenca-Sayausi-Tres Cruces rd., between Sayausi and Quinoas, 3050 m, 16 Aug 1987, Zak 2461 (NY); Cuenca-Bosque de Mozan (ETAPA) rd., 3000-3200 m, 17 Aug 1987, Zak 2478, 2484, 2487 (AAU, NY). BOLIVAR:Chillanes-San Pablo de Atenas-Guaranda rd., between Chillanes and San Pablo, 2300-2500 m, 27 Feb 1987, Zak 1780 (NY); Chillanes-Bucay rd., Hacienda Tiquibuzo, 2100 m, 29 Aug 1987, Zak 2581 (K, NY); San Pablo de Atenas-Chillanes rd., loma de Peresan, 2000-2300 m, 1?50'S, 79?05'W, 31 Aug 1987, Zak & Jaramillo 2623 (K, NY); Chillanes-TambilloTrigoloma rd., vic. San Juan Bampa, 2000 m, 4 Sep 1987, Zak 2729 (NY); Chillanes-Tiquibuzo-AchinBucay rd., vic. Achin, 2200 m, 10 Sep 1987, Zak 2896 (AAU, K, NY). CANAR:Alausi-Cafar rd., side rd. to Oyashig and Hacienda El Carmen, N of Cafar, 32703700 m. 12 Aug 1987, Zak 2396, 2410, 2413 (NY); Cafiar-Azoguez rd., side rd. to Moloboc Grande and Molon Ventana, 3200-3450 m, 14 Aug 1987, Zak 2437, Alausi-Yocon 2440, 2451 (AAU, NY). CHIMBORAZO: Grande-Garcia Moreno, 10 km from town, 2600 m, 9 Aug 1986, Jaramillo et al. 8974b (NY); Riobambard., 2400-2650 m, 12 Feb Cubiji6s-Bafos-Ambato 1987, Zak 1658 (NY); Pallatanga pass-San Juan Llimbe, 2700-2800 m, 26 Feb 1987, Zak 1770, 1772 (NY);

51 Parque Nacional Sangay, Riobamba-Chambo-PungalaAlao rd., 3150 m, 9 Aug 1987, Zak 2359 (NY); La Moya-Achupallas rd., S of Alausi, 10 Aug 1987, Zak 2366 (NY); Alausi-Baguil-Guamote rd., 2800-3200 m, 11 Aug 1987, Zak 2379, 2383, 2388 (AAU, NY). COTOPAXI:Pilalo-Zumbagua rd., 15 km above Pilal6, 3350 m, 0?59'S, 78?58'W, 28 Jul 1980, Holm-Nielsen & Quintana 24626 (NY); W slope of Volcan Cotopaxi, 9700-9900 ft, 0?42'S, 78?28'W, 17 Jul 1978, Webster & Lockwood 22691 (DAV); Latacunga-Angamarca rd., vic. Angamarca, 3100-3600 m, 17 Feb 1987, Zak 1616 (NY). IMBABURA:Imbabura-Mojando, 2300-3000 m, Jul-Aug, Lehmann 4718 (K); Lake Cuicocha, 3000 m, 30 May 1939, Penland & Summers 787 (GH); Lago Cuycocha, ca. 3000 m, 23 Jul 1974, Plowman et al. 3800 (GH, K); Ibarra-Mariano Acosta rd., 2700-3000 m, 1 Nov 1986, Zak 1405, 1408 (NY); Cotacachi-CuicochaApuela-Talba Chupa rd., vic. Tabla Chupa, 2800-3000 m, 6 Dec 1986, Zak 1477 (NY); Ibarra-Zuleta-Otavalo rd., 2800-3100 m, 3 Apr 1988, Zak 3654 (NY); Cotacachi-Apuela rd., 21 km from Cotacachi, Intac valley, 3300 m, 0?20'N, 78?24'W, 11 Aug 1976, 01lgaard & Balslev 8693 (AAU, F, NY); Cotacachi-Apuela rd., 33-36 km from Cotacachi, Intac valley, 2900-2950 m, 0?20'N, 78?26'W, 11 Aug 1976, 0llgaard & Bal,slev 8733 (AAU, F, NY). LOJA: Loma del Oro, 2800-3200 m, Jaramillo et al. 8793 (NY); near Zamora-Chinchipe border, ca. 39 km W of Zamora, 24 km E of Loja on Loja-Zamora rd., ca. 2200 m, 4?00'S, 79?07'W, 5 Feb 1984, Knapp & Mallet 6247 (BH, GH, NY, QCA, QCNE, US); Parque Educacional Recreacional Universidad de m, 4002'02"S, Loja, vic. of Loja, 1200-2300 79?11'87"W, 28 Oct 1994, Knapp et al. 9095 (LOJA, QCNE); rd. to La Toma on slopes of Cerro Villonaco, ca. 20 km W of Loja, 8000 ft, 7 Mar 1965, Knight 579 (K, US); near Sosaranga, between Cariamanga and Macara, 6500 ft, 10 Jun 1966, Knight 1074 (US). NAPO: Quito-Baeza 63.7 km E of Quito at Rio Papallacta, 46.2 km E of Tumbaco, 3180 m, 0?22'S, 78?14'W, 31 VMar 1992, Croat 73294 (QCNE); Quito-Papallacta-Baeza rd., paramo La Virgen, 3900-4100 m, 21 Jun 1987, Zak 2083, 2084 (NY). PICHINCHA:15 km S of Quito along hwy. to Tambillo, 2800 m, 19 May 1939, Asplund 6203 (US); Nono, 2700 m, 1 Jul 1939, Asplund 7430 (K); near Nono, ca. 2700 m, 28 Aug 1955, Asplund 17471 (K); Reserva Geobotanica Pululahua, line from La Ventanilla to Cerro Sincholanua, 3000-3300 m, 0O0'N, 78?30'W, 7 Jun 1987, Ceron M. 1541 (MO, NY); QuitoNanegal rd., via Cotocollao and Nono, 18-21 km NW of Quito, 3100-3200 m, 4 Sep 1976, Croat 38805 (MO, NY); Valle de Lloa, 14 km below Lloa, 2700 m, 0?14'S, 78?38'W, 23 Jun 1979, Holm-Nielsen 18310 (NY); Jamanco, 3500 m, 3500 m, 21 Aug 1975, Little & Campunazo 290 (MO); Hacienda La Campiia, Canton Quito, 2956 m, 12 Sep 1935, Mexia 7664 (GH, K); 8 km W of Aloag on pass to Sto. Domingo, 3000 m, 12 Sep 1984, Whalen 813 (BH, NY, US); 75 km S of Quito on rd. to Santo Domingo via Machachi, 10500 ft, 4 Aug 1980, Wunderlinet al. 8726 (F); Chillogallo-San JuanChiriboga-Ampalme rd., vic. San Juan, 3100-3260 m,

52 0?18'S, 78?39'W, 7 Sep 1985, Zak & Jaramillo 619 (K, NY); old Chillogallo-San Juan-Chiriboga-Empalme rd., km 27, 3180 m, 0?17'49"S, 78?38'47"W, 24 Feb 1986, Zak 967 (K, NY); Calacali-Yunguilla rd., 2800-3100 m, 14 Mar 1987, Zak 1820, 1823 (K, MO, NY); Chiriboga-San Juan rd., 2800 m, 7 Jun 1987, Zak 2040 (NY); Quito-Nono rd., between Nono and Rundopamba, 2700-3000 m, 20 Jun 1987, Zak 2052 (NY); QuitoCalacali-Nebli rd., 2150-2900 m, 27 Jun 1987, Zak 2086, 2090, 2100 (NY); El Pongo, 2800 m, 24 Jul 1987, Zak 2250 (NY), Zak & Jaramillo 2250 (K, MO, NY). PERU. AMAZONAS: Upper slopes of Puma-urcuESE of Chachapoyas, 2700-3000 m, 1 Jun 1962, Wurdack 684 (F, K, NY, US). CAJAMARCA: Otuzco, Rio Pollos, 2600 m, 19 Jun 1950, Anguto 923 (HUT); Chota, Cachil (Miracosta), 2500 m, 18 May 1981, Llatas Q. 707 (HUT); Contumaza, vie. Guzmango, 2500 m, 24 Apr 1978, Sagastegui A. & Mostacero L. 9174 (HUT, NY); km 148 of San Juan-Cajamarcard., E of rd., 2700-2800 m, 2 Jun 1985, Sanchez Vega 3861 (NY); ca. 4 air km ENE of Monteseco, 2400 m, 16 Jun 1987, Santisteban C. & Guevara B. 180 (BM, F, HUT); Huambos, ca. 2000 m, 10 Sep 1956, Soukup 4493 (US); vic. Huancabamba, trail from Las Huaringas to Huancabamba,23 Feb 1981, Davis & Turner 721 (F, NY). LAMBAYEQUE: Ferrefiafe, Incahuasi, ca. 3200 m, 22 Jun 1986, Llatas Quiroz 1907 (F). PIURA: 9 km S of Ayabaca on rd. to Sullana, 2700 m, 23 Sep 1964, Hutchison & Wright 6681 (F, K, MO, NY, US); Ayabaca, Yaurpampa, 2650 m, 26 May 1971, Lopez et al. 7785 (HUT); Tambo between Canchaqua and Huancabamba, May 1958, Soukup 4696 (US).

FLORA NEOTROPICA

occur between Solanum hypaleurotrichum and S. in Ecuador.Specimensof thistypearelisted oblongifolium with S. hypaleurotrichum (see above) andaregenerally in overallmorphologythan closerto S. hypaleurotrichum to S. oblongifolium,butpossess abaxialleaf surfacetrichomesthatareintermediatebetweenthe two species. In lectotypifying Solanumjamesonii, I have chosen Jameson 627 fromthe syntypesat BM as Jameson 625 is a mixed collection with a memberof sect. Torva (Solanum hispidumPers.). Both sheets are annotated in Bitter's hand.

4. Solanum venosum Dunal, Solan. Syn. 19. 1816. Type. Colombia. Caldas: Quindio, Humboldt & Bonpland s.n. (holotype, P-Bonpl. [Morton neg. 22325]; isotypes, P [Mortonneg. 8360, 8361]). Fig. 25 Solanumfulvivillosum Bitter, Repert. Spec. Nov. Pichincha: RegniVeg. 16:99. 1916.Type.Ecuador. Volcan Iliniza, 2700-4300 m, Wagner113 (holotype, M [Mortonneg. 8690]). SolanumhypiodesBitter,Repert.Spec. Nov. Regni Veg. 16: 101. 1919. Type. Colombia. Narifio: Tiquerres,Karstens.n. (holotype,W).

Shrubs or small trees, 1.5-5 m tall; young stems densely pubescentwith congested dendritictrichomes (Christmas-tree-like),the new growthdensely pubescent with loose dendritic trichomes; branches erect, Solanumoblongifoliumis a very common shrubat not glabrate;bark of older stems dark brown. thick, middle to high elevations and usually grows in open Sympodia plurifoliate. Leaves elliptic, widest at the places along roadsides or pastures. It is most closely the adaxial surfaces glabrous with dendritic middle, relatedto S. venosum,with which it is sympatricin at least part of its range. Solanum oblongifoliumdiffers trichomes along the veins or sparsely pubescent with fromS. venosumin its transparenttrichomeswhich are dendritictrichomes,the branchesof the trichomeseither congested (type) or loose, the abaxial surfaces always dendritic, never tree-like as are those of the stems of S. venosum.The flowers of S. oblongifolium densely to sparsely pubescent with reddish dendritic trichomes like those of the upper leaf surfaces, leaf are borne in very large inflorescences; however these areoften not collected as they aredifficultto press.The blades 7-18 (30) cm long, 5-12 cm wide, with 10-20 petals are quitewaxy andthe flowers have a sweet fra- pairsof mainlateralveins,the apexacute,thebase acute grance,unusualin the section (butsee S. validinervium to truncate;petioles 2-5 cm long.Inflorescencestermiin the amblophyllumgroup and S. corumbensein the nal, laterlateral,2.5-12 cm long, manytimesbranched, 5-25-flowered, densely pubescentwith tree-like denleucocarpon group). Leaf base morphologyvariesconsiderablythrough- dritic trichomes like those of the stems;pedicel scars out the rangeof Solanumoblongifoliumandhas no ob- unevenly spaced 1-2 mm apart.Buds globose, later vious geographicalcomponent.Leaves varyfrompeti- ellipsoid,stronglyexsertedfromthe calyxtube.Pedicels olate with the petioles to 3 cm long to sessile with the at anthesis stout, 0.4-1.3 cm long, ca. 1 mm diam., leaf bases somewhat auriculate.In these latterplants, densely pubescent like the rest of the inflorescence. Flowerswiththe calyxtubecampanulate,1-2 mm long, the leaf base has been termed "crotaloid," because when dry it resemblesa rattlesnake'stail (see S. crotalobasis the lobes deltatewith thickenedwhite margins, 1.5-2 Bitter, a synonym of S. maturecalvansBitter of the mm long, densely pubescentwith dendritictrichomes like those of the restof the inflorescence;corollawhite, nigricans group, named for its similar leaf bases). PlantsfromnorthernPeruare generallylargerthan fleshy, 1-1.5 cm diam., lobed ca. 2/3 of the way to the those fromthe restof the species range.Theyhave been base, the lobes planarat anthesis,the abaxialpetal surcalledvar.soukupiiJ.F.Macbr.Somehybridizationmay faces densely pubescentwith minutedensely branched

TAXONOMIC TREATMENT .

53

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?':~i FIG.25 ~~ Solan ~ mvenosm ~~~~.........I Duna.A. olotpenP- onplB.Ecuaor.ipa I. rt~~~~~~~~~~~~~~~~~~~~ur DLUCi: _ ~cr; i::;?~~~~~~~~~~~~~~~~~,__ ....................................................._ ~~~? ? I ~~~~~~~~~~i ~;*................... " _ffFW*

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~~~r,,n~~~~~~~~~~ .................. holotype in P-Bonpl. B. Ecuador. Tipaz 45 rrrutu (BM).~~?ri I*

(NY). CALDAS:Cordillera Central, W slope, SE slope of Volcan Ruiz, Termales, 3400 m, 4 May 1940, Cuatrecasas 9210 (F, US); Cordillera Central, W slope, La Virginia, above La Esperanza, 3400-3600 m, 6 May 1940, Cuatrecasas 9354-B (F); Magana to Quindio pass, old Quindio trail, 3200-3500 m, 2 Aug 1922, Killip & Hazen 9168 (NY, US). CAUCA:Cordillera Central, paramo de Purac6 S of Volcan Purac6, San Francisco, 3400-3450 m, 23 Jul 1943, Cuatrecasas 14678 (F, US); rd. to Purace, Cauca slope, 2400-2600 m, 19 Jul 1948, Garcia Barriga & Hawkes 12795 (US). NARISO: Ipiales rd. from Tuquerres to Ipiales, 2950-3100 m, 28 Jul 1948, Garcia Barriga & Hawkes 13078 (NY, US). VALLEDE CAUCA: Cordillera Central, W slope, Rio Bugalagrande,Barragan, paramo de Bavaya, Corrales, 3450-3520 m, 18-20 Mar 1946, Cuatrecasas 20150 (F); CordilleraCentral,W slope, Distribution (Fig. 26). NorthernColombiato cen- Rio Tulua, between Las Vegas & La Ribera, 3300-3450 tral Ecuador in montane forest and forest margins, m, 25 Mar 1946, Cuatrecasas 20429 (F, US). ECUADOR. BOLIVAR:Santiago-San Vicente-Alizo 2000-3500 m. "San Agustin" rd., Rio Tataguazo, 2500 m, 24 Feb 1987, Selectedspecimensexamined.COLOMBIA.Region Zak 1745 (NY). CARCHf:Paramo del Angel, rd. from El betweenSan Pedroand Belmira,banksof QuebradaEl Angel to Tulcan, 4 km from El Angel, 3160-3280 m, 4 Paramo Jan 1977, Boeke 711 (NY); ca. 2 km along rd. El Angel Diablo,25 Oct 1963,OrozcoI (US). ANTIOQUIA: de Sons6n, toward La Dorada, 2600 m, 20 Jul 1977, to Tulcan, 3150-3300 m, 0?38'N, 77?55'W, 14 May 1973, Albert de Escobar & Escobar U. 502 (NY); Alto de Holm-Nielsenet al 5219 (NY); Loma El Coraz6n(Bretafia), Ventanas,15 km SW of Jardinon way to Riosucio,2400- SE of Huaca, E of Colonia Huaquenia,Rio Minas, 32002800 m, 5?30'N,75?50'W,9 Jun1987,Callejaset al. 3918 3500 m, 0?35'S, 77?42'W, 19 Apr 1989, Tipaz 45 (BM,

trichomes and elongate papillae ca. 0.5 mm long; anthers 2.5-3.5 mm long, 1-2 mm wide, poricidal at the tips, the pores teardropshaped;free portionofthefilaments 0.2-0.5 mm long, the filament tube ca. 0.5 mm long, glabrous;ovary glabrousor with a few dendritic trichomes; style 4-8 mm long, glabrous or sparsely pubescentwith dendritictrichomes;stigmacapitate,the surfaceminutelypapillose.Fruita globose, greenberry, ca. 1 cm diam. (immature?);fruitingpedicels woody, erect, 1-2 cm long, variable within a plant, ca. 2 mm diam.Seeds pale tan to reddish-brown,flattened-reniform with prominent incrassate margins, 3.5-4 mm long, 2.5-3 mm wide, the body flat, the margins minutely pitted. Chromosomenumberunknown.

FLORA NEOTROPICA

54

80?

Solanum venosum is closely related to S. oblongifoliumwith which it is completely sympatric. The species are superficially very similar, with large leaves and complex inflorescences, but differ in trichome color and morphology.The stem trichomes of S. venosum were described as "pilus crassiculis, pubescentibus"by Humboldt, Bonpland, and Kunth (1818). This has caused considerable confusion (see Morton, 1944), but the stem trichomes of S. venosum indeed appearpubescent. They are of the Christmastree-like echinoid type with the branchesoff the main axis of the trichome closely packed and occasionally secondarily branching. This secondary branching is unusualin trichomesin sect. Geminata.Leaf and stem trichomesof S. venosumare red-brownwhile those of S. oblongifoliumare always transparentor dry white.

10? .0 0r

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X

II. Solanum pseudocapsicum species group (S. diphyllum, S. kleinii, S. malacothrix, S. pseudocapsicum, S. spissifolium). Fig. 19D-F Small shrubs, often with woody tap roots, often growing in dryareas;young stems and leaves glabrous or densely pubescent,the trichomeseither simple and uniseriateor dendritic.Sympodialunitsdifoliate,geminate, the minor leaves similar in shape to the major leaves, but usually smaller.Leaves lanceolateto elliptic,glabrousto denselypubescent,thetrichomeslikethose ofthe stems,theapexandbaseusuallyacute.Infloresences opposite the leaves, usually simple; pedicel scars FIG. 26. Distribution of Solanum venosum. closely spaced. Buds globose to obovoid, the corolla somewhatexertedfromthe calyx. Pedicels at anthesis deflexed and the flowers nodding.Flowers white, the QCNE); Tulcan-Tufifio-Maldonado rd., 3 km from calyx lobes leafy andrelativelylarge,the corolla lobes Tufino, 2900 m, 10 Oct 1986, Zak 1370 (NY). COTO- and sepals reflexed at anthesis.Fruit an orange or red PAXI:Pilalo-Latacunga rd., 3400 m, 0?57'S, 78?58'W, 6 soft andjuicy at maturity;fruitingpedicelserect, berry, Jul 1968, Holm-Nielsen & Jeppesen 1406 (NY). IMBAand woody. Seeds flattened-reniform,pale thickened, BURA:Cant6nOtovalo, ParroquiaSan Luis de Quichinche, the surfacespitted. yellowish-brown, San Alberto,3580 m, 0?12'S, 78?30'W, 11 Jul 1989,Moran et al. 46 (QCNE); Ibarra-paramode Jora-Cruz rd., 28003000 m, 19 Mar 1988, Zak 3436 (NY); rd. from Ibarra to paramo de Veracruz, 2800-3000 m, 0?25'S, 78?00'W, 19 Mar 1988, Zak & Jaramillo 3436 (BM, MO, QCNE); Otovalo-Mojanda Casas rd., detour to Haciendi Rubi, 3150 m, 28 Aug 1987, Zak 3508 (AAU, K, NY). PICHINCHA:Hacienda Mi Cielo, 3000 m, 22 Jul 1977, Jaramillo 10 (NY); rd. to Hacienda Mi Cielo, NW slopes of Volcan Pichincha, 3400-3600 m, 0?10'S, 78?30'W, 24 Apr 1987, Zak & Jaramillo 2020 (MO, NY); Quito-Hacienda Mi Cielo Rd., NW slopes of Volcan Pichincha,34003600 m, 20 Apr 1987, Zak 2020 (NY). TUNGURAHUA: San Isidro, cultivated near houses, 3430 m, 1?18'S, 78?45'W, 29 Dec 1982, Brandbyge & Herrera 42034 (U). Local names. Ecuador. Imbabura:furafango, digala. Tungurahua: pululu.

Distribution. Widespreadin drierhabitats,Mexico to Uruguay. Members of the Solanumpseudocapsicum group have traditionallybeen considered as not particularly closely relatedto other species of sect. Geminata(see History;Morphology). The membersof the group are distinguishedby having orangeor redberriesborneon erectfruitingpedicels, flattenedlargeseeds and(often) copious dendritictrichomes.Both S. diphyllumand S. pseudocapsicum are very successful weeds throughout the tropics and have also been cultivated as ornamentals. All of these species are small shrubs, often with woody tap roots, and may persist throughdry or otherwise unfavorableperiods.

55

TAXONOMIC TREATMENT

Key to the species of the Solanumpseudocapsicum species group 1. Mature leaves and stems completely glabrous. 2. Leaves elliptic-lanceolate; 1-2.5 cm diam.; fruit red-orange. Usually cultivated or escaped .................................................................................................................................................. 8. S. p seudocapsic ur 2. Leaves elliptic; corolla 0.7-1 cm diam.; fruit bright orange. Mexico to Costa Rica.............. 5. S. diphyllurn 1. Mature leaves and stems with at least some pubescence. 3. Trichomes simple and uniseriate. Mexico ................................................................................. 7. S. malacothrix 3. Trichomes dendritic. 4. Leaves lanceolate; sympodial units plurifoliate. SE Brazil .................................................. 9. S. spissifolium 4. Leaves otherwise; sympodial units difoliate, geminate. 5. Pubescence reddish-brown, the trichomes 1-2 mm long, covering the entire plant............. 6. S. kleinii 5. Pubescence golden, variously distributed,the trichomes 0.25-1mm long............. 8. S. pseudocap.Licum

5. Solanum diphyllum L., Sp. P1. 184. 1753. Type. America (?), (lectotype, LINN 248.5, designated Figs. 19D,E, 27 by Knapp& Jarvis, 1991). Small straggly shrubs 1-2 m tall; stems glabrous or occasionallyminutelypuberulentwith minutewhite uniseriatetrichomesca. 0.1 mm long, the stemswinged fromthe decurrentleaf bases;barkdarkmaroonbrown, the older stems white lenticellate. Sympodial units difoliate,geminate.Leaveson non-reproductiveshoots andlowerbranchesglabrous,oblanceolate,10-15 x 2.54 cm, the apex acute, the base attenuate;petioles 4-5 mm long, winged fromthe decurrentleaf bases; leaves on the reproductiveshoots glabrous,elliptic to oblong, geminate, widest at orjust distal to the middle; major leaves 4.7-6.8 x 2-2.2 cm, with 4-5 pairs of main lateralveins raisedabove, prominentandpalerbelow, the apex roundedor occasionally acute, the base acute to attenuate,decurrenton the stem andpetiole;petioles25 mm long; minor leaves ovate to obovate, 0.9-2.5 x 0.6-1.4 cm, the apex rounded,the base acuteto attenuate, decurrenton the petiole; petioles 2-3 mm long.Inflorescences opposite the leaves, simple, glabrous,or occasionally minutely glandular,0.3-1.2 cm long, 520-flowered;pedicel scars closely spaced,butnot overlapping.Buds globose, white when young, becoming lavendarandmoreobovoidjustbeforeanthesis.Pedicels at anthesis ca. 5 mm long, taperingfrom the constrictionjust below the calyx tube to a slenderbase ca. 0.5 mm diam.Flowers with the calyx tube ca. 1 mm long, withtwo constrictions, one atthejunctionwiththepedicel and the otherjust below the lobes, the lobes ca. 1 mm long, deltoid,glabrous;corolla white, often tingedwith lavendar,0.7-1 cm across, lobed 3/4 of the way to the base, the lobes reflexed at anthesis,the interpetalarsinuses thin, extendingto the tips of the lobes, the tips of lobes minutely papillose; anthers 1.5-2 x 1 mm, poricidalat the tips, theporesteardropshaped;freeportion of thefilaments ca. 0.5 mm long, the filamenttube ca. 0.1 mm long or less; ovari'glabrous;style 3-4 mm

long, straight,becoming curvedwith age; stigma minutelycapitate,papillose.Fruita globose berry,slightly bilobed from the two locules, especially when young, 0.7-1.2 cm diam.,greenandhardwhenimmature,bright orange and fleshy when ripe;fruitingpedicels

erect,

woody, ca. 1.2 cm long, with a constrictionbetweenthe pedicelandcalyx tube,ca. 1 mm diam.atthebase.Seeds ca. 3 x 2.5 mm, the pale yellow-tan,flattened-reniform, surfacesminutelypitted,the marginsincrassate,paler than the body of the seed. Chromosome number. n = 12 (voucher Knapp 6169, BH 85:234; D'Arcy, 1969).

Distribution (Fig. 28). Mexico to Pacific Costa Rica in drierlowland areasat 0-250 m. Escapedfrom cultivationin manytropicalandsubtropicalareas:e.g., Florida,Java,andthe Antilles. Selected specimens examined. (No cultivated paleotropical specimens are cited). MEXICO. S.loc., 1848-9, Gregg 1008 (MO); s.loc., 1865-66, Hahn s.n. (K); s.loc., s.coll. (MO 276242). CAMPECHE:Campo Forestal Experimental Tropical El Tormento, km 5 of Escarcega-Candelaria rd., 4 Jan 1966, Chavelas P. ES1243 (MEXU). CHIAPAS:1 km N of town of Matamoros, 5 m, 25 Aug 1988, Martinez y Day & TriujilloE. 123 (MEXU); Mojarra, Tonala, 27 Nov 1947, Matuda 17'211 (MEXU); vic. of Tonala, 100 m, 16 Aug 1949, Mattda 18765 (F, MEXU); Venustiano Carranza, above Finca Carmen on rd. from Acala to Fugiltik, 1800 ft, 7 Nov 1967, Ton 3219 (WIS). COLIMA:Manzanillo, 1/2 mi SE of town, 10 m, 8 Aug 1938, Eyerdam & Beetle 8719 (US); Salagua, vic. of Manzanillo, 1 Dec 1925, Ferris 6218 (US); SW outskirts of Colima, 19 May 1973, Hansen et al. 1458 (BH, LL, WIS); Manzanillo, 18 Jul 1932, Howell 10294 (US); Colima, 30 Jun 1892. Jones 341 (MO, US); km 292, 15 ESE of Manzanillo, ca. 0 m, 23 Jul 1957, McVaugh 15681 (US); Colima, Jul 1897, Palmer 34 (US); Manzanillo, 1-31 Dec 1890, Palmer 1001 (K, MO, NY, US); Santiago near Manzanillo, 510 m, 20 May 1939, Stork et al. 25402 (US); Manzanillo. Aug 1866, Theibaut 1036 (P). GUERRERO:Coyaca, 11 Feb 1934, Hinton 6909 (NY, US). HIDALGO:Between Huejutla and Chililuco on rd. to Ixcatlan. 600 ft, 21 May

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loo

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28. Distribution of Solanum malacothrix (open circle) and S. diphyllum (solid circles). FIG.

Coalcoman, Villa 1947, Moore 2925 (BH). MICHOACAN: Victoria, 500 m, 7 Oct 1939, Hinton 13903 (LL, MO, NY, US); Coalcoman, Aquila, 29 Mar 1941, Hinton et al. 15869 (G, LL, NY, US, W), 220 m, 4 Aug 1941, Hinton et al. 15950 (G, LL, NY, US, W); delta of Rio Balsas, 6 Jun 1898, Langlasse 205 (P, US). MORELOS: S side of Cuernavaca, 24 May 1973, Hansen et al. 1489 (US, WIS); Xochitepec, Sep 1933, Lyonnet & Elcoro 1212 (US); valley near Jojutla, 3000 ft, 12 May 1901, Pringle 9410 (US). NAYARIT(by rd.) E of Ciudad Valles on Mex. Hwy. 110, 100 m, 21?58'N, 98?53'W, 23 Mar 1976, Hansen et al. 3853 (LL, U, US, WIS); San Dieguito, 13-16 Jun 1904, Palmer 109 (MO, NY, US); Tamasopo, Sierra Madre Oriental, 400 m, 8-9 Aug 1934, Pennell 17986 (NY, US); 3 km W of Huichihuayan, 250 m, 3 May 1959, Rzedowski 10455 (WIS). TABASCO:1 km NW of Paraiso, on rd. to Barra de Topilco, 21 Nov 1987, Cabrera & Cabrera 14755 (MEXU); N slope of Cerro las Campanas, 3 km E of Teapa, ca. 50 km S of Villa Hermosa, 50-100 m, 9 Aug 1974, Conrad et al. 2795 (MO); Paraiso, km 45 on rd. from Paraiso to Berra de Rupilco, 150 m, 14 Jun 1979, Cowan 2307 (NY); Teapa, along Rio Teapa (Rio Morelia) at Teapa, 19 Sep 1944, Gilly Sr. & Herncndez X. 270 (TEX, WIS); Rio Puyacatengo, rd. to Rancho San Antonio, 14 Feb 1984, Rica A. & Martinez S. 768 (MEXU); 22 mi W traffic circle in Villa Hermosa along Mex. 180, 20 Jul 1971, Spellman et al. 117 (MO). TAMAULIPAS: Tampico, 1827, Berlandier 141 (G-DC, US); Cafion de Galeana, Aug 1938, LeSueur 422 (TEX); along Rio Guayalejo about 1 mi downstream from Colonia Mirador, 24 Dec 1970, Taylor & Taylor 7220 (NY); Tampico, 9 Apr 1931, Viereck 1101 (US); Hacienda Chamal, 25 Jun 1919, Wooton s.n. (US). VERACRUZ:S.loc., Bonpland 4488 (P); s.loc., 16 Jul 1860, Gouin 20 (P); Otatitlan, Playa de Vaca, 15 m, 31 Mar 1967, Calder6n 1344 (MO, NY,

WIS); El Progreso, 70 m, 29 Jul 1969, Calder6n 1958 (MO, NY); INIREB field station at El Morro de la Mancha, 2.4 km E of Hwy. 180 on shore of Golfo de Mexico, ca. 68 km by rd. N of San Jose Cardel and 120 km by rd. S of Nautla, 10-15 m, 19?39'N, 96?25'W, 20 Jul 1978, Cochrane et al. 8596 (BH, TEX, WIS); above San Jose de Gracia, 1 mi S of hwy. between C6rdoba and Veracruz, 750 m, 28 Jun 1977, Croat 39593, 39627 (MO); Alto Lucero, Boca Andrea, 10 m, 19?47'N, 96?25'W, Dorantes 396 (NY); Tezonapa, 240 m, 20 Jun 1973, Gandara & Dorantes 58 (BH); around Jalcomulco, 1?23'N, 96?45'W, 25 Jul 1973, Gdndara & Dorantes 114 (NY); Medellin (?), 25 May 1866, Hahn s.n. (P); Xochicalco, 15-30 Jun 1866, Hahn s.n. (P); Tlacalpan, 23 Apr 1866, Hahn 144 (P); 7 mi NW of Naranjos, 1/2 km W of Mex. Hwy. 180 at the Brecha microwave station, 150 m, 10 Jun 1973, Hansen et al. 1790 (US, WIS); El Mirador, rd. to Yecuatla, 490 m, 28 Apr 1976, Herndndez A. 211 (BH); along the TransIsthmian hwy. (rt. 185) 3 km NE of Minatitlan, 50 m, 1 Aug 1958, King 1015 (TEX, WIS); along rt. 150, just NW of Cuitlahuac, 6 Jun 1960, King 2670 (NY, TEX, US); Laguna Tamiahua, 50 mi S of Tampico, 3 May 1939, LeSueur 549 (TEX); Pital, Nov 1841, Liebmann 1427 (US); ca. 10 km E of C6rdoba on rt. 150, 2000 ft, 24 Mar 1970, Long & Burch 3106 (MO); 11.5 km from Plan de las Hayas, 700 m, 24 Jun 1972, Lot et al. 2021 (MO); Sta. Lucrezia, 28 Jul 1936, Matuda 578 (LL, MO, US); Cuitlahuac, 16 May 1937, Matuda 1437 (MO, US); Dos Rios, 14 Mar 1930, Mell 502 (NY, US); Chalma, 3 km NE of Huejutla on rd. to Platon Sanchez, 250 m, 2011'N, 9?23'W, 22 Jun 1980, Nee & Hansen 18415 (BH); rocky floodplain of Rio Jalcomulco, 400 m, 19?20'N, 96?45'W, 29 Oct 1981, Nee & Castillo C. 22453 (BH); Salto de Eyipantla 5 air km S of San Andr6s Tuxtla, 150 m, 18023'30" , 95?12'30"W,4 Dec 1981, Nee 23643

FLORA NEOTROPICA

58 (BH); ca. 6 km by air S of Tiapacoyanon rd. to Altotonga, 800 m, 19?55N, 97?13'W,11 Jul 1982, Nee & Diggs 24868 (BH, F); Tlacotalpan, 21 May 1894, Nelson 502 (US); vic. of Pueblo Viejo 2 km S of Tampico, 10-25 Feb 1910, Palmer 398 (MO, NY, US); Rio de la Ternera,Oct 1918, Purpus 8208 (NY, US); Plan del Rio, 350 m, 31 Jul 1967, Rosas R. 620 (WIS); Coetzala, rd. to Coetzapotitla, 680 m, 25 Apr 1976, VelasquezL. 227 (K, MO); rt. 180 near jet. with rd. into Catemaco, 24 Jun 1966, Windler & Smith 1013 (MO). YUCATAN: S.loc., Johnson s.n. (NY). GUATEMALA. ALTA VERAPAZ: Margin of swamp below Pantin, 1575 m, 4 Apr 1941, Standley 96018 (F). ESCUINTLA: San Jose, 0 m, 25 Feb 1905, Kellerman 4569 (MEXU, WIS); near San Jose, 0 m, 30-31 Jan 1939, Standley 64167 (US). PETEN: Dolores, km 85, W of Machaquila rd., 22 Jul 1961, Contreras 2649 (MEXU). BELIZE. Burrel Boom, thicket near ferry, 9 Jun 1973, Dwyer 11059 (MO, mixed collection with Solanum nudum). HONDURAS. La Lima, Sep 1929, Johansen 46 (US). COMAYAGUA:Near Rio Comayagua, 15 Oct 1970, Herntndez M. 5369 (MO); Rio Frio near La Libertad, 300 m, 19 May 1956, Molina R. 7002 (US); Rio Humuya W of Comayagua, 650 m, 23 Jun 1964, Molina R. 14314 (NY); Ojos de Agua, banks of Rio Humaya, 30 km N of Comayagua, ca. 350 m, 8 Jan 1981, Nelson et al. 6921 (MEXU). CORTES: Cordillera de Omoa, slopes of San Juan Linfo near Santa Ana, 200 m, 1 Dec 1950, Molina R. 3496 (US). SANTA BARBARA: 20 km NE of Nueva Arcadia on Hwy. 18, 13 Aug 1970, Harmon & Dwyer 3808 (MO); Sesecapa River, near Santa Barbara, 500 m, 22 Aug 1968, Molina R. 22010 (NY). EL SALVADOR. Coastal region, 1923, Choussyi 1602 (US). LA UNION: Laguna de Maquique, 60 m, 18 Vic. Feb 1922, Standley 20974 (NY, US). LA LIBERTAD: of La Libertad, 0 m, 13 Apr 1922, Standley 23211 (NY, US). SAN MIGUEL: Vic. of San Miguel, 110 m, 24-27 Feb 1922, Standley 21104 (NY, US). SAN VICENTE:Vic. of San Vicente, 350-500 m, 2-11 Mar 1922, Standley 21160. 21307 (US). SONSONATE:Izalco, Jul 1923, Calder6n 1700 (NY, US); vic. of Nahulingo, 220 m, 21 Mar 1922, Standdley22011 (US); vic. of Izalco, 19, 24 Mar 1922, Standley 22219 (US). Near Rio Chiquito, El NICARAGUA. CHINANDEGA: Viejo, 27 Dec 1969, Atwood 2644 (MO, WIS); E base of Volcan Coseguina, Jul 1932, Howell 10278 (US); Ameya, 0 m, 19-21 Jul 1923, Maxon et al. 7147 (NY, US); near Rio Chiquito, El Viejo, 27 Dec 1969, Narvdez S. 2652 (MO, NY). LEON: Weeds around Le6n, 20-21 Dec 1975, D'Arcy 10425 (MO). COSTA RICA. GUANACASTE: Culebra, 26 Mar Rd. to Nicoya, 1838, Barclay 2172 (US). PUNTARENAS: Jan 1900, Tonduz 13666 (P). Local names. Mexico. Campeche: tomatillo. Solanum diphyllum is most similar to S. malacothrix, a rare species from western Mexico. The two species share small flowers with reflexed petals at anthesis and berries borne on erect pedicels. Solanum diphyllum differs from S. malacothrix in its glabrous foliage. The mature fruit of S. malacothrix is not known (see discussion under that species).

Solanumdiphyllumis nativeto Mexico andCentral indrierhabitats.Ithasbeen America,whereitis widespread widely cultivated as a foliage shrub,and for its pretty flowers andbrightlycoloredfruits,in tropicalandsubtropicalareaswhere it is hardysuch as Italy, southern France,andFlorida(D'Arcy,1974),theAntilles,Java,and the Philippines,andhas escapedin some of theseareas. Solanumdiphyllumis anattractiveshrub,withitstiny white-purpleflowers andbrightorangefruits.It is selfcompatible,butnotautogamous.All theflowersarelongstyled and the plants set abundantfruitwhen exposed to pollinators.In cultivationthe plantsare bushy,with manyleaves. In theirnativeMexico however,shrubsof S. diphyllumareoftenquitestraggly(Nee, pers.comm.). In Mexico andCentralAmericaSolanumdiphyllum is remarkablyuniformmorphologicallyandis usually easily recognizable.A single geographicalvariantoccursin Pacific coastalChiapas.In the municipalitiesof Arriaga and Cintalapa, near the Mar Muerto on the Pacific coast andsomewhatinland,plantsof a narrowleaved race of S. diphyllumare common. Plants from this areahave leaves thataverage1 cm wide, butotherwise theyconformto theothercharactersofS. diplhyllum. Naming this minorvariantwould obscureits relationship to the rest of the species range. The type specimenof Solanumdiphyllumin LINN is annotated on the back "Solanum lignosum affic. sempervirens,cassinaefoliis. Br."in Linnaeus'shandwriting (Savage, 1945). This is perhaps what led D'Arcy (1974) to speculatethatthe sheetwas collected by PatrickBrowne, as Linnaeusdenotedspecimenshe received fromBrownewith a Br.On these sheets however,the specificnameswerewrittenby Solander(Jackson, 1912).Thetype comes fromthe Cliffordherbarium (Knapp& Jarvis,1991;Savage, 1945).Theprintedvase at the base of the specimen is a characteristicmarkof the sheetsfromthatherbarium.Solanumdiphyllumwas included in the Hortus Cliffortianus(Linnaeus, 1737) as "5. Solanum caule inerme perenni, foliis ovatolanceolatisgeminis,alterominimo."Theprovenanceof the plant was cited as either America or Africa, but LinnaeusemphasizedAmerica,so it is clearhe thought (correctly)thatthe plantwas from America.Solanum diphyllumwas probablycultivated in Clifford's garden;the seeds areeasy to grow andeven in Ithaca,New York(Bailey Hortoriumgarden),and Londongrow to reproductivematurityin one summer. 6. Solanum kleinii L. B. Sm. & Downs, Phytologia 10: 429. 1964. Type. Brazil. SantaCatarina:Mun. CampoAlegre, 4 km S of CampoAlegre on the rd. to Jaraguado Sul, 900-1000 m, 6 Nov 1956, Smith & Klein 7321 (holotype, US; isotypes, HBR-n.v., NY, R-n.v.). Fig. 29

TAXONOMIC TREATMENT

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Shrubs or small subshrubs, ca. 0.5 m tall; young stems and leaves densely pubescentwith dendritictrichomes 1-2 mm long, these dryinggolden or reddish; barkof older stems darkbrown, not glabrate.Sympodial units difoliate, occasionally geminate.Leaves elliptic, widest at the middle, with 4-8 pairsof main lateralveins, glabrousand shiningwith simple uniseriate trichomesalongthe midribadaxially,sparselyto densely pubescentabaxially,the trichomesdendritic,1-2 mm long, like those of the stems; lamina 3-8 x 1.5-3 cm, the apex acute,the base acuteto attenuate;petiole 0.41 mm long, densely pubescentwith long dendritictrichomes like those of the stems and leaves. Inflorescences oppositethe leaves or intemodal,0.5-2 cm long, simple, with 3-5 flowers, densely pubescentwith dendritictrichomes;pedicel scars closely spaced,not overlapping.Buds ellipsoid, not includedin the calyx tube. Pedicels at anthesis nodding, slender, 1.5-2 cm long, ca. 0.5 mm diam. throughout,densely pubescent like therestof the inflorescence.Flowerswith thecalyxtube conical, ca. 2 mm long, the lobes long-triangularwith roundedapices, ca. 3.5 mm long, densely pubescent withdendritictrichomes;corollawhite,2-2.3 cm diam., lobed ca. 3/4 of the way to the base, the lobes planarat anthesis, the abaxial surfaceof the lobes densely papillose, the papillaeoccasionally branched;anthers4-5 x 1.5-2 mm, sagittate at the base (in type specimen), poricidalat the tips,the poresteardropshaped;freeportion of thefilamentsca. 0.5 mm long, the filamenttube ca. 0.5 mm long, glabrous;ovary glabrous or with a few dendritictrichomesat the apex;style straight,5-6 mm long,glabrous;stigmacapitate,thesurfaceminutely papillose. Fruit and seeds not known. Chromosome numbernot known.

FLORA NEOTROPICA

300

500 FIG. 30. Distribution of Solanum kleinii (solid circles) and S. spissifolium(open circles).

lectionsof S. kleiniifromSantaCatarinaarefroma single population.Both collections from SantaCatarinahave woody tap roots attached.This habit is common in the S. pseudocapsicumgroup and may be associated with periodicallydry habitats. 7. Solanum malacothrix S. Knapp,Brittonia38: 89. 1986. Type. Mexico. Guerrero:Distr.Mina, Aguazarca, 5 May 1937, Hinton et al. 10418 (holotype, NY; isotypes, K, LL, MEXU, U, US). Fig. 31

Shrubs2-3 m tall; young stems and leaves densely with soft uniseriate trichomes ca. 0.5 mm Distribution (Fig. 30). In Parana,Santa Catarina pubescent older stems remainingpubescent, not glabrate, long; and Rio Grandedo Sul in southeasternBrazil, in open grayish-brown.Sympodialunitsdifoliate,geminate,the areas at 1000-1125 m. minor leaf often deciduous. Leaves narrowly ovate, Specimens examined. BRAZIL. PARANA:Guara- widestjust proximalto the middle, sparselypubescent puava, Serrada Esperanca, 18 Oct 1960, Hatschbach 7384 above with uniseriate trichomes, these denser on the (US). SAO PAULO: Bananal, Serra da Bocaina, Sertao veins, densely pubescent below with soft uniseriate do Rio Vermelho, 1200 m, 5-6 Oct 1949, Brade & trichomes0.5-0.75 mm long, the epidermisbarelyvisDuarte 20138 (BM, RB). RIO GRANDEDOSUL:Aparados ible in dry specimens;majorleaves 10-16 x 2.5-5 cm, da Serra, 1300 m, 24 Oct 1961, Pabst 6281 (Pereira with 11-14 pairsof main lateralveins, these raisedand 6454) (UPCB). SANTACATARINA: Campo Alegre, 1000 pubescentabove,prominentandyellowish beneath,the m, 17 Oct 1957, Reitz & Klein 5142 (GH, NY, US). apex acuminate,the extreme tip rounded,the base atSolanum kleinii is closely related to Solanum tenuate;petioles channeledabove, 7-8 mm long; minor leaves differing from the majorones only in size, pseudocapsicum, and differs from that species in its more copious, longer dendriticpubescence and larger 6.5-11 x 1.5-3.5 cm, the apex acuminate,roundedat flowers. Fruitsof S. kleinii are not known, but I expect the very tip, the base attenuate;petioles 6-8 mm long. them to be orange-redat maturity,with flattened-reni- Inflorescences opposite the leaves, simple, 1-1.5 cm formseeds. The pubescenceof Hatschbach 7384 from long, 10-30-flowered, pubescentalong the rachiswith Parana is smaller and less dense than that of specimens uniseriatetrichomeslike those of the stems and leaves; from SantaCatarina,but this is probablybecause col- pedicel scars closely spaced,butnot overlapping,occa-

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FLORA NEOTROPICA

sionally ca. 0.5 mm apart.Buds globose, the corolla soon exserted from the calyx and the buds then ovoid to ellipsoid.Pedicels atanthesisdeflexed,8-9 mm long, taperingfrom the calyx tube to a slender base ca. 0.5 mm diam., with a few uniseriatetrichomesca. 0.5 mm long. Flowerswith the calyxtubebroadlycup-like,ca.1 mm long, the lobes broadly deltoid, 0.5-1 mm long, withtuftsof uniseriatetrichomeson thetips of the lobes, otherwisesparselypubescentwith uniseriatetrichomes like therestof the inflorescence;corollaprobablywhite, 0.8-1 cm diam., lobed nearlyto the base, the lobes reflexed at anthesis, the tips and margins of the lobes minutely papillose; anthers ca. 2 x 1 mm, poricidalat the tips, the tips paler andslightly thickened,the pores teardropshaped; free portion of thefilaments ca. 0.5 mm long, the filament tube ca. 0.25 mm long; ovary glabrous;style straight,ca. 5 mm long; stigma a minutely papillose area at the tip of the style. Fruit and seeds not known;fruiting pedicels erect in immature fruit. Chromosomenumbernot known. Distribution (Fig. 28). In the northwesterncomer of the state of Guerrero,on the slopes of the Sierra Madredel Sur in the Rio Balsas drainage,in tropical deciduous forest or dry pine-oak woodland ("bosque tropical caducifolio" or "bosque de Quercus" of Rzedowski, 1978). Selectedspecimensexamined.MEXICO.GUERRERO: Montesde Oca, San Antonio,6 Jan 1937,Hintonet al. 10248 (BM, DS, MEXU,NY, U, US). Solanum malacothrix is most closely related and superficiallyquitesimilarto S. diphyllum,a widespread lowlandMexicanspecies. Solanummalacothrixis distinctive in its long, soft leaf pubescence, which is unusual in the section; while S. diphyllumis completely glabrous.Solanumlasiocladumof Venezuelaalso possesses this type of leaf pubescence. It differs fromS. malacothrixin its apiculatecalyx lobes, largerflowers, and long-petioled leaves with acute apices, and I have not assigned it to any of the species groups. The maturefruitsof S. malacothrixarenot known, but in view of its overall similarityto S. diphyllumI would expect them to be brightly colored, borne on erect pedicels, and ratherfleshy. 8. Solanum pseudocapsicum L., Sp. P1.2: 184. 1753. Type.Madeira.Cultivated,Anon.(lectotype,LINN 248.4, designatedby D'Arcy,1973;confirmedKnapp & Jarvis, 1991, [BH neg. 6792]). Figs. 19F,32 Solanum diflorum Veil., Fl. Flumin. 84. 1829 [1825]. Type. Brazil. Rio de Janeiro: "Habitat campis apricis mediterraneis prope Praedium Boavista" (No specimens extant; lectotype, Vellozo, Fl.

Flumin. Icones 2: fig. 102. 1831 [1827], designated by Morton, 1976). Solanum capsicastrum Link ex Schauer, Allg. Gartenzeitung 1: 228. 1833. Type. Brazil. Specimen not traced. Solanum hygrophilum Schltdl., Linnaea 8: 254. 1833. Type. Mexico. Jalapa, May, Schiede 137 (holotype, B [destroyed]; lectotype, NY, here designated; isolectotypes, G-n.v., GOET-n.v., MO-n.v., P-n.v.). Solanum eremanthum Dunal in DC., Prodr. 13(1): 129. 1852. Type. Brazil. Provincia Rio Grande, 1833, herb. imp. du Bresil, Gaudichaud 619 (holotype, P; isotype, MPU). Solanum ulmoides Dunal in DC., Prodr. 13(1): 130. 1852. Type. Mexico. Pav6n s.n. (holotype, G (Boiss.); isotypes, G [Delessert, F. neg. 34130], M [F neg. 48351], frag. MPU, F). Solanum karstenii Dunal in DC., Prodr. 13(1): 151. 1852. Type. Venezuela. (as Colombia), Karsten s.n. (lectotype, G-DC, here designated (F neg. 6775); isolectotypes, B [destroyed: F neg. 2665], frag. F, G [Morton neg. 8602], frag. MPU). Solanum tucumanense Griseb., Abh. Konigl. Ges. Wiss. Gottingen 24: 254. 1879. Type. Argentina. Tucuman: La Cruz, Lorentz & Hieronymus 49 (lectotype, CORD, designated by Morton. 1976; isolectotype, GOET). Solanum validum Rusby, Mem. Torrey Bot. Club 4: 230. 1895. Type. Bolivia. Cochabamba:Vic. Cochabamba, 1891, Bang 972 (lectotype, NY, here designated; isolectotype, MO, NY, US-n.v.). Solanum diflorum Veil. var. angustifolium Kuntze, Rev. Gen. P1. 3: 225. 1898. Type. Argentina. "Sierra chica de Cbrdoba, Galander, Hieronymus. Uruguay." Not located. Solanum jaliscanum Greenm., Proc. Amer. Acad. Arts 34: 571. 1899. Type. Mexico. Jalisco: Barranca of Guadalajara, 4000 ft, 10 Jun 1898, Pringle 6870 (lectotype, GH, here designated; isolectotypes, BM, CM-n.v., F, GOET-n.v., MEXU, MO, NDG-n.v., NY, PH, UC, US). Solanum diflorum var. pulverulentum Chodat, Bull. Herb. Boissier 1902: 811. 1902. Type. Paraguay. Cordillera: Sapucay, Hassler 1653 (lectotype, G, here designated (Morton neg. 8631); isolectotypes, BM, K, NY, P). [originally and erroneously described as Solanum diphyllum rather than diflorum, later corrected, see Morton 1976]. Solanum dunnianum H. L6v., Repert. Spec. Nov. Regni Veg. 9: 324. 1911. Type. China. Kweichau: Jardin de H. Ke-Chou, et abondant aux rochers de Ting-Mei, Jun 1905, Esquirol 536 (holotype, E, photo A, fide D'Arcy, in litt.). Solanum plurifurcipilum Bitter, Repert. Spec. Nov. Regni Veg. 11: 15. 1912. Type. Mexico. Veracruz: Pacho forest near Jalapa, 4000 ft, 4 Apr 1899, Pringle 8070 (holotype, Vratislava-n.v.; isotypes, BM, F, GH, NY, US).

TAXONOMIC TREATMENT ?

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Solanum ipecacuanha Chodat, Bull. Soc. Bot. Geneve, ser. 2, 8: 158. 1916. Type. Paraguay. Cordillera: Cordillera de Altos, Hassler 1402 (lectotype, G, here designated [Morton neg. 8513]; isolectotypes, BM, K, NY). Solanum ipecacuanha var. calvescens Chodat, Bull. Soc. Bot. Geneve, ser. 2, 8: 158. 1916. Type. Paraguay. Cordillera: Sapucay, Hassler 1653 (lectotype, G, here designated [Morton neg. 8631]; isolectotypes, BM, K, NY, P). Solanum ipecacuanha var. obovata Chodat, Bull. Soc. Bot. Geneve, ser. 2, 8: 158. 1916. Type. Paraguay. Cordillera:Arroyos y Esteros, Balansa 2100 (syntype, G); Paraguari,Balansa 2100a (syntype, G). Solanum capsicastrumLink var. caaguazuense Chodat, Bull. Soc. Bot. Geneve, ser. 2, 8: 159. 1916. Type. Paraguay. Caaguazu: Caaguazu, 19 Sep 1874, Balansa 2097 (holotype, G; isotype, K). Solanum pseudocapsicum L. forma pilulosum Hassl., Repert. Spec. Nov. Regni Veg. 15: 223. 1918. Type. Paraguay. Rojas 390 (holotype, G-n.v.). Solanum pseudocapsicum L. ssp. diflorum (Vell.) Hassl., Repert. Spec. Nov. Regni Veg. 15: 221. 1918. Type. Based on Solanum diflorum Veil. Solanum pseudocapsicum L. ssp. diflorum (Veil.) Hassl. var. typicumHassl., Repert. Spec. Nov. Regni

Veg. 15: 223. 1918. Type. Paraguay. Caaguazu: Caaguazu, Balansa 2097 (lectotype, G, here designated; isolectotype, K). SolanumpseudocapsicumL. ssp. diflorum(Vell.) Hassl. var. sendtnerianumHassl., Repert.Spec. Nov. Regni Veg. 15: 223. 1918. Type. Argentina. C6rdoba: Potrero de Moyano, Sierra Chica de C6rdoba, 28 Jan 1881, Galander s.n. (lectotype, G-DC, designated by Morton, 1976; isolectotype, NY). Solanum pseudocapsicum L. ssp. diflorum (Veil.) Hassl. var. hygrophilum (Schltdl.) Hassl., Repert. Spec. Nov. Regni Veg. 15: 223. 1918. Type. Based on Solanum hygrophilum Schltdl. Solanum pseudocapsicum L. ssp. diflorum (Veil.) Hassl. var. hygrophilumforma calvescens (Chodat) Hassl., Repert. Spec. Nov. Regni Veg. 15: 223. 1918. Type. Based on Solanum ipecacuanha Chodat var. calvescens Chodat, citing the syntype Hassler 1653. Solanum pseudocapsicum L. ssp. diflorum (Vell.) Hassl. var. ambiguum Hassl., Repert. Spec. Nov. Regni Veg. 15: 223. 1918. Type. Paraguay.Hassler 5711 (holotype, G-n.v.). Solanum mexiae Standl., Field Mus. Bot. 4: 261. 1929. Type. Mexico. Nayarit: Ojos de Agua, near Yxtlan del Rio, 1100 m, 23 Sep 1926, Mexia

64

FLORA NEOTROPICA

742 (holotype, F [F neg. 49441]; isotypes, ABrazil, Argentina, and Uruguay, from sea level to 2600 n.v., GH-n.v., MO, NY, UC-n.v., US). m. Widely cultivated throughout the world, often esSolanum pavimentiL. B. Sm. & Downs,Phytologia10: in and subtropical areas. 428. 1964.Type.Brazil.SantaCatarina: Lajeadinho, caped tropical 800 m, 3 Jan 1962, Reitz & Klein 11438 (holoSelected specimens examined. (No cultivated paleotype, US; isotypes, HBR-n.v.,NY, US). tropical specimens cited). MEXICO. HIDALGO: Rancho Smallshrubs,prostrateto 1 m tall;young stems and leaves glabrous to densely pubescent with dendritic trichomes 0.1-0.5 mm long; stems erect or prostrate, soon glabrous;barkof older stems pale golden brown. Sympodialunits difoliate, geminate.Leaves elliptic to narrowlyelliptic, glabrousadaxially,occasionallywith a few dendriticor simpleuniseriatetrichomesalongthe midvein, glabrousto densely pubescentwith dendritic (occasionallysimpleuniseriate)trichomesabaxially,the trichomesgolden when dry,the marginsoften crisped or irregular;majorleaves with 4-6 pairsof main lateral veins, 2.5-9 x 0.7-4.5 cm, the apex acute to rounded, the base acute;petiole 0.2-1 cm long; minorleaves differing from the majorsonly in size, 0.9-3.5 x 0.4-2.7 cm, the apex acute to rounded,the base acute;petiole ca. 0.2 cm long. Inflorescences opposite the leaves, simple, 0.2-1 cm long, with 1-8 flowers, in cultivated forms often with only 1 flower, glabrous to densely pubescent with dendritic trichomes; pedicel scars closely spaced, not overlapping, corky in fruit.Buds ellipsoid,thecorollaincludedin thecalyx lobes.Pedicels at anthesis stout, deflexed, 0.3-1 x ca. 0.5 mm diam., glabrousto denselypubescentwith dendritictrichomes like the stems and leaves. Flowers with the calyx tube conical, 1-2 mm long, the lobes long-triangularwith roundedapices or occasionally spathulate,1.5-4 mm long (to 6-7 mm long in some cultivated specimens), glabrousto densely pubescentlike the restof the plant; corolla white, 1-1.5(2.5 in cultivars)cm diam., lobed 1/2 to 3/4 of the way to the base, the lobes planarat anthesis,densely papilloseon abaxialtips andmargins; anthers3-4 x 1-1.5 mm, darkorange-redin live plants, especially in cultivatedplants,poricidalat the tips, the poresteardropshaped;freeportionof thefilaments0.51 mm long, the filamenttube0.5-1 mm long, glabrous; ovary-glabrous;style 5-6 mm long, glabrous;stigma small-capitate,the surfaceminutely papillose.Fruit a globose berry,green when immature,yellow to deep orange-red when ripe, the pericarpthin and papery; calyx lobes enlarged in fruit, 5-7 mm long;fruiting pedicels erect,woody, 0.8-1 cm long, 0.5-1 mm diam. at the base. Seeds pale tan, flattened-reniformwith incrassatemargins,3-4 x 2.5-3 mm, the surfacesminutely pitted, testal cells long-rectangularin outline. Chromosome number: n = 12 (Gerasimenko & Reznikova,1968(as 2n = 24); Randell& Symon, 1976). Distribution (Fig. 33). In drier microhabitatsin Centraland South America, from Mexico to southern

Viejo, mun. La Mision, 28 Jun 1964, Gonzalez Quintero 1001 (F). JALISCO: Along Mexico 80 from Guadalara to Autlan, 1 km S of Mirador, halfway between Cocula and Tecolotlan,ca., 1800 m, 10 Jan 1979, Iltis & Nee 1402 (F, WIS); Barrancaof Rio Verde, ca. 20 mi N of Tepatitlan on rd. to Yahualica,ca. 1450 m, 27-28 Aug 1958, McVaugh 17425 (NY); Guadalajara, Jul-Oct 1886, Palmer s.n. (NY); Guadalajara, Jul-Oct 1886, Palmer 410 (NY); Tequila, Aug-Sep 1886, Palmer 410 (GH, US); Guadalajara,Jul 1886, Palmer 440 (GH); Barranca,near Guadalajara, 29 Jun 1889, Pringle 2909 (GH, US). MExIco: Temascaltepec, Cuentla, 1960 m, 20 May 1932, Hinton 669 (NY, US); Temascaltepec, Tejupilco, 1340 m, 11 Sep 1932, Hinton 1666 (NY, US); Temascaltepec, Tenayac, 1450 m, 9 Jun 1933, Hinton 4020 (NY, US). MICHOACAN: 1 km SW of Jacona, 1590 m, 16 Oct 1940, Cutler 4063 (F). NAYARIT: Ojos de Agua, near Yxtlan del Rio, 1100 m, 23 Sep 1926, Mexia 742 (F, US). OAXACA: Chiltepec, 16 m, 8 Jun 1967, Martinez Calderon 1419 (NY). PUEBLA: Pahuatan, 16 Dec 1942, Martinez s.n. (US). QUERETARO:S.loc., 1910-13, Bro. G. Arsene (Bro. Agniel) 10578 (US); gardens of Rinconada, 5 Jan 1947, Gregg 51 (MO). SANLUISPOTOSi: Tamasopo, Puerto Verde, 64 km W of Cd. Valles on hwy. to Rio Verde, 900 m, 24 May 1981, Fryxell & Anderson 3587 (NY); Tamazunchale, 24 Nov 1937, Kenolver 824 (F). TAMAULIPAS:Sierrade Guatemala,Rancho del Cielo Biological Station, ca. 3750 ft, 5 Jun 1971, Sullivan 315 (NY); Sierra de Guatemala, between Malacati turnoff and Julilo, on rd. to Rancho del Cielo Biological Station, 4200-4700 ft, 22 Jun 1971, Sullivan 559 (NY). VERACRUZ:Salto de Gato, E of Jalapa, 2 May 1973, Dorantes & Acosta 2031 (F); Cosamaloapan, Playa Vicente, 100 m, 18 Aug 1969, Martinez Calderon 1970 (F); vie. La Calavera, 10 km N of Altotonga (13 km by rd.), on rd. to Tlapacoyan, 1350 m, 19?51'N, 97?13'W, 28 Jun 1980, Nee & Hansen 18644 (F); 1 km above and NW of San Andr6s Tlalnehuayocan, 1700 m, 19?43'N, 96?58'W, 22 Mar 1983, Nee et al. 26177 (F); along Huayacocotla-Zontecomatlan rd., 1 km NE of San Antonio Ixtatetla, 1300 m, 20?42'N, 98?23'W, 27 Apr 1983, Nee & Taylor 26797 (F, NY); 4 km S of Playa Vicente, rd. to Nigromante, 12 Apr 1968, Nevling & Gdmez Pompa 741 (F); Jardin Botanico Clavijero, 1300 m, 19?30'N, 96?55'W, 23 Apr 1978, Ortega O. & Calzada 779 (F); Xalapa, 1300 m, 19?30'N, 96?55'W,22 Jun 1978, Ortega 0. 1357 (F); Alseseca, 1200 m, 9 Jun 1973, VenturaA. 8345 (F); Banderilla, 1500 m, 11 May 1974, VenturaA. 10054 (F); Piletas, San Miguel de Soldado, 1600 m, 22 Apr 1975, VenturaA. 11230 (F); Piletas. 2 km before coming to Banderilla on Perote-Xalapa rd., 1580 m, 10 Apr 194, Zola B. 325 (F). ZACATECAS:13 mi WSW of Jalpa, 6000 ft, 14 Aug 1969, Taylor & Tavlor 6112 (NY, US).

TAXONOMIC TREATMENT Qo^'V

it 'l

65

FIG. Pr

--

.0---------"'""'""'^

Distribution of Solaum

I red byHendnI

- -

----

seudocascum.

.

FIG. 33. Distribution of Solanum pseudocapsicum.

GUATEMALA.

CHIMALTENANGO:Finca La Alameda,

near Chamatenango, ca. 1830 m, cultivated, 11-22 Dec 1940, Standley 80125 (F). GUATEMALA: S.loc., 1939, Aguilar 82 (F). COLOMBIA. San Crist6bal, 2800 m, Nov 1911, Bro. Apollinaire & Bro. Arthur 39 (US); San Crist6bal, Feb 1911, Bro. Ariste-Joseph A330 (US). ANTIOQUIA: San Antonio de Pereira (Rionegro), Nov 1936, Bro. Daniel 971 (US); vic. Medellin, 15 Apr 1927, Toro 171 (NY); Valparaiso, vic. Medellin, 22 Jul 1928, Toro 1360 (NY). CUNDINAMARCA: Bogota,

Ciudad Universitaria,

2620 m, 20 Mar-20 Apr 1946, Duque-Jaramillo 3000 (NY); Bogota, Ciudad Universitaria, Jardin Botanico, 2600 m, 19 Mar 1943, Garcia Barriga 10820 (US); Bogota, 2640 m, Guevara Am6rtegui 26 (US). MAGDALENA: La Paz, 10 Sep 1938, Haught 2329 (A, F, K); near San Juan de Cesar, 200 m, 9 Jun 1944, Haught 4193 (F, GH, K, NY, US); Cerrej6n, ca. 200 m, 21 Aug 1949, Haught 6598 (US). NARINO:Popayan-Pasto rd., near border to Pasto, Gutierrez et al. s.n. (US). PUTUMAYO: Sibundoy, hills N of valley, ca. 2220-2270 m, 28 May 1946, Schultes & Villarreal 7504 (US). VENEZUELA. ARAGUA:Beach at Ocumare, Pittier 14031 (US); near Colonia Tovar, 1854-55, Fendler 990 Vic. Tocuyito, Jun 1941, Saer 807 (K, NY). CARABOBO:

(NY, US). DISTRITO FEDERAL: Caracas, toward El Valle, Dec 1920-Jan 1921, 3000-3500 ft, ca. 10?30'N, 7 Jan 1921, Bailey & Bailey 765 (BH, NY, US); vic. Cristobal Colon, 5 Jan-22 Feb 1922, Broadway 2 (GH, NY), Broadway 763 (GH, NY, US); El Carenero, Jun 1943, Cardona 553 (NY, US); around Caracas, 800-1000 m, 23 Jul 1921, Pittier 9643 (GH, NY); Laguna del Espina, near Caracas, 800-1000 m, 23 Jul 1921, Pittier 9643 (US); near Las Trincheras, Caracas-La Guaira rd., 6 Sep 1925, Pittier 11880 (A, NY, US). FALCON: Near Guaibaca, Mar 1962, Agostini S. 21 (US); Cerro Mampostal, 400 m, 11?27'N, 69017'W, 4 Aug 1977, Gonzalez 990 (MO); Cienaga de Tacarigua, ca. 5 km NW of Tocuyo de la Costa, 17 Apr 1971, Nee & Mori 3951 (US); El Degredo, near Curimagua, ca. 1000 m, 13 Sep 1980, Trujilloet al. 16829 (MY, NY). LARA:Sanare, 1358 m, May 1930, Saer 470 (F); 17-18 km S of Quibor toward Cubiro, 800 m, 9?52'N, 69?37'W, 5 Jul 1974, Steyermarket al. 110052 (F, VEN). MERIDA: San Esteban, 1893-94, Mocquerys s.n. (US); Timotes, 2000 m, cultivated, 23 Jan 1938, Pittier 12689 (NY). MIRANDA: Hacienda Cardenas, 1000 ft, 9 Sep 1929, Holt 599 (F). TACHIRA:Parque Cazadero, Quebrada Cazadero, 16 km NW of San Crist6bal, 400-650 m, 7054'N, 72?18'W, 2 May 1982, Liesner & Guariglia 11698 (MO, NY, VEN);

66 between Las Dantas and Las Adjuntas, 13 km NW of Rubio, 920-950 m, 7?43'N, 72025'W, 27 Jul 1979, Steyermark & Liesner 118839 (MO, NY, VEN). TRINIDAD AND TOBAGO. TRINIDAD: S.loc., s.coll. 1907 (US); s.loc., 19 Dec 1891, Broadway s.n. (NY); Monos, 27 Nov 1915, Broadway 7443 (NY). ECUADOR. AZUAY: Chullubamba, 10 km N of Cuenca, 2350 m, 7 Oct 1981, Dodson & Dodson 11617 (QCNE). LOJA: Cariamanga, 28 Jul 1946, Espinosa & Espinosa 673 (LOJA). PICHINCHA:Vic. Quito, Cotocollao, 2750 m, 17 May 1939, Asplund 6104 (K). cult. PERU. HUANUCO: 3 mi below Ambo, Tomaiquichua, 8500 ft, 19 Sep 1922, Macbride & Featherstone 2439 (F). cult. BRAZIL. S.loc., Gaudichaud 619 (fragment F); Campos de Jordao, Sep 1945, Leite 3624 (A); s.loc., Sellow s.n. (K); Rio Grande,Sellow 154 (GH). DISTRITO FEDERAL: Ca. 5 km SE of Brasilia on rd. to Belo Horizonte, 1000 m, 24 Feb 1966, Irwin et al. 13131 (NY). GoIAs: Rd. to Goiania, 29 Jan 1978, Heringer et al. 16759 (US). MATTO GROSSO: Nossa Senhora do Livramento, near margin of Rio Cotia, 15?46'S, 56?52'W, 6 May 1978, Macedo & Assumpcao 683 (NY); Sta. Liz. de Casalvaseo, Jan 1828, Riedel 1235 (NY). MINAS GERAIS: Caldas, Araujo s.n. (M.N. R de J. 25629) (US); middle slopes of Serra da Piedade, ca. 40 km E of Belo Horizonte, near BR-31, ca. 1800 m, 17 Jan 1971, Irwin et al. 30475 (F, K, MO, NY, US); Caldas, 31 Mar 1847, Regnell s.n. (US); Caldas, 11 Sep 1866, Regnell III 975 (US); Sitio, 3 Dec 1905, Sampaio 367 (US). PARANA: Campo do Tenente, 9 Oct 1908, Dusen 6855 (F, GH, K, MO, NY); Curitiba,7 Sep 1914, Jinsson 899a (GH, NY); Bocaiuva do Sul, Varginha do Carumbe, 23 Jan 1963, Hatschbach 9663 (US); Rio Ipiranga, 8 Oct 1969, Hatschbach 22396 (NY); Rio Negro, 23 Oct 1928, Hoehne 23139 (US); Mandirituba, Cachoiera and vic. to Rio Barigui,ca. 25?45'S,49?15'W,30 Nov 1981,Landrum 3898 (NY); Fazenda Santa Rita, ca. 65 km W of Curitiba on rd. to Ponta Grossa, ca. 25?25'S, 49?50'W, 2 Dec 1981, Landrum 3927 (F, NY); Campina Grande do Sul, Aracatuba, 28 Dec 1966, Stellfeld 1666 (UPCB), Stellfeld 1667 (UPCB, US). RIO DE JANEIRO: Macae, Frade, cult., 18 Oct 1970, Carauta 1223 (US). RIO GRANDE DO SUL: Monte Bonito, Pelotas, 19 May 1954, da Costa Sacco 144 (US); Santa Maria, 18 Nov 1970, Dobereiner & Tokarnia 749 (F); Sao Jer6nimo, CapaoConde rd., P6lo Carboquimico, 22 Sep 1982, Dutra 42 (F); vic. S. Leopoldo, Jul 1941, Eugenio 1713 (NY); s.loc., 1878-1879, Glaziou 11370 (K); Porto AlegrePantano Grande rd., km 17, 14 Dec 1972, Lindeman & Irgang 21046 (U); Vacaria, 14 km from Vacaria on rd. to Bom Jesus, 25 Oct 1961, Pabst 6324, Pereira 6497 (F, US); Guaiba, 30 Oct 1961, Pabst 6408, Pereira 6581 (US); Morro das Abertas, 14 Aug 1979, Soares 131 (F); CaCapuva,22 Sep 1960, Torgo 21254 (US); Farroupilha, Sao Jose, 21 Oct 1984, Wasumet al. 494 (US). SANTA CATARINA: 3 km W of Papanduva along BR-116 to Lajes, 810 m, 10 Mar 1976, Davidse et al. 11066 (MO, NY); Therval, 8 Jun 1911, Dusen 11927 (GH); Sio Jos6 do Cerrito, Lajes, 950 m, 31 Oct 1963, Klein 4315 (US);

FLORA NEOTROPICA Morro do Cambirola,Palhoca, 10 m, 18 May 1971, Klein & Bresolin 9429 (US); Campo dos Padres, 1800 m, 21 Dec 1948, Reitz 2668 (US); Bom Retiro, 950 m, 25 Oct 1957, Reitz & Klein 5461 (US); Vargem Grande, Lauro Miiller, 350 m, 11 Jul 1958, Reitz & Klein 6727 (US); Serra do Matador, Rio do Sul, 700 m, 1 Aug 1958, Reitz & Klein 6837 (US); Rio dos Pocos, Canoinhas, 750 m, 26 Oct 1962, Reitz & Klein 13594 (US); Bom Retiro, between Fazenda Campo dos Padres and Fazenda Santo Antonio, 1400-1650 m, 21 Nov 1956, Smith & Klein 7817 (NY, US); Riozinho, 1000 m, 22 Jan 1957, Smith & Reitz 10289 (K, US); near Xanxere, 700 m, ca. 26?52'S, 52?24'W, 13-14 Oct 1964, Smith & Reitz 12486 (GH, MO, US); Carapichinho, 22 km SW of Sao Joaquim, 1000-1100 m, ca. 28?25'S, 50?01'W, 6 Jan 1965, Smith & Reitz 14387 (NY, US). SAo PAULO: Estacao Alferes Rodrigues, Oct 1899, Edwall 4425 (US); Umuarama, Campos do Jordao, 23 Jan 1935, Kuhlmann s.n. (Hoehne herb. 32487) (US); Parereiros, 40 km from Sao Paulo, 800-850 m, 24 Aug 1979, Mizoguchi 953 (NY); Estaqao Sagrado do Caracao, Linha Sorocabana, 750-800 m, 3 Oct 1979, Mizoguchi 1040 (NY). BOLIVIA. S.loc., Bang s.n. (NY). COCHABAMBA: Brewery near Taquina, 9500 ft, 19 Jan 1949, Brooke 5123 (BM, F, NY); Chocaya, 2600 m, Mar 1939, Cdrdenas 731, 732 (US); 9 km W of Cochabamba, 2600 m, 15 Dec 1942, Cutler 7685 (F, US); 9 km W of Cochabamba, 2600 m, 15 Dec 1943, Cutler 7689 (F); Quillacolla, Bella Vista, 2600 m, 17?20'S, 66?20'W, 1 Dec 1982, Johns 82-60 (F, MO); Cochabamba, Dec 1930, Bro. Julio II7 (US); Cochabamba, 2600 m, 1932, Bro. Julio 11-210 (US); Punata, 6 km NE of Punata, 1 km NE of La Villa, canyon of Rio Pucara Mayu, 2875 m, 17?30'S, 65?47'W, 5 Mar 1988, Nee et al. 36457 (NY); Pocona, 2500 m, 11 Nov 1928, Steinbach 8691 (F, GH, K, MO, NY). LA PAZ: Sirupaya near Yanacachi, 2100 m, 16 Nov 1906, Buchtien 315 (NY); Tunari, 2800 m, Apr-May 1892, Kuntze s.n. (NY); Barrancasde Rio Grande, 28 Jan 1945, Peredo 135 (A, NY). SANTACRUZ:Cordillera, Camiri, 67 km toward Monteagudo, 1150 m, 27 Oct 1983, Beck & Liberman 9808 (NY); Nuflo de Chavez, finca of Don Luis Olivarria, 3 km N of Ascenci6n de los Guarayos, ca. 15?39'S, 63?05'W, 11 Aug 1983, Hopkins et al. 168 (NY); 2 km from center of Concepci6n, 475 m, 16?07'S, 62?02'W, 28 Dec 1986, Nee 33352 (NY); ruderal in town of Concepci6n, 490 m, 16?08'S, 62?02'W, 28 Dec 1986, Nee 33366 (NY); along rd. from Santa Cruz to Samaipata, 1 km SW of Angostura, 650 m, ca. 18?09'S, 63?31'W, 13 Jan 1987, Nee 33477 (NY); 1 km N of center of Cotoca, 350 m, 17?45'S, 62?59'W, 15 Jan 1987, Nee 33529 (NY); along Rio Pirai, 1 km NW of El Torno, 495 m, 17?59'S, 63?23'W, 17 Jan 1987, Nee 33616 (NY); along Rio Pirai, 1 km N of La Guardia, 460 m, 17?53'S, 63?15'W, 20 Jan 1987, Nee 33697 (NY); W side of Santa Cruz, just above floodplain of Rio Pirai, 400 m, 17?47'S, 63?12'W, 22 Jan 1987, Nee 33731 (NY); 2 km W of center of La Belgica, 360 m, 17?34'S, 63014'30"W, 24 Jan 1987, Nee 33775 (K, NY); highest point on Samaipata-Mairanard., 6 km by air WNW of Samaipata and 11 km by rd. SE of Mairana, 1800 m, 18?10'S,

TAXONOMIC TREATMENT 63?55'W, 30 Jan 1987, Nee & Coimbra S. 33877 (NY); 4 km SW of El Trigal on rd. to San Juan del Chaco, S side of Rio Ariruma, 19 km by air NNW of Vallegrande, 1950 m, 18?29'S, 64?07'W, 31 Jan 1987, Nee & Coimbra S. 33931 (K, NY); Quebrada Llullucha, 14 km by air SSE of Mataral, 1550 m, 18?14'S, 64011'W, 1 Feb 1987, Nee & Coimbra S. 33951 (NY); 12 km E of center of Santa Cruz on rd. to Cotoca, 375 m, 17?46-47'S, 63?04'W, 24 Feb 1987, Nee 34244 (NY); Estancia San Rafael de Amboro, 15 km by air SSE of Buena Vista, 375 m, 17?35'S, 63?37'W, 28 Jul 1987, Nee et al. 35404 (MO, NY); 5 km by air SE of Mairana, on rd. to Samaipata at Quebrada Seca, 1550 m, 18?09'S, 63?56'W, 4 Feb 1988, Nee 36172 (NY); 3 km S of Mataral on rd. to Vallegrande, 1425 m, 18?08'S, 64?13'W, 6 Feb 1988, Nee & Saldias P. 36248 (NY); 3 km SE of Cotoca, 350 m, 17?47'S, 62?58'W, 11 Feb 1988, Nee 36281 (NY); 6.5 km NNW of center of Vallegrande, 1950 m, 18?26'S, 64?07'W, 9 Mar 1988, Nee & Solomon 36557 (MO, NY); 5.5 km S of Vallegrande, vic. Santa Rosita, Quebrada Huasacafiada, 2050 m, 18?32'S, 64?06'W, 31 Dec 1988, Nee & Vargas C. 37468 (NY); ca. 15 km NE of Cotoca along hwy. to Puerto Pailas, 325 m, ca. 17?42'S, 62?53'W, 28 Jan 1989, Nee 37754 (NY); Cabezas, La Morita, 23 Jan 1945, Peredo 48 (A, NY); La Quebrada, 405 m, 26 Jan 1945, Peredo 80 (A, NY); ca. 8 km W of San Miguel, before reaching Altamira, ca. 400 m, 27 Apr 1986, Seidel & Beck 211 (NY); Santa Rosa del Sara, 450 m, 15 Oct 1916, Steinbach 3196 (F, GH, NY); Buena Vista, Sara, 500 m, 15 Nov 1920, Steinbach 5101 (F, GH, NY); Santa Cruz, Jardin Botfnico de Santa Cruz (Av. Cotoca at km 12), 1 Dec 1987, Thomas 5554 (NY). TARIJA: Outskirts of Tarija, 27 Feb 1983, Bohs 2076 (GH); Cercado, rd. to Junacas, 4 km from Santa Ana, 1980 m, 15 May 1986, Basti6n 1283 (NY). CHILE. ATACAMA: Copaipo, Vegas de Copaipo, Nov 1886, Gigoux s.n. (GH); San Felipe, Dec 1925, Bro. Claude Joseph 3829 (US). CONCEPCION: Chillanito, Jardin Zoologico, cult., 9 Dec 1957, Junge 3064 (US). SANTIAGO: Comuna de Tiltil, Capilla de Caleu, 900 m, 13 Apr 1969, Plowman 2668 (GH, US). PARAGUAY. Thiaty, in canyon, Jorgensen 4373 (NY); southern Paraguay, 1 Oct 1892, Kuntze s.n. (NY). AMAMBAY: San Juan Caballero, 700-800 m, 19-20 Oct 1979, Mizoguchi & Sano 1129 (NY); Bella Vista, along Rio Apa, 200 m, 23 Mar 1983, Simonis et al. 170 (F, U). CAAGUAZU: In forests, 19 Nov 1874, Balansa 2097 (K); Campo Grande near airport, 19 Jul 1982, Dlouhy & Fernandez Casas 7673 (NY). CENTRAL: Yaguar6n, 1 Feb 1966, Krapovickas et al. 12307 (US); suburbs of Ascunci6n, 16 Mar 1889, Morong 617 (GH, MO, NY, US); Villa Elisa, 12 Oct 1959, Pedersen 5105 (A, K, MO). CHACO: Madrej6n, 20040'S, 59?50'W, 20 Jun 1983, Hahn 1473 (MO, NY, PY); Cacui, Aug 1933, Meyer 2037 (GH). CORDILLERA: Ca. 15 km S-SE of San Bernardino, ca. 40 km E of Ascuncion, 15 Jan 1974, Conrad & Dietrich 2238 (MO); Arroyos y Esteros, Jul 1875, Balansa 2100 (K); vic. Lago Ypacaray,Oct 1913, Hassler 12381 (K, MO, NY, US); Cordillera de los Altos, Jan 1916, Rojas,herb. Corn.Osten.8409 (K). GUAIRA:Villarica,

67 May 1931, Jorgensen 3676 (F, NY, US). NEEMBUCU/ MISIONES: Estero Camba, 8 Nov 1978, Bernardi 18403 HAYES:Villa Rica, Feb 1931, (MO, NY). PRESIDENTE Jorgensen 3676 (K); Rio Pilcomayo, Mar 1888--1890, Morong 1529 (GH, MO, NY, US). SAN PEDRO: Alto Paraguay, Primavera, 20 Jun 1954, Woolston 231 (NY), 8 Aug 1954, Woolston 280 (K, NY), 12 Nov 1958, Woolston 1035 (NY). ARGENTINA. BUENOSAIRES:Paso del Rey, 11 Nov 1944, Alvarez s.n. (A, NY); Pereyra, 13 Feb 1945, Alvarez 518 (NY, 525); Bella Vista, 17 Mar 1945, Alvarez 637 (A), Alvarez 648 (A, NY); J.C. Paz, 19 Mar 1945, Alvarez 669 (A, F); Punta Chica, F.C.C.A., 22 Apr 1945, Alvarez 728 (A, NY); El Palomar, 14 May 1945, Alvarez 761 (NY); Posadas, 29 May 1944, Bertoni 878 (NY); Berisso Las Talas Sefiora, 23 Mar 1945, Boffa s.n. (NY); banks of Rio La Plata, Isla Santiago, 31 Oct 1930, Cabrera 1509 (F); vivero del Delta, Castelar, Jul 1936, Castro s.n. (K); vic. Buenos Aires, 10 Aug 1913, Curran s.n. (US); Echeverry, 29 Oct 1929, Dawson 851 (NY); ca. 10 km of La Plata on NW bank of Rio La Plata, ca. 0 m, 8 Dec 1938, Everdamet al. 23366 (GH, K); La Plata, Punta Lara, Arroyo de Cafias, 34?48'S, 57054'W,Gamundi & Roivainen 1 (US); vic. Buenos Aires, 1891, Hauthal 647 (NY); Partido Candil, Azucenit, 230 m, 23 Jan 1945, Huidrobo 1756 (A); Merlo, Libertad, 18 Oct 1952, Mazzucconi 282 (K); Miramar, Partido General Alavarado, 1 Apr 1951, Pedersen 1046 (U, US); Avellaneda, 15 Apr 1913, Rodriguez 215 (US); La Plata, Punta Lara, 29 Oct 1946, Sparre 7 (K); San Martin, Oct El Totoral,26 May 1901, Vaccaris.n. (US). CATAMARCA: 1943, Bartlett 20257 (GH); El Rodeo, 15 Jan 1911, Castillon 2019 (US); Andalgala, Esquina Grande, 10 Dec 1896, Jorgensen 1808 (US); Agua de Mato, 1300 m, Parodi 14333 (GH); Santa Rosa, Abijilan, 25 Jan 1942, Pierotti 11516 (A, NY). CHACO:Rio Bermejo, 8 km S of Puerto Velez, 14 Nov 1986, Crist6bal et al. 2089 (A, K, NY); Colonia Benitez, Reserva del INTA, 16 May 1979, Ferrucci et al. 108 (K); Cacui, Mar 1933, Me!er 2036 (GH); Colonia Benitez, Jan 1937, Meyer 2735 (A); Resistencia, banks of Rio Negro, 12 Jul 1944, Rojas 11591 (A); Colonia Benitez, 12 Feb 1956, Schulz 9065 (F); Colonia Benitez, 5 Nov 1964, Schulz 14146 (F); Resistencia, Colonia Benitez, 150 m, 12 Dec 1928, Venturi 7894 (F, GH, K, MO, US). CORDOBA:Colanchanga, Sierra Chica de C6rdoba, Jul 1881, Hierony)mus s n. (GH); vic. La Quebrada, Col6n, 13 Oct 1946, Hunziker 6893 (CORD, F); Mercedes, Jofr6, 25 Mar 1945, Ibarrola 2754 (NY); Col6n, Ascochinga, 14 Mar 1944, O'Donell & Rodriguez 401 (A, GH); Rio Ceballos, 15 Mar 1944, O'Donell & Rodriguez 487 (A); between Cerro Blanco and La Hollada (Rancho Alegre), km 767 Ruta Nacional 20, ca. 1300 m, 31?50'S, 64?15'W, 12 Dec 1978, Solomon & Solomon 4079 (MO). CORRIENTES: Ituzaing6, San Pedro, 27?45'S, 56?52'W, 12 Nov 1976, Arbo et al. 1199 (F); Mburucaya, Estancia Santa Maria, 20 Nov 1950, Pedersen 862 (GH, NY, US); Santo Torn6, Estancia Vuelta de Ombf, 3 km SW of Gdor. Virasoro, 10 Jul 1984, Tr]essenset al. 2704 (K); Ituzaing6, 17 km NW of San Carlos, Estancia Rinc6n Chico, 11 Jul 1984,

68 Tressens et al. 2762 (A, K), 14 Feb 1991, Tressens et al. 3902 (U). ENTRE RIOS: Federaci6n, Estancia Buena Esperanza, 20 Oct 1960, Pedersen 6207 (A, K). FORMOSA:Guayalle, Jun 1919, Jorgensen 3341 (GH, US). JUJUY:Santa Barbara, rd. from El Fuerte to Palma Sola, 20 Feb 1972, Cabrera et al. 22299 (NY, US); Arroyo del Medio, 28 Jun 1906, Fries 315 (MO); El Carmen, Dique La Ci6naga, 6 Jan 1971, Krapovickas & Cristobal 17534 (US); Jujuy-Salta boundary, 1972, Morton s.n. (MO, US); between Yala and Lagunas de Yala, 8 Apr 1945, O'Donell 2829 (NY); Sierra de Calilegua, 750 m, 5 Sep 1927, Venturi 5288 (US). MISIONES: Posadas, 29 May 1944, Bertoni 878 (K); Posadas, 7 Dec 1907, Ekman 823 (F, NY, US); Candelaria, Yalelyry, 220 m, 21 Apr 1945, Montes 821 (NY); Loreto, 216 m, 24 Apr 1945, Montes 980 (A); San Ignacio, Teyucuar6, 222 m, 5 Apr 1946, Montes 2103 (NY); Loreto, 6 Sep 1946, Montes 2491 (US); Cainguas, Puerto Rico, 205 m, 12 Nov 1948, Montes 3367 (US); San Ignacio, Naxauguazu, 290 m, 30 Mar 1958, Montes 27586 (NY); San Ignacio, pastures near ruins, 12 May 1969, Plowman 2733 (GH, RB); Candelaria, 11 Jun 1944, Sesmero 74 (US); Caingubs, vic. Arist6bulo del Valle, 29 Jul 1987, Vanni et al. 866 (NY). RIOJA: Capital, Los Duraznillos, km 28, 22 Nov 1941, Alauy 37 (GH), 8 May 1942, Alauy 89 (A). SALTA: Anta, Cevil Sola, 12 May 1944, Huidrobo 515 (NY); Oran, El Naranjo, 22 Jan 1945, Pierotti 18 (NY); Bella Vista, 29 Jan 1945, Pierotti 99 (A, NY); Rio de las Piedras, 20 Nov 1911, Rodriguez 105 (US); El Naranjo, 19 Nov 1974, del V Ruiz et al. 10597 (F); Piquirenda to Aguaray, 500 m, 16 Feb 1925, Schreiter 3790 (US); Alemania, 1400 m, 13 Dec 1929, Venturi 9959 (GH, MO, US). TUCUMAN: Burruyacu,Cerro del Campo, 1800 m, Jan 1918, Bailetti 21 (US); Famailla, Quebrada del Lules, Rio Lules, 4 Mar 1945, Herrera 208 (NY); Tafi, La Toma, 10 Jun 1945, Herrera 636 (A); Cerro de San Javier, 9 Jul 1945, Lillo 49 (A); Siambon, Sierra de Tucuman, 12 Jan 1873, Lorentz & Hieronymus 1016 (B [destroyed: F neg. 2792]); Francas, 800 m, Monetti 19 (US); Rio Chico, El Potrerillo, 21 Nov 1913, Monetti 1535 (US); Cerro del Duraznillos, 24 Feb 1914, Monetti 2008 (US); Ancajulio Las Arquitas, 12 Dec 1944, Olea s.n. (NY); Aconquija, 635 m, Ortiz 35 (A); Tafi, Los Juntos, 1350 m, 6 Jan 1953, Petersen & Hjerting 926 (NY); Quebrada de Lules, 500 m, 9 Jul 1923, Schreiter 9944 (US); Sunchal, 16 Apr 1944, Varelas.n. (A); Tafi, Yerba Buena, 25 Jan 1919, Venturi 105 (GH, US); Cumbre to Taficillo, 1800 m, 4 Mar 1928, Venturi5947 (US); Cerro de Taficillo, 1600 m, 20 Apr 1930, Venturi10427 (GH, MO); Cerro de Aconquija, 20 May 1945, Villa s.n. (NY). URUGUAY. CANELONES:Santa Lucia, Mar 1921, Corn. Osten. 16290 (K). FLORIDA:Rio Tipary below Nausavillagra, 31 Dec 1936, Rosengurtt B726 (US). LAVALLEJA:Boundary of Campos de Loza, SE of Parador Salus, 27 Nov 1943, Bartlett 20944 (NY), Bartlett 20944a (US). MONTEVIDEO: Arroyo Piedras, 5 Feb 1928, Herter 758 (GH, MO, NY, U); Pajas Blancas, 10 Feb 1935, Herter 758d (MO). RIVERA:Near Rivera,

FLORA NEOTROPICA 7 Dec 1943, Bartlett 21097 (GH, NY, US). Rio NEGRO: Estancia Nueva Melhem, 29-30 Dec 1965, del Puerto & Marchesi 5781 (F). SAN JOSE: San Jose, 12 Jan 1935, Legrand 375 (F). Soriano, Juan Jackson, 3 Jan 1941, Gallinal et al. PE-4329 1/2 (US). SORIANO:Juan

Jackson,EstanciaMonz6n-Heber,3 Jan 1941, Gallinal et al. PE-4329 (U).

Local names.Mexico.Oaxaca:hierbaenana;Colombia. (cultivatedform,Bogota:mirto,granode oro);Brazil.Parana:ne-n ka-ni- rary-nh-si~,laranjinha do mato; Uruguay.Soriano:revientacaballos;Argentina.Buenos Aires: revientacaballos;Tucuman:sachaediondilla. Solanumpseudocapsicum is a widespreadand extremelyvariablespecies. Cultivatedformsthataregenknown erallycompletelyglabroushavebeentraditionally as S. pseudocapsicum, while native plants have gone by a varietyof names.Degree of pubescence,however, is a continuousgradientthroughoutthe native and introducedrangeof the species andis not a reliablecharacterwith which to differentiatetaxa in this group. Many of the numerous synonyms of Solanum pseudocapsicumarewell-markedgeographicalvariants thattakenalone withoutconsiderationacrossthe entire species range are distinct. The following are the more clearly markedof these variations. 1. Cultivated specimens are almost always completely glabrous (occasionally with a few dendritic trichomes on the new growth) and they often have large flowers and fruit. Some apparently native collections

from Uruguay,

near Buenos Aires, Argentina, and near Sao Paulo, Brazil, are of this general morphology. The type of Solanumpseudocapsicum was a cultivatedplantcollectedin Madeira(see Knapp & Jarvis, 1991). 2. Plants with broadly elliptic leaves and small fruits from the province of Tucuman, Argentina, have been called Solanumtucumanense. 3. Populations with very dense pubescence from high-elevationBoliviahavebeencalledSolanum validum. 4. In the dry coastal valleys of Venezuela and Colombiaplantswith small flowers, small fruit, and narrow leaves have been called Solanum karstenii. 5. Plantswith themost commonmorphology,found in southernMexico and southernBrazil, have been called Solanum diflorum. Other named populationsaremerelyextremesfromthismean. The originalcollectionof Solanumpseudocapsicumn that ended up in Clifford's garden,to be describedby Linneaus(1737), probablywas broughtto Madeiraby the earlyPortuguesetraders.Many early introductions

TAXONOMIC TREATMENT

69

of Brazilian plants were effected by this route, and it 24 May 1949, Pickel s.n. (SP-56730);Vila Mariana,26 seems likelythatS.pseudocapsicumis anotherexample. Nov 1905, Usteri 16 (herb. Hoehne 15305) (US). Solanum spissifolium is closely related to S. a widespreadweedy species in the pseudocapsicum, 9. Solanum spissifolium Sendtn. in Mart., Fl. Bras. It differs from that species in its narrow, Neotropics. 10: 33. 1846. Type. Brazil. in Brasilia Australiori, lanceolateleaves and somewhatsmallerflowers, fruit, Sellow s.n. (lectotype,K, heredesignated;isolectoand seeds. Solanumspissifoliummay be a geographitypes, B [destroyed:F. neg. 3181], BM). Fig. 34 cal variantof S. pseudocapsicum, but since no recent collections exist andit is easily differentiatedfromthat Subshrubs to shrubs (?), the branches closely I to accordit species statusuntil future species, prefer a stems and packed, giving bushy appearance;young leaves densely pubescent with golden dendritic tri- in-depthstudies areundertaken. The lectotype I have chosen for Solanum chomes 0.5-1 mm long; barkof older stems brown to golden-brown. Sympodial units plurifoliate. Leaves spissifolium was previously the right hand specimen lanceolate,widest at the middle,with 3-4 pairsof main on a composite sheet at K (but now separated,see Fig. lateralveins, glabrousor sparselypubescentadaxially, 34), and was received from B in 1869. The left hand sparselyto densely pubescentwith dendritictrichomes element was also S. spissifolium, but is attributableto abaxially,the trichomes 0.5-1 mm long, denseralong Klotzsch, andthus not type material. the veins; lamina 1-3 x 0.4-0.5 cm, the apex acute to somewhatrounded,the base attenuate;petiole ca. 0.2 mm long. Inflorescencesopposite the leaves or intem- III. Solanum nudum species group (S. acuminatum, odal, simple, densely pubescent with dendritic triS. aphyodendron,S. caavurana, S. campaniforme, chomeslike thoseof the stems, 1-4 mm long, 1-3-flowS. cassioides,S.cordioides,S. daphnophyllum, S.gertii, ered;pedicel scars closely spaced, not overlapping. S. intermedium, S. lasiopodium,S. nudum,S.pseudoBuds ellipsoid, the tips flattened,exsertedfromthe caquina,S. reitzii,S. restingae,S. santosii, S. spirale, lyx tubes, but not exceeding the lobes. Pedicels at anS.symmetricum, S. tepuiense,S. trichoneuron, S. trachythesis slender, nodding, 3-6 mm long, ca. 0.5 mm trichium,S. warmingii). Fig. 35 diam.,densely pubescentwith dendritictrichomeslike those of the rest of the inflorescence.Flowers with the Shrubs, treelets, or small trees; young stems and calyx tubeconical, 1-1.5 mm long, the lobes long-trian- leaves glabrousor pubescent,the trichomes simple or gularwith roundedapices, 2-2.5 mm long, pubescent occasionallyfurcateor dendritic,often only on one side with dendritictrichomes0.5-1 mm long;corolla white of the stem.Sympodialunitsdifoliate,usuallygeminate, (?), 1-1.1 cm diam., lobed nearlyto the base, the lobes usuallyanisophyllous.Leavesellipticto lanceolate,glaplanarat anthesis,the tips and marginsdensely papil- brousor pubescent,often with tufts of trichomesin the lose; anthers 2-2.5 x 1-1.5 mm, poricidal at the tips, axils of the mainveins abaxially,the trichomessimple the poresteardropshaped;free portionof thefilaments or occasionallydendritic,the apex andbase various.Inca. 0.5 mm long, the filament tube ca. 0.5 mm long, florescences opposite the leaves, usually simple, occaglabrous;ovaryglabrous;style straight,ca. 5 mm long, sionallybranched,glabrousor pubescent;pedicelscars sparsely pubescentwith simple trichomesca. 0.3 mm closely spacedandnot overlapping,occasionallymore long; stigma small-capitate,the surfaceminutelypap- widely spaced.Buds globose or ellipsoid, stronglyexillose. Fruita globose, redor orange-redberry,the peri- sertedfromthe calyx tube.Flowers white or greenishcarpthin and shiny, 1-1.5 cm diam.;fruitingpedicels white, sometimes fleshy, the calyx lobes in a few spewoody, erect,ca. 1 cm long, 0.7-1 mm diam.Seedspale cies large and leafy (S. caavurana, S. warmingii),the tan, flattened-reniformwith incrassatemargins,2-2.5 corollalobes planaror slightly reflexedat anthesis;anx 1.5-2 mm, the surfacesminutelypitted,the cells rect- thersusually quite stout.Fruit a green, hardberry,beangularin outline. Chromosomenumbernot known. coming yellowish at maturity;fruitingpedicel woody, deflexed. Seeds flattened-reniform,pale tanto yellow. Distribution (Fig. 30). In southeasternBrazil, Sao Distribution. Widespreadin secondaryhabitats. Paulo state, nearthe coast at sea level. Membersof the Solanumnudumspecies groupare Specimens examined. BRAZIL. S.loc., with no colquite heterogeneous,varyingfrom glabrousto pubeslector on label ("de R. Brown chez M. Banks") (MPU). SAo PAULO:Near Sao Paulo, 19 May 1815, Bowie & centandfromgood-sizedtrees(S.acuminatum)to small Cunninghams.n. (BM); Jabaquara,Dec 1914, Brade 7019 shrubs (S. santosii). Many of the species have a dis(herb. Hoehne 6894) (US); Capital, 18 Nov 1894, Edwall tinctive patternof pubescence on the leaf undersides 2885 (herb. Hoehne 15309) (US); Sao Paulo, Vila Ema, with tufts of trichomes in the axils of the main veins

FLORA NEOTROPICA

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TAXONOMIC TREATMENT

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FIG. 35. Solanum nudum species group. A. S. aphyodendron habit, Costa Rica. B. S. aphyodendron flowers, Ecuador (Knapp & Mallet 6272). C. S. acuminatum flowers, Peru (Knapp & Mallet 6334). D. S. campaniforme habit, Venezuela (Knapp & Mallet 6728). E. S. caavurana calyx, Paraguay (Knapp et al. s.n.). F. S. campaniforme flowers, Venezuela (Knapp & Mallet 6728).

72

FLORA NEOTROPICA

with the midrib. These tufts have been interpretedas domatiaused by mites (O'Dowd & Willson, 1991), but I have only occasionally seen mites in any of the tufts of hairs I have examined closely. A single unusualcollection fromBolivia (Luteyn& Dorr 13722: La Paz, Prov. Sur Yungas, along road throughprimarycloud forest, 7-9.4 km NE of (above) Huancan6, 2286-2499 m, 16?20'S, 67?32'W, 17-18

May 1990) may representa new species belonging to this group.The planthas the semi-umbellateinflorescences and elongate filament tube found in Solanum symmetricum,also fromBolivia, but is densely pubescent with long, white, simple, uniseriate trichomes. Fruitingmaterialof these plantswill be necessaryfor a final analysis andplacement.

Key to the species of the Solanum nudumspecies group 1. Plants glabrous, with no easily visible trichomes on the leaf undersides. 2. Sympodial units difoliate, geminate, usually anisophyllous. 3. Stems strongly winged from decurrent leaf bases; calyx lobes minute; corolla lobes cucullate. ......................................................................................... Bahia, Brazil . 22. S. restingae 3. Stems not winged; calyx lobes petaloid; corolla lobes not markedly cucullate. Minas Gerais, Brazil.......................................................................................................................................... 29. S. warmingii 2. Sympodial units difoliate, usually not geminate, never anisophyllous. 4. Leaves drying pale green; inflorescences generally unbranched;flowers 1.5-2 cm diam., white. S Peru and Bolivia ............................................................................................................ 15. S. daphnoph ll m 4. Leaves drying black; inflorescences many times branched; flowers 6-8 mm diam., greenishwhite. Bahia, Brazil .............................................................................................................. 126. S. co dioides 1. Plants not glabrous, with at least some uniseriate trichomes on the leaves beneath, these occasionally sparse, often in tufts in the vein axils. 5. Plants scabrous; trichomes with broad, multicellular bases, which appear as white specks to the naked eye. Argentina, Paraguay and Argentina..................................................................27. S. trachi,trichium 5. Plants not scabrous, variously pubescent; trichomes uniseriate, simple or dendritic. 6. Trichomes dendritic, some with broad bases; inflorescences internodal or on short shoots. Brazil ......................................................................... 14. S. cassioides 6. Trichomes simple, uniseriate (at least to the naked eye or under a dissecting microscope), without broad bases; inflorescences opposite the leaves. 7. Pubescence dense, covering the leaf undersides. 8. Anthers unequal, 3 long, 2 short; upper leaf surface shiny. Parana and Santa Catarina, Brazil, in Araucaria forests ..... .....................2......................21. S. reitzii 8. Anthers of equal size; upper leaf surfaces sparsely pubescent. E Brazil............. 17. S. intermedium 7. Pubescence generally confined to the axils of the main lateral veins on the leaf undersides, occasionally extending to the lamina. 9. Calyx lobes petaloid, large and scarious-margined. Plants of SE Brazil. 10. Plants drying black; stigma capitate; stems not winged. Widespread in SE Brazil, and in adjacent Paraguay and Argentina................................................................. 12. S. caavurana 10. Plants drying pale sea- or yellowish-green; stigma not capitate; stems winged. Minas Gerais, Brazil ..... .........................................................................................29. S. warmingii 9. Calyx lobes not petaloid, various in shape. 11. Berries reddish-orange at maturity. Old World tropics ........................................24. S. spirale 11. Berries green or greenish-yellow at maturity. New World tropics. 12. Inflorescencesubumbellate,most flowers in the distal 1/4 of the inflorescenceaxis; free portion of the filamentselongate. E Bolivia and W Argentina............25. S. svmmetricum 12. Inflorescence not subumbellate, flowers arising in more than the distal 1/4 of the axis; free portion of the filaments approximately equal to the filament tube. 13. Buds large, ca. 1 cm long; flowers large and usually somewhatfleshy, 1.5-2 cm diam. 14. Stems winged; leaves with tufts of moniliform trichomes in the abaxial vein axils. Venezuela .......................................................................... 26. S. tepuiense 14. Stems not winged; leaves with tufts of simple uniseriatetrichomes in the abaxial vein axils. SierraNevada de Santa Marta,Colombia.......... 18. S. lasiopodium 13. Buds smaller, less than 1 cm long; flowers smaller, not markedly fleshy, usually less than 1.5 cm diam. 15. Calyx lobes spathulate; stems lightly winged; bark green and shiny. Bahia, Brazil ..... .............................................................................................. 23. S. santosii 15. Calyx lobes deltate; stems not winged; bark otherwise.

TAXONOMIC TREATMENT

73 16. Bark white or very pale yellow, even on young stems; abaxial leaf venation pale-colored. 17. Stems and inflorescence axes pubescent with simple uniseriate trichomes 1-1.5 mm long. Parana, Brazil ................................. 16. S. gertii 17. Stems and inflorescence axes glabrous, or at most minutely pubescent when very young. 18. Anthers unequal, 3 long and 2 short; abaxial leaf veins yellow. SE Brazil and Argentina............................................ 20. S. pseudoquina 18. Anthers equal; abaxial leaf veins pale, not bright yellow. 19. Inflorescence 1.2-6 cm long; flowers 1-1.2 cm diam.; calyx lobes deltoid; berries 1-1.2 cm diam. Montane Central America to Bolivia .............................................................. 11. S. aphvodendron 19. Inflorescence 3-8 cm long; flowers 0.6-1 cm diam.; calyx lobes quadrate; berries 7-9 mm diam. S. Bolivia to A rgentina ..............................................................28. S. trichoneuron 16. Bark dark gray or brown, never white, occasionally pale on very young stems, but not as above. 20. Axis of the inflorescence elongate, to 20 cm long; buds ellipsoid; anthers not stout, longer than wide; leaves usually not longer than 5 cm. Widespread, Guianas to SE Brazil ................... 13. S. campaniforrne 20. Axis of the inflorescence not markedly elongate; buds globose until just before anthesis; anthers stout, nearly as wide as long; leaves usually longer than 5 cm. 21. Inflorescences at shoot tips often overtopping the young leaves; flowers 1.2-1.8 cm diam. 22. Axis of the inflorescence glabrous; old inflorescences remaining on lower shoots; calyx lobes not markedly woody when dry; stigma bright green, capitate. E Andes .. 10. S. acuminatum 22. Axis of the inflorescence with a few scattered trichomes; old inflorescences not remaining on lower shoots; calyx lobes appearing woody when dry; stigma not bright green, minutely capitate. Sierra Nevada de Santa Marta, Colom bia .............................................................. 18. S. lasiopodium 21. Inflorescences strictly leaf-opposed, not overtopping the young leaves; flowers 0.5-1.1 cm diam. (can be somewhat larger in high-elevation Venezuela). Widespread, Mexico to Peru and Brazil ...................................................................................... 19. S nudum

10. Solanum acuminatum Ruiz & Pav., Fl. Peruv.2: stems pale grayish-brown,glabrous.Sympodialunits 34. fig. 159a.1799.Type.Peru.Huanuco:adChinchao difoliate,geminate.Leaveselliptic,usuallydryingdark, vicum, Ruiz & Pav6n s.n. (lectotype, MA, here widest at the middle,glabrousandshinyadaxially,with designated; isolectotypes, B [destroyed: F neg. tuftsofuniseriatetrichomes1-1.5 mm long in the axils 2594], MA, MPU). Figs. 35C, 36 of the mainlateralveins abaxially,orwiththe trichomes Solanum pearcei Britton ex Rusby, Mem. Torrey sparselyto densely covering the abaxiallamina,if the Bot. Club 4: 227. 1895. Type. Bolivia. La Paz: abaxiallaminacovered,the trichomesdenseralong the Yungas, 1890, Bang 712 (holotype, NY; isotypes, veins and in the vein axils, the trichomesoccasionally GH, K, MO, NY, US). branched;majorleaves 9-17 x 4-8 cm, with 7-8 pairs Solanum hypomicropogon Bitter, Repert. Spec. Nov. of main lateral veins, these prominentbelow, raised Regni Veg. 18: 56. 1922. Type. Peru. Junin: La above, the apex acute,the base acuteto rounded,occaMerced in Chanchomayo-Tal, 800-1000 m, 12 sionallyoblique;petioles0.7-1.2 cm long;minorleaves Jul 1912, Weberbauer 1876 (holotype, B [defromthe majorones mainly in size, 3-6 x 2differing stroyed: F neg. 2610]; lectotype, MOL, here 3.5 cm, the apex rounded,the base acute;petioles 6-8 designated). mm long. Inflorescences opposite the leaves, often Treelets to trees 20 cm d.b.h., 4-20 m tall; young appearingterminaland overtoppingthe growing tips stems and leaves glabrous, minutely papillose or sparsely of the leaves, 1-5 cm long, usually simple, but occato densely bristly with uniseriate simple trichomes 1sionallyfurcate,5-50-flowered, glabrousandminutely 1.5 mm long, usually dryingblackish;barkof the older papillose at the tips to sparsely or densely pubescent

FLORA NEOTROPICA

74 CLIX.

?+?99 i

4

-

?J;1P,---

a

SOLANUM

awn50LaNU.

. -'--- ~_

-

SOLANUM

///r.

FIG. 36. Solanum acuminatum Ruiz & Pav6n, plate 159a in Ruiz & Pavon's Flora Peruviana.

TAXONOMIC

TREATMENT

75

500~~0

a

p

,

I

i~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~......

~~~~~~~~~~Yl~~~~~~~~~ c~~~~~~J r'~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~..... .........

/

FI.37

isrbtinofSlau

cuiatm(pe

with simpleuniseriatetrichomeslike those of the stems, dryingblack;pedicel scars in pairs, the membersof a pairtouchingbut not overlapping,the pairsca. 0.5 mm apart.Buds globose, the calyx lobes apiculate in bud, the corollaenclosedin the calyx untilquitelate.Pedicels at anthesis 1.1-1.5 cm long, fleshy,whiteon live plants, taperingfrom a basal diam. of ca. 0.5 mm to an abrupt widening at the base of the calyx tube. Flowers with the calyx tube cyathiform, ca. 2 mm long, the lobes roundeddeltoid,apiculate,glabrousto sparselypubescent with simple uniseriatetrichomes, 1.5-2 mm long, the apices with minute uniseriate trichomes;corolla white, 1.2-1.8 cm diam.,lobed ca. 3/4 of the way to the base, the lobes partiallyreflexed at anthesis, the tips and margins of the lobes minutely papillose; anthers shortandstout,ca. 3.5 x 1.75 mm, poricidalat the tips, the poresteardropshaped;free portionof thefilaments 0.25-0.5 mm long, the filamenttube0.25-0.5 mm long; ovary glabrous;style straight, 4-5 mm long; stigma globose, brightgreenon live plants,minutelypapillose. Fruit a globose, greenberry,ripeninga dirtygreenishyellow, ca. 1 cm diam.;fruiting pedicels deflexed, woody, the apical portion expanded,ca. 3 mm diam.,

irls)adS.cavrna(oidcrce)

below this portiontaperinggentlyto a basaldiam.of ca. 1 mm. Seeds pale yellow, flattened-reniform,ca. 3 x 2 mm, the marginsincrassate,the surfacepitted,the pits ca. 0.5 mm diam. Chromosomenumbernot known. Distribution (Fig. 37). FromcentralPeruto northern Bolivia in secondary growth at 1000-3000 m. Selected specimens examined. PERU. Cuzco: Kosiipata, Quitacalzon(QuebradaSta. Alicia), ca. km 163 of Lucre-Paucartambo-Shintuya rd., 1100-1200 m, 13?07'S, 71?15'W, 11 May 1984, Knapp & Mallet 6429 (BH, NY, USM). HUANUCO:Vic. of Tingo Maria, rd. to Fundo San Juan at jct. of Rio Chinchao and Rio Huallaga, 18 Jul 1962, Mathias & Taylor 5918 (F, K, US). JUNiN: Chilpes, 8 km S of Vitoc overlooking Rio Tulumayo, 1420-1700 m, 11?12'S, 75?20'W, 8 Feb 1983, Gentry et al. 40189 (MO, USM); along Rio Colorado, N bank W of Puente Colorado, 12 km N of La Merced, Chanchamayo valley, ca. 850 m, 22 Mar 1984, Knapp et al. 6343 (BH, US, USM); ca. 25 km S of San Ram6n on rd. to Vitoc, ca. 950 m, 23 Mar 1984, Knapp et al. 6350 (BH, US, USM). PASCO:24 km N of Huancabamba, above Rio Pozuzo, 1300-1400 m, 10?18'S,75?32'W,5 Feb 1983, Gentryet al. 40035 (MO, USM); trail between Pozuzo and Yanahuanca, along Rio Pozuzo, ca. 1000 m, 10?05'S, 75?28'W, 14 Mar

76 1984, Knapp et al. 6328 (BH, US, USM); limestone cliffs along Rio Pozuzo on trail from Yanahuancato Chumalli, 800-1200 m, 0?3'S, 75?25'W, 15 Mar 1984, Knapp et al. 6329 (BH, US, USM); trail between Pozuzo and Yanahuanca, along Rio Pozuzo, 800-1000 m, 10?05'S, 75?28'W, 16 Mar 1984, Knapp et al. 6333, 6334 (BH, US, USM); km 15-18 of Villa Rica-Pto. Bermudez rd., ca. 1600 m, 21 Mar 1984, Knapp et al. 6337 (BH, NY); Cueva Grande, estaci6n near Pozuzo, 3500 ft, 23 Jun 1923, Macbride 4794 (F); 5 km SE of Oxapampa, Oswaldo Muller property, 1850 m, 10?36'S, 75?23'W, 9 Apr 1983, D. N. Smith 3655 (MO, NY); trail to summit CordilleraYanachagavia Rio San Daniel, 2500 m, 10?23'S, 75?27'W, 17 Jul 1984, D. N. Smith et al. 7824 (MO, NY). SAN MARTIN: Zepelacio near Moyobamba, 1100-1200 m, Oct-Nov 1933, Klug 3371 (NY); Boca Toma del Shilcayo, N of Tarapotoon Rio Shilcayo, 400 m, 6?29'S, 76?24'W, 12 Aug 1986, Knapp 8009 (MO, NY, USM); San Juan de Pacaizapa, km 72 Tarapoto-Moyobambard., 1000-1050 m, 9 Jun 1977, Schunke V 9667 (MO, U). BOLIVIA. LA PAZ: Mapiri, Jul-Aug 1892, Bang 1526 (US, mixed coll. with type of Solanum lindenii); Sud Yungas, La Paz-Calacoto, 69 km to the E, near Illimani dam, generating station Ikiko, 3100 m, 31 Dec 1980, Beck 3887 (F, NY); 4 km toward Chulumani on old rd. from Unduavi, 3000 m, 20 Nov 1983, Beck 8606 (NY); Unduavi, rd. to Coroico, 10500 ft, 16 Oct 1950, Brooke 6908 (F, NY); Prov. Sud Yungas, Sirupaya near Yanacachi, 2100 m, 16 Nov 1906, Buchtien 325 (NY); Prov. Nor Yungas, Unduavi, 3200 m, Feb 1914, Buchtien 464 (F, GH, K); Unduavi, Feb 1913, Buchtien 5853 (US); Prov. Nor Yungas, along rd. between Caranavi and La Paz, near the summit at jct. to detour to rd. to Chulumane, ca. 2800 m, 29 Nov 1980, Croat 51706 (MO); Prov. Nor Yungas, near Unduavi, 3000 m, 15 Apr 1939, Eyerdam 25142 (F, K); near village of Unduavi, 2850 m, 15 Mar 1984, Fournet 408 (F); Prov. Sud Yungas,ca. 1.8 km WSW of Unduavi, 3300 m, 16?19'S, 67?55'W,29 Oct 1984, Nee & Solomon 30189 (F, K, NY); Prov. Nor Yungas, 2 km by rd. (ca. 1 km by air) NE and below Chuspipata, 2950 m, 16?17'S, 67?49'W, 29 Oct 1984, Nee & Solomon 30214 (BH, K, NY, US); Unduavi, 3000 m, Dec 1855, Pearce s.n. (K); Unduavi, 8000 ft, Oct 1894, Rusby 794 (GH, K, NY, US); Prov. Nor Yungas, 1.6 km below Chuspipata on old rd., 3100 m, 16?18'S, 67?48'W, 10 Aug 1981, Solomon 6018 (MO, NY); Prov. Sud Yungas, 3.1 km SE of Unduavi bridge, on old rd., 3000 m, 16?19'S, 67?53'W, 6 Nov 1982, Solomon 8697 (NY); 2.4 km below Chuspipataon rd. to Chulumani,2950 m, 16?19'S,67?49'W, 4 Mar 1983, Solomon 9675 (MO, NY); prov. Sud Yungas, 1.8 km W of Unduavi on rd. to La Paz, 3300 m, 16?18'S, 67?55'W, 21 Mar 1984, Solomon et al. 11952 (NY); Prov. Murillo, Zongo valley, 25.2 km below the dam at Lago Zongo, 2700 m, 16?08'S, 68?07'W, 19 Jan 1985, Solomon 13096 (MO, NY); 6 km NE of Chuspipata on the rd. to Coroico, 2700 m, 16?17'S,67?48'W,21 Feb 1986, Solomon 14970 (MO, NY, U); 4 km NE of Chuspipata on the rd. to Coroico, 2800 m, 16?18'S, 67?48'W, 21 Feb 1986, Solomon 14983 (MO, NY); valley of Rio Zongo, 23.8 km N of the pass, 2900 m, 16?08'S, 68?07'W, 18 Mar

FLORA NEOTROPICA 1987, Solomon 16400 (MO, NY); 3.5 km NE of

on rd. to Coroico,2900 m, 16?17'S,67?49'W, Chuspipata 21 Jan 1988, Solomon 17607 (K, MO, NY); Rio Aceramana,10,800ft, 24-28 May 1926, Tate711 (NY). Solanum acuminatum is closely related to S. nudum

but has largerflowers and longerpedicels in fruitthan the latterspecies. The plantshave a distinctiveappearance, both in life andon the herbariumsheet. The new growthis usually deep purple,andthe leaves are quite shiny on the uppersurfaces.The inflorescencesremain on the stems afterthe fruitsdrop,anda shoot will often have several pseudoterminalinflorescences with the continuing growth from several generationsovertopping the dry inflorescenceaxes on the lower stem. The leaves usually dry darkin color, and the trichomes in the vein axils of the abaxialleaf surfacesarequitevariable in density and are often branched. NearUnduaviin the Departmentof La Paz, Bolivia, a pubescentformof Solanumacuminatumoccurs.This has been called S. pearcei, but althoughmore densely pubescenton theabaxiallaminae,theseplantsagreewith the type ofS. acuminatumin all otherrespects.The type of S. pearcei is quite variablein leafpubescence, with some of the stems bearingleaves with trichomesconfined to the vein axils and otherswith trichomesover the entireabaxialleaf lamina. The plate in Ruiz and Pavon's Flora Peruviana et

Chilensis(1799; see Fig. 36) shows the inflorescences as pseudoterminal,and is a good match for the specimens at MA. Several specimens of Solanum acuminatumare includedin the type folderat MA, and I have selected the one most closely matchingthe plate as the lectotype. 11. Solanum aphyodendron S. Knapp, Ann. Missouri Bot. Gard. 72: 565. 1985. Type. Panama. Chiriqui:along QuebradaAleman, 8 mi N of Los Planes de Horito, IRHE Fortuna Hydroelectric Project, 1200 m, 8?45'N, 82?12'W, 13 Mar 1982, Knapp, Kress & Hammel 4136 (holotype, BH;

isotypes, K, MO). Figs. 19A,B, 38 Bassoviafoliosa Brandegee,Univ. Calif. Publ. Bot. 6: 373. 1917. Type.Mexico.Veracruz: Zacuapan, Dec 1915, Purpus 7565 (holotype, CAL-n.v.; isotype, MO). Not Solanum foliosum Link.

Solanum nudumof authors, not of Dunal, Solan. syn. 20. 1816. Solanum parcebarbatum of C. V. Morton & Standl.,

Publ. Field Mus. Nat. Hist., Bot. Ser. 18: 1088. 1938. Pro parte,not of Bitter. Shrubsor small trees, 3-15 m tall;young stems and leaves glabrousor with a few scattereduniseriatetrichomes, dryingblackish;barkof older stems greenish-

TAXONOMIC TREATMENT

77

Herbariu Solann

lusaesBritannici

aphyodendron S. Knapp

near Fortuna Dan. PALA&a: Chiriqui; beside Secondary stream. vegetation 8'43'N 82?14'W, 1200 m. Snail shrub at top of steep bank. Pale wood, Arching, spreading bablt. brittle. Sepals tipped purplish. Flowers white, anthers bright yellow. 1 March 1985. R.J. hnpshire

i C. Whbitefoord 200.

5cm.

FIG. 38. Solanum aphyodendron S. Knapp. Panama. Hampshire & Whitefoord 200 (BM).

78

white and rough; stems slightly winged from the decurrentpetiole bases. Sympodialunitsdifoliate, geminate. Leaves narrowly ovate, widest at or just below the middle, dryingpale grayish-green,glabrousabove and below except for tufts of uniseriate trichomes in the axils of the mainlateralveins beneath,the trichomes ca. 1 mm long, white;majorleaves 8.5-15 x 3-6.1 cm, with 8-10 pairs of main lateralveins, these depressed above,raisedandpale greenish-whitebeneath,the apex acuteor acuminate,the base cuneateor slightly attenuate, with a small wing on the petiole; petioles 0.6-1.7 cm long; minor leaves differing from the majorones only in size, 3.2-8.6 x 1-4 cm, the apex acute or acuminate,the base cuneate;petioles 4-8 mm long.Inflorescences opposite the leaves, simple, 1-2.6 cm long, 20-30-flowered, glabrousor occasionally with a few scattereduniseriate trichomes distally;pedicel scars closely spaced,occasionallyoverlapping,beginningca. 1 cm fromthe base of the inflorescence.Buds globose, the corollasoon exertedfromthe calyx, pubescentwith uniseriatetrichomes0.1-1 mm long.Pedicels at anthesis deflexed, 0.7-1.1 cm long, taperingfrom the calyx tube to a slender base ca. 0.5 mm diam.Flowers with the calyx tube ca. 1 mm long, crateriform,the lobes broadlytriangular,0.5-1 mm long, densely to sparsely pubescent with uniseriatetrichomes 0.1-1 mm long, the margins of the lobes white and scarious; corolla white,oftendryingwitha lavendertinge,1-1.2 cm diam., lobed nearlyto the base, the lobes slightly reflexed at anthesis, the tips and margins of the lobes minutely papillose; anthers 2.5-3 x 1-1.5 mm, poricidal at the tips, the pores teardropshaped;free portionofthefilaments ca. 0.25 mm long, the filament tube ca. 0.5 mm long;ovaryglabrous;style5-7 mm long,straight;stigma not distinct fromthe body of the style, minutelypapillose. Fruita globose,brightgreenbery, ripeningyellowgreen, 1-1.2 cm diam., remainingharduntil maturity, then softeningvery quickly;fruitingpedicelsdeflexed, woody, 1.5-2.3 cm long, ca. 1 mm diam. at the base. Seeds tan, flattened-reniform,2.5-3 x 1.5-2 mm, the surfacesminutelypitted,the marginsincrassate,pitting of marginslargerthan that of the body. Chromosome number:n = 12 (voucherKnapp887, Knappetal. 4325, Knapp & Mallet 6791). Distribution (see Fig. 42). Widely distributedin secondgrowthat middle-to high-elevations,800-2500 m, from Mexico to northernBolivia. Selected specimens examined. MEXICO. S.loc., 1865-1866, Hahn s.n. (P); s.loc., Bourgeau s.n. (P); s.loc.,

Sesse, Mocino et al. 1403 (F); s.loc., Sesse et al. 5346 (F); CHIAPAS:Tenejapa,paraje of Koltol Te', 4750 ft, 10 Jul 1964, Breedlove 6121 (DS, US); Paraje of Yehts 'Uk'um, 4500 ft, 23 Nov 1964, Breedlove 7478 (DS); along rd. to Pueblo Nuevo Solistahuacan,1 mi N of Jitotol,

FLORA NEOTROPICA 5200 ft, 12 Feb 1965, Breedlove 8948 (F, MEXU, US); La Trinitaria, Lago de Monte Bello, 25 mi W of La Trinitaria, 5100 ft, 13 Apr 1965, Breedlove 9718 (BM, DS, US, WIS); Paraje of Sibakte'el, 5800 ft, 6 Aug 1966, Breedlove 14733 (DS, MEXU); Jitotol, 7 mi N of Jitotol on a side rd. to an oil well, 6700 ft, 28 Aug 1966, Breedlove 15368 (DS, NY); E of Laguna Tzikaw, Monte Bello National Park, 1300 m, 13 May 1983, Breedlove 35149 (DS); Motozintla de Mendoza, near Niquivil at the jct. with a small side ridge to Cerro Boquer6n, 2600 m, 16 Dec 1976, Breedlove 42767 (MO); near Paraje Kulak'tik, 1675m, 8 Jan 1982, Breedlove & Almeda 56837 (CAS); Uni6n Juarez, Bosque de Pino, 1360 m, 18 Nov 1977, Calzada et al. 3711 (F); Lagos de Montebello, along gravel rd. from blacktop to Dos Lagunas, 3 mi W of Dos Lagunas, 1460 m, 18 Jan 1969, Croat 46671 (BH, MO); along Mexican Hwy. 195, 2 mi N of Colonia San Jos6, 8 mi S of Lay6n, 1500 m, 3-4 Jun 1973, Hansen et al. 1689 (BH, US, WIS); along Mexican Hwy. 195, 2 mi N of Colonia San Jose, 8 mi S of Lay6n, 1800 m, 4 Jun 1973, Hansen et al. 1690 (US, WIS); Volcan Tacana, Chiquihuites, 1920 m, 10 Feb 1969, Herndndez M. 515 (DS, NY); Mt. Ovando, 10 Dec 1938, Matuda 252 (MEXU, US); Mt. Pasitar, 29 Dec 1936, Matuda 465 (MEXU, US); Mt. Tacana, 1000-2000 m, Aug 1938, Matuda 2466 (F, MEXU, US); Mt. Pastal, 12 Apr 1948, Matuda 17683 (F, MEXU); near Finca El Suspiro, 11 Jan 1953, Miranda 7654 (US); Pueblo Nuevo at Simoj6vel, 29 Dec 1959, Miranda 9160 (MEXU, US); near Tumbala, 4000-5500 ft, 20 Oct 1895, Nelson 3332 (US); Finca Mexiquito, Jul 1913, Purpus 6980 (F, NY, US); Cerro del Boquer6n, Jun 1914, Purpus 7318 (F mixed collection with S. narcoticosmum Bitter, MO, NY mixed collection with Lycianthes heteroclita (Sendtn.) Bitter); Ray6n, 9 mi NW of Pueblo Nuevo Solistahuacan along rd. between Rinc6n Chamula and Ray6n, 580 ft, 17?30'N, 92?40'W, 12 Oct 1971, Thorne & Lathrop 46953 (DS, RSA/POM); Tenejapa, paraje Balum K'aual, 8400 ft, 13 Apr 1966, Shilom Ton 811 (DS, F, LL, NY); Tenejapa, paraje of Kulak'tik, 5500 ft, 5 Dec 1966, Shilom Ton 1712 (DS, NY, WIS); Tenejapa, paraje of Mahosik', 4800 ft, 12 Sep 1966, Shilom Ton 1937 (DS, F, US, WIS); Pueblo Nuevo Solistahuacan, along ridge, 6500 ft, 15 Aug 1967, Shilom Ton 2865 (DS, F, WIS); ridge above Pueblo Nuevo Solistahuacan, 6500 ft, 3 Apr 1968, Shilom Ton 3923 (DS, WIS); 19 km NE of Bochil, ca. 1400 m, 25 May 1985, Thomas & Villasehor 3634 (NY). GUERRERO: Las Golondrinas, 29 km Ne of El Paraiso on rd. to Puerto del Gallo, 1810 m, 5 Jun 1983, Martinez & Ramamoorthy 5214 (NY); Lanada, 5 km SW of Taxco, Taxco-Ixcatcopan rd., 1790 m, 7 Jul 1982, Martinez Salas & Soto Niunez 1255 (CAS); 5 km E of Omiltemi, Chilpancingo, 2300 m, 1 Sep 1962, Rzedowski 16019 (US); 25 km NE of Paraiso, toyac, 1640 m, 5 Jun 1983, Soto Ntunez et al. 5235 (NY); 29 km W of Chilpancingo, 1 km W of Omiltemi on rd. to Las Joyas, 2250 m, 6 Jun 1985, Thomas & Contreras 3716 (NY). HIDALGO: Chapulhuacan, 1300 m, 12 Jul 1937, Lundell & Lundell 7192 (LL, NY). JALISCO: Cerro El Almeal, 4-5 km SE of Estaci6n Biol6gica Las Joyas, Sierra de

TAXONOMIC TREATMENT79 ManantlanOccidental, 2050-2100 m, 19?34'N, 104?15'W, 22 Dec 1984, Cochrane et al. 10627 (NY); SW foothills of the Nevado de Colima, ca. 10 mi SW of Atenquique on the Tonila rd., 1600 m, 5 Apr 1951, McVaugh 11801 (NY, US); Sierra del Halo, lumber rd. leaving the Colima hwy. 7 mi SSW of Tecalitan and extending SE toward San Isidro, 2000-2300 m, McVaugh & Koelz 1361 (US); above Ahuacapan, rd. to Corralitos 10-12 mi SSE of Autlan, 1500-1800 m, 29 Sep 1960, McVaugh 19586 (US); 12 mi SW of Chante along rd. to Sierra Manantlan, 5500 ft, 4 Feb 1975, Gentry & Gentry 23510 (US). MEXIco: Temascaltepec, Rinc6n del Carmen, 1460 m, 7 Sep 1932, Hinton 1004 (US); Temascaltepec, Rinc6n, 1960 m, 27 Apr 1933, Hinton 3729 (NY); Temascaltepec, Tejupilco, 1340 m, 9 Apr 1934, Hinton et al. 5753 (LL, NY, US). MICHOACAN: La Tzaracua, 12 km S of Uruapan, 1440 m, 25 May 1980, Soto Nunez & Romdn de Soto 2220 (CAS). MORELOS: Sierra de Ocuila, 18 Sep 1941, Lyonnet 3329 (US). NAYARIT: Along hwy. 28, between Tepic and Jalcatlan (rt. 66 on some maps) at km 14-16, 950-1050 m, 8 Jan 1979, Croat 45256 (MO); OAXACA: 10 mi S of Sola de Vega along rd. to Puerto Escondido, 7000 ft, 8 May 1965, Breedlove 9855 (US); along Hwy. 131 between Oaxaca and Puerto Escondido, 19.5 km by rd. S of Sola de Vega, ca. 2150 m, 16?25'N,97?02'W,25 Jun 1986, Diggs et al. 3973 (NY); 1 km NE of Ayutla, 1825 m, 17?02'N, 96?04'W, 17 Dec 1985, Nee & Martin 32207 (NY). VERACRUZ: Huatuaco, rd. to Tepampa 4 km from the hwy. between Huatusco and Coscomatepec, 1580 m, 14?10'N, 97?00'W, 10 Aug 1979, Avendano & Calzada 411 (F); Orizaba, 5200 ft, 24 Apr 1938, Balls 4312 (BM, US); Orizaba, 1886, Botteri s.n. (US); Orizaba, Botteri 848 (BM, P), Botteri 862 (US); environs of Orizaba, Botteri & Sumichrast 1857 (P); Orizaba, 12 May 1865(6), Bourgeau 240 (US); Coscomatepec, 5 km N of Tetelcingo, 2000 m, 12 Aug 1969, Bretting 13 (F); slopes of San Martin, 183 I'N, 94?49'W, Apr 1971, Calzada 251 (CAS, F); Cerro de Macuiltepetl, Xalapa, 9 Jun 1976, Calzada 2424 (F); Tezonapa, 240 m, 18?39'N, 96?41'W, 20 Jun 1973, Gdndara & Dorantes 147 (F); Ixtoteno, 5 km from Atzalan on rd. to Tlapacoyan, 10 Oct 1969, Lot 238 (F); Jalapa, 6 Sep 1936, MacDaniels 934 (BH, F); Mun. Xico, 6 km NE of Xico, 1500 m, 19?27'N, 97?01'W, 12 May 1973, Mdrquez & Gdndara 101 (F); Orizaba, 1929, Muller 3043 (NY); near Puente de Rieles, 4 km NE of Altotonga (6.5 km by rd.) on rd. to Tlapacoyan, 1750 m, 19?48'N, 97?13'W, 28 Jan 1980, Nee & Hansen 18717 (BH, F); near La Calavera, 10 km N of Altotonga (13 km by rd.) on rd. to Tlapacoyan, 1350 m, 19?51'N, 97013'W, 28 Jun 1980, Nee & Hansen 18643 (BH, BM, F); 8.5 km W of Chocaman, at summit of rd. to Xocotla, 1760 m, 19?01'N, 97?04'W, 18 Nov 1981, Nee 23218 (CAS, NY); 2.5 km (by rd.) E of Ayahualulco and 1.6 km (by rd.) W of Ixhuacan de los Reyes, 1900 m 19?02'N, 97?08'W, 7 Nov 1981, Nee 22959 (F); gorge at Puente Acabaloya, ca. 1 km SE of Xico Viejo and 5 km NW of Xico on trail between the two, 1600 m, ca. 19?27'N, 97?03'W, 31 Mar 1983, Nee & Taylor 26271 (NY); hills above and NE of Xico, 8 km NW of Xico,

1700-1900 m, ca. 19?38'N, 97?04'W, 5 Feb 1984, Nee & Taylor 29390 (NY); 3 km SSW of Zongolica along gravel rd. to Texhuacan, 1350 m, 18?39'N, 97?00'W, 8 Feb 1984, Nee & Taylor 29446 (NY); Jilotepec "El Esquil6n," 1390 m, 7 Jan 1976, Ortega et al. 104 (F, MO); Cerro de la Martinica, NE of Banderilla, 1680 m, 10 Feb 1976, Ortega et al. 152 (F, MO); Jalapa, Jardin Botanico Clavijero, 1300 m, 19?30'N, 96?55'W, 25 Apr 1978, Ortega & Calzada 780, (F); Xalapa, JardinBotanico Clavijero, 1250 m, 19?30'N,96036'W,22 Apr 1981, Ortega 0. 1843 (F); Chiconquiaco, Huerfano 39-A-O, 1400 m, 30 Jul 1967, Rosas R. 579 (U, WIS); s.loc. Scheide 136 (NY); 1 km S of Tequila along main rd. to Zongolica, 1800 m, 18?43'N, 97?04'W, 8 Feb 1984, Taylor & Nee 303 (NY); Los Reyes, 1600 m, 22 Mar 1976, Vacsquez S. 311 (F, NY); Mun. Tlazololapan, Tequila, 1170 m, 29 Mar 1976, VdsquezT 358 (F, NY); Mun. Xonamanca, Zongolica, 1350 m, 20 Mar 1976, Veldsquez L. 63 (F); Mun. Tlazololapan, Tequila, 1170 m, 19 Mar 1976, VelasquezL. 125 (F); Mun. Yecualta,Lomas de Santa Rita, 1350 m, 3 Jun 1971, VenturaA. 3268 (F); Mun. Huatusco, Coscontla, 1340 m, 14 Jan 1972, VenturaA. 4772 (DS, F, RSA/POM); Mun. Jilotepec, Rinc6n del Muerto, 1300 m, 4 Apr 1974, VenturaA. 9834 (DS, F); Mun. Naolinco, Cerro Prieto, 1500 m, 17 Jan 1975, VenturaA. 10820 (F); Totutla, 1250 m, 30 Jul 1976, VenturaA. 13103 (F); Mun. Atzalan, La Florida, 1600 m, 20 Mar 1978, VenturaA. 15102 (F); Xico, PuenteViejo, 1150 m, 1 Mar 1979, Ventura A. 15832 (F); Xalapa, JardinBotanico Rancho Guadalupe, 1300 m, 9 Sep 1976, Vovides 15 (F); near Orizaba, NW of La Perla, 2000 m, 26 Sep 1968, Weaver et al. 1747 (F); Orizaba,1866, Webers.n. (W); 2 km NW of Banderilla, Rancho La Mesa, 1450 m, 19 Jul 1976, Zold B. 512 (F). Near Coban, 1260GUATEMALA. ALTAVERAPAZ: 1440 m, 26 Mar-15 Apr 1939, Standley 69240 (F, JS); near San Jose, SE of Tactic, ca. 1500 m, 30 Mar 1939, Standley 69645 (F); mtns. E of Tactic, on rd. to Tamahi, 1500-1600 m, 9 Apr 1939, Standley 71437 (F); between Coban and Finca Chimot6, near Rubeltein, 800-1500 m, 25 Feb 1942, Steyermark 44158 (F, NY); Coban, 1350 m, Aug 1906, von TuerckheimH 1323 (F, US); 3 km W of San Julian, Tactic, 1600 m, 15?20'N, 90?15'W. 1 Feb 1969, Williams et al. 40415 (BM, F); ca. 5 km E of San Juan Chamelco, 1600 m, 15?26'N, 90?16'W, 8 Feb 1969, Williams& Wilson40759 (F); vic. of San Juan Chaielco, 1400 m, 9 Feb 1969, (?Williamsand?) Wilson40765 (F). BAJAVERAPAZ: Below Patal, 1550 m, 4 Apr 1941, Standley 90958 (F). CHIMALTENANGO: Rd. to Iximche Ruins, Tecpan, 2500 m, 12-23 Jan 1966, Molina R. et al. '6128 Cerro Brujo, vic. of Rio Negro, (F, NY). CHIQUIMULA: below MontaniaMontenegro, near village of Brujo, 15002000 m, 1 Nov 1939, Steyermark 30939 (F, US). EL PROGRESO:Sierra de las Minas, between El Jute de Cobara and Finca Piamontes, 1400-2400 m, 3 Feb 1942, Steyermark 43381 (F, NY). ESCUINTLA:S.loc., 1942, Aguilar 1602 (F); between Santa Maria de Jesus and Palin, 1800 m, 29 Dec 1938, Standley 61307 (F. US). GUATEMALA: Km 28 F.D.R. between San Lucas and Guatemala City, 2000 m, Molina R. et al. 16657 (F, TEX). HUEHUETENANGO: 3 mi NW of Santa Cruz Barillas along

80

FLORA NEOTROPICA

rd. to San Mat6o Ixtatlan, 6000 ft, 7 Aug 1965, Breedlove km NE of Taulabe, 1940 m, 28-29 Feb 1975, Nelson & 11651 (F); canyon of tributary of Rio Blanco 5 km W Vargas 2329 (MO); El Rinc6n, ca. 10 mi W of above Aguacatan, 2000 m, 11 Jan 1974, Molina R. et Siguatepeque, 1400-1500 m, Jun-Aug 1936, Yunckeret al. 30236 (F, MO); between kms 100-107 vic. of Campo al. 6050 (F, MO, NY, U). CORTES:Near Corinto, border de Bolas on way to El Mirador, Sierra Cuchumatanes, with Guatemala 55 km W of Puerto Cortes, 9-11 Aug 2000 m, 12 Sep 1971, Molina R. & Molina 26385 (F, 1975, Nelson et al, 2936 (MO). MORAZAN:Mt. San U); Chiantla, 6700 ft, 15 Dec 1934, Skutch 1949 (F, NY, Juancito, 7000 ft, 19 Jun 1948, Glassman 1666 (F, NY, US); 13 km W of Huehuetenangonear Puente de Xinax6, TEX, WIS); Cerro de Uyuca, 1600-2000 m, 10-20 Mar 1800 m, 30 Dec 1940, Standley 81575 (F, US); Rio Pucal, 1951, Morton 7167 (US); E slopes of Pefia Blanca, Cerro San Juancito, 1900-2000 m, 22 Mar 1951, Morton ca. 14 km S of Huehuetenango, 1780 m, 4 Jan 1941, Standley 82383 (F, US); spring of Rio San Juan near 7244A (BM, US); Cerrode Uyuca, ca. 1600 m, 2 Mar 1947, Standley 4827 (F); Cerro de Uyuca, along trail from Las Aguacatan, 1800 m, 6 Dec 1962, Williams et al. 22501 (F, NY, US, WIS). QUEZALTENANGO: Slopes of Volcan Flores to La Labranza, 1600-1750 m, Oct-Dec 1948, de Zunil, at and above Aguas Amargas, 2430-1850 m, Standley 15651 (F); Mt. Uyuca, 1800 m, 8 Jul 1946, 17 Feb 1939, Standley 65322 (F); above Santa Maria Williams & Molina R. 10029 (F); Mt. Uyuca, 1600 m, 2 Mar 1947, Williams & Molina R. 12096 (F); SW of San de Jesis, 1650 M, 1 Mar 1939, Standley 67194 (F, US); Los Positos SW of San Martin Chile Verde, 1500 m, 8 Juancito, 2000 m, 21 May 1947, Williams & Molina R. Mar 1939, Standley 67901 (F, US); near Vuelta de Tigre 12765 (F). OCOTEPEQUE: Agua Caliente (Guatemalan below Santa Maria de Jesus, ca. 150 m, 11 Mar 1939, border)-Santa Rosa de Copan rd., 18.1 mi E of Santa Fe, 26.8 mi SW of bridge over Rio Huiguito near village Standley 68169 (F, US); Aguas Amargas, W slope of of Cucuyagua Copan, 1800 m, 14?28'N, 89?15'W, 28 Jan Volcan de Zunil, 2450 m, 14 Jan 1941, Standley 83326 (F); along rd. above Santa Maria de Jesuis, 1680 m, 25 1987, Croat & Hannon 63804 (MO, NY); 35 km NE of Jan 1941, Standley 84878 (F, US); El Pocito, S of San Nuevo Ocotepeque, rd. to San Pedro Sula, 1870 m, 12 Martin Chile Verde on rd. to Colomba, 2200 m, 27 Jan Jun 1895, Martinez S. & Tellez 12989, 13016 (NY); 1941, Standley 85008 (F, US); El Pocito S of San Martin Belen Gualcho, 40 km E of Nueva Ocotepeque, 1500 Chile Verde on rd. to Colomba, 2200 m, 27 Jan 1941, m, 29 Jun-3 Jul 1976, Nelson et al. 3628 (MO); around Belen Gualcha, 40 km E of Nueva Ocotepeque, 1500Standley 85018 (F); El Pocito S of San Martin Chile Verde on rd. to Colomba, 2200 m, 27 Jan 1941, Standley 2000 m, 29 Jun-3 Jul 1976, Nelson et al. 3817 (MO). 85019 (F, US); region of Las Nubes, S of San Martin YORO:El Portillo Grande, 4000 ft, Jul 1937, van Hagen Chile Verde, 2250 m, 27 Jan 1941, Standley 85159 (F); & van Hagen 1006 (F, NY); Quebrada Olotillo, 15 km above Mujulia, between San Martin Chile Verde and from Yoro, 1100 m, 8 May 1956, Molina R. 6829 (US). EL SALVADOR. CHALATENANGO: E slope of Los Colomba, 1800 m, 1 Feb 1941, Standley 85452 (F); vic. of Fuentes Georginas, slopes of Volcan de Zunil, 2300Esemiles, 2100-2300 m, 14?21'N, 89?09'W, 1 Apr 1942, 2500 m, 3 Feb 1941, Standley 85865 (F, US); along old Tucker1186 (F, LL). SANTAANA:Along rd. to CerroMonte rd. between Finca Pirineos and Patzulin, 1200-1400 m, Cristo at Los Planes at km 22, 1800 m, 31 Jul 1977, 9 Feb 1941, Standley 86620, 86780 (F, US); W slopes Croat 42332 (MO). SONSONATE: Cerro Verde, 1800 m, of Volcan de Zunil, opposite Santa Maria de Jes6s, 1500 25 Feb 1968, Molina R. & Montalvo 21726 (F, NY). NICARAGUA. GRANADA:Mombacho Volcano, 960 m, 21 Jan 1940, Steyermark35124 (F). SACATEPEQUEZ: Near Pastores, 1560-1650 m, 14 Dec 1938, Standley m, 5 Jul 1923, Maxon et al. 7781 (F, US). JINOTEGA: 59937 (US). SAN MARCOS:Finca Vegel, near Rodeo, Along hwy. 3 from Jinotega to Metagalpa, 5-8 mi SW 900 m, 15 Mar 1939, Standley 68906 (F); barranco of of Jinotega, 1500 m, 7 Aug 1977, Croat 43055 (MO). Rio (Tonana) Suchati between Canjula and La Uni6n COSTA RICA. ALAJUELA: Finca de los Ensayos ca. 11 mi NW of Zarcero, 850 m, 15 Aug 1977, Croat 43541 Juarez, near SE portion of Volcan Tacana, 2000-3000 m, 2 Feb 1940, Steyermark 36278 (F); barrancos S and (MO); upper drainage of Rio Peiias Blancas below W of town of Tajamulco,NW slopes of Volcan Tajamulco, Monteverde Cloud Forest Reserve, 1250-1350 m, 2300-2500 m, 25 Feb 1940, Steyermark 37544 (F). 9?17'N, 84?56'W, 25-26 Feb 1977, Gentry 3812 (F); SANTA ROSA: Jumaytepeque, 6000 ft, Aug 1892, Heyde along Rio Alajuela, Alajuela-Carrizal rd. 5 km S of & Lux 4105 (F, K, MO, NY, US); near El Molino, 600 Carrizal, 1200 m, 24 Mar 1974, Hartshorn 1423 (F, MO); m, 26 Nov 1940, Standley 78379 (US); Volcan Monteverde, rd. to Peias Blancas (over Continental Tecuamburro, along trail to San Francisco Tecuamburro Divide) ca. 1 km below the Divide, Atlantic slope. 1400 on summit of volcano, N of Chiquimulilla, 250-2000 m, 10?25'N, 84?50'W, 13 Apr 1981, Knapp & Mallet m, 20 Dec 1939, Steyermark33149 (F). SUCHITEPEQUEZ:865 (BH, CR, MO); Santa Maria National Park, rd. down SW lower slopes of Volcan de Zunil between Finca de Caribbean slope 1 km E of rd. summit, 4 km W of E Asturias and Finca Alto Mira, NE of Pueblo Nuevo, side of Park, 4 km E of colored house at jct. of rd. to 1000-2000 m, 1 Feb 1940, Steyermark 35349 (F). Hacienda Santa Maria, 600 m, 10?27'N, 85?17'W, 7 Feb HONDURAS. COMAYAGUA:Montafia La Choca, 1978, Liesner 5139 (MO); Zarcero, Palmiras, 6000 ft, Cordillera Comayagua, near Coyocutena, 1200 m, 22 16 Aug 1937, Smith A152 (F, MO, US); region of May 1956, Molina R. 7113 (F); Montafia El Cedral, Zarcero, 4700 ft, 27 Aug 1937, Smith A236 (F, MO); Cordillera Montecillos, 1600 m, 24 May 1956, Molina region of Zarcero, 5500 ft, 6 Jan 1938, Smith Hll (F); R. 7194 (F, US); Farfallon El Pizote, Cerros Cedral 6 San Carlos, Sucre, 1025 m, 2 Mar 1939, Smith H1638

TAXONOMIC TREATMENT (F, US); vic. of Fraijanes, 1500-1700 m, 12-13 Feb 1936, Standley & Torres R. 47454 (US); Naranjo, 5600 ft, 5 Apr 1928, Stork 1852 (F); Alfaro Ruiz, Palmira,7000 ft, 28 Apr 1937, Stork 4158 [Smith 58] (LL); near Rio Tapesco, ca. 10 km N of Zarcero,CordilleraCentral,1600 m, 6 Feb 1965, Williamset al. 28937 (F, MO); Cordillera Centralnear San Juan de Laja ca. 15 km N of Zarcero,ca. 1350 m, 7 Feb 1965, Williams et al. 29033 (F, MO). CARTAGO:Environs of Cartago, 1417 m, Oct 1894, Briolley 8998 (US); Cartago, 1650 m, Dec 1887, Cooper 5868 (F, US); overlooking Rio Grande de Orosi, ca. 3 km SE of Tapanti, 1350 m, 16 Apr 1967, Lent 832 (BM, DS, F, WIS); 5.5 km above Tobosi on rd. to Frailes, 1800 m, 30 Jul 1967, Lent 1158 (F); 5 km SW of Tobosi, 1900 m, 9?50'N, 84?01'W,Lent 3160 (F, MO); El Mufieco on the Rio Navarro, 1400-1500 m, 6-7 Mar 1926, Standley & Torres R. 50939 (US); near La Sierra ca. 25 km S of Cartago, Cordillera de Talamanca, 2000 m, 23 Jan 1965, Williamset al. 28048 (F); GUANACASTE: Along rd. between Santa Elena and Monteverde, ca. 2 mi from Santa Elena-Monteverde jct., 1350 m, 7 Feb 1979, Croat Vic. of Alto la Palma between 47100 (BH, MO). HEREDIA: Paracito and Bajo La Hondura, 1500 m, 14 Jan 1978, Croat 44484 (MO); 3 km N of Salto La Paz and 8 km N of Vara Blanca on rd. toward Puerto Viejo, ca. 1200 m, 25 Dec 1974, Nee et al. 14016 (F, WIS); ParqueNacional Braulio Carillo, Sendero Camino Viejo de las Mulas, Bajo la Hondura, 2 Jun 1984, Sinchez & Zamora 536 (F); Vara Blanca de Sarapiqui, N slope of Central Cordillera, 1500-1750 m, Jul-Sep 1937, Skutch 3286 (MO, NY, Monteverde, pasture edges in village, US). PUNTARENAS: 1400-1500 m, 15 Aug 1976, Dryer 566 (CR, F); Monteverde, 1400-1500 m, 15 Aug 1976, Dryer 607 (CR, F); Monteverde, near the Cloud Forest Reserve, 1520-1560 m, 28 Oct 1976, Dryer 893 (CR, F); ca. 2 km S of village of Monteverde, 1500-1550 m, 10?18'N, 84048'W, 18-21 Mar 1973, Gentry & Burger 2698 (CR, F, MO); ca. 2 km S of Monteverde, 1500-1550 m, 10?18'N, 84?48'W, 18-21 Mar 1973, Gentry & Burger 2719 (CR, F, MO), 25 Feb 1977, Gentry 3793 (F); Finca Las Cruces, 3 km S of San Vito de Java, 4000 ft, 11 Feb 1971, Gillis & Plowman 10129 (F, TEX); Mr. W. James' property, Monteverde, 1500 m, 18 Oct 1963, Jimenez M. 1246 (F); Monteverde, on rd. to Pefias Blancas (over Continental Divide), 1500 m, 10?25'N, 84?50'W, 25 Feb 1981, Knapp 834 (BH, CR). SAN JOSE: Rd. between Alto la Palma and Bajo la Hondura, 1400-1500m, 24 Feb 1978, Almeda & Nakai 3902 (CAS); Tablazo above San Lorenzo de Tres Rios, 1200-1900 m, 9?50'N, 84002'W, 25 Jun 1985, Barringer & Christenson 3304 (NY); Los Angeles de San Ram6n, 1075 m, 20 Nov 1923, Brenes 3947 (CR, F); La Palma, 1250 m, 10 Sep 1924, Brenes 4063 (CR, F); La Palma de San Ram6n, 1050 m, 17 Apr 1927, Brenes 5432 (F, NY); Tablazo, 28 Apr 1946, Echeverria 379 (CR, F); near Quebrada Grande, 3 km NW of Cascajal, 1750 m, 12 Dec 1971, Lent 2295 (CR, F, MO, U); Cerro del Tablazo, 1500 m, 8 Jun 1941, Le6n 802 (F); La Palma, 20 Aug 1970, Schnell 1120 (GH, NY); hills above Aserri, 14 Jun 1955, Schubert & Rogerson 701 (US); vic. of El General, 1190 m, Jan 1936, Skutch

81 2417 (MO, US); vic. of El General, 1430 m, Dec 1936, Skutch 2987 (MO, NY, US); La Hondura, 1300-1700 m, 2-4 Mar 1924, Standley 36199 (US); La Hondura, 12001500 m, 9 Mar 1926, Standley & Valerio 51840 (F, US); Rio Segundo, 2000 m, 10 May 1840, Tonduz1795 (BM, US); Rio Candelaria, 1215 m, Apr 1893, Tonduz 7892 (US); Alto de La Palma, ca. 6 km N of San Jer6nimo, 15 Feb 1974, Utley & Utley 634 (F, NY); 2-5 km SE of Higuito on Calle Tablazo, 8-12 km SE of Desamparados, 1800-1900 m, 5 Sep 1975, Utley & Utley 3047 (CR, F). PANAMA. CHIRIQUI:Trail from Paso Ancho to Monte Lirio, upper valley of the Rio ChiriquiViejo, 15003000 m, 16 Jan 1939, Allen 1492 (F, MO, NY, US); Cerro Colorado, 1500 m, 26 Jul 1976, Antonio 1502 (MO, MY); 2 mi N of Volcan, 29 May 1970, Croat 10465 (F, MO, NY); between Bambito and Cerro Punta, 30 May 1970, Croat 10548 (MO); between Bambito and Cerro Punta, 30 May 1970, Croat 10573 (MO); vic. of Bambito, 30 May 1970, Croat 10626 (MO); between Cerro Punta and Bajo Grande, 22 Jul 1971, Croat & Porter 15998 (MO); ca. 2 km on the Cerro de la Muerte rd., near the Florida State camp, 6000 ft, 19 Apr 1969, Correa A. 1265 (MO); ca. 5.4 km from Hato de Volcan on rd. to Las Lagunas, 26 Apr 1969, Correa A. & Lazor 1480 (MO); between Rio Chiriqui Viejo and Nueva California. 16 Apr 1970, D'Arcy 4244 (MEXU, MO); Nueva California, 16 Apr 1970, D'Arcy 4249 (MO); cafetal beside "Sonrisa Chiricana",Nueva Suiza, 5700 ft, 8 May 1971, D'Arcv 5313 (MO); above Nueva Suiza, 6000 ft, 8 May 1971, D'Arcy 5325 (MO, MPU); above Cerro Punta, 6300 ft, 9 May 1971, D'Arcy 5374 (MO, MPU); N side of Cerro Pando, 6000 ft, 10 May 1971, D'Arcy 5402 (F, MO, WIS); N side of Cerro Pando, 6000 ft, 10 May 1971, D'Arcv 5406 (F, MO); above Boquete, 4500 ft, 12 May 1971, D'Arcv 5443 (F, MO, P); rd. side between Nueva California and Rio Chiriqui Viejo, 7 Aug 1972, D'Arcv & D'Arcy 6492 (F, MO, NY, P, RSA/POM, WIS); El Barni above Boquete, 1200-1800 m, 21 Nov 1975. D'Arcv 9863 (MO); La Popa above Boquete, 1500-2500 m, 20 Mar 1977, D'Arcy 10958 (MO); rd. from Nueva California to Rio Serrano, ca. 7 mi from Rio Chiriqui Viejo, 4200 ft, 7 Apr 1979, D'Arcy et al. 13063 (MO); Bajo Chorro, 6000 ft, 15 Feb 1938, Davidson 296 (F, IMO, US); valley of the Rio Chiriqui Viejo N of Volcan, 52005600 ft, 9 Dec 1966, Duke 9012 (MO); along rd. between Boquete and Cerro Horqueta, 2 Aug 1967, L)Duke A13720 (MO); Cerro Horqueta, 1500 m, 2 Aug 1967, Duke et al. 13690 (MO); NW of Boquete, Cerro Horqueta, 5000-5800 ft, 13 Dec 1966, Dwyer et al. 489 (MO, NY); NW of Boquete, Cerro Horqueta, 5000-5800 m, 13 Dec 1966, Dwyer et al. 555 (MO); Boquete, 5000 ft, 5 Sep 1967, Dwyer 6969 (MO); near Hotel Dos Rios, Boquete, 6 Aug 1970, Dwyer 7638 (MO); Finca Collins, 5000 ft, 7 Aug 1967, Dlwyer 7638 (MO); Cerro Horqueta, 4500-5500 ft, 1 Jul 1968, Dwyer & Lallathin 8759 (MO); Boquete, 20 Mar 1977, Folsom 2172 (MO); Cerro Colorado, 2.2 km from main rd., border with Bocas del Toro, 1700 m, 16 Aug 1977, Folsom 4896 (K, MO, NY); Cerro Hornito, 4500 ft, 8 May 1978, Hammel 3028 (MEXU, MO, NY); between Los Planes de Hornito and

82 Fortuna Lake, trail to Zarzo, 1200 m, 8?41'N, 82?13'W, 17 Mar 1985, Hampshire & Whitefoord 695 (BM, MO, NY); along rd. from Palo Alto to Rio Palo Alto, NE of Boquete, 1700 m, 8?47'N, 82?22'W, 15 Mar 1982, Knapp et al. 4273 (MO); Cerro Punta, 2000 m, 8 Sep 1971, Lao 329 (MO); 1 mi E of CaniasGordas, near Costa Rican border on rd. to Volcan, 800-1200 m, 26 Feb 1973, Liesner 279 (F, MO, NY, US); behind Florida State University cabin, 2.2 mi below Cerro Punta, 6000 ft, 23 Jun 1970, Luteyn 896 (MO); vic. of Boquete, 120 m, 3 Feb-15 Mar 1938, Bro. Maurice 706 (US); N of San Felix on Chiriqui/Bocas del Toro border, on Cerro Colorado copper mine rd. along Continental Divide, 5000-5500 ft, 5 May 1975, Mori & Kallunki 5911 (MO); Bajo Grande, 1-3 km E of town of Cerro Punta, 2000-2200 m, 24 Feb 1974, Nee 9982 (MO); Bajo Grande, 1-3 km E of town of Cerro Punta, 2000-2200 m, 24 Feb 1974, Nee 10000 (MO, MPU); along rd. at Quebrada El Velo, near Finca L6rida, 6 km NW of Boquete, 1700 m, 17 Mar 1974, Nee 10625 (MO); 17 km NE of San Felix on rd. to Cerro Colorado copper mine, 13-14 km by rd. NE of bridge over Rio San Felix, 1000 m, 18-19 Mar 1974, Nee 10755 (MPU, WIS); upper Rio Chiriqui Viejo, vic. of Monte Lirio, 1300-1900 m, 27 Jun-13 Jul 1935, Seibert 167 (MO); Nueva California, just across Rio Chiriqui Viejo, 13 Sep 1972, Tyson 6684 (MO); Cerro Punta, 6025 ft, 28 Jul 1938, White197 (F, MO); vic. of Callej6n Seco, Volcan de Chiriqui, 1700 m, 17 Jul 1940, Woodson & Schery 487 (MO, US). VERAGUAS:Above Santa F6 beyond Escuela Agricola Alto Piedra, 1.8 mi beyond fork in rd. on Pacific slope, Cerro Tute, 5 Apr 1976, Croat 34156 (MO. NY); N of Santa F6 on property of Escuela Agricola Alto Piedra, "Girasol," 16 Oct 1974, Mori & Kallunki 2511 (MO); Cerro Tute, E slopes, 1 km beyond Escuela Agricola Alto Piedra above Santa F6, 9001200 m, 14 May 1981, Sytsma & Andersson 4644 (MO). COLOMBIA. of ANTIOQUiA: Km 10-15 Campamento-Las Brisas rd., 1570-1800 m, 20 Aug 1986, Callejas et al. 2500, 2505 (NY); Parque Nacional Las Orquideas, Finca La Guadulala, quebrada Horacio, affluent of Rio Venados, 1160 m, 6?30'N, 76?30'W, 1 Dec 1986, Callejas et al. 2942 (NY); Jardin-Rio Sucio rd., km 7 near QuebradaBonita, 2200 m, 5?33'N, 75?44'W, 25 Jan 1988, Churchill et al. 15872 (NY); Urrao, Incarnaci6n-Parque Nacional de las Orquideas, ca. 2000 m, 18 Jun 1981, Escobar 1845 (NY); between Rio Guapa and Rio Le6n, 100 m, 18 Mar 1948, Ruiz Landa et al. 118 (US); 4 km from Palmitas, 1700 m, 5 Mar 1949, Scolnik et al. 19An168 (F, US); rd. between Yarumal and Valdivia, region of Ventanas, (Gutierrez et al.) Valdivia 5 (US); rd. between Curamontaand Valparaiso, (Travecedo& Gouzy) Valparaiso3 (US). CAUCA:Popayan, Timbio in Hato-Viejo, 180 m, 14 Jul 1939, Cuatrecasas & Perez-Arbeldez 6070 (F, US); between Popayan and Cajeti, toward Tambo, 1700 m, 30 Dec 1942, Cuatrecasas 13819 (F, US); Cordillera Occidental, E slope, Cuchilla de Tambo, 1750 m, 23 Aug 1949, Idrobo & Ferncindez 246 (US); "La Gallera" Micay valley, near Rio Joaquim, Cordillera Occidental, 1400-1500 m, 2930 Jun 1922, Killip 7846 (NY); Popayan, 1500-2000 m,

FLORA NEOTROPICA Jan-Mar, Lehmann 4770 (BM, F, K, US); Cordillera Occidental, San Antonio to Rio Ortega, 2000-2300 m, 2 Jul 1922, Pennell & Killip 8039 (NY); rd. from Tunia to Popayan, (Gutierrezet al.) Popaydn 1 (US); CUNDINAMARCA:Cordillera Oriental, 7.4 km W of El Salto de Tequendamaon rd. to El Colegio, then turn left on rd. to Victoria and proceed 3.8 km, 2000 m, 21 Jul 1972, Barclay et al. 3599 (US); Anolaima, 14 Jan 1954, David 4852 (US); Santana station, above Sasaima, 160-1700 m, 25-29 Jul 1945, Dugand & Jaramillo 3877 (US); Sasaima, Vereda de San Bernardo, Hacienda "La Victoria,"1700 m, 1 Sep 1946, Garcia-Barriga1213 (US); Cordillera Oriental, Finca "Alto Oscar" 11 km S of La Palma, 5600 ft, 10 Mar 1944, Little 7388 (NY, US). HUILA: Finca Cedral, Quebrada Urraca, Rio Fortalecillas above Vega Larga, 30 km E of Neiva, 1300 m, 2?58'N, 74?58'W, 18 Jan 1943, Fosberg 19765 (NY, US); Finca Merenberg E of Volcan Purac6, near Cauca border, 2300 m, 2?16'N, 76?12'W, 4 Apr 1986, Gentry 53982 (MO, NY). SANTANDER:Between Piedecuesta and La Vega, 2000-2500 m, 19-24 Dec 1926, Killip & Smith 15513 (NY, US); Rio Surata valley above Surata, 2000-2500 m, 5-6 Jan 1927, Killip & Smith 16531 (NY, US); vic. of Charta, 2000-2600 m, 1-11 Feb 1927, Killip & Smith 19089 (NY, US); vic. of Tona, 1900-2100 m, 17 Feb 1927, Killip & Smith 19505 (NY, US). TOLIMA:El Libano above San Jose, 1580-1620 m, 19 Jul 1947, Garcia-Barriga 12239 (US); E slope Cordillera Central, Cerro Purima, 2540 m, 1 Aug 1980, Idrobo et al. 10370 (U). VALLE DE CAUCA:CordilleraOccidental, E slope between Bitaco and Yumbo, 1700 m, 5 Apr 1979, Cuatrecasas & Cuadros 28837 (US); Rio Nima above Tenjo, 1850 m, 2 Oct 1974, Maas & Plowman 1827 (BM, K, MO). VENEZUELA. MERIDA:QuebradaEl Oso, Palmira, Justo Bricefio, 1600 m, 3-11 Oct 1973, L6pez-Palacios & Bautista-Bautista 3481 (MO); TACHIRA:Between Delicias and Villa Paez, 1800-2000m, 3 May 1976, Badillo 7256 (MY); ParqueCazadero,QuebradaCazadero, 16 km NW of San Crist6bal, 650-900m, 7?54'N, 72?18'W, 3 May 1981, Liesner & Guaraglia 11724 (NY, VEN). SUCRE:Cerro TurumiquireN facing slopes above La Trinidad, SW of Cocollar, 2100-2200 m, 5 May 1945, Steyermark 62540 (F, VEN). TRUJILLO:El Alto, ca. 1950 m, 5 Sep 1975, Benitez de Rojas 1977 (MY). ECUADOR. LOJA:Km 25, Loja to San Lucas, 2200 m, 15 Sep 1961, Dodson & Thien 635 (MO, US); Parque Educacional Recreacional Universidad de Loja, vic. of Loja, 2100-2300 m, 4?02.02'S, 79011.87'W,28 Oct 1994, Knapp et al. 9086 (QCNE). MORONA-SANTIAGO: Along Rio Palora 1-4 km upstream from Arapicos, 800-900 m, 14 Apr 1981, Lugo S. 6080 (NY); Colonia Azuay 2 km from Arapicos, 800-900 m, 18 Apr 1981, Lugo S. 6100 (NY). PICHINCHA: Chaupi-Sagcha, Pululagua, ca. 6000 ft, 15 Apr 1951, Bell 440 (BM); just N of Tandapiacross from Rio Toachi (on Sto. Domingo-Quito rd.), ca. 1500 m, 0?20'S, 78?55'W, 11 Feb 1984, Knapp et al. 6272, 6274 (BH, K, QCA, QCNE); Quito-Aloag-Sto. Domingo de los Colorados km 94, 10 km S of rd., W slopes of Volcan El Coraz6n, 1300-1500 m, 0?21'30"S,78?51'15"W, 25 Dec 1986, Zak 1510 (IBE, K, MO, NY); Palltanga-

TAXONOMIC TREATMENT Yungilla-Llimbe rd., banks of Rio Chimbo, 1720 m, 7 Sep 1987, Zak 2805 (K, MO, NY); Quito-Aloag-Sto. Domingo de los Coloradoskm 94, 10 km S of rd., W slopes of Volcan El Coraz6n, 1300-1500 m, 0021'30"S, 78?51'15"W,20 Nov 1987, Zak 2994 (MO, NY). TUNGURAHUA: Cashurcu, 13 km W of Mera on Bafios-Mera rd., 1300-1400 m, 1?25'S, 78?10'W, 22 Jan 1984, Knapp & Mallet 6179 (BH, QCA, QCNE, US); Rio Topo (Rio Toro of maps), above village of Rio Negro, on BafosMera rd., 1300-1400 m, 1?22'S, 78?13'W, 22 Jan 1984, Knapp & Mallet 6181 (BH, K, QCA, QCNE, US). PERU. S.loc., 1840, Mathews 3249 (BM, NY); Ca. 7 km Sascoya, 1840, Mathews 3269 (K). AMAZONAS: above Pedro Ruiz on rd. to Pomacochas, ca. 1500 m, 5?58'S, 77?57'W, 3 Jul 1984, Knapp & Mallet 6562 (BH, K, US, USM); Bongara, Jalca zone 3 km S of Pomacocha, E of Shipasbamba trail, 2400 m, 20 Jun 1962, Wurdack La Mar, ca. 25 km 997 (BM, K, US, USM). AYACUCHO: walking distance SW of Hacienda Luisana and Rio Apurimac, ca. 12 km from Hacienda Santa Rosa, ca. 25 km from Tambo, E massif of Cordillera Central, 1570 m, 12?43'S, 73?50'W, 20 Aug 1968, Dudley 11879 (F); Carrapa between Huanta and Rio Apurimac, 2200 m, 5,6,17 May 1929, Killip & Smith 2306 (NY); La Mar, just below Huanhuachayoon the Capprichio-Puncu trail, W slope of Rio Apurimacvalley, 1590 m, 12?43'S,73?47'W, 15 Jul 1970, Madison 10265-70 (F). CAJAMARCA: Upper Rio Zaia valley, ca. 5 km above Monte Seco, 1450-1500 m, 19 Mar 1986, Dillon et al. 4434 (F, NY); ca. 3 km ENE of Monteseco, 1800 m, 6 Jun 1987, Santisteban C. & Guevara B. 139 (F, NY). HUANUCO: Pachitea, Panao, 2700-2900 m, 3 Mar 1947, Ferreyra 1766 (US); Pachitea, Panao, 4 Mar 1947, Ferreyra 1796 (US); Mufa, ca. 7000 ft, 23 May-4 Jun 1923, Macbride 4027 (F); Huacachi, station near Muia, 6500 ft, 20 May1 Jun 1923, Macbride 4155 (F); rd. from Huanuco to Tingo Maria N of Carpish Pass, 54 km NE of Huanuco, 2310 m, 6 Dec 1981, Plowman & Rury 11164A (F); km 452 of Lima-Tingo Maria rd. near Carpish, ca. 2500 m, 2 Jun 1981, Young& Sullivan 572 (MO, NY). JUNiN: Tarma, at little bridge on Tarma-San Ram6n rd., 1 km above Matichacra, just below Carpapata, E side of Rio Tarma valley opposite mouth of Rio Huasihuasi, 11 km NNE of Palca, 2000 m, 4 Dec 1962, Iltis et al. 322 (K, US, USM, WIS); Pichis trail, between Yapas and Enefias, 1800 m, 28 Jun-8 Jul 1929, Killip & Smith 25618 (US); Schunke hacienda above San Ram6n, 1300-1700 m, Aug-Oct 1923, Schunke AO11 (US); Chanchamayo valley, 1500 m, Nov 1925-1927, Schunke 258 (F); Chanchamayo valley, Dec 1924-1927, Schunke 259 (F); Chanchamayo valley, Sep 1924-1927, Schunke 272 (F); Chanchamayo valley, 1200 m, Jul 1929, Schunke 534 (F); Tarma, Utcuyacu, 1900 m, 26 Feb 1948, Woytkowski 35381 (F). PASCO:Vic. of Oxapampa, ca. 2 km E of village, 1850 m, 10?35'S, 75?35'W, 10 Mar 1984, Knapp et al. 6321 (BH, K, NY, US, USM). PUNO: Sandia, Santo Domingo, 1550 m, 4 Nov 1939, McCarroll 52 (NY). BOLIVIA. COCHABAMBA: On rd. from Comarapa to Cochabamba, 4 km W of Santa Cruz border, 20 km by air and 28 km by rd. NW of Comarapa, 2525 m,

83

17?49'S,64?41'W,10 Feb 1987,Nee & Solomon34049 (NY); Churro,7 km by air ESE of Siberia,8.5 km SW of SantaCruzborder,2600 m, 17?50'S,64?42'W,5 Mar 1988, Nee et al. 36493 (NY); narrowcanyon of Rio MontePuncu,5 km NE of MontePuncu, 10 km by air NW of Epizana,2700-2750 m, 17?33'S,65?16'W, 10 Mar 1988, Nee & Solomon36632 (NY); Sacaba,Cerro de Incachaca,2500 m, 4 Sep 1921,Steinbach5753 (K). LA PAZ:Prov. Murillo, Valle de Zongo, above Jarca, 2100 m, 31 May 1980,Beck3601 (F);Prov.SudYungas, basinof Rio Bopi, San Bartolem6(nearCalisaya),750900 m, 1-22 Jul 1939, Krukoff10452 (F, K, MO, NY, U, US); 36.2 km N of dam at Lago Zongo, 1800 m, 16?05'S,68?03'W,18 Dec 1982, Solomon9163 (MO, NY); Prov.Nor Yungas,Serraniade Bella Vista, 16 km N of Carrasco(37 km N of Caranavi)on raodto Palos Blancos, 1500 m, 15?25'S, 67?34'W, 31 Oct 1984, Solomon& Nee 12649(MO,NY, U); Prov.Nor Yungas, Serraniade Bella Vista,17.6 km N of bridgeat Carrasco, 1600m, 15?40'S,67?28'W,11 Jun 1985,Solomon13981 (MO, NY). SANTA CRUZ: Prov. Valle Grande, Rio

Piraymiri,2900 m, Feb 1956, Cardenas5118 (US). Local names. Mexico. capulinde pajaro,yerba de zopilote; CostaRica. Puntarenas:zorrillo;Guatemala. AltaVerapaz:kaqisakyol(Quecchi);Chiquimula:huele de noche;Quezaltenango: hediondilla;Panama.Chiriqui: hiede hiede; Colombia.Antioquia:chuco amargo. Solanumaphyodendronis most similarandprobably closely relatedto S. trichoneuronof southernBolivia andArgentina,fromwhich it differsin its largerflowers, shorterinflorescences, and somewhat larger berries. Solanumaphyodendron is a commonspeciesof roadsides Central America andnorthernSouthAmerica throughout and it often forms large monospecific stands in open areas.Materialfrom CentralAmerica, Colombia, and Venezuelais quite uniform.Variabilityin pubescence increases in Peruvian and Bolivian specimens, with some specimenshaving shortertrichomeson the calyx lobes and denser tufts of trichomesin the vein axils. In Monteverdede Puntarenas,CostaRica,Solanum bloomsatirregular intervalsthroughout the aphyodendron year (Knapp, 1986a). Data fromherbariumspecimens and observationsof this species in Panamaand Ecuadorindicatethatits floweringbehavioris very muchthe samethroughout itsrange.AtMonteverde, S. aphyodendron is pollinatedby meliponinebees, primarilybyMelipona fasciata (Knapp,1986a).Theoccurrenceof largemonospecific standsof S. aphyodendroncombinedwith the foragingbehaviorofMelipona produceshigh fruitset in this species. The fruitsareeatenby small frugivorous bats and disappearsoon afterthey become ripe. Solanum aphyodendronhas long been confused with S. nudum,to which it is somewhatclosely related, but differs in habitat, flower size and pubescence of flower parts,andin the color andtextureof the barkof older stems. Solanumnudumoccurs in low-elevation

84

FLORA NEOTROPICA

?

~ *OLAUW

A

t .teno1tr CAAV

AWA

aa-.)

B

,

.

e--.

FIG. 39. Solanum caavurana Veil. A. plate 112 of Vellozo's Florae Fluminensis Icones, Vol. 2. B. plate 8 of Martius' Flora Brasiliensis.

thickets, sometimes on beach strands, and has dark brownbarkandglabrous,globose buds.The barkcolor remains consistent even on herbariumsheets, with S. aphyodendronbeing pale white or greenish andS. nudumbeing darkbrown or black. 12. Solanum caavurana Vell., Fl. Flumin. 86. 1829 [1825]. Type.Brazil. Rio de Janeiro:"Undequaque crescit,precipuesylvis excultis"(No specimensextant;lectotype,Vellozo,Fl. Flumin.Icones2: fig. 11. 1831 [1827], heredesignated). Figs. 35E, 39 Solanum acuminatum Ruiz & Pav. var. viridiflorum

Dunalin DC., Prodr.13(1): 147. 1852.Type.Brazil. Bahia: Circa Bahiam, Blanchet 48 (holotype, G-DC [F neg. 6777]; isotypes, BM, K, MPU, NY). Solanum fossarum Dunal in DC., Prodr. 13(1): 145. 1852. Type. Brazil. Bahia: Circa Bahia in fossis, Guillot s.n. (lectotype, P, here designated [F neg.

39170];isolectotypes,P [Mortonneg. 8195], frag. F, MPU). Solanum leucocarpon Dunal var. multiflorum Dunal in DC., Prodr. 13(1): 149. 1852. Type. Brazil. Bahia: S.loc., Blanchet 3537 (lectotype, G-DC, here designated; isolectotypes, F, K, frag. MPU).

Bassovia richardii Dunal var. martii Dunal in DC.,

Prodr.13(1):406. 1852. Type.Brazil.In Brasilia, Martius167 (holotype,P; isotype, BR). Solanum caavurana Veil. forma pauciflora Chodat,

Bull. Herb.Boissier1902:812. 1902.Type.Paraguay. Canendiyu:Prope Jejui guazi, Hassler 5722 (holotype, G-n.v.). Solanum anacamptorhachisBitter, Repert. Spec. Nov.

RegniVeg. 18: 53. 1922.Type.Brazil.Ceara:Bei Ule 9102 (holotype,B [destroyed]; Guaramisanga, lectotype,US, here designated;isolectotype,K). Solanum megalocarpon C. V. Morton, Rev. Argent.

Sp. Solanum106. 1976.Type.Argentina.Corrientes: Mburucuya, EstanciaSantaTeresa,6 Jan1949, Pedersen188 (holotype,US; isotypes,MO, NY). Solanumfoetidumof authors,not of Ruiz & Pav. Shrubsor occasionallysubshrubs1-5 m tall;young stems and leaves glabrous;stems slender,dryingdark; barkof olderstemsbecomingpalerfromstretchingand cracking. Sympodial units difoliate, geminate. Leaves

ellipticto ovate,widest at orjustproximalto themiddle, glabrousabove,pubescentbeneathwithtuftsofuniseriate trichomesca. 0.5 mm long in the axils of the main lateralveins, the leaves often dryingbluish or black;major leaves 8-18.6 x 3.6-7.7 cm, with 8-9 pairsof main

TAXONOMIC TREATMENT

85

Porto Seguro, 22 Mar 1978, Mori et al. 9832 (K, NY); Sta. Cruz de Cabralia,near Est. Ecologica Pau-Brasil (17 km W of Porto Seguro) rd. to Sta. Cruz de Cabralia, 4-6 km E of the station, 18 Oct 1978, Mori et al. 10812 (NY, US); Itamaraju,ca. 5 km W of Itamaraju,20 Sep 1978, Mori et al. 10747 (NY, US); Coroa Vermelha, 16 km N of Porto Seguro, 6 Apr 1979, Mori et al. 11680 (K, NY, RB); Itabuna, km 2 on rd. from Alcoba9a to Prado, 30 Apr 1978, Pinheiro 2158 (US); Ilheos, Sep 1832, Riedel 406 (NY, US); s.loc., Salzmann s.n. (P); in collibus, Salzmann 323 (P); in collibus, 1830, Salzmann 385 (G-DC); Alcobaca, km 6-8 on Rod. BA 001, section from Alcobaca to Caravelas, 16 Sep 1978, dos Santos et al. 3322 (F, US). CEARA: Guarmaranga, 1000 m, Bolland s.n. (K); Sitio Brejo, Aratuba, 17 Oct 1979, Castro & Nunes (E.A.C.) 7120 (NY); residence of Sr. Augusto Alves, Pacoti, 24 Sep 1981, Cavalcante & Bruno (E.A.C.) 10827 (NY). ESPIRITO SANTO: Aracruz, Rio Combios, between Barra do Riacho and Vila do Riacho, near Aracruz Celulose asphalt factory, 11 May 1987, de Lima et al. 2941 (K, NY); Marilandia-Rio Bananal rd., ca. 1 km N of Marilandia, 6 Dec 1994, Pirani et al. 3427 (K). MINAS GERAIS: Estrada de Conceitao, 7 Aug 1933, Mello Barreto 8474 (F); Corocovado, 18 Nov 1928, Brade s.n. (US). Paraiba: Joao Pessoa, Mares, 15 Nov 1990,Agra 1291 (K). PARANA: Porto Camargo to Perola, 22 Jan 1967, Hatschbacl; et al. 4374 (U); Umuarama, Serra Dourada, 19 Jan 1967, Hatschbach 15742 (US); from Porto Camargo to P6rola, 22 Jan 1967, Hatschbach et al. 15846 (F, NY, US, WIS); Xambre, Perola, 23 Jan 1967, Hatschbach & Haas 15846 (F, NY, US). PERNAMBUCO: Caruaru, Reserva Municipal Brejo dos Cavalos, 1100 m, 25 Mar 1994, Borges et al. 34 (BM, PEUFR); Caruaru,Reserva Municipal Brejo dos Cavalos, Fazenda Caruaru,1100 m, 1 Dec 1994, Sales et al. 450 (BM, PEUFR); s.loc., 1838, Gardner s.n. (K). RIO DE JANEIRO: Sao Joao da Barra, 26 Jan 1936, Mello Distribution (Fig. 37). In easternBrazil,Argentina, Barreto 7827 (F); Corcovado, 1857, Casaretto 1845 (G); and adjacentParaguayin pluvial forest, often forming s.loc., 27 Oct 1904, Dusen s.n. (US); s.loc., 1831-33, Gaudichaud 516 (G-DC, P); Gavea, 13 Nov 1868, secondarygrowth thickets, from sea level to 250 m. Glaziou 3075 (P); Morro da Babilonia, Nov 1914, Selected specimens examined. BRAZIL. S.loc., Hoehne 30091 (US); Meyer, Sep 1910, Hoehne 30093 herbier Richard (P); Blanchet s.n. (F); Burchell 898, (US); s.loc., Langsdorffs.n. (NY); Guanabara, Estrada 1078 (P); Martius 167 (BR, K, MO); 1839, Martius 622 de Jacarepagua,16 Apr 1948, Liane et al. 3624 (NY, US), (G-DC); Pohl s.n. (W); Sellow s.n. (F); St. Hilaire s.n. (P). Liane et al. 11019 (G); s.loc., Lund s.n. (NY); Recreio BAHIA: Margin of Rodovia Camacan, Canavieras, 32 do Bandeirantes, 15 Mar 1931, Lutz 519 (F); Restinga km W of Canavieres, 8 Sep 1965, Belem 1753 (NY, US); da Tijuca, 23 Apr 1945, Machado s.n. (F), 22 May 1944, Montiba, Blanchet 3537 (G-DC, P); Alcobaca, 2 km N Machado s.n. (F); Sebastianopolis, Cabo Frio, Ilheos, of city, 8 Dec 1981, Carvalho & Lewis 943 (K); between 1841, Martius 259 (G-DC, NY); Rio Cumprido, Miers 337 (MO); Forte Marechal Hermes, 15 May 1993, Mello Aguida and Porto Seguro, 28 Jun 1962, Duarte 6718 (RB, US); 6 Nov 1963, Duarte 8010 (RB, US); Itabuna, Silva & Pirani 861 (K); G.B. Praia do Grumani. 8 Mar RA. S.C. Cabralia, Res. Bio. Pau-Brasil, 5 Jan 1972, 1974, Occhioni 5848 (F); Guanabara,Jacarepagua,Serra Eupinino 146 (US); Itabuna,km 10-15 on BR 367, Porto do Pan da Fome, Sao Jose on Estrada da Boiuna, 14 Jan Seguro to Eunapolis, 18 Oct 1973, Eupinino 340 (US); 1962, Pabst 6801 (F, NY, US); Guanabara, Sumare, 23 s.loc., Gaudichaud 12 (P); Senhor do Bonfim, Serra da Sep 1958, Pereira et al. 4318 (US); Guanabara,restinga de Jacarepagua, 15 Oct 1958, Pereira et al. 4423 (F, NY, Santana, 650-900 m, 26 Dec 1984, Furlan et al. 36652 (NY); Fonte dos Protomartires do Brasil, Porto Seguro, US); Jacarepagua, Boiuna, 14 Dec 1958, Pereira & 1-10 m, 19?05'S, 16?26'W, 21 Mar 1974, Harley et al. Duarte 4856 (NY, US); Praia de Marica, ca. 0 m, 22?58'S, 17245 (K, MO, NY, U, US); Porto Seguro-Santa Cruz de 42?58'W, 18 Feb 1983, Plowman et al. 12853 (BH, F); Cabralia rd., km 5, 16 Jun 1980, Mattos Silva & Brito s.loc., 1829, Riedel 25 (NY); s.loc., Mar-May 1832, Riedel 841 (NY); Pau-Brasil ecological station, ca. 16 km W of 400 (NY, US); Corcovado, Richard 532 (P); Guanabara,

lateralveins, the apex acute to rounded,this character varyinggeographically,the base acute,winged ontothe petiole; petioles 0.7-1.2 cm long; minorleaves differing from the major ones only in size, 5-5.5 x 2.2-4 cm, the apex acuteor rounded,the base acuteto attenuate;petioles 5-6 mm long. Inflorescencesoppositethe leaves, simple, 1-4.5 m long, 5-20-flowered, glabrous or minutely glandular-papillose;pedicelscars evenly spacedca. 1mmapartintheupper1/2ofthe inflorescence. Buds ellipsoid to ovoid, the calyx split openjust before anthesis.Pedicels at anthesis 1-1.2 cm long, tapering fromthe calyx to a slenderbase ca. 0.5 mm diam.Flowers with the calyx tube 1.5-2 mm long, broadly cupshaped, the lobes broadly triangular,3.5-4 mm long, the marginsscarious,white in drymaterial,the tips minutelypapillose with trichomesless than0.2 mm long; corolla white, 1.6-2 cm diam.,lobed ca. 3/4 of the way to the base, the lobes planaror slightly reflexed at anthesis, the interpetalarsinuses not prominent,tips of the lobes minutelybrownish-papillose;anthersca. 4.5 x 1.5mm,poricidalatthetips,theporesteardropshaped; free portionof thefilaments ca. 0.3 mm long, the filamenttubeextremelyshort,less than0.2 mm long;ovary glabrous;style straight,ca. 1 cm long; stigma capitate, large,minutelypapillose.Fruita globose,greenorgreenish-white berry, 1-1.5 cm diam.;fruiting pedicels slightly deflexed, woody, 1.2-1.7 cm long, ca. 1 mm diam.atthe base;calyx lobes accrescentin fruit,woody, 5-6 mm long, scariousmargined.Seedsreddish-brown, flattened-reniform,ca. 2.5 x 2 mm, the marginspale tan,incrassate,thesurfacesminutelypitted.Chromosome number.n = 12 (voucherKnappet al. 9140)

86 Recreio dos Bandeirantes, Lagoinha das Taxas, 12 Jan 1965. Santos et al. 5326 (US); Corcovado, Nov 1883, Schwacke s.n. (US); s.loc., Oct 1888, Schwacke s.n. (US); Paneiras to Corcovado summit, 465-710 m, 22?56'S, 43?14'W, 17 Nov 1928, Smith 1245 (US); Cabo Frio, 1 Nov 1966, Sucre 1154 (F); Corcovado, 1833, Vauthier 532 (G-DC, P); Lagoinha Sumare,200-300 m, 5 Oct 1927, Zerny s.n. (W). SAO PAULO: Campinas, Mongolinho, Fazenda Santa Elisa, 11 Mar 1983, Brown s.n. (BH). PARAGUAY. S.loc., Andeer s.n. (P); Hassler s.n. (NY); Hassler 1370 (NY). ALTO PARANA:Puerto Gibajas, 4 km S of Puerto Iguaz6, 28 Mar 1970, Krapovickas et al. 15754 (US). AMAMBAY: Cerro Cora, 10 Jul 1978, Bernardi 19088 (G, NY, U); P.J. Caballero, near Tres Palos, 20 Jul 1978, Bernardi 19336 (U). CAAGUAZU: S of Salto del Guaira, Cordillera de Mbaracayu, 30 Oct 1978, Bernardi 18278 (MO); near and S of Yh6, not far from detour to San Joaquin, 13 Dec 1982, Ferndndez Casas et al. 7501 (NY); between Yerbales and Cordillera de Mbaracayu, 1900, Hassler 5179 (K, MPU, NY). CANENDIYU:Ygatimi, Colonia Mboi-jagua, 24?10'S, 54?40'W, 1 Apr 1983, Hahn et al. 1366 (MO, NY, PY mixed collection with S. symmetricum); Reserva Natural del Bosque Mbaracay6, roadside along Rio Jejui-mi E of Horqueta mi, ca. 150 m, 6 Mar 1996, Knapp et al. 9140 (BM, PY). CAPITAL:Aregua, near Lago Ipacaray, 16 Jan 1974, Schinini 7992 (WIS). CENTRAL:Jardin Botanico de Asunci6n, Sep 1971, Schinini 4006 (US). CORDILLERA: Cordillera de Altos, 6 Nov 1902, Fiebrig 376 (F, K), 3 Dec 1902, Fiebrig 558 (F, K); Emboscada, 25 1O'S,57 15'W,27 Aug 1988,Zardini6849 (PY). GUAIRA: Colonia Independencia, 250 m, 25?45'S, 56?13'W, 20 Dec 1986, Schinini & Bordas 24980 (NY, U). MISIONES: Estancia "La Soledad," Isla Carpincho, 26 Apr 1961, Pedersen 5991 (K, US). NEEMBUCUf: Plaine de Piyaru, between Villarica and Paraguari, 11 Feb 1876, Balansa 2122 (P). PARAGUARi: Isla Alta, Tebicuary Mi, 17 Nov 1978, Bernardi 18708 (NY). PRESIDENTEHAYES: Rio Pilcomayo, 11 Jan 1890, Morong 870 (NY). SANPEDRO: Alto Paraguay, Primavera, 20 Sep 1954, Woolston 326 (NY, U, WIS); Alto Rio, Primavera,13 Jan 1957, Woolston 326-B (US). ARGENTINA. CORRIENTES: Itati, Ruta Nac. 12, 47 km E of Itati, (Ea. Santa Catarina), 26 Feb 1977, Ahumada et al. 697 (F, MO); Ituzaing6, Ea. San Pedro, 27?45'S, 56?52'W, 12 Nov 1976, Arbo et al. 1198 (F, MO); General Paz, Poso Florentin, 26 Nov 1945, Ibarrola 3763 (P); Ituzaing6, Estab. Santa Eccla, 15 Mar 1946, Ibarrola 4313 (F); Ituzaing6, Curapaiti, 27 Mar 1946, Ibarrola 4365 (F); Mburucuyb, Estancia Santa Teresa, 10 Feb 1968, Krapovickas & Cristobal 13794 (LL, MO, WIS); Ituzaing6, Isla Apipe Grande, Puerto Mora, 11 Dec 1973, Krapovickaset al. 24276 (MO, WIS); Ituzaing6, ruta 12, 32 km W of Ituzaingb, 14 Apr 1974, Krapovickas et al. 25391 (MO); Estancia Santa Rita, near Rio Parana, 27?03'S, 56?04'W, 5 Mar 1987, Krapovickas et al. 41154 (NY); General Paz, 12 km E of Ita Itat6, shores of Rio Parana,9 Apr 1972, Mroginskiet al. 696 (MO, US, WIS); Carambola,Fortin del Ibera, 9 Nov 1982, Pedersen 13442 (NY); Ituzaing6,mouth of Arroyo Garap6into Rio Parana,

FLORA NEOTROPICA

45 km E of Ituzaing6,11 Dec 1974, Quarinet al. 2811 (MO);San Miguel, 12 km NE of San Miguel, Estancia CuruzfLaurel,14 Jun 1974, Schinini& Gonzalez8348 (MO); Ituzaing6, mouth of Arroyo Garap6into Rio Parana,45 km E of Ituzaing6,24 Apr 1975,Schininiet al. 11106(MO, WIS); San Cosme, Rio Paranaand Arroyo San Juan, 25 May 1975, Schinini & Quarin 11550

(MO); Concepci6n, Rio Santa Lucia, 20 Jan 1950, Schwarz9276 (NY,US); Concepi6n,11 km NW of Santa Rosa, 14 Dec 1977, Tressens863 (WIS); 12 km NE of San Miguel, EstanciaToto-y,27 Feb 1990, Vanniet al. 1445 (U). FORMOSA: Botanical Park of Ing. Lucas

Ruta11,RiachoPilaga,14 Nov 1986,Crist6bal Tortorelli, et al. 2115 (NY); Ruta11, RiachoPilaga,botanicalpark of Ing. Lucas Tortorelli, 19 Apr 1987, Cristobal & Krapovickas 2162 (NY); s.loc., 1918, Jorgensen 2218

(MO, US); Pilcomayo,ruta86, puenteGobernaci6n,16 Apr 1947, Morel 2564 (P). MISIONES: Posadas, Rio Alto

Parana,20 Nov 1907,Ekman818 (US);LeandroN. Alem, 10 km fromCerroAzul, on rd. to Ap6stolesvia Colonia Taranco, 10 Mar 1969, Krapovickas et al. 15038 (MO,

NY, WIS);San Pedro,2 de mayo, 510 m, 22 Apr 1957, Montes 27363 (MO, NY); General Belgrano, Obraje "Caburei,"450 m, 19 Jul 1957, Montes 27475 (NY); Estancia La Soledad, 3 km S of Santiago, 27?10'S, 56?46'W, 3 Feb 1988, Schinini & Vanni26017 (K); San

Ignacio, Santo Pip6, 3 Jul 1947, Schwarz 4691 (P); Candelaria, 22 May 1944, Sesmero 42 (NY).

Local names. Brazil.Amazonas:jurureba(Silva et al., 1977: but surely referringto another species, as Solanumcaavuranadoes not occurin Amazonas);Rio de Janeiro:caavurana.Argentina.Misiones:palo agui. Solanum caavurana is related to S. symmetricum of easternBolivia, Paraguay,andnorthwesternArgentina. Solanum caavurana differs from that species in its large,petaloid calyx lobes, more elongate inflorescences, elliptic to ovoid buds, and largerflowers. The petaloid calyx of S. caavurana is distinctive;the only otherspecieswith suchlargecalyx lobes is S. warmingii, whichis alsorelatedto S. caavurana.Solanumcaavurana has tufts of trichomes in the axils of the main lateral veins of the leaf undersides,andthe leaves usually dry blue-black, while S. warmingiiis less pubescent (but still possesses a few axillarytrichomes)anddriesa pale green or yellowish color. Collections ofS. caavurana fromnorthernBahiaandCearado not dryas blue-black as those fromotherpartsof the species range,andthose plantsalsohavemoreacute,less roundedleaf apicesthan plants from more southernpopulations.The northern populationsare somewhatisolated fromthe mainportion of the species rangeandareperhapsin the process of divergence. These plants have been called S. acuminatumvar. viridiflorumand S. anacamptorhachis. Solanumcaavuranais quite common in the forests of southeasternBrazil,andgrowsin a varietyof habitats. It occasionallyformsdense,foetid,monospecificthick-

TAXONOMIC TREATMENT

87

ets (K. S. Brown, pers. comm.). Vellozo (1829) stated acute;petioles 2-5 mm long. Inflorescences opposite that this species was ubiquitous, especially in second the leaves, often overtoppingthe leaves when nearthe growth.He also reportsseveraluses forS. caavurana. new growth, occasionally branching,long and flexuIt was used as a soap to whiten linen, andas the source ous, 1-10 cm long, 5-50-flowered, sparselypubescent of a blue dye. The vernacularnameandspecific epithet with uniseriatetrichomesca. 0.5 mm long, these more refer to this latteruse (Vellozo, 1829). dense near the distal end; pedicel scars irregularly spaced 0.5-5 mm apart,beginning ca. 1/3 of the way from the base of the inflorescence. Buds elliptic, the 13. Solanum campaniforme Roem. & Schult., Syst. corolla soon exserted fromthe calyx tube, glabrousor Veg. (ed. 16) [Roemer & Schultes] 4: 662. 1819. pubescentwith uniseriatetrichomeslike the rest of the Type.Brazil."inBrasilia,"Hoffmannseggs.n.(holo- inflorescence.Pedicels at anthesis filiform, deflexed, type, B-W [F neg. 2892]). Figs. 35D,F, 40, 41 1-1.3 cm long, taperingfromthe calyx tubeto a threadSolanumcaeruleumVeil.,Fl. Flumin.82. 1829[1825]. like base ca. 0.25 mm diam.Flowerswith the calyxtube Type. Brazil. Rio de Janeiro:"Habitatin silvis a flattened cup, 1-1.5 mm long, the lobes quadratemediterranis" (No specimens extant; lectotype, deltoid,apiculate,the marginshyaline,0.5-1 mm long, Vellozo, Fl. Flumin. Icones 2: fig. 110. 1831 glabrous or sparsely pubescent with uniseriate tri[1827], here designated). chomes;corolla white, 1.2-1.8 cm diam.,lobed nearly Solanum indigoferum A. St.-Hil. in Merat & De to the base, the lobes planaror campanulateat anthesis, Lens, Dict. Sci. Med. 6: 416. 1834. Type. Brazil. the lobes minutely papillose on the tips and margins; Rio de Janeiro: S.Loc., Sellow s.n. (holotype, P; anthers3-3.5 x ca. 1 mm, poricidalat the tips, the pores isotype, BM). teardrop shaped;free portionof thefilaments0.25-0.5 Solanum laxiflorum Sendtn. in Mart., Fl. Bras. 10: mm long, the filamenttube 0.5-1 mm long;ovarygla21. 1846. Type. Brazil. Rio de Janiero: In sylvis caeduis ad Sebastianopolim, Sept, Martius s.n. brous; style straight, 5-7 mm long; stigma capitate, (lectotype, M, here designated; isolectotype, K). minutely papillose. Fruit a globose, green (blue fide Solanum cearense Dunal in DC., Prodr. 13(1): 90. Vellozo) berry,1-2 cm diam.;fruitingpedicels some1852. Type. Brazil. Ceara: Serra de Ararife, Sep what woody, deflexed, 1.5-3 cm long, ca. 1 mm diam. 1838, Gardner 1792 (holotype, G [F neg. 8580]; at the base. Seeds russet brown,flattened-reniform,2isotypes, BM, K, frag. MPU). 4 x 1.5-3 mm, the marginsincrassate,pitted, the surSolanumfalcatum Witasek, Denskr. Akad. Wiss. Wien faces of the body of the seed smooth, the lateral cell 79: 341. 1931. Type. Brazil. Sao Paulo: Prope salto walls of the testa sinuatein outline.Chromosomenumdo Rio 500 v. Wettstein m, grande Paranapanema, ber not known. & Schiffner s.n. (holotype, W; photo in litt.). Solanum microrbitumL. B. Sm. & Downs, Phytologia 10: 426. 1964. Type. Brazil. Santa Catarina: Sao Francisco do Sul, forest, Tres Barras,Garuva, 200 m, 28 Feb 1958, Reitz & Klein 6506 (holotype, US; isotype, HBR-n.v.). Solanum dusenii L. B. Sm. & Downs, Phytologia 10: 426. 1964. Type. Brazil. Parana: Without further locality, 1 Feb 1904, Dusen 3361 (holotype, US; isotype, R-n.v.).

Distribution (Fig. 42). In easternBrazil from the mouthof the Rio Amazonasto the stateof Rio Grande do Sul;also in the GranSabanaregionof Venezuelaand adjacentGuyana;from aroundsea level to ca. 1200 m. Selected specimens examined. VENEZUELA. ANZOATEGUI:QuebradaBonita, tributaryof Rio Querecal

NE of Bergantin, 1200 m, 17 Feb 1945, Stevermark60974

(F, VEN); Ijigua, headwaters of Rio Le6n, NE of

Bergantin, 600-800 m, 27 Feb 1945, Steyermark 61249 Shrubs 1-4 m tall; young stems and leaves sparsely (VEN); Montaiia de las Palomas, tributaryof Rio Neveri, pubescent with uniseriate trichomes ca. 0.5 mm long, between Carmelita and Natalia, NE of Bergantin, 900occasionally glabrous; older stems glabrate, grayish. 1000 m, 9 Mar 1945, Steyermark 61438 (F, VEN). Sympodialunitsdifoliate, geminate.Leaves elliptic to BOLIVAR:Cerro Altamira, Ciudad Piar, ca. 500 m, Apr narrowly elliptic, widest at the middle, glabrous and 1954, Aristeguieta 2227 (NY, VEN); near Sta. Elena de shiny above, pubescentbeneathwith tuftsof uniseriate Uair6n, 800-900 m, 22 Apr 1957, Bernardi 6722 (G, trichomesin the axils of the main lateralveins, the tri- NY); SE of Sta. Elena, 900 m, 23 Apr 1957, Bernardi 6748 (G, NY); "El Paraiso" camp, 48 km NE of Caserio chomes ca. 0.5 mm long, often quite matted and dense; Los Rosos (17 km from Upata on new San Felix-Upata major leaves 5-14 x 2-4.5 cm, with 4-8 pairs of main rd.), 29 Apr-4 Jun 1965, Blanco 103 (VEN); Piar, Calceta lateral veins, these prominent and yellowish beneath, Larga, rd. to San Pedro de las Dos Bocas, NE of Manteco, the apex acute to acuminate, the base acute, slightly de200-290 m, Jul 1978, Delascio Ch. & Liesner 7199 (VEN); current onto the petiole; petioles 0.5-1.5 cm long; miGran Sabana, 11-12 km E of Misi6n Sta. Teresa de nor leaves differing from the major ones only in size, Kavanayen, ca. 1220 m, 5?55'N, 61?45'W, 8 Oct 1984, 1.5-5 x 0.9-3 cm, the apex acute to acuminate, the base Knapp & Mallet 6718, 6719, 6720, 6722 (BH, MY, IJS,

88

FLORA NEOTROPICA

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TAXONOMIC TREATMENT

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FIG. 41. Solanum campaniforme Roem. & Schult., plate 110 of Vellozo's Florae Fluminensis Icones, Vol. 2.

90

FLORA NEOTROPICA

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FIG. 42. Distribution of Solanum aphyodendron (solid circles) and S campaniforme (open circles).

VEN); along small stream between San Francisco de Yuruani and San Ignacio de Yuruani, ca. 5 km S of San Franciscoon rd. to Sta. Elena de Uairen,ca. 860 m, 5005'N, 61?00'W, 9 Oct 1984, Knapp & Mallet 6728, 6729, 6730, 6734 (BH, MY, US, VEN); ca. 30 km W of Sta. Elena de Uair6n on rd. to Icabaru, ca. 900 m, 4?45'N, 61?00'W, 10 Oct 1984, Knapp & Mallet 6735, 6741 (BH, MY, US, VEN); SE of Sta. Elena, 30 May 1946, Lasser 1986 (NY); 20-35 km S of Manteco on rd. to San Pedro de las Dos Bocas, 200 m, 7?10'N, 62055'W, 1-3 Aug 1978, Liesner & Gonzdlez 5902 (BM, VEN); Gran Sabana, San Ignacio de Yuruani,850 m, 5?00'N, 61?10'W,6 May 1988, Liesner 24262 (MO, U); N side of Rio Aponguao, 4200 ft, 2728 Mar 1952, Maguire 33660 (NY); E of Miamo, Altiplanicie de Nuria, 300-500 m, 8 Jan 1961, Steyermark 88179 (K, NY, US, VEN); between San Ignacio de Yuruani and San Francisco de Yuruani, between km 251.5 and 253 S of El Dorado on rd. to Sta. Elena, 1200 m, 4 Jan 1975, Steyermark 111394 (US, VEN); Gran Sabana, 7 km N of the mission Sta. Teresita de Kavanay6n, 1160 m, 20 Feb 1978, Steyermark et al. 115524 (US, VEN); Gran Sabana, Salto de Aponguao, 4205 km NE of the mission Sta. Teresita de Kavanay6n, 1130 m, 22 Feb 1978, Steyermark et al. 115612 (US, VEN). GUYANA. Kato, 650 m, 22 May 1977, Grewal et al. 427 (U). POTARO/SIPARUNI: Pakaraima Mtns., Cipo

Mtn. ca. 2 km from summit escarpment, headwaters of Cipo Creek, 1000 m, 4?54'N, 60?05'W, 26 Jan 1993, Henkel et al. 944 (BM). RUPUNUNI: Roraima, 1842-43, Schomburgk 680, 940 (G, K, P, U); Mt. Roraima, Arapo River, Oct-Jan 1884-1885, im Thurn 24 (K). FRENCH GUIANA. S.loc., Apr 1961, Aubreville 191 (P). BRAZIL. S.loc., Claussen s.n. (W); Lalande s.n. (P); Martius 1253 (B, G-DC, K, M, MO, NY, P); Pohl 3764 (F); Nov-Dec 1877, Regnell 972 (P); Sellow s.n. (B [destroyed: F neg. 2831]), fragment F, P [Morton neg. 8327], K, U); Maragnana,Martius s.n. (M [Morton neg. 8675]). AMAZONAS:Fazenda Dimona of WWF/INPA MCS project, ca. 72 km N of Manaus, 50-125 m, 2?19'S, 60?05'W, 23 Oct 1988, Boom & Pacheco 8518 (U). BAHIA: Porto Seguro, 0 m, 15 Apr 1965, Belem & Magalhaes 853 (NY, US); Santa Cruz Cabralia, 7 Nov 1966, Belem & Pinheiro 2848 (US); Porto Seguro, Estacao Ecol6gica Pau-brasil, CEPLAC, 19 Apr 1982, de Carvalho et al. 1188 (NY); Porto Seguro, BR5, km 5, 20 Aug 1961, Duarte 5976 (RB); Porto Seguro, km 4 of BR 5, 5 Jun 1962, Duarte 6692 (RB, US); Pau Brasil Biological Reserve 17 km W of Porto Seguro on rd. to Eunapolis, 0-20 m, 16?25'S, 39?12'W, 19 Mar 1974, Harley et al. 17198 (K, MO, NY, U, US); Est. Exp. Gregorio Bondar, km 58 of Rod. Belmonte-Itapabi rd.,

TAXONOMIC TREATMENT 16 May 1979, Mattos Silva et al. 360 (F); Santa Cruz de Cabralia, EMBRATEL tower, km 26 of BR 367 (Eunapolis-Porto Seguro), 220 m, 17 Oct 1978, Mori et al. 10766 (NY); Barra do ChoCa-Faz, 3-6 km E of Barra da Choca along Rio Catole, 22 Nov 1978, Mori et al. 11313 (NY); Santa Cruz de Cabralia, km 25.6 of BR 367 (Eunapolis-Porto Seguro), entrance to EMBRATEL tower, ca. 150 m, 4 Jul 1979, Mori et al. 12067 (NY); Sta. Cruz de Cabralia,Res. Biol. da Paubrasil,15 Sep 1971, Santos 1936 (US). CEARA:Chapada do Araripe, 3 Apr 1942, Beserra 437 (NY); Floresta Nac. do Araripe, 20 Nov 1979, Martins & Nunes (E.A.C.) 7525 (NY); Serra da Ibiapaba, Guaraciaba do Norte, 18 Dec 1979, Nunes & Martins (E.A.C.) 7854 (NY). ESPIRITOSANTO:Rod. BR-5, 5 km S of Morro Danta, 9 Aug 1965, Belem 1482 (US); cafezal of Sr. Loureiro, Santa Teresa, 750 m, 26 Jun 1968, Brown s.n. (US); Santa Teresa,Feb 1969, Brown 001, 200 (US); Rancho Alto, mun. Linhares, 7 Dec 1984, Hatschbach & Silva 48690 (NY). MARANHAO: Fazenda Bacaba, Doctor Haroldo, 5 km S of MA 119 from entrance 3 km NW of Lago do Junco, 4?26'S, 44?58'W, 4 Oct 1980, Daly et al. D448 (F, NY); island of Sao Luiz, Estradado Tirical, Feb-Mar 1939, Frdes 11538 (K, MAD, MO, NY, US); Turiba Rosa Santos, S. Joao dos Patos, 14 Dec 1979, Nunes & Martins (E.A.C.) 7791 (NY). MINAS GERAIS: Araponga, 800-1100 m, 21?S, 1 Mar 1924, Bailey & Bailey 1060 (BH); Belo Horizonte, Morro das Pedras, 24 Apr 1933, Mello Barreto 7833 (F); Pocos de Caldas, 9 Jan 1919, Hoehne 2698 (US); Caldas, Dec 1954, Lindberg 170a (BR); Carangola, Fazenda da Gramma, 5 km along trail to Areponga, 920 m, 4 Feb 1940, Mexia 4315 (F, K, MO, NY, US); Caldas, 16 Mar 1868, Regnell III 972 (BR, K, US); s.loc., St. Hilaire cat. cl. 318 bis (P); s.loc., 1845, Widgrens.n. (BR). PARA:W boundary of the I.A.N., Belem, 11 Nov 1942, Archer 7806 (K, US); S forest of the I.A.N., Belem, 16 Nov 1942, Archer 7819 (NY, US); km 18 S of Belem, 17 Nov 1942, Archer 7837 (F, K, NY, US); I.A.N, front, Belem, 2 Mar 1943, Archer 8255 (K, US); vic. of Para, 1908, Baker 77 (RSAIPOM, U); Sao Francisco do Para, across 96, 13 Dec 1978, Bastos et al. 138 (NY); Bel6m, I.A.N. capoeira 157L, 30 Nov 1956, Black 56-18930 (VEN); Santa Isabel do Para, grounds of Molposa factory, 17 Jul 1982, Costich 1082 (BH); Belem, Mar-May 1929, Dahlgren & Sella 387, 655, 716, 753, 772, 782 (F); shores of Lagoa Agua Preta, Belem, 9 Jul 1935, Drouet 2025 (F); Belem, 1 km NE of Santa Isabel, 10 Jul 1935, Drouet 2040 (NY, US); Para, 27 Oct-7 Nov 1929, Killip & Smith 30249 (F, NY, US); Ta pana near Para, 29 Oct 1929, Killip & Smith 30321 (NY, US), Killip & Smith 30346 (F, US); Ilha do Mosqueiro near Para, 3-9 Nov 1929, Killip & Smith 30413 (US), Killip & Smith 30498 (NY, US); upper Cupary River, plateau between the Xingu and Tapajos Rivers, Sep 1931, Krukoff 1179 (NY, P); Curuca, Iririteua, 15 Dec 1978, Nascimento 864 (NY); Rio Jari, Monte Dourado, 1 Jul 1968, Oliveira 4729 (NY, US); Belem, 13 Oct 1957, Pereira 3277 (RB, US); rd. from Belem to Brasilia, 1 Sep 1959, Pereira 5053 (US); BelemBrasilia, hwy., 1 Sep 1959, Pereira 5060 (US); vic. Belem, Sep-Oct 1961, Pires 51941 (NY, U., US); Maraba, N5,

91 aroundlake, 14 May 1982, Secco et al. 184 (NY); Belem, 1/2 km SE of I.A.N. adminstrationbuilding, 31 Dec 1943, A. Silva 5 (US); Belem, on I.A.N. grounds, 20 May 1944. A. Silva 209 (NY, US); Curuia, Abade, 14 Dec 1978, M.G. Silva 4060 (NY); Rio Jari, Monte Dourado, 1 Oct 1968, N.T.Silva 1075 (NY); MaguaryE.F.B.,3 Jul 1923, Snethlage 108 (F); Berlyn, Wullschagel 1528 (W); Utinza Reserve near Belem, 14 Sep 1927, Zernug s.n. (W). PARANA: Therezinain "capoeira,",22Jan 1911,Dusen 11191 (F, NY): Faguariahyra, 13 Jan 1915, Dusen 16336 (NY); Tres Barras, 27 Jan 1916, Dusen 17567 (K, NY); forest near Ivai, 18 km E of Jussara,200 m, 26 Mar 1966, Lindeman & de Haas 768 (NY, U); s.loc., 1879, Schwacke s.n. (US). PERNAMBUCO:Caruaru, Murici, Brejo dos Cavalos, Parque Ecologico Municipal, 1100 m, 25 Feb 1994, Borges s.n. (BM, PEUFR). RIO DE JANEIRO:Organ Mtns., Burchell 2213 (K); Serrados Orgaos, rd. to Campo das Antas, 1150-1250 m, 3 Jan 1971, Carauta 1269 (US); Itatiaya, Marromba, 1000 m, 2 Nov 1827, Ginzburger s.n. (F); s.loc., Glaziou 322 (P); Palmeiras, 13 Jan 1877, Glaziou 8878 (K, P); s.loc., Glaziou 11305 (P); Petr6polis, to Morin, 24 Mar 1879, Glaziou 11376 (K, P); headwaters Rio do Tigre, mun. Cerro Azul, 24 Jan 1980, Hatschbach 42740 (F); Petr6polis, 700 m, 29 Dec 1921, Holwa d& Holway 1434 (US); 17 km from Praca da Parati on rd. from Parati to Cunha, 26 Apr 1972, Kirkbride 1734 (F, MO, NY, US); vic. of Morretes, 2 Mar 1983, KuniYoshi 4646 (NY); Itatiaya, 1200 m, Dec 1892, Kuntze s.n. (NY, US); Sumidouro, Langsdorffs.n. (US); Vehla EstrellaPetr6polis, 1 Jan 1964, Pabst s.n. (US); Guanabara, Jacarepagua,estradada Boiana, 1 Oct 1958, Pereira et a'. 4371 (NY [mixed collection with S. pseudoquina], RB); Serrados Orgaos,25 Dec 1958, Pereira 4866 (US); Parque Estadual do Desengano, Serra da Agulha, fazenda Agulha do Imb6mbetweenSantaMariaMadalenaand SantoAnt6nio do Imbe, ca. 450 m, 21?56'S, 41?52'W, 23 Feb 1983, Plowman & de Lima 12940 (BH, F); Serra Estrella, 23 Feb, Riedel s.n. (NY); Petr6polis, rd. to Fazenda Inglesa, Mar 1951, Rocha e Silva 50 (US); Serra de Mantequeira, Alto do Serra, 1660 m, 22?23'S, 44?46'W, 24 Jan 1987, Salino & Gentry 61 (MO, NY); s.loc., Dec 1836, Sellowi s.n. (B [destroyed:F neg. 2831], frag. F); s.loc., 1838-42, Wilkess.n. (NY). SAOPAULO:Ca. 38 km SW of Jacupiranga along hwy 116 to Curitiba, 680 m, 8 Mar 1976, Davidse et al. 10955 (MO, NY); Alto da Serra, Edwall s.n. (US); s.loc., 2 Oct 1919, Gomes s.n. (US); Alto da Serra, Feb 1920, Hoehne 3670 (US); Alto da Serra, 16 May 1935, Kuhlmann 33208 (US); Alto Tiete, Rio Claro, 19 Oct 1901, Loefgren 5881 (US); Serra da Bocaina, Bocaina, pr. Casa do Peixe, 9 Feb 1959, Pabst 4719 (US); Alto da Serra, 12 Aug 1906, Usteri s.n. (US). SANTACATARINA: Braco Joaquim, Luiz Alves, Itajai, 300 m, 17 Jul 1954, Reitz & Klein 1953 (NY, UPCB); Ibirama, 100 m, 5 Feb 1956, Reitz & Klein 2641 (UPCB); Estrada Dora Francisca, Joinvile, 500 m, 21 Jun 1957, Reitz & Klein 4419 (US); s.loc., 30 Jun 1885, Schwacke s.n. (IJS). Local names. Venezuela. Anzoategui: aguata gallina. Brazil. Bahia: bobola; Para: cachorro, mucura caa-rana; Santa Catarina: joa manso, jua.

92

Solanum campaniformeis a widespread and variable species, andthe following relativelywell-marked geographicalraces exist. 1. Plants from the northernpartof the range in the Guianasandaroundthemouthof theAmazonare glabrouswithsomewhatlargerflowersthanthose from SE Brazil. The type of Solanumcampaniforme comes from among these populations. 2. Collections from Ceara are generally more pubescent than many, and may approach Solanum intermedium, but do not bear the distinctive golden trichomesof thatspecies. 3. Plants from Espirito Santo and Bahia retainthe trichomesof the inflorescence characteristicof more southerly specimens of Solanum campaniforme, but otherwise approachthe type in general morphology. These populations grow in coastal restingaor forest. 4. In the forests of Rio de Janeiro,Parana,and Sao Paulo,plantshavemediumsize flowers,elongate inflorescences, and berries about 1 cm diam. Plants from these populationshave been called Solanum caeruleum. 5. In the drierforests of Minas Gerais,populations consist of plants with larger flowers which often appearblue when dry and longer, more pubescent inflorescences. These have been called Solanumlaxiflorum. 6. Thesouthernmost populationsfromSantaCatarina and Parana have smaller leaves and shorter inflorescences thanplants from morenortherly populations.Intermediatesexist however, and the differences are minute. These plants have beencalledSolanumduseniiandS. microrbitum.

FLORA NEOTROPICA

14. Solanum cassioides L. B. Sm. & Downs, Phytologia10: 430. 1964.Type.Brazil.SantaCatarina: Lauro Muller, Urussanga, Pinhal de Companhia, 300 m, 20 Sep 1958, Reitz & Klein 7202 (holotype, US; isotype, HBR-n.v.). Fig. 43

Shrubletsor shrubs, 0.5-2 m tall; young stems and leavessparselypubescentwithsmalldendritictrichomes, soon glabrate;barkof olderstems darkreddish-brown. Sympodialunitsdifoliate,usuallynot geminate.Leaves elliptic to narrowlyelliptic, widest at the middle, the minor leaves if present not differing in size or shape fromthe majors,3-12 x 1-5 cm, with 5-6 pairsof main lateralveins, the adaxialsurfaceglabrousor with scattered uniseriate simple or dendritic trichomes with swollen reddishbases on the lamina, the abaxial surfaces pubescentwith dendritictrichomes,these denser along the veins, the margins appearingtoothed with smalluniseriateswollen-basedtrichomes,theapexacute, the base attenuate;petioles 0.5-1.2 cm long. Inflorescences intemodal,oftenborneon shortside shoots,0.51.5 cm long, simple,5-8-flowered,pubescentwith dendritic trichomes like those of the stems;pedicel scars closely spaced,not overlapping.Buds ellipsoid to obovate, the corolla long-exserted from the calyx tube. Pedicels at anthesisfiliform, 1.4-2.1 cm long, erector somewhatdeflexed,contractedbelow the calyx tube,ca. 0.5 mm diam. at the base andapex, sparselypubescent with dendritictrichomes.Flowers with the calyx tube broadlyconical, 1-1.5 mm long, the lobes deltoid, 1.52 mm long, with an elongateapicalprojection,sparsely pubescentwith dendritictrichomes,with a tuftof simple uniseriatetrichomes on the projection;corolla white, 2-2.3 cm diam., lobed 2/3 of the way to the base, the lobes campanulateat anthesiswith broadthinmargins, the abaxial surfaces of the lobes pubescent with denThese geographicalracesarenot easily delimitable, dritic trichomes;anthers4.5-5 x 1-1.5 mm, poricidal andthe entirerangeof intermediatesis found.Naming at the tips, the pores teardropshaped;free portion of these would be counterproductiveandserve to confuse thefilamentsminute,0-0.5 mm long, the filamenttube an already complex situationthat merits furtherfield 0.5-1 mm long, glabrous;ovaryglabrousor pubescent study. (only in type);style 0.8-1 cm long, glabrousor pubesSolanumcampaniformehas been collected from a cent(in type only);stigmacapitate,the surfaceminutely varietyof habitatsincludingcoastalforestandcapoeira. papillose.Fruita globose, often apicallypointed,green It appearsto be an opportunisticspecies and grows in (dryingmustardyellow) berry,1-1.5 cm diam.,theperiopen areaswhereverthey are found. Some of the mor- carp very hard and woody; fruiting pedicels woody, phological variationobservedin S. campaniformemay deflexed, 2.5-3.2 x ca. 1 mm diam. at the base. Seeds be the result of ecotypic differences. darkbrownorpaletan,flattened-reniform withincrassate The lectotype I have chosen for the nameSolanum x 2-2.5 2-3 the surfaces mm, margins, minutely pitlaxiflorum, a synonym of S. campaniforme,is most ted. Chromosomenumbernot known. clearlydocumentedandthe firstof the collections cited Distribution (Fig. 44). SE Brazil in Araucauria by Sendtner.The othercollections cited in the original at 300-1200 m. forest, have no and of localities, description thoughspecimens thesearedepositedat M, I preferto use as a lectotypethe Tamandare, Specimens examined. BRAZIL. PARANA: Martiuscollection.Additionalisotypeswill be moreeas- 4 Oct 1908, Dusen 6819 (K); Piraquara, 25 Sep 1909, ily determinedusing this better-documentedspecimen. Dusen 8754 (K); Itaperussiu,880 m, 27 Sep 1914, Jonsson

TAXONOMIC TREATMENT

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94

FLORA NEOTROPICA

..... ~~~~~~~~~~~~~~~~~~~~~~~~~~-... ....... . i.

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FIG. 44. Distribution of Solanum cassioides (solid circles), S. daphnophyllum (open circles), and S. gertii (solid squares).

1005a (K). SANTA CATARINA: Alto da Serra do Espigao, Campos do Areao, 1200 m, 21 Oct 1961, Pabst 6112 (Pereira 6285) (F, US); Serra da Boa Vista, Sao Jose, ca. 1100 m, 4 Feb 1953, Reitz 5469 (US); Serra da Boa Vista, Sao Jose, 1200 m, 4 Feb 1953, Reitz 5493 (US); Barra do Sul, Araquari, 8 Feb 1983, Reitz & Klein 537 (US); Serra da Boa Vista, Sao Jose, 800 m, 14 Oct 1960, Reitz & Klein 10204 (K, US); Campo Alto, Santa Cecilia, 1200 m, 25 Oct 1962, Reitz & Klein 13492 (US); Matos Costa, 1100 m, 27 Oct 1962, Reitz & Klein 13739 (K, US).

Cruz:SanJuan,3200 ft, 23 Mar 1902, Williams221 (holotype, BM; isotype, K). Fig. 45

Shrubsor small trees, 1-4 m tall; young stems and leaves glabrousor minutelypapillose;barkof the older stems deep reddish-purpleand shining, breakingand peelingin transversesectionsoroccasionally(Peru)pale greenish-yellow.Sympodialunitsdifoliate,rarelygeminate,if so membersof thepairof equalsize.Leavesthick andsubcoriaceous-fleshy,obovateto narrowlyobovate, Solanum cassioides is probablyclosely related to widest at orjust distal to the middle, glabrouson both S. trachytrichium,also from southeasternBrazil and surfaces, drying pale green, 7-14 x 2.5-6.5 cm, with adjacent Argentina. It shares with that species the 4-7 pairsof mainlateralveins, these prominenton both hooked trichomeswith mound-likemulticellularbases surfaces,but palerbeneath,the apex acuteto rounded, on the leaf surfaces(see Fig. 9C). InS. cassioides these the base cuneate; petioles 0.4-1.1 cm long, sulcate trichomesare usually confined to the leaf margins,but above. Inflorescences opposite the leaves or pseudoin some collections are found on the adaxial leaf sur- terminal,glabrous, simple, 1-3 cm long, 5-15-flowfaces.Solanumtrachytrichium hasthesetrichomesover ered;pedicelscarsclosely spaced,overlapping,orwidely the entireleaf surface.The two species also sharelarge, spaced ca. 2 mm apart,beginning ca. 2/3 of the way campanulateflowers anddifoliate,non-geminatesym- fromthe base of the inflorescence.Buds globose when young, laterellipsoid and long-exsertedfromthe calyx podial units. Bitter (1919) created subsect. Silicosolanum for tube.Pedicels at anthesis0.9-1 cm long, taperingfrom Solanumtrachytrichiumbased on the roughfeel of the the calyx tubeto a slenderbase ca. 0.75 mm diam.;buds leaves, but the two taxa here are not distinct enough some way or another.Flowers with the calyx tube 1.5from the rest of the S. nudumspecies groupto warrant 2 mm long,the lobes broadlytriangular,0.5-1 mm long, such separation. minutelypapillosewithout,the tips occasionallywith a tuft ofuniseriate trichomesca. 0.25 mm long; corolla white, 1.5-2 cm diam.,lobed3/4 of the way to the base, 15. Solanum daphnophyllum Bitter, Repert. Spec. the lobesplanaratanthesis,thetipsandmarginsminutely Nov. Regni Veg. 18:67. 1922. Type. Bolivia. Santa papillose;anthers5-5.5 x 1-1.5 mm,poricidalatthetips,

TAXONOMIC TREATMENT

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the pores teardropshaped;free portionofthefilaments ca. 0.5 mm long, the filament tube 0.5-1 mm long; ovary glabrous;style straight,7.5-8 mm long; stigma clavate or capitate, minutely white papillose. Fruit a globose,hard,greenberry,1-1.3 cm diam.,in herbarium specimens mustard yellow; fruiting pedicels erect, woody, ca. 1.5 cm long, 1-1.5 mm diam. at the base. Seeds brown,flattened-reniform,3-4 x 3-3.5 mm, the marginsincrassateandyellowish, the surfacesminutely pitted. Chromosomenumbernot known.

.B. more typical specimen with rounded leaf tips,

along oil pipelineca. 5.6 km fromPampaGrande,1600 m, 10 Aug 1991,AcevedoRodriguezet al. 4482 (K, NY); 2000 m, Jun 1944,Cardenas3481 (US);Prov. Samaipata, AndresIbafiez,alongrd. fromSantaCruzto Samaipata, 3 km SW of Angostura,gorge of Rio Pirai, 700 m, ca. 18?10'S,63?32'W,25 Jan 1987,Nee 33828 (NY); Prov. Florida,5 km by air SE of Mairanaon rd. to Samaipata at QuebradaSeca, 1550 m, 18?09'S, 63?56'W,4 Feb 1988, Nee 36155 (IBE, NY); Achira, Samaipata,1500 m, 11 Oct 1928,Steinbach8210 (F, K, MO,NY); Prov. Florida,HierbaBuena, 1300 m, 10 Jun 1966,Steinbach 278 (DAV,MO, NY, US, WIS).

Distribution (Fig. 44). In middle-elevation dry deciduous forests to higher-elevation forests near Solanumdaphnophyllumis an anomalousspecies in steppelandson the eastern Andean slope in southern the S. nudumspecies group.It is probablymost closely Peru and Bolivia, at (700-)1550-2800(-3500) m. relatedto S. restingaeof coastalBahia,Brazil.Solanum examined. Cuzco: PERU. Urubamba, daphnophyllumis unusualin sect. Geminatain growSpecimens MacchuPicchu, above town of Pauqarcancha, 3380 m, ing in dry,deciduoussemi-tropicalforest.It is common 3 May 1982, Peyton & Peyton 147 (MO); Urubamba, where it occurs(M. Nee, pers. comm.). PeruvianmateMacchuPicchu,alongIncatrailin TresPiedrasBlancas, rial from around Macchu Picchu has more elongate 1 km from Huayabamba, 3130 m, 16 Aug 1982, Peyton infloresenceswithmorewidely spacedpedicelscarsthan & Peyton 1026, 1026b (MO). PUNO: Sandia, near San materialfromBolivia. These specimens(Peyton Juan de Oro, Valle de Tambopata,2100-2200 m, 14 May typical & 147, 1026, 1026b)also lackthe redshinybark Peyton 16683 1966, Ferreyra (MO). BOLIVIA. Bank of river at Paquicha,Jul 1865, Pearce of the type; their barkis pale yellow greenish and not s.n. (K). COCHABAMBA:Siberia, 2800 m, Sep 1954, markedlyexfoliating.The Peruvianmaterialmay evenCdrdenas 5601 (US). SANTACRUZ:Pampa Grandeto rd. tuallyneed to be considereda separatetaxon.

96

FLORA NEOTROPICA

The inflorescencein Solanumdaphnophyllumis not strictly leaf-opposed as is the case in most species of sect. Geminata.The inflorescences appearto be short shoots thatare borne terminally(as all inflorescences in Solanum,see Morphology),but arelaterovertopped by the continued growth of axillary shoots; thus they are lateral in flower and fruit, but not always (see S. sessile group for a similar situation).

Distribution (Fig. 44). Atlanticandinteriorforests in the stateof Parana,Brazil.Apparentlya shrubof both the forest interiorand of margins, 800-900 m.

Shrubsor small trees 0.5-2 m tall;young stems and leaves pubescentwith simple uniseriate3-5-celled trichomes 1-1.5 mm long, some with glandulartips;bark of older stems pale greenish-yellow.Sympodialunits difoliate,geminate.Leaveselliptic,widest at orjust below the middle,dryingolive-green,glabrousadaxially, pubescent abaxially with dense tufts of uniseriatetrichomes in the vein axils, these spreadingto the veins andlamina,occasionallywith some trichomesscattered over the lamina;majorleaves 9.5-15 x 3-6 cm, with 67 pairsof mainlateralveins dryingyellowish abaxially, the apex acuminate,the base attenuate;petioles 0.6-1 cm long; minorleaves differingfromthe majorsin size only, 1.5-4.5 x 1-2.5 cm, the apexacute,the base acute; petioles 2-5 mm long. Inflorescences opposite the leaves, simple,0.3-1.5 cm long, 5-7-flowered, the axis sparselypubescentwith simpleuniseriatetrichomeslike those of the young stems;pedicel scars evenly spaced ca. 0.5 mm apart.Budsellipsoid,stronglyexsertedfrom the calyx tube.Pedicels at anthesis filiform, glabrous, 2-2.2 cm long,erectto slightlydeflexed,ca. 1.5mmdiam. at the apex, 0.5 mm diam. at the base.Flowers with the calyx tube conical, 1.5-2 mm long, the lobes quadrate with distinctapicalprojections,0.5-1 mm long, pubescent with scattereduniseriatetrichomesca. 1 mm long anda dense tuftof uniseriatetrichomesca. 0.4 mm long on theapicalprojection;corollawhite, 1.5-1.8 cm diam., lobed nearly to the base, the lobes planar to slightly reflexedat anthesis,the tips andmarginspapillose;anthers3.5-4 x 1.5-2 mm, poricidalat the tips, the pores teardropshaped; free portion of thefilaments ca. 0.1 mm long, the filament tube 0.5-1 mm long, glabrous; ovaryglabrous;style straight,1-1.2 cm long, glabrous; stigma slightly clavateto bilobed,the surfaceminutely papillose.Fruita globose, greenberry,ca. 1.5 cm diam.; firuitingpedicelswoody, deflexed, ca. 2.5 cm long, ca. 3.5 mm diam. at the apex, 1.5 mm diam. at the base. Seeds dark reddish-brown, flattened-reniform with incrassatemargins,ca. 3 x 2 mm, the surfacesminutely pitted. Chromosomenumbernot known.

Solanumgertii is closely relatedto S.pseudoquina, also of southeasternBrazil.It differsfromthatspeciesin its equal anthersand long trichomeson the stems and new growth.It is found within the rangeofS. pseudoquinaandmay be a geographicalisolateof thatspecies.

Specimens examined. BRAZIL. PARANA: Sertan6polis, Fazenda Ferraz, 15 Jun 1994, Colombo et al. 20293 (BM, FUEL); Mun. Morretes, Serra Marumbi, 800-900 m, 25 Feb 1970, Hatschbach 23909 (MBM, NY); Mun. Bocaiuva do Sul, W of Sesmaria, Rio Capivari, 24 Mar 1970, Hatschbach 24053 (BM); near sawmill Brandalize, N of Rio Chopim, ca. 20 km N of 16. Solanum gertii S. Knapp,Brittonia44: 67. 1992. Clevelandia, ca. 700 m, 27 Apr 1966, Lindeman & de Type.Brazil.Parana:Mun.Morretes,Est.Marumbi, Haas 1048 (U); Londrina, Parque Arthur Thomas. 6 Oct 11 Feb 1981, Oliveira 285 (holotype, MBM; 1993, Nishimura et al. 20256 (BM, FUEL); Estaiao isotype, NY). Fig. 46 Marumbi, 8 Feb 1990, Ribas & Cordeiro 252 (BM).

17. Solanum intermedium Sendtn.in Mart.,Fl. Bras. 10:22. 1846. Type.Brazil. MinasGerais:Caxoeira do Campo,Martiuss.n. (lectotype, M, here designated [F neg. 6534]; isolectotype,BR). Fig. 47 Shrubs 2-3 m tall; young stems pilose with uniseriate (often bi-cellular) golden curl-tipped trichomes 0.5-0.75 mm long, these deciduous on older branches;bark of older stems shiny reddish-brown, slightly exfoliating.Sympodialunitsdifoliate, usually geminate.Leaveslanceolate,widest at orjust proximal to the middle,markedlydecreasingin size nearthe tips of stems,glabrousabove,pilose beneathwithuniseriate trichomeslike those of the stems,the trichomesdensest along the veins; majorleaves 5-11 x 1.4-2.8 cm, the apex acuminate,the base acute;petioles 0.4-1 cm long; minorleaves differingfromthemajorsonly in size, 2.24.8 x 1-1.8 cm, the apex acuminate,the base acute;petioles 1-3 mm long. Inflorescencesopposite the leaves or overtoppingthe leaves nearthe shoot tip, occasionally furcate,1-5 cm long,sparselypubescentwithgolden uniseriatetrichomeslike those of the stems andleaves; pedicel scars irregularlyspaced0.5-1.5 mm apart.Buds globose to elliptic, the corolla soon exserted from the calyx tube. Pedicels at anthesis 0.9-1 cm long, tapering fromthe calyx tubeto a slenderbase 0.5 mm diam., deflexed at anthesis.Flowers with the calyx tube ca. 1 mm long, denselyto sparselypubescentwith uniseriate trichomeslike those of the stems and leaves, the lobes deltoid, 1-1.5 mm long, pubescent;corolla white, 1.11.2 cm across, lobed nearlyto the base, the interpetalar sinusesthinandmembraneous, tipsof thelobes minutely at anthers ca. 2.5 mm long, anthesis; papillose, planar

TAXONOMIC TREATMENT

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98

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theterminal0.5 mmpalerandthickened,ca. 1 mmwide, poricidal at the tips, the pores teardropshaped; free portion of thefilaments 0.5-1 mm long, the filament tube ca. 0.5 mm long; ovary glabrous;style 5-6 mm long, straight,clavate; stigma a papillose area on the tip of the style. Fruit and seeds not known. Chromosome numbernot known. Distribution (see Fig. 49). In mountainous and cerradoareasof southernBrazil. Specimensexamined.BRAZIL.S.loc.,(BR,BH3251/ 83-5,6). MINAS GERAIS: S.loc., Claussen s.n. (G-DC, P);

Caxoeirado Campo,1839, Claussen7 (F); s.loc., 1841, Claussen1440 W-7 (P); Caxoeirado Campo,Mar 1839, Martius 58 (BR); Serra do Cip6, Santana do Riacho, near Conceicao do Mato Dentro, Rio San Antonio, 19 Apr

1981, Rossi & Amaral7277 (NY); Lagoa Santa,5 Apr 1864, Warming s.n. (NY). RIO DE JANEIRO: Novo

Friburgo, Nov 1842, Claussen 80 (P (Morton neg. 8216)); s.loc., Lands.n. (NY). Solanum intermediumis relatedto S. reitzii and S. Brazil.Itdifcampaniforme,bothspeciesof southeastern fersfromS. campaniformein its indumentof soft,golden trichomes,andfromS. reitziiin its anthersof equalsize. Solanum intermediumhas only been collected to my knowledge once since the middle 1800s, and may be on thevergeof extinction.Furthercollectingis necessary to determineif this is the case. It may be thatS. intermediumis a cerradoendemic,which might accountfor its patchydistributionandpotentiallyvulnerablestatus. I have lectotypifiedSolanumintermediumwith the firstof the two Martiuscollections cited by Sendtnerin the original description.The specimen is annotatedin Sendtner'shandwritingat M.

99

3-4 cm, with 6-8 pairsof mainlateralveins, impressed above,reddishandprominentbeneath,pubescentin the vein axils with uniseriatetrichomes0.5-1.5 mm long, these extending sparselyto the lamina,the apex acute to sharplyacuminate,the base acute;petioles 1.5-2 cm long; minorleaves differing from the majorones only in size, 2.5-5 x 1.2-3 cm, the apexacute,thebase acute; petioles 5-8 mm long. Inflorescences opposite the leaves, usuallysimple,butin the type specimenfurcate, 1.5-6 cm long, 20-25-flowered, glabrousor with a few uniseriatetrichomes like those of the leaves along the peduncle and at the base of the pedicels, occasionally along the pedicels;pedicel scars corky and expanded, in pairsca. 1 mm apart,the membersof the pairca. 0.5 mm apart,beginning ca. 1/3 of the way from the base of the inflorescence.Buds fleshy (woody in dry material), ovoid, the corolla soon exserted from the thick woody calyx lobes.Pedicels at anthesisthickandfleshy, deflexed, 1.2-1.8 cm long, taperingfromthe calyx tube to a basaldiam.of ca. 0.75 mm.Flowers with the calyx tube ca. 2 mm long, conical, the lobes broadlydeltoid, ca. 1.5 mm long, the marginspale andscarious,the tips minutelypapillose, occasionally with a few uniseriate trichomesca. 0.5 mm long; corolla white, fleshy, 1.51.8 cm diam., planaror slightly reflexed at anthesis, lobed3/4 of theway to thebase,thelobesslightlyreflexed atanthesis,thetipsof thelobesstronglycucullate,the tips and margins densely papillose; anthers 4-5 x 1.5-2 mm, poricidal at the tips, the pores teardropshaped; free portionof thefilaments ca. 0.5 mm long, the filament tube ca. 1 mm long; ovary glabrous;styles 5-6 mm long, glabrous;stigmaminutelycapitate,notclearly differentiatedfromthe style tip. Fruit a globose, green berry, 1-1.2 cm diam.;fruiting pedicels woody, deflexed,ca. 2.5 cm long;calyx lobeswoody in fruit.Seeds yellowish, flattened-reniform,ca. 3 x 2.5 mm, the surfaces coarselypitted,thepits ca. 0.1 mm diam.,the seed marginsincrassate.Chromosomenumbernot known.

18. Solanum lasiopodium Dunal in DC., Prodr. 13(1): 679. 1852. Type. Colombia. Guajira:Sierra Nevadade SantaMarta,Rio Hacha,Taquina,10000 Distribution (Fig. 49). Found only in the Sierra ft, Feb 1842, Linden 1624 (holotype, G-DC [F Nevada de SantaMartain northernColombia, 2000neg. 6785]; isotypes, BM, F, MPU, P [Morton 3000 m. neg. 8235]). Fig. 48 bassovioidesRusby,Descr. S. Amer. Cyphomandra P1. 116. 1920. Type. Colombia.Guajira:Sierra del Libano, 6000 ft, 20 Jan 1902, H. H. Smith 1181 (holotype,NY-n.v.; isotypes, A-n.v., BM, G-n.v., GH-n.v.,K, L-n.v., MPU,P-n.v., S-n.v., fide Bohs, 1994]). U-n.v., US [isotypedistribution Shrubsor small trees, to 6 m tall; young stems and leaves minutely glandularpubescent, soon glabrate; bark of older stems shining, reddish, densely white lenticellate.Sympodialunitsdifoliate,geminate.Leaves ovate to elliptic, widest at or just below the middle, dryingblack above, palerbelow; majorleaves 7-12 x

Specimens

examined.

COLOMBIA.

MAGDALENA:

SierraNevadade SantaMarta,SE slopes Rio Donachui, Cancur6a, 2400 m, 12 Oct 1959, Cuatrecasas & Romero

Castaheda24801 (US); Jabali, HaciendaLa Victoria, San LorenzoMtns., 1690 m, 13 Jun 1944,Kernan11'6 (US); Rio Hacha,SierraNevada, 10,000 ft, Mar 1852, Linden 815 (F, G ( [F neg. 6785], G [F neg. 23125]. P

[Mortonneg. 8234]). Solanum lasiopodiumis most closely relatedto S. nudum, a widespread species of northern South America and the Caribbean.It is very like the highelevation race of S. nudum(S. tovarense), but differs from that set of specimens in its much larger,fleshier

FLORA NEOTROPICA

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TAXONOMIC TREATMENT

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flowers,pubescentnew growthandcalyx lobes, and its ovoid budswiththe corollasoon exsertedfromthe calyx lobes.Solanumlasiopodiummay be a geographicalisolateof its morewidely distributedrelativeS.nudum.The type specimen of Cyphomandra bassovioides (Smith

1181) is almostcompletelyglabrouson the calyces and old growth,butsharestheothercharactersof thisspecies. 19. Solanum nudum Dunal, Solan. Syn. 20. 1816. Type. Mexico. Veracruz:Prope Jalapa,Humboldt & Bonpland s.n. (holotype, P-Bonpl. [microfiche IDC 6209-2:61 1.5]). Fig. 50 Solanum anonaefolium Dunal, Solan. Syn. 15. 1816.

Type. Colombia.Magdalena:Ad fluvium Magdalenae, Humboldt & Bonpland s.n. (holotype, P-

Bonpl.;isotype,B-W-n.v.[F neg. 2888], frag.F). Solanummicranthum Willd.ex Roem.& Schult.,Syst. Veg. (ed. 16) [Roemer& Schultes]4: 663. 1819. Type. Venezuela.Sucre: Cumana,Humboldt& Bonplands.n. (holotype,B-W [F neg. 2903]). Solanum antillarum 0. E. Schulzin Urb.,Symb.Antill. 6: 164. 1909. Type.Cuba.Montede la Clarita,in sylvis, 800 m, Apr 1889, Eggers5039 (lectotype, B [destroyed], US, heredesignated; P). isolectotypes,

Solanum nudum var. micranthum (Willd.) Hassl., Repert. Spec. Nov. Regni Veg 15: 114. 1918. Type. Based on the type of Solanum micranthum, but said by Hassler to be found in Peru and Brazil (No specimens cited). Solanum parcebarbatum Bitter, Repert. Spec. Nov. Regni Veg. 18: 51. 1922. Type. Panama. Panama: Forest on dry limestone around Alajuela, Chagres Valley, 30-100 m, 12-15 May 1911, Pittier 2325 (lectotype, US, here designated [US 676572]; isolectotypes, F, US). Solanum parcebarbatum var. minorifrons Bitter, Repert. Spec. Nov. Regni Veg. 18: 52. 1922. Type. Guatemala. Alta Verapaz: Near Finca Sepacuite, 25 Mar 1902, Cook & Griggs 146 (holotype, US [US 407934]). SolanumsupranitidumBitter, Repert. Spec. Nov. Regni Veg. 18: 69. 1922. Type. Brazil. Amazonas (now Acre): Jurua-Miry,Rio Jurua superior, Aug 1901, Ule 5690 (lectotype, HBG, here designated; isolectotype B [destroyed: F neg. 3785], K-n.v.). Solanum tovarense Bitter, Repert. Spec. Nov. Regni Veg. 18: 65. 1922. Type. Venezuela. Distrito Federal: Prope Colonia Tovar, 2000 m, 1854-5, Fendler 980 (lectotype, G, here designated [Barb.-Boiss., Morton neg. 8562]; isolectotypes, BR, F, G [Deless. F neg. 23163], MO, NY).

102

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4~~~4,. ~n . 50.'Solanum nudum FIG: Dunal. A.'holotype"at P-Bonpl. B.Belize. Whitefoord 9065(BM).

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FIG. 50. Solanum nudum Dunal. A. holotype at P-Bonpl. B. Belize. Whitefoord 9065 (BM).

Shrubs2-5 m tall;young stems andleaves glabrous or minutely puberulentwith appressedunicellular or uniseriatetrichomesca. 0.1 mm long, oftendryingblack; older stems glabrous or with tufts of white uniseriate trichomesnearthenodes;barkof olderstemsdarkbrown or gray,shiny,slightlywingedfromdecurrentleafbases. Sympodialunitsdifoliate,geminate.Leavesovate to elliptic,widest at orjust proximalto the middle,glabrous andshinyabove,withtuftsof whiteuniseriatetrichomes 0.5-1 mm long in the axils of the main lateral veins abaxially,the densityoftrichomes varyinggreatly;major leaves 6.4-17 x 3-9 cm, with 5-10 pairs of main lateralveins, these impressedadaxiallyin the Venezuelan high-elevation race, prominent and yellowish abaxiallyin all specimens,theapexacute,thebase acute, to rounded,somewhatdecurrentonto the petiole; petioles 0.5-2.5 cm long,slightlywingedfromtheleafbases; minor leaves differingfrom the majorones in size and often in shape, elliptic to somewhatorbicular,1.5-6 x 0.9-4.5 cm, the apex acuteto rounded,the base acuteto rounded;petioles 2-6 mm long.Inflorescencesopposite the leaves, simple,glabrousorminutelypuberulentwith thesameappressedtrichomesas thoseof theyoungstems andleaves,0.5-5 cm long, 5-50-flowered;pedicelscars

closely spaced, but not overlapping,ca. 0.5 mm apart. Budsglobose, the calyx lobes appearingas five rounded knobs in young buds, the corolla soon exserted,bright orpale green,the calyx white.Pedicels at anthesiswhite or greenish-whitein live plants, deflexed, 0.7-1.3 cm long,taperingfromthecalyx tubeto a slenderbase0.250.5 mm diam.Flowers with the calyx tube conical, 11.5 mm long, the lobes deltoid, 0.25-1 mm long, glabrous or minutely puberulent; corolla white or greenish-white,0.5-1.2 cm diam.,lobed 2/3 of the way to the base, the lobes slightly reflexed at anthesis, the tips andmarginsof the lobes minutelypapillose;anthers 1.2-2.5 x 1-1.5 mm, poricidalat the tips, theporesteardropshaped;freeportionofthefilaments 0. 1-0.25 mm long,thefilamenttube0.25-0.5 mmlong;ovaryglabrous; style straightor slightly curvedto one side, especially in driedmaterial;stigma small-capitate,minutelypapillose, brightgreenin live plants.Fruita globose, green berry,0.8-1.2 cm diam.;fruitingpedicelsdeflexed and woody, 1-1.7 cm long, ca. 1 mm diam.atthebase.Seeds pale tan, flattened-reniform,3-4 x 1.5-2 mm, the margins incrassateandpaler,the surfacesminutelypitted. Chromosomenumber:n = 12 (vouchers Knapp 887, 4345, Knapp & Mallet 6791, 8480, 8515).

TAXONOMIC TREATMENT103 1972, Webster et al. 17900 (DS, MO). HIDALGO:Between Tamazunchaleand Jacala, 20 Jun 1965, Croat 917 (MO); along hwy. 85 between Tamazunchaleand Jacala, 37 mi W of Tamazunchale near Palomas, 1500 m, 25 Jun 1977, Croat 39320 (MEXU, MO); San Bartolo Selected specimens examined. MEXICO. Tlapacayo, Tutotepec, 1050 m, 5 Jun 1972, Gimate L. 653 (F); 1841-1843, Liebmann 15451 (F); Tehuantepec, Jul 1906, Rancho Viejo, La Misi6n, 28 Jun 1964, Gonzclez Quintero 1017 (F); Santa Maria, Santa Ana, 20 Sep 1964. Gandoger s.n. (MO); s.loc., Kerber 439 (US); s.loc., Sesse et al. 1510 (F); Carmen, Wawra 227 (W). CAMPECHE: Gonzalez Quintero 1624 (F); ca. 43 mi NE of Jacala, 3 Calakmul, 16 km NE of Zoh-Laguna, 262 m, 18?40'32N, Jul 1940, Hitchcock & Stanford 6969 (DS, F, US); km 89?21'16"W,4 Aug 1997, Martinez S. et al. 28037 (BM, 343-344 on hwy. below Chapulhuacan, along Hidalgo-MEXU). CHIAPAS:SW of Pinola, 3750 ft, 28 Jul 1964, San Luis Potosi border,2100 ft, 22 Sep 1949, Moore 5046 Breedlove 6596 (DS, F); 12 mi W of Ocozocoautla, 3100 (BH); 0.5 km E of Rancho Viejo, 1640 m, 4 Jun 1982, 1982, Tenorio L. & Romero de T 552 (NY); Alumbresft, 11 Jun 1965, Breedlove 10273 (DS, LL, US); school house of Pokolum, paraje of Sibanilha', Tenejapa, 5200 Tianguistengo rd., 1 km from Santa Monica detour, 1880 m, 22 Jun 1983, TorresC. & Hernindez 3069 (NY); ft, 15 Jul 1965, Breedlove 11036 (DS, WIS); 3 mi SW of Pinola Las Rosas along rd. to Soyatitan, 4200 ft, 27 7 mi S of Chapulhuacan, Mex. 85, 21 Aug 1971, Wunderlin et al. 1127 (MO). MICHOACAN:Vic. of Jul 1965, Breedlove 11363 (DS, MEXU, NY, US); Morelia, 1950 m, Mar 1909, Arsene 2444 (MO, US), 15 paraje of Kulak'tik, Tenejapa, 6000 ft, 11 Aug 1965, Breedlove 11697 (F); 20 mi S of La Trinitaria along May 1909, Arsene 3074 (MO, US), 21 Aug 1912, Arsene Mex. hwy. 190, 2800 ft, 15 Aug 1965, Breedlove 11797 8701 (NY, US). MORELOS:Cerro Tepoxteco, Tepoztlan, 1500 m, 1 May 1963, Gonzalez Quintero 403 (DS, F); (DS, F, LL, US); 17 mi E of La Trinitaria along rd. to barrancanear Cuernavaca, 5000 ft, 9 May 1898, Pringle Lagos de Montebello, 500 ft, 12 Oct 1965, Breedlove & Raven 12998 (DS, MEXU, NY, US); 32 km N of 6837 (F, K, MO, NY, POM, US). NAYARIT:10 mi SE of Ocozocoautla along rd. to Mal Paso, 2500 ft, 19 Oct Ahuacatlan, on rd. to Barranca del Oro, 1100-1300 m, 7 Jul 1957, McVaugh 15178 (US). OAXACA:10 mi S of 1965, Breedlove & Raven 13590 (DS, F); 1 mi NW of Sola de Vega along rd. to Puerto Escondido, 7000 ft, 8 Comitan, along Mex. hwy. 190, 5500 ft, 6 Nov 1965, Breedlove & Raven 14145 (F, LL, US); Finca Carmen May 1965, Breedlove 9855 (DS, WIS); km 88.5. hwy. 131, 7 Jul 1983, Costich & Baldwin 1518 (BH); 4.5 mi just below paraje of Mahosik', 3800 ft, 9 Aug 1966, Breedlove 14914 (DS, LL, NY); 2-4 km below Ixhuatan S of Valle Nacional above hwy. 175 between Tuxtepec along rd. to Pichucalco, 1200 m, 27 Sep 1971, Breedlove and Oaxaca, 430 m, 29 Jun 1977, Croat 39711 (MO); 19896 (DS, MO); along Comitan River at its sumidero, ca. 11 mi SW of Sola de Vega on rd. to Puerto Escondido, Lagos de Montebello, 42 km NE of La Trinitaria, 1300 2080 m, 14 Aug 1975, Davidse 9684 (MO); La Soledad, m, 23 Oct 1971, Breedlove & Thorne 21173 (DS, MO); 8 Feb 1966, Ernst 2541 (US); 5 km N of village of Matias Ocosingo, rd. from Velasco Suarez to Sintalapa, 400 m, Romero, along Transisthmian hwy. (rt. 185), ca. 50 m. 12 Oct 1976, Calzada et al. 2771 (WIS); 3 km W of 27 Jul 1958, King 795 (TEX, WIS); Tuxtepec, Chiltepec Chiapas-Tabasco border, 19 Aug 1974, Conrad & and vie., ca. 20 m, 16 Jul 1940, Martinez-Calderon 26 Conrad 2986a (MO); 20 air km SW of Tenosique, 150(US). PUEBLA:4 km NE of Villa Juarez on Mex. 130 to 250 m, 19 Aug 1974, Conrad & Conrad 2988 (F, MO); Poza Rica, ca. 1300 m, 20?20'N, 97?56'W, 27 Jun 1969, 5 mi SE of Palenque on rd. to Chancala, Ocosingo and Marcks & Marcks 825 (F, US, WIS); Congregaci6n San Crist6bal de las Casas, 200 m, 4 Jul 1977, Croat Benito Juarez, valley of Rio Portamonedas, ca. 38 km N 40156 (MO); rd. from Palenque to Bonampak, 55 mi of San Pedro Tapanatepec,900 m, 16?43'N, 94?08'30"W, SE of Palenque, 370 m, 5 Jul 1977, Croat 40273 (MO); 15 Aug 1984, Maya J. 426 (NY); Rio Jalatengo, 30 km along Hwy. 199 from Palenque to Ocosingo, 55 mi SW N of Candelaria Loxicha, 1400 m, 14 Jun 1985, Garcia of Palenque, 930 m, 6 Jul 1977, Croat 40356 (MEXU, Mendoza & Torres1646 (NY). QUINTANA ROO: 10 km S of Tres Garantias,on way to Tomas Garrido,26 Apr 1982, MO); along hwy. 199 at Chital, 9 mi SW of Ocosingo, 1190 m, 7 Jul 1977, Croat 40372, 40375 (MO); 6 mi N Cabrera & de Cabrera 2528 (NY); Felipe Carrillo Puerto, of Ocozocoautla along gravel rd. to Apitpac, 1000 m, 9 W of town of Kana, near La Laguna, 21 Aug 1976, Perez Jul 1977, Croat 40572 (MO); 10 km S of Pueblo Nuevo S. 388 (F). SAN LUIS POTOSI: 52 km W of Valles on Solisthuacan, 1860 m, 28 Jul 1965, Lathrop 5996 (US); hwy. 86, ca. 1000 m, 21055'N, 99?25'W, 30 Sep 1965, El Triunfo, Escuintla, 14 Jul 1948, Matuda 18087 (DS, Roe & Roe 2243 (WIS); between Taman and San FranF, MEXU); 4 km SW of Las Rosas on rd. to Venustiano cisco, Tamazunchala,600-900 m, 29 Jun 1959, Rzedowski 11070 (F); La Conchita 2 km NE of Xilitla, 680 m, 28 Carranza, ca. 900 m, 16?20'N, 92?30'W, 8 Aug 1965, Roe et al. 989 (DS, F, WIS); paraje of Mahosik', Oct 1982, Tenorio L. & Romero de T. 2458 (NY). TABASCO:Tenosique, 15 km upriver from ranch of Sr. Tenejapa, 4800 ft, 12 Sep 1966, Ton 1189 (DS, F, US, WIS); E side of Lago Tsikaw in region of Lagos de Angel Zubieta, Punta de Montafia, 4 Jul 1981, Cowan Montebello, 4800 ft, 5 Jul 1967, Ton 2619 (DS, WIS); & Nino 3388 (NY); Mercedes, Balancan, 9-14 May above Mispilla, Venustiano Carranza, 3200 ft, 27 Jul 1939, Matuda 3005 (MEXU, NY); Tenosique, 14-16 1967, Ton 2661 (DS, WIS); 20 mi NW of Ocozocoautla Jun 1939, Matuda 3396 (F, MO, NY); Retiro, Tenosique, on rd. to Malpaso, 1700 ft, 17?00'N, 93?30'W, 16 Aug 19-25 Jun 1939, Matuda 3435 (DS, F, MO, NY); Boca Distribution (Fig. 49). Widely distributed in the Caribbean, Central America, and South America, from sea level to occasionally 2500 m, usually in secondary forest, often growing in dense stands.

104

FLORA NEOTROPICA

Cerro, Tenosique, 1-5 Jul 1939, Matuda 3579 (F, MEXU, Nee 29917 (BH, F, NY); 3 km SW of Campamento La Above Aguacates on rd. to Rancho Laguna, 100 m, 17?16'N,94?32'W,6 Mar 1984, Aee 30019 NY). TAMAULIPAS: del Cielo Biological Station, 4 km NW of G6mez Farias, (NY); 6.5 km W of Tlapacoyan, rd. to Teziutlan, 900 750-800 m, 20 Jun 1982, Diggs & Nee 2425 (BH, F); m, 20 Jun 1970, Nevling & Gomez Pompa 1113 (BH); 10 km NW of El Progreso,which is 18 km NW of Ocampo, Cerro La Martinica, NW of Banderilla, 1680 m, 10 Feb 1450 m, 23? N, 99?30'W, 22 Aug 1941, Stanford et al. 1976, Ortega et al. 152 (NY); 2 km beyond detour, on 1044 (DS, MO, NY); between Rancho del Cielo Bioway to Ingenio La Concepci6n, 1130 m, 25 Jun 1976, logical Station and Old Company clearing, on rd. to Ortega 274, 341 (F, MO); sides of Mexico-Jalapa rd., Julilo, 3730-4000 ft, 15 Jun 1971, Sullivan 470 (TEX); 500 m before Martinica, 1400 m, 19?35'N, 96?56'W, Rancho del Cielo, 11 km W of G6mez Farias, 1190 m, 20 Oct 1977, Ortega Ortiz 722 (F); Jalapa, 19 Jul 1932. 25 Jun 1983, Torres C. 3130 (NY); San Luis, Jul 1930, Plunkett 51 (F); Zacuapan, Jun 1926, Purpus 10766 Viereck 732 (US). VERACRUZ:Tenzonapa, Tlacolulan, (US), Nov 1926, Purpus 10849 (US), May 1936, Purpus 16732 (US); El Marzo NW of Santa Ana Atzacan, 1315 1810 m, 27 Feb 1976, Avendano R. et al. 150 (F, NY); SE side of Laguna Catemaco, above Rio Cuetzalpan, 700 m, 9 Apr 1967, Rosas R. 296 (DS, MO, U, US, WIS); m, 18?23'N, 95001'W, 22 Oct 1971, Beaman 5156 (F, Carta Blanca between Orizaba and C6rdoba, 1100 m, 28 Apr 1967, Rosas R. 396 (DS, LL, U, WIS); Ejido de NY); NE side of Laguna Catemaco, E of Coyame, 450 m, 18?27'N, 95?01'W, 29 Oct 1971, Beaman 5204 (F); Tepetlampa, lower slopes of Cerro de Ejecatepetl, 1214 km NW of Campo Experimental de Hule, El Palmar, Orizaba, Botteri 1094 (F, K, US); Orizaba, Jul 1856, Botteri 1357 (NY, US); Orizaba, 12 May 1865, Boiurgeau Zongolica, 1000-2390 ft, 7 Jun 1944, VeraSantos 3024 2405 (K); region of Orizaba, 12 May 1865, Bourgeau (TEX); C6rdoba, 2700 ft, 20 Aug 1891, Seaton 429 (F); 2408 (NY, P); Misantla, rd. to Buenos Aires, 19?56'N, Orizaba, Rio Jalapilla, ca. 3.5 km from city center Jalapa, 15 Jul 1974, Sohmer 9549 (F); Rio Soloxuchil, 96051'W, 13 Dec 1977, Calzada 4115 (BH); Banderilla to Jilotepec, 1 km from rd., 1530 m, 19?36'N, 96?57'W, Campemento Hnos. Cedillo, 152 m, 17?N 94?37'W, 29 Mar 1974, Vdsquez 161 (MO); Ejido Santa Marta, 8 Nov 1977, Castillo C. 205 (F); Jalapa, Coulter 1238 (K); 3 km WNW of Cuichapa on rd. to Coetzala, 1.5 km Soteapan, 1250 m, 18?25'N, 94?56'W, 11 Nov 1980, SE of Coetzala, 550-700 m, 18?47'N, 96?54'W, 3 Jul Vdsquez B. & Herndndez 21 (F); Cerro Tenango, 1982, Diggs et al. 2726 (BH, F); 2 km beyond El Recreo, Zongolica, 1350 m, 18?40'N, 96?59'W, 4 Mar 1976, rd. to Palma Sola, 740 m, 16 Jun 1972, Dorantes et al. Vdsquez T 222 (F); border with Omealca, Coetzala, 600 737 (DS, F); Salto de Gato, Jalapa, 2 May 1973, Dorantes m, 16 Jul 1976, VdsquezT7456 (F, NY); between Zongolica & Acosta 2012 (F); Cerro de Macuiltepec, N of Jalapa, and Nacaxtla, 1600 m, 10 Nov 1976, Visquez T1556 1500 m, 18 May 1973, Dorantes & Acosta 2083 (LL); (F); La Barranca de Metlac, Fortin, 880 m, 8 Jun 1972, Ventura A. 5510 (DS, NY); El Mirador, Totutla, 1000 Orizaba, 4000 ft, 9 Aug 1924, Fisher 254 (MO, US); m, 26 Apr 1973, VenturaA. 8219 (F); Macuiltepec, 1450 Jalapa, Galeotti s.n. (K); Cordillera, Jun-Oct, Galeotti 1155 (K); 5 km NE of Coscomatepec, 1 km W of turn- m, 24 Apr 1974, VenturaA. 9930 (F); El Tejar, Jalapa, off to La Candelaria,along Mex. hwy. on way to Huatusco, 1300m, 10 May 1974, VenturaA. 10051 (F); El Mirador, 1450 m, 19?06'N, 97?01'W, 3 Jul 1980, Hansen & Nee Totutla, 1000 m, 7 May 1975, Ventura A. 11400 (F); 7604 (F, MO); Temascaltepec, Rinc6n del Carmen, 1460 Naranjo, Naolinco, 1350 m, 1 Jul 1975, Ventura A. 11551 (F); Chapultepec, Coacoatzintla, 1400 m, 4 Nov m, 7 Aug 1932, Hinton 1004 (NY); Papantla, Tajin, 22 1977, VenturaA. 14674 (F); km 2 on rd. from Concha to May 1948, Kelly 323 (BH, WIS); Jalapa, Jun 1838, Linden 234 (K); 11.6 km from Plan de las Hayas, 700 m, La Concepci6n, 1130 m, 25 Jun 1976, Zola B. 454 (F); 24 Jun 1972, Lot et al. 2018 (F); Las Tuxtlas biological Vista Hermosa, Jilotepec, 1050 m, 27 Jul 1976, Zola B. 565 (F); Cerro de Macuiltepetl, 1480 m, 9 Sep 1976, station, 145 m, 23 Jun 1968, Martinez-Calderon 1711 Zola B. 712 (F); rd. to Sumidero, Jalpa, 1210 m, 22 Sep (DS, F, NY, US, WIS); C6rdoba, Aug 1938, Matuda 17002 (F); Orizaba, 1853, Muller 1929 (W); Cascada 1976, Zola B. 753 (DS, F). YUCATAN:Progreso, Sep de Texolo, 3 km SE of Villa Jico (Xico), 1250 m, 19?24'N, 1932, Flores 3 (F); Tuxpena, Campeche, 16 Oct 1931, 97?00'W. 2 Jul 1980, Nee & Hansen 18737 (BH, DS, F, Lundell 838 (DS, F, K, MO, NY, US); Muna, 22-23 Jul MO. U, WIS); 3 km E of Tebanca (3 km E of E side of 1932, Steere 2099 (F). GUATEMALA. Finca Santa In6s, 1700-2000 ft, 21 Lago Catemaco), on rd. to Bastonal, 630 m, 5 Jul 1980, Nee & Hansen 18763a (BH, F); 13 km E of Tebanca Jan 1927, Record & Kuylen G.80 (US). ALTA VERAPAZ: (13 km E of E side of Lago Catemaco), on way to Santa Sebol, about 800 m SEE, 22 Apr 1964, Contreras 4454 Marta, 800-950 m, 5 Jul 1980, Nee & Hansen 18814 (F); near Finca Sepacuite, 7 Apr 1902, Cook & Griggs (BH, DS, F, WIS); 7.5 km N of Huejutla, on rd. to Plat6n 492 (US), 9 Apr 1902, Cook & Griggs 543 (US); along Sanchez and Tempoal, 250 m, 21?12'30"N, 98?23'W, 24 hwy. 14, 5 mi S of Coban, 1300 m, 17 Jul 1977, Croat Oct 1981, Nee 22352 (BH, F); 0-3 km E of Santa Marta, 41375 (MO); Secanquim, trail to Sepacuite, 28 Nov base of Volcan Santa Marta, 1100-1200 m, 18?21'N, 1904, Goll 91 (US); between Semacoch and Panzas, 10 94?52-3'W, 29 Jun 1982, Nee et al. 24694 (BH); along Apr 1905, Goll 209 (NY, US); Coban, May 1882, main gravel rd. of the Uxpanapa region, 5 km NW of El Lehmann 1429 (K, US); rd. between San Pedro Carcha Doce, 100 m, 17?13'N, 94?16'W, 4 Mar 1984, Nee & and Sacoyo6, ca. 1300 m, 11 May 1963, Molina R. & Taylor 29884 (NY); vic. of Campamento La Laguna, Molina 12078 (F, NY); near Coban, 26 Mar-15 Apr 1939, Hidalgotitlan, 100 m, 17?17'N, 94030'W, 4 Mar 1984, Standley 69170 (F, US), Standley 69252 (F), Standley

TAXONOMIC TREATMENT 69539 (F, US); Saquija, NE of Coban, 1200 m, 2 Apr 1939, Standley 70160 (F); near Pancajch6, ca. 360 m, 5 Apr 1939, Standlev 70666 (F); near Tucurf, ca. 450 m, 5 Apr 1939, Standley 70706 (F); above Tamahui,9001200 m, 5 Apr 1939, Standley 70915 (F); near Coban, 1260-1400 m, 26 Mar-15 Apr 1939, Standley 71578 (F); along Rio Carcha, between Coban and San Pedro Carcha, ca. 1360 m, 26-27 Mar 1941, Standley 89995 (F, US); along Rio Polochic, above Tamahf, ca. 1200 m, 10 Apr 1941, Standley 91806 (F); SE of Finca Yalpemech, near Alta Verapaz-Pet6n boundary line, ca. 100-150 m, 23 Mar 1832, Steyermark45208 (F); Coban, 4400 ft, May 1879, von Tuerckheim 436 (F); Coban, 1350 m, Jun 1906, von Tuerckheim111270 (F, LL, MO, NY, US); Cubilqiitz, ca. 350 m, Aug 1900, von Tuerckheim 7754 (MO, NY, US); Coban, 1400 m, Feb 1903, von Tuerckheim8437 (F, K, NY, US); ca. 8 km SW of Coban, 1500 m, 29 Jan 1969, Williamset al. 40264 (F, NY, US). BAJA VERAPAZ: Near and above Santa Rosa, ca. 1500 m, 4 Apr 1941, Standley 91085 (F, US). CHIQUIMULA:Rio Esquipulas, 1100 m, 26 Sep 1971, Molina R. & Molina 26738 (F). GUATEMALA:S.loc., Agzilar 567, 588 (F); km 28 F.D.R. between San Lucas and Guatemala City, 2000 m, Molina R. et al. 16657 (NY); Finca La Aurora, 15 Jun 1923, Ruano 328 (US), 1 Aug 1926, Ruano 573 (US); near Guatemala, 1400 m, Jul 1921, Tonduz624 (US). HUEHUETENANGO: 3 mi NW of Santa Cruz Barillas along rd. to San Mateo Ixtatan, 6000 ft, 7 Aug 1965, Breedlove 11651 (DS, LL, US). IZABAL: Barbasco, Aug 1870, Bernoulli 944 (NY); Los Amates, 160 ft, 6 Feb 1905, Deam 92 (F, MO, NY, US); 2.5 mi N of Rio Dulce on rd. to Pet6n, 8 Aug 1975, Dunn & LeDoux 22015 (MO, NY); Los Amates, 90 m, 15 Mar 1905, Kellerman 5123 (US); Morales, 8 Mar 1907, Kellerman 671? (F); Los Amates, 22 Feb 1908, Kellerman 7535 (NY); Rio Dulce, Livingston, 0 m, Mar 1889, Smith 1839 (F, K, US); vic. of Quirigua, 75-225 m, 15-31 May 1922, Standley 23741, 24188 (NY, US); vic. of Escoba, on bay opposite Puerto Barrios, ca. 150 m, 2 Jun 1922, Standley 24859 (NY, US); vic. of Puerto Barrios, 0 m, 2-6 Jun 1922, Standley 24923 (NY, US); near Entre Rios, ca. 18 m, 30 Apr 1939, Standley 72682 (F); between Milla 49.5 and ridge 6 mi from Izabal, Montafia del Mico, 65-300 m, 2 Apr 1940, Steyermark 38633 (F); between Escobas and Montafia Escobas, across bay from Puerto Barrios, 13 Apr 1940, Steyermark 39261, 39319 (F). PETEN: Vaxactun, 23 Apr 1931, Bartlett 12706 (F, NY); Macanch6, 28 Mar 1966, Contreras 135 (F); Dos Lagunas, El Cedro, km 8, 21 Apr 1969, Contreras 8366 (F, LL); 2 mi S of entrance of Tikal National Park, ca. 500 ft, 19 Jun 1973, Croat 24710 (MO); ca. 5 mi S of Tikal National Park, 19 Jun 1973, Dwver 11266 (F, MO); N shore Lake Pet6n, 19 Jun 1973, Dwyer 11278 (MO, WIS); Carmelita, 22 Jun 1942, Egler 42-196 (F, LL); S of Tikal National Park, 19 Jun 1973, Gentry 8335 (MO); 9 km W of Modesto Mendez, 150 m, 9 Jun 1970, Harmon 2499 (MO); 27 km N of Poptun, 500 m, 17 Jun 1971, Harmon & Fuentes 5732 (MO); Lake Pet6n, 5 May 1933, Lundell 3224 (F); Poptun, 500 m, 11 Nov 1965, Molina R. 15584 (F, NY);

105 0.5 km from Poptin, 14 Apr 1970, Tun Ortiz 945 (F, MO, NY); rd. to El Remate, km 58 in Parque Nacional Tikal, 28 Aug 1970, Tun Ortiz 1250 (F, MO, NY, US); San Luis, 18 Mar 1976, Ventur14 (F). QUICHE:Cunen, Apr 1892, Heyde & Lux 3449 (F, K, US). SACATEPEQUEZ: Los Amates, 15 Feb 1908, Kellerman 7429 (F, NY, US); near Pastores, 1560-1650 m, 14 Dec 1938, Standley 59937 (F). SANTAROSA: Rio Los Esclavos, 2500 ft, Jul 1892, Heyde & Lux 3438 (F, K, US). ZACAPA:Along old rd. to Finca Agua Fria 2 mi N of rt. 9, 41 mi S of turnoff to Peten, near Morales, 150 m, 23 Jul 1977, Croat 41870 (MEXU, MO). BELIZE. El Cayo, 21 Apr 1931, Bartlett 12962 (NY, TEX, US); San Antonio, 6 May 1931, Bartlett 13042 (F); ca. 3 mi S of Granode Oro on rd. between Millionario and La Flor, ca. 1700 ft, 2 Jun 1973, Croat 23397 (F, MO); vic. of Cuevas, S of Millionario, ca. 1900 ft, 2930 May 1973, Croat 23533 (MO); along rd. at Rio Ma Cal, between San Luis and Cuevas, ca. 1050 ft, 28 May 1973, Croat 23533a (MO, WIS); vic. of Cuevas S of Millionario, ca. 1900 ft, 29-30 May 1973, Croat 23551 (F, LL, MO, RSA); Gracie Rock, 1.5 mi S of mi 22 on western hwy., 4-5 Jun 1973, Croat 23903 (MO); along southern hwy., 14.5 mi W of Punta Gorda near jct. with rd. to San Antonio, 11 Jun 1973, Croat 24129 (MO); along trail to Esperanza beginning 1 mi N of Columbia Forest Station, 500-1100 ft, 12 Jun 1973, Croat 24224 (F, MO); vic. of San Jose, Mayan Indian village, 6.7 mi N of Columbia Forest Station, 13 Jun 1973, Croat 24430 (F, MO); near Guatemalanborder, between Benque Viejo and Melchor, 18 Jun 1973, Croat 24609 (MO); along Hummingbird Hwy. S of Belmopan, between mi 30 and 38, 300 ft, 21 Jun 1973, Croat 24830 (MO); 1 mi W of northern hwy. on secondary rd., 1 mi W of Buena Vista, ca. 100 ft, 23 Jun 1973, Croat 24950 (MO); Hummirgbird Hwy. at St. Margaret's Creek, 19 Aug 1969, Dw1ter 9117 (F, MO); Columbia Forest Station, 24 Jul 1970, Dwyer 9893 (MO); at Millionario on rd. to Cuevas. 1900 ft, 30 May 1973, Dwyer 10782 (MO); Millionario, 25 May 1973, Dwyer 10802a (MO, NY); Burrel Boom, 9 Jun 1973, Dwyer 11059 (F); San Jose, 6.7 mi N of Columbia Forest Station, 13 Jun 1973, Dwyer 11185 (F, MO); Belmopan to 10 mi S on Hummingbird Hwy., 6 Jun 1974, Dwyer 12725 (MO); Corozal, 1933, Gentle 29 (F); El Cayo, Vaca, 30 Apr 1938, Gentle 2531 (F. LL); San Antonio, 14 Jul 1945, Gentle 4767 (BH, F, LL); near St. Margaret Creek, Hummingbird Hwy., 28 May 1955, Gentle 8731 (F); Cohune ridge, Sibun River, below the Hummingbird Hwy., 13 Jun 1955, Gentle 8749, 8752 (F); mi 41, Hummingbird Hwy., 12 Dec 1955, Gentle 8984 (F); on Cayo River, 29 Apr 1958, Gentle 9738 (F); S of Millionario, 1900 ft, 29 May 1973, Gentry 7679 (LL, MO, RSA); vic. of Grano de Oro lumber camp, S of Millionario, 1700 ft, 2 Jun 1973, Gentryl 7805 (F, MO); vic. of Columbia Forest Station, N of San Antonio, 300-500 ft, 11-12 Jun 1973, Gentry 8090 (MO); Rio Frio Cave trail, near Augustine, Pine Ridge area, 9 Jul 1972, Huston 634 (MO); Tower Hill, 1928, Karling 14 (F, K, US); mi 50 western hwy., near Roaring Creek, 13 Jul 1970, Long 3216 (MO); Mt. Pine Ridge, 5 mi S of

106 Augustine rd., ca. 35 mi S of San Diego, 13 Jul 1970, Long 3235 (MO); Honey Camp, Aug 1929, Lundell 435 (F, K, NY, US); Roaring Creek, Aug 1929, Lundell 443a (F, US), Lundell 469 (F, NY, US); near Honey Camp, 7 Aug 1930, Meyer 79 (F); Temascal River, May 1894, Moloney 29 (K); Toledo, 30 Mar 1907, Peck 808 (K, NY); Middlesex, 30 Jul 1929, Schipp 312 (F); Middlesex, 200 ft, 8 Nov 1929, Schipp 429 (F, K, MO, NY); 19 Mi, 250 ft, 18 Jun 1932, Schipp 959 (F, K, MO, NY); at Rio Frio Cave near Augustine, ca. 300 m, 5 Jul 1970, Spellman 1580 (MO); 7 mi S of jct. of Hummingbird and western Hwys., N of Roaring Creek, ca. 80 m, 13 Jul 1970, Spellman & Newey 1667 (MO); 2.5 mi S of jct. of Hummingbird and western Hwys., 17 Jul 1970, Spellman 1707, 1766 (MO); ca. 1 mi S of Belmopan, 1/ 2 km E of Hummingbird Hwy., ca. 75 m, 26 Jul 1970, Spellman & Newey 1879 (MO), Spellman & Newey 1889 (MO); 7 mi from jct. of Hummingbirdand western Hwys., N of Roaring Creek, ca. 120 m, 8 Aug 1970, Spellman & Newey 1976 (MO); Caves Branch, along Hummingbird Hwy. between Base Camp and Blue Hole, 26 Jul 1976, Whitefoord 1126 (BM, MO); St. Herman's Hill, track from Hummingbird Hwy. to St. Herman's Cave, 1 Aug 1976, Whitefoord1196 (BM, MO); Cayo, Guacamayo bridge, Cotton Tree Creek, ca. 450 m, 8 Aug 1980, Whitefoord 2050 (BM, F, MEXU, MO); Hummingbird Hwy. E of Over-the-Top Camp, 10 Sep 1980, Whitefoord 2424 (BM, MO); W of Forestry Camp toward San Jose, 5 Aug 1970, Wiley 65 (MO, US), Wiley 67 (MO); s.loc., 10 Oct 1926, Winzerling V-5, tree no. 21 (F, US); Indian Church, Bajo, 20 Sep 1977, Uck 17813 (MO); 60.5 mi N of Belize City on Orange Walk rd., 1/2 mi S of bridge over New River, just N of Carmelita, 4 Sep 1970, Ugent 7 (WIS); 200 yds. S of Blue Hole on Hummingbird Hwy., 28 May 1971, Utley 799 (MO). HONDURAS. Puerto Sierra, 19 Feb 1903, Wilson 479 (F, NY, US). ATLANTIDA: Along Rio Danta, rd. to La Presa, vic. of La Ceiba, 160 m, 25 Apr 1967, Molina R. 20828 (F); vic. of San Alejo, near Rio Alejo, 150270 m, 22-27 Apr 1947, Standley 7928 (F); Lancetilla valley, near Tela, 20-600 m, 6 Dec 1927-20 Mar 1928, Standley 53235 (F, US), Standley 54333 (F), Standley 54361 (F, US), Standley 54761, 55178, 55574 (F, US); above Lancetilla, ca. 90 ft, 10 Jul 1934, Yuncker4518 (F, MO); foothills back of Ceiba, 8 Jul 1938, Yunckeret al. 8301 (F, MO, NY), 6 Aug 1938, Yunckeret al. 8841 (F, MO, NY); ca. 3 mi S of Tela, 200-500 ft, 27 Jul Vic. of 1962, Websteret al. 12583 (WIS). CHOLUTECA: Pespire, 160-200 m, 18-25 Oct 1950, Standley 27197 (F). COLON: Trujillo, Rio Negro, 100-200 ft, 19 Oct El Play6n, 1981, Saunders 641 (MO, NY). COMAYAGUA: San Luis, 3500 ft, 20 May 1932, Edwards P241 (F); Pito Solo, Lake Yojoa, 2000 ft, 19 Aug 1932, Edwards p447 (F, US); near San Jeronimo, valley of Comayagua, 800 m, 13 May 1956, Molina R. 6987 (F); vic. of Comayagua, ca. 600 m, 12-13 Mar 1947, Standley & Chac6n P. 5884 (F); Pitosolo, Lago Yojoa, 500 m, 17 Apr 1945, Valerio R. 2891, 2948 (F); La Misi6n, 600 m, 18 Apr 1945, Valerio R. 2964 (F); Lake Yojoa, 630 m, 9 Aug 1948, Williams & Molina R. 14607 (F, MO,

FLORA NEOTROPICA US); along river at Siguatepeque, 1100 m, 29 Sep-5 Oct 1951, Williams & Williams 18469 (F); river near Siguatepeque, 1050 m, 9 Jul 1936, Yuncker5761 (F, MO, NY, U, US). COPAN:Along Yaragua creek, 1 mi W of Copan ruins, 500 m, 29 Aug 1975, Molina R. & Molina 30840 (F, MO). CORTES: Cuyamel, 3 Mar 1923, Carleton 459 (US); Puerto Cort6s, Finca Las Penas de Ostnmark, 23 Aug 1964, Dickson 1290 (US); Corinto, 55 km W of Puerto Cort6s, 9-11 Aug 1975, Nelson et al. 2918, 2936 (MO); San Pedro Sula, 600-1500 ft, May 1890, Thieme 5374 (F). EL PARAISO:San Juanito and vic., Aug 1960, Pfeifer 2005 (US); Rio Choluteca near Ojo de Agua, 600 m, 31 Dec 1950, Standley 28016 (F); along Rio Choluteca, 600 m, 18 Oct 1951, Standley 28960 (US). INTIBUCA:Cordillera Opacala between Pela Nariz and Calaveras, rd. to La Esperanza, 2000 m, 3 Sep 1968, Molina R. 22640 (F, NY). ISLASDE LA BAHIA: Isla Ruatan, 1885, Gaumer 6 (K), Gaumer 22 (F, K, US). MORAZAN: Santa Lucia, 11 km N of Tegucigalpa, 1400 m, 18 Feb 1984, Berrios 150 (NY); El Picacho, N of Tegucigualpa, 1300 m, 25 Oct 1978, Martinez 124 (MO); along Quebrada Suyapa, 1100 m, 10 Nov 1947, Molina R. 574 (F, US), 20 Dec 1947, Molina R. 711 (F); near Suyapa, 1200 m, 25 Nov-14 Dec 1946, Standley & Williams 1405 (F); vic. of Suyapa 1100-1200 m, SepDec 1948, Standley 12966 (F); Cerro de Uyaca, La Labranza and vic., 1600-1800 m, 2 Jul 1949, Standley 20617 (F). OCOTEPEQUE: 17 km N of Nueva Ocotepeque, 13 Aug 1970, Harmon & Fuentes 3791 (NY). OLANCHO: Vic. of Juticalpa, 380-480 m, 5-16 Mar 1949, Standley 17696 (F); vic. of Catacamas, 450-500 m, 18-26 Mar 1949, Standley 18179 (F). SANTABARBARA:Between Ceguaca and Concepci6n del Sur, 600 m, 21 Aug 1968, Molina R. 21952 (F); La Encenada, margins of Lago do Yojoa, 600 m, 1 May 1984, Paredes 151 (NY). YORO: 3 km NW of Santa Rita on rd. to Negrita, 14 Aug 1970, Harmon & Dwyer 3895 (MO); 7 km NE of Santa Rita on rd. to La Negrita, 14 Aug 1970, Harmon & Dwyer 3899 (MO); La Lima, Sep 1929, Johansen 46 (NY); Quebrada Olotillo, 15 km from Yoro, 1100 m, 8 May 1956, Molina R. 6829 (F); near Rio Aguan, below Coyoles, 29 Jun 1938, Yunckeret al. 8106 (F, MO, NY). EL SALVADOR. CHALATENANGO: E slope of Los Esesmiles, 2100-2300 m, 14?21'N, 89?09'W, 1 Apr Near Col6n, 1942, Tucker1186 (BH, US). LA LIBERTAD: 2000 m, 21 Jan 1949, Williams & Molina R. 15213 (F). SAN VICENTE:Vic. of San Vicente, 400-500 m. 7-14 Feb 1947, Standley & Padilla B. 3600 (F). SANTAANA: Between Matapan & Cerro Monte Cristo, 7 mi NE of Metapan, 1200 m, 30 Jul 1977, Croat 42308 (MO). NICARAGUA. CHONTALES: Chontales, 600 m, Jun 1870, Levy 470 (K); vic. Finca San Pedro de Oluma, on NE flanks of Cerro Oluma, 4 km N of Cuapa, 600 m, 12?18'N, 85?23'W, 22 Sep 1983, Nee 28345 (MO, NY); vic. of La Libertad, 500-700 m, 29 May-i Jun 1947, Standley 8851 (F); vic. of Juigalpa, ca. 160 m, 4-13 Jun 1947, Standley 9439 (F); Chontales, 1867-68, Tate 437 (K). ESTELI: Along Rio Ducuali, vic. of Ducuali, 600 m, 6 Nov 1968, Molina R. 23187 (F, MO, NY); La Gavilana, 940-1000 m, 13?03'N, 86?19'W, 2 Aug 1983,

TAXONOMIC TREATMENT Moreno 21876 (MO, NY); 10 km SSE of Esteli along Hwy. 1, 1000 m, 13?01'30"N, 86?19'W, 1 Sep 1983, Nee 27716 (MO, NY); 14 km S of Esteli, ca. 5 km from main rd. (Hwy. 1) on rd. to San Francisco "TierraBlanca," 1100 m, ca. 12?58'N, 86?19'W, 1 Sep 1983, Nee et al. 27727 (MO, NY); 4 mi S of Esteli, 4 Aug 1971, Wunderlin et al. 444 (F, MO). MANAGUA:Vic. of Managua, Lake Nejapa, 1 Jul 1923, Maxon et al. 7554 (NY, US). MATAGALPA:Along Rio 10-15 km NE of Matagalpa, 600-700 m, 16 Jan 1963, Williams et al. 4011 (F, NY, US); 15 km NE of Matagalpa along Rio Las Cafias, 700800 m, 14 Jan 1965, Williams et al. 27516 (F, MO). RIVAS: Hacienda Fatima, SW of Sapoa, 11?10'N, 85?40'W, 4 Sep 1982, Sandino 3511 (MO, NY). ZELAYA: Near Siuna, Mt. Liveco, Madregava, 7 Jan 1970, Atwood 3236 (F, MO, NY); vic. of La Luz-Siuna, 150-200 m, 14 Mar 1961, Bunting & Licht 638 (F, NY, US); region of Braggman's Bluff, 4 Jan 1928, Englesing 105 (F, K, US); El Recreo, 9 Jul 1948, Long 191 (F); along rio Plata, just past Colonia Rio Plata, ca. 150 m, 11?43'N, 84?26-27'W, 20 Aug 1983, Miller & Sandino 1186 (MO, NY); area of Bahia de Bluefields, Rio Escondido, 0-30 m, 24 Mar 1949, Molina R. 1976 (F); along Rio Grande, 15 m, 21 Apr 1949, Molina R. 2259 (F); near Siuna, 200-500 m, 7 Jan 1970, Narvdez S. 3184 (F, MO, WIS); 0-1 km N of San Martin, ca. 120 m, 11?53'N, 84?21'W, 6 Sep 1983, Nee 27816 (MO, NY); Cerro San Isidro, Rio Kama, Rio Escondido, 0-65 m, 12?05-15'N, 83045'-84020'W, 27 Mar 1966, Proctor et al. 27276 (F, LL, NY); vic. of El Recreo, on Rio Mico, ca. 30 m, 23 Apr-14 May 1949, Standley 19132 (F). COSTA RICA. Aguacate, Oersted 1409 (US, W). ALAJUELA:Los Angeles de San Ramon, 20 Nov 1923, Brenes 3947 (NY); along rd. from Caias to Upala, 13.8 km N of Bijagua, 100-150 m, 26 Jun 1976, Croat 36422 (MO); Palmira, region of Zarcero, 30 Aug 1937, Smith A258 (F, MO); Zarcero, 1500 m, 10 Feb 1948, Smith 48/212 (US); Palmira, 7000 ft, 28 Apr 1937, Smith 4138 (F, NY); Zarcero, 1900 m, 1941, Smith 10051 (F). CARTAGO:Between Jicotea and Rio Pacuare along rd. from Turrialba to Moravia, 650-750 m, 9?50'N, 83030'W, 19--20 Dec 1966, Burger & Ramirez B. 3986 (CR, F, MEXU, MO, NY, U); Turrialba, rd. to Pavones, 750 m, 27 May 1951, Carpenter 259 (US); rd. near Rio Turrialba, 27 Apr 1903, Cook & Doyle 375 (US); Turrialba, 20 Jan 1949, Gregory 1883 (US); Turrialba, 650 m, 13 Oct 1953, Heiser Jr. 3631 (US); lot 5, CATIE, Turrialba, 600 m, Jan 1949, Ledn 1454 (US); lot 11, CATIE, Turrialba, 600 m, Apr 1949, Le6n s.n. (US); vic. of Pejivalle, ca. 900 m, 7-8 Feb 1926, Standley & Valerio 46800, 46839 (F, US); along Rio Attirro ca. 2 km E of La Esperanza, 850 m, 9?47'N, 83?38'W, 1 Jan 1975, Taylor 18009 (NY, US); Turrialba, Oct 1894, Tonduz9018 (US); Rio Tuis, 650 m, Nov 1897, Tonduz 11491 (US); Las Vueltas, Tucurrique, 635 m, Nov 1898, Tonduz 12763 (K, US). GUANACASTE:Hacienda La Pacifica, near Canfas, 19 Jul 1983, Costich & Baldwin 1529 (BH); vic. of Cainas, 18 Aug 1969, Daubenmire 117 (F); La Pacifica, 10 Apr 1973, Gentry & Burger 2929 (F, MO); Finca La Pacifica, edge of Rio Corobici,

107 100 m, 9 Jul 1971, Opler 316 (F), 30 Jun 1972, Opler 919 (F, MO). LIMON:Rd. to La Estrella, 0-5 m, 28 Feb 1958, Cdrdoba 468 (MO); between Rio Bananito and Cahuita, 0-10 m, 9043-53'N, 82?48-59'W, 9 Feb 1977, Gentry 3736a (F); Livingston on Rio Reventaz6n, JulAug 1920, Rowlee & Stork 714 (CU, NY, US); La Columbiana farm of United Fruit Co., ca. 70 m, 6-7 Mar 1924, Standley 36655 (US); Finca Montecristo, along Rio Reventaz6n below Cairo, ca. 25 m, 18-19 Feb 1926, Standley & Valerio 48408, 48426 (US). PUNTARENAS: Near Palmar Norte, 100 ft, 11 Apr 1949, Allen 5261 (F, MO, US); Osa peninsula, Parque Nacional Corcovado, Estaci6n Sirena, near mouth of Rio Sirena, 0 m, 8?30'N, 83?27'W, 13 Jul 1981, Knapp 887 (BI1); Corcovado National Park, between Sirena and Pavo, 010 m, 8?30'N, 83?36'W, 5 Jul 1977, Liesner 2888 (MO); Corcovado National Park, near Pavo, 5 m, 8?30'N, 83?37'W, 7 Jul 1977, Liesner 3035 (MO); 3 km S of Rinc6n, 40 m, 7 Mar 1971, Nee & Mori 3561 (F, MAD, US, WIS); 1 mi S of San Vito de Java, 350 m, 18 Aug 1967, Raven 21917 (CR, MO); Golfo Dulce and Rio Terraba, 30 m, Dec 1947, Skutch 5392 (US). PANAMA. S.loc., Seeman 424 (K); Jan 1860, Hayes 299 (NY), Hayes 753 (NY). CANALAREA: Barro Colorado Island, 1931, Aviles 72 (MO); Madden Forest Rd., 22 Mar 1970, Croat 8958 (MO); NW of Escobal, 30 Sep 1970, Croat 12455 (MO); Pipeline Rd. near gate, 5 Mar 1971, Croat 13940 (MO); Madden Forest, 4 May 1971, Croat 14541 (MO), D'Arcy 5243 (MO, MPU); SE of San Pedro Miguel, Madden Forest, 27 Jul 1972, D'Arcy & D'Arcy 6093 (F, K, MO, NY, RSA, US, WIS); Barro Colorado Island, 9 Apr 1970, D'Arcy 3988 (MO, NY); Barro Colorado Island, creek below Fuertes house, 30 May 1969, Foster 887 (F, MO); Gatun Station, 7 Feb 1860, Hayes 144 (NY); Agua Clara Reservior, near Gatun, 20-30 m, 5 Feb 1911, Pittier 2653 (NY, US); Cruces, Aug 1847, Seemann 424 (K); Barro Colorado Island, 120 m, 18-24 Nov 1925, Standley 40860 (US): Barro Colorado Island, cove N of Drayton House, 14 Feb 1932, Woodworth & Vestal 555 (MO). CHIRIQUi: Along rd. in vic. of branch to Cerro Colorado and Escopeta, above Rio San Felix near town of San F,elix, ca. 13 mi N of Rio San Felix bridge, 800-1200 m, 15 Mar 1976, Croat 33407 (MO); above Paso de la Canoa, 3000 ft, 11 May 1971, D'Arcy 5431 (MO); Cerro Horqueta, 1666-2333 m, 8 Aug 1967, Kirkbride 135 (NY); Boquete, 1400 m, 29 Jul 1972, Maasola 41 (MO); 17 km NE of San Felix on rd. to Cerro Colorado copper mines, 13-14 km NE of bridge over Rio San Felix, ca. 1000 m, 18-19 Mar 1974, Nee 10755 (MO). COCLE: Vic. of El Valle, lower Rio Ant6n, 30 Dec 1936, lllen 101 (MO); vic. of El Valle, 600-1000 m, 8 Dec 1938, Allen 1199 (MO, NY, US); vic. of La Mesa, 11 Feb 1971, Croat 13331 (MO, NY); below Cerro Pil6n, 11 Feb 1971, Croat 13437 (F, MO); La Mesa above E1 Valle, 900 m, 16 Mar 1973, Croat 22960 (F, LL, MO): La Mesa above El Valle, jct. with rd. to Cerro Pil6n, ca. 800 m, 21 Jul 1974, Croat 25420 (MO); El Valle de Anton, 13 Aug 1972, D'Arcy & D'Arcy 6722 (BM, F, K, MO. NY, US, WIS); rd. to Coclesito from Penonom&,ca. 7 km N

108 of Llano Grande,450 m, 11 Oct 1978, D'Arcy & Hammel 12270 (MO); La Mesa N of El Valle de Anton, 400-800 m, 14 Feb 1981, D'Arcy & Sytsma 14667 (MO); Cerro Pilon near El Valle, 700-900 m, 10 Jun 1967, Duke 12058, 12104 (MO, WIS); El Valle de Anton at foot of Cerro Pilon, ca. 2000 ft, 15 Aug 1967, Dwyer & Correa A. 7943 (F, MO, MPU, SCZ), Dwyer & Correa A. 7970a (MO, SCZ); Cerro Pilon, ca. 2700 ft, 14 Sep 1968, Dwyer & Lallathin 8614 (MO, MPU); 7.1 km from El Valle rd. on La Mesa rd., 700-800 m, 25 Jun 1977, Folsom 3906 (MO); forest around Limon, 5 hrs walk N of Alto Calvario, N of El Cop&, 800-1000 M, 10 Oct 1977, Folsom 5849 (MO); above El Valle, 13 Aug 1972, Gentry 5678 (MO); along rd. from La Pineda to El Cope by way of Piedras Gordas, 3000 ft, 20 Apr 1978, Hammel 2601 (MEXU, MO); La Mesa region of Cerro Gaital, 2400 ft, 2 Jul 1978, Hammel 3847 (MEXU, MO, NY); Continental Divide N of Penonome on rd. to Coclesito, 1600 ft, 25-26 Jul 1978, Hammel 4028 (MO); La Mesa, El Valle, 800 m, 25 Apr 1967, Hladik 339 (MO, P); valley and marshes along Rio Anton, El Valle de Anton, ca. 500 m, 2 Feb 1935, Hunter & Allen 368 (MO); El Valle de Anton, La Mesa, ca. 1000 m, 1 Apr 1973, Kennedy et al. 3015 (MO, US); 2 mi N of Cerro Pilon, ca. 900 m, 16 Mar 1973, Liesner 712 (F, MO, NY, US); Boca del Toabr&at confluence of Rio Toabre and Rio Cocle de Norte, 11 Apr 1969, Lewis et al. 1522 (BM, MO), 1570 (MO); El Valle de Anton, 1000-2000 ft, 2-3 Dec 1967, Lewis et al. 2516 (MO); El Valle, 23 Feb 1937, Miller 1835 (US); La Mesa, 4 km N of El Valle, 875 m, 3 Jan 1974, Nee & Dwyer 9161 (MAD, MO, MPU); foot of Cerro Pilon, above El Valle de Anton, 2000 ft, 27 Mar 1969, Porter et al. 4580 (MEXU, MO), 28 Mar 1969, Porter et al. 4604 (MO); La Mesa, 10 Aug 1971, Spellman et al. 591 (F, MO); mtns. above El Valle, 10 Jun 1967, Stimson 5014 (NY, US); between Las Margaritas and El Valle, 15 Jul-8 Aug 1938, Woodson et al. 1276 (F, MO, NY). COLON:To Rio Iguanita,390 m, 7 Apr 1977, D'Arcy 11248 (MO); Rio Iguanita, ca. 2 mi from the sea, 5 May 1979, D'Arcy et al. 13349 (MO); Rio Iguanita, and inland 2 km, 0-50 m, 7 Feb 1980, D'Arcy 14617 (MO). DARIEN: 3.7 mi W of Santa Fe on PanAmerican hwy., 0.7 mi N on logging trail, 20-50 m, 25 Apr 1982, Huft et al. 1948 (MO, NY). PANAMA: Rio Tapia, 7 Dec 1923-11 Jan 1924, Standley 28206 (US). VERAGUAS: 5 mi W of Santa Fe on rd. past Escuela Agricola Alto Piedra on Pacific side of Divide, 800-1200 m, 18 Mar 1973, Croat 23107 (F, K, MO, NY, US); on Caribbean slope above Rio Primero Brazo, 5 mi NW of Santa Fe, 700-1200 m, 18-19 Mar 1973, Croat 23198 (MO); SW side of Cerro Tute, La Cuchilla, ca. 2500 m, 10 Sep 1982, D'Arcy 15012 (MO); CerroTute, 8 Aug 1963, Dwyer 4313 (MO, US); base of Cerro Tute, 6.5 km outside of Santa Fe, 6 May 1977, Folsom 3010 (MO); NW of Santa Fe, 1 km from Escuela Agricola Alto de Piedra, 24 Feb 1975, Mori & Kallunki 4813 (MO, MPU). CUBA. Cumbre, E of Matanzas, 29 Aug 1903, Britton & Wilson 120 (NY); vic. of Herradura,Pinar del Rio, 28-31 Aug 1910, Britton & Earle 6559 (NY, US); Isle of Pines, San Juan, 15-17 Mar 1916, Britton et al.

FLORA NEOTROPICA 15466 (F, NY, US); Gran Piedra, Sierra Maestra, Mar 1949, Clemente 6483 (US); Oriente, Baracoa, Rio Oro, ca. 250 m, 27 Mar 1915, Ekman 5085 (US); Habana, Laguna Ariguanabo, 16 Apr 1922, Ekman 13722 (F, NY); Las Nuifas, Oriente, 28 Dec 1917, Hioram & Bliste 1498 (NY); Isle of Pines, near Caleta Grande, 22 May 1910, Jennings 602 (NY); Havana, Anafe, not far from Ariguanabo, 23 Apr 1914, Le6n & Ekman 4274 (NY), 2 Jul 1914, Le6n 4336 (NY, US); thickets E of Playa de Marianas, Havana, 11 Nov 1915, Le6n 5729 (NY): Loma del Gato and vic., Cobre range of Sierra Maestra, 9001000 m, 11 Jul 14 Aug 1921, Le6n et al. 10013 (NY); Pico Turquino, Sierra Maestra, Jul 1922, Le6n 10959, 11017 (NY); southern Baracoa, Tinias, 17 Jul-4 Aug 1924, Le6n 12272 (NY); Cordillera de la Gran Piedra, Sierra Maestra, 28 Jan 1953, L6pez F 830 (US), 18 Mar 1956, L6pez F 2630 (US); Oriente, rd. from Aserradero San Antonio de los Cumbres to La Bayamesa, crest of Sierra Maestra, 1500-1800 m, 22 Jan 1956, Morton 9313 (US); Oriente, along Rio Peladero, below Aserradero San Antonio de los Cumbres, crest of Sierra Maestra, ca. 1300 m, 23 Jan 1956, Morton 9506 (US); Peninsula de Zapata, 9 Jul 1920, Roig & Cremata 2095 (F); Los Cayuelas, Pinar del Rio, 12 Apr 1924, Roig 3234 (NY); Yumuri River, Jun 1849, Rugel 375 (BM. NY); Maisi to Sabana, Oriente, 14 Dec 1910, Schafer 7944 (NY, US); La Perla, Oriente, 600-660 m, 6-18 Feb 1911, Schafer 8578 (NY, U, US); Firmeza to Gran Piedra, Oriente, 4-5 Mar 1911, Schafer 8998 (NY, US); Sevilla Estate, near Santiago, cafon Rio Ubero, 9 Sep 1906, Taylor 309 (NY); hills near Santiago de las Vegas, 11 Aug 1901, Wilson 1091 (POM). HAITI. Furzy, 1500 m, Jan 1917, Bush 1370 (US); Marne de la Hatte, ca. 800 m, 10 Jun 1917, Ekman 129 (LL); Massif de la Selle, Petionville, Kenskoff, ca. 1250 m, 18 Aug 1924, Ekman 1924 (US); Massif des Matheux, Grands-Bois, ravine Nan-Figues, 1200 m, 15 Mar 1926, Ekman 5717 (US); Massif de la Selle, Marne des Commissaires, Anses a Pitres, Tete de l'Eau, at Rio Pedemales, 26 Aug 1926, Ekman 6756 (F, LL, MO, US); Riviere Glace, 500 m, 5 Aug 1945, Holdridge 2121 (US); vic. of Furcy, 1300 m, 26 May-15 Jun 1920, Leonard 4315 (NY, US); vic. of Marmelade, ca. 800 m, 19 Dec 1925, Leonard 8189 (US); vic. of Dondon, ca. 400 m, Leonard 8587 (US); vic. of Ennery, L'Artibonite, 325-900 m, 20 Jan 1926, Leonard 9040 (F, MO, US); vic. of Plaisance, ca. 400 m, 26 Jan 1926, Leonard 9195 (NY, US); vic. of Port de Paix, 22 Jan 1929, Leonard & Leonard 12257 (US); vic. of St. Louis de Nord, SW of city, 30 Mar-7 Apr 1929, Leonard & Leonard 14448 (NY, US); Port Margot to Correil, 1200-1500 ft,7 Aug 1903, Nash 194 (NY); Camp No. 2, Mt. Maluevre, 2650 ft, 22-23 Jul 1905, Nash & Taylor 1157 (NY); Massif de la Hotte, 13.6 km N of Camp Perrin on rd. from Roseaux to Jeremie, 720 m, 18?23'N, 73?53'W, 15 Nov 1982, Zanoni et al. 24307 (NY). DOMINICAN REPUBLIC. Cordillera Central, Constanza, ca. 1200 m, 9 Nov 1929, Ekman 14061 (NY, US); Barahona,Nizuito, above Paraiso, Jul 1910, Fuertes 433 (F, MO, NY, U, US); La Cienega, S bank of Rio Los

TAXONOMIC TREATMENT Guanos, just above confluence with Rio de la Izquierda, 1100-1200 m, 14 Jul 1967, Gastony et al. 175 (NY, US); El Pico Diego de Ocampo, 1220 m, 31 Aug 1947, Jimenez 4445 (US); Guazumal, Santiago, 10 Sep 1950, Jimenez 2099 (US); La Lomita, ca. 760 m, 7 Apr 1950, Jimenez et al. 2925 (US); Guabito de Yaroa, Yaroa valley, Puerto Plata, 350-500 m, 1 May 1968, Liogier 11016 (NY, US); along Arroyo de la Sal, Loma de la Sal, Jarabacoa, 1100 m, 17-18 Jun 1968, Liogier 11727 (F, NY, US); Ci6naga de Manabao, Los Guanos, Jarabacoa, 100-1300 m, 13 Aug 1968, Liogier 12039 (NY, US); along Rio Sonador, Juan Santiago, near Hndo. Valle, 800-1100 m, 9 Sep 1968, Liogier 12614 (F, NY, US); Casabito, Bonao, 700 m, 21 Jan 1974, Liogier & Liogier 21116 (NY); Loma Isabel de Torres, Puerto Plata, 17 May 1975, Liogier & Liogier 22907 (NY); Los Arroyos, Pedernales, 1600 m, 24-27 Jun 1975, Liogier & Liogier 23259 (NY); E of San Jose de Ocoa, El Manaclar, 1150 m, 18?31'N, 70?27'W, 18 Jun 1978, Mejia 270 (NY); Seibo, rd. from Higuey to Gato, 6 Dec 1909, Taylor 444 (NY, US); near Constanza, 1190 m, Aug 1910, von Tuerckheim 3487 (F, MO, NY, U, US); Lagunas de Cenobi, Sabaneta,vic. of Monte Cristi,21 Aug 1929, Valeur 74 (US); Arroyo Vallecito, 600 m, 29 Dec 1929, Valeur 327 (F, MO, NY, US); s.loc., Jan-Mar 1871, Wright et al. 513 (CU, US); Sierrade Baoruco, 38 km S of Duverge, 5 km S of Aguacate, 1550-1600 m, 18?18'N, 71?42.5'W, 18 Aug 1983, Zanoni & Pimentel 26544 (NY); Sierra de Baoruco, 37.5 km N of Pedernales, 1.5 km N of Puesto Militar Los Arroyos, on Aguacate and Duverge rd., 1240 m, 18?15'N, 71045'W, 19 Aug 1983, Zanoni & Pimentel 26637 (NY); Cordillera Septentrional, on Loma El Mulazo, 1083 m, 19?41'N, 70?58'W, 18 Dec 1984, Zanoni et al. 32786 (NY). PUERTO RICO. Utuado, forest of Rio Abajo, close to old Camp Radley, ca. 800 m, 14 Sep 1987, Acevedo & Chinea 2190 (NY); Luquillo Forest, rt. 191 at km 9.3. 1600 ft, 29 Aug 1967, D'Arey & Woodbur, 1710 (MO); Las Abritas, Rio Abajo, 23 Feb 1940, Holdridge 107 (NY); Pico Guilarte, Adjuntas, 1000 m, 16 Jul 1963, Liogier 9996 (F, NY, WIS); slopes of El Yunque, Luquillo Mtns., 800 m, 13 Sep 1979, Liogier et al. 29312 (NY); Abajo, Utuado, 350 m, 4 Dec 1980, Liogier & Liogier 31276 (NY); Rio Abajo, 300-400 m, 29 Jul 1950, Little 13525 (NY); Caribbean National Forest, Cinchona Experimental Areas, 3500 ft, 17 Aug 1950, Little 13692 (NY); Luquillo Mtns., upper Luquillo forest, 2000-2500 ft, 10 Jul 1966, Little 21615 (NY, US); km 9.3 on rt. 191, E side Luquillo Mtns., 24 Sep 1967, Wagner 1229 (MO, U, WIS), 28 Aug 1968, Wagner1607 (MO, U, WIS); near Jacaya, km 1.6 rd. 144, 18 Jun 1958, Woodburys.n. (NY). VIRGIN ISLANDS. ST. VINCENT: Mt. Saint Andrews, 1800 ft, Dec 1889, Eggers 6742 (US); Grand Bonhomme, 800-930 m, 24 May 1947, Morton 6080 (US); s.loc., Sep 1889, Smith & Smith 495 (NY). GRENADA. ST. ANDREWS:Grand Etang Forest Reserve, ca. 500 m, 8-9 Aug 1959, Websteret al. 9575 (U). COLOMBIA. S.loc., Mutis 3578 (US); s.loc., Triana 2255 (BM, US). ANTIOQUIA: Hacienda Montenegro near La Pintada, ca. 600 m, 20 Oct 1947, Blackman et al.

I09 170287 (US); Chigorod6, Gutierrez V, Facultad 2 (US); Brazuela de Perales, Rio Magdalena, ca. 150 m, 9-10 Jan 1918, Pennell 3695 (NY, US); Fredonia, 25 Jun 1940, Tomds 905 (US). BOYACA: El Umbo region, NW of Bogota, 3000 ft, 29 Sep 1932, Lawrance 485 (NY). Caldas: Pereira, along Rio Otun, 1370-1400 m, 29-30 Aug 1922, Killip & Hazen 11004 (NY); San Jos6, 14001800 m, 3 Sep 1922, Pennell 10252 (NY). CAUCA: Near San Francisco, 1100 m, 23 Sep 1968, Espinal T. & Ramos 2769 (F); La Paila, 12 Feb 1853, Holton 12 (K). CHOCO: Rio Agara, rd. from San Jose del Palmar Norte to N6vita, between La Itlaia (Pueblo Nuevo) and Curund6, 430-450 m, 26 Aug 1976, Forero et al. 2109 (MO, NY); Cerro Torr6, along Rio Surama, 500 m, 21 Feb 1977, Forero et al. 3063 (MO); Rio San Juan, below Noanama, ca. 4?40'N, 76?55'W, 7 Apr 1979, Forero et al. 4850 (MO, NY); Rio Tamana,tributaryof Rio San Juan, between Primavera and Santa Rosa, ca. 5000'N, 76?44'W, 10 Apr 1979, Forero et al. 4934 (MO); Rio Tamana, tributary of Rio San Juan, below Santa Rosa, ca. 5?00'N, 76?42'W, Forero et al. 4960 (MO); PanAmerican Hwy. (in construction), Rio Pat6, ca. 5?35'N, 76?56'W, 21 Apr 1979, Forero et al. 5438 (MO, NY); Chaparraid6, near Tutunendo, ca. 100 m, 26 Apr 1979, Forero et al. 5821 (MO); rd. Tutunendo-ElCarmen, near camp "El 12," upper Rio Atrato, 600 m, 27 Apr 1979, Forero et al. 5878 (MO); Boca Tanando,Rio Atrato, above Quibd6, 23 Mar 1958, Fosberg 39318 (WIS); near Istmina, rd. to Certegui, 75 m, 3 Aug 1944, GarciaBarriga 11209 (US); rd. from Quibd6 to Yuto, 10 km from Quibd6, ca. 350 m, Gutierrez V 2895 (US); vic. of Bahia Solano, 1973, Hill M7 (MO); Rio Tolo, region of Guayabal, SE of Acandi, 28 Mar 197?, Ordofiez & Valencia 040 (MO); Cerro del Torra, E slopes, base of mountain in Rio Negro drainage, 1630 m, 9 Aug 1988, Ramos et al. 1085 (NY); along bank of Rio Virudo, 2-3 km above Virudo, ca. 10 m, 3 Aug 1973, White& Warnrer 98 (MO). CUNDINAMARCA:La Esperanza, 12 Apr 1935, Garcia Barriga 3117 (US); Caparrapi, ca. 1275 m, 813 Jun 1939, Garcia Barriga 7670 (US); old rd. from Guadas to Villeta, between Guadas and Alto de Trigo, 1100-1800 m, 10 Nov 1945, Garcia Barriga 11803 (US); Jerusalem, Jul 1930, Perez-Arbeldez 575 (US); Faraima, Dec 1932, Perez-ArbelIez 2101 (US). HUI A: Rio Mosquera, Mosquera, Caqueta and vic., 6800 ft, 25 Apr 1944, Little 7718 (NY, US); Quebrada de Angeles, above Natagaima, 450-600 m, 24 Jul 1917, Rusbhy& Pennell 279 (MO, NY, US). MAGDALENA: Sierra Nevada de Santa Marta, near Campano, ca. 1300 m, 11 Jan 1948, Barkley & Gutierrez V 1916 (NY, US); between La Gran Via and San Pedro de la Sierra, 1350 m, 10 May 1983, Escobar & Santa 3490 (NY); Sierra Nevada de Santa Marta, Quebrada La Sirena, ca. 1100 m, 10?59'N, 73?59'W, 8 Sep 1972, Kirkbride 2175 (NY); Finca La Granja, base of Cerro Quemado and Cerro San Lorenzo, 200-2300 m, 23 Apr 1959, Romero-Castaneda 7876 (NY); "Cincinnati",lower slopes of Mount San Lorenzo, near Santa Marta, 1300 m, 1932, Seifriz 42 (US); near Cacagualito, 1500 ft, 1898-99, H. H. Smith 1857 (NY). NARINO: Curcuela, just above San Miguel, 8 kin below

110

FLORA NEOTROPICA

& Mallet 6855 (BH, K, MO, MY, US, VEN); Colonia Tovar, Moritz 395, 555 (K); ca. 2 km NW of Estaci6n Biol6gica Rancho Grande on rd. from Maracay to Ocumare de la Costa, 1120 m, 27 Jul 1979, Nee & Whalen 16894 (BH, F, VEN, WIS); Ocumare valley, 800 Oct 1929, Toro 73 (NY). NORTE DE SANTANDER: Samaria, Toledo, 200-2100 m, 29 Oct 1941, Cuatrecasas m, 3 Apr 1926, Pittier 12167 (MO, VEN, US); E of house at Rancho Grande, 1100 m, Jul 1945, Pittier 15196 (US, et al. 12764 (F, U, US); vic. of Pamplona, 2300-2400 m, 27 Feb 1927, Killip & Smith 19779 (NY, US); vic. of VEN); rd. between Maracay and Ocumare de la Costa, Williams 10271 (F, US, VEN). BARINAS:Ca. 8 km N of Toledo, 3-11 Mar 1927, Killip & Smith 20068, 20070 Bumbum, along Rio Bumbum, ca. 300 m, 8?16'N, (NY, US); Culaga valley, near Tapata,N of Toledo, 15002000 m, 3-8 Mar 1927, Killip & Smith 20156 (NY, US). 70?48'W, 25 Oct 1984, Knapp & Mallet 6840 (BH, K, PUTUMAYO: Santa Rosa del Rio Guamues, ca. 300 m, 2 MY, VEN); along Rio Caparo, between Campamento Chacimano and Boca de Graza, E of El Cant6n, 100 m, Dec 1968, Plowman 2099 (F, K, US); Rio Caqueta 12 Apr 1968, Steyermarket al. 102193 (NY, U, US, VEN). downriver from Puerto Lim6n at Yocopena, 300-350 m, 17 Dec 1968, Plowman 2160 (K, US); Valley of Sibundoy, DISTRITOFEDERAL:Colonia Tovar, 1800-2000 m, Dec 2225-2300 m, 29 May 1946, Schultes & Villareal 7679 1924, Allart 447 (NY, US, VEN); between Rosita and (K). RISARALDA:ParqueRegionalUcumari,ca. 22 km ESE Portachuelo, ca. 1700 m, 3 Jun 1972, Benitez de Rojas of Pereira, trail to Ucumari, 2100-2670m, 4?40'N, 1440 (MY, U); near Guarenas, 3500 ft, Dec 1854, Birschel s.n. (K); ca. 12 km E of Colonia Tovar on rd. to 75?35'W, 25 May 1989, Luteyn & Rangel 13105 (BM, NY); Hacienda Alejandria, km La Virginia-Cerrito rd., El Junquito, ca. 2000 m, 10?26'N, 67?08'W, 29 Oct extreme N part of valley of Rio Cauca, ca. 940 m, 7 May 1984, Knapp & Mallet 6866, 6868 (BH, K, MY, US, 1989, Silverstone-Sopkin et al. 5185 (NY). SANTANDER: VEN); La Guaira-Caracas, 7 May 1874, Kuntze 1238 Puerto Wilches and vic., 100 m, 28 Nov-2 Dec 1926, (NY, US); Colonia Tovar and vic., 1700-2300 m, 1921, Killip & Smith 14770 (K, NY, US). VALLEDE CAUCA: Pittier 9350 (NY, US, VEN); near Agua Fria, km 26 of Along carretera al Mar, at Rio Dagua, 15 km from Caracas-Cua rd., 17 Apr 1924, Pittier 11502 (MO, NY, Buenaventura, 6 Nov 1944, Core 1529 (NY, US); beUS, VEN); El Junquito, 2000 m, 29 Dec 1936, Pittier tween Gorgona and Puerto Cabuyo, 1000 m, 4 Jun 1943, 13821 (F, US, VEN); Fila de Agua Negra, Feb 1938, Cuatrecasas 14526 (F, US); Cordillera Occidental, W Tamayo440 (US, VEN); Agua Negra, 8 Mar 1940, Tamavo 1227 (F, US, VEN); Agua Negra, 1200 m, 2 Jan 1940, slope, Rio Sanquinini, left bank, La Laguna, 1250-1400 m, 10-20 Dec 1943, Cuatrecasas15576 (F, US); Rio Calima Williams12466 (F, K, US, VEN). LARA:Terepaima,S side (region of Choc6), La Trojita, 5-50 m, 19 Feb-10 Mar of peak, 20 km S of Cabudare, 1200-1300 m, 3 Aug 1944, Cuatrecasas 16343 (F, US); Cabuyal, ca. 1000 1970, Steyermark et al. 103330 (US, VEN); along rd. m, 20 Jun 1945, Cuatrecasas 19638 (F, US); between leading to Yacambu, 10 km SSE of Sanare, 1640 m, 8 Zaragoza and La Victoria, 950 m, 18 Nov 1946, Cuatre- Aug 1970, Steyermarket al. 103569 (US, VEN); Montafia casas 23002 (F, US); near Puente G. Valencia over Rio Obscura, La Briza, along rd. leading to Yacambi, 10 Cauca, 3 Sep 1968, Espinal T & Ramos 2584, 2597 (F); km NW of El Blanquito, 10 km SSE of Sanare, 1630 m, La Paila, 12 Feb 1853, Holton 1566 (NY); Santa Rosa 8 Aug 1970, Steyermark et al. 103579 (US, VEN); 3-6 to Cisneros, 250-350 m, 10 May 1922, Killip 5348 (NY, km S of Agua Negra, 17-20 km E of Cubiro, 1600US); Guayabal, Zarzal, Cauca valley, 900-920 m, 20 Jul 1700 m, 9?47'N, 69?30'W, 5 Jul 1974, Steyermark et al. 110097 (F, VEN). MERIDA: La Mucuy, 2200 m, 10 Apr 1922, Killip 8345 (NY); Cuchilla, E of Zarzal, 1200-1600 m, 22 Jul 1922, Pennell et al. 8525 (NY); Rio Agua Bonita 1953, Bernardi 411 (K, NY, VEN); Hato Perez, Febto Rio Vieja, E of Zarzal, 1300-1600 m, 23 Jul 1922, Mar 1957, Bernardi s.n. (NY); Tabay, 2200-2300 m, 2 Pennell et al. 8563 (NY, US); Quebradala Canas,Vallejuelos Sep 1930, Gehriger 408 (F, MO, NY, US, VEN); Parque rd. off Cali-Armenia rd., 1100 m, 28 Apr 1987, Restrepo Nacional Sierra Nevada, La Mucuy, ca. 7 km S of Tabay, & Heredia 309 (NY); Rio Calima, Quebrada de la Brea, 2000-2100 m, 8?40'N, 71?08'W, 21 Oct 1984, Knapp 30-40 m, 19 May 1946, Schultes & Villareal 7346 (F, & Mallet 6791, 6793, 6795, 6796, 6797 (BH, K, MY, K, US); Hacienda El Medio, PanAmerican hwy. between US, VEN); 13.4 km S of La Azulita on rd. to Jaji, ca. La Paila and Zarzal, flat part of valley of Rio Cauca, ca. 1850 m, 8?40'N, 71?25'W, 22 Oct 1984, Knapp & Mal975 m, 15 Nov 1986, Silverstone-Sopkinet al. 2521 (MO, let 6805 (BH, K, MY, US, VEN); 27.2 km S of La Azulita, 14.6 km N of Jaji turnoff on La Azulita rd., ca. 2200 m, NY); Jamundi, 1000 m, Jan 1945, von Sneidern 4555 (F); s.loc., 1000 m, 1851-7, Triana s.n. (NY). 8?32'N, 71?16'W, 22 Oct 1984, Knapp & Mallet 6812 VENEZUELA. ANZOATEGUI: Along Rio Cangrejo, (BH, K, MY, US, VEN); 8 km (16 km by rd.) S of La tributary to Rio Zumbador, NE of Bergantin, 500 m, 24 Azulita on rd. to Merida, ca. 1500 m, 3 May 1971, Nee Feb 1945, Steyermark 61171 (F, MY, VEN). ARAGUA: & Mori 4136 (MO, US, VEN, WIS); La Carbonera, Summit of Rancho Grande to the N, ca. 1200 m, 1 May Bosque San Eusebio, 2250-2600 m, 1971, Veillon 23 1971, Benitez de Rojas 928 (MY, U); Parque Nacional (NY, VEN). MIRANDA:Parque Nacional Guatopo, along Henri Pittier, Paraiso trail to Pico Periquito, 1200-1300 hwy. 12, 3 km S of jct. with hwy. 1, 25 Apr 1971, Nee & Mori 4057 (VEN, WIS). PORTUGUESA: La Aparici6n m, 8 Feb 1973, Croat 21396 (MO); ca. 2 km NW of Estaci6n Biol. Rancho Grande on rd. to Ocumare de la de Ospino-Moroturo rd., 200-900 m, 9?31'N, 69?26'W, Costa, 1100 m, 10?21'N, 67042'W, 28 Oct 1984, Knapp 1 Mar 1988, Aymard & Cuello 6580 (NY); "Fundo El Piedrancha, valley of Rio Guabo, 1500 m, 1?11'N, 77?56'W, 1 Oct 1943, Fosberg 21086 (NY, US); GorgonillaIsland, 130-200 m, 8 Feb 1939, Killip & Garcia 33077 (US), Killip & Garcia 33081 (F, US); Tumaco, 5

TAXONOMIC TREATMENT Chaparral," 14 km NE of Guanare, SE of right bank Rio Portuguesa, 180 m, ca. 9?05'N, 69?35'W, 12 Mar 1988, Aymard& Cuello 6593 (NY). SUCRE:Rd. to E from Caruipanoto Maturincito,ca. 100 m, 29 Jan 1981, Benitez de Rojas 2924 (F, K, MY); Peninsula de Paria, Cumbre de la Estrella, W of Manacal, N of El Paujil, 800-850 m, 10?40'N, 62041'W, 17 Oct 1984, Knapp & Mallet 6772, 6773 (BH, K, MY, US, VEN). TACHIRA: Headwaters of Rio Quinimari along Quebrada Agua Negra on trail to Paramo de Judio, 5 km S of San Vicente de la Revancha, 15 km S of Providencia, SE of Santa Ana, 2100-2400 m, 7?25'N, 72?25'W, 23 Oct 1984, Knapp & Mallet 6828 (BH, K, MY, US, VEN); between Michelina and Boca de Monte, W of Zumbador, 2002400 m, 28 Aug 1966, Steyermark & Rabe 96800 (NY, U, VEN); below Paramo de Tama, near the Colombian frontier, above Betania and Tami, near Quebrada Buena Vista, 2300-2450 m, 22-24 May 1967, Steyermark et al. 98714 (US, VEN). YARACUY:8.5-9 km N of Salom, 1200-1300 m, 10?15'N, 68?29'W, 4 Mar 1982, Liesner & Steyermark12342 (MO, NY, U); El Amparo to Candelaria, 7-10 km N of Sanare, 1100-1300 m, 27-30 Dec 1972, Steyermark& CarrenoEspinoza 106797 (MO, US, VEN). ZULIA: Along Rio Yasa, near "Guasama,"above "Kasmera" (Estaci6n Biol6gica de la Univ. de Zulia), SW of Machiques, 500-600 m, 26-27 Aug 1967, Steyermark & Fernandez 99794 (MO, NY, US, VEN). ECUADOR. CARCHI: Environs of Maldonado, 1450-1650 m, 1 Jun 1978, Madison et al. 4895 (F). GUAYAS: Guayaquil, s.coll. (W). ESMERALDAS: Rio Zapallo Grande (tributary of Rio Cayapa), trail to hills 700 m from evangelical station, 200 m, 0?48'N, 78?52'W, 28 Oct 1982, Barfod 41078 (AAU, BM); along Rio Camerones, small tributaryto Rio Cayapa, 50 m, 0?55'N, 78055'W, 8 Oct 1983, Barfod et al. 48042 (AAU, BM); Rio Bolborde, small tributary to Rio San Miguel in upper Cayapa drainage, 300 m, 0?40'N, 78?53'W, 17 Oct 1983, Kvist et al. 48271 (AAU, BM). Los Rios: Rio Palenque Biological Station, km 56 Quevedo-Santo Domingo, 150-220 m, 19 Sep 1972, Dodson 5177 (F, 2-6 km S of Arapicos, 800MO). MORONA-SANTIAGO: 900 m, 6 Apr 1981, Lugo S. 6005 (AAU, NY); along Rio Metzera Grande on Hacienda Sangay (plantation of Compania Ecuatorianadel T6, S.A.) near Palora, ca. 950 m, 1?40'S, 77?58'W, 15 Feb 1984, Knapp & Mallet 6295 (BH, K, QCA, QCNE, US). NAPO: Rd. Baeza-Lago Agrio, 23 km from Baeza at Rio Oyacachi, 1500 m, 0?18'S, 77048'W, 31 Oct 1976, Balslev & Madsen 10507 (F, NY); rd. Baeza-Lago Agrio, 18 km from Baeza, Puente Sardinas Grandes, 1650-1700 m, 0?22'S, 77?49'W, 2 Nov 1976, Balslev & Madsen 10559 (F, MO, NY); Lago Agrio, Dureno, Cofan indigenous community, near Rio Dureno, 350 m, 0?02'S, 76?42'W, 11 Mar-i Apr 1986, Cer6n 260 (MO, NY); Rio Latas, ca. 8 km W of Puerto Misahualli, tributary of Rio Napo, ca. 500 m, 1?02'S, 77?45'W, 23 Jan 1984, Knapp & Mallet 6188, 6190 (BH, K, QCA, QCNE, US); Rio Pano, ca. 10 km W of Tena on rd. to Rio Jatunyacu, new rd. to Salcedo-Napo, ca. 700 m, 1?00'S, 78?55'W, 24 Jan 1984, Knapp & Mallet 6198, 6200 (BH, QCA, QCNE, US); Santa Cecilia,

111 Mufiozlandia, ca. 340 m, 2 Apr 1972, MacBryde & Dwyer 1456 (US); 1 km SW of Baeza, 2000 m, 0?28'S, 77?53'W, 20 Oct 1976, Ollgaard & Balslev 10202 (F. NY); INIAP-PAYAMINOstation, Reserva Floristica "El Chuncho," 250 m, 0?27'S, 77?01'W, 17 Jun 1987, Palacios 1637 (MO, NY); 8 km N of Coca (Puerto Francisco de Orellana), 250 m, 0?24'S, 77?00'W, 8 Apr 1985, Zaruma et al. 61 (BM, MO); Payamino-Loreto rd., 4-6 km from river, 250 m, 0?26'S, 77?02'W, 13 Sep 1986, Zaruma 725 (K, MO, NY). PASTAZA:Hacienda San Antonio del Bar6n von Humboldt, 2 km NE of Mera, 1100 m, 1?27'S, 78?06'W, 5-19 Mar 1985, Baker et al. 5512 (MO, NY); Rio Pastaza at Shell-Palora rd., 1000 m, 1?35'S, 78?03'W, 3 Dec 1984, Jorgensen & Laegaard 56488 (BM, MO, QCA, QCNE); 21 km W of Puyo (just W of Mera) on rd. to Baiios, 100-1100 m, 1?25'S, 78?05'W, 18 Feb 1984, Knapp & Mallet 6307 (BH, K, NY, QCA, QCNE, US); 5 km NE of Mera, rd. to Rio Ansu, 1200 m, 1?26'S, 78006'W, 3 Mar 1985, Neill et al. 5930 (BM, MO, QCNE); vic. of Puyo, 750-1000 m, Sep 1939, Skutch 4530 (K, NY). PICHINCHA:Along rd. between Marianitas and Nanegal, 1200-1250 m, 0?08'N, 78?39'W, 15 Jul 1990, Webster & Rios 28309 (DAV, IBE); Quito-Lloa-Mindo rd., Hacienda El Pedregal, 1650-1850 m, 0?03'S, 78?40'W, 10 Jul 1987, Zak & Jaramillo 2154 (MO, NY). PERU. Cuzco: Quispicanchis, Quincemil, ridges N of the village, 500-600 m, 13?15'S, 70?45'W, 23 Apr 1984, Knapp & Mallet 6385 (BH, CUZ, NY, USM). HUANUCO: Gauso Azul, Rio Pachitea, Oct 1942, Sandeman 3447 (K); Pachitea, Bosque Nacional de Iparia, along Rio Pachitea ca. 20 km from confluence with Rio Ucayali, rd. to Ayamiria, 6 km from Miel de Abeja, 300-400 m, 27 Dec 1966, Schunke V 1428 (F, NY, US); Bosque Nacional de Iparia, along Rio Pachitea, 1 km above Tournevista, ca. 20 km above confluence with Rio Ucayali, near Miel de Abeja, 300-400 m, 15 Feb 1967, Schunke V 1609 (F, US); Bosque Nacional de Iparia, ca. 20 km upriver from Rio Ucayali on Rio Pachitea, Miel de Abeja, 300-400 m, 30 May 1967, Schunke V 2015 (F, MO). LORETO:Maynas, Quistacocha (camino psi granja), 100 m, 29 Sep 1979, Ayala 2009 (AMAZ, MO); near mouth of Rio Napo, 14 Sep 1972, Croat 20152, 20205 (F, MO, NY); Rio Itaya, 20 min (by 40 hp motor) above Iquitos, 120 m, 18 Mar 1977, Gentry et al. 18388 (F, MO, USM); near Brilla Nueva, Borro Indian village on upper Rio Yaguasyacu, tributary of Rio Ampiyacu, 8 Nov 1977, Gentry & Revilla 20441 (AMAZ, MO); trail between Indiana (on Rio Amazonas) and Mazan (on Rio Napo), 130 m, 4 Mar 1979, Gentry et al. 25406 (MO, USM); Quebrada Yanamono, Rio Amazonas above mouth of Rio Napo, 120 m, 14 Nov 1979, Gentry et al. 27993 (MO); Yanamono, Explorama tourist camp on Rio Amazonas between Indiana and mouth of Rio Napo, 120 m, 3?28'S, 72?48'W, 26 Jul 1980, Gentry et al. 29011 (AMAZ, MO. NY); Caserio Alianza, Rio Tamshiyacu, trail toward Rio Manati, 140 m, 4?05'S, 72?58'W, 1 Aug 1980, Gentry et al. 29314 (AMAZ, MO); Yanamono, Explorama tourist camp between Indiana and mouth of Rio Napo, 130 m,

FLORA NEOTROPICA

112 3?28'S, 72?48'W, 18 Feb 1981, Gentry et al. 31347 (AMAZ, MO); Masisea, ca. 275 m, 125 Jul 1929, Killip & Smith 26863 (NY, US); Rio Itaya, Pena Negra, 100 m, 5 Mar 1973, McDaniel & Rimachi Y 16840 (F, IBE, K, NY); Yanamono, Explorama tourist camp on Rio Amazonas between Indiana and mouth of Rio Napo, ca. 80 km NE of Iquitos, ca. 100 m, 3?28'S, 72?48'W, 22 Jul 1984, Knapp 6598 (BH, US, USM); 39 km SW of Sarameriza, ca. 2 km W of Petroperu tower on Loreto/ Amazonas border, 57 km NE of Rio Nieva bridge, ca. 900 m, 4?40'S, 77?36'W, 9 Jun 1986, Knapp & Alcorn 7707 (MO, USM); Rio Nanay, near Shiriara,21 Feb 1969, Plowman 2563 (US, USM); Momoncillo on left margin of Rio Momon, 10 km from mouth of Rio Nanay, 8 Mar 1976, Revilla 300 (AMAZ, F, MO, USM); Rio Momon, near Momoncillo, 16 Nov 1976, Revilla 1834 (AMAZ, USM); Rio Itaya, Moena Cano, Naranjal near confluence with Amazon, 26 Nov 1976, Revilla 1937 (USM); Rio Nanay above Bellavista, Quebrada de Moropan, trail to Caserio de Tres Unidos, 1 Mar 1977, Rimachi Y 2864 (F, MO, NY, USM); Rio Momon, tributaryof Rio Nanay, trail from caserio de Gran, 7 Mar 1978, Rimachi Y 3398 (AMAZ); 3 km from Santa Maria de Nanay on trail to Santa Rosa, 130 m, 8 Mar 1968, Schunke V 2501 (F, NY); Puerto Almendras, Rio Nanay, 122 m, 3?48'S, 73021'W, 3 Nov 1985, Vdsquez & Jaramillo 6874 (K, MO, NY); Amazon river near Iquitos, 120 m, Apr 1930, Williams 8242 (F). MADREDE DIos: Parque Nacional Manu, Rio Manu, Cocha Cashu station, 350 m, Jul 1978, Foster & Terborgh 6497 (F); Explorer's Inn at confluence of Rio Tambopata and Rio Torre, Laguna Chica trail, 170 m, 12?50'S. 69020'W, 15 Jul 1987, Smith et al. 1048 (U, US). SAN MARTiN:Zepelacio near Moyobamba, 11001200 m, Oct-Nov 1933, Klug 3371 (F, K, MO); rd. from Cufiumbuque to Sisas, ca. 1 hr driving from Cufiumbuque (1/3 way to Sisas), ca. 850 m, 6?35'S, 76?39'W, 5 Jun 1984, Knapp et al. 6480 (BH, US, USM); km 2730 Tarapoto-Yurimaguas rd., 650-750 m, 6?25'S, 76?15'W, 9 Jun 1984, Knapp & Mallet 6489, 6492 (BH, US, USM); km 36-41 Tarapoto-Yurimaguas rd., 400600 m, 6?25'S, 76?15'W, 11 Jun 1984, Knapp & Mallet 6498 (BH, US, USM); Convento, trail to Tioyacu and Nuevo Lamas, km 68 Tarapoto-Yurimaguasrd., ca. 270 m, 6?16'S, 76?17'W, 16 Jun 1984, Knapp & Mallet 6517, 6521 (BH, US, USM); km 47 Tarapoto-Yurimaguasrd., ca. 350 m, 6?20'S, 76?18'W, 21 Jun 1984, Knapp & Mallet 6530 (BH, US, USM); km 54 of TarapotoYurimaguas rd., along stream, ca. 350 m, 3 Sep 1986, Knapp 8268 (MO, USM); km 12-16 TarapotoYurimaguas rd., 2250 ft, 23 Aug 1978, Rimachi Y 3844 (AMAZ, IBE, NY, TEX); Fundo Las Flores E of San Juan de Pacayzapa, 800-900 m, 12 May 1973, Schunke V 6245 (NY); San Juan de Pacaizapa, km 72 TarapotoMoyobamba rd., 100-1050 m, 12 Jun 1977, Schunke V 9696 (F, MO, USM); Tarapoto, 750 m, Dec 1929, Williams 5450 (F); Lamas, 840 m, Dec 1929, Williams 6352 (F). UCAYALI:Contamana, trail to Aguas Calientes, 27 Jul 1970, McDaniel et al. 2565 (US); Contamana, 210 m, 10 Oct 1965, Schunke V 918 (F, US); Contamana, 210 m, 15 Dec 1964, Schunke V 6653 (F, US).

BRAZIL.ACRE:Vic. of Porangaba, Rio Jurua-Mirim, 17 May 1971,Maas et al. P13080 (K, NY, P, US, WIS); Cruzierodo Sul, Rio Juruaand Rio Moa, vie. of Serra da Moa, 24 Apr 1971, Prance et al. 12424 (F, K, NY, US, WIS);Rio Acre, SeringalAuristella,Jun 1911,Ule 9758 (K). AMAZONAS:Km 157 Manaus-Itacoatiara rd.,

16 Dec 1974, Gentry& Ramos 13359 (BH, MO);near mouthof Rio Embira(tributaryof Rio Tarahuaca),basin of Rio Jurua,7?30'S,70?15'W,27 Jun 1933,Kruko/f 5043 (F, K, MO, NY, U, US).

Local names. Mexico. Veracruz:huele de noche, sanptipuskat(Totonac).Belize. yerbade barra,hoja de piojillo, diaper wash, lava pafial. Guatemala. Alta Verapaz:sakyol,sac-yol;Guatemala:huis,hojade tinta; Peten:wich tz'on chet.Nicaragua.Zelaya:huelenoche. Panama.Area de Canal:sauco, hoja hedionda. Cuba. ajicillo, tabaco cimarr6n.Dominican Republic. arito, mantequito.Colombia.Antioquia:sauco,zapata;Choc6: sauco, sanco, sauco amargo; Valle de Cauca: lulo, hedionda.Venezuela.Anzoategui:tabacero.Ecuador. Napo:jonchimbachoquinicco(Cofan);zapata,zapatachi (Cayapa); ataqui panga, asna panga, asna panga cushnichinapangayura(Quichua).Peru.Huanuco:ayac mullaca; Loreto: asno panga, sacha conjompe; San Martin:chiric sanangoblanco. Solanumnudumis an extremelycommonshruband small treeof secondaryhabitatsthroughoutlow-elevation Centraland South America, but in some parts of its range (see below) it grows in higher-elevationforests. It often formsdense, monospecific standsin open areas and river banks. Solanum nudumis one of the most commonly collected species of sect. Geminata. Solanumnudumis remarkablyuniformmorphologically, consideringits broadrange.In Venezuela,highelevation specimens of S. nudumhave largerflowers andmore coriaceousleaves thanmaterialfromCentral America and the Caribbean,but plants with all intermediatesin bothof these charactershave been collected ascending the mountains.This large-flowered, highelevation race has been called S. tovarense. Specimensfromthe Amazonbasin,andparticularly those from the areaaroundIquitos,Peru,tend to have much larger leaves than plants collected in Central Americaandthe Caribbean.These Iquitospopulations also have trichomesuniformlydistributedon the inflorescence axis and on the buds. The nameSolanumnudumhas been used in several floras for what is now called S. aphyodendron(see Knapp, 1985). The holotype of S. nudum in the HumboldtandBonplandherbariumat P clearlybelongs to the materialpreviously called S. antillarumso this name commonly used in floristic works in the 1960s and 1970s is a synonym of S. nudum.Furtherdiscussion of the nomenclaturalchanges and characteristics of the type at P can be found in Knapp(1985).

TAXONOMIC TREATMENT .. ::

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FIG. 51. Solanum pseudoquina A. St.-Hil. Lectotype. Brazil. St. Hilaire s.n. (P).:::;:::i::.:':';':

Severallectotypificationproblemsarosewithrespect to the synonyms ofSolanum nudum.As a lectotype of S. antillarum I have chosen one of the many collections cited by Schulz in his original descriptionof the species, Eggers 5039. A duplicateof this collection is found at US, and is in good condition. Others of the syntypes are sterile, and are otherwisenot suitablefor designation as the lectotype. The lectotype chosen for S. parcebarbatum is again one of many collections cited, and is well documentedand in good condition. The specimenat US cited by Bitter(Pittier2325) bears an annotation label in his handwriting. Solanum

tovarense, the high-elevation Venezuelan race, is lectotypified with the first collection cited by Bitter in his description. Several specimens at G have annotations in Bitter'shandwriting,all undated.I have selected the sheet from the Barbey-Boissier herbariumas the lectotype, as duplicatesof it are widely distributed. 20. Solanum pseudoquina A. St.-Hil., PI. Usuel. Bras., part 5: tab. 21, 1. 1825. Type. Brazil. Sao Paulo: "Saint Paul," St. Hilaire s.n. (holotype, P [Mortonneg. 8302]). Figs. 51, 52

14

FLORA NEOTROPICA

./

Pent. Monog. SOLANUIM, INAQUALE (Tab. 16.)

FIG. 52. Solanum pseudoquina A. St.-Hil., plate 116 of Vellozo's Florae Fluminensis Icones, Vol. 2 (as S. inaequale Vell.)

Solanum inaequale Veil., Fl. Flumin. 87. 1829 [1825]. Type. Brazil. Rio de Janeiro: "Habitat silvis maritimis ad ripis vulgo dicti Taguahy"(No specimens extant; lectotype, Vellozo, Fl. Flumin. Icones 2: fig. 119. 1831 [1827], here designated). Solanum flagrans Ten., Sem. Orto Bot. Napoli 12. 1839. Type. Originally from Brazil, cultivated in Naples (no specimen traced). Pionandra flagrans (Ten.) Miers, London J. Bot. 4: 364. 1845. Based on Solanum flagrans Ten. Cyphomandraflagrans (Ten.) Walp., Repert. Bot. Syst. 6: 579. 1847. Based on Solanum flagrans Ten.

Solanum undatifolium Dunal in DC., Prodr. 13(1):

104. 1852. Type. Brazil. Santa Catarina:s.loc., Gaudichaud 162 (holotype,P; isotype,frag.MPU). Solanum ramosissimum Dunal in DC., Prodr. 13(1):

144. 1852. Type. Brazil. Mtns. of Sta. Tereza, Nov 1815, Bowie & Cunningham s.n. (holotype,

BM; isotype, frag. MPU). Shrubsor small trees,2-10 m tall;young stems and leaves minutely glandularpuberulent,soon glabrate; barkof olderstemsyellowish-green,glabrousandshining; barkof large branchesand trunkyellowish-gray,

TAXONOMIC TREATMENT

longitudinallyrugose.Sympodialunitsdifoliate,geminate. Leaves narrowly elliptic, widest at the middle, glabrousabove andbelow except for tuftsofuniseriate trichomes in the axils of the main lateralveins; major leaves 6-13 x 2.5-5.5 cm, with 7-9 pairs of main lateralveins, these prominentandpale yellow below, the midribraisedabove, the apex acute,the base attenuate, only slightly decurrenton the petiole; petioles 1.5-2 cm long; minor leaves differing from the majorones only in size, 4.5-8 x 2-3 cm, the apex acute, the base attenuate;petioles 0.5-1 cm long. Inflorescences opposite the leaves, simple or occasionally furcate, 1-3 cm long,25-40-flowered,theextremetipminutelyglandularpuberulentlike the young leaves;pedicel scars unevenly spaced0.5-1.5 mm apart,beginningnearthe baseof theinflorescence.Budsglobosewhenverysmall, laterellipsoid, the corolla soon exsertedfromthe hyaline calyx. Pedicels at anthesisslender,0.9-1 cm long, taperingfrom the calyx tube to a slender base ca. 0.5 mm diam.Flowerswith the calyxtubeca. 1.5 mm long, taperingto the pedicel, the lobes square,apiculate,the apex with few uniseriatetrichomesca. 0.25 mm long, the marginshyaline;corolla white, 1.2-1.8 cm diam., planarat anthesis, lobed nearlyto the base, the tips of the lobes minutely papillose; anthers dimorphic,falcate, three large ones 3-3.5 mm long, two short ones 2-3 x ca. 1 mm, poricidal at the tips, the pores round and very small, the outer surface of the antherswith thickenedcell walls, appearinggrayishin drymaterial; free portionof thefilaments ca. 1.5 mm long for the 3 largeanthers,0.5 mm long for the remaining2 anthers, the filament tube ca. 1 mm long, ca. 0.5 mm long betweenthetwo shortanthers;ovaryglabrous;stylestraight, ca. 7 mm long;stigmaclavate,minutelypapilloseat the extremetip.Fruita globose,greenberry,1.2-2 cm diam.; fruitingpedicels woody, erect or somewhat deflexed, ca. 1.5 cm long, expandedat the distal end to ca. 3 mm diam., ca. 1.5 mm diam. at the base. Seeds yellowish, flattened-reniform,3.5- x 3-3.5 mm, the margins incrassate,the surfacesminutelypitted.Chromosome

115

.7-.

/n)?

500

/

0

FIG. 53. Distribution of Solanum pseudoquina (solid circles), S. restingae (open circle), and S. santosii (solid square).

65875 (F); Vicosa, Fazenda da Creciuma, 700 m, 15 May 1930, Mexia 4706 (MO, NY, US); Vicosa, Fazenda de Aguada, small valley 0.5 km beyond gate, 700 m, 15 Oct 1930, Mexia 5176 (BH, F, K, MO, NY, TEX, US); Vicosa, Fazenda de Jose Alexandre, 700 m, 29 Dec 1930, Mexia 5462 (BH, F, K, MO, TEX, U, US); Caldas, 20 Oct 1873, Regnell 652 (K); Caldas, 30 Oct 1862, Regnell 973 (U, US); Ouro Preto, 18 Oct 1877, Schwacke 13980 (P). PARANA: Ponta Grossa, 16 Jan 1909, Dusen 7554 (F, K, MO, NY); Antonina, 30 Oct 1909, Dusen 8867 (F, K, NY); Porto Dom Pedro II, 25 Feb 1911, Dusen 11440 (F, NY); Alexandra, 5 Mar 1911, Dusen 11493 (F, K, MO, NY); Serra do Mar, Desvio Ypiranga, 700 m, 22 Aug 1914, Dusen 15445 (F, K, NY); Jaguariahuaga, 13 Jan 1915, Dusen 16336 (F); Iacarehy, 11 Nov 1915, Dusen 17309 (F, K, LL, MO, NY); Curitiba, cultivated in garden of Faculty of Pharmacy, Fob 1964, Filho 382 number not known. (NY, US); Cerro Azul, Morro Grande, 20 Jun 1957, Distribution (Fig. 53). Common in the forests of Hatschbach 3951 (US); San Jose dos Pinhaes, Areia southeastern Brazil, from Espiritu Santo to Santa Branca dos Assis, 15 Oct 1958, Hatschbach 5150 (US); Catarina,and in Argentinaand Paraguay,at 100-800 Morro Grande, Cerro Azul, 3 Oct 1961, Hatschbach 8304 (F, US); Paramagua,Caioba, 5 m, 7 Dec 1963, Hatschbach m, in a variety of habitats. 10773 (US); Pato Branco, Laranjeiras do Sul, 16 Dec Selected specimens examined. BRAZIL. S.loc., 1966, Hatschbach 15520 (F, NY, US); Belizario, Campina Burchell s.n., cat. 4475 (K); s.loc., Martius 261 (K); Grande do Sul, 4 Oct 1967, Hatschbach 17302 (K, s.loc., cultivated in Naples, 1847, Tenore s.n. (G-DC); UPCB); Campina Grande do Sul, sitio do Belizario, 1100 s.loc., s.coll. (G-DC); s.loc., Schott 5421 (F); s.loc., 1832, m, 27 Dec 1967, Hatschbach 18184 (BH, MO, NY, US); Lhotsky s.n. (G-DC); s.loc., 1864, Sellow s.n. (F); s.loc., Castro, Abapa, 14 Oct 1968, Hatschbach 20076 (MO); Sellow s.n. (K, P, U); s.loc., 1815-1817, Sellow s.n. (MO); Tijucas do Sul, Pirai, 25 Oct 1977, Hatschbach 40394 s.loc., Swainson s.n. (K); s.loc., Tweedies.n. (K). MINAS (BH, NY) Lageado Cinco Reis, mun. Jaguariaiva, 7 Dec GERAIS: Sao Juliao, Oct 1945, Jordao 2330 (US); Vicosa, 1988, Hatschbach & Cordeiro 52650 (BM); Villa Velha, Esc. de Agronomiade Vicosa, 2 Oct 1935, Kuhlmann 875 m, 30 Apr 1914, Jdnsson 249a (NY); Mata Costa

116 da Lagoa, 490 m, 21 Jan 1969, Klein & Souza Sob. 8077 (US); Marumbi, Serra do Mar, Nov 1971, Kuniyoshi 3103 (US); Antonina, 15 Nov 1972, Kuniyoshi 3552 (US); Guaraquecaba,3-40 km along rd. to Antonina, 10200 m, 25?15'S, 48?30'W, 14 Dec 1977, Landrum 2898 (NY); vic. of Pontal do Sul, 0 m, 2 Jan 1967, Lindeman et al. 3822 (NY); Guarapuara, 10 km W of Guarapuara, 1100 m, 14 Dec 1965, Reitz & Klein 17653 (F, NY, US, WIS); Curitiba, cultivated, 15 Nov 1954, Stellfeld 547 (NY); Seminario, Curitiba, 28 Oct 1966, Stellfeld 1625 (NY, UPCB, US); Ivahy, 300 m, 12 Feb 1937, Tessmann 6062 (K). RIO DE JANEIRO: Tejuca, Nov 1836, Gardner 237 (K, NY); s.loc., 1834, Gaudichaud 518 (F, G-DC, P, US), Gaudichaud 518bis (G, P); s.loc., Glaziou 8199 (K); near Rezende, 22 Nov 1876, Glaziou 8862 (K, US); s.loc., 20 Nov 1878, Glaziou 11386 (F, K); Santa Teresa, Jan 1839, Guillemin 647 (P); Morro da Babil6nia, Nov 1914, Hoehne 30098 (US); Serra dos Orgaos, Sep 1831, Lhotsky 106 (F, G-DC, MO); Serra de Petr6polis km 50, Nov 1973, Occhioni 5748 (F); Itaguai, 21 Oct 1957, Pabst 4329 (US); Guanabara, Estrada da Boiana, Jacarepagua,1 Oct 1958, Pereira et al. 4371 (F, US [mixed collection with S. campaniforme]); Serra de Itatiaiana, Itamonte, 1900 m, 16 Sep 1961, Pereira 5782 (US); Cabo Frio, Arrarial do Cabo, Pontal beach, 13 Aug 1953, Restinga I-Segadas-Vianna et al. 834 (US); Serra do Estrella, Dec 1832-Jun 1833, Riedel s.n. (NY); s.loc., Oct 1832, Riedel 1079 (NY, US); margins of Rio Itapoana,Oct 1909, Sampaio 1042 (US); s.loc., Schwacke s.n. (US); s.loc., Simon s.n. (P); s.loc., 1816-1821, St. Hilaire Catal. B2 356 (P); Serra dos Orgaos, 1833, Vauthier 529 (G-DC, MPU, P). RIO GRANDE DO SUL: Morro do Subia, 28 Dec 1948, Rambo 39272 (F); near Porto Alegre, 19 Oct 1949, Rambo 43987 (K, US); Barreto Viana p.S. Leopoldo, 24 Oct 1949, Rambo 44128 (US); Lagoa dos Barros near Osorio, 14 Dec 1949, Rambo 44740 (TEX); San Francisco de Paula, 18 Dec 1949, Rambo 44865 (RSA/POM); Toca do Tigre near Itapoan, 11 Oct 1950, Rambo 48933 (MO); Butterbergnear Montenegro, 13 Nov 1950, Rambo 49141 (NY); Porto Alegre, mtns. of Belem, Oct 1896, Reineck s.n. (RSA/POM); s.loc., 1897, Reineck & Czermack 90 (K); Pareci Novo, Montenegro, 50 m, 24 Oct 1945, Sehnem 1639 (NY). SANTA CATARINA: Horto Florestal, INP, Ibirama, 200 m, 2 Dec 1953, Gerieski 66 (NY, US); Mata da Azambuja, Brusque, 50 m, 30 Dec 1949, Klein 116, 249 (US), 17 Feb 1950, Klein 258 (US); Braco Joaquim, Luis Alves, Itajai, 400 m, 13 Jan 1955, Klein 1070 (NY); Morro da Ressacada Itajai, 10 m, 18 Nov 1955, Klein 1773 (K, UCPB); Alto da Serra, Encruzilhada, Lajes, 900 m, 4 Dec 1962, Klein 3180 (US); Ilha da Santa Catarina, Florian6polis, Morro dos Inglezes, 50 m, 23 Nov 1965, Klein & Bresolin 6324 (US); Ilha de Santa Catarina, Florian6polis, Morro do Ribeirao, 250 m, 16 Jan 1967, Klein 7084 (US); Ilha de Santa Catarina, Florian6polis, Saco Grande, 250 m, 23 Nov 1967, Klein & Bresolin 7672 (US); Itajai, Cunhas, 30 m, 27?S, 48?46'W, 13 Nov 1977, Landrum2496 (NY); s.loc., Jun 1868, Mueller 157 (K); Horto Florestal, INP, Ibirama, 250 m, 1 Mar 1954, Reitz & Klein 1610, 1666 (K, US); Braco Serafim, Luis

FLORA NEOTROPICA Alves, Itajai, 100 m, 22 Jan 1948, Reitz C2011 (US); Braco Joaquim, Luis Alves, Itajai, 300 m, 7 Jan 1955, Reitz & Klein 2342 (K, UCPB, US); Horto Florestal, INP, Ibirama, 500 m, 13 Apr 1956, Reitz & Klein 3145 (NY, US); Rancho de Taboa, S. Jose, 500 m, 27 Oct 1957, Reitz & Klein 5543 (US); Barracao, Bom Retiro, 500 m, 27 Oct 1957, Reitz & Klein 5522 (US); Tres Barras, Garuva, S. Francisco do Sul, 10 m, 19 Dec 1957, Reitz & Klein 5727 (US); Estrada Dona Francisca Joinvile, 600 m, 6 Nov 1957, Reitz & Klein 575 (NY, US); Serrado Matador,Rio do Sul, 500 m, 29 Dec 1958, Reitz 6096 (NY, US); Rio do Sul, 400 m, 18 Oct 1958, Reitz & Klein 7394 (US); Pirao Frio, Sambrio, 10 m, 31 Oct 1959, Reitz & Klein 9306 (K, US); Serra do Espigao, Papandura, 1000 m, 24 Oct 1962, Reitz & Klein 13405 (NY, US); Mondai, 250 m, 31 Dec 1963, Reitz & Klein 16697 (NY, US); Catanduvas, 17-19 km W of Joacaba, 700-800 m, 27?03'S, 51?45'W, 15 Dec 1964, Smith & Klein 13944 (MO, NY, US). SAo PAULO: S.loc., s.coll. (US); Serra da Cantareira,Estrada da Pedra Grande, 4 Sep 1978, de Aguiar 5743 (NY); Santo Andre, Paranapiacaba,Estacao Biol6gica, 800 m, 23?46'S, 46?17'W, 30 Jul 1980, Custodio Filho 305 (NY); Oleo, 20 May 1958, Edwall s.n. (US); Ubatuba, top of Morro Escuro, 800 m from Rio Escuro, 7 Nov 1961, Fontella & Moura 98 (US); Ubatuba, near Hotel das Flores, 9 Nov 1979, Gibbs et al. 3484 (NY); s.loc.. 1833, Gaudichaud s.n. (herbier Imperial du Bresil no. 62) (P); Ubatuba, Agriculture Experiment Station, along Rio Comprido, 100 m, 23?30'S, 45?00'W, 8 Jan 1985, Gentry & Zardini 49341 (MO, NY); Alto da Serra, 6 Feb 1920, Hoehne 3669 (US); Parque do Estado, cultivated, 23 Sep 1931, Hoehne (Inst. Biol.) 28274 (BH, F, US); Serra da Cantareira, 28 Mar 1933, Koscinski s.n. (US); Serra da Cantareira, 12 Sep 1933 (fr. 14 Dec 1933), Koscinski 211 (US); Boituva, 21 Dec 1887, Loefgren s.n. (US); Piruibe, 26 Oct 1891, Loefgren & Edwall s.n. (US); Santa Lucia, in a park, 15 Dec 1943, Pickel 1117 (US): Santo Amaro, 10 Oct 1943, Roth 825 (F); Alto da Serra, Oct, Schwebel s.n. (US); Santa Maria da Serra, Fazenda Barreiro Rico, 13 Dec 1977, Tamashiro 4182 (F, NY); s.loc., 1861-62, Weir s.n. (K). PARAGUAY. In regione fluminis Rio Alto Parana, 1909-1910, Fiebrig 6024 (G, US); Rio Yaca, 1900, Hassler 6796 (K, MO, MPU, NY). ALTOPARANA: Centro Forestal Alto Parana, 12 km W of Pto. Stroessner (Cuidad del Este), 25?38'S, 54?42'W, 16 Apr 1986, Brunner 1807 (PY). CANENDIYU: Sierra de Maracayu, Hassler 5096 (G, K, MO, NY). PRIMAVERA: Alto Rio Paraguay, 28 Jan 1956, Woolston 642B (US). SAN PEDRO:Alto Paraguay,Primavera,28 Jan 1956, Wbolston 642 (K, NY, U); Primavera, Alto Paraguay, 15 Jan 1957, Woolston 1415 (K). ARGENTINA. CORRIENTES:Santo Tom6, Establ. las Marias, Ruta Nac. 14, 7 km S of Gdor. Virasora, 2 Dec 1970, Krapovickas et al. 16904 (US). MISIONES: Candelaria, Bella Vista, 6 Nov 1945, Bertoni 2636 (P); El Dorado, 10 Dec 1943, Burkart 14687 (US); vic. of Puerto Aguirre, 100 m, 8-10 Jul 1914, Curran 28 (NY, US); San Ignacio, 15 Mar 1947,Medina 310 (P); Frontesa, Barranc6n Pepiri-Guazu, 5 Nov 1949, Montes 7110

TAXONOMIC TREATMENT117

(US); Posadas,19 Mar 1930, Rodriguez142 (US); San Ignacio, SantoPip6 (PuertoSanto Pip6), 21 Feb 1948, Schulz 7219 (P); Cainguas,PuertoLeoni, 26 Nov 1945, Schwarz 1563 (P); San Antonio, Salto Tabaz,20 Jan 1946,Schwarz1868 (F); Cainguas,Tabay,25 Nov 1948, Schwindt987 (RSA/POM);ParqueNacionalIguazu,rt. 101 betweenArroyoYacuyand PuestoTimb6, 19 Dec 1991, Vanni et al. 2982 (U).

URUGUAY.In hortoBuschentalculta Montevideo, s.coll. (G).

Local names. Brazil. Parana, Sao Paulo: quina; SantaCatarina:canema.

Smalltreesto 8 m tall;youngstemsandleavesdensely tomentosewithuniseriategoldencurl-tippedtrichomes 0.5-1 mm long, these not markedlydeciduouson older stems; barkof older stems yellow-brown and shining. Sympodialunits difoliate, geminate.Leaves elliptic to ovate, widest at or just below the middle, densely tomentosebeneathwith uniseriatecurl-tippedgolden trichomes0.5-1 mm long, especiallyalongthe midriband the 6-8 main lateralveins, glabrousandshiningabove, withuniseriatetrichomeslike those of theundersideson the midribandmainlateralveins;majorleaves 9.9-12 x 4.5-5 cm, the apex acute, the base acute;petioles 1.41.5cm long,denselytomentoselikethestemsandleaves; minorleaves, if present,2.3-5.5 x 1.5-2.4 cm, the apex acute, the base acute;petioles 5-8 mm long. Inflorescences opposite the leaves, or appearingterminaland overtoppingtheshoots,simpleorseveraltimesbranched, 1.5-3.5 cm long, 20-60-flowered, densely tomentose with the same uniseriategolden hairs as the stem and leaves,thesevery densedistally;pedicelscars hiddenin thetrichomes,evenly spacedca. 1 mm apart.Budsellipsoid, the corollasoon exsertedfromthe calyx.Pedicels at anthesis6-8 mm long, erect,taperingfromthe calyx tubeto a slenderbase ca. 0.5 mm diam.,pubescentwith the samegoldentrichomesas the stemandleaves.Flowers with the calyx tube 1-1.5 mm long, pubescentwith golden uniseriatetrichomes,the lobes broadlytriangular to deltoid, 0.5-1 mm long, densely pubescentwith uniseriategolden trichomes;corolla white, 1.2-1.4 cm across, lobed nearly to the base, the lobes planar or slightlycampanulateat anthesis,the interpetalarsinuses membranous,the lobes long-papillosewith a few scattereduniseriatetrichomes0.2 mm long at the tips;anthersclavate,falcate,unequalin size, two smallones ca. 2 mm long, threelargeones ca. 2.5 mm long, all 1-1.25 mm wide, minutelyporicidalatthetips,the poresround andverysmall;freeportionof thefilamentsalsounequal, the two small antherswith the free portionca. 1.5 mm long, the threelarge antherswith the free portionca. 2 mm long, the filamenttube ca. 0.5 mm long;ovaiy glabrous;style 4.5-5 mm long, straightor slightly curved; stigma dark-papillose.Fruit a globose, greenberry,ca. 1 cm diam.;fruitingpedicels deflexed, 2-2.5 cm long. Seedsnotknownfrommaturefruits.Chromosomenumber not known.

Solanumpseudoquina has long been known as S. inaequaleVell., but the epithetpseudoquinahas priority (see above andCarauta,1973, for dates of publication of Floraefluminensis). Solanumpseudoquina is closely relatedto a rarespecies of southeasternBrazil, S. reitzii, with which it sharesdimorphicantherswith small, roundpores. This characterstate is found only in these two species. Solanumpseudoquina is easily distinguishedfromS. reitzii by its largerflowers, glabrous stems, leaves with pubescence only in the axils of the main lateralveins beneath, and the pale yellow venationof drymaterial.Solanumpseudoquinais common in southeasternBrazil.It grows to be a largetreelet and is often locally common where it occurs. In Sao Paulo, St. Hilairereportedthe use of the bark of Solanumpseudoquinaas a febrifuge,hence the specific epithet.A chemical extractof the barkrevealed a high percentageof a bitter substance("principeamer de nature purementvegetale. 1/12"), which was not furtheranalyzed(St. Hilaire, 1825). Two otherspecies of the S. nudumspecies group(S. nudumin Colombia and S. spirale in Laos) are also reportedto be used as febrifuges.Chemicalanalysesmay revealcharactersto furthersupportthe unity of this difficult group. The type specimen of Solanumpseudoquina is in fruit(see Fig. 51), so until a carefulexaminationof the material was made it was not known that this name applied to the same taxon as did S. inaequale. A type specimen does not exist for S. inaequale, but there is no doubt about the disposition of this name since the unequalanthersare so striking.The type specimen of S. pseudoquina is an exact match for materialidentified as S. inaequale from nearVellozo's type locality. The absence of flowers on the type of S. pseudoquina Distribution (see Fig. 60). Foundonly in the mounled botaniststo ignore it as a valid epithet,andhas cretains of Santa Catarinaand adjacent Parana states, ated some confusion in herbaria. southeasternBrazil, inAraucaria angustifolia forests at ca. 800 m. 21. Solanum reitzii L. B. Sm. & Downs, Phytologia Specimens examined. BRAZIL. PARANA: Londrina, 10: 426. 1964. Type. Brazil. Santa Catarina:Alto Floresta do Godoy, 27 Aug 1991, Chagas e Silva 1410 Matador,Mun. Rio do Sul, pinhal, 800 m, 30 Dec (FUEL);Tibagi, Fazendada Barrado Rio Borrinha,4 1958, Reitz 4092 (holotype, US; isotypes, HBR, Dec 1989, Chagas e Silva et al. 7802 (FUEL); Palmeira, NY-n.v.). Fig. 54 Fazenda Capao Bonito, 8 Jan 1991, Chagas e Silva et

FLORA NEOTROPICA

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TAXONOMIC TREATMENT

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FIG. 55. Solanum restingae S. Knapp. (Reproduced with permission from Bulletin of the British Museum, Natural History (Botany) 19: 110, fig. 7. 1989.)

al. 20268 (FUEL); PuntaGrossaopp., 800 m, 14 Nov 1914, Dusen 15986 (F, K); Fda. das Almas, Mun. de Jaguariava, 29 Nov 1968, Hatschbach 20469 (F); Cerro Azul, Cab. Rib. do Tigre, 8 Dec 1983, Hatschbach47843 (NY); Ponta Grossa, parque Vila Velha, 23 Nov 1992, Passos de Lima 11 (FUEL); Segundo Faxinal, trail to Monte Alegre, Mun. Lapa, 30 Nov 1989, Ribas & Silva 197 (BM). SANTA CATARINA: Mafra, PotreiroGrande, 750 m, 11 Dec 1965, Reitz & Klein 17422 (NY, US); Mafra, Potreiro Grande, 750 m, 11 Dec 1965, Reitz & Klein 17424 (F).

Local names. Brazil. Santa Catarina: canema, peloteira. Solanum reitzii is most closely related to S. pseudoquina,

a widespread species of southeastern

Brazil, with which it shares the unusual characterof dimorphic anthers with round, very small pores.Solanum

reitziidiffersfromS.pseudoquinain its dense indument of golden uniseriatetrichomes,and its slightly smaller flowers. Solanumreitzii may be an isolate of the more widespreadand closely relatedS. pseudoquina. Solanumreitzii is one of the most temperatemembers of sect. Geminata,growing in Araucariaangustifolia forests as an understoryshrub.These habitatsare rapidly disappearing;this accounts for the paucity of collections of S. reitzii. 22. Solanum restingae S. Knapp, Bull. Brit. Mus. (Nat. Hist.), Bot. 19: 109. 1989. Type. Brazil. Bahia: Mun. Valenca, km 11 of rd. from Valenca to Povado de Guaibim (litoral), 13 Aug 1980, Hage, de Carvalho& MattosSilva 389(holotype,CEPEC; isotype, F). Fig. 55

120

Shrubs, 2-2.5 m tall; young stems glabrous, the young leaves sparsely red-papillose;older stems yellowish-brown and shiny, strongly winged from the decurrentleaf bases and from the inflorescencebases. Sympodial units difoliate, geminate. Leaves ellipticobovate, widest at the middle or just above, glabrous and shiny adaxially, glabrous abaxially, the margins revolute;majorleaves 11-15.5 x 3.5-7 cm, with 8-9 pairsof mainlateralveins, these yellowish beneath,the apex acute,the base attenuate,decurrentonto the stem; petiole 0.5-1.5 cm long; minor leaves differing from the majorsonly in size, 5-7 x 2-3.8 cm, the apex acute, the baseattenuate;petiole0.4-1 cm long.Inflorescences usually intemodal, occasionally opposite the leaves, simple, completely glabrous, 2-5 mm long, with 3-5 flowers;pedicel scars closely spaced, clustered near the tip of the inflorescence. Buds globose, later longellipsoid, the corolla strongly exserted from the calyx tube,the tips of the lobes cucullate.Pedicels at anthesis erector slightlydeflexed,0.9-1 cm long, filiform,tapering from the base of the calyx tube to a base 0.4 mm diam.Flowers with the calyx tube an open cup, 1.5-2 mm long, the lobes glabrous,minute,if presentbroadly deltate, 0-0.5 mm long; corolla white, perhapssomewhat fleshy, 1-1.5 cm diam., lobed nearlyto the base, the lobes planarat anthesis,the tips of the lobes cucullate andminutelypapillose;anthers3-3.5 x ca. 1 mm, poricidal at the tips, the pores teardropshaped; free portionof thefilaments ca. 0.2 mm long, the filament tube 0.1-0.2 mm long; ovaryglabrous;style erect,5-6 mm long; stigma clavate, the surface minutely papillose. Fruita globose, greenberry,0.8-1 cm diam.;fruiting pedicels woody, deflexed, 1-1.2 cm long, 1-1.5 mm diam.atthebase.Seedspaletan,flattened-reniform, with incrassatemargins, 3-3.5 x 2-2.5 mm, the surfaces minutelypitted.Chromosomenumbernot known. Distribution (Fig. 53). Only known from the restingaarborealforeston sandy soils nearthe coast in southernBahia. Specimens examined. BRAZIL. BAHIA: Mun. Valenca, Valenca-Guaibim rd., km 8 E of Valenqa, 27 Jul 1981, de Carvalho & Gatti 809 (CEPEC, F); Cairu, Ilha

de Tinhare,Fazendados Piloes, ca. 20 m, 20 Nov 1985, MattosSilva & dos Santos 1916 (BM, CEPEC,NY). Solanum restingae is a distinctive species with its strongly winged stems, completely glabrous mature foliage,indistinctcalyx lobes, andcucullatecorollalobe apices.WithintheS. nudumspeciesgroupit is mostsimilarandprobablymostcloselyrelatedtoS. daphnophyllum of easternBolivia. Solanum restingae differs from S. daphnophvllumin its difoliate sympodialunits, larger flowers with strongly hooded corolla lobes, and

FLORA NEOTROPICA

strongly winged stems. Such stronglyhooded corolla lobe apicesarealso foundin S. cucullatum,an unrelated species of sect. Geminata (S. robustifrons species group) from premontaneforests in western Ecuador. The habitatof Solanumrestingaeis disturbedarboreal restingaon white sandnearsea level. The species is apparentlyof extremelyrestricteddistribution,andhas onlybeencollectedin a tinyareaoff theValenca-Giuaibim road and on the Isla de Tinhare. Large areas of the restingasofBahia remainunexplored,however,so more collectionsmay show it to be of somewhatwiderdistribution.Manyinterestingspecieshavebeencollectedfrom the arborealor shrubbyrestingas on white sand soils in Bahia (de Carvalho,pers. comm.; Plowman, 1987). 23. Solanum santosii S. Knapp,Bull. Brit. Mus. (Nat. Hist.),Bot. 19: 106. 1989.Type.Brazil,Bahia:Mun. Ilheus, CEPEC (Centro de Pesquisas da Cacau) inventory area, quadratD, km 22 Ilh6us-Itabuna (BR415), 50 m, 10 Sep 1982, TS. dos Santos 3797 (holotype, CEPEC;isotype, F). Fig. 56 Shrubs, 1.5-3 m tall; young stems glabrousor with a few uniseriatetrichomesalong the angles, soon becoming glabrous,green,andshiny;olderstems slightly winged from the decurrentpetioles. Sympodial units difoliate,usuallygeminate.Leaveselliptic,widestatthe middle,glabrousandshinyadaxially,glabrousabaxially witha few white,simple,uniseriatetrichomesin thevein axils, the trichomesarisingfromthe veins andnot from the leaf lamina,the leaf marginsslightlycrisped;major leaves 12-15 x 4.5-6.5 cm, with 7-8 pairsof main lateralveins, thesedryingyellowish,theapexacuteto acuminate,thebase acute,oblique,minutelydecurrentonto the petiole andstem;petioles 0.8-1.2 cm long, winged fromthedecurrentleafbases;minorleavesdifferingfrom the majorsonly in size, 3.5-5.5 x 2.2-3.7 cm, the apex acute, the base acute;petioles 3-5 mm long. Inflorescences oppositethe leaves, simple, 1-2.5 cm long, 1020-flowered,glabrous;pedicelscars closely spaced,not overlapping.Buds ellipsoid,the tip pointed,the corolla stronglyexsertedfromthe calyx tube.Pedicels at anthesis erector slightlydeflexed, 1.2-1.5 cm long, filiform, taperingfromthe calyx tubeto a base ca. 0.25 mm diam. Flowers with the calyxtubebroadlyconical,very short, ca. 0.5 mm long, the lobes broadlyspathulate,the apex fleshy andenlarged,perhapsknob-likewhen fresh,0.51 mm long, the sinusesrounded;corollagreenor greenish-white,0.8-1 cm diam.,lobednearlyto the base, the lobes planarat anthesis,the tips of the lobes minutely papillose;anthers2-2.5 x ca. 1 mm, slightly sagittate, poricidalat the tips,theporesteardropshaped;freeportion of thefilamentsca. 0.2 mm long, the filamenttube ca. 0.25 mm long; ovaryglabrous;style erect, 5-6 mm

TAXONOMIC TREATMENT

121

FIG. 56. Solanum santosii S. Knapp. (Reproduced with permission from Bulletin of the British Museum, Natural History (Botany) 19: 108, fig. 5. 1989.)

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FIG. 57. Solanum spirale Roxb. A. lectotype in Wallich herbariumat K. B. People's Republic of China. McLaren et al. 197B (BM).

long; stigma minutely bilobed, the surface minutely papillose.Fruit a globose, greenberry,1-1.3 cm diam., fruitingpedicels woody, deflexed, 2-2.5 cm long, ca. 0.5 mm diam. at the base. Seeds dark brown in dry with thickenedmargins,3material,flattened-reniform 4 x 2-2.5 mm, the surfaces minutely pitted. Chromosome numbernot known.

Solanum santosii is most similar to S. warmingii and S. caavurana, also of southeasternBrazil. It differs fromthose species in its much smallerflowers and smallercalyx lobes.BothS. warmingiiandS. caavurana have expanded,petaloid, calyx lobes. Neither of those two species grows in the wet forest of coastal Bahia, the habitatof S. santosii. Solanum santosii has only been collected in the Distribution (Fig. 53). In wet forest ("mata CEPEC forest inventory plot between Ilheus and in near sea level. Known southern Bahia, higrofila") Itabuna.This areais somewhatsecondaryin nature(de only fromthe type locality. Carvalho,pers. comm.) andI suspectS. santosii grows Specimens examined. BRAZIL. BAHIA: Ilh6us, CEPEC(Centrode Pesquisasdo Cacau)inventoryarea, in light gaps in the forestas do many othermembersof quadrat D, km 22 Ilh6us-Itabuna(BR 415), 50 m, 4 Mar the S. nudumspecies group.It is apparentlynot a com1981, Hage, dos Santos & da Vinha 510 (CEPEC, F); mon plant, and it is surprisingthat it had not been colsame locality, 29 Apr 1981, Hage & dos Santos 630 lected before 1978, having been overlooked by the (CEPEC, F); same locality, 12 Jan 1982, Hage & E.B. numerousbotanistswho have worked aroundthe city dos Santos 1596 (CEPEC, F); grounds of CEPEC, km of Ilheus since the early 1800s. 22 of BR-415 from 24 Nov Maas

Ilheus-Itabuna, 1987, et al. 6976 (U); Salvador, Mata de Oitis, Centro Administrativo da Bahia, 10-30 m, 12?57'S, 38?27'W, 5 Jul 1985, Noblick & Britto 4413 (NY); Ilh6us, CEPEC (Centro de Pesquisas do Cacau) inventory area, quadrat D, km 22 Ilh6us-Itabuna (BR 415), 5 Dec 1978, T.S. dos Santos 3405 (CEPEC, F, NY).

24. Solanum spirale Roxb., Fl. Ind. 2: 247. 1824. Type. India. Silhet, Wallichcat. no. 2619a (lectotype, K, here designated; isolectotypes, BM, K, possible frag. MPU [ex G-DC]). Fig. 57

TAXONOMIC TREATMENT

123

SolanumapoenseElmer,Leafl. Philipp.Bot. 2: 730. Distribution (Fig. 58). Paleotropical, in mid-eleva1910. Type. Philippines.Mindanao:Davao Distion forests from southern China to Queensland, trict, Todaya(Mt. Apo), Aug 1909,Elmer11599 Australia. (holotype,NY; isotypes, MO, US). Solanum superficiens Adelb., Blumea 6: 331. 1948.

Specimens examined. BHUTAN. Lingsti, Kusted, 5000 ft, Aug 1915, Cooper 4367 (BM); Shanga, Pumthana, 4000 ft, 28 Aug 1915, Cooper 4723 (BM). BANGLADESH. Sheyrpore, W. Mymensingh, 19 Jul 1867, Clarke 4830 (US); Mymensingh, 11 Jul 1872, Clarke 17271 (US). INDIA. Guahati, Aug 1851, s.coll. (P); Khasia, 5000 ft, Jun 1879, s.coll. (US); Mymersingh, Nalin, 8 Nov 1868, Clarke 8008 (BM); Malmro, Khasia, 3500 ft, 23 Oct 1871, Clarke 15840B (G); Khasia, Mamloo, 3500 ft, Shrubs1-4 m tall;young stems greenandglabrous, 23 Oct 1871, Clarke 15840C (BM); Assam, Guahati, 500 occasionally minutely papillose with rusty brown pa- ft, 18 Feb 1885, Clarke 37217D (G); Khasia, Normal, pillae; bark of older stems dark brown, white- 4600 ft, 24 Aug 1885, Clarke 40047D (BM); Shillong, lenticellate.Sympodialunitsdifoliate,geminate.Leaves 4000 ft, 3 Aug 1886, Clarke 44429C (BM); Shillong, 4000 narrowlyelliptic to elliptic, widest at or just distal to ft, 3 Aug 1886, Clarke 44429E (US); Shillong, 4000 ft, the middle, glabrousabove, with tufts ofuniseriate tri- 3 Aug 1886, Clarke 44429F (G); Assam, Monier Khal chomes 0.5-1 mm long in the axils of the main lateral on the Sonai River, Aug 1903, Gage s. n. (G); Khasia, Herb. Griffith 5902 (BM, G, P); Meghalaya, Mount veins beneath,originatingboth from the veins andthe Khasia, Hooker & Thomson s.n. (BM, G, MPU, NY, P, lamina; major leaves 12-21 x 3.6-8.8 cm, with 5-7 US); Assam, Tirap-NamchikDivide, 16 mi mark on Ledo pairsof mainlateralveins, the apex acuminate,the base Rd., 10 Aug 1945, Juan et al. 94 (G, US); Assam, Mar attenuate;petioles 1.2-2.5 cm long; minorleaves dif- 1861, King s.n. (P); Assam, Walong in jungle, 14 Sep 1950, fering fromthe majorones only in size, 5.3-7.5 x 1.6- Kingdon Ward20203 (BM); Assam, Gaulpara,Jan 1886, 2.8 cm, the apex acuminate,the base attenuate;petioles Mann s.n. (BM, G); Khasia, Mann 115 (G). BURMA. Morth Triangle, Hkinkum, 4000 ft, 29 Jul 0.5-1 cm long. Inflorescencesopposite the leaves, often somewhatintemodal,simple, glabrous,shiny dark 1953, Kingdon Ward 21213 (BM). CHINA. Near the village of Tong ina Tsen, 3 Oct brown(whitein live plants),0.5-6 cm long, 5-30-flow1888, Delomy 4718 (P); Yunnan, Meng-tsze, 22 Jul ered;pedicel scars evenly spaced ca. 1 mm apart,beHenry 11038A (MO, US); Tibet, Lohit valley, ca. 1000 ginning very nearthe base of the inflorescence.Buds m, 30 Jul 1950, Kingdon Ward 19679 (BM); Yunnan, globose to obovoid,white, appearingstrongly5-angled Lichiang Range, 1933, McLaren's Native Collectors from the ridged calyx tube. Pedicels at anthesis de- 197B (BM). flexed, 1.2-1.6 cm long, taperingfrom the base of the THAILAND. Sumka Uma, 2000 ft, 25?57'N x calyx tube to a slenderbase ca. 0.5 mm diam.Flowers 97?49'E, 27 Jul 1939, Kaulbeck 300 (BM); Chiangmai, with the calyxtube 1.5-2 mm long, 5-angled,the angles 5 Apr 1922, herb. Kerr s.n. (BM); Chiangmai, 300 m, extendingdownthepedicelin driedspecimens,the lobes 10 Nov 1914, herb. Kerr 3450 (BM); Peng Zeng, 24-27 Aug 1915, herb. Kerr 3651 (BM); broadlytriangular,apiculate,0.5-1 mm long, minutely Chiangmai, ca. 300 m, 27 Aug 1921, Noi Mao s.n. (BM); Chiangmai, 1.3-1.6 cm on the corolla white, diam., papillose apices; trail from Raheng (Tak) to Pang Ma Kham Pom, lobed ca. 3/4 of the way to the base, the lobes planaror Tak, 15-16 Dec 1920, Rock 981 (US); trail from Nam Dip to sinuses only slightlyreflexedat anthesis,the interpetalar Pang Luang, Doi Paw Waw, and Mesawt, 19-20 Dec membraneous,curledandundulate,the tips of the lobes 1920, Rock 673 (US). minutely papillose; anthers 3-3.5 x 0.75-1 mm, LAOS. Haut Laos, Bonn Gai Pro, 500 m, 11 May poricidal at the tips, the pores teardropshaped; free 1936, Poilane 26074 (P); Naminh, 1903, Spire 1424 portionof thefilamentsca. 0.25 mm long, the filament (P); Nmatha, Jun 1929, Ward8950 (NY). VIETNAM. Valley of Banton, near Yen-Luang, in tube ca. 0.5 mm long; ovaryglabrous;style straight,67 mm long; stigmabilobed, minutelypapillose.Fruit a the villages, Dec 1887, Balansa 3751 (P). INDONESIA. Java, 1835-1863, Junghuhn 404 (L); globose berry, 1.1-1.6 cm diam., dull yellow-orange when ripe;fruitingpedicels deflexed, woody, 1.9-2.3 Java, Mt. Papandayan, s.coll. (L-908.245-262); cm long, ca. 1 mm diam.at thebase.Seedsyellow ortan, Sumatra, Korthals s.n. (L). SULAWESI. Minahassa, N. Celebes, Bojong, 1888, flattened-reniformto nearly roundin outline, 3-3.5 x Warburg 15074 (NY); Minahassa, N. Celebes, Bojong, 2.5-3 mm, the marginsincrassate,paler, the surfaces 1888, Warburg 15075 (NY). pitted, the pits ca. 0.02 mm long. ChromosomenumAUSTRALIA. NEW SOUTH WALES: Mt. Gipps, ber. n = 24 (Randell & Symon 1976; Knappunpubl., Wiangarie State Forest, 2.5 mi N of Cougal, 300-400 voucherBH 81:139,bothfromAustralianpopulations). m, 28?21'S, 152?58'E, 29 Jul 1979, Coveiny & Hind

Type. Indonesia. Java: W. Java, Priangan, KartamanaEstate, 1600 m, 20 Sep 1911, Smith 641 (holotype, L). SolanumcalliumR.J.F.Hend.,Austrobaileya1: 13. 1978. Type.Australia.New SouthWales:ca. 25 km NW of Kyogle,28?27'S,152?42'E,Dec 1968, HendersonH489 (holotype, BRI-n.v.; isotypes, CANB-n.v., K, NSW-n.v.).

FLORA NEOTROPICA

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58. Distribution of Solanum spirale.

10572 (RSA/POM). QUEENSLAND: Lever's Plateau, SE of Rathdowney, property of G. Phelp, 700 m, 28?18'S, 152?55'E, 29 May 1975, Moriarty 1680 (RSA/POM); border with New South Wales, Lever's Plateau, 9 Jul 1974, Swan 68 (MO); pot-grown at W.A.R.I. from seedlings from Henderson 1259, Toonumbar State Forest ca. 26 km NW of Kyogle, 3 Mar 1974, Symon s.n. (MO).

S. spirale is expandingin Australia,perhapsindicative of its being an introduction. Anotherinstanceof a close relativeof an otherwise neotropical group occurringin the Old Worldtropics (in this case in the Society Islands) is Solanum repandumG. Forst.of sect. Lasiocarpa(Whalenet al., 1981; Bruneauet al., 1995). Solanumrepandum,however, is found only in association with humans, and Local names. India. titikuchi (Assam), oko-oing may be a recentintroduction(see discussionin Whalen (Miri), lara-tita (Goalpara), sohjaring, soh-jhari etal. 1981). (Khasia), khlein-deng (Synt.). Thailand.pak dip, puk Solanumspirale is most similar,and perhapsmost dip (Lao), pak dit. Laos. mak dit. closely related,to S. nudum,a widespreadneotropical Solanum spirale is the only paleotropicaland the species (see above). It differs from that species in its only tetraploidmember of sect. Geminata.No speci- largerleaves, inflorescences,buds, and flowers, and in mensof S. spiralehavebeencollectedin theNew World, its brightorangish-redberries.MaterialfromAustralia and its widespread use in India, Thailand, and Laos is tetraploidwith n = 24, but no authenticchromosome pointsto its being eithernativethereor an old introduc- counts exist for S. spirale fromotherpartsof its range. tion. Henderson(1978) speculates on the status of S. The countofn = 12 (Gerasimenko& Reznikova, 1968) spirale in Queenslandand concludes that if it is an in- for plants from Bogor certainlyrefersto S. diphyllum, troduction,it has not spreadas a weedy species should. a widely cultivated species often confused with S. Symon (pers. comm.) however, feels thatthe rangeof spirale. Moreliving collectionsofS. spirale areneeded

TAXONOMIC TREATMENT

to determineits relationshipsto New Worldtaxa.Crossing studies would be very useful in solving this problem. Despite its brightly colored berries,S. spirale is not closely relatedto S. pseudocapsicum or the membersof thatspecies group.Solanumspirale differsfrom those species in its deflexed fruitingpedicels, flower shape, and leaf trichomemorphology. As mentionedabove,Solanumspiraleis widely used and cultivatedin dooryardgardensin India,Thailand, and Laos. No uses appearto have been reportedfrom Indonesiaor Australia.In Assam (Knajilalet al. 1939) the roots are employed as a narcoticand diuretic,and the leaves andberriesareeaten.Berriesareeateneither raw or cooked, while leaves are eaten cooked. In Laos (Poilane 26074) the bark of S. spirale is broken and soakedin cold water,thenused as a febrifugefor adults and infants. Other members of the S. nudumspecies group are also used as febrifuges in tropical America (S. pseudoquina in Brazil, S. nudumin Colombia). Species describedby Roxburghare often difficult to lectotypify,as he did not cite specimensin the original descriptions (Sealy 1956-1957). For many of the species described in Flora Indica (Roxburgh, 1820, 1824), drawingswere made by native artistsandthese arehoused at K. A drawingwas not made forSolanum spirale, so this makes the choice of a lectotype difficult. Many of Roxburgh'sspecimens and types are in the Wallichherbariumat K (Stafleu & Cowan, 1983). I have chosen to lectotypify this species with a specimen (Fig. 57) in the Wallichherbarium(formerlythe herbariumof the East India Company in Calcutta). Roxburghwas the curatorof this herbariumfrom 1793 until 1813, and deposited many specimens there. The lectotypeI have chosen is fromSilhet,the locality cited by Roxburghin theFlora Indica, andis the only specimen of S. spirale fromthatlocality in the catalogof the plantsin the EastIndiaCompany'sherbarium(Wallich, 1830). The sheet matchesthe descriptionvery well and is probablythat collected by Roxburgh. 25. Solanum symmetricum Rusby,Mem. TorreyBot. Club 6: 89. 1896. Type. Bolivia. La Paz: Mapiri, Jul-Aug 1892,Bang 1478 (holotype,NY; isotypes, F, K, MO, US, WIS). Fig. 59 Solanum nudum Dunal var. pseudoindigoferum Hassl., Repert. Spec. Nov. Regni Veg. 15: 113. 1918. Type. Paraguay. Paraguay centralis, Cordillera de Altos, in silvis, 1897, Hassler 3793 (lectotype, G, here designated [Morton neg. 8537]; isotypes, BM, NY). Solanulm versabile C. V. Morton, Rev. Argent. Sp. Solanum 107. 1976. Type. Argentina. Tucumbn: Famailla, Quebradade Lules, 700 m, 13 Feb 1921, Venturi1181 (holotype, US; isotypes, A, SI-n.v.).

125

Shrubs or small trees, 1.5-5 m tall; young stems minutely glandular,slender;older stems glabrous,the bark becoming shiny and deep maroon. Sympodial units difoliate, geminate.Leaves elliptic, widest at the middle, glabrous above, with tufts of uniseriate trichomes 0.5-1 mm long in the axils of the main lateral veins below, the trichomesarisingfromboth the veins andthe lamina;majorleaves 11-15.5 x 4-6.5 cm, with 8-10 pairsof mainlateralveins, these impressedabove, darkandprominentbelow,the apexacuminate,thebase acuteto attenuate,decurrenton the petiole; petioles 11.6 cm long, winged from the bases of the decurrent leaves;minorleaves differingfromthe majorones only in size, 3.8-6.1 x 2-4.2 cm, the apex acute, the base acuteto attenuate;petioles6-8 mm long.Inflorescences opposite the leaves, simple (occasionally furcate), subumbellate,0.8-1 cm long, 5-20-flowered, glabrous or minutely glandular;pedicelscars closely packed in the distal2-3 mm of the inflorescence,all overlapping. Buds globose, the corolla scarcely exserted from the calyx. Pedicels at anthesis 0.9-1.1 cm long, tapering from the calyx tube to a slenderbase ca. 0.5 mm diam. Flowers with the calyx tube somewhat urceolate, ca. 1.5 mm long, the lobes deltoid,ca. 1 mm long, glabrous or with a few minute papillae at the tips of the lobes; corolla white, 1.3-1.8 cm diam., lobed nearly to the base, the lobes slightly reflexed at anthesis, with thin interpetalarsinuses, minutelypapillose with unicellulartrichomeson the tips of the lobes;anthersca. 3 mm long, the terminal0.5 mm paler and thickened, ca. 1 mm, poricidal at the tips, the pores teardropshaped; freeportionof thefilamentsca. 1 mm long, the filament tube less than 0.5 mm long; ovary glabrous; style straight,5-6 mm long; stigma capitate,bilobed, darkpapillose.Fruit a globose, greenberry,ca. 1 cm diam.; fruiting pedicels ca. 1.5 cm long, woody, deflexed, expandedjust below the calyx tube, ca. 1 mm diam. at the base. Seeds brown, ca. 50 per fruit,flattened-reniform, ca. 2 x 1.5 mm, the marginsincrassate,the surfaces minutelypitted.Chromosomenumbernot known. Distribution (Fig. 60). In mid-elevationforestsand the easternAndeanslope in Bolivia andArgentina;also in Misiones, Argentina,Brazil, and Paraguayin hilly areas;400-1000 m. Selected specimens examined. BRAZIL. Mato Grosso: Mun. de Maracay6, fazenda Santo Antonio, prop. of Alfredo Neder, ca. 590 m, 29 Dec 1973, Sucre 10533 (RB). MINAS GERAIS: Pocos de Caldas, rd. to Cristo Redentor, 14 Jan 1980, Krapovickas & Cristobal 35311 (MO). PARANA:Lobato, prop. of Irmaos Ferraz, Faz. Remanso, 23 Jul 1962, Gomes e Mattos 1058 (RB); Maringa, Orto Florestal, 12 Oct 1965, Hatschbach 12938 (F, US), 7 Dec 1965, Hatschbach et al. 13249 (NY, US).

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BOLIVIA. SANTACRUZ: S.loc., 700 m, May 1892, Kuntzes.n. (NY, US). TARIJA:Huacanque,Bermejo-Tarija rd., 119 km S of Tarija (Quebrada Alavachi), 13 May 1971, Krapovickas et al. 18744 (MO); 29.2 km S of Emboraza-Sidras rd. on rd. to Bermejo (12.7 km S of Naranjo Agrio), 22?22'S, 64?29'W, 21-23 Apr 1983, Solomon 9936 (MO, NY); 5.5 km S of Mamora, 1500 m, 22?08'S, 64?40'W, 27 Apr 1983, Solomon 10173 (MO, NY); 21.5 km E of Narvaez on rd. to Entre Rios, 6.9 km W of Entre Rios, 1350 m, 21?29'S, 64?13'W, 5 Oct 1983, Solomon 11064 (MO, NY); hillsides vic. Sidras, 5.5 km N of Emborazi, 950 m, 22?12'S, 64?32'W, 11 Oct 1983, Solomon 11129 (K, MO, NY, U); upper Rio Camnari ca. 5 km from first pass on trail from Sidras to Tariquia,1200 m, 22?10'S, 64?26'W, 13 Oct 1983, Solomon 11163 (MO, NY, U); valley of Rio Chillaguatasbelow Rancho Nogalar on trail between Sidras and Tariquia, 1100 m, 22?05'S, 64?25'W, 14-16 Oct 1983, Solomon 11257 (MO, NY). PARAGUAY.

CAAZAPA: Along Rio Pirapo, Cordi-

llera de Caaguazu, ca. 600 m, 15 Mar 1937, West 8532 (MO). CANENDIYU:Guazi rd. to Puerto Adela, 360 m, 24?30'S, 53?20'W, 16 Dec 1982, Hahn et al. 950 (K).

CENTRAL:Forests near Asunci6n, 22 Mar 1875, Balansa Tobatinear Coacupe, 5 Feb 1903, 2123 (P). CORDILLERA: Fiebrig 864 (F, K, M-n.v. (Morton neg. 8721)). GUAIRA: Villa Rica, 16 Feb 1876, Balansa 2122a (P); Colonia Independencia, 25?40'S, 56?15'W, 15 Apr 1984, Hahn 2206 (K); Villarica, Feb 1931, J6rgensen 3682 (F, NY, US). ITAPUA:Guarapi, Apr 1880, Balansa 3126 (P). ARGENTINA. CORRIENTES:Mburucuya, estancia "Santa Teresa," 20 Feb 1972, Pedersen 10057 (NY); Santo Tome, ruta 37, 5 km E of Gdor. Virasoro, 14 Nov 1974, Schinini & Carnevali 10580 (G). JuJuY: Along rd. to old coffee plantation,Calilegua, 3 Jun 1943, Bartlett 20346 (US); Sta. Barbara, rd. from El Fuerte to Palma Sola, 20 Feb 1972, Cabrera et al. 22320 (US); San Pedro, Sierra Sta. Barbara, 700 m, 5 Oct 1929, Venturi 9563 (MO, US). MISIONES:Posadas, 19 Mar 1930, Rodriguez 142 (F); Guarany, rt. 14, km 304 near Arroyo Chafariz, 4 Mar 1950, Schwindt 3286 (BH). SALTA: Orhn, quebrada of Rio Carapari, 15 Jul 1937, Cabrera 4221 (F, NY); Santa Victoria, rd. from Los Toldos to Lipeo, on the old track to the right, 5 km from Lipeo, 1300 m, 10 Dec 1973, Legname & Cuezzo 9866c (LL); Toldos-Lipeo

128

FLORA NEOTROPICA

rd., 8 km from Lipeo, 1600 m, 2 Oct 1973,Legname&

rest of the species, but definitely representthe same species. They arepossibly relictualin nature. Oran, San Andr6s, 1800 m, 7 Feb 1945, Pierotti 211 The lectotypeI have chosenforSolanumnudumvar. (NY); Oran,Aguaray,500 m, 11 Feb 1940, Schreiter pseudoindigoferumis one of the severalspecimenscited 11213 (F); Oran, 200 m, 8 Jul 1944, Schulz & Varela in the publicationof this name.Hasslerdid not cite any 570 (NY), 362 m, Jun 1944, Schulz & Varela5014 (US); herbaria,butthe specimenat G bearsanannotationlabel Oran,S of Tartagal,Nov 1915, Steinbach 1708 (US); in whatappearsto be his handwriting,andis designated Oran,Rio Balanco,650 m, 13 Nov 1927, Venturi8651 the lectotype.Hassler3 793,the lectotype,is represented (US); Rosariode Lerma,CampoQuijano, 1800 m, 20 Jan 1927, Venturi8229 (US); Guachipes,Alemania,1500 in severalherbaria,andis the second specimencited by m, 5 Dec 1929, Venturi9850 (MO, US); Oran, Rio Hassler.The duplicatesare all in good condition. Pescado9 kmifromFincade Yakulica,rd.to Los Toldos, Cuezzo 9718c (W); Oran, 20 Feb 1943, Meyer 4540 (F);

650 m, 25 Oct 1970, Vervoorst & Cuezzo 7635c (US,

W). TUCUMAN: Capital,HorcoMolle, 730 m, 26?50'S, 26. Solanum tepuiense S. Knapp, Novon 1: 125. 65?15'W,13Oct 1966,Boelckeet al. 5437(US);Monteros, 1991. Type.Venezuela.Bolivar:North-facingslope km 23 in Quebradadel Rio de los Sosa, S of the Indian forest Sororopan,cumbre Sororopan-tepui,1,800 monument, 5 Mar 1959, Diero 381 (SI); Famailla, 33940 (hom, 15 Dec 1952, Maguire & Wturdack Quebradade Lules,Rio Lules,4 Mar 1945,Herrera228 F). Fig. 61 lotype, NY; isotype, (NY); Villa Nogues, 12 Jul 1911, Lizer C. 50 (SI); base

of Cuestade la Puertade SanJavier,18 Jan 1873,Lorentz

Shrub, 2.5 m tall; young stems and leaves with a few yellowishpapillae,otherwiseglabrous,thebranches stout, strongly winged from the decurrentleaf bases; barkof older stems pale green, dull, the wings persisting. Sympodialunitsdifoliate, geminate.Leaves elliptic ("repandlyrugose"fide Maguire& Wurdack),thick and coriaceous, somewhat falcate, glabrousand shining adaxially,glabrousor with tuftsof simpleuniseriate trichomes 0.5-1 mm long, consisting of 5-15 small bead-likecells, at thejunctionof the main lateralveins andthe midribabaxially,the trichomesarisingfromthe veins, not from the lamina;major leaves 8-14 x 3-5 cm, with 7-10 pairs of main lateral veins, these impressed adaxially,prominentandyellowish abaxially, the apex acuteto acuminate,the base acuteto attenuate, (F, US); Burruyacfi,Cerrodel Campo, 1000 m, 7 Jan stronglydecurrentonto the stem;petioles winged from 1929, 'Venturi 7951 (MO, US); Burruyaci, Cerro El the leaf bases, 0.8-2.5 cm long; minor leaves not difNogalito, 1000 m, 5 Apr 1929, Venturi 8795 (US). fering fromthe majorsin shape, 3.5-8 x 1.5-3 cm, the Local names. Paraguay. Itapua: caa-hu (Guarani). apex acute to acuminate,the base acute to attenuate; Tucuman: malfato. petioles winged, 0.4-1 cm long. InflorescencesoppoArgentina. site the leaves, simple, with a few yellowish papillaeat Solanuml svmmetricurn is probably most closely the tips, otherwise glabrous and drying dark brown, relatedto S. caavurana,also of southernSouthAmerica. 1-1.6 cm long, with 10-14 flowers, only one or two Sola/num symmetricunm differs from that species in its at a time; pedicel scars evenly spaced 1-1.5 mm open pedicel scars clustered in the distal 1/3 to 1/4 of the apart,beginning about 1/3 of the way from the base. inflorescence axis, smaller, non-petaloidcalyx lobes, Buds globose when young, laterellipsoid, the corolla more deeply lobed flowers, minute filament tube, and strongly exserted from the calyx tube just before anelongate anthers.The subumbellateform of the inflo- thesis. Pedicels at anrthesisdeflexed, glabrous,fleshy, rescences is distinctive, and makes S. symmetricumn apparentlygreen,stronglytaperedtowardthebase, 1.3easy to distinguish from other members of the S. nu- 1.7cm long,ca. 0.5 mmdiam.atthebase,ca. 2 mmdiam. dum species group. at the apex. Flowers with the calyx tube conical, 1-1.5 Therearerelativelyfew oldercollectionsof Solanum mm long, the lobes broadlydeltatewith an acuteapical from Bolivia, probably due to the col- projection,0.5-1 mm long, the marginsthin andtranssynnmmetrictun lecting deficit in that country which is rapidly being parent,a few golden papillae at the tips of the lobes; redressed.Manycollectionsexist fromArgentinawhere corolla white, somewhat thick and fleshy, 1.5-2 cm the plantis apparentlyquite common. The populations diam.,lobednearlyto thebase,the lobes planarat anthein Misiones and Paraguayare discontinuouswith the sis (?), minutelypapilloseat the tips andmarginsof the & Hierolnv1us 1070 (F, US); Tafi Viejo, 500 m, 20 Feb 1941, Meyer 3912 (F, NY); Tafi, Simabori, Quebrada Grande, 31 May 1947, Melyer12555 (TEX); Chicligasta, Cochuna, 4 Mar 1941, O'Donell 51 (F, US), O'Donell 85 (F, NY); Tafi, rd. from Acheral to Tafi del Valle, Jan 1944, O'Donell 1395 (F); Tafi, Aconquija, 635 m, 1 Apr 1945, Ortiz 5 (NY); Burruyaci, Ampahuasi "El Ojo," 23 Sep 1934, Peirano 9860 (US); Rio Chico, Escaba, between Rio Mora and Arroyo Chico, 600-800 m, 24 Nov 1952. Petersen & Hjerting 623 (MO, NY); Monteros, rd. fi-om Acheral to Tafi, 500 m, 24 Feb 1949, Petersen & Hjerting s.n. (MO); Naranjal, Yerba Buena, 500 m. 18 Sep 1921, Schreiter 9955 (US); Tafi, Yerba Buena, 700 m, 19 Jan 1919, Ventulri 157 (F, US); Famailla, Quebrada de Lules, 600 m, 14 Nov 1926, Venturi2447c

TAXONOMIC TREATMENT

FIG. 61. Sol

epee

129

S. Knapp (Reproduced with permission from Novon 1 126 fig 1 1991.)

FIG. 61. Solanumin tepuienise S. Knapp. (Reproduced

lobes;anthers4-5 x 1.5-2 mm, poricidalat the tips, the poresteardropshaped;free portionofthefilaments ca. 0.5 mm long, the filamenttubeca. 1 mm long, glabrous; ovaty glabrous;style glabrous,7-8 mm long, expanded apically; stigma bilobed, minutely papillose. Fruit a globose, greenberry,hardat maturity,ca. 1.5 cm diam.; fruitingpedicels deflexed and woody, 2-2.5 cm long, ca. 1 mm diam. at the base, 2.5 mm diam. at the apex. Seeds reddish-brown,flattened-reniform,4-4.5 x 2.53 mm, the margins incrassateand paler, the surfaces minutely pitted. Chromosomenumbernot known.

with permission

from Novon

1' 126, fig. 1. 1991.)

Posadas,Bonpland,"Almac6nFinlandesa,"Ekmnan 821 (holotype, S-n.v.). Fig. 62

Small shrubs 0.5-3 m tall; stems slender; young stems sparsely pubescent with minute hooked trichomes, ca. 0.5 mm long, the uni- or bicellular hook borneon a multicellularmound-likebase,thehookoften deciduous.Sympodialunitsdifoliate,usuallynot geminate. Leaves ovate, widest at or just proximal to the middle;uppersurfaceof the leaves coveredwith hooked trichomes 0.5-1 mm long, like those of the young stems, the hooks soon deciduous leaving only the Distribution (Fig. 60). Known only from the type multicellular mound-like bases, which appearto the collection from the top of Sororopan-tepui, a low- naked eye as minute white specks on the upper leaf elevation sandstone table mountain near the Gran surface;lower surfacewith minutehooked trichomes, less than 0.5 mm long, on the veins and in small deSabanaregion of Venezuela. on the lamina;both surfacesscabrous;major pressions Solanum tepuiense is relatedto S. nudum,a wideleaves6.5-14 x 1.9-6.5 cm, theapexacuminate,thebase of Central and northern South America. spreadspecies It differs from S. nudum in its larger flowers and acute,decurrenton thepetiole;petioles0.6-1.4 cm long; minor leaves 2.9-7 x 1.5-4 cm, the apex acuminate, stronglywinged stems. The intemodes in the type colthe base attenuate;petioles 0.9-1.4 cm long. Infloreslection are very shortandareperhapsa distinctivefeatureof S. tepuiense,but may insteadbe relatedto habi- cences opposite the leaves, simple, 0.5-3.5 cm long, tatandthusnot consistent.The trichomesofS. tepuiense few-flowered, sparsely pubescent with uniseriatetriare quite different from those of S. nudum in being chomes less than0.5 mm long;pedicel scars unevenly composed of many small cells. The cells appearbead- spaced 0.5-1 mm apart. Buds globose, glabrous. like and are generally squarein shape. The trichomes Pedicels at anthesis 0.8-1 cm long, taperingfrom the ofS. nudumarecomposedof two or threeelongatecells calyx tube to a slenderbase ca. 0.5 mm diam.Flowers with the calyx tube very short,less than 1 mm long, the and are often branched. lobes deltoid,ca. 2 mm long, the tips tuftedwith minute uniseriatetrichomes;corollawhite,1.6-2 cm across,lobed 27. Solanum trachytrichium Bitter,Repert.Spec.Nov. 2/3 of the way to the base, the lobes planar or camRegniVeg. 16: 10. 1919.Type.Argentina.Misiones: panulateat anthesis,the tips of the lobes minutelypap-

130

FLORA NEOTROPICA

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131

TAXONOMIC TREATMENT

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Estrada Dona Francisca, 500 m, 23 Aug 1957, Reitz & Klein 4034 (K, NY, US), 26 May 1957, Reitz & Klein 4221 (US); Capinzal, 500 m, 21 Dec 1962, Reitz & Klein 14356 (US); Itapiranga,Barra do Macaco Branco, 250 m, ovary glabrous; style 6-7 mm long, straight; stigma not 1 Jan 1964, Reitz & Klein 16779 (US); Itapiranga, Rio markedlylargerthanthestyle,minutelypapillose.Fruita Peperiguacu, Linha Caqueiro, 200-300 m, 27?07'S, globose, greenor greenish-yellowberry,slightlyapicu- 53?47'W, 17 Oct 1964, Smith & Reitz 12669 (US); Rio latefromthepersistentstylebase,0.8-1.3 cm diam.firuit- Uraguai, 4-8 km W of Itapiranga, 150-200 m, 27?00'S, ingpedicelsslightlyexpandedapically,woody,deflexed, 53?45'W, 18 Oct 1964, Smith & Reitz 12683 (US); Rio ca. 2 cm long;calyx lobes slightlyaccrescentin fruit,ca. Uraguai, 6 km W of Itapiranga, 150-200 m, 27?10'S, 2 mm long.Seedspaleyellow or tan,flattened-reniform, 53045'W, 11 Nov 1964, Smith & Klein 13168 (US); Sao ca. 2 x 1.5 mm, the margins incrassate, the surfaces Miguel d'Oeste, above Rio Peperi-guacu, Peperi, 300600 m, 26?32'S, 53?44'W, 13 Nov 1964, Smith & Klein minutely pitted. Chromosomenumbernot known. 13241 (US); gorge of Rio Irani, 12 km S of Fachinal Distribution (Fig. 63). In southeastern Brazil dos Guedes, 500-600 m, 26?57'S, 52?12'W, 9 Dec 1964, (Paranaand Santa Catarina),northeasternArgentina, Smith & Klein 13898 (US); Foz de Iguacu, falls of Iguacu, and Paraguayin secondaryforests and areaswith high 25?42'S, 54?26'W, 10-11 Feb 1965, Smith 15019 (NY, US); s.loc., Tweedie s.n. (K). light intensity, 100-600 m. PARAGUAY. ALTO PARANA:Escuela Tecnica ForSpecimens examined. BRAZIL. PARANA:Rio Ivahy estal, Puerto Presidente Strossner, km 12, 250 m, 27 Jan nearColonia Therezina,22 Jan 1911,Dusen 11143(F, 1982, Fernandez Casas & Molero 5653 (NY); Forestry Technical School. 25 30'S, 54 40'W, 10 Sep 1983, Hahn NY, S-n.v.); Fenix, Irapos, 3 Jun 1963, Hatschbach 10152 (US); forest near Vista Alegre, ca. 15 km N of 1662 (PY). CANENDIYU:Guadalupe, ca. 30 km NW Francisco Beltrao, 15 May 1966, Lindeman & de Haas of Puerto Adela, 360 m, 17 Dec 1982, Schinini 23169 1370 (U); forest along Rio Iguacu, 600 m, 14 Mar 1967, (NY). ITAPUA:Pirap6, CEDEFO, 26 30'S, 56 50'W, 4 Lindeman & de Haas 4889 (U). RIo GRANDEDO SUL: Aug 1984, Hahn & Perez de Molas 2737 (PY). ARGENTINA. MISIONES:San Javier, Acaragua, 1 Schwabenschreib, 25 May 1949, Rambo 41684 (BH); Porto Alegre, Avenida Ernesto Fantoura, Sep 1898, Jul 1946, Bertoni 3124 (TEX); Iguazi, Parque Nacional Reineck & Czermak 604 (P); around city of Irai, 23 Jul Iguaz6, 26 Jul 1976, Cabral 68 (F); Ruta 12, 5 km N of 1960, Torgo 67 (US). SANTACATARINA:Morro do Rio El Dorado, 30 Jun 1986, Ferrucci et al. 451 (NY); Vermelho, 100 m, 27 Jun 1968, Klein & Bresolin 7754 Candelaria, Loreto, Jesuit mines, 19 Jan 1972, Krapovickas & Mroginski 20739 (MO); Iguazi, trail to Isla (US); Tapera, Ribeirao, 250 m, 18 Aug 1970, Klein 8718 San Martin, 9 Dec 1945, Lourteig 1132 (NY, P); lot 106, (US); Itapiranga, 3 Feb 1951, Reitz 3826 (US); Joinvile,

illose; anthersca. 4 x 1.5 mm, poricidalat the tips, the poresteardropshaped;free portionof the filamenttube less than0.5 mm long, the filamenttubeca. 1 mm long;

132

FLORA NEOTROPICA

12 km N of LeandroN. Alem, 18 Aug 1972, Marunak majorones only in size, 3-8 x 1-3.5 cm, the apex acute, 377 (MPU, WIS); Candelaria,SantaAna, 6 Dec 1945, the base acuteto attenuate;petioles 0.5-1 cm long.InMontes1532(DS);Obera,near"PosoHondo,"24 Jul 1970, florescences oppositethe leaves or terminal,oftenoverMroginski73 (US);Obera,14 kmN of CampoRam6n,27 topping the new leaves near the shoot apex, and later Dec 1973,Pire & Mroginski305 (WIS);Establecimiento lateraland leaf-opposed, simple, 3-8 cm long, 10--30OroVerde, SantaInes,Jul 1926,Scalas.n. (NY);Cainguas, 2-3 at a time,glabrousandshiny; 10779 San ruta 2 Sep 1950,Schwarz 12, flowered,bearingonly (TEX); Ignacio, scars pedicel irregularlyspaced 0.5-2 mm apart,bekm 100, 12 Sep 1950,Schwarz10874 (RSA/POM);San 2 cm from the base of the inflorescence. ca. Pedro,ColoniaMonteCarlo,21 Feb 1949,Schwindt1248 ginning Buds when 5 1792 San Jun Schwindt globose young, laterellipsoid with the exPedro, 1949, (TEX); Laharrague, 1 Jun 1949, sertionof the corolla,the corollasoon exserted.Pedicels (RSA/POM);San Pedro,PuertoLaharrague, Schwindt1827 (BH); Iguazu,El Dorado, 13 Oct 1949, at anthesis0.7-1 cm long, taperingfromthe calyx tube Schwindt2814 (RSA/POM);Cainguas,SaltoEncantador, to a slender base ca. 0.4 mm diam. Flowers with the 12Jul 1950,Schwindt 4851(BH);SanIgnacio,3 Mar1914, calyx tube cup-shaped, 1-1.5 mm long, sparsely puVattuone& Bianchi 166 (US). bescentwithuniseriatetrichomesca. 0.5 mm long, these Local names. Brazil.SantaCatarina:canemamirim. often somewhat reddish, the lobes quadratewith terminalcusps,thecuspsdenselypubescentwithuniseriate Solanum trachytrichium is clearly closely related to trichomesca. 0.1 mm long,thelobeswitha few uniseriate S. cassioides, also of southeastern Brazil. S. trachytrichomes ca. 0.5 mm long; corolla white, 0.6-1 cm trichiumis easily distinguishedfromthatspecies by its diam., lobed 3/4 of the way to the base, the lobes relarger,campanulateflowers,shorterinflorescences,and flexed at anthesis, the tips and margins of the lobes scabrousleaves. The scabrousleaf surfacesmake this anthers2.5-3 x 1 mm, poricidalat minutely papillose; one of the easiest species in sect. Geminatato identify the tips, the pores teardropshaped;free portionof the using vegetative characters.The trichomesareunusual 0.1-0.5 mm long, the filament tube ca. 0.5 filaments in the section in having a unicellularhook on a moundmm long;ovaryglabrous;style straight,4.5-6 mm long; likemulticellular base.Indrymaterial,theentiretrichome the surfaceminutely is transparent,andthe bases appearas white specks on stigma capitate,stronglybilobed, Fruit a globose, green berry,woody in dry papillose. the upperleaf surfaces. Their structurewas so differ7-9 mm diam.;fruiting pedicels deflexed ent fromthatof othersolanumtrichomesthatBitterde- specimens, and woody, 1.5-2 cm long, ca. 0.75 mm diam. at the scribed subsect. Silicosolanum, containing only S. Thetrichomesdo not containsilica,but base. Seeds dark brown in dry specimens, flattenedtrachytrichium. reniform,2-3 x 1-1.5 mm, the marginsincrassateand are rough to the touch, thus the name. numpaler, the surfaces deeply pitted. Chronmosome The lectotype of Solanum trachytrichium chosen her not known. by Smith and Downs (1966), Dusen 11143, is superfluous as Bitter clearly designatedEknman821 as the Distribution (Fig. 63). In S Bolivia and NW Artype in his original descriptionof S. trachytrichium. gentinaat 750-1500 m on theeaster slopesof theAndes. 28. Solanum trichoneuron Lillo, Contr.Arb. Argent. 99. 1910. Type. Argentina. Tucuman:Potrerillo, Alpachiri,5 Jan 1910, Venturi410 (holotype,LILn.v.; isotype, A). Fig. 64 Solanumoblongzum of Griseb.,Abh. Kon. Ges. Wiss. Gottingen 24: 253. 1879. Not of Ruiz & Pav. Shrubs to trees, 2-10 m tall, 10-20 cm diam.; young

stems andleaves glabrousandshiny,minutelyreddishresinous; bark of older stems pale yellowish-white. Sympodial units difoliate, geminate. Leaves elliptic,

widest at the middle, glabrous above, with tufts of uniseriatetrichomesca. 0.5 mm long in the axils of the main lateralveins beneath,9-15 x 2.5-6 cm, with 1012pairsof mainlateralveins,theseprominentandsomewhat yellowish beneath,the apex acute,the base acute to attenuate,somewhatdecurrentonto the petiole; petioles 1-1.8 cm long; minor leaves differing from the

Specimens examined. BOLIVIA. SANTA CRUZ: Parque Nacional Amboro, near Cerro Bravo 10 km N of Comarapa, vic. of permanent plot, 2400-2500 m, 17?49'S, 64?32'W, 7-10 Apr 1994, Vargas C. et al. 3108 (BM, NY); Parque Nacional Amboro, Cerro Bravo, near confluence of Rio Alizar and Rio Amparo. 20 km NW of San Juan del Potrero, 200 m, 17?57'S, 64?24'W, 1014 Apr 1994, Vargas C. et al. 3138 (BM, NY). TARIJA:

O'Connor,rd. to Entre-Rios,90 km E of Tarija,1680 m, 10 May 1986, Fournet 686 (NY); Arce, 7.4 km SE of Emboroza on rd. to Bermejo, 1100 m, 22016'S, 64?30'W, 24 Apr 1983, Solomon 10056 (K, MO, NY); O'Connor, 91.4 km E of Tarija-Padcaya rd., on rd. to Entre Rios, 8.6 km NW of Entre Rios, 1600 m, 21?28'S, 64?13'W, 3 May 1983, Solomon 10431 (MO, NY, U); Arce, 2-3 hrs by trail from Sidras on trail to Tariquia, 900-100 m, 22?12'S, 64?32'W, 7 May 1983, Solomon 10561 (MO, NY). ARGENTINA. JUJUY:S.loc., 1913, Schuel 135 (W). SALTA: Rosario de la Frontera, Los Bafos, 900 m, 25 Jul 1929, Venturi 9323 (MO, NY, US). TUCUMAN:Rio

133

TAXONOMIC TREATMENT

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trchoneuron LilloBolivia FIG?. 64. Solanum :.' p_e ~'i .

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Vargas C. et al. 3108 (BM).

134

Cochunato Las Lenguas,Jul 1929,Schreiter9941 (US); Chicliogasta,PuenteRio Cochuna,11 Jul 1929,Schreiter 9943 (US);Tafi,quebrada de Lules,500 m, 12 Aug 1923, Schreiter9952 (US); Tafi, YerbaBuena,700 m, 12 Feb 1919, Venturi209 (US); Tafi, La Hoyada, 1500 m, 3 May 1922, Venturi1840 (F, US); Famailla,Quebrada de Lulo, 600 m, 4 Nov 1923, Venturi2447 (BM); Tafi, Tafi Viejo, 750 m, 18 Aug 1923, Venturi2464 (US); Burroyacu,CerroEl Nogalito, 1200 m, 12 Apr 1929, Venturi8818 (MO, US). Local names. Argentina, Tucuman: hedionilla grande. Solanumtrichoneuronis most similarto andprobably closely related to S. aphyodendronof northern South and Central America. It differs from S. aphyodendronin its longer,often more terminalinflorescences, smaller flowers, quadratecalyx lobes, and smallerberries.Solanumtrichoneuroncan grow to be a large tree in Tucuman,reaching 20 cm diam. The wood, however, is too soft to be of any use commercially. Few collections of S. trichoneuronfromBolivia exist, but the species is common in northwestemmost Argentina. The ranges of S. aphyodendron and S. trichoneuronnearlyabut,andthe species arevery similar.They areperhapsallopatricorparapatricderivatives of a single ancestraltaxon. 29. Solanum warmingii Hiem, Vidensk. Meddel. Dansk Naturhist. Foren. Kjobenhavn 44. 18771878. Type. Brazil. Minas Gerais: Lagoa Santa, Warming 8 (holotype, C-n.v. [F neg. 22929]; isotypes, K, NY, P). Fig. 65 Shrubs(occasionallyquitelax) or smalltrees 1.5-8 m tall; stems glabrous, zigzag and two-winged from the bases of the decurrentleaves, these ridges prominent between nodes; bark pale greenish-yellow, not lenticellate.Sympodialunitsdifoliate,geminate.Leaves ovate to narrowly ovate, glabrous or with a few uniseriatesimple trichomesin the axils of the main lateral veins beneath, these 0.6-1 mm long, geminate, widest just proximal to the middle, slightly bullate above, the marginsrevolute;majorleaves 16.5-20.5 x 5-6.4 cm, with 8-11 pairs of main lateralveins raised andpale yellow on the uppersurface,all venationyellow andprominentbeneath,theapexacuminate,thebase attenuate,decurrenton the petiole and stem; petioles 0.8-1 cm long, narrowly winged from the decurrent leaf base; minor leaves not differing from the majors except in size, on lower stems nearly equal in size to the majors, 3.4-14.5 x 1.8-4.5 cm, the apex acuminate, the base attenuate,decurrenton the petiole; petioles 4-8 mm long, narrowlywinged fromthe leaf base. Inflorescencesoppositethe leaves, occasionallyfurcate,

FLORA NEOTROPICA

2.5-8 cm long, 5-40 flowered,glabrous;pedicelscars evenly spaced ca. 1 mm apartbeginning 1.1-1.2 cm from the base of the inflorescence. Buds ovoid, the corolla not exserted from the calyx until just before anthesis.Pedicels at anthesis 1.45-1.5 cm long, tapering from the calyx tube to a slender base ca. 0.5 mm diam. Flowers with the calyx tube ca. 3.5 mm long, lobes triangular,ca. 5.5 mm long, glabrousor minutely papillose on the tips; corolla white, ca. 2.4 cm diam., lobed 3/4 of the way to the base, the lobes planarat anthesis,tipsof the lobes minutelypubescentwithwhite papillaeanda few uniseriatetrichomes;anthersca. 4 x 1.5 mm, poricidalat the tips, the poresteardropshaped; free portionof thefilaments ca. 0.1 mm long, the filamenttubeca. 1 mm long;ovaryglabrous;style straight, 0.9-1.2 cm long;stigmaclavate,decurrentandminutely papillose. Fruit a globose, whitish-greenberry, 1.6-2 cm diam., appearingorange in dry material;fruiting pedicels woody, deflexed, 2.3-2.5 cm long, 1-1.5 mm diam. at the base, expandedat the apex, and appearing somewhatwinged in dry material,3-4 mm diam.; calyx lobes accrescentin fruit,0.75-1 cm long.Seedslight brown, flattened-reniform,ca. 4 x 2.5 mm, the margins incrassateand tan, the surfaces minutely pitted. Chromosomenumbernot known. Distribution(Fig.63). Foundonlyin SEBrazil,states of Minas Geraisandpossibly adjacentRio de Janeiro, in primaryand secondaryforest at 650-1200 m. Specimensexamined.BRAZIL.S.loc., 1816-1821,St. Hilaire 38, 578 nr 5 (P). MINASGERAIS:Vicosa, 650 m, 27 Feb 1924, Bailey & Bailey 1000 (BH); Hwy. BR-262, 5 km from Realeza, 6 Apr 1984, Hatschbach 47677 (NY);

PonteNova,FazendaVirginha12 km E of PonteNova, 7 Dec 1958, Irwin 2229 (F, NY, TEX, US); Vicosa, agri-

culturalcollege lands,680 m, 10 Jan 1930,Mexia4217 (MO, US), 5 Mar 1930, Mexia 4429a (CAS, NY, US); Vicosa, Fazenda do Paraiso, 730 m, 21 Nov 1930, Mexia 5337a (US); Vicosa, Fazenda de Aguada, 750 m, 27 Nov

1930, Mexia5368a (MO,NY, US); Vicosa, Fazendado Bom Jardim,rd. to Sao Domingo,740 m, 9 Dec 1930, Mexia5392 (CAS, MO, NY, US); Matade Silvicultura, ESF,UCV,Vicosa,9 Nov 1976,Rodrigues990 (RB);s.loc.. 1816-21, St. Hilaire catal. B1 748 (P); Nova Lima, Serra do Curral, Ribeirao da Mutuca, 1200-1350 m, Williams & Assis 6754 (F, MO, NY, US). Rio DE JANEIRO:S.loc., 1816-21, St. Hilaire catal. AI ler partie 565 (P).

Local names. Brazil.MinasGerais:lapavermelha. Solanum warmingii is related to S. caavurana. It shares with S. caavurana petaloid calyx lobes, large elliptic buds, and fleshy flowers. Solanumwarmingii is easily distinguishedfromS. caavuranaby its nearly glabrous leaves that dry pale sea-green or yellowish, its longerfruitingpedicels thatareslightly expandedat the apex, and its winged stem. The large flowers and

TAXONOMIC TREATMENT

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FLORA NEOTROPICA

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petaloidcalyx lobes of S. warmingiiaredistinctiveand make this species easy to identify.Solanumwarmingii is a species of limiteddistribution,andvery few collections exist. The habitatof S. warmingii is cerradoor campolimpo,bothtypesof savannahvegetation.It often grows as a small tree at the edge of pastures.

IV. Solanum leucocarpon species group (S. S. corumbense, S. leucocarpon, S. alatiranmeum, lindenii, S. oblongum). Fig. 66

some populations,the apex andbase various.Inflorescences opposite the leaves or occasionally somewhat intemodal(Solanumalatirameum),simpleorbranched, glabrousor pubescentlike the stemsandleaves:pedicel scars closely andregularlyspaced.Buds ellipsoid, the corolla stronglyexsertedfromthe calyx tube.Flowers white, fleshy,andrelativelylarge,the corollalobes planarat anthesis.Fruit greenor yellowish-greenat maturity;fruitingpedicelserector slightlydeflexed,enlarged just below the berry,woody. Seeds flattened-reniform, the marginsincrassate,paler,reddish-brownto yellowish-tan.

Shrubsor small trees;young stems and leaves glaDistribution. Secondaryhabitats,SouthAmerica, brous or pubescentwith simple, uniseriatetrichomes. Sympodialunitsdifoliate andgeminateor plurifoliate, with a few collections from Panama. The enlargedswellingjust below the berry(see Fig. often anisophyllous. Leaves elliptic to occasionally or with ovate, glabrous variously pubescent simple, 66B) is characteristic of members of the Solanmin uniseriate trichomes, these occasionally dendritic in leucocarpon species group.

Key to the species of the Solanum leucocarpon species group 1. Leaves glabrous, with no visible trichomes on the upper or under surfaces. 2. Leaves very large, 16-35 x 10-14 cm; leaf base attenuate, the leaves sessile; stems winged. SE Brazil .....................................................................................................................................30 . S. alatirame 2. Leaves smaller; leaf base acute to cuneate, the leaves never sessile; stem not winged. 3. Buds and new growth resinous; fruiting pedicels erect. Peru and Bolivia ...........................33. S. lindenii 3. Buds and new growth not resinous, usually glabrous or occasionally sparsely pubescent; fruiting pedicels somewhat deflexed. Isla de Coiba, Panama; Rio Negro basin, Brazil ............ 32. S. leucocarpon 1. Leaves variously pubescent. 4. Trichomes covering the entire abaxial leaf surface. 5. Trichomes mostly dendritic, golden; buds and new growth resinous. Peru and Bolivia .......33. S. lindenii 5. Trichomes mostly simple and uniseriate, some dendritic;buds and new growth sparsely to densely pubescent, not resinous. Bolivia, Brazil, and Paraguay ................................................. 31. S. corumbense 4. Trichomes not covering the entire abaxial leaf surface, usually distributed along the midrib and in the axils of the main lateral veins. 6. Ovary and berry pubescent; abaxial leaf trichomes concentrated along the midrib. Central Peru ............................................................................................................................................ 34 . . o lo ngrn 6. Ovary and berry glabrous; abaxial leaf trichomes in the axils of the main lateral veins and along the midrib. 7. Buds and new growth resinous; fruiting pedicel erect. Peru and Bolivia...................... 33. S. lindenii 7. Buds and new growth not resinous, variously pubescent or glabrous; fruiting pedicels erect or deflexed. 8. Leaf base attenuate;corolla 1.5-1.6 cm diam.; fruiting pedicel 1.5-2 cm long. Santa Cruz, B o liv ia ........................................................................................................................3 1. S. corumbense 8. Leaf base acute to cuneate; corolla usually > 1.5 cm diam.; fruiting pedicel 1-1.5 cm (longer in Isla de Coiba populations only). Widespread..................................32. S. leucocarpon

30. Solanum alatirameum Bitter,Repert.Spec. Nov. Regni Veg. 16: 88. 1919. Type. Brazil. Parana: Carvalho,5 Nov 1911, Dusen 13299 (holotype, Sn.v.; isotypes, F [US neg. 5920], GH, NY).

just abovethemiddle,whenyoung coveredwith minute red glandularpapillae,laterglabrousabaxiallyandadaxially;majorleaves 16-35 x 4-10 cm, with 7-10 pairs of main lateral veins, these drying reddish-brown Shrubs 1-3 m tall; stems glabrous,thick and some- abaxially,the apex acute,the base attenuateandthe leaf what fleshy, erect, stronglywinged fromthe decurrent sessile and decurrentonto the stem; minor leaves difleaf bases, the wings to 5 mm;barkof older stems dark fering fromthe majorsin size only, 11-20 x 2.5-4 cm. reddish-brown.Sympodialunitsplurifoliateor difoliate Inflorescences opposite the leaves or occasionally and geminate.Leaves elliptic to obovate, widest at or somewhatintemodal,3.5-12 cm long, oftenmany-times

TAXONOMIC TREATMENT

137

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FIG. 66. Solanum leucocarpon species group. A. S. leucocarpon flowers, Ecuador (Knapp & Mallet 6744). B. S. leucocarpon fruit, Venezuela (Knapp & Mallet 6836). C. S. oblongum habit, Peru (Knapp & Mallet 6637). D. S. oblongum flowers and young fruit, Peru (Knapp & Mallet 6637).

branched,dryingdark,minutelyred-papillosenearthe tip, with 10-20 flowers;pedicelscars regularlyspaced ca. 3 mm apart.Buds ellipsoid, the corolla stronglyexsertedfromthe calyx tube.Pedicels at anthesiserectto somewhatdeflexed, 1.4-1.7 cm long, ca. 1 mm diam.at the apex, ca. 0.5 mm diam.atthe base.Flowerswith the calyxtubeconical, 1-1.5 mm long,the lobesmereapiculateprojectionsfromthe rim,ca. 0.5 mm long, minutely papilloseat the tips;corolla white or cream,1.2-1.5 cm diam., lobed nearly to the base, the lobes planar(?) at anthesis, the tips of the lobes cucullate and densely papillate;anthers3-4.5 x 1-1.5 mm,poricidalatthetips, the pores lengthening to slits with age; free portionof

thefilaments0.5-1 mm long, the filamenttube ca. 0.25 mm long, glabrous;ovaryglabrous;style 6-8 mm long, somewhatclavate;stigmabilobed,the surfaceminutely papillate.Fruit a globose berry,greenor whitish-green; fruitingpedicels woody, deflexed, 1.5-2 cm long, ca. 1 mm diam.at the base, ca. 3 mm diam.at the apex.Seeds with incrassate, lightreddish-brown,flattened-reniform palermargins,2-2.5 x 1.5-2 mm, the surfacesminutely pitted. Chromosomenumbernot known. Distribution (Fig. 68). Known only from SE Brazil, in primaryforest,often in the lauraceousunderstory of Araucaria forests, at 800-1300 m.

FLORA NEOTROPICA

138 Specimens examined. BRAZIL. PARANA: Quatro Barras, Rio do Corvo, 950 m, 7 Nov 1966, Hatschbach 15086 (NY, US); Piraquera, Novo Tirol, 30 Oct 1967, Hatschbach 17674 (F, UPCB); San Jose dos Pinhaes, Rio Pequeno, 4 May 1972, Hatschbach 29644 (NY). RIO GRANDE DO SUL: S.F. Paula, Rincao dos Kroeff, 800 m, 10 Jan 1964, Sehnem8263 (US). SANTACATARINA: S. Jose, Serrada Boa Vista, ca. 1300 m, 2 Feb 1953, Reitz 5408 (US).

yellowish abaxially, the apex acute to acuminate,the base attenuate,slightlywinged onto the petiole;petiole 0.6-2 cm long; minorleaves differingfromthe majors only in size, 2.5-8 x 1-4 cm, the apex acute to acuminate,the base attenuate;petioles 0.5-1 cm long, winged along their entire length with the decurrentleaf base. Inflorescences opposite the leaves or occasionally intemodal,simple, 5-10-flowered, glabrousor with scatteredsimple,uniseriatetrichomesca. 1 mm long;pedicel Solanum alatirameum is an unusual species in the scars evenly spaced ca. 2 mm apart.Buds ellipsoid, group due to its sessile leaves and apparently terminal stronglyexsertedfromthecalyx tube.Pedicelsat antheinflorescence, but it is probably best placed here until sis deflexed,slender,0.9-1.3 cm long,ca. 0.25 mmdiam., furtherstudies can be done. The calyx with its minute abruptlywideningto the swollen calyx, glabrous.Flowapiculatelobes is similarto thatof S. leucocarpon,as is ers sweetly fragrant(fideNee),withthecalyxtubeswoltheswollenpedicelapexin fruit.SmithandDowns(1966) len, broadlyurceolate,3-4 mm long, the lobes minute, placedS. alatirameumin sect. Anthoresisbased on the deltatewith prominentapicalprojections,0.25-0.5 mm terminalinflorescence.Otherspecimens,however,have long, with tuftsof uniseriatetrichomeson the apiculae; lateraland leaf opposed inflorescences, so this is an- corolla white, 1.5-1.6 cm diam., lobed ca. 3/4 of the othercase of the variabilityin inflorescenceposition in the way to the base, the lobes planarat anthesis,papilspecies of sect. Geminataobscuringtheir affinities. lose at the tips and marginsof the lobes; anthers 4.55 x 1-1.5 mm, poricidalat the tips, the pores lengthento slits with age; free portion of thefilaments ca. 31. Solanum corumbense S. Moore, Trans. Linn. ing 0.5 mm long, the filamenttubeca. 1 mm long, glabrous; Soc., ser. 2, 4: 404. 1895. Type. Brazil. Mato 6-8 mm long; stigma Grosso: Corumba, Jan 1892, Moore 974 (holo- ovary glabrous;style straight, the surfaceminutelypapillose.Fruita globose, capitate, Fig. 67 type, BM; isotype, K-n.v.). green berry,1-2.3 cm diam.;fruitingpedicels woody, Cyphomandra verruculosa Hassl., Repert. Spec. deflexed, 1.2-2 cm long, 0.5-1 mm diam. at the base, Nov. Regni Veg. 9: 118. 1911. Type. Paraguay. the calyxbeneaththefruitstronglyswollenandenlarged. Concepci6n:Zwischen Rio Apa and Rio Aquidaban, Seeds yellowish-tan,flattened-reniform,3.5-4 x 2-2.5 San Luis, Jan 1908/1909, Fiebrig 4461 (lectotype, the surfacesminutelypitted,the marginincrassate mm, BM, here designated; isolectotypes, G-n.v., K). and darker.Chromosomenumbernot known. verruculosum Solanumn (Hassl.) Hassl., Repert. Spec. Nov. Regni Veg. 15: 218. 1918. Type. Based on Cyphomandra verruculosa Hassl. Solanum tumescens S. Knapp, Brittonia 44: 64. 1992. Type. Bolivia. Santa Cruz: Jardin Botanico de Santa Cruz, 12 km E of center of Santa Cruz on rd. to Cotoca, 375 m, 17046'S, 63004'W, 19 Jan 1989, Nee 37620 (holotype, NY; isotypes, ADn.v., CORD-n.v., G-n.v., IBE, Jardin Botanico de Santa Cruz-n.v., K, LPB-n.v., MO, MPU, Pn.v., RSA-n.v., TEX, US, USZ-n.v.).

Shrubs or treelets, 4-5 m tall; young stems glabrous or densely pubescent with small uniseriate trichomes, the young leaves densely red-papillose; bark of older stems pale greenish-yellow, on large stems reddish with large white lenticels. Sympodial units difoliate, geminate. Leaves elliptic to ovate, widest at or just below

the middle, glabrous or sparsely pubescent adaxially, pubescentabaxiallywith tufts of simple, uniseriatetrichomes in the axils of the main lateralveins, these often extendingto theentirelamina,the laminawith dense deposits of crystal sand visible as tiny white specks with a dissecting microscope;majorleaves 7-13 x 36 cm, with 7-9 pairsof main lateralveins, these drying

Distribution (Fig. 68). In subtropicalsemi-deciduous forest,usually in disturbedsituations,Bolivia and adjacentBrazil and Paraguay,300-400 m. Specimens examined. BOLIVIA. LA PAZ: Prov. Inquisivi, canyon of Rio Khatu,2 km by air N of Inquisivi on rd. to Chorocona, 2100-2200 m, 16?53'S, 67?08'W, 12 Mar 1988, Nee & Solomon 36658 (BM, NY). SANTA CRUZ:Prov. Andr6s Ibfaiez, 12 km E of center of Santa Cruz, on rd. to Cotoca, 375 m, 17?46'S, 63?04'W, 21 Jan 1987, Nee 33716, 33723 (NY); Prov. Cordillera, ca. 21 km SE of Palmar del Oratorio, ca. 14 km SE of Rio Chore-Chore (=Rio Pantano), 365 m, 18?02'S, 63?01'W, 22 Jan 1989, Nee 37646 (IBE, K, NY); 1 km SW of Buenavista, 360 m, 17?28'S, 63?40'W, 5 Jun 1991, Nee 40830 (K); Prov. Ichilo, 4 km (by air) SW of Buenavista, Parque Nacional Ambor6, S side of Rio Surutf, 320 m, 17?29'S, 63?41'W,3 Dec 1991, Nee 41859 (BM, NY); Prov. Florida, vie. Puente Las Cruces, along hwy. from Santa Cruz to Samaipata, 6 km W (by air) of Bermejo, 1000 m, 18?08'S, 63041'W, 21 Dec 1991, Nee 42238 (BM, NY); 3 km SW of Angostura, N side of canyon of Rio Pirai, 800-1000 m, 18?10'S, 63?33'W, 27 Dec 1992, Nee & Vargas C. 43346 (BM, NY); 6 km NW of Terebinto on rd. to El Hondo, 450 m, 17?41'S,

TAXONOMIC TREATMENT

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FLORA NEOTROPICA

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63?25'15"W, 15 Jan 1994, Nee 44478 (BM, NY); 7 km by air N of Pampa Grande,just E of Las Juntas on main hwy. from Samaipata to Comarapa, 1240 m, 18?02'S, 64?06'15"W, 28 Jan 1994, Nee & Vargas C. 44705 (BM, NY); 2 km by air NW of center of Bermejo, around Laguna Volcan, 1125-1175 m, 18?07'S, 63?39'W, 24 Dec 1994, Nee 46140 (BM, NY); Bosque de Rio Palmetillas, 400 m, 18 Mar 1917, Steinbach 3265 (K); Rio Piray, 400 m, 27 Feb 1925, Steinbach 7348, 7348a (BM, K); Buenavista, 450 m, 29 Dec 1926, Steinbach 7664 (BM, K); El Estanque, 40 km due S of Vallegrande, 1400 m, 18?56'S, 64?85'W, 24-27 Dec 1990, Vargas C. 843 (BM, NY); San Buenaventura, 1400 ft, 20 Nov 1901, Williams 652 (K). BRAZIL. MATTOGROSSO:Corumba, 18 Dec 1902, Robert 733 (BM, K). MATOGROSSODO SUL: Ca. 15 km from Corumba, 29 Jan 1991, Ratter et al. R6511 (K). PARAGUAY.

CONCEPCI6N: 8.3 km NE of Loreto,

rd. to Paso Barreto, 255 m, 17 Dec 1983, Vanni et al. 391 (PY); between Rio Apa and Rio Aquidaban, Caballero-Cue, Feb 1908/1909, Fiebrig 4726 (BM, G-n.v., K); between Rio Apa and Rio Aquidaban, Fiebrig 4824 (BM, G-n.v., K). Solanum corumbense is closely related to S. leucocarpon, a widespread species of Central and South America. Characters in common are the swollen calyx in flower and fruit and apiculate calyx lobes. When I originally described S. tumescens it was from largely glabrous Bolivian specimens. Since that time many specimens of intermediate pubescence have been collected (e.g., Nee & Solomon 36658) and it is clear the in S. Pubescence two taxa are synonymous.

corumbense is extremely variable across the species range, as it is in S. leucocarpon (see above), and may to some extent be related to the dryness of the habitat. As with many of the species in this group, S. corumbense is superficially similar to species of Solanum sect. Pachyphylla (Cyphomandra, also see Bohs, 1994). The close affinities of this group may lie with either sect. Pachyphylla or with sect. Cyphomandropsis, but this idea remains to be tested.

32. Solanum leucocarpon Dunal in Poir., Encycl., suppl. 3: 756. 1814. Type. probably French Guiana, ile Ste. Marthe, Richard s.n. (holotype, P). Figs. 66A,B, 69 Solanum surinamense Steud., Flora 26: 764. 1843. Type. Surinam. S.loc., Dec 1842, Hostmann 1271 (lectotype, G, here designated [Morton neg. 8646]; isolectotypes, K, MO, NY, P [Morton neg. 8345], U). Solanum coarense Sendtn. in Mart., Fl. Bras. 10: 19. 1846. Type. Brazil. Amazonas: In sylvis ad Coari, provinciae Rio Negro dictae, Nov, Martius s.n. (lectotype, M, here designated [F neg. 6527]; isolectotype, BR). Solanum triste Jacq. var. crassipes Dunal in DC., Prodr. 13(1): 148. 1852. Type. French Guiana. in Guiana gallica, s.coll. (holotype, G). Solanum patellare Van Heurck & Mull. Arg., Observ. Bot. 53. 1870. Type. Peru. San Martin: Prope Tarapoto, 1855-1856, Spruce 4914 (isotypes,

TAXONOMIC TREATMENT

141

:

*L~YI~ :y~rk.:

7:,._..

SICIOMEN

iJeflPL

'F. W. IO4TMNS;

.. . .... 18... .r

: ...,

SURIALam

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=

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=.::=: .,.

.,

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FIG.

Steud.).

69

Solanum

leucocarpon

Dunal,

.."i .

...

Surinam

..

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Hostmann

& Kappler

1471

(BM-isotype

of

. surinamense

142

FLORA NEOTROPICA

AWH,BM, C-n.v. [F neg. 22904], F, G [Morton ing frombelow the calyx margin,coveredwith the same neg. 8545], K, MPU,NY, P [Mortonneg. 8284]). as therestof theinflorescence;corollawhite Solanum brevipedunculatum Rusby, Bull. New York pubescence and waxy, occasionally with a purplishtinge, 1.5-2.8 Bot. Gard.4: 421. 1907. Type. Bolivia. S.loc., cm lobed nearly to the base, the lobes planarat diam., Bang 2525 (holotype, NY; isotypes, BM, F, K, sinusesmembranous,the lobes anthesis,the interpetalar NY, US, WIS). Solanumcoibae Riley, Kew Bull. 1927: 125. 1927. narrowly triangular, papillose to pubescent with Type. Panama. Veraguas: Coiba Island, Sep, uniseriate trichomes 0.5 mm long along the margins Collenette in Riley 473 (holotype, K; isotype, US). and at the tips; anthers orange or deep orangish-yelSolanum eshbaughianum D'Arcy, Ann. Missouri low, 3.5-6 mm long, the terminal0.5-1 mm paler and Bot. Gard.60: 745. 1974 [1973]. Type.Panama. thickened, 1-1.2 mm wide, tightly conniventat antheDarien:Vic. of La Palma,0-50 m, Pittier 6695 sis, poricidalat the tips, the poresteardropshaped;free (holotype, US). portionof thefilaments ca. 0.5 mm long, the filament Solanum extraxillare Klotzsch in R. H. Schomb., FlorafindFaunavon BritischGuiana1154. 1848. tube 0.2-0.5 mm long; ovary glabrous;style straight, Nom. nud. Guyana(as BritishGuiana).Savanne, 7-9 mm long;stigmaclavate,minutelypapillose.Fruit Apr 1843, Schomburgk1263 (B [destroyed],P a globose berry,1-1.5 cm diam.,dirtyyellowish-green whenripe;fruitingpedicelswoody,deflexed, 1-1.5(2.5) [Mortonneg. 8190]). Solanum diphyllum of Pulle, Enum. Vasc. P1. cm long (longestin Isla de Coiba,Panamapopulations), Surinam409. 1906. Pro parte,not of L. ca. 1 mm diam. at the base, greatly expandedto ca. 5 Solanumfoetidumof Pulle, Enum.Vasc.P1.Surinam mm diam. about 5 mm from the apex. Seeds pale yel410. 1906. Not of Ruiz & Pav. lowish-tan,flattened-reniform,3-3.5 x 1.5-2 mm, the Foetid shrubs or small trees, 1-6 m tall; young marginsincrassate,the surfacesminutelypitted.Chrostems andleaves densely hispidulouswith minuteuni- mosomenumber:n = 12 (voucherKnapp6241, 6836). cellular or uniseriatetrichomesto 0.2 mm long; older Distribution (Fig. 70). Widely distributedthroughstemspartiallyglabrate;barkof olderstemsreddish-gray. outtropicalSouthAmericato Panama,in secondgrowth Sympodial units difoliate, geminate, anisophyllous. situationsfrom sea level to 1500 m. Leaves elliptic, widest at the middle, usually glabrous Selected specimens examined. PANAMA. VERAGUAS: above, but occasionally with minute uniseriate trichomes along the veins, pubescence of the leaf under- Isla de Coiba, 16 Aug 1961 Dwyer 1554 (MO, US). COLOMBIA. AMAZONAS:Rio Igara-Parana(afflusides extremely variable, from glabrous to densely ent Rio Putumayo) La Chorrera, land of Witoto-Jito with minuteuniseriatetrichomesalong the hispidulous 16 Jan 1974, Gasche & Desplats 148 (MO); 9 Maguro, to with to dense veins, densely pubescent sparse pu- km from Leticia on Leticia-Tarapara rd., 30 Jun-7 Jul bescence of uniseriate (occasionally branched) tri1964, Leticia 8 (Gutierrez V et al.) (US); Rio Igarachomes 0.5-1 mm long, along the majorveins, but not Parana, Bella Vista, 8 Sep 1973, Sastre 2163 (MO); on the veins themselves,these trichomesin some speci- Leticia, ca. 100 m, 20 Sep 1945, Schultes 6545 (K, US). mens extendingto the lamina;majorleaves 10-19 x 4- ANTIOQUIA: 10.1 km W of San Rafael on Guatap6-San 10.5 cm, with 6-10 pairs of main lateralveins, these Rafael rd., 1220 m, 6?18'N, 75?04'W, 24 Oct 1987, impressedabove,prominentandyellowishbeneath,the Brant & Roldin 1473 (K, MO); 16 km SW of Las apex acute, the base acute to cuneate, often oblique; Partidas de San Luis, on Medellin-Bogota rd., Vereda de Josefina, 800 m, 6?00'N, 74?50'W, 25 Jun 1987, petioles 1.2-3 cm long;minorleaves differingfromthe Callejas et al. 4209 (NY); VeredaManizales, 12 km from majorones in shape and size, the minorleaves orbicu- San Luis on San Luis-San Carlos rd., along Rio x late, 3.5--8 3-6.5 cm, the apex acute, the base Dormilon, 1440 m, 6?05'N, 75?00'W, 26 Jun 1987, rounded,occasionallysomewhatcordate;petioles 0.5- Callejas et al. 4289 (NY); Mulatos, ca. 50 m, 13 Jun 1 cm long. Inflorescencesopposite the leaves, simple, 1946, Haught 4883 (US); thermal springs of Santo 1-4 cm long, 5-15-flowered, occasionally glabrous, Domingo, ca. 1200 m, 7 May 1949, Scolnik et al. more often densely pubescent with minute uniseriate 19An519 (US); valle de Rio Anori, between Dos Bocas trichomes0.1-0.5 mm long;pedicel scars in pairs,the and Anori, along Rio Buenos Aires, 400-700 m, 26 Aprmembers of a pair closely spaced, the pairs 1-3 mm 3 May 1973 Soejarto et al. 3970 (F, MO); vic. of Medellin, 10 Sep 1927 Toro 612 (NY); 10 km from Pto. Valdivia apart.Buds globose when very young, the corolla soon exserted from the calyx, and the bud ellipsoid with a on rd. to Yarumal, Gutierrez et al. Ventanas2 (US); km 4.3 of Juanes-Correg. Alto Samana rd., 0.3 km past rd. pointed tip. Pedicels at anthesis 1.3-1.7 cm long, ta- to top of Puchina dam, 800 m, 6?11'N, 74?52'W, 20 May peringfromthe calyx tubeto a slenderbase ca. 0.5 mm 1988, Zarucchi et al. 6724 (MO, NY). BOYACA:NW of diam. Flowers with the calyx tube 1-3 mm long, gla- Bogota, region of Mt. Chapon, 3500 ft, 8 May 1932, brousor minutely pubescent,the lobes shallowly del- Lawrence 20 (F, K, NY, US). CUNDINAMARCA: From seed toid, 0.1-1 mm long, apiculate,the apiculaeoften aris- of Whalen 518, 2 km past Noracacia on rd. to Honda, ca.

143

TAXONOMIC TREATMENT

o0

~.

,-

?500-

FIG. 70. Distribution of Solanum leucocarpon (solid circles) and S. oblongum (open circles).

600 m, 11 Aug 1981, Knapp 889 (BH); S of La Palma along trail from La Palma to Finca "Alto Oscar," 11 km S, 4800-5000 ft, 10 Mar 1944, Little 7376 (NY, US); trail from Finca "Alto Oscar,"N of La Palma, 5000 ft, 13 Mar 1944, Little 7430 (NY, US). HUILA: Mosquera, Caqueta and vic., Rio Mosquera, 25 Apr 1944, Little 7718 (NY). META: Villavicencio, 20 Jan 1899, Sprague 107 (K). NARINO: Rd. between Nariio and Samana, 2 km from exit to La Dorada,Gutierrezet al. Narino I (US). SANTANDER: Between San Vicente de Chuchuri and El Carmen, 900-1000 m, 28 Feb 1964, Uribe U. 4697 (NY). VENEZUELA. AMAZONAS: Oromana, 750 m, 4?35'N, 64?40'W, 12 Jun 1976, Colchester 2502b (K); Rio Ocamo, among the Yanomani, 1978, Fuentes s.n. (MY); S edge of San Juan de Manapiare airport, Rio Manapiare, ca. 140 m, 5?18'N, 66?04'W, 31 Jan 1977, Huber 473 (VEN). BARINAS: 76 km from Barinas on rd. to San Crist6bal, Ticoporo Forest Reserve, 350 m, 10 May 1964, Breteler 3954 (F, MO, NY VEN); trail from Mesa de Canagua (ca. 8032'N, 70?39'W) to Alto de la Aguada (ca. 8?37'N, 70?40'W), ca. 23 km NW of Curbati, 800-1400 m, 17 Apr 1988, Dorr et al. 4752 (NY); SW of Alto de La Aguada (ca. 8?37'N, 70?40'W) in area known locally as "Montaias de la Palmita," 12001400 m, 19 Apr 1988, Dorr et al. 4882 (NY); outskirts of Barinitas, 31 Jul 1979, Nee 17091 (BH, F, VEN, WIS). BOLIVAR: Piar, 0-3 km SE of SSE corner of Amaruaytepui, 500-700 m, 5?55'N, 62?13'W, 9 May 1986, Liesner & Hoist 20715 (BM, MO); Tfriba, banks of river, ca. 80 m, 6 Jun 1984, L6pez-Palacios et al. 4385 (NY); Chimantf massif, W slopes of Chimantf-tepui (Toronotepui), 1100-1700 m, 30-31 May 1953, Steyermark 75592 (F, NY, VEN); Gran Sabana, Sto. Domingo-Sta.

Elena rd., 5 Mar 1946, Tamayo 3080 (F, US, VEN); La Prisi6n, Medio Caura, 100 m 18 Mar 1939, Williams 11541 (F, US, VEN). Coje des Cerro Azul, Fila de la Blanquera, 28 Feb 1979, Reyes L. & Aquilez G. 264 (VEN). TACHIRA: Las Dantas-Buena Vista-Puente Alianza rd., ca. 12 km S of Las Dantas, 1100-1200 m, 7?40'N, 72?25'W, 24 Oct 1984, Knapp & Mallet 6836, 6837 (BH, K, MY, US, VEN); vic. of Las Minas, N of La Laguna, 16 km SE of Santa Ana, 1150-1250 m, 7?36'N, 72013'W, 28 Jul 1979, Steyermark & Liesner 118906 (VEN). YARACUY:Sierra de Aroa, Cerro Tigre, 10 km by air E of Aroa, Rio Carabobo, 800-1000 m, 10?26'N, 68?49'W, 30 Mar 1980, Liesner & Gonzclez 9680 (VEN); San Felipe Aracal, 12 Apr 1949 Trujillo & Ferndndez 97 (MY). GUYANA. S.loc., 1837, Schomburgk 256 (G-DC, K, P); slopes of Mt. Makarapan,along MakarapanCreek, 250 m, 3?59'N, 58?57'W, 18 Sep 1988, Maas et al. 7518 (K, NY); Marudi Mtns., on top of Mazoa Hill near drilling area, 450 m, 2015'N, 59?10'W, 10 Nov 1982, Stoffers et al. 257 (K, NY, U). BERBICE:Along Berbice-Rupununi cattle trail, upper Ituni Creek, 4 Apr 1919 Abraham 9 Bartica, 31 Jul 1949, Fanshawe 2932 (NY). DEMERARA: (Forest Dept. B.G. 6079) (NY, U, US); Demerara River, Jun 1894, Jenman 6743 (K, NY); along rd. betwen Rockstone and Linden, 10-100 m, 5?59'N, 58?25'W, 23 Jan 1993, McDowell et al. 1810 (U, US); Comaka, Demerara River, May 1923, Persaud 242 (F); Comaka, Demerara River, May 1923, Persaud 242bis (F). ESSEQUIBO:Mazaruni forest station, 17 Aug 1934, Archer 2470 (US); Rockstone, 15 Jul-i Aug 1921, Gleason 588 (NY), Gleason 695 (NY, US); Blue Mountain, 2-5 km NW from shops on rd., 100-200 m, 6?35'N, 58?47'W,

144 18 Apr 1993, Henkel & Williams1934 (U); KamuniCreek, Groete Creek, Essequibo River, 19 Apr 1944, Maguire & Fanshawe 22900 (F, K, NY, U, US). MAZARUNI:Upper Mazaruni River region, vic. Kako, an Akawaio Indian village on Kako River near confluence with Mazaruni River, ca. 500 m, 5?45'N, 60?35'W, 14 Apr 1987, Boom & Gopaul 7290 (NY, U); upper Mazaruni River, 60010'W, 22 Sep-6 Oct 1922, de la Cruz 2092 (F, MO, NY, US), de la Cruz 2147 (BH, F, MO, NY, US); upper Mazaruni River, Kamakusa, 59?50'W, 23-29 Nov 1922 de la Cruz 2773 (F, MO, NY, US), de la Cruz 2872 (NY, US): Kaiteur Falls, Potaro River, 23 Oct -3 Nov 1923, de la Cruz 4428, 4496 (F, K, MO, NY, US); Kartabo Station, jct. of Mazaruni and Cuyuni Rivers, 9 Jul 1924, Graham 93 (US); Kartabo, 11 Dec 1919, Hitchcock 17203 (NY, US); Kalakoon, Mazaruni River, Nov 1886, Jenmanl 2405 (K); Kaburi rd., Bartica, 10 Dec 1923, Linder 47 (NY); along rd. from Ariching airstrip to Mazuruni River, 6?10'N, 60?07'W, 22 Feb 1991, McDowell 4048 (K, U); Cuyuni River, lower Camaria landing, 20 Nov 1929 Sandwith 629 (NY); upper Mazaruni basin, Kamarang River W bank above Utschi mouth, ca. 550 m, 23 Aug 1960, Tillett & Tillett 45772 (NY); Mazaruni station, 8 Jun 1933 Tutin 169 (U, US). NORTHWEST: Waini River, 8?20'N, 59?40'W, 3-18 Apr 1923, de la Cruz 3678 (F, MO, NY, US); WanamaRiver, 7?45'N, 60?15'W, 10-23 May 1923, de la Crutz3860 (F, MO, NY, US). POTARO-SIPARUNI: Upper slopes of Mt. Wokomung, 1540-1600 m, 5?05'N, 59?50'W, 11 Jul 1989, Boom & Samuels 9193 (NY, U); Essequibo River at Karapukari crossing, 60-75 m, 4?40'N, 58?41'W, 19 Apr 1992, Hoffinan et al. 1341 (K, U); Annai-Karupukari rd., 18 km N of Surama village cutoff, 0.1 km W of rd., 80-90 m, 4?14'N, 58?56'W, 29 Apr 1992, Hoffman & Pennington 1495 (U). POMEROON:Waranuiri mission, Moruka River, 23-27 Oct 1922, de la Cruz 2541, 2585 (F, MO, NY, US). RUPUNUNI:Wabewak, Kanuku Mtns., Nov 1948. Forest Dept. of British Guiana (WilsonBrowne) 440 (K, NY); Komo Creek, Takatu River, Dec 1948, Forest Dept. of British Guiana WB 574, 5964 (K, NY, US); Kanuku Mtns., Maipama, Camp 3 on Tsikoma Creek. 160 m, 3?22'N, 59?30'W, 23 Nov 1987, JansenJacobs et al. 1203 (K, NY, U); basin of Rupununi River, Isherton, 2?20'N, 9-15 Nov 1937, Smith 2495 (F, K, MO, NY, U, US); NW slopes of Kanaku Mtns., Moku-Moku Creek, tributary of Takutu River 150-400 m, 31 Mar6 Apr 1938 Smith 3417 (NY, US). SURINAM. Carolina and vic., Marshall Creek, 1317 Dec 1934, Archer 2917 (U, US); Nassau Mtns. Marowijne River 400-550 m 10 Jan 195 Cowan & Lindeman 39207 (NY, U, US); Mapane Creek area near camp 8, 20 Dec 1963, Elburg L.B.B. 9827 (NY, U); Bakhuis between Kabalebo River and Coppename, along Kabalebo and Zandkreek, 22 Dec 1964, Florschutz & Maas 2481 (U); s.loc., Hostmann & Kappler 890 (B [destroyed: F neg. 2700]), BM, F, K, M (Morton neg. 8749), U); Jodensavanne, Mapane Creek area, 19 Dec 1960, Kramer & Hekking 2392 (LL, U, VEN); Sectie O, Paramaraibo-Dam railway, Nov 1941, Krukoff 12301 (NY); near Moengo tapoc, 20 Sep 1948, Lanjouw &

FLORA NEOTROPICA Lindeman 364 (K, NY, U); Wia Wia bank near Grote Zweibelzwamp, 19 Nov 1948, Lanjouw & Lindeman 1162 (NY, U); Nassau, 15 Feb 1949, Lanjouw & Lindeman 2091 (K, NY, U); Jodensavanne-Mapane Creek area. Suriname River, 27 Oct 1953, Lindeman 5000 (MO, U); Brokopondo district, Nat. Res. Brownsberg, 450 m, 16 Feb 1977, Lindeman et al. 28 (K, NY, U, VEN); Nickerie, area of Kabalebo Dam project, km 24, 30-130 m, 4?5?N, 57030'-580W, 3 Sep 1980, Lindemnanet al. 107 (F, NY, U); Brownsberg, trail form guesthouse to Irene falls, 400-450 m, 8 Nov 1974, Maas et al. 2301 (U); Charlesberg Rift, 3 km N of Paramaribo, 8 Apr 1944, Maguire & Stahel 22802 (NY, U); Kamp 8, forestry camp on Mapane Creek, ca. 18 km SE of Jodensavanne, 5?17'N, 54?0'W, 22 May 1971, Nee & Mori 4191 (MO, U, VEN, WIS); surroundings of Blakawatra, camp 8, 60 km SE of Paramaribo,22 Oct 1974, den Outer 852 (U); Saramacca River, 3 Dec 1902, Pulle 32, 128, 228 (U); forest of the station, Groningen, 10 May 1916, Samunels 63 (K, NY, P); Amakakondre, 20 Feb 1985, Sauiain 269 (NY); s.loc., Schimper 890 (W). FRENCH GUIANA. S.loc., Herb. Muis. Paris 284 (P); Jul 1824, Poiteau s.n. (K); 1821, s.coll. (G-DC); Sinnamary,Ste. Elie, km 15, 21 Oct 1981, Billiet & Jadin 1090 (BR, NY, U); Inini, Sinnamary rd. to Ste. Elie, km 22, 23 Oct 1981, Billiet & Jadin 1125 (BR); Montagne de Kaw, 4033'N, 52?09'W, 31 Dec 1987, Crenmeiset al. 9688 (U); rd. between Gallion and Montsiney, just W of Port Inini, 15 Jan 1974, Descoings & Luu 20230 (MPU, P); s.loc., Oct 1919, Godebert et al. 14 (P, U, US); Montagne de la Trinite, NE summit, ca. 1 hr from camp 4, along trail betweem camps 1 and 3, ca. 300 m. 8 Feb 1984, de Granville 6584 (K, NY, U); Montagne de Rorota, 10 Jan 1974, Halle 2199 (MPU); Saul, Monts La Fum6e, 200-300 m, 3040'N, 53?10'W, 24 Jul 1987, Hahn 3607 (NY); Piste Ste. Elie, CD-21, km 15.7, 100 m, 5?00'N, 53?10'W, 15 Sep 1987, Hahn 3791 (NY); station ECLEREX, rd. to Ste. Elie, 5020'N, 53?00'W, 30 Oct 1989, Hoff'5734 (MPU); Montagne Bellevue de l'Inini, 700 m, 7 Sep 1985, de Granville et al. 8079 (K, NY, U); Rio Oiapoque, ca. 2.5 km N of Cachoiera Tres Saltos (border with Terr. Amapa, Brazil), 2?12'N, 52?52'W, 11 Sep 1960, Irwin et al. 48162 (BM, NY); Kaw Mtns, Tresor, Favard Creek, 10-50 m, 9 Feb 1996, JansenJacobs et al. 5169 (U); s.loc., Leprieur s.n. (NY); s.loc. Martin s.n. (P, U); Maroni, 1877, Melinon s.n. (F, K, US); Saul, Mont Galbao, 12 Dec 1976, Mori et al. 8742 (MO, NY); rd. to Brazil (rt. de l'est) 4 km S of the Compte River, ca. 50 km S of Cayenne, 30 Dec 1976, Mori 8841 (MO, NY); s.loc., 1820, Perrothet 217 (GDC); Karouany, 1855, Sagot 454 (K, MPU, NY, U); Maroni River, Pompidou-Pachiton 29 Apr 1975, Sastre et al. 4037 (MO); Piste de Ste. Elie, 17 Jan 1984, Sauvain 14 (NY, U); St. Laurent de Maroni, Sotog., 27 Feb 1950 SetmvForestier Cayenne 5140 (P); Suzine, Feb 1950, Se-. Forestier Cayenne 3702 (P); Montalo, Aug 1950 Serv. Forestier Cayenne 3523 (P); Cayenne rd., km 8, 28 Feb 1955, Serv. Forestier Cayenne 7032 (NY); Cayenne region, along rd. in Forest Macouria, ca. 20 km in from hwy. D5 (Tonate-Montsinery), 1-30 m, 4?57'N, 52?30'W,

TAXONOMIC TREATMENT145 25 Oct 1986, Skog et al. 7045 (NY, U, US); St. Laurent region, Piste de Paul Isnard, 10 km SE of St. Laurent, 5-15 m, 5?25'N, 54?00'W, 17 Nov 1986, Skog et al. 7442 (NY. U); Montagnes de Kaw, Auberge de Brousse des Cascades, 140 m, 4?35'N, 52?17'W, 12 Sep 1987, Weitzman& Hahn 291 (NY, US). ECUADOR.

MORONA-SANTIAGO: Ca. 32.5 km S of

Gualaquiza (ca. 91 km N of Zamora) on rd. to Zamora, ca. 850 m, 3?35'S, 78?30'W, 4 Feb 1984, Knapp & Mallet 6244, 6245 (BH, K, QCA, QCNE, US); along Rio Metzera Grande on Hacienda Sangay, near Palora, ca. 900 m, 1?40'S, 77?58'W, 15 Feb 1984, Knapp & Mallet 6284, 6290 (BH, K, QCA, QCNE, US). PERU. AMAZONAS:Bagua, Puente Almendro I, km 296 of Marafionrd. (from Olmos jct.), ca. 620 m, 5?15'S, 78?20'W, 6 Jul 1984, Knapp & Mallet 6567 (BH, K, US, USM). LORETO:Tamshiyacu, 29 Nov 1980, Ayala et al. 2868 (MO, NY); Rio Javari, Angamo Garrison, 5 Aug 1973, Lleras et al. P17182 (K, MO, NY, US, WIS); Maynas, Rio Nanay vic. of Iquitos, caserio Santa Clara, 25 Oct 1976, Revilla 1647 (F, MO); La Victoria on Rio Amazonas, Aug-Sep 1929, Williams 3087 (F, US). SAN MARTIN:Zepelacio near Moyobamba, 1100-1200 m, Oct-Nov 1933, KlIg 3262 (F, K, USM); Naranjal, trail to Jorge Chavez, km 80 of Tarapoto-Yurimaguas rd., ca. 210 m, 6?15'S, 76017'W, 15 Jun 1984, Knapp & Mallet 6513 (BH, K, USM); trail to television antenna, km 17.5 of Tarapoto-Yurimaguas rd., 2.5 km N of Cataratas de Ahuashiyacu, 850-1200 m, 6?27'S, 76021'W, 7 Sep 1986, Knapp 8279 (MO, NY, USM); Quebrada de Ishichimi, near Tocache, 400 m, 17 Mar 1978, Schunke F 10040 (MO). BOLIVIA. LA PAZ: Prov. Nor Yungas, 1 km W of Yolosa on rd. to Unduavi, valley of Rio Cedro, 1400 m, 16?13'S, 67?45'W, 12 Nov 1987, Solomon 17383 (MO, NY). BRAZIL. S.loc., Allemao 1225 (P); s.loc., de Jussieu catal. no. 6373 (P); North Brazil, 1898, Vaughns.n. (K). AMAZONAS:Km 13 ZF-2, intersects with Manaus-Boa Vista hwy. at km 60, 12 Dec 1981, Costich 1074 (BH); Manaus, Rio Taruma 15 Mar 1937 Ducke 423 (F, K, NY, RB, US); Manaus, Flores, 21 Jan 1963 Fromm 1365 (US); Ducke Reserve near Manaus, 18 Feb 1974, Lawton 1821 (K); Rio Solimoes and Rio Javari, Rio Javari behind Palmeiras Army post, 5?08'S, 72?49'W, 31 Jul 1973, Lleras et al. P16968 (K, MO, NY, US, WIS); Sao Antonio de Iza, 8 Jul 1969, MacDaniel et al. 2374 (US); km 120 Manaus-Caracarai hwy., 12 May 1974, Nelson & Lima P21122 (K, MO, NY, US, WIS); Reserva Florestal Ducke, km 26, 31 Aug 1966, Prance et al. 2131 (NY); Humaita, Rio Madeira km 53 Humaita-Labrea, between Rios Ipixuna and Itaparana 23 Nov 1966, Prance et al. 3238 (F, K, MO, NY, US); Rio Urubu, km 214 ManausItacoatiara, 30 May 1968, Prance et al. 4892 (F, NY, US, WIS); Manaus-Porto Velho hwy. margin of Igap6 Acu at BR 319 crossing, 15 Mar 1974, Prance et al. 20563 (K, NY, U, WIS); Igarape do Tabatinga, 15 Jan 1962, Rodriguez & Chagas 4083 (NY); ManausCaracarai hwy.. km 120, 20 Feb 1974 Steward et al. P20380 (MO, NY, US); Represa de Balbina on Rio

Uatama, ca. 4 km NW of dam on D-1 rd., 1?50'S, 59?32'W, 4 Jul 1986, Thomas et al. 5346 (NY). MARANHAO: Urutawy, Ka'apor indian reserve, basin of Rio Turiacu, 28 Sep 1986, Balee 2666 (K); Fazenda Bacaba, Doctor Haroldo, 5 km S of MA 119 from entrance 3 km NW of Lago do Junco, 4?26'S, 44?58'W, 6 Oct 1980, Daly et al. D523 (K, NY); 281 km from Santa Ines on Santa Ines-Aqailandia rd., (BR 222), ca. 300 m, 4?45'S, 47?10'W, 17 Dec 1978, Jangoux & Bahia 561 (NY); Fazenda Agric. da Varig, L bank of Rio Pindare, 2 Apr 1983, Lobo et al. 344 (K, NY). MATOGROSSO: 110 km from Alto Floresta on rd. to Salto do Rio Apaicas hydroelectric plant, 10?15'S, 56?50'W, 29 Sep 1985. Cid Ferreira et al. 6275 (K, NY); 49.5 km N of base camp (base camp at 12?49'S, 51?46'W), 1 Nov 1968, Haarleyl et al. 10914 (K, NY, U); Aripuana, near Humboldt Centre on rd. to Rio Juruema, 10?12'S, 9?21'W, 8 Oct 1978, Prance et al. 18230 (K, MO, NY, US, WIS). PARk: Altasmira, Estacao Experimental de EMBRAPA, 19 Aug 1978, Bahia 97 (BH, NY); Santarem, 1877-78, Jobert 915 (P); km 113 Belem-Brasilia hwy., 150-500 ft, 7 Aug 1963, Maguirie et al. 56062 (NY, US); Tome Acu, entrance to Fazenda Curiman, 30 Dec 1977, Nascimento 358 (NY); left margin of Rio Sao Manuel, Igarap& Fernando de Noronha, below Cachoeira do Caldeirao, 7 Jan 1952, Pires 3854 (US); BR 16i, Cuiaba-Santarem hwy., km 1131, vic. of Igarape Natal, 15 Nov 1977, Prance et al. 25424 (K, RB); Santar6m, Curua-Una, Prainha, 18 Jan 1979, Santos 553 (NY); Santarem, 1878, Schwacke 915 (US); Rio Jari Monte Dourado, bandeira, 26 Nov 1968, Silva 1441 (NY); Itaituba km 85 on Itaituba-Jacareacanga Parque Nacional do Tapaj6s (IBDF), reserva biol. Ramaldopau rosa, 14 Nov 1978, Silva & Rosario 3723 (BH, NY, U); Obidos, Dec 1849, Spruce 446 (G, K, NY, P); Lageira,airstripon Rio Maicuru, ca. 800 ft, 0?55'S, 54?26'W, 17 Jul 1981, Strudwicket al. 3059 (K, NY); Obidos, Dec 1849, Traill 580 (K). RONDONIA: Km 463, linea 603 of BR 364, Cuiaba--Porto Velho rd., km 48 of Theobroma-Anari rd., Serra sem Calqa, 10?l'S, 62?63'W, 2 Jul 1984, Cid et al. 4964 (NY); km 428, linea 621 of BR 364, Cuiaba-Porto Velho rd., Pedra Branca branch, km 40, 10?11'S, 62?63'W, 3 Jul 1984, Cid et al. 5006 (NY); Porto Velho to Cuiaba hwy. 4 km S of Nova Vida 14 Aug 1968, Forero & Wrigleyl 7069 (F, K, MO, NY, US, WIS); ca. 17 km SW of Ariquemes along rd. to Santa Cruz (BR 421), 25 May 1984, Frame et al. 191 (K, NY); 15 km N of Ariquemes on Hwy. BR 364 and 1.5 km E on "Linea 75," 175 m, 9?47'S, 63?04'W, 12 Mar 1987, Nee 34331 (K, NY); 9.5 km SE of Ariquemes on hwy. BR 364, then 3.6 km E on "Linea 55," 200 m, 10?00'S, 62?59'W, 15 Mar 1987, Nee 34386 (NY); Costa Marques, Rio Cauterinho along BR 429, 200 m, 12?04'S, 63?27'W, 23 Mar 1987, Nee 34456 (K, NY); basin of Rio Madeira, 1 km S of Riberao, rd. of Abuna to Guajara-Mirim, 27 Jul 1968, Prance et al. 6567 (NY, WIS); Porto Velho to Cuiaba hwy., vic. of Sta. Barbara 15 km E of km 117, 14 Aug 1968, Prance & Ramos 6924 (F, NY, P, US, WIS); basin of Rio Madeira, vic. of Sao Lorenco mines, 9?33'S, 65?06'W, 26 Nov 1968, Prance et al. 8878 (F, NY, WIS); Alvorada-

146

PresidenteM6dicird., km 15, 20 Jun 1983, M.G. Silva 6291 (NY); MineragaoTaboca,55 km from Ariquenes nearMassangana, BR 421, 10?1'S,63?20'W,9 Oct 1979, Vieira et al. 338 (MO). RORAIMA:Manaus-Caracaraird.

(BR 174), 7 km S of equator,0?05'S, 60?40'W,14 Jun 1985, Cordeiroet al. 8 (K); Ipiranga,Rio Mucajai,betweenPratinhaandRio Apiau,26 Jan 1967,Pranceet al. 4120 (NY,US, WIS);Serrados Surucucus,S of mission station,2?46'N,36-37? W, 18 Feb 1969, Prance et al. 10044 (NY, WIS);Posto Mucujai,Rio Mucujai,21 Mar 1979, Prance et al. 11152(F, NY, US, WIS);trail from Surucucu(2?53'N,63?36'W)to Uaica(3?33'N,63?11'W) Serrados Surucucus,6 Feb 1971, Prance et al. 13514 (F, NY, US, WIS). Localnames.Colombia.Amazonas:monaigi(WitotoJito). Venezuela. Bolivar: huevo de gato. Surinam. aboepan (njulka), parabita, parabeita, bradebeita, todobita, soekaradang(Arawak),koeroeliwa (Carib), boesibita. French Guiana. mananga(Paramaka),abo pao (Sar.).Brazil. Amazonas:monagui;Mato Grosso: erva livriana.Peru. San Martin:gurac chiric sanango. The flowers of Solanum leucocarpon are reminiscent of those of Cyphomandra (Solanum sect. Pachyphvlla sensu Bohs, 1995; see also Bohs, 1994), and specimens have occasionally been identified as such. Flower size, calyx lobing, and size of the lobe apices vary somewhat throughout the range of S. leucocarpon,andleafpubescence is enormouslyvariable. Solanum leucocarpon is one of the most widespreadand variable species in sect. Geminata.Found from Panamathroughthe Amazon basin, its diverse pubescenceand flower size formshave received many names. These somewhat distinctive local forms are treatedhere as unnamedgeographicalraces, as intermediatesin all variablecharactersexist. Theseracesare briefly describedbelow. 1. In northern South America and the Guianas, plants have large, fleshy flowers and are pubescenton the leaf undersides.The trichomes arelong andslender,uniseriate,andeithersimple or branched,with both types usually occurring on a single leaf. The type specimensof Solanum leucocarpon,S. surinamense,andS. extraxillare come from among these populations. 2. In the areaaroundManausand in the Rio Negro basin, populations of almost completely glabrousplantswith somewhatsmallerflowers are common. These plants have been called Solanum coarense. 3. Populationsfromthe eastern(PeruandEcuador) and southeastern(states of Rond6niaand Mato Grosso) Amazon basin consist largely of plants with smallerflowersandshorterleafpubescence. The leaf pubescence of plants from these populations is generally of tiny, erect, usually

FLORA NEOTROPICA

simple,uniseriatetrichomes.Occasionallysome longertrichomeslike those of Guiananplantsare found along the midribon the leaf undersides. TheseplantshavebeencalledSolanumpatellare. 4. Plants from the mountains of western Guyana have shiny, glabrousleaves andrelativelysmall flowers. Very few collections exist from these glabrouspopulations. 5. Two extreme forms are found in Panama,both known from only one or two collections. Very pubescent plants from Darien province have been called Solanum eshbaughianum, and nearly glabrous ones from Isla de Coiba (Veraguas) have been called S. coibae. The fruiting pedicels of the glabrous Panamanian race aresomewhatlongerthanthose fromother populations,but they have the distinctive distal swelling characteristicofS. leucocarpon. Solanum leucocarpon is a shrub or small tree of secondaryhabitats,and is rarelyfound in the primary forest. In the Guianas it grows along forest edges in savannah,as well in many othertypes of secondaryor highly disturbedhabitats.This is one of the few species of sect. GeminatathatI have seen only in secondaryhabitatsandis also one of the most commonly collected species in the section. Specimenscited as Solanumbrevipedunculatum by Cabrera(1978) from Jujuyprovince, Argentina,may be members of Solanum sect. CyphomandropsisBitter.Membersof this sectionshareelongate,pointedbuds with the S. leucocarpon group, but have elongate inflorescenceswith widely andunevenly spacedpedicel scars and brightlycolored, few-seeded fruits. Section Cyphomandropsisis well developed in drier areas in easternBolivia andnorthwesternArgentinaandis currentlybeing monographedby L. Bohs (see Bohs, 1994 for a preliminarylist). 33. Solanum lindenii Rusby,Mem. TorreyBot. Club 6: 88. 1896. Type. Bolivia. La Paz: Mapiri, JulAug 1892, Bang 1526 (holotype, NY; isotypes, F, K, MO, NY, US). Fig. 71 SolanumpsidiifoliumRusby,Bull. TorreyBot. Club 26: 194. 1899. Type. Bolivia. La Paz: Yungas, 4000 ft, 1885, Bang 2641 (holotype, NY; isotype, NY). Solanum devernicascens Bitter, Repert. Spec. Nov. Regni Veg. 11: 483. 1913. Type. Bolivia. La Paz: Songo, Bang 2250 (holotype, B [destroyed]; lectotype, NY, here designated; isolectotypes, BM, F [F neg. 49369], K, MO, NY, WIS).

Shrubsor small trees, 5-9 m tall; young stems and leaves covered with a pinkish resinous secretion, this

147

TAXONOMICTREATMENT

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cracking and peeling as the parts expand;bark of the older stems reddish-gray,smooth, but still somewhat varnishedandresinous.Sympodialunitsdifoliate,geminate.Leaveselliptic,widest at the middle,glabrousand shiny above, dryingsomewhatdark,glabrousto densely pubescentbeneath,the trichomesuniseriate,golden, and usuallybranched,eitheralong the veins andin the primaryvein axils, or covering the entireundersideof the lamina;major leaves 10-25 x 5-10 cm, with 8-9 pairs of main lateral veins, these impressed above, prominentandgoldenbelow,the apexroundedto acute, the base acute, slightly decurrenton the petiole; petioles 1-1.5 cm long, caniculateabove;minorleaves differing from the majorones only in size, 5-6 x 2.5-3.5 cm, the apex roundedto acute,the base acute;petioles 0.8-1.2 cm long. Inflorescences opposite the leaves, simpleoroccasionallyfurcate,1.5-6 cm long,5-20-flowered,resinouslike the young stems andleaves;pedicel scars in pairs, the members of a pair closely spaced, butnot overlapping,thepairsca. 0.5 mm apart,the scars beginningabout 1/2 way fromthe base of the inflorescence. Buds globose and completely enclosed in the calyx when young, later the corolla exserted and the buds obovoid, the entire bud resinous, otherwise glabrous. Pedicels at anthesis deflexed, 0.8-1 cm long, taperingfrom a base ca. 0.75 mm diam. to the abrupt wideningof the calyx tube.Flowerswith the calyxtube campanulate,1.5-2 mm long, glabrousbutcoveredwith a resinous varnish, the lobes rounded-deltoid, 1-1.5 mm long, covered with the same resinous materialas the tube,minutelypapilloseat the tips;corollawhite or purplish, 1.5-2 cm diam., lobed ca. 3/4 of the way to the base, the lobes planarat anthesis,the tips andmargins of the lobes minutely papillose; anthers 4.5-5 x 1-1.5 mm, poricidal at the tips, the pores teardrop shaped;freeportionofthefilaments0.1-0.25 mm long, thefilamenttube0.5-0.6 mm long;ovaryglabrous;style straight, 6-7 mm long; stigma bilobed, the surface minutelypapillose.Fruita globoseberry,apiculatewith the persistentstyle base, green,ca. 1 cm diam.;fruiting pedicels woody,erector slightlydeflexed,ca. 2 cm long, 0.5-1 mm diam. at the base. Seeds tan, flattened-reniform, ca. 2 x 1 mm, the surfaces minutely pitted, the marginsincrassate.Chromosomenumbernot known. Distribution (Fig. 68). On the easternslope of the Andes in Peruand Bolivia, 1500-3000 m, in forest or secondary growth. La Mar, Specimens examined. PERU. AYACUCHO: along rd. between Machente and Rosario, 1600 m, 23 Sep 1976, Wasshausen & Encarnaci6n 697 (K, US). Cuzco: Paucartambo, Parque Nacional Manu, between Patria and Pillahuanta, 1000-2000 m, 13?08'S, 71?25'W, 13 Nov 1986, Foster et al. 12223 (F, USM). HUANUCO: Cordillera Azul, ca. 42.7 km E of Tingo Maria on rd. to

FLORA NEOTROPICA Pucallpa, 5650 ft, 21 Nov 1979 Davidson & Jones 9367 (F, NY); Chinchao, hill behind restaurant, 1 Aug 1964 Dwyer 6149 (MO); Yanono, 6000 ft, 13-16 May 1823, Macbride 3732 (F); Huallaga between Chaglla and Mufia, 1900 m, 1909-1914 Weberbauer 6709 (F, MOL, US). JUNIN: Carpapata above Huacapistana 2400 m, 7 Jun 1929, Killip & Smith 24364 (NY, US); Huacapistana. 5600 ft, Oct 1943, Sandeman 4515 (K). PASCO: Cani 7 mi NE of Mito, ca. 8500 ft, 16-26 Apr 1923, Macbride 3732 (F). SAN MARTIN: Rioja-Pomacochas rd. below Venceremos, ca. 20 km NW of Rioja near Restuarante El Amigo, 1600 m, 5?45'S, 77?38'W, 8 Feb 1984, Gentry & Smith 45143 (K, MO, NY); Venceremos. near Amazonas border, km 291 of Rioja-Pomacochas rd., 1850 m, 5?45'S, 77040'W, 12 Feb 1984, Gentry et al. 45505 (MO, NY). BOLIVIA. LA PAZ: Prov. Nor Yungas, on rd. N of Caranavi toward Sta. Ana del Rio Beni, 4500-5000 ft 8 Nov 1976, Davidson 4889 (F, US); Prov. Larecaja, Copacabana, ca. 10 km S of Mapiri, 850-950 m, 8 Oct15 Nov 1939, Krukoff 11060 (F, K, MO, NY, U, US); Prov. Nor Yungas, 16 km by rd. (ca. 8 km by air) down and NE from Chuspipata on rd. to Coroico, 2000 m, 16?14'S, 67?47'W, 29 Oct 1984, Nee & Soloimon 30235 (BH, NY); 1-3 km W of Huancane along rd. to Chuspipata, 2000 m, 16?22'S, 67?33'W, 29 Sep 1985, Nee & Solomon 32021 (NY); 16 km E of Chuspipata on rd. to Chulumani, 2150 m, 16?19'S, 67?49'W, 29 Sep 1985, Nee & Solomon 32052 (K, NY); Prov. Nor Yungas, 4.6 km below Yolosa, then 19.1 km on rd. up Rio Huarinilla, 1700 m, 16?12'S, 67?53'W, 12 Nov 1982, Solomon 8841 (MO, NY); Prov. Murillo, 44.3 km N of dam at Lago Zongo, 1500 m, 16?05'S, 68?02'W, 19 Dec 1982, Solomon 9209 (MO, NY); 44 km below dam at Lago Zongo, 1200 m, 16?03'S, 68?01'W, 12-15 Sep 1983, Solomon 10838 (MO, NY, U); 14.3 km SW (above) Yolosa on rd. to Chuspipata, 2000 m, 16?14'S, 67?47'W, 23 Mar 1984, Solomon et al. 12099 (MO, NY); Serrania de Bella Vista, 16 km N of Carrasco, 37 km N of Caranavi, on rd. to Palos Blancos, 1500 m, 15?35'S, 67?34'W, 31 Oct 1984, Solomon & Nee 12630 (K, MO, NY); 45.5 km below dam at Lago Zongo, Rio Zongo valley, vic. of Cahua hydroelectric plant, 1200-1400 m, 16?03'S, 68?01'W, 23 Dec 1984, Solomon 12927 (MO, NY); Prov. Larecaja, valley of Tipuani, 1851, Weddell s.n. (P). Solanum lindenii is closely related to S. oblongurm but differs from that species in its resinous new growth and inflorescences, and in its indument (when present) of mostly branched trichomes. These two species both have fleshy flowers with the petals planar at anthesis, globose, fleshy buds, berries borne on erect fruiting pedicels, and tiny, flattened seeds. The variation in degree of pubescence in Solanum lindenii is extraordinary, but specimens of all intermediates between the two extremes are found. Solanum lindenii is the name that was applied to the glabrous members of the species, while S. psidiifolium was ap-

TAXONOMIC TREATMENT

149

plied to the pubescentindividuals.In Bolivia, both ex- 3/4 of the way to the base, the lobes planarat anthesis, tremes of pubescence are found, but in Peru plants the tips of the lobes slightly hooded andminutelypapusuallyhave a sparseindumentof branchedtrichomes. illose; anthers4.5-5 x 1-1.5 mm, poricidalat the tips, TheinheritanceoftrichometypeanddensityinS. lindenii the poresteardropshaped;free portionof thefilaments ca. 0.4 mm long, the filamenttubeca. 1 mm long;ovary is of interest,as is the patternof trichomeontogeny. densely golden-pubescent(the trichomeswhite in live plants) with uniseriatetrichomes, these of two types, 34. Solanum oblongum Ruiz & Pav.,Fl. Peruv.2: 34, one type very short, ca. 0.05 mm long, the other type fig. 165. 1799. Type. Peru. Huanuco: In Peruvia longer and occasionally branched,0.1-0.5 mm long; nemoribusadPillaotractus,Aug-Sep,Ruiz& Pav6n style straight,7-9 mm long; stigma clavate, minutely s.n. (lectotype, MA, here designated;isolectotype, papillose.Fruit a globose, greenberry,1-1.5 cm diam., B [destroyed:F neg. 2628], MA, frag. F). densely pubescentwith uniseriatetrichomes;fruiting Figs. 66C,D, 72, 73 pedicels erect,woody, ca. 1.5 cm long, ca. 1 mm diam. SolanumoblongumRuiz & Pav. var. abruptumJ.F. at the base. Seeds pale brown,flattened-reniform,ca. 2 Macbr.,Publ. Field Mus. Nat. Hist., Bot. Ser. x 1-1.5 mm, the margins incrassate,the surfaces mi13(5B): 212. 1962. Type. Peru. Huanuco:Vill- nutely pitted. Chromosomenumber. n = 12 (voucher cabamba,haciendaon Rio Chinchao,ca. 6000 ft, Knapp & Mallet 6636). 17-26 Jul 1923, Macbride5205 (holotype, F). Distribution (Fig. 70). In the mid- to high-elevaSolanumtriste of Dunal, Solan. Syn. 21. 1816. Not tion forests of centralPeru, at 1400-3000 m. of Jacq. Shrubs or small trees, 2-4 m tall; young stems densely pubescentwith soft uniseriatetrichomes,0.21 mm long, these golden in dry material,white in live plants; young leaves glabrous; older stems more sparselypubescent,butnot glabrate;barkof trunksgray. Sympodial units difoliate, geminate. Leaves elliptic, widest at the middle, glabrousabove, pubescentalong the midribbeneathwith uniseriatetrichomeslike those of the stems,these occasionallybranched;majorleaves 15-20 x 7-10 cm, with 7-10 pairsof mainlateralveins, these impressed above, prominentand very darkbelow (in dry specimens), the apex acute to acuminate, the base slightly oblique, acute, winged onto the petiole; petioles winged from the decurrentleaf bases, 115 cm long; minorleaves more roundedin outlinethan the major ones, 2-7 x 1.5-5 cm, the apex acute, the base rounded;petioles 3-5 mm long. Inflorescences opposite the leaves, simple, 1-2 cm long, 6-10-flowered, sparsely to densely pubescent with the same uniseriatetrichomes as those of the stems, 0.2-1 mm long, golden in dry specimens, white on live plants; pedicel scars closely spaced, ca. 0.5 mm apart,raised andcorky.Buds globose when young, the corolla soon exserted from the calyx andthe buds becoming obovoid, pubescentwith the sametrichomesas therestof the inflorescence.Pedicels at anthesis0.8-1 cm long, tapering from a slenderbase ca. 0.5 mm diam. to the abrupt wideningof the calyx tube.Flowerswith the calyxtube campanulate,ca. 2 mm long, sparsely to densely pubescentwith uniseriatetrichomes0.1-0.5 mm long, the lobes broadlydeltoid, 0.5-1 mm long, pubescentwith uniseriatetrichomes,these shorterthanthose of the calyx tube, the marginsof the lobes thickenedandwhite; corollawhite, thickandfleshy, 1.6-2.2 cm diam.,lobed

Specimens examined. PERU. HuANUCO:Leoncio Prado, near La Divisoria, 1500-1600 m, 25 Jun 1976. Schunke V 9395 (MO); Pendencia, 900 m, 13 Sep 1962, Woytkowski 7528 (MO). PAsco: Palcazu valley, Rio Pichinaz, along rd. from W of divide between Rio Cacazu and bridge over Rio Pichinaz, 600-800 m, 10?22'S, 75?08'W, 26 Oct 1982, Foster & Smith 9437 (F, USM); km 42.6 on Oxapampa-Pozuzo rd., near Agua Salada, ca. 1500 m, 10?20'S, 75?28'W, 13 Mar 1984, Knapp & Mallet 6324 (BH, K, NY, US, USM); km 1518 Villa Rica-Pto. Benrudez, ca. 1600 m, 21 Mar 1984, Knapp et al. 6339 (BH, NY, US, USM); Oxapampa, Enefias-Alto Yurinaki-La Florida rd., ca. 9 km E of Villa Rica, 1250-1400 m, 10?50'S, 75?15'W, 12 Aug 1984,

Knapp& Mallet6624 (BH, K, NY, US, USM);SanJuan de Cacazu,km 36 on VillaRica-Pto.Bermudezrd. along Rio Chivis, ca. 950 m, 10?38'S, 75?10'W, 13 Aug 1984, Knapp & Mallet 6628 (BH, K, NY, US, USM); 6630 (BH, K, US, USM); ca. 1 km from division of Villa RicaPto. Bermudez rd. and Villa Rica-Palcazu rd., along Palcazu branch,Rio Cacazu, ca. 500 m, 10?30'S, 75?10'W,

15 Aug 1984,Knapp& Mallet6636, 6637 (BH, K, US, USM). SANMARTIN:Near Arroyo Bravo ca. 40 km from Tingo Maria on hwy. to Pucallpa, ca. 1250 m, 1 Nov 1949, 5 Jan 1950, Allard 21767 (US).

Solanumoblongumis closely relatedto S. lindenii of PeruandBolivia, but differs fromthat species in its pubescentovaries,longer,thinnertrichomes,andin its pubescent calyces and inflorescences. Solanrum oblongum,althoughlocally commonwhereI have collected it, has not been collected very often. It is apparently of relativelyrestricteddistributionin centralPeru along small streamsandrivers at middle elevations. The sheet I have selected as the lectotype at MA is thatmost closely matchingtheplatein theFloraPeruviana et Chilensis (Ruiz & Pav6n, 1799).

150

FLORA NEOTROPICA

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FIG. 72. Solanum oblongum Ruiz & Pav6n, plate 165b of Ruiz & Pav6n's Flora Peruviana.

TAXONOMIC TREATMENT .i.'.

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Solanaceae Solanum oblonqum R. & P. PASCO: Oxapampa. ca. 1 km. from division of Villa Rica-Pto. Bermudez road and Villa Rica Palcazu road, on Palcazu branch. Along small to the Rio Cacaszu. quebrada, tributary wet forest and old second Tropical growth. ca. 500 m. 10 30' S X 75 10' W. Shrub or small FIG.

73.

FIG. 73.

Solanum

oblongum

Ruiz

&

Solanum Ruiz oblongum

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Knapp

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2-3 m.

& Mallet

Flowers 6637

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& Knapp & Mallet 6637 (K). Peru. Pav6n. H. BAILY HOORTORlUM L. R:..

(

FIG. 73. Solanum oblongum Ruiz &i Pavbn. Peru. Knapp & Mallet 6637 (K).

white,

FLORA NEOTROPICA

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C^aWm>42 ;s~~~~~~~~~~~~~~~~~~dr g,,_Diii:i FIG. 74. Solanum nutans species group. A. S. malletii shoot, Peru (Knapp & Mallet 6564). B. S. malletii flowers, Peru (Knapp & Mallet 6564). C. S. youngii flowers, Ecuador (Knapp et al. 9117). D. S. youngii plant, Ecuador (Knapp et al. 9117).

V. Solanum nutans species group (S. malletii, S. microleprodes, S. nutans, S. xanthophaeum, S. Fig. 74 youngii).

andsomewhatclaspingto attenuate.Inflorescencesopposite the leaves, simple or occasionallybranched,pubescent like the stems and leaves. Buds globose to elthe corolla strongly exserted from the calyx Shrubs or small trees; young stems and leaves lipsoid, tube. Flowers white or greenish-white, usually denselypubescentwith dendriticor echinoidtrichomes, the corolla lobes planarat anthesis, octhese usually golden. Sympodial units unifoliate heterostylous, casionallypubescentwithdendritictrichomes.Fruitgreen (Solanumnutans)ormorecommonlydifoliateandgemi- and hard at maturity,glabrous or pubescent;fruiting nate, often anisophyllous.Leaves elliptic to obovate, the erector deflexed.Seeds flattened-reniform, pedicels usually glabrousor sparsely pubescent with dendritic incrassateandpaler,pale tanto reddish-brown. margins or echinoidtrichomesadaxially,denselypubescentwith Distribution. Lowlandto highlandforestsandopen dendritictrichomesabaxially,the surfaceoccasionally bullate(S.youngii),theapexusuallyacute,thebasesessile areas,CentralandSouthAmerica(Panamato Bolivia).

TAXONOMIC TREATMENT

15 3

Themembersof theSolanumnutansspeciesgroupare can be difficultto identify(see below),butthe consistent all remarkablysimilardespitetheirextremesof habitat. prescenceof unifoliatesympodialunitsandcomplexdenSolanumnutansis themostvariablespeciesinthegroupand dritictrichomesthatdryyellowishorreddishis distinctive.

Key to species of the Solanum nutans species group 1. Sympodial units difoliate, geminate; ovary glabrous or pubescent. 2. Leaves strongly bullate; ovary glabrous. High-elevation cloud forests, Peru and Ecuador............39. S. ounigii 2. Leaves not bullate; ovary glabrous or pubescent. Lowlands in Central and South America. 3. Stem and leaf trichomes dendritic; flowers pale green; seeds reddish-brown. 4. Ovary and berry glabrous; leaf bases acute; minor leaves elliptic, the apex acute ............................................................................................................................ 3 8. S. xanthoph ae 4. Ovary and berry pubescent; leaves auriculate; minor leaves orbicular, the apex ro und e d .................................................................................................................................... 35. S. alletii 3. Stem and leaf trichomes echinoid; flowers white; seeds tan. Panama to northern Colom bia ........................................................................................................................... 36.. S. icroleprodes 1. Sympodial units unifoliate or difoliate, never geminate; ovary glabrous. Ecuador to Bolivia ..........37. S. nu.tatns

35. Solanum malletii S. Knapp,Novon 2: 344. 1992. Type.Peru.Amazonas:Bagua,QuebradaChinganza, 8 km N ofMuyo, km 472 ofOleoducto Norperuano, 350-400 m, 5?25'S, 78?28'W, 5 Jul 1984, Knapp & Mallet 6564 (holotype, USM; isotypes, BH, F, K, US). Figs. 74A,B, 75

lobes planarat anthesis,the abaxialsurfaceof the lobes densely dendritic-pubescent;anthers2-2.5 mm long, 1.5-2 mm wide, in the type with a few dendritictrichomes on the abaxialsurface,poricidalat the tips, the pores teardropshaped;free portionof the filamentsca. 0.25 mm long, the filamenttubeca. 0.5 mm long;olvanr Slender shrubs to small trees, 1-7 m tall; young densely pubescentwith dendritictrichomesca. 0.5 mm stems and leaves densely pubescentwith dendritictri- long; styles heteromorphic,in short-styledflowers 1chomes, these 0.5-1 mm long, pale golden when dry, 1.5 mm long, in long-styled flowers 5-6 mm long; beige when fresh;barkof older stems reddish-brown, stigma clavate, the surfaceminutelypapillose. Frulita sparsely pubescent with dendritictrichomes.Sympo- globose, greenberry,1-1.2 cm diam.;fruitingpedicels dial units difoliate, geminate, anisophyllous. Leaves woody, erect or deflexed, 0.7-1 cm long, ca. 1.5 mm glabrous and shining adaxially, with a few dendritic diam.at the base. Seeds reddish-brown,flattened-renitrichomes along the main veins, pubescent with den- form with the margins incrassateand paler, 1-3 mm dritictrichomesabaxially,the pubescencedenseralong long, 1.5-2.5 mm wide, the surfaces minutely pitted. the veins; majorleaves elliptic-obovateto obovate, 17- Chromosomenumber not known. 26 cm long, 8-15 cm wide, with 9-13 pairs of main Distribution (Fig. 76). On the W margin of the lateral veins, the apex acute to acuminate, the base Amazon basin in Ecuadorand Peru, in tropical wet sessile andtruncate,occasionallysomewhatauriculate; forest, at 200-1300 m. petiole if presentca. 2 mm long; minorleaves differing Specimens examined. ECUADOR. NAPO:Reserva from the majors in both size and shape, orbicular,37.5 cm long, 3-7 cm wide, the apex rounded,the base Biol6gica Jatun Sacha, Rio Napo, 8 km E of Misahualli, 450 m, 1?04'S, 77?36'W, 21-25 May 1987, Cerd6l M. rounded;petioles 1-2 mm long. Inflorescencesoppo1394 (MO, NY); Parque Nacional Yasuni, Pozo Amo 2. site the leaves, simple, 1-15 cm long, 10--15-flowered trochas de Amosur, 230 m, 0?52'S, 76?05'W, 9-13 Jan at a time, but with up to 100 scars, densely pubescent 1988, Cerdn M. & Coello 3182, 3298 (MO, NY): La with dendritictrichomes;pedicel scars densely spaced, Joya de Sachas, P.N. Yasuni, Oleoducto de Maxus, km not overlapping. Buds globose, the corolla strongly 45, 230 m, 0?45'S, 76?28'W, 8-15 Aug 1993, Dik 194 exserted from the calyx tube, densely pubescentwith (QCNE); Cant6n El Chaco, Rio Granadillo, INCEL camp dendritic trichomes. Pedicels at anthesis somewhat Codo Alto, 1300 m, 0?08'S, 77?28'W, 13-15 Sep 1990. deflexed, strongly contractedbeneaththe calyx tube, Palacios 5561 (QCNE); Reserva Faunistica Cuyabeno. ca. 1 mm diam. at the apex, 0.5 mm diam. at the base, just N of Laguna Grande, 205 m, 0?01'N, 76011'W, 130.7-1 cm long, densely pubescent with dendritic tri- 18 May 1988, Renner 69073B (AAU); Est. Biol. Jatun chomes like those of therestof the inflorescence.Flow- Sacha, near Rio Chinguipino, 380 m, 1?04'S, 77?36'W. 11 Aug 1992, Rueda 1142 (QCNE). ers with the calyxtubebroadlyconical, 1.5-3 mm long, PERU. AMAZONAS:400 m behind La Poza. Rio the lobes triangular,1.5-3 mm long, densely pubesSantiago, 180 m, 22 Aug 1979, Huashikat 130 (MO). cent with dendritic trichomes;corolla pale green, 1- LORETO: Balsapuerto, lower Rio Huallaga, 150-350 m. 1.2 cm diam., lobed ca. 3/4 of the way to the base, the 28-30 Aug 1929, Killip & Smith 28636 (NY. US).

154

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TAXONOMIC TREATMENT

155

Shrubsor small trees, 1-5 m tall; stems and young leaves densely pubescentwith erect,dendritic,densely branched,tree-liketrichomes,these not persistentonto older stems; barkof older stems pale grayish-brown. Sympodial units difoliate, geminate. Leaves elliptic, widest at the middle, glabrous and shining adaxially, with a few scatteredtree-liketrichomesalong the main veins, sparsely to densely pubescent along the veins abaxially, occasionally with a few trichomes on the 00 lamina;majorleaves 10-20 x 5.5-8 cm, with 6-9 pair of main lateralveins, these dryingyellowish abaxially, the apex acuteto acuminate,the base acute, somewhat decurrenton the petiole; petiole 1-1.5 cm long; minor leaves differing from the majorsin size and shape, elliptic or orbicular,2-9 x 1.5-6.5 cm, the apex acute to rounded,the base acute;petiole ca. 0.5 cm long. Inflorescence opposite the leaves, simple or occasionally once branched,1-3(12) cm long, 5-10-flowered,densely pubescentwith tree-like trichomes like those of the stems; pedicel scars closely spaced but not overlapping. Buds globose when young, laterelliptic with the corolla stronglyexsertedfromthe calyx tube.Pedicels at anthesis slender, deflexed, abruptlycontractedbelow the calyx tube,0.6-1 cm long, 0.5 mm diam.,densely pubescentwithdendritictree-liketrichomeslike those of the inflorescence.Flowers with the calyx tube conical, abruptlycontractedto the pedicel, 1.5-2 mm long, the lobes minute,if presentbroadlydeltate,0.1-0.5 mm long, densely pubescent with dendritic, tree-like triFIG. 76. Distribution of Solanum malletii (solid chomes; corolla white, 1-1.3 cm diam., lobed nearly to the base, the lobes planarto slightly reflexed at ancircles) and S. microleprodes (open circles). thesis, the abaxialsurfacepubescentwith tree-like trichomes like those of the restof the inflorescence,these denser along the marginsand at the tips; anthers 2.5Solanummalletii can easily be distinguishedfrom 3 mm long, 1-1.5 mm wide, poricidal at the tips, the othermembersof the S. nutansspecies groupby its orpores teardropshaped;free portionof thefilamentsca. bicularminorleavesandauriculatemajorleafbases. The 0.5 mm long,thefilamenttubeminute,ca. 0.25 mm long, elongate,densely pubescentinflorescencesof this spe- glabrous;ovaryglabrousto pubescentwithdendritictreecies are like those of S. youngii, butS. malletii is easily like trichomes;style glabrous,heteromorphic,in longdistinguishedfromthatspeciesby its non-bullateleaves, styled flowers ca. 6 mm long, in short-styledflowers orbicularminor leaves, and low-elevation habitat. ca. 2 mm long, somewhatclavate;stigma a broadened The presence of short-styled flowers in Solanum areaon the styletip,the surfaceminutelypapillose.Fruit malletii(see Fig. 74B) indicatesthatit is probablyandro- a globose,greenberry,1-1.5 cm diam.;fruitingpedicels monoecious, but this has yet to be tested in the field. woody, erectto somewhatdeflexed, 1-1.5 cm long, 12 mm diam.at the base, swollen below the calyx lobes. with paler,incrassatemarSeedstan,flattened-reniform 36. Solanum microleprodes Bitter,Repert.Spec. Nov. gins, 2.5-3 mm long, 2-2.5 mm wide, the surfacesmiRegniVeg.13:96. 1913(1914?).Type.Panama.Darien: nutely pitted. Chromosomenumbernot known.

800

Boca de Pauarand6, Rio Sambu, 20 m, Feb 1912,

Pittier5686 (holotype,US; isotype,US).

Fig. 77

SolanummicroleprodesBitter var.felicis D'Arcy, Ann. Missouri Bot. Gard. 60: 728. 1974 [1973]. Type. Panama. Chiriqui: vic. San Felix, 0-120 m, Pittier 5173 (holotype, US).

Distribution (Fig. 76). In secondarygrowth from Panamato N coastal Colombia, at 0-500 m. Specimens examined. PANAMA. S.loc., 1852-57, Seemann 1072 (BM). CHIRIQUi: Remedios and vic., 0100 m, Dec 1911-Jan 1912, Pittier 5465 (US). DARIEN:

FLORA NEOTROPICA

156

pov.

DARIE.:

Solanun sioroleprod.s Bitt. det.

W. D'Arcy

of it Tuira to track QuebradaSierpe froam (between Rlos Cube and

Ionusa),

Chevy ExpeditIon

Shrub 3 emtersa flowere whites fruits green.

.................. ..I_

:

Al Gentry no. MNO"i

27 Feb 1972

4433

UD arsIce wOTMICA

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5cm FIG. 77. Solanum microleprodes Bitter. Panama. Gentry 4433 (BM).

(MD)

TAXONOMIC TREATMENT 110 mi. from Bayano bridge, vic. of Cangl6n, trail to S of rd., ca. 50 ft. 14 May 1980, Antonio 4569 (BM, MO); Rio Pirre, 14 Jul 1971, Croat & Porter 15507 (MO); rd. from El Real to Pinogana, 6 Jul 1962, Duke 5152 (MO); Rio Balsas between Rio Areti and Manane, 14 Sep 1965, Duke 8791 (MO); Agua Fria, ca. 8 mi N of Santa Fe, ca. 50 m, 13 Feb 1967, Duke 10109 (MO, US); Rio Tuira, between Rio Punusa and Ri6 Mangle, 1 Oct 1967, Duke 14603 (US); Quebrada Sierpe from Rio Tuira to track of Chevy Expedition, between Rio Cube and Rio Punasa, 27 Feb 1972, Gentry 4433 (MO); Rio Pirre near town of Pirre, 27 Dec 1972, Gentry 6944 (MO); 1.5 mi. W of Santa Fe on PanAmerican Hwy., then 2 mi. N on logging trail, 30-50 m, 26 Apr 1982, Huft et al. 1968 (MO); Rio Pirre, up river from house of Bartolo, 16 Mar 1973, Kennedy 2874 (MO, US); along rd. from Cangl6n to Yaviza, beginning 10 mi. E of Cangl6n, 50 m, 8?15'N, 77?45'W, 7 Mar 1982, Knapp & Mallet 3976 (MO); ridges of Filo del Talo, SW of Canglon, upper reaches of Rio Canglon, 250-300 m, 8?20'N, 77?55'W, 8 Mar 1982, Knapp & Mallet 4005 (MO); vic. Paya, Rio Paya, trail from Paya to Pucro, 12 Jun 1959, Stern et al. 417 (MO). HERRERA:10 km W of Las Minas on rd. to El Toro, 200-600 m, 24 Jan 1981, D'Arcy & Sytsma 14327 (MO); 10 km W of Las Minas on rd. to El Toro, 600 m, 24 Jan 1981, Svtsma & D'Arcli 3240 (MO). PANAMA: 1-2 mi. S of PanAmerican Hwy., 3 mi. E of Cafazas checkpoint, foothills of Serrania de Cafazas, 0-50 m, 8?52'N, 78?15'W. 27 Feb 1982, Knapp 3880a (MO). VERAGUAS:Trail between Canazas and foot of Cordillera Central, headwaters of Rio Cafiazas, 300-600 m, 8 Feb 1937, Allen 202 (MO). COLOMBIA. ANTIOQUIA: Km 4 DabeibaChigorod6, 180-400 m, 30 Jul 1987, Callejas et al. 4775 (MO, NY); near Rio Le6n, ca. 20-30 km upstream and S of mouth, ca. 13 km W of Chigorod6, ca. 100 m, ca. 7?45'N, 76?50'W, 20 Mar 1962, Feddema 1982 (NY); banks of Rio Cauca at Puerto Valdivia, 17-20 Feb 1942, Metcalf & Cuatrecasas 30091 (MO, US); UrabaChigorod6-Malag6n, 10 m, 26 Mar 1986, Renteria et al. 4822 (MO). BOLiVAR: Frasquillo, on Rio Sinu, 5-6 Mar 1918, Pennell 4597 (NY). CHOCO: Parque Nacional de Los Katios, W of Salto de Tilupo, 180 m, 7 Mar 1976, Forero P. & Le6n 146 (MO). MAGDALENA: Near Onaca, 2500 ft, 6 Feb 1998-99, Smith 1154 (F, K, NY). SANTANDER: Camp on Arenosa creek, vic. Barranca Bermeja, Magdalena Valley between Rio Sogamoso and Rio Colorado, 100-500 m, 20 Jul 1934, Haught 1318 (US); San Juan valley, vic. Puerto Berrio, between Rio Carare and Rio Magdalena, 100-700 m, 11 Jun 1935, Haught 1769 (K, US); 22 km S of El Centro, vic. BarrancaBermeja,between Rio Carareand Rio Sogamoso, 100 m, 2 Oct 1936, Haught 2000 (US); Viscaina Creek 31 km S of El Centro, vic. Barranca Bermeja, between Rio Carare and Rio Sogamoso, 100 m, 3 Nov 1936, Haught 2057 (A, K, US). VENEZUELA. ZULIA:Basin of Rio Guasare, 1 km below Destacamento Guasare No. 1 (La Yolanda), L bank of Rio Guasare, 400-500 m, 3 Feb 1983, Bunting & Leon 12858 (NY).

157

Solanum microleprodes is closely related to S. xanthophaeumand S. malletii, both of Peru. It shares with those species a dense indumentof branchedtrichomes and swollen calyx tube base in fruit. The trichomes of S. microleprodesare tree-like or echinoid, with short,congested branches,while those of the two SouthAmericanspecies areloosely dendritic.D'Arcy's var.felicis from Chiriquiprovince has more loosely branched trichomes than material from Darien and Colombia, but specimens from Herreraand Veraguas areintermediatein this character.

37. Solanum nutans Ruiz & Pav.,Fl. Peruv.2: 35, fig. 166a. 1799. Type.Peru.Huanuco:Pillao, Aug-Sep, Ruiz& Pav6n s.n. (lectotype,MA, heredesignated; Figs. 78, 79 isolectotypes, MA, MPU). Solanumbassoviicarpum Rusby,Bull. New YorkBot. Gard. 4: 419. 1907. Type. Bolivia. S.loc., Bang

2524 (lectotype, NY, here designated;isolectotypes, K, NY, US). Solanumhypomalacothrix Bitter,Repert.Spec. Nov. Regni Veg. 18: 70. 1922. Type.Bolivia. Cochabamba:QuebradaPocona, 3000 m, Apr 1911, Herzog 2195 (lectotype,B, here designated). Shrubsor small trees, 3-8 m tall; young stems and leaves denselypubescentwith goldentree-likedendritic trichomes, those of the stems more densely branched and congested, the trichomes0.5-1 mm long; barkof older stems gray to brownish-red. Sympodial units unifoliateoroccasionallydifoliate,not geminate.Leaves elliptic, widest at the middle, 7-20 x 3.5-8 cm, with 10-14 pairsof main lateralveins, these dryingreddish, glabrous and shining adaxially with a few scattered dendritictrichomes on the veins and lamina, densely pubescentabaxially,the trichomesdendriticwith congestedbranches(see discussion),theapexacute,thebase acute;petioles 1-2.5 cm long. Inflorescencesopposite the leaves or interodal, 0.5-1.5 cm long, simple, 510-flowered,denselypubescentwithtree-liketrichomes like those of the stems. Buds globose, the corolla ca. 1/2 way exserted from the calyx tube;pedicel scars closely spaced, not overlapping.Pedicels at anthesis nodding, 0.5-1 cm long, ca. 1 mm diam. at the base, 0.25-0.5 mm in diameterat the apex, densely pubescent like the rest of the inflorescence.Flowers with the the calyxtubeconical, 1-2.5 mm long, the lobesbroadly deltoid, 1-4 mm long, somewhatpetaloidin a few collectionsfromcentralPeru,denselypubescentwith treelike dendritictrichomeslike those of the rest of the inflorescence; corolla white, fleshy (occasionally describedas yellow, but this probablydue to the dense abaxialpubescence),1-1.5 cm diam.,lobednearlyto the base, the lobes planarat anthesis,the abaxialsurfaceof

158

FLORA NEOTROPICA

C.LXVI.

S TOLANUM S/0,A..

N SOLANUM

ari/&w<w.

FIG. 78. Solanum nutans Ruiz & Pav6n, plate 166a of Ruiz & Pav6n's Flora Peruviana.

159

TAXONOMIC TREATMENT

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Jaramillo 2485 (K, NY), 4 Aug 1978, Boeke & Jaramillo 2596 (NY); I km from control post on Loja rd., on way to Yanasacha, 8 Aug 1978, Jaramillo & Boeke 485 (NY); 00 Chinquintad-Represa Electrica-Labrado rd., 13 km from jet., 2780 m, 7 Aug 1986, Jaramillo et al. 8877 (AAU, NY); Cuenca-Bosque de Moza (ETAPA) rd., 3000-3200 m, 17 Aug 1987, Zak 2471 (AAU, NY), Zak 2479 (AAU, NY); Cuenca-Paute-Sevilla de Oro rd., paramos del Castillo, 2500-3000 m, 18 Aug 1987, Zak 2496 (AAU, NY); Cuenca-Giron rd., Nudo de Portete 9 km from Cuenca, 7 km from main rd. on side rd. to Hacienda Patacocha, 2700 m, 20 Aug 1987, Zak 2515 (NY), Zak 2516 (NY). CANAR: Alausi--Cafar rd., detour to Oyashig and Hacienda El Carmen N of Cafar, 3270-3700 m, 12 Aug 1987, Zak 2407 (NY). El, ORO: Track from Chilla to the Antennas, 2900 m, 3 28'029"S, 79 36'181"W, 20 November 1996, Lewis et al. 2841 (K). IIBABURA: Canton Cotacachi, rd. to Apuela, NW of Cuicocha, 2500-2800 m, 0?20'N, 78?26'W, 12 Jun 1992, Cuamacudset al. 146 (QCNE); Cotacachi-CuicochaApuela-San Luis de las Delicias rd., vic. San Luis de las Delicias, 2400-2600 m, 6 Dec 1986, Zak 1468 (NY). LOJA: Catamayo-Catacocha, km 25, turnoff at Las Chincha toward Pifas, km 1.6, 2490 m, 3 57'08"S, 800 79 28'53"W, 13 Dec 1994, Jorgenson et al. 1477 (BM). PICHINCHA: Reserva Geobotanica Pululahua, rd. to Los between Reventaz6n and property of Don Julio Reales, FIG. 80. Distribution of Solanumn nutan(s. Mosquera, 1800-3356 m, 0?05'N, 78?30'W, 24 Oct 1987, Cerdn M. & Benavidez 2540 (NY); km 14, Lloa--Mindo rd., Hcda. Las Palmas of Lcdo. F. Sotomayor, 2900 m, the petals sparsely (northernpopulations) to densely 0?10'S, 78038'W, 9-10 Jul 1988, Jorgensen 65446 (centralPeru)pubescentwith congested dendritictri- (AAU), 7 Feb 1989, Jorgensen 65972 (AAU); 30 km W chomes;anthers3-5 mm long, 1-2 mm wide, poricidal of Aloag on way to Santo Domingo, 2200 m, 12 Sep 1984, at the tips, the pores teardropshaped; free portion of Whaleni816 (BH, NY); 16 km E of Tandapi on Santo Domingo-Quito rd., 6800 ft, 3 Aug 1980, Wunderlinet thefilamentsca. 0.2 mm long, the filamenttube ca. 0.2 al. 8694 of Lloa and Palmira, SW slopes of mm long, glabrous; ovary glabrous or occasionally Volcan (F); valley20-29 km from Pichincha, Quito-Lloa-Mindo pubescent with dendritic trichomes in a few central rd., 2500-3000 m, 0?12'S, 78?39'W, 15 Jun 1986, Zak Peruviancollections;styles heteromorphic,the shorter 1069 (NY), Zak 1070 (NY); Quito-Chiriboga-Empalme styles ca. 2.5 mm long, the longerstyles 5-6 mm long; rd., km 75-85, 1500-1600 m, 0?15'S, 78?50'W, 31 Jul stigma capitate,the surfaceminutelypapillose.Fruita 1987, Zak & Jaramillo 2321 (NY). PERU. AMAZONAS: Prov. Chachapoyas, Balsasglobose, green to greenish-black berry, 1.3-2.3 cm diam.;fruitingpedicels woody, deflexed or somewhat Leimabambard., km 394, 3 Jun 1977, Boeke 1892 (BM, erect, 1.5-2.5 cm long. Seeds reddish-brown,flattened NY); Chachapoyas,1835, Matthews 1513 (K). AYACUCHO: lenticularwith incrassatemargins,4-4.5 mm long, 2.5- Huanta, Choimacota valley, 28 Feb-10 Mar 1926, Weberbauer 7551 (F, GH, NY, US); 4 km above Ocros, 3 mm wide, the surfacesminutelypitted.Chromosome 3800 m, 29 Oct 1935, West 3671 (MO). CAJAMARCA: numbernot known. Colasay, 2700 m, 18 Oct 1961, Wo)'tkowski6947 (MO). Distribution (Fig. 80). In the Andes from S Co- Cuzco: Urubamba, Macchu Picchu, Pacasmayo river lombia to Bolivia, in high-elevation forest and ceja de drainage, 0.2 km N of where Inca trail crosses Rio Pacasmayo, 3220 m, 22 Aug 1982, Pevton & Pevton selva, 1800-3500 m. 1093 (F). HUANUCO:Huanuco, Dombey s.n. (F, P); rd. Selected specimens examined. COLOMBIA. from Huanuco to La Union, km 5-31, vic. Mito, ca. Valle de Sibundoy, 1.5 km S of Sibundoy, 3000 m, ca. 9?55'S, 76?20'W, 17 May 1982, Landrtum PUTUMAYO: 2200 m, 24 Jul 1963, Bristol 1259 (US); Sibundoy, in 4617 (F, NY); Yanahuanca, ca. 10000 ft, 16-22 Jun 1922, Macbride & Featherstone 1214 (F, US); Mito, ca. houseyards and fencerows, 11 Nov 1968, Plowman 2003 9000 ft, 8-22 Jul 1922, Macbride & Featherstone 1473 (K, US). ECUADOR. Cedropamba, 2350 m, 19 Aug 1975, (F); 3-6 mi. NW of Mito, ca. 11000 ft, 10 Aug 1922, Little & Campunzano263 (MO). AZUAY: Cuenca, Haci- Macbride & Featherstone 1925 (F, US); 15 Mi. SE of enda Yanasacha, 3000-3200 m, 22 Jul 1978, Boeke & Huanuco, ca. 10500 ft, 31 May-3 Jun 1922, Macbride

TAXONOMIC TREATMENT & Featherstone 2103 (F, US); Mufa, trail to Tambo de Vaca, ca. 8000 ft, 5-7 Jun 1923, Macbride 4278 (F), Macbride 4315 (F); Tambo de Vaca, ca. 13000 ft, 1024 Jun 1923, Macbride 4416 (F); Huanuco-La Union rd. above town of Mito, 2800-3200 m, 9?45'S, 76?48'W, 20 Apr 1966, Macurdy 1038 (F, US); Guahuncho, 1 Jul 1927, Sawtada52 (F). JUNiN:Mt. Pariahuanca,Matthews 1195 (K); Pintac, 11-12000 ft, Apr 1864, Pearce s.n. (K). SANMARTIN:P.N. Rio Abiseo, near La Playa camp, 2650 m. 7?S, 77?W, 30 Aug 1985, Young 1514 (K, USM). BOLIVIA. LA PAZ: Prov. Murillo, 30.5 km N of dam at Lago Zongo, trail up Rio Jachcha Cruz, 2200 m, 16?07'S, 68005'W, 30 Nov 1982, Solomon 9062 (K, NY, U); Prov. Nor Yungas, 1.2 km E of Cotapata on rd. to Unduavi and Chuspipata, 3100 m, 16?17'S, 67?50'W, 26 Jun 1986, Solomon 15350 (NY). SANTACRUZ: 10 SSE by air of Vallegrande, on rd. from Guadelupe to

Piraimiri,2200 m. 18?34'S,64?03'W,31 Jan 1987,Nee and "El et al. 33920 (IBE, NY); between"Mataralcito" Palmar"on rd. from Vallegrandeto TierrasNuevas, 17 km by air ESE of Vallegrande,2150 m, 18?32'32"S, 63?57'30"W, 30 Dec 1988, Nee et al. 37434 (IBE, NY); Comarapa, 2800 m, 21 Oct 1928, Steinbach 8380 (F,

NY, US); Comarapa,2600 m, 24 Oct 1928, Steinbach 8458 (F. K, NY); Prov. Vallegrande,Huascacafiada,5 km S of town of Vallegrande, 2050 m, 18?31'S, 64?06'W, 5 Aug 1991, Vargas C. & Vargas 1075 (K).

Local names. Colombia. Putumayo: shanzanshu (Kansa, Sibundoy). Peru. Ayacucho: ccormentoy; Huanuco:chuchulate. nutansis a variablespecies,particularly with Solanumw respect to pubescence. Specimens from Peru (including the type) aredensely yellow pubescent,while those from Ecuadorand Colombia tend to be less densely pubescent and the trichomes are more matted and brownish-redin color. The trichomes ofS. nutans are distinctive and distinguish it from all other species in this group.Trichomesof the stems and leaves are dendritic, with very short, congested branches.They approachthe echinoidtrichomesofS. microleprodes,but are more elongate and highly branched. The lectotypechosenat MA is thatspecimenclosely matching the illustration in the Flora Peruviana et Chilensis (Ruiz & Pavon, 1799).

38. Solanum xanthophaeum Bitter, Repert. Spec. Nov. RegniVeg. 16:401. 1920.Type.Peru.Huanuco: HuamaliesbetweenMonzonandRio Huallaga,700 m, Sep, Weberbauer3690 (holotype, B [destroyed: F neg. 2643, F, GH, NY, US], lectotype, G, here designated [Morton neg. 8667 F, GH, NY, US], isolectotype, MOL). Fig. 81 Shrubor smalltreelet2-3 m; stemsandnew growth denselypubescentwithgoldendendritictrichomes,these either loose (ca. I mm long) or compact (ca. 0.5 mm

161

long); barkof older stems darkbrown, the trichomes somewhatpersistent.Sympodialunits difoliate, geminate. Leaves elliptic-obovate, glabrous and shining adaxiallywith a few dendritictrichomesalongtheveins, sparsely pubescent on the lamina abaxially with dendritictrichomeslike those of the stems, densely pubescent on the main lateralveins; majorleaves 10-28 x 49.5 cm, with 4-7 pairs of main lateral veins, these yellowish abaxiallywhen dry,the apex acuteto acuminate, the base attenuate;petioles 0.5-1 cm long; minor leaves differingfromthe majorsin size andshape,elliptic to rounded,2.5-5 cm long, 1.5-3.5 cm wide, theapex acuteto rounded,thebaserounded;petioles3-5 mmlong. Inflorescencesoppositethe leaves, 1-5 cm long, simple, 10-15-flowered, densely pubescent with golden trichomes, these eitherloose or compactlike those of the stems;pedicel scars closely spaced but not overlapping, beginningnearthe base of the inflorescence.Buds globose when young, laterelliptic, the corollastrongly exsertedfromthe calyx tube.Pedicels at anthesisslender, 5-8 mm long, more or less deflexed, apical diam. 0.5-1 mm, basal diam. 0.25-0.5 mm, densely pubescent with golden dendritictrichomes.Flowers with the calyx tube conical, 1-1.5 mm long, the lobes narrowly deltate to triangular,1-2 mm long, densely pubescent with dendritic trichomes like the inflorescence axis; corolla brightyellow-green, 0.7-1 cm diam., lobed ca. 3/4 of the way to the base, the lobes planarat anthesis, the distalportionof the abaxialsurfacedensely pubescent with dendritictrichomes;anthers2-3.5 mm long, 1-1.5 mm wide, poricidalat the tips, the poresteardrop shaped;free portionof thefilaments minute, the filamenttube ca. 0.25 mm long, glabrous;ovaryglabrous; style 4.5-5 mm long, glabrous;stigma capitate, the surface minutely papillose. Fruit a globose, green or bluish-green berry, 1-1.3 cm diam.;fruitingpedicels woody, somewhaterect, 0.5-1 cm long, slightly swollen below the calyx lobes, 2-3 mm diam. at the apex, 1-1.5 mm diam.at the base.Seeds reddish-brown,flattened-reniformwith incrassatemargins,3-4 mm long, 2-3 mm wide, the surfaces minutely pitted. Chromosome numbernot known. Distribution (Fig. 82). The upperHuallagabasin (with one collection from adjacentPasco department) in primaryforest, at 500-800(-2000) m. Specimens examined. PERU. HuANuco: E of Tingo Maria, 700 m, 24 Oct 1962, Schunke V 6193 (K, US). PASCO:Oxapampa, Cordillera Yanachaga, W slope at base of highest peaks, side rd. to NE halfway between Oxapampa and Huancabamba, ridge SE of Cooperativa 3 de Mayo, 2000-2500 m, 10?27'S, 75?26'W, 18-21 Sep 1984, Foster et al. 11240 (F, NY). SANMARTIN:Trail to Shunt&, ca. 10 km W of Tocache Nuevo, E of Rio Tocache, ca. 550 m, 8?14'S, 76?34'W, 13 Dec 1981, Plow-

162

FLORA NEOTROPICA

HERB.HORT.KEW.

-Est of Ti.rQoJar?a, 700 Im lt. shrub 2-53r

high;

flowers

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yellow green ( 2.5 2Y
5cmFIG. 81. Solanum xanthophaeum Bitter. Peru. Schunke V 61939 (K). r24o

FIG. 81. Solanum xanthophaeum Bitter. Peru. Schunke V 6193 (K).

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TAXONOMIC TREATMENT

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163

the type in all othercharacters.Foster et al. 11320 and Schunke V 6193 have the loose dendritictrichomesof the type. This differencemay be due to habitat,with the plantswith more compact trichomesgrowing in more exposed or driersites. I have chosen the G-DC sheet of Weberbauer3690 as the lectotypeofS. xanthophaeumbecausethe Morton photographis widely distributed.Neither the G-DC sheet nor the MOL sheet were annotatedby Bitter. 39. Solanum youngii S. Knapp,Novon 6: 28. 1996. ParqueNacional Type.Ecuador.Zamora-Chinchipe: Podocarpus, Loja-Zamorard. just E of pass, ca. 2800 m, 3?58'S, 79?07'W, 15 Mar 1989, Madsen 85888 (holotype,QCA;isotypes,AAU, BM, LOJA, QCNE). Figs. 74C,D, 83

Shrubs,1.5-2 m tall;young stemsandleaves densely pubescentwith matteddendritictrichomesto 1.5 mm long, the trichomesdryinga rich golden brown;stems thick,erect;barkof maturestems darkgray.Sympodial units difoliate, geminate, occasionally appearingunifoliate throughloss of the minorleaf.Leaves elliptic to widest at or just below the middle, thick and ovate, 800 fleshy, stronglybullate and corrugatedwhen dry,with 5-6 pairsof main lateralveins, sparselyto moderately pubescentwith dendritictrichomesadaxially,densely FIG. 82. Distribution of Solanum xatnthophaeunm pubescent with dendritic trichomes 1-1.5 mm long (open circles) and S. youngii (solid circles). abaxially,the trichomesdenseralong the veins, major leaves 7-27 x 4-18 cm, the apex acute,the base decurrentontothepetiole;petiole 1.2-5 cm long;minorleaves differing from the majorsonly in size, 4-8 x 2-4 cm; man et al. 11372 (F, K, MO); mouth of Rio Mishollo, L petioleca. 1 cm long.Inflorescencesoppositethe leaves, bank of Rio Huallaga, 8 Feb 1971, Schunke V 4718 simple or occasionally branched,0.5-10 cm long, 5(IBE, NY, US); rd. to Pushurumbo, 4-6 km from Palo 60-flowered, but bearing only a few open at a time, Blanco, 500-600 m, 22 Nov 1972, Schunke V 5572 (MO, with dendritictrichomeslike those densely pubescent NY); trail to Santa Rosa, R bank of Rio Mishollo, 350of the stems and leaves;pedicel scars closely spaced, 370 m, 4 Aug 1973, Schunke V 6693 (MO); trail to Shunte, E of Puente Palo Blanco, 500-800 m, 14 Jul obscuredby the dense pubescenceof the inflorescence 1974, Schunke V 7428 (MEXU, MO, MPU, US); E of axis.Budsglobose,laterellipsoid,stronglyexsertedfrom bridge over Rio Uchiza, ca. 400 m, 24 Jul 1974, Schunke the calyx tube. Pedicels at anthesis erect to somewhat V 7739 (MO); Caserio Nueva Uni6n below Puerto deflexed, thick and fleshy, 0.8-1 cm long, pubescent Huicte, R bank Rio Huallaga, 450-500 m, 1 Aug 1974, with dendritictrichomes.Flowers with the calyx tube Schunke V 7948 (MO); Palo Blanco near Fundo of broadly conical, 1-1.5 mm long, the lobes triangular Manuel Aranjo, 700-800 m, 1 Mar 1979, Schunke V and irregularlysplitting, 1-4 mm long, with a tuft of 10884 (MO); trail to Pucayacu, across from the chacra dendritictrichomesat the tips, otherwise sparselypuof Sr. Alfredo Sinarahua, ca. 400 m, 11 Jul 1980, bescent; corolla white, waxy, 1.7-2 cm diam., lobed Schunke V 11992 (IBE). ca. 2/3 of the way to the base, the lobes planarat antheLocal names. Peru. San Martin: ayac mullaca. sis, tips of the lobes with a tuft of dendritictrichomes ca. 0.5 mm long, the marginspapillose;anthers 5-6 x Solanum xanthophaeum is closely related to S. 1-2 mm, poricidalat the tips,the poresteardropshaped; malletii from Ecuador and northern Peru and differs from it in its smaller, brilliant yellow-green flowers, its free portionof thefilaments ca. 0.5 mm long, the filament tube ca. 0.5 mm; ovary glabrous;style 7-9 mm glabrous fruits, and in its golden yellow pubescence. Most of the collections of S. xanthophaeum have more long, glabrous; stigma bilobed, the surface minutely compact pubescence than that of the type, but agree with papillose.Fruita globose, greenberry,1-1.2 cm diam.;

164

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Specimens examined. ECUADOR. LOJA: Parque fruiting pedicels erect and woody, ca. 2 cm long, ca. 1.5 mm diam.at the base, ca. 2.5 mm diam.at the apex, Nacional Podocarpus, Cajanuma, trail to Lagunas del stonecells apparentlypresentin thepericarp.Seedsdark Compadre, 3100-3300 m, 4?05'S, 79?10'W, 13 Jul 1995, Alvarez & Tirado 1528 (BM); Parque Nacional Podobrown,flattened-reniform,2-3 x 1.5-2 mm, the mar- carpus, pass and just E of pass on Loja-Zamora rd., ca. gins incrassateandpaler,the surfacesminutelypitted. 13 km E of Loja, 2600-2800 m, 3?59'S, 79?08'W, 30 Chromosomenumbernot known. Oct 1994, Knapp et al. 9117 (QCNE); Parque Nacional

Distribution (Fig. 82). In wet montane and cloud forest, often in forest patches above timberline, in S Ecuadorand N Peru, at 2500-3500 m.

Podocarpus, S of Loja, Centro de Informaci6n E of Nudo de Cajanama, 2850-2950 m, 4?5'S, 79?10'W, 2122 Feb 1985, Ollgaard et al. 57903 (AAU). ZAMORACHINCHIPE: Loja-Zamora rd., near the pass, 2800-2900

165

TAXONOMIC TREATMENT m, 15 Mar 1989, Romoleroux 814 (AAU). PERU. CAJAMARCA: Prov. Cutervo, Bosque Cutervo, Parque Nacional de Cutervo, NW corner of Cordillera Tarros, Chorro Blanco sector, ca. 10 km WNW of San Andres de Cutervo, ca. 2550 m, 6?12'S, 78026'W, 3 Nov 1990, Dillon et al. 6131 (F); Chota, Bosque El Pargo, between Llama and Huambos, 3010 m, 13 Aug 1994, Leiva G. et al. 1507 (BM); Prov. Chachapoyas, roadside in jalca of Calla-calla, 3300 m, 12 Aug 1998, Sagastegui A. et al. 16102 (BM, F, HAO). SAN MARTiN: Rio Abiseo National Park, Chochos, ca. 3425 m, 7?S, 77?W, 25 Nov 1985, Young2157 (F, NY, USM); Chochos, above timberline, 3350 m, 24 Nov 1985, Young 2425 (K); along trail at lower end of Chochos valley, E side of river, 3100 m, 11 Feb 1986, Young 2650, 2664 (K); forest on edge of Laguna Chochos, 3300 m, 19 May 1986, Young3195 (K); trail between Mirador and Puerto del Monte, 3300-3450 m, 7?S, 77?W, 27 Jun 1986, Young 3833 (BM, K, NY, USM).

The stronglybullateleaves andthe dense dendritic pubescencemakethis species very distinctive.The trichomes are slender and very highly branched, and the

branchesthemselves are very short.Solanumyoungii is closely related to S. nutans, also of the Andean cloud forests, but differs from it in its looser, less tightly

branchedpubescence,bullateleaves anddifoliategeminate sympodial units. The pale coffee-colored pubescence is very similarto thatof S. malletii, but thatspecies has elongate inflorescences and auriculateleaves and is found at low elevations.

VI. Solanum amblophyllum species group (S. amblophyllum, S. barbulatum, S. laurifrons, S. psychotrioides, S. tovarii, S. vacciniiflorum, S. validinervium). Fig. 84 Shrubsor smalltreesof highelevations,oftengrowing in windswept edges of paramos;young stems and leaves glabrous (Solanum laurifrons) to densely pubescent, the trichomes simple and uniseriate.Sympodial unitsplurifoliate,or variouslydifoliate, geminate or not, if geminate,the minorleaves usually similarin size and shape to the major leaves. Leaves elliptic to obovate, often thick andcoriaceous,the apex andbase usuallyacute.Inflorescencesoppositethe leaves, or occasionally lateral or appearing terminal (S. amblophyllum,S. laurifrons),simple or several times branched;pedicel scars usually closely spaced, but in S. amblophyllumwidely spaced ca. 2 mm apart.Buds fleshy,the corollastronglyexsertedfromthe relatively short calyx tube, flattened(S. barbulatum)to pointed (S. vacciniiflorum).Flowers white, fleshy and rather large,thepetalsplanarorcampanulate(S.validinervium) at anthesis.Fruit green or yellowish, hardat maturity; fruiting pedicels deflexed or somewhat erect, woody andoccasionallyrugose.Seeds flattened-reniform, pale tan to reddish-brown. Distribution. Andean,from Colombiato southern Peru.

Dillon etal. 6131, from Cutervo, Peru, has a very large The Solanum amblophyllumspecies group is one andcomplexbranchedinflorescenceandpeculiarverru- of convenience andwill need reassessmentin the light cose berries.The leaves on this specimenarealso much of phylogeneticanalysis.The species all resembleeach largerthanthose from othercollectionsthroughoutthe other,all occurat high elevation,but areplacedhere as a species groupwith some hesitation. range of the species and the plant may be aberrant.

Key to species of the Solanum amblophyllumspecies group 1. Leaves completely glabrous. 2. Inflorescence branched, apparently terminal; leaves drying golden beneath ........................42. S. laurifirols 2. Inflorescence simple, opposite the leaves; leaves not markedly golden beneath ..............41. S. barbulatum 1. Leaves variously pubescent. 3. Sympodial units not geminate; flowers thick and fleshy. Plants growing above tree line. 4. Flowers spicily fragrant, the corolla lobes campanulate; leaves with trichomes along the veins; inflorescence simple, sparsely pubescent...................................................................... 46. S. validinenvilum 4. Flowers not fragrant,the corolla lobes planar; leaves with dense rows of trichomes on either side of the midrib, drying crepe-paper like; inflorescence resinous or glabrous ..............40. S. amblophvllum 3. Sympodial units difoliate, geminate; flowers not markedly thick and fleshy. 5. Leaves with red-papillosepubescenceon both surfaces;largertrichomestranslucent,reddish,usually branched. 6. Bark of older stems peeling; stem pubescence of trichomes with multiseriate bases; fruiting 45. S. vacciniiflor m pedicels deflexed ............................................................................................. 6. Bark of older stems not peeling; stem pubescence of uniseriate trichomes; fruiting pedicels erect ................................................................ ........................................................... 43. S. ps chot ioide s 5. Leaves pubescent. but not red-papillose; larger trichomes whitish, various. 7. Leaves uniformly pubescent over the undersurfaces; stems pubescent . .................... 44. S. tovarii 7. Leaves with trichomes only along the veins or in the vein axils; stems usually glabrous or with a few scattered hairs. 8. Buds flattened apically when young; inflorescence simple ................................41. S. barbulatum 8. Buds globose; inflorescence branched..............................................................40. S. amblophvllum

166

FLORA NEOTROPICA

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D

FIG. 84. Solanum amblophyllumspecies group. A. S. amblophyllumplant, Peru (Knapp et al. 6309). B. S. amblophyllum flowers, Peru (Knapp et al. 6309). C. S. validinervium fruit, Venezuela (Knapp & Mallet 6785). D. S. validinervium flowers, Venezuela (Knapp & Mallet 6785).

40. Solanum amblophyllum Hook. f., Bot. Misc. 2: 231. 1831. Type. Peru.Lima: Obrajillo,in the valley ofCanta, s.coll. (holotype, K). Solanumhypostichopogon Bitter,Repert.Spec. Nov.

crowded at the junctions of the midrib and the main lateralveins, leafmarginsrevolute,blades4.5-13 x 1.73 cm, with 6-10 pairs of main lateralveins, these not visible above, prominentbeneath, the apex rounded, Regni Veg. 16: 95. 1919. Type. Peru. Lima: the base acute,decurrentonto the petiole;petioles 0.6Puente del Inferniello (via Lima-Oroya), 3328 1 cm long. Inflorescencesterminal,laterbecoming latm, 12 Jul 1910, Seler 236 (holotype, B [deeral, occasionally opposite the leaves, branchedsevstroyed:F neg. 2611]). Figs. 84A,B, 85 eral times, 2-5 cm long, 10-25-flowered,butwith only Dense shrubs 1-2 m tall; young stems and leaves a few flowers at a time, glabrous or glandular-resindensely red-papillose, the papillae ca. 0.1 mm long; ous; pedicel scars irregularlyspaced 1-2 mm apart, older stems glabrous;bark of older stems dark gray- beginning ca. 1 cm from the base of the inflorescence. ish-brown.Sympodialunitsplurifoliate.Leavesnarrowly Buds flattenedapicallywhen young, laterglobose, the elliptic, coriaceous,crinkledandglabrousabove in dry corolla soon exserted from the calyx tube.Pedicels at specimens,denselypubescentalongthe midribbeneath, anthesis deflexed, thick and fleshy, 1-2 cm long, tathe trichomes uniseriate, simple, 0.5-1 mm long, pering from the base of the calyx tube to a base 1 mm

TAXONOMIC TREATMENT

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168

FLORA NEOTROPICA

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FIG. 86. Distribution of Solanulm ambloph illunm (solid circle), S. barhublatunm (open circles), and S. lawur'ifons(stars).

Specimens examined. PERU. S.loc., 1839-1840, Gay 1167 (P); ca. 1878, Martinet 277 (P). LIMA:Rio Rimac, above Matucana,3000 m, Apr 1882, Ball s.n. (K); Chancay, Naupay, 2700 m, 22 Mar 1975, Cerrate et al. 251 (USM); Canta,Pucachacanear Huaras,3500 m, 31 Aug 1952, Diaz s. n. (USM); Huarochiri, ca. 4 km E of San Mateo, ca. 117 km E of Lima on rd. to La Oroya, ca. 3600 m, 7 Jul 1981, Dillon 2513 (F, MO, NY, USM); Huarochiri,above Infiernillo, Lima-Huancayo rd., 3300-3500 m, 19 Jul 1950, Ferreyra 7702 (MOL, US, USM); Huarochiri, Infiernillo, 3300-3400 m, 21 May 1953, Ferreyra 9197 (USM); Huarochiri, above San Bartolome, 2900-3000 m, 29 May 1954, Fe-rrera 9699 (USM); Huarochiri, Rio Blanco, 3400-3500 m, 25 Jan 1963, Ferreyra 14855 (USM); 5 km NE of Suchi, ca. 56 km NE of Chosica, on rd. to Huanza, 3300 m, 6 May 1978, Gentr, 21685 (MO, USM); Huarochiribetween San Mateo and Chicla, 30003300 m, 8 Sep 1953, Hjerting & Petersen 1249 (USM); Huarochiri, quebrada de San Mateo, 26 Sep 1863, Isern 2537 Eulalia(F); Huarochiri, Chosica-Sta. Marcapomacocha rd., 117 km E of Lima, ca. 3600 m, 11?40'S, 76?25'W, 8 Mar 1984, Knapp et al. 6309 (BH, US, USM); Viso, ca. 9000 ft, 5-14 May 1922, Macbride & Featherstone 568 (F); San Mateo, ca. 1878, Martinet 590 (P); Canta, along Rio Chillon above Obrajillo, 31003300 m, 13-23 Jun 1925, Pennell 14418 (F, K, NY, US, USM); Tambaraque, ca. 3000 m, May 1938, Sandeman 250 (K); Huarochiri, Zarate, 3090-3150 m, 3 Mar 1977, Valencia 004 (USM); Canta, Huaros, 3600 m, 29 Apr 1973, Vilcapoma S. 162 (USM); Canta, above Obrajillo, 3100 m, 29 Apr 1973, Vilcapoma S. 164 (USM). Solanum amblophyllum is a shrub of open areas

at high elevations and is most similar and probably of northernPeru. It closely related to S. barbulatunm differs from that species in its plurifoliate sympodia, diam.Flo"werswith the calyx tubebroadlycup-shaped, branched inflorescences, larger flowers, and distincca. 1.5 mm long, the lobes rounded-triangular,often tive leaves. The upper surfaces of dried leaves of S. auriculate,1-1.5 mm long, the marginsthickenedand amblophyllumhave an unmistakablecrinkled, crepepaler,glabrous;corollawhite, fleshy, 1.2-1.6 cm diam., paperappearance,makingthis species easy to identify. lobed 2/3 of the way to the base, the lobes planarat The flowers are largeandfleshy but ratherinconspicuanthesis,the tips andmarginsof the lobes densely pap- ous as they hang down amongst the foliage. The triillose; anthers4-4.5 x 1-1.5 mm, poricidalat the tips, chomes on the undersidesof the leaves are quite dense theterminal0.5 mm thickenedandpaler,the porestearand their structureis difficult to determinefrom dried dropshaped;free portionofthefilaments 0.25-0.5 mm material. The trichomes are always simple and long, the filamenttubeca. 0.5 mm long;ovaryglabrous; uniseriate.The of the trichomes is distinctive position stvlestraight,6-8 mm long;stigmacapitate,brightgreen in S. amblophyllum.They occuron the midribon either in live plants, the surface minutely papillose. Fruit a side of the primarynervesof the leaf andarenot strictly globose, greenberry,1-1 .3 cm diam.;fruitingpedicels in the vein axils as in S. barbulatumand members of woody anddeflexed, 1.2-1.5 cm long, ca. 1.5mm diam. the S. nudum species group. at the base. Seeds darkbrownishin drymaterial,greenSolanumamblophyllumhas a limited distribution, ish-tan in live plants, flattened-reniform,3.5-4 x 2but is rathercommon where it occurs. In the area in 2.5 mm, the margins incrassateand somewhat paler, which I collectedthis species, it was a commonstreamthe surfaces deeply pitted. Chromosomenumber.n = side shrubjust below the snowline. Few specimens of 12 (voucherKnapp, Mallet & Smith 6309). S. amblophyllumare found in North Americanor EuDistribution (Fig. 86). On the westernslopes of the ropeanherbaria,but it is well representedin Peruvian Andesin thedepartmentof Lima,Peru,at 2500-3500 m. herbaria.

TAXONOMIC TREATMENT

41. Solanum barbulatum Zahlbr., Ann. K. K. Naturhist. Hofmus. 7: 6. 1892. Type. Peru. Cajamarca:Cutervo, 1879, Jelski 56 (holotype, W [F neg. 33048]; frag. F). Fig. 87 Solanumn ecuadorense Bitter, Repert. Spec. Nov.

Regni Veg. 16: 404. 1920. Type. Ecuador. Pichincha:Umgebungvon Quito, no collector or specimencited. Solantum ecuadorense Bitter var. nervisequum Bit-

ter,Repert.Spec. Nov. RegniVeg. 16: 406. 1920. Type. Ecuador(?), s.loc., Lehmann7817 (holoCuenca, type,B; isotype,US-n.v. ["RioUdushapa, 2000-2200 m, October"fide Nee, in litt]). Solanul ecuadorense Bitter var. modicepilosum Bitter, Repert. Spec. Nov. Regni Veg. 16: 406. 1920.

Type. Ecuador.Pichincha: environs of Quito, Jameson 457 (lectotype, BM, here designated; isolectotypes,G (Mortonneg. 8583), K). Solanuclmecuadorense Bitter var. glabriusculum Bitter, Repert. Spec. Nov. Regni Veg. 16: 406. 1920.

Type. Ecuador.Pichincha: Environs of Quito, weedy areas, 2800 m, Jameson 522 (lectotype,

BM, here designated). Solannumiecuadorense Bitter var. ilinizae Bitter,

Repert. Spec. Nov. Regni Veg. 16: 406. 1920. Type. Ecuador.Pichincha: Vulcan Iliniza, auf Tracht, 2800-4400 m, Dec, Moritz Wagner s.n

(holotype, M [Mortonneg. 8686]). Solannumamblophylltnl of J.F. Macbr., Publ. Field

Mus. Nat. Hist., Bot. Ser. 13 (5B): 196. 1962. Pro parte,not of Hook. f. Shrubs 1.5--5 m tall; young stems and leaves glabrousand shiny,occasionally with a few uniseriatetrichomes ca. 0.3 mm long, the young leaves red-papillose; bark of older stems shiny and whitish or gray. Sympodial units difoliate, geminate. Leaves elliptic, widest at the middle, glabrous above, glabrousto pubescent with uniseriatesimple or branchedtrichomes along the veins and in the vein axils beneath, the trichomes occasionally extending to the lamina; major leaves 2.5-13 x 0.8-3.7 cm, with 8-13 pairs of main lateralveins, these often strongly parallel, prominent above andbeneath,the apex acute, the base acute;petioles 2-5 mm long; minor leaves differing from the majorones only in size, 1.3-3 x 0.6-1.4 mm, the apex acute, the base acute;petioles 2-3 mm long. Inflorescences oppositethe leaves, simple,0.5-2 cm long, 2-8flowered,usuallyglabrous,but occasionallywith a few uniseriatetrichomes along the rhachis;pedicel scars closely spacedbutnot overlapping.Buds flattenedperpendicularto the pedicel when very young, later globose, the corolla soon exserted from the calyx tube. Pedicels at anthesis deflexed, 0.8-1.3 cm long, tapering from the base of the calyx tube to a slender base 0.5-0.75 mm long.Flowers with the calyxtubebroadly cup-shaped,ca. I mm long, the lobes broadlydeltoid,

169

fleshy, appearingwoody in dry specimens, 0.5-1 mm long, glabrous and shiny; corolla white, fleshy, 0.81.2 cm diam.,lobed2/3 of the way to the base, the lobes planaror somewhat deflexed at anthesis, the tips and marginsof the lobes densely papillose; anthers 2.5-4 x ca. 1 mm, poricidal at the tips, the terminal0.5 mm thickened and paler, the pores teardropshaped; free portionof thefilaments0.05-0.1 mm long, the filament tube 0.25-0.5 mm long; ovary glabrous;style straight, 5-6 mm long; stigma a widened area on the tip of the style, the surfaceminutely papillose. Fruit a globose, green berry,1-1.5 cm diam.;fruitingpedicels woody, erector deflexed, 1-1.5 mm long, ca. 1 mm diam.at the base. Seeds darkbrown, flattened-reniform,2-4 x 23 mm, the margins incrassateand paler, the surfaces minutely pitted. Chromosomenumbernot known. Distribution (Fig. 86). In the Andes of southern Colombia,EcuadorandnorthernPeru,at2500-4000 m. Selected specimens examined. COLOMBIA. CAUCA:Paramo de San Pedro, near Totor6, 3200 m. 15 Jan 1969, Espinal T. & Ramos 3407 (F). ECUADOR. Interandine highland, 2800 m, Rimbach 16 (F). AZUAY:Las Cajas near Laguna Llariuco, 3100-3200 m, 2?49'S, 79?11'W, 21 Sep 1983, Boylsen Larsen & Eriksen 45092 (AAU); Cuenca, Rio Udushapa, 2000-2200 m, Oct, Lehmann 7817 (K). BOLIVAR: Guaranda-Riobamba rd., 2800-3200 m, 27 Feb 1987, Zak 1786 (NY); Chillanes-Bucay rd., Hacienda "Tiquibuso," 2100 m, 1?50'S, 79?05'W, 29 Aug 1987, Zak & Jaramillo 2560, 2576 (K, MO, NY), 3 Sep 1987, Zak 2720 (NY), 10 Sep 1987, Zak 2867 (NY). CANAR: Alausi-Cafar rd., detour to Oyashig and Hcda. El Carmen, N of Cafiar, 3270-3700 m, 12 Aug 1987, Zak 2392, 2419 (AAU, NY); El Tambo-Zhud-Oyashig rd., between Zhud and Oyashig, 3200-3300 m, 13 Aug 1987, Zak 2432 (NY); Bosque La Carbonaria, CaiarComunidad Llurac-Rumi rd., 3000 m, 13 Aug 1987, Zak 2435, 2436 (AAU, NY); Moloboc Grande-Molon Ventana rd. off Cafiar-Azoguez rd., between Moloboc and Molon Ventana, 3200-3450 m, 14 Aug 1987, Zak 2438, 2448 (AAU, NY). CARCHI:El Angel, San IsidroChulte rd., 3200 m, 0?36'N, 78?02'W, Palacios & Rubio 7003 (QCNE). CHIMBORAZO:S slopes of Volcan Chimborazo, near Hacienda Chimborazo, 3400 m, 27 Nov 1980, Balslev 1021 (AAU); Riobamba-CubijiesPuela-Bafios-Ambato rd., 2400 m, 19 Feb 1987. Zak 1645 (NY); Parque Nacional Sangay, between Pugala and Alao on Riobamba-Chambo-Pungala-Alao rd., 3100 m, 8 Aug 1987, Zak 2352, 2353a (NY), 9 Aug 1987. Zak 2357 (NY); Pallatanga-Yungilla-Llimbe rd., between Yunguilla and Llimbe, 2300-2500 m, 8 Sep 1987. Zak 2835 (NY). COTOPAXI:Around Pilalo, 2400 m, 0?57'S, 79?02'W, 3 Jul 1968, Holm-Nielsen & Jeppesen 1283 (AAU, NY); between Tambo de Le6n and km 60. 28502900 m, 3 Jan 1987, Jaramillo 9381 (NY); LatacungaAngamarca,vie. Angamarca, 3100-3600 m, 17 Feb 1987, Zak 1611 (NY). IMBABURA: Res. Ecol. Cotacachi-Cayapas.

170

FLORA NEOTROPICA

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TAXONOMIC TREATMENT171 Laguna de Cuicocha, isolte Teodoro Wolff, 3100-3300 Podocarpus, Loja-Zamora rd. just E of pass, 2800 m, m, 0?18'N, 78?22'W, 30 Aug 1991, Penafiel et al. 315 3?58'S, 79?07'W, 15 Mar 1989, Madsen 85897 (AAU, (QCNE); Otovalo-Lago San Pablo track, 2800 m, 0?13'N, LOJA); new rd. from Loja to Zamora, river banks, 200078 13'W, 20 Sep 1984, J0rgensen & Vive 56029 (AAU); 2100 m, 10 Mar 1989, Romoleroux 719c (AAU). Ibarra-Mariano Acosta rd., 2700-3000 m, 1 Nov 1986, PERU. AMAZONAS:Chachapoyas, above LeimeZak 1409, 1410 (NY); Pimanpiro-Chuga-Palmar Chico bamba, km 422, 2700-2800 m, 20 Aug 1963, Ferreyra rd., between Chuga and Palmar Chico, 2800-3000 m, 16 & Acleto 15272 (US, USM); Bongara, 4 km N of Jan 1988, Zak 3358 (NY). LOJA: Parque Nacional Podo- Pomacochas on rd. to Rioja, trail down gorge to W of carpus, Cajanuma, trail to El Mirador, 2800-3200 m, rd., 2150-2200 m, 5?40'S, 77?52'W, 2 Jun 1986, Knapp et al. 7480, 7510 (MO, USM); Leimebamba, 2400 m, 27 4?06.97'S, 79?10.34'W, 20 Oct 1994, Knapp et al. 9050 (QCNE); Parque Nacional Podocarpus, pass and just E Dec 1962, Woytkowski7825 (MO); Bongara, Jalca zone of pass on Loja-Zamora rd., ca. 13 km E of Loja, 2600along Shipasbamba-Pomacocha trail, 2200-2400 m, 29 2800 m, 3?59'S, 79?08'W,30 Oct 1994, Knapp et al. 9119 Jun 1962, Wurdack1116 (NY); Chachapoyas, summit of (QCNE); Parque Nacional Podocarpus, E of Nudo de Cerro Malcabal (Cerro Tumbe), 3-6 km SW of MolinoCajanuma, just N of Centro de Informaci6n, 2900 m, pampa, 2850-2900 m, 20 Jul 1962, Wurdack 1423 (F, 4?05'S, 79?10'W, Madsen 85753 (AAU, QCNE); Parque NY, US). CAJAMARCA:Contumaza, Cachil, CascasNacional Podocarpus, above Nudo de Cajanuma, trail Contumaza, 2200 m, 24 Jun 1982, L6pez M. et al. 9085 to Mirador,2900-3000 m, 4?05'S, 79?10'W, 23 Feb 1989, (BH, MO); San Miguel, Cerro Quill6n, 3150 m, 5 Jul Madsen 85817 (AAU, LOJA, QCNE); Parque Nacional 1986, Mostacero L. et al. 1280 (F, HUT, MO, NY); San Podocarpus, vic. Lagunas de Compadre, ca. 6 hrs walk Miguel, Quishuarpampa(El Tingo-Jalca de las Estacas), 2950 m, 12 May 1977, Sagcstegui A. et al. 8831 (HUT, from Centro de Informacion, 3000-3400 m, 4?10'S, 79007'W. 21 Nov 1989, Madsen & Pedersen 86428 MO); El Tingo (Agua Blanca), 2750 m, 20 Jun 1980, E slopes of paramos de Sagdstegui A. et al. 9512 (HUT, NY); vie. Cajabamba, (AAU). MORONA-SANTIAGO: Matanga ca. 3-40 km S of Sigsig on rd. to Gualaquiza, 2600 m, 18 Feb 1983, Sagdstegui A. et al. 11258 (HUT, 2800-3100 m, ca. 3025'S, 78?48'W, 22 Jan 1985, Luteyn NY); Challuayaco, between La Encanada and Celendin, 3520 m, 3 Jul 1975, Sinchez Vega et al. 1697 (NY); & Cotton 11200 (AAU, NY). NAPO: Reserva Ecol6gica Bajada de la Totorilla, along course of Rio Shitamalca, Cayambe Coca, 5 km from Cuyuja, rd. to Quito, 3150 3450 m, 31 Oct 1992, Sdnchez Vega & Torres6413 (BM, m, 0?22'S, 77?56'W, 30-31 May 1991, Gavilanes et al. F); Chota, pass S of Conchan, 2500 m, 7 Dec 1938, Stork 665-B (AAU), Gavilanes et al. 665-D (AAU); Julio Andrade-San Francisco-Sta. Barbara rd., 2600-3000 m, & Horton 10066 (F, K). HUANUCO:12 mi S of Panao, ca. 10,000 ft, 4-10 Jul 1922, Macbride & Featherstone 28 Dec 1986, Zak 1570 (AAU, NY). PICHINCHA:Vic. 2209 (F); Mufia, trail to Tambo de Vaca, ca. 8000 ft, 5Quito, Guapulo, 2650 m, 26 May 1939, Asplund 6321 7 Jun 1923, Macbride 4274 (F). JUNiN: 26 km NE of (K, NY); slope of Mt. Pichincha above Quito, 3200 m, 2 May 1955, Asplund 16164 (K, US); Cerro Pasochoa, Tarma on rd. to Oxapampa, along Rio Palca, 8850 ft, 12 Nov 1979, Jones & Davidson 9055 (NY); above W slopes, 0?28'S, 78?32'W, 14 Aug 1982, Balslev 2790 Oct 1943, Sandeman 4471 (K). LA (NY, QCA); Potrerillo, Pululagua, ca. 7700 ft, 17 Apr Huacapistana, LIBERTAD: Hacienda Cochabamba, R bank Rio 1951, Bell 573 (BM, NY); Quito-Baeza km 29, 2835 2600 m, 26 Jun 1958, Ldpez & Sagdstegui 4. m, 26 Nov 1976, Boeke & McElroy 310 (NY); rd. from Chusg6n, 2786 (HUT). PIURA: Cuello del Indio, CanchaqueMagdalena to Lloa, 2850 m, 3 Mar 1927, Firmin 11 Huancabamba rd., 3600 m, 2 Sep 1976, Sagastegui .4. E of 3200 23 Jan rd., (US); Quito-Lago Agrio Pito, m, & Cabanillas S. 8565 (BM, HUT). SAN MARTIN:Valley 1980, Gentry 30647 (MO, NY); trail from Nono to San of Rio Apisoncho (Abiseo), 30 km above Jucusbamba, 2804 14 Mexia 7689 Francisco, m, Sep 1935, (K, U); 3600 m, 7?55'S, 77?10'W, 27 Aug 1965, Hamilton & Chillogallo-San Juan-Chiriboga-Empalme rd., vic. San 164 (K); Rio Abiseo National Park, near Gran Holligan 3100-3260 Juan, m, 0?18'S, 78?39'W, 10 Sep 1985, Zak 2700 m, ca. 7?S, 77?W, 22 Jul 1985, Young ruins, Pajaten & Jaramillo 654 (K, MO, NY); Quito-Nono-Tandayapa1240 (NY); Rio Abiseo National Park,near La Playa camp, San Miguel de los Bancos rd., between Tandayapaand Sta. 2650 m, ca. 7?S, 77?W, 30 Aug 1985, Young1513 (NY); Rosa, 2000-2400 m, 17 Jan 1987, Zak 1594 (NY); El Pongo, Rio Abiseo National Park, across river from La Playa 2800 m, 0?15'S, 78040'W, 24 Jul 1987, Zak & Jaramillo 2600 m, ca. 7?S, 77?W, 1 Sep 1985, Young 1534 camp, 2255 (NY), 25 Jul 1987, Zak & Jaramillo 2565 (NY); (NY); Parque Nacional Rio Abiseo, Puerta del Monte, Quito-Chirboga-Emplame rd., side rd. to San Juan-San 3200 m, ca. 7?S, 77?W, 1 Sep 1985, Young 1568 (NY). Jose de la Victoria, near San Jos&de la Victoria, 30003400 m, 24 Dec 1987, Zak 3259 (NY). TUNGURAHUA: Local names. Ecuador. Loja: mataperro; Peru. San Jose de Poal6, watershed of Rio Pisayambo, Cajamarca: choloque de perro. Yanayacu, 2600 m, 1?06'S, 78?29'W, 11 Feb 1991, Ceron Solanum barbulatum is similar morphologically to 13273 (QCNE); Hacienda San Antonio, near Bafios, Dec 1937, Sydow 565 (US). SUCUMBIOS:ENE of Cayambe both S. psychotrioides (Colombia) and S. amblophyllum Mtn., ca. 3200 m, 15 Dec 1961, Cazalet & Pennington (Peru) but differs from those species in its small flow5591 (K). ZAIORA-CHINCHIPE:P.N. Podocarpus, trail ers and short fruiting pedicels. The trichomes in the vein to Mirador, 2800-3200 m, 4?06.97'S, 79?10.34'W, 20 axils of S. barbulatum are thinner and weaker than those Oct 1994, Knapp et al. 9050 (QCNE); Parque Nacional of S. psychotrioides and usually dry white. The bark of

172

FLORA NEOTROPICA

S. barbulatumis usually brightwhite in dry specimens W of Pasto,ca. 2800 m, Oct 1950,Espinosa3062 (NY). and appearsto peel off in thin sections from the stem. Solanum laurifronsis an isolated species in sect. Populationsof S. barbulatumare more pubescent in Geminata,with no apparentclose relatives.It is superPeru and progressively become more glabrous into ficially similarvegetativelyto S. goniocaulonin its glaColombia,but much variationexists and populational brousleaves that dry golden yellow beneath,but is unsamples are not known. relatedto thatspecies and differs from it in possessing flattened-reniform seedsandsympodialunitsthatarenot Solanumlaurifronsis anomalousin this spegeminate. 42. Solanum laurifrons Bitter, Repert. Spec. Nov. cies groupin being completelyglabrous,butshareswith Regni Veg. 16:102. 1919. Type.Colombia.Narifo: the other species of the groupfleshy buds and flowers, exkursionvon Pastozum Rio Patiain derUmbebung flattened-reniform seedsanda tendencyto possesselabovon Pefal, Calderaoder Weg nach Guascaviga, 14 rateinflorescencesandplurifoliatesympodia. Dec 1869, Stibel 402 (holotype, B [destroyed:F neg. 2666]; frag., F). Shrubsof unknownheight;young stems and leaves glabrous, the stems 4-angled and erect; barkof older stems greenish-brown.Sympodiadifoliateor trifoliate, not geminate.Leaveselliptic-obovate,widestjust above the middle,coriaceouswith revolutemargins,glabrous and shiny adaxially,glabrousanddryingmattegolden abaxially, 6-10 x 2-3.5 cm, with 6-7 pairs of main lateral veins, these drying yellow abaxially, the apex acute, the base attenuate;petioles 0.5-0.7 cm long.Inflorescences oppositethe leaves or appearingterminal, 4-10 cm long, many-timesbranched,20-50-flowered, glabrousandshiny;pedicel scars unevenlyspaced0.51.5 mm apart.Buds globose when young, laterelliptic, the corolla strongly exserted from the calyx tube. Pedicels atanthesis filiform,0.8-1.1 cm long, deflexed or erect,ca. 0.5 mm diam., glabrous.Flowers fragrant, with the calvx tube conical, 1-1.5 mm long, the lobes quadrate,0.5-1 mm long, the margins thickened and white in dry specimens, glabrous;corolla white, 1-1.2 cm diam., lobed nearly to the base, tips of the lobes cucullateand minutelypapillose;anthers3-4 x 1-1.5 mm, poricidal at the tips, the pores teardropshaped; free portionofthefilaments ca. 0.5 mm long, the filament tube ca. 0.5 mm long; ovary glabrous;style 5-6 mm long, glabrous;stigma merely a broadenedpapillose areaon the apex of the style.Fruit a globose, green (mustardyellow when dry) berrywith thick pericarp, 1-1.2 cm diam.;fruitingpedicels woody, deflexed or partiallyerect, 2-2.5 cm long, 1-1.5 mm diam. at the base.Seeds pale tan,flattened-reniform with incrassate x 4-4.5 3.3.5 the surfaces mm, margins, minutelypitted. Chromosomenumbernot known.

43. Solanum psychotrioides Dunal, Solan. Syn. 21. 1816.Type.Colombia.adfluviumMagdalenae(prope Tenerife et Mompox fide HBK), Humboldt & Bonplands.n.(holotype,P-Bonpl. [microficheIDC 6209-2:61 I1.2,F neg. 39013, Mortonneg. 22329]; isotype, P [Mortonneg. 8303]). Fig. 88 SolanumgracilescensC. V. Morton,Contr.U. S. Natl. Herb.29: 52. 1944.Type.Colombia. Cauca:Carpinterias,between peaks of Munchique and Altamira,

2450-2500m, 15 Jul 1939,Perez-Arbelhez & Cuatrecasas 6170 (holotype, US; isotype, F [F neg. 49460]).

Shrubsto trees, 3-10 m tall, to 8 cm diam.;young stems and leaves densely red-glandularpapillose, the trichomes (papillae) less than 0.1 mm long, roundish, persistenton olderleaves; stems soon glabrate;barkof older stems rugose and transverselyexfoliating, pale gray.Sympodialunitsdifoliate,occasionallygeminate. Leaves elliptic to obovate, usually not geminate, widest at the middle orjust above, red glandularpapillose on both surfaces, occasionally with a few uniseriate trichomesalongtheveins above,the trichomesbeneath 0.5-0.7 mm long, usually branched,along the veins and in the vein axils, occasionally extending to the laminabeneath,blades3-13 x 2-5 cm, with 8-11 pairs of main lateralveins, these impressed above, prominentandreddishbeneath,the apexacute,thebase acute; petioles 0.7-1.5 cm long; minor leaves when present, differingfromthe majorones only in size, 2.5-4 x 1.52 cm, the apex acute, the base acute;petioles 2-5 mm long. Inflorescences opposite the leaves, simple, 0.51.5 cm long, 2-7-flowered, glandularpapillose, often withuniseriate,usuallybranchedtrichomes0.5-0.7 mm Distribution (Fig. 86). In the southernColombian long alongthe axis;pedicel scars closely spaced,large, Andes,departmentsof HuilaandNarifo, 2800-3000 m. corky,and overlapping,beginningnearthe base of the Cordi- inflorescence;Budsflattenedapicallywhen veryyoung, Specimensexamined.COLOMBIA.HUILA: llera Oriental, 1 km NW and W of Finca Venatanes, 15 later globose, just before anthesis ovoid. Pedicels at anthesisthickandrugoselike the stems, 1-1 .2 cm long, km SE of Gigante, 9000 ft, 14 Sep 1944, Little 8648 deflexed, taperingfrom the base of the calyx tube to a (US). NARINO: Paramo near Naripo, W of Pasto, ca. 3000 m, Oct 1950, Espinosa 3061 (NY); cerca de Narifo, base ca. I mm diam.Flowers with the calyx tube open

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174

andbroadlycup-shaped,1-2 mm long,thelobesbroadly deltoid, 0.5-2 mm long, ratherirregularin shape, red glandularpapillosewith uniseriatetrichomeslike those of the rest of the inflorescence,the entirecalyx appearing woody in dry material;corolla white, fleshy, 1.62.2 cm diam.,lobed nearlyto the base, the lobes planar at anthesis, the tips of the lobes cucullate, the tips and marginsof the lobes densely papillose; anthers 4-6 x 1.5-2 mm, poricidal at the tips, the terminal0.5 mm thickened and paler, the pores teardropshaped; free portion of thefilaments 0.5-1 mm long, the filament tube 1-1.5 mm long; ovary glabrous;style straight,89 mm long; stigma a terminalthickening on the style, minutely papillose. Fruit a globose, green berry, 1.41.7 cm diam.;fruitingpedicels erect and woody, conspicuously rugose, 2-3 cm long, ca. 2 mm diam. at the base. Seeds not known (only one specimen with fruit; seeds appearratherlargethroughthe fruitwall). Chromosome ntimbernot known.

between Santa Maria and paramo de San Antonio, 29003180 m, 1 Jun 1946, Schultes & Villarreal 7793 (K, NY). VALLE DE CAUCA: Cordillera Los Farfallones, Occidental, E slope, Alto de Buey, 3450-3300 m, 13 Oct 1944, Cuatrecasas 18063 (F); Cordillera Occidental, Los Farfallones,extreme N, E slope Alto del Buey, 3300-3450 m, 13 Oct 1944, Cuatrecasas 18078 (F); Cordillera Central, W slope, Rio Bugalagrande, Barragan, paramo de Bavaya, Corrales, 3450-3520 m, 18-20 Mar 1946, Cuatrecasas 20218 (F); Cordillera Central, W slope, Rio Bugalagrande, ridge of Barragan, 3250-3270 m, 12 Apr 1946, Cuatrecasas 20606 (F); Cordillera Occidental. Los Farfallones, Quebrada de Nieves, below La Diamante mine, 2900 m, 30 Jul 1946, Cuatrecasas 21795 (F): Las Amarillas, El Cairo-Rio Blanco rd., Choc6-Valle border at base of Cerro Ingle6s,Cordillera Occidental, Serrania de los Paraguas, 2060-2125 m, 29 Mar 1988, Silverstone-Sopkin et al. 3762 (CUVC, NY). Solanum psychotrioides is probably most closely

relatedto S. vacciniiflorumof high-elevationCostaRica, Distribution (Fig. 90). In the CordilleraOcciden- and to S. validinerviumof high-elevation Venezuela. The trichomeson all threeof these species are of simital and Centralof Colombia, at 2500-3000 m. larmorphologyandstructure:they areoften branched, Specimens examined. COLOMBIA.S.loc., Mutis have weak walls, and are reddishwhen dry.Solanium 1984, 2002 (US). ANTIOQUIA:Jardin-Rio Sucio rd., 13 psychotrioides differs from those species in its dense km fromJardin,2100-2400 m, 5?35'N,75?52'W,8 Jun indumentof papillose trichomes on the new growth, 1987, Callejas et al. 3843 (IBE, MO, NY), Callejas et al. and in its elongate, woody, rugose fruiting pedicels. 3850 CAUCA:Cordillera W (IBE, MO, NY).

Central,

slope,

headwatersof Rio Palo, Quebradadel Raio L6pez,Alto del Duende, 3300-3500 m, 2 Dec 1944, Cuatrecasas 18912 (F); Cordillera Central, headwaters of Rio Palo, quebrada de Santo Domingo, 2700-2800 m, 11 Dec 1944, Cuatrecasas 19147. 19147a (F, NY, US); Cordillera Central, E slope, Moscopan, Rio San Jose, Aguabonita. 2280 m, 30-31 Jan 1947, Cuatrecasas 23538 (F); Popayan to Totor6 rd., Alto de Angosi, 11 km from Totoro, 3200 m, 18 Jul 1948, Garcia Barriga & Hawkes 12736 (US); rd. to La Plata, from Purac6 to Alto de San Rafael, 2660-3450 m, 21 Jul 1948, Garcia Barriga & Hawkes 12851 (US); Popayan, 2800-3100 m, FebMar, Lehmann 6986 (K); km 40-41 NE of Uribe on rd. to Parque Nacional Munchique, 2560 m, 23 Apr 1969, Lute,yn et al. 7384 (MEXU, NY); Cordillera Central, Paleterato Calaguala,3000-3200 m, 17 Jan 1922, Pennell

Solanum psychotrioides is variable in pubescence, with

some populationspossessing only papillatetrichomes, andotherspossessingbothpapillateanduniseriate,both branchedandsimple,trichomeson the new growthand leaf undersides.As with many of the species of sect. Geminata,the usefulness of trichomecharactersis difficult to assess; trichomes are not immediatelyuseful in identificationof some particularlyvariablespecies. The type locality of Solanum psychotrioides is along

the banksof the lower Rio Magdalenain lowland Colombia,butno specimensmatchingthe type have ever been collected fromnearthat locality. The type specimen itself does not have the morphologyof a lowland memberof the section but is more typical of high-elevationplants.Therecentcollectionmostclosely match7100 (NY,US); MountSantaAna,CordilleraOccidental, ing the type specimen is Pennell 7449 fromthe Cordi2700-3000 m, 29--30 Jun 1922, Pennell 7449 (K, US); lleraOccidentalin the departmentof Cauca.Humboldt Las Escalaretas, Moras valley, Rio Paez basin, Tierra and Bonplandmay have used composite localities and Adentro, 2500-3000 m, Feb 1906, Pittier 1370 (US). did visit some highland areas during this portion of CUNDINAMARCA:CordilleraOriental, summit of Cord. de theirvoyage (Sprague,1926). The locality on the label Heliconia, 13 km SE of Gachala, 15 km NW of Medina, of the type specimen is most likely in error. 2930 m, 23 Sep 1944, Grant 10246 (NY); El Cedro, 2600 m, 17 Jul 1943, Huertas & Camargo 750 (F). PUTUMAYO: Valle de Sibundoy, 1 km NW Sibundoy, 2500 m, 29 May 1963, Bristol 1081 (US); El Portachuelo between Valle de Sibundoy and Mocoa, 2600 m, 30 Dec 1940, Cuatrecasas 11476 (F); Rio Putumayo, paramo San Antonio del Bardonillo to Santiago de Sibundoy, 2800 m, 4 Jan 1941, Cuatrecasas 11788 (F, US); rd. from Sibundoy to Pasto,

44. Solanum tovarii S. Knapp,Novon 2: 348. 1992. Type.Peru.Huancavelica:HaciendaTocasbetween Colcabambaand Paucarbamba,3000-3100 m, 10 Apr 1954, Tovar 1932 (holotype, USM; isotype, IBE). Fig. 89

TAXONOMIC

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176FLORA

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cal, 1.5-2.5 mm long, the lobes quadratewith minute apiculae,sparselypubescentwith simple,uniseriatetrichomes, the apiculae with tufts of simple trichomes; corolla white, waxy, 1.3-1.5 cm diam.,lobed 3/4 of the way to the base, the lobes planarat anthesis,the tips and marginsof the lobes densely papillose;anthers4-4.5 x 1.5-2 mm,poricidalatthetips,theporesteardropshaped; free portionof thefilamentsca. 0.25 mm long, the filamenttubeminute,oftenabsent,glabrous;ovaiy sparsely pubescentwith simple uniseriatetrichomesca. 0.5 mm long;style ca. 6 mm long, pubescentat the base;stigma capitate,the surfaceminutelypapillose.Fruita globose, green berry (immature),sparsely pubescent;fruiting pedicels woody,erect,ca. 1.5cm long.Seedsnotknown. Chromosomenumbernot known. Distribution (Fig. 90). In high-elevation savanna areasin the Peruviandepartmentsof Huancavelicaand Junin,at 2700-3300 m. Specimens examined. PERU.

FIG.90. Distribution of Sol s80otioide

HUANCAVELICA:

Tayacaja, Salcabamba, 3250 m, 7 Jan 1939, Stork & Horton 10273 (F, K). JUNiN: Carpapataabove Huacapistana, 2700-3200 m, 7 Jun 1929, Killip & Smith 30680 (NY, US); Mt. Pariahuanca, Matthews 1155 (K).

The placement of Solanum tovarii in the S. amblophyllumspecies groupis provisional,as the fruits FIG. 90. Distribution of Solanum psychotuioides and seeds of this species are not known. Solanum (solid circles), S. tovarii (open circles), S. vacciniiflorum tovarii is morphologically similar to the much more (stars), and S. validinervium (solid squares). widespread S. barbulatumand differs from it in its quadratecalyx lobes, largerflowers, and in its evenly distributedabaxial leaf pubescence. In S. tovarii the Shrubs 1-2 m tall; young stems and leaves densely trichomesareoverthe entireabaxialsurface,while those pubescent with soft, uniseriate, simple (occasionally of S. barbulatumare confined to the main vein axils. branched)trichomes 1-2 mm long; barkof older stems Solanum tovarii is also superfically similar to two greenish-brown,glabrate. Sympodial units difoliate, anomalous specimens (Boeke 892 from Napo, Ecuageminate.Leaves elliptic, thickandsomewhatleathery dor,andNunez&Arque8325 fromCuzco, Peru)which with revolute margins, sparsely pubescent with may representa new taxon (see S. callianthum in S. uniseriate trichomes like those of the young growth nigricans species group).These specimensdiffer from adaxially,the trichomes denser along the main veins, S. tovarii in theirunifoliate sympodial units and elonthe adaxialepidermislarge-celledandcrystalline,more gate, branchedinflorescences. densely pubescentwith uniseriate,simple or branched trichomesabaxially;majorleaves 4-8.5 x 1.5-3.5 cm, with 7-10 pairsof main lateralveins, these dryingyel- 45. Solanum vacciniiflorum Standl. & L. 0. Willlowish, the apex acute, the base acute; minor leaves iams,Ceiba 1:247. 1951.Type.CostaRica.Cartago: differing from the majorsonly in size, 1.5-4.5 x 0.5Bog "ElJardin,"Cerrode las Vueltas,Cordillerade 2.1 cm, the apex acute, the base acute;petiole 2-4 mm Talamanca,2700 m, 30 Mar 1949, Williams16114 long. Inflorescences opposite the leaves, simple, 2-4 (holotype, US; isotype, F). Fig. 91 mm long, densely pubescentwith uniseriatetrichomes like thoseof the stemsandleaves,3-5-flowered;pedicel Largeshrubs to trees, largerin sheltered habitats, scars densely packed, overlapping.Buds globose, the 3-25 m tall;young stems andleaves glabrousor pubescorollaexsertedfromthe calyx tube.Pedicels at anthe- cent with uniseriatetrichomes0.25-0.75 mm long; the sis tapering,0.7-1 cm long, deflexed, sparselypubes- stems occasionally with gnarledtrichomeswith multicent with simple, uniseriatetrichomeslike those of the seriatebases ca. 6 cells wide, 0.25-0.5 mm long, thetips inflorescence,ca. 0.5 mm diam.at the base, 1 mm diam. uniseriate,0.25-0.5 mm long, these hairssoon deciduat the apex. Flowers with the calyx tube broadlyconi- ous; olderstems with the epidermallayerpeeling trans-

TAXONOMIC TREATMENT

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178

FLORA NEOTROPICA

versely in one-cell-layer thick sheets, in dry material appearingwhitish; barkof trunksgrayish.Sympodial units difoliate, geminate. Leaves elliptic to narrowly elliptic, widest at the middle, somewhatbullatein live plants, the margins revolute, the lamina glabrous or minutelyred-papillosealongthe veins above,occasionallywitha few scattereduniseriatetrichomeson theupper leaf lamina,below sparselyto densely pubescentwith weakuniseriatetrichomesprimarilyalongthe veins, but occasionallyon the lamina,the trichomessmall-celled, quiteweak and curling,reddish,0.5-1 mm long, occasionally branched;majorleaves 6-17 x 2-6 cm, with 10-22 pairs of parallelmain lateralveins, these raised above, prominentandreddishbelow, the apex acuteto acuminate,the base acute; petioles 0.6-2.2 cm long; minorleaves differingfromthe majorones only in size, 3-9 x 0.8-4 cm, the apex acute to acuminate,the base acute;petioles5-9 mm long.Inflorescencesoppositethe leaves, simple or occasionally severaltimes branched, 0.8-10 cm long, 2-50-flowered, glabrous or with uniseriatetrichomeslike those of the young stems and leaves; pedicel scars on short inflorescences closely spaced,butnot overlapping,on long inflorescencesca. 1.5 mm apart.Buds ovoid, the corolla soon exserted from the calyx tube, larger buds ovoid with the tip pointed,glabrousor with a few scattereduniseriatetrichomes. Pedicels at anthesis deflexed, 1.2-2 cm long, taperingfromthecalyx tubeto a slenderbase ca. 0.5 mm diam.Flowerswiththecalyxtubebroadlycup-like,0.51 mmlong,thelobesbroadlyrounded-deltoid, 0.5-1 mm long, occasionally with a few uniseriate trichomes, more often glabrous;corolla white, fleshy, 1.5-1.8 cm diam.,lobednearlyto the base,the lobesplanarat anthesis, the tips and margins of the lobes minutely papillose; anthers 4.5-5 x 1-1.5 mm, poricidal at the tips, the pores teardropshaped;free portionof thefilaments minute,ca. 0.2 mm long, the filamenttube ca. 0.5 mm long; ovayvglabrous;style straight,in shortstyle flowers ca. 4 mm long, in long style flowers ca. 8 mm long; stigma small-capitate,minutelypapillose.Fruit a globose, apiculate,green or yellowish-brownberry,ca. 1 cm diam.;fruitingpedicelswoody, deflexed, 1.8-3 cm long, 1-1.5 mm diam. at the base. Seeds pale reddishtan, flattened-reniform, ca. 4 x 3 mm, the margins incrassate,the surfacesminutelypitted. Chromosome number:n = 12 (voucherKnapp 790). Distribution (Fig. 90). In the cloud forests of montaneCosta Rica and Panama,at 2000-3000 m. Selected

specimens

examined.

COSTA RICA.

ALAJUELA: Volcan Poas, 16 Apr 1973, Gentry & Burger

2954, 2858 (CR, F, K, MO); between Poasito and Volcan Poas, 2600 m, 2600 m, 19 Mar 1963, Jimenez 466 (F); Vara Blanca. 1800-2000 m, 6 Sep 1964, Lems 64090605 (NY); Volcan Poas ca. 20 km NNW of Alajuela, 2700 m,

18 Aug 1967, Taylor4508 (MO, NY). CARTAGO: Turrialba, Peralta, 2500-2600 m, 11 Sep 1965, Bernardi 10533 (NY); Cordillera de Talamanca, Finca El Sitio, La Catarata trail, 2650 m, 9?34'30"N, 83?41'10"W, 2 Mar 1996, Gamboa & Picado 345, 355, 381 (INB); RF Los Santos, Sauvegre drainage, Est. Ojo de Agua, 3000 m, 9?37'16"N,83?49'43"W,26 Jun 1997, Gamboa 1586 (INB); finca ca. 0.5 km E of InteramericanHwy. ca. 20 km SE of El Empalme, ca. 2500 m, 15 Jul 1970, Lellinger & lWhite 1153 (F). HEREDIA: 2-2.5 km N of rd. terminus to Volcan Barba, 2600-2700 m, 16 Apr 1971, Almeda 436 (F); El Roble, Volcan Barba, 2750 m, 8 Apr 1951, Le6n 3273 (CR); P.N. Braulio Carillo, 2600 m, 10?08'00"N, 84?07'00"W, 15 Mar 1995, Fernandez 1540 (INB); P.N. Braulio Carrillo, Cerro Guariri trail, 2600 m, 10?07'n, 84007'W, 8 Oct 1989, Rivera 101 (INB); near Los Cartagos, 2400 m, 31 Mar 1963, Skutch 5462 (US); summit of Volcan Barba, ca. 1 km S of Laguna Barba, 2700 m, 24 Apr 1975, Utley & Utley 2247 (DUKE, NY). LIMON:P.N. Cordillerade Talamanca,trail from Quebrada Kuisa to Rio Lori, between Ujarrasand San Jose Cab6car, 2000 m, 9?21'00"N, 83?13'20"W, 16 Mar 1993, Herrera 5903 (INB); P.N. Cordillera de Talamanca, SE slope Cerro Biricuacua, Rio Dapari, between Ujarras and San Jose Cab6car, 2300 m, 9?23'45"N, 83009'25"W, 5 Apr 1993, Herrera & Gamboa 6260 (INB). PUNTARENAS: Parque InternacionalLa Amistad, between Las Nacientes and Quebrada Barranco and Rio Blanco, Finca San Carlos, 2640 m, 9?31'47"N, 83?35'30"W, 9 Apr 1995, Aguilar & Garrote 4032 (INB); Z.P. Las Tablas, Terraba-Sierpe drainage, trail to Cerro Echandi, 2500-2700 m, 9?00'32"N, 82049'41"W, 14 Aug 1997, Alfaro 1359 (INB). SANJOSE:Cerro de la Muerte, ca. 3000 m, 7 Oct 1978, Antonio 681 (HUT); 6 km E of Siberia near Cerro de la Muerte, next to CATIE reserve, finca Cuerici, 2600 m, 9?33'N, 83?42'W, 31 Jul 1992, Haber & Zuchowski 11341 (INB); Cerro de la Muerte, Villa Mills, 2800 m, 26 Feb 1965, Jimenez M. 2987 (CR, NY); Cerrode la Muerte, ca. 3000 m, 30 Jul 1980, Knapp 790 (BH); Cerro de la Muerte, 2000 m, 26 Feb 1966, Molina R. et al. 17908 (F); El Empalme, 14 Nov 1975, Poveda 1200 (CR, MO). PANAMA. CHIRIQUi: Ca. 8 km W of Cerro Punta in vic. of Las Nubes, 6100-6400 ft, 11 Feb 1978, Almeda & Nakai 3527 (CAS); Alto Respinga, above Cerro Punta, ca. 3000 m, 16 Mar 1977, D'Arcv 10691 (BM, MO); W of La Nevera, head of Rio Chiriqui Viejo on Volcan Baru, 10,400 ft, 14 Mar 1979, Hammel & D'Arcy 6393, 6408 (BH, MO); E slope of Volcan Baru, 8500 ft, 19 Mar 1979, Hammel et al. 6559 (BH, MO); Alto Pineda, end of rd. right turnjust before cooperative entranceto Cerro Punta, 8400 ft, 11 Apr 1979, Hammel et al. 6977 (BH, MO). Solanum vacciniiflorum is easily recognized by its large pointed buds, bullate leaves, and weak, reddish pubescence of the leaf undersides. It is most closely related to S. validinervium of montane western Venezuela but differs from that species in its leaf shape, geminate leaf clusters, and pointed flower buds. Solanum vacciniiflorum is one of the largest of the

179

TAXONOMIC TREATMENT

CentralAmericanmembers of sect. Geminata,growing to 25 m tall in sheltered areas. Collections of the species fromPanamahave much longerinflorescences thanthose from Costa Rica but areotherwiseidentical to Costa Rican plants. Trichomesin S. vacciniiflorumarepolymorphicin being uniseriate simple or branched, but whether branchedor simple,they aresmall-celledandweak.The occurrenceof branchedtrichomeson the leafundersides is sporadicin the species; some individualshave many of thetrichomesbranched,whileothershavealluniseriate trichomes.Those individualswith morebranchedhairs arenot frommoreexposedhabitats,so it seemsthe characteris not underenvironmentalcontrol.

glabrous;style straight,5-6 mm long; stigma a papillose area on the blunt tip of the style. Fruit a globose berry,green when immature,translucentyellow when ripe, 8-9 mm diam.;fruiting pedicels erect in young fruit, droopingwhen the fruitmature, 1-1.2 cm long, 0.75-1 mm diam.at the base.Seedspale yellowish-tan, flattened-reniform,3-3.5 x 2-2.5 mm, the margins incrassate,paler,the surfacesminutelypitted.Chromosome number:n = 12 (voucherKnapp& Mallet 6789). Distribution (Fig. 90). In the Andes of Venezuela from the state of Tachirato Trujillo,at 2000-3000 m. Specimens examined. VENEZUELA. LARA: Upper

limit of dwarfwoods betweenLas Sabanetasand ridge top by Lara-Trujilloboundary,above HumocaroBajo, 2600-2800 m, 6 Feb 1944, Steyermark55359 (MY. 46. SolanumvalidinerviumBenitez& S. Knapp,Emstia VEN). TACHIRA: BetweenFatimaandSanVicentede la 34: 6. 1985.Type.Venezuela.Trujillo:Bocon6,cerca Revancha,2760 m, 15 Nov 1973,Ferndndez1974(MY). TRUJILLO: ParamoLa Cristalina,2300 m, 16 Oct 1972, del Paramo La Cristalina,2300 m, 16 Oct 1972, Badillo 5096 (MY);windsweptsummitof old BoconoBadillo 5092 (holotype, MY). Figs. 84C,D, 92 Trujillord.,ca. 54 km W of Trujillo,ca. 2350 m, 9?21'N

Strict,virgatelybranchedshrubs, 1-2 m tall;young stems andleaves densely pubescentwith uniseriatetrichomes 0.2-0.6 mm long; older stems remainingpubescent; barkof the trunksgray.Sympodialunitswith varyingnumbersof leaves, not geminate.Leaves elliptic, widest at the middle, coriaceous, glabrous above, sparsely pubescent beneath along the veins with lax uniseriatetrichomes0.2-0.6 mm long, with 10-14 pairs of mainlateralveins, these impressedabove, raisedand prominentbeneath, the leaf blades 3-9 x 1-5 cm, the apexacuteorrounded,thebase acute;themarginsrevolute;petioles 0.8-3 cm long, slender.Inflorescencesopposite the leaves or intemodal, 2-4 mm long, simple, 2-6-flowered, densely pubescent with lax uniseriate trichomes0.1-0.5 mm long, these occasionallyreddish in dry specimens; pedicel scars closely spaced and overlapping,partiallyobscuredby thepubescence.Buds ellipsoid,stronglyexsertedfromthe calyx tube.Pedicels at anthesis 1-1.1 cm long, sparselypubescentwith lax uniseriatetrichomes,taperingfrom the base of the calyx tube to a slender base ca. 0.5 mm diam. Flowers spicy-fragrant,with the calyx tube cup-shaped, ca. 1 mm long, appearingwoody in dryspecimens,the lobes deltoid,the tips apiculateandtruncate,1-1.5 mm long, sparselypubescentwith uniseriatetrichomeslike those of the rest of the inflorescence; corolla bright white, 1.5-1.8 cm diam.,lobed 1/2way to thebase,cup-shaped at anthesis,the tips of the lobes denselypuberulentwith uniseriatetrichomesandpapillae;anthers3-3.5 x 1-2 mm, the antherbases with two indentationssuch that they appearauriculate,poricidal at the tips, the pores roundand very small; free portionof thefilaments ca. 0.5 mm long, the filament tube ca. 1 mm long; ovary

70?19'W, 20 Oct 1984, Knapp & Mallet 6782. 6785,

6787, 6789 (BH, K, MY, US, VEN); Misisi, below paramoLa Cristalina,on the old Bocon6-Trujillord.. 2150-2470 m, 21 Sep 1972, Ruiz Teran& Ldpez Palacios

7633(MY);old rd.betweenBocon6andTrujillo,between Urbinoand San Rafael, 2300-2500 m, 3-4 Sep 1966. Steyermark & Rabe 97284 (F, NY, VEN).

Solanumvalidinerviumis a tightly branchedshrub of windswept paramoregions, and is quite common whereit occurs.The inflorescencesarenot strictlyleafopposed,but the species belongs in sect. Geminata.Its closest relative is S. vacciniiflorumof montaneCosta Rica. Solanumvalidinerviumdiffers fromthatspecies in its plurifoliatesympodialunits, long petioles, short, broadcorolla lobes, and translucentyellow fruit.The spicy odorof the flowers ofS. validinerviumis unusual in the section; very few of the other species have any flower odor (however see S. lucens of the S. arboreum species group),andnonethatI have collectedhave such a strongsmell. The smell is reminiscentof a mixtureof cloves and cinnamon.

VII. Solanum deflexiflorum species group (S. deflexiflorum, S. imberbe, S. incomptum, S. irregulare, S. sieberi). Fig. 93 Shrubsor small trees; young stems and leaves glabrousto densely pubescentwith simple, uniseriatetrichomes.Sympodialunitsplurifoliate(Solanumimberbe) or difoliate,geminate,often anisophyllous.Leaves lanceolate (S. imberbe)to ovate, glabrousto sparselypubescent abaxially,the apex andbase various.Inflorescences opposite the leaves, simple or once branched,

180

FLORA NEOTROPICA ....... .. ..... oe...

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TAXONOMIC TREATMENT

181

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FIG. 93. Solanum deflexiflorum species group. A. S. sieberi plant, Venezuela (Knapp & Mallet 6706). B. S. sieberi flowers, Venezuela (Knapp & Mallet 6706). C. S. imberbe plant, Panama (Knapp & Kress 821). D. S. imberbe flowers, Panama (Knapp Kress 821). (Knapp & Kress 821).

usually somewhat elongate;pedicel scars evenly or unevenly spaced, overlappingor more commonly not. Buds elongateandellipsoid, stronglyexsertedfromthe calyx tube.Flowers white, lobed nearlyto the base, the petals planar (S. imberbe) or more often strongly reflexed at anthesis; antherselongate relative to flower size. Fruit green andhardat maturity;fruitingpedicels erect or somewhatdeflexed, woody. Seeds ovoid-reniform, pale tan to darkbrown.

Distribution. (Fig. 95) Central America and N SouthAmerica. Membersof theSolanumdeflexiflorum speciesgroup areeasily recognizableby the following suiteof characters: ellipsoid buds, elongate anthers,reflexed petals, usually erectfruitingpedicels, andmore or less slender inflorescencewithusuallyunevenlyspacedpedicelscars. Species of the group form large thickets in open areas andcan be a dominantfeatureof the understory.

FLORA NEOTROPICA

182

Key to the species of the Solanum deflexiflorumspecies group 1. Calyx lobes deltoid, the margins hyaline; leaves geminate or not, glabrous or pubescent along the veins beneath, not in the axils of the main lateral veins. 2. Shrubs of gravel or sand bars in river courses; petals planar at anthesis; leaves not usually geminate, narrow, willow-like, glabrous. Panama to N coastal South America ....................................... 48. S. imberbe 2. Shrubs or small trees of wet sites in dry forest, not of river courses; leaves geminate, elliptic, glabrous or pubescent along the veins beneath; petals strongly reflexed at anthesis. Northern South America to Trinidad ............................................................................................................................................ 5 1. S. sieberi 1. Calyx lobes long-triangular to acuminate; leaves geminate, variously pubescent. 3. Calyx lobes unequal in size, glabrous or minutely puberulent; calyx lobes not woody in fruit; plants drying black. Low elevations, Villavicencio, Meta, Colombia ............................................. 50. S. irregulare 3. Calyx lobes equal, usually pubescent; calyx lobes woody and accrescent in fruit; plants not drying black. Montane forests. 4. New growth glabrous or minutely red-papillose; berries globose. Cordillera Occidental, Co lom bia .............................................................................................................................. 47. S. deflexiflorum 4. New growth variously pubescent; berries umbonate. Montane Costa Rica and Panama......49. S. incoliptum

47. Solanum deflexiflorum Bitter,Repert.Spec. Nov. Regni Veg. 18: 49. 1922. Type. Colombia. Cauca: Sommetde la CordillereOccidentale,2000 m, 7 Nov 1899, Langlasse 40 (holotype,B [destroyed:F neg. 2657]; lectotype, K, here designated;isolectotype, Fig. 94 US). SolanunicalycopogonBitter,Repert.Spec.Nov. Regni Veg. 18: 52. 1922. Type. Colombia. Cauca: In parkartigen Savannengebuschen am oberen Rio Dagua (ein von West-Kordillerezum stillen Ozean abfall ender kleiner Flusse), 1500-2000 m, Jul-

Aug, Lehmann4715 (holotype,B [destroyed:F neg. 2651]; lectotype, F, here designated; isolectotypes, K, US [locality La Pa(o)rquera, Cali, 1400-1500 m, same number]).

Shrubs or small trees 2-4 m tall; young stems and leaves glabrous or minutely red-papillose; older stems glabrate;barkof the older stems andtrunkspale reddish-brown.Sympodial units difoliate, geminate. Leaves ellipticto narrowlyelliptic,the minorleaf often deciduous, glabrous above, pubescent beneath with tufts of white uniseriatetrichomes 0.5-1 mm long in the axils of the main lateralveins, the trichomesarising fromthe veins andthe lamina;majorleaves 8-15 x 37 cm, with 4-6 pairsof main lateralveins raisedabove, prominentand orangish beneath, the apex acute, the base acute, often oblique; petioles 0.8-1.5 cm long; minorleaves differingfromthe majorones only in size, 2-3 x 1.2-1.5 cm, the apex acute, the base acute;petioles ca. 5 mm long. Inflorescencesoppositethe leaves, simple, 0.5-1 cm long, 10-15-flowered, minutelypapillose or with uniseriate trichomes 0.5-1.5 mm long, these varying in density from sparseto dense;pedicel scars closely packed, often overlapping,beginning at the extremebase of the inflorescence.Buds when very young appearingpointed fromthe long-acuminatecalyx lobes,the corollasoon exsertedfromthecalyx lobes,

the buds laterbecoming ellipsoidjust before anthesis. Pedicels at anthesis deflexed, 0.9-1.1 cm long, tapering fromthe base of the calyx tubeto a slenderbase ca. 0.5 mm diam. Flowers with the calyx tube ca. I mm long, obconic, the lobes long-triangularto acuminate, ca. 2 mm long, minutely red-papillose, sparsely to densely pubescentwith uniseriatetrichomeslike those of the rest of the inflorescence;corolla white, 1.4-1.5 cm diam.,lobed ca. 2/3 of the way to the base, the lobes reflexed at anthesis,the tips of the lobes minutelypapillose; anthers 3-3.5 x ca. 1 mm, poricidalat the tips, the poresteardropshaped;free portionof thefilaments ca. 0.4 mm long, the filament tube ca. 0.75 mm long; ovary glabrous;style straight,5-6 mm long; stigma a papillose area on the tip of the style. Fruit a globose, green berry, ca. 1 cm diam.;fruiting pedicels erect, woody, 1.5-2.5 cm long, ca. 1 mm diam. at the base; a calyx lobes woody and persistent in fruit, formnning cup aroundthe base of the fruit.Seeds darkbrown in dry material, ovoid-reniform,ca. 4 x 3 mm, the surfaces minutelypitted.Chromosomenumbernot known. Distribution (Fig. 95). In the CordilleraOccidentalof Colombia and into N Ecuador,from 1000 to 3000 m. Selected specimens examined. COLOMBIA. S.loc., Mutis 80 (US). ANTIOQUIA: Bello, 1500-1800 m, 17 Jun 1930, Archer 153 (US); Fredonia, 1850 m, 3 Aug 1930, Archer 510 (US); Medellin, ca. 1500 m, Feb 1931, Archer 1613 (US); below Bocana, 7 km E of Medellin, 1800 m, 20 Feb 1949, Barkley 19Anl115(F, US); Sonson, Feb 1939, Bro. Daniel 2105 (F); near Medellin, 1560 m, Oct 1945, Diaz 324 (US); rd. from Medellin to Rionegro, 1750 m, 17 Nov 1948, Killip et al. 39918, 39928 (US); near Medellin, 19 Sep 1927, Toro663 (NY); Angelopolis, vic. of Medellin, 22 Jan 1928, Toro 901 (NY); Medellin, 1851-7, Triana s.n. (P); Medellin, Triana 186 (BM). CAUCA: Above Popayan, 1700-2300 m, Nov, Lehmnann

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de/lexitlorum (solid circles), S. incomptum (open circles), and S. irregulare (star). Solanuim

FLORA NEOTROPICA Rio Pichind6, 1800 m, 3 May 1977, Lozano et al. 4 (CUVC); Valle/Choc6borderarea, Serraniade los Paraguas, ca. 21-25 km beyond El Cairo, 1850-2000 m, ca. 4?45'N, 76?20'W, 25 Apr 1989, Luteyn & Giraldo 12672 (CUVC, NY); Alban, 1933, Perez-Arbelez 2190 (US); Hacienda Canada, right bank Quebrada El Miedo, 1030 m, 13 Aug 1987, Restrepo & Heredia 384 (NY); Hacienda El Medio, PanAmerican hwy. between La Paila and Zarzal, ca. 975 m, 16 Nov 1986, Silverstone-Sopkin et al. 2527 (MO, NY); Cerro del Ingles, Cordillera Occidental, Serrania de los Paraguas, I hour by jeep from El Cairo, ca. 2300 m, 1 Jan 1987, Silverstone-Sopkinet al. 2827 (MO, NY); Hacienda El Medio, PanAmericanhwy. between La Paila and Zarzal, ca. 975 m, 16 Apr 1987, Silverstone-Sopkin & Paz 3102 (MO, NY), 23 May 1987, Silverstone-Sopkin & Paz 3132 (MO, NY); Hacienda San Gerardo, near Guabas, ca. 950 m, 19 Dec 1987, Silverstone-Sopkin et al. 3392 (MO, NY) Hacienda El Medio, PanAmerican hwy. between La Paila and Zarzal, ca. 950 m, 21 Mar 1988, Silverstone-Sopkin & Paz 3732 (MO, NY). ECUADOR. CARCHI: El Carmen, Cerro Golondrinas, 2000-2400 m, 0?50'N, 7801 'W, 18-25 Aug 1994, Tirado et al. 1260 (BM). PICHINCHA:Mindo, 26 Jan 1876, Andr-e3835 (K); Calarali, Quito, 2600-2800 m, Jul-Aug, Lehmnann6325 (K). Local names. Colombia. Antioquia: chuchu; Valle de Cauca: guaco.

Solanum deflexiflorum is closely related to S. incomptum of montane Costa Rica and Panama. It differs from that species in its larger flowers, long-acuminate calyx lobes, sparser leaf pubescence, and glabrous K217 (F, K, P, US); Mt. Trompo del Puerco, 2500-3000 The two species share erect fruiting m, 29-30 Jun 1922, Penniell 7512 (NY); Popayan, 1700 young growth. pedicels and woody accrescent calyces. m, 5 Oct 1944, Sneidern 4704 (US). HUILA: Forest Solanum deflexiflorum is quite variable in infloaround Meherenberg, rd. from Popoyan, 2350 m, rescence pubescence, and the two extremes have been D.Arcv et al. 15577. 15579. 15599 (CUVC); Finca Merenberg, Cauca borde E of Leticia, 2275-2300 m, named as two different species. This seems excessive, as intermediates in pubescence density exist, and the 2?016'N, 76012'W, 7 Jul 1984, Gentryi et al. 47709 (CUVC, MO, NY); CordilleraCentral, E slope, Popoyan- two pubescence types and their intermediates occur La Platard., ca. I hr in car E of Leticia, Finca Mehrenberg, together throughout the Cordillera Occidental. Such ca. 2275 m, 7 Jul 1984, Silverstone-Sopkin & Gentry variation in pubescence is common in sect. Geminata. 1875 (CUVC); . VALLE DE CAUCA: Cordillera Occidental, E slope, Rio Cali, Pichind6, between La Marina and La Margarita, 2120-2160 m, 4 Nov 1944, Cuatrecasas 48. Solanum imberbe Bitter, Repert. Spec. Nov. Regni 18674 (F, US); rd. to the sea, 2000 m, May 1939, Veg. 18: 62. 1922. Type. Panama. Colo6n: Dos Drvander 2370 (F, US); Tulua near Cauca, 1000 m, Aug Bocas, Rio Fato valley, 40-80 m, Pittier 4204 (ho1941, Drvander 2490 (F); Hacienda Media Canoa, Rio lotype, US [US-679305]). Figs. 93C,D, 96 Paila between Caserio La Paila and Zarzal, 1000 m, 4?20'N, Solanumn aquiatile C. V. Morton, Contr. U.S. Natl. 76?04'W, 6 Apr 1986, Gentry et al. 54132 (CUVC, MO, Herb. 29: 50. 1944. Type. Colombia. Santander: NY); Bosque de Ema, ca. 60 km N of Cali, Cordillera Camp Aguila, Carare Valley, in the vic. of Puerto E 1700 17 Dec Occidental, slope, m, 3?38'N, 76?33'W, Berrio, 100-700 m, 9 Aug 1935, Haught 1868 1987, Gentriyet al. 59502 (MO, NY); La Cumbre, 1700(holotype, US; isotypes, F [F neg. 49455], K). 1900 m, 11-16 Jun 1922, Hagen 11989 (US); Hacienda of D'Arcy, Ann. Missouri Bot. Solanum intermediumn Himalaya, Bitaco, 1800 m, 11 May 1988, Heredia et al. Gard. 60: 747. 1973. Not of Sendtner. 562 (NY); near Mares on hwy. from Cali to Buenaventura, 1880 m, 22 Jun 1944, Killip et al. 39187 (F, US); Carretera Shrubs or subshrubs growing in river courses, al Mar, W of Cali, CordilleraOccidental, 1600 m, 13 Nov 0.5-2 m tall; young stems slender, glabrous; bark dark 1948, Killip & Lehmann V 39806 (US); Pefias Blancas,

TAXONOMIC TREATMENT

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brown,palerandexfoliatingon olderstems,lenticellate. Sympodial units difoliate, geminate or plurifoliate. Leaves oblanceolate to narrowlylanceolate, variable over the rangeof the species, widest at orjust distal to the middle, with 6-9 pairsof main lateralveins prominent andyellowish beneath,glabrous;majorleaves 814 x 0.8-3.2 cm, the apex acuminate,the base attenuate, decurrenton the petiole; petioles 5-6 mm long, winged from the decurrentleaf bases; minorleaves, if present,muchthe samesize andshapeas themajorones, ellipticto linear,7-8 x 0.8-3 cm, the apex acuteto acuminate, the base acute to attenuate;petioles 3-5 mm long. Inflorescences opposite the leaves or somewhat intemodal,simpleor furcate,0.5-2 cm long, 3-40-flowered, glabrous;pedicel scars evenly spaced ca. 1 mm apart,beginning very close to the base of the inflorescence. Buds globose, soon exserted from the hyaline calyx, the budsthen ellipsoid.Pedicels at anthesis0.81.3 cm long, white, taperingfrom the calyx tube to a slender base less than 0.5 mm diam., erect or slightly deflexed. Flowers with the calyx tube ca. 1 mm long, hyaline, white, the lobes deltoid, apiculate, ca. 1 mm long, minutelypapillose at the tips or glabrous;corolla white, 1-1.2 cm across, lobed 2/3 of the way to the base, the lobes planaror slightly reflexed at anthesis, with scatteredminutepapillae on the tips of the lobes; anthers 2.5-3 x 1 mm, poricidal at the tips, the pores teardropshaped;free portionof thefilamentsless than 0.1 mm long, the filamenttube ca. 0.2 mm long; ovary glabrous;flowers heterostylous,shortstyles 1-1.5 mm long, with indistinctstigmas,long styles 5-6 mm long, straight;stigma small andcapitate,minutelypapillose. Fnruit a globoseberry,whenimmatureslightlyumbonate, 0.8-1.3 cm diam., green, occasionally with a metallic bluishcast;fruitingpedicelswoody,erect,ca. 2 cm long, in age becoming very corky.Seeds brown,ovoid-reniform, ca. 3 x 1.5 mm, the surfaces minutely pitted, nearly smooth. Chromosomenumbernot known. Distribution (Fig. 100). FromE CostaRica and Panamato Colombia,Ecuador,andVenezuelain coastal areas or major river drainages, usually growing on gravel bars or the marginsof rivers, often submerged when rivers are in flood. Very common where it occurs. At the type locality, Colo6nprovince, Panama,it often fills entire gravel islands in the centerof rivers. Selected specimens examined. COSTA RICA. LIMON:Canton de Pococi, R.F. Cordillera de V. Central, in quebrada Molinete, Bosque de Teleferico, 500 m, 10?10'22"N, 83?54'32"W, 22 Oct 1994, Hammel et al. 19630 (INB, MO). PANAMA. CANALAREA: Vic. of Madden Dam, Chagres River, gravelly shores and bars at Landing 2, 15 Aug 1940, Bartlett & Lasser 16778 (MO); near Rio Frijoles, 9 Sep 1972, Gentry 5824 (MO). COLON:Rio

FLORA NEOTROPICA Guanche, 16 Nov 1975, D'Arcy 9725 (MO, NY); ca. 2 km up Rio Guanche from Colo6n-Portobelo hwy., 0-20 m, 9?30'N, 79?40'W, 27 Aug 1980, Knapp & Kress 821, 822 (BH); forest and forest edge from Portobelo hwy. to 4 km up Rio Guanche, 0-50 m, 9?30'N, 79?40'W, 30 Aug 1981, Knapp 981 (MO); 2-4 km up Rio Guanche from the Portobelo hwy., 0-50 m, 9030'N, 79?40'W, 3 Oct 1981, Knapp & Chazdon 1408 (MO); at base of Cerro Bruja, along Rio Escandaloso above Mina Boquer6n # 2, 19 mi E of Translsthmianhwy. on rd. to Salamanca, 150-200 m, 9?50'N, 79?32'W, 28 Mar 1982, Knapp et al. 4445 (MO); trail S of Rio Guanche, on ridge to Cerro Pan de Azucar, ca. 200 m, 20 Sep 1974, Mori & Kallunki 2024 (MO); along Rio Guanche, 0.5-1 km upstream from Puerto Pilon-Portobelo rd., 6 km S of Portobelo. 5-30 m, 25 Sep 1973, Nee 7137 (MO). COMARCADE SAN BLAS:El Real Quebrada, Trapiche, Duke & Bristan 313 (US, WIS); mainland opposite Play6n Chico, 0-3 mi from Caribbean, 4 Oct 1972, Gentry 6410 (MO). DARIEN:Vic. of Boca del Cupe, ca. 40 m, 5 Oct 1938, Allen 905 (MO, NY); Rio Pirre, 16 Nov 1967, Bristan 1482, 1483 (MO); upper Rio Tuquesa, early 1973, Le Clezio 102, 257 (MO); ridges NE of Ensenada Guayabo, on trail to Rio Jaque, 0-300 m, 7?26'N, 78?06'W, 23 Jan 1982, Knapp 3065 (MO); N facing slopes and flatlands of Rio Jaque valley, along Quebrada Luka, 0-300 m, 7?27'N, 78?05'W, 24 Jan 1982, Knapp 3116 (MO); 5 km SW of Pinogana on the Rio Tuira, 15 Nov 1967, Mori 365 (MO, MPU, WIS), 366 (MO, WIS). PANAMA: Along Pan-Am hwy. near Jenine, Rio Cafilta. 24 Sep 1961, Duke 3844 (MO); Pan-Am hwy. at Rio Mamonica, 4 mi beyond Chepo, 11 Sep 1962, Duke 5576 (MO); Rio Cafasas, 100 m, 27 Sep 1967, Duke 14511 (MO, WIS); San Isidro, 2 mi from the Transisthmianhwy., Nov 1974, Garrido 11 (MO); Pacora, Juangil, 6 Oct 1974, Hertentains 9 (MO); along Rio Ipeti near Choc6 village of Ipeti, 4 km S of PanAmericanhwy., 50-100 m, 8?50'N, 77?30'W, Knapp et al. 4563 (MO); Juangil, quebrada 2 km from Rio Pacora, 6 Oct 1974, de la Lastra 11 (F, MO); Chepo near river, 3 Oct 1965, Tyson 1428 (MO); I mi W of El Llano, 29 Sep 1972, Tyson 6843 (MO). COLOMBIA. ANTIOQUIA:Footpath along Rio Anori to Madreseca approx. 8 km upriver from Quebrada Tirana, 28 km SW of Zaragoza, 400-700 m, 19 Mar 1977, Alverson et al. 216 (MO, NY); along Rio Anori, 0-5 km S of Quebrada Tirana, 28 km S of Zaragoza, 400-700 m, 23 Mar 1977, Alverson et al. 248 (MO); 14-17 km NW of Remedios on rd. to Zaragoza, region of Cerro Cabeza, bands of Rio Tucupe, 250-350 m, 7?20'N, 74?30'W, 15 Sep 1987, Callejas et al. 5091 (NY); Callejas et al. 5156 (NY); 8-20 km S of Mutata on rd. to Dabeiba, Alto de Echeverri, 210-230 m, 7?15'N, 76?30'W, 22 Nov 1987, Callejas et al. 5801 (K, NY); Frontino, km 27 of Nutibara-Murri rd., 960 m, 6?40'N, 76?26'W, 21 Sep 1987, Zarucchi et al. 5528 (K, MO). CAUCA: San Juan de Micay, 100 m, 28 Dec 1946, Haught 5401 (US). CESAR:Rio Badillo. Valle du Par,Aug 1844, Purdies.n. (K). CHOCO:Along Rio Truando between Rio Sucio and Esperando, 15 Dec 1966, Duke

187

TAXONOMIC TREATMENT 9088 (WIS); Rio El Valle, between Rio Mutata and Rio Miniquia, at mouth of Rio Miniquia, 7 Jan 1973, Forero & Gentry 740 (MO, NY); region of Rio Pichima, Waunara Indian village, 100 m, 4?25'N, 77?17'W, 19 Nov 1976, Forero 712 (MO); Rio El Valle between Choc6 Indian village (Ijito) and mouth of Rio Mutata, 100 m, 7 Aug 1976, Gentry & Fallen 17326 (MO, NY); Rio Mutata, tributary of Rio El Valle, between base of Alto de Buey and mouth of river, 100-150 m, 9 Aug 1976, Gentry & Fallen 17458 (MO); Rio Nuqui, 50 m, 22 Jan 1947, Haught 5462 (US); Rio Taparal off Rio San Juan, 100 ft, 25 Aug 1962, Hugh-Jones 339 (K, US); Bahia Solano, Quebrada Resquiata, 2 m, 6?16'N, 77?21'W, 2 Jan 1984, Juncosa 1650 (MO. NY); Rio Mecana, 5-10 m, 6016'N, 77021'W, 4 Jan 1984, Juncosa 1686 (NY); Bahia Solano, near Ciudad Mutis, 0-75 m, 21-23 Feb 1939, Killip & Garcia 33608 (US); Virud6, in bed of quebradadel Salto, 0-100 m, 3 Aug 1973, Warner& White 130 (MO, NY); km 27 of Nutibarra-Murri rd., 960 m, 6?40'N, 76?26'W, 21 Sep 1987, Zarucchi et al. 5528 (MO, NY). MAGDALENA:Near La Jagua, 40 km NE of Chiriguana, 20 Aug 1938, Haught 2266 (US); 4 km SW of La Jagua, in bed of Quebrada Sororia, 100 m, 4 Aug 1943, Haught 3590 (F, US). SANTANDER: Quebrada Angulo, 4 km S of Lebrija, 955 m, 7005'N, 73?13'W, 29 Jul 1944, St. John 20600 (US). VENEZUELA. ZULIA: Perija, ca. 60 km S of Machiques near mission Los Angeles del Tukuko, 200 m, 14 Oct 1966, de Bruijn 1178 (K, MO, VEN); basin of Rio Guasare, Hacienda Chiquinquira, left bank of river, 200-250 m, 10?52'10"N, 72?29'30"W, 9 Nov 1982, Bunting et al. 11984 (MO, NY); ca. 55 km SW of Machiques by air, Aricuaisa (Ariguaisa)-Pie de Monte on Rio Aricuaisa, 100-250 m, 9?36'N, 72?54'W, 24-25 Mar 1982, Liesner & Gonzalez 13065 (VEN); Mara, La Trunlitre, Rio Guarar6,50 m, 22 Aug 1957, Medina 744 (VEN); Sierra de Perija, along the Rio Yasa, vic. of "Kasmera" (Biol. Station of Univ. de Zulia), SE of Machiques, 300 m, 23-24 Aug 1967, Steyermark & Fernendez 99627 (NY, US, VEN); Mara, along Rio Cachiri, just N of Hacienda Salamanca, 10 air km SW of El Paraiso, 150 m, 10?46'30"N, 72?20'W, 6 Jun 1960, Stevermark et al. 123459 (NY, VEN). ECUADOR. ESMERALDAS:Creek pouring into Rio Palavi, across from Awa camp on W bank, 200 m, 0?58'N, 78?16'W, 9 Feb 1988, Hoover et al. 3071 (MO, QCA); creek down the Palavica, 200 m from Awa encampment on E bank, 200 m, 1?07'N, 78?37'W, 12 Feb 1988, Hoover et al. 3151 (MO, QCA); up Rio Palavi from Awa camp, 200 m, 1?07N, 78?37'W, 14 Feb 1988, Hoover et al. 4160 (MO, QCA). Solanum imberbe is most similar to S. sieberi of coastal Colombia and Venezuela, particularly to the glabrous populations of that species. It differs from S. sieberi in its non-geminate, usually completely glabrous narrowly elliptic to lanceolate leaves, flowers with the corolla lobes only somewhat reflexed at anthesis, and its corky fruiting pedicels without conspicuous apical swellings. Solanum imberbe is also distinct in habitat

from S. sieberi, growing in river courses. Like other solanums growing in these situations,S. imberbehas narrowleaves, and is subjectto periodic inundation. Leaf shapeis quitevariablein Solanumimberbe. Plants from central Panama have obovate to oblanceolate leaves, but furthereast in Panamaandin northern SouthAmericaplantshave narrowlylanceolateto even linearleaves. The narrowestleaves are found on plantsfromZuliastate,Venezuela,thesebeing the eastern-mostpopulationsof the species. Flower and fruit morphologyareconstantthroughoutthe species range.

49. Solanum incomptum Bitter, Repert. Spec. Nov. RegniVeg. 18:60. 1922.Type.CostaRica. SanJose: Llanos de Alajuelita, 1 Dec 1889, Tonduz1468 (holotype, BR; isotype, US). Fig. 97 SolanumcopeyanumBitter,Repert.Spec. Nov. Regni Veg. 18: 54. 1922. Type. Costa Rica. San Jose: Forets du Copey, 1800 m, Feb 1898, Tonduz 11689 (lectotype,BR, here designated;isolectotypes, B [destroyed:F neg. 2857], BM, US). Solanum incomptumvar. longiuspilosumnBitter. Repert. Spec. Nov. Regni Veg. 18: 61. 1922. Type. Costa Rica. San Jose: La Verbenapres Alajuelita,Jul 1894, Tonduz9096 (holotype,BR: isotypes, BM, US). Solanum incomptumvar. lugens Bitter, Repert. Spec.

Nov. RegniVeg. 18: 61. 1922. Type.CostaRica. SanJos6:Taldes Rio Poas,Massifde Iscasu,2100 m, Pittier 2396 (holotype, BR).

Shrubs 1-4 m tall; young stems glabrousor puberulentwith minuteto 0.3 mm long straw-coloredunicellularor uniseriatetrichomes;older stems glabrous; barkwhite, the outerlayerscrackingandpeeling transversely. Sympodial units difoliate, geminate. Leaves ovateorobovate,usuallygeminate,widest at orjust distal to the middle, the uppersurfaceglabrousor with a few scatteredtrichomes,the lower surfacesparselyto densely pubescentwith uniseriatetrichomes0.5-1 mm long on theveins andlamina,thepubescencevariablein degreeandlength;majorleaves 6.5-17.7 x 2.1-6.9 cm, with 8-1 pairs of main lateralveins impressedabove, midribslightlykeeled, the apex acuteor acuminate,the base acute;petioles 0.8-1.8 cm long; minorleaves 1.35.1 x 0.8-2.9 cm, the apex acuteor acuminate,the base acute;petioles3-8 mm long.Inflorescencesoppositethe leaves, simple,0.3-1 cm long, subumbellate,with most of the flowersin thedistal1/4, 8-10-flowered,occasionwithsimpleuniseriatetrichomeslikethose allypuberulent of the stem and leaves;pedicel scars closely spaced, overlapping.Buds obovoid to ellipsoid, glabrous or pubescentwith simpleuniseriatetrichomes.Pedicels at anthesisdeflexed, 6-8 mm long, slender,abruptlynar-

FLORA NEOTROPICA

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TAXONOMIC TREATMENT

rowingat the base of the calyx tube,ca. 0.5 mm diam.at the base. Flowers with the calyx tube 1-2 mm long, glabrousorpubescentwithuniseriatetrichomeslikethose of the stems andleaves, the lobes triangular,1-1.5 mm long, thetips minutelypapillose;corollawhite or greenish-white, 1-1.3 cm across,lobednearlyto the base, the lobes reflexedat anthesis,the tips of the lobes minutely papillose;anthersca. 3.5 mm long, the terminal0.5 mm paler and thickened,ca. 0.5 mm wide, poricidalat the tips, the poresbecomingslit-likeupon drying;freeportion of thefilamentsca. 0.4 mm long, the filamenttube ca. 1 mm long; ovary glabrous;styles of two different types, shortstyles ca. 1 mm long, long styles ca. 6 mm long,straight;stigma(long-styledflowers)minutelypapillose. Fruit a globose, greenberry,umbonatewith the persistentstyle base, 0.5-1 cm diam.;fruitingpedicels woody anderect,0.8-1.6 cm long, ca. 0.75 mm diam.at the base; calyx accrescentin fruit,formingan open cup 0.7-1.2 cm diam., the calyx lobes 2-5 mm long, stiff andoftenbecomingchartaceous.Seedsfew perfruit(ca. 10), ovoid-reniform,brown,ca. 3.5 x 2.5 mm, seed coat thin,the surfacesminutelypitted.Chromosomenumber not known.

189 7000 ft, 5 Apr 1979, Hammel et al. 6755 (BH, MO); W slopes of Volcan de Chiriqui(Bari), 1500 m, 21 Apr 1975. Mori & Kallunki 5691 (MO); on Boquete trail, 4 km from Cerro Punta, near Paso de Respingo, ca. 2300 m, 22 Apr 1975, Mori & Kallunki 5729 (MO); W side of Volcan Baru, along trail from lava flats to summit, 2000-2500 ft, 22 Jul 1975, Mori & Bolten 7363 (MO); 6 mi NE of El Volcan, 7000-7500 ft, 4 Mar 1962, Tyson 812 (MO): canyon to waterfall, 5 mi NE of El Volcan, 7000 ft, 18 Dec 1970, Tyson 6347 (MO); Rio Chiriqui valley, near Cerro Punto gate, 29 Mar 1938, White 51 (MO, US).

Solanum incomptumis most closely related to S. deflexiflorumof montane western Colombia, and differs from that species in its smallerflowers, longer inflorescences,deltoidcalyx lobes, umbonatefruit,and in its more pubescentnew growth.The umbonatefruit andthe chartaceouscalyx lobes in fruitare distinctive characteristicsof S. incomptum. Solanum incomptumis variable both as to degree of pubescenceandlengthoftrichomes.Thosewith the longest trichomes have been called S. incomptum var. longiuspilosum, and those with the shortest S. copeyanum,but the variationis continuous and even Thevariationmay varietaldistinctionsarenotwarranted. distributionof this species in isothe of result the be Distribution (Fig. 95). Cordillera de Talamanca lated local populationsthroughoutits range. in Costa Rica andPanamafrom 1200 to 2500 m, often were BothSolanumincomptumandS. copeyanumn in the understoryof oak forests. publishedby Bitterin the same paper.I have chosenS. Specimens examined. COSTA RICA. S.loc., Oer- incomptumas the valid name for the species because it sted 1415 (US). ALAJUELA:Cerro Daser, Fila Cedral, is that more commonly in use, even though S. 2000 m, 30 Aug 1971, Burger & Burger 8186 (F, MO). copeyanumappearson an earlierpage in the publicaCARTAGO: Forests of El Copey, 1800 m, Feb 1898, tion. No page priorityrule exists in the Botanical Code Cordillera de Tilaran, Tondutz11723 (US). PUNTARENAS: et al., 1994). Monteverde, Pacific slope below moist forest, 1300 m, (Greuter 10?18'15"N, 84?48'40"W, 14 Apr 1994, Haber 11751 (INB). SAN JOSE: Cordillera de Talamanca, Las Nubes, Sta. Elena, 1210 m, 9?23'30"N, 83?35'50"W,4 Aug 1995, Aliaro 310 (INB); hills above Escaz6, 5000 ft, 2 Oct 1953, Heiser 3586 (CR, US); near San Isidro, 1500 m, Dec 1881, Lehmann 1774 (US); valley of Archangele, Escazu, 1500-1800 m. 11 Dec 1898, Pittier 13060 (US); Z.P. Cerros de Escazfi, Aserri, Cedral and Hierbabuena, 2000 m, 9?50'30"N, 84?07'20"W, 1 Oct 1993, Ramirez & Morales 143 (INB); vic. of El General, 1190 m, Jan 1936, Skutch 2417 (NY); Las Nubes, 1500-1900 m, 20-22 Mar 1924, Standlev 38579 (US); between Aserri and Tarbaca, 1200-1700 m, 6 Dec 1925, Standley 41398 (US). PANAMA. CHIRIQUi: El Barn, SE side above Yen Finca, headwaters of Rio Calga, 7600-8500 ft, 20 Mar 1979, D'Arcy et al. 12737 (BH, MO); Bajo Chorro, Boquete district, 6000 ft, 10 Jan 1938, Davidson 99 (F, US), 26 Mar 1938, Davidson 446 (F, US); valley of Rio Chiriqui Viejo, N of Volcan, 5200-5600 ft, 9 Dec 1966, Duke 9011 (MO); Volcan Baru (E slope) deep draw W of Finca Yen, 8000 ft, 17 Mar 1979, Hanimel et al. 6456, 22 Mar 1979, Hammel et al. 6583 (BH, MO); Llanos E of Hato de Volcan, savannah and woods on lava flow, 6600-

50. Solanum irregulare C. V. Morton, Contr. U.S. Natl. Herb. 29:49. 1944. Type. Colombia. Meta: Puerto Sanchez on Rio Humea, in thickets along river,400 m, 8 Feb 1939, Haught 2587 (holotype, Fig. 98 US; isotypes, F, K, U). Much branchedshrubs 1-2 m tall; young stems hirsutewithuniseriatetrichomes(oftenonly bi-cellular) ca. 0.5 mm long; older stems glabrousexcept for a few trichomes at the nodes; bark dark brown, minutely white lenticellate.Sympodialunitsdifoliate,geminate, strongly anisophyllous. Leaves glabrous or minutely glandular,ovate,widestatorjustproximalto themiddle; majorleaves 11-16.4 x 3.6-8.8 cm, with 9-10 pairsof main lateralveins, raised above, prominentand dark beneath,the apex acuminate,the base cuneate,slightly decurrenton the petiole; petioles 0.6-1.9 cm long; minor leaves differing from the major ones in size and shape, elliptic to orbicular,2.2-7.9 x 1.9-5.5 cm, the

190

FLORA NEOTROPICA

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TAXONOMIC TREATMENT191

apex rounded,the base cuneate;petioles 2-4 mm long. Inflorescences opposite the leaves, simple, 2.5-8 cm long, 10-40-flowered, sparsely pubescent with uniseriatetrichomeslike those of the stems andleaves; pedicel scars evenly spaced 1-2 mm apart,beginning ca. 1.5 cm from the base of the inflorescence.Buds ellipsoid, appearingpointed when young from the longtriangularcalyx lobes, glabrous or minutely puberulent with simple uniseriate trichomes. Pedicels at anthesis deflexed, 8-9 mm long, filiform, ca. 0.2 mm diam.at the base.Flowers with the calyx tubeca. 1 mm long, the lobes triangular,ca. 1.5 mm long, with a few uniseriatetrichomeslike those of the stems andleaves, after fertilizationthe calyx lobes becoming markedly asymmetric, two lengthening (5-6 mm long in very young fruit), two remaining much the same size (ca. 1.5 mm long) andthe fifth of intermediatesize (ca. 3.5 mm long); corolla white, 7-8 mm across, lobed nearly to the base, the lobes reflexedat anthesis,the tips of the lobes minutely papillose; anthers ca. 2 x 0.75 mm, poricidal at the tips, the pores teardropshaped; free portionof thefilamentsless than0.5 mm long, the filament tube ca. 0.5 mm long; ovary glabrous;style ca. 3 mm long, straight,widening slightly to the truncate, minutelypapillosestigma. Fruit andseeds not known. Chromosomenumbernot known. Distribution (Fig. 95). Known only from the foothills of the CordilleraOrientalin Colombia, near Villavicencio. Specimens examined. COLOMBIA. META: Villavicencio,thicket near Guatiguia,500 m, 1-2 Sep 1917, Pennell 1584 (NY); Villavicencio,21 Jan 1889, Sprague89 (K, US). Solanum irregulare is rather isolated in the group,but is most similarto S. sieberi of coastalnorthem SouthAmerica.It shareswith thatspecies ellipsoid buds, simple uniseriatepubesence of the new growth and occasionally leaves, and elongate inflorescences. Furthercollections of this species areneeded to determine its variabilityand fruitstructure.The structureof the fruiting calyx in S. irregulare is unique in sect. Geminata,andit will be interestingto see maturefruiting collections of this species. 51. Solanum sieberi Van Heurck & Miill. Arg., Observ. Bot. 56. 1870. Type. Trinidad and Tobago. Trinidad: S.loc., Sieber 22 (lectotype, GDC, here designated;isolectotypes, G, M [Morton neg. 8720], P). Figs. 93A,B, 99 Solanumnzudum Dunalvar.longifoliumDunalin DC., Prodr. 13(1): 145. 1852. Type. Trinidad and Tobago. Trinidad:S.loc., Sieber 22 (holotype, G-DC;

isotypes, G, M [Mortonneg. 8720], frag. MPU, P).

SolanumcallobotrysPittier,Cat.Fl. Venez.381. 1947. Nom. nud. Venezuela. Portuguesa: Acarigua, en

lugares incultos, Chardon107 (VEN). Solanum micranthumof R. E. D. Baker & N. W. Simmonds, Fl. Trin. Tobago 2: 267. 1953. Not of Willd. ex Roem. & Schult.

Shrubsor small trees, 1-5 m tall; young stems and leaves minutelyred-papillose,glabrousor pubescentwith whiteuniseriatetrichomes0.2-0.75 mm long; older stems glabrate; bark of older stems grayishbrown. Sympodial units difoliate, geminate. Leaves ellipticto ovate,widest at orjust proximalto themiddle, glabrousabove, glabrousor pubescentalong the veins below with uniseriatetrichomeslike those of the young growth;majorleaves 9-16 x 3-9 cm, with 6-8 pairsof main lateralveins, these raised above, prominentand darkerbelow in dryspecimens,the apex acute,the base acute, slightly decurrenton the petiole; petiole 1-1.5 cm long; minorleaves differingfromthe majorones in size and shape, more roundedin outline, 1.5-5 x 1-3 cm, the apex acuteto rounded,the base acute;petioles 0.5-1 cm long.Inflorescencesoppositethe leaves, slender,often once furcate,0.7-6 cm long, 5-50-flowered, glabrous or pubescent with uniseriatetrichomes like those of the young stems and leaves; pedicel scars unevenly spaced, closely spaced to 0.5 mm apart,beginning ca. 1/3 of the way from the base of the inflorescence. Buds globose when very young, the corolla soon exserted from the calyx tube, older buds with a characteristicnarrowly ellipsoid shape, the tips flattened.Pedicels at anthesis7-9 mm long, taperingfrom the calyxtubeto a slenderbase ca. 0.25 mm diam.Flowers sweetly fragrant,with the calyx tube 0.75-1 mm long, broadlycup-shaped,the lobes deltoid,ca. 0.5 mm long, glabrous or pubescent with the same uniseriate trichomes as the rest of the inflorescence, 0. 1-0.75 m long; corolla white, 0.8-1 cm diam., lobed 3/4 of the way to the base, the lobes stronglyreflexedat anthesis, the tips and margins of the lobes minutely papillose; anthers 2.5-3 x 0.75-1 mm, poricidal at the tips, the pores teardropshaped; free portion of thefilaments minute,less than0.2 mm long, the filamenttubeca. 0.5 mm long; ovary glabrous;style straight,5-6 mm long; stigmanotmuchdifferentiatedfromtherestof the style, minutelypapillose. Fruit a globose, green berry,ca. I cm diam.;fruitingpedicelserect,woody, 1-1.7 cm long, ca. 1 mm diam. at the base, ca. 3 m diam.at the apex in dry specimens (with a conspicuousrhomboidalswelling at the apex in live plants). Seeds pale tan, ovoidreniform,ca. 3.5 x 2 mm, the surfacesminutelypitted. Chromosome number: n - 12 (vouchers Knapp & Mallet 6698, 6702, 6710).

FLORA NEOTROPICA

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TAXONOMIC TREATMENT

00f700

FIG. 100. Distribution of Solanium imberbe (solid circles) and S. sieberi (open circles).

Distribution (Fig. 100). Along the northern coast of South America from Colombia to Trinidad and Martinique, in wet microsites in otherwise dry areas, from sea level to 1000 m. Selected specimens examined. WINDWARD ISLANDS. MARTINIQUE:S.loc., Sieber s.n. (AWH, G-DC). COLOMBIA. GUAJIRA: 9 km S of Carraipia,300 m, 4 Jul 1944, Haught 4235 (F, K, NY, US); 17 km S of Carraipia, 300 m, 7 Aug 1944, Haught 4295 (F, K, NY, US). MADGALENA: San Juancito, Sep 1852, Holton 10 (K); Badillo, 7 Sep 1852, Holton 27 (NY); San Juancito, 10 Sep 1852, Holton 557 (NY); Santa Marta, ca. 0 m, 1898-1901, Smith 1191 (BM, CAS, MO, NY, US, W); Parque Nacional Tayrona, mule trail from Pueblita and Calbozo, 250--450 m, 11?19'N, 73?58'W, 26 Oct 1972, Kirkbride 2566 (NY, US); near Fonseca, 200 m, 18 Aug 1944, Haught 4312 (NY, US); Rio Rancheria near Fonseca, 150 m, 2 Oct 1944, Haught 4386 (K, NY, US). VENEZUELA. CARABOBO: Near Puerto Cabello, 0 m, 21 Jun 1913, Pittier 6360 (US); Guaremales along rd. from Puerto Cabello to San Felipe, 10-100 m, 25 Jul 1920, Pittier 8993 (NY, US, VEN). FAI,CON: Between Tucacas and Morrocoy, La Cuevita, 2 m, 6 May 1972, Benitez de Rojas & Ferrari 1373 (MY, NY); between Santa Maria and La Chapa, rd. to Curimagua, ca. 800 m, 14 Jan 1975, Benitez de Rojas 1816 (MY); Sierra San Luis, along old rd. from Curimaguato San Luis, ca. 15 km E of Curimagua, ca. 1000 m, 11?12'N, 69?40'W, 29 Sep 1984, Knapp & Mallet 6689 (BH, K, MY, US, VEN); Sierra San Luis, along access rd. to Hotel Parador,near Curimagua, ca. 1300 m, 10?12'N, 69?40'W,

29 Sep 1984, Knapp & Mallet 6694, 6695, 6696 (BH, K, MY, US, VEN); Cerro Chichiriviche, Parque Nacional Morrocoy, on side rd. 0.7 km W of La Cuevita, 0.4 km E of Lizardo, ca. 110 m, 10?52'N, 68?16'W, 1 Oct 1984, Knapp & Mallet 6697, 6698, 6700 (BH, MY, US, VEN); La Cuevita, 11.5 km E of Parque Nacional Morrocoy entrance on Falc6n 2, S of Sanare, 0-10 m, 10?52'N, 68?16'W, 1 Oct 1984, Knapp & Mallet 6701, 6702 (BH, K, MY, US, VEN); 5.3 km E of main hwy. (Falc6n 2) on rd. to Lizardo, 0-5 m, 10?52'N, 68?20'W, 1 Oct 1984, Knapp & Mallet 6703, 6704, 6705, 6706, 6707 (BH, K, MY, US, VEN); 1/2 km N of bridge over Rio Tocuyo at El Alto, 18 Apr 1971, Nee & Mori 3957, 3958 (MAD, US, WIS); La Luisa, S base of Lomas de Chichiriviche, 9 Sep 1923, Pittier 11132 (NY, US, VEN); S margin of Golfete de Guare, at foot of calcareous hills SW of Chichiriviche at Cueva de los Indios, 1-4 m, 10?545'N, 68? 17--9'W, 31 Aug 1974, Steyermark & Manara 110485 (NY, VEN); between the entrance to Cueva del Toro and Palma Sola, ca. 800 m, 13 Sep 1980, Trujilloet al. 16784 (MY, NY); 12 km SE of Coro, Siburia, 60 m, 7 Feb 1978, Wingfield5085 (MY). LARA: Las Adjuntason rd. to Guarico, ca. 900 m, 7 Dec 1974, Benitez de Rojas 1703 (MY); Canon de Angostura, Rio Yacambf, Sanare, 16 Jun 1974, Fernandez 2228 (MY, NY); near Barquisimeto, Jul 1925, Saer 270 (US, VEN); Paso de Angostura, Yacambf dam, confluence of Quebrada Honda and Rio Yacambf, 500 m, 9?41'N, 69?31-33'W, 28-31 Jul 1973, Steverniark & Carreho Espinoza 107615 (NY, VEN); El Tocuyo, Aug 1937, Tamayo 316 (US, VEN); along Rio Tocuyo, near city of Tocuyo, Apr 1962, Tamayo 4592 (MY, NY); El Tocuyo-Guarico rd., 6 km N of Guarico, 29 Mar 1980, Xena 593 (NY, VEN). PORTUGUESA:Acarigua, 21 Jun 1940, Chardon 107 (US, VEN). ZULIA: Basin of Rios Socuy-Guasare, near Casanara, ca. 5 km S of Carichaunocamp of Carbozulia,100-150 m, 10 Aug 1981, Bunting & Kauffman 10224 (NY); Fuentes de Socuy, near "Los Laureles" and the cave, Sierra de Perija, ca. 615 m, 7 Jul 1976, Trujillo & Ferrari 14042 (MY, NY). TRINIDAD AND TOBAGO. TRINIDAD:S.loc., Bot. Gard. Herb. 2670 (US); Tunapuna dry river, SE of Augustine, Oct 1949, Baker in Trin. herb. 14438 (MO); Government house grounds, 17 Jul 1907, Broadway s.n. (NY); Sans Souci, via Toco, 23 Oct 1932, Broadway 9028 (G, MO); s.loc., 1877-80, Fendler 603 (NY), Fendler 607 (NY); s.loc., Apr 1874, Kuntze 886 (NY); on river banks Sans Souci, 5 Dec 1927, Williams 11889 (NY). Local names. Venezuela. Falc6n: zamurito. Solanum sieberi is most closely related to S. imberbe of Panama, coastal Colombia, and Venezuela, but differs from that species in its elliptic, geminate leaves, flowers with the petals strongly reflexed at anthesis, and its fruiting pedicels with a conspicuous rhomboidal swelling at the apex. Solanum sieberi is polymorphic for the presence of uniseriate trichomes, which if present, are on all parts of the plant. This is reflected in the large number of names given to this species by Pittier (many of which

FLORA NEOTROPICA

194

areonly on herbariumannotationlabels);most of these "taxa"aremerelypubescencevariations.Populationsof thespeciesfromadjacentareasdifferin onlythischaracter, andI do not considerthisa particularlyinformativecharacter in this species. The flowers of S. sieberi have a sweet scent, andthe plantsflower profusely,attracting bees(seeTableIX).Thehabitat a numberof smallgeneralist of S. sieberiis most commonlynearseeps andotherwet places in otherwise rather dry habitats. In Parque Nacional Morrocoy,Venezuela,the plants grew at the edgesof roadsin ditchesof standingwaterandattheedges of mangroveswamps, beyond the reach of salt water. Two Sieber collections are cited in the original description of this species, one unnumbered from Martinique,andthe collection fromTrinidadI have selected as the lectotype. The Trinidadiancollection has been selected as the lectotypebecauseit is betterrepresentedin herbaria,a betterdocumentedcollection, and an annotated specimen exists in G-DC. Henri Van Heurck'sherbariumis housedat AWH,but I have seen orduplicatesofthe Siebercollectionfrom no photographs Martiniquein any United Statesor Europeanherbaria.

VIII. Solanum arboreum species group(S. amnicola, S. anisophyllum, S. arboreum, S. falconense, S. goniocaulon, S. gratum, S. laevigatum, S. lucens, S. plowmanii,S. ramonense,S. ripense,S. roblense, S. tanysepalum). Figs. 101A-D, 102

Shrubs or small trees; young stems and leaves glabrousto densely red-papillose,occasionallypubescent with simple, uniseriatetrichomes (e.g., Solanum falconense). Sympodialunits difoliate and geminate, often stronglyanisophyllous,occasionallyunifoliate(S. laevigatum)or plurifoliate(S. amnicola). Leaves narrowly elliptic to obovate,usuallyglabrouson both surfaces, the apex andbase various.Inflorescencesopposite the leaves or occasionally intemodal, simple, few to many-flowered;pedicel scars closely packed and usually overlapping. Buds various, glabrous or redpapillose.Pedicels at anthesiserector deflexed.Flowers white, small andwith the lobes deflexed at anthesis or largerand fleshy with the lobes planarat anthesis. Fruit usually green and hard at maturity; fruiting pedicels usually erect and thickening at maturity(deflexed in S.falconense, sometimes deflexed from the weight of the fruitin otherspecies). Seeds ovoid-reniform, pale tan to darkbrown, often ratherlarge. Distribution. CentralAmerica to Bolivia, middle to low elevations. Members of the Solanum arboreum species group are glabrous shrubs, usually with strongly anisophyllousgeminateleaves. Therearesome exceptions to this, however: S. amnicola has plurifoliate sympodiaandS. laevigatumunifoliatesympodia.Species in this group all share copiously papillate new growth(althougha few taxa are sometimes glabrous); this often dries reddishin color.

Key to the species in the Solanum arboreumspecies group 1. Sympodial units plurifoliate; leaves not geminate, narrowly obovate to linear, willow-like; shrubs of river courses; inflorescences highly branched. E Peru .................................................................. 52. S. amnicola 1. Sympodial units difoliate; leaves geminate, not willow-like; shrubs or small trees of forests; inflorescences various. 2. Leaves long-petiolate, the petiole clearly differentiated from the lamina, the base oblique. Montane Venezuela. 3. Calyx lobes long-acuminate, equal to or longer than the corolla lobes; axis of the inflorescence absent, consisting of two flowers arising directly from the stem; bark gray, smooth. Cordillera de la Costa, Venezuela .................................................................................................. 64. S. tanysepalum 3. Calyx lobes deltoid, shorter than the corolla lobes; axis of the inflorescence ca. I cm long; bark black, verrucose. Andean Venezuela ......................................................................... 62. S. ripense 2. Leaves not long-petiolate, the petioles various, not markedly differentiated from the lamina, the base not markedly oblique. 4. Flowers small, 0.5-1.2 cm diam., the petals reflexed at anthesis. 5. Calyx lobes long-triangular; nodes swollen; leaves glabrous; shrubs of forest interiors, never rhizomatous. Montane Costa Rica and Panama ........................................ 61. S. ramonense 5. Calyx lobes deltoid (if long-triangular, the leaves sessile and minutely golden-puberulent along the veins beneath); nodes not markedly swollen; leaves glabrous or with minute golden pubescence along the midrib; shrubs or rhizomatous shrublets of low-elevation forests. Central and northern South America ..........................................................54. S. arboreum 4. Flowers larger, 1.2-1.5 cm diam.; the petals generally not reflexed at anthesis. 6. Stems strongly four-angled; leaves large, markedly discolorous, the undersurfaces drying bright golden, the upper surfaces drying bright green. Cloud forests of S Ecuador and N Peru ........................................................................................................................... 56 . S. g oniocaulon 6. Stems and leaves not as above.

195

TAXONOMIC TREATMENT '

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,

B

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FIG. 101. Solanum arboreumand S unifoliatumspecies groups. A S amnicolaplant, Peru (Knapp & Mallet 6651). B. & amnicolaflowers,Peru(Knapp& Mallet 6651). C. S arbomm plan Venezuela (Knapp& Mallet 676!). D. S. arborem young fruit, Venezuela (Knapp & Mallet 6761). E. S. bellum flowers, Ecuador(Knapp & Mallet 6298). F. S. bellum fruit, Ecuador(Knapp & Mallet 6298).

196

FLORA NEOTROPICA

. dark green. ?EasternEcuador lobes long-acuminate;leaves strongly anisophyllous, 7. Calyx

and Peru ...........................................................................................................53. S. anisophyllum 7. Calyx lobes deltoid; leaves not as above. 8. Leaves shiny above; bark shiny and usually dark in color, not dull and grayish. 9. Leaves elliptic, completely glabrous, usually thick and fleshy, the apex usually rounded, the midrib not strongly keeled above; minor leaves not markedly different in shape from the majors; most flowers on an inflorescence opening at the same time, pendant. 10. Leaves drying golden brown. Cordillera de la Costa, Venezuela......... 57. S. gratum 10. Leaves drying with a reddish tinge. NW Peru............................. 60. S. plowmanii 9. Leaves obovate or elliptic, glabrousor densely pubescentabaxially,not fleshy, the apex acute to acuminate,the midrib keeled above; minor leaves orbicular,very different in shape from the major ones; flowers opening one at a time on an inflorescence.

TAXONOMIC TREATMENT

19 7

11. Leaves glabrous; fruit on an erect pedicel, pale green. SW Venezuela and adjacent Colombia ................................................................................... 59. S. l cens 11. Leaves densely pubescent abaxially with simple uniseriate hooked trichomes; fruit on a deflexed pedicel, pale translucent yellow. Sierra San Luis, Venezuela ..................................................... 55. S. falconlense 8. Leaves not markedly shiny above; bark grayish, soft and peeling. 12. Leaves glabrous on both surfaces; new growth sparsely red-papillose. 13. Minor leaves differing from the major ones only in size, not generally in shape, not markedly anisophyllous; calyx lobes minute; shrubs, not rhizomatous. Montane Costa Rica .............................................63. S. roblense 13. Minor leaves orbicular,differing from the major ones in size and shape; calyx lobes various;rhizomatousshrubs.Centraland N South America........... 54. S. arboreium 12. Leaves pubescent abaxially with tufts of simple or branched uniseriate trichomes in the vein axils; new growth densely red-papillose. Montane central Colombia to Peru ...................................................................................................... 58. S. laevigatum

52. Solanum amnicola S. Knapp,Brittonia 38: 295. ca. 0.25 mm long; ovary glabrous;style straight,5-6 1986.Type.Peru.Pasco:km28 Repartici6n-Iscozacin mm long; stigma capitate,bilobed,minutelypapillose. (km 86 Villa Rica-Iscozacin-PuertoMairo),Rio La Fruit a globose, green berry, 1-1.2 cm diam.;fruiting RayanearAmueshacommunityof Laguna,ca. 350 pedicels erect and woody, 1.5-1.7 cm long, ca. 1 mm m, 10?20'S, 75?10'W,22-23 Aug 1984, Knapp & diam. at the base; calyx lobes slightly woody and Mallet 6651 (holotype, BH; isotypes, K, NY, US, accrescentin fruit,3-5 mm long. Seedspale brownishUSM). Figs. O11A,B, 103 green, ovoid-reniform,2-3 x 1.5-2 mm, the surfaces Shrubsgrowing along the rocky,gravelly,or sandy minutely pitted. Chromosomenumbernot known. banksof streamsandrivers,0.5-1.5 m tall;young stems and leaves minutely and densely red-papillose;older stems glabrate;barkgrayish,faintlylongitudinallystriate. Sympodialunitsusually difoliate, geminate,occasionallyplurifoliate.Leavesnarrowlyellipticto obovate, widest at the middle or just above, glabrous above, minutely golden-puberulentalong the veins beneath, the trichomesunicellularandbarelyvisible, blades 1120 x 1-4 cm, with 6-10 pairsof mainlateralveins, these impressedabove, yellowish beneathin dry specimens, the apex long-acuminate,the base attenuate;petioles winged from the decurrentleaf bases, 1.5-2 cm long. Inflorescencesoppositethe leaves or sometimesinternodal, 1-4 cm long, occasionallybranchedonce, bearing 3-5 flowers at a time, butwith many scars,glabrousor minutelyred-papilloseat the tip;pedicel scars closely spaced but not overlapping, beginning at the base of the inflorescence.Budsellipsoideven whenveryyoung, the corollanot earlyexsertedfromthe tube;pedicelsat anthesis deflexed, 0.8-1 cm long, tapering from the base of the calyx tubeto a slenderbase ca. 0.4 mm diam. Flowers with the calyx tubebroadlycup-shaped,1-1.5 mm long, the lobes deltoid, 1-2.5 mm long, the margins white and thickenedin dry material,minutelypuberulent along the margins and tips of the lobes; corolla white,0.8-1 cm diam.,lobednearlyto thebase,the lobes planaror slightlyreflexedatanthesis,thetipsandmargins of thelobesminutelypapillose;anthers2-2.5 x ca. 1mm, poricidalat the tips, theporesteardropshaped;freeportion of the filamentsca. 0.5 mm long, the filamenttube

Distribution (Fig. 106). Onthe banksof streams and rivers in eastern Peru at low elevations, from Amazonas south to Cuzco. Specimens examined. PERU. AMAZONAS: Bagua, slopes of Cerro Unki (Punpuntas), km 44 SaramerizaChiriaco, ca. 52 km NE of bridge over Rio Nieva, ca. 750 m, 4?40'S, 77?38'W, 10 Jun 1986, Knapp et al. 7708 (MO, USM). AMAZONAS/LORETO:Mouth of Rio Santiago, on high bank, Tessmann 4008 (NY); Cuzco: Cerro de Pantiacolla, Rio Palotoa 10-15 km NNW of Shintuya, 500-650 m, 12?35'S, 71?18'W, 8 Dec 1985, Foster et al. 10658 (F, NY); Quispicanchis, environs of village of Quincemil, trail to Rio Nusinescatu, 500-600 m, 13? 15'S, 70?45'W, 22 Apr 1984, Knapp & Mallet 6369 (BH), 24 Apr 1984, Knapp & Mallet 6390 (BH, CUZ, K, US, USM). MADRE DE DIOS: Manu, Aguas Calientes, across and downriver from Shintuya on Rio Alto Madre de Dios, 400-500 m, 12?40'S, 71?15'W, 13 May 1984, Knapp & Mallet 6438 (BH, K, USM).

Solanumamnicolais morphologicallyextremely similarto S. monadelphum,also a shrubof riverbanks in easternPeru.Solanumamnicolacanbe distinguished from S. monadelphumby its slender leaf-opposed inflorescences,long-acuminatecalyx lobes, smallerflowers, and the minute golden trichomes along the veins on the undersideof the leaves. Solanummonadelphum is muchmorecommonthanS. amnicolawherethe two occur in sympatry.Solanumamnicola is known from only a few localities, but is probablyfoundthroughout easternPeruand only rarelycollected.

198

FLORA NEOTROPICA

FIG. 103. Solanum amnicola S. Knapp. (Reproduced with permission from Brittonia 38: 297, fig. 16. 1986.)

53. Solanum anisophyllum VanHeurck& Mull. Arg., rolla.Pedicels at anthesisdeflexed, 1-1.4 cm long, taObserv.Bot. 52. 1870. Type. Peru. San Martin:in peringfromthe calyx tubeto a slenderbase ca. 0.5 mm Peruviaeorientali,propeTarapoto,in montibusse- diam.Flowers with the calyx tube 1-1.5 mm long, the cus flumen Mayo, Spruce 4830 (lectotype, G-DC, lobes long-triangularto deltoid, 0.5-3 mm long, gladesignatedhere;isolectotypesAWH,BM, C [F neg. brous,the marginsof the sinuses white andthickened; 22874], G [Mortonneg. 8503], K, NY, P [Morton corollawhite, ca. 1.8 cm diam.,lobednearlyto the base, neg. 8146]). Fig. 104 the lobes reflexed at anthesis,the tips andbacks of the lobes minutely papillose; anthers 3-4 x ca. 1.5 mm, Shrubs or small trees, 2-4 m tall; young stems poricidalat the tips, the poresteardropshaped;freeporand leaves minutely rusty-papillose;older stems glation of thefilamentsca. 0.25 mm long, the filamenttube brous; bark gray, large white-lenticellate. Sympodial ca. 0.5 mm long; ovaryglabrous;style straight,slightly unitsdifoliate,geminate,stronglyanisophyllous.Leaves clavate,ca. 7 mm long;stigmaa minutelypapillosearea stronglyanisomorphic,glabrouson both surfaces;ma- on the of the club.Fruit a tip globose, greenberry,umjor leaves elliptic, widest at the middle, 12.6-20.8 x bonate when immature, 1.1-1.8 cm diam.;fruiting 4.6-9.3 cm, with 5-10 pairsof mainlateralveins, raised woody, erect, 1.8-2.3 cm, 0.75-1 mm diam.at both above and beneath,the apex acuminate,the base pedicels the base; calyx lobes erectandwoody in fruit,3-4 mm acute to rounded,slightly winged on the petiole; petilong. Seedsunknown.Chromosomenumbernot known. oles 0.9-2 cm long; minorleaves orbicular,2-5 x 1.94.1 cm, the apex acute or obtuse, apiculate, the base Distribution (Fig. 108). In AmazonianEcuador, broadly rounded or cordate; petioles 1-4 mm long. Peru, and Brazil, in wet forests often growing near Inflorescencesoppositethe leaves, simple, 0.3-1.5 cm streamson rocky ground. long, 5-20-flowered, glabrous except for the rustySelected specimens examined. COLOMBIA. papillose distaltips andvery young buds;pedicelscars S.loc.,Andres.n.,K646 Cordillera Oriental, (K).CAQUETA: evenly spaced ca. 0.5 mm apart,beginning ca. 8 mm E slope,banksof Rio Hacha,1000m, 3 Apr 1940,Cuatrefrom the base of the inflorescence.Buds ovoid, rusty- casas 9041 (F,US). VALLE:SantaHelena,aboveTopacio, papillose,the calyx lobes long-acuminatein bud,so that edgeof Farallonesde CaliNationalPark,1930m, 3?30'N, the bud appearspointed, completely enclosing the co- 76?35'W, 10 Dec 1985, Gentry et al. 53005 (MO, NY).

TAXONOMIC TREATMENT W *

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200 ECUADOR. MORONA-SANTIAGO:Vic. of Macuma, path Macuma-Rio Cusutcaime, ca. 50 km NE of Macas, 27 Mar 1973, Lugo S. 3744, 29 Mar 1973, Lugo S. 3779 (GB, MO); along Rio Metzera Grande on Hacienda Sangay (plantationof CompaniaEcuatorianadel Te, S.A.), near Palora, ca. 950 m, 1?40'S, 77?58'W, 15 Feb 1984, Knapp & Mallet 6292 (BH, QCA, QCNE, US). NAPO: Anangu, S bank of Rio Napo 95 km downstream from Coca, ca. 300 m, 0?32'S, 76?23'W, 11-28 Apr 1986, Balslev et al. 62282 (AAU); Lago Agrio, Dureno, indigenous community of Cofan-Dureno, 350 m, 0?02'S, 76042'W,29-31 Dec 1987, Cer6n M. & Cer6n 3072 (MO, NY); Afiangu, NW corner of Parque Nacional Yasuni, close to Rio Napo, ca. 300 m, 0?32'S, 76?22-23'W, 1-30 Oct 1983, Korning & Thomsen 47131, 47209 (AAU); Shushifindi (Nuevo Loja), rd. Coca (Puerto Francisco de Orellana)-Lago Agrio, ca. 50 km NE of Coca, 10 Feb 1973, Lugo S. 3281 (GB, MO); 45 min. walk by trail from Santa Cecelia up Rio Aguarico, 350 m, 20 Mar 1972, MacBrvde & Dwyer 131 (MO, US); Apuya, ca. 10 km S of the Puerto Misahualli turnoff (bridge over Rio Napo), on Puyo-Tena rd., ca. 500 m, 1?07'S, 77?50'W, 23 Jan 1984, Knapp & Mallet 6186 (BH, QCA, QCNE, US); Reserva Biolo6gica Jatun Sacha, 8 km ESE of Puerto Misahualli, ca. 450 m, 1?04'S,77?37'W,3 Jul 1986, Miller et al. 2339 (MO, NY); new rd. on right bank of Rio Napo, 14 km E of Puerto Napo, 9 km E of Atahualpa, 500 m, 1?02'S, 77?40'W, 12 Feb 1987, Palacios & Neill 1547 (MO, NY). PASTAZA:Waorani (Auca) community Centro--Oriente-Tiwaeno, N of village on trail to Rio Tiwaeno, 400-500 m, 10 Aug 1980, Jaramillo & Coello 3382 (NY, QCA, QCNE); Colonia 24 de Mayo, side rd. to rd. Puyo-Puerto Napo, ca. 18-20 km from Puyo, 13 Sep 1968, Lugo S. 403 (GB, MO); Cabeceras on Rio Bobonaza, ca. 12 km from Puyo, 16 Nov 1974, Lugo S. 4596 (GB, MO). PERU. AMAZONAS: Trail N of Cenepa to Tuhushiku Creek, 700-800 ft, 30 Jul 1974, Berlin 1859 (MO). JUNIN: Rio Negro, 800 m, 19 Aug 1960, Wovktoawski5845 (US). LORETO: Above Pongo de Manseriche, streamlet at right bank of Rio Santiago, 250 m, 4 Dec 1931, Mexia 6212a (K, NY, US); above Pongo de Manseriche, right bank of Rio Santiago, 200 m, 10 Dec 1931, Mexia 6247 (BH, F, K, NY, TEX, U, US); mouth of Rio Santiago, Tessmann3968, 4531 (NY). SAN MARTIN:Venceremos, near Amazonas border, km 291 on Rioja-Pomacocha rd., 1850 m, 5045'S, 77?40'W, 10 Feb 1984, Gentry et al. 45319 (MO, NY); Lamas, Alonso de Alvarado, Cerro Blanco on rd. to Moyobamba, 9001000 m, 13 May 1973, Schunke V 6268 (MO); Pedro Ruiz-Moyabamba rd., km 390, Venceremos, 2040 m, 5?50'S, 77045'W, 2-3 Aug 1983, D.N. Smith & Vdsquez S. 4578 (MO, NY); San Roque, 1350-1500 m, Jan-Feb 1930, Williams 7197, 7616 (F). UCAYALI: Middle Ucayali, boca de Yarina, Tessmann 3497 (NY). BRAZIL. ACRE: Near mouth of Rio Macuahan (tributary of Rio Yato), 9?20'S, 69? W, 14 Aug 1933, Krukoff 5510 (F, NY). AMAZONAS: Sao Paulo de Olivenca, near Palmares, 11 Sep-26 Oct 1936, Krukoff 8377 (K, NY).

FLORA NEOTROPICA

Solanumanisophyllumis relatedto S. arboreum, and differs from it in its long-acuminatecalyx lobes and smaller flowers. The long-acuminatecalyx lobes aresuperficiallysimilarto those ofS. tanysepalumfrom montane coastal Venezuela. The two species are not closely relatedwithin the group and can be easily distinguishedby othercharacters.Solanumanisophyllum has elongate inflorescences with small flowers and closely spacedpedicelscarsandorbiculateminorleaves, whereasS. tanysepalumhas sessile inflorescenceswith larger,fleshy flowers and elliptic minorleaves. Solanum anisophyllum is a shrub of primary forest andis a sparsespecies (Rabinowitz, 1981) with few individualsin any given population. I have selectedthe specimenat G as the lectotype of Solanum anisophyllumas the Mortonphotograph (see above) of it is widely distributedand it was one of two (theotherat AWH)citedin the originaldescription.

54. Solanum arboreum Dunal, Solan. Syn. 20. 1816. Type.Venezuela.Sucre:Cumanacoa,Humboldt& Bonplands.n. (holotype,P-Bonpl. [microficheIDC 6209-2:61 1.6]; isotype, B-W [F neg. 2889]). Figs. 101C,D, 105 Solanum dolichostylum 0. E. Schulz in Urb., Symb. Antill. 6: 158. 1909. Type. Trinidad and Tobago. Tobago: In peludos greg. ad Great Dog River. Nov 1889, Eggers 5797 (lectotype, US, here designated; isolectotypes, B [destroyed], MPU). Solanum enchylozumBitter, Repert. Spec. Nov. Regni Veg. 18: 64. 1922. Type. Costa Rica. Lim6n: Forets de Shirores, Talamanca, ca. 100 m, Feb 1895. Pittier & Tonduz 9193 (holotype, BR). Solanum kenoyeri Standl., Publ. Field Columbian Mus., Bot. Ser. 4: 157. 1929. Type. Panama.Canal

Area(CanalZone):BarroColoradoIsland,Lutz Trail, 22 Jul 1927, Kenoyer 515 (holotype, US).

Solanumor'galeC. V. Morton,Contr.U.S. Natl.Herb. 29: 53. 1944. Type. Colombia. Santander: San Fernando between Rio Carare and Puerto Berrio. 300 m, 30 Jun 1939, Haught 2850 (holotype, US; isotype, F). Solanumn ripivagumPittier,Cat. Fl. Venez. 2: 382. 1947. nom. nud. Type. Venezuela.Aragua:Orillas del Rio Chuao, en playas arenosas, 25 m. s.d., Pittier

12182(F, MO,US, VEN). Rhizomatous shrubs to treelets, 0.5-3 m tall; young stems and leaves densely puberulentwith appressed uniseriatetrichomes 0.1-0.3 mm long, these reddishor somewhatgolden;olderstemsglabrate,rather thick;barkof older stems grayishandpeeling. Sympodial units difoliate, geminate.Leaves obovate, sometimes elliptic, widest at the middle or in the distalthird of the leaf blade, glabrouson both surfaces,in Central

TAXONOMIC TREATMENT

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FIG. 105. SolanumarboreumDunal. Holotype in P-Bonpl.

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Americanmaterialoccasionally with scatteredminute uniseriate or unicellular trichomes along the midrib beneath;majorleaves 8-28 x 4-13 cm, with 7-12 pairs of main lateralveins, these impressed above, prominentandreddishin drymaterialbeneath,the apex acute to acuminate,the base acute, often oblique, to somewhat auriculatein Costa Ricanmaterial;petioles 0.3-2 cm long; minorleaves differingfromthe majorones in size and shape, orbicularto elliptic, 1.5-9 x 1.4-6 cm, the apex acute,the base rounded;petioles2-3 mm long. Inflorescences opposite the leaves, simple, 0.2-4 cm long, puberulentwith the same appressed uniseriate trichomes as those of the young stems and leaves, 530-flowered, but only bearing one or two at a time; pedicel scars closely spaced and overlapping,beginning at thebase of the inflorescence.Budsglobose when young, laterellipsoid with the exsertionof the corolla, if the calyx lobes are long-triangular,these appearing somewhatcaudatein bud.Pedicels at anthesisdeflexed, 0.4-1 cm long (rarelyshorter),taperingfromthe calyx to a slender base ca. 0.5 mm diam. Flowers with the calyx tubeconical, 0.75-1 mm long, the lobes variable, deltoid to long-triangular,0.5-2 mm long, glabrousor puberulentwith the appressed,often golden, trichomes of the rest of the inflorescence;corolla white, 0.6-1.5 cm diam., lobed 3/4 of the way to the base, the lobes planaror reflexed at anthesis, the tips and marginsof the lobes minutely papillose; anthers 1.5-4 mm long, poricidal at the tips, the pores teardropshaped; free portionof thefilaments0.25-0.5 mm long, the filament tube 0.25-0.5 mm long; ovary glabrous;style straight, in short-styledflowers 1-1.5 mm long, in long-styled flowers 3-6 mm long; stigma a broadenedareaon the tip of the style, the surfaceminutely papillose. Fruit a globose, greenberry(redfideHaught 2850), 1-1.5 cm diam.;fruitingpedicelserectandwoody,0.8-2 cm long, varying in length even on a single plantand in a single locality,the surfacesroughandrathercorky, 1-1.5 mm diam.at the base. Seeds pale yellowish-tan,ovoid-reniform, 3-4 x 1.5-2.5 mm, the surfacesminutelypitted. Chromosomenumber:n = 12 (vouchers Knapp 802, 4862, 6835).

FLORA NEOTROPICA

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FIG. 106. Distribution of Solanum arboreum (solid circles) and S. amnicola (open circles).

85?20'W, 31 Oct 1987, Herrera 1064 (INB); CARTAGO: 2 km E of Mufieco, 21 Apr 1969, Davidse & Pohl 1687 (F, MO); CATIE, 4 km S of Turrialba,1 Apr 1977, Haber 100 (F, MO); 12 km S of Turrialbaby air,4 km S of Pejibaye along Rio Gato, 700 m, 9?48'N, 83042'W, 16-17 Apr 1983, Liesner 14371 (MO); ca. 8 km S of Cartago by air, Finca El Chaparral, 4.5 km S of bridge of Agua Caliente at Lourdes, 1500 m, 9?49'N, 83?55'W,21 Apr 1983, Liesner & Judziewicz 14633 (MO); Guayacan, ca. 20 km NNE of Turrialba,500 m, 3 Aug 1974, Maas 1111(F, U). HEREDIA: Rio PuertoViejo 2 km upstreamfrom confluence with Rio Sarapiqui, Finca La Selva, 100 m, 10?26'N, 84?00'W, 14-17 Jun 1968, Burger & Stolze 5751 (CR, F, MO, NY); near Tirimbina E of Rio Sarapiqui, 150-250 m, 10?24'N, 84?07'W, Burger & Burger 8085 (CR, F, MO); near Puerto Viejo along rd. near Rio Sucio, 20 m, 27 May 1976, Croat 35765 (MO); Finca La Selva, Puerto Viejo, 90 m, 13 May 1969, Frankie 129 (F, MO); near Rio Puerto Viejo, ca. 2 km upstream of confluence with Distribution (Fig. 106). Widely distributedin Rio Sarapiqui, Finca La Selva, 100-200 m, 10?26'N, northernSouth AmericaandwesternCentralAmerica 84?00'W, 26 Apr 1973, Gentry & Burger 3024 (F, MO); from Costa Rica to Trinidad and Tobago, at low to Finca La Selva, Rio Sarapiqui, ca. 100 m, 20 Aug 1980, Knapp 802 (BH); La Selva, near Puerto Viejo, line B middle elevations, generally in wet places. Washington plot II, 90 m, 29 May 1971, Opler 187 (F); Selected specimens examined. COSTA RICA. La Selva near Puerto Viejo, River Rd. near El Salto, 90 ALAJUELA: ReservaBiolo6gicaMonteverde,Rio Pefias m, 21 Apr 1972, Opler 748 (F); Tirimbina, 700 ft, 29 Blancas, Laguna Poco Sol, 950 m, 10?21'N, 84?40'W, 8 May 1971, Proctor 32123 (LL). LIM6N: Vic. of Lim6n, 0-100 m, 3-7 Sep 1971, Burger & Burger 8442 (F, MO); Aug 1989, Bello 1082 (INB); R.B. Monteverde, Children's Rainforest, Finca Villalobos, 8 km from Volcan SW-most ridge of Cerro Coronel, NW-facing slope, just Arenal, 1000-2000 m, 10?23'N, 84?43'W, 21 Apr 1990, S of Rio Colorado, 10-80 m, 10?40'30"N, 83?39'30"W, Haber & Zuchowski 9866 (INB); Upala, Dos Rios, 5 km 17-18 Sep 1986, Davidse & Herrera 31325 (INB, MO); S of Brasilia, R bank of Rio Pizote, 500 m, 10?55'N, vic. of Portete near Puerto Lim6n, 12 Feb 1965, Godfrey

TAXONOMIC TREATMENT203 66424 (MO); El Progreso, Fila de Matama, Valle de la Estrella, 1350 m, 9?47'18"N, 83?08'45"W, 19 Apr 1989, Herrera & Chac6n 2661 (INB); Guapiles, Los Angeles, San Miguel, Rio Blanco, 700 m, 10?07'30"N, 83?50'45"W, 24 Feb 1990, Herrera & schik 3793 (INB); 2.5 km E of Guapiles, 250 m, 17 Jul 1964, Jimenez M. 2100 (NY); bridge over Rio Toro Amarillo 5 km SW of Guapiles, 300-400 m, 9 Feb 1965, Jimenez M. 2800 (F, NY); Parque Nacional Tortuguero, Est. Agua Fria, 5 km from house, Sendero Las Lomas, 40 m, 10?25'N, 83?34'W, 26 Oct 1987, Robles 1145 (INB); Los Diamantes, 12 May 1956, Schubert 1327 (US); vic. of Guapiles, 300-500 m, 12-13 Mar 1924, Standley 37153, 37171, 37320 (US); Barra del Colorado, N side, 0-2 m, 10?47'N, 83?35'W, 12 Sep 1986, Stevens & Montiel 24192 (INB, MO); forest of Shirores, Talamanca, 100 m, Feb 1895, Tondiuz9189 (US). PUNTARENAS: Reserva Forestal Golfo Dulce, Rio Tigre, Finca of Ram6n Bellido, 2 km W of Tigre, 200 m, 8?31'N, 83?23'W, 8 Sep 1997, Azofeita 398 (INB); R.B. Monteverde, Refugio Veracruz, Finca las Salazar, Rio Veracruz,Pacific slope, 1500 m, 10?15'N, 84?48'W,9 Jan 1990, Bello 1719 (INB); Peninsula de Osa near Golfo Dulce, ca. Pto. Jim6nez, Rio Nuevo, 40 m, 4 Apr 1930, Cujodonti 195 (W); R.B. Monteverde, Penias Blancas river valley, E. Rockwell parcel, 800 m, 10?19'N, 84?43'W, 18 Mar 1989, Haber 9187 (INB); Osa Peninsula, Parque Nacional Corcovado, Rio Rincon just above Cerro de Oro (Cerro Degro of maps), 100-200 m, 8?30'N, 83027'W, 11 Jul 1981, Knapp & Mallet 886 (BH, CR); Alajuela border, 2-5 km E of Monteverde on Continental Divide, 1580-1700 m, 10?18'N, 84?46'W, 4 Mar 1978, Lawton 1244 (F); Parque Nacional Corcovado airfield near Pavo, 5 m, 8?30'N, 83?37'W,7 Jul 1977, Liesner 3020 (MO); N end of Golfito Bay 4 km W of Golfito hospital, 10 m, 5 Mar 1971, Nee & Mori 3549 (F). PANAMA. CANALAREA:Barro Colorado Island, 1931, Aviles 17 (F), 9 Jul 1931, Bailey & Bailey 540 (BH, F), 8-10 Aug 1940, Bartlett & Lasser 16740 (MO), 3 Oct 1968, Croat 6680 (MO), Drayton trail, 7 Oct 1968, Croat 6768 (F, MO, NY); Pipeline Rd. 2.5 mi from gate, 4 Apr 1970, Croat 9352 (F, MO, NY); Barro Colorado Island, 30 Apr 1970, Croat 10119 (MO), Zetek trail 300, 8 Jul 1970, Croat 11151 (F, MO, NY), 31 May 1971, Croat 14851 (MO), Snyder-Molino trail 450, 16 Jun1971, Croat 15002 (MO); Pipeline Rd. 2 mi from Gamboa gate, 8 Aug 1971, Croat 16610 (MO); Barro Colorado Island, Shannon trail, 25 Jun 1972, Croat 17373 (MO), 21 Apr 1970, D'Arcy 4296 (MO); Pipeline Rd. 1/2 mi N of gate N of Gamboa, 2 May 1971, D'Arcv 5208, 5210 (MO); Barro Colorado Island, Zetek trail, 22 Jul 1960, Ebinger 546 (MO, US); along Achiote Rd., 2 mi from Achiote, 23 Jun 1977, Folsom 3869b, 3869c (MO); Madden Forest, Las Cruces trail, 3 Aug 1971, Gentry 1388 (MO); Pipeline Rd., 1 Aug 1972, Gentry 5603 (MO); near Rio Frijoles, 9 Sep 1972, Gentry 5827 (MO); Rio Providencia 3 km SE of Achiote, 5100 m, 4 Dec 1973, Gentry & Nee 8647 (MO); Maumes Station, Aug 1861, Hayes 399 (K); Barro Colorado Island, Barbour-Lathrop trail, 200-300 m, 24 Nov 1948, Killip 39986 (K, US); 2 mi N of Gamboa on Pipeline

Rd., along Rio Frijolito, 0-50 m, 9?10'N, 79?45'W, 26 Apr 1982, Knapp & Schmalzel 4862 (MO); Madden Forest Preserve, Las Cruces trail, 8 Apr 1969, Lewis et al. 5326 (MO); Pipeline Rd. 4 km NW of Gamboa, 50 m, 3 Oct 1973, Nee 7224 (MO); headwaters of Rio Caraya, 6 km E of Gamboa, 100-180 m, 27 Dec 1973, Nee 9017 (MO), Nee 9032 (MO, POM, U, US); along Rio Mendosa above first gorge, 1/2-2 km upstream from Pipeline Rd. bridge, 8 km NW of Gamboa, 100-135 m, 16 Apr 1974, Nee & Smith 11377 (MO); Cuatro Calles hills near Matuchin, 20 m, 23 Jul 1911, Pittier 4048 (US); Barro Colorado Island, Gatun Lake, 17 Jan 1924, Standley 31358 (US), ca. 120 m, Standley 41143 (F, US), Miller trail, Jul 1931, Starry 291 (F, MO); 6 mi N of Gamboa near Rio Frijoles, 6 Oct 1965, Tyson 1456 (MO); 12 mi S of Colon on Rio Providencia, 16 May 1966, Tyson & Blum 3980 (MO). BOCASDEL TORO: 15 km S of town of Changuinola, 900-1500 ft, 13 Dec 1979, Antonio 3119 (MO); ca. 1.5 mi W of Almirante, 16 Oct 1965, Blum 1363 (MO); above RR stop at milla 7.5, 26 Jul 1971, Croat & Porter 16236, 16298 (MO); hillside above Almirante, 28 Nov 1971, Gentry 2707 (MO); Rio Changuinola near Changuinola, 9 Sep 1963, Dwyer 4399 (MO); Chiriquicito to 5 mi S along Rio Guarumo, 5-7 Jun 1967, Lewis et al. 2122 (MO); Almirante near Nigra creek, 15 Aug 1964, McDaniel 5069 (MO); region of Almirante, Jan-Mar 1928, Proctor Cooper 217, 383 (F); Fish Creek Mtns., vic. of Chiriqui Lagoon, 18 Apr 1941, von Wedel 2268 (MO, US), 30 Apr 1941, von Wedel 2367 (MO, US); Changuinola River, vic. of Chiriqui Lagoon, 18 Aug 1941, von Wedel 2603 (MO, US); vic. of Nievecita, 0-50 m, 8-19 Aug 1938, Woodson et al. 1828 (MO, NY). CHIRIQUI: Quebrada Macho, Burica Peninsula, 140 m, 22 Feb 1973, Croat 22117 (F, MO, NY, US); Burica Peninsula, Puerto Armuelles, 3 Feb 1967, Hladik 184 (MO); Burica Peninsula, Quebrada Mellize 6 mi S of Puerto Armuelles, 0-150 m, 5 Mar 1973, Liesner 442 (F, MO, NY, US). COLON:Along Rio Guanche ca. 3-5 mi inland, 10-100 m, 3 Aug 1974, Croat 26208 (MO); lower Rio Guanche, 17 Apr 1974, Dressler 4654 (MO); S approach to Cerro Bruja from Rio Escandaloso, 20 May 1978, Hammel 3217 (MO); ca. 1 km from the Colon-Portobelo rd. on the Rio Guanche, 0-50 m, 27 DE SAN Aug 1980, Knapp & Kress 823 (BH). COMARCA BLAS:SE of Puerto Obaldia, 18 Aug 1971, Croat 16731, 16781 (MO); Rio Cuadi, ca. 46 m, 15 Mar 1968, Duke 15465 (WIS); above Puerto Obaldia, 19 Aug 1971, Gentry 1546 (MO); headwaters of Rio Mulatopo, 17 Aug 1967, Elias 1774 (MO, US); Marraganti and vic., 10200 ft, 3-9 Apr 1908, Williams 791 (NY, US). DARIEN: Vic. of Pinogana, ca. 20 m, 6 Oct 1938, Allen 930 (F, MO, NY, US); N Puerto GuayaboGrande,trail to ridgetop, 200-600 ft, 21 Apr 1980, Antonio & Hahn 4332 (MO); Rio Uruceca, Nov 1967, Bristan 1445 (MO, WIS); vic. of gold mine at Cana, 500-600 m, 26 Jul 1976, Croat 37597 (MO); vic. of airstrip at Cana gold mine, 480 m, 29 Jul 1976, Croat 37961 (MO); 10 km NE of Jaque, headwatersof Rio Pavarando,1400 ft, 31 Jan 1981, D 'Arcy, & Sytsma 14503, 14530 (MO); Cerro Pidiaque, 5 Apr 1966, Duke 8083 (MO, WIS); Quebrada Nigua, below

204 Santa F6, 29 Sep 1966, Duke 8826 (MO, WIS); Agua Fria ca. 8 mi N of Santa F6, ca. 50 m, 13 Feb 1967, Duke 10111 (MO, US, WIS); trail from Rio Pucro to Quebrada Maskia, 22 Jun 1967, Duke 13083 (WIS); Cerro Pirre, 2500-4500 ft, 9-10 Aug 1967, Duke & Elias E13669 (MO); between Paya and Palo de Las Letras, 30 Aug 1967, Duke & Kirkbride14029 (MO); Rio Sabana above Santa F6, 14 Sep 1967, Duke 14080 (MO); Tumaganti, ca. 300 m, 18 Sep 1967, Duke 14145 (MO); Santa F6, ca. 15 m, 20 Sep 1967, Duke 14286 (MO); Rio Pirre,near town of Pirre,27 Dec 1972, Gentry6927 (MO); Cerro Tacarcuna, W ridge, 1550-1700 m, 2 Feb 1975, Gentryi& Mori 14112 (MO); S slope of W-most peak of Tacarcuna massif above Rio Pucuro, 800-1200 m, 23 Jul 1976, Gentry et al. 16920 (MO); E base of Cerro Sapo, 1300 ft, 30 Jan 1978, Hammel 1135 (MO); Cerro Sapo, ca. 2500 ft, 1 Feb 1978, Hammel 1243 (MO); Mannene to mouth of Rio Coasi, 28 Apr 1968, Kirkbride & Bristan 1536 (MO, NY); S of El Real, Alturas del Nique, near Cana mine, 650-800 m, 7?45'N, 77?44'W, 21 Aug 1987, McPherson 11527 (BM, MO); vic. of Cana, 1750 ft, 23 Jun 1959, Stern et al. 498 (MO, US), 24 Jun 1959, Stern et al. 696 (MO, US); trail NW of Cana, 600 m, 28 Jul 1976, Sullivan 683 (MO). HERRERA:10 km W of Las Minas on rd. to El Toro, 200-600 m, 24 Jan 1981, D'Arcv & Sytsma 14346 (MO); hills above Chepo de las Minas, 700 m, 19 Dec 1977, Folsom et al. 6978 (MO). Los SANTOS:Vic. of headwaters of Rio Pedregal, 25 mi SW of Tonosi, 2500-3000 ft, 7 Dec 1967, Lewis et al. 2888 (MO); vic. of Tonosi, rd. between Tonosi and Guanico, 28 Feb 1963, Stern et al. 1886 (MO, US); 10 mi N of Tonosi, 23 Jan 1966, Tysonet al. 2949 (MO). PANAMA:Cerro Campana, 31 Dec 1939, Allen 2088 (MO), 10 Sep 1970, Croat 12072 (MO, US), 16 May 1971, D'Arcy & Dressler 5506, 5493 (MO), ca. 800 m, 29 Mar 1977, D'Arcy 11118 (MO); Isla de Bayano, 12 Feb 1977, Correa A. et al. 2983 (MO); along PanAm hwy. near Jenine, Rio Canita, 24 Sep 1961, Duke 3873 (MO, US); halfway between El Llano and Rio Mamoni, 9-10 Sep 1962, Duke 5562 (MO); headwaters of Rio Corso (off Rio Pacora), ca. 500 m, 9 Jun 1967, Duke 11924 (MO, WIS); Piria-Cafiasas trail near Piria, ca. 100 m, 22 Sep 1967, Duke 14332 (MO, US, WIS); between Cafasas and Sabalo, ca. 100 m, 26 Sep 1967, Duke 14484 (US, WIS); Cerro Campana, 3/4 way to summit, 31 Mar 1969, Dwyer et al. 4701 (MO); 4-5 km E of Rio Ipeti on PanAm hwy., 9 Feb 1977, Folsom & Collins 1691 (MO); stream from Serrania de Maje, 10 Feb 1977, Folsom & Collins 1720 (MO); Cerro Campana, 30 Mar 1977, Folsom et al. 2299 (MO); from Torti to Pilota de Toro, overlooking Torti Arriba, 27 Aug 1977, Folsom et al. 4995 (MO); Pilota de Toro (near Torti), 4 Dec 1977, Folsom et al. 6782, 6797 (MO); Isla Bayano, 31 Jan 1976, Garibaldi 34, 43 (MO); dam site near Rio Bayano beyond Cafita, 1 Sep 1971, Gentry & Tylson1647 (MO); Rio Pasiga, to Rio Maestro, 30 Oct 1971, Gentry 2318 (MO); Altos de Campana, near motel Sulin, 3060 ft, 26 Dec 1977, Mendez 198 (F, MO); Indio, Madden Lake, 13-15 Apr 1937, Miller 2046

FLORA NEOTROPICA (US); ca. 25 km NE of Cerro Azul on Rio Piedras, 550 m, 20-22 Nov 1974, Mori & Kallunki 3312 (MO); jct. of Rios Pacora and Corso to headwaters of Rio Corso, 9 Jun 1967, Oliver 2377 (MO, SCZ); Cerro Campana, Su Lin motel, 25 Mar 1969, Porter et al. 4274 (MO); 2 mi E of El Llano, 15 Oct 1965, Tyson 1746 (MO). VERAGUAS:Coiba, May 1847, Seemann 642 (K). COLOMBIA. Locality unreadable, 400-600 m, 19 Feb 1876, Andre 1796 (K); s.loc., Multis s.n. (US). ANTIOQUIA:Vic. of Planta Providencia 28 km SW of Zaragaza, valley of Rio Anori in area of confluence of Quebrada la Tirana and Rio Anori, 3 km upriver from Planta Providencia, 400-700 m, 23 Mar 1977, Alverson et al. 248 (WIS); Jardin-Riosucio rd., 7 km from Jardin, 2020 m, 5030'N, 75050'W, 7 Jun 1987, Callejas et al. 3714 (NY); 9-20 km SE of Remedios, Remedios-Puerto Berrio rd., Vereda Santa Lucia, 500-620 m, 7?05'N, 74?15'W, 17 Sep 1987, Callejas et al. 5267 (NY): Puerto Valdivia, 2 Feb 1943, Bro. Daniel 3386 (US); near Villa Arteaga, ca. 150 m, 10 Oct 1947, Gutierrez V.& Barklev 17C110 (US); Guapa, 53 km S of Turbo, 50 m, 3 May 1945, Haught 4620 (NY, US); valley of Rio Medellin near Porcesito, 1100 m, 16 Apr 1946, Hodge 6802 (F); near Villa Arteaga, 150 m, 4-8 Aug 1947, Hodge 6979 (US); Rio Anori valley near Planta Providencia, 350600 m, 16 Aug 1976, Shepard 555 ((WIS); km 9 of rd. to Vereda La Hundida, WSW of Ituango, 1700 nm, 7?10'N, 75?47'W, 11 May 1988, Zarucchi & Betancur 6428 (MO, NY). CESAR: Sierra de Perija, E of Manaure, Hacienda Nuevo Horizonte, El Podrido, 1500-1600 m, 15 Nov 1959, Cuatrecasas & Romero Castanieda25386 (US). CHOCO: Quebrada Argueta near Bahia Solano, 18 Apr 1982, Dressler 6038 (BH); upper Rio San Juan, slopes of Cerros la Mojarra, Quebrada El Salado, 80100 m, 26 Jun 1983, Espina et al. 1353 (NY); Rio San Juan below Docord6, 0 m, 4?15'N, 77?22'W, 30 Mar 1979, Forero et al. 4372 (MO); Rio Mutata, tributaryof Rio El Valle between base of Alto de Buey and mouth of river, 100-150 m, 9 Aug 1976, Gentry & Fallen 17461 (MO, NY); trail from coast N of Mecana into Serrania de Baud6, 0-100 m, 6?15'N, 77?25'W, 5 Mar 1983, Gentry & Juncosa 40967 (MO, NY); Bahia Solano, 2 m, 6?16'N, 77?21'W, 2 Jan 1984, Juncosa 1647 (MO, NY); Bahia Utria, along Rio Sampechi, 0 m, 1 Feb 1947, Haught 5515 (US); Parque Nacional Los Katyos, above El Lim6n, 300-380 m, 19 Feb 1976, Le6on434 (MO); Parque Nacional Los Katyos, Cararicas-Quebrada Tembladera, 80-100 m, 9 Dec 1976, Leon 706 (MO). CUNDINAMARCA:Cordillera Central, Quetame, Rio Contador, 1450 m, 21 Sep 1963, Uribe Uribe 4472 (US). MAGDALENA: Poponte, Magdalena Valley, 30 Sep 1924, Allen 756 (MO); Sierra Nevada de Santa Marta, SE slope Rio Donachui, ravine SE of Donachui, 1350-1500 m, 22 Sep 1959, Cuatrecasas & Romero Castahleda 24353 (US); s.loc., Sep 1852, Holton s.n. (K); Sierra Nevada de Santa Marta, N of Finca Cecelia, Quebrada Indiana, ca. 1820 m, 10?59'N, 73?58'W, 31 Aug 1972, Kirkbride 2024 (NY), 3 Sep 1972, Kirkbride 2077 (US); Sierra Nevada de Santa Marta, 4500 ft, 1898-1899, Smith

TAXONOMIC TREATMENT205 1186 (F, NY, US). SANTANDER:Vic. of BarrancaBermeja, Magadalena Valley between Sogomaso and Colorado rivers, Camp Zarzal, 100-500 m, 15 Dec 1934, Haught 1462 (US); vic. of Puerto Berrio, between Carare and Magdalena rivers, San Juan valley, 100-700 m, 15 Jun 1935, Haught 1781 (US); Aguas Blancas creek 24 km S of El Centro, vic. of Barranca Bermeja, Magdalena Valley, between Sogomaso and Carare rivers, 100 m, 10 Oct 1936, Haught 2016 (US); rd. to Barranca, 1073 m, 16 Oct 1977, Renteria et al. 722 (NY); 15 leagues SE of Barranca Bermeja, 3 km from left bank of Rio Op6n, ca. 200 m, 9 Oct 1954, Romero Castaneda 5011 (US); vic. of Tona, 1900-2000 m, 17 Feb 1927, Killip & Smith 19450 (NY, US). VALLEDE CAUCA:Las Delicias, NW of Restrepo, 1000 m, 2 Aug 1962, Robinson 211 (K); Buenaventura, community of San Isidro, 236 m, 3?59'N, 76?57'W, 15 Nov-6 Dec 1979, van Royen et al. 349 (MO, NY); Cerro del Ingles, Cordillera Occidental, Serrania de los Paraguas, 1 hour by jeep from El Cairo, 2400 m, 1 Jan 1987, Silverstone-Sopkin et al. 2875 (MO, NY). VENEZUELA. ANZOATEGUI: Guanta, 20 Aug 1935, Potter 5157 (US). APURE: 25 km E of El Nula, 4 Jul 1983, van der Wei-ff & Gonzalez 4834 (BM). ARAGUA:Parque Nacional Henri Pittier, rd. to Choroni, Rio Romerito, N slope, 550 m, 29 May 1980, Benitez de Rojas 2748 (MY); behind station at Rancho Grande, 1100 m, 23 Apr 1969, Ferrari 762 (MY); Parque Nacional Henri Pittier, behind station at Rancho Grande, 1140 m, 7 Sep 1975, Huber 135 (VEN); ParqueNacional Henri Pittier, Rancho Grande, behind station trail to Pico Guacamayo, 1100-1400 m, 10?21'N, 67?42'W, 27 Oct 1984, Knapp & Mallet 6853, 28 Oct 1984, Knapp & Mallet 6858 (BH, MY, US, VEN); Playa Grande near Choroni, Pittier 14037 (US). BOLIVAR:3-4 km SE of Los Patos, N of Rio Hacha and Rio Supamo, 30 km S of Manteco, 365 m, 9 Aug 1960, Steyermark 87069 (F, NY, US, VEN); Rio Toro (Rio Grande) between Rio Reforma and Puerto Rico, N of El Palmar, 200-250 m, 12 Dec 1960, Stevermark 88007 (NY, U, US, VEN); near El Palmar,ca. 75 m, 8?05'N, 61?50'W,23 Nov 1967, Wessels Boer 2067 (VEN). DELTAAMACURO:E of Rio Grande, directly E of El Palmar,6 Jul 1975, Gentry & Berry 14954 (MO); Serrania Imataca, 10 km S of Cerro Muerto, 275 m, 7 Nov 1955, Wurdack & Monachino 39616 (NY). DISTRITOFEDERAL:Near Colonia Tovar, 1854-1855, Fendler 979 (K, MO, NY). FALCON: Sierra San Luis, ridges around Hotel Parador, ca. 7 km S of Curimagua, 1300-1500 m, 11?10'N, 69?35'W, 28 Sep 1984, Knapp & Mallet 6674 (BH, MY, US, VEN); along old Curimagua-San Luis rd., ca. 15 km E of Curimagua, ca. 1000 m, 11?12'N, 69?40'W, 29 Sep 1984, Knapp & Mallet 6692 (BH, K, MY, US, VEN); Cerro Chichiriviche. above La Soledad, 200 m, 10?51'N, 68?20'W, 5 Sep 1974, Stevermark & Manara 110830 (NY, VEN). LARA: Parque Nacional Yacamb6, 19 Sep 1980, Ferndcndez 3711 (MY); slopes of mountain between Santo Domingo and Los Quebraditos, S of Sabanetas, above Humocaro Bajo, 2430-2745 m, 8 Feb

1944, Steyermark 55407 (F). MERIDA: EstanquezParamo Los Colorados rd., 1500 m, 26 Aug 1981, Aymard 405 (MY); s.loc., 15 Feb 1957, Bernardi s.n. (NY); Quebrada La Lajita, ca. 4 km N of La Azulita, 800 m, 8?43'N, 71?26'W, 1 Jul 1980, Davidse & Gonzalez 18863 (NY, VEN); quebrada just S of La Azulita, ca. 1100 m, 8?40'N, 71?25'W, 22 Oct 1984, Knapp & Mallet 6804 (BH, K, MY, US, VEN); 0.5-2 km above dam site on Rio Guaimaral, 7045'N, 71?29'W, 15 Mar 1981, Liesner & Gonzalez 10615 (VEN); Ejido, 1893-4, Mocquerys 1029 (P, US, VEN); near El Morro, banks of Rio Nuestra Sefiora, 14 May 1964, Trujillo6376 (MY). MONAGAS: Parque Nacional El Guacharo, trail from Cueva El Guacharo to Cerro Negro, 1000-1200 m, 10?ll'N, 63?30'W, 15 Oct 1984, Knapp & Mallet 6756, 6757, 6758, 6759, 6760, 6761 (BH, K, MY, US, VEN). NUEVAESPARTA:NE part of island between Espiritu Santo and Cerro Palma Real, 800 m, 22 Sep 1973, Benitez de Rojas 1666 (MY); Copey, 350 m, Aug 1953, Gines 3944 (US); Cerro Copey, 700 m, 18 Jun 1977, Hoyos 3053, 15 Oct 1977, Hoyos 4183 (VEN); Juan Griego trail, 450 m, 22 Jul 1903, Johnston 70 (F, K, NY, US). TACHIRA: On Las Dantas-Buena VistaPuente Alianza rd., ca. 12 km S of Las Dantas, 11001200 m, 7?40'N, 72?25'W, 24 Oct 1984, Knapp & Mallet 6835, 6838 (BH, K, MY, US, VEN); vic. of Mata Mula, N of Delicias on rd. to Bramo6n,1750 m, 7?39'N, 72?26'W, 26 Jul 1979, Steyermark & Liesner 118719 (VEN); Sierra El Casadero, 13 km N of Rubio, between Las Dantas and Los Adjuntas, 900-1050 m, 7?43'N, 72?23'W, 12 Nov 1979, Stevermarket al. 120120 (VEN); Sierra El Casadero, along hwy. between Las Dantas and Los Adjuntas, 850 m, 7?45'N, 72?25'W, 12 Nov 1979, Steyermark et al. 120149, 120164 (VEN). TRUJILLO: Near Cuicas, ca. 950 m, 26 Apr 1968, Benitez de Rojas 272 (MY); near El Cacao, leaving Cuicas, ca. 900 m, 11 Oct 1971, Benitez de Rojas 1143 (MY); 15 mi inland from La Ceiba, 15 Mar 1931, Reed 961 (US). YARACUY: Sierra de Aroa, 6-7 Jul 1953, Aristeguieta & Pannier 1959 (VEN); near Taria, 75 m, 29 May 1944, Steyermark 56871 (VEN); between Salom and Temerla, 1000 m, 11 Nov 1967, Steyermark et al. 100354 (VEN); Cerro Negro, headwaters of Rio Cocorotico, above San Felipe, 1100-1250 m, 13 Nov 1967, Steyermark& Wessels Boer 100399 (VEN). TRINIDAD AND TOBAGO: S.loc. Herb. Trin. 2779 (NY). TRINIDAD:Maracas falls, 6 Jan 1949. Baker & Simmonds 14337 (MO), 14357 (herb. Trin.) (U); El Tucuche, 2500 ft, 28 Sep 1950, Baker in herb. Trin. 14643 (US); Mount Tocuche, 3-5 Apr 1920, Britton et al. 1339 (NY, US); Maracas waterfall, 10 Apr 1920, Britton et al. 1636 (NY); Maracas near waterfall, 29 Aug 1924, Broadway 5362 (MO, U); heights of Aripo, 1026 Jan 1922, Broadway 9847 (NY); Maracas falls, 11 Jun 1903, Johnston 40 (NY); Maracas waterfall, 100 ft, 24 Feb 1962, Kalloo B1107 (NY). TOBAGO:Castara, 30 Sep 1911, Broadway 4142 (NY); between Parlutivier and Bloody Bay, 14 Nov 1912, Broadway 4271 (NY, U, W); Caledonia, 11 Oct 1937, Sandwith 1731 (K, NY).

206

FLORA NEOTROPICA

Local names. Panama. Panama: nusasapi designateas the lectotype a collection thatI have seen, Merida: Eggers 5797. sailaleta(Kuna).Venezuela.Falc6n:barradera; canelo6n,mamey de la mueuy. Solanumarboreumis a misnomersince this species is usuallya small shrubandis often rhizomatous.I returnedto the type locality to re-collectthis plant,and found plants correspondingexactly to the type in the Humboldtand Bonplandherbarium,but all were rhizomatousshrublets.HumboldtandBonplandmay have madea mistakein transcribingtheirfield notes, or perhapsthis species occasionallydoes attaintreesize in the area.Near CuevaEl Guacharoin Monagas,Venezuela, all of the plants growing in a 3x3 m squareplot were interconnectedby rhizomes.Solanumarboreumis also rhizomatousat Finca La Selva, Costa Rica, and in the CanalArea,Panama.Inalltheseareastheplantsaregrowing in waterloggedor clay soils andthe habitis perhaps correlatedwith this environmentalcondition.The only otherspeciesof sect.GeminataI haveseenwiththishabit is S. robustifronsof easternPeru. Solanumarboreumis extremelyvariablein leaf shape, particularly in the shape of the leaf base. In northernSouthAmerica,the leaves arepetiolate,while in Costa Rica, they are nearly sessile, and the base is somewhatcordate.The charactertransitionis relatively abrupt and occurs in the vicinity of the Canal in Panama.Those collections with nearly sessile leaves often have long-acuminatecalyx lobes, but this character extends into populations with petiolate leaves. The nameS. enchylozumhas been used forthose plants with long-acuminate calyx lobes and sessile leaves. In conditions of severe root crowding (in small pots in the greenhouse) and water stress, individuals of S. arboreurm develop even more extremelysessile leaves and crowded internodes. The morphological differences thereforemay be at least partiallyenvironmentally controlled. Solanum arboreumis most likely related to S. ramonenseof high-elevationPanamaand Costa Rica, sharingwith that species papillate new growth, short inflorescenceswith denselypackedpedicel scars,green berries on erect pedicels, and tiny white flowers with reflexed petals. Both of these species are apparently andromonecious,withbothlong-styledandshort-styled flowers (mentionedby Schulz in his descriptionof S. dolichostylum). Few berries are borne on any given inflorescence, and these are usually nearthe base. I have chosen to lectotypify the name Solanum dolichostylum0. E. Schulz,a synonymofS. arboreum, with the the first of the collections cited by Schulz in the originaldescription,neitherof which arepresently extantat B. I have been unableto locate any duplicates of Hart 6779 from Trinidad,and prefer therefore to

55. Solanum falconense S. Knapp, Bull. Brit. Mus. (Nat. Hist.), Bot. 19: 103. 1989. Type. Venezuela. Falc6n: SierraSan Luis, along access rd. to Hotel Parador,nearCurimagua,1300m, 11 12N, 69?40'W, 29 Sep 1984,Knapp& Mallet6693 (holotype,MY; isotypes,BH, K, NY, US, VEN). Fig. 107 Shrubs, 1-2.5 m tall; young stems and leaves denselypubescentwith simple,uniseriatetrichomesca. 0.5 mm long, these drying reddish, curled at the tips; bark of older stems dark brown. Sympodial units difoliate, geminate. Leaves elliptic, widest at or just above the middle, glabrousandshiny adaxially,pubescent with uniseriatesimpletrichomesabaxially,the trichomes 0.5-1 mm long, with swollen bases, drying reddish,usuallyall orientedin a single direction;major leaves 4.7-12 x 1.8-4.5 cm, with 6-7 pairs of main lateralveins, these dryingreddishbeneath,the midrib prominentlykeeled above,the apex acuteto acuminate, thebaseattenuate;petiole0.3-1.2 cm long;minorleaves differing from the majors in both size and shape, orbicular,0.9-2.3 x 0.8-2 cm, the apex rounded,the base rounded;petiole 1-2 mm long. Inflorescences opposite the leaves, simple, 3-5 mm long, pubescent with simple uniseriate trichomes like those of the young stems andleaves, 4-10-flowered;pedicel scars closely spaced, not overlapping.Buds globose when young, laterellipsoidandstronglyexsertedfromthe calyx tube. Pedicels at anthesis deflexed (?), slender, 0.8-1 cm long, ca. 0.5 m diam. at the apex, ca. 0.25 mm diam. at the base, sparselypubescentwith simple uniseriatetrichomes like those of the stems.Flowers with the calyx tube conical, ca. 1 mm long, pubescent with simple uniseriatetrichomeslike those of the rest of the inflorescence, the lobes triangular,reflexed at anthesis(?), ca. 1 mm long, sparselypubescentwith simpleuniseriate trichomesca. 0.5 mm long;corollawhite,ca. 1 cm diam., lobed nearly to the base, the lobes planarat anthesis (?), sparsely pubescent on the abaxial surfaces with simple uniseriatetrichomes,papillose at the tips of the lobes; anthers 2.5-3 x ca. 1 mm, poricidal at the tips, the poresteardropshaped;free portionof thefilaments ca. 0.25 mm long, the filamenttube ca. 0.25 mm long; ovary glabrous;style straight,glabrous,4-5 mm long; stigmaa broadenedareaon the tip of the style, minutely papillose.Fruita globose berry,greenwhen immature, translucentyellow when ripe, 1-1.5 cm diam.;fruiting pedicels deflexed, woody, 1.5-1.8 cm long, ca. 2 mm diam.attheapex,ca. 1 mmdiam.atthebase.Seedsbright green in fresh material,darkbrown when dry,ovoid-

TAXONOMIC TREATMENT

207

FIG. 107. Solanum falconense S. Knapp. (Reproduced with permission from Bulletin of the British Museum, Natural History (Botany) 19: 104, fig. 1. 1989.)

reniform, imbedded in a firm, fleshy matrix, ca. 3 x ca. 2.5 mm, the surfaces minutely pitted. Chromosome number not known. Distribution (Fig. 108). Only known from the cloud forests of the Sierra San Luis in the state of Falc6n, Venezuela, at 1300-1400 m. Grows in open places in forest, but not in full sun.

along river below waterfall, between Hotel Parador and Curimagua, E of Hotel Parador, 1300 m, 23 July 1967, Steyermark 99457 (NY, US, VEN). Solanumfalconense is closely related to S. lucens

with which it shares shiny, obovate majorleaves with keeledmidribsthatdryrussetbrown.Solanumfalconense differsfromS. lucens in its pubescentleaves andtranslucentyellow berryon a deflexed fruitingpedicel. Specimens examined. VENEZUELA. FALCON: Thebrightgreenseeds in translucentyellow fruits Sierra San Luis, above Sta. Maria, 1000 m, 1 Feb 1979, in sect. Geminataandareprobablyrelated are unusual Flora Falcon (HW, TR, BV, ES) 283 (U); Sierra San to dispersal mechanism. Most species of sect. Luis, ridges around Hotel Parador, ca. 7 km E of Geminatawith greenfruitaredispersedby batsor small Curimagua, 1300-1350 m, 11?10'N, 69?35'W, 28 September 1984, Knapp & Mallet 6676 (BH, K, MY, VEN, rodentsbut those with brightly colored fruit are birdUS), Knapp & Mallet 6683 (BH, K, MY, VEN, US), dispersed. Solanumfalconense may also be bird disKnapp & Mallet 6686 (BH, K, MY, NY, VEN, US); persed,as it possesses relativelybrightlycoloredfruits. near Hotel Parador, 7 km S of Curimagua, La Tablaconfined to the isois Curimagua rd., Sierra San Luis, 1400 m, 29 April 1978, Solanumfalconense apparently Morillo et al. 7223 (VEN); Sierra San Luis, Montana latedcloud forests of the SierraSan Luis in northwestem Venezuela.This is one of the centers of endemism de Parguila, ca. 1300 m, 18 Feb 1980, Sobel et al. 2039 (BH, NY); Sierra San Luis, Montafia de Paraguariba, in Venezueladetailedby Steyermark(1979).

FLORA NEOTROPICA

208

700

.0

00

0.

closely spaced,not overlapping.Budselongateandfalcate when very small, largernot known.Flowers with the calyx tube conical, very fleshy, 2-3 mm long, glabrous,the lobes irregularlydeltate,2-3 mm long, glabrous;corolla white to pinkish-white,fleshy, 1-1.8 cm diam., lobed nearlyto the base, the lobes planarat anthesis, densely papillose along the margins,papillose and cucullate at the tips; anthers 4.5-5 x ca. 1 mm, poricidal at the tips, the pores teardropshaped; free portionof thefilamenitsca. 0.05 mm, the filamenttube ca. 1 mm, glabrous;ovary glabrous;style straight5-6 mm; stigma capitate, the surface minutely papillose, dryingdarkerthanthe style.Fruita globose,greenberry, 1-1.2 cm diam.;fruitingpedicels woody, deflexed, 22.2 cm long, ca. 2 mm diam.at the base. Seeds pale tan, ovoid-reniform,3-4 x 2.5-3 mm, the surfacesminutely pitted. Chromosomenumber-not known. Distribution (Fig. 108). In rainforestin N Peru and S Ecuador,from 2000-3000 m.

FIG. 108. Distribution of Solanurm anisoph v/lum (solid circles), S. Jalconense (solid squares), S. goniocaiulont (open circles),

and S. graturm (stars).

56. Solanum goniocaulon S. Knapp,Novon 2: 343. 1992. Type.Peru.San Martin:Rio Abiseo National Park,hill E of GranPajatenruins,2350 m, ca. 7S, 77?W, 13 Aug 1986, Young 4213 (holotype, HUT; Fig. 109 isotypes, K, NY).

Specimens examined. ECUADOR. LOJA:Cerro de Celica, Celica-Guachanama, km 14-18, 2700 m, 4003'24"S, 79053'41"W, 13 Apr 1994, Jorgen.sen et al. 159 (BM, MO); Cerro de Villonaco, Loja-La Toma, km 13, turnoff toward Chuquiribamba, km 2, Hacienda La Huangora,2640 m, 3056'52"S, 79?15'52"W, 18 Apr 1994, Jorgensen et al. 332 (BM, MO). ZAMORA-CHINCHIPE: Parque Nacional Podocarpus, near pass on Loja-Zamora rd., mule track toward Zamora, 2800-3000 m, 3?59'S, 79007'W, 8 Mar 1990, Madsen 86967 (AAU). PERU.CAJAMARCA: Jaen, Rio Agua Blancanear Rio Jeronga, 8000 ft, 30 Jul 1943, Evinger 461 (US).

Solanumgoniocaulon is a species of uncertain relationships in the S. arboreum species group. The flowers of S. goniocaulon, like the buds (on Evinger 461), are similar to those of S. anisophyllum,another Trees, 7 m tall;young stems and leaves minutely memberof theS. arbor-eum species groupfromthe eastpubescentwith golden uniseriatetrichomesca. 0.5 mm em Andeanslopein PeruandEcuador.Thestronglyfourlong; stems distinctly four-angled,erect;barkof older winged stem andthe distinctiveyellow color of the unstems yellow-gray,glabrate.Sympodial units difoliate, dersides of dried leaves make S. goniocaulon readily geminate.Leaves elliptic, thick and leathery,glabrous differentiatedfromany otherspecies of sect. Geminata. and shiny adaxially, drying dark green, glabrous or Solanumgoniocaulon is superficiallysimilarto pubescentwith uniseriategolden trichomesca. 0.5 mm S. laurifronsof the S. amblophyllumspecies group in long along the midriband main lateralveins abaxially, its angled stems and leaves drying golden yellow bethe undersidesdryingdarkgold; majorleaves 15-28 x neath, but differs from that species in its ovoid-reni5-8 cm, with 8-10 pairs of main lateral veins, these form seeds and difoliate, geminate sympodial units. obscure abaxially, the apex acute, the base attenuate, somewhat winged onto the petiole; petioles 1-1.5 cm long; minor leaves differing from the majors only in 57. Solanum gratum Bitter,Repert.Spec. Nov. Regni. Veg. 18: 59. 1922. Type. Venezuela. Los Andes size, 3-10 x 1.5-5 cm, the apex acute, the base attenu1701(lectotype,BM, (Aragua):ColoniaTovar,Moritz 0.7-1 onto the petiole; petiole ate, somewhat winged heredesignated;isolectotypes,B [destroyed],K). cm long. Inflorescences oppositethe leaves, simple, 13 cm long, sparselypuberulentat the tips, 20-30-flowSolanum avilense Pittier, Cat. Fl. Venez. 382. 1947. nom. nud. Type. Venezuela. Distrito Federal: ered;pedicel scars beginningca. 1/2way fromthe base,

TAXONOMIC TREATMENT

209 .

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332 Um (MO ) OCNE Saciwty

i

210

FLORA NEOTROPICA Cerca de San Isidro de Galipan, Tamayo 336 (F, US, VEN). Figs. 102A, 110

Shrubs or small trees, 2-8 m tall; young stems and leaves glabrous or occasionally minutely papillose; bark of older stems greenish-yellow and shining. Sympodial units difoliate, usually geminate. Leaves elliptic or obovate, widest at orjust distal to the middle, coriaceous, glabrous, and shining; major leaves 7.5-12.5 x 2.8-6.5 cm, with 6-8 pairs of main lateral veins, these prominent beneath, impressed above, the apex rounded or acute, the base acute; petioles 1.1-1.2 cm long; minor leaves 2.1-4.2 x 1.1-2.5 cm, the apex rounded, the base acute; petioles 2-5 mm long. Inflorescences opposite the leaves, simple, glabrous, 0.1-1.7 cm long, 10-30-flowered; pedicel scars closely spaced, overlapping, beginning ca. 1 mm from the base of the inflorescence. Buds glabrous, globose when young, becoming ovoid with age, the corolla exserted from the calyx. Pedicels at anthesis 1.5-2 cm long, deflexed, abruptly narrowing below the calyx tube, then tapering to the slender base ca. 0.5 mm diam. Flowers sweetly fragrant, with the calyx tube ca. 1.5 mm long, campanulate, the lobes deltoid, 1-1.5 mm long, thick-margined, minutely papillose on the tips of the lobes; corolla white, 1.2--I.8 cm diam., lobed 3/4 of the way to the base, the lobes planar or slightly campanulate at anthesis, minutely papillose on the margins and tips of the lobes; anthers ca. 3.5 x 1-1.5 mm, the terminal portion ca. 0.5 mm long, paler and thickened, poricidal at the tips, the pores teardrop shaped; free portion of thefilaments ca. 1.5 mm long, the filament tube ca. 0.5 mm long; ovary glabrous; style 6-7 mm long, straight, clavate; stigma a minute papillose area on the tip of the style. Berries globose, green, 1-1.5 cm diam.; fruiting pedicels woody, erect or deflexed from the weight of the berry, 1.8-3.4 mm long, 0.75 mm diam. at the base. Seeds dark brown in dry material, ovoid-reniform, ca. 3 x 2 mm, the surfaces minutely reticulate. Chromosome number. n = 12 (voucher Knapp & Mallet 6872). Distribution (Fig. 108). Found only in the Cordillera de la Costa in Venezuela from 1600-2000 m, often in paramo. Most collections are from Parque Nacional El Avila, Caracas. Specimens examined. VENEZUELA. ARAGUA: Betwecn Portachueloand La Regresiva del Diablo, Parque Nacional, 1000-1 100 m, 13 Dec 1947, Pittier15660(US, VEN); Parque Nacional, 1947, Pittier & Nakichenovich 15828 (VEN). DISTRITo FEDERAL: Near Galipan, ca. 2000 m, Jan 1958, Aristeguieta 2992 (MY, VEN); Serranias del Avila, ca. 1800 m, May 1958, Aristeguieta 3133 (VEN); Parque El Avila, S slope, 1800 m, 28 Apr 1979, Benitez de Rojas 2507 (MY); near Colonia Tovar, 18541855, Fendler 978 (MO, NY, P); Serranias del Avila, 29

Dec 1943, Lasser 1001 (US, VEN); Caracas, Jul 1842, Linden 42 (G (Morton neg. 8589)); Parque Nacional El Avila, airplane crash site, 3 Sep 1975, Manara s.n. (MO, VEN); Cordillera de la Costa, Los Venados-Boca de Tigre-Teleferico-Galipan, 19 Mar 1971, Mon/illo et a!. 813 (MY, VEN); Boca de Tigre, (Cerros de Avila near Caracas), 1850 m, 26-29 Dec 1918, Pittier 8373 (BH. F, US, VEN); Colonia Tovar and vic., 1700-2300 m, 31 Dec 1921, Pittier 10063 (B [destroyed:F neg. 2701], NY, US, VEN); Avila near Caracas, 2500 m, 10-17 Jan 1958, Smith& Aristeguieta3474 (F, NY, US); Cordillerade Avila, above Caracas, between Los Venados and Guayabo Mocho, 1675-2075 m, 28 Dec 1943, Stevermark 55-56 (VEN); near La Chivera on La Fila de Avila, Cerro Avila, 2040 m, 11 Dec 1976, Steyermark & Huber- 112785 (VEN); San Isidro de Galipan, 25 Dec 1937, Tanmvl,o 336 (F, US, VEN).

Solanumgratumis related to S.plowmaniiof the westernAndeanslopes in Peru.It shareswith thatspecies thick, fleshy leaves thatdry golden brown, fleshy flowers thatapparentlyopen synchronously,andfruiting pedicels deflexed fromthe weight of the berry.The barkof both species is pale and shiny.The leaves ofS. gratum are distinctive and make this species easy to recognizeon herbariumsheets.They arethickandrelatively coriaceous, and dry a rich golden brown. The undersidesdry with a golden tinge. The thick leaves that dry a golden brown are similar to those of S. goniocaulon, but the pale shiny barkand non-angled stems make S. gratum easy to distinguish from that

species. The plate of S. gratum in the Flora del Avila (Steyermark & Huber, 1978) shows many flowers open

synchronously.When in flower S. gratum must be a conspicuous plant. It is not common, even in Parque Nacional El Avila, where it has been collected most often. Most collections are of flowering plants.

58. Solanum laevigatum Dunal,Solan. Syn. 20. 1816. Type. Colombia. S.loc., (crescit juxta pagum Fusagusuga et propterponten Icononensium fide HBK 1819), Sep, Humboldt& Bonplands.n. (holotype,P-Bonpl.[F neg. 38998]; isotype,P [Morton neg. 8226]). Fig. 111 Solanuin obovatum Dunal, Solan. Syn. 21. 1816. Type. Ecuador. Azuay: Cuenca, Hmlniboldt& Bonplanid s.n. (holotype, P-Bonpl.; isotypes, P [Morton neg. 8270], frag. F). Solanum novo-granatense Dunal in DC., Prodr. 13(1): 680. 1852. Type. Colombia. Cundinamarca: Fusagasuga, Linden 834 (holotype, G-DC [F neg. 6780]; isotype, frag. F). Shrubs or small trees, 1-3 m tall; young stems

and leaves felty with dense brownisharachnoidpubes-

TREATMENT

TAXONOMIC

21

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cence, soon glabrate;stems erect and lightly winged; barkof older stems gray-brown.Sympodialunits unifoliate, occasionally difoliateandgeminate.Leaveselliptic to obovate, widest at or just above the middle, somewhatthick and fleshy, glabrousadaxially,pubescent abaxiallywith tuftsof uniseriate,simple,occasionally dendritictrichomesin the axils of the main lateral veins; majorleaves with 5-9 pairsof mainlateralveins, 6-15 x 2-7 cm, the apex acute to acuminate,the base acuteto attenuate;petioles0.5-1 cm long, lightlywinged from the decurrentleaf base; minor leaves if present differingfromthemajorsin size andoccasionallyshape, 3-4 x 1.5-2 cm, the apex acute,the base acute;petioles ca. 0.5 cm long. Inflorescences opposite the leaves, simple, congested, 0.2-1 cm long, 3-15-flowered, glabrousto sparselypubescentwith arachnoidtrichomes; pedicel scars closely spaced,nearlyoverlapping.Buds globose, laterellipsoid to obovoid, the corolla strongly exserted from the calyx tube. Pedicels at anthesis tapering, deflexed, 0.8-1.1 cm long, ca. 0.25 mm diam. at the base, ca. 0.5 mm diam. at the apex, sparselypubescent with arachnoidtrichomes. Flowers with the calyx tube broadly conical, 1-1.5 mm long, the lobes deltate,0.5-1 mm long, sparselyto densely arachnoid-

in P-Bonpl.

B. Colombia.

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Schlim 541 (K).

pubescent;corolla white, 1-1.2 cm diam.,lobed nearly to the base, the lobes planarat anthesis, the tips of the lobes minutelypapilloseandstronglycucullate;anthers 3-3.5 x 1-1.5 mm, poricidalat the tips, the pores teardrop shaped;free portionof thefilaments ca. 0.5 mm long, the filamenttube ca. 1 mm long, glabrous;ovary glabrousor arachnoid-pubescent;style in long-styled flowers 6-6.5 mm long, in short-styled flowers ca. 4 mm long, glabrous;stigma a minutely papillose area on the tip of the style. Fruit a globose, green berry,11.5 cm diam.;fruitingpedicelserect,woody, 1-2.4 cm long, ca. 2 mm diam. at the base. Seeds darkbrown, ovoid-reniform, 3-3.5 x 2-2.5 mm, the surfaces minutely pitted. Chromosomenumbernot known. Distribution (Fig. 112). In cloud forests and forestedges in the ColombianAndes (CordilleraOccidental and Central),950-2800 m. Specimensexamined.COLOMBIA.Ocania,2000 ft, May 1846-1852, Schlim 541 (BM). BOYACA: Raquira, 2250 m, 4 Jun 1989, Castroviejo et al. 10631 (MO). CALDAS: Manizales, rd. to Villa Maria, 2050 m, 17 May 1985, Escobar Cardona 132 (NY); Cordillera

Central,Montele6n,2250 m, 16 Feb 1985, Melida de Fraume& Alvarezy Gallego352 (NY). CAUCA: Cordi-

TAXONOMIC TREATMENT

213

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Roblal, L bank of QuebradaEl Tigre, 2100 m, 6 Apr 1988, Heredia 530 (NY); old Cali-Buenaventura rd., between km 14 and 18, 1900 m, 21 Oct 1987, Ramos 803 (NY); Hacienda El Medio, Carretera Panamericana between Paila and Zarzal, valley of Rio Cauca, ca. 950 m, 21 Dec 1987, Silverstone-Sopkin et al. 3445 (NY). PERU. AMAZONAS: Bagua, Third camp, Cordillera Colan SE of La Peca, 2000-2500 m, 27 Sep 1978, Barbour 3595 (MO). Solanum laevigatum is superficially similar to S. cornifolium (S. nigricans species group), also of the

00

FIG. 112. Distribution of Solanlum laevigatum (solid circles), S. lucens (open circles), S. plowmanii (solid squares), and S. r-amoniense(stars).

ColombianAndes. It differs from it in its barbatevein axilsof theabaxialleaf surfaceandin its generallylowerelevation distribution.The trichomes are a somewhat darkerbrownandnot quiteso dense. Both species have polymorphicglabrousand pubescentovaries, as does anothersympatricspecies, S. callianthum. I have seen no recent collections of Solanum laevigatumfromEcuador,althoughthe type collection of S. obovatumis apparentlyfromsouthernEcuadornear Cuenca(prov.Azuay).Barbour3595, fromAmazonas Peru, is more densely and loosely pubescentthan Colombianmaterial.As more materialfrom Ecuadorand Perubecomes available,these plants (i.e., from Ecuador and Peru) may be shown to be distinct from S. laevigatum s.str.

59. Solanum lucens S. Knapp,Brittonia38: 280. 1986. Type. Venezuela. Apure: Reserva Forestal San llera Central, W slope, headwaters of Rio Palo, Quebrada Camilo, selva siempreverde a lo largo de la de Santo Domingo, R bank, 2700-2800 m, I1 Dec 1944, QuebradaBotina,unos 2 km al suroestedel Caserio Cuatrecasas 19149 (F). CUNDINAMARCA: Barroblanco, San Camilo (El Nula), 250 m, 28 Mar 1968, 2200 m, 4 Feb 1876, Andre 1426 (F, K); Cordillera Oriental, W slope between El Salto and El Colegio, 1680 m, 10 Mar 1940, Cuatr-ecasas 8234-A (F), Cuatrecasas 8238 (US); Fusagasuga, hacienda La Negrita, 1680 m, 31 Dec 1968, Cuatrecasas et al. 26959 (US); Sasaima, Vereda San Bernardo along Rio Dulce, 1660 m, 10 Jan 1946, Garcia Barriga 11895 (US); vic. Fusagasuga, 1780-1850 m, 15 Apr 1946, Garcia Barriga 11956 (US); below Tequedama, rd. to El Colegio near La Herradura, 1900 m, 28 Jul 1962, Uribe Uribe 4068 (US); Mun. Zipac6n, Lagunaseca, 1800 m, 1 Aug 1964, Ur-ibeUribe 4896 (US); Viota, toward Liberia, 1400 m, 30 Dec 1967, Uribe Uribe 6035 (US); Fusagasuga, ca. 1800 m, 10 Nov 1055, Vogel 51 (US). NARINo: Near La Cruz de Pasto, 2000-2500 m, Lehmann 4951 (B [destroyed: F neg., 2680], US). PASTO: Popayan, 1500 m, Jun 1853, Triana 16, 17 (BM). VALLE DE CAUCA: Cordillera Central, W slope, Rio Bugalagrande, Loma de Barragan,Hacienda de San Jose, 2600-2850 m, 15 Mar 1946, Cuiatrecasas 20023 (F, US); Cordillera Occidental, W slope, Finca Torremolinos, km 22 of Cali-Buenaventura rd., 1800 m, 13 Oct 1982, Escobar- et al. 2685 (NY); Hacienda El Entamborado, R bank Quebrada El Tigre, 2200 m, 20 Oct 1987, Heredia 475 (NY); Hacienda El

Steyermark et al. 101421 (holotype, VEN; isotypes,

IBE, US).

Figs. 102B, 113

Shrubs or small trees, 1-4 m tall; young stems andleavesdenselyred-floccose,thetrichomesweakand appressedto the stems, ca. 0.1 mm long, soon deciduous; stems lightlywinged fromthe decurrentleaf bases; barkof older stems deep grayish-red,shiny; branches withthe leavesheld in a singleplane,giving eachbranch a flattenedappearance.Sympodialunitsdifoliate,geminate.Leaveselliptic to obovate,widest at the middleor in the distal third of the blade, glabrous on both surfaces,veryshinyandwiththemidribprominentlykeeled above; majorleaves 8.5-16 x 3-5 cm, with 6-8 pairs of mainlateralveins, these somewhatprominentabove in live plants,notat all prominentin drymaterial,prominent andreddishbeneath,the apex acuteto acuminate, the base cuneateto attenuate,decurrenton the petiole; petioles 0.5-1 cm long, lightly winged; minor leaves differingfromthe majorones in size andusuallyshape, elliptic to rounded,2-4 x 1-2 cm, the apex acute, the

214

FLORA NEOTROPICA

FIG.

113. Solanum lucens S. Knapp. (Reproduced with permission from Brittonia 38: 282, fig. 7. 1986')

FIG. 113. Solanumlucens S. Knapp. (Reproduced with permission from Brittonia 38: 282, fig. 7. 1986.)

base cuneate; petioles 2-3 mm long. Inflorescences opposite the leaves, simple, 2-5 cm long, 3-8-flowered, densely reddish-floccose;pedicel scars closely spaced, often overlapping,beginningat the base of the inflorescence.Buds globose, the calyx appearinghyaline, the calyx completely enclosing the corolla until quite late, after corolla exsertion the buds ellipsoid. Pedicels at anthesis deflexed, 0.9-1.3 cm long, tapering from the calyx tube to a slenderbase 0.25-0.5 mm diam.Flowerswith a fragrancereminiscentof citronella et al. 761-25), the calyx tubebroadlyconi(fide Yillett cal, 1-1.5 mm long, the lobes deltoid, 1-1.5 mm long, glabrous or red floccose, the trichomes often deciduous; corolla white, 1.1-1.3 cm diam., lobed nearly to the base, the lobes planaror slightly reflexed at anthesis, the tips and margins of the lobes minutely papillose; anthers 2.5-3 x 1 mm, poricidal at the tips, the poresteardropshaped;freeportionof thefilaments0.51 mm long, the filament tube 0.5-1 mm long; ovary glabrous;styles straight,in short-styledflowers ca. 1.5

mm long, in long-styled flowers 4-6 mm long;stigma a minutely papillose area on the broadenedtip of the style.Fruit a globose, pale greenberry,0.8-1 cm diam., usually 1-3 berriesper inflorescence;fruitingpedicels erect andwoody, corky and large-lenticellate,1.5-1.7 cm long, often appearinglonger if the fruit is from a terminalflower, 0.75-1 mm diam. at the base. Seeds dark brown in dry material, ovoid-reniform,2-2.5 x 1.5-2 mm, the surfacesminutelypitted. Chromosome numbernot known. Distribution (Fig. 112). In secondary growth of dry forest areas in western Venezuela and eastern Colombia, on the Amazonianslopes of the Andes and in the Tichira depression, from 500-1200 m. Specimens examined. COLOMBIA. NORTE DE SANTANDER: Cordillera El Banco, Oriental, regionof Sanar6, confluenceof Rio Cubug6nandRio Cobaria,320 m, 15 Nov 1941, Cuatrecasas13161 (F, US). VENEZUELA. BARINAS:Ticoporo Forest Reserve, banksof Rio Bumbun,350 m, 8?15'N,70?45'W,

TAXONOMIC TREATMENT 11 Mar 1964, Breteler 3664 (K, US, VEN), 15 Jul 1964, Breteler 4028 (K, MO, NY, US, VEN); near Barinitas, 500 m, 13 Jan 1965, Breteler 4410 (VEN); Reserva Forestal Caparo, Unidad 1, SE of Campamiento Cachicana, E of El Cant6n, 100 m, 9 Apr 1968, Steyermark et al. 101969 (US, VEN); MERIDA: 0.5 km SW of dam site on Rio Caparo, 31 km ESE of Santa Barbara, 100-200 m, 7?41'N, 71?28'W, 9 Mar 1980, Liesner & Gonzalez 9222 (VEN); Tucupido, Rio Camburito to Santa Maria de Caparo, 4 Aug 1973, Lopez Palacios & Bautista B. 3219 (MO). TACHIRA: Ca. 10 km upstream on Rio Frio from confluence with Rio Quinimari, along rd. to El Porvenir and Santa Ana, ca. 600 m, 7?36'N, 72?13'W, 24 Oct 1984, Knapp & Mallet 6830 (BH, K, MY, US, VEN); on Las Dantas-Buena Vista-Puente Alianza rd., ca. 12 km S of Las Dantas (ca. 7 km S of the guardia

post at Buena Vista), 1100-1200 m, 7?40'N, 72?25'W, 24 Oct 1984, Knapp& Mallet 6832, 6834 (BH, K, MY, US, VEN); along Rio QuebradaGrande,at end of rd. past puebloQuebradaGrande(tributaryof Rio Burgua), ca. 375 m, 15 Jan 1976, Tillettet al. 761-25 (MY). Solanum lucens is probably related to S. falconense fromthe SierraSanLuisin thestateof Falc6n, Venezuela.Characterssharedbetween the two species are discussed with the latter.Solanumlucens is easily distinguishedfromS. falconense by its glabrousleaves andpale green berriesborneon erectpedicels. The fragranceof the flowers of S. lucens (citronellafide Tillett et al. 761-25) is unusualin the section. The flowers of S. lucens seemnto open synchronously,with an entire branch or plant being in bud, flower, or fruit at the same

time,a charactersharedwithS. gratumandS.plowmanii.

60. Solanum plowmanii S. Knapp, Bull. Brit. Mus. (Nat. Hist.), Bot. 19: 104. 1989. Type. Peru. Lambayeque:Abrade Porculla,rd. from Olmos to Pucara,km 45 E of Olmos, 1920 m, 13 July 1986, Plowman et al. 14280 (holotype, F; isotypes HUT Fig. 1 14 n.v., K, MO, NY).

Shrubsor small treelets,2-3 m tall, young stems and leaves completely glabrous,dryingdark,stoutand erect; older stems glabrous, the bark pale and shiny. Syvmpodialunits difoliate, usually geminate. Leaves elliptic,widest at the middle,completelyglabrousboth adaxiallyandabaxially,slightlyfleshy;majorleaves 1014.5 x 4-6.5 cm, with 10-12 pairsof mainlateralveins, these drying reddish beneath, the apex acute, the base acute; petioles 1-1.5 cm long; minor leaves differing

from the majors only in size, 2-5.5 x 0.8-2.2 cm, the apex acute,the base acute;petioles 3-5 mm long.Inflorescences opposite the leaves or intemodal,simple, 15 cm long, 5-8-flowered,glabrous;pedicelscars closely spaced, not overlapping, clustered in the distal 1/2 to

1/4 of the inflorescence.Buds globose, laterellipsoid,

215

the corollaexsertedfromthe calyx tube.Pedicels at anthesis deflexed, 1.1-1.4 cm long, fleshy, taperingfrom the calyx tube to a basal diam. of ca. 1 mm. Flowers with the calyx tube conical, 1.5-2 mm long, glabrous, the lobes deltate,the marginsthickened,ca. 1 mm long, glabrous;corolla white, fleshy, 1.7-2 cm diam., lobed 3/4 of the way to the base, the lobes planarat anthesis, the tips of the lobes minutelypapillose;anthers4.5-5 x 1-1.5 mm, poricidal at the tips, the pores teardrop shaped; free portion of thefilaments ca. 0.5 mm, the filament tube ca. 0.5 mm long; ovary glabrous;style erect, 6-7 mm long; stigma slightly clavate, minutely papillose.Fruit a globose, greento yellow-greenberry, 1-1.5 cm diam.;fruitingpedicelswoody anddeflexed, ca. 1 mm diam. at the base. Seeds darkbrown in dry material,tan in freshmaterial,ovoid-reniform,4-4.5 x 3-3.5 mm, the surfacesminutelypitted. Chromosome numbernot known. Distribution (Fig. 112). On the western slopes of the Andes in S Ecuadorand N Peru, in the general areaof the Huancabambadepression,remnantsof dry or moist forest from 1500-3200 m. Specimens examined. ECUADOR. LOJA: Sozoranga, Reserva Natural El Tundo, property of

Fundaci6nArcolris,km 6 along trackfrom SozorangaMacara rd., 1800 m, ca. 4?19'S, 79049'W, 19 Aug 1997, Lewis et al. 3488 (BM, K, LOJA-n.v., QCNE-n.v.). PERU. AMAZONAS: Near Aramango, above village of Muyo, ca. 400 m, 5?28'S, 77?28'W, 8 Jul 1984, Knapp & Mallet 6579 (BH, K, MO, NY, US, USM). CAJAMARCA: La Palma, 10 km NW of Chirinos, 1780 m, 5?25'S, 78?53'W, 4 Feb 1988, Gentri' et al. 61146 (MO, NY); Rio Agua Blanca on Jeronga River, 8000 ft, 30 Jul 1943, Evinger 455 (NY, US); Prov. Santa Cruz, Bosque Monteseco, 1880 m, 20 Jan 1996, Leiva et al. 1746 (BM, F). LAMBAYEQUE: Chinche, Olmos, 1600 m, 2 May 1981, Llatas & Lao 640 (HUT, MO). PIURA: Huancabamba, Canchaque, "Chorro Blanco," 1600 m, 18 Jan 1988, Diaz et al. 2772 (MO, NY); ca. 2 km E of Canchaque en route to Huancabamba, ca. 1450 mn. 5?24'S, 79?36'W, 21 Jul 1991, Dillon & Scinchez V 6317 (BM, F); km 46 Abra Porculla, 1900-2000 m. 28 Jun 1959, Ferreyra 13758 (MO); descending from Ayabaca to Puente de Arraipite, 2000 m, Ochoa 1794 (MOL). Tumbes: Descending Paso del Indio toward Camchaque, 3200 rn, s.d., Ochoa 1788 (US).

Solanumplowmanii is related to S. gratum of the VenezuelanCordillerade la Costa.The two species share fleshy leaves that dry a rich golden brown and fleshy flowersthatapparentlyopen synchronously(see Figs. 102A, 110). The leaf apices of both aregenerally rounded,an unusualcondition in sect. Geminata.The flowers of S. gratum are reportedto be sweetly fragrantbut no such informationexists forS. plowmanii. The isolated forests of the western slopes of the Andes in which Solanumplowmaniioccurs are a very

216

FLORA NEOTROPICA

MT

FIG. 114. Solanutm plowvnanii

S. Knapp. (Reproduced

with permission

from Bulletin of the British Museum,

Natural History (Botany) 19: 106, fig. 3. 1989.)

limited habitatunderthreatof deforestationat present (Sagastegui A. & Dillon, 1991; Dillon, 1993).

61. Solanum ramonense C.V. Morton & Standl., Publ. Field Mus. Nat. Hist., Bot. ser. 18: 1090. 1938. Type. Costa Rica. San Jose: San Rafael de San Ram6n, Brenes 22013 (holotype, F; isotypes, NY). Fig. 115 Solanini lepidocaule Standl. & L. 0. Williams, Ceiba 3: 218. 1952. Type. Costa Rica. Alajuela: San Luis de Zarcero, in cloud forest, 1550 m, I

Jun 1938, Smith NY725 (holotype, F; isotype,

NY, photos US, WIS). Shrubs 1-2 m tall; young stems and leaves glabrous and occasionally minutely papillose, soon glabrous;barkof the olderstemspale gray,lenticellate;the stemsslightlyswollenatthenodes;branchesgentlyarching.Sympodialunitsdifoliate,geminate.Leaveselliptic, glabrous,darkgreen and shiningabove, palerbeneath, widest at the middle;majorleaves 9-15 x 4.2-6.5 cm, with 8-10 pairsof main lateralveins, impressedabove, prominentandpalerbelow,the apexacuminate,thebase attenuate;petioles0.9-1 cm long;minorleavesdiffering

217

TREATMENT

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FLORA NEOTROPICA

218

from the majorones in size and occasionally in shape, usually rounded,1.2-5 x 0.9-3.1 cm, the apex acuteor rounded,the base attenuate;petioles 2-7 mm long.Inflorescencesoppositethe leaves,simple,0.2-0.8(-2) cm long, 5-10-flowered, glabrousor minutelypapilloseon the extremetip;pedicel scars closely spacedandoverlapping,beginningatthebase of the inflorescence.Buds globose when very young, laterbecomingellipsoid,the calyx lobes long-acuminatein bud, minutelypapillose andoccasionallywitha few uniseriatetrichomesca. 0.05 mm long. Pedicels at anthesis 3-5 mm long, deflexed, tapenngfromthecalyx tubeto a slenderbaseca. 0.5 mm diam.Flowers with the calyx tube 0.5-1 mm long, the lobes long-triangular,deflexed, white, 0.5-1 mm long, glabrousat anthesis;corollawhite,5-6 mm diam.,lobed nearlyto thebase, the lobes reflexedat anthesis,the tips andmarginsof the lobesminutelypapillose;anthers1.52 x 0.75-1 mm, poricidalat the tips, the poresteardrop shaped;freeportionof thefilaments0.25-0.5 mm long, the filamenttubeca. 0.25 mm long;ovaryglabrous;style straight,3-3.5 mm long;stigmacapitate,minutelypapillose. Fruit a globose, green berry,1.4-1.6 cm diam., smellingstronglyof wintergreen(methylsalicylate)when ripe;.fruitingpedicelserectandwoody, ca. 1.2 cm long, ca. 1.5 mm diam. at the base. Seeds very few per fruit, large, pale tan, ovoid-reniform,3-5 x 1-2.5 mm, the surfacesminutelypitted.Chromosomenumber:n = 12 (voucherKnapp4270). Distribution (Fig. 112). In primary forest in montane Costa Rica and westem Panamafrom 6002000 m. Selected specimens examined. COSTA RICA. ALAJUELA:

ParqueNacional Rinc6n de la Vieja, La

Siembra trail 6 km above Admin. toward Volcan Sta. Maria, 700 m, 10?47'50"N, 85?18'19"W, 7 Sep 1991, River-a1610 (INB); Tapuco de Zarcero, 1575 m, 15 Jun 1938, Smith NY781 (NY); Alfaro Ruiz, Llano Bonito de Zarcero, 1750 m, 2 Jul 1938, Smith NY821 (NY); Alfaro Ruiz, Tapuco de Zarcero, 1550 m, 26 Oct 1938, Smith NY1288 (NY); San Carlos, San Juan de Lajas, 1375 m, 16 Jan 1939, Smith NY1491 (NY); San Carlos, Pueblo Nuevo, 1100 m, 15 Apr 1939, Smith NY1905 (F, NY, US). CARTAGO: El Mufieco on Rio Navarro, 1400-1500 in, 6-7 Mar 1926, Standley & Valerio 50894 (US). GIJANACASTE:

P.N. Guanacaste,Est. Cacao, 1100 m,

10055'N, 85028'15"W, 10 Dec 1990, Chaivez455 (INB); Los Ayotes near Tilartn, 600-700 m, 21 Jan 1926,

1540-1600 m, 26 Jan 1977, Dryer 1158B (F); 2 km SE of Monteverde, 1500-1550 m, 10?18'N, 84?48'W, 1821 Mar 1973, Gentry & Burger 2681 (F, MO); 0.5 km from station on Sendero Rio, 1500 m, 10?25'N, 84?50'W, 19 Feb 1981, Knapp 829 (BH, CR); along Sendero Rio near Rio Guacimal, 1500 m, 10?25'N, 84?50'W, 30 Apr 1981, Knapp 875 (BH, CR); PI La Amistad, Cord. de Talamanca,Est. Pittier, 1900 m, 9?01'50"N, 82?57'30"W, 1 Aug 1995, Moraga 264 (INB). SAN JoSE: TerrabaSierpe drainage, Est. Sta. Elena, El Llano trail, 1900 m, 9?23'33"N, 83037'19"w, 20 Mar 1997, Alfaro et al. 1109 (INB); Cerros de Zurqui, S facing slope just below main ridge, N of village of San Luis Norte, 17 km NNE of city center of San Jose, 1750 m, 10?03'N, 84001'W, 23 Apr 1975, Crosby 11439 (CR, F, MO); hills around Rio Savegre, ner Finca Zacatales, 2000-2400 m, )'335'N, 83?47'W, 1 Aug 1991, Gaiy et al. 1563 (INB). PANAMA. BOCASDEL TORO: Between Buena Vista coffee finca and Cerro Pil6n on Chiriqui trail, 17 Apr 1968, Kirkbride & Duke 705 (MO); between Qda. Gutierrez and E slope of La Zorra, headwaters of Rio Mali on Chiriqui trail, 18 Apr 1968, Kir-kbride& Dlke 722 (MO). CHIRIQUi: Along rd. between Gualaca and Fortuna dam site, 5.9 mi NW of Los Planes de Hornito, 1370 m, 9 Apr 1980, Antonio 4124 (MO); between Palo Alto and Top of Divide near Cerro Pate Macho, above Rio Palo Alto NE of Boquete, 5400-7100 ft, 18 Mar 1979, DArcy et al. 12644 (MO); along rd. to Fortuna dam site, 22.7 mi beyond the bridge over Rio Esti, 11.8 mi N of Los Planes de Hornito, 10.7 mi N of tunnel jct., 1400 m, 26 Nov 1979, Croat 48665 (BH, MO); along rd. to Fortuna dam site on Rio Chiriqui, 7.7 mi beyond Francisco Linares' lane, 19.2 mi beyond bridge over Rio Esti, 9.1 mi beyond Los Planes de Hornito, 1300 m, 27 Nov 1979, Croat 48740 (BH, MO); along rd. from Gualaca to Fortuna dam site, 5.9 mi NW of Los Planes de Hornito, 1370 m, 8?43'N, 82?17'W,9 Apr 1980, Croat 49879 (MO); 3.5 mi NE of Boquete, end of rd. along Rio Palo Alto, 18 Nov 1978, Hammel 5692 (BH, MO): Cerro Colorado, top Bocas rd., 1500 m, 17-18 Feb 1977, Folsom & Collins 1739 (MO); Cerro Horqueta, 5000-7000 ft, 8 Aug 1967, Kirkbride 135 (MO); trail to Cerro Pate de Macho, headwaters of Rio Palo Alto, above Palo Alto, 1700-2100 m, 8?47'N, 82?22'W, 15 Mar 1982, Knapp et al. 4270 (MO); N of San Felix at ChiriquiBocas del Toro border, on Cerro Colorado copper mine rd. along Continental Divide, 5000-5500 ft. 5 May 1975, Mori & Kallunki 5929 (MO). Solanum ramonense is very similar vegetatively to S. roblense, also of montane Costa Rica, but is easily

distinguishedfrom thatspecies by its minute flowers, triangularcalyx lobes, and largefruitson ratherstocky StandleY& Valerio45514 (US). HEREDIA: VaraBlanca pedicels. Solanum ramonenseis most closely related de Sarapiqui, N slope of Cordillera Central, 1500-1750 to S. arboreumof lowland Centraland northernSouth m, Jul--Sep 1937, Skutch 3280 (K, US); P.N. Braulio Carrilo, Est. Barra, 2300 m, 10?07'222N, 84007'15"w, America,with glabrousleaves, congestedpedicel scars, 24 Jun 1990, Varela87 (INB). PUNTARENAS: Monteverde, and berriesborne on erect pedicels. The ripe fruitwhen split open smells stronglyof above Quebrada Cuecha, 1540-1600 m, 2 Aug 1976, a trait observed for other deep forest wintergreen, 724 (CR, F); S of rd. to reserve, 1520-1580 m, DrYer 26 Jan 1977, Drver 1158A (F); above Quebrada Cuecha, solanums (Knapp & Helgason, 1997). This suggests

TAXONOMIC TREATMENT

an animaldispersalagent,butnone havebeen observed. Fruitsof Solanum ramonensehave been found on the forest floor with the seeds germinating inside (pers. obs.). The fruitshave very few seeds, andthe seeds are quitelarge.Solanumramonenseblooms in the late wet season and early dry season (Knapp, 1986a), an unusualfloweringtime for shrubsin the CostaRica cloud forest. Most other species of sect. Geminata at Monteverdebloom at the end of the dry season andthe beginning of the wet season.

219

ca. 1 mm long, the filamenttubeless than0.5 mm long; ovary glabrous;style straight,5.5-7 mm long; stigma consisting of a papillose area on the tip of the style. Fruit a globose, green berry, 1-1.2 cm diam.;fruiting pedicels 1.5-2 cm long, erect,woody andrathercorky lenticellate like the stems, the base 1-1.5 mm diam. Seeds not known. Chromosome number: n = 12 (voucher Knapp & Mallet 6813). Distribution (Fig. 117). Foundonly in the Venezuelan Andes in the state of Meridaat ca. 2000 m.

Specimens examined. VENEZUELA. LARA: Duaca,1893-4, Mocqueryss.n. (US). MERIDA: 27.2 km S of La Azulita,14.6 km N of Jajiturnoffon La Azulita rd., ca. 2200 m, 8?32'N,71?16'W,22 Oct 1984,Knapp & Mallet 6809, 6813, 6817 (BH, K, MY, US, VEN); Monte Zerpa, 5 km NNE of Merida,2100 m, 30 Dec 1983, Molinari83-2 (MY); La Carbonera,2300 m, 18 & Wiehler106591(MO,US, VEN). Dec 1972,Steyermark Treeletsor small trees, 2-5 m tall, young stems Solanumripense has only been collected a few of dry specimens minutely red-glandular-puberulent, I foundit locally abundantnearLa Carbonera, but times or brown black, soon glabrate;barkof olderstems dark with large corky lenticels 1-2 mm diam. Sympodial Edo. Merida,Venezuela.It grows in cloud forest,often unitsdifoliate, geminate.Leaves elliptic to ovate, wid- in densetangledstandsin old treefalls.The treesdo not est at orjust proximal to the middle, darkgreen, gla- flower profusely; this may be the reason for the paubrous and shining above, glabrous below, slightly city of collections of this species. Solanumripense is paler, the young leaves in dry specimens minutely most similar and probablymost closely related to S. but soon glabrate;ma- tanysepalum,fromwhichit differsin its shorter,fleshier reddish-glandular-puberulent, x 6-12.5 cm, with 5-6 pairs of pri- calyx, longer inflorescences, and larger stature.The jor leaves 12-18 marynerves,these raisedabove, prominentanddarker barkof S. ripenseis distinctivein its blackishcolor and in dry specimens below, the apex acuminate,the base warty texture; this is apparenteven on young twigs. acute, strongly oblique; petioles 1.5-5 cm long; mi- This characteris the only one useful in identifyingsternor leaves 1.2-3 x 0.8-2 cm, the apex rounded or ile specimens of these two species, the leaves of S. acute, the base acute; petioles shorterrelative to leaf ripense and S. tanysepalumbeing nearly identical in size than in the majorleaves, 3-7 mm long. Inflores- shape and texture.In the Flora del Avila (Steyermark cences opposite the leaves, simple, minutely reddish- & Huber, 1978), individuals of S. tanysepalumwere glandular puberulent, 4-6 mm long, 4-6-flowered; identifiedas S. ripense. This is perhapsnot surprising, pedicel scars closely spaced and rathercorky, in age as so few collections of true S. ripense exist and the overlapping, beginning ca. 3 mm from base of the two species are very similar vegetatively. Solanumn inflorescence.Buds ellipsoid-ovoid,completelyclosed ripensedoes not occurin the coastalCordilleraof Venwhen young with one calyx lobe hooded over the rest, ezuela. A label on the isotype specimen in MPU indithe corolla soon exserted from the calyx. Pedicels at that Solanumripense was establishedin cultivacates anthesis erect or deflexed, fleshy, 0.8-1 cm long, tain fromLinden'scollectionin the midLuxembourg tion pering from the calyx tube to a base 0.5-1 mm diam. I have seen no otherindicationthat nineteenth century. Flowers with the calyx tube 1-1.5 mm long, in dry or passed on to any other was cultivated the species specimens appearingwoody, fleshy and pale green in botanic garden. European deltoid, the lobes crateriform, live material, broadly 1.5-2 mm long, the margins thickened and densely papillose; cor-olla white, fleshy, 1.2-1.5 cm diam., 63. Solanum roblense Bitter, Repert. Spec. Nov. lobed nearlyto the base, each petal with a raisedridge Regni.Veg. 18:62. 1922.Type.CostaRica.Heredia: in the center of the adaxial side, the lobes planar or Foretsde L'Alto del Roble, massif de Barba,2000 slightly incurved at anthesis, tips of the lobes cuculm, May 1888, Pittier et Durand 210 (lectotype, late, densely papillose along full marginof the petal; BR, here designated [photos LL, US]). anthers ca. 3.5 x 1-1.5 mm, poricidal at the tips, the Shrubsor small trees, 1-4 m tall; young stems pores teardropshaped; free portion of thefilaments 62. Solanum ripense Dunal in DC., Prodr. 13(1): 680. 1852. Type.Venezuela.Mrida: Rio Cacique, 8000 ft, Jul 1843, Linden 1417 (holotype, G-DC [F neg. 6779]; isotypes, BM, K, frag. MPU, P Fig. 116 [Mortonneg. 831 1]).

NEOTROPICA ~~~~~~~~~~~~~FLORA

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and leaves glabrousor minutely papillose; barkof the olderstems pale, transverselyrugoseandpeeling.Sympodial unitsdifoliate, geminate.Leaves elliptic to narrowly elliptic, widest at the middle, glabrous,the margins lightly crenulate;majorleaves 6.5-15.6 x 2.2-4.6 cm, with 8-12 pairsof mainlateralveins, slightlyraised above, prominentandyellowish beneath,the apex acuminate, the base attenuate;petioles 0.5-1.1 cm long; minorleaves differingfromthe majorones only in size, 2-5.4 x 0.8-2.4 cm, the apex acuminate,the base attenuate;petioles 2-6 mm long. Inflorescences opposite the leaves, simple, glabrous or occasionally minutely papillose at the extreme tip, 0.5-1 cm long; pedicel scars closely spaced and overlapping,beginning at the base of the inflorescence. Buds ovoid, the corollasoonexsertedfromthenearlytruncatecalyxtube. Pedicels at anthesis 0.9-1.1 cm long, deflexed, tapering fromthe base of the calyx tube to a slenderbase ca. 0.5 mm diam. Flowers with the calyx tube 1-1.5 mm long, narrowlyconical, the lobes short deltoid, 0.5-1 mm long, glabrousor occasionally minutelypapillose; corolla white, 1-1.3 cm diam., lobed 3/4 of the way to the base, planarat anthesis (?), the tips and marginsof the lobes minutelypapillose;anthers3-3.5 x 0.75-1.5 mm, the terminalportion ca. 0.5 mm long, paler and thickened, poricidal at the tips, the pores teardrop shaped;free portionof thefilaments ca. 0.1 mm long, the filament tube 0.5-0.75 mm long; ovary glabrous; stylestraight,ca. 5.5 mm long;stigmacapitate,minutely papillose. Fruit a globose, green, glabrousberry,0.81.1 cm diam.;fruitingpedicels erect,woody, 1.6-2 cm long, 1-1.5 mm diam. at the base; calyx lobes slightly

accrescent and woody in fruit, ca. 2 mm long, with thick margins. Seeds tan, ovoid-reniform, ca. 3 x 2 mm, the surfaces minutely pitted. Chromosome number not known. Distribution (Fig. 117). In primary forest in central montane Costa Rica. Most of the collections are from the slopes of Volcan Barba and Volcan Poas from 1800-2000 m. Specimens examined. COSTA RICA. ALAJUELA: SE slope Volcan Poas, near Poasito, 2000 m, 10?10'N, 84?12'W, 16 Apr 1973, Gentry & Burger 2961 (CR, F, MO, U); Palmira, 6000 ft, 5 Oct 1937, Smith A479 (F, US), 21 Apr 1937, Smith 4138 (F); vic. of Fraijanes, 1500-1700 m, 12-13 Feb 1926, Standley & Torres R. 47564 (US). CARTAGO:Between La Georgina and La Giralda, 1850 m, 20 Sep 1966, Jimenez M. 4121 (F, NY). HEREDIA: Pass between Volcan Irazu and Volcan Barba, beyond San Rafael de Heredia, 22 Apr 1969, Davidse & Pohl 1694 (F); Rio Vueltas (upper Rio Patria)on E slope of Volcan Barba, Caribbean side of Continental Divide, 1900 m, 10?06'N, 84?04'W, 1-3 Apr 1973, Gentry & Burger 2863A, 2864 (F, MO); forest of Rio Vueltas, 2100 m, 23 May 1969, G6mez P. 2243 (F, MO); Vara Blanca, 1900 m, 11 Feb 1951, Le6n 3070 (CR); 100 m N of Vara Blanca restaurant, 1920 m, 4 Feb 1966, Jimenez M. 3667 (CR, F, NY); Vara Blanca de Sarapiqui, N slope of Cordillera Central, between Poas and Barba volcanoes, 1950 m, Jan 1938, Skutch 3477 (MO, NY, US); Cerros de Zurqui NE of San Isidro, 2000-2400 m, 3 Mar 1926, Standley & Valerio 50379 (US). SAN JOSE: Villa Mills, Cerro de la Muerte, between Villa Mills and Divisi6n, 2800 m, 26 Feb 1965, Jimenez M. 2978 (CR, F); Parque Nacional Chirrip6, Cord. de Talamanca, trail to Mirador,

222

FLORA NEOTROPICA

FIG. 118. Solanum tanysepalum S. Knapp. (Reproduced with permission from Brittonia 38: 285, fig. 9. 1986.)

2600 m, 9?33'20"N, 83?40'15"W, 18 Aug 1995, Picado & Gamboa 265 (INB); Piedra Blanca, Escaz6, 1900 m, 20 Jan 1935, Solis 163 (CR, F, MO); near Laguna de la Escuadra NE of El Copey, 2000-2200 m, 16 Dec 1925, Standley 41937, 41988 (US); near Finca La Cima, above Los Lotes, N of El Copey, 2100-2400 m, 21-22 Dec 1925, Standley 42566 (US); Tablazo, 1800 m, 23 Jan 1935, Valerio 1044 (F); N of San Isidro del General, 7000-1 1,000 ft, 12 Auf 1971, Vaughnet al. 682 (MO). Solanum roblense is very similar in appearance and is probably closely related to S. ramonense, another montane Costa Rican species. Solanum roblense is easily distinguished from S. ramonense by its larger flowers, longer, thinner anthers with a terminal thickened portion, and by its narrower leaves. The calyx of

S. roblense is truncatein bud, in flower the lobes become short deltoid, and in fruit are woody and accrescent, while the calyx lobes of S. ramonenseare long-triangularin budandflower,andshrivelandbreak off in fruit.

64. Solanum tanysepalum S. Knapp, Brittonia 38: 284. 1986.Type.Venezuela.Aragua:ParqueNacional HenriPittier,Portachueloto Pico Periquitotrail,W of Estaci6nBiol6gica RanchoGrande,premontane to montane rainforest, 1100-1400 m, 10?21'N, 67?42'W,22 Oct 1984, Knapp& Mallet 6856 (holotype, MY; isotypes, BH, K, NY, US, VEN). Figs. 102C,D, 118

TAXONOMIC TREATMENT

223

Solanumripenseof Steyerm.& Huber,Floradel Avila Tovar),N slope Cordillerade la Costa, ca. 1850 m, 10?26'N, 825. 1978, not of Dunal.

Shruibs, 1-2.5 m tall; young stems and leaves minutely red-papillose; bark of older stems grayish, often transversely peeling. Sympodial units difoliate, geminate. Leaves elliptic to ovate, widest at or just proximal to the middle, glabrous on both surfaces, the upper surfaces shiny, the lower matte, drying reddish; major leaves 10-20 x 4-9 cm, with 5-7 pairs of main lateral veins, prominent above and below, the apex acuminate, the base oblique, acute to rounded; petioles 1.5-5 cm long; minor leaves differing from the major ones only in size, 2.5-3.5 x 1.5-2 cm, the apex acute, the base rounded; petioles 3-5 mm long. Inflorescences opposite the leaves, sessile, with 1-2 flowers arising directly from the swollen node; pedicel scars on the node, closely spaced, corky. Buds ellipsoid, the calyx lobes long-acuminate and enclosing the corolla until just before anthesis. Pedicels at anthesis pale green, 1.5-2 cm long, strongly 5-ribbed, ratherthick and fleshy, ca. 1 mm diam. at the base. Flowers with the calyx tube 1-1.5 mm long, 5-ribbed, the lobes extremely long-acuminate, also ridged with the ribs of the pedicel, glabrous; corolla white, 1.2-1.5 cm diam., lobed ca. 3/4 of the way to the base, the lobes planar or with the tips slightly reflexed at anthesis, the tips hooded, the tips and margins of the lobes densely papillose; anthers 3-3.5 mm long, the terminal 0.5 mm paler and thickened, ca. 1 mm wide, poricidal at the tips, the pores teardrop shaped; free portion of thefilaments ca. 0.5 mm long, the filament tube minute, not visible in most specimens; ovaiy glabrous; style straight, 6-8 mm long; stigma capitate, bright green in live plants, minutely papillose. Fruit a globose to ellipsoid, green berry, ca. 1 cm diam., often with the persistent style base on top;fruitingpedicels woody and erect, 2.5-3.5 cm long, ca. 1 mm diam. at the base; calyx lobes persistent and somewhat woody in fruit, 0.5-1 cm long. Seeds not known from mature fruits. Chronmosome number.n = 12 (voucherKnapp& Mal-

let 6856). Distribution (Fig. 1 7). In the cloud forests of the Cordillera de la Costa in Venezuela, from 1000-1700 m. Specimens examined. VENEZUELA. ARAGUA: ParqueNacional Henri Pittier, Portachuelo, 13 Aug 1975, Kalin Arroyo & Aristeguieta 75117 (VEN); Parque Nacional Henri Pittier, Rancho Grande, trail to Pico Guacamayo behind station, 1100-1400 m, 10?21'N, 67?42'W, 27 Oct 1984, Knapp & Mallet 6846, 6848 (BH, MY, US, VEN); DISTRITO FEDERAL: 2 km NW of jct. of rd. between El Junquitoand Tovar and rd. to Carayaca, 1770 m, 13 Nov 1973, Davidse 4036 (VEN); 1 km NW of jct. of rd. between El Junquito and Tovar and rd. to Carayaca, 1820 m, 15 Nov 1973, Davidse & Tillett 4059 (VEN); prope Colonia Tovar, 1854-55, Fendler 975 (K, MO, NY, P); ca. 1 km NW of jct. with El Junquito-Colonia Tovarrd. on rd. to Carayaca(jct. is ca. 16 km E of Colonia

67?08'W, 29 Oct 1984, Knapp & Mallet 6862, 6864 (BH, K, MY, US, VEN); ParqueNacional El Avila, N of Boca de Tigre on N facing slopes of Cordillera de la Costa. 1650-1800 m, 10?35'N, 66?55'W, 31 Oct 1984, Knapp & Mallet 6871 (BH, K, MO, MY, US, VEN); Parque Nacional El Avila, Quebrada Gamboa, 1600 m, 29 May 1976, Manara s.n. (VEN); Papelon, Feb 1938, Tamavo 451 (VEN). FALCON: SierraSan Luis, ridges aroundHotel Parador, ca. 7 km S of Curimagua, 1300-1350 m. 11?10'N, 69?35'W, 28 Sep 1984, Knapp & Mallet 6673, 6687 (BH, K, MY, VEN).

Solanumtanysepalumis closely relatedandvegetatively extremely similar to S. ripense of western AndeanVenezuela.Sterilespecimensarenearlyimpossible to distinguish. Solanum ripense however, is a largetreeletto 10 m tall in high cloudforestabove2000 m, while S. tanysepalumis always a small shrub of forest understory at somewhat lower elevations. Solanum tanysepalumis easily distinguished fromS. ripensewhenreproductiveby its sessile inflorescences, strongly five-ridged pedicels, long-acuminate calyx lobes, andoften elliptic berries.Solanumtanvsepaluni grows in forests with S. ombrophilum(S. confine species group),but is usually the less common where the two occur together.

IX. Solanum unifoliatum species group(S. bahianum, S. bellum, S. cyclophyllum,S. longevirgatum,S. marantifolium,S. triplinervium, S. unifoliatum). Fig. lOlE,F Shrubsor small trees;young stems andleaves glabrous or pubescent,the trichomessimple, golden and uniseriateorfloccose.Sympodialunitsunifoliate.Leaves ellipticto orbicular,glabrousorpubescent,thetrichomes usuallygoldenandsimpleuniseriate,oftenwith strongly parallel secondaryvenation, the margins often erose. Inflorescencesopposite the leaves, simple, congested; pedicel scars closely spaced, often overlapping.Buids globose, very small. Flower-swhite to greenish-white, small(0.5-1.2 cm diam.).Fruitgreenandhard;fruiting pedicelserector slightlydeflexed.Seedsovoid-reniform. Distribution. Choc6,Colombia,W andAmazonian Ecuadorand SE Brazil. Membersof this species groupare concentratedin the Choc6 floristic province, butSolanumbellumnand S. bahianumoccurwell outside it in AmazonianEcuador and coastal Brazil respectively. They are also the most morphologicallydivergentspecies of the species group.Speciesin the groupall haveunifoliatesympodia and relatively elongate inflorescences with tightly packedpedicelscars.Theflowerstendto be rathersmall, andthe fruitingpedicels erect.

FLORA NEOTROPICA

224

Key to the species of the Solanum unifoliatumspecies group 1. Primary leaf veins 3-5 pairs, arising in the basal 1/2 of the blade. 2. Leaves elliptic; secondary leaf venation conspicuously parallel, like the leaves of Melastomataceae. Gorgona Island, Narifio, Colombia and coastal Ecuador ............................................ 70. S. triplinervluin 2. Leaves broadly elliptic to orbicular; secondary leaf venation somewhat parallel. 800-2000 m. ...................... 67. S. c(clophyllioln Choc6, Colombia and adjacent Ecuador...................................... 1. Primary leaf veins 5-20 pairs, not all arising in the lower 1/2 of the blade. 3. Inflorescence axis pubescent. 4. Trichomes of the inflorescence axis minute and floccose; calyx lobes reflexed at anthesis. Bahia, Sabhiaoulnl S. Brazil.65 4. Trichomes of the inflorescence axis golden, simple and uniseriate; calyx lobes not reflexed at 68. S. longeivirgatumn anthesis. Cordillera Occidental, Colombia, ca. 2000 m ...................................... 3. Inflorescence axis glabrous. 5. Main lateral leaf veins ca. 20 pairs, conspicuously parallel; fruiting calyx truncate. 600-2(000 69. S. o1arautiifbliu7ni m, S Colombia, N Ecuador. 5. Main lateral leaf veins 5-10 pairs, not conspicuously parallel; fruiting calyx not trtuncate. the lobes deltate or rounded. 6. Abaxial leaf surface pale green or whitish, the venation obscure; corolla white. E 6;6. S. belito Ecuador ...................................... 6. Abaxial leaf surface not different in color from adaxial surface, the venation prominent: corolla pale green. 7. Calyx lobes deltate to long-triangular, 1-1.5 mm; leaves broadly elliptic. 67. iS. c'1vclopvll'Y111 800-2000 m ................................... 7. Calyx lobes broadly rounded, ca. 0.5 mm; leaves elliptic. Sea level to 150 m ................................................................................................................................71. S. Iun7i/ liatrn1

65. Solanum bahianum S. Knapp, Bull. Brit. Mus. (Nat. Hist.), Bot. 19: 109. 1989. Type. Brazil. Bahia: llheus, CEPEC,cacao plantation,25 Oct 1969, TS. dos Santos 513 (holotype, CEPEC; isotype, Fig. 119 US).

at the tips, the pores teardropshaped; free portion of thefilamentsca. 0.5 mm long, the filamenttube ca. 0.1 mm long;ovaty glabrous,yellow (fidedos Santos513); st'le erect, ca. 3 mm long; stigma capitate,the surface minutelypapillose.Fruit a globose, greenberry,1-1.5 Slubshrubto shrub, 0.5-1.5 m tall; young stems cm diam.;fiuitingpedicels woody, deflexed, 2-2.5 cm long, ca. 1.5 mm diam.at the apex, ca. 0.5 mm diam.at andleaves minutelypuberulentwith tiny,reddish,flocthe base, the calyx lobes to 5 mm long in fruit.Seeds cose, branchedtrichomes, soon becoming glabrous; greenishor darkbrown, ovoid-reniform,4.5-5 x 2.5units older stems glabrous,reddish-brown.Synmpodial 3 mm, the surfaces minutely pitted. Clhromtiosomle nminunifoliate. Leaves elliptic, widest at the middle, combee not known. pletely glabrous both adaxially and abaxially; major leaves 13.5-21 x 6-9 cm, with 8-10 pairs of veins Distribution (Fig. 124). In wet forest (mata dryingpale yellow abaxially,the apexacute-acuminate, higr6fila)nearthe coast in southernBahia.Onlyknown the base acute.Inflorescencesmoreor less oppositethe fromthe type locality. leaves, often somewhat internodal,simple, 1-1.5 cm Specimens examined. BRAZIL. BAHIA: Ilheus, long, with 10-15 flowers, the inflorescence axis mi- CEPEC (Centro de Pesquisas da Cacau) inventory area. nutelypuberulentwith reddishfloccose trichomeslike quadrat D, km 22 llheus-Itabuna (BR 415), 50 m, 8 Jtl those of the young stems and leaves; pedicel scars 1981, Hage & E.B. dos Sanitos 1068 (CEPEC. F), 12 closely spaced in distal 1/2 of the inflorescence.Buds Jan 1982, H/age & E.B. dos Santos 1592 (CEPEC, F), 2 globose, the corolla not long-exserted from the calyx Feb 1983, Hage 1670 (CEPEC, F); same locality, Parque tube.Pedicels at anthesisdeflexed, 6-7 mm long, fili- Zoobotanico, 5 Jun 1986, Hage & Brito 20(53 (NY), 10 fonrm, taperingfromthe calyx tubeto a base ca. 0.1 mm Jun 1986, Hage & (1os Sanitos 2065 (NY). diam. Flowverswith the calvx tube conical, ca. 1 mm Solanumnbahianum is a species of somewhat long, the lobes reflexed at anthesis, membraneous, uncertainaffinities in this group. It clearly belongs in deltate,ca. I mm long, with a few floccose trichomesat this assemblagewith its unifoliatesympodia.tiny glothe tips; corn/llawhite, 7-9 mm diam., lobed nearlyto bose buds, small flowers, and filiform flowering the base, the lobes slightlyreflexed,the tips of the lobes pedicels. Solanum bahianumis unusual in the group minutelypapillose;antheris1.5-2 x ca. I mm, poricidal (and in the traditionalconcept of sect. Geminata:see

TAXONOMIC TREATMENT

225

FIG. 119. Solanu(m hahianlumtS. Knapp. (Reproduced with permission from Bulletin of the British Museum, Natural History (Botany) 19: 111, fig. 8. 1989.)

Morphology;History)in possessing floccose branched trichomeson the new growth and inflorescence axes. These trichomes are common in the S. arboretimspecies group,which I considerto be closely relatedto the unifoliatumgroup (see Knapp, 1986a). The combination of tiny globose buds, reflexed calyx lobes at anthesis, unifoliatesympodia,and floccose trichomeson the new growth is unique in sect. Geminata. Solanumbahianumis superficiallysimilarto the sympatricS. santosii:bothhave small flowers andpendant fruits (Knapp, 1989b). It is not unusual to find two similarspecies of sect. Geminatagrowing sympatrically (Knapp, 1986a, 1986b).

Ecuatorianadel Te, C.A.) nearPalora,selectivelycut woods, ca. 950 m, 1?40'S,77?58'W, 15 Feb 1984, Knapp & Mallet 6298 (holotype, BH, isotypes, F, NY, QCA, QCNE, US). Figs. IOIE,F, 120

Delicateshrubor small slendertreelet 1-3 m tall; young stems completely glabrous, shiny and green; barkof the older stems gray.Sympodialunits unifoliate. Leaves elliptic to lanceolate,widest at the middle, glabrous on both surfaces, the undersurfacespale in bothdryand live plants,8-16 x 2-5 cm, with 7-9 pairs of main lateralveins, these raised above, in dry specimens somewhat prominentbelow, darkish,the rest of the venation obscure, the apex acute, the base acute, decurrenton the petiole;petioles 0.8-2 cm long, longer 66. Solanum bellum S. Knapp,Brittonia38: 294. 1986. on older leaves. Inflorescences opposite the leaves, Type.Ecuador.Morona-Santiago: alongRio Metzera minute,simple, 3-5 mm long, 3-6-flowered, glabrous; Grandeon HaciendaSangay(plantationof Compania pedicel scars closely spaced,butnot overlapping.Buds

226

FLORA NEOTROPICA

FIG. 120. Solanum bellum S. Knapp. (Reproduced with permission from Brittonia 38: 295, fig. 14. 1986.)

globose, when very young minutely hirsute with uniseriatetrichomesca. 0.1 mm long or less, the corolla soon exsertedfromthe calyx.Pedicels at anthesisslender,6-8 mm long,deflexed,ca. 0.3 mmdiam.atthebase. Flowers with the calyx tube not much differentiated from the pedicel, ca. 0.25 mm long, the lobes broadly deltoid,minutelyapiculate,ca. 0.25 mm long, glabrous except for the tips of the lobes; corolla white, minute, 4-5 mm diam., lobed ca. 2/3 of the way to the base, the lobes reflexedat anthesis,the tips of the lobes minutely papillose; anthers 1-1.5 mm long, ca. 0.75 mm wide, poricidal at the tips, the pores teardropshaped; free portionof thefilamentsca. 0.25 mm long, the filament tube ca. 0.25 mm long; ovary glabrous;style straight, in long-styled flowers ca. 3 mm long, ca. 1 mm long in short-styledflowers;stigma a papillose areaon the tips of the style. Fruit a globose, greenberry,ca. 1 cm diam. (immature);fruitingpedicels erect, woody, 2-2.2 cm long, ca. 0.75 mm diam. at the base. Seeds not mature in fruits examined. Chromosomenumbernot known. Distribution (Fig. 124). In middle-elevation forests in eastem Ecuador,one recent collection from adjacentPeru, 900 to 1500 m.

Specimens examined. ECUADOR. MORONASANTIAGO: E of Misi6n Bomboiza, ca. 840 m, 1 Feb 1971,

MacBryde 175 (MO); along Rio Metzera Grandeon Hacienda Sangay (plantation of Compania Ecuatoriana del Te, S.A.) near Palora, ca. 950 m, 1?40'S, 77?58'W, 15 Feb 1984, Knapp & Mallet 6277 (BH, QCA, QCNE, US); Knapp & Mallet 6280 (BH, QCA, QCNE, US); Cordillera de Cutucu, W slopes along trail from Logrono to Yaupi, 1600 m, 2?46'S, 78?06'W, Nov 1976, Madison et al. 3352, 3527 (MO); Rio Cuyes and BomboizaGualaquiza rd., 800 m, 3?25'S, 78?35'W, 1 Nov 1986, Palacios 1461 (MO, NY). NAPO: Huirano, at foot of Volcan Sumaco, Hollin-Loreto rd., 5 km SE of Loreto, ca. 450 m, 0?43'S, 77?20'W, 24 Nov 1989, Hurtado 2694 (QCNE); Reserva Biol6gica Jatuna Sacha, 8 km from Puerto Misahualli, right bank of Rio Napo, 450 m, 1?04'S,77?37'W,1-15 Sep 1987, Palacios 1957 (MO, NY). PASTAZA:Via Auca, 115 km S of Coca, 10 km S of NapoPastaza border,near Rio Tiguiino,320 m, 1?15'S, 76?55'W, 26-31 Jan 1989, Hurtado & Neill 1445 (QCNE); petroleum exploration line Corrientes of UNOCAL, 300 m, 1043'S, 76?49'W, 1-31 Aug 1990, Gudinio555 (QCNE); 17 km N of Palora, ca. 2 km N of Tashapi (Rio Pastaza crossing), 46 km S of Puyo on Puyo-Palora rd., W of rd., ca. 900 m, 1?42'S, 77?52'W, 17 Feb 1984, Knapp & Mallet 6302, 6304 (BH, NY, QCA, QCNE, US).

TAXONOMIC TREATMENT PERU. SANMARTiN: Nearvillageof Palestina,trail to Cueva Palestina, Bosque de Protecci6n, 890 m, 5?54'S, 77?21'W, 1 Nov 1996, Sanchez Vega& Dillon 8400 (BM).

Solanumbellumis most probablyclosely related to S. unifoliatum and S. triplinerviumfrom westem Colombia, with unifoliate nodes and minute flowers, but is the only memberof the species groupto occuron the eastem slope of the Andes. Solanumbellumdiffers fromthose species in its fleshy leaves that drypale on theundersides.Theleaveson dryspecimensareextremely pale on the undersides,and often the secondaryvenation is not at all obvious. Leaves of live plantsdevelop a silvery sheen when growing in full sunlight, and are generally paler than those of other members of sect. Geminata.

67. Solanum cyclophyllum S. Knapp,Bull. Brit.Mus. (Nat. Hist.), Bot. 22: 147. 1992. Type. Colombia. Choc6: San Jose del Palmar, Cerro del Torra,E slope, nearedge of heliport, 1600 m, 26 Aug 1988, Ramoset al. 1603 (holotype,CUVC;isotypes,MO, NY). Fig. 121 Shrubs 1-3 m tall; young stems and leaves glabrous; barkof older stems pale yellowish-gray.Sympodial units unifoliate. Leaves broadly elliptic to orbicular, widest at or just below the middle, glabrous and shining adaxially,glabrousand drying somewhat golden abaxially,with 3-9 pairs of main lateralveins, the secondaryvenationconspicuouslyparallel;lamina 1-21 cm long, 6-19 cm wide, the apex acute,the base acute to rounded;petioles 1-2 cm long.Inflorescences opposite the leaves or sometimes intemodal, simple, 3-10-flowered, glabrousexcept for a few reddishpapillose trichomes on the tips of the axes;pedicel scars closely packed, not overlapping. Buds ovoid with pointed tips, ca. 1/2 exserted from the calyx tube. Pedicels at anthesis deflexed, glabrous,5-7 mm long, ca. 0.5 mm diam. at the base and apex. Flowers with the calyx tubeconical, 1-1.5 mm long, the lobes deltate to long-triangular,1-1.5 mm long, glabrous;corolla greenish-whiteto brightgreen(fideRamoset al. 1603), 0.7-1 cm diam., lobed ca. 3/4 of the way to the base, the lobes more or less planarat anthesis, the tips and marginsof the lobes minutelypapillose;anthers2-2.5 mm long, ca. 1 inm wide, poridicalat the tips, thepores teardropshaped;freeportionof thefilaments0.5-1 mm long, the filament tube 0.25-0.5 mm long, glabrous; ovaryglabrous;style straight,4-5 mm long, glabrous; stigma capitate,the surfaceminutelypapillose.Fruit a globose, greento greenish-whiteberry,1-1.5 cm diam.; fruhitingpedicels woody,erect, 1-1.5 cm long, 1.5-2 mm diam. at the base. Seeds darkbrown, ovoid-reniform,

227

4-4.5 mm long, 2-2.5 mm wide, the surfacesminutely pitted. Chromosomenumbernot known. Distribution (Fig. 124). WesternColombiaand Ecuadorfrom Choc6 to Esmeraldas,800-2000 m, in wet cloud forest. Specimens examined. COLOMBIA. CHOCO: N6vita, N slope of Cerrodel Torra,ridge to W of Rio Surama,trailto Alto de Oso, 600-900 m, 22 Feb 1977, Forero et al. 3175 (COL,MO);Mun.San Jos6 del Palmar,FincaEl Tabor,1650 m, 17 Jan 1983,Franco et al. 1395 (MO);San Jose del Palmar,Cerrodel Torra,base of E slope,rightedge of heliport,2000 m, 31 Aug 1988, Ramos et al. 1663 (CUVC,MO, NY). Valle:Bosquede San Antonio, W of Cali, near television tower, 19502050 m, 15 Jul 1984, Gentry et al. 48121 (MO, NY); old rd. to Buenaventura kms 14-18, above Saladito,ca. 1800 m, 21 Oct 1987,Ramos 803 (CUVC,MO,NY); El Cairo,Cerrodel Ingles, Serraniade los Paraguas,Cordillera Occidental,Valle-Choc6border1 hourby jeep fromEl Cairo,2070-2400 m, 1 Apr 1988,SilverstoneSopkin et al. 3946 (CUVC,MO, NY). ECUADOR. ESMERALDAS: ReservaEtnicaAwa, CerroMataje,200 m, 1?08N, 78?33'W,21 Sep 1992, Aulestia et al. 420 (BM, MO, QCNE). Solanum cyclophyllum is closely related to S. triplinerviumof GorgonaIslandoff the coast of Narifno department.It shareswith thatspecies three-nervedvenation (at the base) and completely glabrous foliage. The regular,deltateto long-triangularcalyx lobes and ovate to roundedleaves readily distinguish it fromS. triplinervium.The parallelsecondaryvenation shared between the two species is much moreprominentin S. triplinervium (see Fig. 125). The only other member of the Solanum unifoliatumspecies groupoccurringat middle to high elevations is S. longevirgatum,which is easily distinguished fromS. cyclophyllumby its golden pubescent stems and young leaves and its erose leaf margins.

68. Solanum longevirgatum Bitter, Repert. Spec. Nov. Regni Veg. 18:67. 1922. Type. Colombia. Cauca:an den Westgehangender West-Andenvon Popayan,2000-2500 m, Jun,Lehmann8511 (holotype, B [destroyed: F neg. 2667]; lectotype, F, here designated). Fig. 122 SolanumcrispulumC.V. Morton,Contr.U.S. Natl. Herb.

29: 48. 1944. Type.Colombia.Cauca:Cordillera Occidental,CerroMunchique.vert.W. en la hoya del Rio Tambito, 2000-2500 m, 16 Jul 1939. Perez-A rbelez

& Cuatrecasas 6252 (holotype,

COL;isotype frag., US). Slender, often unbranchedshrubs ca. 2 m tall; young stems andleaves dryingblack,the stems hirsute

228

FLORA NEOTROPICA

Solanum,cyclophyllum

S. Knapp-

PLAN TAS DE COLOMBIA

FufuIIa~ ~4..AXMK

Dept:

ThaOC

Mumn.; SantJ or,ado I

Ceo eiarrd, Ca 0MO vertl.ente o,ri-ertal, en su.beev 0. del agua qe ale4 del e*rr, aare,n denofr .d'ibet. nuarto. Arbusto ca. 2 m --iVtura1envda verSe cesarhao ria aute y rpAllna verdesccuras; S#fallfl buse d, in coroal blannco v:ctosoe, '.Ataloe verde oa'rIsing vflTeres wmA1e f t c-op4hlldaa 4_la6arvrt ;a a ovrxio, riatilo yOir? ?308; ran; irJtrftC I p ved .l dro, . cr3 semn-is. un bl-arrousco;, Ali_

FIG. 121. Solanum cyclophyllum S. Knapp. (Reproduced with permission from Bulletin of the British Museum, Natural History (Botany) 22: 149, fig. lb. 1992.)

TAXONOMIC

TREATMENT

.

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with uniseriate golden trichomes 0.1-0.5 mm long, young leaves pubescent on the veins beneath;barkof older stems dark greenish-brown, with large white speckles, sparselypubescentwith uniseriatetrichomes. Sympodialunits unifoliate. Leaves ovate to narrowly ovate, pale yellow-green (fide Lehmann8511), widest just proximalto the middle, with 5-8 pairsof main lateral veins, these raised above, the midrib yellow, all venationprominentlyraised andpale yellow beneath, occasionallypubescenton the veins with uniseriatetrichomes like those of the young stems and leaves, 6.516.5 x 2.6-7 cm, the apex acuminate,the base acute, slightly decurrenton the petiole; cells of the upperepidermisin dryspecimensvery large,ca. 0.05 mm diam., bullate; leaf margins erose, especially on the larger leaves; petioles 0.7-1 cm long. Inflorescences opposite the leaves or intemodal, simple, 1.1-1.5 cm long, 5-1 0-flowered,denselyhirsutewith the sameuniseriate golden trichomes as on the young stems;pedicel scars

evenly spaced ca. 1 mm apart,beginning ca. 1/2 way up the inflorescence. Buds globose, the calyx lobes acuminatein bud, fleshy,with uniseriatetrichomeslike those of the peduncle.Pedicels at anthesisdeflexed, 68 mm long, taperingfrom a basal diam. of ca. 0.5 mm to an abruptwidening at the base of the calyx tube. Flowers with the calyx tube 1-1.5 mm long, slightly urceolate, sparsely pubescent with the same golden uniseriatetrichomesas those of the peduncle,the lobes long-triangular,2.5-3.5 mm long, fleshy in bud, later more petaloid, sparsely pubescent with the same trichomes as the rest of the infloresence,papillose on the tips; corolla white, 1.1-2 cm diam.,lobed nearlyto the base, the lobes narrow,lanceolate,with a few scattered uniseriatetrichomeson the outersurface,minutelypapillose on the tips andmargins;anthers3-3.5 mm long, 1-1.5 mm wide at the widest point, the terminalportion ca. 0.5 mm, paler,thickenedandslightlynarrower, ca. 0.75 mm wide, poricidalat the tips, the pores teardropshaped;free portionof thefilaments0.1-0.5 mm long, the filament tube 0.5-1.5 mm long; ovary glabrous;style straight,ca. 4 mm long; stigma decurrent on the style, minutelywhite-papillose.Fruit andseeds not known. Chromosomenumbernot known. Distribution (Fig. 124). Found only on the westernslopes of the CordilleraOccidentalin the province of Cauca, Colombia, from 2000 to 2500 m. Both type collections seem to have been from the slopes of Cerro Munchique,the highest peak in the Cordillera Occidental. Solanum longevirgatum is unusual in the S. unifoliatumspecies groupin possessingcopiousgolden trichomes on the young stems and leaves. It is otherwise most similarto S. unifoliatumof lowland Choc6

FLORA NEOTROPICA

department,sharingwith that species elongate inflorescences and often erose leaf margins.

69. Solanum marantifolium Bitter, Repert. Spec. Nov. Regni Veg. 11: 13. 1912. Type. Colombia. Narifio:An den WestgehangenderWestAndes von Popayan,2000-2500 m, May-Jun,Lehmann8513 (holotype,B-n.v. [perhapsdestroyed:F neg. 2670]; isotype, K). Fig. 123 Coarseherbor few-branchedshrub,0.5-2 m tall; young stems and leaves glabrousto minutely puberulent with golden, erect, uniseriatetrichomes;bark of older stems brown.Sympodialunitsunifoliate.Leaves broadly elliptic, widest at the middle, 16-25 x 11-13 cm, with ca. 20 pairs of closely parallel main lateral veins; laminaglabrousto minutelypuberulentalongthe midrib adaxially, densely puberulentabaxially with golden,erect,uniseriatetrichomeslikethoseof theyoung stems along the main lateralveins andmidrib,the apex acuminate,the base acute, not winged on the petiole; petiole 1.5-3 cm long, minutelypuberulentwith golden, erect, trichomes.Inflorescencesopposite the leaves or somewhatinternodal,simple, 1.5-4 cm long, the flowers borne only in the upperhalf, the non-flower bearing portionglabrousor minutelypuberulent,the flower bearingportionminutelypuberulentwith golden,erect, uniseriatetrichomes ca. 0.25 mm long;pedicel scars closely spacedin the distalone half of the inflorescence, not overlapping.Buds globose with the calyx crumpled at the apex, the corolla not strongly exserted from the calyx tubeuntiljust before anthesis.Pedicels at anthesis deflexed,2-5 mm long, glabrousorpuberulentwith golden trichomeslike those of the rest of the inflorescence. Flowers with the calyx tube conical, ca. 1 mmn long, the lobes lanceolate, 1.5-2 mm long, reflexed (?) at anthesis,minutelypuberulentwith golden trichomes abaxially,denselypapilloseadaxially;corollawhite,56 mm diam.,the lobes somewhatreflexed (?) at anthesis, the tips and margins of the lobes minutely papillose; anthers 1.5-1.6 x 0.9-1 mm, poricidalat the tips, the poresteardropshaped;free portionof thefilaments ca. 0.5 mm long, the filamenttube minute;ovar' glabrous;style straight,2.5-4 mm long; stigma capitate, minutelypapillose.Fruit a globose, green(?) berry,ca. 1.2cm diam.(fideBitter);fruitingpedicels erect,woody, ca. 1.5 mm diam.; calyx lobes in fruitbroadly obtuse (fide Bitter),apparentlybreakingoff at the base, leaving a broadly flattenedcup with an indistinctmargin. Seeds not known. Chromosomenumbernot known. Distribution (Fig. 124). On the western slopes of the Andes in wet forestfromS Colombiato N Ecuador, from 600 to 2000 m.

TAXONOMIC TREATMENT

231

ft_ 17-

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5cm HRBURT.

BOT. REG. KEW.

HRAIIIlUM HJIlI

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FIG. 123. Solanum marantifolium Bitter. Colombia. Lehmann 8513 (K-isotype).

232

FLORA NEOTROPICA

500 A6

FIG. 124. Distribution of Solaniuml bahianirn (solid square), S. bell/um (solid circles), S. cyclophyllurm (open circles), S. longevirgatium (open square), and S. mat antioliumn(solid triangles).

Specimens examined. ECUADOR. Ri6 Nembi, 23 May 1876, Andre 3405 (K). ESMERALDAS:Rio Lita and tributaries (affluent of Rio Mira), ca. 117 km NW of Ibarra,ca. 12 km N of Lita, ca. 600 m, 0?52'N, 78?29'W, 10 May 1987, Dalv & Acevedo R. 5176 (NY).

crassisobtusiusculis."This soundsvery like the appearance of the fruiting calyx I have seen. On the photographof the holotype no fruit is visible.

Solanummarantifoliumwas originallydescribed as being in sect. Pteroidea,an unrelatedgroupof fleshy forest herbs with axillary inflorescences (Knapp & Helgason, 1997). This species is probablyclosely related to S. longevirgatum,also of the western Andean slopes in southernColombia.Solanummarantifolium differs from S. longevirgatumin its larger, minutely pubescentleaves with conspicuousprimaryvenation,its minutelypuberulentinflorescencesbearingflowersonly in the distal half, and in its smaller flowers. Solanum marantijoliumis also similarto S. cyclophyllumandS. unifoliatumof Choc6 departmentof Colombia.It shares with those species prominentprimaryleaf venationand largeleaves.Boththosespecies,however,arecompletely glabrousand have fewer main lateralveins (3-5 forS. cyclophylluni and 5-7 for S. unifoliatumn). The fruitingcalyx of Solanummarantifoliumis unusual in apparentlylacking lobes (Daly & Acevedo R. 5176 fromwesternEcuador)and looking very similar to the calyx of many Lycianthes (L. synanthera group)or Witheringiaspecies. The calyx lobes appear to abscise at the basejust afterthe corollafalls off. This characteristicis uniqueto S. marantifoliumand needs to be investigated further.Bitter, in his descriptionof S. marantifolium,describesthe calyx as "instatufructifero manifesteampliatus,diam. fere 8 mm, lobis satis

70. Solanum triplinervium C. V. Morton,Contr.U.S. Natl. Herb.29: 47. 1944. Type. Colombia.Narifio: east side of GorgonaIsland,50-100 m, 11 Feb 1939, Killip& Garcia33169 (holotype,US; isotypes,BM, COL-n.v., F [F neg. 49470]). Fig. 125 Slendershrubsca. 2 m tall;youngstemsandleaves glabrous;barkgreen, not lenticellate.Sympodialunits unifoliate.Leavesellipticto oblanceolate,glabrous,widest at orjust abovethe middle,27-35 x 9-15 cm, with 3 (or5) pairsof mainlateralveinsarisingca. 3 cm abovethe leaf base,raisedabove,prominentandbrightyellow beneath,the 3 majorveins runningin recurvedarcsto the pointof convergenceatthe leafapex,thesecondaryveins to theprimariesandparallelwitheachother, perpendicular so thattheleaf venationresemblesthatof membersof the theapexlong-acuminate, familyMelastomataceae, thebase acute,slightlydecurrenton the petiole,oblique;petioles 1.5-2.2 cm long, lightlywinged fromthe decurrentleaf bases.Inflorescencesoppositethe leaves, simple, 1-1.6 cm long,ca. 50-flowered,butonly bearing1-4 flowersat a time,glabrous;pedicel scarscloselyspaced,butnotoverlapping.Budsglobose,glabrous.Pedicelsat anthesisca. 1.1cm long,taperingfromthecalyxtubeto a slenderbase ca. 0.5 mm diam.Flowerswith thecalyxtube 1-1.5 mm long, cup-like,the lobes minute,oftenpresentonly as ir-

23 3

TREATMENT

TAXONOMIC

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234

FLORA NEOTROPICA

FIG. 126. Solanum unifoliatum S. Knapp. (Reproduced with permission from Annals of the Missouri Botanical Garden 73: 741, fig. 3. 1986.)

regularities inthecalyxmargin,ca.0.5 mmlong,glabrous; corollawhite, ca. 8 mm across,lobednearlyto the base, the lobesreflexedat anthesis,glabrous;anthersca. 3 mm long, 1 mm wide, minutelygranular,greenish-yellow, poricidalat the tips, the poresteardropshaped;freeportion of thefilamentsless than0.5 mm long, the filament tube ca. 0.5 mm long; ovaryglabrous;style ca. 4.5 mm long, straight;stigmapapillose,butotherwisenot differing fromthe style.Fruita globose,greenberry,0.8-1 cm diam.(immature?), apiculatewiththecorkypersistentstyle base;fruitingpedicelswoody,erect,ca. 1.5cm long,ca. 1 mm diam. at the base. Seeds not known. Chromosome numbernot known.

Reserva Etnica y Forestal Awa, ca. 200 m, 1?21'N, 78?42'W, 17 Mar 1988, J0rgensen et al. 65310 (AAU).

Solanumtriplinerviumis very similarto andperhaps most closely relatedto S. cyclophyllumof coastal ColombiaandEcuadorandto S. unifoliatumof mainland Choc6departmentColombia,anddiffersfromthosespecies in its shorterinflorescencesand in its strikingleaf venation.The species all shareunifoliatesympodiaand ratherlargeleaves.TheleafvenationofS. triplinervium is unmistakable,makingthis one of the easiest species in sect. Geminatato recognize.Solanumtriplinerviumhas morepronouncedparallel venationthanS.cyclophyllum, butindividualspecimenscanbe difficultto identify.The Distribution (Fig. 127): In primaryforest along calyx lobes of S. triplinerviumarealwaysminute,while streamson Isla Gorgona(Colombia) and the adjacent those ofS. cyclophyllumaredeltateto long-triangular. mainlandof Ecuador. Specimens examined. COLOMBIA. NARINO: 71. Solanum unifoliatum S. Knapp, Ann. Missouri GorgonaIsland,7 Aug 1924,Hicks (St. GeorgeExped.) Bot. Gard.73: 739. 1987 (1986). Type. Colombia. 371 (K); GorgonaIsland, 10 Aug 1924, Langsfield(St. Choc6: Municipio de Choco, CarreteraQuibd6GeorgeExped.) 553 (K). Tutenendo 15 km de Quibdo, 45 m, 6 Sep 1976, ECUADOR. CARCHI: Reserva Indigena Awa, Forero & Jaramillo 2544 (holotype, MO; isotype, GualpiAlto, 1800 m, 1?02'N,78?14'W,15-28 Jun 1991, Rubio et al. 1611 (BM, MO, QCNE). ESMERALDAS: COL-n.v.). Fig. 126

TAXONOMIC TREATMENT

235

Shrubsorsubshrubs,1-3 m tall;stems glabrous, 800 lightly winged between the nodes with the decurrent leaf bases; bark light greenish-brown, sparsely lenticellate,in age becomingpalerandexfoliating.Sympodial units unifoliate. Leaves elliptic to narrowlyelliptic, widest at the middle, glabrouson both surfaces, 18.8-27.5 cm long, 4.8-7.3 cm wide, with 5-6 pairs of main lateral veins, only the midrib raised above, prominentbelow, the apex long-acuminate,the base attenuate,winged onto the petiole and the stem; mar0 gins revoluteandslightly undulate,occasionallysomewhat erose; petioles 2-2.6 cm long, winged from the decurrentleaf bases.Inflorescencesoppositethe leaves, 0 simple, 0.6-3 cm long, 20-100-flowered, sparsely papillose distally; pedicel scars closely and evenly 00~~~~~~~ packed,butnot overlapping.Budsglobose, translucent, minutelypapillose. Pedicels at anthesis 5-6 mm long, 0 deflexed, filiform,less than0.25 mm diam.at the base, 00~~~~ abruptlywidening to the calyx tube.Flowers with the calyx tube ca. 0.5 mm long, cup-shaped, the lobes 0 roundedand irregular,ca. 0.5 mm long, minutelypapillose; corollaminute,palegreen,ca. 5 mm diam.,lobed *V 0 3/4 of the way to the base, the lobes reflexed at anthesis, the tips and margins of the lobes minutely papillose; anthersca. 1.5 mm long, 0.5 mm wide, poricidal 0 at the tips, the pores teardropshaped;free portion of thefilaments less than0.1 mm long, the filament tube less than 0.1 mm long; ovary glabrous;style straight, ca. 2.5 mm long, in short-styled flowers ca. 0.5 mm long; stigma clavate, dark-papilloseat the tip. Fruit a globose, green berry,slightly umbonatewhen immature, 0.8-1 cm diam.;fruitingpedicels erect or someFIG. 127. Distribution of Solanum triplinervium what deflexed, woody, 1.8-2 cm long, 0.5-1 mm diam. (solid circles) and S. unifoliatum(open circles). at the base. Seeds (immature?)ovoid-reniform,ca. 2 mm long, 2 mm wide, the surfaces minutely pitted. Chromosomenumbernot known. cence. It is probably most closely related to S. Distribution (Fig. 127). Foundonly in the plu- cyclophyllumand S. triplinervium,both of the Choc6 vial forestin the departmentof Choc6in Colombia,near floristic province. A habitatnote on Gentry& Fallen sea level. All knowncollections are fromthe upperRio 17585 indicatesthatthe vegetationwhereS. unifoliatum Atrato basin. Apparentlya plant of both primaryand occurs ("wet forest on gray sand")is very like that of Amazonianwhite sand regions. secondary forest. Onlytwo flowersarepresenton thetypespecimen, Specimens examined. COLOMBIA. CHOC6: short-styled. Andromonoecy Quibd6, Guayabal, Rio Hug6n, ca. 80 m, 12 Sep 1976, andoneoftheseis apparently Forero & Jaramillo 2808 (MEXU, MO); 7 km W of has not been demonstratedin sect. Geminata,but this Tutenendo on rd. to Quibd6, ca. 100 m, 12 Aug 1976, is one of the many species with short and long-styled Gentry & Fallen 17585 (MO, NY); 11 km S of Quibd6 flowers. The large numbersof fruit set on each infloon rd. to Yuto, ca. 50 m, 7 Jan 1979, Gentry & Renteria rescence of S. unifoliatumseems to indicate that the A. 23726 (MO); rd. from Lloro to Yuto, ca. 2 km E of species is not andromonoecious(see Whalen& Costich, Yuto, ca. 50 m, 18 Jan 1979, Gentry & Renteria A. 24410 1986), but furtherwork is clearly needed.

D0

(MO, NY); Bagad6, near town, 80 m, 5?25'N, 76?25'W, 6 Dec 1983, Juncosa 1485 (MO, NY). Solanum

unifoliatum

is very similar to S.

longevirgatumof high-elevation CordilleraOccidental, but differs from that species in its lack of pubes-

X. Solanum robustifrons species group (S. abitaguense, S. cucullatum, S. heleonastes, S. robustifrons,S. superbum). Fig. 128

FLORA NEOTROPICA

236

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FIG. 128. Solanum robustifronsspeciesgroup.A. S. abitaguense plant,Ecuador (Knapp & Mallkt 6276). B. S. abitaguense flowers, Ecuador(Knapp & Mallet 6180). C. S. cucullatum flowers, Ecuador(Knapp & Mallet 6264). D. S. abitaguense fruit, Ecuador(Knapp & Mallet 6180). E. S. robustifrons flowers, Peru (Knapp & Mallet 6659). F. S. robustifrons fruit, Peru (Knapp & Mallet 6653).

237

TAXONOMIC TREATMENT

Shrubs or small trees, often of wet places; young stems and leaves densely red- or gray-papillose to densely pubescent (Solanum superbum), thick and fleshy. Sympodial units difoliate, geminate or ocasionally not, not strongly anisophyllous. Leaves largeand fleshy, the venationmore or less parallel,the apex andbase acute,the base often decurrent.Inflorescences opposite the leaves, few to many-flowered,the flowers large and fleshy to waxy, the petals planarat anthesis. Fruit green and hardat maturity,often very

large(to 4 cm diam.).Seeds ovoid-reniform,very large, the spiralembryoclearly visible. Distribution. Andeanfoothills,Ecuadorto Bolivia. Membersof theSolanumrobustifronsspeciesgroup have the largestfruitsof any species of sect. Geminata. These have creamy white flesh and are possibly dispersedby small mammalsor bats. Most of the species in this groupgrow alongsmallstreamsandrivercourses in forested areas. However, S. heleonastes grows in gallery forest in very dry areas.

Key to the species of the Solanum robustifronsspecies group 1. Flowers large, greater than 1.5 cm diam., thick and waxy; calyx lobes deltoid; inflorescence simple: berries large, to 4 cm diam. (usually > 1 cm diam.), densely grayish-papillose. 76. S. superbui 2. Pubescence of dendritic trichomes, Bolivia ............................................................. 2. Plants glabrous or with minute papillate trichomes. Peru and Ecuador. 3. Stems strongly winged, bright green; calyx in buds closed until just before anthesis. E Andean 72. S. abitaguense slope. Ecuador and N Peru....................................................... 3. Stems not winged, gray and lenticellate; calyx in buds open long before anthesis. W Ecuador....................................................... 73. S. ciicillaatiloz 1. Flowers smaller, less than 1.5 cm diam., not thick and waxy; calyx lobes long-triangularto acuminate; inflorescence occasionally branched; berries ca. I cm diam., glabrous. 4. Trees; new growth densely reddish-papillose; leaves not markedly shiny above; inflorescence simple; calyx lobes 2-5 mm long. Swampy forest in drier vegetation (Chaco) in Paraguay, Argentina, and Bolivia .......................................................... 74. S. heleonastes 4. Large shrubs or rhizomatous shrublets; new growth sparsely reddish-papillose; leaves shiny above; inflorescence often branchedand massive, the flowers crowded; calyx lobes ca. 3 mm long. E Peru and adjacent Brazil and Bolivia .......................................................... 75. S. roboustifrons

72. Solanum abitaguense S. Knapp, Brittonia 38: 290. 1986. Type. Ecuador.Tungurahua:Cashurcu, 13-14 km W of Mera on Bafios-Mera rd., along stream, cloud forest, 1200-1400 m, 1?25'S, 781 O'W,22 Jan 1984, Knapp& Mallet 6180 (holotype, BH; isotypes, F, K, NY, QCA, QCNE, US). Figs. 128A,B,D, 129 Shrub or sInall tree, 2-6 m tall; young stems and

leaves densely red-papillose,the new growth appearing grayish and matte on live plants;stems fleshy and green, stronglywinged, the wings 2-5 mm broad;bark of older stems soon glabrate,grayish.Sympodialunits difoliate,geminate.Leaveselliptic,widestatthe middle, glabrous and shiny above, paler below, occasionally minutelyred-papilloseon the veins below;majorleaves 20-50 x 10-20 cm, with 15-17 parallelpairs of main lateralveins, the apex acute,the base acuminate,decurrenton the petiole and from there onto the stem; petioles strongly winged from the bases of the decurrent leaf blades, 4-7 cm long; minor leaves differing from the majorones only in size, 15-19 x 9-1 1 cm, the apex acute,the base acuminate,winged onto the petiole;petioles 1.5--2cm long.Inflorescencesoppositethe leaves, simple or very occasionally furcate,0.5-1 cm long, 3-

5-flowered,minutelyred-papilloselike theyoung stems and leaves;pedicel scars closely spaced and overlapping. Buds ovoid, the calyx a completely closed envelope untiljustbeforeanthesis,when the corollaemerges and tears the lobes. Pedicels at anthesis white and fleshy, 1.5-2 cm long, abruptlynarrowingbelow the calyx tube, then tapering to 0.5-1 mm diam. in dry specimens (thicker in live plants). Flower-swith the calyx tube campanulate,3-3.5 mm long, occasionally minutelyred-papillose,the lobes very irregularin shape, 3-5 mm long, the marginsthickandwhite in dryspecimens; corolla white, ratherfleshy, 2-3 cm diam., planarat anthesis,lobednearlyto the base,the corollatube only 1mm long, the lobes wider at the middle than at the base, the tips and margins of the lobes minutely papillose;anthers4-6 x 1-1.5 mm,poricidalatthe tips, the pores teardropshaped,the antherstightly connate at anthesis;freeportionof thefilamentsca. 1 mm long, the filamenttube ca. 0.5 mm long;ovarydensely pinkish-graypapillose,the papillaeminutein flower, growing with the fruit;style straight,ca. 1.3 cm long;stigmna bilobed,the surfaceminutelypapillose.Fruita globose, green berry with creamy white flesh, ca. 4 cm diam. when fresh;maturefruitglabrate,the young fruitcov-

238

FLORA NEOTROPICA

FIG. 129. Solanum abitaguense S. Knapp. (Reproduced with permission from Brittonia 38: 291, fig. 12. 1986.)

ered with minutepinkish-graypapillae, these making the fruitappeargrayish in live plants, pinkish in dried specimens;fruitingpedicels very woody, deflexed, 22.5 cm long, 3-3.5 mm diam. at the base. Seeds pale tan,ovoid-reniform,ca. 5 x 4 mm, the surfacesminutely pitted. Chromosomenumber:n = 12 (voucherKnapp & Mallet 6180).

Misahualli, right bank of Rio Napo, 450 m, 1?04'S, 77?36'W, 8 Nov 1987, Cer6n M. 2647 (MO, NY); along rd. between Baeza and Lago Agrio, 107 km W of Lago Agrio, 1470 m, 19 Dec 1979, Croat 49481 (MO, NY); El Chaco, E of town, 1700-2000 m, ca. 0?18'S, 77?47'W, 15 Mar 1991, Gavilanes & Quezada 482 (AAU); Cascada San Rafael, turnoff (INECEP Campamiento Quijos, Proyecto Coca) 71.3 km NE of Baeza on rd. to Lago Distribution (Fig. 130). In cloud forests of the Agrio (Nuevo Loja), 1200-1400 m, 0?10'S, 77?40'W, E Andean slope in Ecuadorand N Peru, from 1000 to 25 Jan 1984, Knapp & Mallet 6205 (BH, QCA, QCNE, US); 50 km NE of Baeza on rd. to Lago Agrio (Nuevo 1700 m. Loja), at bridge over Rio Azuela, ca. 1400 m, 0?10'S, Selected specimens examined. ECUADOR. 77?40'W, 25 Jan 1984, Knapp et al. 6209 (BH, K, QCA, BOLIVAR: Chillanes to Bucay rd., Hacienda of Sr. QCNE); Knapp et al. 6211 (BH, QCA, QCNE, US); ca. GonzaloGomez, 2100 m, 1 Sep 1987, Zak 2692 (NY). 65 km NE of Baeza on rd. to Lago Agrio (Nuevo Loja), CHIMBORAZO: Riobamba to Pallatanga rd., near along quebrada N of rd., ca. 1300 m, 0?20'S, 77?47'W, Pallatangaand San Juan,2000-2100 m, 25 Feb 1987, 25 Jan 1984, Knapp et al. 6216 (BH, K, QCA, QCNE, Zak 1747 (NY); Pallatanga-SanJuan-Llimberd., 2300- US); Baeza, ca. 1 km SW of village, 2000 m, 0?28'S, 2720 m, 26 Feb 1987, Zak 1756 (MO). LOJA: Parque 77?53'W, 20 Oct 1976, 0llgaard & Balslev 10235 (AAU, EducacionalRecreacionalUniversidadNacionalde Loja, NY); W slopes of Cordillera de Guacamayos, R bank of vic. of Loja, 2100-2300 m, 4?02.02'S,79?11.87'W,28 Rio Cos, 2400 m, 0?30'S, 77?50'W, 17 Oct 1990, Palacios Oct 1994,Knappet al. 9089, 9091 (QCNE);Urituzinga, 6359 (BM, QCNE). PICHINCHA: Reserva Maquipucuna, 2700 m, 27 May 1978, VivarC. et al. 1089 (LOJA). along trail from Hacienda Esparragos to Cerro Sosa, ca. MORONA-SANTIAGO: 31 km S of Indanza(42 km N of 5 cm SE of Nanegal, 1500-1600 m, 0?07'N, 78?38'W, Gualaquiza),ca. 1600 m, 3?15'S,78?31'W,3 Feb 1984, 13 Sep 1989, Websteret al. 27554 (DAV, IBE); Reserva Knapp& Mallet 6229 (BH, K, QCA, QCNE,US); 10 Maquipucuna, Cerro Sosa, ca. 5 air km SE of Nanegal, km N of Indanza(4 km N of jct. with rd. to Macas)on near Base Camp 2, ca. 2000 m, 0?07'N, 78?38'W, 14 Jul rd. to Gualaceo,1700-1900 m, 3?05'S,78?31'W,3 Feb 1990, Websteret al. 28248 (DAV, IBE). PASTAZA:Along 1984, Knapp& Mallet 6236 (BH, QCA, QCNE,US). rd. between Puyo and Bafnosat second bridge W of Mera, NAPO: ReservaBiol6gicaJatunSacha,8 km fromPuerto ca. 3 km W of Mera, 1160 m, 23 Dec 1979, Croat 49712

TAXONOMIC

239

TREATMENT

FIG. 130. Distribution of Solanum abitaguense (solid circles) and S. heleonastes (solid squares).

(MO). TUNGURAHUA: 6.5 km W of Mera on Banios-Mera rd., ca. 1400 m, 1?25'S, 78?07'W, 22 Jan 1984, Knapp & Mallet 6177 (BH, F, QCA, QCNE, US); Cashurcu, 14 km W of Mera on Banios-Merard., 1300-1400 m, 1?25'S, 78?lO'W, 14 Feb 1984, Knapp & Mallet 6276 (BH, QCA, QCNE, US); 2 km NE of Mera, Hacienda San Antonio del Bar6n von Humboldt, 1100 m, 1?27'S, 78?06'W, 18 Mar 1985, Zaruma et al. 20 (BM, MO). ZAMORACHINCHIPE: ParqueNacional Podocarpus,on Vilcabambavallodolid-Zumba raod, ca. 46.3 km S of Vilcabamba,ca. 2800 m, 4?28.39'S, 79?08.90'W, 19 Oct 1994, Knapp et al. 9040 (QCNE). PERU. AMAZONAS: Shillac, N by trail from Pedro Ruiz, 2300 m, 5?49'S, 78?01'W,31 Aug-2 Sep 1983, Smith & VasquezS. 4892 (MO, NY); Bongara, YambrasbambaPomacocha trail between Yambrasbambaand Yanayacu, 1900-2200 m, 24 Jun 1962, Wurdack 1018 (K, US); Sipabamba, Shilla, ca. 1900 m, 5 May 1981, Young & Eisenberg 361 (MO, NY). CAJAMARCA: Jaen, Hacienda Agua Blanca on Jeronga River tributary of Chunchuga River, 8000 ft, 30 Jul 1943, Evinger 446 (US); Evinger 470 (US). SAN MARTiN: Rioja, near km 398 of Carretera Marginalbetween Pomacochasand Rioja, trail to Quebrada Venceremos and Rio Serranoyacu, 1300-1400 m, 5?45'S, 77?30'W, 10 Jul 1984, Knapp & Mallet 6586 (BH, F, K, US, USM); MariscalCaceres, ParqueNacional Abiseo, Las Palmas, Rio Montecristowatershed,2350 m, 17 Aug 1986, Young 3974 (F, HUT, K); Parque Nacional Abiseo, Rio Montecristo watershed, below Gran Pajaten ruins, 27502850 m, 11 Aug 1986, Young 4177 (F, HUT, K).

Solanum abitaguense is closely related to S. cucullatum of western Ecuador. Both species have stronglyparallelprimaryleaf venation, fleshy leaves, and reddish papillose trichomes on the new growth. SolanumabitaguenseandS. cucullatumaredifficultto distinguish on the herbariumsheet, but fresh material is quite distinct. Charactersuseful in distinguishingS. abitaguense are its shorter buds, less fleshy, more deeply lobed corolla, and its strongly winged, green stems. Solanumabitaguense grows in light gaps along streamsin forests,often with its roots in the water.The buds, flowers, fruits, and seeds of S. abitaguense are the largest (barringS. cucullatum)in sect. Geminata. The calyx is peculiar in bud, being a closed structure, even at the tip, untiljust before anthesis:the exsertion of the corolla tearsthe calyx into irregularlobes. This character,in young buds, is sharedwithS. cucullatum. Another peculiar charactershared by these two extremely closely relatedspecies is the indumentof the ovary,andthen the fruit.The ovary in flower is densely coveredwithpinkish-graypapillosetrichomes,which are much more obvious in dry than fresh specimens. The trichomespersiston the fruitbutbecome less dense as the fruitexpands. The trichomes on dry specimens appearsomewhatresinous, but this is not at all apparent in fresh specimens.

240

FLORA NEOTROPICA

FIG. 131. Solanum cucullatum S. Knapp. (Reproduced with permission from Brittonia 38: 293, fig. 13. 1986.)

73. Solanum cucullatum S. Knapp,Brittonia38: 292. 1986.Type.Ecuador.El Oro:onLoja-Machalard.,10 km W of Las Balzas, 11.5km E of Loja-Piias crossroads(31.5 km E of La Avanzadanear SantaRosa), W slope of N portionof the CordilleraLarga,along stream in forest, ca. 700 m, 3?45'S, 79?50'W, 7 Feb

1984,Knapp& Mallet6266 (holotype,BH; isotypes, F, K, QCA, QCNE, US). Figs. 128C, 131 Smalltreelet,2-6 m tall;young stems andleaves densely red-papillose, the papillae persistentonto the older stems; older stems partially glabrate, gray, not winged;barkofthe trunkgray,smooth.Sympodialunits difoliate, geminate. Leaves elliptic, widest at or just proximal to the middle, the upper surfaces appearing matte in live plants, glabrous, minutely reddish-gray papillose along the veins below; majorleaves 20-45 x 14-21 cm, with 7-9 pairs of main lateralveins, these prominentabove and below, the apex acute, the base attenuate,decurrenton thepetiolebutnot ontothe stem; petioles 2.5-3 mm long, winged fromthe decurrentleaf bases;minorleaves differingfromthe majorones only in size, 10-15 x 6-11 cm, the apex acute, the base attenuate,winged onto the petiole; petioles 1-2 cm long. Inflorescencesoppositethe leaves, occasionallyfurcate, 5-7 mm long, 5-20-flowered, densely reddish-gray papilloselike the young stems andleaves;pedicelscars

closely spaced and overlapping, rather corky. Buds densely red-papillose,ovoid with the tip of the closed calyx elongate and flopping over the rest of the bud, appearinghooded, the corolla soon exserted from the calyx, then the bud appearingfive-pointed from the secondaryhoods of the corolla lobes. Pedicels at anthesis white and fleshy, 1.2-1.5 cm long, erect or deflexed, ca. 0.75 mm diam.at the base.Flowers with the calyx tube campanulate,ca. 3 mm long, the lobes long3-5 mm long, denselyred-papillose, triangular-deltoid, the margins thickened and white in dry specimens; corolla white and fleshy, 2-3 cm diam., lobed ca. 2/3 of the way to the base, the interpetalarsinuses andlobe marginsthinanderose, the lobes planarat anthesis,tips of the lobes with two hoods, one arising from the tip itself andthe other(secondaryhood) from a thickened ridgealongthe entireadaxialsurfaceofthe corollalobe; anthers slightly asymmetricin flower, with two held higherthanthe rest,this characternot apparentin dried specimens, ca. 5 x 1-1.5 mm, poricidal at the tips, the poresteardropshaped;freeportionofthefilaments0.51 mm long, the filamenttube 1-1.5 mm long, visible in the flower; ovary densely pinkish-graypapillose, the papillaetiny in flower, but growingwith the fruit;style straight,ca. 7 mm long;stigma a broadenedareaon the tip of the style,minutelypapillose.Fruita globoseberry,

TAXONOMIC TREATMENT

ca. 3 cm diam. (immature),densely pinkish-graypapillose;fruitingpedicels woody, deflexed or erect, ca. 2 cm long, ca. 2 mm diam. at the base. Seeds tan, ovoidreniform,size not measuredas matureones not seen, the surfaces minutely pitted. Chromosomenumber. n = 12 (voucherKnapp & Mallet 6266).

241 Monteseco, 1500 m, 10 Oct 1993, Leiva G. & Lemma

A. 935 (BM).

Solanum cucullatum is closely related to S. abitaguenseand is difficult to tell fromthatspecies on the herbariumsheet. Both species grow along streams in forests,have largeflowers andfruits,anddistinctive fruit pubescence (see discussion for S. abitagutense). Distribution (Fig. 134). In premontaneor monCharactersuseful in distinguishingS. cucullatumfrom tane wet foreston the W slope of the Andes in Ecuador S. abitaguense are its unwinged grayish stems, more and adjacentPeru, from 700 to ca. 2500 m. elongate hooded buds, and its extremely fleshy flowSelected specimens examined. ECUADOR. ers with doubly-hooded petal tips. In live plants the AZUAY: Along Rio PatulbetweenHaciendaYubayand leaves of S. cuicullatum havemattesurfaces,while those Hacienda San Jose de Caimotan, in region of Sanaguin, of S. abitaguensearesomewhatglossy.Theabaxialpetal ca. 850 m, 28 May 1943, Stevermark52710 (MY, VEN). surfaces of the flowers of S. cucullatumndry greenBOLIVAR: Chillanes-Bucayrd., hacienda"Tiquibuco," ish-pink. 2100 m, 1?55'S, 79?05'W, Zak & Jaramillo 2692 (BM, Small individualsof S. cucullatumappearto be MO, QCNE); carretera Pallatanga-Yunguilla-Llimbe, banks of Rio Chimbo, 1720 m, 7 Sep 1987, Zak & female sterile,while largerplantshave hermaphroditic Jaramillo 2830 (BM, MO, QCNE).EL ORO: 11 km W flowers. This does not appear to be the case in S. abitaguense. Many more collections of S. cucullatunm of Pifias on new rd. to Sta. Rosa, 850 m, 8 Oct 1979, Dodson et al. 9080 (F); on Loja-Machalard. 10 km W are needed to establish this as a pattem in the species. of Las Balzas, ca. 11.5 km E of crossroads Loja-Pifias (31.5 km E of La Avanzada near Santa Rosa), W slope of N portion of Cordillera Larga, ca. 700 m, 3?45'S, 79'50'W, 7 Feb 1984, Knapp & Mallet 6264 (BH, QCA, QCNE, US). IMBABURA:Cotocachi, Tablachupa, rd. to Apuela, NE of Cuicocha, along Rio Azapi, 2200-2500 m, 0?20'N, 78?26'W, 11 Jun 1992, Cuarnacds et al. 138 (BM). Los Rios: 10 km S of Quevedo, 150 m, 1010'S, 79?20'W, 9 Jul 1990, Gudino 492 (QCNE). PICHINCHA: Km 20 Calacali-Nanegalito rd., 2000 m, 0002'N, 78?39'W, 19 Jan 1989, Cer6n et al. 5931 (QCNE); old Quito-Sto. Domingo rd., Reserva Forestal La Favorita, near Rio Saloyu, 1600-1800 m, 0?21'S, 78047'W, 8 Feb 1990, Cerbin& Iguago 8611 (QCNE); km 70 old QuitoSto. Domingo rd., 2100 m, 13 Oct 1981, Dodson & Dodson 11854 (QCNE); Reserva Tropical Maquipucuna, 10 km N of Nanegalito, 1200 m, 0?10'N, 78?35'W, 1 Feb 1991, Espinioza 729 (QCNE); Reserva Floristica-Ecol'ogica

"Rio Guajalito,"km 59 of old Quito-Sto. Domingode los Colorados rd., 3.5 km N of rd., W slopes of Volcan Pichincha, 1800-2200 m, 0?13'53"S, 78048'10"W, 31 Aug 1985, Jaranmillo & Zak 590 (AAU), Zak & Jaramillo 590 (K, MO, NY); Reserva Rio Guajalito, near Chiriboga, 1850 m, 5 Jul 1991, van der Werffet al. 12213 (QCNE); old Qtito-Chiriboga-Empalme rd., km 50, Zapadores, along Rio Saloya, 2000-2100 m, 0?13'S, 78048'W, 21 Jul 1987, Zak & Jaramillo 2196 (K, MO, NY); km 30-34, Lloa-Mindo rd., 2100-2400 m, 0?03'S, 78?40'W, Zak & Jaramillo 2109 (BM, MO); QuitoChiriboga-Empalme rd., km 85, "Estacifn Faisanes," 1400-1450 m, 31 Jul 1987, Zak & Jaramillo 2285 (MO, NY); Quito-Tono-Tandayapa-Pto. Quito rd., between Nono and Tandayapa, 1800-2500m, 0?03'S, 78?30'W, 28 Nov 1987, Zak & Jaramillo 3046 (BM, MO). PERU.CAJAMARCA: Santa Cruz, upper Rio Zafia valley, ca. 5 km above Monte Seco on path below campsite, 1450-1500 m, 19 Mar 1986, Dillon et al. 4438 (BM. F, NY); rd. to Chorro Blanco, Bosque de

74. Solanum heleonastes S. Knapp, Ann. Missouri Bot.Gard.72:560. 1985.Type.Argentina.Corientes: Depto. Capital,PuertoItaliaen montescosterosdel Rio Parana,6 Nov 1972, Schinini 5671 (holotype, MO; isotypes, F, MO, WIS). Fig. 132 Shrubs or small trees, l-5 m tall; young stems and leaves densely covered with appresseduniseriate papillatetrichomes,0.05-0.1 mm long, these irregular and floccose in appearance,grayish-brown,the young growth with a grainy texture;barkof the older stems red-brown, shining, glabrate, minutely whitelenticellate.Sympodialunitsdifoliate,geminate.Leaves ovate, widest just proximal to the middle, glabrous above,minutelypapillateon theveins beneath,theveins yellowish, epidermislarge-celledand appearingcrystalline; majorleaves 10.2-18.5 x 5.8-7.9 cm, with 8-10 strong,evenly spaced main lateralveins, these yellow beneath,the secondaryvenationobscure,the apex acuteor acuminate,the base broadlycuneate,strongly oblique;petioles 1.7-3.2 cm long, slightlywingedfrom the decurrentleaf bases;minorleaves differingfromthe majorones only in size, 7-1 1 x 3.4-5.5 cm, the apex acuteor acuminate,the base broadlycuneate,attenuate, oblique; petioles 0.8-1.2 cm long. Inflorescences opposite the leaves, simple, 1-2 cm long, 10-30-flowered,densely coveredwith papillatetrichomesas those of the young stems and leaves; pedicel scars closely and evenly spaced, but not overlapping.Buds at first globose,enclosedin theurceolatecalyx tube,laterovoid and exserted fromthe calyx tube.Pedicels at anthesis deflexed, 1.1-1.4 cm long, taperingfrom the base of the calyx tubeto a slenderbase ca. 0.5 mm diam.,occa-

242

FLORA NEOTROPICA

FIG. 132. Solanum heleonastes S. Knapp. (Reproduced with permission from Annals of the Missouri Botanical Garden 72 561 fig 3 1985)

sionallydistinctly5-angledandalmostwinged(fideNee 43191), minutely white-speckled, sparsely papillate with uniseriatetrichomeslike those of the young stems andleaves.Flowers with the calyxtube 1.5-2 mm long, urceolate in bud, later cup-shaped,the lobes long-triangular,2.5-5 mm long, white speckled,minutelypapillose at the tips; corolla white (blue in Schwarz8182), 1.2-1.5 cm across, lobed nearly to the base, the lobes planar at anthesis, tips of the lobes slightly carinate, minutelypapillose at the tips andthe margins;anthers 3.5-4 mm long, tightly connate in young flowers, 11.5 mm wide, poricidal at the tips, the pores teardrop shaped;freeportionof thefilamentsca. 1 mm long, the filament tube ca. 0.5 mm long; ovary glabrous;style filiform, 5-7 mm long, straight;stigma minutelycapitate, white-papillose. Fruit a globose berry, dry and

woody with very little pulp, 1-1.2 cm diam., mustard yellow in dried specimens, the septum between the carpels dry and papery in dry specimens;fruiting pedicels erect and woody, 1.1-1.4 cm long, ca. 1 mm diam. at the base; calyx lobes becoming woody and reflexed in fruit.Seeds darkbrown,ovoid-reniformbut somewhat flattened, ca. 2 x 1.5 mm, the surfaces minutely pitted. Chromosomenumbernot known. Distribution (Fig. 130). Found in swampy forests on the marginsandislandsof the upperRio Parana, on the borderbetween ArgentinaandParaguay,and in eastem Bolivia, from 100 to 500 m. Specimens examined. BOLIVIA. BENI: Prov. Cercado,Trinidad,W side of city, ca. 200 m, 14?50'S, 64?55'W,6 Jan 1989, Nee 37520 (IBE, NY). SANTA

TAXONOMIC TREATMENT

243

CRUZ:Prov. Andres Ibafiez, 10 km ESE of Don Lorenzo, vic. of Quebrada Caracore on dirt rd. to Estancia Caracore, 310 m, 17050'S, 62?47'W, 20 Dec 1992, Nee 43191 (BM, NY); Prov. Andres Ibaiiez, 3 km W of Puerto Pailas, along old dirt rd. to Santa Cruz and railroad, 290 m, 17?40'S, 62?49'W, 11 Dec 1994, Nee 45901 (BM, NY); Prov. Ichilo, 3 km W of center of Colonia San Juan, 270 m, 17?17'17"S, 63?52'30"W, 17 Dec 1994, Nee 45960 (BM, NY); banks of Rio Surut6, 1300 m, 22 Aug 1917, Steinbach 3381 (NY). PARAGUAY. S.loc., 1853-1856, Palmer s.n. (US). ARGENTINA.CHACO:10 de Mayo, Col. Benitez, Bermejo, Isla Brasiliera, 4 Oct 1964, Schulz 9422 (F, WIS). CORRIENTES:Capital, on the Rio Parana 5 km from Corrientes, 10 Oct 1944, A. T Hunziker 5700 (US); ltuzaing6, Isla Apipe Grande, Puerto Mora, 11 Dec 1973, Krapovickas et al. 24308 (NY); Arroyo Itaimbi, 15 Dec 1944, Sesmero 201 (NY); Capital, Puerto Italia, 27 Jul 1974, Schinini & Gonzalez 9502 (MO, WIS); Ituzaing6, Isla San Martin, 9 Oct 1949, Schwarz 8182 (BH, W). MISIONES:Posadas, Rio Alto Parana, 18 Nov 1907, Ekmnan 820 (MO); Posadas, 17 Apr 1930, Rodriguez 188 (F); Candelaria, San Juan, 216 m, 4 Jan 1947, Schwindt 30 (RSA/POM).

ally glandular;older stems soon quite thick and very fleshy, later woody, hollow when dry, glabrous;bark dark,not stronglylenticellate.Sympodialunitsdifoliate, sometimes geminate.Leaves elliptic or ovate, if geminatethe two leaves not differingmarkedlyin size, widest at orjust proximalto the middle, 18-44.5 x 9.5-2 7.5 cm, main lateralveins in 10-18 pairs, approximately perpendicularto the midrib,but occasionally arching fromit, the veins on the undersurfacespuberulentwith minute papillate trichomes ca. 0.05 mm long or less, these not presenton very large leaves, laminarsurface above glabrous, finely reticulate from the collapsed upperwalls of the largeepidennalcells, the apex acute or acuminate,the base acute,occasionallyoblique;petioles 2-5 cm long. Inflorescences opposite the leaves, stoutandoccasionallymassive, 1-4 times furcate,0.5-3 cm long, 20-50-flowered, glabrousor finely puberulent with minute papillate trichomes like those of the young leaves;pedicel scars evenly spaced ca. 0.5 mm apart,in branchedinflorescences only appearingafter the fork.Budsglobose, the calyx long-acuminate,petaloid in bud.Pedicels at anthesis stronglydeflexed, 1-Solanum heleonastes is closely related to S. 1.3 cm long, taperingfrom the calyx tube to a slender robustifronsof eastern Peru and adjacentBrazil, dif- base ca. 0.5 mm diam.Flowers with the calyx tube l-feringfromthatspeciesin its longercalyx lobes, smaller 1.5 mm long, the lobes narrowlytriangular,3-3.5 mm leaves, andgenerallymorepubescentyoung stems and long, acuteto long-acuminate,glabrous;corolla white, leaves. Solanumheleonastesis usuallya smalltree(but 1-1.2 cm across, lobed 3/4 of the way to the base, the lobes only slightly reflexed at anthesis, the tips of the occasionally a shrub), and S. robustifronsis a small shrub or treelet growing in the understoryof dense lobes minutelypapillose;anthers 1.5-3.5 x 1-1.5 mm, forestat middleelevations.Solanumheleonastesis one poricidal at the tips, the pores teardropshaped; free of the few species of Solanum sect. Geminatagrow- portion of thefilaments ca. 0.1 mm long, the filament ing in swampforestandis apparentlyquitecommonin tube ca. 0.5 mm long; ovary glabrous;style 5-6 mm long, straight;stigma clavate,minutelypapillose.Fruit areas where it is found. a globose, greenberry,appearinghardandwoody with very little pulp in dry material, ratherhard in living 75. Solanum robustifrons Bitter,Repert. Spec. Nov. collections, 1-1 .1 cm diam.;fruitingpedicels woody, Regni Veg. 11:473. 1912. Type. Peru. Loreto: Peruvia erect, 1-1.3 cm long, ca. 1 mm diam.at the base;calyx orientalis, Maynas in silvis humidis obscuris, lobes in fruitbecoming woody and slightly accrescent, Yurimaguas, Jun 1831, Poeppig 2483 (holotype, W 3-4 mm long. Seeds russetbrownin drymaterial,pale [F neg. 33105]; frag. F). Figs. 128E,F, 133 tan in live plants,ovoid-reniform,ca. 2-3 x 1.5-2 mm, Solanumnzamnorense Bitter, Bot. Jahrb.Syst. 50, Beibl. the surfacesminutelypitted,the seeds all imbeddedin 111: 61. 1913. Type. Ecuador. Loja: In silvis the hard placenta. Chromosome number. n = 12 densis prope flumina Savanilla et Zamora, ca. (voucherKnapp & Mallet 6474). 1000-1300 m, Nov 1888, Lehmann 4941 (holotype, B [destroyed: F neg. 2684]). Solanum deflexiflorumBitter var. diversumJ. F. Macbr., Publ. Field Mus. Nat. Hist., Bot. Ser. 13:203. 1962. Type. Peru. Loreto: Creek Carapisa, above Pongo de Manseriche, 215 m, 12 Dec 1931, Mexia 6261 (holotype, US; isotypes, BH, CAS, F, K, MO, NY, TEX, U).

Fleshy, malodorousshrubs 0.5-5 m tall; young stems and leaves puberulentwith minute, uniseriate papillatetrichomesca. 0.05 mm long, these occasion-

Distribution (Fig. 134). In primaryforests of the upperAmazonbasin in Ecuador,Peru,Bolivia, and adjacentBrazil, 100 to 600 m. Quite common in eastern Peru in lighter areasalong streamsand rivers. Selected specimens examined. ECUADOR. MORONA-SANTIAGO: 7-8 km N of Gualaquiza on rd.

to Indanza, 1450 m, 16 Apr 1985, Harling& Andersson 24177 (AAU, NY). NAPO: Reserva Biol6gica JatunSacha, Rio Napo 8 km E of Misahualli, 450 m, 1?04'S, 77?36'W, 19-28 Mar 1987, Cer6n M. 1004 (MO, NY); Proyecto

FLORA NEOTROPICA

244

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TAXONOMIC TREATMENT

QO

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800 FIG. 134. Distribution of Solanum cucullatum (open circles), S. robuistifrons(solid circles), and S. superbum (solid square).

Hidroelectrico Coca, R bank of Rio Quijos, ca. 10 km S of Reventador, 1450 m, 0?08'S, 77?30'W, 6-10 Oct 1990, Palacios 5991 (QCNE). PASTAZA: Petroleum exploration line Villado 2 of ARCO, 400 m, 1?25'S, 77?20'W, 1-18 Dec 1991, Hurtado 3003 (QCNE); Puyo, Sta. Cecilia, 380 m, 1?30'S, 77?27'W, 1 May 1992, Palacios 10111

(QCNE).

PERU. AMAZONAS: Valle de Rio Santiago, QuebradaCaterpiza, 2-3 km behind village of Caterpiza, 200 m, 3?50'S, 77?40'W, 6 Mar 1980, Tunqui 1004 (MO); Bagua, along Rio Santiago 3-5 km above mouth, 250-300 m, 8-13 Oct 1962, Wurdack2202 (F, K, NY, US, WIS). AYACUCHO:Between Huantaand Rio Apurimac, 750-1000 m, 7, 17 May 1929, Killip & Smith 22784 (NY, US). Cuzco: Environs of Quince Mil, trail to Rio Nusinescatu, 500-600 m, 13?l5'S, 70?45'W,22 Apr 1984, Knapp& Mallet 6370, 6373, 6374 (BH, CUZ, K, US, USM); Limonchayoc, ca. 1 km from Cuzco-Pto. Maldonado rd., ca. 16 km E of Quince Mil, Rio Nusinescatu, 400-500 m, 13?l5'S, 70?40'W, 25-26 Apr 1984, Knapp & Mallet 6393, 6395 (BH, CUZ, K, US, USM); Kosfiipata, Quitacalz6n (Qda. Sta. Alicia), ca. km 163 on LucrePaucartambo-Shintuya rd., 1100-1200 m, 13?07'S, 71?05'W, 11 May 1984, Knapp & Mallet 6428 (BH, CUZ, US, USM); km 234 Lucre-Paucartambo-Shintuya rd., ca. I hrs drive W of Shintuya, ca. 600 m, 12?42'S, 71?20'W, 12 May 1984, Knapp & Mallet 6432 (BH,

245 CUZ, US, USM); along Rio Carb6n, near Atalaya, confluence of Rio Carb6n and Rio Alto Madre de Dios, 500-600 m, 13?00'S, 71?07'W, 15 May 1984, Knapp & Mallet 6453 (BH, CUZ, US, USM). HUANUCO: Codo de Pozuzo, trail to NW behind settlement, 450 m, 9?40'S, 75?25'W, 18 Oct 1982, Foster 9262 (F); Codo de Pozuzo, trail E of settlement, 300 m, 10015'S, 74055'W, 22 Oct 1982, Foster 9384 (F); km 11-12 CarreteraMarginal, S of Pucallpa-Tingo Maria rd. at Bosque von Humboldt, 270 m, 8?45'S, 75001'W, 31 May 1983, Gentry & Jaramillo 41361 (MO); Bosque Nacional de Iparia along the Rio Pachitea ca. 1 km above Tournevista and 20 km above the confluence with the Rio Ucayali, 300-400 m, 26 Oct 1967, Schunke V 2282 (F, G, NY, US). JUNIN: Schunke hacienda above San Ram6n, 1400-1700 m, 8-12 Jun 1929, Killip & Smith 24728 (US); Puerto Yessup, ca. 400 m, 10-12 Jul 1929, Killip & Smith 26382 (US); Cahuapanas on Rio Pichis, ca. 340 m, 20-21 Jul 1929, Killip & Smith 26787 (US); Fondo Romero, Pampatigre above Santa Ana (SE of La Merced), 15001700 m, 11?00'S, 75?10'W, 7 Mar 1985, Stein & Todzia 2341 (MO, NY). LORETO: Yurimaguas, lower Rio Huallaga, ca. 135 m, 23 Aug-7 Sep 1929, Killip & Smith 28032 (F, US). MADRE DE Dios: ParqueNacional Manu, Cocha Cashu station, ca. 400 m, 11052'S, 71?22'W, 24 Sep 1982, Emmons 87 (MO); Parque Nacional Manu, on old oxbow lake of Rio Manu, Cocha Cashu station, 13 Aug 1973, Foster 2608 (F, MO, US); Cocha Cashu station, 350 m, Jul 1978, Foster & Terborgh 6534 (F, US); across river from Cocha Cashu camp, Manu National Park, 340-380 m, 23 Oct 1979, Gentry et al. 27180 (MO); Tambopata, Explorer's Inn Tourist Camp at jct. of Rio La Torre and Rio Tambopata, 270 m, 12019'S, 69040'W, 21 Jul 1985, Gentry et al. 51077 (MO, NY); Aguas Calientes, across and downriver from Shintuya on Rio Alto Madre de Dios, 400-500 m, 12040'S, 71?40'W, 13 May 1984, Knapp & Mallet 6437 (BH, CUZ, US, USM); Manu Park, Cocha Cashu uplands, 400 m, 1?45'S, 71?00'W, 4 Sep 1986, Nuniez5993 (MO, NY); Explorer's Inn near confluence of Rio Tambopata and Ri6 La Torre, 39 km SW of Puerto Maldonado, 12?50'S, 69020'W, 1 Oct 1984, Smith 275 (F, NY, US). PASCO: Rio Pichis valley, Puerto Bermudez, 10 km downriver E of island Reserva Musmanqui, Nuevo Hoboken, Dec 1980, Foster 8140 (F); km 15 of Palcazu rd. (km 73 Villa Rica-Iscozacin-Pto. Mairo) along Rio Palcazu, ca. 380 m, 10?21'S, 75010'W, 17-18 Aug 1984, Knapp & Mallet 6646, 6647 (BH, K, US, USM); km 28 Repartici6-Iscozacin (km 86 Villa RicaIscozacin-Pto, Mairo), Rio La Raya, near Ameusha community of Laguna, ca. 350 m, 10?20'S, 75?10'W, 2223 Aug 1984, Knapp & Mallet 6494 (BH, US, USM); Knapp & Mallet 6650 (BH, K, NY, USM); Iscozacin, along Rio Iscozacin, tributaryof Rio Palcazu, ca. 320 m, 10012'S, 75013'W, 27 Aug 1984, Knapp & Mallet 6659, 6660 (BH, K, US, USM); 5 km SE of Oxapampa, 1850 m, 10036'S, 75?23'W, 9-11 Dec 1982,D. N. Smith 2906 (F, MO, NY). SAN MARTiN: Tingo Maria, 2300 ft, 30 Oct 1949, 19 Feb 1950, Allard 21607 (US); Zepelacio

246 near Moyobamba, ca. 1100 m, May 1934, Klug 3632 (F, K, US); Cufiumbuque to Sisas, ca. 1 hr driving time from Cufiumbuque (1/3 of way to Sisas), ca. 850 m, 6?35'S, 76?39'W, 5 Jun 1984, Knappet at. 6474, 6477 (BH, K, US, USM); km 27-30 Tarapoto-Yurimaguas rd., 650-750 m, 6?25'S, 76015'W, 9 Jun 1984, Knapp & Mallet 6494 (BH, US, USM); km 21-22 of TarapotoYurimaguasrd., just N of pass, ca. 1000 m, Knappet al. 7882 (MO, USM); Mariscal Caceres, Tocache Nuevo, swamp edge on rd. to Almendras, 400 m, 7 Aug 1969, Schunke V 3306 (F, NY, US); Tocache Nuevo, NE of Instituto Agropecuario, 20 Apr 1970, Schunke V 3948 (F, G, GH, NY); San Juan de Pacayzapa, E of the bridge on rd. to Moyobamba, 900 m, 7 Apr 1973, Schunke V 5866 (IBE, NY); rd. to Roque, 8 km from San Juan de Pacayzapa, 800-900 m, 4 May 1973, Schunke V 6172 (IBE, NY); Schunke V 6176 (IBE, NY); Rio Mayo, near Tarapoto, Jul-Aug 1856, Spruce s.n. (K); Rio Mayo, Aug 1856, Sprluce s.n. (K); San Roque, 1300-1500 m, 6 Jan 1930, Williams 6952 (F), 13 Jan 1930, Williams 7352 (F). UCAYALI: Banks of Ucayali, 100 S, 1923, Tessmann 3107 (G). BOLIVIA. BENI: Prov. Yacuma, Isla del BosqueEstacion Biol6gica del Beni, plot 7, 220 m, 14?51'S, 66?20'W, 29 Jul 1996, Aymardet al. 11388 (MO). LA PAZ: Prov. Nor Yungas, Serrania de Bella Vista, 16 km N of Carrqasco (37 km N of Caranavi) on rd. to Palos Blancos, 1500 m, 15?35'S, 67?34'W, 31 Oct 1984,

FLORA NEOTROPICA

over. Plants that topple when young may persist as rhizomatousindividualsin these populations.The two formsof S. robustifronsdifferonly in theirstature;they areotherwiseidentical. Solanumdeflexiflorum(S. deflexiflorumspecies group)is a species of high elevations in Colombiaand is not even closely relatedor superficiallysimilarto S. robustifrons.It is unclearwhy MacbridenamedMexia 6261 a varietyof S. deflexiflorum,but sheetsof the type collectionareannotated"speciesnearS.deflexiflorum" by Morton.

76. Solanum superbum S. Knapp, Brittonia44: 62. 1992. Type. Bolivia. Cochabamba:Chapare,at arroyo side at base of colina Limbo, 2100 m, 26 Nov 1966, R.E Steinbach 541 (holotype, F; isotypes, MO-n.v., NY). Fig. 135

Erecttree 5 m tall;young stems thick andsomewhat fleshy, densely pubescentwith soft dendritictrichomes, these l-1.5 mm long, dryinggolden-red;bark of older stems pale reddish-gray,the trichomespersistent.Sympodialunitsdifoliate,geminate.Leavesfleshy, elliptic-ovoid, widest at orjust below the middle, pubescent with simple, rarely furcate, uniseriate, 2-9Solomon& Nee 12651 (MO, NY). celled trichomes adaxially, the trichomes 1-1.5 mm BRAZIL. AMAZONAS: Near mouth of Rio Embira, long, evenly spaced ca. 1 mm apartover the lamina tributaryof Rio Tarauaca,7?30'S, 70?15'W, 26 Jan 1933, surface, denser along the veins, pubescent with soft 5006 (F, G, MO, NY, US); km 1-4 of rd. B6ca dendritictrichomesabaxially,the trichomesca. 1 mm Krlukoff do Acre to Rio Branco, 15 Sep 1966, Pranceet al. 2299 long, denseralong the veins; majorleaves 17-30 x 12(NY, US). 16 cm, with 7-9 pairsof parallelmain lateralveins, the apex acute,the base attenuate,ocasionallyoblique;petLocal names. Peru. Huanuco: tohuecillo; San ioles ca. 2 cm long, winged from the decurrentleaf Martin:tintunasacha. bases; minor leaves differing from the majorsonly in Solanum robustifronswas originally placed in size, 10- 15 x 6-8 cm, the apex acute, the base attenusect. Polvbotryon(correctly Pteroidea) by Bitter, who ate; petioles ca. 1.5 cm long, winged from the decurthought it was closely related to S. anceps and its allies rent leaf bases. Inflorescences opposite the leaves or (Knapp& Helgason, 1997). Solanumrobustifronsdif- intemodal, simple, fleshy, 1.5-3 cm long, 5-1 0-flowfers from membersof that section in its difoliate sym- ered, densely pubescentwith soft dendritictrichomes podial units, leaf-opposedinflorescences,andglobose like those of the stems;pedicel scars closely spaced fruits. The superficial similarity lies in the fleshiness but not overlapping.Buds globose with the corolla inof the plant, most probably a result of its shady, moist cludedin the calyx tubewhen young, laterelliptic with habitat.SolanumrobustiJronsis most closely relatedto the corolla exserted from the calyx tube. Pedicels at S. heleonastesof the swampsalong the Rio Paranaand anthesis thick and fleshy, somewhat deflexed, 1.7-2 easternBolivia. It shareswith thatspecies long-acumi- cm long, ca. 2.5 mm diam.atthe apex,ca. 1.5 mm diam. nate calyx lobes, deflexed floweringpedicels anderect at the base, denselypubescentwith dendritictrichomes fruiting pedicels, and small flowers comparedto the like those of the inflorescence axis. Flowers with the other membersof the group. calyx tube broadlyconical, 3-4 mm long, the lobes irSome collections of Solanumrobustifronshave regular in shape but mostly deltate, 3-4 mm long, portionsof a fleshyrhizomatousrootstockattached.The densely pubescentwith loose dendritictrichomeslike rhizomatoushabit is unusualin sect. Geminata,and I the rest of the inflorescence;corolla white, thick and have seen several populationsof this species contain- fleshy (fide Steinbachthe lobes 2 mm thick), 2-2.5 cm ing individuals of both rhizomatousand erect habits. diam., lobed 2/3 of the way to the base, the lobes perYoungplantshaveweak,fleshy stems,andeasily topple pendicularto the anthercone, densely pubescentwith

TAXONOMIC TREATMENT

247

FIG. 135. Solanum superbum S. Knapp. (Reproduced with permission from Brittonia 44: 63, fig. 2. 1992.)

dendritictrichomesabaxially,denselypapillatewith an occasional dendritictrichomeadaxially;anthers6-7 x 1.5-2 mm, thickenedandverrucoseabaxially,poricidal at the tips, the pores teardropshaped; free portion of the filamentsca. 1 mm long, the filamenttubeca. 1 mm long, glabrous;ovary glabrousor with a few dendritic trichomes;style 0.8-1 cm long, clavate;stigmabilobed and minutely papillose. Fruit and seeds not known. Chromosomenumber not known. Distribution (Fig. 134). Only known from the type locality in the little collected Chapare area of Cochabamba,Bolivia. Solanum superbum is closely related to S. abitaguense and S. cucullatum. It differs from those species in its dense indumentof dendritictrichomesbut like them is unusual in sect. Geminata in possessing largefleshy flowers. Althoughthe fruitsarenot known for S. superbum, I expect them to be large (to 4 cm diam.) like those of its close relatives.

The possession of branched trichomes on all partsof S. superbumwould have excluded it fromsect. Geminatain previous classifications of the group(see Seithe, 1962; D'Arcy, 1974). The polarityof this character is in doubt for all of Solanum, as many groups and species previously thoughtto have only simple or branched trichomes now are found to possess both types (see Morphology).

XI. Solanum narcoticosmum species group (S. dasyneuron, S. foetens, S. narcoticosmum). Fig. 136A Shrubs or small trees, usually of cloud forests; young stems and leaves glabrous or pubescent with simple, uniseriatetrichomes.Sympodialunitsunifoliate or difoliate and geminate, if geminate the minor leaves differing from the majors only in size. Leaves elliptic, ovate or obovate, quite fleshy, drying black,

248

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glabrousorpubescentwith simple,uniseriatetrichomes along the veins and extending to the lamina, the apex andbase various,in Solanumfoetensthe leaves sessile. Inflorescences opposite the leaves, simple or furcate, glabrous or pubescent;pedicel scars closely spaced, occasionally corky. Bludsovoid to globose, strongly exsertedfromthe calyx tube.Pedicels at anthesisusually somewhatdeflexed.Flowerswhite,fleshy,the lobes planarat anthesis.Fruitgreenandhardatmaturity;fruiting /pedicels deflexed, woody, the calyx lobes enlarg-

ing and becoming woody in fruit. Seeds ovoid-reniform, pale tan (only known from S.foetens). Distribution. Montanecloud forests, Centraland northernSouthAmerica. These threespecies are all very similarmorphologically andalthoughthey may not representa monophyleticgroup,areeasy to distinguishfromothermembers of sect. Geminata.The fleshy leaves thatusually dry black are distinctive, as are the fleshy flowers and enlargedcalyx lobes in fruit.

Key to the species of the Solanum nar-coticosmumspecies group 1. Sympodial units unifoliate; leaves sessile, with uniseriate trichomes along the veins and on the lamina 78. S. fbetens beneath. Andean Venezuela ............................................................... 1. Sympodial units difoliate, geminate; leaves petiolate, the trichomes, if present, only along the veins. 2. Leaves glabrous beneath;plants not drying black. Chiapas, Mexico and Guatemala .............79. S. narcoticosonuon 2. Leaves pubescent along the veins beneath, the trichomes uniseriate, simple; plants drying blackish. Volcan Tacana, border of Chiapas, Mexico and Guatemala................................................ 77. S. d(isvneuron

77. Solanum dasyneuron S. Knapp, Ann. Missouri Bot. Gard.72: 563. 1985. Type. Mexico. Chiapas: VolcanTacana,Chiquihuite,2800 m, 27 Mar 1939, Matluda2849 (holotype, F; isotypes, F, MEXU, MO, NY). Fig. 137 Shrlubs2-3 m tall; young stems and leaves glabrousor occasionally minutelyrusty-papillate,drying black and shiny;barkof older stems white, exfoliating in small transverse pieces. Symtipodialunits difoliate,

geminate. Leaves obovate to narrowly obovate, glabrous above, the midrib depressed, the main lateral veins raised, densely pubescent on the primary and secondary veins beneath with uniseriate trichomes 0.25-0.5 mm long, trichomes restrictedto the veins; majorleaves 12.5-17.3 x 5-6.5 cm, with 11-15 pairs of mainlateralveins, the apex acute,the base attenuate, not decurrenton the petiole; petioles 1.5-1.9 cm long; minorleaves differingfromthe majorsonly in size, 2.2-7 x 1.6-2.9 cm, apex acute,the base attenuate;petioles 0.6-1.1 cm long. Inflorescences opposite the leaves, simple or occasionally once-furcate,1-5 cm long, 1020-flowered, glabrous and shining; pedicel scars closely spaced and overlapping, beginning 4-6 mm from the base of the inflorescence.Buds ovoid, fleshy, pointedandconstrictedapically,only partiallyexserted fromthe calyx. Pedicels at anthesis deflexed or somewhat erect, 1.1--1.5 cm long, robust and somewhat fleshy, 0.75-1 mnmdiam. at the base, taperingimperceptiblyto the calyx lobes. Flowers with the calyx tube 1.5-2.5 mm long, conical, the lobes deltoid, apiculate, 1-2 mm long, papillose within, glabrouswithout, the

marginsof the lobes whiteandthickened;corollawhite, thick and fleshy, ca. 1.5 cm diam., lobed nearly to the base, the lobes planarat anthesis,the tips and margins of the lobes minutelypapillose, the tips cucullate;anthers3.5-4 x 1-1.5 mm, poricidalat the tips, the pores teardropshaped;free portionof thefilaments ca. 0.25 m long, the filamenttubeca. 0.5 mm long, dryingblack; ovary glabrous;style ca. 7 mm long, straight;stigmza largeandcapitate,ca. 1 mm diam.,minutelypapillose. Fruit a globose berry,8 mm diam. (immature),green; fruiting pedicels deflexed, woody, 1-1.5 mm diam. at the base; calyx lobes enlarging and becoming woody in fruit,ca. 4 mm long. Seeds not known. Chroniosoinie numbernot known. Distribution (Fig. 138). Endemic to the upper slopes of VolcanTacana,on the borderof Mexico and Guatemala,in wet forest from 2500 to 3000 m. Specimens examined. GUATEMALA. SAN MARCOS:Between La Vega ridge along Rio Vega and NE slopes of VolcanTacana,to 3 mi from GuatemalaMexico border,in vic. of San Rafael,2500-3000m. 20 Feb 1940, Stev:ermnark 36198 (F). Solanum dasyneuronis most closely relatedto S. narcoticosmumof middle-to high-elevationChiapas andGuatemala,anddiffersfromthatspeciesin its larger flowers and its dense pubescence along the veins on the leaf undersides.Solanumni dasYneuronis perhaps derived from isolated populations of the more widespread S. narcoticosmum.It is of extremely limited distributionandhas not been collected recently.

FLORA NEOTROPICA

250

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FIG. 137. Solanum dasyneuron S. Knapp. (Reproduced with permission from Annals of the Missouri Botanical Garden 72: 562, fig. 4. 1985.)

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TAXONOMIC TREATMENT

251

FIG. 139. Solanumfoetens S. Knapp. (Reproduced with permission from Brittonia 38: 276, fig. 3. 1986.)

78. Solanum foetens Pittierex S. Knapp,Brittonia38: 275. 1986.Type.Venezuela.Merida:ca. 1 km SW of Chachopo on rd. to Merida, ca. 2400 m, 8o56N, 62?41'W,21 Oct 1984,Knapp& Mallet 6799 (holotype, MY; isotypes,BH, K, US, VEN). Fig. 139 SolanumsilvicolumC.V. Mortonin Pittier,Cat. Fl. Venez. 2: 382. 1947. Nom. nud. Venezuela. Merida:Paramode la Sal, 2600 m, 21 Sep 1921, Jahn 532 (VEN). The specimen bears a typed annotationlabelso it is possiblethatMortonnever saw the specimenin question.

units unifoliate. Leaves obovate, sessile, widest just distal to the middle, glabrousor sparselypubescenton the veins abovewithuniseriatetrichomes,thetrichomes 0.1-0.7 mm long, densely pubescentbeneathwith soft uniseriatetrichomes 0.1-0.8 mm long, denser on the veins, the trichomesreddish-brownon dry specimens, white on living plants,blades 10-23 x 3.5-10 cm, with 7-10 pairsof mainlateralveins, these impressedabove, prominentanddenselypubescentbeneath,theapexacute, the base attenuateand occasionally somewhatauriculate,wingedontothepetioleandultimatelyontothestem; Shrubs or small trees 1-4 m tall; young stems petioles consisting only of the winged leaf base.Inflopubescent with scatteredto dense, soft uniseriate tri- rescences opposite the leaves, simple, 0.2-2 cm long, chomes, stronglywinged fromthe decurrentleafbases; 10-20-flowered, sparsely to densely pubescent with older stems glabrate,often only partiallyso, remaining the same soft uniseriatetrichomesas those of the young strongly winged even on the trunks;barkof the older stemsandleaves;pedicelscars corkyandraised,evenly stems remaining bright green, the wings becoming spaced ca. 0.2 mm apart.Buds globose, soon becombrittleand breakingoff on very old stems. Sympodial ing ovoid, laterlong-ellipsoid,the corollasoon exserted

2 52

from the calyx lobes. Pedicels at anthesis slightly deflexed, 0.9-1 cm long, taperingfrom the calyx tube to a slender base ca. 0.5 mm diam., pubescent with the same uniseriate trichomes as those of the rest of the inflorescence. Flowers with the calyx tube short, 1-2 mm long, strongly ribbed in dry specimens from the lobe vasculature,the lobes deltoid to long-acuminatetriangular,1.5-2 mm long, the entire calyx pubescent with soft uniseriatetrichomes like those of the rest of the inflorescence, these denser along the veins of the lobes; corolla white, 1.5-1.7 cm diam., lobed 7/8 of the way to the base, the lobes planarat anthesis, the lobes keeled abaxially,with uniseriatetrichomesca. 0.2 mm long along the keel, minutely papillose along the tips and marginsof the lobes; anthers ca. 3.5 x 1 mm, poricidal at the tips, the pores teardropshaped; free portionof thefilaments ca. 0.1 mm long, the filament tubeca. 0.6 mm long;ovaryglabrous;style straight,ca. 7 mnmlong; stigma bilobed, minutely papillose, black in dry specimens, green in live plants.Fruit a globose, green berry,becoming a dirty greenish-yellow when ripe,ca. 1.2 cm diam.;fruitingpedicels woody, erector deflexed, 1.5-1.8 cm long, ca. 1 mm diam. at the base. Seeds pale tan, ovoid-reniform,ca. 3 x 2 mm, the surfaces minutely pitted. Chromosomenumber: n = 12 (vouchers Knapp & Mallet 6788, 6799). Distribution (Fig. 138).IntheVenezuelanAndes in the statesof MeridaandTrujillo,in secondarythickets from 2000 to 2800 m. Specimens examined. VENEZUELA. MNRIDA: Quebrada El Volcan, rd. between Redoma and Pie de la Cuesta, toward Guirigay, 13 Feb 1975, Badillo 6804 (MY): between Redoma and Pie de la Cuesta toward Guirigay, 2400-2500 m, 26 Feb 1975, Badillo 6904, 6906, 6911, 6920, 6937, 6940 (MY); Chachopo, ca. 2000 mn,16 Aug 1972, Benitez de Rojas 1478 (MY); 1 km SW of Chachopo on rd. to M&rida,ca. 2400 m, 8?56'N, 62041'W, 21 Oct 1984, Knapp & Mallet 6801, 6803 (BH, K, MY, US, VEN); s.loc., 1 Mar 1974, Morales Mendez 248 (MY); Chachopo, 25 Jan 1929, Pittier 13291 (K, NY, US, VEN); Piedra Gorda, above Timotes toward El Aguila, 2250 m, 3 Dec 1975, Ruiz Teran & Duigarte 12907 (K, MY); Dtto. Rangel, basin of Rio Aracay, Above Las Piedras, 2550-2700 m, 16 Dec 1972, Ruiz Teran et al. 8261 (K); between Timotes and Paramito, 2285-3500 m, 24 Mar 1944, Steyermark 55696 (MY, VEN); Chachopo, 4 km NE, near small waterfall, 2300 m, 23 Jun 1981, Stevermark & Manara 125203 (MY, VEN). TRUJILLO: Near Visun (Las Mesitas), ca. 2800 m, Aug 1958, Ar-istegutieta3672 (MY, NY); Paramo de la Nariz, between La Pefia and Las Palmas, 2400 m, 1 Dec 1979, Benitez de Rojas 2682 (MY); paramo de Agua de Obispo, 2500 m, 24 Sep 1922, Jahn 1184 (VEN); old rd. from Bocon6 to Trujillo, ca. 51 km W of Trujillo, below summit, 2200-2250 m, 9?21'N, 70019'W, 20 Oct

FLORA NEOTROPICA 1984, Knapp & Mallet 6780 (BH, K, MY, US, VEN):

windsweptsummitof old rd. from Bocon6 to Trljillo, ca. 54 km W of Trujillo,ca. 2350 m, 9?21'N,70A19'W, 20 Oct 1984, Knapp & Mallet 6784, 6786, 6788 (BH,

K, MY,US, VEN); El Lamedero,Tufiame-LasMesitas rd., 2400 m, 19 Sep 1972, Ruiz Terdn & Ritiz Palacios 7566 (MY);betweenLa PefiaandAgua de Obispo,2228 km from Carache, 2400-2500 m, 1 Mar 1971. Stevermark 104982 (US, VEN).

Local names. Venezuela. Merida:borrachero. zamurito. Solanumfoetensis a distinctiveplantandnot easily confusedwith any otherSolanum.Characterswhich immediatelydistinguishS.foetens from its close relatives, S. narcoticosmum and S. dasyneuron,

are its

sessile leaves with denselypubescentundersides,strikingly winged stems, unifoliate nodes, and its deeply lobedplanarcorollas.Intheparamoatthe summitof the old Bocon6-Trujillord.,S.foetens grows with another species of sect. Geminata,S. validinerviumn (S. amblophyllumspeciesgroup),withwhich it is somewhatsimilar morphologically.The two species differ in microhabitat,withS.foetensgrowingin wetter,moreprotected microsites.Flowerodoralso differsbetweenthe species: S.foetens has completely odorless flowers in all sites, andthe S. validiner-vium has flowers with a spicy odor. In the aforementionedsite, the two species bloom at the same time, but no intermediateswere found.

79. Solanum narcoticosmumBitter,Repert.Spec.Nov. RegniVeg. 18:55. 1922.Type.Guatemala.Huehuetenango:Berghangab Jacaltenango,9 Apr 1896,C. & E. Seler (Plant mexic et centr. arneric.) 2627

(holotype, B; isotype, DS).

Fig. 140

Solanim fontiumnStandl. & Steyerm., Field Mus. Nat. Hist., Bot. Ser. 23:234. 1947. Type. Guatemala. Quezaltenango: Fuentes Georginas, western slope of Volcan de Zunil, 2850 m, 4 Mar 1939, Standlev 67473 (holotype, F).

Shrubs or weak trees, 1.5-6 m tall; stems glabrous,darkabove,becominglighterandlenticellatewith age; new growth glabrous.Sympodialunits difoliate, geminate.Leavesquitevariablein size, ellipticto ovate, widest at orjust proximalto the middle, glabrous;major leaves 9.5-22.5 x 2.5-7.8 cm, with ca. 13 pairs of main lateralveins raisedabove, prominentbelow, reddish-brownwhen dry,the apex acute, the base attenuate;petioles 1-3.5 cm long;minorleaves differingfrom the majorsonly in size, 2.2-9 x 0.8-3.5 cm, the apex acute,the base attenuate;petioles 0.5-1.5 cm long.Inflorescences opposite the leaves, usually furcate, 1.54 cm long, 10-50-flowered, completely glabrous,drying black;pedicel scars closely spaced,not overlapping,

253

TAXONOMIC TREATMENT _

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PURPUYSIANAE PLANTS M1!XZCANAE FIG. 140. Solanum narcoticosmum Bitter. Mexico. Purpus 7318 Other sheets of this number are variously (BM). mixed with S. aphyodendron, S. rovirosanum, and S. nudum. See specimens cited for those taxa.

FLORA NEOTROPICA

254

in branchedinflorescences flowers producedonly afterthe division.Budsglobose, the corollaearlyexserted from the calyx. Pedicels at anthesis 1.1-1.3 cm long, taperingfrom the calyx tube to a slender base ca. 0.5 mm diam.Flowers with the calyxtubeca. 1.5 mm long, the lobes 1-1.5 mm long, deltoid, apiculate, the tips minutely papillose; corolla white, ca. 1.4 cm diam., lobed nearly to the base, the lobes planarat anthesis, tips of the lobes minutely papillose; anthers ca. 3.5 x 1-1.5 mm, poricidal at the tips, the pores teardrop shaped;freeportionof thefilamentsca. 1 mm long, the filamenttube ca. 0.5 mm long;ovaryglabrous;style ca. 7.5 mm long, straight;stigma capitate,minutelywhite papillose. Fruit a globose, green (?) berry,0.9-1.2 cm diam. (immature);fruiting pedicels woody, erect to slightly deflexed, 2-2.5 cm long, ca. 1 mm diam.at the base; calyx lobes becoming slightly accrescentin fruit, to 3 mm long, woody. Seeds not known. Chromosome not known. numinber

Madre, 1800-2400 m, 10-18 Dec 1963, Williams et al. 25764 (F); Volcan Tajumulco, 10 km W of San Marcos, Sierra Madre, 2400-2700 m, 3 Jan 1965, Williams et al.

27139 (F, NY, US). Local names. Guatemala. San Marcos: hediondilla. Solanumnarcoticosmumis most closely related to S. dasyneuron,also of high-elevationChiapasand Guatemala.Itdiffersfromthatspeciesin its smallerflowers, completelyglabrousleaves andnew growth,andin its somewhatlower-elevationhabitat.Solanumnarcoticosmumis quitevariableas to leaf shape;those collections with narrowerleaves have been calledS.fiontium. The specificepithetcomes fromthenarcoticodor of the flowers ("narcoticeolens"). In the description, Bittercites the Seler collection as the sourceof this information,but the labels on both the holotype and the isotype bearno mention of any flower odor.

Distribution (Fig. 138). In wet forests (with XII. Solanum nigricans species group(S. callianthuim, Cheiranthodendron andAlnus)athighelevations(2000S. cornifolium, S. maturecalvans, S nigricans,S. oclhro2300 m) in southemMexico (Chiapas)andnorthwestphyllum, S. platycypellon, S. quebradense). em Guatemala,often common where it occurs. Fig. 136B-D Specimens examined. MEXICO. CHIAPAS: Shrubs or small trees; young stems and leaves Along hwy. 119 at Chital,9 mi SW of Ocosingo, 1190 m, 7 Jul 1977, Croat40379 (MO); VolcanTacana,be- usuallypubescentwithusuallyarachnoid,butoccasiontween Talquianand the summit, 1600-2400 m, 19 Jun ally dendritic or simple trichomes. Sympodial units 1985, Martinez S. et al. 13217 (MEXU, NY); Fraylesa, unifoliate, difoliate or trifoliate,or difoliate geminate Siltepec, 11 Mar 1945, Matuda 5255 (F); Cerro del (Solanummaturecalvans,S. nigricans). Leaves ellipBoquer6n, Jun 1914, Purpus 7318 (BM, MO, mixed tic to ovate, usually pubescentabaxially,occasionally collection with S. aphyodendron). glabrous(S.platycypellon),the trichomeslike those of GUATEMALA.CHIMALTENANGO:Region of Las the young stems. Inflorescences opposite the leaves, Calderas,1800-2100 m, 22 Nov 1938, Standley57782 usually3-20-flowered;pedicelscarsclosely spacedand (F); slopesof Volcande Acatenango,aboveLasCalderas, occasionally almost overlapping.Pedicels at anthesis 2400-2700 m, 3 Jan 1939, Standley 61813 (F, US). deflexed,usuallythickandfleshy.Flowers white, thick HUEHUETENANGO: Above San Juan Ixcoy, Sierra de los 2400 m, 4 Aug 1942,Steyerinark50039 and fleshy, the corolla lobes planarat anthesis;ovary' Cuchumatanes, (F. NY). QUEZALTENANGO:Fuentes Georginas, W slope glabrous or pubescent.Fruit green and hard at matuVolcande Zunil,2850 m, 4 Mar 1939,Standley67409, rity, glabrous or pubescent (S. cornifolium);fruiting 67492(F); regionof Azufral,N slopeof Volcande Zunil, pedicels erect or somewhat deflexed, woody. Seeds 2300-2500 m, 3 Feb 1941,Standley85735 (F); Fuentes ovoid-reniform,usually darkreddish-brown. Georginas,slopes of Volcande Zunil, 2300-2500 m, 3 Distribution. Montane forests, from Mexico to Feb 1941, Standley85871, 85928, 85935 (F, US). SAN Hondurasand the Andes from Venezuelato Bolivia. MARCOS: 10 mi S of San Marcosalong rd. from San Rafael, 2100 m, 13 Jul 1977, Croat40993 (BM, MO); The matted, floccose trichomes typical of the along rd. above BarrancoEminencia,2700 m, 14 Mar majority of the taxa in this group are absent from 1939, Standley 68502 (F); Barranco Eminencia, rd. Solanumcallianthum,with dendritic(or occasionally betweenSan MarcosandSanRafaelPie de la Cuesta,in upper part of the barrancobetween Finca Lucha and simple?,Ecuador)trichomes,andS. quebradense,with BuenaVista,2500-2700 m, 6 Feb 1941,Standlei'86356 simple trichomes. These taxa however do share the (F, US); upper S-facing slope of Volcan Tajumulco, ovoid-reniformseeds of the rest of the groupandhave between Las Canojasand top of ridge, 7 mi from San similar overall flower morphology. The inclusion of Sebastian, 3300-3900 m, 16 Feb 1940, Steyermark them here is thus provisional.Solanumlaevigatumof 35837 (F); Aldea Fraternidad,between San Rafael Pie the S. arboreumspeciesgroupis also superficiallysimide la Cuestaand Palo Gordo,W-facingslope of Sierra lar to these species and may belong here.

TAXONOMIC TREATMENT

2 55

Key to species of the Solanum nigricans species group 1. Leaves densely pubescent abaxially, the trichomes simple or dendritic. 2. Trichomes of leaf undersides loose, 0.5-1 mm long; the leaves concolorous; sympodial units unifoliate. 3. Trichomes of leaves dendritic or occasionally simple and uniseriate (Ecuador); leaves rugose ......................................................... 80. S. callianthum 3. Trichomes of leaves simple, uniseriate; leaves not rugose...................................... 86. S. quebradense 2. Trichomes of leaf undersides arachnoid, felty, less than 0.5 mm long; the leaves strongly bicolorous; sympodial units di- or tri-foliate. S Peru to Bolivia .............................. 84. S. ochrophyllurn 1. Leaves variously pubescent or glabrous abaxially, the trichomes never completely covering the abaxial leaf surfaces, always arachnoid and felty. 4. Sympodial units unifoliate. 5. Calyx tube a flat dish, the lobes mere undulations on the rim; mature leaves glabrous. S Bolivia and Argentina...................................................... 85. S. platycypellon 5. Calyx tube conical, the lobes deltate, 1-3 mm long; mature leaves with patches of arachnoid trichomes abaxially in the axils of the main lateral veins. Colombia and Venezuela ............................................................ 81. S. cornifoliun 4. Sympodial units difoliate, geminate. 6. Corolla 1-1.3 cm diam.; pedicels at anthesis 1.1-2.3 cm; fruit usually apically pointed, green or greenish-yellow when ripe. Pine-oak forests, Mexico to Costa Rica .................... 83. S. nigricans 6. Corolla 1.5-2.2 cm diam.; pedicels at anthesis 0.5-1.5 cm; fruit globose, black tinted when 82. S. inaturecalvans ripe. Peru to Bolivia ......................................................

80. Solanum callianthum C. V. Morton, Contr.U.S. Natl. Herb. 29(1): 43. 1944. Type. Colombia. Cundinamarca:CordilleraOriental, Boquer6n de Chipaque, 3100-3200 m, 31 Dec 1939, Cuatrecasas 7898 (holotype,US [US-1795321 ]; isotypes, COL-n.v., F [F neg. 49457], WIS [Nee neg. 6 Apr 1971]). Fig. 141

tubebroadlyconical, 1-1.5 mm long, the lobes deltate, 1.5-2 mm long, somewhatirregularin shape, sparsely to denselypubescentwith dendriticor simpletrichomes; corolla white to pinkish-white,fleshy, 1.5-2 cm diam., lobed nearly to the base, the lobes planarat anthesis, abaxialdistallobe surfacepubescentwith dendritictrichomes 0.25-0.5 mm long; anthers3-4 mm long, 1-2 mm wide,ponicidalatthetips,theporesteardropshaped; free portionof thefilamentsca. 0.25 mm long, the filament tube ca. 0.5 mm long, glabrous;ovary glabrous to denselypubescentwith dendritictrichomes;style 510 mm long, glabrousor pubescent, correspondingto ovarypubescence;stigmabilobed,the surfaceminutely papillose.Fruita globose,greenberry,1.5-1.7 cm diam., occasionally apiculate,the pericarphard and woody; fruiting pedicels woody, erect, 1.5-2 cm long, 1.5-3 mm diam. at the base. Seeds darkbrown, ovoid-reniform, 5-5.5 mm long, 3-4 mm wide, the surfacesminutely pitted. Chromosomenumberunknown.

Treesor shrubs, 1.5-10 m tall; young stems and leaves sparselyto densely pubescentwith lax dendritic trichomes 0.5-1 mm long, not soon glabrescent;the stemsthick anderect;barkof older stems pale grayishbrown. Sympodial units unifoliate or occasionally difoliate, geminate.Leaves elliptic to ovate, widest at orjust below the middle, thick andperhapssomewhat fleshy, with 6-10 pairs of main lateralveins, glabrous or sparsely pubescent with dendritic (or simple uniseriate) trichomes adaxially, sparsely to densely pubescent with dendritic (or simple uniseriate) trichomes abaxially,the trichomesdenseralongthe veins, the lamina 6-16 cm long, 3.5-8.5 cm wide, the apex Distribution (Fig. 144). In cloud forest and acute,thebase acuteto attenuate;petiole 1.8-3 cm long; paramoedges in the CordilleraOrientalof Colombia, minor leaves if present not differing from the majors. departmentsof Cundinamarca, Boyaca, andSantander, Inflorescences opposite the leaves, simple, 0.5-3 cm 2000 to 3500 m (but see discussion). long, 5-10-flowered, densely pubescent with lax triSpecimensexamined.COLOMBIA.S.loc.,Muttis chomes like those of the stems. Pedicel scars closely 3568(US).BOYACA: Valledala Uvita,7 Sep 1938,Cuatrespaced but not overlapping.Buds globose, the corolla casas 1147 (US); SierraNevada del Cocuy, La Cueva near strongly exserted from the calyx tube.Pedicels at an- Finca la Pajarita,3560 m, 18 Mar 1979, Kieft & Becerra thesis deflexed, thick and fleshy, 0.7-1.2 cm long, C. 130 (NY). CUNDINAMARCA: Guasca to Gacheta rd., sparselyto densely pubescentwith trichomeslike those Jan 1920, Ariste-Joseph s.n. (US); camino de Gacheta, of the rest of the inflorescence.Flowers with the calyx 1920, Ariste-Joseph A537 (US); 2 km E of Bogota on

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257

TAXONOMIC TREATMENT

Pasto, 2800 m, Triana 2248 (holotype, W; isord. to Choachi, 2900 m, 8 Mar 1972, Barclay et al. type, B [destroyed: F neg. 2678]). 3232 (US); Macizo de Bogota, Quebradadel Rosal, mutisiiC. V. Morton,Contr.U.S. Natl. Herb.29: Solanum 3000 m, 29 Jun 1939, Cuatrecasas 5695 (F, US); Sibate, 45. 1944. Type. Colombia. s.loc., Mutis 1993 (holo2700-2800 m, 13-15 Oct 1917, Pennell 2429 (US); type, US [US-1563674]). Simijaca, ca. 2000 m, May 1930, Prez Arbeldez254 (US); CerroMonserrate,11000 ft, 12 Jan 1969, PlowShrubs or small trees, 1-5 m tall; young stems man 2204 (COL, GHi,K, S, US); Macizo de Bogota, andleaves denselypubescentwith beige to white arachRetiro, 2600-2700 m, 7 May 1946, Schultes 7224 (F, NY); Chiquinquira, 2500 m, 17 Feb 1950, von1 noid trichomes,appearingfelty,soon glabrescent;stems lightly winged between the nodes, erect;barkof older Sneidern 5824 (NY. S); Bogota, 2700 m, 1851-57, stems pale brownish-gray.Sympodialunitsunifoliate, Vic. Vetas, 3100-3250 Triana 2219 (NY). SANTANDER: m, 16-20 Jan 1927, Killip & Smith 17249 (A, GH, NY, occasionally difoliate and geminate on some young US); W slope of paramode Santurban,towardTona, shoots.Leaveselliptic,somewhatfleshy,with 7-9 pairs 2200-2800 mn,18 Feb 1927, Killip & Smith 19524 (A, of main lateralveins, these impressedabove,brownish GH, NY,US). beneath,glabrousadaxially,glabrousor with a few scatteredpatchesof arachnoidpubescenceabaxially,these Solanurm callianthum is unusual (with S. qluebradense)in the S. nigricans species group in its felty patches denser along the veins, the lamina 6-13 possession of loose ratherthanarachnoidtrichomes.It cm long, 3-7 cm wide, the apex acute, the base acute; shares fleshy flowers and erect frutingpedicels with petiole 0.7-1.2 cm long; minorleaves if presentdifferthe other membersof the S. nigricans species group. ing from the majors only in size, 4-8 cm long, 2-4.5 Threecollections from eastem Ecuadorarepro- cm wide. Inflorescences opposite the leaves, simple, visionallyplacedinSolanumcallianthum,althoughthey 0.5-2.5 cm long, 3-7-flowered, sparsely to densely possess only simple, uniseriatetrichomes ratherthan arachnoid-pubescent;pedicelscars closely spacedbut the more typical dendritictrichomesfound in Colom- not overlapping.Buds globose, later somewhat ellipbian material:Ecuador.Napo: Salcedo-Napo rd., km soid, stronglyexsertedfromthe calyx tube.Pedicels at 55 Rio Tena, 2995 m, 4 Feb 1977, Boeke 892 (BM, anthesis thick and somewhat fleshy, deflexed, 0.5-1 NY): SE of El Paly6n de San Francisco on slopes of cm long, ca. 0.5 mm diam., sparselyarachnoid-pubesCerro Mirador,3300-3700 m, 0035'N, 77?38'W, 28 cent. Flowers with the calyx tube broadlyconical, 1-2 Dec 1980, Holm-Nielsen et al. 29804 (AAU, BM). mm long, the lobes broadly deltate, 1-3 mm long, Morona Santiago: km 22-27 of Cuenca-Gualaceo- sparsely arachnoid-pubescent;corolla white to dirty Sigsig-Chiguindard., 3050-3280 m, 3?09'S,78?39'W, white, fleshy, 1-2 cm diam., lobed nearly to the base, 12 Jul 1995, Al/arez & Tirado 1503 (BM, MO, the lobes planarat anthesis,the abaxialsurfacesof the QCNE). These specimens share rugose leaves, loose lobes sparselyarachnoid-pubescent,the tips and marpubescence, unifoliate sympodia, and fleshy flowers gins papillose; anthers 3-4 mm long, 1-2 mm wide, with the more typical Colombian specimens. Further poricidal at the tips, the pores teardropshaped; free studies of these plants in the field and good fruiting portionof thefilaments ca. 1.5 mm long, the filament collections from the Ecuadorianpopulationsmay help tubeca. 0.5 mm long, glabrous;ovay glabrousor denseto clarify theirrelationships.The Ecuadorianmaterial ly pubescentwith arachnoidtrichomes;stvle 6.5-8 mm long, glabrous;stigma merely a broadenedareaon top may representa new species. A single collection from southernPeru(Cusco: of the style, the surfaceminutelypapillose.Friiit a gloprov. Urubamba, Machupicchu, 88-112 km from bose, green berry,1-1.5 cm diam., appearinggrayishCusco, along Inca trail, 2060-4150 m, 13?09'10"S, green in specimens with pubescent berries;frtitinog pedicels thick andwoody, erect, 1-1.5 cm long, 1-1.5 72031 OO"W,14-22 Oct 1987,Niniez & Arque 8325 mm diam.atthebase,2-2.5 mm diam.atthe apex.Seeds (NY)) may also belong here, but the specimen has apdarkbrown,ca. 10 per fruit,ovoid-reniform,5-5.5 mm parentlybeen collected froma very immatureplantand long, 3.5-4 mm wide, the surfaces minutely pitted. is only in young bud. Chroimosonme numberunknown.

81. Solanum cornifolium Dunal,Solan.Syn. 21. 1816. Type. Colombia. Narinio:Popayan (crescit in convalli

fluminis Caucae inter Palace et Popayan,950 hex

Distribution (Fig. 146). Woods,paramoedges, and cloud forest second growth, 2300-3000 m, from Merida,Venezuela,to Carchi,Ecuador.

Specimens examined. COLOMBIA. Sl.oc., Apolinar-Maria44 (F); Liniden778 (BM); Muitis1980, 2000, 2003, 3565, 3581 (US). CUNDINAMARCA: Boquer6n de Solaonlnu sqiuamdlijerumtn Bitter, Repert. Spec. Nov. Bogoti. 2780 m, 21 Oct 1871, Andre 727 (K): region RegniVeg. 16:97. 1919.Type.Colombia.Narino: of Bogota, Ariste-Joseph s.nf. (US), 4riste-Joseph sn.H. fide HBK 18199),Humboldt & Bonpland s. n. (lecto-

type, P,here designated).

Figs. 136B, 142

2 58

FLORA NEOTROPICA

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(US); Salto de Tequendama, May 1917, Ariste-Joseph A115 (US); Bogota, 1917, Ariste-Joseph A252 (US); camino de GachetA,Jan 1920, Ariste-Joseph A546 (US); rd. to Mundo Nuevo, 12 km E of Calera, 2800 m, 10 Feb 1972, Barclay et al. 3168 (US); Salto de Tequendama, 2500 m, 1-3 Oct 1938, Cuatrecasas 83, 93 (US); Cordillera Oriental, Bogota, Usaqu6n, 2500 m, 9 Apr 1932, Cuatrecasas 3372 (K); Cordillera Oriental, Macizo de Bogota, Quebrada de Chico, 2750-2890 m, 8 Jun 1939, Cuatrecasas 5370 (F, US); Macizo de BogotA, Quebrada del Rosal, 3000 m, 29 Jun 1939, Cuatrecasas 5689 (F, US); Chipaqu6, May 1916, Dawe 195 (K); vic. Bogota, 2620 m, 28 Jan 1944, Dugand 3530 (US); Quebrada El Chic6, above N extremity of Bogota, 3000 m, 3?40'N, 74?04'W, 11 Jul 1943, Fosberg & Villareal 20573 (US); Anolaima, Finca Bethania below Peiia Negra, 2700 m, 29 May 1941, Garcia Barriga & Jaramillo M 10418 (US); Boqueronde Bogota, Goudots.n. (K); Monserrate,Guevara Am6rtegui 255 (US); Guadelupe, Bogota, ca. 2800 m, 5 Oct 1946, Haught 5054 (US); Bogota, Nov 1852, Holton 16 (K); between Cagua and La Caldera, 2 Aug 1943, Huertas & Camargo 872 (F); E edge of Chapinero, Bogota, 8900 ft, 6 Jan 1945, Little & Little 9192 (US); Facatativa, La Vega, Dintel, 2300-2700 m, 4 Jun 1939, Perez Arbeldez & Cuatrecasas 5303 (US); BogotA, 3000 m, 3 Feb 1948, Sandeman 5744 (K); Macizo de Bogota,

ls-pfsottpo

ooye

Quebrada Chic6, 8500-9500 ft, 30 Nov 1952, Schultes 18576 (US); Bogota, Boquer6n, 2800 m, Jan 1925, Schultze 111 (US). META: Cumaral, Jan 1937, Gama B. 5147 (US). PASTO: S.loc., 2800 m, Jun 1853, Triana 47 (BM). PUTUMAYO: Laguna La Cocha, above lake on rd. to Sibundoy near paramo de Bordoncillo, 2800-3000 m, 27 May 1946, Schultes & Villarreal 7522b (US). SANTANDER:W slope of Mount San Vicente, near Charta, 2500-2700 m, 9 Feb 1927, Killip & Smith 18994 (US); 10 Feb 1927, Killip & Smith 19178 (US). VENEZUELA. MtRIDA:Campo Elias, S of Pueblo Nuevo, between Mucusus and Cerro la Becerrera, 20002500 m, 18 Sep 1984, Ortega & Diaz 2135 (MO). TACHIRA: Headwaters of Rio Quinamari, along Quebrada Negra on trail to paramo de Judio (Apure border), 5 km S of San Vicente de La Revancha, 15 km S of Providencia, SE of Santa Ana, 2100-2400 m, 7025N, 72?25'W, 23 Oct 1984, Knapp & Mallet 6821, 6824 (BH, K, MO, MY, US, VEN). ECUADOR. CARCHI: Tulcan-Maldonado rd., ca. 3600 m, 2 Mar 1974, Harling & Andersson 12438 (GB, MO). Solanum cornifolium is morphologically quite similar to S. laevigatum (S. arboreum species group),

TAXONOMIC TREATMENT

259

ally,brownishabaxially,the apex acute,the base acute; petioles 0.7-1.5 cm long; minor leaves differing from the majors only in size, 3-9 cm long, 1.5-5 cm wide, the apex acute, the base acute;petioles 0.3-1 cm long. Inflorescencesoppositethe leaves, simple, 0.3-1.1 cm long, 5-15-flowered, glabrous or sparsely pubescent with arachnoidtrichomes like those of the stems and young leaves;pedicel scars congested,closely spaced, not overlapping.Buds globose when young, later ellipsoid, the corolla strongly exserted from the calyx tube. Pedicels at anthesis deflexed, filiform, abruptly contractedbelow the calyx tube, 0.5-1.5 cm long, ca. 0.5 mm diam. at the base, glabrousor sparsely(densely in Solomon 8488) pubescent with arachnoid trichomes like the rest of the inflorescence.Flower-s with the calyx tubebroadlyconical, 1-2 mm long, the lobes broadly deltate, 1-2 mm long, glabrous or occasion82. Solanum maturecalvans Bitter,Bot. Jahrb.Syst. ally with a few matted arachnoidtrichomes;corolla 50, Beibl. 111: 64. 1913. Type. Peru. Ayacucho: white or greenish-white, in some populations with a above Quinua, 3300-3500 m, 30 May 1910, pale lilac tinge, waxy, 1.5-2.2 cm diam., lobed nearly Weberbauer5543 (holotype, B [destroyed:F neg. to the base, the lobes planar at anthesis, the tips and 2622]; lectotype, F, here designated;isolectotypes, marginsof the lobes denselypapillose;anthers4-6 mm Figs. 136C,D, 143 long, 1-2 mm wide, poricidalat the tips, the porestearGH, US). Mem. Solanuimaureiftlium Rusby, Torrey Bot. Club drop shaped;free portion of thefilaments ca. 0.5 mm nearsnow long, the filamenttube0.5-1 mm long;ovaryglabrous; 6: 87. 1896.Type.Bolivia.Cochabamba: line on Mt. Tunari?, 1891, Bang 1119 (holotype, style 0.6-1 cm long, glabrous;stigma slightly clavate, NY: isotypes, F, GH, K, NY, P). nom. confusum, the surfaceminutelypapillose.Fruit a globose or ovoid type does not match description, see discussion. green berry,often grayish-greenat maturity,the apex Solanurni crotalobasis Bitter, Repert. Spec. Nov. pointedwhen dry, 1-1.5 cm diam., to 2 cm long;firuitRegni Veg. 13: 96. 1914. Type. Bolivia. La Paz: ingpedicels woody, deflexed, 1.5-2.5 cm long, ca. 1.5 Prov. Larecaja, vic. Sorata, Queliguayo, ca. 2800 mm diam. at the base. Seeds darkbrown, ovoid-renirn, Oct 1879, Mandon 413 (holotype, P). Solanunmverniciflor-um Bitter, Repert.Spec. Nov. Regni form, 2.5-3.5 mm long, 2-3 mm wide, the surfaces Veg. 18: 66. 1922. Type. Bolivia. Rio Tocorani, minutelypitted.Chromosomenumber.n = 12 (voucher 2500 m, Ilerzog 2291 (holotype, B [destroyed]; Knapp & Mallet 6351; Roe, 1967a).

also of Colombia,but differs from it in its non-barbate vein axils and denser, more grayish pubescence. The berrypubescence polymorphismis presentat both local andregional scales, with specimens not clearly dividing into regional sets. The type specimen in P is not in the Bonpland herbarium,where other solanumsdescribedby Dunal from Humboldtand Bonpland's collections generally arehoused. However, a sheet in the generalherbarium at P is an exact matchfor the unpublishedNode-Veran drawing made for Dunal's planned second edition of Solanorumsynopsis and is annotatedin Dunal's hand (drawingsheld at MPU, see Knapp& Spooner, 1999). I have designated this sheet at P (Fig. 142) the lectotype of Solanum cornifolium.

no isotypes located). ex descr. Solanurn ochrophyllum Van Heurck & Muill.-Arg. var. schnmidtiiJ. F. Macbr., Publ. Field Mus. Nat.

Distribution (Fig. 144). S Ecuadorto Bolivia and NW Argentina in montane cloud forests from Hist,Bot.Ser.13(5B):213. 1964.Type.Peru.Cuzco: (1000-)2000-3000 m.

Hacienda de Urcos, 18 Sep 1939, Schnmidts.n. (holotype. F; isotype, F). Solanutim kieslingii Cabrera,Hickenia 1(31): 166. 1978.

Selectedspecimensexamined.ECUADOR.LOJA: 16 Nov 1876,Andre4445 (F); between Chuquiribamba, El Tamboand La Toma, 1000-2200 m, 3 Sep 1923, Type:Argentina.Jujuy:Depto.ValleGrande,Abra Hitchcock21346 (NY, US). de Canas, 1700 m, 28 Dec 1977, Kiesling et al. PERU. APURIMAC: NNW of Abancay,km 16 of 1525 (holotype, SI-n.v.). Cuzcord., ca. 3000 m, 19 Dec 1962,Iltis & Ugent638, Shrubsto small trees, 1-10 m tall; young stems 650 (BM, F, US, WIS);km 20 of Cuzcord., ca. 6-7 km andleaves densely arachnoid-pubescent, the trichomes by air NNW of Abancay,ca. 3100 m, 19 Dec 1962,Iltis N mattedandfelt-like,soon deciduous;barkof olderstems & Ugent672 (, BM, F, US, WIS);Ampay,Tamburco, darkgrayish-brownwith largewhite lenticels.Sympo- of Abancay,3800-5235 m, 13?30'S,75052'W,22 Feb 1987, Nunez & Vargas 7225 (MO, NY); Abancay, near dial unitsdifoliate, geminate.Leaves elliptic, widest at CurahuasiandTrancapata, 2800 m, 8 Nov 1938, Vargas the middle, glabrous and shiny adaxially, often con- C. 9621 (CUZ, K, MO). CAJAMARCA: Ca. 17 km S of spicuously bullate, occasionally with a few arachnoid Cajamarca on rd. to SanJuan,just S of AbraEl Gavilan, trichomesalong the main veins, glabrousor pubescent ca. 3030 m, 20 Oct 1984, Dillon & Whalen 4064 (F, K, with arachnoidtrichomes along the veins abaxially; NY); ca. 7 km S of Contumazaen routeto Cascas, ca. majorleaves 4-1 8(-30) cm long, 2-1 1(-28) cm wide, 2170 m, 7?23'S, 78?47.5'W,25 Oct 1990, Dillon, & with 7-12 pairsof main lateralveins, impressedadaxi- SagasteguiA. 6080 (BM, F); Hualgayoc,HaciendaTaulis

FLORA NEOTROPICA

260

_........ ....... .....

Dept. APUR.IY.AOPray. aomeshrub In hedgewowe F:l. l.ot ross violet. Gienies.

-

to t-2 v. tell.

Rtook hedgerow. and thickets (remnanite of former Zealo aloud forest) between fields. With heavily poilarded trees of shrabty Uains, sscailonia, lalcolaria, sclanum, lyoium omaria,tet. lcpatoriue,PaCOLf,lOra, 2~seeAt pa%ur o4fal49 te 'ea&blef ar ; at km. 20 sml chacra" (field) on the.Ouzoo road, ca.6-7 km DNN M. lt.ca.3100 Qair)f rcm *:e. '19. 1962.

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.5cm

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.

.

FIG. 143. Solanum maturecalvans Bitter. Peru. ltifs & Ugent 672 (BM).

'

261

TAXONOMIC TREATMENT

74)0

Mallet 6351 (BH, K, MO, NY, US, USM); Paucartambo, km 130 of Lucre-Paucartambo-Shintuya rd., Rio Alto Madre de Dios drainage, ca. 2300 m, 13012'S, 71?32'W, .0,~~~~7. 17 May 1984, Knapp & Mallet 6463 (BH, GH, K, 0~~~~ MEXU, MO, NY, US, USM); vic. of Ollantaytambu,Rio Vilcanota valley, ca. 2800 m, 13?15'S, 72?20'W, 18 May 0 1984, Knapp & Mallet 6466 (BH, F, GH, MO, NY, US, USM); Urubamba, 67 km from Cusco, from Rio 0 Calicanto to Chacchapata, vic. of Muris, 2700-3020 m, 21 Nov 1987, Nuniez & Bengoa 8662 (MO, NY); Quispicanchis, between Huallahualla & Marcapata, 210 km from Cusco, 2800-4600 m, 13 ?25'S, 70?54'W, 2125 Apr 1988, Nuinez et al. 9060 (MO, NY); Urubamba, Quebrada de Pumahuanca, 2900 m, Mar 1983, Ochoa 15106 (US); Machu Picchu', along Inca trail in Tres 00~~~ Piedras Blancas near Huayabamba, 3330 m, 15 May 1982, Peyton & Peyton 241 (MO, USM); Quillabamba, Santa Teresa, hillside called Mandornilloc, 0.5 km W of La Playa, 2525 m, 9 Sep 1982, Peyton & Peyton 1244 (MO, USM); Urubamba, 3 Jan 1936, Soukup 36 (F); Anta, Huancacalla, 3100 m, Jul 1937, Vargas C. 439 (CUZ, F); Urubamba, Pumahuanca, 2950 m, Jan 1949, Vargas C. 7661 (CUZ, MO); Quispicanchis, between Cachu-pampa and Cgile-chile, 2000-2500 m, 10 Dec 1938, Vargas C. 9692 (A, CUZ, F, K, MO). HUANUCO: Pillao, 2700 m, 14 Feb 1946, Woytkowski 34064 (F). JUNIN: Tarma-San Ram6n rd., rd. to Illic beyond Palca Campalanayoc, 2900 m, 11?20'S, 75?40'W, 11 Jan 1987, Diaz S. & Huanqui 1995 (MO, NY); rd. to Illic (Palca), 2750 m, 13 Jan 1987, Diaz S. & Balde6n 2197 (MO, NY); 26 km NE of Tarmaon Carretera20B to Oxapampa, 8850 ft, 12 Nov 1979, Davidson & Jones 9055 (F, NY); FIG. 144. Distribution of Solanum callianthum vic. of Comas, 3300 m, 8 Jul 1948, Ochoa 531 (US). (open circles) and S. maturecalvans (solid circles). LA LIBERTAD: Santiago de Chuco, Huacas (Cachicadan), 2800 m, 15 Jun 1984, Sagastegui A. et al. 11924 (HUT, NY). PASCO: Oxapampa-Villa Rica rd., 7 km from rd. on Rio Taulis, 2770 m, Sep 1964, Hutchison & von head, 2120 m, 10?36'S, 75?20'W, 4 Jan 1984, Smith & Bismarck 6469 (K, NY); Bosque Cachil, 2570 m, 18 Aug Albain5574 (MO, NY). PIURA: Huancabamba,Aug 1943, 1994, Leiva G. et al. 1553 (BM, F); above Chiquiden, Sandeman 4309 (K). PUNO:Carabaya, Ollachea, across 2850 m, May 1982, Ochoa 14781 (F, US); Cuchero, 1830, Rio San Gaba from town, , 23 Aug 1980, Boeke & Boeke Poeppig 1266 (F); Contumaza, Las Quinuas-EI Mej6n, 3191 (NY); Ollachea to San Gab6n, 1000-2000 m, 173200 m, 14 Jul 1981, Sagastegui A. et al. 10101 (F, 24 Jul 1978, Dillon et al. 1180 (MO). SAN MARTiN: HUT, MO, NY); E of Cajamarca-Pacasmayo rd., de- Prov. Huallaga, below La Morada, 2000-2200 m, 6 57'S, scending Paso El Gavilan toward San Juan, 3000 m, 18 77 32'W, 11 Aug 1997, Quipuscoa S. & Bardales 991 Amy 1986, Sanchez Vega 4048 (NY); Chetilla rt. to (BM, F); Rio Abiseo National Park, top ridges of Cerro Central, 2900 m, ca. 7?S, 77?W, I Aug 1985, Young1406 Llullapuquio, 2650-2750 m, 21 May 1986, Sanchez Vega & Cabanillas S. 4095 (NY); 17 km S of Cajamarca (K, NY); NW corner of Rio Abiseo National Park, S side on rd. to Chilete, just over Paso Gavilan, 20 Oct 1984, of river, Chochos, 3400 m, 7 Jun 1986, Young 3722 Whalen & Dillon 890 (BH, MO, NY). Cuzco: (HUT, IBE, K, NY). BOLIVIA. COCHABAMBA: 80 km from CochaOllantaytambo, ca. 3000 m, 26 Apr 1915, Cook & Gilbamba toward Villa Tunari, 2250 m, 23 Nov 1981, Beck bert 329 (US); 8 May 1915, Cook & Gilbert 587 (US); 11 May 1915, Cook & Gilbert 629 (US), 18 May 1915, 7268 (F); Choro, 10000 ft, 12 Mar 1950, Brooke 6179 (BM, F, NY); Comarapa-Cochabamba rd., 4 km W of Cook & Gilbert 796 (US); Paucartambo valley, Hacienda Capana, Aug 1926, Herrera 1132 (US); Chinceros, border with Depto. Santa Cruz, 20 km by air and 28 km by rd. NW of Comarapa, 2525 m, 17?49'S, 64?41'W, 10 Antakillqa hillside above Incaj mallkin pampa, 3350Feb 1987, Nee & Solomon 34037, 34050 (IBE, NY); 5 3500 m, ca. 13?23'S, 72?02'W, 25 Jan 1982, Davis et al. 1777 (F, GH); Cordillera Ver6nica, 3500 m, 28 Apr 1957, km SE of bridge at L6pez Mendoza, 19 km NW of Hirsch P1024 (F); Urubamba, ca. 4 km N of Ollantay- Epizana, on rd. from Comarapa to Cochabamba, 2900 tambu on rd. to Quillabamba, Rio Vilcanota drainage, m, 17?32'S, 65?22'W, 11 Feb 1987, Nee & Solomon ca. 2700 m, 13?l5'S, 72?20'W, 10 Apr 1984, Knapp & 34091 (IBE, NY); Churro, 7 km by air ESE of Siberia,

FLORA NEOTROPICA

262 8.5 km SW of Santa Cruz border, 2600 m, 17?50'S, 64?42'W, 5 Mar 1988, Nee et al. 36491 (IBE, NY); narrow canyon of Rio Monte Puncu, 5 km NE of Monte Puncu, 10 km by air NW of Epizana, 2700-2750 m, 17?33'S, 65016'W, 10 Mar 1988, Nee & Solomon 36629 (1BE, NY); Chapare, 23.8 km N of Colomi, jct. of rd. to Candelaria, on rd. to Chapare, 2.2 km NW on side rd., upper Rio Cayani, 2700 m, 17?10'S, 65?53'W, 19 Oct 1985, Solomon 14389 (MO, NY); 23.8 km N of Colomi, jct. of rd. to Candelaria,on rd. to Chapare, 7.4 km W on side rd., upper Rio Cayani, 2400 m, 17?09'S, 65?53'W, 19 Oct 1983, Solomon 14415 (MO, NY); 4.5 km SE of Siberia on Cochabamba-Santa Cruz rd., 2800 m, 17?50'S, 64?44'W, 5 Feb 1987, Solomon & King 15949 (MO, NY). LA PAZ: Quime, 100 mi. from Oruro via Eucalyptus, crossing the Quimcruz mtns. by the Tres Cruces pass, 8000 ft, 16'30'S, 67?00'W, 5 Apr 1949, Brooke 5427 (BM); Prov. Nor Yungas, Unduavi, 3150 m. Oct 1931, Buchtien 9030 (NY, US); Prov. Larecaja, vic. Sorata, Queliguayo, 2700-3000 m, Oct 1879, Mandon 418 (BM, K); Rio Zongo valley, 27 km below dam at Lago Zongo, 2600 m, 16?08'S, 68?06'W, 10 Oct 1982, Solomon 8488 (MO, NY, U); Cordillera Real, Nequejahuira, 8000 ft, 15-24 May 1926, Tate 676 (NY). SANTA CRUZ: 32 km from Comarapa on way to Cochabamba, 2500 m, 24 Mar 1981, Beck 6813 (NY); Samaipata-Ayopayo, 2750 m, Mar 1935, Cardenas 3117 (US).

sympodia.Many collections ofS. maturecalvansfrom elsewhere in its range appear to have unifoliate sympodia,butthis is due to the loss of the minorleaf in drying or specimen preparation.These populationsin NW Argentinameritcloser studyandmay also deserve recognitionas a distinct species. Rusby's descriptionof Solanumaureifoliumis clearlya compositeof severalsheets anddoes not completely match the type specimen at NY. The leaf size and leaf base measurementsfit Bang 1119 at NY, the designatedtype, however the numbersof veins in the leaves, inflorescences, flowers, and fruit matchBang 1118, which RusbyhaddescribedasS. clavatuimRusby (Rusby, 1896). This specimen was also a mixed collection, with one elementclearlymatchingthe description writtenby Rusby for S. clavatumand the other a memberof sect. Cyphomandropsisandlatergiven the nameS. confusumC.V.Morton(see Morton,1944;Bohs, 1994). The elementof Bang 1118 to which the description of S. clavatum,andthe descriptionof the flowers, inflorescences, and fruitof S. aureifoliumn refer is the widespread species of sect. Holophylla s.str., S. aligerum Schltdl.

83. Solanum nigricans M. Martens& Galeotti,Bull. Acad. Roy. Sci. Bruxelles 12: 134. 1845. Type. Mexico. Oaxaca:Comaltepeque,3000 ft, Galeotti orccotunya(Ollantaytambu),qosmayllu(Chincheros). s.n. (lectotype, BR, here designated;isolectotypes, Solanum maturecalvans is closely related and G [F neg. 23136], GH, NY, P [Mortonneg. 22291], morphologicallysimilarto S. ochrophyllumof Bolivia US). Fig. 145 and S Peru and S. nigricanis of Mexico and Central Solanum multinervium Dunal in DC., Prodr. 13(1): America.Populationsfromthe Cuzco areahave larger, 127. 1852. Type. Mexico. S.loc., Pav6n s.n. (coll. more conspicuously bullate leaves, but are otherwise Sesse & Mo9ifio) (holotype,G-n.v.; isotypes. F, similar to the rest of the species. Solanum matureG-DC [F neg. 34124]). calvans is easily recognizable due to its conspicuous Solanum brachystachys Dunal in DC., Prodr. 13(1): felty pubescence which appears to slough off in an ir128. 1852. Type.Mexico. Chalco,Andrieux186 (holotype, G-DC [F neg. 6787, NY]; isotypes, regular fashion and the leaves with revolute margins K, M-n.v. [Morton neg. 8674], frag. MPU). narrowingabruptlyto the petiole (the basis for Bitter's Solanum vernicinitensBitter, Repert. Spec. Nov. Regni epithet crotalobasis). Older leaves are more glabrous Veg. 18: 58. 1922. Type. Guatemala.Alta Verapaz: thanyoungerones, thusBitter's choice of specific epiCoban, 1300 m, von Tuerckheimin Donn. Smith thet. Solomon 8488 from the Rio Zongo valley in Bo745 (holotype,B [destroyed];lectotype,US, here livia has huge leaves and is much morepubescentthan designated; isolectotype, K). the rest of the specimens I have seen. It may represent a new species but is only just outside the range of variaShrubsor small trees, 1-10 m tall; young stems tion found in this quite variable species. andleaves densely pubescentwith compressed,arachI have not seen the type specimen of Solanum noid dendritictrichomesor with compactdendritictrikieslingii, fromNW Argentina,nor any of the material chomes, or occasionally glabrous;barkof older stems cited in the original description (Legname& Cuezzo darkbrown,glabrous,with large white lenticels.Sym5954 (LIL, LP), Fabris etal. 3104 (LP), and de la Sota podial units difoliate, geminate.Leaves elliptic to nar4418 (LP)-all fromthe type locality). It appearsto be rowly elliptic, widest at the middle, glabrousand shina synonym of S. maturecalvansfrom the description, ing adaxially, glabrous or occasionally sparsely and fits well within the wide range of variationof the pubescentwith arachnoidor dendritictrichomesalong species. The illustration in the original description the veins; major leaves 5-16 cm long, 3-6 cm wide, (Cabrera,1978), however, is of a plantwith unifoliate with 8-12 mainlateralveins dryingyellowish abaxially, Local names. Peru. Cuzco: urco moyocjaya,

263

TAXONOMIC TREATMENT

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the apex acuteto acuminate,the base acuteto attenuate; petioles 1.5-3 cm long;minorleaves differingfromthe majors only in size, 4-10 cm long, 2-5 cm wide, the apexacuteto acuminate,thebase acuteto attenuate;petioles 1-3 cm long. Inflorescences opposite the leaves, occasionally intemodal, simple, 0.5-3 cm long, 5-15flowered,congested,denselypubescentwith arachnoid or dendritictrichomes;pedicel scars closely spaced, occasionally overlapping.Buds globose when young, laterellipsoid, the corollaexsertedfromthe calyx tube. Pedicels at anthesisdeflexed, 1.1-2.3 cm long, tapering fromthe calyx tubeto a slenderbase ca. 0.5 mm diam., densely pubescentwith trichomeslike those of the rest of the inflorescence or glabrous.Flowers with the catube conical, 1-1.5 mm long, the lobes deltate,0.5Ivvx 1 mm long,denselyto sparselypubescentwithtrichomes like those of the restof the inflorescence;corollawhite, fleshy, 1-1.3 cm diam., lobed nearly to the base, the lobes planar at anthesis, densely papillose along the marginsand at the tips; anthers 3-3.5 mm long, 1-1.5 mmwide,ponrcidalatthetips,theporesteardropshaped; free portionof thefilamentsca. 0.25 mm long, the filament tube ca. 0.5 mm long, glabrous;ovary glabrous; stvle 5-6 mm long;stigmacapitate,minutelypapillose. Fruit a globose or apicallypointedgreenberry,1-2 cm diam.;fruitingpedicels woody, more or less deflexed, 1-3 cm long, ca. 1.5 mm diam. at the base, 2-3 mm diam.at the apex.Seedsdarkreddish-brownor pale tan, ovoid-reniform,3-4 mm long, 2.5-3 mm wide, the surfaces minutelypitted.Chromosomenumbernot known. Distribution (Fig. 146). In cloud forests and pine-oak forests from central Mexico to Honduras, from 1000-3200 m. Selected specimens examined. MEXICO. CHIAPAS: Near boundary of Chamula on rd. to

Zinacantan, 7500 ft, 17 Aug 1964, Breedlove 5060 (US); San Crist6bal-Tenejapa rd., 7500 ft, 4 Aug 1964, Breedlove6796 (F, US); nearboundaryof Chamulaon rd. to Zinacantan,7500 ft, 17 Aug 1964,Breedlove7060 (F, MEXU);crestof ridgeon SanCrist6balde las CasasTenejapard., 8300 ft, 19 Feb 1965,Breedlove9078 (F, US); trailfromTenejapacenterto ColoniaSan Antonio, 7200 ft, 12 Jul 1965,Breedlove10804(F);lakeIk'alNab, on boundarybetweenChamulaandZinacantan on way to ZinacantanCenter, 7800 ft, 21 Sep 1965, Breedlove 12380 (F, US); parajeof Banabil,9100 ft, 10 Oct 1965, Breedlove& Raveni12927 (F, US); NE side of Matsab hill, 9200 ft, 25 Aug 1966,Breedlove15269(NY);Cerro Huitepec(Muk'ta vits), W of SanCrist6balde LasCasas, 2700 m, 5 Dec 1971, Breedlove 23042 (NY); Cerro Huitepec,W of San Crist6balde las Casas,4 Dec 1983, Cabrera & de Cabrera 5998 (NY); along rd. from Motozintlade Mendozato Siltepecvia El Provenir,14.1 mi NW of Motozintla,11 Feb 1976, Croat47306 (MO, NY); along rd. between Motozintla de Mendoza and

Siltepec 21.5 mi NW of Motozintla, 1.3 mi W of El Porvenir, 1770 m, 11 Feb 1976, Croat 47325 (MO, NY); along rt. 190, ca. 5 mi W of San Crist6bal de las Casas. 26 Jun 1960, King 2073 (NY, US); below Zinacantan Center on trail to Ixtapa, 6200 ft, 5 Dec 1966, Laughlin 2923 (F, US); Sierra Madre de Chiapas, Cerro Mozotal, Huixtla-Siltepec rd., along rd. to microwave tower, 28652975 m, ca. 15?28'N, 92 19'W, 19 Jun 1985, Luteyyn& Lebron Luteyn 11627 (MEXU, NY); Mt. Ovando, 17 Dec 1936, Matuda 432 (MEXU, US); Mt. Pasitar, 29 Dec 1936, Matuda 458 (MEXU, US); Saxchanal, Sierra Madre, 2700 m, 1 Jul 1941, Matuda 4289 (F, NY, US); Mt. Pashtal, 2500 m, 12 Apr 1948, Matuida 17678 (F, MEXU); Saxchanal, Sierra Madre, 11 May 1948, Matluda 17817 (F, MEXU); 24 May 1948, Matluda 17866 (F); 8 Jun 1948, Matuda 18665 (F); s.loc., 1913, Purpls 7105 (US); Cerro del Boquer6n, Jun 1914, Purplus 7325 (F, NY, US); Rancho del Boquer6n, Jun 1914, Ptiopius7327 (NY, US); Sumidero of Yochib, paraje Kotolte', 4500 ft, 5 Apr 1966, Ton 748 (F, MEXU, NY); paraje Balum K'anal, 8400 ft, 13 Apr 1966, Ton 812 (F, MEXU, NY); paraje of Matsab, 8900 ft, 28 Sep 1966, Ton 1255 (F, US); paraje of Matsab, 8900 ft, 5 Oct 1966, Ton 1306 (F, NY, US); Cerro Huitz, 11 May 1985, Ton 8126 (MEXU, NY); Cruzton camino Tzonte-Huitz, 20 Apr 1985, Toni 8197 (MEXU, NY). GUERRERO: 6.4 mi SW of Filo de Caballo, 9100 ft, 8 Jul 1987, Anderson sn.. (NY); 30 km NE of Puerto del Gallo, Atoyac-Filo de Caballo rd., 2650 m, 23 Nov 1983, Martinez S. & Baornie5672 (NY); 18 km S of Filo de Caballo, 2680 m, 9 May 1982, Rodriguez B. & Martinez S. 112 (NY); 5 km W of Camotla, 2600 m, 8 Apr 1963, Rzedowski 16419 (US). JALISCO: 15.4 mi SW of Ciudad Guzman, 7500 ft, 2 Aug 1988, Andersoni88-21 (NY); Sierra de Manantlan, Cerro El Almeal, headwaters of Arroyo San Campus, ca. 4-5 km by rd. SE of Estaci6n Biol6gica Las Joyas, 2050-2100 m, 19?34'N, 104?15'W, 22 Dec 1984, Cochranie et al. 10626 (NY); Arroyo de Manantlan. N facing slope of Sierra de Manantlan, ca. 2-3 km S of Rincon de Manantlan, 15 km S of El Chante, 1760 M, 19034'50"N, 104012'55"W, 5 Jan 1979, Iltis et al. 1204 (F); Sierra de Manantlan, Zarza Mora, 2 km E of Las Joyas, 6 km WSW of Rinc6n de Manantlan, 15 km SSW of El Chante, 1900 m, 19?35'15"N, 104015'50"W, 6 Jan 1979, Iltis et al. 1287 (F); Sierra de Manantlai Occidental, 1 km W of Zarza Mora, 1 km SW of Las Joyas, ca. 20 km SSE of Autlan, 1800-2000 m, 19?35'N. 104?18'W, 31 Dec 1983, Iltis & Guzman 29023a (NY); Sierra de Manantlan, Cerro Grande, around Terrero, 2200 m, 19026'40"N, 103057'W, 13 Mar 1987, Iltis et al. 29502 (NY); Sierra de Manantlan Occidental, ca. 2 km NE of Cerro de la Cumbre, 3.5 km SSW of Rinc6n de Manantlan, 16.5 km due S of El Chante, 1950-2100 m, 6 Jan 1980, Kowal 2806 (F); Sierra del Halo, 7 mi SSW of Tecalitlan, on lumber rd. SE to San Isidro, 2000-2200 m, 28-30 Nov 1959, McVaugh & Koelz 1204 (NY, US); Sierra de Manantlan, 15-20 ml SE of Autlan, ca, 2 mi from Aserradero San Miguel Uno, W and S of divide toward Manzanillo, 2250-2400 m, 4-5 Nov 1952, McVaugh 13892 (US); Sierra de Manantlan, between

TAXONOMIC --

TREATMENT -

265 -

~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~~~~ >

7 0 0 - - -- - - -

0~~~~~~~0

00~

~~~~

-------t

FIG. 146. Distribution of Solanum cornifolium (open circles), S. nigricans (solid circles), S. ochrophyllum (open squares), S. platycypellon (solid squares), and S. quebradense (solid triangle).

Aserradero San Miguel Uno and Durazno, near pass to El Fresno, ca. 2000 m, 6 Nov 1952, McVaugh 13945 (US); near Santa M6nica, 1950-2050 m, ca. 20?N, 104?30'W, 12-13 Nov 1952, McVaugh 14044 (US); headwatersof Rio Mascota, 25-30 km airline SE of Talpa de Allende, 1-2 mi N of sawmill La Cumbre and 11-12 mi S of El Rinc6n, 2000-2250 m, 29 Nov 1960, McVaugh 21518 (NY, US); 3-5 mi NW of San Miguel de la Sierra, ca. 40 km airline W of Ayutla, 2000 m, 3 Nov 1962, McVaugh 22031 (NY); Sierra Madre Occidental, Real Alto, trail to San Sebastian, below El Bufa, 2000 m, 27 Feb 1927, Mexia 1772 (NY, US); Puerto El Floripondio, rd. E of microwave station Las Viboras, 2450 m, 26 Jul 1987, Rodriguez C. & Suarez J. 918 (MEXU); 22 km SE of hwy. 110 on lumber rd. leaving hwy. 12 km SSW of Tecalitlan toward San Isidro, ca.

1700 m, 19018'N, 103?05'W, 25-26 Sep 1965, Roe & Roe 2140 (US, WIS). MEXIco: Cajones, Temascaltepec, 2480 m, 22 Dec 1932, Hinton 2327 (F); Temascaltepec, Los Hornos, 2600 m, 17 Apr 1933, Hinton 3701 (F, K, US); Temascaltepec, Cumbre, 2600 m, 21 Apr 1935, Hinton et al. 7479 (NY, US); Cumbre-Pefia Blanca, 29 Mar 1936, Hinton et al. 9004 (F, NY, US); San Rafael, Dec 1933, Lyonnet & Elcoro 1128 (US); Cieneguillas, 14 km N of Temascaltepec, 2300 m, 12 May 1973, Moreno G. 174 (F). MICHOACAN: Mil Cumbres, 8300 ft, 27 Jul 1988, Anderson 88-4 (NY); NW slope of Cerro El Cacique, 19?23'30"N, 100?19'45"W, 25 Jan 1981, Contreras 1305 (F); N slope of Cerro El Cacique, 19024'N, 100018'38"W, 5 May 1981, Contrer-as 1410 (F); Zitacuaro-Macho de Agua, 2420 m, 3 Jun 1938, Hinton et al. 11911 (NY, US); S. Torricillas, 2180 m,

266 15 Dec 1938, Hinton et al. 12747 (F, NY, US); Tancitaro, Uruapan, 2075 m, 17 Oct 1940, Hinton et al. 15537 (F, NY, US); ca. 18 mi S of Patzcuaro, 8900-9000 ft, 2025 Nov 1961, King & Soderstrom 5212 (NY, US); 2 mi W of Tancitaro, 6660 ft, 15 Jul 1940, Leavenworth 249 (F, NY); 6 mi N of Tancitaro, 8500 ft, 25 Jul 1940, Leavenworth 355 (F, NY); Pedregal lava flow 2 mi. S of Tancitaro, 6500 ft, 14 Aug 1940, Leavenworth 553 (F); Las Peras, 38 km E of Morelia, km 272 on Hwy. 15 (Ciudad Hidalgo-Morelia), 2515 m, 13 Sep 1962, Ugent & Flores C. 2011 (US). MORELOS: Valle de Tepeite, Jul 1936, Lyonnet 1492 (US). OAXACA: S peak of Huitepec, near Las Casas, 18 Apr 1945, Alexander 1133 (NY); km 12 of San Andres Yaa detour, rd. to Oaxaca S of Villa Alta, 2300 m, 17 Mar 1982, Cedillo T & Torres C. 1208 (F); 7 km SW of El Punto, Between El Punto and La C'umbre,2000 m, 20 Apr 1982, Cedillo T & Lorence 1272 (F); El Recibimiento, Cuyamecala, Cuicatlan, 1900 m, 20 Jun 1909, Conzatti et al. 2470 (F); Natividad, timber rd. to Llano Verde, ca. 7000 ft, 26 Jun 1983, Costich & Baldwin 1501, 1502 (BH, F); 3 Jul 1983, Costich & Baldwin 1510 (BH, F); Villa Alta-Totontepec, 2520 m, 5 May 1986, Flores M. 1065 (NY); Sierra Mazateca, E of Teotitlan, 7500 ft, 21 Aug 1963, Gentry et al. 10303 (herb. unknown, US?); Puerto de la Soledad, Teotitlan del Camino, 7000 ft, 2 Apr 1972, MacDougall H342 (F, NY); Sierra de Clavellinas, 9000 ft, Oct 1894, Charles L. Smith 775 (NY, US); 2 km NW of Huautla, 1400 m, 18009'N, 96052'W, 6 Jan 1984, Solheim & Reisfield 1240 (NY); 15 km E of Calpulalpan "Llano Verde," 2160 m, 30 Jun 1982, Torres C. & Cedillo T 704 (NY); 13.7 km SE of San Jose del Pacifico, 2605 m, 25 Apr 1983, Torres C. & Cedillo T 2751 (NY); 4.7 km N of Maravillas and 23 km N of Zoogocho, rd. to Talea de Castro, 15 May 1983, Torres C. et al. 2918 (NY); 19.1 km N of Yacochi, rd. to San Andres Yaa, 2340 m, 8 Aug 1985, TorresC. & Torres C. 7112 (NY). PUEBLA: Along Tehuacan-Orizaba hwy. on W slopes below Puerto del Aire, ca. 1800-2200 m, 18 Jul 1961, Smith et al. 3926 (F, NY, US); Tezuitlan, El Carrizal, 1700 m, 31 Jul 1969, VenturaA. 298 (F). VERACRUZ: Rd. from Puerto del Aire to Laguna, 2320 m, 16 Dec 1977, Calzada & Delgado 4180 (F); Rancheria Pozo Panal, rd. to Positos, 2600 m, 19030'N, 97?06'W, 10 Jan 1980, Calzada 5745 (F); Tepanosas, 1420 m, 19026'N, 97001'W, 25 Nov 1977, Castillo C. et al. 232 (F); Cerro de Villa Rica, near Mundo Nuevo, 1600 m, 19047'N, 96?46'W, 7 May 1981, Castillo C. et al. 1827 (F); vic. Tonolaco slopes of Cofre de Perote, 2680 m, 13 Jun 1981, Castillo C. et al. 1942 (F); near Buenavista, rd. from Herradurade Xizo to Corral de Rajas, 2200 m, 7 Aug 1986, Chazaro & Robles 3853 (NY); 14.4 km on rd. from Coscomatepec and Hwy toward Alpatlahua, 5.2 km beyond Alpatlahua, 10 Jun 1983, Cowan 3919 (NY, TEX); Potrero de Garcia, 4 km NE of La Joya, 1700 m, 25 Nov 1975, Marquez R. et al. 442 (F, NY); Coscomatepec, May 1937, MatudaS-143 (US); Maltrata,May 1937, Matuda 1286 (US); BarrancaLa Funda, near Los Laureles, S of Cofre de Perote, 2750 m, 19?20'N, 97?20'W, 7 Oct 1983, ANaraveE & Vasquez B. 1070 (NY); 5.3 km W of Escola on rd. to Jacal. 18.5 km NW of Coscomatepec,2300

FLORA NEOTROPICA m, 19?10'N, 97?11'W, 12 Jan 1981, Nee & Schatz 19709 (F); 9.5 km W of Escola along Coscomaltepec-EscolaJacal rd., 2600 m, ca. 19?07'N, 97?1l'W, 15 Nov 1981, Nee 23188 (F); 9.5 km W of Escola along CoscomatepecEscola-Jacal rd., 2600 m, ca. 19?07'N, 9711 'W, 15 Nov 1981, Nee 23190 (F, NY); 1 km above and NW of San Andres Tlalnehuayocan, 1700 m, 19?34'N, 96?58'W, 22 Mar 1983, Nee et al. 26183 (F, MPU, NY); Puente Acabaloya, ca. 1 km SE of Xico Viejo and 5 km NW of Xico along trail between the two, 1600 m, ca. 19?27'N, 97?03'W, 31 Mar 1983, Nee & Taylor 26305 (NY); 1 km NW of Elotepec along rd. to Chichiquila, 1700 m, 19012'N, 97?02'W, 17 Jan 1984, Nee & Taivlor28900 (F, NY); above and NE of Xico, 8 km NW of Xico, 17001900 m, ca. 19?28'N, 97?04'W, 5 Feb 1984, Nee & Talilor 29388 (F, NY); along hwy. Mex. 150, 0.5 km from Edo. Puebla border and 0.8 km SSW of Puerto del Aire, 2250 m, 18?41'N, 97?21'W, 20 Sep 1986, Nee 33104 (NY); before Acultzingo near border of Puebla and Veracruz,2300 m, 28 Jul 1971, Nevling & Gomez-Pompa 2139 (F); Cumbres de Acultzingo, 20 Apr 1966, Smith & Tejada 4529 (US); Tiapa, 2050 m, 30 Apr 1971, VenturaA. 3536 (F); Tongonapa, 1550 m, 14 Jan 1974, Ventura A. 9503 (F); Acajete, 1800 m, 8 May 1976, VenturaA. 12746 (F). GUATEMALA. Between Jacaltenango and San Martin, 5300-7000 ft, 24 Dec 1895, Nelson 3605 (US); Calderas, 5 Jul 1941, Johnston 1916 (F, US). ALTA VERAPAZ: Banks of Rio Frio, between Tactic and Santa Cruz, 1460 m, 14 May 1963, Molina R. & Molina 12234 (F); Chicoj,NE of Carcha,ca. 1200 m, 2 Apr 1939, Standley 70037 (F); Tactic, 1480 m, 9 Apr 1939, Standlev 71350 (F, US); between Tactic and the divide on rd. to Tamahu, 1500-1600 m, 1-7 Apr 1941, Standley 90682 (F). BAJA VERAPAZ: Uni6n Barrios, NW of km 159, 12 Jun 1975, Lundell & Contreras 19445 (F); Nino Perdido, 19 May 1977, Luindell & Contreras 20920 (F); Nino Perdido, La Cumbre de San Jose, 21 Jun 1977, Lundell & Contreras 21171 (F). CHIMALTENANGO:Tecpan, 6 May 1937, Johnston 653 (F); Tecpan, 9000 ft, 24 Mar 1941, Johnston 1854 (F); Rio Los Indolos, 10 km from Godinez, 2000 m, 21 Sep 1971, Molina R. & Molina 26712 (F, U); Chichavac, 2400-2700 m, Nov-Dec 1930, Skutch 74 (US); Chichavac, 10 Jul 1933, Skutch 383 (US); Cerro de Tecpan, region of Santa Elena, 2400-2700 m, 26 Dec 1936, Standley 61064 (F, US); Cumbre de Chicoy above Tecpan, 3000 m, 15 May 1948, Williams 14221 (F). GUATEMALA: S.loc., 1940, Aguilar 494 (F); Aguilar 691 (F); slopes of Volcdn de Pacaya, between San Francisco Sales and base, 1800-2300 m, 20 Dec 1940, Standlev 80730 (F). HUEHUETENANGO: Along rd. to Huehuetenango,5 mi. S of San Juan Ixcoy, 9200 ft, 4 Feb 1965, Breedlove 8518 (F); Sierra de los Cuchumatanes, trail from San Mateo Ixtatan to aldea of Hok'ante, Yolk'ultak Chemte', 2900 m, 15 May 1965, Hopkins 10 (US); near Todos Santos, 2470 m, 27 Sep 1944, Melhus & Goodman 3621 (F); Sierra de los Cuchumatanes, rd. to San Juan Ixcoy, 3000 m, 18 Nov 1967, Molina R. 21306 (F, NY); between Jacaltenango and San Martin, 5300-7000 ft, 24 Dec 1805, Nelson 3605a (F); from Todos

TAXONOMIC TREATMENT Santos, 10000 ft, 26 Dec 1805, Nelson 3631 (F); San Juan Atitan, 8400 ft, 9 Sep 1934, Skutch 1170 (F, US); vic. Nucapuxlac, 2500 m, 17 Jul 1942, Steyermark48942 (F); Cruz de Lim6n, between San Mateo de Ixtatan and Nuca, Sierra de los Cuchumatanes, 2600-3000 m, 31 Jul 1942, Steyermark 49866 (F); top of Cerro Chemalto, Sierra de los Cuchumatanes, 3.5 mi W of Santa Eulalia, 3100-3150 m, 2 Aug 1942, Steyermark 49945 (F, NY). JALAPA: Volcan Jumay, N of Jalapa, 1300-2200 m, 1 Dec 1939, Stevermark 32346 (F); between Buena Vista and Miramundo, MontainaMiramundo, between Jalapa and Lago Ayarza, 2000-2200 m, 6 Dec 1939, Ste yermark 32825 (US). QUEZALTENANGO: Near Quezaltenango, 29 Oct 1965, Andrews 502 (NY); slopes of Volcan Zunil, at and above Aguas Amargas, 24302850 m, 17 Feb 1939, Standley 65388 (F, US); Volcan de Zufiil, at and above Aguas Amargas, 2430-2850 m, 17 Feb 1939, Standley 65436 (F); above Olintepeque, 2550-2850 m, 20 Feb 1939, Standley 65984 (F, US); Volcan de Santa Maria, above Palojunoj, 2400-3768 m, 6 Mar 1939, Standley 67578 (F); SE of Palestina, along rd. to San Juan Ostuncalco, ca. 2600 m, 21 Jan 1941, Stan7dlex'84361 (F); El Pocito, S of San Martin Chile Verde, on rd. to Colomba, ca. 2200 m, 27 Jan 1941, Standlev,84955 (F, US), 85047 (F, US). QUICHE:S.loc., 1942, Aguilar 1312 (F); Nebaj, ca. 10 km W, 8000 ft, 3 Jul 1964, Contreras 5170 (F). SAN MARCOS: Finca Armenia, San Rafael Pie de Cuesta to Carrizal, past Finca Africa, 1300-1600 m, 9-12 Aug 1980, Dwyer 15337 (MEXU, NY); rd. from Talquian to summit of Volcan Tacana, 2500 m, 19 Oct 1985, Martinez S. et al. 14080 (NY); above Rio Tacana, near San Antonio, ca. 270 m, 22 Feb 1939, Standley 66177 (F); El Boquer6n, near Quezaltenangoborder, ca. 2700 m, 22 Feb 1939, Standley 66306 (F, US); above Barranco Eminencia, ca. 2700 m, 14 Mar 1939, Standley 68503, 68538 (F); Barranco Eminencia, San Marcos-San Rafael Pie de Cuesta rd., between Finca La Lucha and Buena Vista, 2500-2700 m, 6 Feb 1941, Standley 86218 (F, US); Barranco Eminencia, rd. between San Marcos and San Rafael Pie de Cuesta, between Finca La Lucha and Buena Vista, 2500-2700 m, 6 Feb 1941, Standley 86231 (F); Volcan Santa Maria between Santa Maria de Jes6s, Las Mojadas and summit, 1500-3000 m, 12 Jan 1940, Steyermark 33966 (F); Volcan Zunil, above Fuentes Georgina, 2500-3800 m, 22 Jan 1940, Steyermark 34668 (F); Volcan Tajamulco, along rd. between San Sebastian and km 21 and km 8. 8-18 mi. NW of San Marcos, 27003800 m, 15 Feb 1940, Steyermark 35614 (F); Volcan Tacana, along Quebrada Canjula, between Sibinal and Canjula, 2200-2500 m, 18 Feb 1940, Steyermark 36006 (F); Volcan Tacana, Rio Vega near San Rafael and Guatemala-Mexico Border, 2500-3000 m, 20 Feb 1940, Steyermark 36229 (F); NW slopes of Volcan Tajamulco, between towns of Tajamulco and Tecutla, 9 mi S and W of Tajamulco, 1800-2500 m, 27 Feb 1940, Steyermark 36790 (F, US). SOLOLA: 15 km S of Panajachel, S of Lake Atitlatn,ca. 2200 m, 8 May 1972, Burch 5948 (F); 5-10 km W of Los Encuentros, Cerro Maria Tecum, Sierra Madre, 2900-3100 m, 24 Dec 1972, Williams et

267 al. 41731 (F, NY); Sierra Madre, Cerro Maria Tecum, 2-10 km W of Los Encuentros, 2800-3400 m, 7 Dec 1963, Williams et al. 25422 (F, NY, US). SUCHITEPEQUEZ: Volcan Santa Clara, between Finca El Naranjo and upper slopes, 1250-2650 m, 23 May 1942, Steyermark 46753 (F). TOTONICAPAN: Maria Tecun, 3800 m, 21 Nov 1965, Molina R. 15926 (F, NY); cloud forest of Maria Tecun, 3000-3600 m, 12-23 Jan 1966, Molina R. et al. 16367 (F, NY); Maria Tecun, 30003300 m, 15 Jan 1974, Molina R. et al. 30390 (F); Cumbre del Aire, on rd. between Huehuetenango and Sija, 3000-3300 m, 14 Jan 1939, Standley 62657 (F, US); 20 Feb 1939, Standley 65912 (F); 11 Jan 1941, Standley 83096, 83105 (F); region of Chui-quisis, above Totonicapan, on rd. to Desconsuelo, 2500-2800 m, 23 Jan 1941, Standley 84402 (US); SE of Totonicapan, 9600-9800 m, 27 Jun 1962, Webster et al. 11788 (F).

HONDURAS.

COMAYAGUA:

25 km NE of

Comayagua, Quebrada El Agua Helada, 1600 m, 13 Apr 1985, H.J. Ramos 183 (NY). INTIBUCA: Quebrada del Pel6n de Guise, 1600 m, 9 Apr 1956, Molina R. 6392 (F); between Calaveras and El Durazno in Cordillera Opalaca, 1800 m, 12 Mar 1970, Molina R. & Moli/mai 25551 (F, NY, US); vic. of La Esperanza and Intibuca, 1500-1600 m, 31 Jan-12 Feb 1950, Standley 25622 (US). LA PAZ: Montafia Verde, Las Marias Cordillera de Guajiquiro,2100 m, 23 May 1964, Molina R. & Molina 14045(6) (F, NY, US); Montafia Verde, Cordillera Guajiquiro, 1900 m, 23 Mar 1969, Molina R. & Molina 24378 (F, NY). MORAZ.AN:CerroLa Tigra,rd. to El Rosario, 3 Mar 1952, Carlson 2613 (F); Rancho Quebrada, rd. to El Rosario, San Juancito, 3 Mar 1952, Carlson 2629 (F); Cerro Cholombo ("El Tigra"),22 km NE of Tegucigalpa, 7200 ft, 28 Nov 1976, Fryxell 2852 (F); Mt. San Juancito. 6500 ft, 18 Jul 1948, Glassman 1984 (F); ParqueNacional La Tigra, 30 km NE of Tegucigalpa, 1200 m, 22 Mar 1985, L. Martinez 204 (NY); La Vuelta de Caite NE of San Juancito, Montafia La Tigra, 2300 m, 17 Mar 1957, Molina R. 7733 (F, US); Montafia La Tigra, SW of San Juancito, 2000 m, 23 Apr 1964, Molina R. 13741 (F, NY, US); La Tigra, SW of San Juancito, 1800-2100 m, 2 Feb 1966, Molina R. et al. 16996 (F, US); E slopes of Pefia Blanca, San Juancito mtns., 1900-2000 m, 22 Mar 1951, Morton 7244 (F, US); W slopes of Pefia Blanca. San Juancito mtns., 1800-2100 m, 25 Mar 1951, Morton 7457 (F, US); Montafia La Tigra, 20 km NE of Tegucigalpa, 1600 m, 8 Apr 1984, I. Padilla 172 (NY): Parque Nacional La Tigra, 30 km NE of Tegucigalpa, 1000 m, 23 Feb 1985, S. Palma 308 (NY); slopes of Cerro de Uyaca, ca. 1500 m, 25 Nov-5 Dec 1946. Standley & Williams 784 (F); SW of San Juancito, 2000 m, 21 May 1947, Williams& Molina R. 12797 (F); mtns. above San Juancito, 2000 m, 6 Nov 1947, Williams & Molina R. 13320 (US); mtns. above San Juancito, 1600 m, 20 Feb 1948, Williams& Molina R. 13715 (F); mtns. above San Juancito, 2000 m, 25 Mar 1948, Williams & Molina R. 13972 (F); near summit of San Juancito mtns., 2000 m, 22 Mar 1951, Williams 17415 (F, US).

Local names. Mexico. Chiapas: shinte, shintez; Michoacan: hierbaamargosa. Guatemala.Chimaltenango:

268

FLORA NEOTROPICA

duraznillo, huele de noche; Quezaltenango: aurella; with thickenedwhite margins, 1-2 mm long, densely Quiche:tzetze. to sparselypubescentwith arachnoidtrichomes;corolla Solanumnigricansis superficiallysimilarto and white or pale blue (fide Solomon 5973), fleshy, 1.2-2 probablythe sister species of S. maturecalvansof the cm diam., lobed nearly to the base, the lobes planarat Andes, sharingwith that species patchy arachnoidpu- anthesis, abaxial surfacesof lobes sparselypubescent bescence anddifoliate,geminatesympodialunits.Like with arachnoidtrichomes,the marginsandtips densethe rest of the species in this group, S. nigricans oc- ly papillose; anthers 3-5 mm long, 1-2 mm wide, cursat middleto high elevations,where it is apparently poricidal at the tips, the pores teardropshaped; free a common element of the oak-pine forests of southem portionof thefilaments ca. 0.5 mm long, the filament Mexico. The apically pointed fruitand mattedpubes- tube ca. 0.5 mm long, glabrous;ovarv glabrous;stvle cence makeS. nigricanseasily distinguishablefromany 5-8 mm long, glabrous;stigma clavate, bilobed, the surfaceminutely papillose. Fruit an apically pointed, other solanumgrowing in similarhabitats. globose,greenberry, 1-1.5cmdiam.;fruitingpedicels Variationinpubescenceis quitemarkedinSolanum nigricanswith some specimenshavingloose, dendritic glabrous, woody and deflexed, 1.5-3.5 cm long, ca. trichomesthatdryreddish(reminiscentofS. callianthum 1.5 mm diam. at the base, 3-5 mm diam. at the apex. of Colombia),ratherthanthemoretypicalmatted,arach- Seeds tanor reddish-brown,ovoid-reniform,4-4.5 mm noidpubescence.ThetypespecimenofS. brachystachys long, 3-3.5 mm wide, the surfaces minutely pitted. comes from this end of the range of variation,but all Chromosomenumbernot known. intermediatesexist across the species range. Distribution (Fig. 146). Near tree line in the Andes of S Peru and N Bolivia, from 3000-3500 m. 84. Solanum ochrophyllumVanHeurck& Mull. Arg., Observ.Bot. 50. 1870. Type.Bolivia. La Paz: Prov. Larecaja,vic. Sorata,9500-11000 m,Mandon 416 (lectotype, G [Mortonneg. 8538], here designated; isolectotypes, BM, F, G [F neg. 23138], G-DC, GH, K, MPU, NY, P [Mortonneg. 8273]). Not in AWH fide von Steenis. Fig. 147

Specimens examined. PERU. Cuzco: Paucartambo-Tres Cruces, Cerro de Cusilluyoc. 32003400 m, 2-6 May 1925, Pennell 14152 (F, NY). BOLIVIA. S.loc., Bang 1931 (F, GH, K, NY). COCHABAMBA: Prov. Chapare, 23.8 km N of Colomi, jct. of the rd. to Candelaria, on rd. to Chapare, then 2.2 km NW on side rd., upper Rio Cayani, 2700 m, 17010'S, 65?53'W, 19 Oct 1985, Solomon 14377 (MO, NY, U). LA PAZ: Vic. Sorata, Nov 1892, Bang 1630 (A, F, GH, Shrubs or small trees, 2-5 m tall; young stems K, NY, US); Prov. Nor Yungas, Unduavi, 3200 m, Feb and leaves densely pubescentwith felt-like arachnoid 1914, Buchtien 462 (F, K, NY); Not Yungas, Unduavi, trichomes, these drying pale golden brown; bark of 12 Feb 1907, Buchtien 760 (US); Prov. Murillo, Carretera older stems glabrate, grayish, the stems lightly four- Fundamental 3 toward Nor Yungas, ca. 17.3 km NE of angled.Sympodialunitsdifoliateor trifoliate,not gemi- El Cumbre, 4 km SW of Unduavi, 5 Nov 1976, Davidson nate.Leaves elliptic, widest at the middle, stronglybi- 4767a (NY); Sorata-Consata, 3250 m, 7 Mar 1982, colored,glabrousandshinyadaxially,occasionallywith Fernandez Casas & Molero 6558 (NY); Ilabaya on a few arachnoidtrichomes along the main veins, the Sorata-Achacachi rd., ca. 3500 m, 16 Mar 1980, Feuerer abaxial laminacompletely covered with densely mat- 9471a (NY); Prov. Nor Yungas, Unduavi valley, 3530 ted arachnoidtrichomesdryingpale goldenbrown,with m, 6 Mar 1982, Feuerer 10148a (NY); Sorata-Tacacoma 10-16 pairsof main lateralveins, impressedadaxially, rd., 10 km from Sorata, 3300 m, 11 Mar 1982, Feluer-er 10358a (NY); valley or Rio Kuriwaya above Sorata on glabrous,prominentandyellowish abaxially,4-15 cm way to Apacheta, 3440 m, 9 May 1985, Felnerer 23027 long, 1.5-7 cm wide, the apex acute, the base acute, (NY); Sorata, Apr 1919, Gunther 12307 (US); Prov. somewhat decurrentonto the petiole; petiole 0.5-1.5 Inquisivi, along trail between Loma El Abra and Cerro cm long. Inflorescences opposite the leaves or intern- Negro Khota, ca. 6 km N from Inquisivi, 2900-3000 odal, simple, 0.5-1.5 cm long, 5-10-flowered, dense- m, 16?50'S, 67?08'W, 22 Aug 1988, Lewis [8811106, ly pubescentwith mattedarachnoidtrichomes;pedicel [8811128 (MO); Prov. Sud Yungas, ca. 1.8 km WSW of scar-sclosely spaced, not overlapping.Buds globose, Unduavi, 3300 m, 16019'S, 67055'W, 29 Oct 1984, Nee becoming ellipsoid just before anthesis, the corolla & Solomon 30190 (MPU, NY); Unduavi, 10000 ft, Oct strongly exserted from the calyx tube.Pedicels at an- 1885, Ruisby777 (NY); Prov. Nor Yungas, 3.9 km W of thesis ratherthickandapparentlyfleshy,deflexed, 1.3- bridge at Unduavi on old rd., ca. 3400 m, 16'19'S, 67?58'W, 10 Aug 1981. Solomon 5973 (MO. NY); 1.4 1.6 cm long, ca. 1 mm diam.at the base, 2-3 mm diam. km W of bridge at Unduavi on old rd., ca. 3250 m, atthe apex,denselypubescentwith arachnoidtrichomes 16?19'S, 67?52'W, 10 Aug 1981, Solonion 5980 (K. MO, like the rest of the inflorescence.Flowers with the ca- NY); Prov. Murillo, 20.3 km N of dam at Lago Zongo, lvxtubeconical,2-3 mm long, the lobes broadlydeltate 3250 m, 16?10'S, 68'08'W, 5 Mar 1983, Solomon 9720

TAXONOMIC TREATMENT

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270

FLORA NEOTROPICA

FIG. 148. Solanum platycypellon S. Knapp. (Reproduced with permission from Brittonia 44: 66, fig. 5. 1992.)

(MO, NY); Prov. Sud Yungas, 4.7 km SW and above Unduavi, 3500 m, 16?l9'S, 67?56'W, 12 Feb 1984, Solomon & Kuijt 11516 (MO, NY); Prov. Murillo, 2.6 NE and below Pongo, 3500 m, 16?19'S, 67?56'W, 8 Mar

of S. maturecalvans,but the density of trichomes on the lower leaf surfacesandthe non-geminatesympodia areclear differentiatingfeatures. Two specimens were cited by Van Heurck & 1984, Solomon& Stein 11672(MO,NY); Prov.Murillo, Muller in the protologue, one from VanHeurck'sown 20.8 kanN of La Cumbrein valley of Rio Zongo, 3200 m, 16?09'S, 68?07'W, 20 Feb 1987, Solomon 16124 (K, NY, herbarium,now at AWH, and the other from G. No U); valley of rio Zongo, 21.1 km N of the pass, 3200 m, specimenof this numberor species exists at AWH(van 16?09'S, 68?07'W, 4 Apr 1987, Solomon 16441 (MO, Steenis, in litt.), so the G-DC sheetbecomes the logical NY); Cordillera Real, Pongo, 12000 ft, 17 Feb-l Mar choice for a lectotype. 1926, Tate210 (NY); Cordillera Real, top of pass on the Tipuani-Ancoma-Sorata trail down to Sorata, 10000-17200 ft, 30 Apr 1926, Tate 791 (F, NY).

Solanum ochrophyllumis one of the most distinctive species of sect. Geminata with its bicolored leaves and closely and densely mattedpubescence on the abaxialleaf surfaces.Plantsarequitesimilarto those

85. Solanum platycypellon S. Knapp, Brittonia 44: 65. 1992.Type.Bolivia.Tarija:15.8km E ofNarvaez on rd. to Entre Rios (12.5 km W of Entre Rios), 1650 m, 21028'S, 64012'W,4 Oct 1983, Solomon 11045 (holotype,NY; isotype, MO). Fig. 148

TAXONOMIC TREATMENT

Shrubsca. 2 m tall;youngstemsandleavesdensethe trichomesfine, delicateand ly arachnoid-pubescent, dendritic,the leaves and stems soon glabrate;barkof older stems darkreddish-brown.Sympodialunitsunifoliate.Leaivesellipticto narrowlyelliptic,widest at the middle, with 9-12 pairs of main lateralveins (reddish abaxially), glabrous and shiny adaxially, with a few arachnoid trichomes abaxially, especially along the midrib,9-14 cm long, 4-5 cm wide, the apex acute to acuminate,thebase attenuate;petioles 1.5-1.6 cm long. Inflorescences opposite the leaves, simple, 1-1.3 cm long, sparsely pubescent with floccose arachnoidtrichomes alongthe axis, especiallynearthe apex;pedicel scars closely spaced, not overlapping.Buds rounded whenyoung, laterellipsoid,very stronglyexsertedfrom the calyx tube. Pedicels at anthesis slender, more or less deflexed, 1.5-2.5 cm long, ca. 0.5 mm diam., glabrous or occasionally sparsely pubescent with arachnoid trichomes. Flowers with the calyx tube broadly cup-shaped, nearly flat, 1-1.5 mm long, the lobes minute, broadly deltate, 0.25-0.5 mm long, very sparselypubescentwith arachnoidtrichomes;corolla white, 0.8-1.5 cm diam., divided ca. 3/4 of the way to the base, the lobes planarat anthesis,the tips andmargins of the lobes minutelypapillose;anthers3-3.5 mm long, ca. 1 mm wide, poricidalatthe tips, the poresteardropshaped;free portionof thefilamentsca. 0.25 mm long, the filament tube ca. 0.25 mm long; ovary glabrous;style straight,ca. 5 mm long;stigma clavate,the surfaceminutelypapillate.Fruit andseeds not known. Chromosomenumbernot known. Distribution (Fig. 146). In southemBolivia and northwestemArgentina,in subtropicalmoist forest at ca. 1600-1700 m elevation.

271

Shrubsor treelets, 1-2 m tall; young stems and leaves densely pubescent with simple, uniseriate trichomes 0.5-1 mm long, the stems erect;barkof older stemsgrayish-brown. Sympodialunitsunifoliate.Leaves elliptic to narrowlyelliptic, widest at the middle, with 6-9 pairs of main lateral veins, thick and somewhat coriaceous, glabrousadaxially,densely to moderately pubescent abaxiallywith simple uniseriatetrichomes 1-1.5 mm long, these denser along the veins; lamina 4-15 x 1.5-5 cm, the apex acuteto acuminate,the base acute, somewhatdecurrentonto the petiole; petiole 11.5 cm long.Inflorescencesoppositethe leaves, simple, 3-5 mm long, 5-8-flowered, densely pubescent with simple uniseriatetrichomeslike those of the stems and leaves;pedicel scars closely spaced,overlapping.Buds globose, laterellipsoid, stronglyexsertedfromthe calyx tube just before anthesis.Flowers with the calvx tube broadly conical, 1-1.5 mm long, the lobes elongate, deltate, with the tips rounded, 1-2 mm long, sparsely pubescent with simple uniseriatetrichomes; corolla white, 1-1.2 cm diam.,lobed ca. 3/4 of the way to the base, the lobes planar(?) at anthesis,the tips and margins of the lobes densely pubescent with simple trichomes;anthers4.5-5 x 1-1.5 mm, poricidalat the tips, the pores teardropshaped;free portionof thefilaments ca. 0.5 mm long, the filament tube ca. 0.5 mm long; ovary glabrous;style 6-7 mm long; stigma capitate andminutelybilobed,the surfacepapillose.Fruita globose, greenberry,1-1.2 cm diam.;fruitingpedicels erect,woody, 1.7-2.7 cm long, ca. 1.5 mm diam.at the base, ca. 2 mm diam. at the apex. Seeds darkbrown, ovoid-reniform, 3-4 x 2-2.5 mm, the surfaces very smooth, the embryo clearly visible throughthe testa. Chromosomenumbernot known.

Specimensexamined.ARGENTINA.SALTA: San Distribution (Fig. 146). Cloud forests in the Andresbei Oran,24 Sep 1873, Lorentz& Hieronymus VenezuelanAndes in the statesof MeridaandTrujillo, 255 (B [destroyed:F neg. 2765], NY). from 2200-3200m. Solanumplatycypellon is closely related to S. Specimens examined. VENEZUELA.MERIDA: maturecalvans,also of Andean distribution.It differs Monte Zerpa, Paramo de los Conejos, 3100-3200 m, 16 from that species in its unifoliate sympodial units, May 1964, Bernardi 1255 (NY); San Javier valley, along smaller flowers, and flatter, more open calyx tube. Quebrada La Cuesta toward Monterrey, 2500-2650 rn, Solanummaturecalvansis a plantof higherelevations, 10 Dec et al. 48 (NY, US). TRUJILLO: 1983, Weitzmann while S. platycypellonoccurs in subtropicalmoist for- Old rd. from Bocon6 to Trujillo, ca. 51 km W of Trujillo, ests at somewhat lower elevations. below summit, 2200-2250 m, 9?21'N, 70?19'W, 20 Oct The araclnoid pubescence typical of members 1984, Knapp& Mallet 6778 (BH, K, MY, VEN). of the Solanum nigricans species group is very soon Solanumquebradetnse is superficiallyvery simideciduousin S.platycypellon,but is always presenton lar and probablyclosely related to S. callianthum of the new growth and the tips of the inflorescence axes. high-elevation Colombia. The uniseriatepubescence sharedby these two taxa differs from the more common mattedpubescencefoundin othermembersof the 86. Solanum quebradense S. Knapp, Novon 6: 29. S. nigricans species group. Solanumquebradensecan 1996.Type.Venezuela.Mrida: Mucuruba,quebrada be distinguished from S. callianthum by its simple neartown, ca. 2600 m, 25 Jun 1930, Gehriger244 ratherthandendriticpubescence,narrowerleaves, gla(holotype, NY; isotypes, F, MO). Fig. 149 brous ovaries, and smaller, less fleshy flowers. Fruits

272

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stems andleaves, the trichomesdendritic;pedicelscar-s closely spacedor unevenly spacedandca. 5 mm apart. Flowers white, usually somewhat fleshy, the corolla lobes planarat anthesis.Fruit greenor greenish-yellow XIII. Solanum arenarium species group (S. andhardat maturity,the pericarpoccasionallythinand arenariunm,S. gnaphalocarpon, S. smithii, S. brittle in dry specimens, glabrous or densely pubestunariense). Fig. 136E,F cent with dendritic trichomes (S. gnaphalocarpon); fruiting pedicels woody, deflexed. Seeds very large, Small shrubs;young stems and leaves sparsely ovoid-reniform,pale tan or yellowish. to densely pubescent with dendritictrichomes, these Distribution. Andes in Ecuador,Peru,Bolivia, often very densely branched.Sympodialunitsunifoliate or more commonly difoliate and geminate.Leaves and SE Brazil. lanceolateto elliptic, occasionally shiny adaxially,puMembers of the Solanum arenarium species bescent abaxiallywith dendritictrichomes,these con- group are morphologically quite reminiscent of S. fined to the vein axils or densely covering the surface, pseudocapsicum,but the similarityis merely superfithe apex andbase various.Inflorescencesopposite the cial. The large,smooth,pale, andovoid-reniformseeds leaves or occasionally internodal on short shoots are distinctive in this group.All of the species grow in (Solanumn smithii), simple, densely pubescent like the dry forests, often along streamsin gallery forest. of Knapp& Mallet 6778 were very pale green in color. The type collection is the only one bearingflowers.

Key to the species of the Solanum arenariumspecies group 1. Fruit densely pubescent with dendritic trichomes; leaves narrowly elliptic, densely pubescent on both surfaces ............ 88. S. gnaphalocaipon 1. Fruit glabrous or at most with a few scattered trichomes; leaves elliptic to ovate, densely to sparsely pubescent only abaxially. 2. Trichomes covering the abaxial leaf surface; inflorescence pubescent with dendritic trichomes. 3. Sympodial units unifoliate or difoliate, geminate; leaves 7-16 x 3-8.5 cm. SE Brazil.... 87. S. arenariulm 3. Sympodial units plurifoliate, never geminate; leaves 1-11 x 0.7-4 cm. Dry forests of E Bolivia ... 90. S. tunariense 2. Trichomes confined to the axils of the main lateral veins on the abaxial leaf surface; inflorescence S. smithii glabrous .......89.

87. Solanum arenarium Sendtn.in Mart.,Fl. Bras. 10: 26. 1846. Type.Brazil."InBrasiliaaustrali,"Sellow s.n. (lectotype, BR, here designated). Fig. 150 Solanum chloranthum Spreng., Syst. Veg. (ed. 16) 1:

682. 1814. Type.No type materiallocated.Nom. dub. As "chloranthon." Solanum sprengelii

Dunal in DC., Prodr. 13(1): 372.

1852.Type.Brazil.in Brasilia,Zeyhers.n. (Specimen not located: nom. nov. for Solanum chloranthlum Spreng.)

Shrubs 0.5-2 m tall; young stems and leaves densely pubescentwith beige-tan dendritictrichomes 0.25-0.5 mm long, the older stems not glabrate,the young leaves dryingdark;barkof older stems reddishbrown. Symnpodialunits unifoliate, occasionally difoliate,geminate.Leaveselliptic,widestatthe middle, with 7-9 pairsof main lateralveins, glabrousandshining adaxially,with a few scatteredtrichomesalong the midrib, densely pubescent abaxially with congested dendritictrichomes,the laminanot visible throughthe interlockingtrichomes,7-16 x 3-8.5 cm, the apexacute to rounded,the base attenuate;petioles 0.6-1 cm long.

Inflorescencesleaf-opposed,simple,0.2-1 cm long, 26-flowered,denselypubescentwith dendritictrichomes like those of the young stems andleaves;pedicel scars closely spaced,not overlapping.Buds globose, the corolla stronglyexsertedfromthe calyx tube.Pedicels at anthesis slender, 0.7-1 cm long, deflexed, densely pubescentwith dendritictrichomesca. 0.25 mm long. Flowers with the calyx tube conical, 1-1.5 mm long, the lobes deltatewith an apicalknob-likeprojection,11.5 mm long, the lobes pubescent with dendritictrichomes, the midribs densely pubescent, the margins glabrousor sparselypubescent;corollagreenish-white, 6-9 mm diam.,lobed ca. 3/4 of the way to the base, the lobes planar at anthesis, sparsely pubescent on the abaxial surface;anthers 2-2.5 x 1-1.5 mm, poricidal at the tips, the pores teardropshaped; free portion of thefilaments ca. 0.25 mm long, the filament tube ca. 0.25 mm long; ovaryglabrous;style 5-7 mm long, glabrous;stigma a minute papillose area at the tip of the style.Fruita globose,greenbeny, 1-1.5 cm diam.;fruiting pedicels deflexed, woody, 1.3-1.5 cm long, ca. 1 mm diam.at the base.Seeds pale tan,ovoid-reniform,4-4.5

274

FLORA NEOTROPICA

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x 2.5-3 mm, the surfacesminutelypitted.Chromosome tenuate;petioles 3-7 mm long; minorleaves differing from the majorsin size and often shape, elliptic to ornumber unknown. bicular,1-2.3 x 0.7-1.7 cm, the apex roundedto acute, Distribution (Fig. 153). In forest and second the base acute; petioles 1-2 mm long. Inflorescences growth in southem Brazil from Rio de Janeiroto Rio opposite the leaves, simple, 2-9 mm long, 2-5-flowGrandedo Sul, elevation unknown. ered, densely pubescent with closely branched triSpecimensexamined.BRAZIL.Hort.Berol.,coll. chomeslike those of the stemsandleaves;pedicelscars 1837 (B-n.v. (F neg. 2806)); s.loc., Burchell345 (K); closely spaced but not overlapping. Buds globose, later cult. Kew, 1882, Anon. s.n. (K). RIO DE JANIERO: P.N. obovoid, the corolla strongly exserted from the calyx Tijuca, Pedrade Gave, between Iposeir and Chamine tube. Pedicels at anthesis probablydeflexed, thin and Hely, 24 Feb 1978, Carautaet al. 2850 (RB); Rio de tapering,4-5 mm long, ca. 0.5 mm diam. at the base, Janiero, Mar-May 1832, Riedel 399 (NY, US). Rio densely pubescent like the rest of the inflorescence. GRANDE DO SUL: Morro das Abertas, 26 Mar 1980, Flowers with the calyx tube conical, 0.5-1 mm long, Aguiar& Martau248 (F); cultivatedin I.A.S. Botanical Garden,Pelotas,9 May 1955, CostaSacco 391 (F, NY); the lobes triangular,0.1-5 mm long, densely pubesvic. Sao ILeopoldo, Nov 1941,Leite 659 (NY); Rio dos cent with closely brancheddendritictrichomes;corolla white, 5-7 mm diam., lobed 3/4 of the way to the base, Sinos, 8 Nov 1949, Rambo44290 (GH, US). the lobes planarat anthesis, the abaxial surface of the Solanum arenarium is probably most closely lobes pubescentwith minute dendritictrichomes;anrelatedto S. gnaphalocarpon,also of SE Brazil. It difthers 2-2.5 mm long, poricidal at the tips, the pores fers fromthatspecies in its stronglydiscolorouslarger teardropshaped;free portionof thefilaments ca. 0.25 leaves, glabrous fruit, and smaller flowers. The trimm long, the filamenttubeca. 0.45 mm long, glabrous; chomes of the lower leaf surface in S. arenariumare ovarydenselypubescentwith dendritictrichomes;style so closely packed as to obscurethe lamina.Label data 4-4.5 mm long, densely pubescent at the base with indicatethatthe flowers ofS. arenariumarepale green, dendritictrichomes like those of the ovary, glabrous while those of S. gnaphalocarpon, S. smithii, and S. and somewhatclavate distally;stigma a minutepapiltunarienseare white. lose areaon the tip of the style. Fruit a globose, green, The epithet Solanum chloranthumSprengel is densely pubescent berry, 1-1.3 cm diam., the pubesthe oldest availablefor this species, andBitter(1920b) cence consisting of closely branchedtrichomes ca. 1 considered it as such. However the materialcited by mm long, the branchesvery shortandcongested;fruitBitteras type materialwas cultivatedin Berlinafterthe ingpedicels somewhat deflexed, 6-9 mm long, ca. 1 publication date of the epithet. Thus the identity of mm diam. at the base. Seeds pale brown, ovoid-reniSprengel's species is in doubtand the name should be form with somewhat incrassatemargins,3-4 x 2.5-3 considereda nomen dubium. mm, the surface minutely pitted, the cells sinuate in outline. Chromosomenumberunknown. 88. SolanumgnaphalocarponVell.,Fl. Flumin.82. 1829 Distribution (Fig. 153). In woods, secondary [ 1825].As "Solanumgnaphalocarpus." Type.Brazil. growth, and open areas from 700-1300 m. Southern Riode Janeiro:"Crescitcampisapricusmediterraneis" Brazil in Minas Gerais, Rio de Janeiro,and Sao Pauilo (No specimensextant;lectotype,Vellozo,Fl. Flumin. states. Icones2: fig. 91 ("Solanumgnaphalocarpus"). 1831 [1827], here designated). Figs. 151, 152 Specimens examined. BRAZIL. S.loc., Glazioli Shrubs 1-1.5 m tall; young stems and leaves densely pubescentwith pale tan,closely brancheddendritictrichomes0.1-0.25 mm long; barkof olderstems reddish-brown, not glabrescent. Svmpodial units difoliate, geminate. Leaves narrowly elliptic to lanceolate, widest at the middle, sparselypubescentwith dendritictrichomesadaxially,thetrichomesdenseralong the main veins, pubescentwith closely brancheddendritictrichomesabaxially,the trichomes0.25-0.5 mm long, most of the trichomebranchesarisingin the distal 1/3 of the trichome, denser along the veins; major leaves with 9-11 pairs of main lateralveins, 7.5-12 x 1.5-4.5 cm, the apex acute to acuminate,the base at-

s.n. (F, P); s.loc., 1832, Lhotsky s.n. (G (F neg. 6786)). MINAs GERAIS: Serra do Cip6, 3 Feb 1958, Heringer 6334 (US); ca. 27 km SW of Diamantinaon rd. to Gouveia. 1300 m, 15 Jan 1969, Irwin et al. 22053 (F, MO, NY, US); rd. to Sao Miguel, km 4.5, 700 m, 5 Nov 1930, Mexia 5258 (F, MO, NY, US); s.loc., 1 May 1874. Mosen 1997 (MO, S); Lagoa Santa, Warming s.n. (C-n.v. (F neg. 22891), MPU). PARANA:Serra do Itatiaia, ca. 900 m, 23 Oct 1903, Dusen 2180 (GH); Serra Marumbi, trail to Olimpo, Mun. Morretes, 19 Jan 1995, Ribas et al. 775 (MBM, NY) . RIO DE JANEIRO:Therezopolis, 900 m, Sep 1929, Brade9207 (US); Teresopolis, granja Mafer, 28 May 1977, Carvalho 560 (BM, RB); Corcovado, 7 Mar 1880, Glaziou s.n. (F); environs of Rio de Janeiro, 1880, Glaziou 12107 (K); Organ Mtns., Miers 4534 (K); Nova

276

FLORA NEOTROPICA

FIG. 151. Solanum gnaphalocarpon veill plate 91 of Vellozo's Florae Fluminensis Icones, Vol. 2 (as S. gnaphalocarpus Veil.)

TAXONOMIC TREATMENT

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FIG. 153. Distribution of Solanum arenarium (solid triangles), S. gnaphalocarpon (open circles), S. smithii (solid circles), and S. tunariense (solid squares).

Friburgo,MorroCurusu,Pontede Saudade,ca. 1250 m, s.d., Pessoa et al. 160 (BM, RB); Corcovado,Oct 1832, Riedel 1078 (NY, US); Corcovado, Aug 1833, Riedel 1373 (NY,US);Parahyba do Sul, 1881,Schwackes.n. (US); Rio

dasFlores,FazendaSta.Genovesa,500-600m, 7 Oct 1971, Sucre 7771 (RB). SAOPAULO: Sao Josedo Rio Pardo,28 Sep 1889, Loefgren 1423 (Hoehne 15377) (US).

Solanumgnaphalocarponis superficiallysimilar to S.pseudocapsicum(S.pseudocapsicumspeciesgroup) in itsdendriticleafpubescenceandgeneralleafshape,but differs from thatspecies in many characters.Solanum pseudocapsicurm hasflattened-reniform seeds,largerflowers and fruit, and never has orbicularminor leaves. The trichomes on the fruit of Solanum gnaphalocarponaredistinctiveandfoundnowhereelse in sect. Geminata.They aretree-like,but with the axis much elongatedandthe generalaspectof the trichome approaching a stellate type morphology. In fruit, S. gnaphalocarpon is not easily mistaken for any other species of sect. Geminata.

89. Solanum smithii S. Knapp, Novon 2: 346. 1992.

Type. Ecuador.Loja: Ca. 30 km S of Catamayoon rd.to Cariamanga,1900-2000 m, 4?1O'S,79?20'W, 6 Feb 1984, Knapp & Mallet 6252 (holotype, NY; isotypes, AD, BH, F, G, GH, K, MEXU, MO, QCA, QCNE, US). Figs. 136E,F, 154

Shrubsto 1 m; young stems and leaves glabrous to densely red-papillose, the branches erect; bark of older stems darkreddish-brown.Sympodialunits not geminate, difoliate or plurifoliate, with many short shoots. Leaves elliptic, widest at the middle, occasionally thick and somewhatcoriaceous,glabrousadaxially, pubescentwith tufts of dendritictrichomes ca. 0.5 mm long in the axils of the main lateralveins abaxially, with4-6 pairsof mainlateralveins; lamina5-14 x 1.55.5 cm, the apex acute to rounded,the base attenuate, minutely winged onto the petiole; petiole 0.5-1.5 cm long. Inflorescences internodal or on short shoots, simple, glabrous,1-5 cm long, 6-10-flowered;pedicel scars unevenly spaced 3-5 mm apart.Buds elongate and somewhatpointedwhen young, laterellipsoid and strongly exserted from the calyx tube.Pedicels at anthesis thick and fleshy, erect, 0.9-1 cm long, ca. 1 mm diam. at the base, ca. 1.5 mm diam. at the apex. Flowers withthecalyxtubeconical, 1.5-2 mm long,the lobes deltatewith a smallroundedknobat the apex, 1-1.5 mm long, papillose, the margins bearing a few simple or dendritic trichomes; corolla white, fleshy, 1.5-2 cm diam., lobed nearly to the base, the sinuses thin and membranous, the lobes planar at anthesis, tips and marginsdensely papillose;anthers4.5-5 x 2-2.5 mm, poricidal at the tips, the pores teardropshaped; free portion of thefilaments ca. 0.5 mm long, the connective somewhat enlargedat the base, the filament tube

TAXONOMIC TREATMENT

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ca. 1 mm long, glabrous;ovary glabrous;style 6-7 mm long, glabrous; stigma strongly and divergently bilobed, the surfaceminutelypapillose.Fruit a globose, green to greenish-whiteberry, 1-1.2 cm diam.;fruiting pedicels somewhat deflexed, woody, 1-1.3 cm long, 1-1.5 mm diam. at the base, 3-4 mm diam. at the apex, the calyx lobes reflexed in fruit.Seeds pale tan, ovoid-reniform,4-5 x 3-4 mm, the surfacesminutely pitted. Chromosomenumbernot known. Distribution (Fig. 153). In dryforestsandscrublands in the Huancabambadepressionof northernPeru and southernEcuador,from 1900 to 2600 m. Specimensexamined.ECUADOR.AZUAY:W of Gironon Cumbe-Giron-Sta.Isabel-Pasajerd., ca. 1850 m, 3?12.09'S,79?11.12'W, Knapp et al. 9065 (QCNE);vic. of Nab6n,26 Sep 1918,Rose et al. 23045 (US). LOJA: 10 km N of Saraguro,on LagunaHuaycu, 1200-2300 m, 3033'S,79014'W, 17 Mar 1989, Elleman 91699 (LOJA);Catacocha,2050 m, 4 Jul 1946,Espinosa E607 (LOJA); rd.to thecoast,20 Jun1984,Espinosa2110 (LOJA); rd. Loja-Zamora,betweenChinchesand Sambi, 1700-2400 m, 3 May 1974,Harling & Andersson 14257 (GB, on Catamayo-Machala MO);vic. Chaguarbamba rd., ca. 21 km W of Las Chinchas, ca. 1850 m, 3?56.83'S, 27 Oct 1994,Knapp & Mallet 9081 (QCNE); 79036.26'W, Sozoranga,Tacamoros,La Pozo, CruzGrande,2088 m, 8 Aug 1976, Vivar C. et al. 811 (LOJA). PERU. CAJAMARCA: Prov. Cajabamba,vic. of Cajabamba,2600 m, 18 Nov 1983, Sagastegui et al. 11258 (BM, HUT); Condebambavalley, CajabambaCajamarcard., 2100-2600 m, 7033'S,78?09'W,15 Feb & Vasquiez3409 (MO, NY, USM). 1983, Snmith Solanum sinithii is superficially similar to S. barbulatum(S. amblophyllumspecies group), also of highlandEcuadorand Peru,but differs from that species in its ovoid-renifor-mseeds and axillary tufts of dendritictrichomes.The closest relativeofS. smithiiis probably S. tunariense of Andean Bolivia. Solanunm smithii differs fromS. tunariensein having trichomes confined to small tufts in the axils of the main lateral veins on the abaxialleaf surfaces,in its somewhatlarger leaves, and in its longer fruitingpedicels.

FLORA NEOTROPICA

shoots.Leaveselliptic,widestatthemiddle,coriaceous, with revolute margins, 1-11 x 0.7-4 cm, the lamina glabrous or sparsely pubescent with golden dendritic trichomes,the veins densely pubescentwith dendritic trichomes adaxially, densely pubescent with golden dendritictrichomes abaxially, these giving the leaf a yellowish colorbeneath,the apex acute,the base acute; petioles 2-7 mm long, sulcateadaxially.Inflorescences opposite the leaves or intemodal, 0.4-1.2 cm long, simple, 2-8-flowered, densely pubescentwith golden dendritictrichomeslike those of the stems andleaves; pedicel scars closely spacedbutnot overlapping.Buds ellipsoid, laterobovate, stronglyexsertedfromthe calyx tube.Pedicels at anthesisfiliform, 1.2-1.3 cm long, hanging,ca. 1 mm diam. at the apex, ca. 0.5 mm diam. at the base, densely pubescent with golden dendritic trichomes.Flowers with the calyx tube conical, 0.751mm long,the lobes deltateto quadrate,0.5-1 mm long, with a minuteapicalprojection,denselypubescentwith golden dendritictrichomeslike the rest of the inflorescence; corolla white (?), 0.8-1 cm diam., lobed nearly to the base, the lobes reflexedat anthesis,the tips of the lobes minutely papillose; anthers 2.5-3 x ca. 1 mm, poricidal at the tips, the pores teardropshaped; free portionof thefilaments ca. 0.5 mm long, the filament tubeca. 0.5 mm long, glabrous;ovaryglabrous;style ca. 6 mm long, sparselypubescentwith minuteuniseriate golden trichomes, these usually simple but occasionally branched;stigma merely a papillose areaat the tip of the style. Fruit a globose, green (drying mustard yellow) berrywith thinpericarp,1-1.5 cm diam.;fuidtingpedicels deflexed, 1.4-2 cm long, ca. 0.5 mm diam. at thebase.Seedspale tan,ovoid-reniform,3.5-4 x 2.53 mm, the surfacesminutelypitted.Chromosonme numniber unknown. Distribution (Fig. 153). In moist to dry deciduous foreston the eastem Andeanslope in Bolivia, from 900-2550 m.

Specimens examined. BOLIVIA. COCHABAMBA: Sailapata, Ayopaya, 2700 m, Nov 1935, Cardenas 3261 (US). LA PAZ: Coripata,Yungas, 12 May 1894. Banig2188 (BM, K, NY, US); Milluguaya, Prov. Nor Yungas, 900 m, Dec 1917, Buchtien 4166 (GH, NY, US); Inquisixi, 3 km 90. Solanum tunariense Kuntze,Revis. Gen. P1.3(2): toward Circuata,ca. 2200 m, 20 Feb 1981, Beck 4464 (F); 228. 1898.Type.Bolivia.Cochabamba: Tunari-gebirge mule trail between Inquisivi and Rio Khatu bridge, ca. in derCinchonaregion,Apr-May1892,Kuntzes.n. 1-3 km N of Inquisivi, 2400-2550 m, 16?53'S, 67?09'W, 29 Mar 1989, Lewis 35378 (MO, U); Prov. Inquisivi, (holotype,B [destroyed:F neg. 2723]; lectotype,NY, canyon of Rio Khatu, 2 km by air N of Inquisivi on rd. here designated;isolectotype, US). Fig. 155 to Chorocona, 1200-2200 m, 16?53'S, 67?08'W. 12 Mar Shrubs1-3 m tall;young stemsandleaves dense- 1988, Nee & Solomon 36657 (IBE, NY, U).

ly golden pubescent with dendritictrichomes ca. 0.5 In describingSolanumtunariense,Kuntzeindimm long, these persistent;stemsthinanddelicate;bark catedthathe thoughtit was relatedtoS.pseudoctapsicumii, of older stems shiny and darkbrownish-black.Sympo- presumablydueto its possessionof dendritictrichomes. dial units difoliateorplurifoliate,plantswithmanyshort Thetrichomesarelike those ofS. pseudocarpsicum, but

281

TREATMENT

TAXONOMIC

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FLORA NEOTROPICA

282

even sterile specimens are easy to differentiate,due to the smaller,more ovoid leaves of S. tunarienseandthe darkshiny barkof maturestems. Solanum tunariense is most closely relatedto S. smithiiof the Huancabamba depressionin northernPeruandadjacentEcuador,differing from it mainly in the more copious pubescence of the abaxial leaf surfaces.

XIV. Solanum sessile species group (S. chlamydogynum, S. confertiseriatum,S. monadelphum,S. obovalifolium, S. oppositifolium, S. palmillae, S. rovir-osanum, S. sessile, S. triste, S. turgidum). See

Knapp(1991a) for a revision and Knapp (1991b) for a cladistic analysis of the group.

dritic;majorleaves usually differingin size and shape from the minors. Inflorescences overtopping shoots, laterlateral,often leaf-opposed, simple or more often branched,puberulentwith minute golden trichomes. Buds ellipsoid, fleshy. Flowers with the calyx tube abruptlynarrowingto the pedicel;corolla fleshy,white or greenish-white,the lobes planarat anthesis;oMarT andstyle glabrousor pubescent.Fruit a globose, green berry,hard at maturity,bome on erect (occasionally slightly deflexed) fruiting pedicels. Seeds pale tan, ovoid-reniform,the testa thin and minutely pitted; in two species the marginssomewhatincrassate. Distribution. Central America, Andes, Amazon basin.

The species of theSolanumsessile species group Shrubsor trees;youngstemsandleavesminutely are illustratedwith photographsof live plantsin Knapp golden-puberulent or glabrous. Sympodial units ( 199la). Species of the groupall shareminute golden difoliate, geminate, occasionally plurifoliate.Leaves pubescenceon the new growthandinflorescenceaxis, usually large, elliptic or obovate, glabrous or pubes- fleshy flowers (except S. oppositifolium),andusually cent with erect golden trichomes,these simple or den- branchedinflorescences.

Key to the species of the Solanumsessile species group 1. Leaves narrowly obovate or linear; shrubs of river courses, often growing near or in running water. 2. Young stems minutely puberulent; buds ellipsoid, exserted from the calyx tube; corolla 1-1.2 cm 96. S. palmillae diam. Veracruz, Mexico .......................................................... 2. Young stems minutely red-papillose; buds globose, later ellipsoid, enclosed in the calyx until just before anthesis; corolla 1.5-2 cm diam. E Peru.................................................. 93. S. monadelphurm 1. Leaves elliptic to obovate; habit not as above. 3. Stems and new growth with red-papillose trichomes; leaves obovate. 4. Leaves sessile; inflorescencemany-times branched,very large. E Peru, middle elevations..... 98. S, sessile 4. Leaves petiolate; inflorescence branched or occasionally simple. 5. Flowers 2-2.2 cm diam.; fruiting pedicels 1.5-2 cm long; calyx lobes accrescent and woody in fruit. Cordillera de la Costa, Venezuela.................................................... 94. S. obovalifoliun 5. Flowers 1.2-1.5 cm diam.; fruiting pedicels 0.8-1.3 cm long, thick and woody; calyx lobes not accrescent in fruit. Montane Costa Rica ....................................................... 97. S. rovlirosanurn 3. Stems and new growth minutely golden-puberulent; leaves various. 6. Leaves with tufts of trichomes in the axils of the main lateral veins and/or the trichomes along the midrib beneath or over the entire surface beneath, the trichomes usually branched. 7. Plants dioecious (the flowers on a plant all long-styled or all short-styled); flowers 0.8-1 cm diam.; calyx with a strong constriction between the lobes and the tube; calyx lobes reflexed. W Ecuador and NW Peru....................................................... 92. S. confe tiseriatum 7. Plants not dioecious; flowers 1-1.5 cm diam.; calyx not as above. 8. Leaves densely pubescent over the entire abaxial surface. Central coastal Venezuela, Caracas and vicinity................................................. 91. S. chlaniydogvynlm 8. Leaves with trichomes confined to the vein axils of the abaxial surface. E coastal Venezuela and the Lesser Antilles................................................. 99. S. tr-iste 6. Leaves without tufts of trichomes in the axils of the main lateral veins beneath; if pubescent, the trichomes minute and along the veins. 9. Inflorescences many-times branched, often quite dense. 10. Flowers 0.8-1 cm diam., not markedly fleshy. 11. Plants dioecious; ovary and berry densely pubescent; inflorescences many-times branched,especially on male plants. W Ecuador and NW Peru......... 92. S. confertiseriaturn 11. Plants not dioecious; ovary and berry glabrous or sparsely puberulent; inflorescences 95. S. oppositi/oliuni only 2-4 times branched. Amazonia.................................................. 10. Flowers larger, usually greater than 1 cm diam., fleshy.

TAXONOMIC TREATMENT

2 83

12. Inflorescences stout; flowers 0.8-1.3 cm diam.; fruiting pedicels 1.2-1.5 cm long. Mexico to Panama............................................ 97. S. rovirosanum 12. Inflorescences lax and open; flowers 1.2-1.5 cm diam.; fruiting pedicels 1-1.7 cm 98. S. sessile long. E Andean slopes and Amazonia..................................................... 9. Inflorescences simple, or at most once-furcate. 13. Flowers greenish-white, greater than 10 mm diam., each corolla lobe with a medial adaxial ridge, the lobes planar at anthesis; leaves 10-25 cm long. Peninsula de Paria, Venezuela ............................................... 100. S. turgidui 13. Flowers white, 8-9 mm diam., the lobes reflexed at anthesis, without a strongly marked adaxial ridge; leaves 7-10 cm long. Amazonia to the Guianas ............ 95. S. oppositijolium

91. Solanum chlamydogynum Bitter, Repert. Spec. Nov. Regni Veg. 16: 398. 1920. Type. Venezuela. Distrito Federal: Around Caracas, 800-1200 m, 23 Feb 1913, Pittier 5862 (lectotype, US, designated by D'Arcy, 1974 [1973]; isolectotypes, F, NY, US). Fig. 156 Shrubs or small trees, 2-8 m tall; young stems and leaves densely pubescent with golden, branched trichomes0.5-1 mm long; stems stronglywinged, the wings apparenteven in stems 10 cm diam.;older stems glabrate, the bark reddish-brown. Sympodial units difoliate, geminate.Leaves obovate, widest in the distal third, glabrous or with a few trichomes along the midrib adaxially, densely pubescent with golden, branchedtrichomes ca. 1 mm long abaxially; major leaves 12-30 x 7-14 cm, the apex acute, the base attenuate, winged onto the petiole; petioles winged, 11.5 cm long; minorleaves 4-8.5 x 2-5 cm, elliptic, the apex acute, the base acute to attenuate;petioles 0.5-1 cm long. Iniforescencesoppositethe leaves, branched, 2-8 cm long, 10-40-flowered, densely pubescentwith golden, branched trichomes; pedicel scars densely spaced, not overlapping,largely in distal 1/2 of inflorescence.Budsglobose whenyoung, laterellipsoidwith the corolla strongly exserted from the calyx tube. Pedicels at anthesiserector somewhatdeflexed, 1-1.5 cm long, taperingfrom the constrictionat the base of the calyx tube to a base ca. 0.5 mm diam.Flowers with the calyx tube cyathiform, abruptlynarrowedto the pedicel, 1.5-3 nun long, the lobes deltoid,1-2 mm long, denselypubescentwith golden,branchedtrichomeslike those of the rest of the inflorescence; corolla white, fleshy and somewhatwaxy, 1.5-2 cm diam., lobed ca. 3/4 of the way to the base, the lobes planarat anthesis, the abaxialsurfacespubescentwith branchedtrichomes and minutely papillose; anthers 4-5 x 1.5-2 mm, poricidal at the tips, the pores teardropshaped; free portion of thefilaments 0-0.5 mm long, the filament tube ca. 0.5 mm long; ovary densely pubescent with golden, branchedtrichomes;style straight,pubescent with trichomeslike those of the ovary;stigma capitate, brightgreen in live plants, minutely papillose.Fruit a

globose, green berry,sparselypubescentwith golden, branchedtrichomes, 1-1.5 cm diam.;fruitingpedicels woody, erectto somewhatdeflexed, 1.5-2 cm long, ca. 1 mm diam.at the base.Seedspale tan, ovoid-reniform or somewhat flattened,2-3 x 1.5-3 mm, the surfaces minutelypitted.Chromosomenumber.n = 12 (voucher Knapp & Mallet 6867). Distribution (Fig. 158). In the Cordillerade la Costa near Caracas,Venezuela,from 1500-2000 m. Specimensexamined.VENEZUELA.S.loc.,6 Jul 1891, Eggers 13294 (F, GH); s.loc., Linden 238 (F). ARAGUA: Colonia Tovar, 1800-2000 m, Dec 1924, Allart 327 (US); between Las Moras and HaciendaEl Atravesado, rd. to Colonia Tovar, ca. 1700 m, 12 Jan 1975, Benitez de Rojas 1800 (MY, NY); Colonia Tovar, 1854-55, Fendler 985 (G (Morton neg. 8507), GH, K); Colonia Tovar, 1854-55, Fendler 2096 (GH, K). DISTRITo FEDERAL: Quebrada de las Comadres, near Las Mostazas, 1000 m, Nov 1924, Al/art 237 (A); Caracasand vic., 30003500 ft, ca. 10?30'N, 19 Dec 1920, Bailey & Bailey 292 (BH, US); 21 Dec 1920, Bailey & Bailey 377 (BH, US); 5 Jan 1921, Bailey & Bailey 716 (BH, US); 21 Jan 1921, Bailey & Bailey, s.n. (BH); Caracas, Urb. Santa Maria, near Parque Nacional El Avila, 4 Oct 1980, Benitez de Rojas 2833 (MY, NY); vic. Caracas, Nov 1854, Buschel s.n. (K); Caracas,26 Dec 1912, Gollmers.n. (B [destroyed: F neg. 2687]); ca. 4 km NW of jct. with El JunquitoColonia Tovar rd. on rd. to Caracaya,jct. is ca. 16 km E of Colonia Tovar, N slope, Cordillera de La Costa, ca. 1750 m, 10?26'N, 67?08'W, 29 Oct 1984, Knapp & Mallet 6860 (BH, K, MY, NY, US, VEN); ca. 12 km E of Colonia Tovar on rd. to El Junquito,ca. 2000 m, 101 8'N, 670 lOW, 29 Oct 1984, Knapp & Mallet 6867 (BH. K, MY, NY, US, VEN); Caracas, 1842, Linideni238 (G (Morton neg. 8574)); in JardinBotanico, Caracas, 10 Aug 1979, Nee 17319 (BH, F); El Tas6n near El Valle, 12 Nov 1912, Pittier 9906 (GH, US); Caracas, 11 Mar 1922, Pittier 10225 (BH, GH, US); Antimano, 19 Mar 1927, Pittier 12286 (A, US); Ingomar(Tumerito), 900 m, Pittier 14176 (US); Monte Grappa 1 km before El Junquito, 1950 m, 27 Feb 1979, Plowman 7767 (F, K, U).

Solanumchlamydogynum is a distinctivespecies, with its densely pubescent leaf undersides and large waxy flowers. It is quitecommonalong roadsidesnear

NEOTROPICA ~~~~~~~~~~~~~FLORA

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TAXONOMIC TREATMENT

285

Caracas.The winged stems of S. chlamydogynumare of the restof the inflorescence,the corollasoon exserted strikingandthewings arepersistenton quitelargestems fromthe calyx.Pedicels at anthesisca. 5 mm long, slen(fide Nee 17319). Solanum chlamydogynumwas de- der, ca. 0.5 mm diam. at the base, abruptlybroadening scribedby Bitteras in sect. Anthoresisdue to the pres- to the calyx tube,erectto slightlypendant.Flowerswith ence of branchedtrichomes.It clearlybelongs in the S. the calyxtubecyathiform,1.5-2 mm long, in live specisessile species group, as it shares all the synapo- mens with a strong constrictionat the top of the tube where it meets the lobes, lobes roundeddeltoid, 0.5-1 morphies of the group (see Knapp, 199lb). Solanumchlamydogynumis most closely related mm long, the edges white and thickened in dry specito S. triste with which it shares branchedtrichomes, mens, puberulentwith minute trichomeslike those of white-edged calyx lobes, and seeds with incrassate the inflorescence;corolla white, 0.8-1 cm diam.,lobed margins.Solaniumchlamydogynumis always found at 3/4 of the way to the base, the lobes planaror slightly higherelevationsandin wetterhabitatsthanis S. triste. reflexed at anthesis, the tips and marginsof the lobes densely papillose; anthers ca. 3 x ca. 1 mm, poricidal at the tips, the pores teardropshaped; free portion of 92. Solanum confertiseriatum Bitter, Repert. Spec. thefilaments minute, the anthersnearly sessile on the Nov. Regni. Veg. 11: 490. 1913. Type. Ecuador. filamenttube,the filamenttube ca. 0.4 mm long;ovary Manabi: El Recreo, in sylvis, Jan 1897, Eggers densely pubescentwith golden, simple or dendritictri15499 (lectotype, M, designatedby Knapp, 1991a; chomes (usually correspondingto the type found on Fig. 157 therestof theplant);style in short-styled(male)flowers isolectotypes, F, K, US). Solanurtm polvasterGilli,FeddesRepert.94: 321. 1983. 1-1.5 mm long, in long-styled (female) flowers 5-6 Type. Ecuador.Chimborazo:Wald bei El Corazon, mm long, straight;stigmaclavate,decurrenton the style, minutelypapillose.Fruit a globose, greenberry,1-1.5 1480 m, 28 Jun 1975, Gilli 253 (holotype, W). Solanlin rovirosanumof A. H. Gentry& Dodson, cm diam.,pubescentwithminutegoldentrichomesoften Selbyana 4: 554. plate 261A. 1978. Not of nearly invisible to the naked eye, these trichomes Donn. Sm. simple,uniseriateor dendritic;fruitingpedicelswoody Solanurn arboreurm of A. H. Gentry & Dodson, andrathercorky,erect, 1-1.5 cm long, 1-1.5 mm diam. Selbyana 4: 550. plate 258B. 1978. Not of at the base. Seeds ovoid-reniform,with thintestae,pale Dunal. tannish-yellow,ca. 3.5 x ca. 2.5 mm, the surfacesmiBushy shrubs to small or medium size trees, 1- nutely pitted. Chromosomenumber:n = 12 (vouchers 10 m tall, young stemsminutelypuberulentwith golden Knapp & Mallet 6220, 6226, 6228). trichomes,theseuniseriateordendritic,0.05-1 mm long; Distribution (Fig. 158). In wet and dry forests olderstems glabrate,reddish.Sympodialunitsdifoliate, as a secondgrowthshrubor treein westem Ecuadorand geminate.Leaves elliptic to obovate, widest at or just northwestemPeru, from sea level to 500 m elevation. above the middle, glabrous and shining above, paler Selected specimens examined. ECUADOR. S.loc., and often pubescent on the veins, in the axils of the 22 Jul 1893, Eggers 14844 (F, K), 16 Aug 1893, Eggers veins, or over the entiresurfacebeneath,the trichomes 15122(F, K). CARCHI:N of San Marcos, 660 m, 1?07'N, simpleanduniseriateor dendritic(thosespecimenswith 78?20'W, 17 Jan 1983, Barfod41441 (AAU, BM); San only dendritictrichomesnearlyalways with the pubes- Marcos, along trail going by Rio San Juan, 660 m, 1?07'N, cence over the entire undersurfaceof the leaf); major 78?20'W, 2 Mar 1983, Barfod 41601 (AAU, BM); San leaves 11-20 x 7.5--9 cm, with 10-1 1 sets of main lat- Marcos valley, 600 m, 1?07'N, 78?20'W, 20 Nov 1983, eral veins, these impressed above, pale yellowish be- Kvistet al. 48695 (AAU, BM); above San Marcos de los neath, the apex acute or acuminate,the base acute (in type specimen) to attenuate,but not decurrenton the petiole; petioles 1-1.5 cm long, slightly channeled above; minor leaves differing from the majorsonly in size, 4-9 x 2-5.5 cm, the apex acuteto acuminate,the base acuteto slightly attenuate;petioles 0.7-1 cm long. Itiflorescencesopposite the leaves, two to many-times furcate,often extremelylargeandcomplex, 1.5-10 cm long, 5-120-flowered, puberulentwith the samegolden trichomesas the young stems;pedicelscars closely and evenly spaced,butnot overlapping,0.5-0.75 mm apart, beginning 1--5cm from the base of the inflorescence. Budsovoid,puberulentwiththe sametrichomesas those

Coaiqueres on trail toward Gualpi Bajo, ca. 1000 m, 1?06'N, 78?17'W, 7 Feb 1985, Ollgaard et al. 57352 Trail El Coraz6n-Ram6n Campana, (AAU). COTOPAXI: 1-3 km N of El Coraz6n, 12-1300 m, 18 May 1980, Harling & Andersson19246 (GB). ESMERALDAS:San Lorenzo, 18 Jul 1964, Jativa & Epling 825 (US); 3 km E of Quininde, ca. 65 m, 10 Apr 1943, Little 6225 (F, K, MAD, NY, US). GuAYAs:HaciendaLa Mina,at Rio Boliche, ca. 20 km E of Duran, 120 m, 2?25'S, 30 Sep 1955. B5cher et al. 212 (K); Balao, Jan 1892, Eggers 14219 (M); Capeira, 22 km N of Guayaquilon rd. to Daule, 20-150 m, 2?00'S, 79?58'W,14 Jul 1986, Gentrv& Dodson 54808 (MO, NY); vic. Samborondon, 10-60 m, 1?57'S, 79?43'W, 5 Feb--27 Apr 1980, Matthewson 125B (AAU); near Guayaquil, Dec 1926. Mille 970 (NY, QCA); Bosque Protector Cerro

NEOTROPICA ~~~~~~~~~~~~~FLORA

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Solanm confetiseriaum

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TAXONOMIC

287

TREATMENT

700

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Colorado community "Congoma Grande" at km 23 of Santo Domingo-Puerto Lim6n rd., 100 m, 0?21'S, 79?22'W,12-15 Jul 1982, Kvist 40630 (AAU); near Mindo, 3 Nov 1937, Sydow 300 (US). PERU. TUMBES: Bosque Nacional de Tumbes, near Campo Verde, 600-800 m, 21 Dec 1967, Simpson & Schunke V 440 (F, NY, US, USM), Simpson & Schunke V 460 (F, NY, US, USM).

Local names. Ecuador.Carchi:zapata;Los Rios: hoja de burro;Pichincha:wa pubancura (Colorado).

00

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Solanum

confertiseriatum

is related to S.

rovirosanumof CentralAmericaandMexico. Solanum confertiseriatum is easily distinguished from S. rovirosanum by its elliptic leaves, more complex branched inflorescences, smaller flowers with more prominentconstrictionsat thejunctionof the calyx tube and lobes and at the base of the calyx tube, reflexed calyx lobes, andits pubescentberries(occasionallywith branched trichomes). Solanum confertiseriatum grows

to be a medium-sizedtreelet,usuallywith a single trunk. Treesgrowing in the open have foetid smelling leaves, while those growing in the forest have little leaf odor. Solanum confertiseriatum is extremely variable

FIG. 158. Distribution of Solanum chlamydogynum (solid circles), S. confer-tiseriatum(open circles), and S. nmonadelphum(solid squares).

Blanco, 15 km W of Guayaquil, Cerro Mirador de los Monos, 200 m, 2 10'S, 79 58'W, 26 Feb 1992, Rubio & Palacios 2435 (BM). Los Rios: Km 170-175 Sto. Domingo-Quininde, 300 m, 2 Sep 1949, Acosta Solis 13665 (F), 5 Sep 1949, Acosta Solis 13789 (F); Rio Palenque Biological Station, km 56 Quevedo-Sto. Domingo, 150-220 m, 3 Sep 1972, Dodson & McMahon 5069 (US), 17 Sep 1973, Dodson & Tan 5351 (US), 7 Mar 1975, Dodson 5833 (MO, US), 13 Feb 1974, Gentry 9915 (MO), 22 Feb 1974, Gentry 10115 (MO); Centro Cientifico Rio Palenque, km 56 Sto. Domingo-Quevedo, 150-300 m, 30 Jan 1984, Knapp & Mallet 6219, 6220, 6221, 6226, 6228 (BH, K, QCA, QCNE, US); Cant6n Vinces, Hacienda Santa Lucia, 50 m, 19 Oct 1934, Mexia 6568 (CAS, BM); trail Quevedo-Naranjal, 60 m, 7 Nov 1934, Mexia 6670 (CAS, BM). MANABi: Due E of Hacienda Napo (Moteles del Playa) at N edge of San Vicente, 3-4 km E of ocean ca. 6 km N of Bahia Caracas,ca. 50 m, 0030'S, 80022'W, 15 Oct 1980, Croat 50700 (MO, NY); El Recreo, 16 Aug 1893, Eggers 15122 (F, BM, NY), 15 Jan 1897, Eggers 15556 (F), 20 Jan 1897, Eggers 15585 (B (F neg. 2655), F, K); Montecristi, Cerro Montecristi, E slopes above town, 300-400 m, 18 Jul 1986, Plowman & Alcorn 14354 (F, K, NY). PICHINCHA: Sto. DomingoRosa Zarate (Quininde), Rio Blanco, 180 m, 9 May 1968, Harling et al. 9308 (GB); Rancho Grandeto 43 km on rd. to Quinind6, 6 Apr 1943, Little 6180 (K, MAD, US); in

in both degree and type of pubescence, and can be either almost completely glabrous (with a few minute uniseriatetrichomes on the inflorescence) or densely pubescentwith dendritictrichomes.Plantswith intermediatedegrees of pubescence also exist. The density and branchingof the trichomesseem to vary together, and appear to correspond to a gradient in aridity in coastal Ecuadorand Peru.Forestsare generallywetter in northemEcuadorandbecome drieras one proceeds south fromthe equator.Those specimens of S. confertiseriatumfromTumbes,Peru,arethe most pubescent and the trichomes are almost exclusively dendritic. Labels also indicatethe plants come from dry areas. 93. Solanum monadelphum Van Heurck & Mull. Arg.,Observ.Bot.42. 1870.Type.Peru.SanMartin: Prope Tarapoto, 1855-1856, Spruce 4051 (lectotype, G, designatedhere;isolectotypes,AWH-n.v., Fig. 159 BM, C, G-DC, K, MPU). Small shrubs of river courses, 1-1.5 m tall; young stems puberulent with appressed reddish uniseriatepapillatetrichomesca. 0.1 mm long; young leaves glabrousandsomewhatshiny,occasionallywith a few trichomes like those of the young stems; olders stems glabrate,the barkreddish-gray.Sympodialunits plurifoliate.Leaves bornein whorl-like clusters along the stems, lanceolateto oblanceolate,very rarelygeminate,widest at or distalto the middle, glabrouson both surfaces,sessile, 6-20 x 0.8-4.5 cm, with 6-8 pairsof mainlateralveins, these prominentandyellow beneath, the apex acute,the base sessile, attenuateanddecurrent

FLORA NEOTROPICA

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TAXONOMIC TREATMENT

onto the stem.Inflorescencesterminal,laterbecoming lateral,often many-timesbranched,glabrous,1.5-5 cm long, 5-20-flowered;pedicel scars irregularlyspaced 0.5-1.5 mm apart.Budscompletelyenclosedin thecalyx until quite late, when young appearingsomewhatcaudate from the elongate calyx tube, later globose, then ellipsoid after corolla exsertion. Pedicels at anthesis thick, erect, andwhite, 0.7-1.1 cm long, taperingfrom the base of the calyx tube to a base 0.5-1 mm diam. Flowers with the calyx tube cup-shaped,constrictedat the base, 1.5-2.5 mm long, the lobes irregularlydeltoid, 2.5-3 mm long, glabrous and appearingwoody in dry material,fleshy in live plants;corolla white and fleshy, 1.5-2 cm diam.,lobed 2/3 of the way to thebase, the lobes planarat anthesis, the tips and distal 1/3 of the marginsof the lobes denselypapillose;anthers3.54.5 x 1-1.5 mm, poricidal at the tips, the pores teardropshaped;freeportionof thefilaments0.5-0.75 mm long, the filamenttube 1-1.5 mm long;ovaryglabrous; stlle straight,7-9 mm long; stigma merely a widened areaon the tip of the style, the surfaceminutelypapillose. Frutita globose, greenberry,becomingyellowish when ripe, 0.8-1 cm diam.;fruitingpedicels woody, erect, 0.5-1.2 cm long, 1-2 mm diam. at the base; calyx lobes accrescentandwoody in fruit,ca. 5 mm long. Seeds darkbrownin drymaterial,pale tanin live plants, ovoid-reniform, 3-3.5 x 1-1.5 mm, the surfaces minutely pitted. Chromosomenumbernot known. Distribution (Fig. 158). Throughout eastern Peru along river courses, from 250 to 500 m.

289 600-800 m, 17 Oct 1984, Maas et al. 6055 (NY, U, USM). SAN MARTiN: Boquer6npass, 92 km fromTingo Maria on hwy.to Pucallpa,ca. 400 m, 16 Dec 1949,5 Jan 1950, Allard21749 (F); km 27-30 Tarapoto-Yurimaguas rd., 650-750 m, 6?25'S,76?15'W,9 Jun 1984,Knapp& Mallet 6490 (BH, K, US, USM); km 36-41 TarapotoYunmaguas rd.,alongRio Cachiyacu andRio Tiyacu,400600 m, 6?25'S,76?15'W,11 Jun 1984,Knapp& Mallet 6496 (BH, K, US, USM); km 50 Tarapoto-Yurimaguas rd., 300-400 m, 6?20'S,76018'W,12 Jun 1984,Knapp& La Mallet6502 (BH,K, US, USM);Pongode Cainarache, Perla,km 60 Tarapoto-Yurimaguas rd.,ca. 300 m, 6018'S. 76 18'W,12 Jun1984,Knapp& Mallet6506(BH.K, US. rd., trail to Rio USM); km 40 Tarapoto-Yurimaguas TiriyacuandRio Cashiyacu,eventuallyto salt mineson 24 Apr 1986, Rio Cashiyacu, 300-400m, 6025'S,76?15'W,

Knapp& Mallet 7199, 7200 (MO, USM); MariscalCaceres,

V Rio Sion,SW of caserioof Sion,22 Oct 1969,Schlunke 3549 (F); Rio Cainarachenear bridge for TarapotoYurimaguasrd., ca. 30 km NE of Tarapoto,ca. 550 m, 6?30'S,76?20'W,12 Feb 1985,Stein& Todzia2154 (MO, NY); ChilcayonearTarapoto,Dec 1902, Ule 6632 (K); Tarapoto,360-900 m, 26 Dec 1929, Williams6734 (F). Solanum monadelphumis closely related to S. sessile, also of easternPeru,but is easily distinguished from that species by its narrow,willow-like leaves, plurifoliatesympodialunits, andriverbedhabitat.The buds of the two species are nearly identical, and the inflorescence structure is quite similar. In the area aroundQuincemil (Peru, Cuzco, Quispicanchis), intermediatesbetween the local formof S. sessile andS. monadelphumarecommon.The intermediatesare usually found growing in the water with typical S. monadelphumplants.This could representlocal variation in S. monadelphumor hybridizationbetweernthe two species. Solanumn monadelphumandS. sessile are sympatricin otherpartsof theirranges,but I have seen intermediatesonly in the Quincemilarea. Solanummonadelphulnis one of the most ecologically striking members of sect. Geminata.It is a shrubof river beds, growing on gravel or sand banks, and is subject to periodic inundation,particularilyin the wet season, when the plantsmay spendmany (lays underwater.The narrowleaves of S. rnonadelpliluni are relatedto this habitat.Threeotherspecies also growing in river beds, but not necessarily closely related to S. monadelphum(S.palmillae,S. imberbeof theS. deflexiflorum species group,S. amnicola of the S. arboreurm species group), sharethis leaf shape.Solanummicna-

Specimens examined. PERU. Cuzco: Quispicanchis, Quince Mil, trail to Rio Nusinescatu, 500-600 m, 13?15'S, 70?45'W, 22 Akpr1984, Knapp & Mallet 6377 (BH, CUZ, K, US, USM); ridges N of Quince Mil, 500-600 m, 13?l5'S, 70?45'W, 23 Apr 1984, Knapp & Mallet 6387, 6389 (BH, CUZ, US, USM); along Rio Sausipata, near village of Cadena,ca. 2(1km W of Quince Mil on rd. to Cuzco, 9001000 in, 13?20'S, 70?50'W,29 Apr 1984, Knapp & Mallet 6413 (BH, CUZ, US, USM); Quince Mil, TanamayoChicoNusinescatu, 500-700 m, 21--28 Jan 1949, Var-gasC. 7721 (CUZ); Quispicanchis, Pte. Huacyumbe,Quince Mil, 500700 in. 21-28 Jan 1949, Vargas C. 7792 (CUZ); Quispicanchis, between Quince Mil and San Lorenzo, 700 Sumrn, 25 Jul 1957, VargasC. 11750 (CUZ). HUANUCO: mit of La Divisoria, rd. between Tingo Mariaand Pucallpa, 170 m, 18 Jan 1987, Diaz & Baldeon 2277 (MO, NY, USM). JUNiN: Puerto Yessup, ca. 400 m, 10-12 Jul 1929, Killip & Smith26400 (F, US). PASCO:Ca. I km from didelphum is common where it occurs, forming dense vision of Villa Rica-Pto. Bermudez rd. and Villa Ricaknee-high thickets along riverbanksin local areas. Palcazu rd.. Palcazu branch, Rio Cacazu, ca. 500 m, 10030'S, 75010'W. 15 Aug 1984, Knapp & Mallet 6638 (BH, US, USM); kin 28 Repartici6n-Iscozacin(km 86 Villa Rica-lscozacin-Pto. Mairo), Rio La Raya near Ameusha 94. SolanumobovalifoliumPittierex Benitez,Emstia19: 17. 1983.Type.Venezuela.Aragua:SelvasdelVallede community of Laguna, ca. 350 m. 10?20'S, 75010'W. 22El Medio,Chuao,600 m, 15 Mar 1926,Pittier12127 23 Aug 1984, Kniapp & Mallet 6652 (BH, US, USM). PITNO:Rio Sangabin, in vic. of Sangaban(LanlacuniBajo), Fig. 160 (holotype,VEN;isotype,MO).

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TAXONOMIC TREATMENT

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FIG. 161. Distribution of Solantim obovalijolium (open circles) and S. oppositifolium (solid circles).

Shrubs or small trees 2-7 m tall; young stems and leaves minutely puberulentwith tiny reddish trichomes, these soon deciduous; older stems glabrate; barkof older stems andtrunksgray and shiny.Sympodial units difoliate, geminate. Leaves obovate, rather fleshy, widest distal to the middle, glabrous on both surfaces,or occasionally with a few minuteuniseriate trichomes near the base of the midriband petiole beneath; major leaves 20-40 x 10-15 cm, with 11-12 pairsof main lateralveins, these orangeandprominent beneath in dry material,the apex acute to acuminate, the base long attenuate;petioles 1.5-2 cm long, slightly winged from the decurrentleaf bases; minor leaves differing from the majorones only in size, 7-10 x 3-5 cm, the apexacute,thebase attenuate;petioles0.7-1 cm long.Inflorescencesoppositethe leaves, simpleor more often furcate,0.5-5 cm long, 10-50-flowered,minutely puberulentwith the same reddishtrichomesas those of the young stems and leaves; pedicel scars closely spaced, but not overlapping.Buds fleshy, globose, the corolla soon exsertedfromthe calyx, the buds then ellipsoid, the calyx appearingwoody in dry material. Pedicels at anthesiswhite, ratherfleshy,deflexed, 0.91.1 cm long, tapering from the calyx tube to a basal diam. of ca. 0.75 mm. Flowers with the calyx tube cyathiform,ca. 2 mm, the lobes deltate, 1-2 mm, glabrous;corolla white, fleshy, 2-2.2 cm diam., lobed ca. 3/4 of the way to the base, the lobes planarat anthesis,

the tips of the lobes hooded, the tips and marginsminutely papillose; anthers 4-4.5 x 1.2-1.5 mm, poricidal

at the tips, the pores teardropshaped; free portion of thefilaments 1-1.5 mm long, the filamenttube ca. 0.5 mm long; ovaryglabrous;style straight,5-7 mm long; stigma a minutelywhite papillose areaon the tip of the style.Fruita globose,greenbenry,1-1.3 cm diam.;fruiting pedicels deflexed, woody, 1.5-2 cm long, ca. 1.5 mm diam. at the base; calyx lobes persistentand woody in fruit,ca. 3 mm long.Seeds darkbrown,ovoid-renifonn, ca. 4 x 2.5 mm, the surfaces minutely pitted, nearly smooth. Chromosome number not known.

Distribution (Fig. 161). In the Cordillerade la CostaandtheAndesof Venezuela,growingin lightgaps from 600 to 1000 m. Specimens examined. VENEZUELA. ARAGUA: Pico Periquito, Parque Nacional Henri Pittier, Apr 1959, Aristeguieta 3875 (VEN); Alto de Choroni, 1000-1400 m, 8 May 1949, Badillo 1911 (MY); Rancho Grande, 800-1000 m, 12 Apr 1962, Badillo 3848 (MY); Parque Nacional Henri Pittier, rd. to Choroni, N slope, 950 m, 29 May 1980, Benitez de Rojas 2746 (MY), 5 Jul 1983, Benitez de Rojas 3139 (MY); Parque Nacional Henri Pittier (Rancho Grande), la Cumbre de Rancho Grande trail, 1300 m, 10?21'N, 67?39'W, 17 Apr 1979, Davidse et al. 16724 (VEN); Rancho Grande, 1150 m, 10 May 1951, Garcia 48 (VEN); between Portachuelo and Pico Periquito, 10 Apr 1980, Manara s.n. (VEN-176417, VEN-

FLORA NEOTROPICA

292 176416); Parque Nacional Henri Pittier, Rancho Grande, trail to Pico Guacamayo behind station, 1100-1400 m, 10?21'N, 67?42'W, 27 Oct 1984, Knapp & Mallet 6851 (BH, MY, US, VEN); Alto de Rancho Grande, 1200 m, 2 Apr 1926, Pittier 12150 (F, G, K, MAD, MO, NY, US, VEN); Parque Nacional Rancho Grande, 1400 m, 24 Apr 1937, Pittier 13990 (F, US, VEN); W of El Portachuelo de Rancho Grande, 1300-1800 m, 17 Apr 1947, Pittier & Nakichenovich 15335 (US, VEN); Parque Nacional Henri Pittier, on peak between hotel and la Toma de Agua, 1100 m, 31 May 1966, Steyermark et al. 95822 (NY, VEN); Parque Nacional Henri Pittier, headwaters of Rio Grande del Medio, near Quebrada Rio Hondo between Tremaria and La Regresiva del Diablo on Fila Alta de Choroni, 1000 m, 30 Apr 1972, Ste.vermark & Carreno Espinoza 105843 (NY, U, US, VEN); cumbre de Choroni, 1200 m, Apr 1953, Truijillo 1703 (MY). CARABOBO: Headwaters of Rio San Giun above los Tanques and la Toma, between Quebrada de los Verros, S of Borburata, 750-900 m, 29 Mar 1966, Ste-vermark & Steyermar-k 95281 (F, NY, US, VEN). TRUJILLO: 10 km from Biscucuy, 850-900 m, 9022'46"N, 70002'04"W, 12 May 1975, Graterol et al. 25 (MY). YARACUY: Sierra de Aroa, Cerro Negro, 8 km SWv'of San Felipe, 1200-1800 m, 10017'N, 69?01'W, 1-2 Apr 1980, Liesner & Gonzalez 9928 (VEN). Solanum obovalifolium is most closely related

to S. sessile of the Amazonbasin,particularlyto higherelevation populations of that species. Solanum obovalifoliumgenerally lacks the erect uniseriatetrichomes so characteristicof other members of the S. sessile species group, but agrees with them in several other characteristics, such as fleshy flowers, large leaves, andbranchedinflorescences. The constriction just below the calyx lobes is less pronounced in S. obovalifoliurm than in othermembersof the group. Most of the collections of Solanum obovalifolium

arefromthe stateof Aragua,butthis in partreflectsthe intensity of collecting in this area. Solanum obovalifolium is a light gap colonizer and probablyoccurs at low densities throughoutthe coastal ranges of Venezuela, and into the Andes of Trujillo and Mrida. In ParqueNacionalHenriPittier,Aragua,S. obovalifoliurm grows in dense stands in old light gaps under closed canopy cloud forest.

95. Solanum oppositifolium Ruiz & Pav., Fl. Peruv. 2: 35. fig. 168a. 1799. Type. Peru. Junin: Vitoc, Tafalla s.n. (lectotype, MA [F neg. 29729]; isolectotypes, K, MA, frag. F). Fig. 162 Solanum urceolatium Pers., Syn. P1. 1: 223. 1805. Type. Based on S. oppositifolium Ruiz & Pav. "Dantur plures species ubi flores fol. sunt oppositi, hinc nomen minus congruum mutavi." Solanum schizopodilum Sendtn. in Mart., Fl. Bras.

10: 46. 1846. Type. Brazil. Para: In sylvis ad Para,Aug, Martiuss.n. (lectotype,M, designated by Knapp, 1991a [F neg. 6543]). Solanum viliflor-um Sendtn. in Mart., Fl. Bras. 10:

46. 1846. Type. Brazil. Amazonas: In sylvis Japurensibus, prov.ad Rio Negro dictae,Martiu.s s.n. (lectotype,M, designatedby Knapp,1991a). Solanum confine Dunal var. curi-tumJ. F. Macbr.,

Publ. Field Mus. Nat. Hist., Bot. Ser. 13(5B): 202. 1962. Type. Peru. Loreto:Mishuyacunear Iquitos, 100 m, Jan 1930, Klug 834 (holotype. F; isotype, NY). Solanumpuber-ulobaSteyerm.,Bol. Soc. Cienc.Nat. 26: 444. 1966. Type. Venezuela. Bolivar: Wooded ridge of Fila de la Danta, between km 125 andkm 127, betweenLuepa campamento and Cerro Venamo, 1200 m, 15-17 Apr 1960. Steyermark & Nilsson 185 (holotype. VEN; isotypes, MO, NY). Solanumconfertiseriatumof Vasquez Martinez,Fl. Res.

Biol. Iquitos693. 1997.Not of Bitter. Shrubs or small trees, 1-5 m tall, occasionally quitesmall(ca. 30 cm) in inundatedforests;youngstems andleaves densely hirsutulouswith erectuniseriatetrichomes, the trichomes to 0.2 mm long; older stems remainingpuberulent,the new growth soon glabrous except on the veins; bark of the older stems grayishbrown. Sympodial units difoliate, geminate. Leaves

obovate to elliptic, widest at or above the middle, glabrous and often shiny above, glabrous or hirsutulous on the veins beneath,the trichomeserectanduniseriate, ca. 0.2 mm long, the undersides paler, especially in plantsfromsouthernpopulations;majorleaves 7-30 x 3.5-12 cm, with 7-9 pairs of main lateralveins, these slightly impressedabove, yellowish beneath,the apex acuteto acuminate,the base cuneateto attenuate;petioles 0.5-1.1 cm long; minorleaves differing from the major ones only in size (but in one Guyanese collection the minorleaves orbicular),2.5-1 1 x 1.3-4.5 cm., the apex acuteto acuminate,the base cuneateto attenuate;petioles 5-8 mm long. Inflorescencesoppositethe leaves, occasionallyterminal,but in such cases becoming lateralandoppositethe leaves as shoot growthcontinues, simple or many-times furcate, 1-12 cm long. 5-50-flowered, commonly with only a few flowers open at a time, densely hirsutulous with tiny erect uniseriatetrichomeslike those of the young stems and leaves; pedicel scars in pairs, the members of a pair closely spaced,the pairsevenly spacedca. 1 mm apart. Buds ellipsoid, the corolla soon exserted from the calyx tube. Pedicels at anthesis deflexed, 4-6 mm long, taperingfrom the calyx tube to a slender base ca. 0.5 mm diam., densely hirsutulouswith tiny uniseriatetrichomes. Flowers with the calvx tube narrowly campanulate, 1-1.5 mm long, hirsutulous,the lobes deltoid, hyaline, 0.2-1 mm long, densely hirsutulouswith

293

TAXONOMIC TREATMENT

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294

FLORA NEOTROPICA

Carlos, 4 km S of Solano, 120-140 m, 2 Feb 1977, Morillo & Hagesawa 5132 (MY, VEN); between Cafio Chola and Solano, 1-4 km S of Solano, 120 m, 8 Feb 1977, Morillo 5507 (MY, VEN); behind "Tobagan de la Selva" camping area, 35 km S of Puerto Ayacucho, 85 m, 21 Feb 1979, Plowman 7724 (F); Atabapo, 15 km SE of San Fernando de Atabapo, 110 m, 3?50'N, 67?40'W, 10-16 Jan 1988, Stergios et al. 11514 (NY); Atures, vic. of and upstream from damsite, N bank Rio Cataniapo, 45 km SE of Puerto Ayacucho, 100 m, 5035'N, 67'15'W, 13 May 1980, Steyermark et al. 122393 (NY); Atures, between Las Lomas and dam site, SE of Puerto Ayacucho, l00 m, 3 May 1981, Steyermarket al. 122431 (VEN); slopes of Mount Duida, 4000 ft, Aug 1928-Apr 1929, Tate 920 (NY); Tamatama, Alto Orinoco, 125 m, 6 Dec 1942. Williams 15847 (US, VEN); Yavita, 128 m, 24 Jan 1942, Williams 13959 (US, VEN). BOLiVAR: Slopes going up to Gran Sabana, km 128 of El Dorado-Sta. Elena de Uairen rd., ca. 10 km S of Salto El Danto. ca. 1300 m, 6?25'N, 62?25'W, 11 Oct 1984, Knapp & Mfallet 6743 (BH, K, MY, US, VEN); entrance to Gran Sabana, km 128 of El Dorado-Sta. Elena rd., 18 Jan 1973, Morillo Distribution (Fig. 161). Widely distributedin et al. 2900 (VEN); Roraima, Rio Cuyuni drainiage km the Amazonbasin,fromGuyanato southemPeru,from 117.5-132.5 S of El Dorado, 865-1300 m, 26-27 Dec sea level to 800 m. This species grows in a variety of 1970, Steyermark et al. 104491 (VEN); near El Palmar. ca. 75 m, ca. 8?05'N, 61?50'W,23 Nov 1967. WesselsBoer habitatsfrom inundatedforest to uplandrainforest. 2067 (NY). DELTA AMACURO: El Palmar-Raudal trail. Selected specimens examined. COLOMBIA. upper Rio Toro drainage, Serrania Imataca, 270-470 ANIAZONAS: Parque Nacional Natural Amacayacu, m, 7 Nov 1955, Wurdack & Moanachino 39616 (NY). Centro Admin. Mata-mata (Inderena), 100m, 3?47'S, GUYANA. MAZARUNI:Takatu Creek to Puruni 70?15'W, 15 Feb 1991, Rudas & Joaquin 1295 ,4 Mar River, Mazaruni River, 24 Oct 1944, Fanshawe F2058 1991, Rudas et al. 1450 (BM, MO); P.N.N. Amacayacu, (K, NY, US). Quebrada la Mufiequa, 100 m, 3?06'S, 70?03'W, 18 Jun FRENCH GUIANA. Saul, trace Limonade, 11 1991, Rudas et al. 2113 (BM, MO); Rio Igaraparana, Dec 1976, Cremers3834 (NY, P), Cremers3934 (U); Saul, between QuebradaMenaje and Rio Putumayo, 150-170 m, ORSTOMtrail to crique Limonade, 3 km from the village, 15-17 Jun 1942, Schultes3986 (K); Leticia, 14 Oct 1946, 3 Nov 1974, de Granville2276 (U); Sail, crique Limonade, Schultes & Black 46--205 (US). 2.5 km from village, 12 Oct 1972, de Granville B4560 VENEZUELA. AMAZONAS: Casiquiare,ca. 12 km (NY, P); Saul, Sophie airport, 7 Sep 1962, HallW801 SE of San Fernando de Atabapo, 110 m, 3?50'N, (NY, P, US); Sail, Limonade circuit, 16 Jan 1974, Hall 67?47'W, 10-16 Jan 1988, Aymard et al. 6400 (NY); 2241 (MPU), 12 Feb 1978, Halle 2672 (MPU). San Carlos del Rio Negro, ca. 20 km S of confluence of ECUADOR. NAPO: Rio Cuyabeno, from 2 hours Rio Negro and Brazo Casiquiare, 4.3 km NNE of San upstream from outlet in Rio Aguarico to Puerto Bolivar, Carlos on Solano rd., 119 m, 1?56'S, 67?03'W, 12 Nov 300 m, 0?15'S, 76?0-10'W, 17 Aug 1981, Brandbyge et 1969, Clark 7246 (MO, NY); Atabapo, N part of Siera al. 33621 (K, NY); Nuevo Rocafuerte, 1.5 km N of Parima,between Shimada-Wochi and Rio Matacuni, 820Caserio Quichuan, 200-230 m, 26 Feb 1981, Jaramillo 100 m, 3?55'N, 64?40'W, 8 Nov 1983, Huber & Colchester & Coello 4282 (AAU). PASTAZA: Ceilan, trail from 8396 (NY); 4 km E of San Carlos de Rio Negro, 120 m, Ceilan to Rio Cononaco, N side of Rio Curaray,200 m, 1?56'S, 67004'W, 23 Nov 1977, Liesner 3779 (MY, 1036'S, 75040'W, 6 Jun 1980, Brandbvge & Asanza C. VEN); 4 km NE of San Carlos de Rio Negro, ca. 20 km 31669 (AAU). SUCUMBIOS: Along rd. between tLago S of confluence Rio Negro and Brazo Casiquiare, 120 Agrio and Coca, 6 km S of Lago Agrio, 355 m, 0?02'S, m, 1056'N, 67?03'W, 4 Apr 1979, Liesner 6123 (MO, 76?43'W, 29 Feb 1992, Croat 72494 (QCNE). VEN), 9 Apr 1979, Liesner 6428 (MO, VEN), 1 May PERU. AMAZONAS: E of Huampami, 70)0-800 ft, 1979, Liesner 7120 (MO, VEN); Cerro Sipapo (Paraque), 13 Dec 1972, Berlin 538 (MO); Huambisa, Rio Santiago, 125 m, 25 Jan 1949, Maguire & Politi 28620 (NY); QuebradaCaterpiza, 2-3 km behind village of Caterpiza, Cerro(le la Neblina, Rio Yatua, 140-1700 m, 24 Dec 1953, 200 m, 3?50'S, 77040'W, 20 Dec 1979, Tunqui 383 Maguire et al. 38653 (NY); Cerro Neblina, Rio Yatua, (MO). HUANUCO: Pachitea, Sungaro W of Puerto Inca, between Camps 3 and 4, 900-1000 m, 8 Nov 1957, along hwy. construction 10 km W of Sungaro river Maguire et al. 42009 (NY); 2 km from San Carlos on crossing, 250-300 m, 9?22'S, 75'00'W, 14 Sep 1982, rd. to Solano, 27 Mar 1971, Manara 164 (MY, VEN); Foster s.n. (F, USM); Tingo Maria,valley of Rio Huallaga, San Carlos-Solano rd., between Canio Chola and San ca. 700 m, 24 Oct 1938, Stor-k& Hortoni 9486 (F, K).

erectuniseriatetrichomes;corolla white or greenishwhite, 8-9 mm diam.,lobednearlyto thebase, the lobes reflexed at anthesis, the tips and marginsof the lobes puberulentwith golden uniseriatetrichomes;anthers 2-2.5 x 0.75-1 mm, the terminal 0.2 mm paler and thickened, poricidal at the tips, the pores teardrop shaped;free portionof thefilaments0.1-0.3 mm long, the filamenttube 0.2-0.5 mm long; ovary glabrousor pubescentwith golden simple uniseriate(occasionally branched)trichomes,these, if present,persistingonto the berry;style in short-styledflowers ca. 1.5 mm long, in long-styled flowers 3-4 mm long, straight;stigma slightly clavate, minutely papillose. Fruit a globose, green berry,glabrous or pubescent, 1-1.5 cm diam.; fruitingpedicels woody, erect, 0.6-1.5 cm long, 1-1.5 mm diam. at the base, the apex slightly expanded,ca. 3 mm diam.Seedspale brownish-tan,ovoid-reniform,34 x 2-2.5 mm,the surfaces minutely pitted. Chromosome number. n = 12 (voucherKnapp& Mallet 6618).

TAXONOMIC TREATMENT LORETO: Maynas, Rio Ucayali, 28 Jan 1973, Ayala 231

(AMAZ); Maynas, Isla Lupuna, 11 Jul 1974, Ayala 622 (AMAZ, MO); Rio Corriente, Shiriyacu on rd. to Forestales (38 km), 25 Nov 1979, Ayala 2325 (AMAZ); Maynas, Caballo Cocha, Ramon Castilla, 110 m, Ayala et al. 3287 (AMAZ); 7 km SW of Iquitos, 31 Jul 1972, Croat 18632 (MO); Maynas, Padre Isla, Rio Amazonas in front of Iquitos, ca. 120 m, 27 Feb 1978, Diaz & Jaramillo 12 (AMAZ, MO); Maynas, Pefia Negra, on rd. from Iquitos past Quisto Cocha, ca. 150 m, 4 Dec 1976, Davidsoni 5277 (MO, NY); lower Rio Mom6n, tributary of Rio Nanay, near Iquitos, 6 Dec 1979, Davidson & Jones 9613 (F, NY); Rio Nanay, Caserio Mishana 30 km SW of Iquitos, 4 km S of Mishana, 15 Aug 1980, Foster 4301 (F, USM); Maynas, E of Puerto Alegria on quebrada leading to Bella Vista, peninsula between Rio Amazonas and Rio Javari, 100 m, 15 Mar 1977, Gentry & Daly 18364 (AMAZ); Maynas, Rio Yavari, Emilia (Peruvian village above Atalaia del Norte), ca. 120 m, 22 Nov 1977, Gentry & Revilla 20779 (MO); vic. of Mishana, between Rio Nanay and Rio Itaya, ca. 130 m, 29 Nov 1977, Gentry, et al. 20975 (MO); outskirts of Iquitos, Rio Amazonas between Puchana and Santa Clara de Nanay, ca. 120 m, 3 Feb 1977, Gentry et al. 21610 (MO); Maynas, mouth of Rio Nanay near Iquitos, 120 m, 4 Mar 1979, Gentry et al. 25378 (AMAZ, MO); Maynas, Quebrada Yanamono, Explorama tourist camp, Rio Amazonas above mouth of Rio Napo, ca. 130 m, 13 Nov 1979, Gentry et al. 27985 (MO); Maynas, trail to Rio Itaya, left bank of Rio Amazonas just below mouth of Rio Ucayali, 4?20'S, 72?30'W, 12 Oct 1980, Gentry et al. 30049 (MO); Yanomono, Explorama Tourist Camp, 120 m, 3?28'S, 72?48'W, 19 Feb 1981, Gentry et al. 31487 (MO, NY); Mishana, Rio Nanay halfway between Iquitos and Santa Maria de Nanay, 140 m, 3?50'S, 73030'W, 25 Feb 1981, Gentry et al. 31740 (MO); Mishana, Rio Nanay halfway between Iquitos and Santa Maria de Nanay, 140 m, 3?52'S, 73?30'W, 31 Dec 1982. Gentry & Enmmons38797 (MO); Iquitos, ca. 100 m, 3-11 Aug 1929, Killip & Smith 26981 (F, NY, MO), Killip & Smith 27054 (US); Puerto Arturo, lower Rio Huallaga below Yurimaguas, ca. 135 m, 24-25 Aug 1929, Killip & Smith 27836 (US); Santa Rosa, lower Rio Huallaga below Yurimaguas, ca. 135 m, 1-5 Sep 1929, Killip & Smith 28717 (NY, US); Mishayacu near Iquitos, ca. 100 m, 24-28 Sep 1929, Killip & Smith 29878 (NY, US); Mishayacu, near Iquitos, ca. 100 m, Oct-Nov 1929, Klug 8 (F, NY, US); Florida, Rio Putumayo at mouth of Rio Zubineta, ca. 200 m, MarApr 1931, Klug 2063 (BM, F, K, NY), May-Jul 1931, Klug 2214 (BM, F, K, NY, US); Yanamono, Explorama tourist camp on Rio Amazonas between Indiana and mouth of Rio Napo, ca. 80 km NE of Iquitos, ca. 100 m, 3?28'S, 72?48'W, 22 Jul 1984, Knapp & Mallet 6594, 6599 (BH, K, NY, USM), 23-27 Jul 1984, Knapp & Mallet 6618 (BH, K, US, USM); Rancho Indiana, overflowed left bank of Rio Maranion, 110 m, 24 Jan 1932, Mexia 6421 (BM, F, K, NY, U, US); Rio Nanay, Zamito, ca. 100 m, 19 Feb 1969, Plowman 2498 (K, US); Rio Itaya near Palo Seco, 14 May 1977, Revilla et al. 2547,

295 2579 (AMAZ, MO); Santa Maria de Nanay, NW of Rio Nanay, ca. 130 m, 27 Feb 1968, Schunke V 2418 (F., US); Maynas, San Alejandro, 1 hr from mouth of Rio Napo, left bank, 3?20'S, 72?40'W, 30 Mar 1980, Vasquez & Jaramillo 003 (AMAZ, NY); Recreo, Rio Manati NEf of Iquitos, ca. 115 m, 3?42'S, 72?50'W, 22 Dec 1980, Vasquez& Jaramillo 1147 (MO); Maynas, Puerto Alianza (Qda. Tamshiyacu), 160 m, 4?08'S, 72?55'W, 29 May 1981, V6squez & Criollo 1870 (MO); Llachapa, Explor Napo, Rio Napo, ca. 130 m, 3?18'S, 72?55'W, 19 Jan 1983, Vasquez & Jaramillo 3748 (MO, NY); Maynas, Alpahuayo, IIAP station, 20 Oct 1984, Vasquez & Cuiollo 5799 (MO, NY); Yanamono, Explorama Lodge camp, 106 m, 3?30'S, 72?50'W, 17 Apr 1985, Vasquez & Jaramillo 6368 (MO, NY); Estaci6n Biol6gica Rio Blanco, 150 m, 4?20'S, 72?45'W, 16 Sep 1985, Vdisqueet al. 6735 (MO, NY); Puerto Almendras, Rio Nanay, 122 m, 3?48'S, 73?25'W, 2 Oct 1986, Vasquezet al. 8080 (MO, NY); along Rio Itaya, May 1929, Williams57 (F); Timbuchi on the Rio Nanay, 1 Jul 1929, Williams 1164 (F); Pr6 on the Rio Amazonas, 1 Aug 1929, Williamns 1993 (F); Caballo Cocha on Rio Amazonas, 5 Aug 1929, Williams2095 (F, US), Williams2224 (US); Paraiso, Alto Rio Itaya, 145 m, 30 Sep 1929, Williams3285 (F); Iquitos. 120 m, 23 Mar 1930, Williams 7912 (F), 30 Mar 1930, Williams8066 (F), 6 Apr 1930, Williams8184 (F). MADRE DE DIos: Parque Nacional Manu, Cocha Cashu station, ca. 400 m, I 1?52'S, 71?22'W, 12 Nov 1982, Em7mous 140 (MO); Parque Nacional Manu, Cocha Cashu station on old oxbow lake of Rio Manu, 16 Aug 1973, Foster2636 (F, MO, US); Shintuya, ca. 1 km up small stream from Rio Madre de Dios, 8 Aug 1974, Foster et al. 3140 (F, MO); Rio Manu, vic. of Cocha Cashu station, 27 Oct 1976, Foster & Terborgh5143 (F, MO); Cerrode PantiacoLla, Rio Palotoa 10-15 km NNW of Shintuya, 500-650 m, 12035'S, 71018'W, 8 Dec 1985, Foster 10815 (F, USM); Pantiacolla, serrania across Rio Madre de Dios from Shintuya, 480-840 m, 29 Oct 1979, Gentry et al. 27'20 (MO); Manu park, Cocha Cashu uplands, 400 m, 11045'S, 71000'W, 18 Aug 1986, Nuinez2800 (MO, NY); Tambopata, Concepci6n to Lago Sandoval, 350 m, 15 Jan 1967, Vargas C. 18561 (CUZ, US). PASCO: Km 1518 of Villa Rica-Pto. Bermudez rd., ca. 1600 m, 21 MIar 1984, Knapp et al. 6339 (BH, K, USM); km 15 of Palcazu rd. (km 73 of Villa Rica-Iscozacin-Pto. Mairo r(l.) along Rio Palcazu, ca. 380 m, 10?21'S, 7510'W, 17--18 Aug 1984, Knapp & Mallet 6643 (BH, K, US, USM); km 28 Repartici6n-Iscozacin (km 86 Villa Rica-Iscozacin-Pto. Mairo) Rio La Raya near Ameusha community of Laguna, ca. 350 m, 10?20'S, 75?10'W, 22-23 Aug 1984, Knapp & Mallet 6656 (BH, K, US, USM); Iscozacin, forests near PEPP camp, Rio Iscozacin, tributary of Rio Palcazu, ca. 320 m, 10?12'S, 75?13'W, 27 Aug 1'984, Knapp & Mallet 6657 (BH, K, USM). SAN MARTiN: Tarapoto, s.coll. (K); Rio Canuto, "Curarelandia"property of J. Schunke V. near km 23 of Tocache NuevoPuerto Pizana, ca. 475 m, 8?06'S, 76?36'W, 19 Dec 1981, Plowman & Schunke V 11510 (F); Mariscal Caceres, rd. to Pueblo Viejo de Tocache, 8 Feb 1970, Schunke V 3738 (F, NY, US); Mariscal Caceres, Quebrada de Saule

296 Grande,right bank of Rio Huallaga, 13 Sep 1970, Schunke V 4391 (F, NY, US); Caniutomleft bank of Rio Huallaga, 10 Oct 1970, SchunkeV 4503 (F, NY, US); rd. to Cachiyacu, 5 km from Puerto Pizana, 7 Oct 1971, Schunke V 5042 (F, K, US); Juan Guerranear Tarapoto,720 m, 29 Dec 1929, Williams6860 (F); San Roque, 1350-1500 m, 9 Jan 1930, Williams 7217 (F, US), 9 Feb 1930, Williams 7655 (F). UCAYALI:Coronel Portillo, 4-5 hrs above Caserio Abujao, on Rio Abujao, ca. 250 m, 8?15'S, 73?45'W, 16 Dec 1978, Diaz et al. 589 (MO); just above Pucallpa on Rio Ucayali, ca. 180 in, 8?26'S, 74?36'W, 2 Nov 1946, Fosberg 28946 (WIS); upper Ucayali, Cumaria, Tessmann 3355 (B [destroyed], F, NY); Pau-Cocha (Pucallpa), 200 m, 5 May tkowski6291 (MO). 1961, Wfoy BOLIVIA. PANDO: Manupiri, ca. 15 km S and 5 km E of San Silvestre, Reserva Manupiri-Heath, 200 m, 11?55S, 68?36'W, 25 Sep 1991, Pero, et al. 118 (MO). BRAZIL. ACRE: Sena Madureira,4 km from right bank of Rio laco, 5 Oct 1980, Cid & Nelson 2788 (NY); basin of Rio Purus, near mouth of Rio Macauhan (tributary of Rio Yaco), 9?20'S, 69? W, 8 Aug 1933, Kr-ukoff 5371 (NY); Rio Jurua between Mundurucus and Tatajuba, 13 May 1971, Maas et al. P128 76 (K, US, WIS); vic. of Aldota, Rio Jurua-Mirim, 24 May 1971, Maas et al. P13296 (F, K, NY, US, WIS); E of Rio laco, 10 km above Sena Madureira,4 Oct 1968, Prance et al. 7814 (NY, US, WIS). AMAZONAS: Maraa, right bank of Rio Japura, Lago do Macaco, 31 Oct 1982, Amaral et al. 266 (NY); banks of Rio Negro, at Isla Macara, near mouth of Rio Padarauiri,ca. 0?03'S, 64? W, 15 Jan 1946, Cardona 1300 (F, VEN); Fonte Boa, Rio Solimoes near mouth of Rio Jurua, 2-3?S, 65-66?W, 11 May 1986, Cid et al. 7363 (NY); S shore of Rio Negro, ca. 5 mi above confluence with Amazon, 27 Jun 1982, Costich 1078 (BH); Rio Negro, near oil refinery of Manaus, 3?10'S, 60?03'W, 14 Dec 1977, Keel & Guedes 372 (NY); basin of Rio Jurua, Marapata, 25 May 1933, KrukofJf 4569 (F, K, MO, NY, U); Sao Paulo de Olivenra, basin of creek Belem, 26 Oct-11 Dec 1936, Krukoff 8953 (F, K, MO, NY, , UUS); Rio Javari, behind army post at Palmeiras, 5008'S, 72049'W, 1 Aug 1973, Lleras et al. P1 7024 (NY, WIS); Rio Curuquete, Providencia, 27 Jul 1971, Prance et al. 14630 (NY); Rio Acari, 11 Mar 1945, Proctor Cooper 15 (NY); Rio Solim6es, Santo Ant6nio do IvA, mouth of Auati-Parana, 15 Oct 1968, Silva 2061 (NY); Manaus, Jan 1851, Spulce 1262 (K). PARA&:Shore of Rio Tocantins 2 km S of Baiao, 3 Jul 1935, Dr-ouiet1979 (F, NY, US); Rio Trombetas, Ilha Jacitara, 4 km SE of Oriximina, 60 m, 14 Jun 1980, Martinelli 6982 (K, NY, RB); Lageira, airstripon Rio Maicuru, ca. 800 m, 0?55'S, et al. 3319 (F, NY). 54?26'W, 20 Jul 1981, Stru&dwick RORAIMA: Between Maita (3?20'N, 63?34'W) and ParamiteriIndian village (3?25'N, 63?03'W), 18 Feb 1971, Prance et al. 10601 (NY, US, WIS). Local names. Venezuela. Amazonas: araniagato, parcha,tupirito,tupido de galapago. Peru. Loreto:mananoey (Huitoto), ocuerilla, sacha conjompe. Parcha is the common name for species of Passiflora in Venezuela and is almost certainly a labeling error.

FLORA NEOTROPICA

Solanumoppositifoliumis most closely related to S. confertiseriatumof westernEcuadorandPeru(see Knapp, 199lb). It differs from that species, however, in its Amazonian distribution,smaller leaves and inflorescences, and in its probably andromonoecious (ratherthan dioecious) breeding system. These sister taxamay have speciatedallopatrically(Knapp,199lb). Solanum oppositifolium is a widespread and variablespecies, andthe following aresome of its more well-markedgeographicalraces. 1)PlantsfromcentralandsouthernPeruandadjacent Brazil have simple inflorescences and pubescent berries.The leaf undersidesof plants from thesepopulationsdrya pale, goldenbrown.Ruiz andPav6n'stype specimenof Solaniumoppositifolium came from among these populations. 2) In the inundatedforests of the Amazon basin, plantsoften have long, branchedinflorescences, glabrous berries, and usually leathery, larger leaves. This race has been called Solanlini schizopodium(an adultplant) andS. viliflorumlnz (a tiny plant with its first inflorescence). Vasquez (1997) misidentified Iquitosplants of this race as S. confertiseriatuni, which only occurs on the western slopes of the Andes. 3) A higher elevation, isolated race on the slopes of the Guyana sandstone plateau has simple inflorescences, glabrous berries, and usually leatheryleaves. This form is relatively limited in distribution and has been called Solanum puberuloba. Manyotherformsmorelocalin distributioncould be described.It would be counterproductiveto provide formalnamesforthis mosaic of variation,as even these well-markedgeographicracesarenot easily delimitable entities. Specimens exist thatbridge all the gaps and a complex patternof variation exists in which several geographicalnormsappear.

96. Solanum palmillae Standl., Publ. Field Mus. Nat. Hist., Bot. Ser. 8: 149. 1930. Type. Mexico. Veracruz:In rocky gulches, Rancho Palmilla, Apr Fig. 163 1929, Purpus 13014 (holotype, F). Shrubs(?), height unknown; young stems sparselypuberulentwith uniseriatepapillatetrichomes ca. 1 mm long, soon becoming glabrate;barkof older stems dark,lenticellate. Sympodialunits plurifoliate. Leaves linear,widest at orjust proximalto the mniddle, glabrous,8.5-23 x 0.9-1.8 cm, with 8-1 0 pairsof main lateralveins, these prominentandyellowish below, the apex long-acuminate,roundedat the very tip, the base attenuate,decurrenton the petiole; petioles 3-8 mm

TAXONOMIC

297

TREATMENT tt|

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FLORA NEOTROPICA

long, winged from the decurrentleaf bases. Inflorescences oppositethe leaves, occasionally appearingterminal, furcate, 0.8-2 cm long, 10-30-flowered, minutely puberulentwith uniseriatepapillate trichomes 0.05-1 mm long;pedicel scars unevenly spaced, not overlapping,0.05-1 mm apart,slightly raisedfromthe surfaceof thepeduncle.Buds ellipsoid,the corollasoon exsertedfromthecalyx,puberulentwiththesameminute trichomes as those of the inflorescence. Pedicels at anthesis ca. 6 mm long, abruptlynarrowingat the base of the calyx tube,with a slenderbase ca. 0.75 mm diam., erect or slightly deflexed. Flowers with the calyx tube 1-1.5 mm long, abruptlynarrowingto the pedicel, the lobes widely deltoid, apiculate, 0.5-1 mm long, minutely papillose at the tips; corolla white (?), 1.1-1.2 cm diam., lobed nearlyto the base, planarat anthesis, the interpetalarsinuses very thin, the tips of the lobes minutelypapillose;anthersca. 3 x 0.5-1 mm, poricidal at the tips, the pores teardropshaped; free portion of thefilamentsca. 0.5 mm long, the filamenttube ca. 0.5 mm long;ovaryglabrous;stylestraight,4.5-5 mm long; stigma clavate,minutelypapilloseat the tip of the club. Fruita globose,6 mm diam.(immature),green(?) berry; fruitingpedicels woody, erect, ca. 1.1 cm long, 1-1.5 mm diam. at the base; calyx lobes becoming woody and slightly accrescent in fruit, 3-4 mm long. Seeds not known. Chromosomenumbernot known. Distribution (Fig. 165). Known only from the stateof Veracruzin Mexico, in rocky or sandyareason the eastem slope of the Sierra Madre Oriental. The species is known from so few collections that little is known aboutits habitatpreferencesand distribution. Specimens

examined.

MEXICO. VERACRUZ:

Barrancade Panoaya,Jul 1919,Purpus8415 (BM, NY, P); Arroyo,Camar6n,Oct 1926,Purpus11049(F); close to water,RanchoPalmilla,Sep 1933,Purpus16177(DS). Solanumpalmillae is superficiallyvery similar to S. imberbe(S. deflexiflorumspecies group) of eastern Panama and northern South America, S. inonadelphum of eastern Peru and Brazil, and S. amnicola (S. arboreumspecies group)of eastem Peru. All of these species have narrow,willow-like leaves andbranchedinflorescences.The similaritymost probably is dueto convergenthabitat;thesearespeciesgrowing in the bottoms of, and along, swift-flowing watercourses which often flood. Solanumpalmillae is most closely relatedto S. rovirosanum,also of Mexico, and may be derived from that species (Knapp, 199 lb).

97. Solanum rovirosanum Donn. Sm., Bot. Gaz. 48: 397. 1909.Type.Guatemala. AltaVerapaz: Cubilqiiitz, 350 m, Aug 1904, von Tuerckheim8716 (lecto-

type, US [US-1324746], designated by D'Arcy, 1974; confirmed by Knapp, 1991a; isotype, US [US-1324747]). Fig. 164 CyphomandrahomalophyllaStandl., Trop. Woods 10:50. 1927.Type.Panama. Bocasdel Toro:Almirante region, 1927, Cooper & Slater 39 (holotype, YU-n.v.; isotypes,F, US). SolanumschippiiStandl.,Publ.CarnegieInst.Wash. 461: 85. 1935. Type.Belize. In open forest near Machala,15 m, 26 Aug 1933, Schipp 544 (holotype, F [F neg. 44855]). Solanuim brenesiiC. V. Morton& Standl.,Publ.Field Mus.Nat.Hist.,Bot.Ser.18: 1076.1938.Type.Costa Rica. SanJose:San JuancercaSan Ram6n,3 Jan 1933, Brenes 16859 (holotype, F).

Shrubsor small trees, 2-10 m tall; young stems and leaves sparsely to densely puberulentwith erect uniseriatetrichomesca. 0.1 mm long, these golden in dry specimens;older stems glabrate,the barkdarkreddish-brown;bark of the trunks grayish-brown.Sympodial units difoliate, geminate.Leaves obovate to elliptic,widestdistalto themiddle,often in the distalthird of the leaf blade, glabrousabove, usually glabrousbeneath,occasionally puberulentalong the veins, the trichomeserect,uniseriate0.1-0.5 mm long;majorleaves 9-35 x 3.5-17 cm, with 7-10 pairsof mainlateralveins, these slightly raisedabove, prominentandpale yellow in dryspecimensbeneath,the apexacute,the base acute to attenuate, oftenwingedontothepetiole;petioleswinged from the decurrentleaf bases, 1.5-3 cm long; minor leaves differingfromthe majorones in size and sometimes in shape, occasionally more rounded than the majorleaves, 5-17 x 2.5-1 1 cm, the apexacute,thebase acuteto attenuate;petioles0.8-1 cm long.Inflorescences oppositethe leaves, simpleor many-timesfurcate,1.58 cm long, 5-25-flowered, glabrousto densely puberulent with erect uniseriate trichomes like those of the young stems and leaves; pedicel scars evenly and closely spaced,not overlapping.Budswhen young globose, the corollasoon exserted,the buds laterellipsoid, the exsertedportionof the corollaequalto the lengthof the calyx tube.Pedicels at anthesiswhite in live plants, deflexed, 0.7-1 cm long, taperingfrom the constriction at the base of the calyx tube to a slender base ca. 0.5 mm diam. Flowers with the calvx tube narowly campanulate,1-2 mm long, the lobes deltoid,0.5--1mm long, glabrousor puberulentwith the same uniseriate trichomesas those of the rest of the inflorescence;corolla white, 1.2-1.5 cm diam., lobed ca. 3/4 of the way to the base, the lobes planarat anthesis, the tips and marginsof the lobes puberulentandminutelypapillose; anthers3-4 x 1-1.5 mm, poricidalat the tips, the pores teardropshaped;free portion of thefilaments 0-0.25 mm long, the filamenttube 0.5-1 mm long;ovary glabrous or densely pubescentwith golden uniseriatetri-

299

TAXONOMIC TREATMENT

_~~~

.o

FXPLANTISGUTA MALAI-r S NSNAVISKI1 cO TAXIS.N (AIIIIS JONICAR(URTIIJ,lS I' QV'AR

DEPT,ALTAVERAPAZ. FLORAVONGUATEMALA.

FIG.

164. Solanum rovirosanum Donn. Sm. Lectotype at US.

300

FLORA NEOTROPICA

I

N ~~~~~~~~~~~~~~~~CZ-C

-

0

0

~

~

~

*

1900 FIG. 165. Distribution of So/anutmpa/millae (open circle) and S. rovirosanuim (solid circles).

chomes; stvle straight, in long-styled flowers 7-8 mm long, 1-1.5 mm long in short-styled flowers; stigma capitate, minutely papillose. Fruit a globose berry, glabrous or sparsely puberulent, pale lemon-yellow when ripe, otherwise green, 1-1.5 cm diam.;fruitingpedicels thick and woody, erect, occasionally deflexed from their own weight, 0.8-1.3 cm long, the length quite variable even within an individual, 1.5-2 mm diam. at the base, ca. 5 mm diam. at the tip. Seeds pale tan, ovoid-reniform, 2.5-3 x 2-2.5 mm, the surfaces minutely pitted. Chromosome number: n = 12 (vouchers Knapp 804, 844, 848, 861). Distribution (Fig. 165). From southern Mexico to western Panama and northwestern Colombia, sea level to 1500 m, in both primary and secondary forests. Selected specimens examined. MExIco. S.loc., Galeotti 7027 (F); 1791, Haenke 1153 (F). CHIAPAS: Cerro del Boquer6n, Jun 1914, Purpus 7318 (US); Palenque archeological site, 3 mi S of Palenque, ca. 500 ft, 27 Jun 1970, Thorne & Lathsrop 40528 (DS, LL, RSA). DISTRITO FEDERAL: Temascaltepec, Pungarancho, 14 Jan 1933, Hinton 3145 (F, NY). OAXACA: S.10C., 184143, Liebmann 11907 (US). TABASCO: Chinantla, Jul 1844, Galeotti 1225M (US); Sta. Maria Chimalapa, Santa Maria, disturbed areas in village, 300 m, 16?54'30"N, 91?41'W, 15 Jun 1984, Herncandez G. 137 (NY); La Esperanza, Comaltepec, Ixtlan, ca. 5000 ft, 18 Apr 1970, MacDougall s.n. (NY); San Isidro, Balancan, 7-11 Jun 1939, Matucda3359 (LL, MO, NY, US); Mayito, 29 Jul 1889, Rovir-osa 544 (NY, US). BELIZE. Cayo, Blue Hole section, Hummingbird hwy., 27 Sep 1955, Gentle 8884 (F); Machaca, ca. 50 ft, 26 Aug 1933, Schipp 1221 (F, MO, NY).

GUATEMALA. ALTA VERAPAZ: Between Sachi and Sacacac, 150-180 m, 20 Mar 1942, Stevermark 45128 (F). PETEN: Rio San Luis, at km 118 rd. to Poctun, ca. 200 m, 3 Dec 1970, Tun Ortiz 1456 (F, NY). NICARAGUA. San Juan de Nicaragua, 11 May 1841, Friedrichsthal 504 (W). Rio SAN JUAN: Sabalo, 3 km from jct. of Rio Sabalo and Rio San Juan, 40-50 m, 26 Sep 1982, Araquiistain 3195 (MO); near Cano Chontalefio 20 km NE of El Castillo, 200 m, 18-21 Apr 1978, Neill & Vincelli 3540 (MO). ZELAVA: Ca. 6 km upriver from Barra de Punta Gorda, S side, 8-10 m, 11030'N, 83049'W, 30 Sep 1981, Stevens 20738 (MO). COSTA RICA. ALAJUELA: Los Angeles de San Ram6n, 6 Mar 1929, Brenes 6723 (F, NY); La Palma de San Ram6n, 10 Aug 1928, Brenes 6261 (F, NY); near San Ram6n, 21 Jan 1932, Brenes 15012 (F, NY); San Pedro de San Ram6n, La Catarata, 29 Jan 1936, Brenes 21434 (F, NY); San Pedro de San Ram6n, 29 Jan 1936, Brenes 21435 (CU, F, NY); Quebrada Marin, ca. 7 km E of Ciudad Quesada, 500 m, 10022'N, 84024'W, 17-18 May 1968, Burger & Stolze 4942 (NY); Finca los Ensayos, 11 mi NW of Zarcero, 900 m, 15 Aug 1977, Croat 43517 (MO); Finca la Constancia, Buena Vista, San Carlos, 850 m, 3 Mar 1963, Jimenez 441 (F, NY); between Finca de los Ensayos and Zapote, Alfaro Ruiz, 1450 m, 4 Mar 1963, Jimenez 451 (CR, F, NY); Finca la Constancia, Buena Vista, San Carlos, 850 m, 30 Mar 1963, Jimenez 580 (F, NY); between Ciudad Quesada and Florencia, San Carlos, 600 m, 22 Jan 1966, Jimenez 4024 (F, NY); rd. from Monteverde to Pefias Blancas, 1400 m, 10?25'N, 84050'W, 13 Apr 1981, Knapp & Mallet 860, 861 (BH, CR); Quebrada Guillermina, N side of Volcan Arenal, 500 m, 10?29'N, 84?42'W, 21 Apr 1973, Lent et al. 3383 (CR, F, MO); N side Volcan Arenal, 800 m, 10028'N, 84042'W, 11 Apr 1974, Lent 3859 (F, MO); 4 km SE of Fortuna,400-500 m, 10029'N,84?43'W,

TAXONOMIC TREATMENT 29 Apr 1983, Liesner et al. 15219 (MO); Zarcero, 16 Jan 1938, Smith H140 (F); Alfaro Ruiz, Zarcero, 1500 m, 10 Feb 1948, Smith 201 (US); San Carlos, Villa Quesada, 650 m, 29 Mar 1939, Smith F1840 (F, NY, US); Zarcero, 6500 ft, 8 Apr 1937, Smith 10/Stork 4110 (F, NY); Rio Barranca, near San Ram6n, 1300-1400 m, 5 Apr 1913, 'Tonduz17688 (F, US); San Ram6n, 1300-1400 m, 10 May 1913, Tonduz 17696 (US); between Angeles de Norte and Balsa de San Ram6n, 7-18 km N of San Ram6n, 550-1050 m, 8 Aug 1975, Utley & Utley 2765 (F, MO); ca. 11 mi S of Quesada, 4850 ft, 12 Jul 1962, Websteret al. 12197 (US); CordilleraCentralnear San Juan de Laja, ca. 15 km N of Zarcero,ca. 1350 m, 7 Feb 1965, Williams et al. 29020 (F, NY). CARTAGO:Cerro de la Carpinteria, 1500-1850 m, Feb 1924, Standlev 35732 Near Laguna de Arenal, 600 m, 26 (US). GUANACASTE: Jun 1930, Brenes 12656, 13146 (F); Hacienda Quebrada Azul, Volcn Tenorio, 400-600 m, 18 Feb 1930, Dodge & Thomas 6185 (F, MO); gorge below Tilaran, rd. to Hacienda Quebrada Azul, 19 Feb 1930, Dodge 6197 (MO); Cerro de San Jose, 760-800 m, 15 Feb 1930, Dodge & Thomtas6383 (MO); vic. of Tilaran, 500-650 m, 10-31 Jan 1926, Standlevl& Valerio 44495 (US); La Tejona N of Tilaran, 600-700 m, 25 Jan 1926, Standley & Valer-io45910 (US); vic. of Tilaran, 500-650 m, 1031 Jan 1926, Stanidlei'& Valerio46636 (US); Arenal, 600 m, 21 Aug 1923, Valerio 96 (US); Cord. de Tilaran, La Sierra,Campos de Oro, jct. Rio Canuasand Rio Abangares, 1000 m, 10?22'N, 84?55'W, 31 Mar 1993, Bello 4966 (INB); Parque Nacional Rincen de la Vieja, trail to volcano, 800--1200 m, Bittner & Venschatt 1936 (INB); P.N. Rincn de la Vieja, Tempisque watershed, Est. Las Pailas, trail to volcano summit, 1200 m, 10?47'50"N, 85021'10"W, 27 Jun 1997, Ramirez & Rojas 470 (INB). HEREDIA:ILaSelva, Rio Puerto Viejo ca. 2 km upstream from confluence with Rio Sarapiqui, 100-200 m, 26 Apr 1973, Gentry & Burger 3015 (F, MO); Barranca del Rio Sarapiqui, Colonia Virgen de Socorro, 700 m, 3 Feb 1984, Gomez Laurito 9858 (F); between Santo Domingo and Rio Virilla, 1110 m, 9?58'N, 84?05'W, 6 Jan 1974, Lenit3752 (CR, F, MO); Finca La Selva, OTS Field Station on Rio Puerto Viejo just E of confluence with Rio Sarapiqui,CentralTrail,Line 2500, ca. 100 m, 1 Aug 1980, MacDougal 1006 (DUKE, NY); La Selva near Puerto Viejo, Arboretum II, 90 m, 7 Aug 1971, Opler 361 (F); La Selva, 3 km SE of Puerto Viejo, 45 m, 8 Aug 1972, Opler 1509 (F, MO), 30 Aug 1973, Opler 1874 (F, MO); Cerros de Zurqui NE of San Isidro, 2000-2400 m, 3 Mar 1926, Stand/lev& Valerio 50253 (US). LIMON: SWmost ridge of Cerro Coronel, NW-facing slope, just S of Rio Colorado, 10-80 m, 10?40'30"N, 83039'30"W, 1718 Sep 1986, Davidse & Herrera 31331 (INB); near Rio Catarata, between Bribri on the Rio Sixaola and Caribbean coastal plain, 50-100 m, 9?37'N, 82?49'W, 10 Feb 1977, Gentrn3732 (CR, F, MO); nearFincaPais, Talamanca, ca. 50 m, 5 Jul 1983, Gomez Laurito 9501 (F); near Guapiles, 250 mn,17 Jul 1964, Lent 61 (F, LL, MO); El Carmen, Siquirres,40 m, 10?13'N, 83?32'W, 19 Dec 1973, Lent 3678 (CR, F, NY); P.N. Tortuguero,Est. Agua Fria, 3 km S of the station, 40 m, 10?25'N, 830341W,22 Oct 1987, Robles 1113 (INB); Suerre and Dos Bocas, Rio

301 Parismina and Rio Reventazon drainages, 0 m, 3 Oct 1951, Schank & Molina R. 4245, 4290 (F, US). PUNTARENAS: Rio Nuevo, near Puerto Jimenez de Osa., 10-15 m, 4 Apr 1930, Brenes 12204, 12214 (F): E of Las Cruces, 5-6 km S of San Vito property of R. Wilson, 1100-1200 m, 8?47'N, 82?58'W, 15-16 Jan 196?, Burger & Matta U. 4465 (CR, F, MO, NY, U); between Rio Coton and Rio Negro, ca. 15 km from Sabalitc, 1200-1300 m, 8?53'N, 82?54'W, 17-18 Jan 1967, Burger & Matta U. 4520 (CR, F, MO); 2-8 km N of Cafias Gordas, 1000-1 100 m, 26 Feb 1973, Busey 652 (MO); Rio Coto Brus near Cot6n, 23 km N of La Umnin, Panama border, 9 Aug 1974, Croat 26629 (MO); 5 krn W of Rincon de Osa, Osa peninsula, 50-200 m, 8'42'N, 83?31'W, 24-30 Mar 1973, Gentry & Burger 27788(F, MO), Gentry & Burger 2839 (F, MO, NY); Finca Las Cruces, near San Vito, 4000 ft, 19 Aug 1977, Gilbert 10 (LL, MO); Monteverde, Wallace farm, 1400 m, 10`25'N, 84?50'W,12 Mar 1981, Knapp 844 (BH, CR); Monteverde. Rockwell farm, edge of plateau, 1200 m, 10c25'N. 84?50'W, 19 Mar 1981, Knapp 847 (BH, CR); Monteverde Cloud Forest Reserve, S slopes, 1500 m, 10c25'N, 84?50'W, 8 Apr 1981, Kniapp & Mallet 848 (BH. CR,; Monteverde, Rockwell farm, plateau edge, 1200 m. 10?25'N, 84?50'W, 16 Apr 1981, Knapp & Mallet 869 (BH, CR); ca. 3 km up Rio Agua Buena, Rincon de Osa, 20-150 m, 12 Feb 1974, Liesner 2078 (CR, MO); Osa Peninsula, Parque Nacional Corcovado, near Pavo, 5 ni. 8?30'N, 83?37'W, 7 Jul 1977, Liesner 3037 (MO); at NW end of Golfito Bay, ca. 4 km W of hospital in Golfito, 10 m, 5 Mar 1971, Nee & Moni 3549 (MAD, US, WIS); Osa Peninsula,W of Rincon de Osa, 100 ft, 7 Aug 1967, Raven 21581 (CAS, CR, F, NY); above Laguna, San Vito de Java. 16 Mar 1956, Schulbert1228 (US). SAN Jost: Below La Palma along Rio Claro (upper Rio La Hondura), trail to Guapiles, 1000 m, 10?03'N, 83?58'W, 1 Jan 1967, Burger 4156 (F, MO, NY); Bajo La Hondura below La Paloina NE of San Jer6nimo, 1000-2000 m, 10?03'N, 83"58'W, 23 Oct 1975, Burger et al. 9419 (F); basin of El General, 675-900 m, May 1940, Skutch 4943 (F, MO, NY, US). PANAMA. BoCAS DEL TORO: Rio Terebe just below Puerto Palenque, 350 ft, 12 Apr 1968, Kirkbria'e & Duike 555 (MO, NY); 4 mi N of Almirante, 13 Aug 1964, McDaniel 5097 (MO); s.loc., 30 Jul 1940. von Wedel 226, 241 (MO); Water Valley, 12 Sep 1940, von Wedel731 (MO, US); vic. of ChiriquiLagoon, 8 Oct 1940, von Wedel 1085 (MO, US). CHIRIQUi: S.loc., Hart CL 120 (K). PANAMi: Campo Tres, 3 mi NE of Alto de Pacora, 500-800 m, 10 Mar 1973, Croat 22742 (F, MO, US); Cerro Jefe, ca. 800 m, 9?12-13'N, 79?22'W, 9 Jan 1971, Webster et al. 16471 (MO). COLOMBIA. CESAR: Sierra Nevada de Santa Marta, SE slopes, environs of Sagrome, valley of the Donachui, land of the Ika, ca. 2000 m, 9 Jan 1977, Davis 571 (K, US). C6RDOBA: Mun.Tierralta,Rio Esmeralda above confluence with Rio Sinu, 180 m, 27 Jul 1986, Bernal et al. 1178 (MO). In the cladogram of the sessile species groulp, Solanum rovirosanum is part of a trichotomy in the

302

oppositifolium subclade (Knapp, 1991b). Solanum rovirosanium itself has no automorphiessuggesting it is close to the stem. Species such as this have been hypothesized as ancestralspecies in some groups(see Kitchinget al., 1998), but aremore commonlyreferred to as stem species (Forey, 1992) or have recentlybeen termedplesiospecies (Olmstead, 1995). Fruit pubescence varies geographically in Solanumrovirosanum,with populationsfrom Mexico andGuatemalahavingpubescentovariesandberries,and populationsfrom Costa Rica and Panamahaving glabrousovariesandberries.The significanceof this characteris not yet understood.The same type of polymorphismoccursin S. sessile of the Amazonbasin.Material with glabrousbemreshas been called S. brenesii. The two races areotherwisenot differentfromone another. Themoreeasterlypopulations(i.e., CostaRica,Panama, and Colombia) of S. rovirosanummay be specifically distinct,but furtherfield studyis necessary,particularly in the northernpartof the species range. In Costa Rica, Solanum rovirosanumgrows in both primaryand secondaryforest, and the two forms are quite differentmorphologicallyandareoften identified as different species. The primaryforest plants, growing under closed canopy, have large, obovate leaves, and often have simple inflorescences. Those plants frompastureedges have smaller,often elliptic, rathercoriaceous leaves, and larger,usually branched inflorescences. These differences are striking at first glance, butupon closer examination,the two fonns are obviously the same species. Forest and secondary growthplants arevisited by differentbees, andbloom at different times in the site where I studied them (Monteverde, see Knapp 1986a). Secondary growth plants arevisited primarilyby Melipona spp., and forest plantsby bumblebees (see TableIX). The peaks of flowering time differ for the two forms, but the populationsstudiedwere at the extremesof an elevationand ariditygradient,so this is perhapsnot surprising.Plants betweenthetwo studypopulationsareperhapsin bloom in the interveninginterval,thus maintaininggene flow in this species; however the situationmay meritcloser investigation.

FLORA NEOTROPICA (no specimen cited, see discussion). Solanum dibrachiatum Van Heurck & Mull. Arg., Observ. Bot. 59. 1870. Type. Peru. San Martin: Prope Tarapoto, 1855-1856, Spruce 4250 (lectotype, G, designated here (Morton neg. 851 1); isolectotypes, AWH, BM, C-n.v. [F neg. 23201], G-DC, K, NY, P [Morton neg. 8180]; frag. F). Solanum marmellosanum Bitter, Repert. Spec. Nov. Regni Veg. 16: 89. 1919. Type. Brazil. Amazonas: Rio Madeira, Santa Maria de Marmellos, Mar, Ule 6109 (lectotype, HBG, designated here; isolectotype, B [destroyed]). Pheliandra herthae Werderm., Notizbl. Bot. Gard. Berlin 15: 54. 1940. Type. Ecuador.Pastaza:Mera, ca. 1000 m, 10 Nov 1938, Schult-e-Rhonhof 2973 (holotype B, no duplicates or photos located, destroyed?). Solanum grandifolium C. V. Morton, Contr. U.S. Nat. Herb. 29: 51. 1944. Type. Colombia. Vaupes: San Jose de Guaviare,240 m, 4 Nov 1939. Cuatrecasas 7394 (holotype,US; isotype, COL-n.v.).

Shrubsto small trees, 2-8 m tall; young stems and leaves glabrousto minutelypuberulentwith erect uniseriatetrichomes ca. 0.1 mm long, these golden in dryspecimens;older stems glabrate,occasionallywith some scattereduniseriatetrichomes;barkof olderstems reddish-brown,latergrayish.Sympodialunitsusually difoliate,geminate.Leavesvery large,obovate,widest distalto the middle,usuallyin the distalthird,glabrous above, occasionally with minuteuniseriatetrichomes along the veins beneath,this conditionextremelyvariable; majorleaves 15-40 x 9-20 cm, with 9-1 1 pairs of mainlateralveins, these raisedabove,prominentand golden beneath,the apex acute,the base variable,from sessile and slightly cordateto acute; petioles 0-1 cm long, oftenbroadlywingedwith the decurrentleaf base; minor leaves differing from the major ones in shape and size, orbicularto obovate, 2.5-13 x 1.5-9 cm, the apex acute,the base roundedto acute;petiolesca. 5 mm long. Inflorescences terminal and overtopping the leaves at the shoot tips, later lateral and opposite the leaves,complexandmany-timesfurcate,6-25 cm long, 20-100-flowered, sparselyto densely puberulentwith erectuniseriatetrichomeslike those of the young stems and leaves, the trichomes0.1-0.3 mm long, golden in dryspecimens;pedicelscars irregularlyspaced 1-5 mm 98. Solanum sessile Ruiz & Pav.,Fl. Peruv.2: 35, fig. apart,corky and slightly raisedfromthe inflorescence 167a. 1799. Type. Peru. Huanuco:Mufia,Ruiz & axis. Buds globose and completely enclosed in the caPaiv6ns.n. (lectotype, MA, designated by Knapp, lyx when quiteyoung, laterthe corollaexsertedandthe 199la; isolectotypes, MA, frag. MPU). Fig. 166 buds long-ellipsoid, the corolla ca. 2 times longerthan the calyx tube.Pedicels at anthesisdeflexed,white and Solanum pulchrum Dunal in Poir., Encycl. Meth., Botanique, suppl.3: 750. 1814.Type.Brazil.S.loc., fleshy, 0.6-1.1 cm long, taperingfrom the abruptnarno collectorcited(holotype,P [Mortonneg. 8305]). rowing below the calyx tube to a slender base ca. 0.5 Solanum pulchrum Dunal var. peruvianum Dunal in mm diam.Flowerswith the calyxtubethickandwoody, DC., Prodr. 13(1): 96. 1852. Type. Peru. Loreto: fleshy, white or greenish-whitein live plants, broadly Ad ripas Amazonumet Huallagae,Poeppig s.n. cup-shaped,2.5-3.5 mm long, the lobes broadly del-

TAXONOMIC TREATMENT

303

CLXVII

. . . . _.. ...._... .-_---. ..

7;*

a SOLANUM

SOLANUM

FIG. 166. Solanum sessile Ruiz & Pav6n, plate 167a of Ruiz & Pav6n's Flora Peruviana.

FLORA NEOTROPICA

304

toid, occasionallyapiculate,the marginsthickenedand whitish in dry material,glabrousor minutely puberulent with erect uniseriatetrichomes ca. 0.1 mm long; corolla white and fleshy, 1.5-2.5 cm diam., lobed 2/3 of the way to the base, the lobes planarat anthesis,the tips and margins of the lobes densely papillose with occasionaluniseriatetrichomes;anthers3.5-A x 1-1.5 mm, the ten-ninal0.5 mm thickenedandpaler,poricidal at the tips, the pores teardropshaped;free portion of thefilaments 0-0.5 mm long, the filament tube 0.5-1 mm long; ovary glabrous or pubescent with minute, golden, uniseriatetrichomesca. 0.1 mm long; style in long-styledflowers0.8-1 cm long, in short-styledflowers ca. 1 mm long, straight;stigma capitate,minutely papillose.Frulita globose, greenberry,1-1.5 cm diam., in dryspecimenswoody andhard;fruitingpedicels erect and woody, 1-1.7 cm long, occasionally deflexed due to the weight of the fruits and the size of the inflorescence, 1.5-2 mm diam. at the base, thickerat the apex, ca. 3 mm diam.; calyx lobes slightly accrescent and woody in fruit.Seeds pale tan, ovoid-reniform,2.5-4 x 2--3 mm, the surfacesminutely pitted.Chromosome number:n=i 12 (vouchersKnapp& Mallet 6207, 6316, 6391, 6569). Distribution (Fig. 168). On the eastem slope of the Andes from Colombia to Bolivia, extending into AmazonianBrazil. A species of wide altitudinalrange, from 100m in theAmazonbasinto 1800m on the slopes of the Andes in centralPeru and Venezuela. Selected specimens examined. VENEZUELA. BARINAS: Ca. 8 km N of Bumbum, alongRio Bumbum, ca. 300 in, 8?16'N, 70?48'W,25 Oct 1984, Knapp& Mallet

6839 (BH, F, K, MO, MY,NY, VEN);Knapp& Mallet 6841 (BH, F, K, MY, NY, VEN). MERIDA: Dtto. Libertador, caminoentreEl Morroy Aricagua,La Loma de la Vagabunda,cerca El Morro,2400-2500 m, 23 Jul 1974, Badillo 6533 (MY); Dtto. Andres Bello, San Eusebjo, 2250 m, 9 Aug 1978, Benitez de Rojas 2401 2200 m, 15 May 1953, Little15168 (MY);La Carbonera, (VEN):La Carbonera, ca. 20 km NW of Ejido,2600 m, 2 Oct 1953, Little 15590 (VEN); La Carbonera,carretera via Azulita, 17 Feb 1979, Marcano Berti & Salcedo M.

5-979 (MER);Dtto.CampoElias,al S de PuebloNuevo, betweenMucususandCerrola Becerrera,2000-2500 m, 18 Sep 1984, Ortega & Diaz 214 (PORT);Dtto. Libertador, Asentamiento Prado Verde, 2020 m, 4 Mar 1976, Qliintero & Herniandez 231 (MER); Dtto. Arzobispo Chac6n,paramo de Canagud,2187 m, 6 Apr 1966,Ruiiz Terdn2979 (MER);Dtto.RivasDavila, betweenla Playita

and la M, above Bailadores, 2300-2600 m, 26 Sep 1977, Ruiiz Teran et al. 14332 (MERF); along quebrada

of Cuestadel Barroand Mesa del Trapiche,tributaryto Rio Capuri,between Canaguaand El Molino, 25302715 m, 11 May 1944, Stevermark 56488 (F, VEN); La Carbonera,Finca San Isidro, Merida-La Azulita rd., 2100-2200 m, 17 Dec 1984, Weitzmann 196 (NY).

COLOMBIA. CAQUETA: Puerto Lara, Rio Orteguaza,Laralandia,24 Jan 1969, Plowman & Kennedy 2270 (F, K, NY, US). PUTUMAYO: Rio Putumayo, above Puerto Ospina, toward La Loma, 230--250 m, 19 Nov 1940, Cuatrecasas 10661 (F); rd. between Mocoa and Villa Garz6n, Gutierrez et al. Mocoa 2 (US); Umbria, 325 m, 0?54'N, 76?10'W, Dec 1930, KlIng 1858 (BM, K); Sta. Rosa del Rio Guarmes, trail to San Antonio, ca. 300 m, 1 Dec 1968, Plowman 2088 (GH, K). ECUADOR. NAPO: Payomino project of Min. de Agricultura, 200 m, 0?26'S, 77?01'W, 25 Feb 1980. Brandbyge & Asanza C. 30016 (NY); on rd. to Auca oilfield, between Auca 5 and Auca 6, 200 m, 0035'S, 76?54'W, 26 Feb 1980, Brandbvge & Asanza C. 30038 (NY, U); downstreamRio Napo, ca. 3 km from Coca, 300 m, 0?27'S, 76?56'W, 26 Mar 1980, Brandbvge & Asaniia C. 30346a (NY); Rio Wai si aya a northerntributaryof Rio Aguarico, ca. 6 km upriverfrom San Pablo, 300 m, 0015'S. 76?21'W, 10 Aug 1980, Brandbyge & Asan-a C, 32745 (NY); Reserva Biol6gica Jatun Sacha, Rio Napo, 8 km E of Misahualli, 450 m, 1?04'S, 77?36'W, 19-28 Mar 1987, Ceron 1034 (MO, NY); along rd. between Tena and Puyo, 39.9 km N of Puyo, 800 m, 22 Dec 1979. C7roat 49635 (MO, NY); Rio Jivino, Limoncocha, 15 Mar 1968, Harling et al. 7745 (NY); Misahualli, confluence Rio MisahualliRio Napo, 500 m, 1?03'S, 77?41'W, 13-14 Aug 1979, Holm-Neilsen et al. 19147 (NY); Misahualli, confluence Rio Misahualli-Rio Napo, 500 m, 1?03'S, 77041W, 13--14 Aug 1979, Holm-Nielsen 19265 (NY); Cascada San Rafael (Coca Falls) turnoff 71.3 km NE of Baeza on rd. to Lago Agrio (Nuevo Loja), 1200-1400 m, 0010'S, 77?40'W, 25 Jan 1984, Knapp & Mallet 6203 (BH, K, QCA. QCNE, US); CascadaSan Rafael (Coca Falls), 71.3 km NE of Baeza on Baeza-Lago Agrio rd., 1200-1400 m, 0?10'S, 77?40'W, 25 Jan 1984, Knapp & Mallet 6207 (BH, NY, QCA. QCNE, US); on Baeza-Lago Agrio rd., 50 km NE of Baeza, at bridge over Rio Azuela, ca. 1400 m, 0?10'S. 77040'W, 25 Jan 1984, Knapp et al. 6210 (BH, QCA, QCNE, US); Aniangu,NW corner of Parque Nacional Yasuni, 300 m. 0?32'S, 76?22'W, 1-30 Apr 1985, Korning & Thomsen 47177 (AAU); Aiiangu, NW corner of Parque Nacional Yasuni, 260-360 m, 0?33'S, 76?22'W, 1-30 Apr 1985, Korning& Thomsen58696 (AAU); San Pablo de Aguarico, wa:ya creek, 21 Jan 1984, Lescur-e2127 (NY); San Pablo de Aguarico, Rio Shushufindi, 8 Feb 1984, Lescurse 2167 (NY); Reserva Biol6gica Jatun Sacha, Rio Napo 8 km below Misahualli, 450 m, 104'S, 77?36'W, 24 Aug 1986. Neill & Cer6n 7245 (MO, NY); Rio Arajuno, Sola Cocha. 500 m, 1?07S, 77?36'W, 23-27 Oct 1985, Palacios et al. 868 (MO, NY); 8 km downriver and 1 km S from Misahualli on Rio Napo, 500 m, 1?04'S, 77037'W7,30 Oct 1985, Palacios & Neill 912 (MO, NY); Estaci6n Experimental INIAP-Napo, San Carlos, 8 km S of Los Sachas, 350 m, 0?20'S, 76?56'W, 8 Sep 1986, Palacios & Neill 1188 (MO, NY); Reserva Biol6gica JatunSacha, rightbank Rio Napo 8 km downriverfrom Misahualli,450 m, 1?04'S, 77036'W, 13 Feb 1987, Palacios & Neill 1541 (MO, NY); 8 km N of Coca (Puerto Francisco de Orellana), 250 m, 0?24'S, 77?00'W,8 Apr 1085, Zarumnia et a!. 70 (MO, NY). PASTAZA:Curaray,Jesus Pitishka,ca. 200 mn19 Mar 1980,

TAXONOMIC TREATMENT Hai-ling & Andersson 17440 (NY); Curaray,N bank 2 km W of school, 250 m, 1?22'S,76?58'W,18 Mar 1980, HolmNielsen et al. 21888 (K, NY, U); Rio Villano, 260 m, 1?25'S, 77?02'W, 24 Mar 1980, Holm-Nielsen et al. 22682 (NY); Pacayacu on Rio Bobonaza, ca. 16 km NW of Sarayacu, 10 Aug 1979, Lugo S. 5255 (NY). ZAMORA-CHINCHIPE: Outskirts of Zamora, ca. 1000 m, 4?05'S, 78?55'W, 4 Feb 1984, Knapp & Mallet 6241 (BH, K, QCA, QCNE, US). PERU. Pariahuanca, Mathews 842 (BM, K, NY). AMAZONAS: Village of Kusi, Rio Numpatakai (trib. Rio Cenepa), 1100-1300 ft, 10 Mar 1973, Berlin 926 (F, LL, NY); Caterpiza, W bank of Rio Santiago, 19 Sep 1979, Huashikat 710 (MO); Bagua, QuebradaChinganza, 8 km N of Muyo (km 472 of Oleoducto Norperuano), 350-400 m, 525'S, 78?28'W, 5 Jul 1984, Knapp & Mallet 6563 (BH, K, NY, US, USM); trail on N bank of Rio Imaza (Chiriaco), N of village of Chiriaco, ca. 350 m, 5?09'S, 78?18'W, 6 Jul 1984, Knapp & Mallet 6569 (BH, K, NY, US, USM); 6570 (BH, US, USM); Pongo de Manseriche, Rio Santiago, 1924, Tessmann 3871 (G); Quebrada Caterpiza, 2-3 km behind village Caterpiza, Rio Santiago valley, 200 m, 3?50'S, 77?40'W, 20 Feb 1980, Tunqui925 (MO); Bagua, left bank of Rio Mara-nonopposite Quebrada Mirana (opposite km 277 of Marafion rd.), 425-450 m, 16 Sep 1962, Wurdack2016 (F, K, USM). AYACUCHO: Ccarrapa,between Huanta and Rio Apurimac, 1500 m, 5, 6. 17 May 1929, Killip & Smith 22365 (NY, US). Cuzco: La Convenci6n, slopes and banks of Rio Mapitunuari, 2 hrs walk from Luisiana,Rio Apurimacand Indianencampment, ca. 780 m, 12?38'S, 73?41'W, 29 Jul 1968, Dudley 11361 (F); Pilcopata-Salvaci6n rd., near Madre de Dios border,ca. 450 m, 13 Oct 1979, Gentty et al. 26718 (MO); Quispicanchis, environs of Quince Mil, trail to Rio Nusinescatu, 500-600 m, 13?15'S, 70?45'W,22 Apr 1984, Knapp & Mallet 6372, 6376 (BH, CUZ, K, US, USM); Quince Mil, ridges N of village, 500-600 m, 13?15'S, 70?45'W, 23 Apr 1984, Knapp & Mallet 6378 (BH, CUZ, K, US, USM); environs of Quince Mil, trail to Rio Nusinescatul,500-600 m, 13?l5'S, 70'45'W, 24 Apr 1984, Kniapp & Mallet 6391 (BH, CUZ, K, US, SUM); Limonchayoc, ca. 1 km from Cuzco-Pto. Maldonado rd., ca. 16 km E of Quince Mil, 400-500 m, 13'15'S, 70?40'W, 25-26 Apr 1984, Kniapp& Mallet 6401 (BH, CUZ, US, USM); vic. of village of Pilcopataalong Rio Pilcopata,700800 m, 13?05'S, 71?10'W, 10 May 1984, Knapp & Mallet 6421 (BH, K, NY, US, USM); Kosfiipata,Quitacalz6n(Qda. Sta. Alicia), ca. km 163 on Lucre-Paucartambo-Shintuya rd., 1100-1200 m, 13?07'S,71?15'W, 11 May 1984, Knapp & Mallet 6426 (BH. CUZ, US, USM); along Rio Carb6n nearAtalaya.confluence of Rio Carb6nand Rio Alto Madre de Dios, 500-600 m, 13000'S, 71'07'W, 15 May 1984, Knapp & Mallet 6454 (USM); Quince Mil, 27 Aug 1969, Lockwood 574 (CUZ, MO, USM); La Convenci6n, Valle de Santa Ana, above Quillabamba,rd. to Potrero, 4800 ft, 18 Jan 1975, Plowman & Davis 4763 (K, MO); Paucartambo,Sta. Isabel, Kosfiipata, 1320 m, 5 Dec 1947, Vargas C. 6765 (CUZ); Quillabamba, QuellounoChirumbia, 200-2400 m, Dec 1947, Vargas C. 11405 (CUZ); Paucartambo,Atalaya, Kosfiipata, 750 m, 22 Nov 1962, Vargas C. 13993 (CUZ, WIS); Paucartambo,

305 Montefiesa, Kosfiipata, 800-900 m, 28 Sep 1966, Vcirgas C. 17808 (CUZ, US). HuANuco: Tingo Maria,Rio de Pavas, 13 Mar 1977, Boeke 1246 (BR); rd. to Aucayacu, ca. 3 km from jet. with rt. 16, 15 km N of Tingo Maria, 2200 ft, 9 Nov 1975, Davidson 3435 (MO); Pachitea, Dantas, km 42 CarreteraMarginal,22 Jan 1987, Diaz & Balde6n 2306 (MO, NY); Puerto Inca to Sungaro, foot trail between Rio Pachitea and Rio Sungaro, 250-300 m, 9?18'S, 75?00'W, 13 Sep 1982, Foster 8747 (F); Codo de Pozuzo, trail S of settlement to river, 450 m, 9?40'S, 75025'W,21 Oct 1982, Foster 9374 (F); Cushi, ca. 5000 ft, 19-23 Jun 1923, Macbride 4815 (F); rd. from Huanuco to Tingo Maria, N of Carpish pass, 54 km NE of Huanuco, 2310 in, 6 Dec 1981, Plowman & Rurv,11164 (F); Tingo Maria, Jardin Botanico, Ave. Pimental 358, seed from forest reserve S of town, 670 m, 7 Dec 1981, Plowman & Ramirez R. 11186 (F); Pachitea, Bosque Nacional Iparia, along Rio Pachitea near camp Miel de Abeja, 1 km above Toumevista, ca. 20 km above confluence with Rio Ucayali, 300-400 mn.26 Oct 1967, Schunke V 2269 (NY); Bosque Nacional Iparia, along Rio Pachitea near town of Puerto Inca, trail to Quebrada de la Viuda, 8 km from Puerto Inca, 400-500 m, 12 Dec 1968, Schunke V 2852 (F, G); Mufia, 2600 m, 1909-1914, Weberbauer6719 (F, MOL); Pillao, 2700 m, 2 Mar 1946, Woytkowski34168 (F); Huacapistana,20002100 m, 25 Jun 1948, Ferreyra 3599 (USM), 26 Jun 19'48, Ferreyra 3661 (USM); below Huacapistana, 1700-1800 m, 21 Sep 1955, Ferreyra 11235 (USM); above Huacapistana, 2100-2200 m, 24 Sep 1955. Fcrre)ra 11412 (USM); Huacapistana,1800-2400 m, 5-8 Jun 192.9, Killip & Smith 24111, 24327 (F, US). JUNIN: Rd. from Tarma to San Ram6n, 11?10'-11?40'S, 75'20'-75045'W, 21 May 1981, Sullivan et al. 1096 (MO, NY); La Merced, 2000 ft, 10-24 Aug 1923, Mathews 5443 (BM, F): Chanchomayo valley, 1500 m, Sep 1924-7, Schunke 32?1 (F); 15 km E of Carpapataon rd. to San Ram6n, 29 Sep 1984, Whalen842 (BH). LORETO:Ripas Huallagae, 1830, Dias 2093 (F); Varaderode Maranfrom Rio Amazonas to Rio Napo, 21 Aug 1972, Croat 19372 (F); Alto Amazonas, Chambira near Yurimaguas, 170-200 m, 14 Sep 1948, Ferr^eyra4943 (MOL, US, USM); Yanamono, Exploramna tourist camp, Rio Amazonas above mouth of Rio Napo. 120 m, 23 Mar 1982, Gentry et al. 36627 (MO), 3?38'S, 72'50'W, 24 Jan 1983, Gentrv et al. 39679 (MO), 27-28 Dec 1982, Gentrv & Emmons 38717 (MO); Yurimaguas, lower Rio Huallaga,ca. 135 m, 23 Aug-7 Sep 1929, Killip) & Smith 27624, 27672 (US); Puerto Arturo, lower Rio Huallaga below Yurimaguas,ca. 135 m, 24-25 Aug 1929, Killip & Smith 27920 (US); between Yurimaguas and Balsapuerto(lower Rio Huallaga), 135-150 m, 26-31 AugJ 1929, Killip & Smith28087 (F, US); Santa Rosa, lower Rio Huallaga below Yurimaguas,135 m, 1-5 Sep 1929, Killip & Smith 28717, 28983 (US), Killip & Smith 28992 (F); San Antonio on Rio Itaya, ca. 110 m, 18 Sep 1929, Killip & Smith 29492 (NY, US); Soledad on Rio Itaya, ca. 110 m, 20-22 Sep 1929, Killip & Smith 29564, 29712, 29779 (F, US); Balsapuerto, ca. 220 m, Feb 1933, Klug 2883 (F, NY, US); Yanamono, Explorama tourist camp on Rio Amazonas between Indiana and mouth of Rio Napo, ca. 80 km NE of Iquitos, 3?28'S, 72?48'W,27 Jul 1984. Kniapp

306 6605 (BH, K, US, USM); Intuto, Rio Tigre, 160 m, 9 Aug 1968, McDaniel 10845a (IBE, NY); above Pongo de Manseriche right bank of Rio Marafion, 200 m, 24 Dec 1931, Mexia 6338 (BH, CAS, F, K, MO, NY, TEX, U, US); Rio Amazonas, Quebrada de Yanayacu, 22 Mar 1977, Rimachi Y 2900 (F, NY, USM); Yanamono, Explorama Lodge, 50 mi NE of Iquitos, ca. 106 m, 3?30'S, 72'50'W, 25 Nov 1981, Vasquez & Jaramillo 2714 (MO, NY); Paranapura,Yurimaguas,lower Rio Huallaga, 155-210 m, Oct-Nov 1929, Williams4596 (F); SantaRosa, Yurimaguas, lower Rio Huallaga, 155-210 m, 8 Nov 1929, Williams 4764 (F), 11 Nov 1929, Williams 4932 (F); Yurimaguas, Puerto Arturo, lower Rio Huallaga, 155-210 m, 14 Nov 1929, Williams4983 (F), 19 Nov 1929, Williams5246 (F); Alto Amazonas, along Rio Marafionnear Teniente Pinglo, just above Pongo de Manseriche, 250-300 m, Wurdack 2126 (K, NY). MADRE DE Dios: Cocha Cashu station,350 m, I Oct 1980, Foster 5441 (F); Parque Nacional Manu, Rio Manu. Cocha Cashu station,400 m, 15 Feb 1977, Foster & Terborgh6085 (F, USM); Parque Nacional Manu, Cocha Cashu station, Rio Manu, 350 m, 2 Feb 1981, Foste;' & Wright 8257 (F); Parque Nacional Manu, Rio Cumerjali, 11?49'S, 71?32'W, 23 Oct 1986, Foster & dAchille 11997 (F, USM); along Rio Shintuya near Shintuya, S bank of Rio Alto Madre de Dios, ca. 500 m, 12?40'S, 71?15'W, 12 May 1984, Knapp & Mallet 6430 (BH, CUZ, K, US, USM); Lucre-Paucartambo-Shintuya rd., ca. 1 hr. drive W of Shintuya,ca. 600 m, 12 May 1984, Knapp & Mallet 6434 (BH, CUZ, US, USM); Aguas Calientes, across and downriverfrom Shintuyaon Rio Alto Madrede Dios, 400-500 m, 13 May 1984, Knapp & Mallet 6442 (BH, CUZ, K, NY, US, USM). PASCO:Chontabamba valley, 2--8 km W of Oxapampa, 1800-1850 m, 10?31'S, 75?30'W, 2 Feb 1983, Gentry et al. 39917 (MO); Oxapampa, 1720 m, 19 Sep 1948, Infantes 1479 (USM); pasturesin vic. of Oxapampa,ca. 2 km E of town, ca. 1850 m, 10(35'S, 75?35'W, 10 Mar 1984, Knapp et al. 6316 (BH, K, US, USM); environs of Villa Rica, ca. 1450 m, 21 Mar 1984, Knapp & Mallet 6341 (BH, K, US, USM); Enefias-Alto Yurinaki-La Florida rd., ca. 9 km E of Villa Rica, 1250-1400 m, 10?50'S, 75?15'W, 12 Aug 1984, Knapp & Mallet 6625 (BH, K, USM); vic. of Huancabamba, Hacienda Yanachaga, 1850 m, 10?22'S, 75?32'W, 24 May 1982, D. N. Smith et al. 1676 (F, MO, NY); valley of Rio San Alberto, E of Oxapampa,2200 m, 10-34'S, 75?22'W,4 Jul 1984, D. N. Smith & Pretel 7573 (F, MO, NY). SAN MARTiN: Tarapoto, 19 Mar 1970, Fernando T7et al. 2441 (US); valley of Rio Huallaga, 29 km S of Tarapoto near El Abra, 450-540 m, 6?40'S, 76?20'W, 5 Feb 1984, Gentry & Smith 44989 (MO, NY); Venceremos, near Amazonas border, km 291 RiojaPomacocha. 1850 m, 5?45'S, 77?40'W,9 Feb 1984, Gentry & Smith 45265 (MO, NY); Zepelacio near Moyobamba, 1200--1600 m, Jan 1934, Klug 3491 (F, K); ca. S km N of Tarapotoalong Rio Shilcayo, ca. 400 m, 6?30'S, 76?22'W, 7 Jun 1984, Knapp & Mallet 6483 (BH, K, NY, US, USM); Tarapoto,Rio Cumbasa,ca. 350 m, 6?31'S, 76?22'W,8 Jun 1984. Knapp & Mallet 6487 (BH, K, NY, US, USM); km 36-41 Tarapoto-Yurimaguas rd., Rio Cachiyacu & Rio Tiyacu, 400-600 m, 6?25'S, 76?15'W, 11 Jun 1984, Knapp

FLORA NEOTROPICA & Mallet 6497 (BH, K, NY, US, USM); Naranjal, trail to Jorge Chavez, km 80 Tarapoto-Yurimaguas rd., ca. 210 m, 6?15'S, 76?17'W, 15 Jun 1984, Knapp & Mallet 6514 (BH, K, US, USM); Convento, trail to Tioyacu and Nuevo Lamas, km 68 Tarapoto-Yurimaguas rd., ca. 270 m, 6?16'S, 76?17'W, 16 Jun 1984, Knapp & Mallet 6516 (BH, US, USM); 6520 (BH, K, NY, US, USM); ca. km 62 TarapotoYurimaguasrd., along Rio Yuracyacu,ca. 260 m, 6018'S, 76?18'W,23 Jun 1984, Knapp & Mallet 6539 (BH, K, NY, US, USM); hills above Chazuta, on Rio Huallaga, 200400 m, 6?36'S, 76?ll'W, 30 Jun 1984, Knapp & Mallet 6545 (BH, K, NY, US, USM); 3-7 km W of Bueno Aires on rd. to Santo Tomasand San Antonio de Paujilzapa,along QuebradaPaujilzapa,200-300 m, 6?47'S, 76?27'W, 13 Apr 1986, Knapp et al. 7041 (MO, USM); hills above Chazuta, ridge to W of Quebrada Chazuta, 200-300 m, 6?34'S, 76?12'W,21 Sep 1986, Knapp8353 (MO, USM); Tarapoto, 1835, Matthews 1512 (K); Tocache Nuevo, Isla de Caserio de Cedro,rightbank of Rio Huallaga,18 Oct 1970, Schunke V 4514 (F, NY); near Tarapoto, Ule 6632 (frag. F); Rio Abiseo National Park, near Gran Pajaten ruins, 2600 m, ca. 70S, 770W, 23 Jul 1985, Young1280 (HUT,NY); Tarapoto, 750 m, 4 Dec 1929, Williams 5505 (F); Huingillo, 500 m, Woytkowski 7188 (US). UCAYALI: Miraflores, 30 km above Pucallpa on Rio Ucayali, 180 m, 8?26'S, 74?05'W,3 Nov 1947, Fosberg 28975 (MO); along Ucayali, ca. 100 S, 1923, Tessmann 3494 (G); Leoncio Prado, Yarinacocha, 200 m, 8?20'S, 74?35'W, 14 May 1984, Visquez 4977 (MO, NY). BOLIVIA. COCHABAMBA:Between Villa Tunariand Puerto Villarroel near Chimore, 380 m, 30 Dec 1982, Ferndndez Casas 7965 (NY). LA PAZ: Vic. of Rurrenabeque, 1000 ft, 26 Nov 1921, Cardenas 1198 (NY). PANDO: Prov. Nicolas Suarez, ca. 20 km from Cobija toward CastroErifia,9 Jan 1983, FernandezCasas & Susanna 8109 (NY); Prov. Manuripi, Conquista on Rio Madre de Dios, 150 m, 3 Feb 1983, Fernandez Casas 8595 (NY). SANTA CRUZ:Espejillos, 20 km SW of Santa Cruz, 550 m, 17059'S,63021'W,16 Nov 1993,Billiet & Jadin 6179 (MO); along rd. from Santa Cruz to Samaipata,3 km SW of Angostura, gorge of Rio Pirai, 700 m, ca. 18?10'S, 63?32'W, 25 Jan 1987, Nee 33792 (NY); Prov. Ichilo, between Rio Chonta (Rio Ancho) and Rio Saguayo, Parque Nacional Ambor6, 425 m, 17?39'S, 63042'W, 22 Jan 1988, N\ee & Saldias P. 36043 (NY); Prov. Florida, 5 km by air SE of Mairana on rd. to Samaipata, at Quebrada Seca, 1550 mn, 18?09'S, 63056'W,4 Feb 1988, Nee 36153 (K, NY); valley of Rio Bermejo, near Bermejo, 700 m, 18?10'S, 63?32'W, 8 Dec 1988, Nee 37057 (NY); Parque Nacional Amboro, along Rio Saguayo 1 km NE of entrance to first Andean foothills, 400 m, 17?39'S, 63?43'W, 19 Dec 1988, Nee 37259 (NY); Avenida Kennedy, ca. 0.5 km E of Rio Pirai, W side of Santa Cruz, 420 m, 17?47'WS,63013'W, 27 Jan 1989, Nee 37747 (NY); Prov. del Sara, 100-1500 m, 18 Oct 1916, Steinbach 3091 (NY). BRAZIL. ACRE: Near mouth of Rio Macuahan, tributaryof Rio Yaco, 9?20'S, 69? W, 8 Aug 1933, Krukoff 5357 (F, K, U, US); near mouth of Rio Macuahan, tributary of Rio Yaco, 9?20'S, 69?W, 5 Sep 1933, Keukoff5797 (K, NY); vic. of Porangaba, Rio Jurua-Mirim, 17 May

TAXONOMIC TREATMENT

et al. P13083 (NY); vic. of km 7 Sena 1971, MWaas Madureira-RioBrancord., 28 Sep 1968, Prance et al. 7647(NY);SanFrancisco,Rio Acre,May 1911,Ule 9720 (US). AMAZONAS: Rio Solim6es between Manausand Boca do Paranado Ariabu,23 May 1941, Manacapuru, Duckes.n. (RB); along Rio Solim6es,Codajaz,19 May 1933, Krukoff4511 (G, K, NY); Ilha Aramaqi,almost oppositeTabatinga,Rio SolimoesandRio Javari,24 Jul 1973, Pranceet al. 16799 (NY); Rio Solim6es,Rio Ica, LagoCarucaca,IgarapePauleta,23 Feb 1977,Pranceet atl.24552 (NY, U); confluence of Rio Negro and Rio Solimoes,May 1851,Spruce1541 (BM,G, K); Sao Francisco, Jun 1911, Ule 9760 (K). Local names. Peru.Amazonas:maikuanim,mun sauk (Aguaruna); San Martin: sacha congompe, sanango. With the exception of Solanumoppositifolium, S. sessile is the most widespread and variable of the species of this group. It is most closely related to S. obovalifoliurn(Cordillerade la Costa, Venezuela)and S. monadelphum (E Andean slope, Peru). Solanum sessile differs from the formerin its much more complexly branchedinflorescences, larger,fleshier flowers, and large, obovate, usually sessile leaves. It differs from S. inonadelphum in its larger leaves, inflorescences, and flowers. Solanum sessile rarely grows in the riverbedhabitatof S. monadelphum.In the forests of Quincemil(Dept. Cuzco, Peru),intermediates between S. sessile and S. monadelphumoccur (see discussion of S. monadelphum).The most distinctive featuresof S. sessile are its large, branchedinflorescences, and large, fleshy flowers. Solanum sessile is a widespread species, with several well-marked geographical races. These are briefly describedhere. 1) PopulationsfromnearMufia,in middle-elevation centralPeru,havestronglysessileleaves,congested inflorescenceswith little or no axis between the flowers,andvery large,fleshy flowers.Ruiz and Pav6n's collection of Solanum sessile is from this area, an isolated Andean valley in the departmentof Huanuco. 2) InareasofcentralPeru(notincludingtheabovearea nearMufia),plantsarequitesimilarin appearance to the preceeding form, but the inflorescences areopenandmany-timesbranched, withsomewhat smallerflowers. Most collections frommiddleto high-elevation Peru are of this form. populationsinPeru, 3) Morenorthern,lower-elevation Ecuador,Colombia, and Venezuela consist of plantswith sessile leaves, the two leaves of the geminatepairmarkedlydifferingfromoneanother, elongate, branchedinflorescences, and pubescentberries.These populationshave been called PheliandraherthaeandSolanumgrandifolium.

3 07

4) Low-elevation Brazilianpopulationswith open branched inflorescences and glabrous berries are of two types: a) those with sessile leaves have been calledSolanummarmellosanum;and b) those with petiolate leaves have been called S. pulchrum. 5) Plants from southeastern Peru and adjacetit Brazil are almost completely glabrous, have sessile, geminate leaves, and glabrousberries. These populations grade into and are perhaps hybridizingwith Solanummonadelphumn. 6) Plants from high-elevation Venezuela have coriaceous,sessileleavesthatareoftensomewhat auriculate,andlarge open inflorescences. Manyotherformsof even morelocal occurrence couldbe described.Togive formalnamesto theelements of this complex mosaic of variationwould be counterproductive,andwould only serveto confuse an already confusingsituation.Theabovewell-markedformsgrade intoone another,andthe collectionsampleis quitesmall for this forest species. Although perhapsdifficult fcor the field botanist,it is more realisticphylogenetically to leavethis complexpatternunderthenameofSolanurn sessile until detailed work is done on the complex. Individuals of Solanum sessile are most likely andromonoecious,as they bear both long- and shorrtstyled flowers. Few berriesare set on any given inflorescence, lending supportto this hypothesis.The syrrtpodialstructureof S. sessile is quitevariable(see above anddescription).Some plantshave difoliate,geminate sympodia, while others appear to have plurifoliate sympodia (for discussion of sympodial types, see Morphology). This difference largely depends on the partof theplanttakento makethe specimenin question. is taken,thesympodia If onlythemostrecentinflorescence appearplurifoliate,andthe inflorescenceterminal,with no shoot growth continuing. On the other hand, if a lower node is taken, the inflorescence appearsto be oppositetheleaf,andthe sympodiumdifoliateandgeminate.Solanumsessile is a good exampleof the difficulties involved in using inflorescenceposition to identify species in sect. Geminata.This single species has beeni placed in two subgeneradue to this variability.

99. Solanum triste Jacq., Enum. Syst. P1. 15. 1760. Type.Martinique.adripasfruticosasfluviiDivaPetri, (lectotype,Jacquin,Select. stirp.amer.hist. fig. 40. Fig. 167 1763, designatedby Knapp, 1991a). Solanzumcataractae Britton in R. E. D. Baker& N. W. Simmonds, Fl. Trin. Tobago 2(4): 267. 1953. Type. Trinidad and Tobago. Trinidad: Maracas waterfall, 10 Apr 1920, Britton et al. 1641 (holotype, herb. Trin.-n.v.; isotypes, NY, US).

308

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167 Solnu

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TAXONOMIC

309

TREATMENT

Shrubs or small trees, 1-7 m tall; young stems and leaves glabrous or minutely papillose, soon glabrous;brancheserect, green; young leaves pubescent along the midribbeneath;barkof older stems grayishbrown. Sympodial units difoliate, geminate. Leaves elliptic to obovate,widest at orjust distalto the middle, glabrousabove, sparselyto densely pubescentbeneath withuniseriatesimpleorbranchedtrichomesin the axils of the main lateral veins or along the midrib, the trichomes golden and usually branched,0.5-1 mm long; major leaves 10-30 x 7-14 cm, with 10-12 pairs of main lateralveins, these dryinggolden brownbeneath, the apex acute,the base acuteto attenuate;petioles 1.52.5 cm long, lightly winged from the decurrentleaf bases;minorleaves differingfromthe majorones only in size, but occasionally in shape, orbicularto elliptic, 3-19 x 2-9 cm, the apex acute,the base acuteto attenuate;petioles 0.4-1 cm long.Inflorescencesoppositethe leaves, 1.2-6 cm long, usually branchedin the distal 1/3, densely puberulentwith erectuniseriatetrichomes 0.1-0.5 mm long, these either simple or branched; pedicel scars evenly spaced ca. 0.5 mm apart.Buds globose when very young, the corolla soon exserted, the budsthenellipsoid, densely puberulent.Pedicels at anthesisdeflexed, white, 1-1.3 cm long, taperingfrom the constrictionat the base of the calyx tube to a slenderbase ca. 0.5 mm diam., sparselyto densely puberulentwith trichomeslike those of the restof the inflorescence.Flowerswiththecalyxtubebroadlycampanulate, 1-1.5 mm long, with a strongconstrictionat the base, this not apparentin drymaterial,the lobes broadlydeltoid with roundedtips, 0.5-1 mm long, with a constriction at the junction of the lobes and the tube, densely golden-puberulent;corolla white, fleshy, 1-1.5 cm diam.,lobed 3/4 of the way to the base, the lobes planar at anthesis, the margins and tips of the lobes densely golden-puberulentlike the rest of the inflorescence; anthers 3.5-4 x 1-1.5 mm, poricidal at the tips, the pores teardropshaped; free portion of the filaments minute, absent to 0.2 mm long, the filament tube 0.50.7 mm long; ovary densely golden-pubescent,the trichomes uniseriate, simple or branched, 0.2-0.6 mm long;style straight,5-6 mm long, pubescentat thebase; stigma capitate,the surfaceminutelypapillose.Fruit a globose, greenberry,1-1.5 cm diam.;fruitingpedicels woody, erect or slightly deflexed at fruit maturity,11.5 cm long, ca. 1 mm diam. at the base. Seeds dark brownin drymaterial,ovoid-reniform,ca. 2.5 x 2 mm, the marginssomewhatincrassate,the surfacesminutely pitted. Chromosomenumber:n = 12 (voucherKnapp & Mallet 6750).

700

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FIG. 168. Distnibutionof Solanum ses.sile(solid circles), S. triste (open circles), and S. turgidum(solid square).

Specimens examined. S.loc., De Ponthieu s.n. (3M); Jurian herb. 160.999 (P-Juss.). MARTINIQUE. S.loc., Duss 366 (F, MO); s.loc., Dec 1867, Hahn 318 (BM, F, MPU); s.loc., Siebet- 54 (F, G, MO); s.loc., von Rohr 92 (BM). VENEZUELA. ANZOATEGUI: Guanta, Jun 1943, Cardona572 (US, VEN); banks of Rio Neveri, 27 Dec 1941, Tamayo2063 (VEN). BOLIVAR: Caicara, 25 May 1952, Smith234 (US). MONAGAS: Aparicio, banks of Rio Aragua on Maturin-Caripe rd., ca. 250 m, 27 Mar 1970, Benitezde Rojas 851 (MY). SUCRE: Vic. of Las Piedras, ca. 800 m, 10 Mar 1977, Badillo et al. 7313 (MY); Rio Cocollar, I May 1983, Cumana1610 (MY); foothills of Serrania de Turimiquire, SW of Cumanacoa, ca. I10km SW of San Lorenzo along Rio Manzanares, 400-500 m, 10?l5'N, 63?55'W, 14 Oct 1984, Knapp& Mallet 6749, 6750, 6751, 6752, 6753, 6755 (BH, K, MY, US, VEN). TRINIDAD AND TOBAGO. TRINIDAD: Maracas Falls, rocky gorge to left of falls, 7 Mar 1993, Johnston 372, 373 (BM, TRIN-n.v.); s.loc., 1826, Sieber 309 (F, MO). Distribution (Fig. 168). In dry forest and secWINDWARD ISLANDS. DOMINICA: Delices, SE ondarygrowthin coastal easternVenezuela,Trinidad, coast, track to Belvedere Estate, 250 m, 26 Sep 1983, Martinique,and Dominica, from sea level to 200 m. 3769 (BM). White~foord

FLORA NEOTROPICA

310

FIG. 169. Solanum turgidum S. Knapp. (Reproduced with permission from Brittonia 38: 289, fig. 11. 1986.)

Local names. Venezuela.Anzoategui:tabaquero. Solanumtristeis relatedto S. turgidumof higher elevations on the Pariapeninsula (Sucre, Venezuela), from which it differs in its white flowers, pubescent leaves, longer fruiting pedicels, and second growth habitat (see Knapp, 199 lb). Solanum triste is a rank second growth weed, and grows in large stands at the edges of pastures.The leaves arequite foetid smelling, andthis characteristicis reflectedin the commonname ofthis plantin Venezuela.NearCumanacoa,Venezuela, the leaves of S. triste are fed upon by the larvaeof the ithomiine butterflyPteronymialatilla (see TableXI). Thesecaterpillarswere quitecommonwhereI collected them, and defoliated plants ofS. triste.

100. Solanum turgidum S. Knapp,Brittonia38: 288. 1986. Type. Venezuela. Sucre:Peninsulade Paria, Cumbrede Estrella,W of Manacal,N of El Paujil, low cloud forest on steep ridges, 800-850 m, 10040'N,6241 'W, 17 Oct 1984, Knapp & Mallet

6765 (holotype, MY; isotypes, BH, K, NY, US, VEN). Fig. 169 Small trees, 3-6 m tall; young stems and leaves minutely golden-puberulentwith erect uniseriate trichomes less than 0.1 mm long; stems of live plants green, not winged; barkof older stems grayish-brown, smooth. Sympodial units difoliate, geminate. Leaves obovateto elliptic, widest at orjust distalto the middle, glabrouson both surfaces,but occasionally with a few minuteuniseriatetrichomesalong the midribnearthe petiole; majorleaves 10-25 x 5-15 cm, with 5-7 pairs of main lateralveins, these prominentandoccasionally yellowish beneath,the apex acute, the base acute, not decurrentonto the petiole;petioles 1-2 cm long; minor leaves differingfromthe majorones in size and shape, elliptic to orbicular,5-9.5 x 3.5-8.5 cm, the apex acute to rounded,the base acute;petioles4-6 mm long.Inflorescences appearingterminalon the terminalnode due to the slow expansion of leaves, but on other nodes opposite the leaves, ratherstout, simple, but occasion-

TAXONOMIC TREATMENT

ally once-furcate, 1.5-4 cm long, 5-30-flowered, minutelygolden-puberulent likethestemsandyoungleaves, the trichomesdensernearthe distal end;pedicel scars closely spaced, but not markedlyoverlapping,beginning very nearthe base of the inflorescence.Buds globose when young, laterellipsoid with the exsertion of the corolla, the corolla stronglyexserted, 3-4 times as long as the calyx tube.Pedicels at anthesis erect, pale green,5-7 mm long, taperingfromthe base of the calyx tubeto aslenderbaseca. 0.5 mm diam.Flowerswiththe calyxtubebroadlycup-shaped,appearingwoody in dry specimens, 1-1.5 mm long, the lobes broadlydeltoid, 0.3-1 mm long, edged white, the tips minutely papillose, the entire calyx puberulentwith the same erect golden uniseriatetrichomesas the rest of the inflorescence;corolla white with a greencenter,each lobe with a greenabaxialstripeanda single medialadaxialridge, the ridge apparenteven in dry specimens, 1-1.2 cm diam.,lobednearlyto thebase,the lobesplanarat anthesis, the tips of the lobes slightly cucullate,the tips and marginspuberulentwithminuteuniseriatetrichomeslike those of the buds; anthers 2.5-3.5 x 0.75-1 mm, poricidalat the tips, the terminalportionthickenedand paler,theporesteardropshaepd;freeportionof thefilameilts0.25-0.5 mm long, the filamenttubeca. 0.25 mm long; ovaIrydensely golden-pubescentwith uniseriate trichomes0.14.3 mm long;stylestraight,5-6 mm long; stigma minutely capitate, the surface minutely papillose. Fruiita globose berry,tingedpurplishwhenyoung, green when ripe, 1-1.2 cm diam., the surfacedull due to the continuedpresence of minuteuniseriategolden trichomes,these not deciduous,0.1-0.3 mm long;fruitingpedicels swollen, woody, and erect, 8-9 mm long, 1-2 mm diam.atthebase.Seeds(immature),darkbrown in dry specimens, ovoid-reniform,2-2.5 x 1-1.5 mm, the surfaces minutely pitted. Chromosomenumber.n 12 (voucher Knapp& Mallet 6765).

311

chlamydogynum,both also of coastal Venezuela (see Knapp, 1991b). They share several homoplastic synapomorphies:branchedpubescence, thickenedcalyx lobes, and incrassate seed margins. Solanurnm turgidumis easily distinguished from S. triste or S. chlamydogynum by its greenish flowers, glabrous leaves, andshort,stoutfruitingpedicels. The trichomes on the style base of S. turgidumappearuniseriateand simple under the dissecting microscope, but when viewed at high magnificationswith the SEM, it is clear thatat least some of themarefurcate(see Fig. I 1).This possession of both branchedand simple trichomes is common in the S. sessile species group. Solanum turgidumgrows in low cloud forest, underclosed canopy.Theplantshave a clumpecldistribution,and apparentlyare growing in old light gaps in the forest. Solanum turgidumis common where it ciccurs, but is uncommonon a regional scale.

XV. Solanum confine species group (S. capillipes, S. confine, S. cruciferum,S. darienense, S. dissirnile, S. erosomarginatum,S. hypocalycosarcum,S. leptopodum,S. leptorhachis,S. morii,S. nematorhachis, S. ombrophilum, S. pastillum, S. per-tenlue,S. stipulatum,S. tenuiflagellatum,S. tuerckhelnii, S. valerianum,S. yanamonense). Figs. 170, 171

Delicate shrutbsof archinghabit, usually growilng in the primary forest understory; young stems anid leaves variously golden pubescent or glabrous.Svmiipodial unitsunifoliateor difoliate,geminate,sometimes anisophyllous.Leaves elliptic, the marginsusually undulate,giving the leaves on live plantsa ruffledappearance. Inflorescencesopposite the leaves, simple, delicate andslender,sessile to extremelyelongate;pedicel scars usually widely andunevenly spaced.Buds ellipsoid to globose. Pedicels at anthesis deflexed. Floviers white to greenish-white, usually quite small, thie Distribution (Fig. 168). Known only from the lobes deflexed at anthesis.Fruitusuallyhardandgreen ridges of the Peninsulade Pariain easternVenezuela, at maturity;fruitingpedicels usually much longer and in cloud forest at ca. 800 m. thicker,especiallyattheapex,thanthefloweringpedicels, Specimens examined. VENEZUELA. SUCRE: deflexedand hangingbetweenthe leaves.Seeds ovoi(lPeninsula de Paria, Cumbre la Estrella, W of Manacal, N reniform,tanto darkbrown,the surfacesminutelypitted. of El

Paujil, 800-850 m, 10?40'N, 62?41'W, 17 Oct 1984, Knapp & Alallet 6763, 6767 (BH, MY, NYUS, VEN); Cerro de Rio Arriba, W slopes of Cerro Humo along Rio Santa Isabel, above Santa Isabel, 600-700 m, 9 Aug 1966, Stelvermark& Rabe 96175 (NY, US, VEN); headwater tributaries of Rio Cumana, below trail between El Paujil and El Brasil, 850-900 m, 10?39-40'N, 62?43'W, 21 Feb 1980, Stever-market al. 121482 (VEN), Steyerinarket al. 121522 (VEN); between Tacarigua and headwaters of Rio TacariguaE of Cerro Humo, N of Rio GrandeArriba, 560 m, 10(41-42'N, 62?36-37'W, 23 Feb 1980, .Stevernmark et al. 121616 (VEN). Solanium turgidum is related to S. triste and S.

Distribution. CentralAmerica,the Andes, Amnazon basin and SE Brazil, primaryforests. Steyermark 90111 (MO, VEN) from Parque Nacional Guatopo in coastal Venezuela is a potential new species in this group, but the material is scanty enough to put off describing it until better specimens are available.It differs from otherspecies in the group in its apparentlyorbicularminorleaves and relatively short fruitingpedicels. It most closely ressembles S. dissimileandthese specimensmay representa disjunct and somewhataberrantpopulationof that species.

312

FLORA NEOTROPICA

A

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FIG. 170. Solanum confine species group. A. S. confine branch, Peru (Knapp & Mallet 6589). B. S. confine flowers, Peru (Knapp & Mallet 6649). C. S. dissimile flowers, Venezuela (Knapp & Mallet 6825). D. S. hypocalycosarcum flowers, Ecuador (Knapp & Mallet 6765). E. S. leptorhachis inflorescence, Ecuador (Knapp & Mallet 6160). F. S. leptorhachis flowers, Ecuador (Knapp & Mallet 6160).

TAXONOMIC TREATMENT

E

3 13

FR

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~~~~~~~~~~~~~~~~AmB

FIG. 171 Solanum confine species group A S leptopodum branch Peru (Knapp & Mallet 6608). B. S. leptopodum flowers, Peru (Knapp & Mallet 6608). C. S. pastillum flowers, Costa Rica (Knapp 840). D. S. tenuifiagellatum fruit, Venezuela (Knapp & Mallet 6859). E. S. stipulatum plant, Brazil (Bovini et al. 814). F. S. stipulatum fruit Brazil (Bovini et al 814).

3 14

FLORA NEOTROPICA

Key to the speciesof the Solanumconfinespeciesgroup 1. Sympodial units always difoliate; leaves geminate on reproductive shoots. 2. Nodes crowded; bark white and peeling; stems strongly winged; major leaves obovate to linear; 115. S. stipulatlrn berry slightly to markedly turbinate. SE Brazil....................................................... 2. Nodes spaced; bark brown or gray; stems not markedly winged; major leaves not as above; berry globose. 3. Plants densely pubescent; leaf margins erose or entire and undulate. 4. Leaf margins erose; minor leaves orbicular. Andean Venezuela...........106. S. erosomarginatum 4. Leaf margins entire; minor leaves not orbicular, similar to majors in shape. 5. Buds globose; calyx lobes with a globose apical projection. French Guiana......... 110. S. morli 5. Buds conspicuously pointed; calyx lobes with an elongate apical projection. N Peru 103. S.cr. cifeuerum ..........................................P. 3. Plants glabrous or only minutely puberulent; leaf margins undulate or entire, never erose. 6. Minor leaves minute and stipule-like, usually less than 1 cm long. E Peru and Amazonian Brazil ............................................ 108. S. leptopodlun 6. Minor leaves larger, usually more than 1 cm long, not minute and stipule-like. 7. Pedicel scars closely and evenly spaced; pedicels purplish; inflorescence congested: stems minutely puberulent. W Ecuador.........................................107. S. kvpocalycosarctrm 7. Pedicel scars widely and unevenly spaced; pedicels green; inflorescence elongate; stems glabrous. 8. Calyx lobes knob-like swellings on the rim of the calyx tube; minor leaves orbicular to elliptic; leaves not drying golden beneath; bark pale green. Montane Costa Rica ...................................... 113. S. pastilll/mi 8. Calyx lobes deltoid, not as above; minor leaves not differing from the major ones in shape; leaves drying golden beneath; bark dark red, exfoliating. Mexico to Costa Rica ........................................................... 117. S. tUe1ckheiinlii 1. Sympodial units always unifoliate on reproductive shoots. 9. Pedicel scars closely spaced; inflorescence not elongate, rather stout. Montane coastal Venezuela .................................................. 112. S. ombr-ophilium 9. Pedicel scars widely and irregularly spaced; inflorescences and pedicels filiform. 10. Inflorescences long and slender, to 30 cm long; petals planar to somewhat reflexed at anthesis. 11. Plants sparsely to densely pubescent along the veins of the leaf undersides; calyx lobes deltate to long-triangular. 12. Calyx lobes long-triangular, reflexed at anthesis; fruiting pedicels 3-3.5 cm long. Cordillera de la Costa, Venezuela.......................................... 116. S. tenuiflagellatum 12. Calyx lobes deltate, not reflexed at anthesis; fruiting pedicels 1.5-2 cm long. Choc6, Colombia ..........................................l 1. S. niematorhachis 11. Plants glabrous or minutely puberulent; calyx lobes rounded, minute. W Ecuador ............................................ 109. S. leptorhachis 10. Inflorescences not extremely elongate, usually less than 2 cm long; petals reflexed (usually strongly) at anthesis. 13. Flowers minute, 4-5 mm diam.; greenish-white; extreme leaf tips blunt; venation arching, converging near midrib. Mouth of Rio Napo, Peru...................................... 119. S.' ananionense 13. Flowers larger, 0.8-1 cm diam.; white; leaf apices and venation not as above. 14. Stems with 1-1.5 mm long, straw-colored trichomes, not usually with shorter pubescence. Trinidad and Central America. 15. Leaves firm and shiny above; main lateral veins joined in a series of prominent arches; trichomes sparse on veins beneath. Trinidad and Paria peninsula,

Venezuela.......................................

101. S. capillipes

15. Leaves membranous; the venation not as above; trichomes long and dense oni the veins beneath. SW Costa Rica and adjacent Panama............................. 118. S. v7alerianirni 14. Stems with shorter pubescence, or minutely puberulent, if longer very dense and dark, usually puberulent along the veins. 16. Inflorescences 1-3-flowered; less than I cm long, never longer; stems and veins of the leaf undersidesminutely puberulent.Montane Costa Rica and Panama......... 114. S. per-tenue 16. Inflorescences to 15-flowered; longer, to 2.5 cm long; pubescence various. 17. Pedicels in fruit only 1-1.5 times their length in flower; calyx lobes minute; leaves 5-O0 x 1 .5-3 cm. Andean Venezuela and adjacent Colombia. 105. S. dissi-nile

TAXONOMIC TREATMENT

3 15

17. Pedicels in fruit 2-3 times their length in flower; calyx lobes and leaves larger. 18. Leaf bases obtuse, slightly oblique; leaf undersides glabrous; inflorescence 104. S. darienense 4-5-flowered. Darien, Panama...................... 18. Leaf bases acute, not oblique; leaf undersides puberulent to pubescent on 102. S. coqifine veins; inflorescence 4-15-flowered. E Peru .

101. Solanum capillipes Britton, Bull. Torrey Bot. Club 48: 338. 1922 [1921]. Type. Trinidad and Tobago. Trinidad: Ortoire River, Guayaguayare Rd., Britton et al. 2521 (holotype, NY; isotypes, K, US). Fig. 172 Shrubsto 1.5 m tall;young stemspubescentwith uniseriatecurly-tippedtrichomesca. 0.5 mm long, not winged from the leaf bases; older stems glabrate,the barkgrayish-green.Sympodialunitsunifoliate.Leaves elliptic, widest at or just below the middle, glabrous and shining above, glabrous or pubescent along the veins beneath, 9.2-18 x 3-6.5 cm, with 7-9 pairs of main lateralveins, these raisedandjoined in a series of prominent arches above, prominent and reddish beneath,the apex acute,the base acute,slightly decurrent on the petiole; petiole 4-8 mm long. Inflorescences opposite the leaves, simple, 0.8-3 cm long, 5-8-flowered,sparselypubescentwith uniseriatetrichomeslike those of the stems;pedicel scars unevenly spaced0.52 mm apart,raised from the inflorescence axis. Buds globose when young, laterellipsoid with the exsertion of the corolla. Pedicels at anthesis deflexed, filiform, 0.8-1 cm long, taperingfrom the calyx tube to a slender base ca. 0.25 mm diam., sparsely pubescent with uniseriate trichomes. Flowers with the calyx tube broadlyconical, 0.5-1 mm long, the lobes deltoid,hyaline, 0.25-0.5 mm long, sparsely pubescent with uniseriatetrichomeslike those of the rest of the inflorescence, minutely papillose on the tips of the lobes; corolla white, 6-8 mm diam., lobed ca. 3/4 of the way to the base, the lobes reflexed at anthesis, the tips and narginsof the lobes minutelypapillose;anthers 1.5-2 0.75--1 mm, poricidal at the tips, the terminal0.05 mnm palerandthickened,theporesteardropshaped;free portionof thefilaments0.25-0.3 mm long, the filament tube 0.25-0.3 mm long; ovary glabrous;style straight, 3.5-5 mm long: stigma clavate and bilobed, minutely papillose.Fruit a globose, greenberry,0.6-1 cm diam. (immature);fruitingpedicelsdeflexed, woody, 2-2.5 cm long, 0.5-0.75 mm diam. at the base. Seeds not known. Chromosomenumbernot known. Distribution (Fig. 174). In forests of southem Trinidadandthe Peninsulade Pariaof Venezuela,from sea level to 800 m. Specimens examined. VENEZUELA. SUCRE: Peninsula de Paria, Cumbre de las Estrellas W of Manacal, ca. 15 air km NW of Irapa E of El Sanche, N

of El Paujil, 800-830 m, 30 Nov 1979, Steyer-mark & Liesner 120749 (VEN). TRINIDAD AND TOBAGO. TRINIDAD: Central Range Reserve, 1 Aug 1973, Adams 13387 (BM);

Moruga,La FortunaTrace,14 Feb 1916,Broadway-v 7909 (NY); EdwardSt., Moruga,Clubbes.n. (BM). Solanum capillipes is extremely similar to and probablyclosely related to S. valerianumof lowland westernCostaRica andPanama,sharingwith thatspecies long, curly-tippedtrichomes.Solanumcapillipes differs fromS. valerianumin its somewhat less membranous, shiny leaves, longer inflorescences, shorter stem pubescence, and largerflowers.

102. Solanum confine Dunal in DC., Prodr. 13(1): 137. 1852. Type. Peru. "Perou", Poeppig 2279

(holotype, G-DC [F neg. 6782]; isotype, F). Figs. 170A,B. 173 Shrubsor small trees 1.5-5 m tall; young stems sparselyto denselypuberulentwithminuteuniseriateand unicellulargolden trichomes0.05-0.25 mm long; bark ofthe olderstemsgray-brown,sparselypuberulent; stems lightly winged fromthe decurrentleaf bases;branches spreadinghorizontally,giving the shruba flattenedaspect. Sympodialunitsunifoliate, except on non-reproductivebranches.Leaveselliptic,geminateonly on nonreproductivenodes, widest at orjust below the middle, glabrous above, glabrous or puberulenton the veins beneathwith the same minuteuniseriateand unicellular trichomes as those on the young stems, these occasionally sparse;majorleaves 8-15 x 3.5-6 cm, withl 9-1 1 mainlateralveins,impressedabove,prominentand yellowish beneath,the veins convergingnearthe margins of the leaf, the apex acute, the base acute, slightly decurrentonto the petiole; petioles 0.6-1 cm long: minor leaves differing from the majorsonly in size, 1.6-6 x 0.7-2.5 cm, the apex acute,the base acute;petioles 2-3 mm long.Inflorescencesoppositetheleaves,simple, 0.5-2.5 cm long, 4-15-flowered, minutelypuberulent with uniseriateandunicellulartrichomeslike those of the stems and leaves;pedicel scars irregularlyspaced 1-1.5 mm apart,beginning ca. 3 mm from the base of the inflorescence.Buds obovoid, flattenedapically.the corollaearlyexsertedfromthe calyx, when very young the buds globose and minutelypapillose or with a few shortunisefiatetrichomes.Pedicels at anthesisdeflexecl.

FLORA NEOTROPICA

316

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TAXONOMIC TREATMENT

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1984

FLORA NEOTROPICA

318

7-9 mm long, filiform, taperingfromthe calyx tube to a threadlikebase ca. 0.25 mm diam.Flowers with the calyx tube broadly conical, 1-1.5 mm long, the lobes deltoid, 0.5-1 mm long, minutelypapillose on the tips or with a few uniseriatetrichomeslike those of the rest of the inflorescence; corolla white, 1-1.4 cm across, lobed nearly to the base, the lobes planar or slightly reflexedat anthesis,minutelypapilloseon thetips of the lobes; anthers3-3.5 x 1-1.5 mm, poricidalat the tips, the pores teardropshaped;free portionof thefilaments 0.25-0.5 mm long, the filamenttube0.25-0.5 mm long; ovaryglabrous;style straight6-8 mm long;stigmacapitate, the surface minutely papillose. Fruit a globose, green berry, 1.2-1.5 cm diam.;fruiting pedicels deflexed,woody,widenedat the apex, 0.5-0.75 mm diam. at the base. Seeds darkbrown, ovoid-reniform,2.5-3 x 1.5-2 mm, the surfaces deeply pitted. Chromosome number. n = 12 (voucher Knapp & Mallet 6649).

.0

Distribution (Fig. 174). On the easternslope of the Andes in Peruand N Bolivia, from 200 to 2200 m. Specimens examined. ECUADOR. NAPO:Proyecto Hidroelctrico Coca, R bank of Rio Quijos, ca. 10 km S of Reventador, 1500 m, 011'S, 77?39'W, 3-5 Oct 1990, Palacios 5942 (QCNE). PASTAZA: Mera, ca. 1100 m, 27-28 Mar 1968, Harling et al. 7870 (GB, MO); 1020 km N of Canelos, 12 Nov 1974, LugoS. 4574 (MO); Parayacu, ca. 10 km E of Canelas, 8 Nov 1974, LlugoS. 4520 (GB, MO). PERU. Peruvia subandina, Poeppig s.n. (P); San Jacinto, ca. 1878, Martinet 1257 (P). HUANuco: Near Puerto rd. to Monz6n, 650-700 m, 11 Jun 1958, Fer-reyra 13201 (USM); vic. of Tingo Maria, 24 Jun 1961, Mathias & Taylor 5400 (F); Pachitea, Bosque Nacional de Iparia, along Rio Pachitea I km from Tournavista, 20 km from confluence with Rio Ucayali, 300-400 m, SchuinkeV 1556 (F, US); Leoncio Prado,W of Tingo Maria, 680-700 m, 8 Nov 1971, SchunkeV 5127 (F, K, NY, US). LORETO: Left bank of Rio Chanuzi at jct. with Rio Huallaga, 170-200 m, 17 Sep 1948, Ferlevra 5047 (MOL, US, USM); Yurimaguas, lower Rio Huallaga, 135 m, 23 Aug-7 Sep 1929, Killip & Smith 27610 (F, NY, US); Puerto Arturo, lower Rio Huallaga,below Yurimaguas,135 m, 24-25 Aug 1929, Killip & Smith 27861 (F, NY, US). PASCO:Km 1518 on Villa Rica-Pto. Bermudez rd., ca. 1600 m, 21 Mar 1984, Knapp et al. 6335 (BH, K, US, USM); Oxapampa, km 15 Repartici6n-Iscozacin(km 73 Villa Rica-IscozacinPto. Mairo), along Rio Palcazu, ca. 380 m, 10?21'S, 75?10'W, 17-18 Aug 1984, Knapp & Mallet 6648, 6649 (BH, K, MEXU, US, USM). SANMARTiN:Rio Intiyacu,S of Bellavista, ca. 250-260 m, 29 Jul 1970, Chrostowski 70-399 (AAU); between Juanjui and Tingo de Saposoa, 200-300 m, 5 Sep 1948, Ferreyra 4794 (MOL, US, USM); Bellavista, 200-300 m, 7 Sep 1948, Ferreyr-a4800 (MOL); Rioja, km 398 of CarreteraMarginalbetween Pomacochas and Rioja, trail to Quebrada Venceremos and Rio Serranoyacu,1300-1400 m, 5?45'S, 77?30'W, 10 Jul 1984, Kniapp& Mallet 6587, 6589 (BH, K, MEXU, US, USM); Tarapoto, 1835, Mathews 1508 (BM, K, P); Tarapoto,

FIG. 174. Distribution of Solanum capillipes (solid and S. c-onfine

squares), S. dat-ienenise (open circles), (solid circles).

1855-6, Spruce 4161 (K, NY, P); Tarapoto, Pongo de Shilcayo, Oct 1902, Ule 6482 (K); Tarapoto, 750 m, 4 Dec 1929, Williams 5527 (F); Tarapoto, upper Rio Huallaga, 360-900 m, 13 Dec 1929, WilliamKs6179 (F); Juan Guerra, near Tarapoto, 720 m, 30 Dec 1929, Williams 6921 (F). BOLIVIA. LA PAZ: Prov. Murillo, 30.5 km N of (below) dam at Lago Zongo, trail up Rio Jachcha Cruz, 2200m, 16?07'S, 68?05'W, 30 Nov 1982, Solomon 9()52 (MO0, N Y).

Local names. Peru. San Martin: chirisanago chico, chirapa sacha. Solanum confine is most closely related to S. pertenue of montane Central America, and to S. yanamonenise of lower-elevation eastern Peru. It differs from those species in its taller stature, larger flowers, and shorter fruiting pedicels. The foliage of S. conJine is foetid when the plants are growing in sunny areas. The corolla lobes are not strongly reflexed at anthesis. This species is one of the few in this species group to grow in rank secondary growth as well as in primary forest. Solanum confine is a variable species, with more pubescent individuals occurring at higher elevations. Solomon 9()52 was said to be growing as an epiphyte in moist forest. The type of S. confine at G-DC does not bear any specific locality information, but an isotype found at F bears the locality information "Maynas, 1831I,"and the same collecting number as the holotype. The specimen at F is an exact match for that at G-DC, and I am

TAXONOMIC TREATMENT

3 1S'

sure they came from the same plant. Maynas, now a province of the departmentof Loreto, Peru, was the general name for the eastern slopes of the Andes in northernPeru, so the type could have been collected anywhere in thatregion.

Specimens examined. ECUADOR. LOJA:Celica-Zapotillo rd., km 6-10, 1800-1900 m, 10 Apr 1980, Harling & Andersson 18128 (GB); Cerro Sozoranga, Colaisaca-Utuana, km 0.5, 2430 m, 4?19'S, 79?41'18"W, 28 Apr 1994, Jorgensen et al. 617 (BM, MO, QCNE). PERU. CAJAMARCA: Prov.Contumaza, Bosque Cachil, 2650 m, 14 May 1994, Sagastegui A. et al.

103. Solanum cruciferum Bitter,Repert. Spec. Nov. Regni Veg. 16: 402. 1920. Type. Peru. Cajamarca: Chugur,NW of Hualgayoc,2700-2900 m, 22 May 1904, [Weberbauer 4092 (holotype, B [destroyed: F neg. 2603]; lectotype, MOL, here designated; isolectotype,G [Mortonneg. 8579]). Fig. 175

Masamerich between tambos of Calabaza and Atac, 2900

15274 (BM, F, HAO). JUNIN:

Jauja, valley of Ric,

m, 8 May 1913, Weberbauer6678 (F, MOL,US). Solanumcruciferumhas an apparentlyscatterect distributionand is only known from a few collections. The tiny, almost sessile inflorescences are similar to those of S. pertenue of Central America, but S. cruciferum differs from that species in its difoliate sympodia, smallerflowers, andmuch shorterand less copious pubescence. The bud shape of S. cruci/tirum. ellipsoid with an elongate, nipple-like projection, is distinctivein the S. confinespecies group.Bitter'sepi-. thet refers to the apparently4-lobed corollas of this species he saw on the type specimen. That sheet is unfortunatelydestroyed, but the specimen I ha-vese-lected as the lectotypehad a few 4-lobed andmostly 5-lobed flowers. This aberrationis occasionally found in other solanums as well.

Shr-iubsto 1 m tall; young stems and leaves denselypubescentwith golden,erect,uniseriate,simple trichomes0.1-0.2 mm long, these not soon deciduous; bark of older stems pale grayish-white. Synmpodia difoliate, geminate.Leaves elliptic, dryinggolden beneath, glabrous adaxially, pubescent along the veins with simpleuniseriatetrichomeslike those of the stems; majorleaves 2-9) x 0.8-3.5 cm, with 7-9 pairsof main lateral veins, these joined into a series of prominent archesca. 2 mm fromthe leaf margin,the apex acuteto acuminate, the base acute; petioles 0.4-1.5 cm long; minorleaves differingfromthe majorsonly in size, 15 x 0.5-3 cm, the apex acute to acuminate,the base acute;petioles 0.3-1 cm long. Inflorescencesopposite 104. Solanum darienense S. Knapp, Ann. Missouri Bot.Gard.73:738. 1986(1987).Type.Panama.Darien: the leaves or offset slightly, 1-2 mm long, simple, 2-3CananearRio Setigandi,540-580 m, 18 Apr 1980, flowered, pubescentwith golden erect trichomes like Gentrvet al. 28541 (holotype, MO). Fig. 176 those of the stems; pedicel scars closely spaced, not overlapping.Buds ellipsoid, apically pointed, the coShrubswith foetid foliage, 1-1.5 m tall; young rolla long-exserted from the calyx tube. Pedicels at stems sparsely hispidulous with erect uniseriate trianthesis filiform, 1.4-1.6 cm long, deflexed, ca. 1 mm chomes ca. 0.1 mm long, these often only on one side diam.attheapex,ca. 0.25 mmdiam.atthebase,sparsely of the stem;young leaves glabrous;stemswinged frorrm pubescentwith golden uniseriatetrichomeslike those the decurrentleaf bases; barkof older stems reddishof the rest of the inflorescence. Flowers with the calyx golden and shiny. Sympodialunits unifoliate. Leave,s tubebroadlyconical, 0.5-1 mm long, the lobes deltoid, elliptic to ovate, not geminateexcept on non-reproduc1-1.5 mm long, with an elongatedapicalprojectionca. tive nodes,widest at orjust proximalto the middle,gla0.5 mm long, sparselypubescentwith goldentrichomes brouson both surfaces,occasionallyminutelypuberulike those of the pedicels; corolla white, 1-1.5 mm lent along the veins beneath, 1-14 x 3-4.5 cm, with diam., lobed nearly to the base, the lobes strongly re7-8 pairs of main lateralveins, these not prominently flexed at anthesis,the abaxialsurfaceof the lobes denseraisedabove,prominentandyellowishbeneath,theapex ly pubescentwith minutegolden trichomes,the tips of acute to acuminate,the base truncate;petioles wingecl the lobes cucullate and densely papillose; anthers 3from the decurrent leaf bases, ca. 1 mm long. In/flor-es3.5 x 1-1.5 mm, poricidal at the tips, the pores tearcences opposite the leaves, simple, threadlike,0.5-1.5 dropshaped;free portionofthefilaments ca. 0.25 mm cm long, 4-5-flowered, minutely puberulent with long, thejilamentstubeca. 1 mm long, glabrous;ovary erect uniseriate trichomes like those of the stems: glabrous; style 4.5-5 mm long; stigma bilobed, the pedicel scars evenly spaced 1-2 mm apart, slightly surface minutely papillose. Fruit a globose, green (?) raised. Buds globose when young, hispidulous with berry,ca. 1 cm diam. (immature);fruitingpedicelsdeuniseriatetrichomeslike those of the rest of the infloflexed,woo(ly andexpandedat the apex, 1.5-2 cm long, rescence,the corollasoon exsertedfromthe calyx tube ca. 0.5 mm diam. at the base. Seeds not known from makingthe budsellipsoidto obovoid.Pedicels at anthematurefruit. Chromosomenulmbernot known. sis filiform,0.6-1 cm long, taperingfromthe calyx to a. Distribution (Fig. 178). In thick, moist woods slenderbase ca. 0.25 mm in diam., sparselypuberuleni; in S EcuadorandN centralPeru,from2700 to 2900 m. with uniseriatetrichomes.Flowers with the cal x tutbe

320

FLORA NEOTROPICA

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175. Solanum cruciferum Bitter. Ecuador. Jorgensen et al. 617 (BM).

TAXONOMIC TREATMENT

321

FIG. 176. Solanum darienense S. Knapp. (Reproduced with permission from Annals of the Missouri Botanical Garden 73: 739, fig. 1. 1986.)

broadly conical, ca. 0.5 mm long, the lobes deltoid, 0.25-0.5 mm long, the marginspaler,the lobes andtube minutelyhispidulouswith uniseriatetrichomesca. 0.1 mm long; corolla white, 5-7 mm diam., lobed nearly to the base, the lobes reflexed at anthesis, the tips and margins of the lobes minutely papillose; anthers ca. 1.5 x 1 mm, poricidal at the tips, the pores teardrop shaped; free portion of the.filaments 0.25-0.3 mm long, the filament tube ca. 0.1 mm long; ovary glabrous; style straight, 3-3.5 mm long; stigma a slight broadeningat the top of the style, minutely papillose. Fruit a globose, green berry, ca. 1.5 cm diam.;fruitingpedicels deflexed, woody, 1.8-1.9 cm long, expanded at the apex, 0.5-0.75 mm diam. at the base. Seeds darkbrown in dry material,ovoid-reniform,33.5 x 2-2.5 mm, the surfaces minutely pitted. Chromosome number not known. Distribution (Fig. 174). In the low mountains of extremeeastem Panamaand adjacentColombia. Specimens examined. PANAMA.DARIEN: Vic. of airstripat Canagold mine,480 m, 29 Jul 1976,Croat

37963 (MO);vic. of Cana, 1750 ft, 23 Jun 1959,Stern et al. 477, 661 (MO, US). COLOMBIA. ANTIOQUIA:

Mun. Frontino, Nutibara,headwatersof Rio Cueva, 1880 m, 18 Nov 1986, Sdnchezet al. 579 (NY); km 17 of rd. NutibaraLa Blanquita,regionof Murri,1860m, 6045N, 76?24'W, 3 Nov 1988, Zarucchi et al. 7065 (MO).

Solanum darienense is related to S. confine, a species of the Andean foothills in eastem Peru, andto S.pertenueof montaneCostaRicaandwestem Panama. Solanumdarienenseis distinct fromthese two species in its reddish-goldenbark, lightly winged stems, obtuse, slightlyobliqueleaf bases, andgenerallyglabrous leaves.

105. Solanum dissimile C. V. Morton, Contr. U.S. Natl. Herb. 29: 52. 1944. Type. Colombia. Norte de Santander:Loso and vic. (N of Toledo), 22002400 m, in woods along stream, 6-7 Mar 1927, Killip & Smith 20415 (holotype, US; isotypes, F,

K, NY).

Figs. 170C, 177

322

FLORA NEOTROPICA

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FIG. 177. Solanumdissimile C.V. Morton. Colombia. Killip & Smith20415 (US-holotype).

323

TAXONOMIC TREATMENT

Smallshrubsor treelets, 1-4 m tall;young stems slender and zigzag, densely pubescent with golden, curling,uniseriatetrichomesca. 0.1 mm long;branches arching and flattened, giving the plant a pagoda-like aspect, older plants occasionally with erect branches; bark of older stems glabrate, reddish-brown, not lenticellate. Sympodialunits unifoliate, not geminate except rarely on lower non-reproductive branches. Leavesellipticto narrowlyelliptic,widestat the middle, 5-10 x 1.5-3 cm, the apex acuminate,the base acute, then decurrenton the petiole; laminawith 5-7 pairsof main lateralveins, glabrousandwith the midribraised above, occasionally a few uniseriatetrichomespresent on the prominentyellowish veins below; petioles very short, winged, 2-4 mm long. Inflorescences opposite the leaves, simple, 2-3 mm long, 4-10-flowered, pubescentwithuniseriatetrichomeslikethoseof the stems; pedicel scars corky and raised, evenly spaced ca. 0.5 mm apart.Buds obovoid, becomingmore elongateand ellipsoid with age, the corolla soon exserted from the minute calyx tube. Pedicels at anthesis filiform, 8-9 mm long, taperinggently fromthe calyx tubeto a basal diam. of less than0.5 mm.Flowers with the calyx tube only a slight widening of the pedicel, 0.5-1 mm long, the lobes minute,deltoid, 0.25-0.5 mm long, minutely papillose on the tips, or occasionally with a few uniseriate trichomes;corolla white, 0.8-1 cm diam., lobed 3/4 of the way to the base, the lobes strongly reflexed at anthesis,the tips of the lobes minutelypapillose; anthersca. 2.5 x 1 mm, poricidalat the tips, the pores teardropshaped;free portionof thefilamentsca. 0.25 mm long, the filamenttube narrow,0.5 mm long; ovaryglabrous;style straight,3-3.5 mm long;stigma a papillose area on the widened tip of the style. Fruit a globose, green berry,ca. 1 cm diam.;fruitingpedicels deflexed, slightly woody, 1.2-1.5 cm long, ca. 0.5 mm diam. at the base. Seeds tan, darkbrown in dry material, ovoid-reniform,2.5-3 x 2-2.5 mm, the surfaces minutelypitted.Chromosomenumber:n= 12 (voucher Knapp & Mallet 6817).

800

00

W~~~~~~ A 0

FIG. 178. Distribution of Solanum cruciferum (solid circles), S. dissimile (open circles), S. erosomarginatum (solid square), and S. hypocalycosarcum (solid triangles).

ca. 7 km E of Jaji, I km W of tumoff to La Mesa, 1500 m, 29 Jul 1979, Nee & Whalen17041 (BH, F, K, VEN, WIS); Merida, La Pedregosa,2000 m, 10 Mar 1969, Oberwinkler & Oberwinkler 14771 (VEN); Libertador, El Maciegal, above La Pedregosa, 10 km NW of Merida, 1800 m, 23 Apr 1977, Ruiz Tera~n& Ruiz Perez 13470 (MY); Monte Zerpa, 10 Feb 1984, Schwarzkopf 17 (MY); along quebradaof Cuesta del Barroand Mesa del Trapiche,tributary to Rio Capuri,between Canaguaand El Molino, 25303715 m, 11 May 1944, Steyermark 56479 (MY, VEN). TACHIRA:Headwaters of Rio Quinimari along Quebrada Agua Negra on trail to Paramo de Judio, 5 km S of San Vicente de la Revancha, 15 km S of Providencia, SE of Distribution (Fig. 178). In the Andes of eastern Santa Ana, 2100-2400 m, 7?25'N, 72?25'W, 23 Oct 1984, Colombia and Venezuela,on both sides of the Tachira Knapp & Mallet 6823, 6825 (BH, K, MY, US, VEN); bedepression,in forestsalong streams,from 1500 to 3000 low paramode Tama, between Betania and Tama, 2700 m, 13 Jul 1944, Steyermark 57188 (F, VEN); headwaters of m elevation. Rio Quinimari,along quebradaLas Copas, below Pefia de Specimens examined. VENEZUELA. MERIDA: Pata de Judio (below paramo de Judio), 15 km S of San 27.2 km S of La Azulita, 14.6 km N of Jaji turnoff on La Vicente de la Revancha,30 km S of Alquitrana,SW of Sta. Azulita rd., ca. 2200 m, 8?32'N, 71?16'W, 22 Oct 1984, Ana, 2350-2400 m, 14 Jan 1968, Steyermarket al. 100929 Knapp & Mallet 6810 (BH, MY, US, VEN); Chorreralas (MY, US, VEN); along QuebradaAzul, S of El Reposo, 14 Gonzalez, 4 km E of Jaji turnoff,2 km W of La Mesa turn- km S of Las Delicias, 2150-2300 m, 7?32'N, 72?24'W,22off, on Merida-LaAzulita rd., ca. 1600 m, 83 I'N, 71?12'W, 23 Jul 1979, Steyermark & Liesner 118491 (VEN). 22 Oct 1984, Knapp & Mallet 6814 (BH, K, MY, US, Local names. Venezuela. Merida: mirra. VEN), 6815, 6817, 6818 (BH, MY, US, VEN); La Isla, Solanum dissimile is most closely related to S. Jaji, Finca of Eleazar Davila, 1650 m, 6 Jan 1968, L6pezPalacios 1885 (NY); waterfall along Meridato Azulita rd., erosomarginatum, also of Andean Venezuela. It differs

324

FLORA NEOTROPICA

S

FIG. 179. Solanum erosomarginatumS. Knapp. (Reproduced with permission from Brittonia 38: 274, fig. 1. 1986.)

fromthatspeciesin its usuallyunifoliatesympodia,more congested inflorescences, smallerflowers, less pubescent foliage, and non-erose leaf margins. Individuals of S. dissimile grow to be small trees along streamsin the cloud forest,andare amongthe largestmembersof the S. confine species group. Solanum dissimile, like other members of the group, grows in closed or partially closed canopy forest.Solanumdissimile is quite common where it occurs, and is a majorcomponentof the forest understory.

106. Solanum erosomarginatum S. Knapp,Brittonia 38: 273. 1986.Type.Venezuela.Trujillo:Oldrd.from Bocon6 to Trujillo,ca. 51 km W of Trujillobelow summit,cloudforest,2200-2250 m, 9021N, 70?19W, 20 Oct 1984, Knapp& Mallet 6779 (holotype,MY; Fig. 179 isotypes, BH, MO, US, VEN). Slender shrubs, 1-1.5 m tall; young stems and leavesdenselypubescentwith uniseriatetrichomes0.50.75 mm long, these becoming stiff and curly tipped on older stems;largerstems glabrate,the barkreddishgray.Sympodialunitsdifoliate,geminate.Leavesovate, widestjust proximalto the middle, the marginssome-

what erose, glabrousor with a few trichomesalong the veins and lamina above, densely pubescent beneath with uniseriatetrichomes0.5-1 mm long, these denser along the veins; lamina6-13 x 3-6 cm, with 5-6 pairs of main lateralveins, these strongly impressedabove, prominentandhairybeneath,the apex long-acuminate, the base acute;petioles winged fromthe leaf bases, 6-9 mm long;minorleaves differingfromthe majorones in shapeandsize, orbicular,1.2-2.5 x 1.2-2.5 cm, the apex acute,thebase rounded;petioles2-4 mm long.Inflorescences oppositethe leaves, slender,simple, 0.8-3.5 cm long, 3-15-flowered, bearingonly one or two flowers ata time,denselypubescentnearthebasewithuniseriate trichomeslikethoseofthe stems,thesebecomingsparser distally;pedicelscars irregularlyspaced1-1.5 mm apart, beginning ca. 1/3 of the way from the base. Buds globose when young, laterellipsoid with the exsertionof the corolla.Pedicels at anthesis 1.2-1.5 cm long, tapering fromthecalyxtubeto a slenderbaseca. 0.5 mmdiam. Flowers with the calyx tube broadlyconical, ca. 1 mm long,the lobesdeltoid,occasionallyapiculate,0.5-1 mm long, sparselypubescentwith uniseriatetrichomes0.50.75 mm long; corolla white, 1-1.3 cm diam., lobed nearlyto the base, the lobes stronglyreflexed at anthe-

TAXONOMIC TREATMENT

sis, the tips andmarginsminutelypapillose andwith a few uniseriatetrichomes ca. 0.5 mm long; anthers 22.5 x ca. 1 mm, poricidalat the tips, the pores teardrop shaped;free portion of thefilaments 0.5-1 mm long, the filamenttube0.5-1 mm long, slightlypleatedin the sinuses; ovary glabrous;style straight,5-6 mm long; stigma clavate, the surfaceminutelypapillose.Fruit a globose, greenberry,1-1.2 cm diam.;fruitingpedicels deflexed, slightly woody, ca. 2 cm long, ca. 0.75 mm diam.at the base. Seeds not known. Chromosomenumber: n = 12 (voucher Knapp & Mallet 6779). Distribution (Fig. 178). Known only from the type locality in the state of Trujilloin the Venezuelan Andes at 2200 m in forest understory. Specimensexamined.VENEZUELA.TRUJILLO: Oldrd.fromBocon6to Trujillo,ca. 51 km W of Trujillo, below summit,2200-2250 m, 9?21'N,70?l9'W,20 Oct 1984, Kntapp& Mallet 6777 (BH, MY, US, VEN).

32 5

anduniseriatetrichomes;pedicelscars in distinctpairs, the members of a pair touching, but not overlapping, the distancebetweenpairsca. 1 mm.Buds globose, the corolla soon exserted from the calyx, obovoid with corolla exsertion, minutely pubescent with erect unicellulartrichomes,the young budsblack in driedmaterial. Pedicels at anthesis deflexed, 0.8-1 cm long, taperinggentlyfromthe calyx to a basaldiam.of 0.25-0.5 mm.Flowerswiththecalyxtubebroadlyconical,barely differentiatedfrom the pedicel, in live plantspurpleat the base, ca. 1 mm long, the lobes deltoid, ca. 0.5 mm long, minutelypapillose on the tips; corolla white, ca. 1 cm diam., lobed ca. 2/3 of the way to the base, the lobes stronglyreflexed at anthesis,minutelypapillose on the tips and marginsof the lobes; anthers ca. 2.5 x 1 mm, poricidal at the tips, the pores teardropshaped; free portionof thefilaments ca. 0.3 mm long, the filamenttubeca. 0.3 mm long;ovaryglabrous;stylestraight, ca. 5 mm long; stigma globose, pale papillose in dry material,bright green in live plants. Fruit a globose, greenberry,ca. 1.1 cm diam.;fruitingpedicelsslightly woody, deflexed, greatly expanded and purple at the apex,this characteristicnot obvious in drymaterial,ca. 2 mm diam.atthe apex(dryspecimens),ca. 1 m1mdianm. at the base. Seeds tan, ovoid-reniform,ca. 2.5 x 2 mn, the surfaces minutely pitted, the margins somewhat incrassateanddarkerthanthe seed body.Chromosome number:n = 12 (voucher Knapp & Mallet 6265).

Solanumerosomarginatumis closely relatedto S. dissimile,also of AndeanVenezuela,butdiffersfrom thatspecies in its larger,darkergreen leaves with conspicuously erose margins, smaller stature, difoliate, geminate sympodia, longer inflorescences, and larger flowers. Leaf size in S. erosomarginatumvaries considerably,but the consistent presence of minor leaves is enough to distinguish this species from any other closely related to it. Solanum erosomarginatumis a delicate shrubof forest understoryand, althoughrare Distribution (Fig. 178). In rainforests on the distributionally,is rathercommonwhere it does occur. westem slopes of the Andes in Ecuador,from sea level to ca. 600 m.

Specimens examined. ECUADOR. BoliVAR: Charquiyacu,lower W slopes of CordilleraOccidental, 600 m, 4 Oct 1943, Acosta Solis 6120 (F). EL ORO: Along Rio CristalnearMontalvo,70 m, 8-11 Jul 1962, Jdtiva & Epling 49 (NY); on Loja-Machalard., 10 k.m W of Las Balzas, 11.5 km E of Loja-Pifiascrossroads Delicate shrubs 1-3 m tall; young stems and (31.5 km E of La Avanzadanear SantaRosa), ca. 700 leaves minutely pubescent with erect unicellular and m, 3?45'S,79?50'W,7 Feb 1984, Knapp& Mallet 6265 uniseriatetrichomesca. 0.01Inm long, these not soon (BH, MO, QCA, QCNE,US); between Portovelo and deciduous;brancheswith a flattenedaspect;barkof the Zaruma,640--1150m, 22 Aug 1943, SteylermiartA53986 olderstemspale, dryinggrayor white.Sympodialunits (NY). GUAYAS: Milagro,30 Jun-2 Jul 1923, Hitchcock usually difoliate, geminate. Leaves narrowly ovate, 20241 (NY, US); Naranjito8 km to E along railway, widest in the proximal 1/3 of the blade, glabrous on Hcda.San Antonio,50 m, 2?10'S,79?22'W,3 Jul 1976. both surfaces, the main lateral veins prominent and Olgaard & Bals/ev 7519 (AAU). Los Rios: Montalvo, yellowishbeneath;majorleaves 10-15 x 3.5-5 cm, with foothills of Andes, ca. 40 km E of Babahoyo, 1045,"S 7-8 pairsof mainlateralveins, the apex acuteto acumi- 79?17'W,30 Mar-2 Apr 1973,Holin-Nielseni et at. 2617 nate,thebase truncateto rounded,decurrenton the short (AAU, F, MO, NY). petiole; petioles minute, ca. 5 mm long; minor leaves With its closely spaced pedicel scars, geminate differingfromthemajorsin size andshape,elliptic,2.5- leaves,andratherstout(fortheS. confinespeciesgroup) 4.5 x 1.2-2.5 cm, the apex acuteto acuminate,the base inflorescences, S. hypocalycosarcumis related to S. truncateto rounded;petioles 3-5 mm long. Inflores- pastillum and S. tuerckheimii of montane Central cences opposite the leaves, simple, 0.8-1.5 cm long, America.It sharesslightlyincrassateseed marginswith 10-14-flowered, minutely pubescentwith unicellular S. leptorhachis,also of westernEcuador,butis otherwise 107. Solanum hypocalycosarcum Bitter, Repert. Spec. Nov. Regni Veg. 11:489. 1913. Type. Ecuador. Guayas:Balao, Feb 1892, Eggers 14405 (hoFigs. 170D, 180 lotype, M [F neg. 2663]).

FLORA NEOTROPICA

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notatall similarto thatspecies.Thetinyerecttrichomesof the stems and inflorescences, and their absence from the leaves, are characteristicofS. hypocalycosarcum. The leaf shapeofS. hypocalycosarcum,narrowlyovate and widest in the lower 1/3, is also quite distinctive. TherelationshipsofS. hypocalycosarcum areenigmatic, and many more collections are needed to assess its variabilityand to furtherinvestigate charactersinformative as to its position in the section. Solanum hypocalycosarcum is not a common plantandis knownfromonly a few collections.It grows nearstreamsin the rainforestsof southwestemEcuador, and apparentlyis not found furthernorthin the forests of the Choco floristicprovince.WhereI have collected this species I only found one plant, indicatingperhaps that it has small populationsizes where it does occur.

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tium flum. Solim6es, Jan 1851, Spruce 1700 (lectotype, G-DC, here designated [F neg. 23127]; isolectotypes, AWH, BM, K, MO, NY [US neg. 2435]). Figs. 171A,B, 181 Solanum loretoanum Bitter, Repert. Spec. Nov. Regni Veg. 18: 70. 1922. Type. Peru. Loreto: Yurimaguas, E. Ule 6277 (lectotype, HBG, here designated; isolectotype, B [destroyed: F neg. 2619]).

Much-branched shrubs ("herb" on Kruckoff 8365, but distinctly woody) 1-4 m tall; young stems glabrousor pubescentwith soft, pale yellow uniseriate trichomes 0.1-0.5 mm long; bark of the older stems reddish-brown, smooth and shining, sparsely lenticellate and occasionally pubescentwith the same trichomes as the young stems; branchesarching, giving the shrub a flattened aspect. Sympodial units difoliate, geminate,stronglyanisophyllous.Leaves el108. Solanum leptopodum Van Heurck & Mull. liptic to ovate, glabrousabove, glabrousor somewhat Arg.,Observ.Bot. 57. 1870.Type.Brazil.Amazonas: pubescent beneath, if pubescent, with soft uniseriate Ad orammeridionaleumflum. Amazonum,ad os- trichomes like those of the young stems on the midrib

TAXONOMIC TREATMENT z"D' 'S;S

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and principal veins, trichomes 0.1-0.75 mm long on the midrib, shorteron the smaller veins; majorleaves ellipticto ovate,widest at orjust below the middle,9.519.6 x 3.6-8.5 cm, with 10-1 1 main lateralveins, impressed above, prominent,reddish-brownand occasionallypubescentbeneath,the apexacuminate,thebase acute; petioles 3-5 mm long; minor leaves orbicular, 0.3-1.9 x 0.3-1.9 cm, the apex rounded,the base acute; petioles 0-2 mm long. Inflorescences opposite the leaves, simple, slender,0.7-3 cm long, 5-20-flowered, but only bearing a few flowers at a time, glabrous or pubescentwith soft uniseriatetrichomeslike the stems andleaves;pedicelscars evenly spaced0.5-2 mm apart, beginning 5-9 mm fromthe base of the inflorescence. Buds globose, pubescentwith soft uniseriatetrichomes 0.05-0.5 mm long, occasionally glabrous.Pedicels at anthesis deflexed, 0.7-1 cm long, filiform, tapering from the calyx tube to a threadlikebase ca. 0.25 mm diam. Flowers with the calyx tube 0.5-1 mm long, broadly campanulate,lobes deltoid, 0.5-1 mm long, pubescent with the same trichomes as the buds, occa-

sionally glabrous;corolla white, 6-7 mm across,lobed nearlyto the base, the lobes stronglyreflexed at anthesis, minutelypapillose on the marginsandbacks of the lobes; anthersca. 2 x 1.25 mm, poricidalat the tips, the pores teardropshaped;free portionof thefilamentsca. 0.5 mm long, the filamenttubeca. 0.25 mm long;ovary glabrous;style 4-5 mm long; stigma minutely white papillose,notmarkedlydifferingin shapefromthe style. Fruita globose,greenbeny,mustardyellowon herbarium specimens, 1-1.2 cm diam.;fiuiting pedicels woody, deflexed, 2.1-2.6 cm long, ca. 2.5 mm diam.apically,1 mm diam.at the base.Seedsbrown,ovoid-reniform,ca. 3.5 x 2.5 mm,the surfacesminutelypitted.Chromosome number:n = 12 (voucherKnapp& Mallet 6610). Distribution (Fig. 183). Upper Amazon basin from Manaus, Brazil (type), to Peru, Ecuador, and Colombiain primaryforest and second growth, 100 to 500(-1900) m. Selected specimens examined. COLOMBIA. AMAZONAS:

Loretoyacu river, 100 m, Oct 1945,

32 8 Schultes 6681 (K, US), Oct 1946, Schultes & Black 8413 (US). CAQUETA: Rio Dedo, Dec 1930, Uribe P s.n. (US). META: Sierra de la Macarena, Central Mtns., collecting ridge, 1450 m, 21 Jan 1950, Philipson et al. 2184 (BM). PUTUMAYO: Rio Putumayo, Pifiufia Negro, 240 m, 20 Nov 1940, Cuatrecasas 10698 (F, US); Rio Putumayo near Ospina, 230 m, 24 Nov 1940, Cuatrecasas 10784A (US); Umbria, 325 m, 0?54'N, 76?10'W, Jan-Feb 1931, Kling 1913 (F,MO, NY, US); Rio Caqueta. downriver from Puerto Lim6n, 300-350 m, 20 Dec 1968, Plowman 2179 (K, US). ECUADOR. MORONA-SANTIAGO: Palora,Sinai, 820 m, 2?10'S, 78?05'W, 5 Nov 1991, Guidifo et al. 1593 (QCNE); along Rio Metzera Grande on Hacienda Sangay (plantation of Compania Ecuatoriana del Te, S.A.), near Palora, ca. 950 m, 1?40'S, 77?58'W, 15 Feb 1984, Knapp & Mallet 6286, 6289, 6294 (BH, K, QCA, QCNE, US). NAPO: Afiangu on S bank of Rio Napo 2 hours below Coca, 300 m, 0?30'S, 76?25'W, 9 Apr 1982, Balslev 2386 (NY); Lago Agrio, Parroquia Dureno, indigenous community Cofan-Dureno, 350 m, 0?02'S, 76?42'W,29-31 Dec 1987, Ceron M. & Cer6n 3051 (BM, MO, QCNE); Parque Nacional Yasuni, petroleum exploration line Daimi II of CONOCO, 230 m, 1002'S, 76?10'W, 27 May 1988, Hurtado et al. 40 (QCNE); trail to Palma Roja, 28 Apr 1986, Jaramillo 8517 (AAU); Campanococha,L bank Rio Napo, trail from Campanococha to Dayono. 360-400 m, 22 Aug 1980, Jaramiiillo & Coello 3738 (MO, NY); Afiangu, NW corner Parque Nacional Yasuni, close to Rio Napo, ca. 255 m, 0?32'N, 76022'W, 1-30 Oct 1983, Kor-ning & Thomnsen47061 (AAU, QCNE); San Pablo de Aguarico, 21 Jan 1984, Lest ure 2106 (NY); Shushufindi (Nueva Loja), rd. Coca (Puierto Francisco de Orellana)-Lago Agrio, ca. 50 km NE of Coca, 10 Nov 1973, Liogo S. 3308 (GB, MO); Santa Cecilia, Mufiozlandia on Rio Aguarico, island in 1461 (MO); near river, 2 Apr 1972, MacBryde & Dwv_ver Tena. 400 m, 2-11 Apr 1935, Mexia 7149 (F, UC, US); Napo rd., Limoncocha, Sep 1969, Mowvbray 69966 (MO); R margin of Rio Quijos, Finca La Ave Brava of Segundo Pacheco, 1800-1900 m, 0WI2S, 77?39'W, 710 Sep 1990, Palacios 5335 (QCNE); Rio Aguarico, Shushufindi, 244 m, 8 Mar 1975, Vickers 232 (F). PASTAZA: Cabeceras on the Rio Bobonaza, ca. 12 km E of Puyo, 16 Nov 1974, Logo S. 4609 (GB); 3-4 km E of Puyopungu. 28 Sep 1976, Lugo S. 5037 (GB); 17 km N of Palora, ca. 2 km N of Tashapi(Rio Pastaza crossing), 46 km S of Puyo, along Puyo-Palora rd., ca. 900 ni, 142'S, 77?52'W, 17 Feb 1984, Knapp & Mallet 6301 (BHI, K, QCA, QCNE, US); Rio Chico, affluent of Rio Pastaza, village of Rio Chico, 10 km S of Puyo, 3 km S of Tarqui, 1000 m, 1'30'S, 77?55'W, Aug 1979, Shemlluck& Ness 198 (F). PERU. LORETO: Rio Ampiyacu, environs of Pucuraquillo, 8 Mar 1981, Davis et al. 785 (F, K, NY, US); Maynas, 183 1, Dia: 2476B (W); Yanamono tourist camp, Rio Amazonas above mouth of Rio Napo, 130 m, 3028'S, 72050'W, 27-28 Dec 1982, Gentrv & Emnmlotns 38738 (MO); Yanamono, Explorama Tourist Camp on

FLORA NEOTROPICA

Rio Amazonas,130 m, 3?25'S, 72?50'W,12 Jun 1986, and Gentryet al. 54278 (MO,NY); betweenYurimaguas Balsapuerto,lower Huallagabasin, 135--150m, 26-31 Aug 1929, Killip & Smith 28163 (F, NY); Yanamono,

Exploramatouristcampon Rio AmazonasbetweenIndianaandmouthof Rio Napo, ca. 80 km NE of Iquitos, ca. 100 m, 3?28'S, 72?50'W, 27 Jul 1984, Knalpp 6608,

6610, 6612 (BH, K, US, USM);Rio Nanay,Samilo,ca. 100 m, 20 Feb 1969, Plowman2523 (GH, K); Rio Yaguasyacu,affluentof Rio Ampiyacu,Brillo Nuevo and vic., 2?40'S, 72?00'W, 12 Apr 1977, Plowman ct al. 6843

(F); Yurimaguas,May 1855, Spruce 3877 (K); Yanamono ExploramaTouristCamp,halfwaybetweenIndiana and mouthof Rio Napo, 130 m, 1 Aug 1988, v,an der Werff et al. 9903 (MO, NY); lower Rio Hluallaga, Yurimaguas, 155-210 m, 22 Oct 1929,Wi/lit,ms 3851

(F); lower Rio Huallaga,Fortaleza,Yurimaguas,155210 m, 30 Oct 1929,Williams 4377 (F). San Martin: Convento,trail to Nuevo Lamas,km 68 of TarapotoYurimaguasrd., 200 m, 6?16'S,76017'W 10 Aug 1986. Knapp 7966 (MO, NY, USM). SAN MARTiN: Km 40 Tarapoto-Yurimaguas and Rio rd., trail to Rio TirnyacL Cashiyacu,eventuallyto salt mines on Rio Cashiyacu, 300-400 m, 6?25'S, 76?15'W, 24 Apr 1986, Knapp & Mallet 7211 (MO, USM);Convento,trail to Nuevo Lamas, km 68 of Tarapoto-Yurimaguas rd., ca. 200 rn, 10 Aug 1986,Knapp7966 (MO, USM);Pongode Aguirre, along Rio HuallagadownstreamfromChazuta,ca. 200 m, 6?25'S, 75?25'W, Jan 1987, Knapp & Ltallet 8549a (F, K, USM). UCAYALI:Pucallpa, kmn8, 20)0 ':1, 6 Jun 1960,Woytkowski 5774 (US). BRAZIL. AMAZONAS: SaoPaulode OlivenSanear Palmares,11 Sep-26 Oct 1936, Krukoff 8365 (F, MO, NY, U, US).

Local names. Colombia. Putumayo. salado sacha. Ecuador. Napo: huiarapanga, nuairapuhana panga (Quichua),oyo ha'o (Secoya). is most similar to S. of the rainforest of SE Brazil andto S. constipulatumn fine, also of easternAmazonia.It shareswithS. confine soft, golden pubescence, somewhatelongate inflorescences, and fruiting pedicels twice the lengt:hof the floweringpedicels.WithS. stipulatuntit sharesstrongly anisophyllousgeminateleaves andcompletelyorbicular,oftenappearingstipule-like,minorleaves.Theleaves Solanum leptopodum

of both S. leptopodum and S. stipulatunmare veiy shiny

and darkgreen above and paler beneath. Solanum leptopodum is variable in pubescence throughoutits range. Specimens from Peru and Colombia have been called S. loretoanuim1and are nearly glabrous,but intermediatesexist between these nearly glabrousforms andthe more pubescenttype specimen from near Manaus, much furtherdown the Amazon. Thetrichomesin all the specimensareof the sametype: soft, yellow uniseriatetrichomes,which varyin length in differentareas. Some populationsof S. leptopodum

329

TAXONOMIC TREATMENT

fromeasternEcuador(particularlyin the Rio Aguarico basin) have narrower leaves than the more typical plants.This characterhowever, gradesinto the typical leaf shape, and no other differences between the two forms exist. Solaniumleptopodumis one of the few species of sect. Geminatafor which indigenoususes have been reported.In the Rio Shushufindiregion of eastem Ecuador,the Secoyapeopleuse the leavesofS. leptopodum as a treatmentfor "crybaby."The leaves are crushed into lukewarmwater,and the infantis bathedwith the infusion (Vickers & Plowman, 1984). In this same general area of eastern Ecuador,but among Quichua speaking people, the plant is used for the treatmentof "malviento," a broadgeneral illness category,and for sleeplessness in children (Balslev 2386). In both of these treatments,bunchesof leafy branchesarewaved over the afflicted person. Balslev (pers. comm.) has often seen S. leptopodumcultivated in dooryardgardens among these people. In lowland Colombia, a decoction of leaves is used to treatskin irritations(Plowman 2179).

1.5-2 cm long, taperingfrom the calyx tube to a fillformbase ca. 0.2 mm diam.Flowerswith the calyxtube broadly conical, ca. 1 mm long, glabrous or minutely puberulent,lobes rounded,0.5-1 mm long, papillose at the tips; corolla pale greenish-white,translucent,5-8(1 1) mm diam., lobed ca. 2/3 of the way to the base, the lobes planaror slightly reflexed at anthesis,papillose at the tips; anthersca. 2.5 x 1 mm, poricidalat the tips, the poresteardropshaped;free portionof thefilaments less than 0.5 mm, the filament tube ca. 0.5 mm long;ovaryglabrous;style3-4 mm long, straight;stign2a brightgreenin live specimens,small-capitate,minutely papillose.Fruit a globose berry,0.8-1 cm diam.,greeniish-yellowwhenripe;fruitingpedicelgreatlyelongated and hanging, 3.5-4 cm long, tapering from a basal diam.of ca. 0.5 mm to an expandedapex 3-4 mm diarn. (thismuchmoreapparenton live specimens).Seetdspale tan,ovoid-reniform,ca. 3 x 2 mm, the surfacesminutely pitted,the marginssomewhatincrassate.Chromosomie number. n = 12 (voucherKnapp & Mallet 6160). Distribution (Fig. 183). In wet forestat low and middle elevations on the westem slope of the Andes from southem Colombia to northemPeru, from 50 to 2000 m, usually growing as an understoryshrub. Selected

examined. COLOMBIA. specimens 109. Solanum leptorhachis Bitter, Repert. Spec. NARINo: La Planada, trail to El Hond6n, 6-12 km SW Nov. Regni Veg. 18: 50. 1922. Type. Ecuador. of La Planada, 1750-1800 m, 1?04'N, 78'02'W, 5 Jan Manabi: "In schattigen Waldernbei El Recreo," 1988, Gentry et al. 60401(MO).VALLE DE CAUCx: Near Apr, Eggers 15743 (holotype, M [Morton neg. Yatacue, Alto Anchicaya, near CVC hydroelectric plant, 8707]). Figs. 170E,F, 182 valley of Rio Dagua, 710-800 m, 3?38'N, 76045'W. 16 Solan ini inlacrotonuni of A. H. Gentry& Dodson, Jul 1984, Gentry & Monsalve 48193 (CUVC, MO, NY). ECUADOR.AZUAY:Jesus Maria-Molleturo rd., Selbyana 4: 554, plate 260A. 1978. Not of ca. 12 mi from Guayas border, ca. 1200 m, 16 Jul 1977, Bitter.

Slender shr-lubswith arching, planarbranches, 1-2 m tall, young stems zigzag, glabrousor puberulent with minuteuni- orbicellulargoldentrichomesca. 0.01 mm long; older stems glabrate,greenish-yellow.Sympodial/units unifoliate,not geminateexcepton thebasal, non-reproductivenodes. Leaves elliptic, widest at the middle,glabrousandmatteabove,glabrousorminutely hispidulouson the veins beneath,the blades 9-15(26) x 4-6(14) cm, with 8-10 pairsof mainlateralveins,these parallelto one anotherandat rightangles to the midrib, the apex acuminate,the base obtuse, then sharplyattenuate,decurrenton the petiole andstem;petioles 0.51.7 cm long, slightly winged from the decurrentleaf bases.Inflorescencesoppositethe leaves, long andslender, filiform, 1-8 cm long, bearing 1-2 flowers at a time, but with up to 25 pedicel scars, glabrousor minutelypuberulentwith trichomeslike thoseof theyoung stems;pedicel scars widely and unevenly spaced, 3-7 mm apart.Bludsglobose, glabrousor minutelypuberulent, greenish-white, becoming obovoid just before anthesis.Pedicels at anzthesisthreadlikeand hanging,

Boeke & Loyola 2179 (NY). CARCHI: N of Carmlen, rd. to Chical, 2000-2200 m, 0?17'N, 78?13'W, 10 Feb 1992, Palacios et al. 9818 (QCNE).COTOPAXI:Tenefuerte, Rio Pilal6, km 52-53 Quevedo-Latacunga, 750-1300 m, 21 Feb 1982, Dodson & Gentry 12804 (MO, QCNE); Quevedo-Latacunga rd., 2-4 km W of Pilalo, 73--75 km NE of Quevedo, 2200-2300 m, 25 Dec 1978, Luteynii & Lebrdn-LuteLVn 6490 (NY). ESMERALDAS: Vic. of L-ita on lbarra-San Lorenzo R.R., 550-650 m, 9 Jun 1978. Madison et al. 5060 (F), 11 Jun 1978, Madisoni et al. 5205 (F). GUAYAS:Cerro Manglar Alto, 1200 ft. 14 May 1923, Anthony& Tate26 (US). IMBABURA: Lita, 540 m, 28 May 1949, Acosta Solils 12610 (F); Cart6n Cotacachi, ParroquiaApuela, Cuellage, 1600 m, 0015'N, 78X25'W, 10 Jul 1992, Tipa: & Aulestia 1672 (QCNE). Los Rios: Km 170-175 Sto. Domingo-Quinind6 rd., 300 m, 5 Sep 1949, Acosta Solis 13791 (F): Baba, 0-5 m, 1?47'S, 79?40'W, Sep 1993, Corniejo 397 (QCNE); Rio Palenque Biological Station, km 56 Quevedo-Santo Domingo, 150-220 m, 7 Aug 1975, Dodsoni 5933 (US); Hacienda Las Balsas near Montalvo, Bosque de Oro, 300400 m, 27 Jul 1962, Jcativa& Epling 218 (NY): Centro Cientifico Rio Palenque, km 56 Sto. Domingo-Quevedo, 150-300 m, 30 Jan 1984, Knapp & Mallet 6222, 6223

FLORA NEOTROPICA

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FIG. 183. Distribution of Solanuim leptopodum (solid circles), S. leptorhachis (open circles), S. morii (solid square), and S. nematorhachis (solid triangles).

(BH,K, QCA, QCNE, US). PICHINCHA:20 km W of Santo Domingo de Los Colorados, 1000 m, 1I Jun 1961, Cazalet & Pennington 5297 (K, NY, US); Reserva Forestal ENDESA, Rio Silanche, km 113 of Quito-Puerto Quito rd., W slopes, 10 km N of main rd., 650-700 m, 0005'N, 79002'W, 7 Dec 1984, Jaramillo 7412 (AAU); Reserva Floristica-Ecol6gica "Rio Guajalito," km 59 of old Quito-Sto. Domingo de los Colorados rd., 3.5 km NE of the rd.,W slopes of Volcan Pichincha, 1800-2200 m, 0?13'53S, 78048'1OW,12 Aug 1985, Jaramillo & Zak 8035 (AAU, K, MO, NY); Tinalandia, km 112 QuitoSto. Domingo, 500-1000 m, 0020'S, 78050'W, 15 Jan 1984, Knapp & Mallet 6157, 6160 (BH, K, QCA, QCNE, US); Reserva Floristica-Ecol6gica "Rio Guajalito," km 59 of old Quito-Sto. Domingo del los Colorados rd., 3.5 km NE of rd.,W slopes of Volcan Pichincha, 0013'53"S, 78048'10"W, 13 Aug 1986, Zak 1113 (AAU); Quito-Chiriboga-Empalme, km 59, 15 km NW of the rd., 1700-2000 m, 23 Sep 1986, Zak 1278, 1283, 1285 (NY); Reserva Floristica-Ecol6gica "Rio Guajalito," km 59 of old Quito-Sto. Domingo de los Colorados rd., 3.5 km NE of rd.,W slopes of Volcan Pichincha, 0?13'53"S, 78048'10"W, 21 Dec 1986, Zak 1483 (AAU, NY); Bosque Protector La Perla, km 41 Sto. DomingoQuininde, 220 m, 0?0-02'S, 79022'21-32"W, I Jul 1990, Zak et al. 5406, 9 Aug 1990, Zak et al. 5544, 11 Aug 1990, Zak et al. 5619 (all QCNE). PERU. TUMBES:Zarumilla, Bosque Nacional de Tumbes, near Campo Verde, 600-800 m, 19 Dec 1967, Simpson & Schulnke V 429 (F).

Solanumleptorhachisis most closely relatedto S. tenuiflagellatumof montanecoastalVenezuela,sharing with that species extremely long inflorescences, flowers with the corolla lobes planaror only slightly reflexed at anthesis,and fruitingpedicels threeto four times theirlengthat anthesisandgreatlyexpandedand corky at the apex. Plantscollected at higherelevations (Boeke& Loyola2179, Luteyn& Lebr6n-Luteyn6490, Zak 1113, 1283, 1285) have larger,apparentlythicker leaves andlargerflowersthanmoretypicalplants.This species is a relatively common component of the understory,oftengrowingin old lightgapsandalongsmall streamsunderclosed canopy. InTinalandia,nearSantoDomingo,Ecuador,the leaves of Solanum leptorhachis are fed upon by the larvaeof the ithomiinebutterflyPteronymiaglauca (see TableXI), and the plants are often severely defoliated by these caterpillars. The specimen collected in Tumbes, Peru (Simpson& Schunke V 429), consists of a few apparently virally infected sucker shoots with very small, deformedleaves and very shortinflorescences.

110. Solanum morii S. Knapp, Brittonia 44: 61. 1992. Type. French Guiana. Saul, on short cut to airport,ca. 200 m, 3?37'N, 53?12'W,28 Aug 1988, Mori et al. 19213 (holotype, NY; isotypes, IBE, Fig. 184 MO, P).

332

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TAXONOMIC TREATMENT

Shrubs1-3 m tall;young stemsandleaves hispid withgolden,curling,uniseriatetrichomesca. 1mm long; barkof older stems reddish-brown;the stems arching, slender.Sympodialunitsdifoliate,geminate.Leavesnarrowly elliptic-ovate,widest at orjust below the middle, glabrous adaxially, pubescent with golden curling uniseriatetrichomesalong the veins abaxially,the trichomes denseralong the midrib,the marginsundulate; major leaves 6-15 x 2-4 cm, with 7-9 pairs of main lateralveins, these dryinggolden-redabaxially,joining in a series of archesnearthe marginof the lamina,the apex long-acuminate,the base attenuate;petiole ca. 0.5 cm long;minorleaves differingfromthe majorsonly in size, 2-4 x 1-2 cm, the apex long-acuminate,the base attenuate;petiolesca. 0.3 cm long.Inflorescencesopposite the leaves, simple, slender,glabrousor with a few minute golden trichomes near the tips, occasionally sparselypubescentwith curlingtrichomeslike those of the stems, 0.5-4 cm long, with up to 25 scarsbut bearing only 1 or 2 flowers at a time, these soon deciduous; pedicel scars evenly spaced ca. 1 mm apart.Buds globose, the corollaexsertedfromthe calyx tube.Pedicels (itanthesisfiliform,deflexed, glabrous,0.8-1 cm long, ca. 0.5 mm diam. at the apex, ca. 0.25 mm diam. at the base. Flowers with the calvvxtube conical,the tube0.5I mm long, the lobes broadlydeltatewith a swollen apical projection,ca. 0.5 mm long, glabrous;corollawhite or greenish-white,0.8-1 cm diam.,lobed3/4 of the way to thebase, the lobes reflexedat anthesis,minutelypapillose at the tips;anthers2.5-3 x 0.75-1 mm, poricidal at the tips, the poresteardropshaped;freeportionof the filamentsca. 0.5 mm long, the filamenttubeca. 0.5 mm long, glabrous;ovary glabrous;style glabrous,heteromotphic,in short-styledflowers1.5-2 mm long,included withinthe anthercone, in long-styledflowers4-4.5 mm long,exsertedfromtheanthercone;stigmaminutelypapillose.Fruita globoseberry,greenandoccasionallywhite at thebase,thepericarphardat maturity; fruitingpedicel deflexed, 2--2.5 cm long, ca. 0.5 mm diam. at the base, ca. 2.5 mm diam.attheapex.Seedsovoid-reniform,3.54 x 2-2.5 mm, pale tan,the marginsslightlythickened, the surfacesminutelypitted. Distribution (Fig. 183). Central Surinamand the centralplateauof FrenchGuiana,with most collections fromnearSaul, at ca. 200 m. Usually collected in secondary growth forest. Specimens examined. SURINAM. Nassau, forest at border of Plateau A above camp km 10.5, between ferrite boulders, ca. 500 m, 18 Mar 1949, Lanjouwt,& Lindemnan2819 (U). FRENCH GUIANA. Grand Inini, R bank toward Degrad Four-mi,250 m, 12 Aug 1985, de Granville et al. 7446 (B, BR, CAY, MO, NY, P, U); Montagne Bellevue

333 d'Inini, central plateau, W extremity, 700 m, 10 Sep 1985, de Granville et al. 8120 (B, CAY-n.v., INPA-n.v., MG-n.v., MO, NY, P, U); Mont Galbao,E sector, 550 m, 3036'N, 53017'W, 12 Jan 1986, de Granville et al. 8629 (B, CAY-n.v., NY, P, US); Saul, crique Cochon, 7 Mar 1975, Hal/e 2377 (MPU); Saul, Belizon rd., 6 Mar 1976, Ha/l 2406 (MPU); Saull, 18 Oct 1986, de Foresta 666 (NY, U); Sau], in vic. of airport, 200-250 m, 3?37'N, 53?12'W, 30) Aug 1987, Marshall & Rombold 236 (IBE, NY); Saul, Circuit Petit Boeuf Mort, 7 Dec 1976, Mori et al. 8702 (MO): Saul, base of Mont Galbao, 13 Dec 1976, Mori et al. 8762 (NY); Saul, Montws la Fumee, 200-400 m, 3?37'N, 53?12'W, 2 Nov 1982, Mori & Boom 15149 (NY, U); Sail, 12. Aug 1984, Privost 1613 (MPU, NY); Saul, circuit Petit Boeuf Mort, 18 Mar 1985, Prevost 1815 (NY); Saul region, along trail 14 (Trace Belvedere Nord) and 16 (Jct. La DouaneBelvedere), SE of Saul beyond airportnear CriqueCochorn, 200-210 m, 3?34'N, 53?llW, 30 Oct 1986, Skog et al. 7143 (NY, US); 2 km N of Sail, trail to Boeuf Morte, pait of Rt. de Belizon, 3?37'N, 53?12'W, 31 Oct 1986, SkAogct al. 7187 (NY, US). Solanum morii is closely relatedto S. capillipes of Trinidadand the adjacentPariaPeninsulain Venezuela. It differs from that species in its consistently difoliate, geminate sympodial units and its smaller leaves with long-acuminateapices. It appearsto be a plant of disturbedareasand secondaryvegetation.

111. Solanum nematorhachis S. Knapp, Bull. Bril. Mus. (Nat. Hist.),Bot. 22:147. 1992. Type.Colombia. Choc6: SanJose del Palmar,Cerrodel Torra,E slope between heliportand summit,2000-2400 mrt, 28 Aug 1988, Silverstone-Sopkin et al. 4846(holo-

type, CUVC; isotypes, MO, NY).

Fig. 185

Shrubs, 1-3 m tall; young stems and leaves densely pubescentwith golden, curl-tippedtrichomes ca. 0.5 mm long; barkof older stems darkbrown, glabrous or only sparsely pubescent; brancheshorizontally arching. Sympodial units unifoliate. Leaves ellip-

tic, with 8-9 pairsof brochidodromus mainlateralveins, glabrousbut with a few scatteredgolden uniseriatetrichomes along the midribadaxially,sparselyto densely pubescentwith golden trichomesalong the main veins abaxially, 8.5-17 x 2.5-8 cm wide, the apex acuminate, the base acute, not winged onto the petiole: petiole ca. 0.5 mm long. Inflorescencesoppositethe leaves or intemodal, simple, glabrousto sparselypuberulent at the tip, 6-17 cm long, 30-60-flowered, but bearing only two orthreeflowersat a time;pedicel scars widely andunevenly spaced 4-5 mm apart.Buds globose, the corollaexsertedfromthe calyx tube.Pedicels at anthesis deflexed, glabrous, filiform, 5-8 mm long, ca. 0.2.5

mm diam.at the base.Flowers with the calvxtubeconical to somewhatcyathiformandswollen (CerroIngles),

334

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Fm. 185. Solanumnematorhachis S. Knapp. (Reproduced with from Bulletin the ofo permission Britisherient Mu-*ntal,er4ke seum, NaturalHistory(Botany) 22:de152,cm3b. 1992.)o fig.;iud

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TAXONOMIC TREATMENT

335

FIG. 186. Solanum ombrophilum S. Knapp. (Reproduced with permission from Brittonia 38: 283, fig. 8. 1986.)

0.5-1.5 mm long, the lobes deltoid, ca. 0.5 mm long, glabrous or sparsely pubescent with uniseriate trichomeslike thoseofthe stemsandleaves;corollawhite, 7-8 mm diam.,lobed nearlyto the base, the lobes more or less reflexed at anthesis, minutely papillose at the tips and margins; anthers 1.5-2.5 x ca. 1 mm wide, poricidal at the tips, the pores teardropshaped; -free portionof thefilaments ca. 0.5 mm long, the filament tube ca. 0.25 mm long; ovary glabrous;style straight, glabrous, ca. 4 mm long; stigma capitate,the surface minutelypapillose.Fruit a globose, greenberry,ca. 1.5 cm diam. when fresh, 1-1.2 cm diam. dried;fruiting pedicels woody, deflexed, 1.5-2 cm long, ca. 1.5 mm diam. at the base, 4 mm diam. at the apex. Seeds dark brown, ovoid-reniform,3.5-4 x 2.5-3 mm wide, the surfacessmoothto minutelypitted.Chromosomenumber not known. Distribution (Fig. 183). In the CordilleraOccidental of Colombia on the Choco/Valle border, from 1500 to 2400 m. Specimensexamined.COLOMBIA.CHoc6: San Jose del Palmar,Cerrodel Torra,W slope, below the heliportat base of cerro,Rio Negro drainage,1680-1940 m, 10 Aug 1988, Ramoset al. 1111(CUVC,MO, NY). VALLE DE CAUCA: S slope of CerroIngles (Cordillera Occidental, Serraniade los Paraguas), 1 hour in jeep

from El Cairo, 2260 m, 5 Jan 1987, Silverstone-Sopkin et al. 2975 (CUVC, MO, NY).

Solanum nematorhachis is related to S. tenuiflagellatum and S. leptorhachis,both of which share extremelylong inflorescenceswith this species. It differs from those species by the charactersin the key. Solanumnematorhachisoccursat higherelevationsthan S. leptorhachis, the only other similar species with which it is potentially sympatric.

112. Solanum ombrophilum Pittier ex S. Knapp, Brittonia38: 282. 1986. Type. Venezuela.Aragua: ParqueNacional HenriPittier,Portachueloto Pico Periquito trail, 1100-1400 m, 10?21'N, 79?42'W, 28 Oct 1984,Knapp& Mallet 6857(holotype, MY, isotypes, BH, K, US, VEN). Fig. 186 Solanum arboreum of Steyermark & Huber, Flora del Avila 824. 1978. Pro parte. Not of Dunal.

Slender, single-stemmed shrubs, 1-2 m tall; young stems and leaves glabrousor minutelyred-papillose; stemswinged fromthe decurrentleafbases; bark of older stems pale grayish-red.Sympodial units unifoliate.Leavesovateto narrowlyelliptic,thinandmembranous,widest at or proximalto the middle, glabrous on both surfaces, 10-30 x 4-15 cm, with 9-11 pairsof

FLORA NEOTROPICA

336 700

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FIG. 187. Distribution of Solanum ombrophilum (solid circles) and S. pastillum (open circles).

Parque Nacional Henri Pittier, Maracay-Ocumare rd., RanchoGrande,Pico Paraiso,1200 m, 21 Jan 1969, Bunting 3304 (MY); Alto de Choroni, 1200 m, 15 Apr 1969, Ferrari 738 (MY); ParqueNacionalHenriPittier,Fest 21, 39 (VEN); Parque Nacional Henri Pittier, Portachuelo, 31 Sep 1975, Kalin Arroyo & Aristeguieta75108 (VEN); ParqueNacional Henri Pittier, Rancho Grande,trail to Pico Guacamayobehind station, 1100-1400 m, 10?21'N, 67?42'W, Knapp & Mallet 6842, 6843, 6845, 6854 (BH, MY, US, VEN); Knapp & Mallet 6844, 6847, 6849 (BH, K, MY, VEN); rd. between Maracay and Ocumare, El Portachuelo, 1100 m, 8 May 1925, Pittier 11814 (MO, VEN). DISTRITO FEDERAL: Ca. 12 km E of Colonia Tovar on rd. to El Junquito, 2000 m, 10?26'N,67?08'W,29 Oct 1984, Knapp& Mallet 6865 (BH, K, MY, US, VEN); El Yagrumalon rd. to Colonia Tovar, 1900 m, 23 Sep 1936, Pittier 13721 (K, VEN); Agua Negra, 8 Mar 1940, Tamayo 1228 (US, VEN). SUCRE:Parque NacionalPeninsulade Paria,fromRio GrandeArribato guard post on La Pava trail, S. slope, 760-1050 m, 21 May 1994, Benitez de Rojas et al. 5122 (MO); Peninsula de Paria,between Manacaland Los Pocitos de Santa Isabel, 25 km NE of Irapa,Wof Cerro Humo, 700-900 m, 10?40'N,62?48'W, 9-10 Jul 1972, Dumont et al. VE-7526 (VEN); Cerro de Humo or Terr6n de Azucar, NE of Irapa, 21 Mar 1977, Fernindez 3144 (MY); Peninsula de Paria, Cerro Patao, N of Puerto Hierro, NE of Guiria, 850-860 m, 20 Jul 1962, Steyennark & Agostini 91158 (K, US, VEN); Peninsula de Paria, Cerro de Humo, slopes facing La Roma and Irapa, between summit and lake, NW of Irapa, 1060-1273 m, 3 Mar 1966, Steyermark94962 (US, VEN); Peninsulade Paria, Cerro de Humo, S-facing slopes between Laguna and La Roma, NW of Irapa,4 Mar 1966, Steyermark95002 (F, NY, US, VEN); Peninsulade Paria,Cuchilla de Cueva de Tigre, headwatersof Rio Nuevo, W of Cerro Humo, 750-850 m, 9-12 Aug 1966, Steyermark & Rabe 96410 (US, VEN); Distribution (Fig. 187). In the Coastal CordiPeninsula de Paria, above Las Melenas N of Rio Grande lleraof Venezuela,fromthe stateof Yaracuyto the Paria Arriba on trail to Tacarigua, 850-1000 m, 10?41-42'N, peninsula at 1000-1900 m. 62036-37'W,22 Feb 1980, Steyermarket al. 121564 (VEN); Sierrade Aroa, 1740 m, 18 Dec 1952,Aristeguieta YARACUY: Specimens examined. VENEZUELA.ARAGUA: ParqueNacional HenriPittier,trail from hotel to Pico & Foldats 1504d (VEN); Sierra de Aroa, Cerro Negro, 8 Periquito,28 Oct 1963,Agostini& Farinas 103 (VEN); km SW of San Felipe, 1200-1800 m, 10?17'N,69?01'W,1-

main lateralveins, these raised above, prominentand reddish-goldbelow in dry specimens, the apex longacuminate,the base acute, stronglydecurrentonto the petiole and then onto the stem; petioles narrowly winged, 0.8-3 cm long. Inflorescences opposite the leaves or intemodal,simple, 0.5-2 cm long, 2-5-flowered, glabrousor minutelypapillose, the papillae soon deciduous;pedicel scars closely spaced, but not overlapping,beginning near the base of the inflorescence. Buds translucentand fleshy, globose, the corolla soon exserted from the hyaline calyx tube. Pedicels at anthesis deflexed, translucentgreen, 0.8-1 cm long, taperingfromthe calyx tube to a slenderbase ca. 0.5 mm diam. Flowers with the calyx tube broadly and shallowly cup-shaped, 0.5-1 mm long, the lobes broadly deltoid, 0.254.5 mm long, hyaline in dry specimens, glabrous;corolla white or greenish-white, 0.8-1 cm diam., lobed 3/4 of the way to the base, the lobes reflexed at anthesis,the tips of the lobes minutelypapillose; anthers 1.5-3 x 1-1.5 mm, poricidal at the tips, the terminal0.1 mm palerandthickened,the poresteardrop shaped; ovary glabrous;style straight, 4-6 mm long in long-styled flowers, 1-1.5 mm long in shortstyled flowers; stigma small-capitate,minutely papillose, drying dark.Fruit a globose, green berry,0.8-1 cm diam.;fruitingpedicels elongate, slightly woody, deflexed, 2-3 cm long, ca. 0.75 mm diam. at the base; calyx lobes woody and somewhat accrescent in fruit, 3-4 mm long. Seeds pale tan, ovoid-reniform,ca. 4 x 2 mm, the surfacesdeeply pitted.Chromosomenumber: n = 12 (vouchersKnapp & Mallet 6842, 6847, 6857).

TAXONOMIC TREATMENT

337

FIG. 188. Solanum pastillum S. Knapp. (Reproduced with permission from Annals of the Missouri Botanical Garden 72: 559, fig. 1. 1985.)

2 Apr 1980, Liesner & Gonzalez 9850 (VEN); 7.5 km N of Knapp,Holbrook&Put 6063(holotype,BH;isotypes, Salom, 1200-1300 m, 10?15'N, 68?29'W, 4 Mar 1982, Figs. 171C, 188 CR, F, K, MO, NY, US). Liesner & Steyernark 12374 (VEN), Liesner & Steyermark Slender archingshrubs, 1-3 m tall; stems gla12392 (MO, NY, VEN); El Amparo to Candelaria,7 krnN of Salom, 1220-1250 m, 17-19 Jul 1972, Steyermark brous, green, horizontallyspreading,winged between 106203 (NY, US, VEN). the nodeswiththe decurrentleafbases;barkof the older

stems pale green, white-lenticellate.Sympodial units difoliate, geminate.Leaves elliptic-ovate, widest at or just proximal to the middle, dark green and slightly bullateabove, pale sea-greenbeneath,glabrous;major leaves 11.2-22 (25) x 3.4-6.6 (11.3) cm wide, with 68 pairs of main lateral veins, raised above, paler and prominentbeneath,the apex long-acuminate,the base decurrenton the petiole, attenuateto cuneate;petioles 0.5-1 cm long; minor leaves elliptic or occasionally orbicular,differing from the major ones in size and shape, 2.1-6 x 1.1-3.5 cm, the apex long-acuminate, the base attenuate(or cuneate);petioles 3-5 mm long. Inflorescences opposite the leaves, simple, glabrous, minutelywhite-lenticellate,1.7-7 cm translucent-green, long, slender and pendulous, 5-50-flowered;pedicel scars irregularlyspaced, 1-3 mm apart,beginning ca. 113. Solanum pastilium S. Knapp,Ann. MissouriBot. 1 cm from the base of the inflorescence.Buds conical, Gard.72: 558. 1985. Type.CostaRica. Puntarenas: the corollasoon exsertedfromthe globose calyx lobes, Campbell's Woods,1450m,20Aug 1982, glabrous. Pedicels at anthesis deflexed, 1.4-1.5 cm Monteverde,

Solanum ombrophilumis similar and closely relatedto S. pastillum of montaneCosta Rica, but differs from it in its unifoliatenodes, deltoid calyx lobes, andmore closely packedpedicel scars. The flowers of S. ombrophilumvary considerably in size depending on their sex, with male (short-styled) flowers being muchsmallerandhavingshorteranthersthanhermaphroditic flowers. Flowers of both types can be borne on the same plant, and flowers with intermediate style lengths occasionally occur. A given individual, however, is most often either short-styledor long-styled. Solanum ombrophilumis quite common where it occurs, and at the type locality is an importantcomponent of the forest understory.

338

long, taperingfromthe calyx tube to a slenderbase ca. 0.5 mndiam. Flowers with the calyx tube ca. 1.5 mm long, broadlycup-shaped,translucent-green,the lobes on fresh specimens consisting of five orbiculateprojectionsca. 0.5 mm diam. on the rim of the calyx tube, in dryspecimensof irregularlyshapedprojections0.51 mm long fromthe rimof the calyx tube,glabrousand translucentgreen;corolla greenish-white, 1.1-1.2 cm across, lobed ca. 3/4 of the way to the base, the lobes reflexed at anthesis, the tips of the lobes carinate-

FLORA NE-OTROPICA

Aug 1995, Grayumet al. 10959 (INB).

PUNTARENAS:

Sendero El Brillante, Monteverde Cloud Forest Reserve, 1450-1500 m, 13 May 1976, Dryer 81 (F); S of the rd., Monteverde Cloud Forest Reserve, 1520-1580 m, 16 Jun 1976, Dryer 208 (F); Monteverde, near the Ventana(Continental Divide), 1500-1600 m, 12 Jul 1976, Drver 426 (F); S of rd., Monteverde Cloud Forest Reserve, 15201580 m, 28 Aug 1976, Dryer 683 (F); S of rd. to Reserve, Monteverde Cloud Forest Reserve, 1520-1580 m. 28 Aug 1976, Dryer 684 (F); Monteverde Cloud Forest Reserve, 1/2 km from the station along the Sendero Rio, 1500 m, 10?25'N, 84?50'W, 19 Feb 1981, Kniapp 830) (BH); hoodled, minutely papillose; anthers 2.5-3.25 x 1.251.5 mm, poricidalat the tips, the poresteardropshaped; Monteverde Cloud Forest Reserve on Sendero Pantanoso free portionof the filaments ca. 0.5 mm long, the fila- near the Continental Divide, 1600 m, 10?25'N, 84?50W. 11 Mar 1981,Knapp840 (BH). SAN JOSE: Santiago de ment tube ca. 0.5 mm long; ovaiy glabrous;style 5-6 San Ram6n, 1150 m, 29 Jul 1937, Brenes 2.2615 (F); 5 mm long, straightor curved in dry specimens;stigma mi S of Santa Maria, 6800 ft, 5 Feb 1928, Stonrk1757 (F). capitate,ininutelypapillose.Fruit a globoseberry,greenSolanumpastillum is relatedto S. tuterck-heimii, ish-yellow at maturity, ca. 50-seeded, 1-1.5 cm diam., also of montanecentralCostaRica (butextendinginto usually only one or two from an inflorescence;fruiting pecdicels 1.9-2.6 cm long, woody, deflexed, 4-7 mm Nicaragua,Guatemala,and Mexico). The two species

share the understoryhabitat, long inflorescences of greenish-white flowers with reflexed petals at anthesis, and hard green fruits which become yellow and soft upon maturity. These two species and S. ombrophilumof montanecoastalVenezuelaareunusual in the S. confine species group in having ratherstout Distribution (Fig. 187). Foundonly in the cloud inflorescences and large flowers, but otherwise they forests of montane S Nicaraguaand Costa Rica from conforn to thecharactersof therestof the speciesgroup, (75--)1000 to 1700 m. particularlyin theirseed and leaf morphology. Solanumpastillumnblooms at the beginning of Specimens examined. NICARAGUA. ZELAYA: Cerro Saslaya. 20 km W of Siuna, 1100-1400 m, 5 May the wet season at Monteverde de Puntarenas,Costa 1977, Neill 1881 (MO). 3 May 1978, Neill 3830 (MO); Rica, the type locality, and is pollinated primarilyby Cerro la Pimienta #1, summit, ca. 900-980 m, 13?45'N, bumblebees, BombusephippiatusSay (Knapp,I986a). 84?79'W, 13 Apr 1979, Pipolr' 5136 (MO). The local distributionpatternof S. pastillliianis typical COSTA RICA. ALAJUELA: Rerserva Biol6gica of many of the forest understory species of sect. Monteverde, Cord. de Tilaran, valley of Rio Petias Geminata. Small clumps of three to five individuals Blancas, 1600 m, 10?19'15"N, 84?46'30"W,22 Jul 1993, Bello 5154 (INB); Llanura de San Carlos, 17 km E of grow at widely spaced intervalsin the understory,ofFortuna, flac. Platanar, 75 m, 10?27'N, 84?30'W, 9 Dec ten at the edges of old gaps caused by tree or branch & Brad/o4d 11574 (INB); Llanura de San falls. The inflorescencesofS. pastilluniaremany-flow1993, Habe/w Carlos, 100 m, 21 Feb 1966, Molina R. et al. 17659 (F); ered, but only a few fruits per inflorescence develop. Pefias Blancas trail, 3.5-5 km ESE of Monteverde, ca. Whetherthis is due to poor pollination success or to 145(1 m. 17 Aug 1976, Kennedi' & Guiindon 3713 (F); fruit abortionis not known. The presence of a single Monteverde, on rd. to Pefias Blancas (over Continental fruiton a long inflorescencecausesthe effective pedicel Divide) ca. 1/2 km below the Divide on Atlantic slope, lengthof the fruitto be extremelylong. This may make 1450 in, 1(0)25'N, 84?50'W, 13 Apr 1981, Knapp& Mallet the fruitmore visible to small frugivorousbats, known 864 (13H);Alfaro Ruiz, Guadelupe de Zarcero, 1550 m, to take the fruitsof this and at least threeotherspecies 1 Jul 1938, Sinith NY810 (F, NY); Alfaro Ruiz, La Pefia of sect. Gerniiata in Monteverde(Dinerstein, 1983). de Zarcero, 1400 m. 11 Jul 1938, Smith NY904 (F, NY);

diam. at the apex, the base slender, ca. 0.5 mm diam. Seecls pale brown or tan, ovoid-reniform,3.5-4 x 22.5 mm,the surfacesminutelypitted,the pits very shallow, the seeds appearingsmooth. Chromosomenumber: n = 12 (vouchersKnapps.n., Knappet al. 6063).

Alfaro Ruiz, Zapate, 1425 m, 10 May 1940, SmithP2663 (F); Alfaro Ruiz, 5 mi S of Zarcero, 1500 m, 17 Jun 1941, Smiiith2789 (F). CARTAGO: El Mufieco, 1300 m, 9?44'N, & GraoYum 16599 (INB); 83?49'W, 15 Nov 1987, HamnmuFel El Mufieco on Rio Navarro, 1400-1500 m, 6-7 Mar 1926, Standley & TonresR. 51016 (US), Standlev & Torres R. 51040 (US). LIMON: Cord. de Talamanca,headwaters of QuebradaKak6beta,below divide between Rio Xikiari and Rio Boyei, 900-1000 m, 9?47'N, 83?20'30"W, 14

114. Solanum pertenue Standl.& C. V. Morton,Publ. FieldMus.Nat.Hist.,Bot. Ser. 18: 1089. 1938.Type. Costa Rica. San Jose: SantaMariade Dota, 15001800 m, Dec 1925, Standley 42852 (holotype, F; isotype, US [F neg. s.n.]). Fig. 189 Shrubs1-2 m tall;youngstemsandleaves dense-

TAXONOMIC

TREATMENT

33 9

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340

ly puberulentwith minute uniseriatetrichomes 0.050.2 mm long, these often soon deciduous;older stems glabrous,lightly winged fromthe decurrentleaf bases; barkgrayish-red,not lenticellate; branchesspreading horizontally,giving the shrubsa flattenedaspect.Sympodial units unifoliate.Leaves elliptic to narrowlyelliptic, not geminate, except occasionally on the lower stems, widest at the middle, glabrousabove, glabrous or densely pubescentalong the veins beneath,with 57 pairsof mainlateralveins, these slightlyraisedabove, prominentandyellow on leaf undersides;lamina7-13 x 2-4.5 cm, the apex acuteto acuminate,the base acute, decurrentonto the petiole;petioles 2-5 mm long.Inflorescences opposite the leaves, simple, 0-5 mm long, 1-3-flowered, glabrous or minutely puberulentwith erectuniseriatetrichomeslike those of the stems.Buds globose when young, the corolla soon exserted from the calyx tube and then the buds obovoid, flattenedat the apex, glabrousor minutelypuberulent.Pedicels at anthesis deflexed, 1.1-1.4 cm long, filiform, tapering gently fromthe calyx tube to a base ca. 0.25 mm diam. Flowers with the calyx tube narrowly conical, 0.5-1 mm long, the lobes deltoid, glabrous,ca. 0.5 mm long, glabrous or minutely and sparsely puberulent with uniseriate trichomes ca. 0.1 mm long; corolla white, 0.7--l cm diam., lobed 3/4 of the way to the base, the lobes stronglyreflexed at anthesis,the tips of the lobes papillose;anthers2-2.5 x ca. 1 mm,poricidalatthetips, the pores teardropshaped;free portionof thefilaments 0.25-0.5 mm long, the filamenttube0.1-0.5 mm long; ovary glabrous;style straight, 5-6 mm long; stigma bilobed,minutelypapillose.Fruita globoseberry,green or yellowish-green when ripe, 1-1.5 cm diam.;fruitingpedicels deflexed, woody, slightly expandedat the apex, 2-2.2 cm long, ca. 0.75 mm diam. at the base. Seeds pale tan, darkin dry material, ovoid-reniform, 2.5-3 x 1.1.5 mm, the surfacesminutelypitted.Chromosome nunmber not known. Distribution (Fig. 192). In montaneNicaragua, Costa Rica, and Panama,from 700-2000 m, in understory of primarypremontaneor montanecloud forest. Specimens examined. NICARAGUA. ZELAYA: La Posolera,5 km W of Waslala,El Tuma-Waslalard., ca. 700 m. 13017'N, 85?24'W, 22 Dec 1982, Moreno19118

FLORA NEOTROPICA Quebrada Cuecha, Monteverde, Cordillera de Tilaran, 1540-1600 m, 2 Sep 1976, Dryer 729 (CR, F), 18 Mar 1977, Dryer 1229 (CR, F); Monteverde Cloud Forest Reserve, 0.5 km from station along Sendero Rio, 1500 m, 10?25'N, 84?50'W, 19 Feb 1981, Knapp 828 (BH, CR); Monteverde Cloud Forest Reserve along Sendero Pantanoso near the Continental Divide, 1600 m, 10?25'N, 84?50'W, 11 Mar 1981, Knapp 841 (BH, CR); Monteverde Cloud Forest Reserve, along Sendero Rio, near Rio Guacimal, 1500 m, 10?25'N, 84'50'W, 30 Apr 1981. Knapp876 (BH, CR); R.B. Monteverde. Rancho Alegre trail, 1000 m, 10?17'N, 84?45'W, 4 Apr 1995, Martinez 391 (INB). SAN JoSE: Forests of Rancho Flores, 2043 m, 22 Feb 1890, Tonduz 2094 (US). PANAMA.

CHIRIQUi: Forest behind Vivero For-

estal, 9 mi N of Planes de Hornito, IHRE FortuinaHydroelectric Project, ca. 1000 m, 8?45'N, 82?121W, 13 Mar 1982, Knapp et al. 4098 (MO).

Solanumpertenueis related to S. confine of eastem Peru, with which it shares short, golden urniseriate trichomes,white flowers, andoccasionally foetid foliage. It differsfromthatspecies in its tiny inflorescences andmore elongate fruitingpedicels. The inflorescence in S.pertenueis reducedandin the type specimenbears only a single flower.Most plants of S. pertenuehave 2 or 3 buds in each inflorescence, but in Monteverde, Costa Rica, the thirdof these usually does not open. If one of the first two buds is damaged or removed, the thirdbud opens and produces a flower. In Monteverde,Costa Rica, where I stucliedthe reproductiveecology of thisandsix otherspeciesof sect. Geminata(Knapp,1986a),S. pertenueis a shrubof the edges of treefall gaps, and grows under closed or nearly

closed canopyforest.The flowersarepollinatedlargely by workersof Bombusephippiatus,but small halictid bees also visit the flowers (see TableIX). The leaves of S. pertenueare fed upon by the larvaeof the ithomiine butterflyPteronymiaartenna(see TableXI).

115. Solanum stipulatum Vell., Fl. Flumin. 87. 1829 [1825]. Type.Brazil.Rio de Janeiro:"Habitatcampis apricismediterraneis"(No specimensextant;lectotype, Vellozo, Fl. Flumin. Icones 2: fig. 117. 1831 [1827], here designated). Figs. 171E,F, 190, 191

(MO).

Solanum riyulare

COSTA RICA. ALAJUELA: Bijagua, El Pil6n, headwatersof Rio Celeste, 700 m, 10?49'N,84?27'W,

10: 25. 1846. Type. Brazil, Rio de Janeiro, in ripa rivuli prope Mandioccam ad Montes dos Orgaos, locis sylvis umbrosis, a Septembri as Octobrem florens, Martius s.n. (lectotype, M, designated by Smith & Downs, 1964 [Morton neg. 8734]). SolanumnundulatlumDunal in DC., Prodr. 13(1):138. 1852. Type. Brazil. in Brasilia, LhotsAivsnii. (holotype, G-DC [F neg. 6781]; isotype, G, MPU). Solanumnapodum Dunal in DC., Prodr. 13(1): 139.

16 Nov 1987, Herrera 1307 (INB). LIMON: Cord. de Talamanca,N flank of Fila de Matama, in headwaters of Rio Boyei, 1200-1300 m, 9?45'00"N, 83?19'00"W, 18

Aug 1995, Grayum11072(INB). PUNTARENAS: Parque Internacional La Amistad, Cord. de Talamanca, Est. Pittier, trail to Cerro Gemelo, 1680 m, 9001 '30"N, 82?57'40"W, 30 Jan 1995, Alvarado 21 (INB); above

Mart. ex Sendtn. in Mart., Fl. Bras.

TAXONOMIC

TREATMENT

341

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FIG. 190. Solanum stipulatuim Vell., plate 117 of Vellozo's

1852. Type. Brazil. Rio de Janeiro: S.loc., Sellow 133 (lectotype, BM, here designated).

Flor-ae Fluminensis

lcones,

Vol. 2.

Pedicels at anthesis ca. 1 cm long, very slender,tapering from the calyx tube to a slender base ca. 0.5 mm Subshrubs or shrubs 0.5-2 m tall; stems gla- diam.Flowers with the calyx tube ca. 1.5 mm long, the brous,corky,pale and lenticellate; intemodes short, 1 lobes short-triangularto deltoid, ca. 1 mm long, glacm or less, strongly winged from the bases of the de- brous;corolla white, ca. 1 cm across,lobed ca. 1/2way currentleaves, these wings not continuous,but begin- to the base, the lobes planarto somewhat deflexed at ning anew each leaf Sympodialunits difoliate, gemi- anthesis, the tips of the lobes minutely papillose; annate. Leaves obovate to oblanceolate to almost linear, thers ca. 2.5 x 0.5 mm, poricidal at the tips, the pores quite variable in shape, widest in the distal 1/3 of the teardropshaped;free portionof thefilamentsca. 1 mm blade, glabrous;major leaves 11-20.5 x 1.7-5.5 cm, long, the filamenttube ca. 1 mm long; ovary glabrous; with 7-10 pairs of main lateral veins, raised but not style ca. 5 mm long, straight;stigma slightly clavate, prominentboth above and beneath, the apex acute to not markedlylargerthanthe style, minutelypapillose. acuminate,the base attenuate,taperingto anddecurrent Fruit a turbinate,greenberry,6-10 mm diam.,palerat on the stem;petioles indistinct;minorleaves oblong or the distal end, often very pointed;fruitingpedicelsdeorbiculate, 2.5-3.2 x 1.1-2.5 cm, the apex rounded, flexed, 1.2-1.4 cm long, not markedlywoody, 0.5 mm the base attenuate,taperingto the stem;petioles indis- diam.atthebase.Seedsgreenwhen fresh,paletanwhen tinct. Inflorescences opposite the leaves, filiform, dry,ovoid-reniform,ca. 10 per berry,3-4 x 2-2.5 mm. simple, 1-2 cm long, 3-8-flowered, bearing flowers Chromosomenumbernot known. only in the distal third,glabrous;pedicel scars evenly Distribution (Fig. 192). SE Brazil, in coastal spaced 2-3 mm apart,slightly corky.Buds obovoid to rainforestoftengrowingin rockycreekbeds, from 100fusiform, the corolla soon exserted from the calyx. 700m.

FLORA NEOTROPICA

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343

TAXONOMIC TREATMENT

5O0

(solid circles), S. stipulatum (solid squares), S. per-tenue FIG. 192. Distribution of Solanulm tenuiflagellatum (bicolored circle). (solid triangles), and S. vanamonense (open circles), S. valeiianum

Specimens examined. BRAZIL. S.loc., Claiussen s.n1. (P). PARANA:Morretes, Usina Eletrica Marumbi, 4

Jan 1966, Hatschbach et al. 13435 (NY, UPCB, U, US); Campina Grande do Sul, Rio Pardinho, 21 Nov 1965, Hatschbach 13163 (US); Morretes, Cadeado, 30 Nov 1966, flatschbach 15316 (US); Piraquara,Veu de Noiva, 850 m, I Dec 1970, Hatschbach 25718 (NY); Guaraquecabas, Serra Negra, 100 m, 10 Dec 1970, Hatschbach 25798 (BH). RIo DE JANIERO: Parque Nacional da Tijuca, near Vista Chinesa, 125 m, 23 Jun 1995, Bovini, Knapp & Marqluete813 (BM, RB); horto, perto do IMPA, Bovini, Knapp & Marquete 814 (BM, RB); s.loc., 24 Jan 1932, Br-ade 11315 (US); Serra dos Orgaos, valley near Fazenda Bonfim, ca. 1600 m, 21 Jul 1971, Lindemnan & Bar-cia 6391 (U); Terez6polis, Parque Nacional Serra dos Orgaos, Rio Beijaflor, 1100 m, 20 Oct 1977, Maas & Ma,tinelli 3354 (F, NY); Terezopolis, Parque Nacional Serra dos Orgaos, along Rio Beijaflor, 1000 m, 2 Feb 1983, Mar-tinelli& Simoanis9063 (U); Mun. Novo Friburgo, Macae de Cima, Fazenda Sophronites,

1200 rn, 17 Jul 1987, Pessoa et al. 211 (BM, RB); Serra dos Orgaos, 30 Nov 1943, Pereira 236 (US); Serra dos Orgaos, 14 Oct 1962, Per-eira 7212 (F, US); Mandiocca, Pohl s.n. (BR); Rio Mandiocca, Feb 1832, Riedel 403 (NY, US); s.loc., Oct 1932, Riedel 1076 (NY, US); Petr6polis, Rocio, ca. 700 m, 16 Mar 1968, Sucr-e & Braga 2470 (BM, RB); Mana, 1897, Ule s.n. (US). SANTACATARINA: Itajai, Braco Joaquim, Luis Alves, 300 m, 20 Aug 1954, Reitz & Klein 2067 (US), 4 Nov 1954, Reitz & Klein 2234 (US); Sao Franciscodo Sul, Tres Barras,Garuva,250 m, 7 PicinNov 1957, Reitz & Kleini5636 (US). SAO PAUILO: guaba, BR 101, near a waterfall, 2 Oct 1975, Araujo et al. 838 (BM, RB). Local names. Brazil. Santa Catarina: canema minm. Solanum stipulatum is morphologically most similar to S. leptopodum of Amazonia, sharing with that species difoliate, geminate sympodial units with markedly smaller minor leaves. It is convergent with

344

other species of sect. Geminata growing in river courses (see S. amnicola, S. imberbe, and S. monadelphum)but this similarleaf shape is relatedto habitat,not to any realindicationof relationshipamong these species. Solanumstipulatumis an easily recognized species, with its short internodes, striking oblong-lanceolate leaves, and white bark. Sendtner,in his treatmentof SolanumforFlora Brasiliensis (1846), chose to ignore the Vellozo name, because he felt the plate in the Florae Fluminensis (Vellozo, 1831)was a poorone ("iconmala").Theplate is stylized, as areall the plates in Vellozo'sFlorae Fluminensis, but it clearlyrepresentsthe sametaxonas that namedby Sendtner.The name was actually proposed by Martius,andperhapsSendtnerinventeda reasonto ignorethe Vellozo name, so as not to offend his patron. The Vellozo plate shows a plant with winged stems withshortintemodesandoblanceolateto obovateleaves, bothcharacteristic featuresofS. stipulatum,buttheleaves are not geminate, as they always are in S. stipulatum. The inflorescencesof the plantin the platearedepicted as axillaryandone-flowered,whilethoseofS. stipulatum are leaf-opposed and 3-8-flowered. In describingS. aipodulm,Dunalcited a Bowie andCunnninghamspecimen in "herb.Banks,"now housed at BM. There are no sheets of S. stipulatum collected by Bowie and Cunninghamat BM, but a sheet collected by Sellow (see above) has a label in Dunal's hand, indicatinghe saw thismaterial.ThedescriptionofS. apodummatches this sheet, and so is here designatedas the lectotype. Severalvarietalandsubspecificepithetswerenoted on specimensby Bitter(in sched.)butneverpublished. These in general were based on the extremes of leaf morphology in the species, which is quite striking.

116. Solanum tenuiflagellatum Bitter ex S. Knapp, Brittonia38: 286. 1986. Type. Venezuela.Aragua/ Distrito Federal: Prope Colonia Tovar, 1854-55, Fendler- 977 (holotype, G [Morton neg. 8560]; isotypes, K, MO, NY, P). Figs. 171D, 193

FLORA NEOTROPICA

winged from the decurrentleaf base; minor leaves, if present,not differingfromthe majorones except occasionally in size. Inflorescences opposite the leaves or slightlyintemodal,very long andslender,3--30cm long, occasionally irregularlybranched,only bearinga few flowers at a time, but with 10-100 scars of previously bome flowers, sparsely pubescentwith uniseriatetrichomes like those of the leaves andstems;pedicelscars raised,widely andunevenly spacedalong the inflorescence, 0.5-1 cm apart.Buds globose, the corolla soon exsertedfromthe calyx lobes, the calyx lobes reflexed. Pedicels at anthesisdeflexed, 1-1.2 cm long, filifonrn. taperingfromthe calyx tube to a slenderbase ca. 0.25 mm diam.,sparselypubescentwith uniseriatetrichomes ca. 0.5 mm long. Flowers with the calvx tube short obconic, ca. 1 mm long, the lobes long-triangular,0.51 mm long, denselypubescentwith uniseriatetrichomes like those of the rest of the inflorescence,the trichomes denser on the tube than on the lobes; corollaC white, 0.7-

1 cm diam., lobed ca. 3/4 of the way to the base, the lobes planarat anthesis,or very slightly reflexed at the tips, the tips of the lobes minutely papillose; anthers ca. 2 x 0.75-1 mm long, poricidalat the tips, the pores teardropshaped;free portionof thefilamients 0.5-0.7 mm long, the filament tube extremely short, less than 0.1 mm; ovary glabrous;style straight,ca. 4 mm long, dryingblack;stigma clavate,minutelypapillose.Fruiiit a globose, greenberry,ca. 1 cm diam.;fruitingpedicels greatlyelongated,deflexed, causing the inflorescence to be pulleddown straight,3-3.5 cm long, ca. 0.75 mm diam.at the base, expandedat the apex, ca. 3 mm diam. Seeds pale tan, ovoid-reniform,ca. 3 x 2 mm, the sur: = 12 faces minutely pitted. Chromosomenumber. (voucherKnapp & Mallet 6859). Distribution (Fig. 192). In the Cordillerade la Costa of Venezuela, from 1000 to 1750 m. Specimens examined. VENEZUELA. S.loc.. 1856-57, Fenidler 2608 (K). ARAGUA: Colonia Tovar. May, Moritz 1645 (B, K, P, W). DISTRITO FEDERAL: Ca. 4 km NW of jct. with El Junquito-Colonia Tovar rd. on rd. to Carayaca (jct. is ca. 16 km E of Colonia Tovar), N slope Cordillera de la Costa, ca. 1750 m, 10?26'N, 67?08'W, Kniapp& Mallet 6859, 6861 (BH, K, MY, US, VEN); Parque Nacional El Avila, N of Boca de Tigre on N facing slopes of Cordillera de la Costa, 1650-1800 m, 10?35'N, 66?55'W, 31 Oct 1984, Kniapp & M,allet 6870 (BH, MO, MY, US, VEN); Topo Tacahuamaco (Tamanaco),quebradaRio Grande,(9 km E of los Tanques de Electricidadde Caracas), S of CamuriGratnde,N slope Cordillerade la Costa, 900-1000 m, 16 Jul 1973. .Vfor-illo et al. 3303 (MY, VEN).

Shrubs or small trees, 1-5 m tall; young stems andleavesdenselypubescentwithuniseriatecurlytipped trichomes 0.5-0.8 mm long; older stems persistently pubescent;barkof older stems gray.Sympodialunits unifoliate, occasionally difoliate and geminate on the lower stems. Leaves elliptic, widest at the middle, sparselyto denselypubescentwith uniseriatetrichomes 0.5-0.8 mm long along the veins beneath;majorleaves 4-17 x 2--9cm, varyinggreatlyin size on a single stem, with 5-7 pairs of main lateralveins, the midribraised Solaniumtenuiflagellatumis most closely related above, the apex acuteto acuminate,the base acute,de- to S. leptorhachis of western Ecuador, sharing with currenton the petiole;petiole 0.4-1.5 cm long, slightly thatspecies an extremelyelongateinflorescence,flow-

TAXONOMIC

TREATMENT

34 5

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ers with the petals held planarat anthesis,andpedicels 117. Solanum tuerckheimii Greenm. in Donn. Sm., Bot. Gaz. 37: 212. 1904. Type. Guatemala. Alta that in fruitare nearly four times their length at antheVerapaz: Cubilqiiitz, 350 m, Jul 1903, von sis. Solanum tenuiflagellatumis easily distinguished Tuerckheim 8492 (holotype, US). Fig. 194 fromS. leptorhachisby its muchlongerinflorescences, Solanum tenuilamellosum Nov. Bitter, Repert. Spec. copious leaf pubescence,acuminatecalyx lobes which Regni Veg. 18: 57. 1922. Type. Costa Rica. arerecurvedat anthesis, and greenish flowers. The inAlajuela: Collines de Piedades pres San Ram6n, florescences of S. tenuiflagellatumare the longest in ilOOm, 4 Jun 1901, Brenes 14384 (holotype, the S. confine species group, and also in the section. B-n.v. [F neg. 2860]). They are threadlikeand often become tangled in the leaves of the branches below them. Solanum Shrubs, 1-3 m tall; young stems and leaves glatenufilagellatumhas only been collected a few times brous or minutely papillose; bark of older stems reddespiteits strikinginflorescencemorphology.Whereit dish and peeling in longitudinal strips, the strips reddoes occur, often several plants are found in the area, dish and translucent. Sympodial units difoliate, but it seems to have a ratherpatchydistributionover its geminate.Leaveselliptic,widest at themiddle,glabrous on both surfaces, paler beneath in live plants, becomgeographicalrange.

FLORA NEOTROPICA

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TAXONOMIC TREATMENT

ing golden crystalline when dry, shiny above; major leaves 9-17 x 3-6 cm, with 5-9 pairs of main lateral veins, these raised above, obscure below, the apex roundedor acute, the base acute, decurrenton the petiole; petiole ca. 1 cm long; minor leaves differing from the major ones only in size, 2-5 x 1-3 cm, the apex roundedor acute,the base acute;petioles 5-8 mm long. Iniflorescencesopposite the leaves, simple, 1-3 cm long. 5-15-flowered, completely glabrous;pedicel scars evenly spaced ca. 1.5 mm apart,beginning 0.51 cm fromthe base of the inflorescence.Buds globose, the corolla soon exserted from the calyx. Pedicels at atithesiswhite, 0.6-1 cm long, deflexed, taperingfrom the calyx tube to a slenderbase ca. 0.5 mm diam.,completely glabrous.Flowerswith the calyxtube0.5-1 mm long, glabrous,the lobes deltoid, 0.5-1 mm long, minutely papillose on the tips, both the tube and lobes white in living plants; corolla white, 7-9 mm diam., lobed nearlyto the base, the lobes reflexed at anthesis, the tips of the lobes minutelypapillose;anthers2-3 x 0.75-1 mm, poricidal at the tips, the pores teardrop shaped;free portionof thefilaments ca. 0.5 mm long, the filament tube less than 0.25 mm long; ovary glabrous; stvle straight, ca. 3 mm long; stigma bilobed, minutelypapillose. Fruit a globose, green berry,ca. 1 cm diam.;fruitingpedicels woody, deflexed, 1.9-2.2 cm long, ca. 0.75 mm diam. at the base; calyx lobes persistentandwoody in fruit.Seeds yellowish, ovoidreniform,ca. 3 x 2.2 mm, the surfacesminutelypitted, the marginsslightly incrassate.Chromosomenumber: n - 12 (voucher Knapp 872). Distribution (Fig. 197). In the understory of premontanecloud forest from Mexico to Costa Rica, from 100 to 1500 m.

85056'W, Stevens 9163 (MO). MATAGALPA: Cordillera Darienense between Santa Lastonia and Disparate de Poter, 1300-1500 m, 25 May 1974, Molina R. & Molina 30478 (F, MO); El Picacho E of Santa Maria, Cordillera Darienense, 1600 m, 26 May 1974, Molina R. & Molina 30540 (F, MO); Fuente Pura, Matagalpa-Jintega rd., 1400-1480 m, 13?00'N, 85?55'W, 1 Dec 1982, Morenio 18919 (MO). COSTA

RICA.

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San

Ram6n, Apr 1902, Brenes s.n. (NY); Cuenca de Tarcoles, San Juan, raod to Los Angeles, 1100 m, 10?06'N, 84?28'W, 25 May 1997, Rodriguez 2210 (INB). GUANACASTE:Upper slopes of Cerro San Jose de Libano. 500-960 m, Dodge et al. 6380 (MO); Parque Nacional Rinc6n de la Vieja, Hac. Sta. Maria, from Mirador SE to Volcan Sta. Maria, 850-950 m, 10?48'N, 85'19'W, 11 Aug 1987, Herrera 705 (INB); P.N. Guanacaste, Estaci6n Mengo, 1100 m, 10?55'-56'N, 85?28-29'W, 15 Jul 1989, INBio II 196 (INB); P.N. Guanacaste, Cord. Guanacaste, Est. Cacao, 1100 m, 10?55'38"N, 85?29'38"W, 3 Jan 1990, Maass 9 (INB); Parque Nacional Rinc6n de la Vieja, track from Estaci6n Biol6gica Santa Maria to Poilas, 880 m, 10?46'N, 85017'W, 19 Jun 1996, Short & Stafford 139 (BM, INB). PUNTARENAS:E of Monteverde, 1300-1450 m, 10?18'N, 84?48'W, 29 Oct-2 Nov 1975. Burger & Baker 9580 (F); Monteverde, 1400-1500 m. 15 Aug 1976, Dryer 604 (F), 15 Jun 1977, Dry'er 1452 (F); Monteverde, in village near Rio Guacimal, 1300 m, 10?25'N, 84?50'W, 29 Apr 1981, Knapp 872 (BH); lower Monteverde community, Quebrada Guacimal, 1300-1400 m, 23 May 1977. Lawton 1183 (F). SAN JosE: Hills of San Pedro de San Ram6n, 1050-1075 m, 19 Jul 1925, Brenies 4268 (F); Piedades de San Ramon, 900 m, 21 Jul 1925, Brenes 4271[22021] (CR, F), 4 Aug 1925, Brenes 4346 (F). Solanum tuerckheimii is related to S. pastillum.,

also of montaneCostaRica. It is distinctfromthatspe.cies in its rough, shaggy, red bark,its smaller flowers Selected specimens examined. MEXICO. and deltoid calyx lobes, andby the golden cast its leaf VERACRUZ: Hidalgotitlan,km 9-10 on Hnos. Cedillo- undersides take on when dry. In Monteverde, Costa PanchoVillard., 120m, 17?00'N,94?00'W,13 Aug 1974, Rica, S. tuerckheimiiis a shrub of forest understory, Doiantes3482(MO);Hnos.Cedillo-A.Melgarrd., 150m, occurringin drierforest thanmost of the otherspecies 17016'N,94036'W,24 Aug 1974, Vasquez1006 (BH, F). at the site (see Knapp,1986afor descriptionof the site). BELIZE. CAYO: Chiquibul,San Pastor,540 m, The flowers of S. tuerckheimiiin Monteverdeare pol16043'N,88059'W,3 Sep 1997,Mlonro1860 (BRH,BM, linated by Bombus ephippiatus,but the plants do not MO, MEXU, EAP,NY). set large numbersof fruit. Visits by Bonmbusare rare. GUATEMALA. ALTA VERAPAZ: Chahal3 km NWN of airport,200 m, 5 Oct 1968, Contreras 7829 (F); Chahal,3 km fromEl Mayo,7 km W of airport,14 Oct 1968, Contreras7923 (F); Choma,900 ft, 1 Aug 118. Solanum valerianum C. V. Morton & Standl., Publ. Field Mus. Nat. Hist., Bot. Ser. 18: 1097. 1920, Johnson467 (F, US). HUEHUETENANGO: Vic. of 1938. Type. Costa Rica. Puntarenas:Playa Blanca, Ixcan, Sierra de los Cuchumatanes, 200 m, 24 Jul 1942, Golfo Dulce, Feb 1933, Valerio424 (holotype, F; 49436 (F). Ste-vermark NICARAGUA. JINOTEGA: Matagalpa-Jinotega isotype, CR). Fig. 195 rd., km 16, 1200-t400 m, 25 May 1980, Moreno 568 (MO); region of Las Mercedes, sierra E of Jinotega, 1200-1500 m, 3 Jul 1947, Standle y 10687 (F, NY), Siandle-v 10736 (F, US); along Hwy. 3 ca. 1.9 km NW of Aranjuez r(d. entrance, 1460-1480 m, 13?02'N,

Shrubs1-2 m tall;young stemsandleaves dlensely long-pubescentwith uniseriatetrichomes0.5- 1 mmn long, these straw-coloredand curly-tipped;the stems strongly winged from the decurrent leaf bases; older

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TAXONOMIC TREATMENT

stems remaining pubescent, the bark grayish-green. Svmpodial units unifoliate, not geminate, except on non-reproductivenodes.Leavesnarrowlyelliptic,widest at the middle, glabrous above, glabrous or longpubescent along the veins and occasionally on the laminabeneath,thetrichomes0.5-1 mm long,theblades 9-18 x 2.5-4 cm, with 7-8 pairs of main lateralveins, these not strongly converging, not raised above, pale yellow andprominentbeneath,the apex acute,the base attenuate,winged onto the petiole; petioles 0.3-1 cm long, winged from the decurrentleaf bases. Inflorescences oppositethe leaves, simple, 5-7 mm long, 3-6flowered, glabrousor sparsely pubescent with strawcoloredcurly-tippedtrichomes;pedicelscars widely and evenly spacedca. 1 mm apart,slightlyraised.Buds globose when young, laterellipsoidto obovoid,the corolla soon exsertedfromthe calyx tube.Pedicels at anthesis ca. 8 mm long, filiform,taperingfromthe calyx tubeto a slenderbase ca. 0.25 mm diam.Flowerswith thecalyx tube broadly cup-shaped,ca. 0.5 mm long, the lobes deltoidto triangular,0.25-1.5 mm long, sparselypubescent with uniseriatetrichomeslike the rest of the plant; corolla white (?), 0.8-1 cm diam., lobed nearlyto the base, the lobes perhapsreflexedat anthesis,the tips and marginsof the lobes minutely papillose;anthers ca. 2 1 mm, poricidalat the tips, the poresteardropshaped; free portionof thefilaments ca. 0.25 mm, the filament tubeminuteandveryshort;ovaryglabrous;stylestraight, ca. 3 mm, glabrous;stigma capitate.Fruit a globose, greenberry,1-1.5 cm diam.;fruitingpedicelsdeflexed, somewhatwoody, 1.8-2 cm long, 0.5-0.75 mm diam. at the base. Seeds darkbrown in dry material,ovoidreniform, 3-3.5 x 2-2.5 mm, the surfaces minutely pitted. Chromosomenumbernot known.

349 PANAMA. CHIRIQUi: Along rd. from Puerto Armuelles to San Bartolo Limite, ca. 7 mi W of Puerto Armuelles, 120 m, 19 May 1976, Croat 35078 (MO, NY).

Solanumvalerianumis relatedto S. capillipesof Trinidadand Venezuela.Solanum valerianumdiffers fromS. capillipesin its denser,longerstempubescence, smallerinflorescences,andin its leaf venation.The main lateralveins of S. valerianumdo not prominentlyconverge on one another(i.e., not markedlybrochidodro-mous sensu Hickey, 1974) as do those of S. capillipes The type specimen of'S. valerianum is in very poor con-

dition, with only one damaged flower on the collection. Collections from the Carararegion in western Costa Rica have shortertrichomesthanthe type.

119. Solanum yanamonense S. Knapp,Brittonia38: 298. 1986. Type.Peru.Loreto:Maynas,Explorama TouristCamp on Rio Amazonas between Indiana andmouthof the Rio Napo, ca. 80 km N of Iquitos, non-inundatedtropicalrainforest,ca. 100m, 3?28'S, 72?48'W,23-27 Jul 1984, Knapp 6613 (holotype, BH; isotypes, NY, US, USM). Fig. 196

Small shrubs, 0.3-1 m tall; young stems and leaves densely andminutelypuberulentwith uniseriate trichomesca. 0.05-0.1 mm long; the stems not becorning glabrate,not winged; barkof older stems grayishred;branchesarchedand flattened,giving the shruba planaraspect.Sympodialunitsunifoliate,not geminate except on non-reproductivenodes. Leaves narrowly elliptic to narrowlyovate, widest at orjust proximalto the middle, glabrousandshiny above, densely pubeiulent along the veins beneathwith trichomeslike those of the young stems,the leaves 5.5-9 x 1.3-2.5 cm, with 8-9 pairs of main lateral veins, these converging at Distribution (Fig. 192). In forest and second about the halfway point of the lamina, slightly raised growth in Nicaragua, W Costa Rica, and adjacent above,prominentbeneath,the apexlong-acuminate,the Panama,from sea level to 200 m. extreme tip truncateand rounded,the base truncate; petioles minute, 1-2 mm long, winged from the leaf Specimens examined. NICARAGUA. CHONTALES: base. Inflorescences opposite the leaves, threadlike, S.loc., 1867-1868, Tate 275 [181] (BM, K). ZELAYA: El Escobillo, 2 km from Colonia Serrano, rd. to Yolaina, simple, 1-1.5 cm long, 4-6-flowered, but only one floweropen at a time, denselypuberulentwith the same 80-100 m, 11034-35'24'W, 29 Jul 1982, Sandino 3321 (MO). golden, uniseriatetrichomesas the stems, ca. 0.05 inm COSTA RICA. PUNTARENAS: Parque Nacional long;pedicel scars in pairs, the membersof a pair ca. Corcovado, Peninsula de Osa, Est. Los Patos, near Rio 0.5 mm apart,the pairs ca. 2 mm apart.Buds globose Rinc6n, 200 m, 8?34'N, 83031'W, 27 Dec 1993, Aguilar when young, the corolla soon exsertedfromthe calyx, et al. 2867 (INB); Reserva Biol6gica Carara, Quebrada the buds laterobovoid.Pedicels at anthesisthreadlike, Monas, 50 m, 9?48'N, 84?35'W, 8 Jan 1990, Zuiniga48 deflexed, 6-7 mm long, taperingfrom the calyx to a (INB). SAN JOSE: CararaNational Park, near Rio Carara, filiform base ca. 0.1 mm diam.Flowers with the calyx 120 m, 9?46'N, 84?32'W, 1 Apr 1993, Gentr-y et al. tube broadlyconical,ca. 0.5 mm long, the lobes minute, 79264 (INB): Z.P. La Cangreja, coastal slopes of Valle broadly deltoid, often only tiny projectionsof the tube, de Parvita, Mastatal de Puriscal, 200 m, 9?40'48"N, ca. 0.1 mm long, glabrous;corolla greenish-white,4-5 84?22'25"W, 28 May 1994, Morales 2815 (INB); R.B. mmdiam.,lobednearlyto thethebase,thelobesstrongly Carara, near Puesto Carara, 200 m, 9046'40"N, 84?32'10"W, 9 May 1990. Zuihga 194 (INB). reflexed at anthesis, the tips and margins of the lobes

350

FLORA NEOTROPICA

FIG. 196. Solanum yanamonense S. Knapp. (Reproduced with permission from Brittonia 38: 299, fig. 17. 1986.)

minutely papillose; anthers 1-1.2 x 0.75-1 mm, poricidal at the tips, the pores teardropshaped;ovary glabrous;style straight,2-2.5 mm long; stigma clavate, the surface minutely papillose. Fruit a globose, green berry,becoming yellowish-green when ripe, 11.3 cm diam.;fruiting pedicels slightly woody, deflexed, ca. 2 cm long, slightly expanded at the apex, 0.25-0.5 mm diam. at the base. Seeds darkbrown in drymaterial,tan in freshmaterial,ovoid-reniform,ca. 3 x 1.5 mm, the surfacesminutelypitted.Chromosome number:n = 12 (voucher Knapp & Mallet 6613). Distribution (Fig. 192). In the Amazon basin of eastem Peru, only known from the type locality on the Rio Amazonas nearthe mouth of the Rio Napo, at ca. 120 m.

smallerstature,distinctiveleaf venationwith the main lateral veins arching and converging again very near the midrib,andin its minute,greenishflowers. The leaf apex of S. yanamonensediffers fromthatof all closely relatedspecies in being long-acuminatewith a truncate tip. Solanumyanamonenseis a small understoryshrub of terrafirme primaryforest and it grows abundantly in clumpswhere it occurs.Its rangemay be muchmore extensive,butbecauseof its small,inconspicuousgreen flowers and fruit, it is not likely to be collected often; however it should be looked for in AmazonianBrazil. Solanumyanamonenseis not treatedby Vasquez (1997), andhe apparentlydoes not considerit as a synonym of any otherspecies occurringin the Iquitosarea.

Specimensexamined.PERU. LORETO: Maynas, XVI. Solanum dolosum species group (S. dolosum, S. gonyrhachis, S. habrocaulon)

Explorama Tourist Camp on Rio Amazonas between Indiana and mouth of Rio Napo, ca. 130 m, 3?28'S, 72?48'W, 26 Jul 1982, Gentry et al. 37969 (MO), noninundated forest, ca. 100 m, 23-27 Jul 1984, Knapp 6609, 6616 (BH, K, NY, US, USM).

Solanumyanamonense is similar and probably closely relatedto S. confine, also of AmazonianPeru, but the latter species is found in the foothills of the Andes and not in the true Amazonian lowlands. Solanumyanamonense differs from S. confine in its

Spindlyshrubs or hemiepiphytes;young stems and leaves variouslypubescent,stronglyzigzag.Sympodial unitsunifoliate.Leaveslinearto ovate,theapexlong-acuminate,thebaseacuteto rounded,themarginsof theapex withminuteciliatetrichomes.Inflorescencesoppositethe leavesorintemodal,stronglyzigzag;pedicelscarswidely andunevenlyspaced.Budsgloboseorellipsoid.Pedicels at anthesisdeflexed.Flowerswhite,ca. 0.8 cm diam.,the

TAXONOMIC TREATMENT

35 1

lobes reflexedat anthesis.Fruit a globose, greenberry; A single collection from the PeruvianAndes (Pasco: prov. Oxapampa,4-5 km N of Mallapampa,2400 m, fruitingpedicelsdeflexed.Seedsnot known. 10002'S,75?45'W,22 Jan 1984,D. N. Smith& J. CanneR Distribution. Montane cloud forests, Colombia 5793) may representa new species in this group, but to Bolivia. thematerialis scantyandthe duplicateI haveseen (MO-) Membersof the Solanumdolosumspecies group only bearsundeveloped fruits. It differs from the speare all very rare,primaryforest shrubs.They are mor- cies includedherein its distinctivebranchedpubescence, phologically most similar to and probablyclosely re- whichis goldenin dryspecimens,andin its fruits,whicl lated to members of the S. confine species group, but dry a brightgolden color. More collections of this indifferin theirciliate leaf apicesandplanarleaf margins. terestingplantareclearlyneeded.

Key to the species of the Solanum dolosum species group 1. Stems and leaves glabrous; inflorescences 1-2-flowered, 0.1-0.5 cm long........................ 122. S. habrocauloa 1. Stems and leaves pubescent, the trichomes golden and uniseriate; inflorescences more than 1-2-flowered. 2. Leaves lanceolate or linear; inflorescences to 1.5 cm long. Cordillera Occidental, Colombia ................................................................... 120. S. dolosuln 2. Leaves ovate or lance-ovate; inflorescences 2-6.5 cm long. E Andes, Bolivia ....... 121. S. goiiyrhachis

120. Solanum dolosum C. V. Morton ex S. Knapp, Ann. Missouri Bot. Gard. 73: 742. 1987 (1986). Type. Colombia. Valle de Cauca:CordilleraOccidental,LaCumbre,1600-1800 m, 14-19 May 1922, Killip 5705 (holotype, US; isotypes, NY, COLn.v.). Fig. 198

sis, minutelypapilloseon the tips andmargins;anithers 1.5-2 mm long, the terminalca. 0.2 mm thickenedand paler, 1-1.5 mm wide, poricidal at the tips, the pores teardropshaped;freeportionof thefilaments0.5--1mmn long, the filamenttube ca. I mm long;ovaryglabrous; style straight,ca. 4 mm long; stigma not distinguishShrubsor climbing shrubs,0.5-2 m tall, the ul- able fromthe restof the style, minutelypapilloseon the timatebranchletsslender;young stems and leaves hir- extremetip. Fruit a globose berry,glabrous,only imsute with uniseriategolden trichomesca. 0.1 mm long; maturefruitseen;fruitingpedicel elongate. Seeds not barkof the olderstemspartiallyglabrate,golden green; known. Chromosomenumbernot known. stems strongly winged from the decurrentleaf bases, Distribution (Fig. 197). On the western slopes and also from de novo wings arising ca. 1 mm below of the CordilleraOccidentalin ColombiaandEcuador, the inflorescence. Sympodialunits unifoliate. Leaves from 1200-2000 m. lanceolateto linear,widestjust proximalto the middle, Specimens examined. COLOMBIA. VALLE DE 5.5-11.2 x 1.4-1.7 cm, with 6-10 pairsof main lateral CAUCA: Cordillera Occidental,W slope, Rio Digua, right veins, these indistinctabove except for the raisedmid- bank between Queremaland La Elsa, 1160-12.00 In, rib,prominent,yellowish,andpuberulentwithuniseriate 27, 29 Mar 1947, Cuatrecasas 23897 (F, US); 13osque trichomes0.05-0.1 mm long, the apex long-acuminate, de San Antonio, W of Call near television antenna. 1950the extreme tip blunt and rounded,the base acute, de- 2050 m, 15 Jul 1984, Gentry et al. 48143 (MO, NY); La currenton the petiole and stein;the leaf marginserose, Cumbre, Cordillera Occidental, 1600-2100 m, 25-27 Sep ciliate near the apex; petioles winged, 1-5 mm long. 1922, Killip 11589 (NY, US); Cerro del Ingl6s, CordiInflorescences opposite the leaves or occasionally in- llera Occidental, Serraniade los Paraguas, 1 hour by jeep temodal,simple,0.5-1.2 cm long, 2-6-flowered,dense- from El Cairo,south slope, 2260-2300 m, 3 Jani 1987, ly to sparselyhirsutewith uniseriategolden trichomes; Silverstone-Sopkinet al. 2930 (MO, NY). ECUADOR.CARCHI: Valle de Maldonado, km 71 pedicel scars irregularlyspaced 0.5-2 mm apart,beon Tulcan-Maldonado rd., 1200-2200 m, 0?45'N, ginning 2-3 mm from the base of the inflorescence. 78?06'W, 20 May 1973, Holm-Nielseni et al. 5959, 6014 Buds globose. Pedicels at anithesis4-6 mm long, de- (AAU), 6035 (AAU, MO, NY). PICHINCHA: Reserva flexed, taperingfrom the calyx tube to a slender base Geobotanico Pululahua, between Reventaz6n and ca. 0.5 mm diam., sparsely pubescent with uniseriate Chaupisacha, 2070-2300 m, 0?05'N, 78?30'W. 9 Dec trichomes.Flowers with the calyx tube ca. 1 mm long, 1988, Cer6n & Igluago 5700 (MO); Reserva Geobotanico sparselypubescent,the lobes broadlydeltoid, swollen Pululahua, Calacali, Chaupisacha, 1800-2000 m, andknob-like,0.5-1 mm long, sparselypubescentwith 0001'N, 78035'W, 29 Jul 1989, Ceron 7172 (K. MO, golden uniseriatetrichomes,minutelypapillose at the QCA, QCNE). tips;corolla white, 8-9 mm diam.,lobed 3/4 of the way Solanum dolosum is most closely related to S. to the base, the lobes long-acuminate,reflexedat anthe- gonyrhachis from high-elevation eastern Bolivia. It

352

FLORA NEOTROPICA

1700

0 0

~

0

90

1'4~~~~~~~~~~~6

FIG. 197. Distribution of Solanum tuer-ckheimii(open circles), S. dolosum (solid circles), S. habrocaulon (solid square) and S. gonyrhachis (bicolored circle).

differs from that species by the charactersdetailed in the key. Plantsof S. dolosumhave been said to be hemiepiphytic, but I suspect thatthis small shruboccasionally merely grows in moss on trunksor fallen logs in its cloud forest habitat.

archingand slender;barkof older stems gray.Sympodial units unifoliate.Leaves narrowlyovate, widest in the proximal third of the blade, shiny above, with uniseriatetrichomes like those of the stems along the midrib,glabrousbeneath,the blades 6-9 x 2-2.5 cm, with 4-6 pairsof mainlateralveins, these notprominent above, except the keeled midrib,prominentandbrown 121. Solanum gonyrhachis S. Knapp,Bull. Br. Mus. below (dry specimens), the apex long-acuminate,ciliNat. Hist. (Bot.) 19: 105. 1989. Type. Bolivia. La ate, the extreme tip rounded,the base truncate,somePaz: Prov. Nor Yungas, 2 km by rd. (ca. 1 km by whatcordate;petioles minute, 1-1.5 mm long, sparsely air)NE andbelow Chuspipata,2950 m, 16?I7'30"S, pubescentwith uniseriatetrichomes like those of the 67?49'W,29 Oct 1984, Nee & Solomon30221 (ho- stems.Inflorescencesoppositethe leaves or intemodal, lotype NY; isotypes BH, K). Fig. 199 filiformandsimple,2-6.5 cm long, zigzag at thepedicel scars, 5-15-flowered, with only 2 or 3 flowers on the Spindlyshrubsor hemiepiphytes,branchesca. I inflorescenceat a time, sparselypubescentwith simple, m long;young stemsandleaves sparselypubescentwith uniseriatetrichomes0.5-1 mmlong;pedicelscars evenly lax uniseriatesimple trichomes0.5-1 mm long, these spacedca. 5 mmapart,theaxis of theinflorescencebendsparserbut still presenton olderbranches;the branches ing ca. 80-90? at each scar.Buds globose when young,

TAXONOMIC TREATMENT

353

FIG. 198. Solanum dolosum S. Knapp. (Reproduced with permission from Annals of the Missouri Botanical Garden 73: 743, fig. 4. 1986.)

laterellipsoid or obovoid, sparselypubescentwith the uniseriatetrichomes.Pedicels at anthesisfiliform,purplish, deflexed, 0.9-1 cm long, taperingfromthe calyx tube to a slenderbase less than0.5 mm diam.Flowers with the calyx tube conical, 1-1.5 mm long, strongly5ribbed,the ribsblackin drymaterial,sparselypubescent with uniseriatesimple trichomesca. 0.5 mm long, the lobes deltoid,apiculate,0.5-1 mm long, sparselypubescentwith the trichomeslike those of the tube,the tips of

the apiculaedenselypapillose;corolla white, 0.8-1 cm diam.,lobednearlyto thebase,thelobesplanarorslightly reflexed at anthesis, the tips and marginsof the lobes minutelypapillose;anthers2-2.5 x ca. 1 mm, poricidal at the tips, the poresteardropshaped;freeportionof the filaments 0.5-0.6 mm long, the filament tube 0.1-0.2 mm long; ovary glabrous;style 3-5 mm long; stigma capitate,the surfaceminutelypapillose.Fruitandseeds not known. Chromosomenumbernot known.

354

FLORA NEOTROPICA

FIG. 199. Solanum gonyrhachis S. Knapp. (Reproduced with permission from Bulletin of the British Museum Natural History (Botany) 19: 107, fig. 4. 1989.)

Distribution (Fig. 197). Known only from middle to high elevations in NW Bolivia, in cloud forest, perhapsoccasionally growing as a hemiepiphyte.

lected by Rusby at the falls of Madeirain westernBrazil is surely a labeling error.The plant is definitely S. gonyrhachis, but the habitatof the lower Rio Madeira Specimensexamined.BOLIVIA.LA PAz: Prov. is very different from the cloud forests of the type loNor Yungas,1.9 km NE of Chuspipata on rd. to Coroico, cality (Nee, pers. comm.). Labeling errorsoften oc2900 m, 16?18'S, 67?48'W, 20 Feb 1986, Solomon curredwith plantssent back fromearlybotanicalexpe14933 (MO, NY). ditions to South America, and Rusby did pass through BRAZIL.ACRE:Falls of Madeira,October1886, habitatverylikethatofthe typelocalityon his way down Rusby 2606 (NY). the easternslopes of the Bolivian Andes. Solanumgonyrhachisis a rarespecies, only collected from a single locality. This sparse occurrence (sensu Rabinowitz, 1981) is common in the S. confine 122. Solanum habrocaulon S. Knapp,Novon 6: 31. 1996. Type. Peru. Amazonas:Bongara,4 km N of and S. dolosum species groups. Pomacochason rd. to Rioja, trail down gorge to W Both Solanumgonyrhachisandits close relative of rd., 2150-2200 m, 5 40'S, 77 22'W, 2 Jun 1986, S. dolosumhave ciliate leaf apices, an unusualcharacKnapp et al. 7507(holotype, USM; isotypes, MO, ter in sect. Geminata. These species, with S. habrocaulon,areclosely relatedto the membersof the NY). Fig. 200 S. confine species group, but are segregated here beSmall shrubs with arching flattened, winged cause they are so distinctive morphologically. Label informationof thetype specimenindicatesthattheplant branches,ca. 1.5 m tall; young stems and leaves comwas growing as a hemiepiphyte,a habitalso attributed pletely glabrous,a few papillae present in the axils of the new leaves; barkof older stems darkbrown.Symto S. dolosum (see above). The locality informationon the specimen col- podial unitsunifoliate.Leaves narrowlyelliptic to lan-

355

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356

FLORA NEOITROPICA

spaced3-4 mm apart.Buds globose when young, later obovate, the corolla strongly exserted from the calyx tube. Pedicels at anthesis somewhat deflexed, 1-1.2 cm long, densely pubescentwith dendritictrichomes, tapering from an apical diam. of ca. 1 mm to a basal diam. of ca. 0.5 mm. Flowers with the calyx tube conical, 1-1.5 mm long, the lobes long-triangular,1-2 mm long, densely pubescent with minute denclritictrichomes; corolla white, 1.3-2 cm diam., lobed ca. 1/2 of the way to the base, the lobes planar at anthesis, sparselypubescentabaxiallywith dendritictrichomes; anthers3.5-5 mm long, 1-1.5 mm wide, the lower 1/3 (ca. 1mm)of theanthersac apparentlysterileandempty, Distribution (Fig. 197). Known only from the giving the anthersan overall obovate shape, poricidal type locality in montaneN Peru,at ca. 2200 m. at the tips, the pores teardropshaped; free portion of Solanumhabrocaulonis distinguishedfromthe thefilaments 0.5-1 mm long, the filament tube 0.25othermembersof the S. dolosum species groupby its 0.5 mm long, glabrous;ovary glabrous or with a few completely glabrous foliage, strongly winged stems, dendritictrichomesnearthe apex;style 0.8-1 cm long, andminute inflorescences. Membersof this groupare glabrousorwith a few dendritictrichomesnearthebase; all apparentlyrare,but since they arevery inconspicu- stigma clavate,the surfacesminutelypapillose.Fruit a ous plants it is possible that they are merely globose, occasionallypointed,green(?)berry,1.5-2 cm undercollected. diam.;fruitingpedicels more or less deflexeci,woody, 1.8-2 cm long, ca. 1 mm diam. at the base, ca. 3 mm diam. at the apex. Seeds rusty brown, flattened-reniINCERTAESEDIS form with somewhat incrassate margins, 2 5-3 mm 123. Solanum delitescens C. V. Morton,Rev. Argent. long, 2-2.5 mm wide, the surfaces minutely pitted. Sp. Solanum168. 1976. Type.Argentina.Tucuman: Chromosomenumbernot known. Depto. Famailla,Quebradade Lules, rd. to Dique, Distribution (Fig. 202). In the mountainsof NW 600 m, 12 Aug 1923, Venturi2462 (holotype US; isotypes BM, F, GH, LIL-n.v., NY). Fig. 201 Argentina in the provinces of Jujuy, Salta, and Solanumdelitescensvar.amphidoxum C. V. Morton, Tucuman,from 500-2600 (4040) m.

ceolate, widest just below the middle, with 7-8 pairs of main lateralveins, glabrouson both surfaces,6-9 x 1.5-2.7 cm, the apex acuminate, the ultimate tips rounded,marginsof the tips minutely ciliate, the base attenuate,decurrentonto the petiole; petiole 2-3 mm long. Inflorescences opposite the leaves, simple, 1-2 mm long, 1-2-flowered,glabrous;pedicelscars closely spaced, almost sessile. Buds andflowers not known. Fruit a globose, green berry,0.7-1 cm diam. (immature);fruitingpedicels pendant,somewhatwoody, 22.5 cm long, ca. 0.5 mm diam. at the base, ca. 2 mm diam. at the apex. Seeds not known.

Rev. Argent. Sp. Solanum 171. 1976. Type. Argentina. Tucuman: Burruyacu, Puente del Rio Calera, 500 m, I Nov 1939, Venturi9967 (holotype US; isotype GH).

Subshrubs to shrubs, 0.5-1 m tall, from woody rootstocks;stems thick, erect, densely pubescentwith dendritictrichomesca. 0.25 mm long, stronglywinged fromthe decurrentleaf bases, the wings to 5 mm deep; barkof older stems pale yellowish-brown.Sympodial unitsdi- or trifoliate,usuallynot geminate.Leavesobovate, widest above the middle, densely pubescentwith dendritictrichomeslike those of the young stems, later glabrousor sparselypubescentadaxially,sparselypubescent abaxially,the dendritictrichomesdenseralong the veins; major leaves 8.5-37 cm long, 4.5-13 cm wide, with 6-9 pairsof main lateralveins, these drying somewhatyellowish abaxially,the apex acute,the base sessile, decurrenton the stem;minorleaves not differing fromthe majorleaves;petioleabsent.Inflorescences oppositethe leaves or intemodal,3-10 cm long,densely pubescent with dendritictrichomes, simple or manytimes branched,10-20-flowered;pedicel scars evenly

Specimens examined. ARGENTINA. JujuY: Tumbaya, Volcan, 2600 m, 5 May 1929, Venturi 9532 (US). SALTA:PefiasBlancas,Sierradel Caj6n.4040 m, 16 Mar 1914, Rodriguez 1421 (NY); Rosario de la Frontera,Los Batios,1000 m, 28 Jul 1929, Vienturi9414 (F,GH,NY,US).TucUMA&N: Cadillal,Oct 1915,Castillon

s.n. (Lillo87881)(A); Cuestade SanPablo,800 m, 8 Sep 1909, Lillo 9739 (US); Aconquija,15 Oct 1939,Me.yer 2982 (A); ParqueAconquija,500 m, 14 Aug 1941,,Meter 4315 (A, F); ParqueAconquija,14 Aug 1941,Meyer4440 (A); Chicligasta,Rio Cochuna,1200 m, 28 Nov 1937., O 'Donell s.n. (Lillo 18159) (GH); San Javier-Villa

Vasquesrd., 12 Jun 1945,Ortizs.n. (A, NY); Burroyacu, La Ramada,470 m, 21 Aug 1932, Peirano s.n. (herb. Lillo 18155) (GH);Tafi,ParqueAconquija,ca. 1200 rn, 4 Nov 1952, Peterseni & Hjerting 562 (NY); hills of Potrero de Las Tablas, 700 m, 29 Oct 1919, Schreiter

856 (US); Tafi, Raco, 1000 m, 19 Oct 1930, Schbeiter 6491 (A); Tafi,Quebrada de Lules,500 m, 12 Aug 1923, Schbeiter 9982 (A, US); Burroyacu, valley or Rio Loro, 550-700 m, 2 Sep 1951, Sleurner 2088 (US); Tafi.

Siamb6n,1200m, 15 Oct 1925,Venturi 3923 (F,GH,US); Burroyacu,CerroEl Nogalito, 1500 m, 12 Apr 1929, Vtenturi8896 (US).

357

TAXONOMIC TREATMENT

..~~~~~~~~~~~~~~~~~~~~~~.. . ........

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5cm FIG.201. Solanum delitescens C.V. Morton. Argentina. Venturi 2462 (BM-isotype).

35 8

FLORA NEOTROPICA

1X500

-----

-

--

FIG. 202. Distribution of Solanum delitescens (open circles), S. lasiocladum (bi-colored circle), S. mapiricum (solid circle) and S. cordioides (solid square).

Solanumdelitescensis of uncertainaffinities,and could be an isolated relativeof the section in its broadest sense. The antherstructureof the plants is peculiar (see Morton, 1976) and unlike anything else in the genusSolanum,with the narrowbase being completely empty of pollen. Its relationshipsmay be with members of sect. Cyphomandropsis.

trichomes of varying lengths, 0.1-1 mm long; older stems remainingpubescent and golden; barkof older stems reddish. Sympodial units difoliate, geminate. Leaveselliptic,widest at the middle,shiny andsparsely pubescentabove with erectuniseriatetrichomesca. 0.5 mm long, the trichomes denser along the veins, the upperepidermislarge-celled,denselygoldenpubescent beneathwith uniseriatetrichomes of varying lengths, these 0.1-1 mm long, denser along the veins; major 124. Solanum lasiocladum S. Knapp, Brittonia 38: leaves 7.5-15 x 3-6.5 cm wide, with 8-9 pairsof main 278. 1986. Type.Venezuela.Anzoategui:Libertad, lateralveins, these raisedandpubescentabove, promiridges and tops of MontafiasNegras, along Sucrenent and densely pubescentbeneath,the apex acute to Anzoateguiborder,20 airkm NE of Bergantin,NE acuminate,the base acute,not decurrenton the petiole; of Buenos Aires, Serraniade Turimiquire,summit petioles 1-3 cm long; minor leaves differing from the elfin forest, 2350 m, 10?04'30"N,64?1lW, 28 Nov majorones mainlyin size, butoccasionallyequalin size 1981, Davidse & Gonzalez19541 (holotype,VEN; to the majorleaves, 5.5-10 x 2.2-6 cm wide, the apex isotype, MO). Fig. 203 acute, the base acute; petioles 1-3 cm long. InfloresShrubs or small trees, 8-9 m tall; young stems cences opposite the leaves, simple or once-furcate, 1and leaves densely pubescent with golden uniseriate 5 cm long, 3-10-flowered, densely pubescent with

TAXONOMIC TREATMENT

359

FIG. 203. Solanum lasiocladum S. Knapp. (Reproduced with permission from Brittonia 38: 279, fig. 5. 1986.)

golden uniseriate trichomes like those of the young stems and leaves; pedicel scars slightly raised and corky, in pairs, the membersof a pair closely spaced, the pairs ca. 1 mm apart,the scars not obscuredby the pubescence. Buds elliptic, densely golden pubescent, the trichomes denser along the ribs of the lobes, the corollalate exsertedfromthe tube.Pedicels at anthesis deflexed,ca. 9 mm long,taperingfromthe conicalcalyx tubeto a slenderbase ca. 0.5 mm diam.,pubescentwith golden trichomes of varying lengths, these 0. 1-1 mm long. Flowerswith the calyxtubeconicallycup-shaped, 1.5-2 mm long, strongly5-ribbed,the sinuses hyaline, the lobes narrowlytriangular,appearingas mere extensions of the ribs of the tube,0.5-1 m long, the entire

calyx goldenpubescentwith uniseriatetrichomes,these denseralong the ribs and lobes; corolla white, 1.2-1.5 cm diam., lobed nearlyto the base, the lobes somewhat reflexed at anthesis, the tips and marginsof the lobes densely pubescentwith -shortuniseriatetrichomesca. 0.2 mm long; anthers 3-3.5 mm long, 1 mm wide, poricidal at the tips, the pores teardropshaped; free portionof thefilaments ca. 0.5 mm long, the filament tube ca. 1 mm long; ovary glabrous;style straight,66.5 mm long;stigma a broadareaon the tip of the style, minutely papillose. Fruit a globose, green berry,ca. 1 cm diam.;fruitingpedicelswoody, deflexed or appearing somewhat erect when the fruit is immature, 1.51.8 cm long, ca. 1 mm diam.at the base, sparselypubes-

FLORA NEOTROPICA

360

FIG. 204. Solanum mapiricum S. Knapp. (Reproduced with permission from Brittonia 38: 300, fig. 18. 1986.)

centwithgoldenuniseriatetrichomes.Seedsdarkbrown, ovoid-reniform(?),ca. 2 mm long, 1 mm wide (immature),the surfacesminutelypitted.Chromosomenumber not known. Distribution (Fig. 202). Known only from the summitof the Serraniade Turimiquireon the borderof SucreandAnzoateguiin Venezuelaat2350 m elevation. ANZOATEGUI: Specimensexamined.VENEZUELA. Woodedsummitof CerroPeonia(Cerrode los Pajaritos) above SantaCruz,headwatersof Rio Manantiales,E of Bergantin, 2350 m, 21 Mar 1945, Steyermark61673 (MY, VEN). Solanum lasiocladum is quite similar to S. malacothrix, from Guerrero in western Mexico. It shareswith thatspeciesthe dense indumentof uniseriate trichomes,but differs in its largerminorleaves, longer inflorescences, long-petiolateleaves, and largerflowers. The summitof the Serraniade Turimiquirehasonly been ascended by botanists a few times, and twice S. lasiocladumhas been collected, but no mention of its local abundanceis madeon eitherlabel.The Serraniade Turimiquireis isolatedandis oftencoveredwith clouds.

125. Solanum mapiricum S. Knapp,Brittonia38: 299. 1986. Type. Bolivia. La Paz: Prov.Larecaja,Tuiri,

nearMapirion left bankof Rio Mapiri,490-750 m, 12-30 Sep 1939, Krukoff 10898 (holotype, US; Fig. 204 isotypes, F, K, MO, NY, U). Shrubs or small trees 2-6 m tall; young stems and leaves glabrousor minutely white-papillose;bark of older stems deep shiny maroon,becoming grayish on much largerstems.Sympodialunitsusually unifoliate. Leaves elliptic to narrowlyelliptic, widest at the middle, glabrousabove, dryingpale green, pubescent with tufts of uniseriatetrichomes 0.1-0.5 mm long in the axils of the main lateral veins beneath, these trichomes sparseand from the veins, not the lamina,the leaves 4-11 cm long, 1.5-3.5 cm wide, with 7-10 pairs of main lateralveins, these not prominent,often yellowish andshiny beneath,the apex acuteto acuminate, the base acute to obtuse;petioles 3-7 mm long. Inflorescences opposite the leaves, occasionally somewhat intemodal,simple,0.2-1.1 cm long, 5-7-flowered, glabrousexcept for the buds;pedicelscars closely spaced, overlappingor with the edges just meeting, beginning nearthe base of the inflorescence.Buds globose, densely to sparselypubescentwith whiteuniseriatetrichomes ca. 0.25 mm long, laterelliptic with the exsertionof the corolla. Pedicels at anthesis ca. 1 mm long, deflexed, taperingfrom the calyx tube to a slenderbase ca. 0.25 mm diam.Flowerswith thecalyxtubecup-shaped,ca. 1

TAXONOMIC TREATMENT

36 1

mm long, densely to sparselypubescentwith uniseriate trichomes, the lobes deltoid, 0.5-1 mm long, pubescent with the same trichomesas the calyx tube;corolla white, 1-1 .2 cm diam., lobed ca. 3/4 of the way to the base, the lobes reflexed at anthesis, the tips and margins of the lobes minutelypapillose occasionally with a few uniseriatetrichomes;anthers ca. 3 mm long, 1 mmwide,poricidalatthetips,theterminal0.5 mm paler and thickened, the pores becoming slit-like upon drying;freeportionof thefilamentsminute,0-0.1 mm long, the filamenttube 0.5-1 mm long;ovaryglabrous;style straight, 6-7 mm long; stigma capitate, bilobed, the surfaceminutelypapillose. Fruit a globose berry,6-8 mm diam. (immature),green;.fruitingpedicelswoody, slenderand erect, 1.5-1.7 cm long, ca. 0.75 mm diam. at the base. Seeds frommaturefruitsnot known. Chromosome nutmbernot known.

not geminate, occasionally appearing plurifoliate. Leaves elliptic, widest at the middle, glabrouson both surfaces, drying black, the undersurfacepaler in livl plants, the leaves 4-11 x 1.5-3.5 cm wide, with 5-7 pairsof mainlateralveins, these dryingdarkerbeneath, the apex abruptlyacute, the base attenuate,decurren.t ontothe distal 1/3 of thepetiole;petioles 2-3.5 cm long. Inflorescencesintemodal,many-timesbranched,6-1 1cmn long, 30-100-flowered, completely glabrous;pcdicel scars widely and regularyspaced ca. 0.5 mm apart,in upperpartof each terminalinflorescencebranch.Bud's globose, laterellipsoidwith the exsertionof the corolla, glabrousanddryingblack.Pedicels at anthesis6-8 mm long, deflexed, taperingfrom the calyx tube to a slenderbase ca. 0. 5 mm diam.Flowers with the calvx tube cup-shaped, 1.5-2 mm long, glabrous, the lobes broadly deltoid with apiculate tips, 0.5-1 mm long, papillate on the apiculae; corolla white or greenishDistribution (Fig. 202). All collections arefrom white, 6-8 mm diam., lobed ca. 3/4 of the way to the the region of Mapiri, in the lowlands of northernBo- base, the lobes reflexed at anthesis, the tips and marlivia, from 400-800 m elevation. gins of the lobes minutelypapillose, the tips cucullate; anthers 1.5-2.5 x ca. 1 mm, poricidal at the tips, the Specimensexamined.BOLIVIA. LA PAZ:Mapiri region,San Carlos,850 m, 18 Feb 1927,Buchtien1273 pores teardropshaped;free portionof thefilamcntsca. (US); Mapiriregion, San Carlosnear Sarampiuni,600 0.5 mm long, the filament tube ca. 0.5 mm long; ovarv m, 3 Mar 1927,Buchtien1274 (NY,US); Mapiriregion, glabrous; style straight, 3-3.5 mm long; stigma clavSan Carlos, 850 in, 24 Apr 1927, Biuchtien1275 (US) ate,thesurfaceminutelypapillose.Fruita globoseber.y, San Antonio near Mapiri,850 m, Dec 1907, Buchtien 1-1.2 cm diam., green;fruitingpedicels woody, slen1435 (US); Larecaja,Copacabana(about 10 km S of der and erect, 1-1.2 cm long, ca. 1 mm diam. at the Mapiri), 850-950 m, 8 Oct-15 Nov 1939, Krukoff base. Seeds darkbrownwith palermargins,flattened11250 (F, K, MO, NY, US). reniform,2-3 x 3-4 mm, the surfaceminutelyandreguSolanunmmapiricum is very similar to both S. larly pitted. Chromosomenumbernot known. daphnophyllumand S. symmetricum,also of Bolivia. Distribution (Fig. 202). In SouthernAtlanticwet It differs fromS. daphnophyllumin its pubescentvein in coastal Bahia, only known from the Reserva forest axils and buds, and closely packed pedicel scars; and from S. svmmetricumin its minute free portion of the do Mico-Leao in the muncipio of Una at sea level or filaments,small flowers, pubescentbuds,andelongate slightly above. flowering axis. The affinities of this rare species are Specimens examined. BRAZIL. BAHIA: Mun. Una, not clear at present;more collections anddetailedfield ReservaBiol6gicado Mico-leao(IBAMA),entrancefrom km 46 of Ilheus-Unard., 15?09'S,39'05'W, 8--12Mar notes concerninghabitand habitatareneeded. 1993, Amor-iniet al. 1071 (NY-n.v. fide M. Nee), 14-15 Apr 1993, Amorimn et al. 1223 (NY-n.v. fide M. Nee); NEW SPECIES ADDED IN PROOF Mun. Ilheos, between Bom Gosto and Oliven,a, 12 Mar 1943, Froes 20023 (K, NY); Mun. Una, Reserva Biol6gica 126. Solanum cordioides S. Knapp, sp. nov. Type. do Mico-leao (IBAMA), entrance on km 46 of BA-00 1, Brazil. Bahia: Mun. Una, Reserva Biol6gica do llheus--Una, 15 09'S, 39 05'W, 7 Jan 1993, Jar-dinie al. 102 (NY-n.v. fide M. Nee); Mun. Una, Reserva Biol6gica Mico-leao (IBAMA), entranceon km 46 of BA001, Ilheus-Una, 15?09'S, 39?05'W, 7 Jan 1993, do Mico-leao (IBAMA), entrance on km 46 of BA-0)01, Jardim et al. 12 (holotype, CEPEC;isotypes, BM, llheus-Una, near Picado do Principe, 15?09'S, 39?05'W, K, NY). Figs. 205, 206 20 Apr 1996, Jaidim & SantAna 790 (CEPEC,NY); Mlun. Ilhus, new rd. to Povado do Vila Brasil, entrance on km Species Solano acliminatoRuiz & Pav6n affinis, 28 of Ilheus-Una rd., 3 km along branch, 13 Jan 1985, sedfoliisutrinque glabrisnitidis,inflorescentiis ramosissimis, Mattos Silva et al. 1820 (NY-n.v. fide M. Nee); NMun. floribus viridi-albisparvis, seminibusfuscis, differt. liheus, rd. from Olivenmato Una, 18 km S of Olivenoa, ca. 0 m, 21 Apr 1981,Mori et al. 13705 (NY-n.v. fide M. Shrubs or small trees to 8 m tall; young stems Nee); Mun. Una, Reserva Biol6gica de Una (Reserva de and leaves glabrousand shining, dryingblack;barkof Mico-leao), 4.1 km W of rd. from Ilheus to Una, 5.6 km older stems grayish.Svmpodialunitsusually difoliate, W of Reserve gate, 15010'50"S, 39003'40"W,6 Feb 1994,

362

FLORA NEOTROPICA

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DOUBTFUL

3 63

NAMES

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Thomas et al. 10310 (BM, CEPEC, NY); Mun. Uru9aca, 7.4 km N of SerraGrandeon rd. to Itacar6,FazendaLagoa do ConjuntoFazenda Santa Cruz, 14?25'24"S,39?03'38"W, 9 Feb 1994, Thomas et al. 10347 (NY-n.v. fide M. Nee); Mun. Una, 5 km S of Una on rd. to Canavieras, then 0.8 km SW of hwy., 15?19'58"S, 39?03'18"W, 18 Nov 1995, Thomas et al. 11045 (BM, CEPEC, NY).

which it has been misidentifiedin the past.It appearsto be endemicto the Una Reserve,which is home to many otherspecies endemic to the Atlanticcoastal forests of Bahia andEspirituSanto(Thomaset al., 1998).

DOUBTFUL NAMES

Solanumcordioidesis a very distinctivespecies, Solanum caudatumDunal in DC., Prodr. 13(1); 140. with its shiny elliptic leaves and small greenish-white 1852. Type. Peru. S.loc.,Pavon s.n. (holotype, G). flowers. It is most similar and probablyrelated to S. The type specimen in G is very fragmentaryand acuminatum of central Peru, with which it shares I am unable to match it to any Solanaceae I know. branched,usually intemodalinflorescences,and short, stout anthers.It differs from S. acuminatumin being Solanum linkianumRoem. & Schult., Syst. Veg. (ed. completelyglabrous.Thespeciesis namedforits marked 16) [Roemer& Schultes] 4: 601. 1819. Type. Bralikeness to some species of Cordia(Boraginaceae),for zil, Anon. (B?).

3 64

This name certainlybelongs in synonymyunder Solanumpseudocapsicum,but as I have tracedno type material,I leave it as doubtfulfor the present. Solanumcormanthum Vell.,Fl. Flumin.86. 1829 [1825]; Fl. Flumin. Icones 2: fig. 113. 1831 [1827]. Type. Brazil. Rio de Janeiro:royal estates of SantaCruz. It is unfortunatethatnone of Vellozo'stype specimens survive to enable unequivocal identification of the names in the Florae Fluminensis.The taxon represented in this plate is probably a species of either AurelianaorAthenaeaas it has axillaryclustersof flowers,ratherthana typicalcymoseSolanuminflorescence. However, the anthers are those of a Solanum, and Sendtner.who firstdescribedthe generaAurelianaand Athenaea, considered this species to be a memberof the genus Solanum. Without a specimen to resolve these apparentambiguities, it seems reasonableto regard Vellozo's Solanum cormanthumas a name of uncertainaffinity. SolanumglomuliflorumSendtn.in Mart.,Fl. Bras. 10: 24, tab. 3, fig. 11-15. 1846. Syntypes.Brazil. Serra d'Estrella,Schotts.n.(notfound);"Brasiliaaustralis," Sellows.n. (probableisosyntype,B [F neg. 2823]). This nameis of uncertainapplication.In describing SolanumglomuliJorum,Sendtnercites it as being glabrouswith leaf-opposedinflorescencesandflowers havingequalanthers.Dunal(1852) citedtwo additional specimens,Lhotskys.n. andBlanchet209. The Lhotsky sheet is a specimen of S. pseudoquina, but I have been unableto find the Blanchetsheet. Materialidentifiedat Kew as S. glomuliflorumhas axillary inflorescences, and is otherwise a good matchfor Sendtner'sdescription. Sellow s.n. at B (now destroyed) and labeled "SolanumglomuliflorumSendtn.?"(F neg. 2823) is a plantwithleaf-opposedinflorescencesandshinyleaves, but appearsto be a specimen of S. pseudoqtuina. Solanum lacteum Vell., Fl. Flumin. 82. 1829 [1825]; Fl. Flumin. Icones 2: fig. 93. 1831 [1827]. Type. Brazil, Rio de Janeiro. The plantrepresentedin plate 93 in Vellozo also (see above) is most likely a species of Aureliana or Athenaea;it too has axillary clusters of flowers. Like the plate depictingSolanumcormanthum,the characters areambiguous,andthereforeVellozo's S. lacteum too should be regardedas a name of uncertainaffinity. NeitherS. lacteumnor S. cormanthuim aremembersof sect. Gemiinata. The characters differentiating AurelianaandAthenaeaare:calyces accrescentwith 5 principalnerves and 10 secondarynerves (Athenaea) vs. calyces not accrescentwith only 5 principalnerves (Aureliana, see Barboza& Hunziker, 1989; Hunziker & Barbosa, 1991, for revisions of the two genera).

FLORA NEOTROPICA

EXCLUDED SPECIES Species included here are those which have in the past been associated with the members of sect. Geminata:eitherthesetaxahave beenconsideredmembersof the section (see History),or they have been consideredrelatedto species treatedin this monograph. Solanum ardisioides Blume, Bijdr. 696. 1826. Type. Indonesia.Java,Mt. Salak,Blumes.n.(holotype,L). This specimenis a plantof the genus Tetr-ardisia in the Myrsinaceae. Solanum brevifolium Dunal, Sol. Synop. 22. 1816. Type.Ecuador.Pichincha:"adarborespropeurbem HarraeQuitensis,"Humboldt& Bonplands. n. (holotype, P-Bonpl.). Withclusteredaxillaryinflorescencesandsmall, orbicular leaves, this species is a member of sect. Anarrhicomenum Bitter,a smallgroupof predominantly Andean subshrubsand scandentvines. Theirrelationships areprobablywith membersof subgen.Petota s.l. SolanumconanthumDunal in DC., Prodr.13(1): 127. 1852.Basedon S. calvcinumNees, Trans.Linn.Soc. 17:60. 1834. Type."Solanipubescentisvar..?Herb. Madr.,Wall. Catal. Suppl. n. 237 (ad jecto tamen exemplo uno Sol. pubescentis genuini), ex Herb. Heyn. (Wall. lc. n. 246)." AlthoughI havebeenunableto findthetype sheet of thisname,fromthedescriptionit is clearlyreferableto SolanumpubescensWilld.,anAfricanspeciesof uncertainrelationships in subgen.Leptostemonum (see Whalen, 1984).Dunalalso didnot see a specimen,butcitedin full Nees' description(Dunal, 1852). Solanumn pubescens is the type species of sect. Anisantheruiim Bitter,and is distinctivein havingone elongate,curvedanther,a featurementionedin the descriptionof S. calvcinum. SolanumdelicatulumL. B. Sm. & Downs, Phytologia 10: 424. 1964. Type. Brazil. SantaCatarina:Itajai, Cunhas, 10 m, 8 Feb 1955,Klein 1131 (holotype, HBR-n.v.; frag. US). Thisspeciesprobablybelongsto sect.,Solanum with its intemodal inflorescences and somewhat "prickly" stems.Thetypespecimen(inphoto)is quitewoodyhowever,andthe taxonis of somewhatuncertainaffinities. SolanumdistichumSchumach.& Thonn.,in Schumach., Beskr.Guin.P1.122 [142]. 1827(1828--1829).Type. Guinea, Aquapim, Thonning s.n. (C-n.v., fide Hepper, 1976, 3 sheets). Bitter(1923) andHeine(1963) bothincludedthis taxon as a variety of the widespread and variable SolanumindicumL. (= SolanumanguiviLam.),a member of subgen. Leptostemonum,the Solanum anelivi

EXCLUDED SPECIES

365

group of Whalen (1984). Members of this group are occasionally unarmed,which is surelywhat led Dunal to include S. distichumin his groupLeiodendra.

and its relationshipsprobablylie with membersof the Solanumellipticumgroupof Australia(Whalen,1984). The species is a Hawaiianendemic.

Solanum.filiformeRuiz & Pav., Fl. Peruv. 2: 31, tab. 159, fig. 6. 1799. Type. Peru. Lomas de Atiquipa, Tafallas. n. (holotype, MA-n.v.). This species is a memberof subgen.Petota sect. Basarthrum(Bitter) Bitter (equivalent to some members of series CaripensiaCorrell, 1962) and is related to the widespreadSolanum caripense Dunal.

SolanumrancaguenseDunalin DC., Prodr.13(1): 1 C0. 1852.Type.Chile,Prov.O'Higgins,Rancagua,Bertero 633 (holotype,P [Mortonneg. 8308]; isotypes, B [F neg. 2742], MO, P [Mortonneg. 8309]). This species is a member of sect. Solanuin and is a synonymof eitherS.furcatumDunal(fide M. Nee, in litt.) or S. sublobatum Willd. (annotation by .J. Edmondson type dated 23 Aug 1971).

Solanum isabellei Dunal in DC., Prodr. 13(1): 153. 1852. As "isabelli." Syntypes. Uruguay. Monte- SolanumreceptumVanHeurck& Mull. Arg., Observ. video: Montevideo, Isabelle s.n. (G-DC [F photo Bot. 46. 1870. Syntypes. Mexico. Veracruz:prope 6774] IDC microfiche 800-61.2074:11.2, G Jalapem, Schiede s.n. (AWH); prope Orizabem, [Morton photo 8601]); Uruguay. Montevideo: Botteri 852 (BM, G-DC, MPU). Montevideo, Gay 1828 (P). AlthoughVanHeurck& Muiller Argoviensis(1870) Thisspeciesis a memberof sect.Solanumandis a thoughtthis species closely relatedto otherspecies consynonymofS. chenopodioidesLam.(fideM.Nee, inlitt.). sidered to be membersof sect. Geminata,it is clearly referabletoSolanumumbellatumMill.,a commonweedy SolanummembranaceumWall. ex Nees, Trans.Linn. species of sect. Brevantherum(see M. Nee, 1993). Soc. 17:41. 1834.Syntypes.India,Wallichcatalogue 2625 (BM, K, G-DC). The Wallich epithets pub- Solanum reductumC. V. Morton, Rev. Argent. Sp. lished in the Catalogue(183 1) arewithoutdescripSolanum171. 1976.Type.Argentina.Tucuman:CerTo tion andthusarenominanuda.Dunal(Dunalin DC., del Campo,Burruyacui,1500 m, 20 Feb 1930, VenProdr.13(1): 143. 1852) also publisheda combinaturi 10179 (holotype, US; isotypes, A, LIL, NY). tionof Wallich'scataloguename.Bothof thesecomSolanumreductumis a herbaceousplant with a binations arebased on WallichCat. 2625, andthus woody rootstock,reminiscentof some membersof sect. should be consideredhomotypic. The correctcita- Cyphomandropsis. Theinflorescencesareelongate,and tion for this name therefore should be Solantum the pedicel scars are slightly raised, like those of S. membr-anaiceum Wall. ex Nees. andrelatives.M. Nee (in litt.)placesthis speflaccidumn Specimens bearingthe Wallich catalogue num- cies in sect. Cyphomandropsis,but its affinities may ber 2625 in both BM and K are plants of Lycianthes lie either with sect. Solanum or with "dulcamaroid" bigeminiata(Nees) Bitter(see Bitter,1919b), a member taxa.Bohs (1994) does not includeit in herlist of meiof a complexanddifficultgroupof OldWorldLycianthes, bers of sect. Cyphomandropsis.The new growth and which some authorsconsidercongenericwithSolanum inflorescences are covered with tiny dendritichairs. (see Symon, 1985). These specimens have five elongate calyx lobes, and the inflorescences are axillary, SolanumrefractifoliumSendtn.in Mart.,Fl. Bras. 10: 31. 1846. Type. Brasil australiore,Sellowt,/-f19/ clearly excluding them from sect. Geminata. (holotype,B-n.v. [perhapsdestroyed:F neg. 2846]). Solanlummor-tonianunm Standl.,Publ. Field Mus. Nat. With its cordate, densely glandularpubescent Hist., Bot. Ser. 18: 1568. 1938. Type. Costa Rica. leaves, shiny fruitswith few seeds and thin pericarps, Alajuela: Zarcero, 1590 m, 20 Nov 1937, Austin andslightlyraisedpedicelscars,Solanumrefractifolium SmithA615 (holotype, F; isotype, US). is a member of a group containing S. flaccidum, S. Specimens of Solanum mortonianumrepresent boerhaviifolium,S.jasminoides,andothersimilartaxa. plants of a species of Cuatresia, with longitudinally dehiscing anthers, a distinct corolla tube, and leaves Solanum(?)serratumDunal in DC., Prodr.13(1): 146. with an offset midvein. For a synopsis of the genus 1852. Type. Brazil. Rio de Janeiro: Serra dos Cuatresia see Hunziker(1987). Orgaos, Lhotsky 124 (holotype, G-DC [F photo 6778], IDC microfiche 800-61.2073:111.2). Solanumn nelsoni Dunal in DC., Prodr. 13(1): 123. amendedto S. serratifoliumDC., Prodr.13(1):680. 1852. Type. Hawaii, Nelson s.n. (holotype, BM). 1852. "S. ? serratum,lege: serratifolium,nam sp. differt a S. serratoBlancoi. (A.DC.)" This species belongs to subgen.Leptostemonum. It is a scandent, unarmedshrubwith orbicularleaves The type specimen of this plant is not a member

FLORA NEOTROPICA

366

of the Solanaceae. It has altemate, simple, estipulate Solanum argentinum Bitter & Lillo, Repert. Spec. Nov. Regni Veg. 12: 547. 1913. Syntypes[notlectoleaves with serratemargins,andhas a single, detached typifiedby Morton,1976]. Argentina.Cuestade Sa. fruit. The fruit is dark brown and woody, and has a Luisina,Lorentz& Hieronymus648 (CORD).Prov. single carpelwith severallarge(ca. 0.75 cm), pale seeds. Catamarca:Catamarca,Hieronymus& Lorentz479 The fruitingpedicel is articulateabove the base. (CORD). Prov. C6rdoba: Villa Maria, Stuckert Solanum triquetrumCav., Icon. P1. 3: 30, fig. 259. 14860 [14850?] (B, CORD, LIL); C6rdoba, 1795. Type. Cultivated at Madrid, from Mexico, Stuckert15164 (B, CORD, LIL);Villa Maria-Rio Anon. (MA?). Tercero,Stuckert16820 (B, CORD);Cruzdel Eje, Stuckert 17237; Villa Maria, Stuckert 19'987(B, This species is a scandentshrubor semi-woody CORD);EstanciaRio Primero,Stuckert21484 (B, vine from northem Mexico and the southem United CORD), 21528 (B, CORD [Morton, 1976, has States.It has lobateleaves andbrightred,juicy berries, "21258]). Prov.Santiago:Cerrodel Tableaclo,Saile It the has of sect. Dulcamara. and is clearly a member Echegaray s.n. (herb. Hieronymus). Prov. distinctivepedicel bases of thatgroup,which aresomeTucuman:Tucuman,Hieronymus& LorentLs.n. (B, what elevated (see Knapp, 1989). frag.F); Tucuman,bei derBajada,Lorentz6 7 (herb. Hieronymus); Burruyacu, Stuckert 7996 (?); SolanumuporoDunal in DC., Prodr.13(1): 138. 1852. Type. Tahiti, Morrenhout s.n. (holotype, G-DC Burruyacu,Stuckert 13156; Burruyacu, Stuckert microfiche 19721. [lDC 800-61.2072:111.4]). Solanum argentinum var. chroniotrichnov Bitter, This species belongs to subgen. Leptostemonum, Repert. Spec. Nov. Regni Veg. 12: 549. 1913. and has also been called Solanumanthropophagorum Type. Argentina. Jujuy: S.loc., Stuckert 21337 Seem. (see Whalen, 1984). It is a member of the S. (holotype, B [destroyed]; neotype, CORD, desdunalianumgroup (Whalen, 1984), which are all unignated by Morton, 1976). armedtaxa occurringfrom Australiato Polynesia. The following taxa belong to Solanum sect. Holophyllas.str.A preliminarylist of species andsynonymy is given here,but is in no way exhaustive.There are several new taxa in this group which are not yet describedandthe circumscriptionof the grouphas not yet been finalized. The groupis currentlyunderstudy by L. Bohs (UT) and S. Knapp (BM).

Solanum canoasense L. B. Sm. & Downs, Fl. Ilustr. Catar., Pt. 1, Solanac. 104. 1966. Nom. nov. based

on S. cataractae L. B. Sm. & Downs, Phytologia 10: 427. 1964. Type. Brazil. Est. Santa Catarina, Mun.Bom Retiro,Rio Canoas,Campodos Padres, 1300-1400 m, 22 Nov 1956, Smith & Klein 7843 (holotype, US; isotypes, HBR-n.v., R-n.v.). Solanum evonymoides Sendtn. in Mart., Flora 24, Beibl.

sect. Holophylla(G. Don) Walp.,Repert.Bot. Soltanurn 2 (5): 87. 1841. Type. Brazil. Bahia:"Ilheos",NoSyst. 3: 51. 1844.as subsectionin G. Don, Gen. Syst. vember, Martius 625 (B [F neg. 2817], BM, K, 4: 422. 1838. Lectotype species (designated by MO, NY, P [Morton neg. 8189]). Seithe, 1962):SolanumcervantesiiLag.(= Solanum pubigerumDunal). Superfluouslylectotypifiedby Solanumpubigerum Dunal, Hist. Solanum 160, tab. 6. 1813. Type. Cultivated in Montpellier, of unspeciD'Arcy (1972) with SolanumpulverulentumPers. fied origin, no specimens cited, butDunal s.n. (MPU (see Knapp, 1989). Solanumnaligerum Schltdl., Linnaea 19: 301. 1847. Type.Mexico.Michoacan:Angangueo,Oct,Schiede s.n. (holotype, HAL?). SolanumpterocladumVan Heurck & Mull. Arg., Observ.Bot. 44. 1884.Type.Bolivia.La Paz:Prov. Larecaja,Sorata, 3000 m, May 1858, Mandon 415 (syntypes AWH, G-DC [Mortonphoto 8550], NY). Solanum manicatumBitter, Bot. Jahrb.Syst. 50, Beibl. 11 1: 63. 1913. Type. Peru. Ayacucho: Prov.

Huanta.rd. to TamboaboveOsno,Rio Apurimac, 2600-2700 m, Weberbauer 5643 (holotype, B [destroyed: F neg. 2620]; isotype, MOL). Slanumndotanuin C. V. Morton & Standl., Publ. Field Mus. Nat. Hist., Bot. Ser. 18: 1079. 1938. Type. Costa Rica. San Jos6: Laguna de La Chonta, NE of Santa Maria de Dota, 2000 m, Standley42265 (holotype. US).

[Morton photo 22273]) would be a possible neotype; the label states grown at "Jardin," determined by Dunal 1851 as "Solanum cervantesii Lag. pubigerum Dun."). Solanumncervantesii Lag., Nov. Gen. Sp. 10. 1816. Type. Cultivated in Madrid, seeds from Mexico. No specimens located. Solanum modestum Roem. & Schult., Syst. Veg. (ed. 16) [Roemer & Schultes] 4: 663. 1819. Type. Mexico, s.coll. (B-W-n.v.). Solanumndivaricatum M. Martens & Galeotti, Bull. Acad. Roy. Sci. Bruxelles 12(1): 136. 1845. Type. Mexico. Oaxaca: Pefioles and Cerro San Felipe, Galeotti 1163 (lectotype, BR, designated by M. Nee, 1993; isolectotypes, BR, K). Solanuni dichotomum M. Martens & Galeotti, Bull. Acad. Roy. Sci. Bruxelles 12(1): 138. 1845. Type.

EXCLUDED SPECIES Mexico. Oaxaca: Yavezia, Galeotti 1227 (holotype, BR). Solanum martensii Dunal in DC., Prodr. 13(1): 140. 1852. Based on Solanum dichotomum M. Martens & Galeotti.

Solanumpabstii L. B. Sm. & Downs, Phytologia 10: 427. 1964. Type.Brazil, Est. SantaCatarina,Muns. Lajes and Sao Joaquim,Rio Lavatudo, 1050 m, 22 Oct 1961, Pabst & Pereira 6200 (holotype, US [US-2423786]; isotype, B). The following three species belong to a distinct species group distinguished from others in the nonspiny solanums by their unequal,pointed anthersand their unusualgrowth habit. In the absence of a formal sectional name for this group I have called it the Solanum thelopodium species group (see Knapp, 2000). Bitterproposed(in herbariumannotations)naming the taxa as a new genus in Solanum, but they fall well within the overall variabilityin Solanumandthis is not warranted.Membersof the groupalso have distinctiveovoid seeds with curiouslythickenedlateralcell walls somewhat like those of sect. Pteroidea (Knapp & Helgason, 1997; see Morphology).

367

opolis,"Schott5431 (W);Brazil,Sellows.n. (BM?T. SolanumcordovenseSess6 & Mofiio, Fl. Mex., in La Naturalezaser. 2, 2, Suppl.part5: 55. 1894. Type. Mexico. Veracruz:C6rdoba,Sesse & Moifio sn.. (MA?). SolanumdidynumDunal, Hist. Nat. Solan. 236. 1813. Type. Brazil. Based on descriptionin Vellozo, no herbariumspecimens known. Solanum didynumDunal var. subglabrum Dunal in DC., Prodr. 13(1): 125. 1852. Syntypes. BraziL Bowie & Cunninghams.n. (BM). Solanum didynumDunal var. tomentosumDunal in DC., Prodr. 13(1): 125. 1852. Syntypes. Brazil, Martius 1261 (G-DC [IDC microfiche 80061.2071:111.5],P [Mortonneg. 8184]; isosyntypes B [F neg. 2813]); Minas Gerais, 1841, Claussen 1428, no. 5 (P [Mortonneg. 8185]); Brazil,Sellowlt s.n. (not seen by Dunal). Solanum didynumDunal var. subvirgatumWitasek, Denkschr.Akad. Wissensch. Wien 79: 338. 1910. Type.Brazil.Est. Sao Paulo,betweenS. Amaroand BarraMansa,Itapecirica,800-900 m, Jun,Wettstein & Schiffners.n. (W?).

SolanlumdimorphandrumS. Knapp, Bull. Nat. Hist. Mus., London,Botany 30: 21. 2000. Type. Colombia. Magadalena:SierraNevada de Santa Marta, Valparaiso,"4500ft,"28 February1899,H H. Smith Solanum distichophyllumSendtn. in Mart., Fl. Bras. 10: 35, tab. 3, fig. 19. 1846. Syntypes. Brazil. 1190 (holotype, NY; isotypes, GH, K, MO, US). Amazonas: ["Prov.Rio Negro"], Maribi ditionis SolanumnionarchostemonS. Knapp,Bull. Nat. Hist. Japurensis,Martius s.n. (M [Mortonneg. 8682]); Mus., London, Botany 30: 23. 2000. Type. EcuaRio Amazonas,Poeppig (2613) (B [F neg. 2815], dor. Pastaza: Puyo, Comunidad Santa Cecilia, M [Mortonphoto 8415], P [Mortonneg. 8187]). Villano, 380 m, 1?30'S, 77?27'W, 1 May 1992, Palacios 10117 (holotype, QCNE; isotype, MO). SolanumedwardsiiStandl.,Trop.Woods37: 31. 1934. Type. Honduras.Comayagua:Temagua,2000 ft, 6 Solanum thelopodiumSendtn. in Mart., Fl. Bras. 10: Aug 1933, Edwards 639 (holotype, F [F neg. 46. 1846. Type. Brazil. Amazonas: "in sylvis ad 49327]; isotypes, A?, GH, K, NY, US). lacum Teffe, prope Rio Catual,prov. Rio Negro," November, Martius 2903 (lectotype, M [F neg. Solanum eriocalyx Dunal in DC., Prodr. 13(1): 124. 6545], designated by Knapp,2000). 1852. Type. Brazil, 1834, Lund 348 (holotype, GDC [F neg. 6788] IDC microfiche 800The following names refer to members of 61.2071:111.3).Not of Dunal, 1813. Solanumsect. Extensum,a distinctivegroupwith leafopposed inflorescences and copious stellate pubes- Solanum extensumBitter, Repert. Spec. Nov. Regni Veg. 13:94. 1914.Type.Panama.Darien:Paca,near cence. Members of this group also usually have enCana, 15 Apr 1908, Williams704 (lectotype, US, largedcalyces in both flower and fruit and soft, juicy, designated by D'Arcy, 1973; isotype, NY). brightly colored berries. The group is currentlyunder study by L. F. Carvalho (RB). SolanumgemellumMart.ex Sendtn.in Mart.,Fl. Bras. 10:28. 1846. Syntypes.Brazil.MinasGerais:CachoSolanum sect. ExtensumD'Arcy, Ann. Missouri Bot. eira do Campo, Claussen s.n.; Brazil, Sellowvs.n. Gard. 59: 268. 1972. Type species: Solanum extensuimBitter. Solanumgracillimum Sendtn. in Mart., Fl. Bras. 10: SolanumconcinnumSchottex Sendtn.in Mart.,Fl. Bras. 36, tab. 3, fig. 47-51. 1846. Type. Brazil, Sellow 10: 36, tab. 3, figs. 20-24. 1846. Syntypes. Brazil. s.n. (holotype, B [F neg. 2819]; possible isotypes, Rio de Janeiro: Serra Grande, "Prov. SebastianBM, K).

36 8

Solanum inelegans Rusby, Mem. TorreyBot. Club 4: 229. 1895.Syntypes.Bolivia.LaPaz:Yungas,10,000 ft, 1890, Bang 709 (BM, MO, NDG-n.v., NY, US, WIS). Bang 715 (BM, MO, NDG-n.v., NY, US). Solanum isodynanumSendtn. in Mart., Fl. Bras. 10: 33. 1846. Type. Brazil, Sellow s.n. (holotype B [F neg. 2826]; probableisotype, K). Solanium/undelliiStandl.,Publ.Field ColumbianMus., Bot. Ser. 8: 42. 1930. Type. Belize. Cayo Distr., RoaringCreek, Aug 1929, Lundell 324 (holotype, F [F neg. 49436]). Solanum megalochiton Mart.,Flora 21, 2, Beibl.: 63. 1838. Type. Brazil. Rio de Janeiro: "Sebastionopolis," Nov 1817, Martius s.n. (holotype, M [Mortonphoto 8713]).

FLORA NEOTROPICA

woods, Salseiro, 750 m, 15 Sep 1962,Klein?3006 (holotype, US; isotype, HBR-n.v.). The following two taxa belong to a distinct species group distinguishedfrom othersin the non-spiny solanumsby their axillary or terminalinflorescences, large, pentagonal flowers, and poricidal anthers not lengtheningto slits. In the absenceof a formalsectional name for this group I have called it the Solanuln havanense species group. havanense Jacq., Enum. Syst., pl. 15. 1760. Solanumn Type.Jamaica,Jacquin s.n. This species is found on Cubaand Jamaica,and was classified as a memberof the group"Meiomeris" by Dunal (1852). The showy purple flowers are very distinctive,but its relationships,otherthan to its clear sister species Solanumtroyanum,are unclear.

SolanummegalochitonMart.var. villoso-tomentosum Dunal in DC., Prodr. 13(1): 124. 1852. Syntypes. SolanurntroyanurnUrb., Symb. Antill. 5: 487. 1908. Syntypes.Jamaica:Troy,2000 ft, 8 May 1903,HarBrazil,Sellows.n. (BM);Brazil.Rio de Janeiro:Serra ris 8530; Troy, 1500 ft, 22 Apr 1904, IHarris8681; dos Orgaos,Lhotsky 164 (G-DC [IDC microfiche Holly Mount, Mt. Diablo, 3000 ft, 19 Aug 1905. 800-61.2071:111.2]); Sao Paulo, Lund 33 (G-DC 9000. Harris [IDC microfiche 800-61.2071:111.2]);Prov.Minas is clearlyrelatedto S. havanense, Solanumtroyanurn Gerais, Claussen 567 (G); Rio de Janeiro:Novo buthasmuchlargerflowersandleaves.Thebarkof older Friburgo,Claussen 198 (P [Mortonneg. 8247]). stems is shiny andwhite, ratherthangray andrough. SolanumperattenatumI. M. Johnst.,Sargentia8: 265, tab. 17, fig. 2. 1949. Type.Panama.Panama:Perlas The two species that follow, both Jamaican Archipelago,Gulf of Panama,SanJose Island,near endemics, are also obviously relatedto each other,but the Guard House, 26 Dec 1945, Johnston 909 their wider relationships are not clear. I have called (GH, MO, US). them the Solanumacropterumspecies group for simplicity, but resolution of their affinities awaits phyloSolanunipruriens Dunal in DC., Prodr. 13(1): 120. genetic analysis. 1852. Syntypes. Brazil. Sao Paulo: Jaragua,1815, acropternum Griseb.,Fl. Brit.W.Ind.437. 1862. Bowie & Cunninghams.n. (BM); circa S. Carlos, Solanumn Syntypes.Jamaica,Waterss.n.; Jamaica,Wilsons.fn. Lund 34 (G-DC). This is a species of wide distributionon Jamaica Solanum rufescens Sendtn. in Mart.,Fl. Bras. 10: 39. andis foundon limestoneatlow elevations.Itis verysimi1846. Syntypes. Brazil. Rio de Janeiro: Serra larto S. punctulatum,but differs in its glabrousleaves. d'Estrella, Pohl s.n. (M [Morton neg. 8736]); Brasilia australis, Sellow s.n. (B [F neg. 2848], K, SolanumpunctulaturnDunalin DC., Prodr.13(1): 122. P [Mortonneg. 8312]). 1852. Type. Jamaica,Wiless.n. (holotype, BM). The possession of lepidote scales in this species Solanum schwackeanum L. B. Sm. & Downs, is unusual and has led M. Nee (in litt.) to suggest that Phytologia 10: 428. 1964. Type. Brazil. Santa it is from subgen. Leptostemonumn. derived I-tshares Catarina:nearBlumenau,1884,Schwacke175 (hogeneralinflorescencemorphologyandflower structure lotype, US; isotype, R). with Solanum acropterum, however, and is probably

Solanum solum J. F. Macbr., Field Mus. Nat. Hist., closely relatedto it.Solanurnpunctulatumis anendemic Bot. Ser. 13(5-B,1): 220. 1962. Type. Peru. Junin: of cloud forests andof relativelyrestricteddistribution Hacienda Schunke, La Merced, Macbride 5764 on Jamaica. (holotype, F). Solanum subsylvestris L. B. Sm. & Downs, Phytologia 10: 429, t. 2, fig. 12-15. 1964. Type. Brazil. Santa Catarina: Canoinhas, border of

ACKNOWLEDGMENTS I wouldlike to thankall of themanypeoplewho have helped me duringthe course of this study,particularly

369

LITERATURE CITED

those who have guided and helped me to find elusive solanumsin the forests of Centraland SouthAmerica. Theirnamesaretoo manyto mentionhere,buttheirpatient help has been invaluable.This study would have neverbeen possiblewithoutthe unflaggingsupportand calm guidanceof my majorprofessor,the late Michael D. Whalen,andthe moralsupportof the late TimPlowman.Financialassistancewas providedby AAUW,the OTS Jessie Smith Noyes Foundation,the H.E. Moore Jr. Endowment Fund of the L.H. Bailey Hortorium, SigmaXi, the NSF DoctoralDissertationImprovement Program(BSR 8202773), the CornellUniversityAgriculturalExperimentStation,the DarwinInitiativeforthe Survival of Species of the U.K. government(Department of the Environment,Transportandthe Regions), andthe enhancementgrantof TheNaturalHistoryMuseum in London. The following institutionsand individuals helped with field work:TropicalScience Center,SanJose,CostaRica;Serviciode ParquesNacionales, San Jos6, Costa Rica; Organizationfor TropicalStudies, Costa Rica; RE.NA.RE., Panama; Smithsonian Tropical Research Institute, Panama; Ministerio de Agricultura,Ecuador;PontificiaUniversidadCat6licade Ecuador,Quito, Ecuador;HerbarioNacional de Ecuador, Quito, Ecuador;Ministeriode Agricultura,Peru; Museo Nacional de HistoriaNacional (JavierPrado), Lima, Peru;InstitutoNacional de Parques,Venezuela; UniversidadCentralde Venezuela,Maracay,Venezuela; Bob and KerryDressler, Robin Foster, Dave Roubik, Bob Schmalzel, Jan Salick, GerardoLamas M., Rina Ramirez,the late David N. Smith, Blanca Le6n, Olga Villagarcia,Sr.Turpo,PeterWilson,Leo Roth,Teresita Iturriagade Capiello,CarmenEmiliaBenitez de Rojas, andespecially JamesMallet.Duringthe preparationof my dissertation,fromwhich this monographis derived, MarciaWaterway,BethDickson,BrianChabot,andthe lateGeorgeEickwortwere helpfulcritics.I thankCarol Kalafatic,Bente StarckeKing, EdwardRooks, MargaretTebbs,andMickyLuckforillustrations,andthepublicationscited in the figurelegendsforpermissionto reproduce illustrationspreviously published elsewhere. MamenPefiatranslatedthe abstract;Mike Gilbertgenerouslyhelpedwith the formattingof the keys; the PhotographicUnit of The NaturalHistoryMuseum,especially HarryTaylorandPatHart,providedmuchneeded advice and assistance with plate production.The late William G. D'Arcy, Michael Nee, the late Tim Plowman, Carlos Ochoa, Jack Hawkes, Greg Anderson, LilianMentz,LuciaFreirede Carvahlo,andLynnBohs have all sharedinsights into variousaspects of the taxonomyandbiology of Solanumwithme. CarmenEmilia Benitez de Rojas, Lynn Bohs, and Michael Nee carefullyandthoroughlyreviewedthemanuscript; theircomments andcorrectionshave been invaluable.All errors or omissions,however,aremy own. Finally,I thankthe

curatorsof the following herbariawho have made genierousloans of theirspecimens,and/orhave allowedme to use theirfacilities:A, AAU, AMAZ,B, BH, BM, BR, CAS, CEPEC, COL, CR, CU, CUVC, CUZ, DS, ECON, F, FCQ, FUEL, G, GB, G-DC, GH, HAO, HUT, INB, K, L, LL, M, MA, MAD, MBM, MEXUJ, MO, MOL,MPU, MY,NY, P,POM,PY,QCA, QCNE, RB, RSA, SCZ, SI, TEX, U, UPCB, US, USM, VEN, W, WIS. Abbreviations are the standard ones of Holmgrenet al. (1990) andareusedthroughoutthetext.

LITERATURE CITED Anderson, G. J. 1975. The variation and evolution of selected species of Solanum section Basarthrunm. Brittonia 27: 209-222. . 1979. Dioecious Solanunmspecies of hermaphroditic origin is an example of broad convergence. Nature 282: 836-838. & P.G. Gensel. 1976. Pollen morphology and the systematics of Solanum section Basarthrurn.Pollen & Spores 18: 533-552. & D. A. Levine. 1982. Three taxa constitute the sexes of a single dioecious species of Solanulm.Taxon 3 1: 667-672. & D. E. Symon. 1989. Functional dioecy and andromonoecy in Solanum. Evolution 43: 2t)4-2 19. Barboza, G. & A. T. Hunziker. 1989. Estudios sobre Solanaceae. XXIX. Sinopsis taxon6mica de Athenaea. Bol. Soc. Argent. Bot. 26: 91-105. Bates, H. W. 1863. Contributions to an insect fauna of the Amazon valley. Lepidoptera:Heliconiidae. Trans. Linn. Soc. London 23: 685-687. Bawa, K. S. 1980. Evolution of dioecy in flowering plants. Ann. Rev. Ecol. Syst. 11: 15-39. & J. H. Beach. 1981. Evolution of sexual systems in flowering plants. Ann. Missouri Bot. Gard. 68: 254-274. Beccaloni, G. W. 1995. Studies on the ecology and evolution of Neotropical ithomiine butterflies (Nymphalidae: Ithomiinae). Ph.D. thesis, Imperial College, University of London. Bell, A.D. 1991. Plant form. Oxford University Press, Oxford. & T. D. Dines. 1995. Branching patterns in the Solanaceae. Pp. 157-172 in P. C. Hoch & A. G. Stephenson (eds.), Experimental and molecular approaches to plant biosystematics. Missouri Botanical Garden, St. Louis. Bernardello, L. M. & G. J. Anderson. 1990. Karyotypic studies in Solanumnsection Basarthruni (Sollanaceae). Amer. J. Bot. 77: 420-431. Bertin, R. I. 1982. The evolution and maintenance of andromonoecy. Evol. Theory 6: 25-32. Bitter, G. 1913. Solana nova vel minus cognita. VII. Repert. Spec. Nov. Regni Veg. 11: 481-491. . 1916. Solanaceae andinae. Bot. Jahrb. 54 Beibl. 119: 5-17. . 1919a. Solana nova vel minus cognita. XVII. Repert. Spec. Nov. Regni Veg. 16: 10-15. 1919b. Die Gattung Lycianthes. Vorarbeiten zu einer Gesamtschrift. Abh. Naturwiss. Vereins Bremen 24(2): 292-520. . 1920a. Solana nova vel minus cognita. XVIII. Repert. Spec. Nov. Regni Veg. 16: 79-103.

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FLORA NEOTROPICA of tomatoes, potatoes, and pepinos (Solanaceae). Amer. J. Bot. 80: 676-688. Sprague, T. A. 1926. Humboldt & Bonpland's itinerary in Colombia. Kew Bull. 1926: 23-30. Spruce, R. 1908. Notes of a botanist on the Amazon and Andes. A. R. Wallace (ed.). Macmillan, London. Stafleu, F. A. & R. S. Cowan. 1983. TL 2--Taxonomic literature. Vol. 4: P-Sak. Regnum Veg. 110. Stearn, W. T. (ed.). 1968. Humboldt, Bonpland. Kunth and tropical American botany. J. Cramer, Stuttgart. Steele, A. R. 1964. Flowers for the king. The expeditions of Ruiz and Pav6n and the flora ol Peni. Duke University Press, Durham. Steenis, C. G. G. J. van. 1981. Rheophytes of the world. Sijthoff & Noordhoff, Alphen aan den Rijn, The Netherlands. Steyermark, J. A. 1979. Plant refuge and dispersal centres in Venezuela: their relict and endemic element. Pp. 185-221 in K. Larsen & L. B. Holm-Nielsen (eds.), Tropical botany. Academic Press, London. . 1982. Relationships of some Venezuelan forest refuges with lowland tropical floras. Pp. 182-220 in G. T. Prance (ed.), Biological diversificatio-n in the Neotropics. Columbia University Press. New York. & 0. Huber. 1978. Flora del Avila. Sociedad Venezolana de Ciencias Naturales,Caracas. Symon, D. E. 1970. Dioecious Solanuins. Taxon 19: 909-910. . 1979a. Sex forms in Solanum (Solanaceae) and the role of pollen collecting insects. Pp. 385-397 in J. G. Hawkes, R. N. Lester & A. D. Skelding (eds.), The biology and taxonomy of the Solanaceae. Academic Press, London. . 1979b. Fruit diversity and fruit dispersal in Solanuin in Australia. J. Adelaide Bot. (3ard. 1: 321-3 3 1. Thomas, W. W., A. M. V. de Carvalho, A. M. A. Amorim, J. Garrison & A. L. Arbalaez. 1998. Plant endemism in two forests in southern Bahia, Brazil. Biodiv. Conserv. 7: 311-322. Tingey, W. M. & R. W. Gibson. 1978. Feeding and mobility of the potato leafhopper impaired by glandular trichomes of Solanumn berthauiltii and S. polvadeniumn. J. Econ. Entomol. 71: 856-858. Utech, F. H. & S. Kawano. 1975. Spectral polymorphisms in angiosperm flowers determined by differential ultraviolet reflectance. Bot. Mag. (Tokyo) 88: 9-20. Van Heurck, H. & J. Mueller-Argoviensus. 1870. Observationes botanicae. Antwerp and Berlin. Visquez Martinez, R. 1997. Fl6rula de las reservas biol6gicas de Iquitos, Peru. Monogr. Syst. Bot. Missouri Bot. Gard. 63. Vellozo, J. M. da Conceisao. 1829 [1825]. Florae fluminensis. Typographia Nacionali, Rio de Janeiro. . 1831 [1827]. Florae fluminensis icones 2: figs. 1-156. Senefleder, Paris. Vickers, W. T. & T. Plowman. 1984. Useful plants of the Siona and Secoya Indians of eastern Ecuador. Fieldiana, Bot. n.s. 15: 1-63. Wakhloo, J. L. 1975a. Studies on the growth, flowering and production of female sterile flowers as affected by different levels of foliar potassium in Solanum sisymbrifoliumnLam. I. Effect of potassium content of the plant on vegetative growth and flowering. J. Exp. Bot. 26: 425-432. 1975b. Studies on the growth, flowering and production of female sterile flowers as affected by

NUMERICAL LIST OF TAXA different levels of foliar potassium in Solanum sisymbrifolium Lam. IL. Interaction between foliar potassium and applied gibberellic acid and 6furfurylaminopurine. J. Exp. Bot. 26: 433-440. . 1975c. Studies on the growth, flowering and production of female sterile flowers as affected by different levels of foliarpotassiumin Solanum sisymbrifolium Lam. 111. Interaction between foliar potassium and applied daminozide, chlormequat chloride, and chlorflurecol-methyl. J. Exp. Bot. 26: 441-450. Wallich, N. 1828-1849. A numerical list of the plants in the East India Company's herbarium. London. (IDC 1049). Walpers, W. G. 1844. Solanaceae. In Repertorium botanices systematicae 3: 38-100. Watrous, L,. & Q. Wheeler. 1981. Polarization of character states. Syst. Zool. 30: 1-11. Weber, C. A. 1928. Georg Bitter (obituary). Ber. Deutsch. Bot. Ges. 46: 148-156. Whalen, M. D. 1979. Taxonomy of Solantim section Androceras. Gentes Herb. 11: 359-426. 1984. Conspectus of species groups in Solanum subgenus Leptostemontum.Gentes Herb. 12: 179-282. & G. J. Anderson. 1981. Distribution of gametic self-incompatibility and infrageneric classification in Solanini. Taxon 30: 761-767. & D. E. Costich. 1986. Andromonoecy in Solanutni.Pp. 284-302 in W. G. D'Arcy (ed.), Solanaceae: biology and systematics. Columbia University Press, New York. & C. B. Heiser Jr. 1981. Taxonomy ,-of Solanitnt section Lasiocarpa. Gentes Herb. 12: 4 1--1 2 9. Wheelwright, N. T. & G. H. Orians. 1982. Seed dispersal by animals: contrasts with pollen dispersal, problems with terminology, and constraints on coevolution. Amer. Naturalist 119: 402-413. , W. A. Haber, K. G. Murray & C. Guindon. 1984. Tropical fruit-eating birds and their food plants: a survey of a Costa Rican lower montane forest. Biotropica 16: 173-192. Wiley, E. 0. 1981. Phylogeny: the theory and practice of phylogenetic systematics. John Wiley & Sons, New York. Wille, A. 1963. Behavioral adaptations of bees for collecting pollen from Cassia flowers. Rev. Biol. Trop. 11: 205-210. Willson, M. F. 1983. Plant reproductive ecology. John Wiley & Sons, New York. Young, A. M. 1972. On the life cycle and natural history of Hyimenitisnero (Lepidoptera: Ithomiinae) in Costa Rica. Psyche 79: 284--294. . 1973. The life cycle of Dircenna relata (Ithomiidae) in Costa Rica. J. Lepidol. Soc. 27: 258-267. . 1974a. On the biology of Godyris zavaleta caesiopicta (Lepidoptera: Nymphalidae: Ithomiinae). Entomol. News 85: 227-238. 1974b. Notes on the biology of Pteronymia notilla (Ithomiidae) in a Costa Rican mountain forest. J. ILepidol. Soc. 28: 257-268. . 1974c. A natural history account of Oleria zelica pagasai (Lepidoptera: Nymphalidae: Ithomiinae) in a Costa Rican mountain forest. Stud. Neotrop. Fauna 9: 123-140. 1977. Notes on the biology of Hypothryis euiclea in Costa Rica (Lepidoptera: Nymphalidae: Ithomiinae). Pan-Pacific Entomol. 53: 104-113.

375 . 1978. Notes on the biology of the butterfly Hypoleriacassotis (Bates) (Nymphalidae: Ithomiinae) in Northeastern Costa Rica. Brenesia 14/15: 97-108. & M. W. Moffett. 1979. Studies on the population biology of the tropicalbutterflyMechanitis isthmia in Costa Rica. Amer. Midl. Naturalist 101: 309-319. Zavada, M. S. & G. J. Anderson. 1997. The wall arid aperture development of pollen from dioeciouls Solanum appendiculatum: what is inaperturate pollen? Grana 36: 129-134.

NUMERICAL LIST OF TAXA I. Solanum oblongifolium species group 1. Solanum clivorum S. Knapp 2. SolanumhypaleurotrichumBitter 3. Solanumoblongifoliumn Dunal 4. Solanum venosumDunal II. Solanum pseudocapsicum species group 5. SolanumdiphyllumL. 6. Solanum kleinii L. B. Sm. & Downs 7. Solanum malacothrix S. Knapp 8. Solanumpseudocapsicum L. 9. Solanumn spissifolium Sendtn. III. Solanum nudum species group 10. Solanum acuminatumRuiz & Pav. 11. Solanum aphyodendronS. Knapp 12. Solanumcaavurana Vell. 13. SolanumcampaniformeRoem. & Schult. 14. Solanuni cassioides L. B. Sm. & Downs 15. SolanumdaphnophyllumBitter 16. Solanumgertii S. Knapp 17. Solanum intermediumSendtn. 18. Solanum lasiopodiumDunal 19. Solanum nudumDunal 20. SolanumpseudoqulinaSt.-Hil. 21. Solanum reitzii L. B. Sm. & Downs 22. Solanum restingae S. Knapp 23. Solanum santosii S. Knapp 24. Solanumspirale Roxb. 25. SolanumsymmetricumRusby 26. Solanum tepuiense S. Knapp 27. Solanumtr-achytrichium Bitter 28. SolanumtrichoneuronLillo 29. Solanuin warmingiiHiem IV. Solanum leucocarpon species group 30. Solanum alatirameumBitter 31. Solanum corumbense S. Moore 32. Solanum leucocarpon Dunal 33. Solanum lindenii Rusby 34. Solanum oblongum Ruiz & Pav. V. Solanum nutans species group 35. Solanumn malletii S. Knapp 36. Solanumn m1icroleprodes Bitter 37. Solanuni nutans Ruiz & Pav. 38. SolanunixanthophaeumBitter 39. Solanuin voungii S. Knapp

376

VI. Solanum amblophyllum species group 40. Solanum amblophyllumHook. f. 41. Solanum barbulatumZahlbr. 42. Solanum laurifronsBitter 43. Solanumpsychotrioides Dunal 44. Solanum tovarii S. Knapp 45. Solanumvacciniiflorum Standl.& L. 0. Williams 46. Solanum validinerviumBenitez & S. Knapp VII. Solanum deflexiflorum species group 47. SolanumdeflexiflorumBitter 48. Solanum imberbeBitter 49. Solanum incomptumBitter 50. Solanum irregulare C. V. Morton 51. Solanum sieberi Van Heurck& Mull. Arg. VIII. Solanum arboreum species group 52. Solanum amnicola S. Knapp 53. SolanumanisophyllumVan Heurck& Mull.Arg. 54. Solanum arboreumDunal 55. Solanumfalconense S. Knapp 56. Solanumgoniocaulon S. Knapp 57. SolanumgratumBitter 58. Solanum laevigatumDunal 59. Solanum lucens S. Knapp 60. Solanumplowmanii S. Knapp 61. Solanum ramonense C. V. Morton & Standl. 62. Solanurmripense Dunal 63. Solanum roblense Bitter 64. Solanum tanysepalumS. Knapp IX. Solanum unifoliatum species group 65. SolanurmbahianumS. Knapp 66. Solanum bellum S. Knapp 67. SolanurmcyclophyllumS. Knapp 68. Solanum longevirgatumBitter 69. SolanummarantifbliumBitter 70. Solanum triplinerviumC. V. Morton 71. Solanum unifoliatuniS. Knapp X. Solanum robustifrons species group 72. Solanurmabitaguense S. Knapp 73. Solanunm cucullatumS. Knapp 74. Solanum heleonastes S. Knapp 75. Solanum robustifronsBitter 76. Solanum superbumS. Knapp XI. Solanum narcoticosmum species group 77. SolanurndasynleuronS. Knapp 78. Solanumfoetens S. Knapp 79. Solanum narcoticosmurm Bitter XII. Solanum nigricans species group 80. SolanurmcallianthurnC. V. Morton 81. Solaiuin cornifoliumDunal 82. SolanurmmatuirecalvansBitter

FLORA NEOTROPICA

83. Solanum nigricans M. Martens& Galeotti 84. SolanumochrophyllumVanHeurck& Mill. Arg. 85. Solanumplatycypellon S. Knapp 86. Solanum quebradense S. Knapp XIII. Solanum arenarium species group 87. Solanum arenarium Sendtn. 88. SolanumgnaphalocarponVell. 89. Solanumsmithii S. Knapp 90. Solanum tunarienseKuntze XIV. Solanum sessile species group 91. SolanumchlamydogynumBitter 92. SolanumconfertiseriatumBitter VanHeurck& Miill.Arg. 93. Solanummonadelphum 94. SolanumobovalifoliumBenitez 95. SolanumoppositifoliumRuiz & Pav. 96. Solanumpalmillae Stand]. 97. Solanum rovirosanumDonn. Sm. 98. Solanumsessile Ruiz & Pav. 99. Solanumtriste Jacq. 100. Solanum turgidumS. Knapp XV. Solanum confine species group 101. Solanumcapillipes Britton 102. Solanumconfine Dunal 103. SolanumcruciferumBitter 104. Solanum darienense S. Knapp 105. Solanum dissimile C. V. Morton 106. Solanum erosomarginatumn S. Knapp 107. SolanumhypocalycosarcumBitter 108.SolanumleptopodumVan Heurck& Miu-I.Arg. 109. SolanumleptorhachisBitter 110. Solanum morii S. Knapp 111. Solanum nematorhachisS. Knapp 1 2. Solanum ombrophilumn S. Knapp 13. Solanumpastillum S. Knapp 114. Solanumper-tenueStandl. & C. V. Morton 15. SolanumstipulatumVell. 116. SolanumtenuijiagellatumS. Knapp 117. SolanumtuerckheimiiGreenm. 18. SolanumvalerianumC. V. Morton& Standl. 119. SolanumyaniamonenseS. Knapp XVI. Solanum dolosum species group 120. Solanum dolosum S. Knapp 121. Solanumgonyrhachis S. Knapp 122. Solanum habrocaulon S. Knapp Incertaesedis 123. Solanumdelitescens C. V. Morton 124. Solanum lasiocladum S. Knapp 125. Solanum mapiricumS. Knapp New species added in proof 126. Solanum cordioides S. Knapp (Solanuin nudumspecies group)

I TNT OF FXSICCATAFE

LIST OF EXSICCATAE Abraham, A. A.. 9, 97 (32). Acevedo R., P. & D. Chinea, 2190 (19). Acevedo R., P. et al., 4482 (15). Ackland, J., 134 (8). Acosta Solis, M., 5320 (73); 5334 (92); 5459 (41); 5530 (3); 5625 (109); 5951 (3); 6120 (107); 6588, 6614 (41); 6673 (3); 6781, 6782, 6791 (41); 7784 (3); 7942 (41); 8081 (3); 8692, 8937 (41); 10140 (3); 10168 (41); 10379 (3); 10505 (4); 10579 (3); 11256 (41); 12610 (109); 13367 (3); 13534 (41); 13537 (3); 13665, 13789 (92); 13791 (109); 14426 (41); 14769 (37). Agostini, G., 21 (8). Agostini, G. & Farinas, 103 (112). Agra, M. F., 646, 1291 (12). Aguilar, 567, 588 (19). Aguilar Zepeda. J. A. & S. Martinez y Day, 334 (19). Aguilar, I., 82 (8); 494 (83); 505 (8); 691 (83). Aguilar, J.. 1602 (11). Aguilar, J. I., 1312 (83). Aguilar. R., 292, 1665 (97); 3826 (11); 4794, 4797 (19). Aguilar, R. & 0. Garrote, 4032 (45). Aguilar, R. et al., 2867 (118); 5481 (54). Ahumada et al., 697 (12). Alauy, R., 37, 89 (8). Albuquerque, B. W. & Lima (1970), 360 (95). Alexander, E. J., 1133 (83). Alexiades, M. & V. Pesha, 208 (75). Alfaro, E., 252, 265 (97); 310 (49); 816 (11); 1359, 1620 (45). Alfaro, E. & M. Segura, 1376 (61). Alfaro, E. et al., 1109 (61); 1208 (45); 1475 (11). Allard, H. A., 21607 (75); 21749 (93); 21767 (34). Allart, A., s.1l, 237, 237a, 327 (91); 447 (19). Allemao, F. Freire A. e Cysneiros, 1225 (32). Allen. P. H., 101 (19); 202 (36); 756 (54); 905 (48); 930 (54); 1199 (19); 1492 (11); 2088 (54); 5261, 5818 (19). Angulo, L., 515 (97). Anguto, 923 (3). Anon. Students at WIS, s.n. (89). Anthony. H. & G. Tate, 26 (109). Antonio C'., H., 53 (43). Antonio, T., 681 (45); 1502 (I 1); 3119 (54); 4124 (61); 4569 (36, mixed collection with 54). Antonio, T. & W. Hahn, 4332 (54). Apolinar (Apollinaire)-Maria, Bro., 31 (3); 33 (41); 44, 539 (81). Apollinaire. Bro. & Bro. Arthur, 39 (8). Aranda (coll. Nicaragua), 74 (19). Araquistain. M., 3195 (97). Araquistain, M. & P. P. Moreno, 2446, 2539, 2545 (19). Araquistain, M. et al., 1727 (19). Araujo, D., 25629 (8). Araujo, D. et al., 838 (115). Araya, F., 154 (54); 167 (97). Araya, F. & J. Corrales, 401 (54). Araya, F. et al., 255 (97). Arbo, M. M. et al., 1198 (12); 1199 (8); 6192 (12). Archer, W. A., 153, 510, 1613 (47); 2470, 2917 (32); 7806. 7819, 7837, 8255 (13). Arenas. 1238 (8). Arias, J. C., 39 (11). Aristeguieta, L., 2227 (13); 2529 (3); 2992, 3133 (57); 3672 (78); 3875 (94). Aristeguieta, L. & Foldats, 1504d (112).

Aristeguieta, L. & Pannier, 1959 (54). Ariste-Joseph, Bro., s.n. (81); s.n. 1919 (3); Jan 1920 s.n. (80); A115 (81); A119, A139 (3); A252 (81); A330 (8); A537 (80); A546 (81). Arsene, Bro., 2444, 3074, 8701 (19). Arvigo, R., 46 (19). Arvigo, R. & G. Shropshire, 255 (19). Arvigo, R. et at., 503 (19). Ascencio R., J. et al., 1321 (5). Asplund, E., 5746 (92); 5888 (41); 6104 (8); 6203 (3); 6321 (41); 7430 (3); 7845 (11); 16164 (4hl; 17471 (3). Atwood, C., 2644 (5); 3236, 5178 (19). Aubreville, A., 75 (32); 191 (13). Aulestia, C. & E. Aulestia, 895 (19). Aulestia, C. & M. Aulestia, 1368 (67). Aulestia, C. & A. Grijalva, 1172 (67). Aulestia, C. et al., 420 (67). Aulestia, M. & J. Andi, 391 (53). Avendano R. et al., 150 (19). Avendano, S. & Calzada, 411 (11). Aviles, S., 17 (54); 72 (19). Ayala, F., 231, 622 (95); 2009 (19); 2325 (95). Ayala, F. et al., 2868 (32); 3287 (95). Aymard, G., 405 (54); 2387 (19); 4468 (51). Aymard, G. & N. Cuello, 6580, 6593 (19). Aymard, G. et al., 2627 (51); 3629, 4509 (19); 6400 (95); 11388 (75). Azofeita, A., 398 (54). Badillo, V., 1911, 3848 (94); 5092, 5096 (46); 6804, 6904, 6906, 6911, 6920, 6937, 6940 (78); 7256 (1 ). Badillo, V. et al., 7313 (99). Baenitz, C. G., 4 May 1902 (8). Bahia, R. P., 97 (32). Bailetti, E., 21 (8). Bailey, L. H., 682 (5). Bailey, L. H. & E. Z. Bailey, s.n., 292, 377 (91); 540 (54); 716 (91); 765 (8); 1000 (29); 1060 (13). Baker, C. F., 77 (13). Baker, M. et al., 5512 (19). Baker, R. E. D., [in herb. Trinidad 14438], 14643 (51). Baker, R. E. D. & N. W. Simmonds, 14337, 14357 (54). Balansa, B., 2097, 2099, 2100 (8); 2122 (12); 2122a, 2123, 3126 (25); 3751 (24). Balee, W., 2666 (32). Ball, J., s.n. coll. 1882 (40). Balls, E., 4312 (11). Balslev, H., 1021 (41); 2386 (108); 2790 (41). Balslev, H. & R. Alarc6n, 3062 (108). Balslev, H. & E. Madsen, 10281 (75); 10507, 10559 (19). Balslev, H. et al., 62282 (53); 84536 (35). Bang, A. M., 972, 1096, 2511 (8). Bang, M., s.n. (8); 712 (10); 1119 (82); 1478 (25); 1526 (mixed collection 10 & 32); 1630, 1931 (84); 2188 (90); 2250 (33); 2524 (37); 2525 (32); 2641 (33). Barbour, K. & W. Johnson, 95078 (19). Barbour, K. et al., 94091 (19). Barbour,P., 2699 (72); 2854 (102); 2871 (72); 3595 (58). Barchet, J. P., 507 (8). Barclay, A S., 3748 (3). Barclay, A. S. et al., 3168 (81); 3232 (80); 3599 (Il). Barclay, G. W., 2172 (5). Barclay, H. G. et al., 7926 (3). Barfod, A., 41078 (19); 41441, 41601 (92). Barfod, A. & H. Balslev, 41321 (2). Barfod, A. et al., 48042 (19). Barkley, F. A., 19AnI15 (47).

378 Barkley, F. A. & Gutierrez V., 1916 (19). Barr & Wiedhopf, 66-159 (5). Barrier, S., 1903 (95). Barringer, K., 3026 (19). Barringer, K. & Christenson, 3304 (11). Barringer, K. & J. G6mez-Laurito, 2570 (97). Bartlett, H. H., 12706, 12962, 13042 (19); 20257 (8); 20346 (25); 20944, 20944a, 21097 (8). Bartlett, H. H. & T. Lasser, 16740 (54); 16778 (48). Bastion, E., 1283 (8). Bastos, A. M. et al., 138 (13). Bayliss, R. D. A., 4594 (8). BD (at MPU, coll. 1986), 190 (32). Beainan, J., 5156, 5204 (19). Beck, S. G., 3601 (11); 3887 (10); 4464 (90); 4852 (33); 6813, 7268 (82); 7811, 8606 (10). Beck, S. G. & G. Liberman, 9808 (8). Belem, R. P., 1482 (13); 1753 (12). Belem, R. P. & Magalhaes, 853 (13). Belem, R. P. & R. S. Pinheiro, 2848 (13). Bell, D. & S. Wiser, 88-35 (75). Bell. P. R., 440 (11); 573 (41). Bello, E., 1082, 1719 (54); 4966 (97); 5154 (113); 5402 (11). Belshaw, C. M., 3130a (98). Benitez de Rojas, C. E., 272 (54); 851 (99); 928 (19); 1143 (54); 1440 (19); 1478 (78); 1666 (54); 1703 (51); 1780 (19); 1800 (91); 1816 (51); 1977 (11); 2027 (3); 2235 (19); 2401 (98); 2507 (57); 2682 (78); 2746 (94); 2748 (54); 2833 (91); 2924 (19); 3139 (94). Benitez de Rojas, C. E. & G. Ferrari, 1373 (51). Benitez de Rojas, C. E. et al., 5122 (112). Bennett, B., 3468 (19); 4450, 4484, 4499 (98). Bennett, B. et al., 4443 (19). Benoist, M. R., 2021, 2503 (3); 3633 (2). Bensman, R., 183 (108). Berlandier, J. L., 141 (5). Berlin, B., 468 (98); 538 (95); 601 (98); 699 (95); 926, 1508 (98); 1859 (53). Bernal, R. et al., 1178 (97). Bernardi, A. L., 15 Feb 1957 (54); 2-3-1957 s.n., 411 (19); 1255 (86); 3323 (91); 6722, 6748 (13); 10533 (45); 10894, 10927 (3); 18278 (12); 18403 (8); 18708, 19088 (12); 19336 (25, mixed with 12). Bernardi, A. L. et al., 16838 (33); 17182 (62). Bernoulli, C. G., 944 (19). Berrios, J. M., 150 (19). Berry, P., 3654 (3). Bertoni, M. de B., 878 (8); 2636 (20); 3124 (27). Besse, L. et al., 114 (92); 2312 (37). Besserra, 437 (13). Betanicur,J. & S. Churchill, 2058 (47). Betancur, J. et al., 5204 (35). Billict, F. & B. Jadin, 1090, 1125, 1578 (32); 6179 (98). Bird, R. 1191, (37). Birschel, Dec 1854 (19). Bis Ram, 313, 2315 (8). Bittner, J., 1537 (97); 1899 (11). Bittner, J. & B. Venschott, 1936 (97). Black, G. A., 56-18930 (13). Blackman, C. et al., 170287 (19). Blackmore, S. & M. Chorley, 3704 (19). Blanchet, J. S., s.n., 48, 1169, 3537 (12). Blanico, C., 103 (13). Blum, K. E., 1363 (54). Bocher, T. W. et al., 212 (92).

FLORA NEOTROPICA Boeke, J. D., 711 (4); 892 (cf. 80); 1246 (98); 1892 (37); 2089 (82). Boeke, J. D. & S. Boeke, 3191 (82). Boeke, J. D. & J. Jaramillo,2485 (37); 2530 (2); 2596 (37). Boeke, J. D. & H. Loyola, 2179 (109). Boeke, J. D. & J. B. McElroy, 310 (41). Boelcke, 0. et al., 5437 (25). Boerboom, J., LBB-8750 (32). Boffa, P., s.n. (8). Bohs, L., 2027 (11); 2076 (8); 2421 (19); 2424 (11); 3010 (54). Bolland, G., s.n. (12). Bonpland, A. J. A., s.n. (3); 4488 (5). Boom, B. M. & D. Gopaul, 7290 (32). Boom, B. M. & M. Pacheco, 8518 (13). Boom, B. M. & G. J. Samuels, 9095, 9193 (32). Borges, s.n. 13 Feb 1994 (13). Borges et al., 34 (12). Borsboom, N. W. J., LBB-12047 (32). Botteri, M., s.n. (11); 844 (8); 848, 862 (11); 1094, 1357 (19). Botteri, M. & Sumichrast, s.n., 1857 (11). Bourgeau, E., 240 (11); 2405, 2408 (19). Bovini, M. et al., 813, 814 (115). Bowie, J. & A. Cunningham, May 1815 s.n. (9); Nov 1815 s.n. (20). Boyle, B. et al., 2671 (11). Boysen Larsen, B. & B. Eriksen, 45092 (41). Boysen Larsen, B. et al., 45513 (19). Brade, A. C., 18 Nov 1928 (12); 7019 (9); 9207 (88); 11315 (115). Brade, A. C. & A. Duarte, 20138 (6). Braga, P. I. S., 2393 (12). Braga, P. 1. S. & D. Sucre, 312 (115); 1502 (12). Brandbyge, J. & E. Asanza C., 30016, 30038, 30346a (98); 31669 (95); 32745 (98). Brandbyge, J. & F. Herrera, 42034 (4). Brandbyge, J. et al., 33378, 33621 (95). Brant, A. & F. J. Roldan, 1473 (32). Breedlove, D. E., 5060 (83); 6121 (11); 6596 (19); 6796, 7060 (83); 7478 (11); 8518 (83); 8948 (11); 9078 (83); 9718 (11); 9855 (mixed collection II & 19); 10273 (19); 10804 (83); 11036, 11363 (19); 11651 (mixed collection 11 & 19); 11697, 11797 (19): 12380 (83); 14733 (11); 14914 (19); 15269 (83); 15368, 15388 (11); 19896, 20343 (19); 23042, 24386 (83); 25248 (5); 25769 (83); 26133, 26500, 27001 (19); 27285 (5); 27950 (19); 28398 (5); 29055, 29066 (19); 35149 (11); 36859 (5); 42762 (83): 42767 (11); 47337, 50313, 51992, 52159, 56178 (19). Breedlove, D. E. & F. Almeda, 56837 (11); 58283 (83). Breedlove, D. E. & E. McClintock, 34269 (19). Breedlove, D. E. & P. H. Raven, 12927 (83); 12998. 13590, 14145 (19). Breedlove, D. E. & R. H. Thorne, 21173 (19). Brenes, A. M., s.n. Apr 1902 (117); 3947, 4063 (11); 4268, 427 1, 4346 (117); 4856 (54); 5432 (I I ); 6261, 6723, 12204, 12214, 12656, 13146 (97); 14384 (117); 15012, 16859, 21434, 21974, 21978 (97); 22012 (11); 22013 (61); 22615 (113). Breteler, F. J., 3664 (59); 3954 (32); 4410 (59). Bretting, P. K., 13 (11). Breyer-Brandwyk, M., Jan 1937 s.n. (8). Brigada Dioscoreas, 3850 (19). Briolley, 8998 (11). Bristan, N., 1445 (54); 1482, 1483 (48). Bristol, M. L., 1081 (43); 1259 (37).

LIST OF EXSICCATAE Brito & da Vinha, 5 (13); 182, 250 (12). Britton, N. L. & S. F. Earle, 6559 (19). Britton, N. L. & P. Wilson, 120 (19). Britton, N. L. et al., 1339, 1636 (54); 1641 (99); 2521 (101); 15466 (19). Broadway, W. E., 19 Dec 1891 (8); 17 Jul 1907 (51); 2 (8); 763 (8); 4142, 4271, 5362 (54); 7443 (8); 7909 (101); 9028 (51); 9847 (54). Brokaw, N. V. L. & M. Schulze, 31 (19). Brooke, W. M. A., 5123 (8); 5427, 6179 (82); 6908 (10). Brown, C. A., 1492 (8). Brown, K. S., 26 Jun 1967, 001, 200 (13). Brown, K. S. et al., 11 Mar 1983 (12). Bruijn, J. de, 1178 (48). Brunner, D. R., 1807 (20). Buchtien, O., 315 (8); 325 (10); 462 (84); 464 (10); 760 (84); 1273, 1274, 1275, 1435 (125); 1876 (33); 4166 (90); 5854 (10); 9030 (82). Bunting, G. S., 3304, 4710 (112). Bunting, G. S. & K. Kauffman, 10224 (51). Bunting, G. S. & R. Le6n, 12858 (36). Bunting, G. S. & Licht, 638 (19). Bunting, G. S. et al., 11984, 12026 (48); 12095, 12277, 12498 (19). Burch, D., 5948 (83). Burchell, W. J., s.n. (20); 898 (12); 972A-2 (13); 1078 (12); 1345 (87); 1872 (20); 2213 (13); 4475 (20); 9663, 9842-2, 9995-3 (13). Burger, W. L., 4156 (97). Burger, W. L. & T. Antonio, 10955 (97); 10963 (54); 11137, 11225 (97); 11246 (54); 11280 (97). Burger, W. L. & Baker, 9580 (117). Burger, W. L. & K. Barringer, 11534 (45). Burger, W. L. & M. Burger, 8085 (54); 8186 (49); 8442 (54). Burger, W. L. & G. Matta U., 4465, 4520 (97). Burger, W. L. & Ramirez B., 3986 (19). Burger, W. L. & R. Stolze, 4942 (97); 5751 (54). Burger, W. L. et al., 9419 (97); 11400, 11454 (45). Burkart, A. E., 14687 (20). Burnham, R. J. & R. A. Spicer, 124 (19). Busey, P., 652 (97). Bush (Haiti 1917), 1370 (19). Cabral, 68 (27). Cabrera, A. L., 1509 (8); 4221 (25). Cabrera, A. L. et al., 22299 (8); 22320 (25). Cabrera, E., 471 (19); 3784 (83); 12349, 16435, 16458 (19). Cabrera, E. & H. de Cabrera, 2528, 3107, 4748, 4777, 5198 (19); 5998 (83); 11421 (19); 14755 (5). Cabrera, E. & Cortes, 225 (19). Cabrera, E. & Durran, 471 (19). Cabrera, E. et al., 5682, 16536 (19). Cabrera, 1. & E. Hoyos, 2769 (19). Calder6n. S., 25, 1344, 1700, 1958 (5). Callejas, R., 3164 (43). Callejas, R. & 0. Escobar, 3304 (47). Callejas, R. et al., 2500, 2505 (11); 2842 (54); 2942 (11); 3714 (54); 3843, 3850 (43); 3918 (4); 3921, 3943 (47); 4209, 4289 (32); 4775 (36); 4885 (54); 5091, 5156 (48); 5267 (54); 5801 (48); 8511 (47). Calzada, J. I., 251, 2424 (11); 4115 (19); 4896 (5); 5745 (83); 12491 (5). Calzada, J. I. & R. Delgado, 4180 (83). Calzada, J. I. et al., 2771 (19); 3711 (11); 18749, 18925 (19). Camp. W. H., 2704 (3).

379 Campos, G. et al., 2865 (19). Campos V., A. et al., 349 (8). Carauta, J. P. P., 1223 (8); 1269 (13). Cardenas, M., 731, 732 (8); 1198 (98); 3117 (82); 3261 (90); 3481 (15); 5118 (11); 5601 (15). Cardona, F., 553 (8); 572 (99); 1300 (95). Carleton, M. A., 459 (19). Carlson, M. C., 2613, 2629 (83). Carnevali, R., 2857 (8). Carpenter, J., 259 (19). Carranza, E., 3334 (8). Carvajal, A., 348 (11). Carvalho, A. de & Gatti, 809 (22). Carvalho, A. de & G. Lewis, 943 (12). Carvalho, A. de et al., 1188 (13); 1291 (12). Carvalho, L. D'A. F. de, 560 (88). Casaretto, G., 1845 (12). Castaneda, R., 2458 (3). Castillo C., G., 205 (19). Castillo C., G. et al., 232, 1827, 1942 (83). Castillon, L., s.n. [herb. Lillo 87881] (123); 2019 (8). Castro, A. J. & E. Nunes, 7120 (12). Castro, R., s.n., 52341 (8). Castroviejo, S. et al., 10631 (58). Cavalcante, F. S. & F. Bruno, 10827 (12). Cazalet, P. C. D. & T. D. Pennington, 11 Jun 66, 5297 (109); 5470, 5511 (2); 5591 (41); 7644 (19). Cedillo T., R. & D. Lorence, 1272 (83). Cedillo T., R. & R. Torres C., 1208 (83). Cer6n, C. (= Cer6n M., C. E.), 260 (19); 879 (98); 1004 (75); 1034 (98); 1394 (35); 1541 (3); 2647 (72); 5955 (98); 6028 (19); 7172 (120); 13273 (41). Cer6n, C. & G. Benavidez, 2540 (37). Cer6n, C. & M. Cer6n, 3051 (108); 3072 (53). Cer6n, C & F. Coello, 3182, 3298 (35). Cer6n, C. & M. Factos, 7576 (53). Ceron, C. & F. Hurtado, 6517 (102). Ceron, C. & C. Iguago, 5425 (53); 5503 (98); 5700 (120); 8611 (73). Cer6n, C. et al., 2183 (53); 5746 (120); 5931 (73), 7908 (109); 13228 (41). Cerrate, E. et al., 251 (40). Chac6n, A., 1090 (97). Chacon G., I., 784 (54); 1074 (97). Chagas e Silva, F., 1410 (21). Chagas e Silva, F. et al., 7802, 20268 (21). Chan, C., 2414 (19). Chardon, C. E., 107 (51). Charles, S., 775 (83). Charpin, A. & F. Jacquemond, AC13510 (3). Chase, A., 7535 (5). Chatterjee, A. C., s.n. (24). Chavarria, U., 659, 871 (19). Chavelas P., J., ES-1243 (5). Chavelas P., J. & L. A. Perez J., 281 (8). Chavez, C., 455 (61); 456 (97). Chazaro, M. & P. Camarillo, 3877 (19). Chazaro, M. & R. Chazaro, 6517 (83). Chazaro, M. & L. Robles, 3853 (83). Chiao, C. Y., 11 Aug 1927 (8). Ching, R. C., 6241 (8). Ching-en Chang, 9259 (5 - cult.). Choc & Stewart, 43 (19). Choussy, F., 1602 (5). Christensen, E., 33-4 (11). Chrostowski, M. S., 70-399 (102). Chuah, M. S., 130 (32).

380 Chung, H. H., 8031 (8). Churchill, S. et al., 15872 (11). Cid, C. A. & B. W. Nelson, 2788 (95). Cid, C. A. et al., 3917, 4964, 5006, 6275 (32); 7363 (95). Clark, H., 7246 (95). Clark, J. L. & T. Nfiez, 1558 (92). Clark, J. L et al., 1094 (53); 1402 (3). Clarke, C. B., 4830, 8008, 15840B, 15840C, 17271, 37217, 40047D, 44429D, 44429E, 44429F (24). Clarke, D. & B. Hoffman, 617 (32). Claude Joseph, Bro., 3829 (8). Claussen, P., s.n. (17, 115); 5 (13); 7,1440 W-7 (17). Clemente, Bro., 6483 (19). Clewell, A., 3997 (83). Clubbe, C., s.n. 20 Aug 1980 (101). Cochrane, T. S., 12582 (83). Cochrane, T. S. et al., 8596 (5); 10626 (83); 10627 (11). Cogollo, A. et al., 3466 (120); 3667 (11); 3858 (120). Colchester, M. E. M., 2506b (32). Collenette, C. L. in J. Riley, 473 (32). Colombo, L. R. et al., 20293 (16). Conrad, J. & Conrad, 2986a, 2988 (19). Conrad, J. & W. Dietrich, 2238 (8). Conrad, J. et al., 2795 (5). Contreras, E., 135 (19); 225 (5); 1305, 1410 (83); 2649 (5); 4454 (19); 5170 (83); 7829, 7923 (117); 8376 (83); 11126, 11456 (19). Conzatti, C. et al., 2470 (83). Cook, A. Carter, 579 (8). Cook, 0. F. & Doyle, 375 (19). Cook, 0. F. & G. B. Gilbert, 329, 587, 629, 796 (82). Cook, 0. F. & Griggs, 146, 492, 543 (19). C'ooper, G. P., 5868 (11). Cooper, G. P. & G. Slater, 39 (97). Cooper, R. & N. Nickerson, 6167 (8). Cooper, R. E., 4367, 4723 (24). Cordeiro, I. et al., 8 (32). Cordero, G., 241 (97). Cordoba, J., 284 (97); 468 (19). Core, E., 1529 (19). Cornejo, X., 397 (109); 1191 (11). Correa A., M., 1265 (11). Correa A., M. & Lazor, 1480 (11). Correa A., M. et al., 2983 (54). Correll, D. S. & H. B. Correll, 12545 (8). Cory, V. L., 49995 (8). Costa, M. A. S. et al., 64 (19). Costa Sacco, J. da, 144 (8); 391 (87). Costich, D. E., 1074 (32); 1078 (95); 1082 (13). Costich, D. E. & D. Baldwin, 1501, 1502, 1510 (83); 1518, 1529 (19). Costich, D. E. & M. D. Whalen, 1078 (32). Coulter, T., 1238 (19). Coveny & Hind, 10572 (24). Cowan, C. P., 3388 (19); 3919 (83). Cowan, C. P. & Espinosa, 1492 (5). Cowani, C. P. & Mazana, 2307 (5). Cowan, R., 2307 (5). Cowan, R. & J. C. Lindeman, 39207 (32). Cowgill, W. H.. 1086 (8). Cremers, G., 3834, 3934 (95); 4291 (32). Cremners,G. et al., 9688 (32). Cristobal, C. L. & A. Krapovickas, 2162 (12). Cristobal, C. L. et al., 1257, 2089 (8); 2115 (12). Croat, T. B., 917, 2471 (19); 6680, 6768 (54); 8958 (19); 9352, 10119 (54); 10465, 10548, 10573, 10626 (11); 11151, 12072 (54); 12455, 13331, 13437, 13940,

FLORA NEOTROPICA 14541 (19); 14851, 15002, 16610, 16731, 16781, 17373 (54); 17489 (95); 18039, 18051 (98): 18632 (95); 19372 (98); 20047 (95); 20152, 20205 (19); 20690, 20817 (98); 21396 (19); 22117 (54); 22742 (97); 22960, 23107, 23198, 23397, 23533, 23533a, 23551, 23903, 24129, 24224, 24430, 24609, 24710, 24830, 24950, 25420 (19); 26208 (54); 33.407 (19): 34156 (11); 35078 (118); 35765 (54); 36422 (19): 37597, 37961 (54); 37963 (104); 38711 (19); 38805 (3); 38827 (109); 39320 (19); 39593. 39627 (5); 39711, 40156, 40273, 40356, 40372, 40375 (19); 40379 (79); 40379 (19); 40482 (83); 40572 (19); 40737 (83); 40993 (79); 41057 (83); 41375 (19): 41743 (83); 41870, 42308 (19); 42332, 43055 (11); 43517 (97); 43541, 44484, 45256, 46671, 47100 (11); 47306 (83); 47325 (mixed collection 19 & 83); 48665, 48740 (61); 49481 (72); 49635 (98); 49712 (72); 49841 (75); 49879 (61); 50700 (92); 51128 (98); 51706 (10); 57850 (98); 60666, 60718 (19); 60742 (54); 61609 (41); 69564, 69658 (47); 71254 (19); 71312 (67); 72494 (95); 72946 (11); 72963 (92); 73294 (3). Croat, T. B. & D. P. Hannon, 63804, 64047B (11); 64047C (83); 64246 (11). Croat, T. B. & D. M. Porter, 15507 (36): 15998 (11); 16236, 16298 (54). Croat, T. B. & J. Watt, 70578, 70584 (47). Croat, T. B. & G. Zhu, 76680 (19); 77057 (54). Crosby, M., 11439 (61). Cruz, J. S. de la, 2092, 2147, 2541, 2585. 2773, 2872, 3678, 3860, 4428, 4496 (32). Cuamacas, B. et al., 138 (73); 146 (37). Cuatrecasas, J., 83, 93 (81); 240 (3); 1147 (80); 1162, 1285 (3); 3372 (81); 5237 (3); 5370 (81); 5467 (3); 5689 (81); 5695 (80); 6700 (3); 7394 (98); 7898 (80); 8234-A, 8238 (58); 8915 (98); 9041 (53); 9210, 9354-B (4); 10388 (3): 10661, 10695 (98); 10698, 10784-A (108); 10878 (80); 10978 (108); 11476, 11608, 11788 (43); 13161 (59); 13819 (11); 14526 (19); 14678 (4); 15576, 16343 (19); 18063, 18078 (43); 18674 (47): 18912, 19147, 19147a (43); 19149 (58); 19638 (19); 20023 (58); 20150 (4); 20218 (43); 20429 (4); 206t)6, 21795, 21839 (43); 23002 (19); 23538 (43); 23897 (120). Cuatrecasas, J. & Cuadros, 28837 (11). Cuatrecasas, J. & H. Garcia Barriga, 9728 (3). Cuatrecasas, J. & R. Jaramillo, 12006 (3). Cuatrecasas, J. & E. Perez Arbelaez, 6070 (11). Cuatrecasas, J. & R. Romero Castaneda, 24353, 25386 (54); 24801 (18). Cuatrecasas,J. et al., 12596 (3); 12764 (19); 26959 (58). Cufodontis, G., 195 (54). Cumana, 1610 (99). Curran, H. M., s.n. (8); 28 (20). Custodio Filho, 305 (20). Cutler, H. C., 4063, 7685, 7689 (8). D'Arcy, W. G., 3988 (19); 4244, 4249 (11); 4296, 5208, 5210 (54); 5243 (19); 5313, 5325, 5374, 5402,5406 (11); 5431 (19); 5443 (11); 5943 (54); 6492 (11); 9725 (48); 9863 (11); 10425 (5); 10691, 10915 (45); 10958 (11); 11076 (45); 11118 (54); 11248, 14029, 14046, 14049 (19); 14077 (98); 14617 (19); 14835 (3); 14872, 14892 (19); 14990 (61); 15012 (19): 15057 (61); 15918 (11); 18000 (83). D'Arcy, W. G. & J. D'Arcy, 6093, 6722 (19). D'Arcy, W. G. & R. Dressler, 5493, 5506 (54). D'Arcy, W. G. & B. Hammel, 12270 (19).

LIST OF EXSICCATAE D'Arcy, W. G. & K. Sytsma, 14327 (36); 14346, 14503, 14530 (54); 14667 (19). D'Arcy, W. G. & R. 0. Woodbury, 1710 (19). D'Arcy, W. (G.et al., 12644 (61); 12737 (49); 13063 (11); 13349 (19); 15577, 15579, 15599 (47); 15918 (61). Dahlgren, B. E. & E. Sella, 387, 655, 716, 753, 772, 782 (13). Daly, D. C. & P. Acevedo R., 5176 (69). Daly, D. C. et al., D448 (13); D523 (32). Daniel, Bro., 971 (8); 1697 (3); 2105 (47); 3386 (54); 3415 (3). Darwin, S. P., 1210 (8). Daubenmire, P., 117 (19). David, 4852 (1 1). Davidse, G., 4036 (64); 9684, 36352 (19). Davidse, G. & A. Gonzalez, 18863 (54); 19541 (124). Davidse, G. & G. Herrera,31173, 31325 (54); 31331 (97). Davidse, G. & J. S. Miller, 27148 (95). Davidse, G. & R. Pohl, 1687 (54); 1694 (63). Davidse, G. & S. L. Tillett, 4059 (64). Davidse, G. et al., 5748 (53); 10955 (13); 11066 (8); 16724 (94); 25488 (45); 28289, 29907 (19); 37278 (19). Davidson, C., 296 (11); 3435 (98); 4767a (84); 4889 (33); 5277 (95). Davidson, C. & J. Jones, 9055 (82); 9367 (33); 9613 (95). Davidson, M. E. S. (Mrs. Terry), 99, 446 (49). Davis, E. W. & E. Turner, 721 (3). Davis, E. W. et al., 785 (108); 1777 (82). Davis, T. III, 571 (97); 733, 785 (5). Dawe, M. T., 195 (81). Dawson, G., 651 (8). Deam, C. C., 92 (19). Degen, R., 2088 (8). Degener, 0., 7372, 7373, 7374, 35981 (8). Degener, 0. & H. Weibke, 2123 (8). Delascio Ch., F., 9206 (8). Delascio Ch., F. & R. Liesner, 7199 (13). Delgado S., A. & R. Hernandez, 88 (8). Delomy, 4718 (24). Dereims, s.n. (90). Descoings. B. & Ch. Luu, 20215, 20230 (32). Dias (coll. 1830), 2093 (98). Diaz (coll. 1831), 2476B (108). Diaz et al., 589 (95). Diaz, B., 324 (47). Diaz P., S., 2709 (32); 2709 (19). Diaz S., C., sn. (40). Diaz S., C. & S. Balde6n, 2197 (82); 2277 (93); 2306 (98). Diaz S., C. & S. Huanqui, 1995 (82). Diaz S., C. & N. Jaramillo, 12 (95). Diaz S., C. et al., 2772 (60); 4110 (35); 4507 (98). Dick, C., 111 (32). Dickson, 1290 (19). Diero, 381 (25). Diggs, G. & M. Nee, 2425 (19). Diggs, G. et al., 2726 (19); 3973 (11). Dik, A., 194 (35); 236, 524 (98). Dillon, M. O., 2513 (40). Dillon, M. 0. & A. Sagastegui A., 6080 (82). Dillon, M. 0. & 1. Sanchez Vega, 6317 (60). Dillon, M. 0. & M. D. Whalen, 4064 (82). Dillon, M. 0. et al., 1180 (82); 4434 (11); 4438 (73); 6131 (39); 6432, 6491 (41). Dlouhy, C. & J. Fernandez Casas, 7673 (8). Dobereiner & Tokarnia, 749 (8). Dodge, C., 6197 (97).

381 Dodge, C. & Thomas, 6185, 6383 (97). Dodge, C. et al., 6380 (117); 9080 (73). Dodson, C. H., 5177 (19); 5833 (92); 5933, 13660 (109). Dodson, C. H. & P. M. Dodson, 11304, 11365, 11501 (92); 11617 (8); 11854 (73); 11880, 11979 (92). Dodson, C. H. & A. Gentry, 9541 (92); 9568 (109); 10005, 10085 (92); 10238 (11); 10345 (109); 12243 (11); 12637 (92); 12804 (109). Dodson, C. H. & McMahon, 5069 (92). Dodson, C. H. & Tan, 3531 (92). Dodson, C. H. & L. B. Thein, 512 (92); 629 (3); 635 (11); 1277, 1648, 1794 (92). Dodson, C. H. et al., 7994, 8659 (92); 9080 (73). Dombey, P., s.n. (37). Donselaar, J. van, 2813 (32). Doppelbauer, H. & U. Doppelbauer, 513 (82). Dorantes, J., 396 (5); 3482 (117). Dorantes, J. & M. Acosta, 2012 (19); 2031 (8); 2083 (19). Dorantes, J. et al., 737 (19). Dorr, L. J. & L. C. Barnett, 6235 (11). Dorr, L. J. & I. Valdespino, 6408 (37). Dorr, L. J. et al., 4752, 4882 (32). Dorrier Smith, G., 371 (27). Dressler, R., 4654, 6038 (54). Dreyer, D., 317B (19). Drouet, F., 1979 (95); 2025, 2040 (13). Drummond, B. A. III, 7305 (19); 7335 (108); 7336 (98); 7346 (53). Dryander, E., 2370, 2490 (47). Dryer, V., 81, 208, 426 (113); 566 (11); 604 (117); 607 (11); 683, 684 (113); 724 (61); 729 (114); 893 (11); 1158A, 1158B (61); 1229 (114); 1452 (117). Duarte, A. P., 5976 (13); 8010 (12); 8855 (13). Duarte, L., 6692 (13); 6718, 8010 (12). Ducke, A., 423 (32). Dudley, T. R, 11361 (98); 11879 (11). Duefias, R., 60 (82). Dugand, A., 3530 (81). Dugand, A. & Jaramillo, 3877 (11). Duke, J. A., 3844 (48); 3873 (54); M3919 (5); 5152 (36); 5562 (54); 5576 (48); 8083 (54); 8791 (36); 8826 (54); 9011 (49); 9012 (11); 9088 (48); 10109 (36); 10111, 11924 (54); 12058, 12104 (19); 13083 (54); 13720, A13720 (11); 14080, 14145, 14286, 14332, 14484 (54); 14511 (48); 14603 (36); 15465 (54). Duke, J. A. & N. Bristan, 313 (48). Duke, J. A. & T. E. Elias, 13669 (54). Duke, J. A. & J. Kirkbride Jr., 14029 (54). Duke, J. A. et al., 13690 (11). Dumont, M. et al., VE-7526 (112). Dunn & LeDoux, 22015 (19). Duque-Jaramillo, J. M., 2897 (3); 3000 (8). Dusen, P., 27 Oct 1904 (12); 2180 (88); 3361 (13); 6819 (14); 6855 (8); 7554 (20); 8754 (14); 8867 (20); 11143 (27); 11191 (13); 11440, 11493 (20); 11927 (8); 13299 (30); 15445 (20); 15986 (21); 16336 (mixed collection 13 & 20); 17309 (20); 17567 (13). Duss, P. A., 366 (99). Dutra, R. L., 42 (8). Dwyer, J. D., 1554 (32); 4313 (19); 4399 (54); 6149 (33); 6969, 7638 (11); 9117, 9893, 10782, 10802a (19); 11059 (mixed collection 5 & 19); 11185, 11266, 11278, 12725 (19); 15337 (83). Dwyer, J. D. & M. Correa A., 7943, 7970a (19). Dwyer, J. D. & B. Lallathin, 8614 (19); 8759 (11). Dwyer, J. D. & B. MacBryde, 9680 (98). Dwyer, J. D. et al., 489, 555 (11); 4701 (54).

3 82 Earle, F. S., 9 May 1896 (8). Earle, F. S. & L. M. Underwood, 16 May 1896 (8). Ebinger, J., 546 (54). Echeverria, J., 379, 1014, 28097 (11). Edwall,G., 20 May 1958 (20); s.n. (13); 2885 (9); 4425 (8). Edwards, J. B., P241, P447 (19). Eggers, H. F. A. Baron von, 5039 (19); 5797 (54); 6742 (19); 13294 (91); 14219 (92); 14405 (107); 14844, 15122, 15140, 15499, 15556, 15585 (92); 15743 (109). Egler, W. A., 42-196, 129 (19); 818 (12); 820 (74); 821 (27); 823 (8). Ekman, E. L., 1924, 5085, 5717, 6756, 13722, 14061 (19). Elburg, 9827 (32). Elias, T., 1774 (54). Elleman, L., 91699 (89). Ellenberg, H., 1884 (5). Elmer, A. D. E., 11599 (24). Elmore, F. H., 017 (5). Eminons, L., 87 (75); 140 (95). Encarnaci6n, F., 26221 (19). Englesing, F. C., 105 (19). Eriksen, B. & B. Boysen Larsen, 45692 (3). Ernst, W., 2541 (19). Escobar Cardena, F., 132 (58). Escobar. L. A. de & P. Velasquez S., 7475 (37). Escobar, L. Albert de, 1845 (11). Escobar, L. Albert de & L. Escobar U., 502 (4). Escobar, L. Albert de & J. Santos, 3490 (19). Escobar, L. Albert de et al., 2685 (58); 4465 (43). Espina, J. et al., 1353 (54); 3845 (48). EspinalT., S. & J. Ramos,2584, 2597, 2769 (19); 3407 (41). Espinosa (coll. Colombia), 3061, 3062 (42). Espinosa, R., E607, 607, 2110 (89). Espinosa, R. & A. Espinosa, 673 (8). Espinoza, R., 192, 280, 1168 (97). Espinoza, S., 340 (53); 729 (73). Esquirol, J., 536 (8). Estrada, U. et al., 26 (19). Eugenio, J., 1713 (8). Eupinino, A., 146, 340 (12). Evans, R. et al., 2634 (32). Everaarts, A. P., 595, 758 (32). Evinger, E. L., 446 (72); 455 (60); 461 (56); 470 (72). Ewan, J. A., 15650 (3). Eyerdam, W. J., 25142 (10). Eyerdam, W. J. & A. A. Beetle, 8719 (5); 27705 (25). Eyerdam, W. J. et al., 23366 (8). Fabris, H. A. et al., 3104 (82?). Fagerlind, F. & G. Wibom, 947 (41). Fanshawe, G. B., F2058 (95); 2932 (32). Feddema, C., 1982 (36). Fendler, A., 603, 607 (51); 975 (64); 977 (116); 978 (57); 979 (54); 980 (19); 985 (91); 990 (8); 2096 (91); 2608 (116). Fernandez Casas, J., 7965, 8595 (98). FemrnndezCasas, J. & J. Molero, 5653 (27); 5657, 5857 (25); 6558 (84). Fernandez Casas, J. & Susanna, 8109 (98). Fernandez Casas, J. et al., 7501 (12). Fernandez, A., 1540 (45); 1974 (46); 2228 (51); 3144 (112); 3711 (54). Fernandez, A. & L. E. Mora, 1436 (3). Fernando T. et al., 2441 (98). Ferrari, G., 679 (51); 738 (112); 762 (54). Ferreyra, R., 1766, 1796 (11); 3059 (1); 3599, 3661 (98); 4794, 4800 (102); 4943 (98); 5047 (102);

FLORA NEOTROPICA 7702, 9197, 9699 (40); 11235, 11412 (98); 13201 (102); 13758 (60); 14855 (40); 16683 (15). Ferreyra, R. & C. Acleto, 15272 (41). Ferris, R. S., 5408, 6218 (5). Ferrucci, S. et al., 108 (8); 451 (27). Fest, 21, 39 (112). Feuerer, T., 9471a, 10148a, 10358a, 23027 (84). Feuillet, C., 62 (32). Fiebrig, K., 376, 558 (12); 864 (25); 4461, 4726, 4824 (31); 5122 (12); 6024 (20). Filho, 382 (20). Firmin, G., 11 (41); 195 (4); 384, 502 (3). Fishbein, M. et al., 1809 (8). Fisher, G. L., s.n. (5); 254 (19). Fleming, V., 90 (3). Fletes, E., 138 (97). Flora Falc6n, 283 (55). Flores, G., 3 (19). Flores, J. S., 10394 (83). Flores, K., 100 (97). Flores & Tello, 2214 (98). Flores Franco, G. et al., 2867 (11). Flores M., A., 1065 (83). Florschtitz, J. & P. J. M. Maas, 2481 (32). Florschiitz, J. & P. A. Florschiitz, 347 (32). Folsom, J., 2172 (11); 3010 (19); 3869b, 3869c (54); 3906 (19); 4896 (11); 5849 (19); 9371, 9877 (54). Folsom, J. & L. Collins, 1691, 1720 (54); 1739 (61). Folsom, J. et al., 2299 (54); 4995, 6782, 6797, 6978 (54). Fonnegra, R. et al., 4045, 4141, 4150, 4212 (47). Fontella & Moura, 98 (20). Forbes, C. N., 60H (8). Forero, E. & A. Gentry, 740 (48). Forero, E. & Jaramillo, 2544, 2808 (71). Forero, E. & Wrigley, 7069 (32). Forero, E. et al., 2109, 3063 (19); 3175 (67); 4372 (54); 4850, 4934, 4960, 5438, 5821. 5878 (19). Forero, L. E., 712 (48). Forero P., L. E. & H. A. Le6n, 146 (36). Forest Dept. British Guiana, 6079, F2932, F483, WB 440, WB574, WB5694, 5964 (32). Foresta, H. de, 666 (110). Forther, G. et al., 10123 (11). Fosberg, F. R., 19765 (11); 21086 (19); 28946 (95); 28975 (98); 39318 (19). Fosberg, F. R. & M. Villareal, 20573 (81); 20583 (3). Foster, R. B., 14 Sep 1982 (95); 886 (54); 887 (19); 2303 (54); 2608 (75); 2615, 2636, 4041A, 4301 (95); 5441 (98); 8140 (75); 8747 (98); 8803 (95); 9262 (75); 9374 (98); 9384 (75). Foster, R. B. & B. d'Achille, 11997, 12070 (98). Foster, R. B. & D. Janson, 8238 (98). Foster, R. B. & H. Kennedy, 1257 (54). Foster, R. B. & D. N. Smith, 9437 (34). Foster, R. B. & J. Terborgh, 5143 (95); 6085 (98); 6497 (19); 6534 (75). Foster, R. B. & Wright, 8257 (98). Foster, R. B. et al., 3140 (95); 10658 (52); 10815 (95); 11240 (38); 12223 (33). Fournet, A., 408 (10); 686 (28). Frame, P. et al., 191 (32). Franco, P. et al., 1395 (67). Frankie, G., 129 (54). Frei, B., 137 (8). Friedrichsthal, E. von, 504 (97). Fries, R. E., 315 (8). Froehner (Peru 1977), 79 (98).

LIST OF EXSICCATAE Fr6es, R. de L., 11538 (13); 20023 (126). Fromm, R., 1365 (32). Fryxell, P., 2852 (83). Fryxell, P. A. & W. R. Anderson,3479, 3506 (5); 3587 (8). Fuentes, s.n. coll. 1978 (32). Fuentes, Z., 151 (11); 207, 359 (97). Fuentes, Z. & F. Morales, 667 (97). Fuertes Loren, Pere M. D., 433 (19). Furlan et al., 36652 (12). Gage, A. T., Aug 1903 (24). Galender, C., s.n., 28 Jan 1881 s.n. (8). Galeotti, H., s.n. (83); 1155 (19); 1225M (97); 12250 (5); 7027 (97). Gallinal, 11. J. P. et al., PE-4329, PE-4329.5 (8). Gama B., H., 5147 (81). Gamboa, B., 1586 (45). Gamboa, B. & A. Picado, 345, 355, 381, 726 (45); 1039 (11). Gamundi, I. & H. Roivainen, s.n. (8). Gandara, J. M. & J. Dorantes, 58, 114 (5); 147 (mixed collection II & 19). Gandoger, Abbe Michel, Jul 1906 (19). Garcia, 48 (94). Garcia-Barriga, H., 1213 (11); 3117, 7670 (19); 10820 (8); 11209 (19); 11621 (3); 11803 (19); 11895, 11956 (58); 12104 (3); 12239 (11). Garcia-Barriga, H. & J. G. Hawkes, 12736 (43); 12795 (4); 12851 (43); 13078 (4). Garcia-Barriga, H. & R. Jaramillo M., 10418 (81). Garcia-Barriga, H. et al., 12937 (43). Garcia-Mendoza,A. & R. Torres, 1527, 1545, 1646 (19). Gardner, G., s.n. coll. 1838 (12); 237 (20); 1792 (13). Garganta, M. de, 936 (3). Garibaldi, C., 34, 43, 133 (54). Garrido, 11 (48). Gasche & Desplats, 148 (32). Gastony, G. J. et al., 175 (19). Gaudichaud,C., 12 (12); 162 (20); 516 (12); 518, 518bis (20); 619 (8); 918 (13); (herb.Imp. du Bresil 62) coll.1833 (20). Gaumer, G. F., 6, 22 (19). Gavilanes, M. & A. Castellanos, 619 (2). Gavilanes, M. & G. Quezada, 473 (19); 482 (72). Gavilanes, M. et al., 665-B, 665-D (41). Gay, C., 1167 (40). Gay, II. et al., 1563 (61). Gehriger. W., 244 (86); 408 (19);1930 (3). Gely, A. & C. Ducatillon, 126 (32). Gentle, P. C., 29, 2531, 4767, 8731, 8749, 8752 (19); 8884 (97); 8984, 9738 (19). Gentle, P. H., 5098, 6646 (19). Gentry, A., 1388, 1546, 2318, 2707 (54); 4433 (36); 5603 (54); 5678 (19); 5824 (48); 5827 (54); 6410 (48); 6927 (54); 6944 (36); 7679, 7805, 8090, 8335 (19); 9915, 10115 (92); 21685 (40); 30647 (41); 37969 (119); 48121 (47); 48143 (120); 53982 (11); 54132 (47); 73069 (92). Gentry, A. & J. Aronson, 25091 (95). Gentry, A. & P. Berry, 14954 (54). Gentry, A. & D. C. Daly, 18364 (95). Gentry,A. & C. H. Dodson, 39574 (95); 54807, 54808 (92). Gentry, A. & L. Emmons, 38738 (108); 38797 (95). Gentry, A. & M. Fallen, 17326, 17458 (48); 17461 (54); 17585 (71). Gentry. A. & N. Jaramillo, 41361 (75). Gentry, A. & C. Josse, 72362 (92). Gentry, A. & A. Juncosa, 14112, 40967 (54).

383 Gentry, A. & A. Montsalve, 48193 (109). Gentry, A. & M. Mori, 14112 (54). Gentry, A. & M. Nee, 8647 (54). Gentry, A. & R. Ortiz, 74192 (98). Gentry, A. & J. Ramos, 13359 (19); 13365 (32). Gentry, A. & E. Renteria A., 23726, 24410 (71). Gentry, A. & J. Revilla, 16594, 16595 (98); 20441 (19); 20779 (95). Gentry, A. & D. N. Smith, 44975 (102); 44989 (98); 45096 (102); 45143 (33); 45265 (98). Gentry, A. & A. Tyson, 1647 (54). Gentry, A. & R. Vasquez, 42268 (108). Gentry, A. & E. Zardini, 49341 (20). Gentry, A. et al., 16920 (54); 18388 (19); 20303 (83); 20975, 21610, 21793, 25378 (95); 25406 (19); 26718 (98); 27180 (75); 27320, 27985 (95); 27993 (19); 28541 (104); 29011, 29314 (19); 30049 (95); 31347 (19); 31487, 31740 (95); 36627 (98); 37091 (119); 37978, 39679, 39917 (98); 40035, 40189 (10); 42796 (108); 45319 (53); 45505 (33); 47709 (47); 48121 (67); 51077 (75); 51753, 51799 (25); 53005 (53'i; 54278 (108); 59502 (47); 60401 (109); 61146 (60); 64196 (53); 65383 (47); 72065, 73447 (98); 77641 (32); 79264 (118). Gentry, J. L., 3732 (97); 3736a (19); 3793, 3812 (11). Gentry, J. L. & W. L. Burger, 2636 (63); 2681 (61); 2698, 2719 (11); 2788, 2839 (97); 2858 (45): 2863A, 2864 (63); 2929 (19); 2954 (45); 2961 (63); 3015 (97); 3024 (54). Gentry, J. L. & Gentry, 23510 (11). Gerieski, 66 (20). Gibbs, P. E. et al., 3484, 6652 (20). Gigoux, E. E., Nov 1886 s.n.(8). Gilbert, L. E., 10 (97). Gillespie, L. & H. Persaud, 1544 (32). Gilli, A., 253 (92). Gillis, W. T. & T. Plowman, 10129 (11). Gilly, C. E. & E. Hernandez X., 270 (5). Gilmartin, A. J., 510 (92). Gimate L., J., 653 (19). Gines, 1583 (78); 3944 (54). Ginzburger, A., 2 Nov 1827 (13). Giraldo Canas, D. A., 248 (19). Glassman, S. F., 1666 (11); 1984 (83). Glaziou, A. F. M., s.n. (88); 322 (13); 890, 3075 (12); 8199 (20); 8853 (12); 8862 (20); 8878 (13); 9999 (98); 11305, 11360 (13); 11370 (8); 11376 (13); 11386 (20); 12107 (88). Gleason, A. H., 588, 695 (32). Godebert et al., 14 (32). Godfrey, R., 66424 (54). Goes, 0. C. et al., 1192 (20). Goldman, E. A., 922 (8). Goll, G. P., 91, 209 (19). Gollmer, J., 26 Dec 1854 (91). Gomes, 2 Oct 1919 (13). G6mez, A. et al., 23 (47). G6mez Laurito, J., 9501 (97); 9634 (61); 9858 (97). G6mez P., L. D., 2243 (63). Gonzalez, A., 990 (8). Gonzalez, J., 696 (19). Gonzalez, J. et al., 762 (54). Gonzalez, J. C. & R. Villacorta, 61 (5). Gonzalez, R., 38 (97). Gonzalez-Espinosa, M. et al., 1598 (11). Gonzalez M., E., 236 (53); 374 (8). Gonzalez Quintero,L., 403 (19); 1001 (8);1017, 1624 (19).

384 (iornes e Mates, 1058 (25). Cioudot, J., s.n. (81, 47); s.n. coll. Mar 1844 (54); 2 (81); 7 (58); 20 (47). Gouin, 20 (5). Graham, E. H., 93 (32). Granados, J., 63 (19). Grandez, C. et al., 1091 (95). Grant, M., 10246 (43). Granville, J.-J. de, 2276 (95); 2379, 3660, B4253, B4625 (32); B4560 (95); 4948 (110); 6584, 8079 (32). Granville, J.-J. de et al., 7446 (110); 8079 (32); 8120, 8629 (110). Graterol et al., 25 (94). Grayum, M., 1938, 2180 (97); 7244 (61); 10870 (97); 10959 (113); 11072 (114). Grayumn,M. & A. Badilla, 10267 (11). Grayumn,M. & A. Bien, 1849 (97). Grayumn,M. & J. Dickie, 6568 (45). Grayum, M. & B. Hammel, 5390 (117). Grayumn,M. et al., 8017 (97). Gregg, J., 51 (8); 1008 (5). Gregory, D., 1883 (19). Grewel, M. S. & R. Persaud, 87 (32). Grewel, M. S. et al., 427 (13). Grijalva, A. & N. Andi, 156 (98). Grijalva, A. & G. Grefa, 101 (19). Grijalva, A. & E. Gudino, 32, 47 (19). Grijalva, A. et al., 420 (95); 483 (67). Grosourdy, R. de, coll. 1862, s.n. (91). Grubb, P. J. et al., 413 (3); 797 (80). Gudito, E., 435 (2); 492 (73); 555 (66). Gudino, E. & G. Grefa, 1803 (98). Giudijio, E. & S. Papa, 1907 (53). Gudino, E. et al., 806 (98); 1090 (66); 1593 (108); 2099 (53). Guerrero-Nufno, J. J., 1133 (8). Guevara (Honduras 1983), 169 (5). Guevara Am6rtegui, B., 26 (8); 255 (81). Guillemin, A., 647 (20). Guillot. J., s.n. (12). Guizar N., E., 302 (8). Gunther, E., 5841, 12307 (84). GutierrezV., G., Facultad2 (19); 114, 250 (3); 2895 (19). Guti6rrez V., G. & F. Barkley, 17C110 (54). Guti6rrez V., G. et al., s.n. (8); Narifno 1 (32); Popa 1 (11); Ventanas 2 (32); Vald 5 (11); Leticia 8 (32); Mocoa 2 (98). Haber,Wm., MV-34, MV-35, MV-36 (117); MV-56, MV57, MV-58, MV-61 (113); MV-62, MV-63, 100 (54); MV-104 (61); MV-llO, MV-ill (114); 652 (11); 9187 (54); 11751 (49); 11884 (61); 11885 (114). Haber, Wm. & J. Bradford, 11574 (113); 11592 (54). Haber, Wm. & R. Cruz, 10694 (97). Haber, Wm. & W. Zuchowski, 9866 (54); 11341 (45); 11643 (11). Hacnke. T., 1153 (97); 2299 (92). Hage, J. L., 1670 (65). Hage, J. L. & H. S. Brito, 2053 (65). Hage, J. L. & E. B. dos Santos, 630 (23); 1068, 1592 (65): 1596 (23); 2065 (65). Hage, J. L. et al., 389 (22); 510 (23). Hagen, V. W. F. H. von, 11989 (47). Hagen, V. W. F. H. von & C. von Hagen, 1006 (11). Hahn, L., 1865-66 s.n., 25 May 1866 s.n., 144 (5). Hahn. Wim., 690, 824 (12); 1165 (25); 1473 (8); 1662 (27); 2053, 2206 (25); 3607, 3791 (32). Hahn, Wn'm. & Perez de Molas, L., 2737 (27); 4325 (32).

FLORA NEOTROPICA Hahn, Wm. & S. Tiwari, 5115 (32). Hahn, Wm. et al., 950 (25); 1366 (12). HaIIe, F., 0010 (32); 801 (95); 2199 (32); 2241 (95); 2253 (32); 2377, 2406 (110); 2597 (32); 2672 (95). Hamilton & Holligan, 164 (41). Hamilton, C. & H. Stockwell, 1441 (54). Hamilton, C. et al., 963 (11). Hammel, B., 1135, 1243 (54); 2601 (19); 3028 (11); 3217 (54); 3847, 4028 (19); 5692 (61); 6142 (11); 6150 (54); 6167 (97); 8136, 8760, 9021, 9063 (54); 9068, 9316 (97); 9364 (54); 10056, 10525 (97); 11725, 11831, 12034 (54); 12102, 12121 (97); 12484 (54). Hammel, B. & W. G. D'Arcy, 6393, 6408 (45). Hammel, B. & M. Gavita, 19567 (97). Hammel, B. & M. Grayum, 16599 (113). Hammel, B. & J. Trainer, 12681 (54). Hammel, B. et al., 6456 (49); 6491 (54); 6520 (11); 6559 (45); 6583, 6755 (49); 6977, 7031 (45); 18307 (97); 19630 (48). Hampshire, R. & C. Whitefoord, 695 (11). Hampshire, R. J. et al., 486 (19); 573 (83); 714 (11). Hansen, B. & M. Nee, 7604 (19). Hansen, B. et al., 1458, 1489 (5); 1689, 1690 (11); 1790, 3853 (5); 7943 (109). Harley, R. M. et al., 10914 (32); 17198 (13); 17345. 25163 (12). Harling, G. & L. Andersson, 11760 (19); 11937 (75); 12114 (2); 12178 (4); 12293 (68); 12438 (81); 13637 (41); 13683 (3); 13940 (98); 14257 (89); 14263 (3); 16090, 16101 (92); 16486 (102); 17097 (53); 17440 (98); 18048 (92); 18128 (103); 19014 (92); 19186 (41 ); 19246 (92); 19380 (109); 24177 (75). Harling, G. et al.. 7219 (95); 7745 (98); 7870 (102); 7873 (19); 9032 (2); 9308 (92): 9357 (37); 9388. 9673 (73); 10218, 10327 (3). Harmon, W. E., 2499 (19). Hannon, W E. & J. D. Dwyer, 3808 (5); 3895, 3899 (19). Harmon, W. E. & Fuentes, 3791, 5732 (19). Hart, CL120 (97). Hart, J., 862 (3); 1130 (37); 6779 (54). Hartshorn, G., 1423 (11). Hassler, E., s.n. (8, 12); 1370 (12); 1402, 1653, 2595, 3676 (8); 3682, 3793 (25); 4373 (8): 5096 (20); 5179 (12); 5711 (8); 5722 (12); 6419. 6555 (8); 6796 (12); 7604, 12381, 12420 (8). Hatschbach, G., 3951, 5150 (20); 7384 (6); 8304 (20); 9663 (8); 10152 (27); 10773 (20); 11389 (13); 12938 (25); 13163 (115); 15086 (30): 15316 (115); 15520 (20); 15742 (12); 17302 (20); 17674 (30); 18184, 20076 (20); 20327 (13); 20469 (21); 22396 (8); 23909, 24053 (16); 24434 (13); 25299 (12): 25718, 25798 (115); 29644 (30); 40394 (20): 42740 (13); 47677 (29); 47843 (21). Hatschbach, G. & J. Cordeiro, 52650 (20)). Hatschbach, G. & Haas. 15846 (12). Hatschbach, G. & J. Silva, 48690 (13). Hlatschbach, G. et al., 4374 (12); 132449(25); 13435 (115); 15846, 57992 (12). Haught, O., 1318 (36); 1462 (54); 1769 (36); 1781 (54); 1868 (48); 2000 (36); 2016 (54); 2057 (36); 2266 (48); 2329 (8); 2587 (50); 2850 (54); 3089 (92); 3099, 3384 (19); 3590 (48); 4193 (8); 4235, 4295, 4312, 4386 (51); 4620 (54); 4883 (32); 5054 (81); 5183 (43); 5401, 5462 (48); 5515 (54); 5590, 5894, 6036 (3); 6598 (8); 6604 (3). Haushal, R., s.nl. (8). Hauthal, R., 647 (8).

LIST OF EXSICCATAE Hawkes, J. G., 179 (43). Haxaire, Cl., 702, 957 (32). Hayes, S., 144, 299 (19); 399 (54); 753 (19). Heath, 70 (11). Heath, M. & A. Long, MA22 (19); 1066 (61). Heiser, C. B. Jr., 5012, 6114 (3): 3586 (49);. 3631 (19); 6014 (2). Hekking, W. H. A. & J. Schulz, 1219 (32). Henderson, H489 (24). Henkel, T. W. & R. James, 3602 (32). Henkel, T. W. & R. Williams, 1934 (32). Henkel, T. W. et al., 944 (13); 5018 (32). Henry, A., 11038A, 12254A (24). Hensold, N. & G. McPherson, 1025 (I 1). Herb. Bradeanum, 22451 (14). Herb. Chapman, coll.FL (8). Herb. Griffith, 5902 (24). Herb. Herter, 83037, 95797 (8). Herb. Hoehne, F. C., (= Brade 7019) 6894, (= Usteri 16) 15305, (=- Edwall 2885) 15309 (9); 15365, 32487 (8). Herb. Hookerianum, s.n. (91). Herb. Kerr, 5 Apr 1922 s.n., 3450, 3651 (24). Herb. Leiden, 908245-262 (24). Herb. Mus. Paris, 284 (32). Herb. Richard, s.n. (12). Herb. Rio de Janeiro, 25704 (8). HIerb.Sartwell, 1-. P., s.n1.(8). Heredia, M. D., 419 (47); 475, 530 (58). lleredia, M. D. et al., 562 (47). Ileringer, E. P., 6334 (88); Ileringer, E. P. et al., 16759 (8). Hernandez A., 211 (5). Hernandez G., 1H., 137 (97). Hernandez M., R., 515 (11); 5347 (8); 5369 (5). Hernandez M., R. & H. Puig, 255 (5). lHernandez M., R. & P. Tenorio L., 7100 (8). Hernandez M.. R. & Vasquez de Hernandez, 668 (5). Hernandez M., R. et al., 3965, 3985, 6091, 6382 (8). Hernandez Xolocotzi, E. & E. J. Sharp, X-118 (8). Herrera, 208 (8); 228 (25); 636 (8). Herrera, F. L.. 1132 (82). Herrera, G.. 71)5 (117); 734 (97); 1064 (54); 1131 (97)); 1194 (54); 1307 (114); 5903 (45). Herrera, G. & A. Chac6n, 2661 (54). Herrera, G. & W. Gamboa, 6260 (45). Herrera, (G. & S. Schik, 3793 (54). Hertentains, 9 (48). 1-lerter,W. G., 758, 758d (8). Herzog, T., 2195 (37); 2291 (82). Heyde. A. & E. Lux, 3438. 3449 (19); 4105 (11). Hicks, J. B., 371 (70). Hieronymus, G. H. E. W., sn.. 29 Sep 1878, s.n. Dec 1881 (8). Hieronymus, G. H. E. W. & P. G. Lorentz, 1016 (8). Hill, S. R. et al., 13149, 13154 (32). Hinton, Gi. B., 669 (8); 1004 (mixed collection 11 & 19); 1666 (8); 2827 (83); 3145 (97); 3701 (83); 3729 (11); 4020 (8): 6909, 13903, 15869, 15950 (5). Hinton, G. B. et al., 5753 (11); 7479, 9004, 11911 (83); 10248, 10418 (7); 15537 (83). Hioram, Bro. & Bro. Bliste, 1498 (19). Hirsch, R., P1024 (82). Hitchcock, A. S., 14315, 14499, 15165 (8); 17203 (32); 20241 (107); 20592 (92); 20786 (2); 20806 (4); 21 346 (82); 21633 (37). Hitchcock. C. L. & L. R. Stanford, 6969 (19).

385 Hjerting, J. P. & Petersen, 1249 (40). Hladik, 184 (54); 339 (19). Hodge, W., 6802, 6979 (54). Hoehne, F. C., 2698 (13); 3669 (20); 3670 (13); 23139 (8); (Inst. Biol) 28274 (20); 30091, 30093 (12); 30098 (20). Hoff, M., 5215, 5734 (32). Hoffman, B. & T. Pennington, 1495 (32). Hoffman, B. et al., 1341 (32). Hoffmannsegg, G., Baron von, s.n. (13). Holdridge, L. R., 107, 2121 (19); 6773 (97); 6870 (I 1). Holland, D. L., 22 (19). Holm-Nielsen, L. B., 18310 (3); 19265 (98). Holm-Nielsen, L. B. & H. Balslev, 23714 (2); Holm-Nielsen, L. B. & S. Jeppesen, 96 (92); 444 (19); 1164 (37); 1283 (41); 1406 (4). Holm-Nielsen, L. B. & F. Quintana,24161 (92); 24626 (3). Holm-Nielsen, L. B. et al., 2617 (107); 2621 (92); 5219 (4); 5959 (120); 5973 (68); 6014, 6035 (120); 6308. 6311 (3); 17400 (2); 19147 (98); 21028 (66); 21326 (108); 21850 (19); 21888 (98); 22619 (108); 22682 (98); 25689, 25773, 29305 (19); 27374A (41), 27857 (92); 29804 (cf. 80). Holt, E. G., 599 (8). Holton, 1. F., s.n. (54); 10 (51); 12 (19); 16 (81'; 27. 557 (51); 1566 (19). Holway, E. W. D. & M. E. Holway, 1434 (13). Hooker, J. D. & Thomson, s.n. (24). Hoover, W. S. et al., 3071, 3151 (48); 3803 (73); 4160 (48). Hopkins, M. J. G. et al., 168 (8). Hopkins, N. A., 10 (83). Hort. Huber, 597, 640 (8). Hort. Kew, 56-H-1870 (8). Hostmann, F. W., 1271 (32). Hostmann, F. W. & Kappler, 890 (32). House, P. R., 1897 (19). Howard, R. A., 12273, 15737 (19). Howard, R. A. et al., 16154 (19). Howell, J., 10278, 10294 (5). Hoyos, S., 3053, 4183 (54). Huashikat, V., 130 (35); 525 (95); 710, 1279, 2098 (98). Huber, O., 135 (54); 473 (32). Huber, 0. & M. Colchester, 8396 (95). Hudson, J., 860 (98). Huertas, G. & L. Camargo, 750 (43); (C.M.F.) 872 (81). Huft, M., 1921 (19). Huft, M. & K. Barringer, 2021 (63). Huft, M. et al., 1948 (19); 1968 (36). Hugh-Jones, 339 (48). Huidobro, A. M., 515 (8). Huidobro, R., 1756 (8). Humbles, J., 6087 (4). Humboldt, F. W. 11. A. & A. J. A. Bonpland, s.n. (4.3, 81); s.n. 1816 (3). Hunter, A. A. & P. H. Allen, 368 (19). Hunziker, A. T., 5700 (74); 6893 (8). Hunziker, J. H., 1532 (8). Hurtado, F., 586, 1149 (53); 2694 (66); 3003 (75). Hurtado.,F. & P. Mena, 132, 135 (53). Hurtado, F. & D. Neill, 1445 (66). Hurtado, F. et al., 40 (108); 41, 104, 114 (53); 2771 (19). Huston, 634 (19). Hutchison, P. C. & J. K. Wright, 6681 (3). Hutchison, P. C. & K. von Bismarck, 6469 (82). Huttel, C., 889 (41). Ibarra Contreras, G., 1305, 1410 (83). Ibarrola, T. S., 3763, 4313, 4365 (12).

3 86 Idinael, Bro., 8 (3). Idrobo, J. & A. Fernandez P., 246 (11). Idrobo, J. et al., 10370 (11); 10515, 10701 (47). Iltis, H. H. & R. Guzman, 29023a (83). Iltis, H. H. & M. G. Iltis, E-161, 187, 231 (92); 536 (41). Iltis, H. H. & M. Nee, 1402 (8). Iltis, H. H. & D. Ugent, 638, 650, 672 (82). Iltis, H. H. et al., 322 (11); 1204, 1287, 29502 (83). INBio II, 196 (117). Infantes V., J., 1479 (98). Irvine, D., 133, 473 (19). Irwin, H. S., 2229 (29). Irwin, H. S. et al., 13131 (8); 22053 (88); 30475 (8); 48162 (32). Isern, J., 1441 (41); 2537 (40). Jacobs (DUKE), 2364 (54); 2905 (97). Jahn, A., 296 (91); 532 (78); 533, 766 (3); 1184 (78). Janieson, Wm., 457, 522 (41); 625, 627, 23119 (3). Jangoux, J. & R. P. Bahia, 561 (32). Jansen-Jacobs, M. J. et al., 1203, 5169 (32). Jansson, G., 249a (20); 899a (8). Jaramillo, J., 10 (4); 5318 (41); 6491, 6966, 7412 (109); 8517 (108); 8655 (37); 8910 (2); 9381 (41). Jaramillo, J. & J. D. Boeke, 485 (37); 549, 2119 (2). Jaramillo, J. & F. Coello, 2848, 3382 (53); 3738 (108); 4282 (95); 4392 (108); 4490, 4571 (53). Jaramillo, J. & E. Proafio, 1997 (2). Jaramillo, J. & V. Zak, 590 (73); 654 (41); 8016 (73); 8035 (109); 8093 (37). Jaramnillo,J. et al., 1738 (4); 1833, 8793, 8873 (3); 8877 (37); 8910 (2); 8974 (3). Jardim, J. G. & S. C. de Sant'Ana, 790 (126). Jardim. J. G. et al., 12, 102 (126). Jativa, C. & C. Epling, 49 (107); 134 (92); 218 (109); 281 (73); 825 (92). Jelski, 56 (41). Jenikins, P., 28 Jul 1990 (8). Jenman, G. S., 2405, 6306, 6743, 7907 (32). Jennings, 0. E., 602 (19). Jimenez, D., 441, 451, 580, 4024 (97). Jimenez Almonte, J. de J., 2099, 4445 (19). Jimenez Almonte, J. de J. et al., 2925 (19). Jimenez M., A., 466 (45); 1246 (11); 2100, 2800, 2806 (54); 2978 (63); 2987 (45); 3667, 4121 (63). Jobert, 915 (32). Johansen, D. A.. 46 (mixed collection 5 & 19). Johns, T., 82-60 (8). Johnson, E. P., s.n. (5). Johnson, J. C., 467 (117). Johnston, A., 372 (99). Johnston, J. R., 40, 70 (54); 653, 1854, 1916 (83). Jones, J., 341 (5). Jones, J. & C. Davidson, 9055 (41). Jones, M. E., 343 (7). Jonsson, G., 1005a (14). Joor, Dr. J. F., 15 Sep 1868 s.n., 6 Jun 1874 s.n. (8). Jordao, 2330 (20). Jorgensen, P., 1808 (8); 2218 (12); 3341, 3676 (8); 3682 (25); 4373 (8). Jorgensen, P. M., 65446, 65972 (37). Jorgensen, P. M. & S. Laegaard, 56488 (19). Jorgensen, P. M. & Vive, 56029 (41). Jorgensen, P. M. et al., 159, 332, 348, 371 (56); 559, 616 (3); 617 (103); 704 (37); 1060 (3); 1346 (41); 1377, 1457 (56); 1477 (37); 65310 (70); 91926 (2); 92969 (11). Juan, G. E. et al., 94 (24).

FLORA NEOTROPICA Julio, Bro., II. 7, 210 (8). Juncosa, A., 1485 (71); 1647 (54);1650, 1686 (48). Junge, E., 3064 (8). Junghuhn, F. W., 404 (24). Jurgensen, C., 790 (83). Jussieu, A. de, 6373 (32). Kalin Arroyo, M. & L. Aristeguieta, 75108 (112); 75117 (64). Kalloo, M. B., B1107 (54). Kappelle, M. & M. Monge, 5159 (63). Karling, 14 (19). Karsten, H., s.n. (8). Kattan, G., 32 (11). Kaulback, R., 300 (24). Keefe, K., 24 (54). Keel, S. & Guedes, 372 (95). Kellerman, W. G., 4569 (5); 5123, 671?, 7535 (19). Kelly, 323 (19). Kelly (DUKE), 323 (11). Kennedy, H., 2874 (36). Kennedy, H. & W. Guindon, 3713 (113). Kennedy, H. et al., 3015 (19). Kenoyer, L., 515 (54); 824 (8). Kerber, A., 439 (19). Kernan, C., 81 (54); 116 (18). Kieft, E. G. B. & A. Becerra G., 130 (80). Kiesling, R. et al., 1525 (82?). Killip, E. P., 5348 (19); 5705 (120); 7846 (11); 8345 (19); 11589 (120); 39986 (54). Killip, E. P. & Garcia, 33077, 33081 (19); 33169 (70); 33608 (48). Killip, E. P. & T. E. Hazen, 9168 (4); 11004 (19). Killip, E. P. & Lehmann V., 39806 (47). Killip, E. P. & A. C. Smith, 2306 (11); 14770 (19); 15513 (11); 15762, 15850 (3); 16531 (11); 16947, 16983, 17156, 17302, 17351, 17389 (3); 17249 (80); 17863, 17887, 18711 (3); 18994 (81); 19089 (11); 19178 (81); 19249 (3); 19450 (54); 19505 (11); 19524 (80); 19779, 20068, 20070 (19); 20154 (3); 20156 (19); 20415 (105); 22365 (98); 22784 (75); 22933, 24111, 24327 (98); 24364 (33); 24466 (44); 24728 (75); 25618 (11); 26382 (75); 26400 (93); 26787 (75); 26863 (19); 26981, 27054 (95); 27610 (102); 27624, 27672 (98); 27836 (95); 27861 (102); 27920 (98); 28032 (75); 28087 (98); 28163 (108'); 28636 (35); 28717 (95); 28717, 28983, 28992, 29492, 29564, 29712, 29779 (98); 29878 (95); 30249, 30321, 30413, 30498 (13); 30680 (44). Killip, E. P. et al., 39187, 39918, 39928 (47). King, R. M., Mar 1861 (24). King, R. M., 426 (5); 795 (19); 1015 (5); 2073 (83); 2670 (5). King, R. M. & T. R. Soderstrom, 5212 (83), King, S., 512 (32). Kingdon Ward, F. F., 8950, 19679, 20203, 21213 (24). Kirkbride, J. Jr., 135 (19); 135 (61); 1734 (13); 2024, 2077 (54); 2175 (19); 2566 (51). Kirkbride, J. Jr. & N. Bristan, 1536 (54). Kirkbride, J. Jr. & J. A. Duke, 555 (97); 705, 722 (61). Klein, R. M., 116, 249, 258, 1070, 1773. 3180 (20); 4315 (8); 7084 (20); 8718 (27). Klein, R. M. & A. Bresolin, 6324, 7672 (20); 7754 (27); 9429 (8). Klein, R. M. & Sousa Sob., 8077 (20). Klug, G., 8, 834 (95); 1858 (98); 1913 (108); 2063, 2214 (95); 2629, 2883 (98); 3262 (32); 3371 (10); 3371 (19); 3491 (98); 3632 (75).

LIST OF EXSICCATAE Knapp, S., 790 (45); 802 (54); 828 (114); 829 (61); 830 (113); 834 (11); 840 (113); 841 (114); 844, 847 (97); 872 (117); 875 (61); 876 (114); 887 (19); 889 (32); 981 (48); 3065, 3116 (48); 3880a (36); 6598 (19); 6599 (95); 6605 (98); 6608 (108); 6609 (119); 6610, 6612 (108); 6613, 6616 (119); 7966 (108); 8009 (10); 8268 (19); 8279 (32); 8583 (98); 9073 (3); 9075 (11). Knapp, S. & R. Chazdon, 1408 (48). Knapp, S. & W. J. Kress, 821, 822 (48); 823 (54); 4346 (61). Knapp, S. & J. Mallet, 848, 860, 861 (97); 864 (113); 865 (11); 869 (97); 886 (54); 3976, 4005 (36); 6157, 6160 (109); 6177 (72); 6179 (11); 6180 (72); 6181 (11); 6186 (53); 6188, 6190, 6198, 6200 (19); 6203 (98); 6205 (72); 6206 (19); 6207 (98); 6216 (72); 6219, 6220, 6221 (92); 6222, 6223 (109); 6226, 6228 (92); 6229, 6236 (72); 6241 (98); 6244, 6245 (32); 6247 (3); 6252 (89); 6264, 6266 (73); 6271 (3); 6276 (72); 6277, 6280 (66); 6284 (32); 6286, 6289 (108); 6290 (32); 6292 (53); 6294 (108); 6295 (19); 6298 (66); 6301 (108); 6302, 6304 (66); 6307 (19); 6324 (34); 6341 (98); 6351 (82); 6369 (52); 6370 (75); 6372 (98); 6373, 6374 (75); 6376 (98); 6377 (93); 6378 (98); 6385 (19); 6387, 6389 (93); 6390 (52); 6391 (98); 6393, 6395 (75); 6401 (98); 6405 (75); 6413 (93); 6421, 6426 (98); 6428 (75); 6429 (10); 6430 (98); 6432 (75); 6434 (98); 6437 (75); 6438 (52); 6442 (98); 6453 (75); 6454 (98); 6463, 6466 (82); 6483, 6487 (98); 6489 (19); 6490 (93); 6492 (19); 6494 (75); 6496 (93); 6497 (98); 6498 (19); 6502, 6506 (93); 6513 (52); 6514, 6516 (98); 6517 (19); 6520 (98); 6521, 6530 (19); 6539, 6545 (98); 6562 (11); 6563 (98); 6564 (35); 6567 (32); 6569, 6570 (98); 6579 (60); 6586 (72); 6587, 6589 (102); 6594, 6599 (95); 6609, 6616 (119); 6618, 6620 (95); 6624 (34); 6625 (98); 6628, 6630, 6636, 6637 (34); 6638 (93); 6643 (95); 6646, 6647 (75); 6648, 6649 (102); 6650 (75); 6651 (52); 6652 (93); 6656, 6657 (95); 6659, 6660 (75); 6673 (64); 6674 (54); 6676, 6683, 6686 (55); 6687 (64); 6689 (51); 6692 (54); 6693 (55); 6694, 6695, 6696, 6697, 6698, 6700, 6701, 6702, 6703, 6704, 6705, 6706, 6707, 6710 (51); 6718, 6719, 6720, 6722, 6728, 6729, 6730, 6734, 6735, 6741 (13); 6743 (95); 6749, 6750, 6751, 6752, 6753, 6755 (99); 6756, 6757, 6758, 6759, 6760, 6761 (54); 6763 (100); 6765 (107); 6765, 6767 (100); 6772, 6773 (19); 6777 (106); 6778 (86); 6779 (106); 6780 (78); 6782 (46); 6784 (78); 6785 (46); 6786 (78); 6787 (46); 6788 (78); 6789 (46); 6791, 6793 (19); 6794 (86); 6795, 6796, 6797 (19); 6801 (78); 6804 (54); 6805 (19); 6809 (62); 6810 (105); 6812 (19); 6813 (62); 6814, 6815, 6817 (105); 6817 (62); 6818 (105): 6821 (81); 6822 (3); 6823 (105); 6824 (81); 6825 (105); 6828 (19); 6830, 6832, 6834 (59); 6835 (54); 6836, 6837 (32); 6838 (54); 6839 (98); 6840 (19); 6841 (98); 6842, 6843, 6844, 6845 (112); 6846 (64); 6847 (112); 6848 (64); 6849 (112); 6850 (64); 6851 (94); 6853 (54); 6854 (112); 6855 (19); 6856 (64); 6857 (112); 6858 (54); 6859 (116); 6860 (91); 6861 (116); 6862, 6864 (64); 6865 (112); 6866 (19); 6867 (91); 6868 (19); 6870 (116); 6871 (64); 7041 (98); 7199, 7200 (93); 7211, 8549a (108); 9081 (89). Knapp, S. & R. Schmalzel, 4862 (54). Knapp, S. et al., 4098 (114); 4136 (11); 4270 (61); 4273 (11); 4445 (48); 4563 (48); 6063 (113); 6209 (72); 6210 (98); 6211, 6216 (72); 6272, 6274 (11); 6309

3 87 (40); 6316 (98); 6321 (11); 6328, 6329, 6333, 6334 (10); 6335 (102); 6337 (10); 6337 (11); 6339 (95); 6343, 6350 (10); 6474, 6477 (75); 6479 (102); 6480 (19); 7477, 7478, 7480 (41); 7507 (122); 7510 (41):, 7707 (19); 7708 (52); 7882 (75); 9040 (72); 9050i (41); 9065 (89); 9085 (2); 9086 (11); 9089, 9091 (72); 9094 (2); 9095 (3); 9096 (89); 9114 (72); 9117 (39); 9119 (41); 9130, 9140 (12); 9143 (8); 9144 (12). Knight, D. H., 579, 966 (3); 1070 (89); 1074, 1157 (3). Koelz, W., 18-19 5 33, 4495 (8). Kohn, E., 1153 (98); 1274 (19). Korning, J. & Thomsen K., 47061 (108); 47131 (53); 47177 (98); 47209 (53); 58696 (98). Korthals, P. W., s.n. (24). Koscinski, M., 28 Mar 1933 s.n., 211 (20). Kowal, R. R., 2806 (83). Koyama, T. & M. T. Kao, 8923 (5). Kramer, W. U. & W. H. A. Hekking, 2156, 2283, 2392 (32). Krapovickas, A. & C. L. Cristobal, 13794 (12); 17534 (8); 35311 (25); 44495 (27). Krapovickas, A. & L. Mroginski, 20739 (27). Krapovickas, A. & A. Schinini, 31028, 31523 (8). Krapovickas, A. et al. 15038, 15754 (12); 16904 (20); 18744 (25); 24276 (12); 24308 (74); 24766 (25'i; 25824, 27446, 28294 (8); 41154, 25391 (12); 12307, 12568 (8). Kress, W. J., 77-827 (61). Kress, W. J. & T. P. Prinzie, 94-4585 (97). Krukoff, B., 5043 (19); 1179 (13, mixed collection with 32); 4511 (98); 4569 (95); 5006 (75); 5357 (98); 5371 (95); 5510 (53); 5797 (98); 8365 (108); 8377 (53); 8953 (95); 10452 (11); 10898 (125): 11060 (33); 11250 (125); 12301 (32). Kuhlmann, J. G., 65875 (20). Kuhlmann, M., 23 Jan 1935 (8); 33280 (13). Kummrow, R., 1258 (20). Kuniyoshi, Y. S., 3103, 3552 (20); 4646 (13). Kuntze, 0. (= C. E. 0.), s.n. (8); 4-5/1892 s.n. (90); May 1892 s.n. (25); Dec 1892 s.n. (13); 886 (51); 1238 (19); 2177 (91). Kvist, E. P., 40630 (92). Kvist, E. P. et al., 48695 (92); 48271 (19). Lalande, s.n. (13). Land, s.n. (17). Landrum, L. R., 2496, 2898 (20); 3898, 3927 (8); 3983 (20); 4617 (37). Langlasse, E., 40 (47); 205 (5). Langsdorff, G. H., s.n. (13). Lanjouw, J., 1184 (32). Lanjouw, J. & J. C. Lindeman, 1162, 2091 (32); 2819 (1 10). Lao, J., 329 (11). Lasser, T,. 1001 (57); 1986 (13). Lastra, de la, 11 (48). Lathrop, E., 5694 (11); 5996 (19). Laughlin, R. M., 1844, 2923 (83). Lawesson, J. E. et al., 39474 (53); 39691 (98); 43453, 44433 (95). Lawrance, A., 485 (19). Lawrence, 20 (32). Lawton, 1821 (32). Lawton, R., 1183 (117); 1244 (54). L.B.B. (Lands Bosbeheer Suriname), 8750, 11221, 12047, 13662, 14669 (32). Le Clezio, 102, 257 (48). Leavenworth, Wm. C., 249, 355, 553 (83).

388 Leeuwenberg, A. J. M., 11608 (32). Legname, V. (R.) & A. R. Cuezzo, 5954 (82?); 9718c, 9866c (25). Legrand. C.D., 375 (8). Lehmann, F. C., K217 (47); K230 (8); 1429 (19); 1774 (49); 2594 (3); 4715 (47); 4718 (3); 4770 (11); 4941 (75); 4951 (58); 6325 (47); 6986 (43); 7817 (41); 8511 (68); 8513 (69). Leite, J. E., 659 (87). Leiva G., S., 1772 (82). Leiva G., S. & J. Guevara, 1131a (3). Leiva G., S. & P. Lezama A., 935 (73). Leiva G., S. et al., 1402 (11); 1479 (41); 1507 (39); 1553 (82); 1746 (60). Lellinger, D. & J. White, 1153 (45). Lent, R., 61 (97); 832, 1158, 2295, 3160 ( 1); 3678, 3752, 3859 (97). Lent, R. et al., 3383 (97). Le6n (coll. Colombia 1976), 434, 706 (54). Le6n, B. & K. Young, 2137, 2510 (37); 2521 (3). Le6n. Bro., 4336, 5729, 10959, 11017, 12272 (19). Le6n, Bro. & E. L. Ekman, 4274 (19). Le6n, Bro. et al., 10013 (19). Le6n, J., Apr 1949 (19); 802 (11); 1454 (19); 3070 (63); 3273 (45); 4379 (11). Le6nard, E. C., 4315, 8189, 8587, 9040, 9195 (19). Le6nard, E. C. & Le6nard, 12257, 14448 (19). Leprieur, F. R., s. n. (32). Lescure, J. P., 105 (32); 2106 (108); 2127, 2167 (98). LeSueur, D. H., 422, 549 (5). Levy, P., 470 (19). Lewis, G. P., 3225 (41). Lewis, G. P. et al., 2627 (3); 2841 (37); 3488 (60). Lewis, M., 35012 (82); [88]1106, [88]1128 (84). Lewis, W. H. et al., 1522 (19); 2122 (54); 2516 (19); 2888, 5326 (54); 5570 (19). Lhotskv, J., s.n. (88, 115); 106 (20). Liane et al., 3624, 11019 (12). Liebmann, F. M., 1427 (5); 11907 (97); 15451 (19). Liesner, R,. 279 (11); 442 (54); 712 (19); 2078 (97); 2888 (19); 3020 (54); 3037 (97); 3779 (95); 5139 (11); 6123, 6428, 7120 (95); 14371 (54); 24262, 24404 (13); 25824 (95). Liesner, R. & A. Gonzalez, 5902 (13); 9222 (59); 9680 (32); 9850 (112); 9928 (94); 10615 (54); 13065 (48). Liesner, R. & M. Guariglia, 11698 (8); 11774 (11). Liesner, R. & B. Holst, 20715 (32). Liesner, R. & E. Judziewicz, 14633 (54); 14970 (1 1). Liesner, R. & J. A. Steyermark, 12342 (19); 12374, 12392 (112). Liesner, R. et al., 12580 (98); 15219 (97). Lighthipe, Rev. L. H., 25 Feb 1890 (8). Lillo, M., 9739 (123). Lima, H. C. de et al.. 2941 (12). Lindberg. 170a (13). Lindemnan,J. C., 6028, 6139 (32); 8076 (8). Lindenian, J. C. & J. Barcia, 6391 (115). Lindeman, J. C. & J. H. de Haas, 182, 272 (20); 386 (115); 768 413); 1370 (27); 3822 (20); 4374, 4889 (27); 13249 (25). Lindeman, J. C. & B. E. Irgang, 21046 (8). Lindeman, J. C. & J. P. Lescure, 6802 (32). Lindeman, J. C. et al., 28, 107 (32); 3822 (20). Linden, J. J., 234 (19); 238 (91); 778 (81); 815 (18); 834 (58); 1417 (62); 1624 (18). Linder, D. H., 47 (32). I-INN (Herb. Linnaeus), 248.4 (8); 258.4 (5).

FLORA NEOTROPICA Liogier, A. H., 9996, 11016, 11727, 12039, 12614 (19). Liogier,A. H. & Liogier, 21116, 22907, 23259. 31276 (19). Liogier, A. H. et al., 29312 (19). Little, E. L. Jr., 6180, 6225 (92); 7376 (32); 7388 (11); 7430, 7718 (32); 8648 (41); 13525, 13692, 21615 (19). Little, E. L. Jr. & G. Campunazo, 12, 253 (72); 263 (37); 290 (3). Little, E. L. Jr. & R. R. Little, 9192 (81); 9236, 9237 (3). Lizer, C., 50 (25). Llatas Quiroz, S., 707 (3); 1178 (1); 1907, 3172 (3). Llatas Quiroz, S. & J. Lao, 640 (60). Lleras, E. et al., P16968 (32); P17024 (95); P17182 (32). L. M. A., 502 (83). Lobo, M. G et al., 344 (32). Lockwood, T., 574 (98). Loefgren, A., 21 Dec 1887 (20); 1423 (Herb. Hoehnel5377) (88); 5881 (13). Loegren, A. & G. Edwall, 26 Oct 1891 (20). Lojtnant, B. et al., 12606 (2). Long, 3216, 3235 (19). Long & Burch, 3106 (5). Long, L. E., 191 (19). Longfield, C., 553 (70). L6pez, A. & A. Sagastegui A., 2786 (41). L6pez, A. et al., 7785 (3). L6pez, R. & I. Martinez, 307 (11). L6pez Figueiras, M., 830, 2630 (19); 8713 (3). Lopez Garcia, J. L. & F. Ramos M., 6, 117 (8). L6pez M., A. et al., 9085 (41). L6pez Palacios, S., 1885 (105). L6pez Palacios, S. & B. Bautista 3219 (59); 3481 (11). L6pez Palacios, S. et al., 4385 (32). L6pez Prez, J., 523 (11). Lorentz, P. G., 122 (8). Lorentz, P. G. & G. H. E. W. Hieronymus, 49 (8); 255 (85); 1016 (8); 1070 (25). Lot, E., 238 (11). Lot, E. et al., 2018 (19); 2021 (5). Lourteig, A., 1132 (27). Lowe, J., 4023 (95); 4118 (32). Lowe, Rev. R. T., s.n. (8). Lowrie, S. et al., 341, 569 (32). Lozano C., G., 4 (47); 3456 (43); 5035 (70). Lugo S., H., 159, 226 (19); 248, 402 (98); 403 (53); 416, 447 (98); 558, 611 (41); 839, 980, 1006 (19); 1122, 1139 (11); 1442, 1458, 1474 (19); 1514 (72); 1647, 1803, 1872, 1937, 2018 (19); 2094, 2134 (98); 2236 (19); 2238 (75); 2317, 2493, 2533 (98); 2674, 2697 (19); 2802 (98); 2805, 2844 (19); 3126 (98); 3128 (19); 3135 (98); 3182 (19); 3281 (53); 3308 (108); 3634 (66); 3671 (19); 3744 (53); 3751 (35); 3753 (98); 3779 (53); 3784 (95); 3829, 3880 (19); 3939 (66); 4015, 4158, 4217 (19); 4339 (98); 4418, 4435 (19); 4520, 4574 (102); 4596 (53); 4609 (108); 4630. 4680, 4951, 4965 (19); 5037 (108); 5255 (98); 6005 (19); 6080, 6100 (11); 7219, 7435 (95). Lund, P. W., s.n. (12). Lundell, C. L., 435, 443a, 469, 838, 3224 (19). Lundell, C. L. & E. Contreras, 19445 (83); 20088 (19); 20456, 20669 (11); 20920, 21171 (83). Lundell, C. L. & Lundell, 7192 (11). Luteyn, J. L., 896, 3640 (11); 4057 (19); 9215 (51). Luteyn, J. L. & Cotton, 11200 (41). Luteyn, J. L. & J. Giraldo, 12672 (47). Luteyn, J. L. & M. Lebr6n-Luteyn, 6490 (109); 11627 (83).

LIST OF EXSICCATAE Luteyn, J. L. & 0. Rangel, 13105 (19). Luteyn, J. L. et al., 4744 (3); 4897 (98); 7384 (43); 7444 (47); 8359 (108); 8657 (53); 9035 (98); 12322 (67). Lutz, B. M. J., 519 (12). Lyonnet, E., 1492 (83); 3329 (11). Lyonnet, E. & J. Elcoro, 1128 (83); 1212 (5). Maas, P. J. M., 1111 (54). Maas, P. J. M. & Martinelli, 3354 (115). Maas, P. J. M. & T. Plowman, 1827 (11). Maas, P. J. M. et al., 2301 (32); 6055 (93); 6976 (23); 7007 (13); 7518 (32); P12876 (95); P13080 (19); P13083 (98); P13296 (95). Maasola, 41 (19). Maass, G., 9 (117); 61 (19). Macbride, J. F., 3732 (33); 4027, 4155 (11); 4274 (41); 4278, 4325? 4416 (37); 4794 (10); 4815 (98); 5205 (34). Macbride, J. F. & Featherstone, 568 (40);1214, 1473, 1925, 2103 (37); 2209 (41); 2439 (8). MacBryde, B., 175 (66); 777 (75). MacBryde, B. & J. D. Dwyer, 131, 1331 (53); 1455 (108); 1456 (19); 1461 (108). MacDade, L., 692, 712 (48). MacDaniel, S. et al., 2374 (32). MacDaniels, L. H., 934 (11). MacDougal, J., 1006 (97); 1036 (54); 1446 (11). MacDougall, T., 18 Apr 1970 (97); H342 (83). Macedo, NM.& S. Assumpq5o, 683 (8). Machado, D., s.n. (20). Machado, O., 22 May 1944 s.n., 23 Apr 1945 s.n. (12). MacOwan. P., 108 (8). Macurdy, A. C., 1038 (37). Madison, MN.T., 10265-70 (11). Madison, M. T. et al., 3352, 3448, 3527 (66); 3630, 4895 (19); 5060 (109); 5083 (69); 5205 (109). Madrigal, B. & J. Chillambo, 827 (19). Madrigal, B. et al., 554, 804 (19). Madsen, J. E., 85753, 85817 (41); 85888 (39); 85897 (41); 86967 (56). Madsen, J. E. & H. Balslev, 84352 (3). Madsen, J. E. & A. B. Pedersen, 86428 (41). Magaia, M. A. & R. Gonzalez, 16 (5). Maguire, B., 33660 (13). Maguire, B. & G. B. Fanshawe, 22900 (32). Maguire, B. & L. Politi, 28620 (95). Maguire, B. & Stahel, 22802 (32). Maguire, B. & J. J. Wurdack, 33940 (26). Maguire. B. et al., 38653, 42009 (95); 56062 (32). Mahrt, M. et al.. 159 (8). Manara,B., 3 Sep 1975 (57); 29 May 1976 (64); 164 (95). Mandon, G., 178 (8); 413 (82); 416 (84); 418 (82). Mann, G., Jan 1886 s.n., 115 (24). Marcks & Marcks, 825 (19). Marin, A. L., 60 (97). Marin, F., 1775, 2355 (98). Marnn, G. & B. Jimenez, GM439 (12). Mann, G. et al., s.n. (8). Marquez & J. M. Gandara, 101 (11). Marquez R., W. et al., 442 (83). Marshall, N. & J. Rombold, 236 (110). Marshall, N. T. et al., 289 (19). Martin, J., s.n. (32). Martin, R. et al., 1617, 1739 (95). Martinelli, G. & E. Simonis, 9063 (115). Martinelli, G. et al., 6982, 6988 (32). Martinet (at P, J. B.?), s.n., 361(805) (8). Martinet, M., 277, 590 (40); 1257 (102).

389 Martinez, 124 (19). Martinez, K., 391 (114). Martinez, L., 204 (83). Martinez, M., 16 Dec 1942 (8). Martinez A., G., 148 (8). Martinez Calder6n, G., 25 (5); 26 (19); 1419 (8); 1711 (19); 1958 (5); 1970 (8). Martinez S., E. M., 7406, 7433 (19); 7886 (97); 7946, 8469, 9051, 9670, 10759, 11038, 12099 (19); 14080 (83); 15209, 17256 (19). Martinez S., E. M. & G. Aguilar, 12364 (19); 12380 (97). Martinez S., E. M. & D. Alvarez M., 27432, 27470 (19) Martinez S., E. M. & F. Barrie, 5672 (83). Martinez S., E. M. & T. Ramamoorthy, 5214 (11). Martinez S., E. M. & A. Reyes, 20430 (83). Martinez S., E. M. & J. Soto N., 1255 (11). Martinez S., E. M. & 0. Tellez, 12989, 13016 (il). Martinez S., E. M. et al., 13217 (79); 19465, 19667 (83); 28037 (19). Martinez y Day, S. D. & R. Trujillo E., 123 (5). Martins, A. & Nunes, 7525 (13). Martius, K. F. P. von, s.n. (13, 17, 32, 95, 115); s.i. Aug (95); s.n. Sep (13); 58 (17); 167, 259 (12); 261 (20); 622 (12); 1253 (13). Marunak, V., 377 (27). Mathews (= Matthews), A., s.n. (44); 446 (32); 842 (98); 1155 (44); 1195 (37); 1508 (102); 1512 (98); 1513 (37); 1968 (10); 3249, 3269 (11); 5443 (98). Mathias, M. & D. Taylor, 5059 (75); 5400 (102): 5918 (10). Matthewson, K., 125B (92). Mattos Silva, L. et al., 360, 930 (13); 1820 (126). Mattos Silva, M. A. & H. S. Brito, 841 (12). Mattos Silva, M. A. & T. S. dos Santos, 1916 (22). Matuda, E., S-143 (83); 252, (11); 432, 458 (83); 465 (11); 578 (5); 1101, 1286 (83); 1437 (5); 2466 (11); 3005 (19); 3359 (97); 3396, 3435, 3597 (19); 4289 (83); 5181 (97); 5255 (79); 15255 (83); 16584 (5); 17002 (19); 17211 (5); 17678 (83); 17683 ( 11); 17817, 17866 (83); 18087 (19); 18665 (83); 18765 (5); 37596 (8). Maurice, Bro., 706 (11). Maxon, W. R. et al., 7147 (5); 7554 (19); 7781 (Hi). Maya J., S., 426 (19). Mazana & Gonzalez, 16 (5). Mazzucconi, V. J., 282 (8). McCarroll, D., 52 (11). McDaniel, S., 5069 (54); 5097, 5153 (97); 10845a (98). McDaniel, S. & M. Rimachi Y., 16840 (19). McDaniel, S. et al., 2565 (19). McDonald, A., 1141 (19). McDowell, T., 675 (97); 4048 (32). McDowell, T. et al., 1810, 1913 (32). McLaren, H. D. et al., 197B (24). McPherson, G., 11527, 11595 (54); 13218 (58). McVaugh, R., 11801 (11); 13892, 13945, 14044 (83); 15178 (19); 15681 (5); 17425 (8); 19586 (11); 21518, 22031 (83). McVaugh, R. & W. N. Koelz, 1204 (83); 1361 (11). Medina (coll. Misiones, Argenitan), 310 (20); 744 (48). Meier, W., 6, 308 (57). Mejia, M., 270 (19). Melhus, 1. E. & G. J. Goodman, 3621 (83). Melida de Fraume & Alvarez y Gallego, 352 (58). Melinon, M., coll. 1877, s.n. (32). Mell, C. D., 502 (5). Mello Barreto, 7827 (12); 7833 (13); 8474 (12).

39( Mello Silva, R. & J. R. Pirani, 861 (12). Mena, P. et al., 2681 (41). Mendez, 198 (54). Mendez, P. et al., 174, 235 (67). Mendez G., A., 8266, 8365 (83). Mennega, A. M. W., 310 (32). Metcalf, R. D. & J. Cuatrecasas, 30091 (36). Mexia, Y., 742 (8); 1772 (83); 4217 (29); 4315 (13); 4429a (29); 4706, 5176 (20); 5258 (88); 5337a, 5368a, 5392 (29); 5462 (20); 6212a (53); 6241 (95); 6247 (53); 6261 (75); 6338 (98); 6421 (95); 6568, 6670 (92); 7073 (19); 7149 (108); 7171 (19); 7414, 7664 (3); 7689 (41). Meyer, W. C., 79 (19). Meyer, T., 2036, 2037, 2735 (8); 2982 (123); 3912 (25); 4315, 4440 (123); 4540, 12555 (25). Miers, J., s.n. (115); 337 (12); 1209 (8); 3188, 3968 (12); 4534 (88). Mille, L., 15, 970 (92). Miller & Torres C. (DUKE), 2999 (19). Miller, C'apt.,s.n. (19). Miller, G. S., 1835 (19). Miller, J. S., 2046 (54). Miller, J. S. & J. Sandino, 1186 (19). Miller, J. S. et al., 2339 (53); 2631A, 2683 (83); 2714 (11); 2718 (83). Miranda, F., 1594 (19); 6774 (5); 7654, 9160 (11). Mizoguchi, K., 953, 1040 (8); 1466 (20); 2633 (8). Mizoguchi, K. & T. Sano, 1129 (8). Mocquerys, A., coil. 1893-94 s.n. (62); 1893-94 s.n. (8); 1029 (54). Mogenson, B., 1058 (19). Molina R., A., 574, 711, 1976, 2259 (19); 3496 (5); 6392 (83); 6829 (mixed collection 11 & 19); 6987 (19); 7002 (5); 7113, 7194 (11); 7738, 13741 (83); 14314 (5); 15584 (19); 15926 (83); 20828 (19); 21306 (83); 21952 (19); 22010 (5); 22640, 23187 (19). Molina R., A. & A. R. Molina, 12078 (19); 14045, 24378, 25551 (83); 26385 (11); 26712 (83); 26738 (19); 30478, 30540 (117); 30840 (19). Molina R., A. & Montalvo, 21726 (11). Molina R., A. et al., 16128 (11); 16367 (83); 16657 (19); 16996 (83); 17659 (113); 17908 (45); 30236 (11); 30390 (83). Molinari, 83-2 (62). Moloney (May 1894), 29 (19). Monetti, L., 19 (8); 1535, 2008 (8). Monro, A. K., 1860 (117). Montes, J. E., 821, 980 (8); 1532 (27); 2103, 2491, 3367 (8); 7110 (20); 27363, 27475 (12); 27586 (8). Moore, H. E. Jr., 2925 (5); 5046 (19). Mora, G., 158, 467, 605 (11). Mora, L. E., 288 (41). Mora Olivo, A., 236 (5). Moraga, M., 264 (61). Morales, C., 33 (97). Morales, J. F., 88 (61); 93 (11); 1677 (97); 2815 (118). Morales, J. F. & J. F. Corrales, 6028 (97). Morales Mendez, A., 248 (78). Moran, J. et al., 46 (4); 57 (2). Morawetz, W., 12-311075 (20). Morawetz, W. & B. Wallnofer, 13-41085 (98); 1109888 (95). Morel, I., 2564 (12). Morel, T., 2799, 4172 (8); 5938 (25). Moreno, 818 (19). Moreno, A., 66 (97); 67 (54).

FLORA NEOTROPICA Moreno, P. (not P.P.), 818 (19). Moreno, P. P., 568 (117); 897, 942, 972, 2031, 9428, 9644, 10149, 10235, 11632, 11710, 11728, 11734, 11775, 13635, 13828, 13937, 14409, 15698, 17420 (19); 18919 (117); 19118 (114); 19457, 21876, 22450, 22983, 23427, 25027 (19). Moreno, P. P. & W. Robleto, 20553, 20962 (19). Moreno, P. P. & J. C. Sandino, 7855 (19). Moreno, P. P. et al., 24782 (19). Moreno G., S., 174 (83). Mori, S., 365, 366 (48); 8841 (32). Mori, S. & A. Bolten, 7363 (49). Mori, S. & B. Boom, 15149 (110). Mori, S. & C. Gracie, 22469 (95). Mori, S. & J. Kallunki, 2024 (48); 2511 (11); 3312 (54); 4813 (19); 5691, 5729 (49); 5911 (IF); 5929 (61). Mori, S. et al., 8702 (110); 8742 (32); 8762 (110); 9832, 10747 (12); 10766 (13); 10812 (12); 11313 (13); 11680 (12); 12067 (13); 13705 (126); 19213 (110). Moriarty, 1680 (24). Morillo, G., 5507 (95). Morillo, G. & Hagesawa, 5132 (95). Morillo, G. et al., 813 (57); 2900 (95); 3303 (116); 7223 (55). Moritz Wagner, s.n. (41); 113 (4). Moritz, J. W. K., 395, 555 (19); 1645 (116); 1701 (57). Morong, T., 617 (8); 870 (12); 1529 (8). Morton, C. V., s.n. (8); 6080 (19); 7167 (11); 7244 (83); 7244a (11); 7457 (83); 9313, 9506 (19). Mosen, T., 652 (20); 979 (8); 1927 (87); 1997 (88). Mostacero L., J. & J. Guerra L., 63 (1). Mostacero L., J. et al., 1280, 3325 (41). Mowbray, R. N., 69966 (108); 703135 (53). Mroginski, L., 73 (27). Mroginski, L. et al., 696 (12). Mueller, F., coll. 1853 s.n. (83);157 (20). Muller, C., 1929 (19); 3043 (11). Murphy, H., 498 (19). Murphy, N., 1142 (97). Murphy, N. & Jacobs, 1288 (97). Musei Bot. Stockholm, 1997 (88). Museo Nacional do Rio de Janeiro, 22835, 25660 (88). Mutis, C., 3587 (98). Mutis, J. C., 80 (47); 1980 (81); 1984 (80); 1993, 2000 (81); 2002 (80); 2003 (81); 2011 (3); 3565 (81); 3568 (80); 3578 (19); 3581 (81). Narave F., H. & F. Vazquez B., 1070 (83). Narvaez S., 2652 (5); 3184 (19). Nascimento, 0. C., 358 (32); 864 (13). Nash, G. V., 194 (19). Nash, G. V. & N. Taylor, 1157 (19). Navarro, E., 262, 562 (97). Navarro, E. & A. Picado, 598 (11). Nee, L., s.n. (92). Nee, M., 7137 (48); 7224, 9017, 9032 (54); 9982, 10000, 10625, 10755 (11); 10755 (19); 15006, 16070 (8); 17091 (32); 17319 (91); 17855 (19); 22127 (8); 22352 (19); 22959 (11); 23188, 23190 (83); 23218 (11); 23643 (5); 25476 (8); 27648 (117); 27716, 27816, 28345, 29917, 30019 (19); 33104 (83); 33352, 33366, 33477, 33529, 33616, 33697 (8); 33716, 33723 (31); 33731, 33775 (8); 33792 (98); 33828 (15); 34244 (8); 34331, 34386, 34456 (32); 36153 (98); 36154 (25); 36155 (15); 36171 (25); 36172, 36281 (8); 37057 (98); 37226 (25); 37259 (98); 37378 (25); 37520 (74); 37620, 37646 (31); 37747 (98); 37754 (8); 38426 (10); 38887 (19); 40830, 41859 (31); 42404

LIST OF EXSICCATAE (32); 42238 (31); 43191 (74); 44478 (31); 45901, 45960 (74); 46140 (31); 46416, 46432 (74). Nee, M. & Castillo C., 22453 (5). Nee, M. & G. Coimbra S., 33960 (10); 33960 (25); 33877, 33931, 33951 (8). Nee, M. & G. Diggs, 24504 (19); 24868 (5). Nee, M. & J. D. Dwyer, 9161 (19). Nee, M. & B. F. Hansen, 18415 (5); 18643 (11); 18644 (8); 18717 (11); 18737, 18763a, 18814 (19). Nee, M. & Martin, 32207 (11). Nee, M. & J. B. Miller, 27589 (19). Nee, M. & S. Mori, 3549 (54); 3549 (97); 3561 (19); 3951 (8); 3957, 3958 (51); 4057, 4136 (19); 4191 (32). Nee, M. & M. Saldias P., 36043 (98); 36248 (8). Nee, M. & G. Schatz, 19709 (83). Nee, M. & Sebastian, 28481 (19). Nee, M. & Smith, 11377 (54). Nee, M. & J. C. Solomon, 30189 (10); 30190 (84); 30214 (10); 30221 (121); 30235, 32021, 32052 (33); 34037 (82); 34049 (11); 34050, 34091 (82); 36557 (8); 36629 (82); 36632 (11); 36657 (90); 36658 (31). Nee, M. & K. Taylor, 26271 (11); 26305 (83); 26797 (8); 28900, 29388 (83); 29390, 29446 (11); 29884 (19); 36632 (11). Nee, M. & I. Vargas C., 37468 (8); 38269 (37); 38296 ( 1)-);43346, 44705 (31). Nee, M. & M. D. Whalen, 16894 (19); 17041 (105). Nee, M. et al., 14016 (11); 24694 (19); 26177 (8); 26183 (83); 26668 (5); 27727 (19); 33920 (37); 35404, 36457 (8); 36491 (82); 36493 (11); 37434 (37). Neill, D.. 1312 (19); 1881 (113); 1913, 3704 (19); 3830 (113); 4259 (19); 6021 (98). Neill, D. & C. E. Ceron M., 7245 (98). Neill, D. & P. C. Vincelli, 3540 (97). Neill, D. et al., 5930 (19); 6021 (98). Nelson (colt. 1895), 502, 2665 (5). Nelson, B. W. & Lima, P21122 (32). Nelson, B. W. et al., 693 (32). Nelson, C., 3305 (83); 3332 (11). Nelson, C. & Vargas, 2329 (l1). Nelson, C. et al., 2918, 2936 (mixed collection 11 & 19); 3628 (11); 3684 (19); 3817, 4051 (11); 5832, 6636 (19); 6921 (5). Nelson, E. W., 3605, 3605a, 3631 (83). Nevling, L. 1. & A. G6mez Pompa, 741 (8); 1113 (19); 2139 (83). Neyraut, E. J., 197 (5). Niemeyer, E., 155 (3). Nishimura, J. K. et al., 20256 (16). Noblick, L. R. & I. C. Britto, 4413 (23). Noi Mao, 27 Aug 1921 (24). Nuesser, 21 (25). Nunes & Martins, 7791, 7854 (13). Nunez, P., 5800 (95); 5993 (75); 7136 (11); 12971 (98). Nuinez, P. & N. Alanya, 13324 (98). Nuiez, P. & J. Arque, 8325 (cf. 80). Nuinez. P. & E. Bengoa, 8662 (82). Niifiez, P. & W. Galiano, 14637 (98). Nuifiez, P. & L. Quifnones, 6921 (75). Niunez, P. & L. Vargas, 7225 (82). Nuinez, P. et al., 9060, 9083 (82); 11083 (75). Obando, N., 43 (19); 60 (54). Obando, N. et al., 185 (97). Oberwinkler, B. F. & Oberwinkler, 14771 (105). Ocampo, M., 61 (97).

391 Occhioni, P., 5748 (20); 5848 (12). Ochoa, C., 531 (82); 1788, 1794 (60); 14781, 15106 (82). O'Donell, C. A., s.n. [herb. Lillo 18159] (123); 51, 85, 1395 (25); 2829, 2989 (8). O'Donell, C. A. & J. M. Rodriguez, 401, 487 (8). Oersted, A. S., 1409 (19); 1415 (49). Oldeman, R., B.845.A, 2269 (32). Olea, D., s.n. (8). Oliveira, E. de, 4729 (13). Oliveira, P. I., 210 (13); 285 (16). Oliver, R., 2377 (54). 0llgaard, B. & H. Balslev, 7519 (107); 8693, 8733 (3); 10202 (10); 10235 (72). 0llgaard, B. et al., 39034 (53); 57352 (92); 57903 (39); 98669 (67). Olsen, J. & L. Escobar, 490 (41). Opler, P., 187 (54); 316 (19); 361 (97); 748 (54); 919 (19); 1509, 1874 (97). Orcutt, C., 5215 (5). Ord6inez & Valencia, 040 (19). Orea L., L., 599 (5). Orozco, A., 1 (4). Ortega, 274, 341 (19). Ortega et al., 104 (11); 152 (mixed collection 11 & 19). Ortega, F. & D. Diaz, 2135 (81). Ortega O., R. V., 722 (19); 1357 (8); 1843 ( 11). Ortega O., R. V. & J. 1. Calzada, 779 (8); 780 (11). Ortiz (Argentina 1945), s.n. (123); 5 (25). Ortiz, F., 15 (19); 35 (8); 856, 911, 1194, 160(0, 2162 (19). Ortiz, J. J. & M. C. Herrera, 814 (5). Ortiz, L., 234 (3). Osten., C., 16290 (8). Outer, R.W. den, 852 (32). Pabst, G. F. J., 1 Jan 1964 (13); 4329 (20); 4719 (13); 6112 (14); 6801 (12). Pabst, G. F. J. & E. Pereira, 3712 (13); 6324, 6408 8). Pachano, A., 125 (2). Padilla, I., 18 (83); 150, 152 (37); 172 (83); 1121 (3). Palacios, W., 1461 (66); 1637 (19); 1957 (66); 2214, 4202 (53); 5335 (108); 5561 (35); 5581 (98); 5942 (102); 5991 (75); 6146 (19); 6338 (53); 6359 (72); 7089 (53); 10111 (75). Palacios, W. & D. Neill, 912, 1188, 1541 (98); 1547 (53). Palacios, W. & D. Rubio, 7003 (41). Palacios, W. & G. Tipaz, 10534, 10604 (2). Palacios, W. & M. Tirado, 13173 (41). Palacios, W. et al., 372 (53); 850, 868 (98); 6974 (2); 9254 (108); 9818 (109). Palma, S., 308 (83). Palmer, E., s.n. (8); 1853-56 s.n. (74); coll.1889 (8); 34, 109, 398 (5); 410, 440 (8); 1001 (5). Paredes, E. J., 151 (19). Parker, R. N., 23 Dec 1903 (8). Parodi, L. R., 14333 (8). Passos de Lima, I., 1I (21). Pavon, J., s.n. (8). Paz, N., 155 (67). Pearce, R., Apr 1864 (37). Peck, M. E., 808 (19). Pedersen, T. M., 188 (12); 862, 1046, 5105 (8); 5991 (12); 6207 (8); 10057 (25); 13442 (12). Pedersen, T. M. & J. H. Hjerting, 926 (8). Pedley, L., 2520 (8). Peirano, s.n. [herb. Lillo 18155] (123); 9860 (25). Pefiafiel, M., 196 (3).

3 92 Pefiafiel, M. & M. Tamayo, 268 (41). Pefiafiel, M. et al., 228, 277, 315 (41); 894 (3); 911 (41). Penland, C. W. & H. Summers, 625 (3); 626 (2); 787 (3). Pennell, F. W., 1584 (50); 1941, 1962 (3); 2429 (80); 3695 (19); 4597, 4623 (36); 7100, 7449 (43); 7512 (47); 10252 (19); 14152 (84); 14418 (40); 17986 (5). Pennell, F. W. & E. P. Killip, 8039 (11). Pennell, F. W. et al., 8525, 8563 (19). Penneys, D., 607 (97); 658 (117). Pennington, T. D., 5297 (109). Peredo, Y., 48, 80, 135 (8); 337 (31). Pereira, E., 3277, 4866, 5036, 5053, 5060 (13); 5782 (20); 6285 (14); 7212 (115). Pereira, E. & L. Duarte, 4856 (12). Pereira, E. & G. F. J. Pabst, 2876 (13); 6497, 6581 (8). Pereira, E. et al., 4318 (12); 4371 (mixed collection 13 & 20); 4423 (12). Pereira, M. A., 236 (115). Perez, E. & E. Garcia, 1483 (8). P6rez Arbelaez, E., 254 (80); 575, 2101 (19); 2190 (47). Perez Arbelhez, E. & J. Cuatrecasas, 5303 (81); 5318 (105); 6170 (43); 6252 (68). P6rez G6mez, M., 129 (11). Perez S., 388 (19). Perla, J., 36 (83). Perrothet, G. S., 217 (32). Perry, A. et al., 118 (95). Persaud, A. C., 242, 242bis (32). Pessoa, S. de V. A. et al., 137 (13); 160 (88); 211 (115). Petersen, E. & J. P. Hjerting, 24 Feb 1949 s.n. (25); 562 (I123); 623 (25); 926 (8). Peyton, B. & S. T. Peyton, 241 (82); 147, 1026, 1026b (15); 1093 (37); 1244 (82). Pfeifer, H. Wm., 2005 (19). Philipson, W. R. et al.. 2184 (108). Picado, A. & B. Gamboa, 265 (63). Pickel, B., 1117 (20). Pierotti, S. A., 18, 99 (8); 211 (25); 11516 (8). Pinheiro, R. S., 2158 (12). Pipoly, .J. J., 3722, 3936, 4207, 5045, 5145 (19); 5136 (113); 15658 (95). Pirani, J. R. et al., 3028 (13); 3427 (12). Pire & L. Mroginski, 305 (27). Pires, J. M., 3854 (32); 51941 (13). Pittier, H., 210 (63); 1370 (43); 2325 (19); 2396 (49); 2653 (19); 4048 (54); 4204 (48); 5173, 5465, 5686 (36): 5862 (91); 6360 (51); 6695 (32); 8373 (57); 8993 (51); 9340 (19); 9643 (8); 9906 (91); 10063 (57); 10225 (91); 11132 (51); 11502 (19); 11814 (112); 11880 (8); 12127, 12150 (94); 12167 (19); 12182 (54); 12286 (91); 12689 (8); 12915 (3); 13060 (49); 13291 (78); 13513 (94); 13821 (19); 13990 (94); 14031 (8); 14037 (54); 14176 (91); 15196 (19); 15660 (57). Pittier, H. & M. Nakichenovich, 15335 (94); 15828 (57). Pittier, H. & A. Tonduz, 9193 (54). Plowman, T., 2003 (37); 2088 (98); 2099, 2160 (19); 2179 (108); 2204 (80); 2205 (3); 2461, 2498 (95); 2523 (108); 2563 (19); 2668, 2733 (8); 3199, 3800, 3833 (3); 6035 (98); 7724 (95); 7767 (91); 13696 (32). Plowman, T. & P. Alcorn, 14354 (92). Plowman, T. & E. W. Davis, 3770 (41); 4748 (80); 4763 (98). Plowman, T. & H. Kennedy, 2270 (98). Plowman, T. & C. de Lima, 12940 (13). Plowman, T. & Ramirez R., 11186 (98).

FLORA NEOTROPICA Plowman, T. & Rury, 11164 (98); 11164a (11). Plowman, T. & J. Schunke V., 11510 (95). Plowman, T. et al., 6843 (108); 11372 (38); 12853 (12); 14280 (60). Plunkett, 51 (19). Poeppig, E., s.n. (98); 1266 (82); 2279 (102); 2483 (75). Pohl, J. E., col. Mandio (115); 3764 (13). Poilane, E., 26074 (24). Poiteau, Jul 1824 s.n. (32). Pollard, C. L. & W. R. Maxon, 264 (8). Poortmann, H., 81 (3); 173 (82); 221 (73). Porter et al., 4274 (54); 4580, 4604 (19). Potter, 5157 (54). Poveda, L., 1200 (45). Prance, G. T. & J. Ramos, 6924 (32). Prance, G. T. et al., 1152, 2131 (32); 2299 (75); 3238, 4120, 4892, 6294, 6567 (32); 7647 (98); 7814 (95); 8878, 10044 (32); 10601 (95); 12424 (19); 14630 (95); 15314 (32); 16799 (98); 18230, 20563 (32); 24206 (95); 24552 (98); 25424, 30235 (32). Prevost, M., 117 (110); 192, 503 (32); 1613, 1815 (110). Pring, G. H., 213 (80). Pringle, C. G., 2909 (8); 6837 (19); 6870, 8070 (8); 9410 (5). Proanio, E., 1997 (2). Proctor, G. R., 15 (95); 217, 383, 32123 (54l. Proctor, G. R., et al. 27276 (19). Puerto, del & Marchesi, 5781 (8). Pulle, A., 35, 128, 352, 453 (32). Purdie, W., s.ni. (48). Purpus, C. A., 6980 (11); 7105 (83); 7318 (mixed collection 11, 79 & 97); 7325, 7327 (83); 7565 (11); 8208 (5); 8415 (96); 10766, 10849 (19); 11049, 13014, 16177 (96); 16732 (19). Quarin, C. et al., 1539, 2480 (8); 2811 (12). Quelal, C. & J. Luteyn, 486 (67). Quelal, C. & G. Tipaz, 220 (92). Quelal, C. et al., 144 (67); 220 (92); 409 (53). Quesada, Fco. & M. Segura, 698 (97). Quintero, A., s.n. (47). Quipuscoa S., V. & J. Bardales, 991 (82)_ Quipuscoa S., V. et al., 129 (82). Ramamoorthy, T. P. et al., 2857, 3022 (19). Rambo, B., 29224, 39272 (20); 41684 (27); 43987, 44128 (20); 44290 (87); 44740, 44865, 48933, 49141 (20). Ramirez, J. G., 4463 (120). Ramirez, J. G. et al., 4115 (54); 4389 (62). Ramirez. L. & J. G. Ramirez, 2820 (36). Ramirez, M., 69 (97). Ramirez, V. & Fco. Morales, 143 (49). Ramirez, V. & A. Rojas, 470 (97). Ramos, H. J., 183 (83). Ramos, J. E., 803 (67). Ramos, J. E. et al., 1050 (67); 1085 (19); 1111 (111); 1II IA, 1401 (19); 1603, 1663 (67). Rangel, 0. et al., 5377 (62). Ratter, J. A., R4579, R4593 (19); R6511 (31). Raven, P. H., 21581 (97); 21917 (19). Razo Zarate, R. & F. Ramos M., 99 (8). Record, S. J. & Kuylen, 80 (19). Reed, 961 (54). Regnell, A. F., 17 Mar 1831 (8); 652 (20); 972, 111972 (13); 973 (20); 111975 (8). Reineck, E. M., Oct 1896 (20). Reineck, E. M. & J. Czermak, 90 (20); 604 (27). Reitz, P. R., C2011 (20); 2668 (8); 3826 (27); 4092 (21); 5408 (30); 5469, 5493 (14); 60)96 (20).

LIST OF EXSICCATAE Reitz, P. R. & R. M. Klein, 537 (14); 1610, 1666 (20); 1953 (13); 2067, 2234 (115); 2342 (20); 2641 (13); 3145 (20); 4034, 4221 (27); 4419 (13); 5142 (6); 5461 (8); 5522. 5543 (20); 5636 (115); 5727, 575? (20); 6506 (13); 6727, 6837 (8); 7202 (14); 7394, 9306 (20); 10204 (14); 11438 (8); 13405 (20); 13492 (14); 13594 (8); 13739 (14); 14356 (27); 16697 (20); 16779 (27); 17422, 17424 (21); 17653 (20). Renner, S. S., 69073B (35). Renteria, E., 637 (11). Renteria, E. et al., 722 (54); 4822 (36). Restinga I Segadas Vianna et al., 834 (20). Restrepo, C. & M. D. Heredia, 309 (19); 384 (47). Restrepo, C. et al., 77 (19). Revilla, J., 51 (95); 300 (19); 1647 (32); 1834. 1937 (19); 2293 (95). Revilla, J. et al., 2547, 2579 (95). Reyes Garcia, A. & M. Sousa S., 2047 (19). Reyes L. & Aquilez G., 264 (32). Ribas, 0. S. & J. Cordeiro, 252 (16). Ribas, 0. S. & J. M. Silva, 197 (21). Ribas, 0. S. et al., 775 (88). Richard, A., 532 (12). Richard, P., s.n. (32). Rico A., L. & E. M. Martinez S., 768 (5). Riedel, L., Feb 1823 s.n. (115); Feb 1823 s.n. (13); Apr 1823 s.n. (115); coll. 1832-33 s.n. (20); 25 (12); 399 (87); 400 (12); 403 (115); 406 (12); 1076 (rnixed collection 20 & 117); 1078 (88); 1079 (20); 1235 (8); 1373 (88). Rimachi Y., M., 251 (98); 975 (95); 2864 (19); 2900 (98); 3398, 3844 (19); 9854 (95). Rirnbachi,A., 7 (4); 16, 172, 427 (41). Rio, S., 315 (19). Rios, P., 56, 134, 361 (97). Risso, J. L., 525 (8). Rivera, G., 1(11 (45); 1133, 1215 (97); 1610 (61). Rivera, G. & C. Achamm, 1258 (97). Roasa R., 1378 (5). Robert, A., 733 (31). Robinson, J. W. L., 211 (54). Robles, R., 1113 (97); 1145 (54); 1931, 2159, 2839 (97). Robleto, W., 1286 (19). Rocha e Silva, 50 (13). Rock, J. F. C.. 673, 981 (24). Rodrigo, A. P., 3415 (8). Rodriquez, 142 (20). Rodriguez & Chagas, 4083 (32). Rodriguez, A., 322 (19); 514 (54); 1748 (19); 2210 (117). Rodriguez, D., 105 (8); 142 (25); 188 (74); 215 (8); 1421 (123). Rodriguez, D. & E. Martinez S., 112 (83). Rodriguez, G.. 144, 365 (97). Rodriguez, G. et al., 232 (19). Rodriguez C. & Suarez J. (MEXU), 918 (83). Roe, K. E. & E. Roe, 2140 (83); 2243 (19). Roe, K. E. et al., 989 (19). Roig y Mesa, J. T., 3234 (19). Roig y Mesa, J. T. & Cremata, 2095 (19). Rojas, F., 8409, 11591 (8). Rojas, S., & Z6fiiga, R., 188 (19). Rolda?n, F. J., 1415, 1416 (43). Roldan, F. J. et al., 418 (51). Rombouts, H. E., 730 (32). Romero-Castaneda,R., 5011 (54); 7876 (19); 4204 (98). Romoleroux, K., 714 (72); 719a (41); 814 (39). Romoleroux, K. & R. Valencia, 1303 (19).

3 93 Rooden, J. van et al., 349 (54). Rosas R., M., 223 (11); 296, 396 (19); 579 (11); 620, 1378 (5). Rose, J. N. et al., 14362 (5); 22912 (3); 23045 (89). Rosengurtt, B., B726 (8). Rossi & J. Amaral, 7277 (17). Roth, L., 825 (20). Rovirosa, J. N., 544 (97). Rowlee, W. W. & H. E. Stork, 714 (19). Ruano, J. M., 328, 573 (19). Rubio, D., 2086 (73); 5276 (2). Rubio. D. & W. Palacios, 2435 (92). Rubio, D. & C. Quelal, 544 (3). Rubio, D. et al., 897 (92); 1611 (67); 1611 (70): 1700 (67); 1902, 1973 (92); 2179 (67). Rudas, A. & F. de A. Joaquin, 1295 (95). Rudas, A. et al., 1450, 2113, 2533 (95). Rueda, R., 1142 (35). Rugel, F. 1. X., 375 (19). Ruiz, J., 1231 (95). Ruiz, J. & J. Campos, 1744 (95). Ruiz, J. & W. Mel6ndez, 1340 (95). Ruiz, M. M., 12 (19). Ruiz, R. & S. Rengifo, 1040 (11). Ruiz, T. del V. et al., 10597 (8). Ruiz L., H. & J. Pav6n, s.n. (34, 37, 83). Ruiz Landa, et al., 118 (11). Ruiz Teran, L. & J. A. Dugarte, 12488 (3); 12907 (78). Ruiz Teran, L. & S. L6pez Palacios, 7633 (46); 7566 (78). Ruiz Teran, L. & Ruiz Perez, 13470 (105). Ruiz Teran, L. et al., 8261 (78); 12545 (19). Rusby, H. H., 777 (84); 794 (10); 2606 (121). Rusby, H. H. & F. W. Pennell, 279 (19). Rutkis, E., 459 (19). Rzedowski, J., 10455 (5); 11070 (19); 16019 (11); 16419 (83). Sabajo. P. H., LBB-11221 (32). Saer, J. d'Herguert, 270 (51); 470, 807 (8). Sagastegui A., A., 7755 (1); 14206 (82). Sagastegui A., A. & Cabanillas S., 8565 (41). Sagastegui A., A. & S. Leiva G., 14847 (82). Sagastegui A., A. & J. Mostacero L., 9174 (3). Sagastegui A., A. & C. Tellez A., 12702 (82). Sagastegui A., A. et al., 8831, 9512 (41); 10101 (82); 11144 (1); 11258 (41); 11258 (89); 11924 (82); 14749 (41); 14977 (82); 15274 (103); 15572 (41); 16102 (39). Sagot, P., 454 (32). Salino, A. & A. Gentry, 61 (13). Salle, C., s.n. (83). Salvin, O., Aug 1873 S.n. (19). Salzmann, P., s.n., 323, 385 (12). Sampalo, A., 142 (20); 367 (8). Samuels, J. A., 63 (32). Sanchez & N. Zamora, 536 (11). Sanchez et al., 579 (104). Sanchez Vega, I., 3861 (3); 4048, 4095 (82); 4510 (1). Sanchez Vega, I. & M. Cabanillas, 4095 (82). Sanchez Vega, 1. & M. 0. Dillon, 8400 (66); 8406, 8549, 8574 (98). Sanchez Vega, I. & J. Torres, 6413 (41). Sanchez Vega, 1. & M. Zapata, 9149 (19). Sanchez Vega, 1. et al., 1217 (1); 1697, 3791 (41); 5960 (1); 6044 (41). Sandeman, C., 250 (40); 3447 (19); 4309 (82); 436-9 (98); 4471 (41); 4515 (33); 5744 (81).

394 Sandino, J. C., 9, 12, 2385, 2688, 2703, 2876 (19); 3321 (118); 3511, 4811, 5134 (19). Sandwith, N. Y., 629 (32); 1731 (54). Santisteban, J. & B. Guevara, 139 (11); 180 (3). Santiz Cruz, E., 24 (19). Santos et al., 5326 (12). Santos, M. dos, 11 (12). Santos, M. R., 553 (32). Santos, T. S. dos, 513 (65); 1936 (13); 3322 (12); 3405, 3797 (23). Sarukhan. J., 59 (8). Sastre, C., 2163 (32); 6292 (95). Sastr6, C. et al., 4037 (32). Sauliere, Rev. A., rec.K 1914 (8). Saunders, 641 (19). Sauvain, M., 14, 269 (32). Sawada, M., 52 (37). Scala, A. C., Jul 1926 (27). Schafer, J. A., 7944, 8578, 8998 (19). Schallert, P. O., 3765 (5). Schank & A. Molina R., 4245, 4290 (97). Scheinvar, L., 56 (8). Schiede, C. J. W., 136 (11); 137 (8). Schiefer, H., 443, 904 (3). Schimper, 890 (32). Schimpff, H., 226 (3). Schinini, A., 4006 (12); 4143 (8); 5671 (74); 7992 (12); 10738, 12614 (8); 23169 (27). Schinini, A. & 0. Ahumada, 12713 (8). Schinini, A. & E. Bordas, 20622, 24980 (12). Schinini, A. & R. Carnevali, 10580 (25). Schinini, A. & Gonzalez, 8348 (12); 9502 (74). Schinini, A. & C. Quarin, 11550 (12). Schinini, A. & R. Vanni, 26017 (12). Schnell, R. A. A., 1120 (11). Schomburgk, R., 256 (32); 680, 940, 1253 (13); 1263 (32). Short, M. J. & P. J. Stafford, 139 (117). Schott, H., 5421 (20). Schreiter, R., 856 (123); 3790 (8); 6491 (123); 9941, 9943 (28); 9944 (8); 9952 (28); 9955 (5); 9982 (123); 11213 (25). Schubert, B., 1228 (97); 1327 (54). Schubert, B. & Rogerson, 701 (11). Schuel, A., 135 (28). Schultes, R. E., 3986 (95); 6545 (32); 6681 (108); 7224 (80); 11615 (3); 18576 (81). Schultes, R. E. & Black, 8413 (108). Schultes, R. E. & I. Cabrera, 26030 (3). Schultes, R. E. & M. Villarreal, 7346, 7396 (19); 7504 (8); 7522b (81); 7679 (19); 7793, 7854 (43). Schultze, A., 105 (3); 111 (81). Schultze-Rhonhof, H., 2973 (98). Schulz, A. G., 6849, 9065, 14146 (8). Schulz, J. P., 7219 (20); 8248, 9422 (74). Schulz, J. P. & Varela, 570, 5014 (25). Schunke, C., AIOI, 258, 259, 272 (11); 321 (98); 534 ( 11). Schunke V., J., 918, 1428 (19); 1556 (102); 1609, 2015 (19); 2269 (98); 2282 (75); 2418 (95); 2501 (19); 2852 (98); 3306 (75); 3549 (93); 3738 (95); 3948 (75); 4391, 4503 (95); 4514 (98); 4718 (38); 5042 (95); 5127 (102); 5572 (38); 5866, 6172, 6176 (75); 6193 (38); 6245 (19); 6268 (53); 6653 (19); 6693, 7428, 7739, 7948 (38); 8393 (98); 9395 (34); 9667 (10); 9696 (19); 10040 (32); 10884, 11992 (38). Schwabe, W. & W. Kailing, 10 Oct 1978 (83).

FLORA NEOTROPICA Schwacke, C. A. W., s.n. (20, 88); coll. 1879 s.n. (13); Nov 1883 s.n. (12); 30 Jun 1885 s.n. (13); Oct 1888 s.n. (12); 915 (32); 13980 (20). Schwarz, G. J., 1563, 1868 (20); 4691 (12); 8182 (74); 9276 (12); 10779, 10874 (27). Schwarzkopf, 17 (105). Schwebel, E., coll. Oct s.n. (20). Schwindt, R., 30 (74); 987 (20); 1248. 1792, 1827. 2814 (27); 3286 (25); 4851 (27). Scolnik, R. et al., 19Anl68 (11); 19An519 (32). Seaton, H. E., 429 (19). Secco, R. S. et al., 184 (13). Seemann, B. C., 424 (19); 642 (54); 874 (41); 1072 (36). Segura, M., 241 (19). Sehnem, A., 1639 (20); 8263 (30). Seibert, R. J., 167 (11). Seidel, R. & S. G. Beck, 211 (8). Seifriz, W. E., 42 (19). Seler, G., 236 (40). Sellow, F., s.n. (9, 12, 13, 20, 87); coll. 1815-17, coll.1864 (20); 133 (115). Sequeira, V. & M. Torres, 60 (2). Serv. forestier Cayenne, 3523, 3702, 5140, 7032 (32). Servin, B., 506-982 (8). Sesmero, E., 42 (12); 74 (8); 201 (74). Sesse, M. de et al., 1403 (11); 1510 (19); 5346 (11). Seymour (coll. Nicaragua), 5512 (19). Sharp, A. J. & F. Miranda, 3429 (8). Shemluck, M. & Ness, 198 (108). Shepard, 555 (54). Sieber, F. W., 22 (51); 54, 309 (99). Silva, P. (Para 1969), 2286, 2470 (32). Silva, A., 5, 209 (13). Silva, M. G., 2061 (95); 4060 (13); 6291 (32). Silva, M. G. & C. Rosario, 3723 (32). Silva, M. G. & R. Souza, 2602 (32). Silva, N. T., 1075 (13); 1441 (32). Silverstone-Sopkin, F. A. (= P.A.), 1186 (120). Silverstone-Sopkin, F. A. & A. Gentry, 1875 (47). Silverstone-Sopkin,F. A. & N. Paz, 3102, 3132, 3732 (47). Silverstone-Sopkin, F. A. et al., 2521 (19); 2527 (47); 2711 (67); 2827 (47); 2868, 2875 (67); 2930 (120); 2975 (111); 2991, 2994 (67); 3392 (47); 3445 (58); 3762 (43); 3888 (120); 3946 (67); 4846 (111); 5185 (19). Sima, P. et al., 1664 (5). Simon, s.n. (20). Simonis, J. E. et al., 170 (8). Simpson, D. R. & J. Schunke V., 429 (109); 440, 460 (92). Skog, L. et al., 7045 (32); 7143, 7187 (110); 7442 (32). Skutch, A. F., 74, 383, 1170 (83); 1949 (11); 2417 (49); 2987 (11); 3280 (61); 3286 (11): 3477 (63); 4530 (19); 4943 (97); 5392 (19); 5462 (45). Sleumer, H., 2088 (123). Smith & L. Aristeguieta, 3474 (57). Smith & Smith (Virgin Islands, 1889), 495 (19). Smith (coll. 1948), 48/212 (19). Smith (coll. 1941), 10051 (19). Smith, A. (Austin), 10 (97); HIl (11); H140 (97); A152 (11); 201 (97); A236 (11); A258 (19); A479 (63); NY725, NY781 (61); NY810 (113); NY821 (61); NY904 (113); NY1288, NY1491 (61); H1638 (11); F1840 (97); NY1905 (61); 1916 (54); P2663, 2789 (113); 4100 (97); 4138 (19); 4138 (49); 4138 (63). Smith, A. C., 2495, 3417 (32). Smith, C. E. Jr. & G. Tejada, 4529 (83).

LIST OF EXSICCATAE Smith, C. E. Jr. et al., 3926 (83). Smith, C. L., 775 (83). Smith, D., 83, 546 (97). Smith, D. N., 2906 (75); 3655 (10). Smith, D. N. & J. Alban, 5574 (82). Smith, D. N. & A. Pretel, 7573 (98). Smith, D. N. & R. Vasquez, 3409 (89); 3570 (41). Smith, D. N. & R. Vasquez M., 3409 (41). Smith, D. N. & S. Vasquez S., 4578 (53); 4892 (72). Smith, D. N. et al., 1676 (98); 7824 (10); 9045 (82). Smith, H. H., 234 (99); 1154 (36); 1181 (18); 1186 (54); 1191 (51); 1857 (19). Smith, J. D., 1839 (19). Smith, J. J., 641 (24). Smith, L. B., 1245 (12); 15019 (27). Smith, L.B. & R. M. Klein, 7321 (6); 7817 (8); 13168, 13241. 13898 (27); 13944 (20). Smith, L. B. & P. R. Reitz, 10289, 12486 (8); 12669, 12683 (27); 14387 (8). Smith, S. F., 275 (75). Smith, S. F. et al., 1048 (19). Sneidern, K. von, 4555 (19); 4704 (47); 5824 (80). Snethlage, E., 108 (13). Soares et al., 9148 (20). Soares, Z., 131 (8). Sobel, G. L. & J. Strudwick, 2180 (3). Sobel, G. L. et al., 2039 (55). Sodiro, Fr., s.n. (72). Soejarto, D., 424 (3). Soejarto, D. et al., 3970 (32). Sohmer, S. H., 9549 (19). Sol Sanchez, A., 833 (97). Solano, J., 279 (97). Solano, J. et al., 107 (97). Solheim, S. & S. Reisfield, 1240 (83). Solis R., A., 163 (63). Solomon,. J. C., 5973, 5980 (84); 6018 (10); 8488 (82); 8697 (10); 8841 (33); 9052 (102); 9062 (37); 9163 (11); 9209 (33); 9675 (10); 9720 (84); 9856 (8); 9936 (25); 10056 (28); 10164, 10173 (25); 10371 (8); 10431, 10561 (28); 10838 (33); 11045 (85); 11129, 11163, 11257 (25); 12927 (33); 13096 (10); 13981 (11); 14377 (84); 14389, 14415 (82); 14933 (121); 14970, 14983 (10); 15350 (37); 16124 (84); 16400 (10); 16441 (84); 17383 (32); 17607 (10); 19177 (97). Solomon, J. C. & R. M. King, 15949 (82). Solomon, J. C. & J. Kuijt, 11516 (84). Solomon, J. C. & M. Nee, 12630 (33); 12649 (11); 12651 (75). Solomon, J. C. & A. Solomon, 4079 (8). Solomon, J. C. & B. A. Stein, 11672 (84). Solomon, J. C. et al., 11952 (10); 12099 (33). Sota, E. R. de la, 4418 (82?). Soto Nunez, J. C. & A. Roman de Soto, 2220 (11). Soto Nuniez, J. C. & S. Aureoles Conejo, 7409 (83). Soto Nufiez, J. C. et al., 5214, 5235 (11); 8503, 9345, 13466 (19). Soukup, J., 36 (82); 3465 (41); 4336 (3); 4493, 4696 (3). Sousa, M., 255, 1537 (8); 11266 (19). Soza (coll. Nicaragua) et al., 150, 181 (19). Sparre, B., s.n. (8). Spellman, D., 1580, 1707, 1766 (19). Spellman, D. & Newey, 1667, 1879, 1889, 1976 (19). Spellman, D. et al., 117 (5); 591 (19). Sperry (DUKE), 792 (97). Spire, C. J., 1424 (24). Sprague, T. A., 89 (50); 107 (32).

395 Spruce,R., Aug 1856 (75); 446 (32); 1262 (95); 1541 (98); 1642 (95); 1700 (108); 3877 (108); 4051 (93); 4161 (102); 4250 (98); 4830 (53); 4914 (32); 5117 (19). St. Hilaire, A. de, s.n. (12, 20); 38 (29); 318bis (13); 356, 396 (20); 565, 578nr.5, 748 (29). St. John, H., 20600 (48). Stafford, P. J. et al., 295, 305, 338, 357 (83). Standley, P. C., 4827 (11); 7928, 8851, 9439 (19); 10687, 10736 (117); 12234 (83); 12966 (19); 15651 (1]); 17696, 18179, 19132, 20617 (19); 20974, 21104, 21160, 21307, 22011, 22219, 23211 (5); 23741, 24188, 24859, 24923 (19); 25622 (83); 27197, 28016, 282(16, 28960 (19); 31358 (54); 35732 (97); 36199 (11); 36655 (19); 37153, 37171, 37320 (54); 38579 (49); 40860 (19); 41143 (54); 41398 (49); 41937, 41988. 42566 (63); 42852 (114); 53235, 54333, 54361, 54761, 55178, 55574 (19); 57782 (79); 61064 (83): 61307 (11); 61813 (79); 62657 (83); 64167 (5); 65322 (11); 65388, 65436, 65912, 65984, 66177, 66306 (83); 67194 (11); 67409, 67473, 67492 (79); 67578 (83); 67901, 68169 (11); 68502 (79); 68503, 68538 (83); 69170 (19); 69240 (11); 69252, 69539 (1 9); 69645 (11); 70037 (83); 71060, 70666, 70706, 7095 (19); 71350 (83); 71437 (11); 71578, 72682 (19); 78379 (11); 80125 (8); 80730 (83); 81575, 82383 (11); 83096, 83105 (83); 83326 (11); 84361, 84402 (83); 84878 (11); 84955 (83); 85008, 85018, 85019 (11); 851)47 (83); 85452 (11); 85735 (79); 85865 (11); 85871, 85928, 85935 (79); 86218, 86231 (83); 86356 (79); 86620, 86780 (11); 89995 (19); 90682 (83); 90958 (11); 91085, 91806 (19); 96018 (5). Standley, P. C. & Chac6n P., 5884 (19). Standley, P. C. & Padilla B., 3600 (19). Standley, P. C. & R. Torres R., 47454 (11); 47564 (63); 50939 (11); 51016, 51040 (113). Standley, P. C. & M. Valerio, 44495 (97); 45514 (61); 45910, 46636 (97); 46800, 46839, 48408, 48426 (19); 50253 (97); 50379 (63); 50894 (61); 51840 (I 1). Standley, P. C. & L. 0. Williams, 784 (83); 1405 (19). Stanford, L. R. et al., 1044 (19). Starry, E. D., 291 (54). Steele, E. S., 11 Nov 1900 (8). Steere, W., 2099 (19). Stein, B. & C. Todzia, 2154 (93); 2341 (75). Steinbach, J., 1708 (25); 3091 (98); 3212 (31); 3265 (31); 3381 (75); 5101 (8); 5753 (11); 7348, 7348a, 7664 (31); 8210 (15); 8380, 8458 (37); 8621 (10); 8691, 35436 (8). Steinbach, R. F., 278 (15); 541 (76). Stellfeld, C., 547, 1625 (20); 1667 (8). Stergios, B. et al., 11514 (95). Stern, W. L. et al., 417 (36); 477 (104); 498 (54); 661 (104); 696, 1886 (54). Stevens, F. L., 229 (92). Stevens, W. D., 2824, 3639, 7095, 7222, 8119, 8666 (19); 9163 (117); 9700 (19); 11681 (117); 12180, 12281, 12573, 12971, 13102, 18965 (19); 20738 (97). Stevens, W. D. & A. Grijalva, 15211 (19). Stevens, W. D. & W. Hahn, 18952 (19). Stevens, W. D. & E. Martinez S., 25689 (83); 25839, 25865 (19). Stevens, W. D. & 0. Montiel, 17480, 21455(19); 241192, 24435 (54). Stevens, W. D. & P. P. Moreno, 8666, 12281, 16740, 17598, 19211, 19288, 19367, 19492, 20981, 21278 (19); 25439 (11). Stevens, W. D. et al., 16740 (19). Steward, W. C. et al., 8, P20380 (32).

396 Stewart, R. R., 10088 (8). Steyermark,J. A., 30939 (11); 32346, 32825 (83); 33149 (11); 33966, 34668 (83); 35349 (11); 35614 (83); 35837 (79); 36006 (83); 36198 (77); 36229 (83); 36278 (11); 36790 (83); 37544 (11); 38633, 39261, 39319 (19); 43381, 44158 (11); 45128 (97); 45208 (19); 46753, 48942 (83); 49436 (117); 49866, 49945 (83); 50039 (79); 52566, 52710, 52943 (73); 52955 (41); 53986 (107); 54697 (53); 55056 (57); 55359 (46); 55407 (54); 55696 (78); 56479 (105); 56488, 56489 (98); 56871 (54); 57188 (105); 60974 (13); 61171 (19); 61249, 61438 (13); 61673 (124); 62540 (11); 75592 (32); 87069, 88007 (54); 88179 (13); 94962, 95002, 95003 (112); 99036 (19); 99457 (55); 101421 (59); 106203 (112); 111394 (13); 140982 (78). Steyermark, J. A. & G. Agostini, 91158 (112). Steyermark, J. A. & V. Carreno Espinoza, 105843 (94); 106797 (19); 107615 (51). Steyernnark,J. A. & Fernmndez,99627 (48); 99794 (19). Steyermark, J. A. & 0. Huber, 112785 (57). Steyermark, J. A. & R. Liesner, 118491 (105); 118496 (3); 118719 (54); 118839 (8); 118906 (32); 120749 (101). Steyermark, J. A. & B. Manara, 110485 (51); 110830 (54); 125203 (78). Steyermark, J. A. & 1. Narbaiza, 126491 (51). Steyermark, J. A. & S. Nilsson, 185 (95). Steyermark,J. A. & M. Rabe, 96175 (100); 96410 (112); 96800 (19); 96867 (3); 97284 (46). Steyermark. J. A. & C. Steyermark, 95281 (94). Steyermark, J. A. & J. G. Wessels Boer, 100399 (54). Steyermark, J. A. & H. Wiehler, 106591 (62). Steyermark, J. A. et al., 95822 (94); 98413 (3); 98714 (19); 100354 (54); 100734 (3); 100929 (105); 101571 (98); 101969 (59); 102193, 103330, 103569, 103579 (19); 104491 (95); 110052 (8); 110097 (19); 115524, 115612 (13); 120120, 120149, 120164 (54); 121482, 121522 (100); 121564 (112); 121616 (100); 122393, 122431 (95); 122867 (51); 123459 (48). Stimnson,W. R., 5014 (19). Stockdale, F. A., 8760, 8790a (32). Stoffers, A. L. et al., 257 (32). Stork, H. E., 1757 (113); 1852 (11); 4110 (97); 4158 (11). Stork, H. E. & 0. B. Horton, 9486 (95); 10066 (41); 10273 (44). Stork, H. E. et al., 25402 (5). Strudwick, J. J. et al., 3059 (32); 3319 (95). Stubel, A., 402 (42). Sucre, D., 1154 (12); 7771 (88); 8258 (13). Sucre, D. & P. I. S. Braga, 2470 (115); 4020 (13); 4616 (12). Sucre, D. et al., 5697 (12). Sugden, A., 1049 (54). Sullivan, G., 470 (19); 683 (54). Sullivan, G. et al., 1096 (98). Sullivan, J. R., 315, 559 (8). Swan, 68 (24). Sydow, H. von, 300 (92); 565 (41); 886 (98). Symnon,D. E., 3 Mar 1974 (24). Sytsma, K. & L. Andersson, 4644 (11). Sytsmna,K. & W. G. D'Arcy, 3240 (36). Tadeu de Aguiar, 5743 (20). Tafalla, J., s.n. (95). Tafalla, M., s.n. (95). Tamashiro, 4182 (20). Tamayo, F., 316 (51); 336 (57); 440 (19); 451 (64); 1227 (19); 1228 (112); 2063 (99); 2391 (3); 3080 (32); 4592 (51).

FLORA NEOTROPICA Tate, G. H. H., 210 (84); 676 (82); 711 (10): 791 (84); 920 (95). Tate, R., 275[181] (118). Taylor, C.. 205, 1134 (11); 1918, 2064 (5); 2178 (19): 2377 (83); 3793 (54); 3862 (63). Taylor, C. & Skotak, 4510 (19). Taylor, J. & C. Taylor, 6112 (8); 7220 (5). Taylor, K. & M. Nee, 303 (11). Taylor, N., 309, 444 (19). Taylor, R. J., 4508 (45); 18009 (19). Teixeira, L. 0. A. et al., 420 (32). Tellez, 0. & E. Cabrera, 2331 (19). Tellez, 0. & J. L. Villasefior R., 6736 (83). Tellez, 0. et al., 5259 (61). Tenore, M., (cult. Naples 1847) s.n. (20). Tenorio L., P. & R. Hernandez M., 145 (8). Tenorio L., P. & C. Romero de T., 552, 2403, 2458 (19). Tenorio L., P. et al., 14627 (19). Tenuissen, M., LBB-14669 (32). Tessmann, G., 3107 (75); 3355 (95); 3494 (98); 3497 (53); 3871 (98); 3968 (53); 4008 (52); 4531 (53). Theibaut, 1036 (5). Thieme, C., 5374 (19). Thien, L. B. & C. H. Dodson, 635 (11). Thomas, J. H. & E. Contreras, 3716 (11). Thomas, J. H. & J. L. Villasenior, 3634 (11). Thomas, R. D., 80102 (8). Thomas, W. W., 5554 (8). Thomas, W. W. et al., 5346 (32); 10098 (13); 10310, 10347, 11045 (126). Thorne, R. F. & E. Lathrop, 40528 (97); 46953 (11). Thurn, E. F., 24 (13). Tillett, S. S., 673-284 (72); 761-25 (59). Tillett, S. S. & C. L. Tillett, 45772 (32). Timana, M., 1811, 1902, 2365 (75). Timana, M. & A. Rubio, 2049 (75). Timana, M. & P. Smith, 1367 (75). Tipaz, G., 45 (4); 57 (2). Tipaz, G. & C. Aulestia, 1672 (109). Tipaz, G. et al., 313, 1393, 1726 (67). Tirado, M. et al., 509, 755 (92); 1260 (47). Todzia, C., 1155 (97). Todzia, C. & J. Grimes, 2491 (2). Tomas, Bro. A., 905 (19). Ton, A. S.. 748 (83); 811 (11); 812 (83); 1189 (19); 1255, 1306 (83); 1712, 1937 (11); 2366 (83); 2619, 2661 (19); 2865 (11); 3219, 3386 (5); 3923 (11); 4281 (5); 5924, 5928, 8126, 8197 (83). Tonduz, A., 624 (19); 1468 (49); 1795 (11); 2094 (114); 7892 (11); 9018 (19); 9096 (49); 9189 (54); 11491 (19); 11689, 11723 (49); 12763 (19): 13666 (5); 17688, 17696, 17762 (97). Torgo, F., 67 (27); 21254 (8). Toriz A., G., 423, 678 (8). Toro, R. A., 73 (19); 171 (8); 612 (32); 663, 901 (47); 1360 (8). Torres C., R., 3130 (19). Torres C.. R. & R. Cedillo T., 704, 2751 (83). Torres C., R. & H. Hernandez, 3069 (19). Torres C., R. & M. L. Torres C., 7112 (83). Torres C., R. et al., 2918 (83). Tovar, O., 1932 (44). Tracevedo & Gouzy, Valp 3 (11). Traill, J. W. H., 580 (32); 585, 586 (95). Treacy, J. & J. Alcorn, 581 (108). Trelease, Wm., 614 (8). Tressens, S. G., 863 (12). Tressens, S. G. et al., 2704, 2762 (8); 4976 (13).

LIST OF EXSICCATAE Triana, J., s.n. (2); 1851-57 (47); 1851-57 (19); s.n. 1854 (3); 16, 17 (58); 47 (81); 186 (47); 2216 (58); 2219 (80); 2243 (2); 2248 (81); 2255 (19). Trujillo, B., 1703 (94); 6376 (54); 8172 (3). Trujillo, B. & Fernandez, 97 (32). Trujillo, B. & G. Ferrari, 14042 (51). Trujillo, B. et al., 16784 (51); 16829 (8). Tsiang, Y., 6481, 9807 (8). Tucker, G., 2438, 2442 (5). Tucker, J. M., 1186 (11). luerckheim, H. von, 436 (19); in Donn. Smith 745 (83); 111270 (19); 111343 (11); 112279 (114); 3487, 7754, 8437 (19); 8492 (117); 8716 (97). Tun Ortiz, S., 945, 1250 (19); 1456 (97). Tunqui, S.. 383 (95); 866, 925 (98); 1004 (75). Tutin, T.G., 169 (32). Tyson, E. L., 812 (49); 1428 (48); 1456, 1746 (54); 6347 (49); 6684 (11); 6843 (48). Tyson, E. L. & K. E. Blum, 3980 (54). Tyson, E. L. et al., 2949 (54). Ucan, E., 4106 (19). Uck, 17813 (19). Ugent, D., 7 (19). Ugent, D. & R. Flores C, 2011 (83). Ule, E., coll.1897 (115); 5690 (19); 6109 (98); 6277 (1()8); 6482 (102); 6632 (93); 6632 (98); 9102 (12); 9720 (98); 9758 (19); 9760 (98). Uribe P., C., Dec 1930 s.n. (108). Uribe Uribe, L., 4068 (58); 4472 (54); 4697 (32); 4896, 6035 (58). Usteri, A., 12 Aug 1906 (13); 16 (9). Utley, J., 799 (19). Utley, J. & K. Utley, 634 (11); 2247 (45); 2765 (97); 3047 (11); 3712, 4574 (45); 4595, 5231 (97). Utley, K., 5718 (97); 6060, 6107 (19). Vaccari, A., s.n. (8). Valencia, N., 004 (40). Valerio R., 2891, 2948, 2964 (19). Valerio, M., 96 (97); 424 (118); 1044 (63). Valeur, E. J., 74, 327 (19). Valverde, F. M., 508 (92). Vanni, R. et al., 391 (31); 866 (8); 1445 (12); 2982 (20). Varela, G., 87 (61). Varela, L. A., s.n. (8). Vargas, H. & D. Sandoval, 279, 461 (72). Vargas, O., 156 (97). Vargas C., C., 439, 1262 (82); 6765 (98); 7661 (82); 7721, 7792 (93); 9621, 9692 (82); 11405 (98); 11750 (93); 13993, 17808, 18450 (98); 18561 (95). Vargas C., I. G., 312 (37); 823 (8); 843 (31). Vargas C., 1. G-T.& L. Vargas, 1075 (37). Vargas C., I. G. et al., 2098 (98); 3108, 3138 (28); 3173 (25). Vargas Ruiz, E. & G. J. Martin, 142 (8). Vasquez, 161 (19); 1006 (117). Vasquez B. & Hernandez, 21 (19). Vasquez S., 311 (11). Vasquez T., 222 (19); 358 (11); 456, 556 (19). Vasquez, R., 4977 (98). Vasquez, R. & G. Criollo, 1870, 5799 (95). Vasquez, R. & N. Jaramillo, 3, 1147, 1216 (95); 1949 (98); 2714 (98); 3748, 6368 (95); 6874 (19); 9541, 10491 (95). Vasquez, R. et al., 6498, 6735, 8080 (95); 16815 (98). Vattuone, I. C. & Bianchi, 166 (27). Vaughan, J. et al., 205 (5). Vaughn. Gwynne, coll. 1898 (32).

3 97 Vaughn, J. et al., 682 (63). Vaureck (Paraguay 1981), 273 (8). Vauthier, 529 (20); 532 (12). Veillon, J.P., 23 (19). Velasquez L., C., 63, 125 (11); 227 (5). Ventur, 14 (19). Ventura A., A., 5510 (19). Ventura A., F., 298 (83); 3268 (11); 3536 (83); 3608 (19); 4772 (11); 5510, 8219 (19); 8345 (8); 9503 (83); 9834 (11); 9930, 10051 (19); 10054 (8); 10820 (11); 11230 (8); 11400, 11551 (19); 12746 (83); 13103 (11); 14674 (19); 15102, 15832 (11). Ventura, E. & E. L6pez, 2294 (11); 8941 (8). Ventura, E. et al., 2933 (11). Venturi, S., 105 (8); 157 (25); 209, 410 (28): 1181 (25); 1840, 2447 (28); 2447c (25); 2464 (28); 2762, 3923 (123); 5288, 5947, 7894 (8); 7951, 82293, 8651, 8795 (25); 8818 (28); 8896 (123); 9323, 9328 (28); 9414, 9532 (123); 9563, 9850 (25); 9959 (8); 9967 (123); 10427 (8). Vera Santos, J., 3024 (19). Vervoorst, F. B. & A. R. Cuezzo, 7635c (25). Vickers, W., 232 (108). Vidal Senege, M., s.n. (41). Vieira, G. et al., 338 (32). Viereck, H. L., 732 (19); 1101 (5). Vilcapoma S., G., 162, 164 (40). Villa, E., s.n. (8); 37 (25); 49 (8). Villacorta, R. & M. Villacorta, 2131 (5). Villagran, E., 199 (83). Villaneuva, R., 778 (19). Villegas F., E., 371 (8). Villegas F., E. et al., 231 (8). Vincent, K. A., 4063 (8). Vivar C., F. et al., 644 (92); 811 (89); 1089 (72). Vogel, S., 51 (58). Vorides, 15 (11). Vreden, C., LBB-13662 (32). Wagner, R. J., 1229, 1607 (19). Wallich cat. no., 2619a (24). Wallnofer, B., 13-151088 (98); 14-1388 (93). Wallnofer, B. & R. Fernandez, 14-21287 (93). Wallnofer, B. & M. Henzel, 11-19488 (52). Warburg, O., 15074, 15075 (24). Warming, E., s.n. (88); 5 Apr 1864 s.n. (17); 8 (29). Warner, R. N. & White, 130 (48). Wasshausen, D. & F. Encarnaci6n, 697 (33). Wasum, R. et al., 494 (8). Watson, H. C., 144 (8). Wawra, H., 227 (19). Weaver, R. Jr. et al., 1716 (83); 1747 (11). Weber (coll. Mexico), coll.1866 (11). Weberbauer,A., 1876 (10); 3690 (38); 4092 (103); 5543 (82); 6678 (103); 6709 (33); 6719 (98); 7551 (37). Webster, G. L., 21965, 23277 (119). Webster, G. L. & T. Lockwood, 22691 (3). Webster, G. L. et al., 9575 (19); 11788 (83); 12197 (97); 12583 (19); 16471 (97); 17900 (19); 23277 (119); 27554, 28248 (72); 28309 (19); 31931 (37). Weddell, M., s.n. 1851 (33). Wedel, H. von, 731, 1085 (97); 2268, 2367, 2603 (54). Weir, J. R., 1864-65 s.n. (20). Weitzman, A., 196 (98). Weitzman, A. & W. Hahn, 291 (32). Weitzmann, A. L. et al., 48 (86). Wells, M. J., 3971 (8). Werff, H. van der & A. Gonzalez, 4834 (54).

3 98 Werff, H. van der et al., 9903 (108); 12213(73); 13121(75). Wessels Boer, J. G., 2067 (mixed collection 54 & 95). West, J., 3671 (37); 8532 (25). Wettstein, R. von & Schiffner, s.n. (13). Whalen, M. D., 813 (3); 816 (37); 842 (98). Whalen, M. D. & M. 0. Dillon, 890 (82). White, S. & Warner (Colombia 1973), 98 (19). White, C. T., 1266 (8). White, G., 51 (49). Whitefoord, C., 1126, 1196, 2050, 2424 (19); 3769 (99); 8073, 9109 (19). Widgren, J. F., coll. 1845 (13). Wilbur, R., 24394 (97); 24506 (63); 24507, 26036 (19); 26247 (11); 28735 (45); 28859, 29710, 31392, 31659, 31966, 32492, 32506 (11); 33450, 37849 (54); 39982 (97); 40221 (54). Wilbur, R. & F. Almeda, 16869 (11). Wilbur, R. & J. L. Luteyn, 11680 (19). Wilbur, R. & D. E. Stone, 8543, 8581, 10089 (11). Wilbur, R. & J. Teeri, 13094 (61). Wilbur, R. & Wilbur, 1425, 1817 (19). Wilbur, R. et al., 13041, 15269, 15422 (11); 15676 (19); 23521 (45). Wiley, J. R., 65, 67 (19). Wilkes, Capt., 1838-42 (13). Williams & Assis (Minas Gerais), 6754 (29). Williams, L. O., 791 (54); 11541 (32); 14221 (83); 16114 (45); 17415 (83). Williams, L. 0. & A. Molina R., 10029, 12096, 12765 (11); 12797, 13320, 13715, 13972 (83); 14607, 15213 (19). Williams, L. 0. & R. P. Williams, 18469 (19). Williams, L. 0. & Wilson, 40759, 40765 (11). Williams, L. 0. et al., 4011 (19); 22501 (11); 25422 (83); 25764, 27139 (79); 27516 (19); 28048, 28937 (11); 29020 (97); 29033 (11); 40264 (19); 40415 (11); 41731 (83). Williams, LI., 57, 1164, 1993, 2095, 2224, 2245 (95); 2992, 3087 (32); 3285 (95); 3518 (98); 3851, 4377 (108); 4596, 4764, 4932, 4983, 5246 (98); 5450 (19); 5505 (98); 5527, 6179, 6260 (102); 6352 (19); 6734 (93); 6860 (95); 6921 (102); 7197 (53); 7217 (95); 7352 (75); 7616 (53); 7655, 7912, 8066, 8184 (95); 8242, 10271, 12466 (19); 13959, 15847 (95). Williams, R. O., 11889 (51). Williams, R. S., 221 (15); 652 (31). Wilson, P., 479, 1091 (19). Wilson-Browne, G., WB440, WB574 (32). Windler, D. R. & Smith, 1013 (5). Wingfield, R., 5085 (51). Winzerling, H. W., V-5 tree 21 (19). Woodbury, R. O., 18 Jun 1958 s.n., Apr 1960 s.n., Dec 1960 s.n. (19). Woodson, R. E. Jr. & R. Schery, 487 (11). Woodson, R. E. Jr. et al., 1276 (19); 1828 (54). Woodworth, R. H. & P. A. Vestal, 555 (19). Woolston, A. L., 231, 280 (8); 326, 326B (12); 642, 642B (20); 1035 (8); 1500, 1501, 1502 (12). Wooton, E. O., 25 Jun 1925 s.n. (5). Woytkowski, F., 5774 (108); 5845 (53); 6291 (95); 6947 (37); 7188 (98); 7528 (34); 7825 (41); 34064 (82); 34168 (98); 35381 (11). Wright, C. et al., 513 (19). Wullshlagel, H. R., 1528 (13). Wunderlin,R. et al., 444, 1127 (19); 8694 (37); 8726 (3). Wurdack, J. J., 684 (3); 997 (11); 1018 (72); 1116, 1423 (41); 2016, 2126 (98); 2202 (75).

FLORA NEOTROPICA Wurdack, J. J. & J. V. Monachino, 39616 (54). Xena, N., 593 (51). Ybarrola, F., 2754 (8). Yepes Agredo, S., 434 (13). Young, H. J. & D. A. Stratton, 208 (66). Young, K., 1240 (41); 1280 (98); 1289 (37); 1406 (82); 1513 (41); 1514 (37); 1534, 1568, 1692 (41); 2157, 2425, 2650, 2664 (39); 2866 (41); 3195 (39); 3722 (82); 3833 (39); 3974, 4177 (72); 4213 (56). Young, K. & M. Eisenberg, 361 (72). Young, K. & G. Sullivan, 572 (11). Young, K. & F. Watson, 3196 (37); 3401 (41). Yuncker, T. G., 1799 (8); 4518, 5761 (19). Yuncker, T. G. et al., 5761 (19); 6050 (11): 8106, 8301, 8841 (19). Zak, V., 967 (3); 1069, 1070 (37); 1113, 1278 (109); 1281 (19); 1281 (37); 1283, 1285 (109); 1370 (4); 1373 (80); 1384 (2); 1405, 1408 (3); 1409, 1410 (41); 1454 (2); 1465, 1468 (37); 1477 (3); 1483 (109); 1510 (11); 1570 (41); 1586 (2); 1594 (41); 1606 (2); 1611 (41); 1616 (3); 1627, 1629 (2); 1636 (11); 1645 (41); 1655 (80); 1657 (2); 1658 (3); 1659, 1742 (2); 1745 (4); 1747, 1756 (72); 1770, 1772, 1780 (3); 1786 (4 1); 1820, 1823 (3); 1847 (2); 2020 (4); 2040, 2047, 2052, 2083, 2084, 2086, 2090, 2100, 2250 (3); 2321 (80); 2352, 2353a, 2357 (41); 2359, 2363, 2366, 2367, 2377, 2379, 2383, 2388 (3); 2392 (41); 2396 (3); 2407 (37); 2410, 2413 (3); 2419, 2432, 2435, 2436 (41); 2437 (3); 2438 (41); 2440 (3); 2443 (2); 2448 (41); 2451, 2461 (3); 2471 (37); 2478 (3); 2479 (37); 2484, 2487 (3); 2496 (37); 2503, 2504 (3); 2511 (2); 2514 (41); 2515, 2516 (37); 2581 (3); 2692 (72); 2720 (41); 2729, 2741 (3); 2805 (11); 2835, 2867 (41); 2896 (3); 2994 (11); 3052 (41); 3052 (37); 3259, 3358 (41); 3436, 3508 (4); 3654 (3); 3684 (2). Zak, V. & J. Boeke, 374, 485 (37). Zak, V. & J. G6mez, 1436 (2). Zak, V. & J. Jaramillo, 590 (73); 619 (3); 654 (41); 2020 (4); 2109 (73); 2154 (19); 2196 (73); 2219 (19); 2250 (3); 2255, 2265 (41); 2285 (73); 2321 (37); 2333 (19); 2560, 2576 (41); 2581, 2623 (3); 2652, 2656 (41); 2692 (73); 2720 (41); 2729 (3); 2741 (4); 2805 (11); 2830 (73); 2835, 2867, 2890 (41); 2896 (3); 2994 (11); 3046 (73); 3052 (37); 3136 (19); 3259, 3323, 3358 (41); 3436 (4); 3508, 3684 (2). Zak, V. et al., 5305 (92); 5406, 5544 (109); 5548 (92); 5619 (109); 5640 (92). Zamora, N. et al., 2477 (97). Zanoni, T., 20537 (19). Zanoni, T. & R. Garcia, 36948 (19). Zanoni, T. & M. Mejia, 16566 (19). Zanoni, T. & J. T. Mickel, 25569 (19). Zanoni, T. & J. Pimentel, 26544, 26637 (19). Zanoni, T. et al., 20333, 24235, 24307, 25662, 26766, 32786, 37715 (19). Zarucchi, J. L. & J. Betancur, 6388 (4); 6428 (54). Zarucchi, J. L. et al., 5528, 5538 (48); 6724 (32); 7065 (104). Zaruma, J., 725 (19). Zaruma, J. et al., 61 (19); 70 (98). Zehyer, s.n. (87). Zerny, H., 14 Sep 1927 (13); 5 Oct 1927 (12). Zola B., M. G., 325 (8); 454 (19); 512 (11); 565, 712, 753 (19). Zomer, H., 329 (19). Zuiniga, R., 48, 194 (118). Zuniga Donaire, R., 19 (19).

399

INDEX OF SCIENTIFIC NAMES

INDEX OF SCIENTIFIC NAMES Synonyms are in italics. Names in boldface are those accepted as membersof Solanumsect. Geminatas.l in this monograph.Primarypage referencesare in boldface. An asterisk(*) indicatesa page having an illustration or map. Aculeata 4 Amegilla 25 Anthophora 25 Apis mellifera 25 Araucaria 24. 119, 137 Arawacus 36 Aspidomorpha 31 Asteraceae 32 Athenaea 364 Augochlora 25 Augochlorella 25 Augochloropsis 25, 28 Aulacorynchus prasinus 29 Aureliana 364

Leptostemonum 4, 5, 22, 29 Lycaenidae 36 Lycianthes bigeminata 365 Lycianthes sanctae-clarae 10 Lycopersicon 5 Lycopersicon esculentum 12

Bassovia foliosa 76 Bassovia richardii var. martii 84 Bombus 25, 26, 27, 28, 338, 340, 347 Boraginaceae 32

Neocorynura 25, 28 Nomia 25 Notoxaea 25

Maecolapsis 3 1* Maurella 5 Mcclungia 32, 33*, 34 Melipona 25, 27, 28, 83, 302 Melissodes 25 Mesoxaea 25 Myrsinaceae 364

Oleria 35 Oxaea 25

Caenaugochlora 25, 28 Cassidinae 32 Caupolicana 25 Centris 25, 26 Ceratinia 32, 33*, 34, 35, 36, 37* Ceratiscada 32, 33*, 34, 36 Cestrum 32 Colletes 25 Columba fasciata 29 Coptocycla 31 Cuatresia 365 Cyphomandra5, 10, 12, 22, 28, 39, 140 Cvphomandra bassovioides 99, 101 Cvphomnandraflagrans 114 Cvphomandra homalophylla 298 Cv)phomandraverruculosa 138

Pachystemonum 4, 5 Pheliandra herthae 38, 302, 307 Pheliandra 38 Pionandra flagrans 114 Prittwitzia 32, 34, 35, 36 Protandrena 25 Protoxaea 25 Psaenythia 25 Pseudaugochloropsis 25 Pseudocapsica undatifolium 38 Pseudocapsica 38 Pteronymia 32, 33*, 34, 35, 36, 310, 331, 340 Ptiloglossa 25

Diabrotica 31, 32 Dodecatheon 26

Rekoa 36 Rubiaceae 27

Epicharis 25 Episcada 32. 33*, 34, 35, 36 Euglossa 25 Eulaema 25 Eumolpinae 31*, 32 Evylaeus 25 Exomalopsis 25, 28

Scada 33*, 34, 35 Senecio 32

Galerucinae 31 Godyris 32, 33*, 35, 36 Halictus 25 Hoffmannia 27 Hylaeus 25 Hypoleria 36 Hypomenitis 35 Hypothyris 35 Inermia 4 Labidomera 31 Lasioglossum 25, 28 Lepidota 5 Leptinotarsa decemlineata 31, 32

Solanum chorus subg. Solanum 4 Solanum chorus subg. Stellatipilum 4, 5 Solanum grad. ambig. Anthopleuris 5, 38 Solanum grad. ambig. Anthoresis 5 Solanum grad. ambig. Bassovioides 5 Solanum grad. ambig. Indubitaria 5 Solanumgrad.ambig. Leiodendra5,6,38 Solanum grad. ambig. Lepidota 5 Solanum grad. ambig. Oppositifolia(um) 5, 38 Solanumgrad.ambig.Pseudocapsica(um) 5, 38 Solanum sect. Acanthophora 5 Solanum sect. Allophylla(um) 5, 8, 10, 39, 404 Solanum sect. Anarrhicomenum 364 Solanum sect. Androceras 5 Solanum sect. Anisantherum 364

Solanum sect. Anthopleuris 38 Solanum sect. Anthoresis 138, 285 Solanum sect. Basarthrum 5, 365 Solanumsect. Brevantherum 5, 11,365 Solanum sect. Cemuum5, 8,40, 403 Solanum sect. Cyphomandropsis 8, 22, 39, 146, 358, 365, 404 Solanum sect. Dulcamara 366 Solanum sect. Extensum 8, 40, 367 Solanum sect. Holophylla 5, 8, 10, 11, 40, 262, 366 Solanum sect. Indubitaria 8 Solanum sect. Lasiocarpa 5, 124 Solanum sect. Leiodendra(on) 5, 7, 38, 365 Solanum sect. Lepidota(um) 5, 7, 8, 40, 403 Solanum sect. Leptostemonum 5 Solanum sect. Maurella 5 Solanum sect. Oppositifolium 7, 17, 38 Solanum sect. Pachyphylla 5, 8, 12, 22, 28, 39, 140, 403 Solanum sect. Pachystemonium 5 Solanum sect. Petota 5, 22 Solanum sect. Pseudocapsicum 7, 8, 38 Solanum sect. Pteroidea 5, 10, 29, 246, 367 Solanum sect. Solanum 5, 22, 364, 365 Solanum sect. Torva 52 Solanum sect. Tuberarium 5 Solanum subg. Archaeosolanumr6 Solanum subg. Bassovia 6 Solanum subg. Brevantherum 6, 7 Solanum subg. Leptostemonum 4, 6, 364, 365, 366, 368 Solanum subg. Lyciosolanum 6 Solanum subg. Minon 4, 6, 38 Solanum subg. Petota 364, 365 Solanum subg. Potatoe 5, 6 Solanum subg. Solanum 6, 7 Solanum subsect. Indubitaria 7, 17, 20 Solanum subsect. Micranthes 5, 38 Solanum subsect. Oppositifolium38 Solanum subsect. Silicosolanum 7, 38, 94, 132 Solanum subsect. Triplinervium 38 Solanum abitaguense 11, 14, 16*,23, 236*, 237, 238*, 239*, 241, 257 Solanum acropterum species group 39, 40, 368 Solanum acropterum 368 Solanum acuminatum 6, 8, 16*, 25, 71*, 73, 74*, 75*, 363 Solanum acuminatum var. v'iridiflorum 84, 86 Solanum aethiopicum 6

400 Solanum alatirameum 136*, 140* Solanum aligerum 262, 366 Solanum amblophyllum species group 11, 15, 40, 41, 165, 166*, 169, 252, 280 Solanum amblophyllum 23, 165, 166*, 167*, 168* Solanum amnicola 10, 41, 194, 195*, 197, 198*, 202*, 289, 344 Solanumnanacamptorhac his 84, 86 Solanum anguivi 364 Solanum anisophyllum 196, 198, 199*, 208* Solanurn anonaefoliumn 6, 101 Solanum anthropophagorum 366 Solanumnantillarum 101, 112 Solanum aphyodendron 7, 8, 23, 28, 29, 34, 71*, 73, 76, 77*, 90*, 112, 134 Solaniuni apoduim 340, 344 Solanurn apoense 123 Solanuni aquatile 184 Solanum arboreum species group 12, 14. 15, 17, 22, 29, 41, 194, 195*, 196*, 289, 298 Solanum arboreum 7, 8, 10, 13*, 14*, 19*, 23, 31, 34, 194, 195*, 197, 200, 201*, 202*, 218 Solanum ardisioides 364 Solanum arenarium species group 12, 15, 19, 22, 41, 248*, 273 Solanum arenarium 273, 274*, 278* Solanum argentinum 8, 366 Solan7urnlargentin uni var. chroniotrichumn366 Solanurmalureifolium 259, 262 Solanum bahianum 224, 225*, 232* Solanum barbulatum 165, 168*, 169, 170*. 176, 280 Solanlrm bassoviicarplum 157 Solanum bellum 195*, 224, 225, 226*. 232* Solanum boerhaviifolium 365 Solaiurnl brachystachys 20, 262, 289 Solanuirm brenesii 298, 302 Solanum brevifolium 364 Solanurmbrevipedunculatum 142, 146 Solanum caavurana 34, 71*, 72, 75*, 84*, 122, 128, 134 Solanurmcaavurana formapauciflora 84 Solanurtm caeruleum 87, 89, 92 Solanum callianthum 176, 213, 255, 256*, 261*, 268, 271 Solanurnm callium 123 Sol/a7lllnucallobotrvs 191 Solan urmcal/vcopogon 182 Sola111inu calysinumn364 Solanum campaniforme 11, 16*, 23, 34, 71*, 73, 87, 88*, 89*, 90*, 99 Solanum canoasense 366 Solanum capillipes 314, 315, 316*, 318*, 333, 349 So/anurn capsicastrurn 62 Solanion capsicastrurn var. caagua-zense 63 Solanum caripense 365 Solanum cassioides 72, 92, 93*, 94*, 132

FLORA NEOTROPICA Solanum cataractae 307, 366 Solanum caudatum 363 Solanum cearense 87 Solanum cervantesii 366 Solanum chenopodioldes 365 Solanum chlamydogynum 10, 17, 41, 282, 283, 284*, 287*, 311 Solanum chloranthum 273, 275 Solanum clavatum 262 Solanum clivorum 43, 45*, 46* Solanum coarense 140, 146 Solanulm coibae 142, 146 Solanum conanthum 364 Solanum concinnum 367 Solanum confertiseriatum 17, 23, 30, 41, 282, 285, 286*, 287* Solanum confine species group 10, 12, 14, 15, 25, 32, 36, 41, 311, 312*, 313*, 314 Solanum confine 23, 312*, 314, 315, 317*, 318*, 321, 340, 350 Solanlrm confine var. curtum 292 Solanum confusum 262 Solanurm copevanurm 187, 189 Solanum cordioides 72, 358*, 361, 362*, 363* Solanum cordovense 367 Solanum cormanthum 364 Solanum cornifolium 213, 248*, 255, 257, 258*, 265* Solanum corumbense 52, 136, 138, 139*, 140* Solanurmcrispllumn 229* Solanurn crotalobasis 52, 259, 262 Solanum cruciferum 314, 319, 320*, 323* Solanum cucullatum 14, 23, 140, 236*, 237, 239, 240*, 245*, 247 Solanum cyclophyllum 224, 227, 228*, 232*, 234, 235 Solanum daphnophyllum 72, 94*, 95,*, 120, 361 Solanum darienense 315, 318*, 319, 321* Solanum dasyneuron 25, 249, 250*, 252, 254 Solanum deflexiflorum species group 15, 41, 179, 181*, 289, 298 Solanum deflexiflorum 182, 183*, 184*, 189, 246 Solanum deflexiflorurnvar. diversum 243, 246 Solanum delicatulum 364 Solanum delitescens 356, 357*, 358* Solan um delitescens var. amphidoxuim 356 Solanuim devernicascens 146 Solanurmdibrachiaturn 302 Solanurn dichotomum 366, 367 Solanum didynum 367 Solanum didynum var. subglabrum 367 Solanum didynum var. subvirgatum 367 Solanum didynum var. tomentosum 367 Solanurn difloruln 62, 68

Solanurm difloriumiivar. angutstifolium 62 Solanum diflorum var. pulveritlentum 62 Solanum dimorphandrum 367 Solanum diphyllum 5, 6, 22, 23, 28, 39, 44*, 55, 56*, 57*, 62, 124 Solanum dissimile 23, 312*, 314, 321, 322*, 323*. 324, 325 Solanum distichum 364, 365 Solanum disticophyllum 3, 367 Solanum divaricaturm366 Solanum dolichostvl/um 200, 206 Solanum dolosum species group 10, 14, 15, 40, 350 Solanum dolosum 351, 352*, 353*, 354 So/lanu77ldotanumn366 Solanum dulcamara 6 Solaniunmduinalianiuo,u group 366 Solanurni dunnianuirnn 62 Solanuoti dusenii 87, 92 Solanum ecuiador-ense 169 Solanumnecuadorense var. glab;iuscudlum169 Solanum ecuadorenise var. illini:ae 169 Solanunmecutadorense var. mIodicepilosurm 169 Solanlinmecuadorense var. nervisequurnm169 Solanum edwardsii 367 Solanurni ellipticion1 group 365 Solanurn enchylozumn200, 206 Solanlim ereinanthlini 62 Solanum eriocalyx 367 Solanum erosomarginatum 23, 314, 323*, 324* Solanulm eshbauighianurnm 142, 146 Solanum evonymoides 366 Solanum extensum 367 Solanumnextraxillare 142, 146 Solaniuni falcatunm87 Solanum falconense 197, 206, 207*, 208*, 215 Solanum filiforme 365 Solanum flaccidum 365 Solaiurni flagrans 115 Solanum foetens 10, 23, 40, 248*, 249, 250*, 251*, 254 Solanuin foetidumn 6, 84, 142 Solanurnm fontiumi 252 Solanurm JOssarurln 84 Solanurni fulvivill//osu 52 Solanum furcatumn365 Solanum gemellum 367 Solanum gertii 73, 94, 96, 97* Solanum glomuliflorum 364 Solanum gnaphalocarpon 14, 19, 273, 275, 276*, 277*, 278* Solanum goniocaulon 172, 194, 208*, 209* Solanum gonyrhachis 351, 352*, 354* Solanurn gracilescens 172 Solanum gracillimum 367

401

INDEX OF SCIENTIFIC NAMES Solanurmgrandlfolium 302, 307 Solanum gratum 23, 196*, 208*, 211*, 215 Solanum habrocaulon 351, 352*, 354, 355* Solanum havanense species group 39, 368 Solanum havanense 368 Solanum heleonastes 237, 239*., 241, 242* Solanum liispidum 52 Solani rn hygrophilum 62 Solanum hypaleurotrichum 23, 43, 44*, 46*, 47* Solanurm hypiodes 52 Solanum hypocalycosarcum 23, 312*, 314, 323*, 325, 326* Solanurm hvpornalacophvllurn 48 Solanlunz hypomalacothrix 157 So/lanurn hypomicropogon 73 Solan urmhY'postichopogon 166

Solanum leucocarpon 10, 12, 13*, 16*, 17, 23, 38, 136, 137*, 138, 140, 141*, 143* var. leucocarpon Solanum multiflorum 84 Solanum lindenii 17, 136, 140*, 146, 147*, 149 Solanum linkianum 363 224, Solanum longevirgatum 227, 229*, 232*. 235 Solanum loretoanuni 326, 328 Solanum lucens 34, 196*, 197, 207, 213*, 214* Solanum lundellii 368 Solanum lycopersicum 12

Solanum malacothrix 55, 57*, 58, 60, 61* Solanum malletii 152*, 153, 154*, 155*, 157, 163 Solanum manicatum 366 Solanum mapiricum 358*, 360* 224, Solanum marantifolium Solanum imberbe 10, 181*, 182, 184, 230, 231*, 232* 185*, 193*, 289, 298, 344 Solanum marginatum 30 Solanionniinaequiale 114, 117 Solanum marmellosanun? 302, 307 Solanum incomptum 13*, 182, 184*, Solanum mnartensii367 187, 188* Solanurniinconiptlunvar. longuiispilosum Solanum maturecalvans 20, 23, 52, 248*, 255, 259, 260*, 187 261*, 268, 270, 271 Solanuoi incomptum var. lugens 187 Solanum megalocarpon 84 Solanumnindicum 364 Solanum megalochitoni 368 Solanurlniindigojerum 87 Solanum megalochiton var. villosoSolanum inelegans 368 tomentosum 368 Solanum intermedium 25, 72, 92, 96, Solanum membranaceum 365 98*, 101* Solanum mexiae 63 Solanurn ipecacuanha 63 Solanum micranthurn 101 Solan uinripecacluanha var. calvescens Solanum microleprodes 153, 63 155*, 156*, 161 Solanurn ipecacuanha var. obovata 63 Solanlum microleprodes var. filicis Solanum irregulare 11, 182, 184*, 155, 157 189, 190* Solanum microrbitum 87, 92 Solanum isabellei 365 Solanumnmodestum 366 Solanum isodynanum 368 Solanum monadelphum 10,41,197, Solanlrm jaliscanumn 62 282, 287*, 288*, 298, 307, 344 Solanunmjamesoni 48 Solanum monarchostemon 367 Solanurn jasminoides 365 Solanum morii 10, 314, 331*. 332*, 333 Solanntm karstenii 62, 68 Solanum mortonianum 365 Solanum kenoyeri 200 Solanumnmultinerviuni 262 Solanurn kieslingii 259 Solanurn? muitisii 257 Solanum kleinii 55, 58, 59*, 60* Solanuin lacteum 364 Solanum laevigatum 197, 210, 212*, 213*, 258 Solanum lasiocladum 358*, 359* Solanum lasiopodium 72, 73, 99, 100*, 101* Solanum laurifrons 165, 168*, 172, 208 Solanurnm laxiflorum 87, 92 Solanurm lepidocaule 216 Solanum leptopodum 13*, 23, 313*, 314, 326, 327*, 331*, 343 Solanum leptorhachis 14, 23, 34, 312*, 314, 329, 330*, 331*, 335, 344, 345 Solanum leucocarpon species group 12. 15, 40, 136, 137*

Solanum narcoticosmum species group 15, 40, 41, 247, 248* Solanum narcoticosmum 249, 250*, 252, 253* Solanum nelsoni 365 Solanum nematorhachis 314, 331*, 333, 334* Solanum nigricans species group 12, 14, 15, 41, 248*, 254 Solanum nigricans 12, 20, 255, 262, 263*, 265* Solanum nigruin 27 Solanum nitidum species group 5, 8, 40, 404 Solanum novo-granatense 210 Solanum nudum species group

12, 14, 15, 25, 32, 40, 41, 69. 71*, 361 Solanum nudum 5, 6, 13*, 14*, 16*, 21*, 23, 29, 34, 35, 37*, 38, 73,76, 83, 84, 99, 101*, 102*, 124, 129 Solanum nudumvar. longifblium191 101 Solanum nudumvar. micranthumn Solanum nudum var. pseudoindigoferum 125, 128 Solanum nutans species group 12, 14, 15, 19, 40, 41, 152* Solanum nutans 40, 152, 153, 157, 158*, 159*, 160*. 165 Solanum oblongifolium species group 10, 11, 15, 40, 43. 44* Solanum oblongifolium 6, 10, 23, 43, 44*, 45, 48, 49* 50*, ;4 var. Solanurm oblongifoliuim soukupii 48 Solanum oblongum 6, 14, 19*, 21*, 23, 136, 137*, 143*, 148, 149, 150*, 151* Solanunr oblonggumvar. abrupttur 149 Solanum obovalifolium 17, 282, 289, 290*, 291*, 307 Solanum obovatum 210, 213 Solanum ochrophyllum 255, 262, 265*, 268, 269* xar. Solanurm ochrophyll/um schmidtii 259 Solanum ombrophilum 23, 314, 335* ,336*, 338 Solanum oppositifolium 6, 10, 23, 282, 283, 291*, 292, 293*, 307 Solanum orygale 200 Solanum pabstii 367 Solanum palinacanthum 30 Solanum palmillae 282, 289, 296, 297*, 300* Solanurn parcebarbaturm101, 113 Solan um parcebarbaturn v/ar. minorifrons 101 Solanum pastillum 11, 23, 26*, 27, 28, 31, 35, 313*, 314, 326. 336*, 337*, 347 Solanurmpatellare 140, 146 Solanum pavimenti 64 Solanum pearcei 73, 76 Solanum perattenatum 368 Solanum pertenue 28, 31*, 35*, 314, 318, 319, 321, 338, 3;9*,

343 * Solanum platycypellon I 1. 255, 265*, 270* Solanum plowmanii 196, 210, 213*, 215, 216* Solanurn pl/urifiircipilurn 62 Solanum polyaster 285 Solanum pruriens 368 Solanum pseudocapsicum species group 11, 14, 15, 22. 29, 39, 40, 44*, 54, 55, 278 Solanum pseudocapsicumn 5, 6, 7, 8, 12, 17, 2 3, 2 8, 3 5, 3 6 3 8 39, 44*, 55, 60, 62, 63*, 65*,

402

FLORA NEOTROPICA

69, 125, 278, 280 Solanum pseudocapsicum forma pilulosum 63 Solanum pseudocapsicum ssp. diflorum 63 Solanum pseudocapsicum ssp. diflorum var. ambiguum 63 Solanuin pseudocapsicum ssp. diflorurmvar. hygrophilum 63 Solanuin pseudocapsicum ssp. diflorurm var. hygrophilum forma calvescens 63 Solanum pseudocapsicum ssp. diflorum var. sendtnerianum 63 Solanum psetrdocapsicumssp. diflorum var. typicum 63 Solanum pseudoquina 12, 35, 38, 73, 96, 113*, 114*, 115*, 119, 364 Solanum psidiifolium 146, 148 Solanum psychotrioides 165, 169, 172, 173*, 176* Solanum pterocladum 366 Solanum pteropodum 48 Solanum puberuloba 292, 296 Solanum pubescens 364 Solanum pubigerum 366 Solanum pil/chrum 302, 307 Solanumpulchrum var.peruvianum 302 Solanum pulverulentum 366 Solanum punctulatum 368 Solanum quebradense 265*, 271, 272*

255, 257,

Solanum ramonense 23, 3 1, 194, 213*, 216, 217*, 222 Solaurumramosissimum 114 Solainum rancaguense 365 Solanum receptum 365 Solanum reductum 365 Solanum refractifolium 365 Solanum reitzii 12, 25, 38, 72, 99, 117, 118*, 127* Solanum repandum 124 Solanum restingae 72, 95, 115*, 119* Solanum ripense 23, 35, 194, 219, 220*, 221*, 223 Solanum ripivagurm200 Solanum rivnylare 340 Solanum roblense 197, 218, 219, 221* Solanum robustifrons species group 15, 17, 22, 41, 235, 236* Solanum robustifrons 23, 35, 206, 236*, 237, 243, 244*, 245* Solanum rovirosanum 13*, 23, 28, 30, 31, 32, 35, 282, 283, 287, 298, 299*, 300* Solanum rufescens 368 Solanum santosii 72, 115*, 120, 121*,

225 Solanum scabrum 6 Solanum schippii 298 Solanum schizopodium 292, 296 Solanum schwackeanum 368 Solanum(?) serratum 365 Solanum serratifolium 365 Solanum sessile species group 10, 11, 12, 14, 15, 21, 22, 30, 41, 96, 282 Solanum sessile 6, 11, 16*, 23, 27*, 30, 38, 282, 283, 289, 292, 302, 303*, 309* Solanum sieberi 23, 28, 35, 181*, 182, 187, 191, 192*, 193* Solanum silvicolum 251 Solanum smithii 41, 248*, 273, 275, 278*, 279* Solanum solum 368 Solanum spirale 14, 22, 23, 28, 29, 39, 40, 72, 122*, 124* Solanum spissifolium 55, 60*, 69, 70* Solanum sprengelii 273 Solanum squamuliferum 257 Solanum stipulatum 12, 313*, 314, 328, 340, 341*, 342*, 343* Solanum sublobatum 365 Solanum subsylvestris 368 Solanum superbum 237, 245*, 246, 247* Solanum superficiens 22, 123 Solanum supranitidum 101 Solanum surinamense 140, 146 Solanum symmetricum 72, 86, 125, 126*, 127*, 361 Solanum tanysepalum 11, 22, 23, 94, 194, 196*, 200, 219, 221*, 222* Solanum tenuiflagellatum 14*, 15, 23, 25, 35, 313*, 314, 331, 335, 343*, 344, 345* Solanum tenuilamellulosum 345 Solanum tepuiense 72, 127*, 128, 129* Solanum thelopodium species group 8, 15, 39, 367 Solanum thelopodium 367 Solanum tovarense 99, 101, 112, 113 Solanum tovarii 165, 174, 175*, 176* Solanum trachytrichium 11, 19*, 38, 72, 94, 129, 130*, 131* Solanum trichoneuron 8, 10, 73, 83, 131*, 132, 133* Solanum triplinervium 38, 224, 227, 232, 233*, 235* Solanum triquetrum 366

Solanum triste 6, 10, 23, 36, 282, 285, 307, 308*, 309*, 311 Solanum triste var. crassipes 140 Solanum troyanum 368 Solanum tucumanense 62, 168 Solanum tuerckheimii 10, 23, 28, 314, 326, 345, 346*, 352* Solanum tumescens 138, 140 Solanum tunariense 273, 275. 278*, 280, 281* Solanum turgidum 21*, 23, 30, 283, 309*, 310* Solanum ulmoides 62 Solanum umbellatum 365 Solanum undatifolium 114 Solanum undulatuni 340 Solanum unifoliatum species group 12, 15, 40, 195*, 223 Solanum unifoliatum 224, 227, 230, 232, 234*, 235* Solanum uporo 366 Solanum urceolatum 6, 292 Solanum vacciniiflorum 19, 23, 165, 174, 176*, 177*, 179 Solanum valerianum 314, 315, 343*, 347, 348* Solanum validinervium 11, 19*, 52, 166*, 174, 176*, 178, 179, 180*, 252 Solanum validum 62, 68 Solanum venosum 52, 53*, 54* Solanum verniciflorum 259 Solanum vernicinitens 262 Solanum verruculosum 138 Solanum versabile 125 Solanum viliflorum 292, 296 Solanum warmingii 72, 86, 122, 131*, 134, 135* Solanum xanthophaeum 161, 162*, 163*

153,

Solanum yanamonense 23, 25, 314, 318, 343*, 349, 350* Solanum youngii 152*, 153, 155, 163*, 164* Solanum zamorense 243 Spinosa 4 Stenotritus 25 Sturnira ludovici 29 Tetrardisia 364 Tityra semifasciata 29 Tournefortia 32 Trigona 25 Xylocopa 25, 26

403

APPENDICES

APPENDIX I Solanumsection CernuumCarvalho& Sheph.(accepted names only; see Carvalho, 1996, for complete treatment) 1. Solanum caldense Carvalho 2. Solanum castaneumCarvalho 3. Solanum cernuumVell. 4. Solanumoliveirae Carvalho 5. SolanumpachinatiumDunal 6. Solanum leucodendronSendtn. 7. Solanum sooretanum Carvalho 8. Solanum vellozianumDunal

APPENDIX II Solanum section Lepidotum (Dunal) Seithe (accepted names only; see Carvalho, 1996, for complete treatment) 1. Solanum argenteumDunal 2. Solanum carautae Carvalho 3. Solanum cinnamomeumSendtn. 4. Solanum davidsei Carvalho 5. Solanumhatschbachii Carvalho 6. Solanum lepidotumDunal var. lepidotum 7. SolanumlepidotumDunalvar. lepidiochlamys Carvalho 8. SolanumlepidotumDunalvar.trianaeCarvalho 9. Solanumsellowii Dunal 1o. SolanumsteyermarkiiCarvalho 11. Solanum swartzianumRoem. & Schult. ssp. swartzianumvar. swartzianum 12. Solanum swartzianumRoem. & Schult. ssp. swartzianumvar. sordidum Sendtn. 13. SolanulmswartzianumRoem. & Schult. ssp. chrysophyllum(Dunal) Carvalho 14. SolanurmswartzianumRoem. & Schult. ssp. (Dunal) Carvalho argyrophyvllum

APPENDIX III Solanum section Pachyphylla Dunal (from Bohs, 1994, 1995; accepted names only; names in both Solanum and Cyphomandragiven) 1. SolanumbetaceumCav. (Cyphomandrabetacea (Cav.) Sendtn.) 2. SolanumcacosmumBohs (Cyphomandrafeotida Bohs) 3. Solanum cajanumense Kunth (Cyphomandra cajanumensis(Kunth)Walp.) 4. Solanum calidum Bohs (Cyphomandrapilosa Bohs) 5. Solanum circinatum Bohs ssp. hartwegii (Qyphomandrahartwegii (Miers) Walp.)

6. Solanum circinatumBohs ssp. ramosa Bohs 7. Solanum corymbiflorum(Sendtn.) Bohs ssp. corymbiflorum(Cyphomandracorymbiflora Sendtn.) 8. Solanum corymbiflorum(Sendtn.) Bohs ssp. mortonianum(L. B. Smith & Downs) Bohs 9. Solanum diploconos (Mart.) Bohs (Cvphomandradiploconos Mart.) 10. SolanumdiversifoliumDunalssp. chloranthum (Rusby) Bohs 11. SolanumdiversifoliumDunalssp. diversifolium (Cyphomandradiversifolia(Dunal)Bitter) 12. Solanum endopogon (Bitter) Bohs ssp. endopogon (Cyphomandraendopogon Bitter) 13. Solanum endopogon (Bitter) Bohs ssp. guianensisBohs benensis 14. SolanumexiguumBohs (Cyphomandra Britton) hvpo7nalaca 15. SolanumfallaxBohs(Cyphomandra Bitter) dolicho(Cyphomandra 16. SolanumfortunenseBohs carpaBitter) calycina 17. SolanumlatiflorumBohs (Cyphomandra Sendtn.) 18. Solanum maternumBohs 19. Solanum melissarum Bohs (Cyphomandra divaricata (Mart.)Sendtn.) 20. Solanum obliquum Ruiz & Pav6n (Cyphomandraobliqua (Ruiz & Pav6n) Sendtn.) 21. SolanumoccultumBohs (Cyphomandrastellata Bohs) 22. Solanumovum-fringillae(Dunal)Bohs (Cvphiomandraovum-fringillaeDunal) 23. Solanum oxyphyllum C. V. Morton (Cyvphomandrafragilis Bohs) 24. SolanuimparalumBohs (CyphomandraheterophyllaTaubert) 25. SolanumpendulumRuiz&Pav6n(Cyphomandra pendula (Ruiz & Pav6n) Sendtn.) 26. Solanum pinetorum (L. B. Smith & I)owns) Bohs (CyphomandrapinetorumL. B. Smith & Downs) 27. Solanumpremnifolium(Miers) Bohs (Cyphomandrapremnifolia (Miers) Dunal) 28. Solanum protanthum Bohs (Cyphomandlra oblongifolia Bohs) (Standl.& Steyerm.)Bohs 29. Solanumrojasiantum (Cyphomandra rojasianaStandl.& Steyerm.) 30. Solanum roseum Bohs (Cyphomandra acuminata Rusby) 3 1. Solanumsciadostylis (Sendtn.) Bohs (Cyphomandra sciadostylis Sendtn.) 32. Solanum sibundoyense (Bohs) Bohs ( Cvphomandrasibundoyensis Bohs)

FLORA NEOTROPICA

404

12. Solanumpelagicum Bohs 33. SolanumsycocarpumMart.& Sendtn.(Cypho13. SolanumstuckertiiBitter mandrasycocarpa(Mart.& Sendtn.)Sendtn.) 34. Solanum tegore Aubl. (Cyphomandrategore (Aubl.) Walp.) APPENDIX V 35. Solanum tenuisetosum(Bitter) Bohs (CyphoSolanumsectionAllophyllum(Child)Bohs (accepted mandra tenuisetosumBitter) 36. Solanum tobagense (Sandwith)Bohs (Cypho- names only; see Bohs, 1990, for completetreatment) mandra tobagensis Sandwith) 1. Solanumallophyllum(Miers) Standl. 37. Solanum unilobum (Rusby) Bohs (Cypho2. Solanum mapirienseBitter mandra uniloba Rusby) 3. SolanummorellifoliumBitter

APPENDIX IV

APPENDIX VI

Solanumsection Cyphomandropsis Bitter(accepted Solanum nitidumspecies group (accepted species names only; see Bohs, 2001, for complete treatment) only; see Knapp, 1989, for complete treatment) 1. Solanum amotapense Svenson 1. Solanum crispumRuiz & Pav6n 2. Solanum confusumC. V. Morton 2. Solanum cutervanumZahlbr. 3. SolanumcylindricumVell. 3. Solanum imbaburenseS. Knapp 4. Solanumfallax Bohs 4. Solanum leiophyllumBenth. 5. Solanumfusiforme L. B. Smith & Downs 5. SolanummacbrideiHunz. & Lallana 6. SolanumglaucophyllumDesf. 6. Solanum muenscheriStandl. & Steyerm. 7. Solanum hibernumBohs 7. SolanumnitidumRuiz & Pav6n 8. Solanum hutchisonii (J. F. Macbr.)Bohs 8. Solanum ruizii S. Knapp 9. Solanum luridifuscescensBitter 9. SolanumstenophyllumDunal 10. Solanum luteoalbumPers. 10. Solanumstorkii C. V. Morton& Standl. 11. Solanum matadori L. B. Smith & Downs