Rhodostemonodaphne (Lauraceae)

Santiago Madriñán

A phylogenetic analysis based on nuclear ITS and plastid trnK intron sequences confirms that Dahlgrenodendron, Sinopora, Triadodaphne, and Yasunia are members of the Cryptocarya group, as expected from morphology. Dahlgrenodendron from South Africa is sister to Aspidostemon from Madagascar. Triadodaphne inaequitepala is nested within Endiandra (both from Australasia), and Yasunia from South America is nested among South American Beilschmiedia species. Sinopora is a member of the Beilschmiedia clade, but its precise position is still uncertain. Among large genera of the group, Cryptocarya is clearly monophyletic, and Endiandra appears to be as well, if T. inaequitepala is included. Beilschmiedia is paraphyletic with respect to (at least) Potameia and Yasunia. Most wellsupported clades within genera are geographically homogeneous, except a clade including the Chilean Cryptocarya alba and two New Caledonian species. Both Beilschmiedia and Cryptocarya have reached the Americas more than once. Four-locular anthers are plesiomorphic in the Cryptocarya group; two-locular anthers have arisen by fusion of the two pollen sacs of a theca. In the plesiomorphic fruit type, the ovary is completely enclosed in receptacular tissue; a superior fruit, seated free on its pedicel, is a synapomorphy of the Beilschmiedia clade.

"Background and aims – The collections of Lauraceae in the Van Heurck Herbarium (AWH), which is currently on permanent loan to BR, have remained unknown by most of the specialists and by the general public up until the present. The taxonomic status of its one hundred and forty nine specimens of Lauraceae is here presented. Methods – Standard herbarium taxonomy practices were used in conjunction with the literature available. Specimens from fifty eight different herbaria were studied personally, or by checking their holdings available in the internet, or from digital images received from the curators. Key results – The specimens of Lauraceae from AWH are distributed in twenty four genera and seventy eight species with taxonomic status resolved, excepting for three specimens pertaining to species name of status uncertain (one), or to unresolved determination (two). From them, fifty three specimens are nomenclatural types of any sort, which corroborate the relevance of exsiccates acquired by Van Heurck, even for this relatively small set of specimens for the family. Fifteen lectotypifications and one neotypification of species names are proposed here."

An investigation of a questionable species of the genus Alseodaphne led to the discovery of a new genus Alseodaphnopsis H. W. Li & J. Li, gen. nov., separated from Alseodaphne Nees, and a new species Alseodaphnopsis ximengensis H. W. Li & J. Li, sp. nov., endemic to Yunnan province, China. This new species is characterized by having big, axillary, paniculate inflorescences, as well as large, subglobose fruits. Based on DNA sequence data from two gene regions (nuclear ribosomal ITS and LEAFY intron II), we investigate its phylogenetic position within the Persea group. Phylogenies using maximum parsimony (MP) and Bayesian inference (BI) support the recognition of Alseodaphnopsis as a distinct genus but do not resolve well its relationship within the Persea group. The new genus is circumscribed, eight new combinations for its species are made, and a description and illustration of the new species are provided.

Organization for Flora Neotropica Rhodostemonodaphne (Lauraceae) Author(s): Santiago Madriñán Source: Flora Neotropica, Vol. 92, Rhodostemonodaphne (Lauraceae) (Aug. 24, 2004), pp. 1-102 Published by: New York Botanical Garden Press on behalf of Organization for Flora Neotropica Stable URL: http://www.jstor.org/stable/4393932 . Accessed: 02/04/2011 19:33 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 service that helps scholars, researchers, and students discover, use, and build upon a wide range of content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship. For more information about JSTOR, please contact support@jstor.org. New York Botanical Garden Press and Organization for Flora Neotropica are collaborating with JSTOR to digitize, preserve and extend access to Flora Neotropica. http://www.jstor.org FLORA NEOTROPICA MONOGRAPH 92 RHODOSTEMONODAPHNE(LAURACEAE) SANTIAGOMADRINAN [ - -4 - 4---^ " - Jg -- N _ TROPICOF CANCER FLORAL NEOTROPICA Publishedfor Organization for Flora Neotropica by The New York Botanical Garden Bronx, New York Issued 24 August 2004 ? 2004 by The New York BotanicalGarden All rightsreserved. Published by The New York BotanicalGardenPress Bronx, NY 10458 InternationalStandardSerial Number0071-5794 The paperused in this publicationmeets the requirementsof the AmericanNational Standardfor InformationSciences - Permanenceof Paperfor Publicationsand Documents in Librariesand Archives, ANSI/NISO (Z39.48-1992). Printedin the United States of Americausing soy-based ink on recycled paper. MetLife Foundation is a LeadershipFunderof The New YorkBotanicalGardenPress. Library of Congress Cataloging-in-Publication Data Floraneotropica.- Monographno. 1 - New York:Published for Organizationfor FloraNeotropicaby The New York BotanicalGarden,1968v.: ill.; 26 cm. Irregular. title. Eachissuehasdistinctive no.40. andclassifiedin LCbeforemonograph catalogued Separately ISSN0071-5794= Floraneotropica. 1. Botany - Latin America - Classification - Collected works. 2. Botany - Tropics - Classification - Collected works. 3. Botany Classification- Collected works. I. Organizationfor FloraNeotropica. II. New York BotanicalGarden. QK205.F58 581.98'012-dcl9 Libraryof Congress [8508] ISBN0-89327-455-0 03040506070809 10 / 9 8 7 6 5 4 3 2 1 85-647083 RHODOSTEMONODAPHNE(LAURACEAE) SANTIAGO MADRINAN CONTENTS Abstract/Res men ...............................................................................................................2 Introduction................................................................................................. . .............. 2 Taxonomic History .............................................................................................................3 Morphology ................................................................................................. . .............. 7 Habit..............................................................................................................................7 Architecture...................................................................................................................8 Indument .......................................................................................................................8 Leaves............................................................................................................................9 Inflorescences................................................................................................................9 Flowers........................................................................................................................ 10 Pollen........................................................................................................................... 10 Fruits ........................................................................................................................... 10 Distribution.......................................................................................................................11 Systematic Position .......................................................................................................... 18 Species Groups................................................................................................................. 19 19 ............... The Rhodostemonodaphnelaxa Group ................................. The Rhodostemonodaphnescandens Group............................................................... 21 23 The RhodostemonodaphnekunthianaGroup ........ ................................ 23 ................ The Rhodostemonodaphnegrandis Group............................. Species incerta sedis ...................................................................................................24 The Rhodostemonodaphnegrandis Complex ................................................................ 24 Morphometric Study ...................................................................................................26 Methods .................................................................................................................26 Results....................................................................................................................26 Conclusions............................................................................................................29 Species Concepts..............................................................................................................29 Systematic Treatment .......................................................................................................30 30 .......................... Interpretationof Descriptions........................... 31 ........ ................................ to the of Key Species Rhodostemonodaphne Acknowledgments ............................................................................................................95 LiteratureCited.................................................................................................................95 Numerical List of Taxa ....................................................................................................97 List of Exsiccatae .............................................................................................................97 Index of Local Names .................................................................................................... 100 Index of Scientific Names .............................................................................................. 100 2 FLORANEOTROPICA ABSTRACT Madriiian,S. (Departamentode Ciencias Biologicas, Universidadde los Andes, Apartado Aereo 4976, Bogota, D.C., Colombia).Rhodostemonodaphne (Lauraceae).FloraNeotropica Monograph92: 1-102. 2004.-A revision of the genus Rhodostemonodaphneis presented.The genus comprises 41 species of dioecious Lauraceaewith stamensbearing four locelli situatedin a shallow archtowardsthe apex of the anthers.A taxonomichistory of the genus is given. The variation in vegetative and reproductivemorphology is discussed. A brief discussion on the distributionof the species is offered. The systematic position of the genus is discussed, and a subdivisioninto informalspecies groups is suggested. The variationof charactersused in the segregationof the species comprisingthe R. grandis complex is presentedin detail. The species concept used in the delimitationof species in this treatmentis providedand contrastedwith species concepts currentlyused by other workersin the Lauraceae.Generic and species descriptionsare provided,with a list of exsiccatae, species illustrations,and distributionmaps. Sixteen new species are describedand four new combinationsare proposed. RESUMEN Madrifinn,S. (Departamentode Ciencias Biol6gicas, Universidadde los Andes, Apartado Aereo 4976, Bogota, D.C., Colombia).Rhodostemonodaphne(Lauraceae).FloraNeotropicaMonograph92: 1-102. 2004.-Se presentauna revision del generoRhodostemonodaphne. El genero comprende 41 especies de Lauraceae dioicas con los estambres presentandocuatro locelos situados en un arco suave hacia el apice de las anteras.Se resefiala historia taxon6micadel genero. Se discute la variaci6nobservadaen la morfologia vegetativay reproductiva.Se ofrece una corta discusi6n acercade la distribuci6nde las especies. Se discute la posici6n taxon6micadel genero y se sugiereuna subdivisionen gruposde especies informales.Se presentaen detalle la variaci6nde los caracteresusados en la segregaci6n de las especies comprendidasen el complejo R. grandis. Se provee el concepto de especies utilizado en la delimitaci6nde las especies en este tratamiento,y es contrastadocon conceptos de especie usados en la actualidadpor trabajadoresde las Lauraceae. Se proveen la descripciondel genero y de las especies, con una lista de exsicados, ilustraciones de las especies y mapas de distribuci6n.Diez y seis nuevas especies son descritasy cuatronuevas combinacionesson propuestasen este trabajo. vised; for the other 14, generic relationshipsor even basic morphology and distribution are almost unThe Lauraceaeare a large family of mainly panOf the remaining28 medium-sized to large known. tropical distribution,although some species are re9 have recently been revised. An estigenera, only stricted to the subtropicsand a few reach temperate 2170 of the total) belong to generawith mated (80% latitudes. The family has the reputationof being a 100+ species. reidentification with difficult family, species very Generic circumscriptionin the Lauraceaeis based stricted only to specialists. Furthermore,generic ciron a limited numberof charactersused in combinacumscriptionandrelationshipsarepoorlyknown.ReThis has resultedin polythetictaxa,where(with tions. cent estimatesof the size of family give a totalof 2750 few no one characteris necessaryandsufexceptions) species distributedin 52 genera(Rohwer,1993a).The for ficient group membership,and charactersfound on sizes of is based of total number species, however, the individualgenera and most of these (particularly in a particulargroup are found in several other such the large ones with 100+ species) have not been re- groups. Generaof the Lauraceaehave been traditionvised during the last century.Furthermore,of the 52 ally circumscribedprimarily on androecialconfigucurrentlyrecognized genera 14 are monotypic, 7 are ration and the fruiting receptacle. In a much-quoted statement,Kostermansnoted: "Apparentlythe comditypic, and4 morehave fewer than 10 species eachthatis, 50% of the generainclude only 43 species. Of bination of charactersis more or less indefinite and these 24 small genera only 10 have been recentlyre- almost all combinationsare represented.We may as- INTRODUCTION TAXONOMICHISTORY sume either that missing combinationsare extinct or thatthe potentialitythatthey will develop is still present" (Kostermans,1957: 29). However,Rohwer suggested that,in fact, "manyof these combinationsexist but have not yet been found" (Rohwer, 1993a: 373). Indeed, new plants with unknown combinations of characterssuggesting new genera are being found, e.g., a new species includedin the presentmonograph as R. penduliflora.Due to the lack of sufficientmaterial for a descriptionof a new genus I have included it in Rhodostemonodaphneas a species incertasedis. Modem monographicwork in the Lauraceaebegan with Klaus Kubitzki'sinterestin the family in the 1970s. Since then, two FloraNeotropicaMonographs were produced(Kubitzki, 1982; Renner, 1982-Aniba and Aiouea, respectively), along with a recently publishedrevision of the genus Licaria (Kurz,2000). Henk van der Werff's work on the family, startingin the 1980s, has increased interest in the family. Van der Werff published monographs of two mediumsized Lauraceae genera-Mezilaurus (1987) and Pleurothyrium(1993)-as well as many new species of various other genera. Jens G. Rohwer, a former studentwith K. Kubitzkiat Hamburg,also published works on the Lauraceae.These include a monograph of Nectandra (Rohwer, 1993b) and a synopsis of the large genus Ocotea and related genera (Rohwer, 1986), among many others. This recent work in the Lauraceaehas increased both collectors' awareness of Lauraceaespecimens and students'interestin their biology and taxonomy. As a result, 17 new generahave been describedsince the last generaltreatmentof the family by Kostermans (1957), and a large number of species have been addedto the family.The last four genericmonographs of Lauraceaehave increasedin averagethe numberof species by 50%. These new species are for the most partbased on novel materialrecently collected. The genus Rhodostemonodaphnehad been known as a distinctentity since very early on as the subgenus Synandrodaphneof Nectandra (Meissner, 1864). It was reinstatedby Rohwer and Kubitzkiin 1985, and 8 species were transferredby Rohwer in 1986. Various authorshave since published three new species (Baitello & Coe-Texeira1991; van der Werff, 1991b; van derWerff& Vicentini,2000), andthreeadditional species have been transferred(van der Werff, 1989b; Chanderbali,2004). As a result of the work carried out towardsthe completionof the currentmonograph, 7 new species were published (Madrifiin, 1996), 16 additionalnew species are here described,and 4 new transfersare necessary. The bulk of this revision is 3 based on a doctoral thesis submitted in 1996, and therefore very few specimens collected since 1995 have been included in the present work. This may affect the species distributionranges given. The assemblage of species included here in Rhodostemonodaphneis a highly heterogeneousone, containing various species groups that may be more closely relatedto species in othergenerathanto each other. Furthermore,species currently assigned to other genera, in particularEndlicheria and Ocotea, may in fact belong to the species groups included in this treatment.Due to their lack of agreementon the charactersthat define the genus Rhodostemonodaphne,andagreementwith definingcharactersof the othergenera,these species have been left in theircurrently assigned genera. TAXONOMIC HISTORY The genus Rhodostemonodaphnewas originally described as Synandrodaphneby Carl Friedrich Meissner in his monographof the family Lauraceae for De Candolle's ProdromusSystematis Naturalis Regni Vegetabilis(1864). Includedin it were two species, S. laxa and S. antillana. The formerwas a new species based on a single flowering individualfrom Quindioin Nueva Granada(Colombia).The latterwas a new name for the then relatively well known Jamaican species, Nectandra staminea Griseb., of which Meissnernotedthatits flowerstructurediffered entirely from that of the genus Nectandra. The generic charactersof Synandrodaphneas defined by Meissner were "hermaphroditic?" flowers, nine fertile stamensconnatein a ring at the base (the characteron which the generic name was based), and anthers with four locelli in superposed pairs. That questionmarkseems to be an indicationof Meissner's uncertaintyabout the sexuality of the plants (he describedthe ovary as "sterile?"andhadnot seen fruits) ratherthanthe occurrenceof bothhermaphroditicand dioecious species in the genus; Synandrodaphnelaxa is now known to be dioecious, while S. antillana is indeed hermaphroditic.The genus was placed in the tribe Oreodaphneaewith 11 other genera, including Endlicheria,Nectandra,Ocotea, and Pleurothyrium. Henri Ernest Baillon in his Histoire des Plantes (1870) followed Meissner's treatment,accepting the genus andits position withinthe family.In the generic description(a near literal transcriptionof Meissner), characterssuch as hermaphroditicflowers (withouta question mark), antherswith superposedlocelli, and liguliform staminodiaall refer to S. antillana rather than to S. laxa. 4 In Bentham and Hooker's Genera Plantarum, George Bentham (1880) included the genus in the tribe Perseaceae[sic] (a broadtribalcircumscription comprising three of Meissner's tribes and including more or less the same genera that Meissner [1864] recognized). His description of the genus, although somewhat refined, changed little in substance from the one providedby Meissner. The charactersrepresentative of S. laxa given by Bentham are turbinatecampanulate(here referredto as trumpet-shaped)receptacle, stamens of outer two whorls basally connate, with dorsal squamulateglands (these are in fact partsof the deeply dissected glands of whorl III, which in turn were described as reduced or "nearly obsolete"). Also included are those charactersmentioned above found only in S. antillana. However, BenthamexcludedS. antillana, arguingthatit did not agree with the generic characters.Instead,he considered it similar to Oreodaphnefragrans Meissner, or possibly belonging to eitherthe genus Persea or Ocotea (Oreodaphnefragrans and S. antillana are now synonymsof the older nameNectandrastamineaGriseb. -Ocotea staminea (Griseb.)Mez [Mez, 1889]). Thus, the genus was left as monospecific with a description that partly described the single species, S. laxa, but also includedcharactersnot found in it. Ferdinand Albin Pax, who monographedthe family for Engler and Prantl's Natiirlichen Pflanzenfamilien (1891), providedan abbreviateddescriptionalong the lines of Bentham's. This taxonomic history of Synandrodaphne, where generic descriptions in the different family treatmentsare mere transcriptionsof those provided by the initial workers, is not uncommon. Here, as elsewhere,little or no corroborationof characterswas sought,in particularof the floralcharacters,which are importantbut difficult to observe. Furthermore,we see that changes in circumscriptionof the genera, added subsequentto the original descriptionand accepted by latermonographers,very often had little or no effect on the generic descriptions,leaving generic concepts at best ill-defined. Carl ChristianMez was a notable exception. His excellent monograph of the American Lauraceae (1889) remains, after more than a century, a most valuable(and the only!) comprehensivespecies work for the studyof Lauraceaefromthis region.Of course, there have been some generic name changes, species transfers,and a considerableincrease in the number of taxa since then. Mez's description of S. laxa [Nectandra laxa (Meissner) Mez], based on original observationsof the type specimen at Kew, confirmed the staminatecondition of the flowers and thus the FLORA NEOTROPICA dioecious natureof the species. Furthermore,no mention is made of "squamulae"between the anthersof whorls I and II, and the absence of staminodia of whorl IV is correctly noted, contraryto the notion of previous authors. Mez (1889) treatedSynandrodaphneas a subgenus of Nectandracharacterizedby dioecious flowers. He included Synandrodaphnein Nectandra for two reasons. First, he thought it was clearly associated with Nectandra, with N. arottiana linking it to the next subgenus,Eunectandra.Second, he arguedthat the dioecious condition did not by itself constitutea generic characteras evidencedby the inclusion of dioecious and hermaphroditicspecies in Ocotea. Endlicheria, the only dioecious genus he recognized in the Perseeae,was separatedby at least one othercharacterfromits closest genera(Aniba,Aiouea, and Ocotea). In Nectandra subgen. Synandrodaphne,Mez included four additionalspecies. Two of these, N. velutina and N. anomala, were new. The othertwo species were N. dominicana(Meissner)Mez (originally described by Meissner as the penultimatespecies of the large, mainly dioecious Oreodaphne,althoughhe had qualified his placement with a question mark), and N. arottiana. Nectandraarnottianasensu Mez, based on PleurothyriumchrysothyrsusMeissner,was erroneouslyattributedto Nees, whose name Nectandra arnottianarepresentsa differentspecies (see discussion underRhodostemonodaphnekunthiana). A few otherdioecious species were addedto Nectandra in the following years. Mez (1920) in one of his numerousAdditamentamonographica,described N. dioica, with no subgeneric placement indicated. Noel Yvri Sandwith, who visited British Guiana in 1929 as partof an OxfordUniversityexpedition,collected and describedNectandrapraeclara (Sandwith, 1932). He explicitly placed it in subgen. Synandrodaphne.Years later he describedN. miranda (Sandwith, 1952), pointing out its unique characters(see species description),but arguedfor its placementin Nectandra on its agreement in staminal characters (antherlocelli in an arch) with other species in that genus. Albert CharlesSmith (1935) describedN. superba from a pistillateindividual,not noting the sterile anthers. Andre Joseph Guillaume Henri Kostermanswas one of the most prolific studentsof the Lauraceaein the 20th century;however, he did little species-level work in Nectandra and its allies. He transferredtwo species of dioecious Lauraceaeinto Nectandra(Kostermans, 1936). Nectandra grandis (Mez) Kosterm. had originallybeen placed by Mez (1889) in the two- TAXONOMICHISTORY locellate Endlicheria(with a questionmark,as he had seen only a fruiting specimen). The other species, Nectandrakunthiana(Nees) Kosterm.,was based on AcrodiclidiumkunthianumNees. Acrodiclidiumwas a genus comprising hermaphroditicspecies with anthers with two apical locelli, most of which are now included in Licaria. ChristianGottfriedDaniel Nees von Esenbeck (1836) had described A. kunthianum from a pistillate specimen with flowers and developing fruits (apparentlyonly the fruitswere availableto Nees; these, nevertheless,have remnantsof the staminodes). Thus, Nees could not confirm the unisexuality of the flowers (even with fresh flowersit is often hardto distinguishthe staminodesof pistillateflowers from fertile stamens).He then describedthe species, stating that the type did not have perfect flowers, although he had not seen flowers ("Vidiin Herb.Kunthiano absque floribus perfectis... Flores non vidi" [Nees, 1836: 269]). Nevertheless, his observationof the stamens as being two locellate was erroneous. Meissner (1864) treatedA. kunthianumas a synonym of AydendronaciphyllumNees. Aydendronis another genus of hermaphroditic,two-locellate species, now consideredto be a synonymof Aniba. Thus, its placement in that genus was also incorrect. Mez (1889) clarified the dioecious nature of the species and its four-locellatecondition, and placed it in Ocotea, reinstating the earlier epithet kunthianum.Three other heterotypic synonyms of Nectandra kunthianahave been named (van der Werff, 1984). Mez (1903) described Ocotea stenoneura,citing two fruitingcollections from two differentspecies. One of the syntypes, treated as a paratypeof 0. cooperi C. K. Allen by Allen (1945), is conspecific with Nectandra kunthiana. Caroline K. Allen (1964) also described Pleurothyriumcowanianum from a pistillate individual, erroneously citing and illustrating the presence of pairedglands at the base of all nine anthers,thus fitting the then current definition of Pleurothyrium. Nevertheless,she noted thatit closely resembledNectandrameyerianaLasser,a thirdsynonym of N. kunthiana. Nectandra meyerianawas properlyassigned to subgen. Synandrodaphneby its authorTobiasLasser (1948). Kostermans'shigher-level work had profoundrepercussions for the circumscriptionand delimitation of all genera. In his proposed system of the family (Kostermans,1957) he includedNectandraandPleurothyriumas subgenerawithin Ocotea, which resulted in an unmanageablegenus of approximately700 binomials. In his view, the differencesin position of the locelli in the antherswhich distinguishedNectandra and Ocotea was "certainlynot of generic value, as 5 intermediatecases are not uncommon"(Kostermans, 1957: 38). His approach,however, was not new. As early as 1791, Schreber(in the 8th edition of Linne's Genera Plantarum)had merged the two genera, and since then various authorshave done so in different ways (see Rohwer, 1993b). One novel solution was thatof Howard(1981) who, when preparingthe treatment of the family for the Floraof the LesserAntilles, placed the species of the two genera intermingledin alphabetical order by epithet. Luciano Bernardi (1962) discussed the American Lauraceaewith special referenceto theVenezuelanspecies. His treatment follows essentially that of Mez (1889) with inclusion of some of the changes proposed by Kostermans (1957), and adding some of the species describedafter Mez's treatment.Noteworthy,if only as a source of amusement,is Bernardi'sessay on the genus Nectandra (1962: 173-181). Here, in a miscellaneous "philosophical"discourse on variation,Bernardiexplained the numerous difficulties he encountered when workingon this genus. Indeed, the distinction between Nectandra and Ocotea is not all thatclear-cut.The separationof Nectandrafrom other four-locellategenerain the family dates from the work of Nees (1833, 1836). In the key to the tribes of the family in his SystemaLaurinarum, Nees used the position of the antherlocelli to separate the tribes with paniculateinflorescences.Plants with locelli arrangedin one row nearthe apex were keyed out as Nectandra; those with locelli in two superposed pairs were placed in a numberof othergenera, including Ocotea. Linked to this distinctionwas the shape of the anthers:plantswith wide, sessile anthers belonged to either Nectandra or Dicypellium, while those with a discrete filamentcomprisedthe remaining tribes (a printing error in the published key [p. ix], where the couplet separatingNectandrafromDicypellium is not indented, obscures the hierarchyof the previouskey character). Species with the locelli arrangedin ways that appearintermediatebetweenthe linearandbijugate(i.e., four locelli in two superposedpairs) conditions are not uncommon.Many of these have at various times been treatedas belonging to Ocotea and later transferred to Nectandraor vice versa. In the recent revision of Nectandra (Rohwer, 1993b), 54 binomials were excluded from Nectandra and assigned to species in Ocotea. A large numberof these belong to the 0. helicterifoliagroup (Rohwer, 1991). Jens G. Rohwer and Klaus Kubitzki (1985) proposed-as had earlier been suggested by Allen (1966)-the reinstatementof the three genera of the Ocotea complex (Nectandra, Pleurothyrium, and 6 Ocotea). They concluded that the position of the locelli on the anthersis an extremely useful character in separatingthese genera when used in combination with other characters.Taking into account variable characterssuch as the length of the filaments, the length/widthratio of the anthers, form of the apex, indumenttypes, and total length of the stamens,they established five evolutionary lines within the complex. From these they concluded that the linearly arranged locelli, characteristic of Nectandra, have evolved independentlyat least three times from an ancestralbijugatecondition. Additionally,two of the five evolutionary lines consisted of dioecious taxa, while the other three are hermaphroditic.The three groups with linear locelli were Nectandra,to a large extent coinciding with the hermaphroditicsubgen. Eunectandraof Mez (1889); some species of the dioecious Ocotea cernua group; and Synandrodaphne. Nectandra dominicana (which had already been transferredto Ocotea by Howard [1981], but previously included in Synandrodaphneby Mez) was placed as a derived member of the Ocotea cernua group. Ocotea licanioides A. C. Sm. was placed as a species intermediatebetween Synandrodaphneand the basal membersof the Ocotea cernua group. Synandrodaphnewas then reinstated,but, in a note added in proof, the new name Rhodostemonodaphnewas proposeddue to the conservedstatusof the nameSynandrodaphneGilg (Thymelaeaceae).Due to the presence of intermediateformsbetween the dioecious and hermaphroditicgroups of Ocotea, the 0. cernua group was not elevated to generic rank. Rhodostemonodaphnewas thus definedas a genus of dioecious plants, with a stronglyreducedovary in staminate flowers, distinctly fleshy, hairy or rarely glabrousstamens, with locelli in a horizontalline or shallow arch near the apex, and staminodiaof whorl IV lacking. Nectandra was distinguishedby its hermaphroditicflowers, often papillose inner surfaceof the tepals and stamens, anthers often with an acute apex, the locelli occupying the whole anther or located towardsthe base, andpresenceof staminodiaof whorl IV. Rohwer (1986) furtherdeveloped the concept of Rhodostemonodaphnein a synopsis of Ocotea s.l., noting some similaritieswith the four-locellateforms of the hermaphroditicgenus Urbanodendron.Ocotea dominicana was placed outside of the 0. cernua group due to incongruences in inflorescence morphology.The formaltransferto Rhodostemonodaphne of eight species was made, althoughmost had already been associatedwith Synandrodaphneor were known to be dioecious species of Nectandra. Two of these, FLORANEOTROPICA Rhodostemonodaphnelaxa and R. anomala, make up the R. laxa group; Ocotea licanioides A. C. Sm. and 0. macrophyllawere also consideredto belong to this group, but no formal transferwas made (0. macrophylla is now associatedwith the Nectandra-likespecies of Ocotea, the 0. helicterifolia group (Rohwer, 1991); 0. licanioides is here transferredto Rhodostemonodaphne). Nectandra dioica, which had been placed under synonymy with N. grandis by Kostermans(1936), was reinstatedas Rhodostemonodaphne dioica with N. superba as a synonym. Nectandragrandis was transferredto Rhodostemonodaphne carryingthe synonym suggested by Kostermans, Nectandra praeclara. The combinationRhodostemonodaphnekunthianawas made, maintaining the synonymy mentionedabove. Nectandramiranda was also transferred,Rohwer (1991) pointing out its similarityto specimensnow assigned to Paraia bracteata (Rohweret al., 1991). Finally,two transfers,the resultof new associationswith the genus, were made: these were Rhodostemonodaphneceliana and R. mirecolorata, two dioecious species from the Guayana highlands, originally described in Ocotea by Allen (Rohwer, 1986). Rohwer (1986) had seen only fruitingmaterialof Ocotea macrocalyx(Meissner)Mez, but on the basis of the descriptionby Mez (1889) he suggested thatit belonged to Rhodostemonodaphne.Later,most of the material belonging to this species was annotatedas "Rhodostemonodaphne macrocalyx(Meissner)comb. ined." by Rohwer. Meissner (1864) originally describedthis species in the dioecious, two-locellategenus Goeppertia(now a synonymof Endlicheria)with a question mark (he had seen only a fruiting specimen). Mez (1889) transferred it to Ocotea after having examined flowering specimens, describing their filamentousstamenswith long anthersand subextrorselower locelli. Henk van der Werff (1989b) transferredOcotea steyermarkianaAllen, anotherspecies from the Guayana highlands,to Rhodostemonodaphne.He also describedR. synandra,a species vegetativelyvery similar to R. kunthiana,but with very different flower morphology;later (van der Werff & Vicentini,2000) he described R. recurva. Joao Batista Baitello and Beulah Coe-Teixeira(1991) describedR. capixabensis, a very distinctivespecies from the coastal forests of the Brazilianstate of Espirito Santo. When I started the revision of Rhodostemonodaphnein 1991, it included 11 species, one suggested transfer,and one unpublishedcombination.I here reinstatetwo previouslydescribedspecies, R.praeclara (Sandwith) Madriinanand R. velutina (Mez) Madri- MORPHOLOGY fin; the formerhad been synonymizedwith Nectandra grandis by Kostermans (1936), the latter suggested as a synonym of R. laxa by Rohwer (1986). Five new species from the vicinity of Sail, French Guiana,had to be described prior to the completion of this revision (two more from NE South America were included;Madrifinn,1996), in orderto makethe names availablefor the publicationof S. Mori's Guide to the VascularPlants of CentralFrenchGuiana: Vol II (Madrifinn,2002). Sixteen species, the great majority of which are from newly availablematerial,are here described as new and four additionaltransfers are proposed. Andr6 S. Chanderbali,while working on a revision of Endlicheria(2004), noticed the fourlocellate condition of the staminodesof two species in that genus and transferredthem to Rhodostemonodaphne: R. debilis (Kosterm.)Chanderbali,and R. juruensis (A. C. Sm.) Chanderbali;these areincluded in this treatment. 7 the trunkhave been reportedfor some of these species. The distinctionbetween small trees and shrubsis obviously vague. In designatinga small individualas a tree or a shrubI am guided by the terminologyused by the collectors and the general appearanceof the herbariumsample. Lax, thin twigs are taken as suggestive of a shrubbyhabit. The smaller tree species (i.e., ca. 10 m tall) such as Rhodostemonodaphnedebilis, R. crenaticupula,R. frontinensis, R. laxa, and R. capixabensismay also be called shrubs.A few species appearto be consistently shrubby;these are R. avilensis, R. curicuriariensis,R. licanioides, R. miranda, R. parvifolia, and R. steyermarkiana.Among the shrubs, a very characteristicgrowth form is that of the scandentshrubsas illustratedby R. celiana and R. scandens. The scandent shrubs have thin, long, little-branchedaxes, and grow interwovenwith or reclining on the neighboringvegetation.These species grow on sandstone outcrops where the vegetation tends to be of low stature.Putting togetherinformation availableon habit and habitat,and guided by the MORPHOLOGY morphology of the twigs, it seems likely that R. deHABIT bilis, R. crenaticupula,R. curicuriariensis,R. licanThe informationon the habitof the species of Rho- ioides (in one label described as having drooping dostemonodaphnehas, for the most part,been taken branches),R. miranda,andR. tumucumaquensis, may from herbariumlabels. The patchiness and unrelia- also be scandent shrubs.These and the "true"scanbility of this informationwarnsagainstoverstatingthe dent species all tend to have undulate to bullate significance of these data. Notwithstanding,collec- leaves. The scandent, shrubbyhabit of Endlicheria tion label data can be very informativewhen inter- vinotinctaC. K. Allen is remarkablysimilarto thatof preted carefully with prior knowledge of the living Rhodostemonodaphneceliana, andboth species grow plants. on the sandstone formations of the Guayana highAll species of Rhodostemonodaphneare dioe- lands. Other similarities in indument and inflorescious. All fertile plants seen (live or dry) had flowers cences, and especially their similar glandular outwith only one set of functional sex organs. Pistillate growths on the stamens, call for a close relationship flowers sometimeshave well-developedstamenswith between these species (Rohwer et al., 1991). Neverlarge locelli, but I have neverseen pollen withinthem. theless, because of theirdifferencesin locelli number Staminateflowers sometimes have rudimentaryova- they are placed in different genera. Rhodostemonories, but these lack a stigma and an ovule. No indi- daphnesteyermarkiana,a thirdspecies fromthe Guacation of sex change is known. Three trees collected yana highlands,has similar leaf characteristicsto R. twice with roughly one-year intervals had pistillate celiana andR. scandens. In label notes it is invariably flowers and fruits both times (BAFOG361-M, 7425; described as a shrub,without mention of a scandent Kroll-Saldania134, 326; KurzB-9, B-28-14). habit. However,the branchingdynamics of this speThe majority of the species of Rhodostemono- cies differs somewhat from that of the two scandent daphne are medium to large trees 10-30 m tall and species in that the flushes produce lateral branches ca. 20 cm in diameterat the base. The maximumre- regularly,unlike the long unbranchedflushes of R. corded height is 45 m for an individualof R. kunthi- celiana and Endlicheriavinotincta. ana (although15-30 m is more common for thatspeEven when available,the informationon barkand cies; one tree I collected was a canopytree 30 m tall). wood charactersprovided by the collectors is very Seven otherspecies areknown to attain30 m or more; inconsistentin terms of the descriptionof the surface these include R. elephantopus,which has the maxi- sculpturing,outerandinnerbarkcolors andthickness, mum recordedtrunkdiameterfor the genus, at 80 cm. and wood odor and color. I have neverthelessadded Buttressesca. 70 cm tall and spanningca. 30 cm from it to the species descriptionswhen available.The co- 8 FLORA NEOTROPICA pious, clear mucilage exuding from the barkof Rho- and flushing dynamics that stand out from all others dostemonodaphnependuliflorais uniquein the Laur- thus far known in the genus. The growing axes do not aceae so far as I know. An observationcommonly show a marked resting period. Nevertheless, there made in field notes for R. kunthianais that its wood seems to be a certain periodicity in growth: zones characteristicallyoxidizes darkorange to black upon along the main axis where lateral branchesare produced are followed by zones with no lateralbranches exposure. or inflorescences. The branches thus tend to be arARCHITECTURE rangedin pseudo-whorls.These branchesare for the The architectureof Rhodostemonodaphneappears most partmore vigorous than the parentaxis, resultto conformto the models previouslyreportedfor other ing in a chandelier-likearrangement. The terminalbudsof Rhodostemonodaphne arealspecies of Lauraceae, i.e., those of Rauh, Attims, covered with The can be and Roux et Most ways cataphylls. cataphylls Massart, (Halle al., 1978). species in the family have monopodial trunks with spiral elongated and pointed, giving the bud a slender ap(generally 2/5) phyllotaxy, continuous or rhythmic pearance,or thickenedand blunt. The cataphyllsare growth, and mostly orthotropicbranches,these also mostly caducous,but sometimes, as in the case of R. with spiralphyllotaxy(opposite-leafedspecies are al- licanioides, R. miranda,andR. scandens, they persist ways present in small genera such as Anaueria, As- on the twigs throughat least two growing seasons. pidostemon, Caryodaphnopsis,Chlorocardium,and INDUMENT Dahlgrenodendron,and are common in CinnamoMost species in the genus have some type of inmum, but are otherwise rare in the Lauraceae).The branchescan be secondarilyplagiotropicas they be- dument,althoughnearlyglabrousor soon glabrescent come parallelto the groundwith age. Floweringdoes individualsare found in many species. In the two spenot influence the architectureof the plants. A report cies described as glabrous, Rhodostemonodaphne of distichous leaf arrangementin the branchesof an longipetiolata and R. pendulifiora,the buds at least unidentifiedspecies of Cryptocaryafrom New Cal- have some type of hair covering. The hairsin Rhodostemonodaphne,as in all Lauredonia (Halle et al., 1978, citing Veillon, 1976, unavailable to me) is very unusualand suspect (the same aceae, are invariably unicellular and simple. They condition reportedfor Cassythafiliformis L. is prob- generally have a conspicuous lumen, but solid hairs ably due to the extreme morphologyof this parasitic are also present. The indumentmay soon disappear, or persist through more than two growing seasons. genus). monoThe buds and expanding leaves are mostly covered the Lauraceae are consistently Although mowith their shoot is far from being appressed-ascendinghairs.The hairscan remain podial, morphology notonous. In Rhodostemonodaphne,as well as else- straight and appressed on the twigs and leaves, rewhere in the family, some species have a markedly sulting in a sericeous indument,as in species of the rhythmicgrowth (e.g., R. anomala), while in others R. grandis complex, or become erect, or be straight growth is more or less continuous with no apparent to crisped. Depending on the length and density of rest (e.g., R. avilensis), as evidenced by morphologi- the hairs, the indument of twigs and leaves can be cal indicators such as scale leaf scars, and/or short describedas puberulous,tomentose,pubescent,or veinternodalregions. A characteristicfeatureof R. ru- lutinous.Indumentterminologyfollows thatproposed fovirgata is the production of whole flushes, with by Hewson (1988). Hair coloring varies among the leaves, branches,andinflorescences,priorto the elon- species, and even within a species, from translucent to whitish, yellowish, brownish,or reddish. gation and lignificationof the main axis. The indument provides importantcharactersin Differencesin lateralbranchpositionin the flushes of rhythmicallygrowingplants,areimportantin char- distinguishing species or even groups of species in acterizing species, groups of species, or even genera Rhodostemonodaphne.In the R. grandis complex, all (cf. Endlicheria subgenus Ampelodaphneand Acti- species have a characteristicsericeousindument,connodaphne).In Rhodostemonodaphnethe majorityof sisting of long, straight, appressed, yellowish to the species have basitonic branching, where the brownishhairs, giving a silver to golden appearance. branchesof a new flush are producedfrom the axils This indumentcan neverthelessbe caducous and noof cataphyllsor basal foliage leaves. The resultis of- ticeable only on young plant parts. In other cases, ten pseudo-verticillatebranching,as observed in R. however,indumenttype can be variablewithin a spelaxa, R. leptoclada, R. scandens, and R. penduliflora. cies, or even within an individual. Rhodostemonodaphnekunthianais remarkablein Rhodostemonodaphnekunthianahas architecture MORPHOLOGY 9 some specimensof R. licanioides. Apices arerounded to acute, and are most often mucronateto acuminate. The leaf marginis mostly flat along its length, but in some species it is recurvedat the base. Rhodostemonodaphne revolutifolia is distinctive in that the leaf margins of the mature leaves are strongly revolute along their entire length; R. recurva generally has leaves with recurvedmarginnearthe base. A few collections of R. macrocalyxfrom the southernextreme of its range have unusuallylobed leaves. The venationis mostly eucamptodromous,the secondary veins looping near the marginbut fading before connecting with the next secondary vein. Towards the leaf apex the secondary veins tend to be more strongly connected than elsewhere. Only Rhodostemonodaphnescandens and R. licanioides are strictly brochidodromous, with all the secondary veins connected near the margin. In some species thereis a slight tendencytowardsan acrodromousvenation, where the basalmost secondary veins are strongerthanthe rest. In a numberof species some of LEAVES the secondaryveins bifurcatein midlamina.In a few The leaves of Rhodostemonodaphneare spirally species the tertiary veins are distinctly scalariform, R. arrangedin a 2/5 phyllotacticpattern.Suboppositeor runningperpendicularto the secondaryveins (e.g., are randomin but most kunthiana), they in species elsewhere as found pseudo-verticillate leaves, Lauraceae(e.g., Endlicheriasubgen.Ampelodaphne, reticulate. Actinodaphne,etc.), are not normally observed, alINFLORESCENCES though some specimens of R. scandens tend to have The inflorescencesare mostly located towardsthe pseudo-verticillateleaves at the end of the flushes. base of the flushes, but mesotonic or acrotonicinfloas the as thick The petioles can be robust (nearly to adaxterete or and are borne rescences, as well as those found along the whole on) slender, twig they Some canaliculate. to flattened species length of the flush, also occur. In the majorityof the nearly ially have what I refer to as a "pulvinate"petiole. This species the inflorescencesare erect. Pendulousinfloconsists of a short, thickened, terete petiole and is rescences (e.g., Rhodostemonodaphnependuliflora, mir- as observed in the living specimen) are also present foundcharacteristicallyin Rhodostemonodaphne found in anda. Pulvinate petioles are often species in the genus, althoughthese may be under-reported. with a shrubbyhabit and in plants growing on white As a rule the inflorescenceshave a determinateaxis and cymose second order branches.The numberof sands or sandstones. Leaf blade consistency can vary greatly within second-orderbranches and the degree of branching species and even individuals.A tendency to have an varies both between and within species. The stamiundulate or distinctly bullate lamina is common in nate inflorescences usually have higher orders of some species, but in all species wherea bullatelamina branching than the pistillate ones. Second-order has been observed,flat laminae are also present.The branchesaremostly scatteredalong the main axis, but lamina is usually elliptic, and it varies in size from in a few species they are clustered apically (e.g., R. less than 10 cm in length, as constantlyfound in Rho- morii, R sordida). Inflorescencesvary from "spicoid" dostemonodaphneavilensis or R. parvifolia, to 40 cm thyrses, with no more than three branchorders (i.e., long as in R. penduliflora.Abnormallylarge leaves, the main axis plus constrictedlateral simple cymes, as often found in sterile collections, are most likely e.g., R. miranda),to lax, thyrsoidinflorescenceswith takenfrom stumpsproutsor saplings.The base of the up to eight branchorders(e.g., R. macrocalyx).Bracts lamina can be distinctlyroundedto acute or cuneate. and bracteolesare mostly caducous,but in some speA conspicuouslycordatebase, rarein the family as a cies (e.g., R. revolutifolia)they are persistent.In genwhole, is found in the only known collection of R. eral, the inflorescences have the same type of indumiranda; minutely cordate bases are also found in ment as the twigs. terms of the variationof indumenttypes present.As is discussed in detail in the species discussion, indument of R. kunthianacan vary from a covering of short and crisped hairs to long and curvedto straight hairs. Although the former are generally associated with plantscollected in lowlandforests, and the latter with collections from montaneforests, this patternof variation is not clear cut (differences in indument types observed within an individualmay also correspond to different growth stages of the plant). Hair color here also varies from light yellow to reddish brown. Rhodostemonodaphnedioica is unusualin thatthe indumenton the undersideof the leaves consists of two types of hairs,erect and appressed.Anothercomparablecase, where the indumentis mixed, is Nectandrareticulataand some of its allies (J. G. Rohwer, pers. comm.). On the twigs, however, the indument dioica consists only of long, of Rhodostemonodaphne straightto curved, erect hairs. FLORANEOTROPICA 10 FLOWERS On the basis of their externalmorphology,staminate and pistillate flowers are indistinguishable.The flowers usually have a conspicuouspedicel, although nearly sessile flowers are found in R. scandens andR. miranda. The pedicel is usually of equal width throughout,but in a numberof species it is trumpetshapedandgraduallymergeswith the receptacle(e.g., R. laxa). The receptacles themselves are mostly distinct from the pedicel and obconical, although in a few species they are almost nonexistent,while in others they are well-developed, globose, and often constrictedat the place of tepal inception.The indument of the pedicels and receptacle as well as that of the abaxial surface of the tepals is mostly similar to that of the twigs, althoughit is generally shorter. The generalarrangementof flowerpartsis typical of the LauraceaetribePerseeae(see Fig. 1). Thereare two alternatingwhorls of tepals, nine stamensin three alternatingwhorls (labeledI, II, and III, from without inwards;in a few species a fourth,staminodialwhorl [IV] is present),and a unicarpellatepistil. The tepals of the two whorls are equal to subequal in size. At anthesis they are mostly broad, and spreadingor recurved;small, distinctlyerect tepals arecharacteristic of R. morii alone. A number of species have lightcolored tepals, rangingfrom green to creamto white, but yellow, salmon, or distinctly red tepals are not uncommon. The adaxial surface of the tepals is glabrous, hairy, or (in R. penduliflora)papillose. Variationin stamen morphologyis extensive and comparableto that observedin the vegetativeorgans. As with most Lauraceae,the stamens of Rhodostemonodaphneof whorls I and II are similar,and very differentfrom those of whorl III. In most species the anthersof the outertwo whorls arethickenedandsessile, with four locelli arrangedin an archtowardsthe apex. Filiform stamens are characteristicof R. macrocalyx; stipitate, laminar anthers are found in R. capixabensis;and the straplike(loriform)stamensof R. synandraare uniqueto that species. Rhodostemonodaphne recurva, R. cyclops, R. morii, and R. penduliflora vary in locelli number,while in R. revolutifolia, the only known staminateindividual, has in the outerwhorl two-locellateanthers,andthose of the only known pistillate individual have four vestigial locelli. In the four-locellatespecies, the mediallocelli are always introrse,while the two exmedial ones can be introrse,or in R. kunthiana,approachinga latrorse position. Most species have glabrous anthers, but sometimes with a few hairs at the base; those of R. kunthianaand allies are hairy, while R. penduliflora is unique in having distinctly papillose anthers.The stamensof whorl III are mostly columnar,with a pair of glands at the base, and have four locelli; in most species the lower locelli are extrorse,while the upper ones are either extrorse or latrorse. Only in R. penduliflorais there a reductionof locelli numberto two and in some cases one; furthermore,these locelli are introrse(note however,thatthe only known specimen of R. penduliflorais pistillate). The size, shape, and degree of fusion of the glands at the base of the stamens of whorl III varies greatly.Two species, R. peneia and R. praeclara, consistently lack glands on whorl III, while in the flowers of R. longipetiolata with doubled whorls (see discussion underR. longipetiolata) they were also absent. Two species, R. celiana and R. penduliflora,have paired glands at the base of the stamensof whorlsI andII;these, however, are not distinct and globularas are those commonly found at the base of the anthersof whorl III. Glands in whorls I and II are otherwisevery rarein the Lauraceae and their presence in these species needs to be confirmed.Staminodiaof whorl IV are generallyabsent, except in three species, R. celiana, R. elephantopus, and R. miranda, where they are sometimes present. In pistillate flowers, the locelli of the staminodia of all whorls vary from fully developedto completely lacking. In all cases where the locelli were opened there was no pollen. The pistil in staminateflowersis sometimes fully developed but lacks an ovule; more often it is reducedto a filiformappendageor is completely absent.When present,it is often obscuredby the partiallyfused stamens of whorl III. In pistillate flowers the ovary is ovoid or globose, and is either enclosed by the receptacle or is borne on top of it surroundedonly by the staminodesof whorl III. POLLEN In a contributionto the pollen morphologyof neotropicalLauraceae(Raj & van der Werff, 1988), two species of Rhodostemonodaphnewere studied (R. grandis and R. kunthiana).The pollen grainsof these species are apolar,spheroidal,21-24 gm diam., and inaperturate.The exine is 0.5-1 gm thick and is covered with monomorphicvestigial spinules, scattered 1-2 aimapart.The spinules in both species are plain, lacking the usual twisted strands observed in other closely related genera (e.g., Ocotea, Nectandra, and Pleurothyrium). FRUITS The pedicels in fruit vary from evenly thickened to graduallyenlarged,forming a trumpet-shapedcujupule, with the exception of Rhodostemonodaphne ruensis with a spreadingdisklike cupule and R. antioquiensis with a reflexed cupule. When dry the DISTRIBUTION ?-F0 with paired glands FIG. . ower diagramof a typical Lauraceaeof the tribe Perseeae. Tepal Tcpal Ovary and Ovule Ovary and Ovule FIG. 1. Flower diagramof a typical Lauraceaeof the tribe Perseeae. cupules are commonly coriaceous and have a smooth surface. Normally the margin is undulateand the tepals persist in fruit as cupular appendages.The cupules of species in the R. grandis-complexand those of R. kunthianadiffer from the rest in that they are distinctly pedicellate, woody when dry, and warty or lenticellate (in R.kunthiana and R. synandra), and have a straightmargin.The single-seeded berries are nondescriptand elliptic to ovoid. The great majority of the field notes reportthe ripe fruits as having a red cupule and blackish berry, as is common for most Lauraceae.The fruits of the species of Rhodostemonodaphnefor which these are known are shown in Figures 2 and 3. DISTRIBUTION Ourknowledge of neotropicalfloristics is farfrom complete. Althoughplantsfrom better-collectedareas may be well-representedin herbaria,the numberof FLORA NEOTROPICA 12 F H "H G 2 cm FIG. 2. Fruits of Rhodostemonodaphne(drawnto scale from dried specimens). A. R. laxa (Vargaset al. 1053). B. R. cf. anomala (Hage et al. 2357). C. R. macrocalyx(Pereira 714). D. R. licanioides (Vdsquezet al. 644). E. R. miranda (Fanshawe 2863). F. R. scandens (Daly et al. 3837). G. R. parvifolia (Rodrigues5717). H. R. crenaticupula(Vdsquez& Jaramillo9564). I. R. negrensis (Nascimentoet al. [INPA/WWF]1201.288). J. R. celiana (Maguire & Maguire35340). K. R. steyermarkiana(Steyermarket al. 109172). 13 DISTRIBUTION B E /C F G cm FIG. 3. Fruits of Rhodostemonodaphne(drawn to scale from dried specimens). A. R. antioquiensis (Cogollo & Estrada290). B. R. capixabensis(Pereiraet al. 2883). C. R. rufovirgata(MutchnickD.966). D. R. morii (Mori et al. 15334). E. R. kunthiana(Kayap279). F. R. synandra(Tillett672-83). G. R. grandis (Schulz8264). H. R. saulensis (Granville5077). I. R. praeclara (Marcano-Berti254), berry approximate.J. R. peneia (KurzB-9), berry approximate. 14 FLORANEOTROPICA recent floras and monographsof neotropical plants remains relatively low compared to those for other regions of the world. The neotropicalLauraceae(if not the tropicalLauraceaeas a whole) are a good example of a little-knowngroup.This is in partbecause they are mostly tall trees with small flowers, and so are often undetectedby collectors. Furthermore,even if trees are located by means of their aromaticwood, they are frequentlysterile because floweringperiods are short. Sterile collections are mostly indeterminable and are often set aside in piles in herbaria,andare rarelyincorporatedinto monographicor even floristic works. Thus, not only is distributionalinformation from herbariumspecimens fragmentary,but also a greatnumberof species tend to be known from a limited numberof individuals,making inferences about their distributiondifficult. Twenty-threeof the 41 species here treated are knownfrom fewer than 10 collections (TableI). These generally come from a single locality, or if from differentlocalities they are often representedby a single individualin each. The fact thatmany species of Rhodostemonodaphneare known from a single locality may be reason to believe that there is a high degree of localized endemismin the genus-alteratively, it may be the productof poor sampling.Unfortunately, in most cases it is virtuallyimpossible to distinguish between the two possibilities. The interpretationof gaps in the distributionof a species or pairsof species is similarly hampered.Gaps may representdisjunct distributions,or be an artifactof the lack of collections in the interveningareas. Lastly, relativelyhigh species diversity in one area may be the productof high collecting densities ratherthanbiodiversity"hot spots,"as has been shown for the Amazonianflorain general (Nelson et al., 1990). The biogeographicregions I have adoptedare for the most part congruent with previously proposed ones (Schnell, 1987, and references therein). For Amazoniain particularI referto the subdivisionsproposed by Prance (1976), with a few modifications. Unlike Prance,I treatthe northern"AtlanticCoastal" and northern"Jari-Trombetas" subdivisionsof Amazonia together, as the "northeasternSouth America region", comprising northeastern Venezuela, the Guianas, and northernAmapa, Brazil. The region is separatedfrom the Amazonian basin proper by the drainagedivide of the Tumucumaquerangeandother minor mountain ranges separatingBrazil from the Guianas.This and the presence of various species of TABLE I Habitatand distributionof Rhodostemonodaphnespecies No. of known collections Species Habitatand distribution R. anomala Atlantic rain forest, SE Brazil R. antioquiensis Lower montaneforest, C Cord., Colombia 2 Localized R. avilensis Cloud forest, "El Avila,"Venezuela 2 Localized R. capixabensis Restinga,EspirituSanto, SE Brazil 7 Limited R. celiana Tepui,Yutaje/Coro-Coromassif, Venezuela 9 Localized R. crenaticupula Non-inundatedrain forest, E & C Amazonia 32 R. curicuriariensis Sandstone?Curicuriarimtns., Brazil 2 Localized R. cyclops Upper montaneforest, Andes, Ecuador 4 Localized R. debilis Non-inundatedrain forest, NW Peru 1 Localized R. dioica Non-inundated?rain forest, SE Amazonia 3 Limited R. elephantopus Non-inundatedrain forest, FrenchGuiana 3 Localized R. frontinensis Cloud forest, Antioquia,W Cord., Colombia 1 Localized R. grandis Non-inundatedrain forest, NE South America 2 (3?) 45 Coverage1 Limited Limited Limited 15 DISTRIBUTION No. of known collections Species Habitatand distribution R. juruensis Premontaneforests, E slopes of N Andes R. kunthiana Variousforest types, Meso and South America R. laxa Upper montaneforest, C Cord., Colombia 11 R. leptoclada Non-inundatedrain forest, FrenchGuiana 2 (1 indiv.) Localized R. licanioides Non-inundatedwhite sand forest, NW Peru 13 Localized R. longiflora SW Amazonia?,Bolivia 1 Localized R. longipetiolata Non-inundatedrain forest, W Ecuador 1 Localized R. macrocalyx Atlantic rain forest, SE Brazil R. miranda Non-inundated?rain forest, W Guyana 1 Localized R. mirecolorata Tepui, Karowtipumtn., W Guyana 1 Localized R. morii Non-inundatedrain forest, E Guianas R. napoensis Rain forest, E Amazonia,Ecuador R. negrensis Non-inundatedwhite sand forest, NW Brazil R. ovatifolia Upper montaneforest, Andes, Venezuela 1 Localized R. parvifolia Non-inundatedrain forest, C Amazonia,Brazil 4 Localized R. penduliflora Non-inundatedrain forest, W Colombia 1 Localized R. peneia Non-inundatedrain forest, C Amazonia,Brazil 9 Limited R. praeclara s.str. Non-inundatedrain forest, N South America 14 Limited R. praeclara s.l. Non-inundatedrain forest, N South America 32 Widespread R. recurva Non-inundatedrain forest, C Amazonia,Brazil 7 Localized R. revolutifolia Non-inundatedrain forest, FrenchGuiana 2 Localized R. rufovirgata Non-inundatedrain forest, Guianas 12 Limited R. sailensis Non-inundatedrain forest, FrenchGuiana 12 Localized R. scandens Non-inundatedrain forest, Guianasand Amapa 10 Limited R. sordida Non-inundatedrain forest, C W Amazonia 2 Limited R. steyermarkiana Tepui, E VenezuelanGuayanahighlands 9 Limited R. synandra Non-inundatedrain forest, E Andeanfoothills 5 Limited R. tumucumaquensis Non-inundatedrain forest, E Guayanahighlands 4 Localized R. velutina Upper montaneforest, E Cordillera,Colombia 1 Localized Coverage' 5 Widespread 147 Widespread 37 Limited Limited 11 Limited 2 Limited 13 Limited limited= occupying localized= narrowendemic,i.e.,confinedto oneortwoadjacent degreesquares; categories: 'Coverage two or moredegreesquares,non adjacentbut withinone biogeographical region;widespread= foundin two or more regions biogeographical 16 Rhodostemonodaphnerestricted to this region warrant its recognition as a separatebiogeographicalregion, for the purpose of this discussion. The eastern Andeanfoothills, althoughpartof the Amazon basin, have often been treatedas a separateentity because of their peculiarbiota (Hueck, 1966; Cabrera& Willink, 1973; Schnell, 1987). Again, I treat this region separatelyfrom Amazonia, for there are various species of Rhodostemonodaphnethat seem to be characteristicof this region. Rhodostemonodaphneis restricted to the New Worldtropics.All but one species have relativelylimited distributionranges within South America. Only R. kunthianais widespreadin South Americaand extends northof the isthmusof Panamainto CostaRica. The majorityof the species have distributionsthatare highly congruent with some of the major biogeographicregions of South America (TableII; see also species distributionmaps). The largest concentrationof species is found in the lowlands and highlands of northeasternSouth America, comprisingeasternVenezuela,the Guianas, and northeasternBrazil, as well as in central Amazonia. A total of 13 species are found in each of these regions, 12 of which (Rhodostemonodaphnekunthiana excluded) are restrictedto each region. The surroundingsof Saul in centralFrenchGuianaappearto be an area with particularlyhigh species diversity; seven of the 13 species found in the region have been collected in this relatively small area, and four have not been found elsewhere. Ten of the 13 are represented in French Guiana as a whole. Nevertheless, these high figures may result from recent intensive collecting in the countryby The New YorkBotanical Gardenand the CentreORSTOM. Within northeasternSouth American the species are distributedin a variety of habitats.Only five species are found in the northerncoastal plains and none is endemic there. Most species are found in the interior highlands. Among these, two species, Rhodostemonodaphne scandens and R. rufovirgata, have similar disjunct distributions,with collections from both Guyanaand FrenchGuiana,but not reportedfor Suriname (this may be due to lower collecting densities in the interiorof Suriname).A few species appear to be restricted to the higher elevations of the interior and the Tumuc-Humac (French spelling) range on the border with Brazil. Rhodostemonodaphne tumucumaquensis,found on the Amazonian side of the range and on Mount St. Marcell, an isolated mountainin southernFrenchGuiana,is one of these. Rhodostemonodaphnemiranda and R. mire- FLORA NEOTROPICA colorata, both restricted to the interior of Guyana, appearto be associatedwith sandstoneformationsof the Guayanahighlands. Two species are restrictedto the VenezuelanGuayanahighlands.These areRhodostemonodaphne steyermarkiana,from the Yutaje/Coro-Coromassif in the Gran Sabana, and R. celiana, from the Auyan tepuf in southeastVenezuela.Although somewhatdifferent architecturally(see Morphology,above), both species are low shrubsas is typical of the vegetation of the summitsof these tabletopmountains. The region with the second highest species diversity is the Amazon basin proper.The variable Rhodostemonodaphnepraeclara s.l. is found throughout the region, including the "Xingu-Madeira"subdivision, where no otherspecies of Rhodostemonodaphne is found. Another species with a broad distribution, but less so and restrictedto the westernpartof Amazonia, is R. crenaticupula.The remainingAmazonian species have been collected either from a single locality or from largerareas that are presumablycharacterized by a particular habitat. Rhodostemonodaphne negrensis is found along the Rio Negro and in the vicinity of Manaus,andis apparentlyassociated with white sand forests of "blackwater"rivers. Another species commonly collected in white sand forests is R. licanioides, knownfrom the vicinity of Iquitos. Rhodostemonodaphnecuricuriariensisis known only from two collections from the Curicuriaririver, a tributaryof the Rio Negro. Althoughpreciselocality data for this species is unavailable, it seems to be associatedwith the sandstonemountainof Curicuriari from that locality. Rhodostemonodaphnepeneia and R. recurva are known primarilyfrom the vicinity of Manaus. Rhodostemonodaphnedioica has been collected in a small area in southwest Amazonia at the Bolivia-Brazil-Peru border. Rhodostemonodaphne kunthianais spreadacross the Amazon basin, but in localized patches within the region, such as the "Southwest,""Solim6es-Amazonas,"and "Manaus" subdivisions,and in the vicinity of the Amazon river delta. The widespreadRhodostemonodaphnekunthiana is best represented,both in terms of numberof collections and continuityof distribution,in what I here call the easternAndean foothills. There too is where the greatestdiversityof forms of this variablespecies are found. Rhodostemonodaphnesynandra, a close relative of R. kunthiana,is restrictedto this region, being found in four localities from Colombiasouthto Peru,in areaswhereR. kunthianais also present.Rhodostemonodaphnenapoensis and R. longipetiolata, DISTRIBUTION 17 TABLE II Species of Rhodostemonodaphneby biogeographicregion NortheasternSouth America R. elephantopus,R. grandis, R. kunthiana,R. leptoclada, R. miranda,R. mirecolorata,R. morii, R. praeclara s.str.,R. revolutifolia,R. rufovirgata,R. sailensis, R. scandens, R. tumucumaquensis VenezuelanGuayanahighlands R. celiana, R. steyermarkiana Amazonia R. crenaticupula,R. curicuriariensis,R. debilis, R. dioica, R. kunthiana,R. licanioides, R. longiflora,R. negrensis,R. parvifolia, R. peneia, R. praeclara s.l., R. recurva,R. sordida EasternAndean foothills R. juruensis, R. longipetiolata,R. napoensis, R. synandra North Andeanhighlands R. antioquiensis,R. avilensis, R. cyclops, R.frontinensis,R. kunthiana,R. laxa, R. ovatifolia, R. velutina North Pacific lowlands and Mesoamerica R. kunthiana,R. penduliflora Atlantic rain forest R. anomala, R. capixabensis,R. macrocalyx also presumedrelativesof R. kunthiana,arerestricted to Napo provinceof Ecuador,also in the Andeanfoothills. Rhodostemonodaphnelongifloraand the southernmostrepresentativesof R. kunthianaare the only representativesof the genus from the Bolivian lowlands east of the Andes. The strictly Andean species of Rhodostemonodaphne (R. kunthiana excluded) are found in the northernAndes in Ecuador,Colombia,andVenezuela, and are associated with cloud forests at elevations greaterthan 1500 m. The southernmostmemberis R. cyclops, from Ecuador.In Colombia,wherethe Andes trifurcateinto threecordillerasseparatedby deep valleys, each cordillerahas a separate species of Rhodostemonodaphne;these are R. frontinensis,R. laxa, andR. velutina,from the CordillerasOccidental,Central, and Oriental,respectively.In the VenezuelanAndes R. ovatifolia is present.Anothercloud forest species, but in an area isolated from the core of the Andes, is R. avilensis, known from mountainsnear Caracasin Venezuela. Rhodostemonodaphneantioquiensisis includedin this region as an exception,for it is more of a lowland species found at ca. 500 m on the foothills of the CordilleraCentralof Colombia. A populationof R. kunthianafrom the province of Oxapampain Pasco, Peru, characterizedby a long and dense indument, reaches elevations above 2000 m. Only two species are found west of the Andes. One of them is the ubiquitousRhodostemonodaphne kunthiana.Concordantwith a common biogeographical pattern,this species is found east of the Andes only to the north of the Huancabambadeflection in northernPeru. The other species is R. pendulifora, the only species of the genus from the Pacific lowlands of the Choc6 region in Colombia. Rhodstemonodaphnekunthianais also, as mentionedabove, the only species of the genus with representativesin Mesoamerica. Finally,threespecies arefound in the Atlanticrain forest of southeasternBrazil.The morewidespreadof the threeis Rhodostemonodaphnemacrocalyx,which has a rangethatextendsfromthe southerntip of Bahia to just south of the tropic of Capricornin Parana. Rhodostemonodaphneanomala is known from two collections fromthe mountainsof Petr6polis,nearRio de Janeiro, and possibly from a fruiting collection from southernBahia. Rhodostemonodaphnecapixabensis is restrictedto the coastal sand forests ("restingas")of Espirito Santo. A list of species by country, indicating each country's endemics, is provided (TableIII). FLORA NEOTROPICA 18 TABLE III Species of Rhodostemonodaphneby country' Costa Rica R. kunthiana Panama R. kunthiana Colombia R. antioquiensis, R. frontinensis, R. kunthiana,R. laxa, R. penduliflora, R. synandra, R. velutina Venezuela R. avilensis, R. celiana, R. kunthiana,R. ovatifolia, R. praeclara s.l., R. steyermarkiana Guyana R. kunthiana,R. miranda, R. mirecolorata, R. praeclara s.str.,R. rufovirgata,R. scandens Suriname R. grandis, R. kunthiana,R. morii, R. rufovirgata FrenchGuiana R. elephantopus, R. grandis, R. kunthiana,R. leptoclada, R. morii, R. revolutifolia, R. rufovirgata,R. saiilensis, R. scandens, R. tumucumaquensis Ecuador R. cyclops, R. juruensis, R. kunthiana,R. longipetiolata, R. napoensis, R. praeclara s.l. Peru R. crenaticupula,R. debilis, R. juruensis, R. kunthiana,R. licanioides, R. praeclara s.l., R. sordida, R. synandra Brazil R. anomala, R. capixabensis, R. crenaticupula,R. curicuriariensis, R. dioica, R. grandis, R. kunthiana,R. macrocalyx, R. negrensis, R. parvifolia, R. peneia, R. praeclara s.l., R. recurva, R. scandens, R. sordida, R. tumucumaquensis Bolivia R. dioica, R. juruensis, R. kunthiana,R. longiflora 'Namesin boldfacetypeindicatecountryendemics vided into various unrelatedclades, most notablythe Old Worldspecies of this genus falling outsidea clade Efforts towardour understandingof phylogenetic defined by the presenceof a 16-bprepeatin the trLrelationshipswithin the Lauraceaehave recentlybentrnF (Chanderbaliet al., 2001). Withinthis last region efited from the analysis of molecular sequence data is clade a group containing the dioemonophyletic Both et al. (2001). by Rohwer(2000) andChanderbali of cious Ocotea plus the dioecious generaEnspecies these studies have confirmedthe recognition of natand dlicheria Rhodostemonodaphne. ural supragenericgroups such as the Cryptocaryeae Among the dioecious species of the Ocotea com(van der Werff& Richter,1996), includingthe CrypEndlicheria and Rhodostemonodaphnehave tocarya and Beilschmiedia groups of Rohwer plex, been traditionally separatedon the basis of the twowood of (1993a), previously recognized on the basis of the formerand the four-locellate locellate anthers moresult of these A novel anatomy(Richter,1981). lecularphylogenies is the inclusion of the Laureae,a ones of the latter. However, generic delimitation within the Lauraceaerelying on locelli numberhas previously well-established group characterizedby been questioned (Rohwer et al., 1991). The four loinvolucrateinflorescences, within the Perseeae, with celli in Rhodostemonodaphne,arrangedin an archtoPerseeae-Laureae this thyrsoid inflorescences. In clade is a well-supportedgroup,the Ocotea complex, wardsthe apex of the anther,has been used to distincontainingmost of the generapreviouslyincludedin guish it from the dioecious species of Ocotea where the Ocotea and Aniba subgroupsof Rohwer (1993a). the locelli are arrangedin two superposedpairs, as The polyphyleticnatureof various generawithin this well as to suggest a close relationshipwith the hercomplex (e.g., Nectandraand Ocotea) is highlighted. maphroditicNectandra,where the locelli are also arOcotea, a large genus awaitingformalrevision, is di- rangedin an arch. SYSTEMATICPOSITION SPECIESGROUPS The molecular analyses clearly link species of Rhodostemonodaphnewith Endlicheria in a monophyletic clade (Chanderbaliet al., 2001), leading to the recognition of the "Endlicheria-Rhodostemonodaphne alliance" (Chanderbali, 2004). The most probable evolutionary scenario within this group is one where Rhodostemonodaphneis paraphyletic,defined by two plesiomorphic characters(dioecy and four-locellateanthers)from which parallelreductions in locelli numberhave produceda polyphyletic twolocellate Endlicheria. Transferof the Rhodostemonodaphne-like species of Endlicheriato Rhodostemonodaphneor vice versa will be imminentonce the phylogeneticrelationships among the species are more clearly established.Until then it is preferable,for the sake of identification,to keep the two genera separate. SPECIES GROUPS Below I presenta tentativegroupingof the 41 speThese groupsarebased cies of Rhodostemonodaphne. on overall morphologicalappearance,and are given only as an attemptto summarizesalient morphological similarities observed among the species of the genus. The groups have been formedby a process of "chaining,"whereby a few species are taken as representatives of the "core" of a group (the group's namesake and closely related species), with the remaining species in the group being linked to it throughone or more of its membersand by different characters. This method produces heterogeneous groups, not defined by single characters,and where species placed far apartin the linear sequence may have little in common but are linked by intermediate species. Thus, no formal taxonomic category is attributed to them, nor do they necessarily represent monophyletic groups. The groups and their species are listed in TableIV. The first two groups share common fruit morphology. In both groupsthe cupules have an undulate margin or are distinctly lobed, as the result of the tepals being persistentin fruit. Note that of the total 25 species includedin these two groups, the fruits of only 14 of them are thus far known (see Table IV); however, the species are individuallylinked to other species by differentcharacters. LAXA THE RHODOSTEMONODAPHNE GROUP The Rhodostemonodaphnelaxa group consists of plants that are generally medium-sizedtrees (R. avilensis and R. capixabensis are small trees to shrubs). 19 They have, for the most part, an indumentthat consists of long, erect yellowish hairs (R. avilensis is nearly sericeous, with white appressedhairs, and R. juruensis is a low-elevationrepresentativeof this species groupbearinga sparseindumentexpectedof species in warmerclimates).The indumentis mostly persistent throughout, but even the underside of the leaves (R. longiflora and R. capixabensis) or the young twigs (R. avilensis) can be soon glabrescent. In the majorityof the species, some of the secondary veins are distinctlyforked.When dry, the color of the leaves is usually greenish yellow, but darker-drying leaves are characteristicof R. avilensis, R. juruensis, and R. cyclops. The inflorescences are thyrses with few branchorders,located along the whole length of the flush.In both R. avilensis andR. capixabensis,the inflorescencebranchesare long, giving the whole inflorescence a more lax appearance.There is considerablevariationin the flowersof the species included in this group. Those of R. laxa and R. velutina are the tepals arethick andadaxiallyglatrumpet-shaped; brous, and at anthesisthe tepals remainerect.Both R. cyclops and R. frontinensis have very similar flower structurethat is somewhat different from the other species in the group. Their flowers are also trumpetshaped,althoughnot as markedlyso as in the preceding two species, and at anthesisthe pointedtepals are spreading,forming a distinctivesix-pointedstar.The adaxial surface of the tepals is partly covered with hairs. The only species with conspicuously reflexed tepals are R. anomala, R. dioica, and R. macrocalyx; here the tepals are densely covered with yellowish hairs on both surfaces.Rhodostemonodaphneavilensis and R. capixabensishave a similarfloralstructure which is intermediatebetween that of R. laxa and R. frontinensis,with tepals that are erect to spreadingat anthesis and that have some hairs on the adaxialsurface. In androecialmorphologythere is also great variation within the group. The stamensof whorls I and II have sessile and trapezoidanthersin the firstthree species. Rhodostemonodaphnefrontinensis, R. juruensis,andR. cyclops have columnarstamens(staminodes of pistillate flowers for R. juruensis), with an inconspicuousfilament, and there is some indumentum; both features link them to the R. kunthiana group. The stamens of R. anomala and R. dioica are thick and completely glabrous,except for a few hairs near the transition with the receptacle at the base. Rhodostemonodaphnemacrocalyx is unique among all species in the genus with its long, filiformstamens, and inflexed anthers with lateral lower locelli. The laminarstipitatestamensof R. avilensis andR. capix- 20 FLORA NEOTROPICA Table IV (informal)' Species groupsin Rhodostemonodaphne Rhodostemonodaphnelaxa-group R. laxa * R. mirecolorata R. rufovirgata* R. velutna R. celiana * R. ovatifolia R. steyermarkiana* R. longiflora. l R. leptoclada *R.juruernsis R. R. frontinensis parvifolia R. antioquiensis* R. R.J cyclops ic~yclops R. dioica Rhodostemonodaphnekunthiana-group R. kunthiana kt * R. R. anomala * synandra* R. R. macrocalyx*. I'~~~ * R. capixabensis B. capixabensis *~R. R. avilensis /?~R. napoensis longipetiolata R. morii * Rhodostemonodaphnescandens-group R. scandens * R. licanioides * R. elephantopus Rhodostemonodaphne grandis-group R. miranda* R. grandis * R. revolutifolia R. crenaticupula* R. debilis * . . . -. R. curicuriariensis R. saiilensis * R. recurva * . recurve R. tumucumaquensis * R. sordida *- ^^ *Species R..n, negrensis Madrifian*Species * ?R.praeclara praeclara * R.. peneia peneia * incerta sedis icersedis Rhodostemonodaphne penduliflora 'verticallines connect species with "stronger"linkages (thicknessof line indicativeof "strength"of linkage). An asterix(*) indicatesthose species with knownfruits SPECIESGROUPS abensis are intermediatebetween those of R. macrocalyx and the other species. The few fruitsknown all have long, thickenedpedicels graduallymerging with the cupule. The tepals persist on the cupule margin, either as deep lobes which are often reflexed(R. laxa andR. macrocalyx), or as small processes (cf. R. anomala); the cupule of R. capixabensis has an undulatemargin.The cupule of R.juruensis is spreading,forminga disk unlikeany other known species in the group. Althoughthe data availableare still insufficientto hypothesize the relationshipsamong the species in this group, there are groups of species where sisterspecies relationshipsare highly likely. Rhodostemonodaphnelaxa and R. velutina,andR. ovatifolia are in many respects very similarand withoutdoubtclosely related-Rohwer (1986) even suggested putting R. laxa and R.velutinaunder one binomial; I prefer to maintainthem separateon the basis of differencesin the densityof indument,leaf anatomy,andgeography. The second pair of closely relatedspecies is Rhodostemonodaphnecyclops and R.frontinensis.These species have similarleaves, inflorescences,and flowers (including androecial morphology). Rhodostemonodaphnefrontinensis has some similarities with R. laxa, and the two are found in the northernAndes, thus forminga link betweenthe two species pairs.The androecialmorphology described above points to a relationshipwith the R. kunthianagroup. However, the fruits of R. juruensis with their disklike cupules are unlike those found in the R. kunthianagroup.Anotherspecies collected in the same locality as R.frontinensis, and with very similar gross morphological features,is Endlicheriacolombiana (Meissner)Mez. The third example of probable sister-species relationshipsin this group is that of Rhodostemonodaphne anomala and R. dioica. These two species have very similar flowers but differ in size. Their wide, thickened,glabrousanthersare almost identical, and the indumenton the tepals is very similar.The disjunct distributionpatternof this species pair-one is found in western Amazonia, and the other in the Atlantic rainforest of Brazil-is commonin manyplant groups. Rhodostemonodaphnelongiflora, Bolivia, morphologicallyintermediatebetween R. dioica and R. laxa, links the southwesternAmazonianspecies to the Andean ones. The othertwo species from the Atlanticrainforest are somewhat isolated from the core of the R. laxa group, characterizedby the three species pairs described above. Rhodostemonodaphnemacrocalyx is vegetatively very similarto R. dioica or R. anomala, but its filiform stamenshave no parallelin the genus. 21 Rhodostemonodaphnecapixabensis, with its spathulate stamens, may be an intermediatecondition towardsfiliformstamens.Rhodostemonodaphne avilensis is placed in this groupbecause of its similarityin inflorescence, flower, and androecialmorphology to R. capixabensis;vegetatively,however,R. avilensis is isolatedin the genus, anda close relationshipbetween it-growing as it does in the northernextremeof the Andes-and a species of the Atlantic rain forest is unlikely. THE RHODOSTEMONODAPHNE SCANDENS GROUP The R. scandens group, including 15 species, is the largestof the groupsdiscussed here, and it is also very heterogeneous.Most species are small trees to shrubs, or scandent shrubs, and have reddish indument. The first seven species form a seemingly cohesive unit held togetherby relativelyshortandthickened petioles (which I refer to as "pulvinate"),and inflorescenceswith few branchorders.The remaining eight species are linked to the first seven and to each other,eitherindividuallyor in pairs,by differentcharacters. The first three species in the group are scandent shrubs or shrubs with drooping branches.The main characteruniting these species is their persistentcataphylls, which can be found on the old twigs, even aftertwo growthperiods. These three species have a characteristiclong, dense, persistentindumenton the twigs. The inflorescencesaremostly spicate,although in R. licanioides the inflorescencesrangefromspicate to much-branched.Rhodostemonodaphnescandens and R. licanioides appearto be sister-species. Both species have leaves with thick, coriaceousblades,that are often undulateor bullate; the margin is in some specimens slightly recurved,and the secondaryveins are brochidodromousthroughout.Anatomically,the two species have a distinctive thick spongy mesophyll, composed of columnarcells arrangedin a lax reticulum.In both species the adaxial side of the tepals is nearly glabrous,as are the stamens, although there are considerabledifferences in both shape and position of the antherlocelli. A few specimensof R. licanioideshave leaf blades with a rounded or even minutely cordate base, approachingthe characteristicallycordatebases found in R. miranda.The blades of R. mirandadiffer from the precedingtwo species in thatthey are thinner,the venation is eucamptodromous,and the secondary veins are not parallel,with the lower veins being oriented at a more acute angle than the rest. Rhodostemonodaphnerevolutifolia, with its yel- 22 lowish indument,much-branchedinflorescences,and two locellate anthers,is quite distinct. Nevertheless, it has pulvinatepetioles, and leaves similarin texture to those of R. licanioides andR. scandens; the former sometimes has revolute leaf marginslike those of R. revolutifolia. Rhodostemonodaphne crenaticupula andR. debilis both sharea numberof characterswith the firstthree species of this group.The indumenton the twigs, although not as dense, is also reddish. It too has pulvinatepetioles, but the leaves are considerably different. The inflorescences have few but somewhat elongated branches.While the stamens of both R. revolutifolia and R. crenaticupula are glabrous and similar to those of the first three species, the tepals are adaxiallyhairy. Rhodostemonodaphnecuricuriariensisand R. tumucumaquensisare small trees or shrubs.They share with the above-mentionedspecies the pulvinatepetioles, yet their leaves, unlike any of those mentioned above, are thinner and nearly glabrous on both surfaces. As in the first three species of the group, the leaf blades are undulateto nearly bullate. The fruits of these two species are unknown,and at least in R. tumucumaquensis,the margin of cupule must be straightsince the tepals abscise. The twigs of R. curicuriariensis are glabrous,while those of R. tumucumaquensis have a sparse reddish indument as in R. crenaticupula.The adaxial surface of the tepals and stamens of both R. curicuriariensisand R. tumucumaquensis are glabrous, as in the first three species mentioned. The next species, R. negrensis,is a small tree, and the petioles are somewhatthickenedbut not conspicuously pulvinate. The inflorescences are muchbranchedand the flowersare large.The leaves in size, texture,and type of tertiaryvenationare very similar to those of R. napoensis, a species which, on the basis of other characters,I include in the R. kunthiana group.Nevertheless,the cupules of R. negrensishave a deeply undulatemargin, as in the most part of the species included here. The next two species, R. mirecolorataand R. rufovirgata, are vegetativelyindistinguishableandhave a leaf morphology similar to R. sailensis, a species of the R. grandis group.Yet the red, densely tomentose indument in all plant parts links them to other species in this group, and places them far from the sericeous species of the R. grandis group.The flower size and stamen type of these two species are very different.The stamensof R. mirecoloratapoint in the direction of the R. laxa group, while those of R. rufovirgata,with their latrorselower locelli, are typical of the R. kunthianagroup. The fruits of R. rufo- FLORA NEOTROPICA virgata have woody cupules with a straightmargin, which also point in the directionof the R. kunthiana group. Rhodostemonodaphneceliana and R. steyermarkiana also have the deep reddishindumentand lobed cupules that place them in this group. They both inhabit the sandstonemountaintops of the Venezuelan Guayana,where low scrubbyvegetationis common. The habit of R. celiana (scandentshrubs)seems to be differentfrom that of R. steyermarkiana(collector's notes consistently reportit as being an erect shrub). Notwithstandingtheir differences in habit and other morphologicalfeatures,they may constitutea pair of closely related species. Endlicheria vinotinctaC. K. Allen is a anotherspecies from the sandstonemountains of Venezuelaand Colombia,identicalin habitto R. celiana, with similar leaf morphology, but with two-locellate anthers. The relationships among all three species should be investigatedfurther. Rhodostemonodaphneleptoclada and R. parvifolia also have yellowish brown indument, and the only fruit known, that of R. parvifolia, has a deeply lobed cupule. Both species are thin-branchedshrubs with small leaves; their small flowers have a minute receptacle.They aresimilarto the very heterogeneous Endlicheria gracilis Kostermans,as well as to the monotypic genus Phyllostemonodaphne.Rhodostemonodaphneantioquiensisis placed here for lack of a betterplace to put it. It too has thin, reddishtomentose twigs, and leaves similar to those of R. leptoclada. Yet its fruits, which have reflexedcupules, are unlike any other in the genus, and are very similarto those found in species of Endlicheria,in particularE. rubrifloraMez. Confirmationthatthe fruitsbelong to this species is still needed. Biogeographically the R. scandens group is largely restrictedto northeasternSouth America and centralAmazonia, with the exception of the puzzling R. antioquiensis from the CordilleraCentralof Colombia. Eight species are found in the Guianas/Guayana highlands, and five are distributedin central Amazonia.However,the firstseven species, which on the basis of morphologyform a more cohesive unit, are not restrictedto a single biogeographicalregion. On the contrary,hypothesizedsister-speciesrelationships, such as betweenR. licanioides andR. scandens, cut across these boundaries;the formerspecies has a localized distributionnear Iquitos, Peru, the latteris widespreadwithin northeasternSouth America. The apparentdisjunctionsseen in both R. scandens andR. rufovirgata,where no species are reportedfor Suriname, may be due to relatively low collection densities in the interiorof that country. SPECIESGROUPS The species in the next two groups lack, as far as the informationis available,the undulateto lobed cupules characteristic of the preceding two groups. While the R. grandis group is very cohesive, due to its conspicuous sericeous indument,the species composition of the R. kunthianagroup is very heterogeneous. The two groupsare not closely relatedto each other,nor to eitherof the two precedinggroups,apart from all sharingthe charactersthat define the genus. 23 stamens,often with latrorselower locelli. Yetit stands apartfrom the other species in the group on account of its small, thinly chartaceousleaves and its very peculiar urceolate flowers with minute erect tepals. The fruitshave a cupule with a straightmargin,but it appearsnot to be woody, as in R. kunthianaand R. synandra;the surfaceof the cupule is smooth. The last species in this group, Rhodostemonodaphne elephantopus,has very little in common with the other membersexcept that the flowers with deep THE RHODOSTEMONODAPHNE KUNTHIANA receptacles are similar to those of R. kunthiana;the stamens of whorls I and II, althoughcompletely glaGROUP The species in this group are only weakly linked brous, have latrorse lower locelli. The hairs of the buds are whitish, straight,and appressed by a few characters.The majorityof the species in- vegetative cluded here have an indumentthat consists of short, (see the R. grandis group). The small, elliptic to obovate leaf blades with cuneatebases can be compared crisped, reddish hairs. Also, the stamens of most of the species are hairy,at least the filamentor the basal only with those of R. avilensis (R. laxa group). Its fruits are unknown. portionof the anthers.The anthersof whorls I and II While Rhodostemonodaphnemorii and R. elehave latrorse,or nearly latrorse,lower locelli. Howarerestrictedto SurinameandFrenchGuiphantopus there are various even within a sinever, exceptions, the remainderof the species (apartfromthe wideana, to the In conditions described. R. kunthigle species, ana, for example, while the hairs of most collections spread R. kunthiana) are all found in the eastern seen are reddish,short,andcrisped,particularlythose Andeanfoothills. However,the numberof collections from the Amazonian lowlands, some of the collec- availableof eitherR. longipetiolataand R. napoensis tions from the Andes and Costa Rica have yellowish, (one and two collections, respectively)is too small to infer much about their distribution.Nevertheless,R. long, straightto curvedhairs. The first two species, R. kunthianaand R. synan- synandra,with only five collections known, but from dra, are vegetatively very similar and are without a broadrange, is found only in this region. For a dedoubt closely related. Their fruits have large woody tailed account of the distributionand variationof R. see the discussion under its species decupules, often with a rough, warty surface,and a thin kunthiana, scription. straightmargin,similar to species in the genus Pleurothyrium.The distinctly scalariformtertiaryvenaTHE RHODOSTEMONODAPHNE GRANDIS tion of these two species has no parallelin the genus, GROUP but weakly scalariformvenationis presentin the next The Rhodostemonodaphnegrandis group is the species, R. napoensis, as well as in R. negrensis of the R. scandens group. productof the segregationof the differentformscomRhodostemonodaphnenapoensis, although very prising the R. grandis complex (see below). All spesimilar to R. negrensis, has the indumenttypical of cies have a conspicuously sericeous indumentumon this group(the hairsof R. negrensisare straight).Fur- all parts,althoughthis may be soon glabrescent.The thermore,the stamens of R. napoensis have latrorse fruits are also very similar. The cupules are mostly lower locelli, while those of R. napoensis have con- thickened and woody, have a conspicuous pedicel, spicuouslyintrorse(whorlsI andII) lower locelli. The and are relatively large for the genus. Rhodostemoncupules of R. negrensis are deeply lobed; unfortu- odaphne grandis differs from the other three species by its largeleaves with manyparallelsecondaryveins, nately, those of R. napoensis are unknown. The only specimen available of Rhodostemono- althoughits floralmorphologyis similarto that of R. daphne longipetiolata is completely glabrous. The sailensis. Rhodostemonodaphnepeneia, R. recurva, flowershave the stamensof the outertwo whorls var- and R. praeclara sharethe same floral morphology. In indumentand fruit morphology,this group reiously doubled and fused. Notwithstanding,the stamens are very similarto those of R. kunthiana,in that sembles various species of Endlicheria, particularly they arehairyandhave moreor less columnaranthers, E. metallica and allied species, more than any other and latrorselower locelli. species of Rhodostemonodaphne.However, their Rhodostemonodaphnemorii is placed in this four-locellate anthersplace them in Rhodostemonogroup on the basis of its crisped indumentand hairy daphne. 24 FLORA NEOTROPICA NortheasternSouth Americais the region with the in Guyana,andN. dioica Mez, from the Acre riverin highest species representationin this group. Both eastern Brazil, to synonymy underN. grandis. KosRhodostemonodaphnepraeclara s.str.and R. grandis termansbased his decision on the apparentsimilarity have wide, but non-overlapping, distributions in of the flowers of the former,downplayingthe impornortheasternSouth America;the formeron the west- tance of the distinctiveindumentumcharactersof the ern side in Venezuela and Guyana, the latter to the latter-Nectandra dioica has an indumentconsisting east in Suriname,FrenchGuiana,and Amapa,Brazil. of two types of hairs (straight and appressed, and Rhodostemonodaphnesaulensis is localized in central crisped and erect), while all the hairs of N. grandis French Guiana and sympatric with R. grandis s.str. are straightand appressed. After the segregationof the dioecious species of Rhodostemonodaphnepraeclara s.1. has the widest distribution,spreadacrossthe Amazonianbasin.Rho- Nectandra into Rhodostemonodaphne, Rohwer dostemonodaphnepeneia and R. recurva are re- (1986) transferredNectandra grandis to Rhodostestrictedto centralAmazonia, where they are sympat- monodaphnegrandis, keeping Nectandrapraeclara ric with partof R. praeclara s.1. as a synonym. He neverthelessrecognizedN. dioica as a distinct species, making the new combination SPECIES INCERTASEDIS Rhodostemonodaphnedioica (Nectandra superba Rhodostemonodaphnependuliflorais a very odd A. C. Sm., based on a single specimen, also from species indeed. Charactersof rare occurrencein the Acre, was synonymizedwith R. dioica). family or unique to it are the clear, mucilaginousexApproximately150 specimens of Lauraceaehave udate, papillose tepals, glands on the outer staminal been annotated by specialists as Rhodostemonowhorls, and introrse anthers of whorl III. It shares daphnegrandis. These come from nonfloodedforests with other species of Rhodostemonodaphnedioecy of the Amazon river basin, and the north-draining and predominantlyfour-locellatestamens (the varia- lowlandsandmid-elevationsof easternVenezuelaand ble locelli numberobserved may be due to the fact the Guianas.This widespread,highly variabletaxon that the only known specimen is pistillate). Within I here referto as the "R.grandis complex."The great RhodostemonodaphneI can suggest no clearrelation- majorityof specimens have a sparse to dense, short, ship. One species of Endlicheria,also fromthe Pacific ascending appressed, silver-sericeous indument, coast of Colombia,E. brownianaMez, is vegetatively mainly on the twigs, but also present on the abaxial similar,but the flowers are completely different,not leaf surface, inflorescences, and flowers. This induonly in having two-locellate anthersbut also in their ment and a series of leaf characters(e.g., the firsttwo size and shape. Other species with papillose flowers secondaryveins arising from or near the base of the are found in Nectandra, but the stamens of R. pen- lamina and soon joining the margin) are the most dulifiora are very differentfrom any stamens in that strikingvegetativecharactersof the complex. Almost genus, andNectandracomprisesonly hermaphroditic half of the totalnumberof specimensaresterile,while species. Unfortunatelythe wood anatomyof this spe- one-fourthhave only fruits.The flowers of the fertile cies is nondescriptand typical of the various genera specimens are constantin their externalmorphology. of the Ocotea group (H. Richter,pers. comm.). I am All of them have a spherical receptacle that is displacing this species in Rhodostemonodaphne,largely tinctly constricted at the tepal inception. Closer into avoid erecting yet anothermonotypicgenus in the spection of their flowers, however, reveals the presence of highly distinctive floral traits, both varying family. among and constant within differentgroups of individuals. THE RHODOSTEMONODAPHNE After dissecting flowers from all fertile individuGRANDIS COMPLEX als, a total of seven different flower morphologies Mez, in his Lauraceae Americanae (1889), pub- were found. Seven specimens had flowers with clalished the name Endlicheria?grandis based on a sin- vate, two-locellate anthers typical of the genus Engle fruiting collection made by Melinon from the dlicheria, and in fact belong to this genus. These can French Guiana side of the Maroniriver. Kostermans be grouped into three differentspecies. Three fertile (1936), after having seen fertile material from the collections (Oliveira(INPA)59389 & 59887 andZak Guianas and the Amazon basin that was similar to & Espinoza 4671) have flowers with erect tepals and Melinon's specimen, correctlytransferredE. grandis all fertile whorls with clavate, two-locellate anthers. Mez to Nectandra (as N. grandis). He also reduced They match the type of E. metallica Kosterm., a bothN. praeclara Sandwith,from the Essequiboriver poorly known taxon from easternAmazonian,in both THE RHODOSTEMONODAPHNE GRANDISCOMPLEX vegetative and floral morphology. Some specimens annotatedby van der Werffas E. metallica, however, are quite different from the type (cf. R. Vdsques & Jaramillo9180 and van der Werff& Jaramillo9815) and have now been described as a new species of Endlicheria,E. chrysovelutinaChanderb.Two other fertile collections (Grdndezet al. 1612 and Prance et al. 14013) have flowers with two-locellate anthers, and are again differentfrom those of E. metallica and E. chrysovelutina;these have been recentlynamedas E. ruforamulaChanderb.They can be distinguished from both groups by more or less lanceolate leaves thatdry light greenadaxiallyandhave stronglyacrodromous venation and golden-brown indument. In these charactersthey resemble E. bracteolata (Meissner) Allen, a species close to E. metallica. These three species of Endlicheria are part of a larger assemblage of taxa with sericeous indument that includes E. krukovii(A. C. Sm.) Kosterm.,E. Ihotzkyi Mez, and E. sericea Nees. The remainingfloweringspecimensclearlybelong to Rhodostemonodaphne.They are dioecious and their anthers are four-locellate, with the locelli arranged in an arch near the apex of the anthers.The species can be divided into four groups on the basis of floralmorphology.These groupsandthe characters distinguishing them are presented below and summarizedin TableV. 25 Rhodostemonodaphnegrandis s.str. The remaining specimens from French Guiana and Suriname match the type of R. grandis in many respects,especially in leaf morphology, where, compared to all other specimens, the number of secondary veins is high, rangingfrom 6 to 10 pairs. Their flowers differ from those restrictedto Sail in that the anthersare not overlapping and the glands are considerably smaller and not dissected. Their tepals are yellow at anthesis. Rhodostemonodaphne saiilensis. Collections from the surroundingsof Sail in centralFrenchGuiana have flowers with wide, overlappinganthersof whorls I and II and large, dissected, pairedglands on whorlIII, unlike any otherflowersin the complex (the size of the glands, as far as I can tell, is not dependent on degree of maturationof flowers as is common in most Lauraceae).Theirtepals are reddishat anthesis. Jens Rohwer had annotatedmost of these specimens as an undescribedspecies relatedto R. grandis. Rhodostemonodaphne praeclara. A group of specimens from western Guyana and eastern Venezuela match the type of Nectandrapraeclara. Their flowers are peculiarin thatnone of the specimensexamined have glands at the base of the stamens of whorl III, a rare condition in Lauraceaeas a whole. The tepals are thinnerthan those of the two groups described above and their three, parallel traces are TABLE V Comparisonof the species in the Rhodostemonodaphnegrandis complex (R. recurvaexcluded) R. grandis s.str. R. peneia R. saiilensis R. praeclara s.1. Twig diameter(mm) 4.8-7 2.8-4.4 2.2-2.8 4-5(-6.4) Leaf width (cm) 5-12(-21) (2-)4-7 3-7 3-6(-9) Leaf base angle (?) 80-140 40-80 50-100 50-90 Pairs of secondary 6-10 (3-)4-5(-6) 5-7(-8) 4-6 Tepal color (when fresh) Yellow Red Green Yellow Androeciumin receptacle Included Included Exserted Exserted Glands of whorl III Small, simple Large,divided Absent Absent Locelli of whorl III Latrorse/extrorse Latrorse/extrorse Apical Geographicrange Suriname,French Guiana,NE Brazil FrenchGuiana veins CentralAmazonia (Manaus) Latrorse/extrorse NE Venezuela, Guyana,Amazonia 26 FLORA NEOTROPICA conspicuous when light is shone through them. At anthesisthe tepals are yellow, althoughthey often dry reddish.The leaves of these specimens are very uniform, being chartaceousand slightly obovate with an obtuse, mucronate to shortly acuminate apex. Furthermore,their twigs are notably thick. Some specimens from throughoutthe Amazon basin have identical flowers, but in their vegetative morphologythey are quite variable;they are here referredto as Rhodostemonodaphneaff. praeclara. Rhodostemonodaphnepeneia. A group of specimens from the vicinity of Manaus has flowers like those of N. praeclara, with thin tepals and lacking glands. Yet their inflorescencesand flowers are much smaller,and the anthersof whorl III are uniquein this complex in having locelli located towardsthe apex as opposed to their strictly lateralposition in other species in the complex. Their tepals are greenish at anthesis. The recently named Rhodostemonodaphnerecurva (van der Werff & Vicentini, 2000), a species with odd locelli numberwhich initially promptedme to determinethe specimens as an undescribedspecies of Endlicheria,is withoutdoubta memberof the Rhodostemonodaphnegrandis group. Its exclusion from the following morphometricstudy does not alter the results presentedhere. species of Endlicheria.Frequencydistributionswere approximately unimodal, with a tendency to be skewed towardsthe lower values. Therewas no strong bivariate concordance of characters.A summary of the measurementstaken and values for each species is presentedin TableVII. Multivariateanalysis using principalcomponents analysis (PCA), based on a correlationmatrixof standardizedmedians, was used to detect any clustersestablished by interactionsbetween all characters.The computer program NTSYS-pc version 1.80 (Rohlf, 1994) was used in all analyses. Results In a firstPCA analysis using 11 charactersand the complete set of specimens, the first and only significant principalcomponent(PC 1) accountingfor 45% of the variation,separatedthose specimensbelonging to Rhodostemonodaphne grandis s.str. and Endli- cheria sp. 1 from the rest (Fig. 4). However,all were linked by the intermediatespecimens of E. aff. bracteolata and E. metallica. The main cluster was com- posed of specimens of the remaininggroups. Rhodostemonodaphne grandis s.str. is, in fact, morphologicallyquite distinct from any otherspecies in the complex as shown in a second PCA where the specimens belonging to Endlicheria are excluded MORPHOMETRICSTUDY Methods In an attempt to find vegetative differences that may supportthe groups described above on the basis of floralmorphology,I selected and measured10 leaf charactersand twig diameterin a quantitativestudy of morphometricvariation.The leaf characterswere petiole length and width, leaf lamina length and width, acumenlength,laminabase angle, laminaapex angle, numberof pairs of secondary veins, angle of divergenceof secondaryveins, chordalangle (the angle made by a line from the point of insertion of the secondaryto the place it comes nearestto the margin). The characterswere chosen on the basis of their observed variation in the total sample. Furthermore, these are charactersthat can be readily observed in sterile specimens. Twenty-nine fertile specimens representingeach of the four species of Rhodostemonodaphneand the three species of Endlicheria discussed above were chosen for the analysis (TableVI). All the characters measured showed continuous variation. The specimens correspondingto Rhodostemonodaphne grandis s.str. often formed a cluster at the higher end of the scale, but intermixedwith specimens of either of the vI t g v o o? o \ vV o FirstPrincipal Component FIG.4. Principalcomponentanalysis of the Rhodostemonodaphnegrandis complex. All species of the R. grandis complex (including 3 species of Endlicheria) included. R. sailensis (circle);R. grandis (emptysquare);R.praeclara s.str. (triangle);R. peneia (rhomboid);R. praeclara (upsidedown triangle);E. metallica (bottom-left-shadedsquare);E. sp. 1 (bottom-right-shadedsquare);E. aff. bracteata (topright-shadedsquare). THE RHODOSTEMONODAPHNE GRANDISCOMPLEX 27 TABLE VI Specimens examined for morphometricstudy of the Rhodostemonodaphne grandis complex Species Collector and number Endlicheriaruforamula Grandezet al. 1612 Endlicheriaruforamula Prance et al. 14013 Endlicheriametallica Krukoff4932 Endlicheriametallica Oliveira(INPA)59639 Endlicheriametallica Zak & Espinoza 4671 Endlicheriachrysovelutina R. Vdsquez& Jaramillo9180 Endlicheriachrysovelutina van der Werff& Jaramillo9815 praeclara s.l. Rhodostemonodaphne T D. Pennington& Tenorio10760 Rhodostemonodaphne praeclara s.l. Prance et al. 14616 Rhodostemonodaphne praeclara s.1. J. L. dos Santos (INPA)1202.5028 praeclara s.1. Rhodostemonodaphne M. F FESilva et al. 1411 praeclara s.l. Rhodostemonodaphne M. G. Silva & Bahia 3604 Rhodostemonodaphnepraeclara s.1. Sperling et al. 5928 Rhodostemonodaphnegrandis BAFOG 7425 Rhodostemonodaphnegrandis BW 3204 Rhodostemonodaphnegrandis Mori et al. 23998 Rhodostemonodaphnegrandis Schulz (LBB)7419 Rhodostemonodaphne peneia Aluisio 122 peneia Rhodostemonodaphne KurzB-28-14 peneia Rhodostemonodaphne Rodrigues6976 peneia Rhodostemonodaphne Rodrigues 7009 praeclara s.str. Rhodostemonodaphne Fanshawe 130 = FD 2739 praeclara s.str. Rhodostemonodaphne Sandwith387 Rhodostemonodaphnepraeclara s.str. Steyermark111392 praeclara s.str. Rhodostemonodaphne Steyermark60403 Rhodostemonodaphnesaiilensis Mori et al. 14787 Rhodostemonodaphnesaulensis Mori et al. 15014 Rhodostemonodaphnesailensis Mori et al. 20744 Rhodostemonodaphnesailensis Mori et al. 20904 TABLE VII Summary of measurementsused in morphometricstudy of the Rhodostemonodaphneg E. m R. saiilensis R. grandis R. praeclara s.str. R. peneia R. praeclara s.I. (2.4-)3.1(-4) (4.8-)5.3(-5.6) (4-)4.8(-6.4) (1.6-)2.4(-2.8) (3-)3.6(-4.2) (2.6-) Petiole length (cm) (1.5-)2.4(-4.1) (2.4-)3.4(-4.4) (1.2-)2.1(-2.8) (1.2-)2.2(-3.1) (1.3-)2.1(-3.5) (1.8-) Petiole width (mm) (1.6-)2.4(-3.8) (2.8-)3.6(-4.2) (2.2-)3(-3.6) (1.2-)2(-2.4) (1.6-)2.4(-3.6) (2-)2 Leaf length (cm) (9.5-)16(-21) (14-)21(-30) (7.8-)13(-19) (8.8-)15(-22) (7.2-)13(-20) (9.2- Leaf width (cm) (2.8-)4.6(-7) (5.6-)9.3(-14) (3.3-)5.5(-8.2) (2.4-)5.4(-7.1) (3.1-)5.5(-8.3) (4.2 Acumen length (cm) (0-)0.5(-1.9) (0-)0.6(-1.8) (0-)0.1(-0.6) (0-)0.6(-1.8) (0-)0.3(-1.2) (0.2 (3-)4(-5) (5-)7(-9) (4-)5(-6) (4-)6(-7) (4-)6(-9) (4 (30-)42(-60) (40-)50(-60) (30-)41(-50) (30-)45(-60) (30-)45(-60) (40- (10-)18(-30) (20-)30(-40) (20-)21(-30) (10-)24(-30) (20-)21(-30) (20- Base < (?) (40-)55(-80) (70-)90(-110) (50-)72(-90) (50-)68(-100) (50-)68(-100) (60-) Apex < (0) (50-)75(-110) (80-)99(-130) (90-)113(-130) (60-)90(-130) (80-)96(-120) (80-) Twig diameter(mm) 2nd # 2nd. < (?) Chordal< () 'Measurementstaken individually from specimens in Table VI and summarized here by species; (min-)average(-max) SPECIESCONCEPTS 29 (Fig. 5). Thus the apparentmorphologicalcontinuity between Rhodostemonodaphnegrandis s.str. and the rest of the groups is due to the mistakeninclusion of species of Endlicheria in the Rhodostemonodaphne grandis complex. Rhodostemonodaphnepraeclara, R. peneia, and R. saiilensis cannot be furtherdifferentiatedthroughPCA with the charactersused. Conclusions From the previous analyses the following conclusions can be drawn. Specimens of Endlicheriametallica, E. aff. bracteolata, and Endlicheria sp. 1 are vegetatively intermediatebetween Rhodostemonodaphnegrandis s.str. and the remainingspecies of the R. grandis complex. Rhodostemonodaphnegrandis s.str. is vegetatively distinctwithin the R. grandis complex (excluding Endlicheria spp.). It is restrictedto the lowlands of northeasternSouth America (Suriname, French Guiana,andAmapa,Brazil). Althoughno single character is diagnostic for this species, it can be distinguished from the other species of the complex by differencesin medianleaf width, medianbase angle, and median numberof secondaryveins. Rhodostemonodaphnesaiilensis, the only species found in sympatry with R. grandis s.str., can be sharplydistinguishedfrom it on the basis of vegetative characters(see Fig. 5). These include twig diameter,numberof secondaryveins and base angle. 0 a 0 0o oD 0 o V 0oo?? v oV ~VA 19~ A v A FirstPrincipal Component FIG. 5. Principalcomponentanalysis of the Rhodos- temonodaphnegrandis complex. Species of the R. grandis complex,excluding species in Endlicheria.R. saiilensis (circle); R. grandis (square);R. praeclara s.str. (triangle); R. peneia (rhomboid);R. praeclara (upside-downtriangle). Rhodostemonodaphnepeneia, from centralAmazonia, cannot be distinguished vegetatively from specimens of R. praeclara s.l. Nevertheless, it has a distinct floral morphology. Rhodostemonodaphnepraeclara s.str. constitutes a very homogeneous group both vegetatively and in its floral morphologywhen consideredin a restricted sense. It is representedby the specimens from the mid-elevationsand lowlandsof easternVenezuelaand western Guyana. Finally, a numberof specimens of Rhodostemonodaphne praeclara s.l., although identical in floral morphology to R. praeclara s.str., are vegetatively quite variable. These are distributedthroughoutthe Amazon basin. They may be part of R. praeclara, constitutea separatevegetativelyvariabletaxon, a series of homogeneous taxa closely relatedto R. praeclara, or even be the productof a single or multiple hybridizationevents between R. praeclara and R. peneia. Due to the lack of evidence to supportany of these possibilities, I have designatedthe specimensof R. praeclara from the Amazon basin as R. praeclara s.l. This way, the identities of both R. peneia and R. praeclara are maintained,and the close relationship of the Amazonian specimens with the latter on the basis of their sharedfloralmorphologyis pointedout. SPECIES CONCEPTS The species here treated,be they old andtaxonomically stable entities, new circumscriptionsof seemingly cohesive taxa, or new species from recently availablematerial,have all been delimited on the basis of morphological discontinuities. However, the data on which these discontinuitiesare based are inevitably of a circumstantialand fragmentarynature. They are circumstantialdue to the impossibility of visiting even a small percentage of the localities where the plants are known to occur. So this revision, like most others,relies heavily on specimenscollected by other botanists as part of floristic surveys. They are of a fragmentarynaturebecause the collections at a particularlocality are often made from single individuals, and at best, consist of a only few flowering or fruitingtwigs. Thus, an indicationof the extent of populationor the degree of individualvariationcannot be inferred, and any interpretationof character variationbased on this poor sampling must be done carefully. All species describedarepresumedmonophyletic, i.e., including only members related by descent and closer to each otherthanto the membersof any other species. The methodology used in detecting species is basically whatis commonly referredto as the "mor- 30 FLORANEOTROPICA phological species concept,"where consistent differences in morphology,separatedby gaps, are considered evidence of the existence of separatespecies. No one characterwas used as sole indicator of species status, but mainly a combinationof specific characters, or overall appearance.While one species may be diagnosedby leaf shape and indument(e.g., Rhodostemonodaphneovatifolia), anothercan be based on floral characters(e.g., R. negrensis). Some species are strikingly superficiallysimilar. One such group is formed by the Rhodostemonodaphne grandis complex. Here a previously widespreadandvariable"species"was interpretedas being composed of various species based on gaps in the distributionof characters(see "The Rhodostemonodaphne grandis Complex"). Phenology and habitatpresence, as far as the informationwas available,were checked for each species and groups of species, so as to obtain some idea of the ecological natureof the species. Subspecificrankshave been intentionallyavoided, the level of resolutionon the relationshipsof the organisms being very low. The species describedhere are the smallest entities separableby morphological gaps. However, I do not rule out the possibility that nonreticulatinglineages exist within some of the species here described. SYSTEMATIC TREATMENT INTERPRETATIONOF DESCRIPTIONS The species descriptionshave been generatedentirely using the DELTAprogram(DEscriptionLanguage for TAxonomy,Dallwitz et al., 1993). Thus all descriptions are strictly parallel and follow identical formattingparameters.In cases of missing or inapplicable data, the relevant charactershave been left out for ease of reading. All measurementsare taken from dry herbarium materialexcept for those of flower parts,which were taken from rehydratedflowers. When measurements are approximatethey are indicated with a "ca."preceding the value, otherwise normal ranges and in some cases extreme values (in parenthesis)are provided.Variationin characters,whereintermediatesare observed, is indicated by "to";where gaps between the characterstates are present,"or"is used. A question markindicatesthatthe informationwas not available to me directly and I have inferredthe character state. The colors describedfor differentorgans(e.g., leaf lamina, indument,tepals, anthers,etc.) are those observed in the dried specimens. In many cases these differ considerablyfrom colors observedin freshmaterial which have sometimes been recordedon collection labels and given here separately. The use of basitonic, mesotonic, or acrotonic for branchesor inflorescencesrefers to their relative position along the length of a flush. Twig diameteris measuredto the nearestmillimeterat approximately 5 cm below the terminalbud. Indumentterminology follows thatproposedby Hewson (1988). In all cases where a type of indument is mentioned, details of hair type and persistence are added. Length of the longest hairs is given to the nearest tenth of millimeter. Bud dimensions are given to the nearestmillimeter, the diameterreferring to the widest part of the bud. Petiole length is given to the nearest millimeter; width to the nearest tenth of millimeter.Leaf blade texture,as coriaceous or chartaceous,reflects a relative subjectiveindicationof bladethicknesswhen dry. Blade length and width are measuredto the nearest centimeter;length does not include the acumen.Leaf characters,including overall shape, base and apex shape, and venation follow to a great extent the terminology of Hickey (1973). Leaf base and apex angles are given to the nearest ten degrees. Acumen/ mucro length is given to the nearest millimeter.Angles of divergence of the secondary veins are measuredfor the firstcentimeterof the vein, to the nearest five degrees. Chordalangles of secondaryveins, the angle made between the primaryvein and a straight line tracedbetween the origin of the vein andits point nearestto the margin,are measuredto the nearestfive degrees. Peduncle length denotes the total length of the main axis of the inflorescence;it is measuredto the nearestcentimeter.The hypopodiumis the partof the pedunclebelow the firstlateralbranch;lengthis given to the nearestcentimeter,width to the nearesttenthof millimeter.Inflorescencebranchordersare given assumingthe peduncleto be a first-orderbranch.Length of the longest second-orderbranch is given to the nearest millimeter.Bracts and bracteoles, when persistent, are measuredto the nearesttenth of millimeter. Pedicel diameter is measured at the base of the pedicel and is given to the nearesttenthof millimeter. Flower receptacledimensionsare given to the nearest tenth of millimeter; receptacle length is measured from where the pedicel shows an abruptincrease in diameterto the base of the outer tepals. Outertepal length and width are measuredto the nearesttenthof mm. Stamenand staminodedimensionsare given approximatelyin tenthsof millimeter.Pistil dimensions SYSTEMATICTREATMENT are given as total length, and length and width of the ovary to the nearesttenth of millimeter. Fruitpedicel length and width (nearthe base), cupule length and width, and berrylength andwidth are measuredto the nearestmillimeter. Descriptiveinformationextractedfrom labels and field notes is provided in a separateparagraph.This includesplanthabitand size, buttress,bark,andwood characters.Colors of organs observed on fresh material are also providedhere and often differfromthose observed on the dried specimens. Measurementdata takenfrom field labels have been transformedto metric when necessary. 31 chartaceousto membranaceous,elliptic, ovate, obovate to oblong, the outerand innerones mostly equal, erect, spreadingto recurved,hairy or glabrous(papillose); stamens of whorls I & II spathulate,capitate, filiform (loriform), or anthers sessile, thickened or laminar,the two whorls mostly equal, glabrousto puberulous (papillose); anthersmostly 4-locellate, but 2- or 3-locellate ones present;locelli apical in a shallow archto nearlybijugate,introrseor the lower pair latrorse;glands mostly absent; whorl III columnar, capitate or filiform (loriform); anthers mostly 4locellate, but 2- or 3-locellate ones present;the upper locelli mostly latrorseand lower ones extrorseor all locelli extrorse (introrse); glands mostly present, globularor variouslyfused in a ring; whorlIV mostly absent or staminodial;pistillode present or absent. Bot. Rohwer & Kubitzki, Rhodostemonodaphne Pistillate inflorescences: similarto the staminateones Jahrb.Syst. 107: 135. 1985; Rohwer, Mitt. Inst. often with fewer branchordersand flowers. Pisbut Allg. Bot. Hamburg20: 82. 1986. Type. Rhodostillate flowers: ovary filiform to ovoid, or globose, temonodaphnelaxa (Meissner)Rohwer. mostly glabrous. Fruits: pedicels gradually to Meissner,in A. L. de Candolle,Prodr. Synandrodaphne abruptlyenlargingto form the cupule; cupule hemi15(1):176. 1864,nonGilg(1915),nom.conserv. to Nectandra subgen. Synandrodaphne(Meissner) Mez, spherical trumpet-shaped(rarelyreflexed),smooth, wrinkled,or tuberculate;marginwith truncateto deep Jahrb.Konigl.Bot.Gart.Berlin5: 400. 1889. lobes, undulateor straight;tepals persistingor caduDioecious trees, shrubs, or scandent shrubs: cous; berry elliptic (ovoid). branchesbasitonicto mesotonic,in axils of cataphylls or foliage leaves, scattered(pseudo-verticillate);twigs KEY TO THE SPECIES OF terete (angular); indumentpresent (or absent), perRHODOSTEMONODAPHNE sisting to soon falling; terminalbuds plump or slenThis key includes the 41 taxa in this treatment,as der; cataphylls caducous (or persisting).Leaves spimetallicaKosterm.andallied spewell as Endlicheria ral, with 2/5 phyllotaxis;petioles robust, slender, or the basis of vegetativecharacters, on which have, cies, pulvinate, adaxially flattenedor terete; blades coriawith species of the R. granconfused been commonly ceous to chartaceous,flat, undulateto bullate;outline Lauraceae specimen dis flowering Any complex. elliptic, ovate, obovate to oblong; base rounded,obto identified Rhodostemonodaphne using the generic tuse, acute, cuneate to cordate (decurrent); apex van der Werff la) may be iden(199 by key provided muor to acuminate acute attenuate, rounded,obtuse, will Note that R. here. to tified revolutifolia species euveins to revolute; cronate;marginplane secondary its because to the out in generic key Endlicheria, camptodromous(brochidodromous),(forked); terti- key rememUsers should two locelli. have anthers outer ary veins random-reticulateor scalariform;indument ber that measurementsof the length of the leaf blade present or absent, persisting to soon falling. Staminate inflorescences: basitonic, mesotonic, acrotonic, do not includethe lengthof the acumen.An indication or along whole length of flush, pendulous or erect, of geographicalrange is given for most species, prispicatewith up to 2 branchordersto lax, thyrsoidwith marily as a confirmatory character.Colors, when 5(-8) branchorders,the branchesscattered(clustered used, arethose observedon the driedspecimens.Only apically); bracts and bracteoles caducous or persist- where the species are endemic to readily distinctive ent, adaxially hairy or glabrous. Staminateflowers: geographicalareasor habitats(e.g., tepuis), or in they pedicel diametereven throughoutto increasing api- case of well-known disjunct distributions(e.g., Ancally; receptacle obconical to flat, or globose (con- des/easternAmazonia-Atlanticrain forest), is geogstrictedat place of tepal inception);tepalscoriaceous, raphyused as a key character. 1. Twigs, leaves, and inflorescencessilver to golden sericeous, the hairs always straightand appressed(look at young organs under25 X magnification). 2. Leaf lamina soon glabrescent;matureleaf blades normally <10 cm long, the base cuneate, the secondaryveins almost perpendicularto primaryvein. 32 FLORANEOTROPICA 3. Large trees; leaf venation flat and inconspicuouson both surfaces;lowlands, FrenchGuiana .................................................................................................................................................35. R. elephantopus 3. Shrubs;leaf venationraised on both surfaces (at least when dry); cloud forests, Venezuela....... 12. R. avilensis 2. Indumenton leaf lamina persistingat least on lower surfaceof currentflush; matureleaf blades >(7-) 10 cm long, the base acute to rounded,the secondaryveins forming an acute angle with the primary vein. 4. Twigs stout, normally >5 mm diam. Secondaryveins normallymore than 7 pairs. 5. Tepals erect; anthers2-locellate; C and W Amazonia.................................................E. metallica & spp. aff. 5. Tepals spreading;anthers4-locellate; NE South America ........................................................ 36. R. grandis 4. Twigs slender,normally <5 mm diam. Secondaryveins normallyfewer than 7 pairs. 6. Anthersof whorl III with conspicuouspairedglands (tepals on fresh materialred); French Guiana.......................................................................................................................................37. R. saiilensis 6. Anthersof whorl III with pairedglands minute or lacking (tepals on fresh materialgreen or yellowish). 7. Leaf marginrecurvedat base; Manausand vicinity........................................................... 39. R. recurva 7. Leaf marginplane throughout. 8. Inflorescences<11 cm long, the pedicels ca. 2.4 mm long; tepals green; Manausand vicinity ..............................................................................................................................40. R. peneia 8. Inflorescencesto 16 cm long, the pedicels ca. 5.6 mm long; tepals yellowish; NE Venezuela,Guyana,and Amazonia.............................................................................38. R. praeclara 1. Twigs, leaves, and inflorescencesnot sericeous, either glabrous,puberulous,tomentoseor pubescent,the hairs variously colored, straightto crisped, erect to spreadingto appressed. 9. All plant organs (except terminaland axillary buds) glabrous,or soon glabrescent. 10. Petioles distinctly pulvinate;leaf blades undulateto bullate;Curicuriaririver,Brazil... 19. R. curicuriariensis 10. Petioles not distinctly pulvinate;leaf blades flat. 1. Twigs normally >3 mm diam.; leaf blades normally >12 cm long. 12. Twigs drying black; inflorescencespendulous;tepals and stamenspapillose; lowland forests, Pacific coast of Colombia...........................................................................41. R. penduliflora 12. Twigs drying yellow to brown;inflorescenceserect; tepals and stamens not papillose; E Andean foothills ................................................................................................32. R. longipetiolata 11. Twigs normally <3 mm diam.; leaf blades normally <12 cm long. 13. Secondaryveins to 6 pairs, arising at an acute angle to the primaryvein; submontane forests, Colombia ...................................................................................................28. R. antioquiensis 13. Secondaryveins normallymore than 6 pairs, almost perpendicularto primaryvein. 14. Large trees; leaf venation flat and inconspicuouson both surfaces;lowlands, ................................... 35. R. elephantopus FrenchGuiana ............................... 14. Shrubs;leaf venationraised on both surfaces(at least when dry); cloud forests, 2. R. avilensis Venezuela........................................1.................. 9. All plant organs with some kind of indumentpersistingon currentflush (if sericeous, see couplet 1). 15. Plants of the summitsof Tepuis in the VenezuelanGuayanahighlands. 16. Scandentshrubs;leaf apex attenuate,lacking a mucro;inflorescencespendulous;Auyan-tepuf .................................................................................................................................................24. R. celiana 16. Erect shrubs;leaf apex not attenuate,with a fine, sharpmucro;inflorescenceserect; Coro.................. 25. R. steyermarkiana Coro-Yutajemassif ................................................... 15. Plants not of the summitsof tepuis in the VenezuelanGuayanahighlands. 17. Secondaryveins mostly straightand parallel;tertiaryveins distinctly scalariform,ca. 3 mm apart. 18. Stamenslaminar(loriform),those of whorls II and III partiallyfused in a ring; E Andean foothills ...........................................................................................................30. R. synandra 18. Stamens columnar,those of whorls II and III not fused; widespread........................29. R. kunthiana 17. Secondaryveins mostly arched,if parallelthen not conspicuously so due to arching;tertiary veins random-reticulateor if weakly scalariformthen ca. 6 mm or more apart. 19. Indumentof twigs and inflorescencesred to darkreddishbrown;leaf blades below drying reddishor brownish. 20. Cataphyllspersistingon maturetwigs (for at least two flushes). 21. Leaf bases distinctlycordate;secondaryveins fading before connecting with adjacentsecondaryveins, except towardsthe apex; Guyana....................... 15. R. miranda SYSTEMATICTREATMENT 33 21. Leaf bases not distinctly cordate;secondaryveins distinctlyloop-connected. 22. Scandentshrubs;tepals ca. 5 mm long; NE South America .............. 13. R. scandens 22. Shrubsto small trees; tepals to 2 mm long; W Amazonia............... 14. R. licanioides 20. Cataphyllssoon caducous or persistingonly on currentflush. 23. Leaves broadlyelliptic, normally >9 cm wide. 24. Inflorescencesnormally >16 cm long; outer antherswith lower locelli latrorse;AmazonianEcuador..............................................................31. R. napoensis 24. Inflorescencesto 16 cm long; outer antherswith all locelli introrse;Rio Negro and C Amazonia........................................................................21. R. negrensis 23. Leaves narrowlyelliptic, normally <9 cm wide. 17. R. crenaticupula . 25. Bracts persistentat anthesis;C Amazonia .................................. 25. Bracts soon caducous. 26. Flowers >7 mm diam.; Guyana.............................................22. R. mirecolorata 26. Flowers <6 mm diam.; Guianas...............................................23. R. rufovirgata 19. Indumentof twigs and inflorescences(and flowers) yellow or yellowish brown;leaf blades below usually drying greenish. 27. Flowers urceolate;tepals at anthesis erect, leaving a small opening;E Guianas.....33. R. morii 27. Flowers not urceolate;tepals at anthesis spreadingor if erect then stamensclearly visible. 28. Flowers minute, ca. 2 mm diam.; inflorescenceswith secondarybranches clusteredat the tip of the peduncle;C Amazonia ........................................ 34. R. sordida 28. Flowers >2 mm diam.; inflorescenceswith secondarybranchesscattered throughoutthe peduncle. 29. Leaves glabrousor nearly so; inflorescenceswith up to 3 branchorders. 30. Leaf blades narrowlyelliptic, >15 cm long; Curicuriaririver,Brazil .......................................................................................... 9. R. curicuriariensis 30. Leaf blades (broadly)elliptic to ovate, to 15 cm long. 31. Venationbrochidodromous,all secondaryveins similar; Tumucumaquerange .............................................. 20. R. tumucumaquensis 31. Venationweakly acrodromous(sub-triplinerved),the basalmost pair of secondaryveins strongerthan the rest. 32. Leaf blades to 8 cm long; C Amazonia...................... 27. R. parvifolia 32. Leaf blades normally>8 cm long; FrenchGuiana... 26. R. leptoclada 29. Leaves nearly always with some type of conspicuous indument,at least on lower surface;inflorescencesnormallywith more than 3 branch orders,at least towardsthe base. 33. Bracts persistentat anthesis. 34. Matureleaves with conspicuouslyrevolutemargin;French 16. R. revolutifolia Guiana...................................... ............... 34. Matureleaves with flat margin. 35. Leaves above light green to bluish green, below olivegreen to greenishbrown, the veins distinctyellowish; fruits small, pedicel to 10 mm long; C Amazonia .............................................................................. 17. R. crenaticupula 35. Leaves brown, the veins concolorous;fruits large, pedicel 20 mm long or longer, C Amazonia .......................................R. debilis 33. Bractscaducous before anthesis. 36. Plants of the Atlantic rain forest or restingain SE Brazil. 37. Flowers >7 mm diam.; antherssessile, broad,and thick; SE Brazil..........................................................................9. R. anomala 37. Flowers <6 mm diam.; stamens filamentous,filiform,or spathulate. 38. Stamens filiform;widespreadwithin the Atlanticrain forest ................................................................. 10. R. macrocalyx 38. Stamens spathulate;restingasof Espirito Santo ........................................................................ 11. R. capixabensis 36. Plants of Amazonia or the Andes. 39. Flowering pedicels sharplyseparatedfrom the receptacle; SW Amazonia. 34 FLORA NEOTROPICA 40. Leaf blades nearly glabrousbelow or with short, erect hairs only; inflorescencesnormallylonger than the leaves; Bolivia .......................................................4. R. longiflora 40. Leaf blades below with mixed indumentconsisting of both appressedand erect hairs;inflorescencesshorter than the leaves; Peru-Brazil-Bolivia border............... 8. R. dioica 39. Floweringpedicels graduallyenlargingto form the receptacle(trumpet-shaped);plants of the N Andes and E Andean slopes. 41. Leaves broadlyelliptic (nearlyorbicular),ovate or obovate (ca. 2x longer than wide). 42. Antherswith variablelocelli number,the two median locelli often vestigial or fused; Andes of Ecuador..............................................................7. R. cyclops 42. Anthersfour-locellate. 43. Secondaryveins 8-10 pairs, brochidodromous;Andes of Colombia ............................................................6. R. frontinensis 43. Secondaryveins 10-11 pairs, eucamptodromous;Andes of Venezuela ............................................................... 3. R. ovatifolia 41. Leaves normallyelliptic (ca. 3x longer than wide). 44. Leaf blades below distinctly velutinous;margin recurvedat base; CordilleraOriental,Colombia ..........................................................................2 . R. velutina 44. Leaf blades below puberulousto tomentose to rarelyvelutinous;marginplane throughout. 45. Secondaryveins (6-)7(-9) pairs, cupule hemispherical;CordilleraCentral,Colombia ........................................................................1. R. laxa 45. Secondaryveins 3-4(-5) pairs, cupule flat (disklike);E Andean slopes .................. 5. R. juruensis mary vein above slightly raised, below prominent; secondary veins (6-) 7 (-9) pairs, equidistant,eucamptodromous,above slightly raised, below prominent, diverging at ca. 70?, straightto evenly arching (forked), chordal angle 40-50?, the angle uniform along blade length; tertiary veins above and below raised, random-reticulate;higher-orderveins above andbelow slightlyraised;surfaceabove shinyyellowish brown in young leaves, opaque brown in older leaves, the veins slightly lighter,below dull yellowish Trees: branchesbasitonic,in axils of cataphyllsor brownin young leaves, faintly reddishbrownin older basal foliage leaves, often pseudo-verticillate;twigs leaves, the veins lighter; indumentabove puberulous angular,soon becoming terete, 3-6 mm diam.; epi- to absent, the primary and secondary veins tomendermis brownish to black, barely visible due to in- tose, caducousby next flush to persistingfor at least dumentcover; terminalbud plump, 3-7 X 2-3 mm; two flushes,below puberulousto velutinous,the hairs cataphylls caducous; indumentpubescent, persisting dense to sparse, to 1 mm long, straightto curved, for at least two flushes, the hairs dense, to 0.3(-0.9) erect, yellowish, denseron the veins, persistingfor at mm long, straightto curved,erect, yellowish. Leaves: least two flushes. Staminate inflorescences: along petioles robust,0.5-1.3 cm X 1-3 mm, adaxiallyflat- whole length of flush, pendulous,peduncles 3-11 cm tened;blades chartaceous,flat, narrowly(broadly)el- long, the hypopodia 2-6 cm X 1-1.5 mm, branch liptic, (5-) 10-14 (-18) X (2-) 3-5 (-7) cm; base orders 4, the second-orderbranches4-7, dispersed, acute to obtuse, ca. 100?;apex acute, 70-120?, ulti- lowest branchto 1.5(-4) cm long, color andindument mately acuminatefor up to 0.5 cm; marginplane;pri- of all axes as on twigs; bractsandbracteolescaducous 1. Rhodostemonodaphne laxa (Meissner) Rohwer, Mitt. Inst. Allg. Bot. Hamb. 20: 83. 1986. Synandrodaphnelaxa Meissner,in A. L. de Candolle, Prodr. 15(1): 176. 1864. Nectandra laxa (Meissner)Mez, Jahrb.Konigl. Bot. Gart.Berlin 5: 400. 1889. Type. Colombia. Quindio: Quindfo Pass, Nov 1827 (d fl), Goudots.n. (holotype:K [photo NY neg. 8502]; isotypes: FI-W-n.v., G [frags. ex FI-W et P], P). Fig. 6A SYSTEMATICTREATMENT 35 A i B i/^ iD-- .~~~~~::I ._ r;~~~~~ ."'. zl _ _: ., 1 _ i'~ .ye.;'.~:i (ojo Ltta ?-t~~~~~~~~~~~~~~s ~ dq' Terdn Dugarte 15884). D. R. ~ longilora FIG. 6. A. Rhodostemonodaphne (Pearce ~ =' "a't : l.a srm.>.'?ii;/ /;...' ~ ~ ~ ~ ) 26X) laxa (Franco et al. 3956). B. R. velutina (Triana 2040/3). C. R. ovatifolia (Ruiz- 36 (not seen). Staminateflowers: pedicels ca. 5.6 X 1.2 mm, the diameter gradually increasing apically; receptacle obconical, ca. 3.2 X 3.5 mm; tepals coriaceous, ovate, ca. 5 X 4 mm, at anthesis erect to spreading, black, adaxially glabrous; stamens of whorlsI and II, the antherssessile, chubby,trapezoid, ca. 1.4 X 1.2 mm, with a few hairsat base, the locelli 4, apical, in a shallow arch, introrse,the glands absent; whorl III columnar,ca. 1.6 X 0.8 mm, with a few hairs at base, the locelli 4, the upperpairlatrorse, the lower pair extrorse, the glands fused, forming a continuousring protrudingbetween the anthersof the outer whorls; whorl IV absent; all stamens blackish; pistillode filiform, ca. 1.7 X 1 mm, glabrous.Pistillate flowers: pistil ca. 3 X 1.5 mm; ovary ovoid to globose, ca. 1.5 mm long, glabrous.Fruits: pedicels to 45 X 5 mm, graduallyenlargingto formthe cupule; cupule hemispherical, to 7 X 19 mm, smooth, the margin undulate,tepals persisting;berry elliptic, to 35 X 21 mm. Field notes. Trees to 8 m tall and 21 cm diam.; wood aromatic(tangerineodor); leaves below bluish green. Receptacle yellowish green;tepals cream;stamens/staminodescream. Cupule yellow to reddish. Distribution (Fig. 7). Along CordilleraCentralof Colombia (N Andes) from Quindio (E slope) to Antioquia; growing in upper-montanecloud forest, at 2000-3010 m. Flowering October-Februaryduring the dry season. Immaturefruits have been collected in Marchandfound ripe by the new floweringseason. FLORANEOTROPICA been made between these two areas, it is presumed that the species can be found in the interveningdepartmentsof Risaraldaand Caldas. Rhodostemonodaphnelaxa is most closely related to R. velutina,from which it differs primarilyby the denser indument (see discussion under the latter). Rhodostemonodaphnefrontinensis may also be closely related.It differs vegetativelyin the size and shape of the leaves, those of R. frontinensis being broadlyelliptic to obovate. Specimens of this species were repeatedly misidentified as Ocotea declinata (Meissner) Mez by C. K. Allen. Ocotea declinata ( 0. deflexaRohwer, 1986), a species from Bahia, is indeed vegetatively very similar, althoughI have seen only photographs of this species. Nevertheless, its inflorescenceswith theirpersistingbractsare very differentfrom those of R. laxa. 2. Rhodostemonodaphne velutina (Mez) Madriinan,comb. nov. Nectandra velutina Mez, Jahrb. Konigl. Bot. Gart.Berlin 5: 401. 1889. Type. Colombia. Norte de Santander:Ocafa, Andes de Ocafia, May 1851 (d fl), Triana2040/3 = 1034 [= Schlim451] (holotype:P; isotypes: B [frag.ex P], COL-n.v., K). Fig. 6B Trees: branchingunknown;twigs terete,ca. 6 mm diam.; epidermisbrownish,barely visible due to indument cover; terminalbud plump, ca. 6 X 4 mm; cataphyllscaducous;indumentvelutinous,persisting for at least two flushes,the hairsdense, to 1 mm long, Additional specimens examined. COLOMBIA. ANto erect, yellowish. Leaves: petioles TIOQUIA: Medellin-Rionegrord.,below the summitof Santa straight curved, X 3-4.5 mm, adaxially flattened; Elena, 2 Apr 1949 (fr), Barkley et al. 389 (NY, US); San robust, 1-1.5 cm blades coriaceous, flat, elliptic, 10-21 X 5-8 cm; Pedro (Y imm fr), Bro. Daniel 1439 (GH); Mar 1938 (Y imm fr), Bro. Tomas163 (US). QUINDio:Salento, La Co- base obtuse to rounded, 80-110?; apex obtuse, 70100?; margin recurved at base; primary vein above cora, Estaci6n La Montafia,28 Sep 1992 (st), Betancur et al. 3756 (COL,GH);Calarca,QuindioPass,W slope,hwy. slightly raised, below prominent;secondaryveins 7above Calarca,22 Nov 1944 (Y fl, fr), Fosberg22333 (US); 10 pairs,equidistant,weaklybrochidodromous,above Salento, Estaci6n Navarco, 24 Sep 1992 (d fl), Franco et flat,below raised,divergingat 65-70?, evenly arching al. 3956 (COL, GH, MO); Salento, La Cocora,Estaci6nLa (forked), chordalangle 45?, the angle uniformalong Montafia,28 Sep 1992 (st), 4013 (COL, GH); Salento, Es- blade length; tertiaryveins above flat to slightly imtaci6n Navarco, 23 Sep 1992 (d fl), Rangel et al. 9008 below slightly raised, random-reticulate; pressed, (COL, GH, MO); (st), Rangel et al. 9051 (COL, GH); Salveins above inconspicuous, below higher-order ento, Acaime, Jul 1993 (fr), Vargaset al. 1053 (MO). slightly raised; surface above greenish to reddish Rhodostemonodaphnelaxa can be characterized brown,below reddishbrown;indumentabove puberby its yellow tomentose indumentpersistingon most ulous, the veins tomentose, caducous by next flush, parts of the plant (except for the inflorescences and below velutinous, the hairs dense, to 1 mm long, flowers, which are glabrous), leaves drying light straight to curved, erect, golden to reddish brown, olive-green, few-flowered, lax inflorescences, and denseron the veins, persistingfor at least two flushes. Staminate inflorescences: basitonic, pendulous, pefleshy, trumpet-shapedflowers. The specimens from Antioquiahave largerleaves duncles 5-11 cm long, the hypopodia3.5-6 cm X ca. than those of Quindio. Although no collections have 2.5 mm, branch orders 3(-4), the second-order SYSTEMATICTREATMENT 37 ' ' 10 7 S 0 - Rh. anomaS V Rh. avilens i 1000k. . 1 V ? * | Rh capixabensis l .. / II- \7 -1 Rh. Cyclops * Rh. dioica Rh. I frontinensis in tRh. juruensis dg r A Rh. longiflora Rh.macrocalyx ---Rh. -W ovatifolic^ -a,. m u o se \\\c \ Rd .. 1^ _ _2-. + Rh. velutina ) i ' i. . l; ' X ' s / gu/ | V~ \> i" / .- -- V|a 0^^ 0 5J1 ~ -- FIG. 7. Distributionmap of species in the Rhodostemonodaphnelaxa group. Each symbol representsone or more collections found in a degree-square. branches 4, dispersed, lowest branch to 2 cm long, color and indumentof all axes as on twigs, epidermis drying black; bracts and bracteoles caducous (not seen). Staminateflowers: pedicels ca. 6 X 1 mm, the diametergraduallyincreasingapically;receptacleobconical, ca. 1.5 X 4.5 mm;tepals coriaceous,obovate, ca. 2.8 x 3 mm, at anthesiserect to spreading?,black, adaxiallypuberulousto tomentose;stamensof whorls I and II, the antherssessile, chubbytrapezoidto ovate, ca. 1.2 X 1 mm, glabrous, the locelli 4, apical, in a shallow arch, introrse, the glands absent; whorl III columnar,ca. 1.2 X 0.8 mm, glabrous, the anthers trapezoid, the locelli 4, the upper pair latrorse, the lower pair extrorse,the glands absent (? or fused in a ring); whorl IV absent;all stamensreddishblack;pistillode absent.Pistillate flowers andfruits unknown. Field notes unavailable. Distribution (Fig. 7). Known only from the type collected on the CordilleraOriental of the Andes in northernColombia near the town of Ocafa; growing in cloud forest at ca. 2000 m. The floweringcollection was made in May towardsthe beginning of the rainy season. Rhodostemonodaphnevelutina is clearly most closely relatedto R. laxa, a cloud forest species of the CordilleraCentralof the Andes. The two species are very similar in habit, leaf size and shape, and flower morphology.The most strikingdifferenceis the dense velutinous indumentof R. velutinacovering all plant parts, including inflorescence axes and flowers. Further differences are its leaf margins recurved at the base, and an anatomicalcharacter-tabular upperepidermis (in R. laxa it is isodiametric) with a thicker cuticle. The flowering times of the two species are different, R. laxa flowering November-February,R. velutinain May. Mez (1889) consideredNectandra velutina a separatespecies fromN. laxa, on the basis of the different FLORA NEOTROPICA 38 indument.Rohwer (1986) proposedit as a likely synonym. Even though we know nothing about the variation of indumentcharacterfor R. velutina,none of the 11 known specimensof R. laxa show this extreme velutinous indument. In Colombia the Cordillera Orientaland the CordilleraCentralof the Andes are separatedby a wide, deep valley (the Magdalenariver valley). The species may be found on the unexplored mountaintopsof the Serraniade San Lucas, a lowlying mountainrangebetween the CordilleraOriental and the CordilleraCentral;if so, continuitybetween the two known ranges would be more likely. There are no collections of this species in the south of the country where the two Cordillerasmeet-although far from completely known, the Colombianmontane flora is one of the best collected in the country.It is highly unlikely that these populationsare reproductively linked, and this separationmay be quite oldthe Andeanuplift startedin the Tertiarymore than 10 m.y.b.p.-unless they are the result of a recent event of long distance dispersal.Until more collections are available,I preferto keep them as separatespecies. lowish brown,persistingfor at least two flushes. Staminate inflorescences: along whole length of flush, erect,peduncles7-12 cm long, the hypopodia4-6 cm X 1-2 mm, branch orders 4, the second-order branches4-5, dispersed,lowest branchto 3 cm long, color and indument of all axes as on twigs; bracts caducous; bracteoles caducous. Staminate flowers: pedicels 3 X 1.2 mm, the diametergraduallyincreasing apically;receptacleobconical, ca. 3 X 5 mm; tepals coriaceous, ovate, ca. 3 X 2.5 mm (inner whorl slightly smaller), at anthesis erect to spreading,reddish brown,adaxiallytomentose;stamensof whorls I and II, the antherssessile, chubby trapezoid,ca. 2.5 X 1.8 mm, glabrous,the locelli 4, apical,in a shallow arch, introrse,the glands absent;whorl III columnar, ca. 2.5 x 1 mm, glabrous,the locelli 4, the upperpair latrorse,the lower pair extrorse,the glands globose, more or less fused, ca. 1 mm diam.;whorl IV absent; all stamensreddish;pistillode filiform,ca. 2 X 1 mm, glabrous.Pistillate flowers andfruits unknown. Field notes. Trees to 4-5 m tall. Tepalsred. Distribution (Fig. 7). Known only from the type specimen collected on the Cordillerade Mrida, the 3. Rhodostemonodaphne ovatifolia Madrifian,sp. northerncontinuation of the Colombian Cordillera nov. Type. Venezuela.Tachira:Dtto. Jauregui,La Orientalof the Andes, in Venezuela.Growing at ca. Grita-ElRosal rd., 8 Oct 1978 (J fl), Ruiz-Terdn 2500 m in cloud forest; flowers found in October. & J. A. Dugart 15884 (holotype:MO). Fig. 6C Etymology. Named for its ovate leaves [Lat.ovalaxaeet R. velutinaeaffinis,sed Rhodostemonodaphne foliis ovatisbenedistincta. Trees: branchingunknown; twigs angular,soon becoming terete, 5 mm diam.; epidermis brownish, barely visible due to indument cover; terminalbud plump, 8 X 4 mm; cataphylls caducous; indument tomentose, caducous after one flush, the hairs dense, to 1 mm long, straightto curved, erect, brownishto reddish. Leaves: petioles robust, 1-1.5 cm X 2-4 mm, terete; blades coriaceous, flat, ovate, 12-16 X 6-9 cm; base rounded(flat to almost cordate), 150180?; apex acute to attenuate, 100-120?, acuminate for up to 2.5 cm; marginplane; primaryvein above slightly raised,below prominent;secondaryveins 1011 pairs, equidistant, eucamptodromous, above slightly raised, below raised, diverging at 55?, abruptlyarchingnear margin(forked),chordalangle 40?, lowest pair more obtuse than rest; tertiaryveins above inconspicuous, below raised, randomreticulate; higher-orderveins above inconspicuous, below slightly raised;surfaceabove yellowish brown, below reddishbrown;indumentabove tomentose,the primaryand secondaryveins densely tomentose, caducous after one flush, below tomentose, the hairs dense to sparse, to 0.8 mm long, straight,erect, yel- tus, + folium, leaf]. Rhodostemonodaphneovatifolia has distinctly ovate leaves; only R. dioica has a leaf shape that approachesthis ovate outline. Unusually for the genus, the stamens of both outer whorls have a pair of distinct lateralappendages,similarto the glandsof whorl III. The general shape of the stamens of Rhodostemonodaphneovatifolia resembles that of R. laxa and R. velutina,but the tepals lack the distinct thick medial ridge of these two species. Externallythe flowers are very similarto those of R. mirecolorata. On the basis of both vegetative morphology and biogeographythis species forms part of the core of the R. laxa group (e.g., R. laxa and R. velutina).The twigs and leaves aredensely coveredwith a yellowish brown indument,characteristicof the species of this group, and like them it is a high-altitudespecies of the N Andes. 4. Rhodostemonodaphne longiflora Madrifiin, sp. nov. Type. Bolivia. Vicinity of Santa Cruz, Jul 1865 (d fl), Pearce 26-X (holotype:K). Fig. 6D SYSTEMATICTREATMENT 39 Ex affinitateR. laxaesed inflorescentiis longioribuset staminibus laminaribus stipitatisbenedistincta. the northof the country in Deptos. Beni and Pando; flowering in July during the dry season. Trees: branching unknown; twigs angular, soon becoming terete,4-5 mm diam.;epidermisblack;terminalbud plump,ca. 1 X 1 mm; cataphyllscaducous; indument puberulous, caducous by next flush, the hairs sparse,to 0.2 mm long, straightto curved,erect, yellowish to reddish. Leaves: petioles robust, 1-1.5 cm X 2-2.5 mm, adaxially flattened;blades coriaceous, flat, elliptic, (6-)8-13 X 3-6 cm; base acute to obtuse, 60-110?; apex obtuse, ca. 100?; margin plane; primaryvein above flat, below raised;secondary veins 4-6 pairs, equidistant,weakly brochidodromous,above flat,below raised,divergingat 50-55?, evenly arching(forked),chordalangle ca. 20?,the angle uniformalong blade length; tertiaryveins above flat, below slightly raised, weakly scalariform to random-reticulate;higher-orderveins above flat, below slightly raised;surfaceabove greenishbrown,the veins darker,below yellowish green, the veins reddish; indumentabove puberulous,the primary vein tomentose,caducousby next flush,below puberulous, the hairs isolated, to 0.3 mm long, straightto curved, erect, golden yellow to reddish, caducous by next flush. Staminate inflorescences: basitonic to mesotonic, pendulous?,peduncles (3-)11-15 cm long, the hypopodia (0.5-)3-4 cm x ca. 2 mm, branchorders 5, the second-orderbranches5-14, dispersed,lowest branchto ca. 2 cm long, all axes black, indumentof all axes as on twigs; bracts and bracteolescaducous (not seen). Staminateflowers: pedicels 10 X 0.6 mm, the diameter even throughout;receptacle obconical, ca. 1.5 X 4.8 mm; tepals membranaceous,ovate, ca. 1.8 X 1.4 mm, at anthesiserect to spreading,reddish to black, adaxially glabrous;stamens of whorls I and II spathulate,chubby,the anthersovate, ca. 0.8 X 1 mm (whorl II slightly smaller), glabrous, the locelli 4, apical, in a shallow arch, introrse,the glands absent; whorl III columnar,ca. 0.9 X 0.5 mm, glabrous, the antherswith a medial adaxialprocess, oblong, the locelli 4, the upper pair latrorse,the lower pair extrorse, the glands globular,ca. 0.6 mm diam.; whorl IV absent;all stamensreddish;pistillode absent.Pistillate flowers andfruits unknown. Etymology. Named for its inflorescences,unusually longer than the leaves [Lat. longus, long + flos, flower]. Rhodostemonodaphnelongiflora is characterized by the long inflorescences, relative to the length of the leaves. They become glabrousby next flush and are black drying, contrastingwith the greenish- to yellowish-dryingleaves; they also appearto be pendulous. The leaves are very similarto those of R. laxa from the mountainsof Colombia,to which it may be closely related. It neverthelessdiffers in flower morphology, the anthersof R. laxa being sessile, while those of R. longifloraare borne on a conspicuousfilamentousportion. Mez determinedthis specimen as Nectandra anomala, a species from the Atlantic forest of Brazil. Rhodostemonodaphneanomala has a denser,yellowish indumentand larger(ca. 16 X 8 cm) leaves. 5. Rhodostemonodaphne juruensis (A. C. Sm.) Chanderb.,Fl. Neotrop.Monogr.91: 1-140. 2004. Anibajuruensis A. C. Sm., Phytologia 1(3): 116. 1935. Endlicheriajuruensis(A. C. Sm.) Kosterm., Recueil Trav. Bot. Neerl. 34: 542. 1937. Type. Brazil: Amazonas, near mouth of Rio Embira (tributaryof Rio Tarauaca),Jun 1933 (Y fl, imm fr), Krukoff4775(holotype:NY-n.v.; isotypes:An.v., F-n.v., G-n.v., K-n.v., MO, U-n.v.). Fig. 8 Trees: branchesbasitonic,in axils of cataphyllsor basal foliage leaves; twigs angularand remainingso for at least two flushes, 2.5-3 mm diam.; epidermis black;terminalbuds slender,ca. 5 X 2 mm;cataphylls caducous; indumentpuberulous,caducous after one flush, the hairs dense, to 0.1 mm long, straight,appressed, reddish.Leaves: petioles slender, 1-2 cm X 1-1.5 mm, terete; blades chartaceous,flat, elliptic, (6-)9-18 X 3-6 cm; base acute, 50-80?; apex acute, 50-80?, acuminate for up to 2.5(-3) cm; margin plane; primaryvein above slightly impressed,below prominent;secondary veins 3-4(-5) pairs, equidistant,brochidodromous,those towardsapex occasionField notes. Trees to ca. 13 m tall. Tepals green- ally forked, above flat, below slightly raised, diverging at 40-60?, archingnear base, chordal angle 20ish? 30?, the angle uniform along blade length; tertiary Distribution (Fig. 7). Known only from the type veins above and below slightly raised, randomcollection attributedin the label to the vicinity of reticulate to scalariform;higher-orderveins above SantaCruz, Bolivia, at ca. 300 m-it is doubtfulthat and below slightly raised; surface above dark shiny Pearce collected in Santa Cruz de la Sierra or any- brown, below light dull brown; indumentabove abwhere in Depto. de SantaCruz;there are otherlocal- sent, below puberulous,the hairs sparse, to 0.3 mm ities named SantaCruzin the Amazonianlowlandsin long, straight,erect, yellowish, persistingfor at least FLORA NEOTROPICA 40 two flushes. Staminateplant unknown.Pistillate inflorescence: acrotonic, erect (curved upwards), peduncles 4-8 cm long, the hypopodia2-4 cm X 1-1.5 mm, branchorders3, the second-orderbranches5-6, dispersed,lowest branchto 2 cm long, all axes blackish, sparselypuberulous,the hairsreddish;bractsand bracteoles caducous (not seen). Pistillate flowers: pedicels ca. 2 X 0.1 mm, the diametergraduallyincreasingapically;receptacleobconical,ca. 2 X 2 mm; tepals coriaceous, ovate, ca. 1 X 1 mm, at anthesis erect to spreading, black, adaxially patchy puberulous; staminodes of whorls I and II spathulate,the anthers reniform, ca. 0.3 X 0.3 mm, glabrous, the locelli 4, apical, in a shallow arch,introrse(outertwo often latrorse),the glands absent; whorl III capitate, ca. 0.3 X 0.4 mm, glabrous, the locelli 4, the upper pair latrorse,the lower pairextrorse,the glands forming a continuoussegmentedring; whorl IV absent;all stamensblackish;pistil ca. 1.5 X 0.5 mm; ovary globose, ca. 0.8 mm long, glabrous.Fruits: pedicels to 20 X 3 mm, abruptlyenlarging to form the cupule; cupule spreadingto 15 mm diam., smooth,the margin undulate;berry elliptic, to 30 X 20 mm. Field notes. Medium-sized to large canopy trees (9-)20-25 m tall and 20-60 cm diam. Innerbarkdark brown,aromatic.Leaves shiny green.Flowerscreamy green or white with reddish tinge. Fruits with dark reddishorange cupule and black matureberries. koa$ _nL_ Lni@~S:~ s ~ .gr :07 _hhj-I;C FIG. 8. Rhodostemonodaphnejuruensis (Krukoff4775). Distribution (Fig. 7). Apparentlywidespreadon the slopes of the Andes from Ecuador to Bolivia at frontinensis Madrifidn, 6. Rhodostemonodaphne Colombia. nov. Antioquia: Mun. Fronsp. Type. tino, Murr rd. 15 km W of Nutibara (Altos de Cuevas), 18 Oct 1987 (6 fl), Brant & Martnez Additional specimens examined. ECUADOR. CHAR1387 (holotype: HUA; isotypes: AAU, HBG, K, CHI:Tulcan,parroquia Chical,sectorGualpimedio,Reserva MO, NY).F. Fig. 9A indigena Awa, 23-27 May 1992 (Y fl), Tipaz, Quelal & rd. new 1104 MORONA-SANTIAGO: Cantincuz (MO); Along Rhodostemonodaphnelaxae, R. cyclopi et Endlicheriae Mendez-Morona,16 Aug 1989 (Y fr), van der Werff&Gud- colombianae similis, a primariafoliis majoribuset stamiino 11136 (MO). nibus verticillorumI et II latioribus, a secunda ramulis toPERU. SANMARTiN:Prov.MariscalCaceres,Distr.To- mentosis, a tertiariaindumentonec adpresodiffert. cache Nuevo, camino a Santa Rosa, 5 Aug 1973 (9 fl), Trees: branches basitonic, in axils of cataphylls; Schunke6726 (MO). BOLIVIA. LA PAZ: FranzTamayo, Serranfade Chepite, CampamentoSismico de Texaco, 3-8 Apr 1992 (Y fl), epidermis brownish to black, barely visible due to indument cover; terminal bud plump, 7-9 x 4-6 mm; Killeen 3832 (MO). cataphylls caducous; indument pubescent, caducous Local names. Ecuador: yalte; Peru: moena. after one flush, the hairs dense, to 0.8 mm long, This species is a close relative of the Andean spestraight, erect, yellowish. Leaves: petioles robust, 1.5-2.6 cm X 2.4-3 mm, adaxially flattened; blades cies comprising the Rhodostemonodaphne laxa group R. R. R. R. chartaceous, flat, broadly elliptic to obovate, 8-20 X velutina, laxa, (R. ovatifolia, cyclops, dioica, and R. longiflora). It shares with these the 3-12 cm; base obtuse to rounded, minutely decurrent, 80-150?; apex obtuse to rounded, 80-120?, ultimately black-drying flowers with obconical receptacles, and for up acuminate androecium forming a ring. plane; primary primary 1.2 cm; cm; margin margin plane; to 1.2 acuminate for up to (350-)700-1000 m; in premontane to montane wet forests. Flowers April to August, with fruits ripe by August. 41 SYSTEMATICTREATMENT WJ. 3637521 NS 1589286 mor! ::R~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~DI^:Y~ -9. O:...k?L. " E.c: . k;. . : ' i.:>E-, . . ,ND KFAV. ,. tl(c. . , me . ... ... ..... . 'f l "" i.~r-- ' ",1-ml" * * t . .q,, -..e. 1, .no....-; ~ t) - ..:..i ........ i :d :. m.:; FIG. 9. A. Rhodostemonodaphnefrontinensis (Brant & Martinez1387). B. R. cyclops (Jaramillo& Grijalva 13643). C. R. dioica (Ule 9402). D. R. anomala (Glaziou 13150). 42 vein above flat, below prominent;secondaryveins 810 pairs, equidistant, brochidodromous,above flat, below raised, diverging at 50-60?, abruptlyarching near margin (forked), chordal angle ca. 30?, lowest pair more obtuse than rest; tertiaryveins above flat, below slightly raised, random-reticulateto scalariform; higher-orderveins above flat, below slightly raised; surface above olive-green to brown, below dark yellowish brown; indumentabove puberulous, the primary and secondary veins tomentose, below tomentose, the hairs dense, to 1 mm long, straightto curved, erect, yellowish, denser on the veins, caducous after one flush? Staminate inflorescences: basitonic, erect?,peduncles5-12 cm long, the hypopodia 1-4 cm X 1.4-2 mm, branch orders 5, the secondorderbranches4-8, dispersed,lowest branchto 3 cm long, all axes blackish, puberulous;bracts and bracteoles caducous (not seen). Staminateflowers: pedicels ca. 5.6 X 1.2 mm, the diametergraduallyincreasing apically;receptacleobconical, ca. 2.4 X 3.5 mm; tepalscoriaceous,ovate, ca. 2.5 X 1.5 mm, at anthesis erect to spreading, black, adaxially patchy puberulous; stamens of whorls I and 11,the antherssessile trapezoid, ca. 1.6 X 1.2 mm (whorl II slightly smaller),glabrous,the locelli 4 (the lower pair much largerthan the upperpair), apical, in a shallow arch, introrse,the glands absent;whorlIII columnar,ca. 1.4 X 0.8 mm, glabrous,the locelli 4, the upperpair latrorse, the lower pair extrorse (the upper almost introrse), the glands fused, forming a continuousring protrudingbetweenthe outerwhorls;whorlIV absent; all stamens yellowish brown to black;pistillode filiform, ca. 1.2 X 0.4 mm, glabrous.Pistillateflowers andfruits unknown. FLORA NEOTROPICA the indumenton the twigs is shorter,the inflorescence axes are thicker, and the flowers have a more elongated, trumpet-shapedreceptaclethanin R.frontinensis. Rhodostemonodaphne frontinensis is also similar to R. laxa andR. velutina,two closely relatedspecies also from the Andes of Colombia. However, the leaves of these two species are narrow-ellipticas opposed to the wide-ellipticones of R.frontinensis.Furthermore,the flowers of both R. laxa and R. velutina have narrowanthersof whorls I & II as opposed to the wide ones of R. frontinensis. In both vegetativeand inflorescencecharacters,as well as gross floral structure,R. frontinensis is very similarto Endlicheriacolombiana(Meissner)Mez, a species of upper-montaneforest in Colombia, also found in the same general locality as R. frontinensis. Nevertheless,E. colombiana has appressed,straight, ascendinghairs, and flowers with 2-locellate anthers. 7. Rhodostemonodaphne cyclops Madrifiin, sp. nov. Type. Ecuador.Pichincha:Rio GuajalitoReserve, 5 Jul 1991 (d fl), van der Werf, B. Gray, G. Tipas & J. Campaha 12190 (holotype: MO; isotypes: AAU, COL, GH, HBG, K, NY, QCA). Fig. 9B Foliis formaet indumento R. laxae similis,sed ramis crassisin sicconigris,antherarum locellissuperioribus plerdiffert. umquevestigialibus Trees: branchesbasitonic,in axils of cataphyllsor basal foliage leaves; twigs angularand remainingso for at least two flushes, 3-6 mm diam.; epidermis terminalbud plump, ca. 1 X 1 mm; cataphylls black; Field notes. Trees to 6 m tall. Inflorescenceaxes caducous; indumentpuberulous,caducous after one yellowish green; receptacle yellowish green; tepals flush, the hairs dense, to 0.3 mm long, straight to cream. crisped, erect, yellowish. Leaves: petioles robust, Distribution (Fig. 7). Known only from the type 1.5-3 cm X 2-3 mm, adaxiallyflattened;blades charfound on the northernpartof the CordilleraOcciden- taceous, flat, broadlyelliptic, 10-20 X 6-13 cm; base tal of the Colombian Andes, at 1850 m. The type obtuse to rounded,90-150?; apex obtuse to rounded, specimen was found growing at the marginof upper- 120-140?, (mucronate);marginplane; primary vein montane forest. Found flowering in October at the above flat, below prominent; secondary veins 4-7 height of the rainy season. pairs, equidistant,brochidodromous,above slightly raised, below prominent, diverging at 50-60?, Etymology. Named after the Mun. of Frontinoin abruptlyarchingnear margin(forked),chordalangle the departmentof Antioquia,Colombia. 35-40?, the angle uniformalong blade length;tertiary This is a species that closely resembles Rhodos- veins above slightly raised, below raised, randomtemonodaphnecyclops from the cloud forests near reticulate to scalariform; higher-orderveins above Quito, Ecuador.The indumenton all plant parts, in- and below slightly raised; surface above dark green cluding the puberulousadaxial surface of the tepals, to black, below brownishgreen; indumentabove abandleaf shape, size, consistency,andvenationof both sent, the primaryand secondaryveins tomentose,bespecies are very similar.Nevertheless, in R. cyclops low tomentose, the hairs sparse, to 0.6 mm long, SYSTEMATICTREATMENT straightto crisped, erect, yellowish brown,persisting for at least two flushes. Staminate inflorescences: basitonic to mesotonic, erect (curved upwards),peduncles 5-17 cm long, the hypopodia2-8 cm X 1.52.5 mm, branch orders 5(-6), the second-order branches2-8, dispersedor clusteredapically,lowest branchto 4 cm long, all axes blackish, sparsely puberulous;bracts caducous (not seen); bracteolespersistentor caducous,to 3 mm long, adaxiallyglabrous. Staminate flowers: pedicels ca. 5.6 X 0.8 mm, the diametergraduallyincreasingapically;receptacleobconical, ca. 2.5 X 3.2 mm; tepals coriaceous, ovate, ca. 2.5 X 2 mm, at anthesis spreading,black, adaxially patchy puberulous;stamens of whorls I and II spathulate,the anthersreniform,ca. 1 X 1.8 mm, puberulous,the locelli 3 to 4, apical, in a shallow arch, introrse,the glands absent;whorl III capitate,ca. 1 X 0.8 mm, puberulous,the locelli 4, the upper pair latrorse,the lower pairextrorse,the glands fused, forming a continuous ring; whorl IV absent; all stamens immersedwithin blackish;pistillode teardrop-shaped, the receptacle, lacking a stigma, ca. 2 X 1 mm, glabrous.Pistillateflowers: pistil ca. 3.2 X 2 mm; ovary globose, ca. 2.4 mm long, glabrous.Fruits unknown. 43 Rhodostemonodaphnecyclops sharesa numberof characterswith R. frontinensis, to which it may be closely related.They have similar,black-dryinginflorescences, and flowers with a long receptacle and spreading,pointed tepals with a few adaxialhairs. 8. Rhodostemonodaphne dioica (Mez) Rohwer, Mitt. Inst. Allg. Bot. Hamb.20: 83. 1986. Nectandra dioica Mez, Repert.Spec. Nov. RegniVeg. 16: 308. 1920. Type.Brazil.Acre:Rio Acre, Sao Francisco, Aug 1911 (d fl), Ule 9402 (holotype: B [photo F neg. 3753]; isotypes: G, K, US-n.v. [photo NY neg. 7037]). Fig. 9C Nectandrasuperba A. C. Sm., Phytologia 1: 120. 1935. Type. Brazil: Acre, Basin of Rio Purus,near mouth of Rio Macuahuan,3 Sep 1933 (Y fl), Krukoff5757 (holotype: NY [photo NY neg. 7022]; isotypes: A, B X2 [photoNY neg. 7716], BM, F [photo F neg.], G, HBG [frag.], K, M, S, U, US). Trees: branchesbasitonic,in axils of cataphyllsor basal foliage leaves; twigs angularand remainingso for at least two flushes, ca. 5 mm diam.; epidermis black, barely visible due to indumentcover; terminal Field notes. Trees to 20 m tall, alreadyflowering bud slender,ca. 1 X 1 mm; cataphyllscaducous;indumentpubescent to tomentose, caducous after one when 12 m tall; wood aromatic.Tepals green. flush, the hairs dense, to 1 mm long, crisped, erect, Distribution (Fig. 7). Knownonly from the provyellowish. Leaves: petioles robust, 1.5-2.7 cm X 2ince of Pichincha,Ecuador.Three of the four known 3.5 mm, adaxially flattened;blades chartaceous,flat, specimens were collected in the "Rio Guajalito"re- broadlyelliptic (ovate), 8-18 X 3-8 cm; base obtuse, serve near Quito, at 1850-2000 m. Found in cloud minutelydecurrent,60-110?; apex acute,60-100?, ulforest and adjacentpastures. Flowers June-Septem- timately acuminate for up to 0.5(-1) cm; margin ber, during the dry season. plane; primaryvein above flat, below raised;secondAdditional specimens examined. ECUADOR.PI- ary veins 5-7 pairs, equidistant,eucamptodromous, CHINCHA:Rio Guajalito Reserve,29 Jun1991(d fl), Jar- above flat, below slightly raised,divergingat 50-55?, amillo & Grijalva13643 (NY); 9 Aug 1991 (9 fl), Jaramillo evenly arching(forked),chordalangle 20-30?, the an& Grijalva13676(QCA,NY);Quito-Nono-Tandayapa-Los gle uniform along blade length; tertiary veins flat andthe"Y"betweenMindo above, raisedbelow, random-reticulate to scalariform; Bancosrd.,betweenTandayapa andLos Bancos,7 Sep 1986(9 fl), Zak1163 (F, K, MO, higher-orderveins above and below slightly raised; NY,S, US). surface above dull to shiny, yellowish green to yellowish brown,below light yellowish brownto creamy Etymology. The specific epithetalludesto the variable numberof antherlocelli due to fusion of the two white; indumentabove absent, the primary vein tomentose,below tomentose,the hairsdense, to 0.5 mm upperones or theirvestigial nature,reminiscentof the ascending, yellowish, as one-eyed giant (often depicted with vestigial lateral long, straight, appressed, well as to 1.5 mm long, curved to crisped, erect and eyes) of Greek mythology. yellowish, persisting for at least two flushes. StamiRhodostemonodaphnecyclops is a very distinct nate inflorescences: along whole length of flush, species. It has thick but not very dense-wooded erect,peduncles5-12 cm long, the hypopodia2-5 cm branches; the leaves are coriaceous, dark green to X 1-2.4 mm, branch orders 6, the second-order greenish-brown-drying,and the inflorescences long branches 6-10, clustered apically, lowest branch to and stout. All flowers examined had variable anther 1.2(-3.5) cm long, color and indumentof all axes as locelli numberin whorls I & II, where the upperlo- on twigs; bracts and bracteolescaducous (not seen). Staminateflowers: pedicels ca. 2.4 X 1 mm, the dicelli were either present,vestigial, or one missing. 44 ametereven throughout;receptacleobconical, ca. 1.2 X 2 mm; tepals coriaceous, ovate, ca. 2.5 X 2.5 mm (inner whorl slightly smaller), at anthesis spreading to recurved,reddish,adaxiallytomentose;stamensof whorls I and II minutely spathulate, the anthers broadlyelliptical retuse, ca. 1 X 1.2 mm, with a few hairs at base, the locelli 4, apical, in a shallow arch, introrse,the glands absent;whorlIII columnar,ca. 1.2 X 0.8 mm, with a few hairs at base, the locelli 4, the upperpair latrorse,the lower pairextrorse,the glands globular,ca. 0.5 mm diam.; whorl IV absent;all stamens reddish;pistillode filiform, ca. 0.6 X 0.2 mm, glabrous.Pistillateflowers: pistil ca. 2.2 X 1.5 mm; ovary ovoid, ca. 1.2 mm long, glabrous.Fruits unknown. FLORA NEOTROPICA visible due to indumentcover; terminalbud plump, 4-7 X 3-6 mm; cataphylls caducous; indumentpubescent, caducous after one flush, the hairs dense, to 1.5 mm long, straight to curved, erect, yellowish. Leaves: petioles robust,1.5-3.5 cm X 2-4 mm, adaxially flattened;blades chartaceous,flat, broadlyelliptic, (8-)14-20(-22) X (4-)7-8(-12) cm; base acute to obtuse, 70-110?; apex acute to rounded,60-130?, ultimatelyacuminatefor up to 2 cm; marginminutely recurved;primary vein above flat to slightly raised, below prominent;secondary veins 7-10 pairs, equidistant, eucamptodromous,above flat, below raised, diverging at 40-60?, (straight to) evenly arching (forked), chordal angle 25-30?, the angle uniform along blade length; tertiaryveins above flat, below raised, random-reticulate;higher-orderveins above Field notes. Trees to 40 m tall and 61 cm diam.; below flat, slightly raised; surface above olive-green buttressesvery slenderand steep; outerbarksmooth, to below brown, light olive-greento yellowish brown; grey-brownwith patchesof grey;innerbarkca. 7 mm indumentabove puberulous,the primaryand secondflecked wood red thick, cream, red-brown; cream, veins tomentose, caducous by next flush, below purple heartwood,wood non-aromatic.Pedicels yel- ary pubescent, the hairs sparse, to 1 mm long, curved, low to reddish;tepals yellow. erect, yellowish, denseron the veins, persistingfor at Distribution (Fig. 7). Found in the states of Acre least two flushes. Staminate inflorescences: meso(Brazil)andPando(Bolivia), nearthe Brazil-Bolivia- tonic, pendulous?,peduncles 6-11 cm long, the hyPeru border, at 100-250 m, in rain forest. Flowers popodia 2-5 cm X ca. 2.8 mm, branchorders3, the August-September,at the beginningof the rainysea- second-orderbranches7-10, dispersed,lowest branch son. to 2 cm long, color and indumentof all axes as on Additionalspecimensexamined.BOLIVIA.PANDO: twigs; bractscaducous (not seen); bracteolespersistPuertoOro,74 kmSWof Cobija,14Aug 1988(c fl),R. T ent or caducous, to 3 mm long, adaxially hairy.Staet al. 123 (K,MO). minate flowers: pedicels ca. 13 X 1.6 mm, the Pennington diametergraduallyincreasingapically;receptacleobRhodostemonodaphnedioica is a very distinctive conical, ca. 1.2 X 4.8 mm; tepals coriaceous, ovate, species. Its mixed indumenton the undersideof the ca. 5 X 4 mm (inner whorl slightly smaller), at anleaves, consisting of both appressed-ascendingand thesis spreadingto recurved,reddishblack, adaxially erect hairs, is unique in the genus. It has relatively stamens of whorls I and II, the anthers tomentose; large flowers intermediatein size between those of R. sessile, chubby, broadlyellipticalretuse,ca. 1.6 X 2.2 sailensis and R. anomala. mm II (whorl smaller), glabrous,the locelli slightly Nectandra dioica was incorrectly synonymized with N. grandis by Kostermans(see discussion of R. 4, apical, in a shallow arch, introrse,the glands absent; whorl III columnar,ca. 2 X 1.4 mm, glabrous, grandis). Rohwer (1986) reinstatedit, addingN. suthe locelli 4, the upper pair latrorse,the lower pair perba as a synonym. extrorse, the glands globular,deeply folded, yellowish, ca. 1.2 mm diam.; whorl IV absent;all stamens reddish black; pistillode minute, ca. 1.2 X 0.8 mm, 9. Rhodostemonodaphne anomala (Mez) Rohwer, sparsely hairy.Pistillateflowers andfruits unknown Mitt. Inst. Allg. Bot. Hamb.20: 83. 1986. Nectan(but see discussion). dra anomalaMez, Jahrb.Konigl. Bot. Gart.Berlin 5: 401. 1889. Type.Brazil. Rio de Janeiro:Petr6pField notes. Largetrees to 20 m tall. Tepalsgreenolis, Alto do Imperador,7 Mar 1881 (6 fl), Glaish white. ziou 13150 (lectotype,here designated:B [photos: B neg. 1232/2; F neg. 3745; NY neg. 8329]; isolectotypes: C, G, K [photo NY neg. 8503], P). Distribution (Fig. 7). Knownfrom type specimen Fig. 9D and a sterile collection both from nearthe city of PeTrees: branchesbasitonic?,in axils of cataphylls; tr6polis in the state of Rio de Janeiroin southeastern twigs terete, 3-5 mm diam.; epidermis black, barely Brazil at ca. 850 m, in remnantsof Atlanticrainforest SYSTEMATICTREATMENT 45 (see discussionfor possible rangeextension).Flowers in March,at the height of the rainy season. of the lamina);in R. anomala the secondaryveins are oriented at a more obtuse angle. Rohwer (in sched.) designatedthe duplicateat B Additional specimens examined. BRAZIL. BAHIA: as the holotype of Nectandra anomala based on its Mun.Una,Mico-LeaoBiologicalReserve,11Mar1993(fr), the first of three duplicates cited by Mez, the being et al. 2357 as (MO)[annotatedRhodostemonodaphne Hage cf. anomalavel sp. aff.,see discussionbelow].RIODEJA- other two being the ones deposited at K and the NEIRO: 1948(st), G6is& Octavio Warmingherbarium(incorporatedin C). Quitandinha, Petr6polis, 110 (MO). One collection from the state of Bahia (Hage et al. 2357) has some featuressimilarto those of the two Rhodostemonodaphneanomala is apparently a known anomala. specimensof Rhodostemonodaphne large tree. The twigs have a large numberof scalethe is in are there remnants fruit, Although specimen leaf scars and the indumentcolor changes on either side of the scales, indicative of a markedrhythmic of anthers attachedto the cupule. These are sessile and thick, with four locelli arrangedin an arch, simgrowth.Its thick, yellowish, densely pubescentto velutinous twigs, and olive-green- to yellowish-brown- ilar to those of R. anomala. From these old anthersit drying leaves are very distinctive,althoughsimilarto is difficult to tell if they were fertile or not, and thus those of R. laxa from the N Andes. For characters the dioecious condition expected of RhodostemonodistinguishingR. anomala from otherspecies of Rho- daphne can not be confirmed.Leaf size, shape,numdostemonodaphneof the Atlanticforest of Brazil, see ber and orientation of the secondary veins, and TableVIII. higher-ordervenationall agree well with R. anomala. Rhodostemonodaphneanomala is very similar to The pedicels are ca. 15 X 5 mm, and graduallyenOcotea spixiana (Nees) Mez in the type of indument, large to form a hemispherical,fleshy, tuberculatecushape and texture of the leaves, and flower size, but pule ca. 23 X 12 mm with an undulatemargin and the two species differ in androecialmorphology-O. persisting tepals; the berry is elliptic, 25 X 17 mm. the of locelli has that spixiana bijugate typical genus. However,the leaves of this specimen are thickerand Vegetativelythey can be distinguishedby the reddish darkerthan those of the known specimens of R. anbrown color of the dry leaves of 0. spixiana and the omala, and the indument consists of shorter and acute angle of the secondaryveins (at least at the base sparserhairs thanthose of R. anomala (butotherwise TABLE VIII Comparisonof the species of Rhodostemonodaphnefrom the Atlantic coast of Brazil R. anomala R. capixabensis R. macrocalyx Habit Largetrees Small trees Small trees Twig diameter(mm) 3-4.5 Ca. 3 1.8-4 Hairs Long, yellow Short,yellow Short,yellow to red Petiole length (cm) 1.5-3.5 0.6-1.7 1.0-2.5 Leaf shape Broadly elliptic Broadly elliptic to obovate Narrowlyelliptic to elliptic Secondaryveins (pairs) 7-10 (3-)4(-5) (4-)6(-9) Inflorescences Branchorders Indument 3 Tomentose 4-5 Puberulousto absent 2-5(-8) Tomentoseto puberulous Bracteoles Caducous Caducous Persistentto caducous Pedicel length (mm) Ca. 13.0 7.0-12.0 Ca. 5.6 Receptacle width (mm) Ca. 4.8 Ca. 2.2 Ca. 2.4 Stamens Sessile, thick Spathulate Filiform Geographicrange (state) Rio de Janeiro,Bahia? Espirito Santo Bahia-Parana 46 it is very similar). Because of these differences and the provenanceof this specimen-ca. 1200 km from Petropolis-I am hesitantto include it in R. anomala. It may be a new species closely relatedto R. anomala, but until furthermaterial is available it can not be described. Alternatively,R. anomala may have once had a distributionthat extended from Petropolisto S Bahia (if not beyond). The diverse Atlantic forest wherethese plantsgrow is unfortunatelydisappearing at an alarmingrate. Of the total forest coverage existing in S Bahia in 1945, today only 6% remains, mostly in small isolated patches (W. Thomas, pers. comm.). 10. Rhodostemonodaphne macrocalyx (Meissner) Rohwer ex Madrifiin, comb. nov. Goeppertia? macrocalyxMeissner,in A. L. de Candolle,Prodr. 15(1): 174. 1864. Ocotea macrocalyx(Meissner) Mez, Jahrb.Konigl. Bot. Gart.Berlin5: 367. 1889. Type. Brazil. Without locality, 1836 (fr), Sellow s.n. [197fide Mez] (holotype: B [photos: B neg. 5268, NY neg. 8382, F neg. 3678]; isotypes: F Fig. 1OA [frag. ex B], HBG [frag. ex B]). Trees: branchesbasitonic,in axils of cataphyllsor basal foliage leaves; twigs terete, 2-4 mm diam.;epidermisbrownishto black; terminalbud plump, ca. 1 X 1 mm; cataphylls caducous; indumenttomentose, caducousby next flush, the hairsdense, to 1 mm long, curved to crisped, erect, brownish.Leaves: petioles slender, 1-2.5 cm X 1.4-2.4 mm, adaxiallyflattened; blades chartaceous,flat, narrowlyelliptic, sometimes lobed!, (6-)7-13(-23) X (1-)2-4(-8) cm; base acute to obtuse, minutelydecurrent,40-60(-100)?; apex attenuateto acute to rounded,30-50(-110)?, ultimately acuminatefor up to 2.5 cm; margin plane; primary vein above flat to slightly raised, below prominent; secondaryveins (4-)6(-9) pairs,equidistant,eucamptodromous,above flat to slightly raised, below prominent, diverging at 40-60?, evenly arching (forked), chordalangle 20-30?, the angle uniformalong blade length; tertiary veins slightly raised, below raised, random-reticulateto scalariform;higher-orderveins above slightly raised, below raised; surface above green to brown,below light brown,often whitish;indumentabove absent,the primaryvein tomentose,below tomentose, the hairs isolated, to 1 mm long, curved, erect, yellowish to yellowish brown, denser on the veins (with inconspicuoushair domatia),persisting for at least two flushes. Staminate inflorescences: mesotonic to acrotonic, erect, peduncles 216 cm long, the hypopodia 1-5 cm X 0.5-2 mm, branchorders2-5(-8), the second-orderbranches4- FLORANEOTROPICA 10, dispersed,lowest branchto 2(-5.5) cm long, color and indument of all axes as on twigs; bracts soon caducous, to 3.5 mm long, adaxially glabrous;bracteoles soon caducous,to 1.5 mm long, adaxiallyhairy. Staminate flowers: pedicels ca. 5.6 X 0.8 mm, the diametereven throughout;receptacleobconical,ca. 2 X 2.4 mm; tepals coriaceous, elliptic to ovate, ca. 3 X 2.5 mm, at anthesis spreadingto recurved,reddish to salmon to brown, translucent,adaxially puberulous; stamens of whorls I and II filiform, the anthers oblong inflexed, ca. 1.8 X 0.44 mm, glabrous, the locelli 4, in two almost superposedpairs,introrse,the glands absent;whorl III filiform, ca. 1.8 X 0.4 mm, glabrous,the anthersoblong, the locelli 4, the upper pair latrorse,the lower pair extrorse,the glands globular to folded, ca. 1 mm diam. (protrudingbeyond outer whorls); whorl IV absent; all stamens reddish; pistillode absent.Pistillateflowers: pistil ca. 2.4 X 1 mm; ovary ovoid, ca. 1.5 mm long, glabrous.Fruits: pedicels to 18 X 3 mm, graduallyenlargingto form the cupule; cupule trumpet-shaped,to 13 X 17 mm, smooth, the margin undulate to straight,tepals persisting to caducous;berry elliptic, to 23 X 14 mm. Field notes. Trees to 15 m tall and 25 cm diam., already flowering when 6 m tall. Tepals cream to greenish.Berry greenish. Distribution (Fig. 7). This species is the most southerly member of the genus. It is found on the Atlantic coast from Bahia to Paranaat 25? S. The plants grow in coastal forest and adjacentmountains from sea level to 800 m (occasionally reaching ca. 1500 m in the Mantiqueirarange). The bulk of the floweringspecimenswere collectedDecember-April, during the main months of the rainy season, with a few flowering collections in the northernpart of the range made as late (early) as September.The few fruitingspecimens have been collected in June-July, at the beginning of the dry season. Additionalspecimensexamined.BRAZIL.BAHIA: rd., ca. 65 km NW of Prado, 18 Sep 1978 Prado-Itamaraju (d fl), Mori et al. 10669 (HBG, K, NY); Guaratinga,Eunapolis, Itabela, 4 Jul 1970 (d fl), T S. dos Santos 885 (HBG, SP). ESPiRITO SANTO:Santa Teresa,EstaqaoBiol- 6gica da CaixaD'agua,14 Aug 1984 (fr),Hoffinann178 (MO). PARAN;AAA:Rio da Divisa, Mun. Guaratuba,14 Mar 1962 (Y fl), Hatschbach8936 (F, HBG, MO, S, SP); Serrinha,Mun. Guaraquecaba,11 Jan 1968 (Y fl), Hatschbach 18272 (AAU, B, C, F X2, G, INPA, K, MO, NY X2, RB, US); Cacatu, Mun. Antonina, 29 Jan 1968 (c fl), Hatschbach 18487 (NY X2, S, US); Serra de Araraquara, Mun. Guaratuba,15 Mar 1696 (d fl), Hatschbach21269 (C, MO); Bocai6va do Sul, Parquedas Lauraceas,24 Jan 1990 (6 fl), Hatschbach & Cordeiro53737 (MO); 28 Dec 1994 SYSTEMATICTREATMENT 47 ' . 't v:: , W r; c,:v~' eea6 A~~~~~~~~~~~~'i 70: 0t:; l~uc,. 05Xi'ii^ 00B0 9^'8 :'1~~~~~~~~~~~~~~~~~~~r ty tO-.h>:-_,;0., *,ES706 ~~:..::. .....:..;. <<B' _:. 0 ~i eo~~kr . ' -"'?li"~~~~~~~~~~"~~~l m~~~~~ PLD:*M 1Q1; lib ~,.z, ' ~ rCC ' ~ ~ "-CC7': :'d -D a swwwo '': ~ n" ':: rt .' - an( ~ ;d ~ aS ~ ~ S~ ' ~ I Cl?C' ?10 r' wo n<a ...................... K ~,.x. '.,, ' >',.x.- 'S .?.~* i~;Lr . ^y:-.I:I: dr .............. ... - "; V ' ^'; t9S''i i ; Ii ; A 1- N_ 0Ln .wow~-6: r /70) (~i~fVUI D. R. s'a:dens . -a/. Day -v"~~~~~~~~~~. ~~~~~~i i museu v<w ~~ r i :; _ . ...... sr , esE . ......:......: A. Rhds?ondpn FiG~~~~~~~~~~~~~~ 10s. w. mcoay s ' SaPO&O 2~~~~~~~~~~~~~~~~~~~ t.7em~lVntu7t.s1Y.lm f0 ff ;00;;000f:;\ g X~~ * : r-^*-. ................. 7. (Prirs rSE; Cv. R.aei/n*i. .fi;f *~~~~~~~~~~~~~~~~~ _ k* k-^ *_ 00:-: - I;: : -';i:0:: -: : ............................. r /02]. (W~kinD B. tRP., caia~?sis * e; pro, _.ielW _ N, . t w O.*4* Oim b rup'P1Pn CLL* ,: SS : sb @ he~*.~rrsau, ? d * _ ::: *,w ,.- .' b~rl WaJ , ,,: ;i s : 'M >'._t* *le i s: t* _ ., *_ze-. yp~tJol i h _ g b I . ..... S {?l, *FA * l_|e ,E 'tiL - ffl rr mg O 'W: _ ,, ~ 0r10 U*A. ~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~ . ...................... .... ...... ............... ... ..... ................ FIG. 10. A. Rhodostemonodaphne macrocalyx (Pereira 714). B. R. capixabensis (Weinberg 102). C. R. avilensis (Meier 1790). D. R. scandens (Daly et al. 3837). 48 FLORANEOTROPICA (d fl), Hatschbach & J. M. Silva 61392 (MO). Rio DEJANEIRO:Sumar6-Tijucard., summit of Serra da Carioca, 1 Feb 1929 (d fl), Bandeiras.n. (F X2, G, RB); Rio de Janeiro (9 fl), Binot 110 (M); Tijuca (d fl), Boissier 12 (G); Vista Chinesard., Gavea, 15 May 1929 (cd fl), Duarte & Bandeira 263 (F, RB X2); Rio de Janeiro,Redentorrd., 10 Jul 1961 (d fl), Duarte 5605 (F, INPA, K, RB); Petr6polis(st), Glaziou 822 (C); Rio de Janeiro(6 fl), Glaziou824 (C, K, NY); (9 fl), Glaziou 1290 (C, K); Petr6polis,2 Feb 1880 (9 fl), Glaziou 12119 (A, B [photo B neg. 7714], BM, C X2, F x3, G, K, NY, ); 20 Apr 1880 (9 fl), Glaziou 12128 (A, B [photoB neg.7713],C, F, G, K, NY X2); 19 Jan1884(d SerradosOrfl), Glaziou15367(C, G, K, NY);Petr6polis, gaos, 7 Apr 1891 (d fl), Glaziou 17742 (B, C X2, F, G, K x2, US); Serrade Maca6 (6 fl), Glaziou 18454 (A, BM, C, G, K, NY); Tijuca?(fr), Grahams.n. (K); Gravata,Pai Ricardo, 20 Dec 1926 (6 fl), Horto-Florestal(RB)102874 (F, NY); 30 Dec 1927 (d fl), Horto-Florestal(RB)110103 (F, NY);VistaChinesard., 1 Jul 1958(fr),Lieneet al. 3970 (HBG);Rio de Janeiro,Corcovado(st), Nadeauds.n. (P); rd.,summitof SerradaCarioca,22 Jun1978 Sumar6-Tijuca (9 fl), Occhioni8580 (B);Petr6polis,Serrados Orgaos,4 Apr 1953 (9 fl, fr), E. Pereira 714 (F, RB); Sumar6,23 Feb 1959 (d fl), E. Pereira & Duarte 4481 (HB); 24 Feb 1959 ( fl),E. Pereira&Duarte4520(HB,NY);RiodeJaneiro, Redentorrd. (J fl), E. Pereira & Duarte 4535 (B, F, HB, fideMez](d RB);(9 fl),4536(HB);"Brasilia" [Esperanqa fl), Riedel s.n. [770 fide Mez] (B, G X2, K); Teres6polis, FazendaBoa F6, PicadaDavis,9 Apr1943(d fl), Velloso 382 (HBG). SAO PAULO:Serra da Mantiqueira,16 Mar 1939 (6 fl), M. Kuhlmann& Gehrt (SP)40043 (G). Local names. Canelo batallia, canela cedro. Rhodostemonodaphne macrocalyx is character- ized by its flowerswhich have elongated,filiformstamens, inflexed antherswith introrseto latrorsedehiscence, and very large glands of whorl III which protrudebeyond the outer whorls. In all these charactersit resembles various species of the genus Pleurothyrium.However,it appearsto be most closely related to Rhodostemonodaphne capixabensis also from the Atlantic coast of Brazil (see TableVIII). There is a markednorth-southvariationgradient, most pronounced on the southern extratropicalextreme.In the northernmostpartof the range,two collections from the state of Bahia (Mori et al. 10669 and T S. dos Santos 885), have larger, narrowly ellip- tic, long-acuminateleaves, with few, branchedsecondary veins, and profusely branchedinflorescences, allowing the distinction with the sympatricRhodos- tiqueira range in the state of Sao Paulo is indistinguishable from the collections from Rio de Janeiro. Furthersouth the various collections by Hatschbach differfrom the rest in a numberof characters.Hatschbach 8936 has largelong-acuminateleaves andmuchbranchedinflorescencessimilarto the specimensfrom the northernextremeof the range. Two inland specimens from Bocaiiva do Sul (Hatschbach& Cordeiro 53737 and Hatschbach & J. M. Silva 61392), have smallerleaves with a high numberof secondaryveins (to 8). Three coastal specimens collected on mountain slopes (Hatschbach18272, 18487, and 21269) differ from the rest in having widely elliptic leaves, with more obtuse bases and apices, and conspicuously branched secondary veins. The collections are unusual in thatmost of the leaves show varyingdegrees of lobing, associated with the forked secondaries, ranging from a slight apical asymmetryto one conspicuous lateral lobe. This lobing of the leaves is known only in two distantgenera of Lauraceae(i.e., Sassafras and Lindera). These specimens may represent a separateentity from the core of Rhodostemonodaphnemacrocalyx.One collection from Espirito Santo, . A. Silva 345, is here postulated as a hybrid between R. capixabensis and R. macrocalyx (see discussion underR. capixabensis). Vegetatively Rhodostemonodaphnemacrocalyx resemblesEndlicheriapaniculata (Spreng.)Macbr.,a species inhabitingthe same general area. The indument and leaf shape and size are very similar.They can be distinguishedby the fewer, more acute secondaryveins and the two-locellate anthersof E. paniculata. Mez (1889) includedGoeppertiareflectens(Nees) Meissner in the synonymy of Ocotea macrocalyx (Meissner)Mez; Meissnerhad misidentifiedGraham s.n. as that species. However,Grahams.n. belongs to Rhodostemonodaphnemacrocalyx.An invalid name, "Mespilodaphnefloribunda Meissner,"anothersynonym cited by Mez in the same work and attributed to manuscriptannotationsin variousherbaria,can be referredto the specimenRiedels.n. (given the number 770 by Mez), which I have seen annotatedby Meissner as such (cf. G [2 sheets], and K); it belongs to R. macrocalyx. temonodaphne capixabensis (see discussion under R. capixabensis). The majorityof the specimens examined come from the state of Rio de Janeiro.In general they have small, elliptic to narrowlyovate, or attenuate leaves, and inflorescences with few orders of branching.The only known specimen from the Man- 11. Rhodostemonodaphne capixabensis Baitello & Coe-Teix.,RevistaBrasil.Bot. 14:79, fig. 1. 1991. Type. Brazil. Espirito Santo:Villa Velha, Interlagos II, Oct 1980 (9 fl, fr), Weinberg102 (holoFig. 1OB type: SPSF; isotype: VIES-n.v.). SYSTEMATICTREATMENT Trees to shrubs: branches basitonic, in axils of cataphylls;twigs angularandremainingso for at least two flushes, ca. 3 mm diam.;epidermisbrownish;terminalbudplump,ca. 5 X 3 mm;cataphyllscaducous; indumenttomentose,caducousby next flush,the hairs dense, to 0.8 mm long, curved, erect, brownish. Leaves: petioles robust, 0.6-1.7 cm X 1.2-1.8 mm, adaxially flattened;blades chartaceous,flat, broadly elliptic to obovate, (4-)7-9(-11) X (2-)3-6 cm; base obtuse to rounded,70-110?; apex obtuse to rounded, 90-140?, (mucronate);margin plane; primary vein above flat to slightly raised, below prominent;secondary veins (3-)4(-5) pairs, equidistant,eucamptodromous, above flat, below raised, diverging at 5055?, abruptlyarching near margin (forked), chordal angle 20-25?, the angle uniform along blade length; tertiary veins above flat, below slightly raised, random-reticulate;higher-orderveins above flat, below slightly raised; surface above olive-green to greenishbrown,below light green to brown,conspicuously glaucous;indumentabove absent,the primary vein puberulous,below puberulous,the hairs sparse to isolated, to 0.3 mm long, straightto curved,erect, yellowish, caducous afterone flush. Staminate inflorescences: mesotonic to acrotonic, erect, peduncles 7-13 cm long, the hypopodia 3-7 cm X ca. 1 mm, branch orders 4-5, the second-orderbranches 4-6, dispersed, lowest branch to 4(-7) cm long, all axes darkbrown, puberulousto glabrous;bractsand bracteoles caducous (not seen). Staminateflowers: pedicels 7-12 X ca. 0.8 mm, the diametergraduallyincreasing apically; receptacle obconical, ca. 2 X 2.2 mm; tepals chartaceous,ovate, ca. 2.5 X 2 mm, at anthesis erect to spreading,yellowish brown, translucent, adaxiallypuberulous;stamensof whorls I and II spathulate,the anthersoblong inflexed, ca. 1.6 X 0.8 mm, glabrous, the locelli 4, apical, in a shallow arch, introrse, the glands absent; whorl III filiform, ca. 2.4 X 0.8 mm, glabrous,the anthersoblong, the locelli 4, the upper pair latrorse,the lower pair extrorse, the glands globular,ca. 1.5 mm diam.; whorl IV absent;all stamensreddish;pistillode filiform,ca. 1.5 X 0.3 mm, glabrous.Pistillate flowers: pistil ca. 2.5 X 1.2 mm; ovary ovoid, ca. 1.5 mm long, glabrous. Fruits: pedicels to 15 X 3 mm, graduallyenlargingto form the cupule;cupule trumpet-shaped,to 13 X 20 mm, smooth, the marginstraight,tepals caducous; berry elliptic, to 25 X 20 mm. 49 Distribution (Fig. 7). This species is known from only three localities in the state of Espirito Santo on the Atlanticcoast of Brazil. It grows on sandy,coastal forest (restinga)to 50 m. Flowers September-October, at the beginning of the rainy season. Fruits are ripe February-March. Additional specimens examined. BRAZIL. ESPiRITO SANTO:Linhares,CVRD Forest Reserve, 22 Oct 1981 (d 21 Sep 1982 fl), Folli334 (MOx2); VillaVelha,Interlagos, (d fl), O.J. Pereira& Fontenella181 (SPSF,VIES-n.v.); VillaVelha,Jacaranema, 21 Oct 1987(9 fl, immfr), O.J. Pereira1205 (SPSF,VIES-n.v.);Conceicaoda Barra,27 Feb 1994 (fr), O.J. Pereiraet al. 2883 (SPSF);Linhares, CVRDForestReserve,23 Sep 1982(9 fl), I. A. Silva345 I, (CVRD-n.v.,MO X2, SPSFX2);VillaVelha,Interlagos 01-A(MBML-n.v., Nov 1980(d fl), Weinberg MO). Local names. Canelado nativo,oliveira-da-praia. Rhodostemonodaphnecapixabensis has broadly elliptic, often mucronate,stronglychartaceousleaves that are glaucescent abaxially,and slender,profusely long-branched,glabrous inflorescences, unlike any other species in the genus. It can be distinguished from the only other species of Rhodostemonodaphne with overlappingrange,the apparentclose relativeR. macrocalyx,by a numberof vegetativeandandroecial characters.In the northernpartof its rangeR. macrocalyx has elliptic to narrowlyelliptic, long-acuminate leaves. The stamensin R. capixabensisarespathulate, with the locelli arrangedin an arch, while in R. macrocalyx they are filiform with bijugate anthers(see TableVIII). The paratypeI A. Silva 345 may in fact be a hybrid betweenRhodostemonodaphne capixabensisand R. macrocalyx.It differsfrom all the othercollections of R. capixabensis in a variety of characters,these being intermediatebetween the two species. The indumentis darkgolden to reddish;the leaves are narrower, being elliptic, and are not glaucous; the secondary veins are oriented at a shallower angle (ca. 30?), and the apex of the laminahas a longer mucro. The stamens are nevertheless spathulate, agreeing with those of other collections of R. capixabensis. Due to the lack of matureflowerson the only specimen availableto me of O. J. Pereira 1205 (SPSF), I could not confirm the observation by Baitello and Coe-Texeira(1991) that this plant had at least two flowerswith staminatecharacters.While it is common to find pistillate individualswith well-developed deField notes. Trees to shrubsto 8 m tall and 12 cm hiscing anthers,I have never seen pollen within the diam., already flowering when 4 m tall; outer bark locelli of these anthers, or even after forcing open rough; leaves below glaucous (bluish). Inflorescence anthers that have not dehisced. Although there are axes greenish;receptaclepurple;tepals reddish.Cu- reportsof findingboth staminateandpistillateflowers in one individualof an otherwisedioecious species of pule red; berry greenishbrown. 50 Lauraceae,this conditionis extremelyrare(cf. Ocotea tenera Mez & J. D. Sm.; Wheelwright & Bruneau, 1992). Rhodostemonodaphnecapixabensis is one of the few species for which informationon its conservation status is available.Not only does this species have a limited distributionrange, but it also seems to be restricted to a very specific habitat.In both the collection labels and the species description (Baitello & Coe-Teixeira, 1991) it is mentioned that the sandy coastal forests on which it grows are being depleted at an alarmingrate primarily due to housing development projects. For the above reasons it should be considered as a species threatenedwith extinction, and conservationefforts should be encouraged.The authorsalso mentionWeinberg'ssuggestion(in Weinberg, 1984, unavailableto me) that the species is of horticulturalinterest for its decorativequalities as a small shade tree, with shiny, greyish leaves and colorful infructescences. FLORA NEOTROPICA dulous?, peduncles 1-5 cm long, the hypopodia 1-3 cm X ca. 0.5 mm, branchorders3, the second-order branches2-4, clusteredapically,lowest branchto 1.2 cm long, color and indumentof all axes as on twigs; bractsand bracteolescaducous(not seen). Staminate flowers: pedicels ca. 4 X 0.4 mm, the diametereven throughout;receptacle obconical, ca. 1.6 X 2 mm; tepals chartaceous,ovate, ca. 1.5 X 1 mm, at anthesis spreading,reddish black, adaxially puberulous;stamens of whorls I and II spathulate,the anthersroundish, ca. 1.2 X 0.8 mm, glabrous,the locelli 4, apical, in a shallow arch, introrse,the glands absent;whorl III capitate,ca. 1.6 X 0.8 mm, glabrous,the anthers globose, the locelli 4, the upperpairlatrorse,the lower pair extrorse (the upper almost introrse),the glands globular,ca. 0.6 mm diam.; whorl IV absent;all stamens yellowish brown;pistillode teardrop-shaped, ca. 1.5 X 0.6 mm, glabrous.Pistillate flowers: pistil ca. 1.5 X 0.8 mm; ovary ovoid, ca. 1.2 mm long, glabrous. Fruits unknown. Field notes. Shrubs to 5 m tall and 9 cm diam. Inflorescenceaxes purple;tepals yellowish green. 12. Rhodostemonodaphne avilensis Madriinain, sp. nov. Type. Venezuela.Dtto. Federal:El Avila National Park, El Avila ridge, 3 Mar 1992 (d fl), Meier 1790 (holotype:MO). Fig. 10C Distribution (Fig. 7). Both collections are from the CordilleraEl Avila, a mountain range between Caracasand the Caribbean,at ca. 1900 m, in cloud forest. Flowers in March,during the dry season. foliorumR.elephantopodi accedit Formaet magnitudine Additional specimen examined. VENEZUELA. sed staturaminore,venationefoliorumconspicua. El Avila National Park,El Avila ridge, 3 DTTO.FEDERAL: Mar1992(9 fl), Meier1797(MO). Shrubs: branchesbasitonic to mesotonic, in axils of cataphylls or basal foliage leaves; twigs angular, Etymology. Named after the "El Avila" National soon becoming terete, ca. 2 mm diam.; epidermis Park. black;terminalbud slender,ca. 4 X 1 mm; cataphylls Rhodostemonodaphneavilensis is a small tree to caducous;indumentsericeous to absent,caducousby large shrub,with a dense, shiny green foliage, purple next flush, the hairs sparse, to 0.3 mm long, straight, peduncles, and yellowish tepals. The small leaves appressed,ascending,silver.Leaves: petioles slender, with cuneate bases, obtuse apices, and conspicuous 0.7-1 cm X 0.7-1.2 mm, adaxially flattened;blades venation are quite characteristic.In leaf shape and chartaceous,flat, narrowlyelliptic to obovate, 2-9 X size R. avilensis is similarto R. elephantopus,butthat 1-4 cm; base acute, minutelydecurrent,40-70?; apex species is a large lowland forest tree, and has inconobtuse to rounded, 70-130?; margin plane; primary spicuous venation. Furthermore,the inflorescences vein above slightly raised, below raised; secondary and flowers of each species suggest differentrelationveins 4-10 pairs, equidistant, brochidodromous, ships, those of R. avilensis andR. capixabensisbeing above and below slightly raised, divergingat ca. 80?, similarwhile those of R. elephantopusandR. kunthistraight,the angle uniformalong bladelength;tertiary ana show a partialresemblance. veins above and below slightly raised, randomThe leaves of R. avilensis are similar to those of reticulate; higher-order veins above and below Ocotea congestifolia Lasser from the same general slightly raised; surface above shiny greenish brown, area.The leaves of 0. congestifoliaare,however-as below dull greenish brown; indumentabove absent, the epithet implies-clustered at the tips of the twigs below minutely sericeous, the hairs isolated, to 0.6 and less rigid than those of R. avilensis. The low statureof the plant, with its dense, evermm long, straight, appressed, ascending, silver, denser on the veins, caducous by next flush. Stami- green, shiny foliage, and contrastingcolors of its innate inflorescences: along whole lengthof flush,pen- florescences and (presumably) infructescences, are SYSTEMATICTREATMENT 51 spreading, reddish brown, adaxially glabrous; stamens of whorls I and II, the antherssessile, chubby, trapezoid,ca. 2 X 2.4 mm (whorl II slightly smaller), glabrous, the locelli 4, apical, in a shallow arch, introrse,the glands absent;whorl III columnar,ca. 2 X 1.2 mm, glabrous,the locelli 4, the upperpairlatrorse, 13. Rhodostemonodaphne scandens Madriinain, the lower pair extrorse,the glands often fused to adBrittonia48: 63, fig. 9. 1996. Type. French Gui- jacent glands and stamensof outerwhorls, forminga ana. Mont Bellevue de l'Inini, 3 Sep 1985 (d fl), continuousring, ca. 1.2 mm diam.; whorl IV absent; de Granville,Allorge, Cremers,Gorts-vanRijn & all stamensreddish;pistillode absent (minute,linear, Kodjoed-Bonneton8023 (holotype: U; isotypes: ca. 0.3 mm long, hairy). Pistillate flowers unknown. B-n.v., CAY-n.v.,MO, P-n.v., US). Fig. 1OD Fruits: pedicels to 13 X 3 mm, graduallyenlarging to form the cupule;cupule hemispherical,to 12 X 14 Scandent shrubs: with long, slenderand sparsely mm, smooth (chartaceous),the marginundulate,tebranchedshoots; branchesbasitonic, in axils of catpals persisting;berry elliptic, to 12 X 15 mm. aphylls, pseudo-verticillate;twigs terete, 2-3 mm Field notes. Scandent shrubs, sprawling over diam.; epidermis brownish,barely visible due to indument cover; terminalbud plump, ca. 4 X 5 mm; neighboringvegetation,to 3-5 m tall and 5 cm diam. cataphyllspersistingon currentflush (and for at least (trunkrhomboidin cross-section);buttressesabsent. three flushes), to 6 mm long; indumentpubescent, Tepals whitish. Cupule red. persistingfor at least two flushes, the hairs dense, to Distribution (Fig. 11). FrenchGuianaandthe ad1.2 mm long, straight, erect, reddish. Leaves: jacent state of Amapa,Brazil, and Guyana,at 50-750 (pseudo-verticillateat flush apex); petioles pulvinate, m, and growing in non-inundatedmoist forest. Flow0.4-0.7 cm X 2-3.2 mm, terete; blades coriaceous, ers August-October,concurrentwith new vegetative flat to bullate, narrowly elliptic, (9-)13-19(-28) X flushes, during the dry season. Immaturefruits pres(4-)6-9 cm; base acute to obtuse, 50-110?; apex ob- ent late October, by maturingin Decemberat the betuse, 50-110?, ultimately acuminate for up to 1.5 ginning of the rainyseason; old cupules can be found (-2.4) cm; margin minutely recurved;primary vein as late as May. above slightly raised and rounded,below raised and Additionalspecimensexamined.GUYANA.RUPUlongitudinally furrowed; secondary veins (7-)8-11 NUNI: IwokramaReserve, 22 Nov 1995 (imm fr), Clarke (-12) pairs, indistinguishablefrom tertiary veins at 647b (MO-n.v.); KanukuMts., Two Head Mtn., 25 Jan1994 base and apex of lamina, equidistant,brochidodrom(9 fl, fr), Jansen-Jacobset al. 3364 (MO, U); Iwokrama ous, above and below slightly raised to inconspicu- reserve,Mt. Daniel, 27 Feb 1995 (fr), Mutchnick945 (MO, ous, diverging at 60-70?, straight,the angle uniform NY, US-n.v.). along blade length; tertiary veins above and below FRENCH GUIANA: Crique Arataye, Nouragues stainconspicuous to slightly raised, random-reticulate; tion, 11 Oct 1988 (st), Riera 1710 (CAY,MO, P, VI). BRAZIL. AMAPA:Macapd,Rio Araguari,12 Dec 1984 higher-orderveins above and below inconspicuous; surfaceabove olive-greento brown,below brown;in- (fr),Dalyet al. 3837(GH,MO,NY);Macapa,122kmNW dumentabove absent,below tomentoseto puberulous of P6rto Grande, 1 Jan 1985 (fr), Daly et al. 4032 (MO, or glabrous,the hairs sparse,to 0.5 mm long, straight, NY); Oiapoque-Calcoenerd., 17 km SSE of Oiapoque, 3 Dec 1984 (fr), Mori et al. 17169 (MO, NY); Macapa, Rio erect, reddish brown to yellowish brown, persisting 30 Sep 1961 (fr), Pires et al. 51370 (NY); Araguari, for at least two flushes. Staminate inflorescences: in rd., km 4, 22 Sep 1957 (c fl), A. Silva axils of foliage leaves of current flush or previous Oiapoque-Caloene 486 (NY). flushes, along whole length of flush (in axils of catRhodostemonodaphnescandens, although a very aphylls), pendulous,peduncles 1-10 cm long, the hypopodia to 3 cm X 1-1.8 mm, branchorders 2(-3), distinct species on accountof its scandenthabit,bulthe second-orderbranches 5-11, dispersed, lowest late leaves, andlargeflowers,sharesa numberof charbranchto 0.2 cm long, color and indumentof all axes acters with R. licanioides and R. miranda(for distinas on twigs; bractspersistent,to 4 mm long, adaxially guishing featuressee Table IX). In the discussion in the originalpublication(Madglabrous;bracteolespersistent,to 3 mm long, adaxially glabrous.Staminateflowers: pedicels ca. 1.5 X rifian, 1996), I suggested that the five species from 1.1 mm, the diametereven throughout;receptacleflat, the Guayanahighlands with scandenthabit (R. celica. 0.8 x 3.3 mm; tepals chartaceous,ovate, ca. X 5 ana, R. steyermarkiana,R. leptoclada, R. scandens, 3.5 mm (inner whorl slightly smaller), at anthesis and E. vinotincta)may constitute a group of closely characteristics of high ornamental value. Efforts should be made to bring this plant into cultivation, where it may serve as a beautiful garden plant for high-elevationcities. 52 FLORA NEOTROPICA 3X a)X0 _ ?o ~ ~ ~ ~ ~ ~ .0. FIG. 11. Ditribuion ma of prt ofthe spcies n theRhodostemonodaphne , l~~~~I 'allfM m.t 11? \it MD ,,~~~ ~ scandens groupl 0 g o R. cuncuiarensis X R. licanioides.... *R. mr * R. nges A R. revolutifolia-* R-scei-dens V R. tumucumaquensis FIG. 1 1. Distributionmap of partof the species in the Rhodostemonodaphne scandens group.Each symbol represents one or more collections found in a degree-square. relatedspecies. After having looked at the genus as a whole, it seems more likely that the scandent habit shared by these species is the result of convergence ratherthanindicatingcommon ancestry(see "Species Groups"). The Guyanan specimens cited here have been made available to me after the original publication of the species. They are similar in their bullate leaves to the specimens from the Brazilian state of Amapa'and extend the range of the species considerably. 14. Rhodostemonodaphne licanioides (A. C. Sm.) Madrinian,comb. nov. Ocotea licanioides A. C. Sm., Bull. TorreyClub 58: 107. 1931. Type. Peru. Loreto:Mishuyacu,nearIquitos,May 1930 (d fl), Klug 1506 (holotype:NY; isotype: US). Fig. 12A Endlicherialoretensis0. C. Schmidt,Repert.Spec. Nov. Regni Veg. 31: 178. 1933. Type. Peru. Loreto:Mis- huyacu, near Iquitos, Apr 1930 (d fl), Klug 1258 (holotype:F-n.v. [photos:B neg. 7712; G neg. no.?]; isotypes: B [frag. ex F], G [frag.ex F], NY). Shrubs to scandent shrubs: with long, slenderand sparselybranchedshoots; branchesbasitonic,in axils of cataphyllsor foliage leaves; twigs terete, 3-4 mm diam.; epidermis brownish, barely visible due to indument cover; terminal bud plump, ca. 6 X 3 mm; cataphyllspersistingon currentflush (and for at least three flushes), to 6 mm long; indumentpubescent, persisting for at least two flushes, the hairs dense, to 0.8 mm long, straight,erect, brownish.Leaves: petioles robust,0.5-1.5 cm X 0.3-0.5 mm, terete;blades coriaceous to chartaceous,flat to bullate,elliptic, 1532 X 7-13 cm; base obtuse to rounded(almost cordate), (80-)120-140(-180)?; apex obtuse to rounded, (70-)100-140?, acuminate for up to 3 cm; margin plane to recurved;primaryvein above raised, below prominent; secondary veins (7-)9-11(-14) pairs, equidistantto closer together towardsthe apex, bro- SYSTEMATICTREATMENT 53 TABLE IX Comparisonof Rhodostemonodaphnelicanioides, R. miranda, and R. scandens R. licanioides R. miranda R. scandens Habit Shrubsto small trees with Shrubs droopingbranches Scandentshrubs Leaf shape Elliptic to oblong Oblanceolate Narrowlyelliptic Leaf base Obtuse to roundedto inconspicuously cordate Cordate Obtuseto rounded Laminaundulation Flat to bullate Undulate Flat to markedlybullate Secondaryveins, angle of Obtuse divergencew/ primary Basalmostpair acute, be- Obtuse to almost perpencoming obtuse apically dicular Indumentbelow Tomentoseto pubescent Absent Tomentoseto pubescentor absent Inflorescences Paniculate Racemose Racemose Anthers Stipitate Spathulate,laminar Sessile, thickened Locelli arrangement Almost bijugate In a shallow arc Horizontally Geographicrange Loreto, Peru Guyana Guyanato Amapa',Brazil chidodromous, above impressed, below raised, diverging at 55-65', evenly arching,chordalangle ca. 300, lowest pair more obtuse than rest; tertiaryveins above flat, below slightly raised, random-reticulate; higher-order veins above inconspicuous, below slightly raised; surface above shiny brown, below olive-green to brown, the veins lighter; indument above absent,the primaryvein basallytomentose,below tomentoseto pubescent,the hairs sparse,to 1 mm long, straight,erect, yellowish brown, denser on the veins, persisting for at least two flushes. Staminate inflorescences: basitonic, erect?, peduncles 5-19 cm long, the hypopodia 1-8 cm X ca. 1 mm, branchorders 4, the second-orderbranches to 14, dispersed, lowest branchto 0.5-3.5 cm long, color andindument of all axes as on twigs; bracts persistent, to 4 mm long, adaxially glabrous; bracteoles persistent, to 2 mm long, adaxiallyglabrous.Staminateflowers:pedicels ca. 2.5 X 0.7 mm, the diametereven throughout; receptacleobconical, ca. 0.8 X 1.8 mm; tepals membranaceous,elliptic, ca. 1.6 X 1.3 mm, at anthesis spreading, reddish, adaxially glabrous; stamens of whorls I and II spathulate,laminar, the anthers oblong, ca. 0.8 X 0.8 mm, glabrous,the locelli 4, in two almost superposedpairs, introrse,the glands absent; whorl III columnar,ca. 1.4 X 0.5 mm, with a few few hairs at base, the anthersoblong, the locelli 4, the upperpair latrorse,the lower pair extrorse,the glands globose, ca. 0.8 mm diam.; whorl IV absent; all stamensreddish;pistillode absent.Pistillateflowers: pistil ca. 1.6 X 0.6 mm; ovary ovoid, ca. 0.8 mm long, glabrous.Fruits: pedicels to 10 X 3 mm, often thickening well below the bracteoles, graduallyenlargingto form the cupule;cupule trumpet-shaped,to 1 X 1 mm, smooth, the marginundulateto straight, tepals persistingor caducous;berry elliptic, to 3 X 2 mm. Field notes. Small trees to shrubs(with drooping branches, scandent?) to 2-4(-10) m tall; bark unknown. Tepals cream to white. Cupule red; berry green, ripeningblack. Distribution (Fig. 11). Rhodostemonodaphnelicanioides is endemic to the Departamentoof Loreto in PeruvianAmazonia. All known collections come from a small areaalong the Nanay rivernearthe town of Iquitos. Growingin non-inundatedprimaryforest, at ca. 100 on white sand ("varillal,""campinarana"), m. Flowers February-June,at the height of the rainy season; fruitsJuly-November(fruitscan be found on the plants as late as the following flowering season), duringthe dry months. FLORA NEOTROPICA 54 14?~~~~~~~~~~~~~~~~~N 3;1816 _ _ 79 ^s5r13 .. _~~~~~~~~~~~~~~~~~~~~~ .~~~~~~~0 -A"r|s A,' _S_0~~~~~~~~~~~~~~~~~~~~~~ 13 w"..0 ' S -Am-0. iSV s: a.,e *O n*i~~~~~~~~~~~~~~~~~~~VTA ~~~~~~~~~~~~~~~~~~cs 71 4' NV ;235'89 '0~~~~~~~~~~I _ f " '' ,0~~~~~~~~~~ ' ~~~~~~~~~~~~~~~~~~~~~~~ w ui'''. FIG tioi (Sbte 1. A. 350. n Rhdstemooap Dw ... R. ceaicup Licnoie la(Pane et ,;sqe et a'l. 2115 al 64., B. R . miad;Fnh w 283. *C.' ). R.< i _t rv u- --2' SYSTEMATICTREATMENT Additional specimens examined. PERU. LORETO: Mishuyacu, near Iquitos, Feb 1932 (d fl), Klug 2556 (A, NY, S, US); Maynas, Iquitos, PeniaNegra rd., 9 Nov 1982 (fr), Rimachi 6430 (US); Maynas, Iquitos, Varillal rd., Jul 1985 (fr), Rimachi 7849 (US); Maynas, Rfo Nanay, 26 Oct 1980 (fr), R. Vdsquezet al. 644 (F, MO); Maynas, Recreo, Maniti, 17 Oct 1983 (fr), R. Vdsquez& N. Jaramillo 4488 (F, MO); Maynas, Mishana, Rio Nanay, 21 Jul 984 (fr), R. Vasquezet al. 5296 (MO, NY); Maynas,Iquitos,Rfo Nanay, Puerto Almendras, 19 Feb 1985 (fr), R. Vdsquez& N. Jaramillo 6222 (NY); 3 Jan 1987 (fr), R. Vdsquez & N. Jaramillo 8727 (HBG, MO); Maynas,Iquitos,Rfo Nanay,Nina rumi, 6 Mar 1987 (d fl), R. Vdsquez& N. Jaramillo 8929 (MO); Maynas,Iquitos-Nautard., 13 Apr 1988 (fr), R. Vdsquez & N. Jaramillo 10578 (MO); Maynas, Alpahuayo, 3 Nov 1990 (fr),R.Vdsquez& N. Jaramillo14533 (MO);Maynas, Mishana, Rfo Nanay, 18 Aug 1988 (fr), van der Werif et al. 10188 (HBG, MO); Manfinfaon the upperRio Nanay, Jun 1929 (d fl), Williams1118 (NY). Local name. Muena (= moena) negra. Rhodostemonodaphne licanioides is a very distinctive species, yet is most similar to R. scandens and R. miranda, two species from the Guianas (for differentiating characters see Table IX). The three species share a number of vegetative characters. All are small shrubs (one report of "drooping branches" in R. Iicanioides may be indicative of a scandent habit). The persisting cataphylls in the three species are very characteristic. The leaves, although different in size, are very similar in shape and texture; both R. licanioides and R. scandens can have flat to bullate laminas, while the lamina of R. miranda is undulate. Some specimens of R. licanioides have nearly cordate bases similar to those of R. miranda. One distinctive anatomical character, the mesophyll consisting of a lax aerenchyma, is also present in all three species. The inflorescences and flowers of the three species are nevertheless very different. The only inflorescences of R. scandens seen are little branched and have few, large flowers with fleshy sessile anthers characteristic of the genus Rhodostemonodaphne. The inflorescences of R. miranda are also small and little branched, but the flowers are smaller and the stamens spathulate and laminar. The inflorescences of R. Iicanioides are, longer, lax, profusely branched (paniculate), and have small flowers and stipitate stamens with large anther cells. The inflorescence morphology of R. licanioides resembles that of species of Endlicheria subgen. Ampelodaphne, this being the reason why 0. C. Schmidt described it as Endlicheria loretensis. The type of Endlicheria loretensis and one other specimen (Klug 2556), both have leaves with a thinner, membranous, strongly bullate lamina and longer, laxer inflorescences than those of the other 55 specimensseen. Based on the extremevariabilityseen in the two other scandentspecies, where completely flat coriaceous leaves as well as distinctly bullate, membranaceousones can be found in the same species, I am considering all these specimens to belong to the same species. Furtherstudy may indicate that they belong to two separate species, whereuponE. loretensis would become the basionym of the new combination. The anthers of all flowering specimens seen are four-locellate, with the locelli in almost superposed pairs, much like those found in many species of Ocotea. Schmidtwas well awareof the four-locellateanthers, but supported its placement in Endlicheria based on the fact that in at least one species in that genus, Endlicheriaanomala (Nees) Meissner,the anthers of whorl III are four-locellate. This species could well be placed in either of the two generato which it has previouslybeen associated, Ocotea and Endlicheria,and it standsout in Rhodostemonodaphnemainly because of its inflorescence morphology. Initially, I was inclined to follow Schmidt and place it in Endlicheria, where a new combinationwould have to be made, but I refrained from doing so because of its overall vegetative similarity to Rhodostemonodaphnescandens and R. miranda. I excluded it from Ocotea for the same reason. 15. Rhodostemonodaphne miranda (Sandwith) Rohwer,Mitt. Inst. Allg. Bot. Hamb.20: 85. 1986. Nectandra miranda Sandwith, Kew Bull 7: 256. 1952. Type. Guyana.Mazaruni-Potaro:Mazaruni River, IssineruCreek, 4 Mar 1949 (Y fl bud, fr), Fanshawe 2863 = FD 6010 (holotype: K [photo NY neg. 8491]; isotypes: FDG, K X2). Fig. 12B Shrubs to scandent shrubs: with long, slenderand sparselybranchedshoots; branchingunknown;twigs terete, ca. 4 mm diam.; epidermisbrownishto black, barely visible due to indument cover; terminalbud plump,ca. 3 X 2 mm;cataphyllspersistingon current flush, to 6 mm long; indumentpubescent,persisting for at least two flushes or caducous after one flush, the hairs dense, to 1.5 mm long, straightto curved, erect, reddish.Leaves: petioles pulvinate,ca. 8 cm X 3.5 mm, terete; blades thinly chartaceous,flat, narrowly elliptic to obovate, 17-29 X 6-10 cm; base cordate,ca. 2300; apex acute, ca. 700, ultimatelyacuminate for up to 1.7 cm; marginplane; primaryvein above slightly raised, below raised; secondaryveins 6-8 pairs, equidistant (except basalmost two pairs, which are very close together and distant from the rest), eucamptodromous,above slightly raised,below 56 raised, diverging at 50-60', evenly arching, chordal angle 20n30?, lowest pair more acute than rest (ca. 300); tertiaryveins above flat, below slightly raised, random-reticulateto scalariform;higher-orderveins above inconspicuous, below slightly raised; surface above brownish green, below light brown; indument above and below absent, the veins with sparse, appressed, ascending, to 1 mm long, brown hairs. Staminateplant unknown.Pistillate inflorescences:basitonic, from axils of cataphylls before elongation of new buds, pendulous?,peduncles 6-10 cm long, the hypopodia 3-4 cm X ca. 1.5 mm, branchorders2(3), the second-orderbranches10-14, dispersed,color andindumentof all axes as on twigs; bractspersistent, to 6 mm long, adaxially glabrous;bracteolespersistent, to 2.4 mm long, adaxially glabrous. Pistillate flowers: pedicels inconspicuous, the diameter even throughout;receptacle flat, ca. 1.5 X 3 mm; tepals coriaceous, ovate, ca. 3 X 2 mm (innerwhorl slightly smaller), at anthesis erect?, reddishbrown, adaxially glabrous;staminodesof whorls I and II, the anthers, sessile, chubbytrapezoid,ca. 1.2 X 1.2 mm, glabrous, the locelli 4, apical, in a shallow arch, introrse,the glands absent;whorl III columnar,ca. 1.4 X 1 mm, glabrous, the locelli 4, the upper pair latrorse, the lower pair extrorse, the glands fused at base of stamen; whorl IV staminodial;staminodes,minute covered with long hairs; all staminodesreddish brown; pistil ca. 2 X 1 mm; ovary globose, ca. 1.2 mm long, glabrous. Fruits: pedicels to 6 X 6 mm, gradually enlargingto form the cupule;cupule trumpet-shaped, to 4 X 10 mm, smooth, the margin undulate,tepals persisting;berry elliptic, to 20 X 9 mm. FLORA NEOTROPICA dostemonodaphne miranda appears to be most closely relatedto R. licanioides and R. scandens (see discussion underR. licanioides and TableIX). Both vegetativelyas well as in inflorescencestructure, this species is very similar to Paraia bracteata Rohwer, H. G. Richt & van der Werff, from central and easternAmazonia,the only known species of this recently described genus; prior to the descriptionof this new genus, the similarities between the specimens of this species namedin herbariaas "Nectandra paraensis Coe-Texeira" and Rhodostemonodaphne mirandahad alreadybeen noted by Rohwer (1986). Furthermore,Paraia bracteata shows cryptic dioecy (Rohwer et al., 1991). However, the outer stamens with stipitate,incumbentantherswith (sub)apicallocelli, and the characteristic protruding style and stigma of P bracteata(this last featurewas not mentioned in the original description),are unlike any observed in Rhodostemonodaphne.I was unable to find the holotype-the only sheet with complete fruitsof Nectandramirandaat K and thus could not check for the presenceof an incipient double rim in the cupule as was describedfor Paraia bracteata. 16. Rhodostemonodaphne revolutifolia Madrifian, Brittonia 48: 55, figs. 5, 6. 1996. Type. French Guiana.CriqueArataye,Nouraguesstation,8 Aug 1990 (d fl), Sabatier 3500 (holotype: NY; isotypes: K, MO). Fig. 12C Trees: branchesbasitonic, in axils of cataphylls; twigs terete, 2-5 mm diam.; epidermisblack, barely visible due to indumentcover; terminalbud plump, Field notes. Shrubs to scandent?shrubs to 4 m ca. 6 X 4 mm; cataphyllspersistingon currentflush, tall and 3 cm diam. Tepals off-white. Cupule red; to 6 mm long; indumentpubescent, persisting for at berry black. least two flushes, the hairs dense, to 1 mm long, Leaves: petioles to Distribution (Fig. 11). Knownonly from the type straight crisped, erect, yellowish. X 1.2-2.8 mm, terete;blades cm 0.5-1.9 pulvinate, found in westermGuyana,at 200 m, in rainforest, on flat to bullate, narrowly elliptic, 6red lateriticsoil. As with othershrubbyspecies of the chartaceous, X 3-7(-19) cm; base acute, 40-60; apex 18(-22) Guayanahighlands,the presence of both flowers and attenuate, 70-80, ultimately acuminate for up to fruits on the same individual,indicates that this spe1.3(-1.5) cm; margin revolute;primary vein above cies does not have a narrowdelimited flowering seaslightly raised and rounded, below prominent;secson. ondary veins 3-6(-7) pairs, equidistant,eucamptoRhodostemonodaphnemiranda is a most distinc- dromous,above slightly raised, below prominent,ditive species. It bearsglabrous,chartaceous,nearlyob- verging at 55-60(-70)?, evenly arching, chordal ovate, cordateleaves, with the first two pairs of sec- angle 20-30o, the angle uniform along blade length; ondaryveins leaving the midribat a more acuteangle tertiary veins above and below slightly raised, random-reticulate;higher-orderveins on both sides and furtherapartthan the rest. Unfortunatelythis species is known from a single slightly raised; surface above yellowish brown in collection of a pistillate individual, and the flowers younger leaves, olive-green in older leaves, below are in bud. Hence there is little informationavailable greenishbrown,slightly glaucous,the veins yellowish regardingthe androecialstructureof the species. Rho- to brown; indumentabove absent, the primary and SYSTEMATICTREATMENT secondary veins tomentose, below pubescent, the hairs dense, to 1 mm long, straight,erect, yellowish, denseron the veins, persistingfor at least two flushes. Staminate inflorescences: basitonic, pendulous, peduncles 12-17 cm long, the hypopodia5-9 cm X 11.2 mm, branchorders 5, the second-orderbranches 4-5, dispersed,lowest branchto 4.5 cm long, all axes greyish to blackish, puberulous;bracts persistent,to 0.9 mm long, spoon-shaped,reflexedadaxiallyhairy; bracteolespersistent,to 0.7 mm long, adaxiallyhairy. Staminateflowers: pedicels ca. 5.6 X 1 mm, the diameter even throughout;receptacle flat, ca. 2 X 4.8 mm; tepals chartaceous,ovate, ca. 4.2 X 4.2 mm (inner whorl slightly smaller), at anthesisforminga cup ca. 6 mm diam. erect, reddishgray, adaxiallytomentose; stamens of whorls I and II, the antherssessile, chubby,broadlyelliptical, ca. 1.8 X 3.2 mm, with a few hairs at base, the locelli 2, apical, introrse,the glands absent;whorl III columnar,ca. 1.2 X 1 mm, with a few hairs at base, the locelli 4, the upperpair latrorse,the lower pair extrorse,the glands globular, ca. 1.2 mm diam.; whorl IV absent;all stamens reddish;pistillode absent (minute,to 0.6 mm long). Pistillateflowers: pistil ca. 1.7 X 0.8 mm; ovary ovoid, ca. 0.8 mm long, glabrous.Fruits unknown. Field notes. Trees to 12-15 m tall and 10-12 cm diam.;buttressessmall;outerbarksmooth (thin,fragile, finely fissuredand chipped),brownishred in section; inner bark ca. 3 mm thick, somewhat fibrous, pale yellow; wood yellow, wood aromatic.Tepalsyellowish. Distribution (Fig. 11). Known from two collections from central French Guiana, at 100-350 m, in rain forest. Both flowering collections were made in the month of August, at the beginningof the dry season. Additionalspecimenexamined.FRENCHGUIANA: abovePetitSaut,28 Aug 1993 (Y fl), FleuveSinnamary, Moriet al. 23416(CAY-n.v.,GH,MO,NY,P). VegetativelyRhodostemonodaphnerevolutifoliais a very distinctivespecies with conspicuouslyrevolute leaves, a rare condition in the family. The inflorescences are distinctly cymose, with persisting, recurved bracts and bracteoles. The most striking featureof the species are the two-locellateouterstamens of the staminateflowers,characteristicof Endlicheria. However, the vestigial four-locellate staminodia of the pistillateflowersplace it in Rhodostemonodaphne. 17. Rhodostemonodaphne crenaticupula Madriiinn, sp. nov. Type. Brazil. Amazonas:Mun. Ma- 57 naus, Ducke Forest Reserve, 31 Aug 1986 (d fl), Prance, Pena, Ramos & Monteiro2115 (holotype: MO; isotype: K). Fig. 12D Cupulacrenatainsignis;quoadcharacteres vegetativos Endlicheriae spruceiaccedenssedfoliis suprahaudnitidis, infrapubescentibus et staminibus quadrilocellatis differt. Shrubs to trees: branches basitonic, in axils of cataphylls; twigs terete, 1-2 mm diam.; epidermis brownish;terminalbud plump, ca. 3 X 1 mm; cataphylls persisting on current flush, to 4 mm long; indument pubescent, caducous after one flush, the hairs sparse,to 0.5 mm long, straightto curved,erect, brownish.Leaves: petioles pulvinate,0.3-1 cm X 11.8 mm, terete;blades chartaceous,flat, narrowlyto broadly elliptic, 6-18(-22) X 2-9 cm; base obtuse, 50-120?; apex obtuse, 60-80?, ultimately acuminate for up to 2.5 cm; marginplane; primaryvein above and below raised;secondaryveins 6-7 pairs,equidistant, brochidodromous,above slightly raised, below raised, diverging at ca. 600, evenly arching, chordal angle 25-30?, the angle uniform along blade length; tertiary veins above inconspicuous, below slightly raised,random-reticulateto scalariform;higher-order veins above inconspicuous,below slightly raised;surface above light green to bluish green, below olivegreen to greenish brown, the veins yellowish; indumentabove absent, the primaryvein tomentose, below pubescent,the hairs isolated, to 0.5 mm long, straight,erect, yellowish to yellowish brown, denser on the veins, persisting for at least two flushes. Staminate inflorescences: basitonic, erect, peduncles212 cm long, the hypopodia 1-5 cm x 0.5-1 mm, branchorders3, the second-orderbranches5-7, dispersed, lowest branchto 1.5 cm long, color and indumentof all axes as on twigs; bractscaducous,to 4 mm long, adaxially hairy (glabrous);bracteolespersistent, to 1 mm long, adaxiallyhairy (glabrous).Staminateflowers: pedicels ca. 4 X 0.6 mm, the diameter even throughout;receptacleobconical, ca. 1.6 x 2.5 mm; tepals membranaceous,obovate, ca. 3.2 X 2.5 mm (innerwhorl slightly smaller),at anthesisspreading, yellowish brown, translucent,adaxially puberulous; stamens of whorls I and II minutely spathulate to lacking a distinct filament,the anthersbroadlyelliptical, ca. 1.2 X 1.4 mm, glabrous, the locelli 4, apical, in a shallow arch, introrse,the glands absent; whorl III columnar,ca. 1.2 x 1 mm, with a few hairs at base, the locelli 4, the upperpairlatrorse,the lower pair extrorse,the glands globular,ca. 0.5 mm diam.; whorl IV absent; all stamens reddish;pistillode filiform, ca. 0.5 x 0.4 mm, basally hairy.Pistillateflow- 58 FLORA NEOTROPICA so .~~~~~~~~~~~~~~~~~~~ 70407 .. ......o. 40 .- ar fteseisi FIG13-itiuinmpo A Rh. antioquienssf Y Rh. debilis h Rhodostemonodaphne scandens group.Ecsyblrpent ar *Rh. celiana * IRz.crenaticupula ____ *Rh. leptoclada_ R.mirecolorata X Rh.parvifolia -. Rh. rufovirgata 0 Rh. steyermarkiana FIG. 13. Distributionmapof partof the species in the Rhodostemonodaphne scandens group.Each symbol represents one or more collections found in a degree-square. ers: pistil ca. 2 X 1 mm; ovary globose, ca. 1.2 mm long, hairy.Fruits: pedicels to 10 X 3 mm, abruptly enlarging to form the cupule; cupule hemispherical, to 5 X 12 mm, smooth, the marginundulate,tepals persisting;berry elliptic, to 22 X 13 mm. Field notes. Shrubsto trees to 6(-8) m tall and 5 cm diam., alreadyfloweringwhen 1 m tall; outerbark smooth?, dark brown; inner bark ca. 3 mm thick, salmon red; wood creamy yellow. Inflorescenceaxes red; tepals red. Cupule red. Distribution (Fig. 13). Most specimens are from the vicinity of Manaus, in central Brazilian Amazonia, but three specimens from the upperJuruaand Purus rivers (southwest of Manaus), one from the mid-Rio Negro, and four from the departmentof Loreto in Peru indicate that this species may be more widespreadin the Amazon basin. It grows on various soil types in non-inundatedrain forest at 25-100 m (to 300 m in Peru). Flowering begins in June, con- current with the onset of the dry season, and lasts until September. The fruits are found ripe October-January, during the rainy months. Additional specimens examined. PERU. LORETO: Maynas,Iquitos,nearLago Zdngaro,22 Oct 1964 (fr), Dodson 2850 (MO, US); Requena,JenaroHerrera,Rio Ucayali, 27 Jun 1986 (9 fl, fr), R. Vdsquezet al. 7689 (MO X2); Requena,JenaroHerrera,Sapuena,14 Sep 1987 (fr), R. Vdsquez & N. Jaramillo 9564 (MO). UCAYALI: Aguaytia, 18 Oct 1972 (fr), Schunke-V5406 (F, NY). BRAZIL. AMAZONAS:Mun. Manaus,Reserve 1501, 21 Nov 1988 (fr), Boom et al. 8572 (MO); Manaus-Caracarai rd., km 97, 3 Sep 1979 (d fl), Cid et al. 981 (MO); Mun. Manaus,edge of Igarapede Passarinho,25 Jul 1956 (d fl), D. F Coelho (INPA)3990 (NY); Mun. Manaus,BR 17, km 9, 1 Aug 1956 (d fl), D. F Coelho (INPA)4025 (NY); Mun. Manaus,Reserve 3402, 25 Feb 1992 (fr), Dick 6 (US); basin of Rio Jurua,near mouth of Rio Embira,30 Jun 1933 (d fl bud), Krukoff5098 (BM, F, K, NY X2, US); Mun. Manaus, Fazenda Dimona, 27 Nov 1989 (fr), Kukle 123 (MO, NY); Mun. Manaus, Reserve 1501, 11 Nov 1988 (fr), Mori & SYSTEMATICTREATMENT 59 Fernandez-da-Silva19744 (MO, NY); Mun. Manaus, Fazenda Dimona, 12 Feb 1992 (fr), Nee 42516 (NY); ManausAleixo rd., 30 Aug 1973 (d fl), Prance etal. 18761-B (MO); Manaus-Caracaraird., km 28, 10 Nov 1966 (fr), Prance et al. 3047 (NY); Manaus-Manacapuru rd., km 25, 3 Jan 1967 (fr), Prance et al. 3886 (NY, US); Mun. Manaus, Taruma rd., 1 Oct 1967 (fr), Prance et al. 3919 (NY X2); basin of Rio Purus,3 km SW of Labrea,28 Oct 1968 (fr), Prance et al. 7993 (NY); Manaus-P6rtoVelho rd., km 246, 4 km S of Igap6 Aqu, 14 Mar 1974 (fr), Prance et al. 20525 (NY); Manaus-Caracarai rd., km 60, INPA ExperimentalReserve, 30 Aug 1974 (d fl), Prance et al. 21690 (MO); ManausCaracaraird., km 64, 18 May 1984 (d fl bud), Renner 991 (US); Mun. Manaus,Ducke ForestReserve, 30 Jun 1993 (Y fl), Ribeiro et al. 943 (INPA, MO); 4 Jul 1993 (d fl), 1003 (INPA, MO); UpperRio Negro, Ilha das Flores, 14 Jan 1960 (fr), Rodrigues & L. Coelho 1469 (NY); Manaus-Itacoatiara rd., km 70, 11 Nov 1960 (fr), Rodrigues 1911 (NY); Manaus-Itacoatiara rd., km 80, 9 Jun 1961 (st), 2752 (NY); Mun. Manaus,Ducke ForestReserve, 17 Jul 1963 (d fl bud), Rodrigues5369 (F, NY X2); Manaus-PortoVelho rd., 14 Jul 1972 (d fl), M. Silva et al. 744 (MO); Manaus-Caracarai rd., km 40, 10 Dec 1973 (fr), Steward & Ramos P20098 (MO); (d fl), P20100 (MO); Manaus-Caracarai rd., km 127 (fr), Stewardet al. P20308 (MO). RoRAIMA:Serrinha,Rio Mucajaf,31 Jan 1967 (fr), Prance et al. 4211 (NY). Etymology. The epithet refers to the strongly lobed cupule with rounded, woody persistenttepals [Lat. crenatus, with roundedteeth + cupula, cup]. FIG. 14. Rhodostemonodaphnedebilis (Killip 20400). Local names. Brazil: louro branco,louro preto. Rhodostemonodaphnecrenaticupulais easily recognizable by its tepals which are deep red at anthesis and persist in fruit as conspicuous, rounded, woody projections.The dry leaves have a dark lamina that contrastswith the light venation. Vegetatively this species can easily be confused with Endlicheriasprucei (Meissner) Mez, which has an overlappingrange.Both species have slendertwigs with similarindumentum,and leaves similarin shape and size. Nevertheless, at least when dry, the adaxial leaf surfaceof E. sprucei is conspicuouslysmoothand shiny, while that of R. crenaticupulahas a raised venation, and is dull. 18. Rhodostemonodaphne debilis (Kosterm.) Chanderb.,Fl. Neotrop.Monogr.91: 1-140. 2004. Endlicheria debilis Kostermans, Recueil Trav. Bot. Neerl. 34: 555. 1937. Type.Peru.Loreto:Balsapuerto(lower Rio Huallaga basin), 28-30 Aug 1929 (fr), Killip & Smith 28400 (holotype: NY; isotype US-n.v.). Fig. 14 Shrub to tree: branchesbasitonic, in axils of cataphylls; twigs terete, 1-2 mm diam.; epidermis brownish;terminalbud unknown;cataphyllspersisting on currentflush, to 4 mm long; indumentpuberulous, caducous after one flush, the hairs sparse,to 1 mm long, straight to curved, appressed ascending, brownish. Leaves: petioles slender, 1-1.5 cm X 1 mm, terete;blades chartaceous,flat, narrowlyelliptic, 9-15 X 2.5-4.5 cm; base acute, 70 100'; apex acute, 5070 ,ultimately acuminate for up to 1.5 cm; margin plane; primary vein above and below raised; secondary veins 5-8 pairs, equidistant, brochidodromous, above slightly raised, below raised, diverging at ca. 600, evenly arching, chordal angle ca. 300, the angle uniform along blade length; tertiary veins above and below slightly raised, random-reticulate to scalari- form; higher-orderveins above and below slightly raised;surfaceabove and below brown,the veins similar; indument above absent, the primary vein puberulous at base, below puberulous, the hairs sparse, to 0.3 mm long, straightto curved,appressed, brownish, somewhat denser on the veins, persisting for at least two flushes. Staminate plant unknown. Pistillate inflorescence: basitonic, erect?, peduncle 4 cm long, hypopodiumca. 4 cm X 2 mm, branching unknown, color and indument as on twigs; bracts not 60 FLORA NEOTROPICA seen. Pistillate flowers unknown.Fruits: pedicels 25 X 5 mm, abruptlyenlarging to form the cupule; cupule hemispherical,18 X 13 mm, smooth, the margin undulate, tepals persisting; berry elliptic, 28 X 18 mm. Field notes. Tree 6-7.6 m tall. Cupule bright red; berry green. Distribution (Fig. 13). Knownonly from the type collection from the upperAmazon. It grows in dense forest at 150-350 m. Maturefruits found in late August. I have not seen the four-locellate staminodes reported and described by Chanderbali(2003). Rhodostemonodaphnedebilis is indeed very similarto R. crenaticupula.It can be distinguishedfrom the latter by its puberulousindumentwith shorter,sparse, appressed hairs, slender petioles, its brownish drying leaves with inconspicuousvenation, and largerfruits with the persisting tepals slightly distinguishableon the undulatingmarginof the cupule. 19. Rhodostemonodaphne curicuriariensis Madriniain,sp. nov. Type. Brazil. Amazonas: Rio Negro, above mouthof Curicuriaririver,24 Oct 1932 (Y fl), Ducke 25674 (holotype: U). Fig. 15 Frutex insignis, petiolis crassis in sicco nigrescentibus, foliis glabrisin sicco flavoviridibus,bullatis,venationesupra elevata a congeneris diversa. Scandent? shrubs: branchingunknown;twigs angular and remainingso for at least two flushes, ca. 3 mm diam.;epidermisyellowish to brownish;terminal bud slender, 4 X 2 mm; cataphylls caducous; indument tomentose, caducous by next flush, the hairs sparse, to 0.4 mm long, straight,appressed,ascending, yellowish. Leaves: petioles robust,ca. 1.5 cm X 3.5 mm, conspicuously blackened, terete; blades chartaceous,flat to undulateto bullate, narrowlyelliptic, 17-31 x 7-9 cm; base acute, ca. 700; apex acute to obtuse, 70-900, acuminate for up to 2 cm; marginplane; primary vein above and below raised; secondary veins 7-9 pairs, with distinct intersecondaries, equidistant,brochidodromous,above slightly raised,below raised,divergingat ca. 600, evenly arch- FIG. 15. Rhodostemonodaphne (Ducke 25674). curicuriariensis soon glabrescent,the hairs isolated, to 0.4 mm long, straight,appressed,ascending, yellowish. Staminate plant unknown.Pistillate inflorescences: acrotonic?, pendulous?,peduncles ca. 15 cm long, the hypopodia ca. 5 cm XI mm, branch orders 3, the second-order branches 11, dispersed, lowest branch to ca. 1.7 cm long, color and indument of all axes as on twigs; bractscaducous, to 3 mm long, adaxiallyhairy;bracteoles persistent, to 0.8 mm long, adaxially hairy. Pisillate flowers: pedicels ca. 2.8 X 0.6 mm, the di- ametereven throughout;receptacleobconical, ca. 1.2 X 2.4 mm; tepals membranaceous,elliptic to oblong, ca. 2 X 1.6 mm, at anthesis spreading,reddishto yellowish brown, adaxially glabrous; staminodes of whorls l and HI,the antherssessile chubby,trapezoid, ca. 1 X 1.4 mm (whorl II slightly smaller),glabrous, ing, chordal angle ca. 350, the angle uniform along blade length; tertiaryveins above slightly impressed, the locelli 4, apical, in a shallow arch, introrse,the below slightly raised,random-reticulate; higher-order glands absent;whorl III columnar,ca. 1.2 X 1.2 mm, veins above inconspicuous,below slightly raised;sur- glabrous,the antherstrapezoid,the locelli 4, the upper face above shiny light brownishgreen,below greenish pair latrorse, the lower pair extrorse, the glands (if brown; indument above absent, below puberulous, present) each pair fused to each other and the base of 61 SYSTEMATICTREATMENT the stamen which is thickened;whorl IV absent; all acute to obtuse, 70-100; apex obtuse to acute, 70staminodesreddish brown to black; pistil ca. 2.5 X 1000, ultimately acuminatefor up to 1.5 cm; margin 1.6 mm; ovary globose, ca. 1.6 mm long, glabrous. plane; primaryvein above and below raised;secondary veins 6-8 pairs, equidistant,brochidodromous, Fruits unknown. above and below raised, divergingat 55-650, evenly Field notes. Shrubs?Tepals green. arching, chordal angle ca. 300, the angle uniform Distribution (Fig. 11). Known from only two along blade length; tertiary veins above impressed, specimens from the Curicuriarimountains near the below raised, random-reticulate;higher-orderveins mouth of the river with the same name, a tributaryof above inconspicuous, below slightly raised; surface the Rio Negro. Growingin non-inundatedforest at ca. above dull dark green, below shiny light green; in100 m. The two flowering specimens were collected dumentabove absent,the primaryvein tomentose,bein October (ca. 50 years apart),at the beginning of low minutely puberulous,the hairs isolated, to 0.2 mm long, straight, erect, yellowish, denser on the the dry season. veins, caducous after one flush. Staminate infloresAdditional specimen examined. BRAZIL. AMAbasitonic,pendulous,pedunclesto 5 cm long, cences: river,CuZONAS: Rio Negro,abovemouthof Curicuriari 735 (NY). the hypopodiaca. 3 cm X ca. 0.4 mm, branchorders mtns.,24 Oct 1976(Y fl), Nascimento rucuriari 3, the second-orderbranchesca. 5, dispersed,lowest Etymology. Named afterthe Curicuriarimountain branchto 0.5 cm long, color and indumentof all axes rangeand riverwhere the two known specimenswere as on twigs; bracts and bracteoles caducous (not collected. seen). Staminateflowers: pedicels ca. 2.4 X 0.6 mm, diametereven throughout;receptacleflat, ca. 0.8 the Rhodostemonodaphnecuricuriariensishas leaves X 2 mm; tepals membranaceous,elliptic, ca. 2.5 X with distinctly blackened petioles, similar to those at anthesis spreading, reddish, somewhat 1.2 mm, found in Endlicheria rubrifioraMez and some other adaxially glabrous;stamens of whorls I translucent, leaves are conspicuously species of Lauraceae.The shiny and glabrous above and the yellow veins are and II spathulate,the anthersreniform,ca. 0.8 X 0.6 raised from an impressedgroove. The two specimens mm, glabrous,the locelli 4, apical, in a strong arch, cited, although undoubtedlyconspecific, have very introrse,the glandsabsent;whorlIII columnar,ca. 1.2 different leaf blades. In the type these are flat to X 0.8 mm, glabrous,the locelli 4, the upperpair laslightly undulate,while in Nascimento 735 they are trorse,the lower pairextrorse,the glandsfused, formdistinctly bullate. This situationis not uncommonin ing a continuousring; whorl IV absent; all stamens yellowish; pistillode absent. Pistillate flowers: pistil the genus (see "Morphology"). ca. 2.5 X 1.5 mm; ovary globose, ca. 1.5 mm long, glabrous.Fruits unknown. tumucumaquensis 20. Rhodostemonodaphne sp. nov. Type. Brazil. Amapa':Upper Madriniain, Rio Jari, 19 Aug 1993 (J fl), de Granville,Acevedo,Boyer & Hollenberg12377 (holotype:MO; isotypes: BBS-n.v., CAY-n.v., P-n.v., US-n.v.). Fig. 16A Field notes. Shrubs to 3 m tall and 3 cm diam. Tepalspink to red. Distribution (Fig. 11). UpperJaririveron the Tumucumaque mountain range on the Brazil/French Guianaborder,at 380-670 m; a specimen from Mt. St. Marcel (500 m) in southernFrenchGuiana,may this species. An understoryrainforest Rhodostemonodaphneleptocladae sine dubio proxima, also belong to soils on granitic outcrops. of well-drained species necnon sed foliis majoribusbullatis,petiolisbrevioribus, statimdiagnos- Flowersin August, at the beginningof the dry season. perianthiiset androeciiscircumscissilibus cenda. Additional specimens examined. FRENCH GUITrees: branchesbasitonic?,in axils of cataphylls; twigs terete,ca. 2 mm diam.;epidermisbrownish;terminal bud slender, ca. 3 X 2 mm; cataphylls caducous; indumentpubescent,persistingfor at least two flushes, the hairs dense, to 0.5 mm long, straight, erect, yellowish. Leaves: petioles pulvinate, 0.5-0.8 cm x 1.5-2.5 mm, terete;blades thinly chartaceous, flat to bullate, broadlyelliptic, 7-15 X 3-7 cm; base ANA: Mont St. Marcel, 26 Mar 1976 (? fl), Sastre 4551 (CAY,P-n.v.). BRAZIL. AMAPA: UpperRio Jari,20 Aug 1993 (Y fl), de Granvilleet al. 12383 (CAY-n.v., K-n.v., MO, NY-n.v., U-n.v., US); 25 Aug 1993 (d fl), de Granvilleet al. 12457 (CAY-n.v.,MO,US). Etymology. Named after the Tumucumaque (Tumuc-Humacin FrenchGuiana)mountainrangeon 62 FLORA NEOTROPICA V (617 0' 040600419 Y-* to' FI. 6 A Rooseondahe umcmaunss ~ ~ ~ Ganileeta ~ ~ 138) C. R. mirecoorata (Tillet & Tillet4580). B. R. ufovirgata Mo et t 513) ~ ~ B . egesi ~ -' (ienii 0 no; t t62) SYSTEMATICTREATMENT 63 the borderbetween the Brazilianstates of Amapa'and Para'and Surinameand FrenchGuiana. mm long, straight,erect, yellowish, persisting for at least two flushes. Staminate inflorescences: along whole length of flush, erect to pendulous?,peduncles Rhodostemonodaphne tumucumaquensis has 4-16 cm long, the hypopodia 1-5 cm X 1.5-2.5 mm, bright-green-drying,undulate to somewhat bullate branchorders 3-6, the second-orderbranches7-12, leaves. The four specimens cited here were originally included in the description of R. leptoclada Madri- dispersed,lowest branchto 1(-3) cm long, color and niain.Unlike R. leptoclada, which has green flowers, indumentof all axes as on twigs; bractsandbracteoles the flowers of R. tumucumaquensisare red. As orig- caducous (not seen). Staminateflowers: pedicels ca. inally described and illustratedthe pistillate individ- 4 X 0.8 mm, the diametereven throughout;receptacle uals of R. tumucumaquensishave theirtepals and sta- obconical, ca. 1.5 X 3.2 mm; tepals coriaceous, obmens dehiscing in a ring after anthesis, while in R. ovate, ca. 4 X 3.2 mm (innerwhorl slightly smaller), leptoclada they are not. The collection from Mt. St. at anthesis spreadingto recurved,reddish,somewhat Marcel (initially though of as intermediatebetween translucent,adaxially puberulousto tomentose; stathe two above mentionedspecies) has flower buds at mens of whorls I and II spathulate,the antherstrapa very early stage of development;it has reddishto- ezoid, ca. 1 X 1.4 mm, with a few hairs at base, the mentose twigs and leaves drying darkerthan R. lep- locelli 4, apical, in a strong arch,introrse,the glands toclada, and it very likely belongs to R. tumucuma- absent;whorl III columnar,ca. 1.4 X 0.8 mm, with a few hairs at base, the locelli 4, the upperpairlatrorse, quensis. the lower pair extrorse, the glands globular,ca. 1.2 mm diam.;whorl IV absent;all stamensreddish;pistillode filiform, ca. 0.5 X 0.2 mm, hairy. Pistillate sp. 21. Rhodostemonodaphne negrensis Madrifnan, flowers: pistil ca. 3.2 X 1.2 mm; ovary ovoid, ca. 1.6 nov. Type. Brazil. Amazonas: Manaus-Caracarai mm long, hairy.Fruits: pedicels to 1 X 5 mm, gradrd., km 60, Sep 1974 (d fl), D. F Coelho (INPA) to form the cupule; cupule trumpet46810 (holotype:INPA;isotypes: F, U). Fig. 16B ually enlarging shaped, to 10 X 20 mm, wrinkled,the marginunduSpecies foliis magnis discoloribussupraplerumquema- late, tepals persisting;berry elliptic, to 25 X 15 mm. culatis, venatione infra conspicue pallida et cupula tepalis persistentibusa congeneris diversa. Field notes. Trees to 12(-15) m tall and 15(-20) cm diam., already flowering when 5 m tall (but see Trees: branching unknown; twigs angular, soon discussion); outer bark smooth (with minute scales becoming terete, 4-5 mm diam.; epidermisbrownish and irregularlyfissured);inner bark ca. 4 mm thick, to black, barely visible due to indument cover; teryellowish brown;wood light yellowish brown, wood minal bud plump,ca. 5 X 3 mm; cataphyllspersisting axes reddish;tepals orangeto Inflorescence aromatic. on currentflush or caducous, to 1.2 mm long; indument puberulous,caducous after one flush, the hairs purple.Cupule red. dense, to 0.2 mm long, straight, erect, brownish. Distribution (Fig. 11). Known only from the Rio Leaves: petioles robust, 1.3-2.5 cm X 2.2-5 mm, Negro basin, at 50-150 m; it grows in non-inundated adaxially flattened;blades chartaceous,flat to undu- white-sand forest (caatinga). Flowers June-Novemlate, broadlyovate to elliptic, 15-34 X 6-19 cm; base ber, throughthe dry season. FruitsOctober-January, obtuse to rounded, 60-160o; apex acute to rounded, at the beginning of the rainy season. 90-140o, ultimatelyacuminatefor up to 2.5 cm; marAdditional specimens examined. BRAZIL. AMAgin plane; primary vein above flat to raised, below prominent;secondary veins 6-10 pairs, equidistant, brochidodromous,above raised to impressed,below raised, diverging at 45-65?, evenly arching, chordal angle 20-30(, the angle uniform along blade length; tertiaryveins above impressed,below slightly raised, random-reticulateto scalariform;higher-orderveins above flat to slightly raised,below slightlyraised;surface above olive-green to light brown, often mottled, the veins darker,below yellowish brownto brown,the veins lighter; indument above puberulous, the primary and secondary veins tomentose, caducous by next flush, below puberulous,the hairs sparse,to 0.5 Rio Negro, Sao Gabrielda Cachoeira,18 Oct 1987 (fr), Daly et al. 5457 (K, MO, NY); Curicuriaririver,mouth, 13 Oct 1932 (d fl), Ducke 25681 (U); Rio Uaupes, Ipanore, 17 Nov 1987 (d fl), Kawasaki 179 (F X2, MO, NY, US); basin of Rio Jurua,near mouth of Rio Embira,27 Jun 1933 (st), Krukoff5038 (NY); Mun. Manaus,FazendaEsteio, 21 Nov 1984 (fr), Nascimento et al. (INPAIWWF)1201.288 (MO); Rio Cuieiras,above mouthof Rio Brancinho,11 Sep 1973 ( Y fl bud), Prance et al. P.17721 (MO);Mun. Manaus, Ducke Forest Reserve, 6 Sep 1966 (d fl), 2181 (MO); Camandos-UapesAirportrd., 30 Oct 1971 (d fl), Prance et al. 15860 (MO, NY); Manaus-Caracarai rd., km 60, INPA ExperimentalReserve, 15 Aug 1974 (d fl bud), Prance et ZONAS: FLORANEOTROPICA 64 al. 21660 (MO);31 May 1975 (fr), Pranceet al. 23447 (NY);Mun.Manaus,DuckeForestReserve,14 Aug 1965 (d fl), Rodrigues7029 (G, NY); 19 Jul 1994 ( fl), Vicentini et al. 816 et al. 628 (INPA,MO);30 Jan1995(fr),Vicentini (INPA,MO). Etymology. Named after the Rio Negro. Rhodostemonodaphnenegrensisis a low tree,with large leaves thataremottled(at least when dry), often glaucous abaxially, and with conspicuously lightcolored secondary and tertiary venation. Its large flowers are similar only to those of R. dioica and R. sauilensis,yet in addition to other vegetative differences, the indumentof the former is yellow tomentose, with appressedand erect hairs, while that of the latteris silver sericeous. A collection from the upperRio Cuieras(a tributary of the Rio Negro) from an open forest (Prance et al. P.17721) is includedhere with hesitation.It was taken from a low shrub2 m tall; its leaves, although similarin shapeandcolor to those of othercollections of the species, are completely glabrous,have shorter petioles, and secondaryvenationwith the loops more distant from the margin. The buds of the pistillate flowers are much smaller than the buds of the only other known pistillate floweringindividual(Vicentini et al. 628), the staminodes of whorls I and II have their lower locelli latrorse, and the pistil is globose with a very short style. Having seen only pistillate material in bud, I cannot be sure of its identity, yet based on its foliar charactersand geographicaldistribution it may in fact belong to R. negrensis. The collection Kawasaki 179, from the Uaupes river, a tributaryof the Rio Negro, differs from the other collections in that it has more angulartwigs, sparser indument, and broader, nearly glabrous leaves. The flowers, however, are like the other collections of this species. 22. Rhodostemonodaphne mirecolorata (C. K. Allen) Rohwer,Mitt. Inst. Allg. Bot. Hamb. 20: 86. 1986. Ocotea mirecolorata C. K. Allen, Mem. New York Bot. Gard. 12(3): 116, fig. 24. 1965. UpperMazaruni Type. Guyana.Mazaruni-Potaro: River basin, Karowtipu,1 Oct 1960 (d fl), Tillett & Tillett 45580 (holotype: NY; isotypes: G, K, Fig. 16C NY). the hairs dense, to 1 mm long, straight to curved, erect, reddish.Leaves: petioles slender, 1-1.7 cm X 1.4-2.2 mm, terete;blades chartaceous,flat,narrowly ovate to elliptic, 4-12 X 2-4 cm; base acuteto obtuse, 60-1 100;apex attenuateto acute, 35-90?, ultimately acuminatefor up to 1.5 cm; margin plane; primary vein above raised,below prominent;secondaryveins 3-6 pairs, equidistant,eucamptodromous,above flat, below prominent,divergingat 50-60, abruptlyarching near margin, chordal angle 15-20?, the angle uniform along blade length; tertiary veins above inconspicuous, below raised, random-reticulateto scalariform;higher-orderveins above inconspicuous, below slightly raised; surface above brown in young leaves, shiny olive-green in older leaves, below brown; indumentabove puberulous,the primaryand secondaryveins puberulous,caducousby next flush, below tomentose, the hairs dense, to 1 mm long, curvedto crisped,erect, reddishbrown,denseron the veins, persisting for at least two flushes. Staminate inflorescences: along whole length of flush,erect,peduncles 4-9 cm long, the hypopodia 1-3 cm X 1-2 mm, branchorders3-4, the second-orderbranches47, dispersed,lowest branchto 1.5 cm long, color and indumentof all axes as on twigs; bractsandbracteoles caducous (not seen). Staminateflowers: pedicels ca. 6 X 1.6 mm, the diametereven throughout;receptacle flat, ca. 3.5 X 7.2 mm; tepals coriaceous, ovate to obovate, ca. 4 X 3.5 mm, at anthesiserect to spreading, reddishblack, adaxiallypatchy puberulous;stamens of whorls I and II, the antherssessile, chubby, trapezoid,ca. 1.6 X 1.6 mm, glabrous,the locelli 4, apical, in a shallow arch, introrse,the glands absent; whorl IIIcolumnar,ca. 2.4 X 1.2 mm, glabrous,the locelli 4, the upper pair latrorse,the lower pair extrorse,the glands globular,ca. 1 mm diam.;whorl IV absent; all stamens reddish;pistillode absent.Pistillateflowers andfruits unknown. Field notes. Trees to 30 m tall and 60 cm diam. Inflorescenceaxes green;tepals light yellowish green; stamens/staminodesgreen, the locelli flaps yellow. Distribution (Fig. 13). Knownwith certaintyonly from the type from the Karowtipumountain(adjacent to the Roraima-Iluchain of the Guayanahighlands). Range may be extended east to Guyana and French Guiana with the inclusion of two specimens of uncertain identification(see below). Grows in mixedevergreenforest. Flowers during the month of OctoTrees: branchesbasitonic, in axils of cataphylls; ber at the beginning of the rainy season. twigs angular,soon becoming terete,ca. 4 mm diam.; Rhodostemonodaphnemirecoloratais apparently epidermisblack,barelyvisible due to indumentcover; terminalbud plump, ca. 1 X 2 mm; cataphyllscadu- a tall tree, with red tomentose twigs, the indument cous; indumentpubescent, caducous after one flush, tuming black on older growth.The leaves are elliptic SYSTEMATICTREATMENT 65 to narrowlyelliptic, stronglydiscolorous,dark,shiny green above, lighter,matte green below and with distinctive dense, crisped, red indument(this indument also covers the upper surface of the very young leaves). Among the species with red tomentose twigs treatedhere, Rhodostemonodaphnemirecolorataresembles R. rufovirgata the most. Yet the inflorescences and flowers of the two species are completely different, those of R. rufovirgata being muchbranched and the flowers considerably smaller.Vegetativelyit is almost impossible to tell the two apart. Two sterile specimens here treatedas R. rufovirgata, Mutchnick966 and Sabatier & Prevost 3710, could just as well be placed in R. mirecolorata. 1.8 mm, branchorders 5, the second-orderbranches (3-)7-10, dispersed, lowest branch to 3 cm long, color and indumentof all axes as on twigs; bractsand bracteoles caducous (not seen). Staminateflowers: pedicels ca. 2.4 X 1 mm, the diametereven throughout; receptacle obconical, ca. 1.6 X 3.6 mm; tepals chartaceous,ovate, ca. 1.5 X 1.5 mm (inner whorl slightly smaller), at anthesis erect, reddish brown, adaxiallytomentose;stamens of whorls I and II minutely spathulateto lacking a distinctfilament,the anthers reniform,ca. 1 X 0.8 mm, glabrous,the locelli 4, vertically elongate, apical, in a shallow arch, introrse,the glands absent;whorl III columnar,ca. 1.4 X 0.6 mm, glabrous,the locelli 4, the upperpair latrorse,the lower pairextrorse,the glands globular,ca. 0.6 mm diam.; whorl IV absent; all stamens reddish brown;pistillode absent.Pistillateflowers unknown. 23. Rhodostemonodaphne rufovirgata Madriiain, Fruits: pedicels to 20 X 4 mm, graduallyenlarging Brittonia48: 58, fig. 7. 1996. Type. French Gui- to form the cupule; cupule hemispherical,to 6 X 1 ana. Saul, Mont La Fumee, 24 Sep 1982 (d fl), mm, smooth to wrinkled, the margin straight,tepals Boom & Mori 1811 (holotype: NY; isotypes: caducous;berry elliptic, to 20 X 15 mm. Fig. 16D CAY-n.v., K, MO). Field notes. Trees to 20-27(-40) m tall and 25Trees: branchesbasitonic, in axils of cataphylls, 35(-80) cm diam.; buttressessmall or absent; outer precocious, the whole flush and branchesbeing pro- bark rough, gray, brown, pockmarked,with easily duced before elongation and lignification;twigs an- falling scales, tan underneath;inner bark ca. 8 mm gular, soon becoming terete, 2-4 mm diam.; epider- thick, orange;wood yellow. Receptacleyellow-green; mis black, barely visible due to indument cover; tepals yellow-brownto orangish;flowers with musty terminalbud plump, ca. 4 X 4 mm; cataphyllsper- aroma.Cupule orange;berry ripeningpurple. sisting on currentflush, to 6 mm long; indumenttoDistribution (Fig. 13). Originallydescribedfrom mentose, persistingfor at least two flushes, the hairs central FrenchGuiana,a recent collection from Guydense, to 0.5 mm long, curved to crisped, erect, reddish. Leaves: petioles slender, 1.3-2.5 cm X 1.3-2 ana extendingits rangeconsiderably;at 75-300 m, in mm, terete;blades chartaceous,flat,narrowlyelliptic, rain forest. Flowers September-October,during the (7-)10-18(-22) X 2-6(-9) cm; base acute, 60-100?; dry season. apex attenuate,70-80o, ultimately acuminatefor up Additionalspecimensexamined.GUYANA.Ruputo 2.2(-3) cm; margin minutely recurved;primary NUNI: Iwokrama reserve, Mt. Daniel, 27 Feb 1995 (fr), vein above slightly raised and rounded,below raised Mutchnick966 (MO, US-n.v.). and longitudinallyfurrowed;secondary veins 4(-5) Nassau Gebergte, 24 Feb SURINAME. MAROWUNE: pairs, equidistant,eucamptodromous,above flat, be- 1949 (st), Lanjouw& Lindeman2300 (NY). FRENCH GUIANA: Sail, Mont La Fumee, 17 Oct low raised, diverging at 40-65?, evenly arching, chordalangle 25-35?, the angle uniformalong blade 1982 (d fl), Mori et al. 15103 (CAY,NY); Fleuve Sinnalength; tertiary veins above flat, below raised, mary,above Petit Saut, 27 Aug 1993 (st), Mori et al. 23393 random-reticulateto scalariform;higher-orderveins (CAY-n.v.,NY);Sail, EauxClaires,5 Sep1994(d fl),Mori above flat, below slightly raised; surface above red- et al. 23832 (CAY-n.v.,GH,NY); St. Elie rd.,Interfleuve dish brown in young leaves, olive-green in older leaves, below whitish to cream colored in young leaves, dark brown in older leaves; indumentabove absent, the primary and secondary veins tomentose, below tomentose, the hairs dense, to 0.5 mm long, curved to crisped, erect, whitish, denser and reddish on the veins, caducous by next flush. Staminate inflorescences: along whole length of flush, erect, peduncles 6-14 cm long, the hypopodia3-5 cm x 1.2- 2 Oct 1989 (st), Prevost & SabaSinnamary-Counamama, tier 2920 (CAY-n.v., MO, U-n.v.); Crique Arataye, Nouragues station, 4 Nov 1987 (st), Sabatier & Prevost 1975 (B-n.v., CAY-n.v., K-n.v., MO, P-n.v., U-n.v.); St. Elie rd., InterfleuveSinnamary-Counamama,6 Aug 1991 (st), Sabatier & Prevost 3710 (MO); Crique Passoura,littoralregion, 9 May 1992 (st), Sabatier & Prevost 4014 (B-n.v., CAY-n.v., MO, P-n.v.); CriqueArataye,SautsParare,6 Sep 1977 (st), Sastre 5901 (CAY,P-n.v.); Saiil, Eaux Claires, 9 Aug 1993 (st), van der Werifet al. 12954 (MO-n.v., NY). FLORA NEOTROPICA 66 Local name and use. French Guiana, Bala-oiu (Boni). Used for making canoes. Rhodostemonodaphnerufovirgatais distinctivein its flushingpattern-new flushes togetherwith leaves and lateral branches are producedbefore elongation and lignificationof the twigs. During flushes all plant organsare covered with a conspicuousred indument. The small-leafed specimens of this species are vegetatively indistinguishablefrom those of R. mirecolorata; the larger-leafedspecimensresemblethose of R. crenaticupula. However, the flowers of both these species are very differentfrom those describedabove for R. rufovirgata. 24. Rhodostemonodaphneceliana (C.K. Allen) Rohwer,Mitt. Inst. Allg. Bot. Hamb.20: 85. 1986. Ocotea celiana C. K. Allen, Mem. New YorkBot. Gard. 10(5): 106, fig. 57. 1964. Type. Venezuela. Amazonas: Atures, SerraniaYutaje,Cerro CoroCoro, Rio Manapiare,2 Mar 1953 (? fl, fr), Maguire & Maguire 35450 (lectotype here desigFig. 17A nated, NY [photo NY neg. 5938]). Scandent shrubs: with long, slenderand sparsely branchedshoots; branchesbasitonic to mesotonic, in axils of basal foliage leaves; twigs terete, 2-4 mm diam.;epidermisbrownishto black,barelyvisible due to indument cover; terminal bud plump, ca. 4 X 2 mm; cataphylls caducous; indumentpubescent, persisting for at least two flushes, the hairs dense, to 1 mm long, curved to crisped, erect, reddish.Leaves: petioles pulvinate, 0.5-1.5 cm X 1-2.5 mm, terete; blades coriaceous, flat to undulate to bullate, narrowly ovate to elliptic, 3-11 x 1-5 cm; base rounded to cordate,90-1900; apex attenuateto acute, 40-70?, ultimately acuminate for up to 1.5 cm; margin recurvedto revolute;primaryvein above slightlyraised, below prominent;secondary veins 8-11 pairs indistinguishable from tertiary veins towards the apex, equidistantto furtheraparttowardsthe apex, brochidodromous, above impressed, below prominent,diverging at 60-70?, evenly arching,chordalangle 30650, the angle increasingapically;tertiaryveins above slightly impressed, below raised, random-reticulate; higher-orderveins above slightly impressed, below slightly raised; surface above yellowish green to greenish brown, below reddish brown to reddish black; indumentabove absent, the primary vein tomentose, below pubescent,the hairsdense, to 0.6 mm long, straightto curved, erect, reddish,persistingfor at least two flushes. Staminate inflorescences: basitonic to mesotonic, pendulous, peduncles 4-16 cm long, the hypopodia 2-7 cm X 0.5-1 mm, branch orders 4, the second-orderbranches4-7, dispersed, lowest branchto 1(-3) cm long, color and indument of all axes as on twigs; bracts caducous (not seen); bracteolespersistentor caducous,to 2 mm long, adaxially hairy.Staminateflowers: pedicels ca. 3.2 X 0.8 mm, the diametereven throughout;receptacleflat,ca. 1.2 X 2.2 mm; tepals membranaceous,elliptic, ca. 3 X 2.4 mm, at anthesis erect to spreading, reddish brown, adaxiallytomentose;stamensof whorls I and II, the antherssessile, laminar,obovate, ca. 1.4 X 1.2 mm, with a few hairs at base, the locelli 4, apical, in a shallow arch, introrse,the glands globularto flattened, ca. 0.3 mm diam.; whorl III columnar,ca. 1.4 x 0.6 mm, with a few hairs at base, the locelli 4, the upperpair latrorse,the lower pairextrorse,the glands globular,ca. 0.6 mm diam.;whorlIV staminodial,columnar;all stamensreddishyellow; pistillode absent. Pistillateflowers: pistil ca. 2.5 X 1.4 mm; ovary globose, ca. 1.6 mm long, glabrous(with a few hairs at base). Fruits: pedicels to 10 X 3 mm, graduallyenlargingto form the cupule;cupule trumpet-shaped,to 4 X 7 mm, smooth, the marginundulate,tepals persisting; berry elliptic, to 13 X 9 mm. Field notes. Scandent shrubs, sprawling over neighboring vegetation up to 2(-4) m tall; wood white; leaves above dull to dark green. Receptacle greenish brown?; tepals yellowish; stamens/staminodes greenishwhite. Cupulered. Distribution (Fig. 13). Endemic to the YutajeCoro-Coro massif, a tepui (sandstone mountain) reaching to 2400 m, on the western edge of the Venezuelan Guayana.This species grows on the rocky summits and slopes between 1200 and 2200 m, in dense, low (ca. 6 m tall), wet, evergreen, uppermontane forest (Huber, 1995). All collections (flowering and fruiting)have been made in Februaryand March,at the beginning of the rainy season. Additional specimens examined. VENEZUELA. Atures,Serrania Yutaje,RioCoroCorovalley, 3 Mar1987(cd fl), Holst & Liesner3287 (MO,NY);Atures, SerranIa Yutaje,Rio Coro-Coro valley,6 Mar1987(d fl), Holst& Liesner3324(F,MO,U, US);Atures,SerranIaYutaje,summit,E sector,21 Mar1988 (d fl), Huber12603 deYutaje,CerroYutaj6,RioMana(MO);Atures,Sarrania AMAZONAS: piare, 17 Feb 1953 (d fl), Maguire & Maguire35320 (NY); (9 fl), 35320[A] (NY,W);Atures,Serrania Yutaje,Cerro Coro-Coro,Rio Manapiare,2 Mar 1953 (d fl), Maguire & Maguire35450[A] (F [photoF neg. 58176],US); Atures, Sarraniade Yutaje, Cerro Yutaj6, Rio Manapiare, 17 Feb 1953 (9 fl),),Maguire & Maguire35290 (G,GH,NY);(9 fl, fr), Maguire & Maguire35340 (FHO, NY). 67 SYSTEMATICTREATMENT A /_ - 1 ~ If FIG. 17. A Rhodostemonodaphne R. leptoclada (Mori et al. 14889). U - ~ ats5*s1<* Tt>% Nt$k^51*...................... celiana D. R. parvifolia ~ :: _ra __._ ~ (Maguire (Rodrigues ~ & Maguire 5435). ~ 35340) ~ ~ ~ B. R. steyermarkiana ~ ~ (Maguire 32753). C. 68 This is a most distinct species both in termsof its vegetative morphology and flower structure.It is a low, scandent shrub with thin, long, little-branched stems that grow supported by adjacent vegetation. This growth habit is sharedwith a few other species in the genus. The reddish pubescence on the twigs, inflorescences,and undersidesof leaves is also found in a few other species, generallyin those restrictedto the sandstone mountains of the Guayanahighlands. Its hard, brittle, undulate to bullate leaves with recurvedmargins,and anatomicaldetails are also quite characteristic.Its flowers are unlike any other in the genus, apart from those of R. penduliflora; the stamens of all three outer whorls have a pair of glands at the base. Allen (1964) stated that the outer two whorls of stamens in the staminateindividualsseemed sterile, thus resembling the genus Dicypellium;however, in Dicypellium the flowers are hermaphroditic.In the specimens examined, some of the staminateflowers had stamens of the outer whorls with incompletely developed anthers,but the great majorityof flowers seen had fertile anthersin all threeouterwhorls.Allen also mentions that all stamens having glands is akin to the conditionfound in the genus Pleurothyrium.In fact, Pleurothyriumhas glands only on the anthersof whorl III, as in most other Lauraceae,but they are often greatly enlargedand sometimes fuse to form a continuous ring (see Rohwer, 1986, and van der Werff, 1993). Urbanodendrondoes have glandson all staminal whorls, and the anthers are mainly fourlocellate, but the flowers are hermaphroditic. Allen cited Maguire & Maguire35450 as the type of Ocotea celiana, and noted details of the staminate and pistillate flowers, and fruits. Both twigs on the sheet Maguire & Maguire35450 at NY are from pistillate individuals;the one on the left has flowers,and the one on the right an infructescencewith a fruit(the fruit is now included in the envelope, but a NY photographof the specimen made some years ago shows it attached!).No isotypes were designated,but I was able to examinetwo duplicateswith the same number from F and US. Both are staminateindividuals.It is probablethatthe type collection is a mixed collection, and that the species was described prior to distribution of the duplicates. Allen cited the holotype as having staminateflowers and fruits;Maguire & Maguire 35450 at NY now consists of only pistillate individual(s?). Lectotypificationis needed because of the two elements in Maguire & Maguire 35450 (as mentionedin the discussion of R. capixabensis, it is unlikely that one individualwill have both staminate and pistillate flowers); I designate the pistillate ele- FLORA NEOTROPICA ment as the holotype. I have addedthe suffix A to the collection number to representthe staminateduplicates at F and US. One othercollection (Maguire& Maguire35320) is also mixed. The sheet at NY has both staminateand pistillate individuals, while the sheet at W has only one staminatetwig. I have added the suffixes A & B to the numberto refer to the staminateand pistillate elements. 25. Rhodostemonodaphne steyermarkiana (C. K. Allen) van der Werff, Ann. Missouri Bot. Gard. 76: 474. 1989. Ocotea steyernarkiana C. K. Allen, Acta Bot. Venez. 2: 216. 1967. Type. Venezuela. Bolivar:Auyan-tepui,summit,7 May 1964 (Y fl, fr), Steyernark93476 (holotype:NY [photo NY neg. 6678]; isotypes: G, K, P, VEN-n.v.). Fig. 17B Shrubs: branchesbasitonic to mesotonic, in axils of cataphyllsor basal foliage leaves; twigs terete, 25 mm diam.; epidermis black, barely visible due to indumentcover; terminalbud plump, ca. 1 X 1 mm; cataphylls caducous; indumentpubescent,persisting for at least two flushes, the hairs dense, to 1.2 mm long, straightto curved, erect, reddish.Leaves: petioles slender, 0.7-1.2 cm X 1-2 mm, terete; blades coriaceous, flat, narrowlyto broadly elliptic, 3-7 X 1-5 cm; base obtuse to rounded,60(1400; apex attenuate to acute, 40-100', mucronateto acuminatefor up to 1 cm; marginplane to revolute;primaryvein above flatto impressed,below slightly raisedto prominent; secondaryveins 4-7 pairs, equidistant,brochidodromous, above flat to impressed, below slightly raised,divergingat 60-800, straightto evenly arching, chordalangle 50-60?, the angle uniformalong blade length; tertiaryveins above and below inconspicuous to flat, random-reticulate;higher-orderveins above and below inconspicuous; surface above yellowgreen, below reddishbrown;indumentabove absent, below pubescent, the hairs dense, to 1 mm long, straightto curved,erect, reddish,denseron the veins, persisting for at least two flushes. Staminate inflorescences: along whole length of flush, erect, peduncles 2-6 cm long, the hypopodia1-3 cm X 1-1.5 mm, branchorders 3, the second-orderbranches3-6, dispersed, lowest branchto 1 cm long, color and indument of all axes as on twigs; bractspersistentor caducous, to 0.8 mm long, adaxially hairy; bracteoles persistentor caducous,to 4 mm long, adaxiallyhairy. Staminateflowers: pedicels ca. 3.2 X 0.8 mm, the diametereven throughout;receptacleflat, ca. 1 X 3.2 mm; tepals membranaceous,obovate, ca. 2.5 X 2 SYSTEMATICTREATMENT mm, at anthesisspreadingto recurved,reddishbrown, adaxiallypuberulous;stamens of whorls I and II spathulate,the anthersreniform,ca. 1.2 X 1 mm, with a few hairs at base, the locelli 4, apical, in a shallow arch, the upper pair introrse,the lower pair latrorse, the glands absent; whorl III capitate, ca. 1.2 X 0.6 mm, with a few hairs at base, the anthersglobose, the locelli 4, the upper pair latrorse, the lower pair extrorse, the glands globular,ca. 0.7 mm diam.; whorl IV absent;all stamensreddish;pistillode absent.Pistillateflowers:pistil ca. 2.5 X 1.8 mm;ovary globose, ca. 1.5 mm long, glabrous.Fruits: pedicels to 7 X 4 mm, graduallyenlargingto form the cupule; cupule trumpet-shaped,to 10 X 13 mm, smoothto wrinkled, the marginundulate,tepals persisting;berry elliptic, to 15 X 10 mm. Field notes. Shrubsto 2(-4) m tall, alreadyflowering when 1 m tall; leaves above darkgreen, leaves below pale green. Receptacle brownishgreen; tepals yellow-green; stamens/staminodesred?;ovary green. Cupule spreading, coral red; berry pale green with pale spots. Distribution (Fig. 13). Known from the summits of four sandstone mountains-Auyan-tepui, Cerro and CerroJauaGuaiquinima,Cerro Sarisariniama, of the VenezuelanGuayanahighlands, at 1400-200 m. The vegetation is shrublandand meadow on peat accumulationsor rocky soils (Huber, 1995). Apparently with no defined flowering period; the presence of ripe fruits and open flowers at the same time suggests a continuousflowering regime. Additional specimens examined. VENEZUELA. BOLiVAR: Auyan-tepuf,summit, 27 May 1986 (d fl bud), Holst 3028 (MO);29 Feb 1988(Y fl), Huber 12557 (MO, RioParagua, 25 Dec 1952 MYF-n.v.);CerroGuaiquinima, ( fl), Maguire32753 (NY); 7 Jan 1952 (st), Maguire33017 (NY); Auyan-tepui, summit, 10 May 1964 (d fl), Steyermark 93595 (NY, VEN-n.v.); Meseta de Jaua, Cerro Jaua, summit, 22 Mar 1967 (d fl), Steyermark97853 (NY, US); Meseta de Jdua,Cerro Sarisarinfama, summit, 16 Feb 1974 (st), Steyermarket al. 109118 (G, MO). Rhodostemnonodaphne steyermarkianais a very distinct species. Like many upper-montaneplanttaxa of the Guayanahighlands,it is a low shrubwith most organs covered with a dense reddish indument that turnsblack on older branches.It neverthelessis erect, freestanding,andprofuselybranched,unlikethe scandent, little-branchedspecies R. celiana, R. scandens, and R. vinotinctathat occur in the same area. Two specimens from Auyain-tepuf,Steyermark 93935 and S-eyermarket al. 116012, included in the descriptionof Rhodostemonodaphnesteyermarkiana 69 by Allen (1967) are excluded from this species. The leaves of these specimens are characteristically ascending-imbricate,the petioles are shorter,and the venation is slightly acrodromouswith the basalmost pair of secondary veins strongerthan the rest. The anthershave four locelli arrangedin two pairs, indicating a closer affinity to species of Ocotea than to Rhodostemonodaphne.Vegetativelythese specimens resemble Ocoteaferruginea (Meissner)Mez, a highaltitudespecies from the Andes of Peru, or 0. rotundata van der Werff,a recently describedspecies from Ecuador-indeed, some specimens of R. steyermarkiana have been determinedby Bemardi as 0. ferruginea. However, 0. ferruginea has short, crisped hairs, as opposed to the long straightto curved ones of these specimens.These specimensmay in fact represent a differentspecies from eitherof the ones mentioned above. 26. Rhodostemonodaphne leptoclada Madrifinan, Brittonia 48: 49, fig. 3 (A, B, C & D [5 only]). 1996. Type.FrenchGuiana.Saul, MontLa Fumee, 10 Sep 1982 (d fl), Mori, Boom & Keller 14899 (holotype:NY; isotypes: CAY-n.v., MO). Fig. 17C Trees: branchesbasitonic,in axils of cataphyllsor foliage leaves, pseudo-verticillate?;twigs terete,ca. 1 mm diam.; epidermisbrownish,barely visible due to indumentcover; terminalbud plump, ca. 4 X 2 mm; cataphyllscaducous; indumenttomentose, persisting for at least two flushes, the hairs dense, to 0.5 mm long, straightto curved,erect, brownish.Leaves: petioles slender, 0.6-1.2 cm X 0.8-1.4 mm, terete; blades thinlychartaceous,flat,broadlyelliptic, (4-)811 X 1-6 cm; base acute to obtuse, minutely decurrent, 70-130o; apex acute, 70-1 100, ultimately acuminate for up to 1.5 cm; marginplane; primaryvein above slightly raised and rounded,below raised and longitudinally striate; secondary veins 3-5 pairs (slightly triplinerved,indistinguishablefrom tertiary veins towardsapex), closer togethertowardsthe apex, eucamptodromous,aboveshallowlyimpressed,below raised, diverging at 45-60?, evenly arching, chordal angle 20-30?, lowest pair more acute than rest; tertiary veins above inconspicuous,below inconspicuous to slightly raised, random-reticulate;higher-order veins above and below inconspicuous;surfaceabove shiny yellow in young leaves, olive-green in older leaves, the primary vein slightly lighter, below dull yellow-green in young leaves, darkerin older leaves, the veins slightly lighter;indumentabove absent,the primaryvein puberulous,below minutelypuberulous 70 to glabrous, the hairs isolated, to 0.5 mm long, straight,spreading,ascending, translucentto yellowish brown, denser on the veins, caducous after one flush. Staminate inflorescences: basitonic to mesotonic, pendulous?, peduncles 1-6 cm long, the hypopodia 1-4 cm X 0.2-0.4 mm, branchorders3, the second-orderbranches3-5, dispersed,lowest branch to 0.7 cm long, color and indumentof all axes as on twigs; bractscaducous (not seen); bracteolespersistent or caducous, to 1 mm long, adaxially hairy.Staminate flowers: pedicels ca. 3.2 X 0.4 mm, the diameter even throughout;receptacle flat, ca. 1 X 2.3 mm; tepals membranaceous,ovate, ca. 3 X 1.2-1.6 mm (innerwhorl slightly smaller),at anthesisspreading, yellowish, translucent,adaxially tomentose;stamens of whorls I and 11,the anthers,sessile, laminar obovate, ca. 1.4 X 1.6 mm, glabrous, the locelli 4, apical, in a strong arch (whorl II in two almost superposedpairs), introrse,the glands absent;whorl IIl capitate,ca. 1.6 X 0.6 mm, glabrous,the anthersoblong, the locelli 4, the upper pair latrorse,the lower pair extrorse,the glands flattenedand fused, forming a continuousring; whorl IV absent; all stamens yellowish brown;pistillode absent.Pistillateflowers and fruits unknown. Field notes. Trees to 10 m tall and 10 cm diam. Tepals green; stamens/staminodesyellow-green. Distribution (Fig. 13). Knownonly from the type locality, in moist forest, at ca. 300 m. Flower buds were collected in August; the flowers were open in September,at the beginning of the dry season. Additional specimens examined. FRENCH GUIANA:Sail, MontLa Fumee, 18 Aug 1982(d fl), Mori& Boom14737(CAY,NY). FLORANEOTROPICA R. parvifolia these are soon glabrescent.The leaves of R. parvifolia are considerablysmaller than those of R. leptoclada and are glabrous, yet both species have thin, long petioles, a minutelybut conspicuously decurrentlamina, slenderinflorescences,and flowers that are very much alike in both perianthand androecial characters.These similaritiessuggest a close relationshipbetween these two species. As originally noted, the collections from the Tumucumaquerange have shorter,thickerpetioles, and larger bullate leaves than those of either Rhodostemonodaphne leptoclada or R. parvifolia. Furthermore, R. leptoclada has thick, adaxially tomentose tepals, whereas the Tumucumaque specimens are thinnerand adaxially glabrous.The presence of persistent tepals on the cupule of the fruitingR. parvifolia suggests that the yet-unknownfruits of R. leptoclada may have persistent tepals-the tepals and androeciumof the Tumucumaquespecimens dehisce in a ring afteranthesis,and so the cupule presumably has a straightmargin. Based on these differences I have described the specimens from the Tumucumaquerange and Mt. St. Marcel as a new species, R. tumucumaquensis. 27. Rhodostemonodaphne parvifolia Madri-nan, sp. nov. Type. Brazil. Amazonas: Mun. Manaus, Ducke Forest Reserve, 15 Aug 1963 (d fl), Rodrigues 5435 (holotype:NY). Fig. 17D Ex affinitateR. leptocladae, sed staturaminore,ramulis tenuioribus,et foliis parvis subtusvenationeplanadistincta. Shrubs: branches basitonic?, in axils of cataphylls; twigs terete, ca. 2 mm diam.; epidermis Rhodostemonodaphneleptoclada can be recog- brownish;terminalbud slender,ca. 3 X 1 mm; catanized by its slender brown-tomentosetwigs, light- phylls caducous; indument puberulous to absent, green-dryingleaves with white ascending hairs, and caducous after one flush, the hairs dense, to 0.2 mm small green flowers. In the original description of long, straight,spreading,ascending, golden. Leaves: Rhodostemonodaphneleptoclada (Madriniain, 1996), petioles slender, 0.6-1 cm X ca 0.6 mm, adaxially I includedthreecollections (de Granvilleet al. 12377, flattened;blades chartaceous,flat, broadly ovate to 12383 & 12457) from the Tumucumaquerange (N elliptic, 3-8 X 1-5 cm; base obtuse to rounded,minAmapa, Brazil on the borderwith FrenchGuiana)as utely decurrent,90-110'; apex acute to obtuse, 50conspecific with the individual from La Fumee. I 800, ultimately acuminate for up to 2 cm; margin noted considerabledifferencesbetweenthembutcited plane; primaryvein above and below slightly raised; the collection from Mt. St. Marcelas an intermediate. secondary veins (I-)2-3(-4) pairs (strongly tripliThese specimens are now excluded from R. lepto- nerved),closer togethertowardsthe apex, brochidodclada and are considered a new species, R. tumucu- romous, above impressed, below slightly raised, divergingat the firstpairca. 400 (minorsecondaryveins maquensis(see above). A new species from Manaus, Rhodostemono- 600), evenly arching, chordal angle the first pair ca. daphne parvifolia, is very similar to the type of R. 250 (minorsecondaryveins 400); tertiaryveins above leptoclada.They both have slendertwigs, althoughin inconspicuous, below flat to impressed, random- SYSTEMATICTREATMENT reticulate;higher-orderveins above and below inconspicuous;surfaceabove yellowish to greenishbrown, below light yellowish brown;indumentabove absent, the veins tomentose, below minutely puberulous,the hairs isolated, to 0.2 mm long, straight, spreading, ascending,yellowish brown,caducousafterone flush. Staminate inflorescences: basitonic,erect, peduncles ca. 5 cm long, the hypopodiaca. 3 cm X ca. 0.4 mm, branch orders 2, the second-orderbranches 3?, dispersed, lowest branch to 0.5 cm long, color and indumentof all axes as on twigs; bractsand bracteoles caducous (not seen). Staminateflowers: pedicels ca. 2.5 X 0.6 mm, the diametereven throughout;receptacle flat, ca. 0.5 X 1.4 mm; tepals membranaceous, elliptic, ca. 2 X 1 mm, at anthesisspreading,yellowish, translucent, adaxially tomentose; stamens of whorls I and II, the antherssessile, laminar,obovate, ca. 1.2 X 1 mm, glabrous,the locelli 4, apical, in a shallow arch, introrse, the glands absent; whorl III columnar,ca. 1.2 X 0.6 mm, glabrous,the locelli 4, the upper pair latrorse, the lower pair extrorse, the glands fused, forming a continuous ring; whorl IV absent; all stamens reddish brown;pistillode absent. Pistillate flowers unknown.Fruits: pedicels to 14 X 3 mm, graduallyenlargingto form the cupule;cupule trumpet-shaped,to 5 X 7 mm, smooth, the margin undulate,tepals persisting;berry elliptic, to 25 X 15 mm. Field notes. Trees to 6 m tall and 4 cm diam., alreadyfloweringwhen 3 m tall. Tepalsgreen.Cupule red; berry green, ripeningblack. Distribution (Fig. 13). Known only from the Ducke forest reservenearManaus,at ca. 80 m, where it grows in non-inundatedrain forest on sandy to loamy soils. Flowers in August (one collection). FruitsDecember-April, during the rainy season. Additional specimens examined. BRAZIL. AMA- DuckeForestReserve,31 Dec 1963 ZONAS: Mun.Manaus, (fr), Rodrigues5649 (NY); 20 Feb 1964 (fr), Rodrigues 5717 (NY);15 Apr1964(fr),Rodrigues5763 (NY). Etymology. Named for its small leaves [Lat.parvus, small + folium, leaf]. Rhodostemnonodaphne parvifolia is a very distinct species. It is a small shrubwith slendertwigs, and its leaves are small, chartaceous,yellowish to greenish brown, stronglyacrodromous(triplinerved),andlong acuminate.It is closely relatedto R. leptoclada,both having similarflowers,yet the smallerstature,thinner twigs, smallerleaves with stronglyacrodromoussecondaries, and inconspicuous tertiary venation of R. parvifolia, warrantits recognition as a distinct spe- 71 cies. Vegetatively it is indistinguishable from the widespread and heterogeneous Endlicheria gracilis Kosterm. 28. Rhodostemonodaphne antioquiensis Madrifain, sp. nov. Type. Colombia. Antioquia: Mun. San Francisco, Corr. Aquitania, El Topacio, 5 Mar 1990 (d fl), Cardenas,Ramirez& Ciro2522 (holotype: COL; isotypes: JAUM,MO). Fig. 18A Ramulis gracilibus et floribus receptaculisqueminutis cum R. leptoclada congruens, sed indumento ramulorum haud tomentosofoliisque penninerviisdiffert. Trees: branchesbasitonic,in axils of cataphyllsor basal foliage leaves; twigs angular,soon becoming terete, 1-3 mm diam.;epidermisyellowish to brownish; terminalbud slender, ca. 5 X 2 mm; cataphylls caducous; indumentpuberulousto absent, caducous by next flush, the hairs sparse, to 0.2 mm long, straight, spreading, ascending, yellowish. Leaves: petioles slender, 1-1.5 cm X 1-1.6 mm, adaxially flattened;blades coriaceous, flat, broadlyovate to elliptic, 5-12 X 2-6 cm; base obtuse to rounded,minutely decurrent,80-120o; apex acute to obtuse, 601300, ultimately acuminatefor up to 1 cm; margin plane; primary vein above slightly raised, below raised; secondary veins 4-6 pairs, equidistant, eucamptodromous,above slightly raised, below raised, diverging at 50-60', evenly arching, chordal angle 20-25?, the angle uniformalong blade length;tertiary veins above and below slightly raised, randomreticulate; higher-order veins above and below slightly raised; surface above shiny yellowish green, inconspicuously black-dotted, below dull greenish yellow, minutely but conspicuously black-dotted (blackenedsubsidiarycells); indumentabove absent, the primary and secondary veins tomentose, below puberulous, the hairs isolated, to 0.1 mm long, straight, spreading, ascending, yellowish, caducous by next flush. Staminate inflorescences: mesotonic, erect, peduncles 2-7 cm long, the hypopodia 1-3 cm X 0.8-1 mm, branch orders 5, the second-order branches 4-8, dispersed, lowest branch to 1.8 cm long, color and indument of all axes as on twigs; bractsand bracteolescaducous (not seen). Staminate flowers: pedicels ca. 4 X 0.3 mm, the diametereven throughout;receptacleobconical, ca. 0.8 X 1.2 mm; tepals membranaceous,ovate, ca. 1.8 X 1.2 mm, at anthesis spreading,yellowish, translucent,adaxially puberulous;stamensof whorls I and II spathulate,the anthersroundish,ca. 1 X 0.8 mm, glabrous,the locelli 72 FLORA NEOTROPICA :, ^~~~~~~~~~~~- t tRS sxtsom k No - i " ' w* 40833161 B AR.ON -'#C, iS~~~~~~~~~~~~~~~~~~~~.~~~m gf A"t .00"U, FIG. synandra (Nil ~~ ~ ~ ~ ~ ~ '3S .>x ~ ~ ~ 1. A.> Rhdse Plcis ondpn 19) ' s aniqini ir333R . napoes . ist (Qea rdna (C 3 t2). et al 252) B. R. kutin (M di .-1. %.m 6. .......* C. R. SYSTEMATICTREATMENT 73 4, apical, in a shallow arch, introrse,the glands ab- specimens as part of a single new species, distinct sent; whorl III capitate,ca. 1.2 X 0.6 mm, glabrous, from either E. rubriflora or Rhodostemonodaphne the anthersglobose, the locelli 4, the upper pair la- leptoclada. trorse,the lower pairextrorse,the glands globular,ca. 0.5 mm diam.;whorl IV absent;all stamensyellowish brown;pistillode absent.Pistillate flowers unknown. 29. Rhodostemonodaphne kunthiana (Nees) RohFruits: pedicels to 10 x 2 mm, abruptlyenlargingto wer, Mitt. Inst. Allg. Bot. Hamb. 20: 84. 1986. form the cupule; cupule reflexed,to 1 X 3 mm, wrinAcrodiclidiumkunthianumNees, Systema Laurikled, the marginundulate,tepals caducous;berry elnarum269. 1836. Ocotea kunthiana(Nees) Mez, liptic, to 23 X 16 mm. Jahrb.Konigl. Bot. Gart.Berlin 5: 291. 1889. Nectandra kunthiana(Nees) Kosterm.,Meded. Bot. Field notes. Treesto 10-12(-25) m tall and 18 cm Mus. Utrecht25: 19. 1936. Type. FrenchGuiana. diam.; outer barkrough, brown;innerbarkca. 1 mm Withoutlocality, Dec 1824 (Y fl, imm fr), Poiteau thick, yellow; wood light yellow, wood aromatic; s.n. (holotype: B; isotypes: B, G, K-n.v. [photo twigs red. Inflorescenceaxes red; tepals yellow. CuFig. 18B NY neg. 8523]). pule red; berry green. Distribution (Fig. 13). Knownfrom only two collections from the western slopes of the Cordillera Centralof the Andes in Colombia at 925 m; in submontanerain forest on limestone. Flowers in March, towardsthe end of the dry season, fruits ripe in December. Additionalspecimenexamined.COLOMBIA.AN17Dec TIOQUIA:Mun.SanLuis,trailto veredaLaJosefina, 1982 (fr), Cogollo& Estrada290 (JAUM). Etymology. Named after the Colombian department of Antioquia. Rhodostemonodaphneantioquiensis has slender, glabrous twigs and minutely puberulous, penninerved, light-yellow-green-drying leaves that are somewhat glaucous adaxially. It has small flowers with a flat receptacle, similar to those of R. leptoclada, althoughthe latterspecies has a dense, crisped indumenton the twigs and an almost triplinervedvenation. The cupule of the fruiting specimen (Cogollo & Estrada 290) is reflexed, unlike those known in any other species in the genus Rhodostemonodaphne.Cogollo & Estrada290 lacks flowers, and so its conspecificity with the type of R. antioquiensisis not certain. Its fruitsare very similarto those of Endlicheriarubrifiora, a wide-rangingspecies found in Peru, Venezuela, and Colombia, and one specimen of E. rubriflora (Cogollo & Brand 394), was collected in the same generallocality of Rhodostemonodaphneantioquiensis. The leaves of both the type R. antioquiensis and Cogollo & Estrada 290 are smaller, and have longer,thinnerpetioles thanthose of Endlicheriarubrifiora. In flower, the two species are very different; the tepals of E. rubrifloraare glabrous and dry red, and the anthersare two-locellate. Thus, I am guided by vegetative similarities when designating these Aydendronaciphyllum Nees, Linnaea 21: 496. 1848. Type. Guyana. West Demerara-EssequiboCoast: Pomeroon River, above mouth of TapakumaRiver, Sep 1843 (d fl), Schomburgk1424 (holotype:B; isotype: F [frag. ex B?], K). PleurothyriumchrysothyrsusMeissner, in A. L. de Candolle,Prodr.15(1): 169. 1864. Type. Peru.Amazonas: Chachapoyas,1840 (fl), Matthews3031 (holotype: K; isotype: BM X2, G [frag. ex K], OXF). NectandraarnottianaMez non Nees, Jahrb.Konigl. Bot. Gart. Berlin 5: 402. 1889. Based on Pleurothyriumchrysothyrsus(see discussion below). Ocotea cooperi C. K. Allen, J. Arnold Arbor. 26: 335. 1945. Type. Panama.Chiriquf:Bocas del Toro, Changuinolavalley, 1927 (Y fl), G. P. Cooper& Slater 96 = (Y)10277 (holotype: F; isotypes: G [frag. ex F], GH, K, MAD, US). Nectandra meyeriana Lasser, Bol. Soc. Venez. Cienc. Nat. 11: 184, fig. 1948. Type.Venezuela.Aragua: Henri Pittier National Park, 29 Oct 1946 (Y fl), Pittier 15274 (holotype:VEN-n.v.; isotypes: G, US). PleurothyriumcowanianumC. K. Allen, Mem. New YorkBot. Gard. 10(5): 121, fig. 59. 1964. Type.Venezuela. Amazonas: Cerro Huachamacari,Rio Cunucunuma, 4 Dec 1950 (Y fl), Maguire, Cowan & Wurdack29830 (holotype:NY; isotype: F). Trees:branchesbasitonic,in axils of cataphyllsor basal foliage leaves; twigs angular,soon becoming terete,7-12 mm diam.;epidermisblack;terminalbud plumpto slender,ca. 2 X 1 mm; cataphyllspersisting on currentflush or caducous, to 1.5 mm long; indumentpubescentto tomentoseor puberulous,caducous after one flush, the hairs dense to sparse, to 1.2 mm long, curved to crisped, erect, reddish to golden. Leaves: petioles slender to robust, 1-2.5(-5) cm X ca. 5 mm, adaxiallyflattened;blades chartaceous,flat, narrowlyto broadlyovate to elliptic or oblong, (10-) 15-30(-40) X 5-10(-19) cm; base acute to obtuse or rounded,80-110(-140)?; apex acute to obtuse, (40-) 74 70-90?, ultimatelyacuminatefor up to 3.5 cm; margin plane; primary vein above flat to impressed, below prominent;secondary veins 10-16(-20) pairs, equidistant, eucamptodromous(apically brochidodromous), above flat to impressed, below prominent, diverging at 45-600, abruptly arching near margin, chordal angle 35-400, the angle decreasingapically; tertiary veins above flat to slightly raised, below raised, conspicuously scalariform;higher-orderveins above slightly raised, below raised;surfaceabove olive-greento brown,the veins darker,below darkgreen to light brown; indumentabove absent, the primary vein tomentose, below puberulousto pubescent, the hairs dense to sparse, to 1.8 mm long, straight to curved to crisped, erect, yellowish to ferruginous, denseron the veins, persistingfor at least two flushes. Staminate inflorescences: along whole length of flush, erect, peduncles 6-23 cm long, the hypopodia 1-6 cm X 1-4 mm, branchorders 3-6, the secondorder branches 8-16, dispersed, lowest branch to 3(-7) cm long, color and indumentof all axes as on twigs; bractsand bracteolescaducous(not seen). Staminateflowers: pedicels ca. 4 X 1 mm, the diameter even throughout;receptacle globose, ca. 2.8 X 1.5 mm, constrictedat the place of tepal inception;tepals chartaceousto coriaceous, ovate, ca. 2 X 1.6 mm, at anthesisspreading,reddishbrown,adaxiallyminutely puberulous;stamensof whorls I and II spathulate,the anthersreniform,ca. 1.2 X 1.2 mm, puberulous,the locelli 4, apical, in a shallow arch, the upperpair introrse,the lower pairlatrorse,the glandsabsent;whorl III capitate,ca. 1.4 X 0.8 mm, puberulous,the locelli 4, the upperpair extrorse,the lower pair latrorse(to extrorse), the glands globular, ca. 0.8 mm diam.; whorl IV absent;all stamensbrownish;pistillode filiform, ca. 2 X 0.8 mm, glabrous.Pistillate flowers: pistil ca. 3.2 X 2 mm; ovary ovoid to globose, ca. 2.4 mm long, glabrous.Fruits: pedicels to 15 X 3 mm, abruptlyenlargingto form the cupule; cupule hemispherical,to 15 X 20 mm, smooth to tuberculate,the marginstraight,tepals caducous;berry elliptic, to 30 x 20 mm. Field notes. Trees to 15-30(-45) m tall and 2040(-50) cm diam., already flowering when 7 m tall, on maturetrees the crown formed by whorled horizontal branches;buttressesca. 50 cm tall; outerbark smooth (papillose), grey; inner bark ca. 7 mm thick, grainy,mottledyellow and tan;wood yellow to white, rapidly oxidizing orange to black, heartwoodblack, wood aromatic;leaves above shiny green, the veins yellow, leaves below creamywhite to brownishgreen. Pedicels yellow-brown; tepals greenish white; sta- FLORANEOTROPICA mens/staminodes whitish, the glands green; ovary stigma white. Cupule orange-red. Distribution (Fig. 19). Widespreadfrom Costa Rica to central Bolivia, including the Amazon river basin, and the Guianas; mostly on non-inundated, lowland rain forest, but also in the Andes in uppermontaneforest to 2350 m. Flowering specimens can be found throughoutthe year at different localities, but locally, flowering seems to be restricted to the onset of the dry season. In the Amazonbasin the main dry season occurs towards the middle of the year, while in CentralAmerica, northernSouth America, and in the Andean region it is centered on the first months of the year. Fruitingnormally occurs one or two months afterflowering.The seeds germinateimmediately afterdroppingto the ground. Additional specimens examined. COSTA RICA. ALAJUELA:MonteverdeBiological Reserve, Rio Pefias Blan- cas, 25 Feb 1987(9 fl), Haber& Cruz6791 (CR-n.v.,F, MO). CARTAGO:Turrialba,in forestbehindCATIE,18 Mar 1987 (9 fl), Davidse et al. 25600 (MO); (9 fl), Gomez & Herrera23064 (MO).HEREDIA: LaSelva,5 Dec 1980(st), Hammel 10494 (MO); (fr, seedling), Hammel 10693 (MO); 22 Feb 1982 (9 fl), Hammel 11226 (MO); 10 Jan 1973 (9 fl), Hartshorn1082 (MO); 12 Jul 1973 (fr), Hartshorn1256 (MO);27 Feb 1982 (d fl), Perry s.n. (MO,NY). LIMON: Cordillerade Talamanca,Hitoy CerereBiological Reserve, Cerro Bob6cara, 15 Feb 1989 (d fl), Herrera & Chacon 2397 (B, BM, F, GH, MO); Cerro Coronel, E of Laguna Danto, 16 Mar 1987 (st), Stevens et al. 24903 (MO). PUNTARENAS:Cant6n de Osa, Rancho Quemado,Rinc6n, Rio Riyito, 15 Jul 1990 (fr), Herrera 4263 (MO); Corcovado Sirena National Park,Anacardiumtrail, 22 May 1989 (fr), Kernan & P. Phillips 1105 (MO). SAN JOSE:El General basin, Mar 1940 (d fl), Skutch4757 (A, F-n.v., G [frag.ex F], MO, NY-n.v., US). PANAMA. BOCASDEL TORO:Almirante, Cricamola valley, 13 Feb 1928 (d fl), G. P Cooper 498 = (Y)12116 (F-n.v., FHO, G [frag.ex F], K, MAD, S). COLOMBIA. AMAZONAS:PuertoNarinlo,Amacayacd National Park, 25 Feb 1993 (d fl), Madriniin 716 (COL, FMB, GH, MA); Soratama,Rio Apaporis,between Rio Pacoa and Rio Kananari,20 Aug 1951 (9 fl), Schultes & Cabrera 13687 (COL,U, US); Leticia, nearairporton roadside to Tarapaca,8 Sep 1963 (9 imm fr), Soejarto & Cardozo 626 (COL, F, GH, US). ANTIOQUIA: Guatap6,Santa Rita, Finca Montepinar,23 Jul 1987 (9 imm fr), Albert de Escobar et al. 7835 (HUA, MO X2); San Luis, Piedra del Castrill6n, 12 Aug 1987 (9 fl, imm fr), Daly & Betancur 5358 (HUA, MO, NY); Anorn,between Providenciaand Alhibe, 20 Feb 1976 (6 fl), Soejartoet al. 4461 (F, HUA); (6 fl), Soejarto et al. 4502 (F, HUA, NY). CHOc6: Riosucio, Los Katios National Park, 29 Sep 1979 (st), Barbosa 1229 (FMB X3). GUAJIRA: SierraNevada de SantaMarta,Mingueo, bosque La Cueva, 21 Aug 1986 (st), Gentry & Cuadros 55420 (MO, JBGP). META:TinigiuaNational Park, Si- 75 SYSTEMATICTREATMENT R 0 IID _'0 ZID D 4? Rlwdostem~~~~Rh onodaphnekunthianagru proparte * @0 .2 - - 0 '. OR.kunthiana o R.synandra FIG. 19. Distributionmap of partof the species in the Rhodostemonodaphnekunthianagroup. Each symbol represents one or more collections found in a degree-square. erra de la Macarena,Mt. Chamusa,Jul 1993 (fr), Stevenson 500 (MO). Upper Rfo Orinoco, 1951 VENEZUELA. AMAZONAS: (c fl), Croizat917 (NY); Atabapo,Salto Yureba,CanloYureba, Bajo Ventuari, 24 Oct 1981 (d fl), Delascio & Gudnches 10930 (MO); Rfo Negro, Cerro de La Neblina, Rfo Mawarinuma,5 Feb 1984 (d fl), Liesner 15655 (AAU, F, MO, NY, US); Rfo Negro, Cerro de La Neblina, 21 Mar 1984 (fr), Liesner & Stannard 16935 (MO, NY, US); (fr), Liesner & Stannard 16949 (AAU, F, MO, NY, US). ARAGUA: Henri Pittier National Park,9 Aug 1983 (d fl), Field 154 (NY); Nov 1952 (6 fl), Lanes 31819 (K); 5 Apr 1959 (6 fl), Trujillo4122 (U); 30 Nov 1938 (d fl), Williams& Alston 114 (BM); 29 Nov 1938 (T fl), 10718 (F, NY, VENn.v. ); (c3 fl), Williams& Alston 10728 (VEN-n.v. ). BARINAS: Rfo Acequia, SW of jct. of rd. to Ciudad Bolivar, 25 Aug 1966 (d fl), Steyermark& Rabe 96512 (NY); Rfo Zulia, Los Diques, Santa Barbarade Barinas, 31 May 1989 (fr), Valverde& Pefia 1138 (MO). BOLiVAR: Dtto. Cedenio, Along Rfo Mawela, tributaryof the Erebato, 16 Mar 1992 (fr), Boom & Marin 10536 (MO, NY); Dtto. Cedenlo,Rfo Nichare, May 1988 (fr), Elcoro 324 (MO); Dtto. Cedenio, Serraniade Maigualida,20 km E of San Jose de Kayama, Apr 1989 (9 imm fr), Ferndndez5476 (MO, NY); Tabaro river,Dedemai camp, 23 Nov 1991 (d fl), Goldstein& Salas 97 (MO); (fr), Salas TT-205 (MO); woods 3-4 km SE of "Los Patos"N of Rio Hacha and N of Rfo Supamo, 30 km S of El Manteco, 9 Aug 1960 (fr), Steyermark87031 (G, NY); CerroVenamo, 8 Jan 1964 (9 imm fr), Steyermarket al. 92862 (F, G, K, NY X2, P,US). FALCON:CerroLa Mina, Mun. Jacura,Dtto. Acosta, 14 Nov 1979 (fr), Marcano-Berti et al. 448-979 (HBG). MERIDA: El Vigia, Canio Amarillo, 28 Jan 1955 (fr), Bernardi 1862 (G X2, NY). MIRANDA: Cerros del Bachiller, 20 Mar 1978 (fr), Steyermark& Davidse 116883 (MO). ZULIA:Dtto. Bolivar, between El Pensado (town some 4 km E from Quir6s)and Las Tres Marfas, 8 km E from El Pensado, 16 Feb 1980 (fr), Bunting& Stoddart 9056 (NY); Sierrade Perija,QuebradaPerija,tributary of Rfo Tokuku,SW of Los Angeles de Tokukumission, SW of Machiques,29 Aug 1967 (d fl), Steyermark99866 (HBG, NY). GUYANA. MAZARUNI-POTARO: Upper Mazaruni River Basin, Mt. Ayanganna,16 Aug 1960 (fr), Tillettet al. 45660 (NY X2). RuPuNUNI:Wabuwak,KanukuMtns., Oct 76 1948 (6 fl), Wilson-Browne379 = FD 5793 (FDG,NY X2). WITHOUT LOCALITY: 1841 (6 fl), Schomburgk798 (BM, G X2, MO, NY, P). SURINAME. MAROWIJNE:UpperLitany,3 Aug 1993 (stam fl), de Granvilleet al. 12022 (CAY-n.v., MO). SAREmmaKeten, 19 Mar 1922 (fr), Gonggrijp& StaAMACCA: hel 188 (U). STATE NOT INDICATED:Laida, 10 Aug 1908 (9 fl, imm fr), Tresling343 (U). FRENCH GUIANA: Tumuc Humac, 26 Aug 1972 (6 fl), de Granville B-4469 (CAY, HB, NY, U X2); Rivi&re Comte, between Roche Fendee & Be1izon,7 Feb 1973 (fr), de GranvilleB-4667 (CAY X2, NY, P, U); Saut Ananas, 12 Aug 1981 (d fl), de Granville4819 (MO, U); Cayenne, 1804 (fr), Martins.n. [65 in BMJ (BM X4, MO); Saiil, Mont La Fumee, 1 Jun 1986 (d fl bud),Mori et al. 18234 (CAY,MO, NY, U); Saul, Eaux Claires, 26 May 1992 (fl bud), Mori et al. 22324 (MO, US); 28 Oct 1992 (9 imm fr), Mori et al. 22670 (MO); Riviere Comte, Crique Galibi, on CriqueRupert, 22 Jul 1967 (fl bud), OldemanB.1119 (CAY,U); Riviere Comte, between Roche Fendee and Belizon, 21 Jul 1965 (st), Mori et al. 1448 (US). ECUADOR. EL ORO: Hacienda Ingenio, 15 km S of Piedras, 20 Jun 1943 (st), Little 6646 (F, NY). ESMERALDAS:Quininde, Rfo Guayllabamba,5 Oct 1965 (fr), Little & Dixon 21238 (MO, NY X3, US X3); Anchayacu, Eloy Alfaro, Mayronga, 15 Apr 1994 (fr), T D. Penningtonet al. 14950 (MO); San Lorenzo,parroquiaAlto Tambo,sectorEl Cristal, 13 Apr 1992 (d fl), Tipazet al. 794 (G, MO X2). Los Rios: Quevedo,Rio Palenque,2 Oct 1976 (st), Dodson & Gentry 6326 (MO, QCA); (st), Dodson & Gentry 6339 (MO); (d fl), Dodson & Gentry6412 (AAU, MO, NY, QCA X2); 7 Oct 1976 (d fl), Dodson & Gentry 6564 (MO); 5 Mar 1977 (fr), Dodson & Gentry 6658 (AAU, F, MO); 5 May 1983 (6 fl), Dodson & Dodson 13675 (F, MO); 19 Jun 1991 (d fl bud), Palacios & Freire 7412 (MO); 15 Jul 1991 (d fl), van der Werff et al. 12358 (MO). MORONAPachicutza,14 Sep 1975 (9 fl), Littleet al. 355 SANTIAGO: (COL, MO, US); Taisha, 16.7 km NE, 14 Aug 1976 (6 fl), Ortega 005 = 056 (US X2); Taisha,23 Sep 1976 (st), 211 (US); Montalvo, 1 Jul 1989 (6 fl), Zak & Espinoza 4452 (MO). NAPO:Yasuni, 16 Jan 1988 (fr), Ceron 3435 (AAU, C, HBG, INPA,MO, NY, QCA, US); San Jose de Payamino, 27 Apr 1984 (st), Irvine 996 (F); Estaci6n Biol6gica Jatun Sacha, 3 Jul 1987 (6 fl), Palacios 1689 (HBG, MO, NY, QCA, US); Archidona, 20 Mar 1989 (fr), Palacios 4044 (MO); CantonEl Chaco, Las Palmas,Finca Carmita,12 Oct 1990 (6 fl), Palacios 6247 (MO); (9 fl), Palacios 6251 Pastaza,1 Aug 1990 (d fl), Gud(MO, NY-n.v.). PASTAZA: ifio 533 (MO); Rio Tigiiino, 7 Jan 1989 (fr), Hurtadoet al. 1293 (MO, NY). PERU. AMAZONAS: Chachapoyas,Rodriguezde Mendoza, Cochamal,Tinas,Yanamonte,3 Jul 1991 (d fl), D(az et al. 4532 (MO); Rio Santiago,Caterpiza,26 Oct 1979 (fr), Huashikat 1070 (HBG, MO); QuebradaChigki Shinuk, 31 Jan 1973 (fr), Kayap 279 (F, MO); Bongara, 31 Aug 1983 (9 fl, imm fr), D. N. Smith & S. Vdsquez4905 (MO X2); Bagua prov., Marati6nriver, Kusu-Chapicreek, Feb 1995 (sterile), R. Vdsquezet al. 19537 (GH, MO-n.v.); (sterile), La R. Vdsquezet al. 19541 (GH, MO-n.v.). CAJAMARCA: FLORANEOTROPICA Palma, 4 Feb 1988 (st), Gentry et al. 61109 (MO); (Y fl, fr), Gentry et al. 61141 (AAU, F, MO, NY). JUNiN: Santiago, Rio Tambo,18 Apr 1981 (c fl), Reynel-R.197 (F,MO). LORETO:QuebradaTahuayoabove QuebradaTamishiyaco, 27 Aug 1972 (c fl), Croat19790 (AAU, G, F, MO X2, NY, U); Yurimaguas,Rio Huallaga, 11 Feb 1924 (c fl), J. G. Kuhlmann20050 (U); Maynas, Alpahuayo, 14 Nov 1984 (fr), R. Vasquezet al. 5950 (F, HBG, MO, NY); Maynas, Iquitos, Rio Nanay, PuertoAlmendras, 18 Sep 1988 (6 fl), R. Vasquez& N. Jaramillo 11035 (HBG, MO). MADRE DE Dios: Tambopata,29 May 1987 (st), Gentry& N. Jaramillo 57819 (MO X2); Manu, 18 Aug 1989 (Y fl), Ntiuiezet al. 11424 (GH, HBG, MO); Tambopata,7 Jul 1989 (c fl), Phillips et al. 182 (MO); Manu,Puesto Pakitza,4 Oct 1987 (st), Sobrevila et al. 1871 (MO). PASCO:Palcazu, 20 Jun 1198 (Y fl), Hartshornet al. 2630 (MO); Oxapampa,Palacazu, 22 Sep 1986 (st), Pariona & Ruiz 963 (MO); Oxapampa, Bermiudez,10 Nov 1980 (fr), Reynel-R. 39 (MO); Oxapampa, 2 Dec 1982 (fr), D. N. Smith 2886 (MO, NY); 23 Aug 1967 (Y fl), E. Vdsquez119 (K X3, MO X2); (Y fl), 120 (MO, US); Oxapampa,Villa Rica, 27 Feb 1986 ( fl), van der Werifet al. 8292 (MO, NY). SAN MARTiN: Rioja, Venceremos, 10 Feb 1984 (st), Gentry et al. 45294 (MO); (Y imm fr), Gentry & D. N. Smith45342 (MO, NY); 5 Aug 1983 (Y imm fr), D. N. Smith & S. Vdsquez4615 (MO); 7 Aug 1983 (Y imm fr), D. N. Smith & S. Vdsquez4720 (K, MO X2); Mariscal Caceres, Rio Abiseo, 17 Aug 1986 (c fl), Young4072 (F, MO). BRAZIL. AcRE: Brasileia, 2 Jun 1991 (? fl), Daly et al. 6830 (MO, NY); Rio Branco, 7 Jun 1991 (Y fl), Daly et al. 6898 (MO X2, NY); Brasileia, 1 Nov 1991 (fr), Daly et al. 7072 (MO, NY); Sena Madureira,30 Sep 1968 (fr), Prance et al. 7675 (NY); Brasileia,24 Jul 1991 (Y imm fr), Saravia & Rego 1321 (MO, NY). AMAZONAS: Mun. Manaus, Fazenda Dimona, 29 Nov 1989 (fr), Kukle 142 (GH X2, MO, NY, US). PARA:Altamira,5 Nov 1985 (Y imm fr), Balee & B. G. Ribeiro 1848 (MO); Anajas,Ilha de Maraj6, 31 Oct 1984 (Y imm fr), Sobel et al. 4923 (MO). RoNDONIA: Ribeirao, basin of Rio Madeira,25 Jul 1968 (c fl), Prance et al. 6432 (HBG, MO, NY X2, US); Ariquemes, 18 May 1982 (d fl), Teixeiraet al. 578 (F, GH, K, MO, Rio Uraricoera,13 Mar 1979 (fr), Pires et NY). RORAIMA: al. 16962 (NY X2). BOLIVIA. BENI: Prov. Ballivian, 21 Apr 1991 (d fl), Killeen et al. 2885 (GH, LPB-n.v., MO-n.v.); Prov. Ballivian, arroyo San Bernardino,13 Feb 1992 (fl bud), Killeen & Krudenky3611 (MO); Prov. Ballivian, Mar 1990 (st), D. N. Smith et al. 14354 (MO). COCHABAMBA:Prov. Carrasco, Valle del Sacta, 27 Oct 1987 (st), Beck 13676 (MO); 14 Oct 1991 (fr), Killeen et al. 3530 (MO); 11 Jun 1989 (cd fl), D. N. Smithet al. 13691 (MO, NY-n.v.). LA PAZ: Franz Tamayo,Serraniade Chepite, 3 Apr 1992 (st), Killeen 3836 (MO); Prov. Sur Yungas, 1 Jul 1939 (Y imm fr), Krukoff 10173 (A, F, G X2, MO, NY, S, US); Prov. Nor Yungas, 1 Nov 1984 (Y imm fr), Nee & Solomon30297 (AAU, F, MA, MO, NY, SP,U, US). PANDO: Rio Madera,W bank,opposite Abuna (Brazil), 9 Jul 1968 (9 imm fr), Prance et al. 5709 (AAU, HBG X2, K, MO, NY X4, U, US). SANTA CRUZ: SYSTEMATICTREATMENT 77 Prov.Ichilo,Ambor6NationalPark,2 Oct1991(st),I. Var- inflorescences with branches up to three orders of branching(see below). The flowers are very uniform, Local names and uses. Costa Rica: quisarra with acute, wedge-shaped,spreadingtepals, at antheamarilla,quisarranegra; Panama:sweetwood, yaya; sis resemblinga six-pointedstar.The glands of whorl Colombia: canelo; Venezuela:canau-yek (Arekuna); III can be very large,protrudingbeyondthe outertwo Suriname:pisie oema; Ecuador:jigua, jigua amarilla, antherwhorls. In the genus Pleurothyriumall anthers yana quillu caspi yura (= blackening yellow-wood were believed to bear a pair of glands, these often tree, Payamino),yacu ahua yura (= water ahua tree, fusing and forming a glandulardisk, hence the synibid.), wanchuctar;Peru: ishpingo, moena, moena onyms P. chrysothyrsusMeissnerandP. cowanianum amarilla de hoja ancha, roble, roble amarillo, roble C. K. Allen (cf. Allen, 1964, fig. 59) above. Note, playa, tinchi, yuwich; Bolivia: laurel,laurelamarillo, however, that the glands of Pleurothyriumare, as in negrillo; Brazil: aiju'i-heter (Arawete), lapamaen most otherLauraceae,presentonly in whorl III (Roh(Uaicd-Mucajail,louro bofo. The wood is often used wer, 1986; van der Werff, 1993). Although the species is readily distinguishedprifor construction.Irvine (Irvine996) reportsits being commonly used for making canoes, paddles, and as marily on the basis of vegetative characteristicsand firewoodin San Jos6 de Payaminoin AmazonianEc- the very constantfloral morphology,a numberof local variants can be identified. These, however, are uador. linkedby intermediates,thusmakingtheirrecognition Rhodostemonodaphne kunthiana characteristi- as distinct segregatesimpossible. cally has light yellow to whitish wood that rapidly The more distinct local variantsare those plants oxidizes upon exposureto air.The flushingdynamics found in the lowlandsof the Amazonbasin. Typically are also quite distinctive. Flowering individuals do the Amazonianindividualshave a very short,crisped, not show as clear a seasonal break in growth as is dense indument. The staminate inflorescences are characteristic of other species. Flowering occurs much reducedboth in length (rarelyexceeding 12 cm) along the length of the flush from axils of leaves and and branch order number (generally three; if four, cataphyllsof currentand precedingflush. The strong then restrictedto basalmostbranches).These differlateral branches overtoppingthe apical meristem is ences are most striking when comparingspecimens anotherfeaturenot common in the genus. from the eastern Andean foothills with those of the There is strong heteroblastyin this species. Sap- Amazonianlowlands. ling, stump-sprout,and understoryleaves arebroadly In Ecuadorthe separationbetween the two variant elliptic, large (reachingthe upperlimits of the range populationsis most pronouncedin the two versants in both length and width), membranaceous,nearly of the Andes (cf. collections from Rio Palenquewest glabrouswhen expanded,with widely spaced, slowly of the Andes and from JatunSacha to the east). Furarching secondaries, and weakly percurrenttertiary thersouth,althoughthereareno collections fromwest venation. Canopy leaves, on the other hand, are gen- of the Andes (probablya reflection of the drier clierally narrowly elliptic to long-oblong, coriaceous, mate in that region of Peru), the distinctionbetween often with some kind of indument, with narrowly the plants found at higher elevations on the Andean spaced and for the most partstraightsecondaries,and foothills and those of the Amazonian lowlands is maintained.Furthermore,those plantsfromelevations strongly percurrenttertiaries. It is also very probable that leaf morphology is above 1000 m in the states of Amazonas,San Martin, influenced by different ecological conditions where and Pasco represent a third variant with a lanose the plantsgrow. The broadlyelliptic leaf morphology (densely long-pubescent) indument (cf. Diaz 4532, is in general more common in montane or lowland Gentry & Smith 45342, Matthews 3031 [type of P. alluvial soils, presumably high in nutrient content, chrysothyrsus Meissner], D. N. Smith & Vdsquez while the narrowlyelliptic type is often found on sites 4615, 4720, 4905, E. Vdsquez119, 120, and Young where relativelyimpoverishedsoils are likely. 4072). All along the western fringe of the Amazon The inflorescences are strongly dimorphic. Sta- basin only a few intermediatesarefound (Little6646, minate inflorescences are larger and more profusely Palacios 6247, 6251), whereas towardsthe norththe branched than the pistillate ones, generally having Amazonian form gradually intergrades with the branches of fourth order (fifth-orderbranches ob- longer-hairedform with longer, laxer inflorescences. If segregationinto separatespecies were to be jusserved in the lower branches of vigorous inflorescences), although staminate individuals with the tified, I would then consider the specimens from narrowly elliptic leaf type often have reduced Costa Rica, Panama, eastern lowlands of northern gas et al. 1101 (MO). 78 South America, and Andean slopes (excluding those typified by Pleurothyriumchrysothyrsus Meissner, see below) as conspecific. The correctnamewould be Ocotea cooperi C. K. Allen. The lanose specimens from high elevations (above 1000 m) on the E slopes of the PeruvianAndes cited above, would be typified by Pleurothyriumchrysothyrsus.All otherspecimens from the Amazonianlowlands, and those from N Colombia, N Venezuela,and the Guianas,would be typified by AcrodiclidiumkunthianumNees, i.e. Rhodostemonodaphnekunthiana. Van der Werff (199lb) describedRhodostemonodaphnesynandraon the basis of four specimens (one staminate,three pistillate) from the western edge of the Andes and adjacent lowlands of Ecuador and Peru. These are vegetatively indistinguishablefrom the broadly elliptic leaf form of R. kunthiana.However, at the one locality where two specimensof these two species have been collected (the JatunSacha biological station in AmazonianEcuador;Neill & Palacios 7129 = R. synandraand Palacios 1689 = R. kunthiana),the representativeform of R. kunthiana has narrowlyelliptic leaves. Moreover,the inflorescences of R. synandraare peculiarin thatthe axes of all orders are much reducedin length. This, coupled with the largerflowers, results in a much constricted, apparentlydensely floweredinflorescence.The flowers are also very characteristicin having long filaments, those of whorls II and III being partiallyfused into a tube-hence the nameR. synandra.This unique combinationof charactersand lack of intermediates warrantsthe recognitionof R. synandraas a distinct species, althoughit is clearly very close to the variable R. kunthiana. In CostaRica, R. kunthianacan be easily confused with Ocotea stenoneuraMez & Pittier.The name 0. stenoneura is based on two syntypes, J. J. Cooper 10217 (now paratypeof 0. cooperi C. K. Allen-R. kunthiana)and Tonduz13377 (lectotype of 0. stenoneura) (Allen, 1945). Ocotea stenoneurahas a long, decurrent,narrowlyrevoluteleaf base which gives the impression of a long petiole-as in Rhodostemonodaphne kunthiana. Sterile materialfrom the westernlowlands of Colombia with dark-dryingleaves and ferruginousindumenton the lower leaf surfacehas repeatedlybeen identifiedas Rhodostemonodaphnekunthiana.I agree with Rohwer'sannotationsthat the venationis "completely wrong"in that this materiallacks the typical scalariformpatternof R. kunthiana.These specimens probablybelong to Pleurothyriumglabritepalumvan der Werff. FLORANEOTROPICA PleurothyriumchrysothyrsusMeissner, based on Matthews 3031, was erroneously synonymized with Nectandra arnottiana Nees by Mez (1889). The identity of N. arnottiana Nees (-0. arnottiana (Nees) van der Werff) based on Matthews 1429 was clarified by van der Werff (1989a). Nectandra chrysothyrsus (Meissner) Bentham, also cited as a synonym of N. arnottiana Nees by Mez, is an invalid name because the combination was not clearly stated (Rohwer, 1993b). 30. Rhodostemonodaphne synandra van der Werff, Ann. Missouri Bot. Gard. 78: 422, fig. 7. 1991. Type. Ecuador. Napo: Estaci6n Biologica Jatun Sacha, 10 Feb 1986 (d fl), Neill & Palacios 7129 (holotype: MO; isotypes: AAU, F, GH, HBG-n.v., K, MO, QAME-n.v.). Fig. 18C Trees: branching unknown; twigs angular, soon becoming terete, ca. 8 mm diam.; epidermis black, barely visible due to indument cover; terminal bud plump, ca. 1 X 1 mm; cataphylls caducous; indument puberulous, caducous after one flush, the hairs dense, to 1 mm long, straight to curved, erect, golden. Leaves: petioles robust, 2.5-4.2 cm X 0.3-0.6 mm, adaxially flattened; blades chartaceous, flat, broadly obovate to elliptic or oblong, 16-29(-39) X 6-1 1(17) cm; base acute to obtuse, 80-130o; apex acute to obtuse, 80-120o, ultimately acuminate for up to 1.5(2.5) cm; margin plane; primary vein above slightly impressed to flat, below prominent; secondary veins (12-)14-17 pairs, equidistant, brochidodromous, above flat, below prominent, diverging at 45-60?, abruptly arching near margin, chordal angle 45-60?, the angle uniform along blade length; tertiary veins above slightly raised, below raised, conspicuously scalariform; higher-order veins above slightly raised, below raised; surface above shiny brown, below dull creamy brown; indument above absent, the primary vein puberulous, below tomentose, the hairs dense, to 0.4 mm long, straight to curved, erect, yellowish, denser on the veins, persisting for at least two flushes. Staminate inflorescences: along whole length of flush to acrotonic, pendulous, peduncles 10-20 cm long, the hypopodia 3-6 cm X ca. 3.5 mm, branch orders 3, the second-order branches 13-18, dispersed, lowest branch to 3.5 cm long, color and indument of all axes as on twigs; bracts caducous (not seen); bracteoles persistent or caducous, to 2 mm long, adaxially glabrous. Staminate flowers: pedicels ca. 1.2 X 1.4 mm, the diameter even throughout; receptacle globose, ca. 4 X 3.6 mm, constricted at the place of tepal inception; tepals coriaceous, ovate, ca. 2.2 X 1.8 mm, SYSTEMATICTREATMENT at anthesis spreading,yellowish brown, adaxiallypuberulous;stamens of whorls I and II loriform,the anthers oblong, ca. 1.6 X 0.6 mm, puberulous,the locelli 4, apical, in a shallow arch, the upper pair introrse, the lower pair latrorse, the glands absent; whorls II and III fused for two thirdsof their lengths into a tube;whorl III loriform,ca. 2.4 X 0.8 mm, puberulous, the locelli 4, the upper pair latrorse, the lower pair extrorse, the glands minute, ca. 0.4 mm diam.; whorl IV absent; all stamens reddish brown; pistillode filiform, ca. 1 X 0.3 mm, glabrous.Pistillateflowers: pistil ca. 4 X 1.6 mm; ovary globose, ca. 2.8 mm long, glabrous. Fruits: pedicels to 10 X 5 mm, abruptlyenlarging to form the cupule; cupule hemispherical,to 15 X 25 mm, smooth, the margin straight,tepals caducous; berry elliptic, to 40 X 22 mm. Field notes. Trees to 30 m tall and 24 cm diam.; buttressespresent;innerbark ca. 8 mm thick. Tepals green. Distribution (Fig. 19). Eastem foothills of the Andes and adjacentlowlands in Colombia, Ecuador, and Peru; at 250-1600 m, in low-montaneand lowland rain forest. Flowers collected in October,January, and February.Ripened fruits found in January togetherwith flower buds. Additional specimens examined. COLOMBIA. Mt. Chamusa, META:Tiniguianational park,la Macarena, 22 Mar1990(d fl), Stevenson126 (GH). PERU.AMAZONAS: Cenapa,Bagua,26 Jan1967(Y fl, Puerto Inca, Llullapifr), Tillett672-83 (GH). HUANUCO: 642(MO). chis,DANTAS,2 Oct1989(Y fl),Kroll-Saldaha CordilleraAzul, Fundo Cinchona,km 209 rd.,9 Oct 1944(Y fl),Hodge&Lescano Huanuco-Pucallpa I (F). UCAYALI: Local names. Peru: moena negra, and moena amarilla. Rhodostemonodaphne synandra is unique in having the stamensof whorls II and III fused for twothirdsof theirlengths into a tube. This featureis most strikinglyseen in the only staminatespecimenknown (Neill & Palacios 7129), althoughit is also evidentin the pistillate flowers. Furthermore,the stamensof all three whorls have straplike(lorate)filaments.The inflorescencesof R. synandraare peculiarin having all branchaxes reduced,thus producingan elongatedinflorescence with lateral clusters of flowers. Vegetatively, however,R. synandracannot be distinguished at first sight from the yellowish, densely pubescent forms of R. kunthianafrequentlyfound on the E Andean slopes in Ecuadorand Peru; the yellow-brown 79 pubescencecited by van der Werffin the original descriptionis also found on severalothervariantsof R. kunthiana,in particularthe CentralAmericanpopulations as well those east of the Andes in Ecuador. The variationin leaf shape observedin the few specimens available falls well within the range of R. kunthiana;R. synandra has leaves that in size are comparableto the largerleaves recordedfor R. kunthiana. 31. Rhodostemonodaphne napoensis Madriiain, sp. nov. Type. Ecuador.Napo: Aguarico, 27 Nov 1991 (d fl), Quelal 312 (holotype: MO; isotype: Fig. 18D GH). Rhodostemonodaphnekunthianaeet R. negrensisimilis, abillainflorescentiis pendulisomnibuselongatis,abhacantherarum locellislatrorsisdiffert. Trees: branching unknown; twigs angular,soon becoming terete, ca. 3 mm diam.; epidermis black, barely visible due to indument cover; terminalbud plump?,ca. 1 X 1 mm; cataphyllspersistingon current flush, to 0.8 mm long; indumentpuberulous,caducous after one flush, the hairs dense, to 0.3 mm long, straight,erect, reddish.Leaves: petioles robust, 0.7-1.2 cm X 2.5-3.5 mm, adaxiallyflattened;blades thinly chartaceous,flat, broadlyelliptic, 17-27 X 911 cm; base obtuse to rounded,120-140?; apex acute to obtuse, 80-130?, ultimately acuminatefor up to 2 cm; marginplane;primaryvein above flat to slightly raised,below raisedto prominent;secondaryveins 810 pairs, equidistant, brochidodromous,above impressed to flat, below raised, diverging at 45-65?, evenly arching,chordalangle 20-35?, the angle uniform along blade length;tertiaryveins above flat,below slightly raised, random-reticulateto scalariform; higher-orderveins above flat, below slightly raised; surfaceabove grayishbrown,below brown;indument above absent, the primary and secondary veins puberulous,below puberulous,the hairs isolated, to 0.4 mm long, straight,erect,reddishbrown,denseron the veins, caducous after one flush. Staminate inflorescences: basitonic to mesotonic, pendulous?,peduncles 16-30 cm long, the hypopodia5-8 cm X 1.5-2 mm, branchorders4, the second-orderbranches 1419, dispersed,lowest branchto 1.5 cm long, color and indumentof all axes as on twigs; bractsandbracteoles caducous (not seen). Staminateflowers: pedicels ca. 4.8 X 1 mm, the diametereven throughout;receptacle obconical, ca. 2 x 3.1 mm; tepals coriaceous, obovate, ca. 3.5 X 2.5 mm, at anthesisspreading,reddish brown,adaxiallypuberulous;stamensof whorls I and 80 FLORA NEOTROPICA l10= i |9 Rh odaphn ;t00010- ktng 10 R. elepha topus A R. longipetiolata 0 l R . naponsi -Unrelatedseis X R. penduliflora FIG. 20. Distributionmap of partof the species in the Rhodostemonodaphnekunthianagroup and R. penduliflora. Each symbol representsone or more collections found in a degree-square. 11,the antherssessile, oblong, ca. 1.2 X 1.2 mm, with a few hairs at base, the locelli 4, apical, in a strong arch, the upper pair introrse,the lower pair latrorse, the glands absent; whorl III columnar,ca. 1.4 x 1 mm, with a few hairs at base, the locelli 4, the upper pair latrorse,the lower pair extrorse,the glands globular, ca. 0.8 mm diam.; whorl IV absent;all stamens reddish;pistillode absent.Pistillateflowers andfruits unknown. Field notes. Trees to 20 m tall and 20 cm diam., alreadyflowering when 10 m tall. Tepals yellow. Distribution (Fig. 20). Napo department,AmazonianEcuador,at 200-250 m, in primaryrainforest, on boggy soils. Flowering November,during the dry season. Additional specimen examined. ECUADOR. NAPO: Orellana,3 Nov 1989 (d fl bud), Gudifio143 (MO). Etymology. Named afterthe Napo riverandprovince in Amazonian Ecuador. The most characteristicfeature of Rhodostemonodaphne napoensis is its inflorescences; they are longer thanthe leaves, most likely pendulous,andthe secondarybranchesare numerousand little branched. Vegetatively it is very similar to R. negrensis, from the Rio Negro, which can also have elongated inflorescences. However,R. napoensis has yellow flowers while R. negrensis has orange to purple ones; furthermore,the anthersof whorls I and II in R. napoensis have conspicuously latrorse lower locelli, while those of R. negrensis are strictly introrse.Rhodostemonodaphnenapoensis and R. negrensismay, in fact, form a link between the R. scandens andR. kunthiana species groups. 32. Rhodostemonodaphne longipetiolata Madriiain,sp. nov. Type. Ecuador.Napo: Estaci6nBiol6gica JatunSacha, 21 Sep 1990 (d fl), Palacios & Iguago 4454 (holotype: MO; isotypes: GH, MO). Fig. 21A 81 SYSTEMATIC TREATMENT W_2 7 .03 b~~~~~~~~~QAW, 9% , - _~~~~~~~~~~~~~.4 ... __s~~~~~~~~~~~~~~~~~.~ _ E _ - E +v^S |iV _ r? A,'T;; r _~~~~~~~~~~~~~~~~~~~io _ i~~~~~~~~~~~~~~~~~~~~~~~~~~ 3O (Vzuz&Jrmllo 206)D R eleh_antopus(oietal. 1916) W Y*OC4 0199' '%' k. 82 FLORANEOTROPICA Antherismonstrosissed floribusaliterR. kunthianaeaccedens, ramulisfoliisque glabris et petiolis longis bene distincta. Trees: branching unknown; twigs terete, ca. 5 mm diam.; epidermis yellowish; cataphylls caducous; indument absent. Leaves: petioles robust, (2-)3-4 cm X ca. 2.2 mm, adaxially flattened; blades coriaceous, flat, elliptic, (8-)12-18 X (4-)7-9 cm; base acute, 70-90; apex obtuse to rounded, 70-120 (acuminate for up to 1 cm); margin plane; primary vein above flat, below raised; secondary veins 6-8 pairs, equidistant, weakly brochidodromous, above flat, below raised, diverging at 40 60?, evenly arching, chordal angle 15-20?, the angle uniform along blade length; tertiary veins above flat, below slightly raised, random-reticulate; higher-order veins above and below flat to minutely raised; surface above yellowish brown, below light yellowish brown; indument above absent, below glabrous. Staminate inflorescences: basitonic?, erect, peduncles 2-6 cm long, the hypopodia 0.5-3.5 cm X 1.4 mm, branch orders 4. the second-order branches 3-7, dispersed, lowest branch to ca. 1.5 cm long, color and indument of all axes as on twigs; bracts and bracteoles caducous (not seen). Staminateflowers: pedicels ca. 1.4 X 0.5 mm, the diameter even throughout; receptacle obconical, ca. 2.2 X 1.2 mm; tepals membranaceous, ovate, ca. 1.4 X 1.2 mm, at anthesis spreading?, yellowish brown to black, adaxially papillose; stamens of whorls I and II, the anthers sessile, chubby, trapezoid, ca. 0.8 X 0.7 mm, whorl I usually doubled and variously fused with stamens of whorls II and III (see discussion), papillose, the locelli 4, in two almost superposed pairs, the upper pair introrse, the lower pair latrorse, the glands absent; whorl IIl columnar, ca. 1 X 0.8 mm, papillose, the anthers trapezoid, the locelli 4, the upper pair introrse, the lower pair latrorse, the glands absent; whorl IV absent; all stamens reddish; pistillode absent. Pistillate flowers and fruits unknown. glabrous, yellowish-drying twigs, long petioles and yellowish-brown-drying,glabrousleaves. Both inflorescence position and flower morphologyof the only known specimen seem to be anomalous. The inflorescences arisein clustersof differentdegreesof elongation from the axils of foliage leaves, each terminatedby an apparentlyfunctionalterminalbud. They are similarto the inflorescencesfound in variousgenera of the Laureae(e.g., Lindera,Litsea), where the inflorescences are produced on indeterminateshort shoots. This condition is rare in, if not absent from, the Perseeae. Most likely these are the result of expression of axillary buds as vegetative shoots after abortionof the terminalbud, with the productionof inflorescences from their onset. All of the various flowers seen had aberrantandroecialconfiguration.In most of the flowers the stamens of whorl I were duplicatedresultingin a whorlof 6 stamens.These were situatedopposite the outer petal whorl. Each stamen had four locelli. The stamens of whorl II were not differentfrom the "normal"condition found in other species of the genus. Those of whorlIII,however,had introrse-latrorseanthers as opposed to the extrorselatrorseones found elsewhere in the genus. In a few flowers two and more anthersfrom the same or differentwhorlswere fused togetherformingan aberrant double anther.Because only one collection of this species is known, nothing can be said aboutthe constancy of these anomalous forms. Most likely these are the result of some externalinfluencein the development of the inflorescences and flowers-perhaps linked to the fact that the tree is supposedly a forest tree now living in open pastures. Nevertheless, the vegetative charactersused in distinguishingthis species are not unusualin the Lauraceaeand constitutea unique combinationsufficient to distinguish it from any other species in the genus. It resembles R. frontinensis in the shape of the flowers. The stamens are very similarto those of R. kunthiana. Field notes. Trees to 15 m tall. Distribution (Fig. 20). Known only from the type specimen from the Jatun Sacha biological station in Amazonian Ecuador at 400 m. The tree was growing in pastures within rain forest on red soils. The flowering specimen was collected in September towards the end of the rainy season. Etymology. Named for its long petioles [Lat. longus, long + petiolatus, petiolatel. Rhodostemonodaphne longipetiolata can be distinguished vegetatively from any other species by its 33. Rhodostemonodaphne morii Madriinan,Brittonia 48: 52, fig. 4. 1996. Type. French Guiana. Sail, Mont La Fumee, 8 Sep 1989 (d fl), Mori, Fitzgerald, Gebhards & N. Stevens 20850 (holotype: NY; isotypes: CAY-n.v., MO, U, US). Fig. 21B Trees: branchesmesotonic, in axils of basal foliage leaves; twigs terete, 1-3 mm diam.; epidermis black; terminalbud plump, ca. 3 x 2 mm; cataphylls caducous; indument puberulous, caducous by next flush, the hairs sparse, to 0.5 mm long, straight to SYSTEMATICTREATMENT crisped, erect, brownish. Leaves: petioles slender, (0.1-)0.7-1.2 cm X 0.8-1.6 mm, terete;blades thinly chartaceous, flat, broadly elliptic, (4-)9-17 X 2-5 (-7) cm; base obtuse, 50-80'; apex attenuate,70-90o, ultimately acuminatefor up to 1.8(-3.2) cm; margin plane; primary vein above slightly raised and rounded,below prominent;secondary veins (2-)3-5 pairs, often number of secondary veins on the two halves of lamina differing by one, equidistant,brochidodromous, above slightly raised, below prominent, diverging at (45-)55-65?, evenly arching, chordalangle 20-35?, the angle uniformalong blade length; tertiaryveins above flat to slightly raised,below raised, random-reticulateto scalariform;higherorder veins above and below slightly raised; surface above olive-green in young leaves, dark green to black in older leaves, inconspicuouslyblack-dotted, below olive-green,slightly glaucous,the veins brown; indumentabove absent, the primary and secondary veins tomentose,below puberulous,the hairsisolated, to 1.5 mm long, curved, erect, yellowish brown,persisting for at least two flushes. Staminate inflorescences: basitonicto acrotonic,pendulous?,peduncles 4-13 cm long, the hypopodia 1-6 cm X ca. 0.8 mm, branch orders 5(-6), the second-orderbranches3-5, clustered apically (scattered),lowest branchto 4 cm long, color and indument of all axes as on twigs; bractsand bracteolescaducous (not seen). Staminate flowers: pedicels ca. 4.8 X 1 mm, the diametereven throughout;receptacleglobose, ca. 1.6 X 4 mm, constricted at the place of tepal inception;tepals chartaceous, ovate, ca. 0.9 X 1.2 mm, at anthesis erect, blackish, adaxially puberulous;stamens of whorls I and II capitate,the antherstrapezoid,ca. 0.8 X 0.64 mm (whorl II slightly smaller), with a few hairs at base, the locelli 2 to 3, apical, in a shallow arch, introrse,the glands absent;whorl III columnar,ca. 1.2 X 0.8 mm, with a few hairs at base, the locelli 4, the upperpair latrorse,the lower pairextrorse,the glands globular,ca. 0.3 mm diam.; whorl IV absent;all stamens reddish; pistillode absent. Pistillate flowers: pistil ca. 1.6 X 1 mm; ovary ovoid, ca. 1 mm long, glabrous. Fruits: pedicels to 10 X 5 mm, abruptly enlarging to form the cupule; cupule hemispherical, to 9 X 20 mm, smooth to tuberculate,the margin slightly undulate,tepals persisting;berry elliptic, to 18 X 15 mrn. Field notes. Trees to 35 m tall and 35(-45) cm diam., already flowering when 18 m tall; buttresses thick,low or absent;outerbarksmooth(with incipient sculpturing),purplishgray;innerbarkca. 8 mm thick, 83 yellow orange;wood yellow orange, wood aromatic. Tepals cream. Cupule red. Distribution (Fig. 20). Known from four localities in northeasternSouth America, two from coastal forests in Suriname at near sea level, and two from centralFrenchGuianaat ca. 300 m. FlowersAugustOctober,concurrentwith new vegetativeflushes,during the dry season. Immaturefruitsalreadypresentby late September,maturingin December at the beginning of the rainy season. Old cupules can be found as late as May. Additional specimens examined. SURINAME. MABoven-Tapanahony,Awalape creek, Jan 1961 (st), R. Elburg (LBB)8914 (U); Moengo, Rikanau,29 May 1954 (st), Lindeman(LBB)5884(NY); 2 Jun 1954 (st), Lindeman (LBB)5959 (NY); TapanahonyRivier, 4 km above mouthof PaloemeuRivier,Aug 1959 (st), Schulz(LBB)8178 (AAU, NY, U). SARAMACCA: SaramaccaCamp,Coisepyba Rivier, 29 Jul 1962 (st), Allen 67 (NY). FRENCH GUIANA: Mont Alikene, 29 Sep 1960 (Y fl, imm fr), Irwin 48584 (NY, US); Sail, Mont La Fumee, 2 Oct 1982 (Y fl), Mori et al. 15031 (CAY,NY); 3 Oct 1982 (d fl), Mori et al. 15044 (CAY,MO, NY [spirit coil.]); 23 Oct 1982 (fr), Mori et al. 15122 (CAY,MO, NY, US); 6 Dec 1982 (fr), Mori & Boom 15334 (CAY,GH, MO, NY [with color photo], US); 25 May 1986 (fr), Mori & T D. Pennington 18147 (CAY,MO, NY). ROWLJNE: Local name. Suriname:pisie. moriiis characterizedby its Rhodostemonodaphne dark brownish-green- to black-drying, membranaceous leaves, and tepals thatare erect at anthesis.The latter featureis unique in the genus. Locelli number in staminal whorls I and II is variable, as in R. napoensis. 34. Rhodostemonodaphne sordida Madrinfan,sp. nov. Type. Peru. Loreto: Maynas, Iquitos, Quistococha, 28 Apr 1989 (d fl), R. Vdsquez & N. Jaramillo 12066 (holotype: MO; isotype MOn.v.). Fig. 21C Rhodostemonodaphnemorii nisi tepalis expansis et reflexis similis. Trees: branchesbasitonic, in axils of cataphylls, pseudo-verticillate;twigs terete, 3 mm diam.;epidermis black; terminal bud plump, 4 X 3 mm; cataphylls caducous; indumentpuberulous to tomentose, caducous after one flush, the hairs dense to sparse, to 1 mm long, straight to crisped, erect, yellowish to reddish. Leaves: petioles slender to robust, 0.5-1.2 cm X 2-3 mm, terete; blades chartaceous, flat, elliptic, 84 5-12 X 4-6 cm; base rounded to obtuse, 90-150'; apex roundedto acute, 60-100', acuminatefor up to 2.5 cm; marginplane;primaryvein above raised,below prominent;secondary veins 4-6 pairs, equidistant, brochidodromous,above flat, below raised, diverging at 600, evenly arching,chordalangle 300, the angle uniformalong bladelength;tertiaryveins above inconspicuous, below slightly raised, randomreticulate; higher-orderveins above inconspicuous, below slightly raised; surface above light to dark brown (mottled),below darkbrown;indumentabove absent, the primary vein tomentose, below puberulous, the hairs sparse, to 1 mm long, straight to curved, erect, yellowish brown, denser on the veins, caducous after one flush. Staminate inflorescences: acrotonic, erect, peduncles 4-8 cm long, the hypopodia 1.5-5 cm X 1-1.5 mm, branch orders 6, the second-orderbranches2-8, dispersedto clusteredapically, lowest branch to 0.6-2 cm long, color and indumentof all axes as on twigs; bractsandbracteoles caducous (not seen). Staminateflowers: pedicels 1.2 X 0.4 mm, the diametereven throughout;receptacle obconical to flat, ca. 1 X 1.5 mm; tepals chartaceous, ovate, ca. 1 X 1.2 mm, at anthesisrecurved,yellowish brown,adaxiallypuberulous;stamensof whorlsI and II capitate,the anthersoblong, ca. 0.8 X 0.5 mm, with a few hairs at base, the locelli 4, apical, in a strong arch, introrse,the glands absent;whorl III columnar, ca. 1.2 X 0.5 mm, with a few hairs at base, the locelli 4, the upperpair latrorse,the lower pair extrorse,the glands globose, ca. 0.6 mm diam.; whorl IV absent; all stamensreddishbrown;pistillode absent.Pistillate flowers andfruits unknown. Field notes. Trees to 15-18 m tall and 20 cm diam. Tepals green, maturingred. Distribution (Fig. 20). Known from the vicinity of Iquitosin Peruandof Manausin centralAmazonia, at ca. 100 m, in non-inundatedrain forest. Flowers found in April and May. Additional specimen examined. BRAZIL. AMAZONAS: Mun.Manaus,DuckeForestReserve,18May1965 (d fl), Rodrigues6938 (G, NY). Etymology. The epithet refers both to the indument on all plant parts and to the color of the dried leaves, which togethergive the specimens a dirty appearance[Lat. sordidus, sordid]. Rhodostemonodaphnesordida is similar and possibly closely related to R. morii. Both species have similarindument,leaf texture,venation,inflorescence position on the flush, and inflorescencemorphology. They differ, however,in the color of the indumentof FLORANEOTROPICA the leaves, that of R. sordida being yellowish brown, and especially in the shape of the flowers,those of R. morii being urceolate, with minute, erect tepals and those of R. sordida having larger, spreading to reflexed tepals. Rhodostemonodaphnesordida is found in western Amazonia, while R. morii is a species of the Guianas. 35. Rhodostemonodaphne elephantopus Madritahn,Brittonia48: 46, fig. 2. 1996. Type. French Guiana.Saul, Mont La Fumee, 1 Oct 1989 (d fl), Mori, Gebhards,Fitzgerald & Settle 20774 (holotype: NY; isotypes: CAY-n.v., GH, MO). Fig. 21D Trees: branchesbasitonicto mesotonic,in axils of cataphyllsor basal foliage leaves; twigs angularand remainingso for at least two flushes, 1-3 mm diam.; epidermisbrownishto black;terminalbud slender,ca. 5 X 4 mm; cataphyllscaducous;indumentsericeous to glabrous,caducousby next flush, the hairs sparse, to 0.5 mm long, straight,appressed,ascending,silver. Leaves: petioles slender,0.2-1.3 cm X (0.8-)1.5-1.8 mm, adaxiallyflattened;blades chartaceous,flat,narrowly ovate to obovate, (2-)4-10 X 1-4 cm; base cuneate to acute, minutely decurrent,50-70?; apex obtuse, 80-110?, mucronate; margin minutely recurved;primaryvein above and below flat to slightly raised; secondary veins (4-)6-7(-9) pairs, equidistant,brochidodromous,above and below flat, diverging at 60-80?, straight,the angle uniformalong blade length; tertiary veins inconspicuous, randomreticulate; higher-orderveins visible only in transmitted light; surfaceabove and below brown,the primary veins above slightly darker, inconspicuously black-dotted,below primary vein lighter at base of lamina, darkerapically; indumentabove absent, below minutely puberulous,silver. Staminate inflorescences: along whole length of flush, erect, peduncles 2-8 cm long, the hypopodia 1-5 cm X ca. 0.8 mm, branchorders4, the second-orderbranches5-7(-8), dispersed, lowest branch to 1.5 cm long, color and indumentof all axes as on twigs; bractsandbracteoles caducous (not seen). Staminateflowers: pedicels ca. 2.4 X 0.8 mm, the diametereven throughout;receptacle globose, ca. 2 X 2 mm, constrictedat the place of tepal inception; tepals chartaceous,ovate (margin slightly recurved), ca. 1.6 X 1.5 mm (inner whorl slightly larger),at anthesis spreading,reddish,adaxially papillose; stamens of whorls I and II capitate, filamentlong, the anthersreniform,ca. 1.4 X 1.4 mm (whorl II slightly larger),glabrous,the locelli 4, apical, in a shallow arch, introrse, the glands absent; SYSTEMATICTREATMENT whorl III filamentous, capitate, ca. 1.4 X 0.6 mm, with a few hairs at base, the anthers trapezoid, the locelli 4, the upper pair latrorse, the lower pair extrorse, the glands globular, ca. 0.8 mm diam.; whorl IV absent (staminodial, staminodes columnar); all stamens yellowish brown; pistillode teardrop-shaped, ca. 1.4 X 0.4 mm, glabrous. Pistillate flowers: pistil ca. 1.5 X 0.7 mm; ovary ovoid, ca. 1 mm long, glabrous. Fruits unknown. 85 b _ _: I i ~~~~~~~~~~~~~~~~~ _~~~~~~~~~~~~o Field notes. Trees to 40 m tall and 80 cm diam.; buttresses thick, low, symmetric; outer bark smooth, peeling in large plates towards base, orange, with red concentric bands; wood aromatic. Tepals green. Distribution (Fig. 20). Known only from central French Guiana in the vicinity of the hamlet of Saul, at ca. 300 m, where it grows in non-flooded moist forest. Flowers August-September, at the end of the rainy season. Additional specimens examined. FRENCH GUIANA: Sail, Mont La Fum6e, 27 Aug 1988 (d fl), Mori et al. 19196 (CAY,MO, NY); ( fl), Mori et al. 19199 (CAY, MO, NY [spiritcoil.]). Rhodostemonodaphne elephantopus has pointed terminal buds covered with silver, appressed hairs; it is otherwise conspicuously glabrous. It has small leaves with a rounded, mucronate tip. It is vegetatively very similar to R. avilensis, Ocotea ceanothifolia (Nees) Mez, and 0. congestifolia Lasser, particularly in leaf size and shape. It differs from Rhodostemonodaphne avilensis both in habit (R. avilensis is a small shrub), and in its less conspicuous venation. Ocotea ceanothifolia has smaller inflorescences and shorter pedicels, while 0. congestifolia differs most obviously in its leaves, which are conspicuously clustered at the tips of the branches. 36. Rhodostemonodaphne grandis (Mez) Rohwer, Mitt. Inst. Allg. Bot. Hamb. 20: 84. 1986. Endlicheria grandis Mez, Jahrb. Konigl. Bot. Gart. Berlin 5: 124. 1889. Nectandra grandis (Mez) Kostermans, Meded. Bot. Mus. Utrecht 25: 17. 1936. Type. French Guiana. Fleuve Maroni [St. Laurent?], 1863 (fr), Melinon 1863 [1865 in B] (lectotype, here designated: B; isolectotypes: NY, P). Fig. 22 Trees: branches basitonic, in axils of cataphylls; twigs angular, soon becoming terete, 5-7 mm diam.; epidermis black, barely visible due to indument cover; terminal bud slender, ca. 9 X 6 mm; cataphylls caducous; indument sericeous, caducous after one flush, FIG. 22. Rhodostemonodaphnegrandis (BW 6908). the hairs dense, to 0.2 mm long, straight,appressed, ascending, golden to silver. Leaves: petioles robust, 1.9-4.4 cm X 2-3.8 mm, adaxially flattened; blades chartaceous,flat, broadly elliptic, 10-33 X 5-12 cm (sapling leaves to 47 X 21 cm); base obtuse to rounded,70- 10O; apex rounded,80-1400, mucronate to acuminate for up to 1.8 cm; margin plane to recurved;primaryvein above flat and as wide as petiole at base, slightly raised and narrowtowardsapex, below prominent;secondary veins 6-10 pairs, equidistant, eucamptodromous,above flat to slightly raised, below prominent,diverging at 40-60', abruptlyarching near margin,chordalangle 15-30', the angle uniform along blade length; tertiaryveins barely distinguishable from higher-ordervenation, above slightly raised, below raised, random-reticulateto scalariform; higher-orderveins above and below slightly raised;surfaceabove darkbrownin young leaves (the veins silver), brownto olive-greenin older leaves, the veins darker,below yellowish brown to silver; indument above absent,the primaryvein silver sericeous, below minutely sericeous, the hairs dense to sparse, to 0.28 mm long, straight,appressed,ascending, yellowish, the veins glabrescent,caducousafterone flush 86 FLORANEOTROPICA to persistingfor at least two flushes.Staminate inflorescences: along whole length of flush, erect, peduncles 7-17 cm long, the hypopodia 2-7 cm X 1.6-2 (-3) mm, branch orders (4-)5-6, the second-order branches7-13, dispersed,lowest branchto 2.5(-3.5) cm long, color and indumentof all axes as on twigs; bractsand bracteolescaducous (not seen). Staminate flowers: pedicels ca. 4.8 X 1 mm, the diametereven throughout;receptacle globose, ca. 2.8 X 1.6 mm, constrictedat the place of tepalinception;tepalschartaceous, ovate, ca. 2.4 X 2 mm (inner whorl slightly smaller),at anthesis spreadingto recurved,yellowish brown,adaxiallypuberulous;stamensof whorlsI and II minutely spathulateto lacking a distinct filament, the anthers ovate, ca. 1 X 1 mm (whorl II slightly smaller),with a few hairsat base, the locelli 4, apical, in a shallow arch, introrse,the glands absent;whorl III columnar,ca. 1.2 X 0.6 mm, with a few hairs at base, the locelli 4, the upper pair latrorse,the lower pair extrorse,the glands globular,deeply furrowedto divided, ca. 0.4 mm diam.; whorl IV absent; all stamens yellowish brown;pistillode filiform, ca. 0.4 X 0.1 mm, hairy at base. Pistillateflowers: pistil ca. 1.6 X 0.8 mm; ovary ovoid, ca. 1 mm long, glabrous. Fruits: pedicels to 14(-16) X 8 mm, abruptlyenlarging to form the cupule; cupule hemispherical,to 17 X 25 mm, walls thicktuberculate,the marginstraight, tepals caducous;berry ovoid, to 35 X 22 mm. Field notes. Trees to 30 m tall and 40 cm diam., already flowering when 15 m tall; buttresses low; outer bark rough, warty,with horizontalscars; lenticels 4-5 mm across, darkbrownor gray and spotted; inner bark ca. 15 mm thick, fibrous, yellow-brown, becoming reddishbrown;wood white, with unpleasant smell or non-aromatic;leaves above dark green, the primary vein yellow-green, below light green to whitish, the veins yellow-green. Inflorescence axes whitish green; pedicels reddish; tepals yellow; stamens/staminodesred; ovary stigma white when receptive. Cupuletinged with red. Donselaar 2425 (AAU); Brownsberg,Bosch ForestReserve, 6 Sep 1915 (o' fl), Stahel & GonggrijpBW 629 (MO, U); (c fl bud), Stahel & GonggrijpBW 670 (MO, U); Jodensavanne, Mapane Kreek, 12 Jul 1962 (st), Allen 41 (NY); (st), 51 (NY); 16 Jan 1963 (fr), Boerboom (LBB)9584 (K, MO, NY, U); 1963 (st), J. P. Elburg(LBB)9470(U X2); 30 Apr 1953 (st), Lindeman(LBB)3738 (NY, U); 3 Oct 1953 (st), Lindeman(LBB)4826 (U); (st), Lindeman(LBB)4841 (MO, U); 3 Oct 1955 (st), Lindeman(LBB)6763 (NY); 16 Jan 1963 (d fl), Schulz(LBB)7419(C X3, NY, U, US); 1956 (st), Schulz (LBB)7521 (MO, U); (fr), Schulz (LBB)8264 (NY X2, U); Oct 1960 (J fl), Schulz (LBB)8455 (F, MO, NY X2, U); Aug 1961 (st), Schulz (LBB)9023 (MO, U). Awalapecreek,Feb 1961 MAROWIJNE:Boven-Tapanahony, (st), R. Elburg (LBB)8802 (U); Moengo, Rikanau, 1 Jun 1954 (st), Lindeman (LBB)6031 (U). NICKERIE:Kayser Mountains,5 Nov 1976 (Y fl), Mori & Bolten 8612 (HMB, Emma Keten, 30 Jul 1959 (st), Daniels NY). SARAMACCA: & Jonker805 (U). FRENCH GUIANA: Acarouanyrd., km 3, 3 Oct 1957 (Y fl), BAFOG361-M (CAY,NY, U); Mana rd., Place No. 4, CarreauNo. 42, 3 Feb 1956 (fr), BAFOG7241 (CAY,NY, U); Acarouanyrd., km 3, 29 Mar 1956 (fr), BAFOG 7425 (CAY,NY, U); Crique Arataye, Saut Parare,23 Aug 1983 (st), Barrier 4003 (B-n.v., CAY, COL-n.v., IAN-n.v., Kn.v., MO, NY-n.v., U-n.v., US-n.v.); CriqueArataye,Sauts Parare,2 Dec 1985 (st), Barrier 5153 (CAY,MO); Cayenne rd., km 11.51, 14 Sep 1955 (d fl), Bena BAFOG 307-M (CAY,NY, U); Riviere Comt6,right flank,600 m from Saut Boeuf, 17 Jan 1957 (st), Bna 1300 (CAY,U); Gourdonville, 27 Sep 1914 (st), Benoist 1620 (NY); Saul, Eaux Claires, 10 Sep 1994 ( fl, fr), Mori et al. 23998 (CAY-n.v., GH, NYn.v.); Saul, Boeuf Mort,27 May 1971 (st), OldemanB.3923 (MO, US); St. Elie rd., km 16 Chablis "CEC",8 May 1982 (st), Riera 488 (MO); CriqueArataye,Sauts Parare,29 Oct 1983 (fr), Sabatier 573 (CAY);Saul, Layon Roche Bateau, 10 Mar 1985 (st), Sabatier 1061 (CAY); Crique Arataye, SautsParare,17 Oct 1987 (st), Sabatier1473 (CAY);Crique Arataye, Nouragues station, 28 Oct 1987 (st), Sabatier & Prevost 1870 (MO X2); Fleuve Sinnamarybasin, PetitSaut, 7 Sep 1991 (st), Sabatier & Prevost 2193 (CAY-n.v., MO, P-n.v., U-n.v.); St. Elie rd., Interfleuve SinnamaryCounamama,5 Aug 1991 (st), Sabatier & Prevost 3683 (CAY-n.v., MO). BRAZIL. AMAPA: Rio Falsino, Reserve INCRA, 22 Aug 1983 (st), Campbellet al. 14647 (MO); Macapa, 13.6 km SSW of Cupuxi, 5 Jan 1985 (fr), Mori & J. Cardoso 17717 (F, MO, NY); Oiapoque-Caloene rd., 110 km NW of Caloene, 9 Dec 1984 (fr), Rabelo & R. Cardoso 2941 (GH, MO, NY, US). Distribution (Fig. 23). NortheastemSouthAmerica in Suriname, French Guiana, and the Brazilian stateof Amapa',from sea level to ca. 600 m in lowland rain forest. Flowers (June-)September-November (-January), during the dry season. Fruits (SeptemLocal names. Suriname: papaja-pisi, grootber-)December-February(-March),at the beginning bladige-zwarte[harde]-pisi, watjarang (Caribbean). of the rainy season. Additionalspecimensexamined.SURINAME.BROKOPONDO:Brownsberg,Bosch ForestReserve, 10 Sep 1917 (d fl), BW3204 (U); 27 Oct 1921 (d fl), BW5546 (U); 20 Jul 1925 (cd fl), BW 6908 (U); Brownsberg,8 km ESE of village, 24 May 1965 (st), van Donselaar 1891 (U); (st), van French Guiana: baaka-apisi, gueli[geli]-apisi maccan); cedre jaune, cedre noir. (Para- Rhodostemonodaphne grandis as circumscribed here in the strict sense constitutes only a minor part of R. grandis s.l., a widespread, vegetatively very var- 87 SYSTEMATICTREATMENT so . so 60 so 40. ~~ ~~~~0 9 500 10D006 10~~~~~~~. Rhodseonodaphne gra 'is group & * sr.A Rh.praeclara 20 A Rh.praeclara s. str.0\q UI Rh.recurva1}! C iZ_ m 1 -I1 FIG. 23. Distributionmap of species in the Rhodostemonodaphnegrandis group. Each symbol representsone or more collections found in a degree-square. iable but intergradingcomplex that includes at least six species in the genera Endlicheria and Rhodostemonodaphne(see "The Rhodostemonodaphnegrandis complex" section above). This complex is characterized by silver sericeous indument on most organs, with usually large leaves, althoughthey vary greatly in size and shape. Their inflorescencesare at first sight very similar,but careful dissection reveals considerable floral differences between the different taxa. Vegetatively Rhodostemonodaphnegrandis s.str. can be distinguished from the other species of the complex by its leaves that are on averagewider,have more obtuse bases, and high number of secondary veins. Othercharactersused to distinguishthe species in the R. grandis complex are listed in TableV. Rhodostemonodaphnegrandis is found growingin sympatrywith one other species of the complex, R. saulensis. Nevertheless, the differences between R. grandis and R. saiilensis are sharp. Rhodostemono- daphne saulensis has leaves with an acute, often revolute leaf base, and the lowest numberof secondary veins in the complex (3-6). Furthermore,the tepals of R. grandis are yellow, while those of R. saiilensis are red. Furthercollections may show range overlap with the widespread R. praeclara. This variable species has on averagesmallerleaves thanthose of R. grandis and fewer secondary veins (only 4-6). Furthermore, the absence of pairedglands on the stamensof whorl III in R. praeclara is a very distinctivedifferencebetween them. A large numberof collections of this species have been made from saplings. The extreme variabilityof sapling leaves often makes identificationimpossible. Thus the determinationof saplings as Rhodostemonodaphne grandis is only tentativeand based on other information,such as being collected in the same localities where adult R. grandis are found or having the same local names. Most of the specimens from 88 FLORA NEOTROPICA the Amazonianlowlands previouslydeterminedas R. grandis in fact belong to Endlicheria metallica and Rhodostemonodaphnepraeclara. The great majority of these are in fruit, which is indistinguishablein the three species. Mez (1889) originally described this species as Endlicheria?grandis [sic], based on a single fruiting collection. He placed it at the end of his subgenus Euendlicheria,where he included a numberof sericeous species, including Endlicheria sericea Mez, and a now synonym of E. sericea, E. guadaloupensis Mez. Kostermans(1936: 17) afterhaving seen flowers with the typical Nectandra-likefour-locellateanthers transferredEndlicheria grandis to Nectandra as N. grandis. He also synonymizedN. praeclara Sandwith and N. dioica Mez with N. grandis. He did not, however, note the absence of glands from N. praeclara. In the discussion of N. dioica he dismissed the relevance of the indument type, but as Rohwer (1986) correctly stated, in Rhodostemonodaphne dioica alone the indument consists of two types of hairs, appressedand erect. 37. Rhodostemonodaphne sauilensis Madrinakn, Brittonia48: 60, fig. 8. 1996. Type. French Guiana. Sail, Mont La Fumee, 1 Oct 1982 (d fl), FIG.24. Rhodostemonodaphnesaiilensis (Mori et al. Mori, Boom, Prance & Boeke 15014 (holotype: 20797). NY; isotypes: CAY,K, MO, U, US). Fig. 24 Trees: branches basitonic, in axils of cataphylls; twigs angular,soon becoming terete, 3-4 mm diam.; epidermisblack,barelyvisible due to indumentcover; terminal bud slender, ca. 5 X 3 mm; cataphylls caducous; indumentsericeous, caducous by next flush, the hairs dense, to 0.2 mm long, straight,appressed, ascending, golden to silver. Leaves: petioles slender, 1.5-3.2 cm X 1.4-2.4 mm, adaxiallyflattened;blades chartaceous,flat,narrowlyelliptic to obovate,(9-)1417(-21) X (2-)4-7 cm; base cuneate to acute, decurrent, 40-80?; apex acute, 50-1 10?,ultimatelyacuminate for up to 2.5 cm; margin at base recurved; primaryvein above flat, below prominent;secondary veins (3-)4-5(-6) pairs, equidistant,eucamptodromous, above slightly raised, below prominent,diverging at (35-)45-55?, abruptly arching near margin, chordal angle 10-20(-30)?, lowest pair more acute than rest; tertiary veins above and below slightly raised, random-reticulateto scalariform;higher-order veins above flat, below slightly raised; surfaceabove darkbrownto black, below darkbrown,slightly glaucous; indumentabove absent, below minutely sericeous, the hairs sparse, to 0.2 mm long, straight,appressed, ascending, yellowish brown, caducous after one flush. Staminate inflorescences: along whole length of flush, erect, peduncles 5-1 1 cm long, the hypopodia 2-6 cm X 0.8-1.2 mm, branchorders3(4), the second-order branches (6-)7-9(-10), dispersed, lowest branch to 2 cm long, color and indument of all axes as on twigs; bracts and bracteoles caducous (not seen). Staminateflowers: pedicels ca. 5.6 X 0.8 mm, the diametereven throughout;receptacle globose, ca. 2.4 X 2.4 mm, constricted at the place of tepal inception;tepals chartaceous,ovate, ca. 2 X 2 mm, at anthesis recurved, reddish, adaxially puberulous;stamens of whorls I and IH, the anthers sessile, roundish, ca. 1.4 X 1.4 mm, glabrous, the locelli 4 (median ones vestigial), apical, in a shallow arch, introrse,the glands absent;whorl III columnar, ca. 1.6 X 1 mm, glabrous, the locelli 4, the upper pair latrorse, the lower pair extrorse, the glands minute, ca. 0.2 mm diam.; whorl IV absent; all stamens reddish; pistillode absent (minute, linear, ca. 0.3 mm long, hairy). Pistillate flowers: pistil ca. 2 X 1 mm; ovary ovoid, ca. 1.2 mm long, glabrous. Fruits: pedicels to 10 X 7 mm, gradually enlarging to formnthe cupule; cupule hemispherical, to 15 cm 22 mm, SYSTEMATICTREATMENT 89 smooth to wrinkled, the margin straight, tepals caducous; berry elliptic, to 30 X 15 mm. Some flowers of the type specimen of R. saiulensis have anthersof whorl I with only the two lower locelli fully developed, while the upperones are vestigial. Field notes. Trees to (15-)25-30(-42) m tall and (28-)34(-40) cm diam.; buttresses small or absent; outer bark smooth, gray, with hoops and prominent 38. Rhodostemonodaphne praeclara (Sandwith) lenticels; inner bark ca. 6 mm thick, creamy orange, Madrifian, comb. nov. Nectandra praeclara Sandgranular; wood bright yellow; leaves below light with, Kew Bull. 46: 224. 1932. Type. Guyana. green to whitish,the veins distinctlyred. Pedicels redMazaruni-Potaro:Essequibo River, Wallaba fordish; tepals red with whitish margin;stamens/stamiest, MoraballiCreek, 7 Oct 1929 (6 fl), Sandwith nodes deep red to purple; ovary yellowish green, 387 (holotype: K; isotypes: F [frag. ex G], G stigma white. Cupule red. [photo F neg. 27606], K, P, S [photo NY neg. Fig. 25 8558], U, US). Distribution (Fig. 23). Known only from the vicinity of the town of Saul in central French Guiana, Trees: branches basitonic, in axils of cataphylls; at 250-410 m in non-inundatedrain forest. Flowers twigs angular, soon becoming terete, 4-5(-6) mm late August-October, during the dry season. Fruits diam.;epidermisbrownishto black, barelyvisible due ripe by April, at the height of the rainy season. to indument cover; terminal bud slender, ca. 7 X 3 Additional specimens examined. FRENCH GUI- mm; cataphylls caducous; indumentsericeous, caduANA: Sauil,Mont La Fumee, 16 Sep 1982 (st), Boom & cous by next flush, the hairs dense, to 0.3 mm long, Mori 1722 (CAY,NY); May 1982 (seedling), de Granville straight, appressed, ascending, golden to silver. 5077-A (CAY); 6 Apr 1982 (fr),), de Granville 5077 Leaves: petioles robust, 1.5-2.8 cm x 2.2-3.6 mm, (CAY); 28 Aug 1982 (d fl), Mori & Boom 14787 (clonotypes CAY,K, MO, NY, U, US); 9 Sep 1982 (d fl), Mori et al. 14896 (CAY,NY, U, US); 28 Aug 1989 (Y fl), Mori & Wightman20744 (CAY,MO, NY, U); 19 Mar 1989 (Y fl, fr), Mori et al. 20797 (CAY-n.v., GH, MO, NY, US); 15 Sep 1989 (Y fl), Mori et al. 20904 (CAY-n.v., MO, NY, US); Saul, SentierBotanique, 17 Aug 1993 (fr), Mori et al. 23321 (NY); Saul, Eaux Claires, 4 Sep 1994 (Y fl), Mori et al. 23821 (CAY-n.v., GH, NY-n.v.); 10 Sep 1994 (d fl), Mori et al. 23872 (CAY-n.v., GH, NY-n.v.). SBIAHSCI HUSluM 3811144 B 27W.1980 UTREr HI ' The specimenshere describedas R. saulensis have been previously identifiedeither as R. grandis or as a new species of close affinity to it (J. G. Rohwer, in sched.). Detailed analysis of vegetativeand reproductive morphology of R. grandis have resulted in the segregationof various distinct groupswithin this variable complex (see "TheRhodostemonodaphnegrandis complex"above).Rhodostemonodaphnesaiilensis can be distinguishedreadily from the other species in the R. grandis complex by its darkbrownto blackish dry leaves (with reddish venation when fresh), few (4-5) pairs of lateral veins, arching near the margin to form an acute chordal angle (<250), and margin often recurvedat base. The flowers, with their wide, laminar,overlappinganthersof whorls I andII in both IAA staminateand pistillate individuals, and their red tepals, are also diagnostic within the R. grandis complex. Rhodostemonodaphnegrandis, the only other species from this complex in French Guiana, has 59 pairs of lateralveins, their chordal angle is greater than 250, and the flowers are yellow to cream (see FIG.25. Rhodostemonodaphne praeclara s.str.(Steyermark 111392). TableV). 90 adaxiallyflattened;blades chartaceous,flat, narrowly elliptic to obovate, 7-19 X 3-6(-9) cm; base cuneate to acute or obtuse, minutely decurrent,50-90?; apex rounded, 100-130', (mucronate);margin plane; primary vein above flat, below prominent;secondary veins 4-6 pairs, equidistant, eucamptodromous, above slightly raised, below raised, divergingat 45600, abruptlyarchingnear margin,chordalangle 20300, the angle uniform along blade length; tertiary veins above and below slightly raised, randomreticulate to scalariform;higher-orderveins above and below slightly raised; surface above and below brown to olive-green; indumentabove absent, below minutely sericeous, the hairs sparse,to 0.3 mm long, straight, appressed,ascending, yellowish to translucent, caducous after one flush. Staminate inflorescences: along whole length of flush, erect, peduncles 6-16 cm long, the hypopodia3-8 cm X ca. 1.6 mm, branch orders 4, the second-orderbranches 10-14, dispersed, lowest branchto 2 cm long, color and indumenton all axes as on twigs; bractsand bracteoles caducous (not seen). Staminateflowers: pedicels ca. 5.6 X 0.8 mm, the diametereven throughout;receptacle globose, ca. 2 X 2.4 mm, constrictedat the place of tepal inception; tepals membranaceous,ovate, ca. 2 X 2 mm (inner whorl slightly smaller),at anthesis recurved, reddish, adaxially puberulous;stamens of whorls I and IH,the antherssessile, oblong, ca. 0.6 X 0.8 mm (whorl II slightly smaller), glabrous,the locelli 4, apical, in a shallow arch, introrse,the glands absent; whorl III columnar,ca. 1.2 X 0.6 mm, glabrous,the anthersglobose, the locelli 4, the upperpair latrorse, the lower pair extrorse, the glands absent; whorl IV absent; all stamens reddish;pistillode absent. Pistillateflowers: pistil ca. 2.6 X 1.4 mm; ovary ovoid, ca. 1.6 mm long, glabrous.Fruits: pedicels to 12 X 6 mm, abruptlyenlargingto form the cupule; cupule hemispherical,to 15 X 23 mm, tuberculate, the marginstraight,tepals caducous;berryunknown. FLORANEOTROPICA the Gran Sabana and the two Marcano-Berticollections from the Imatacarange). Flowers SeptemberNovember,January-February, duringthe dry season. FruitsMarch-April,at the beginningof the rains;cupules from old fruitshave been found in June. Additional specimens examined. VENEZUELA. AMAZONAS:Atabapo,Rio Paru,Oct 1989 (fr), Delgado 741 (MO).BOL; AAIVAR: Sucre,Helipuerto 23, May 1990(fr), Delgado1038(MO,NY);ImatacaForestReserve,LaQuebradacamp, 23 Jun 1964 (fr), Marcano-Berti248 (F, G X2, HBG X3, MER, MO, NY, US); 24 Jun 1964 (fr), Marcano- Berti254 (F, HBGX2, K, MER,MO X2, NY,U X2, US X2); Quebrada O-paru-md, betweenSantaTeresitade Kavanay6nandRioPacairao, 20 Nov 1944(9 fl),Steyermark 60403 (F X2);betweenSanIgnacioandSanFranciscode 4 Jan 1975 (d fl), Steyermark Yuruani, 111392(F, HBG, NY,U). DELTA AMACURO: E of RfoGrande,E-NEof El Palmar,neartheborderof Bolivar,Sep 1965(d fl),Blanco 278 (G, HBGX2, MO X2, NY);AntonioDiaz,8 kmN of Wausa,rightbankof Rio Amacuro,Feb 1987(9 fl), Ferndndez3915 (HBG, MO X2, NY). GUYANA. MAZARUNI-POTARO:Demerara-Essequibo RR,AnarikaLine,nearMonkeyCreek,21 Mar1910(fr), Anderson62 (K);EssequiboRiver,Wallabaforest,MoraballiCreek,4 Nov 1938(d fl),Davis 596 = FD 2716 (FHO, U); 13 Oct 1938 (d fl), Fanshawe 130 = FD 2739 (FHO, G,NY,S);Essequibo River,Wallaba forest,Winiperu Creek, 13 Apr 1940 (fr), Fanshawe450 = FD 3186 (FDG); Esse- quiboRiver,Wallabaforest,Moraballi Creek,29 Oct 1929 (d fl), Sandwith535 (K). Local names. Venezuela:laurel grande;Guyana: shirua, buradie (Arawak); broad-leavedsoft silverballi. Rhodostemonodaphnepraeclara is another species segregated from the R. grandis complex (see "The Rhodostemonodaphne grandis complex" above). As delimitedhere, it can be recognizedby its coriaceous, slightly obovate, mucronateleaves and absence of pairedglands on the stamensof whorl III. It is found in a fairly circumscribedarea-easterm VenezuelaandadjacentGuyana-the only memberof Field notes. Trees to 30 m tall and 30 cm diam., the R. grandis complex growing there. A specimen already flowering when 9 m tall; buttressesca. 100 from Mt. Roraima (Steyermark59001), previously cm tall; outer bark smooth, dark; wood aromatic; identifiedas R. grandis, is vegetatively similarto the leaves above darkgreen, the veins yellow, leaves beupper-montanecollections of R. praeclara, but it has low light green to glaucous. Inflorescenceaxes rederecttepals,andclavate,sessile, two-locellateanthers, dish; pedicels reddish; receptacle reddish; tepals as in some Amazonianspecimens determinedas Encreamy yellow; stamens/staminodesgreen. Cupule dlicheria metallica Kosterm. reddishgreen to purple. Eighteen specimens previously identifiedas RhoDistribution (Fig. 23). Northeast Venezuela to dostemonodaphnegrandis, with variable vegetative north-central Guyana, at 250(-700) m, in non- morphology and collected throughoutthe Amazon inundated,lowlandrainforest;also in uplandrainfor- river basin, still present an unsolved problem in deest, to 1200 m (the two Steyermarkcollections from limitation. These differ from R. praeclara s.str. not SYSTEMATICTREATMENT only vegetatively,but also are found beyond the limited geographicalrange where the well-defined morphological group describedabove is found. However, the twelve fertile specimens have flowers identicalto those of R. praeclara, including stamens of whorl III which are devoid of glands. I have annotatedthe Venezuelan/Guyananspecimens as R. praeclara s.str. The remainingheterogeneousgroup has been annotated as R. praeclara s.l. and the specimens in it are listed below. These specimens to a certainextent link R. saiilensis, from central French Guiana, R. peneia from Manaus and its surroundings,and R. praeclara from the Venezuela-Guyanaborder.It is because of these specimens, togetherwith the fruiting/sterilematerial of Endlicheria metallica misidentified as Rhodostemonodaphne grandis, that the R. grandis complex came to be recognized as a widespreadvariable taxon. For a list of charactersdistinguishingR. praeclara s.str., R. praeclara s.l., and other species of the R. grandis complex see TableV. 91 (F, GH, MO, NY, US); Mun. Jiraparana,Gleba G.km 2, 29 Mar 1983 (d fl), Ferreira-P.-E83-31 (HBG). 39. Rhodostemonodaphne recurva van der Werff, Novon 10: 291. 2000. Type. Brazil. Amazonas: Mun. Manaus, Reserva Florestal Ducke, 5 Aug 1994 (d fl), Vicentini et al. 653 (holotype:INPAn.v.; isotypes: MO, SP-n.v.). Fig. 26 Trees: branches basitonic, in axils of cataphylls; twigs angular,soon becoming terete,ca. 3 mm diam.; epidermisbrownishto black, barely visible due to indument cover; terminal bud plump, ca. 5 X 3 mm; cataphylls caducous; indument sericeous, caducous by next flush, the hairs dense, to 0.25 mm long, straight,appressedascending to erect, golden yellow. Leaves: petioles slender, 1.5-3 cm X 1.5-2.5 mm, adaxially flattened;blades chartaceous, flat, widely elliptic to oblong (obovate), 6-14 x 3-5.5 cm; base cuneate to acute, minutely decurrent,65-80?; apex rounded,80-100o, mucronateup to 1 cm long; margin Specimens annotated as Rhodostemonodaphnepraeclara s.I. COLOMBIA. AMAZONAS: Soratama, Rfo Apaporis, between Rfo Pacoa and Rio Kananarf,31 Aug 1951 (d fl), Schultes & Cabrera13805 (BM, C, GH, NY, US, U). VENEZUELA. Atures,SerraniaYutaj6,Rio Manapiare, 18 Jun 1976 (fr), Berry 2255 (G, MO, MYF, NY, U); DepartmentAtabapo,Pandamoriver plain, Feb 1990 (fr), Ma0~~~~~~~~~~~~~~~~~ 0417177! rin 805 (MO). BOLiVAR: Cerro Cuquenan,S of Mun. Ur4 daneta, Dtto. Riosucio, 16 Apr 1986 (fr), Herndndez & Dezzeo 235 (HBG, US). ECUADOR. MORONA-SANTIAGO:Valleyof Rio Zamora, ca. 50 km NE of Yatanza, 17 Nov 1982 (Y fl), T D. Pennington & Tenorio 10760 (QCA). NAPO: Yasuni river and Jatun-Cochalakes, 29 Sep 1988 (cd fl), Ceron & Gallo 5170 (MO). PERU. AMAZONAS: Baguaprovince,Maranionriver, Kusu-Chapi creek, Feb 1995 (sterile), R. Vdsquez et al. 19409 (GH, MO-n.v.); (sterile), R. Vdsquez et al. 19472 (GH, MO-n.v.). BRAZIL. Basin of Rio Jurua,near mouth of Rio Embira, 10 Jun 1933 (d fl), Krukoff4747 (A, F, G, K, M, MO, NY, S, SP, U, US); 4 Jul 1933 (Y fl), 5171 (A, BM, F X2, G, M, MO, NY, S, SP, U, US); Mun. Manaus,Fazenda Esteio, 17 Sep 1980 (d fl), J. L. dos Santos (INPAI WWF)1202.5028(MO); Serrada Neblina, 22 Dec 1965 (Y imm fr), N. T Silva & Brazdo 60720 (MO, NY X2); Mun. Manaus, Ducke Forest Reserve, 5 Apr 1994 (fr), Vicentini et al. 468 (INPA, MO). PARA:Serra dos Carajds,Serra do Norte, 30 May 1983 (d fl), M. F F Silva etal. 1411 (HBG); Jacunda,Rio Tocantins,Rio Cajazeiras,16 May 1978 (Y fl), M. G. Silva & Bahia 3604 (HBG, MO, NY); Serrados Carajas,AMZA camp 4-Alfa, 6 Jun 1982 (Y fl bud), Sperling et al. 5928 (MO). ROND6NIA: Espigao do Oeste, BR 364, FIG. 26. Rhodostemonodaphnerecurva(Vicentiniet al. 653). estradada FUNAI, km 5, 20 Jun 1984 ( fl), Cid et al. 4658 FLORA NEOTROPICA 92 recurved at base or raised; primary vein above flat, below prominent;secondaryveins 3-4 pairs,equidistant, eucamptodromous,above flat, below slightly raised,divergingat ca. 300,graduallyarching,chordal angle ca. 200, the angle increasingtowardsapex; tertiary veins above and below slightly raised, randomreticulate to scalariform; higher-orderveins above andbelow flat;surfaceabove shiny brown,below dull brown(grayishon young leaves); indumentabove absent, below minutely sericeous, the hairs sparse, to 0.4 mm long, straight,appressed,ascending,yellowish to translucent, persistent. Staminate inflorescences: along whole length of flush, erect, peduncles (4-)5-7 cm long, the hypopodia2.5-3.5 cm X ca. 1 mm, branchorders4, the second-orderbranches3-5, dispersed,lowest branchto 2.5 cm long, all axes color indumentas on twigs; bractsandbracteolescaducous (not seen). Staminateflowers: pedicels ca. 2.5 X 0.6 mm, graduallyenlargingto form an obconicalreceptacle ca. 1.5 X 2 mm; tepals membranaceous,ovate, ca. 1.7 x 2 mm (inner whorl slightly larger), at anthesis spreading, yellowish, adaxially puberulous; stamens of whorls I and II, the antherssessile, globose, ca. 0.8 X 1.3 mm (whorl II slightly smaller), glabrous,the locelli (1,) 2 (3, 4), apical, in a shallow arch, introrse;whorl III columnar,ca. 1 X 0.7 mm, glabrous,the antherscolumnar,the locelli 4, the upper pair latrorse, the lower pair extrorse, the glands 2, minute ca. 2 mm broad, black; whorl IV absent; all stamens reddish; pistillode minute, elongate, ca. 2 mm long. Pistillate plant not seen. Field notes. Trees 17-25 m tall and 30-35 cm diam.; trunkstraight;outer bark smooth with inconspicuouselliptic to roundishlenticels arrangedin vertical lines and alternatingwith longitudinalfissures; <1 mm thick; inner bark two-layered, the external beige, homogeneous, the internal mixed dark and light beige; wood whitish yellow. Inflorescenceaxis green with reddish branchlets and pedicels; tepals whitish green adaxially,yellow green abaxially. Distribution (Fig. 23). Rhodostemonodaphnerecurva is only known from the surroundingsof Manaus. It grows on terrafirme in yellow clay soils. Additional specimens examined. BRAZIL. AMA- ZONAS: Distrito agropecuariode SUFRAMA, BR-174, km 72, FazendaDimona, 6 Ago 1982 (d fl), Mackenzie(INPAI WWF2108/35) (MO); ibid. km 64, Fazenda Esteio, 2 Oct 1980 (cd fl), J. L. dos Santos (INPA/WWF1202/5767) (MO); ibid. s.d. (d fl) s.coll. (INPAIWWF1202/3094)(MO). When I first saw the specimens that now represent Rhodostemonodaphnerecurva,I determined them as an undescribed species of Endlicheria due to their two-locellate anthers. The plants however share a numberof characterswith the species in the Rhodostemonodaphnegrandis complex, such as the sericeous indument,andthe flowerswith spreadingtepals and broad sessile anthers. 40. Rhodostemonodaphne peneia Madrinian,sp. nov. Type. Brazil. Amazonas: Mun. Manaus, Ducke Forest Reserve, 14 Jul 1965 (d fl bud), Rodrigues 6976 (holotype: INPA; isotypes: G, Fig. 27A HBG). praeclarae et R. sauilensisimilis, Rhodostemonodaphne binis a hacstaminibus exsertis,eis verticilliIIIbasiglandulis indestitutis,ab illa foliorumformaet textura,ab ambabus minoribus differt. floribusque florescentiis Trees: branchesbasitonic,in axils of cataphyllsor basal foliage leaves; twigs angular, soon becoming terete, 2-3 mm diam.; epidermisbrownishto black; terminalbud slender, ca. 6 X 3 mm; cataphylls caducous; indumentsericeous to absent, caducous by next flush, the hairs sparse,to 0.2 mm long, straight, appressed,ascending,silver.Leaves: petioles slender, 1.3-2(-3.1) cm X 1.2-1.8 mm, adaxially flattened; blades chartaceous,flat, narrowlyelliptic, 9-19(-22) X 3-7 cm; base acute, minutely decurrent,50-100?; apex obtuse, 70-120?, mucronateto acuminatefor up to 1.3 cm; marginplane; primaryvein above slightly raised, below prominent; secondary veins 5-7(-8) pairs, equidistant,eucamptodromous,above flat, below raised, diverging at 35-55?, evenly arching, chordalangle ca. 200, the angle uniformalong blade length;tertiaryveins above flat,below slightly raised, random-reticulateto scalariform;higher-orderveins above and below slightly raised; surface above and below olive-green to dark brown, inconspicuously black-dotted;indumentabove absent,below minutely sericeous, the hairs sparse, to 0.2(-l) mm long, straight, appressed, ascending, yellowish, caducous after one flush. Staminate inflorescences: along whole length of flush, erect?, peduncles 3-11 cm long, the hypopodia 1-5 cm X 0.8-1.2 mm, branch orders 4(-5), the second-orderbranches (6-)8-9(11), dispersed, lowest branch to 1.2(-2.4) cm long, color and indumentof all axes as on twigs; bractsand bracteoles caducous (not seen). Staminate flowers: pedicels ca. 2.4 X 0.6 mm, the diametereven throughout; receptacleglobose, ca. 1.2 X 2 mm, constricted at the place of tepal inception;tepalsmembranaceous, obovate, ca. 1.5 X 1 mm (inner whorl slightly smaller), at anthesis recurved,reddishbrown, adaxi- SYSTEMATICTREATMENT ally puberulous;stamensof whorlsI and II spathulate, the anthers reniform, ca. 0.8 X 1 mm (whorl II slightly smaller), with a few hairs at base, the locelli 4, apical, in a shallow arch, introrse,the glands absent; whorl III columnar,ca. 1.4 X 0.6 mm, with a few hairs at base, the anthersglobose, the locelli 4, nearly apical, but clearly derived from the upper latrorsepair,and the lower pairextrorse,the glands absent; whorl IV absent; all stamens yellowish; pistillode absent.Pistillateflowers: pistil ca. 2.5 X 1 mm; ovary ovoid, ca. 1.2 mm long, glabrous. Fruits: pedicels to 15 X 6 mm, abruptlyenlargingto form the cupule; cupule hemispherical,to 25 X 30 mm, smooth to tuberculate,the marginstraight,tepals caducous; berry unknown. Field notes. Trees to 18(-25) m tall and 25(-40) cm diam., already flowering when 10 m tall; wood yellow. Tepals yellow; stamens/staminodesgreen. Distribution (Fig. 23). The Ducke forest reserve nearManausin the Brazilianstateof Amazonasat ca. 100 m; one specimen (CampbellP22290) from the Rio Trombetasin the state of Para'at 200 m. It grows in non-inundatedrain forest near streams.In the Manaus areafloweringJune-Septemberat the end of the rains and beginning of the dry season; in Para'flowering in May, also towardsthe end of the rainyseason. 93 both species is also indicative of their close affinity. Rhodostemonodaphnepeneia has thinnerleaves, although the range of variationin R. praeclara is quite broad,and the inflorescencesand flowersof R.peneia are considerablysmaller than those of R. praeclara. For othercharactersused in distinguishingthe species of the R. grandis complex see Table V. The whitish indumenton the reproductivepartsgives them a quite characteristic greyish appearance, reminiscent of some species of Endlicheria. 41. Rhodostemonodaphne penduliflora Madrifian, sp. nov. Type. Colombia.Cauca:PuertoLopez de Micay, Rio Naya, 1 km upstreamfrom vereda"El Carmen,"2 Apr 1992 (Y fl), Cogollo, Devia & Madrinidn5159 (holotype:COL;isotypes: C, GH, HBG, JAUM,JBGP,K, MO, NY, P, QCA, TULV, U). Fig. 27B Exudatomucilaginoso maximeinsignis,a congeneribus inflorescentiis pendulis,staminibus omnibusglanduliferis et eorumverticilliIIIlocellisintrorsisdivergens. Trees: branchesbasitonicto mesotonic,in axils of foliage leaves, pseudo-verticillate;twigs angularand remaining so for at least two flushes, ca. 10 mm diam.;epidermisblack; terminalbud slender,ca. 6 X 3 mm; cataphylls caducous; indumentminutely puAdditional specimens examined. BRAZIL. AMAcaducous by next flush, the hairs isolated, berulous, 2 ZONAS: Mun. Manaus, Ducke Forest Reserve, Sep 1968 to 0.1 mm long, straight, appressed, ascending, in(Y fl), Aluisio 122 (HBG, INPA, M, US); 6 Aug 1980 (Y fl), Kurz B-28-14 (MO); (Y fl), Kurz B-28-14 (MO); conspicuously golden. Leaves: petioles robust, 1.5-2 Manaus-Caracarai rd., km 115, Rio Urubd, 13 Aug 1974 (od cm X 2.5-5.5 mm, adaxially flattened;blades coriafl bud), Prance et al. 21642 (HBG X2, K, MO, NY, US,); ceous, flat, broadly elliptic to obovate, (8-)15-28(Mun. Manaus, Ducke Forest Reserve, 15 Jun 1994 (d fl), 38) X (6-)10-16(-21) cm; base obtuse, 70-110?; Ramos & C. E da Silva 2834 (INPA,MO); 11 Aug 1965 ( apex obtuse to rounded,110(150?, (mucronate);marfl), Rodrigues 7009 (INPA); 23 Mar 1995 (d fl), Vicentini gin plane; primaryvein above flat, below prominent; & C. F da Silva 963 (INPA, MO). PARA:Rio Trombetas, secondary veins 7-10 pairs, equidistant,brochidodkm 109, access rd. from CachoeriaPorteirato Perimetraldo romous, above raised, below prominent,divergingat Norte, 27 May 1974 (d' fl), Campbellet al. P22290 (HBG 60-70?, straight to evenly arching, occasionally X2, NY, U). forked,chordalangle ca. 450, the angle uniformalong Etymology. Named after the Greek river god Pe- blade length; tertiaryveins above slightly raised, beneus, fatherof Daphne,in allusionto the riverinehab- low raised, random-reticulateto scalariform;higheritat of the plant, as also its local name [Lat.Peneius, order veins above and below slightly raised; surface belonging to Peneus]. above andbelow yellowish green to yellowish brown; indumentabove andbelow absent,the veins with isoLocal name. Louro do baixio. lated, ca. 0.2 mm long, appressed,ascending,yellow Rhodostemonodaphne peneia is one of the species hairs. Staminateplant unknown.Pistillate infloressegregated from the R. grandis complex (see "The cences: basitonicto mesotonic,pendulous,peduncles 9-27 cm long, the hypopodia5-12 cm X 1.5-3 mm, Rhodostemonodaphne grandis complex" above). It is vegetatively very similarto R. praeclara, with a dis- branchorders5, the second-orderbranches6-8, distributionrange that overlaps with that of R. peneia; persed, lowest branchto 1.5(-4.5) cm long, color and their membranous tepals, slightly exserted anthers indumentof all axes as on twigs; bractsandbracteoles and absence of glands on the stamensof whorl III in caducous (not seen). Pistillate flowers: pedicels ca. 94 FLORA NEOTROPICA A V? 04671774 B FIG. 27. A. Rhodostemonodaphne peneia (Ramos & C F da Silva 2834). B. R. pendulflora 2.4 X 1.5 mm, the diametereven throughout;receptacle obconical, ca. 1.5 x 3.6 mm; tepals coriaceous, obovate, ca. 3.5 X 3.5 mm (inner whorl slightly smaller), at anthesis recurved, yellowish to reddish brown, adaxially papillose; staminodes of whorls I and 11,the antherssessile, chubby,broadlyelliptical, ca. 1.2 x 2 mm, papillose, the locelli 2-3-4, apical, in a shallow arch, introrse,the glands minute, ca. 0.6 mm diam.; whorl 111columnar,ca. 1 x 3 mm, papillose, the antherstrapezoid,the locelli 1-2, introrse, the glands laminar, each pair fused and enveloping the staminodeon threesides, ca. 0.6 mm diam.;whorl IV absent;all staminodesyellowish to reddishbrown; pistil ca. 1.5 x 1 mm; ovary globose, ca. 1.2 mm long, papillose. Fruits unknown. Field notes. Trees to 24 m tall and 30 cm diam.; buttresses ca. 60 cm tall, 30 cm broad; outer bark smooth (with horizontal furrows), shiny black; inner bark ca. 5 mm thick, yellow-brown, with copious clear mucilaginous exudate; light yellow, wood nonaromatic;twigs green; leaves above green, the veins yellow, leaves below light green to yellowish green. Inflorescence axes reddish; pedicels reddish; tepals yellowish green; stamens/staminodes green; ovary green, stigma green. (Cogollo et al. 5159). Distribution (Fig. 20). Known only from the type found in the Pacific coast of Colombia, on the border between the Cauca and Valle departments, in the Choc6 biogeographic province, at near sea level. It grows in disturbedforest, on the slope of a hill near a riverbank. Flowers in Marchtowardsthe end of the dry season. Etymology. The specific epithet alludes to the pendulous inflorescences. [Lat. pendulus, pendulous + flos, flower]. Local names. Jigua pava,jigua baboso. Rhodostemonodaphnependuliflora is a very distinct species. Its thick, almost glabrous,black-drying twigs, and large, coriaceous, glabrous, yellow to brown green-drying leaves are unlike those of any other Lauraceae known to me. It is also unique in having featuresnot found in any othermemberof the genus: copious, clear, mucilaginousexudatefrom the bark, papillose tepals, staminodes and pistil, paired glands in all three staminalwhorls, and introrsestaminodes of whorl III. These featuresare found individually in othergenera;the papillose tepals and stamens are a feature common in the genus Nectandra; the LITERATURECITED 95 presence of glands in all staminal whorls is known only in two small genera,Chlorocardiumand Urbanodendron, and one species of Rhodostemonodaphne (R. celiana (C. K. Allen) Rohwer); and the introrse anthersof whorl III-the normin the subfamilyLaureae sensu Rohwer (1993a)-are known from only a few species in the Perseeae, the subfamily to which Rhodostemonodaphnebelongs (e.g., Ocotea staminea (Griseb.)Mez). This combinationof charactersplace R. penduliflora in an isolated position within Rhodostemonodaphne. In the meantime,since staminateindividuals in which the staminal charactersmay be better expressed, are unknown, I place this species with the otherdioecious species thathave fourlocelli arranged in an arch, i.e., Rhodostemonodaphne.The wood anatomyis nondescript(H. Richter,pers. comm.). ACKNOWLEDGMENTS I am grateful to the Fundaci6n "Universidadde Los Andes, New York"for financialsupportduringa crucialstage of my studies.Field workwas madepossible thanksto financialsupportfrom the AtkinsFund for TropicalBotany,graduatestudentresearchgrants from the Departmentof OrganismicandEvolutionary Biology of HarvardUniversity,the TinkerField Research Grantfrom the Committeeof Latin-American and IberianStudies, HarvardUniversity,and the Garden Club of America Grant for Tropical Botany (World Wildlife Fund). I am especially indebted to Cesar EduardoBarbosa, who introducedme to the study of systematicbotany and who was an infallible field companion during most of my field trips in Colombia.I also want to thankJulio Betancur(COL), Alvaro Cogollo (JAUM),Hermes Cuadros(formerly at JBGP), andWilson Devia (TULV)for theirhelp in the field. I am gratefulto PeterF. Stevens for his continuous support and for sharing his experience and knowledge. Many thanks to P. Barry Tomlinsonand David Baum for their advice and suggestions for improvement of many aspects of the work. I wish to thankHenk van der Werff(MO) and Jens G. Rohwer (Hamburg)for their help in many aspects of the production of this work. At HarvardI am indebted to many people and institutionsthat have providedresources, support,and most importantlytheir friendship, specially JudyWarnement,Directorof the Harvard Botany Libraries; Emily Wood and Michael Canoso, GustavoRomero, and Shiu-YingHu in particularwere not only helpful in the whole process,but also four very special friends, as were all colleagues from the graduateschool, particularlyStuartJ. Davies. Many institutionswere very kind in providing loans of their material for study even for extended periods of time. In lieu of citing all the acronymshere I thankthem throughtheircuratorscollectively.I specially want to thank the directorsand staff of those herbariathat I visited, BM, COL, K, MO, NY, and US for their hospitality.In particular,I thankthe former Directorsfor Scientific Researchat NY, Enrique Forero and Scott Mori, for generously providing funds for a visit. Also at NY, I want to thankJames L. Luteyn for many detailed and helpful comments on the manuscript.Workon the final stages of preparation of the manuscriptwas possible thanks to a TinkerFoundationFlora NeotropicaFellowship and a Mellon FoundationKew Latin AmericanResearch Fellowship.GustavoRomerodid a wonderfuljob digitizing the illustrations;gracias! LITERATURE CITED Allen, C. K. 1945. Studies in the Lauraceae, VI: preliminary survey of the Mexican and Central American species. J. Arnold Arbor. 26: 280-434. . 1964. Lauraceae. Mem. New York Bot. Gard. 10: 44-123. . 1966. Notes on Lauraceae of tropical America I. 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(Lauraceae), sensu lato. Mitt. Kostermans,A. J. G. H. 1936. Studiesin SouthAmerInst. Allg. Bot. Hamburg 20: 1-278. ican Malpighiaceae,Lauraceae,and Hernandiaceae, . 1991. Borderline cases between Ocotea, Necespeciallyof Surinam.J. H. de Bussy,Amsterdam. tandra, and Phoebe (Lauraceae): the "marginal"spe. 1957.Lauraceae.Commun.ForestRes. Inst.57: cies of the Ocotea helicterifolia-group including 1-64. the 0. heydeana-group. Bot. Jahrb. Syst. 112: 365Kubitzki,K. 1982.Aniba.Fl. Neotrop.Monogr.31: 1397. 84. .1993a. Lauraceae. Pp. 426-437 in: K. Kubitzki Kurz, H. 2000. Revision der GattungLicaria (Laura(ed.), The families and genera of vascular plants. Vol. ceae). Mitt.Inst.Allg. Bot. Hamburg28/29:89-221. 2. Springer-Verlag,Berlin. Lasser, T. 1948. PlantaenovaeVenezuelanae.Bol. Soc. . 1993b. Nectandra. Fl. Neotrop. Monogr. 60: 1Venez.Ci. Nat. 11: 181-198. 333. Madrifiain,S. 1996. New species of Rhodostemonoda.2000. Toward a phylogenetic classification of SouthAmerica. phne (Lauraceae)fromnortheastern the Lauraceae: evidence from matK sequences. Syst. Brittonia48: 45-66. Bot. 25: 60-71. . 2002. Rhodostemonodaphne Rohwer & Ku& K. Kubitzki. 1985. Entwicklungslinien im bitzki.Pp. 381-383 in: S. A. Mori,G. Cremers,C. A. Ocotea-Komplex (Lauraceae). Bot. Jahrb.Syst. 107: Gracie,J.-J. de Granville,S. V. Heald, M. Hoff & 129-135. J. D. Mitchell(eds.), Guideto the vascularplantsof CentralFrenchGuiana.Part2: Dicotyledons.Mem. , H. G. Richter & H. van der Werff. 1991. Two new genera of neotropical Lauraceae, and critical reNew YorkBot. Gard.76(2). marks on the generic delimitation. Ann. Missouri Pp. 1-260 Meissner,C. F. 1864.Rhodostemonodaphne. Bot. Gard. 78: 388-400. in: A. P. de Candolle(ed.),Prodromussystematisnaturalisregni vegetabilis.Vol. 15, part 1. Masson, Sandwith, N. Y. 1932. Contributions to the flora of tropical America: XII [contd.]. Kew Bull. 5: 209-220. Paris. . 1952. Contributions to the flora of tropical Mez, C. 1889. LauraceaeAmericanae.Jahrb.Konigl. America: LIV [contd.]. Kew Bull. [7]: 251-263. Bot. Gart.Berlin5: 1-556. . 1903. Additamenta monographica, 1903 Schnell, R. 1987. La flore et la vegetation de l'Amerique tropicale. Masson, Paris. [contd.].Bull. Herb.Boiss. Ser. II, 3: 224-238. . 1920. Additamentamonographica,1919. Re- Schreber, J. C. D. von. 1791. Genera Plantarum. Varrentrapp& Wenner, Frankfurt. pert.Spec. Nov. RegniVeg. 16: 317-332. Nees von Esenbeck,C. G. D. 1833.Revisiolaurinarum; Smith, A. C. 1935. Plantae Krukovianae: Lauraceae. Phytologia 1: 115-122. adjectisunt generumet specierumcharacteres.LinVeillon, J. M. 1976. Architecture vegetative de quelques naea8: 36-51. arbres de l'archipiel Neo-Caledonien. Ph.D. disserVeit,Berlin. . 1836. Systemalaurinarum. tation, Universite des Sciences et Techniques du LanNelson, B. W., C. A. C. Ferreira, M. F. da Silva & gedoc, Montpellier. M. L. Kawasaki. 1990. Endemismcentres,refugia LIST OF EXSICCATAE 97 Weinberg,B. 1984. Restingas:origem,estruturae pro. 199lb. New speciesof LauraceaefromEcuador cessos. CEUFF,Niteroi. andPeru.Ann. MissouriBot. Gard.78: 409-423. Werff, H. van der. 1984. Notes on neotropicalLaura. 1993. A revisionof the genus Pleurothyrium ceae. Ann. MissouriBot. Gard.71: 1180-1183. (Lauraceae).Ann.MissouriBot. Gard.80: 39-118. . 1987. A revision of Mezilaurus(Lauraceae). & H. G. Richter. 1996. Towardan improved Ann. MissouriBot. Gard.74: 153-182. classificationof Lauraceae.Ann.MissouriBot.Gard. . 1989a. A new combinationin Ocotea (Laura83: 409-418. ceae). Ann. MissouriBot. Gard.76: 1175. & A. Vicentini.2000. New speciesof Lauraceae . 1989b.Novelties in the LauraceaefromVenefromCentralAmazonia,Brazil.Novon 10:264-297. zuelanGuayana.Ann. MissouriBot. Gard.76: 462- Wheelwright, N. T. &. A. Bruneau. 1992. Population 475. sex ratiosand spatialdistributionof Ocotea tenera . 1991a. A key to the generaof Lauraceaein the (Lauraceae) treesin a tropicalforest.J. Ecol.80:425New World.Ann. MissouriBot. Gard.78: 377-387. 432. NUMERICAL LIST OF TAXA 1. Rhodostemonodaphne laxa (Meissner) Rohwer 2. R. velutina (Mez) Madriinin 3. R. ovatifolia Madrina'n 4. R. longiflora Madrinain 5. R. juruensis (A. C. Sm.) Chanderb. 6. R. frontinensis Madriniin 7. R. cyclops Madrifiin 8. R. dioica (Mez) Rohwer 9. R. anomala (Mez) Rohwer 10. R. macrocalyx (Meissn.) Rohwer ex Madrinian 11. R. capixabensis Baitello & Coe-Teix. 12. R. avilensis Madrifiin 13. R. scandens Madrinian 14. R. licanioides(A. C. Sm.) Madrifian 15. R. miranda (Sandwith) Rohwer 16.R. revolutifolia Madrifiin 17. R. crenaticupula Madrifian 18. R. debilis (Kosterm.) Chanderb. 19. R. curicuriariensis Madrifian 20. R. tumucumaquensis Madrifiin 21. R. negrensis Madrifnin 22. R. mirecolorata (C. K. Allen) Rohwer 23. R. rufovirgata Madriiain 24. R. celiana (C. K. Allen) Rohwer 25. R. steyermarkiana (C. K. Allen) van der Werff 26. R. leptoclada Madriiiin 27. R. parvifolia Madriniain 28. R. antioquiensis Madrifidn 29. R. kunthiana (Nees) Rohwer 30. R. synandra van der Werff 31. R. napoensis Madrifiain 32. R. longipetiolata Madrinian 33. R. morii Madrifian 34. R. sordida Madrifian 35. R. elephantopus Madrifian 36. R. grandis (Mez) Rohwer 37. R. sauilensis Madrifian 38. R. praeclara (Sandwith) Madrifian 39. R. recurva van der Werff 40. R. peneia Madrifnan 41. R. penduliflora Madrifian LIST OF EXSICCATAE Albert de Escobar, L. (et al.), 7835 (29). Allen, C. K. (et al.), 41, 51 (36); 67 (33); 330 (Paraia bracteata). Aluisio, J., 122 (40). Ancuash, E., 133, 283 (Endlicheria metallica). Anderson, C. W., 62 (38). BAFOG, 361-M, 7241, 7425 (36). Balee, W. L. (et al.), 1848 (29); 2315 (Endlicheria metallica). Bandeira, M. C., s.n. (10). Barbosa, C., 1229 (29). Barkley, F. A. et al., 389 (1). Barrier, S., 4003, 5153 (36). Beck, S. G., 13676 (29). Bena, P., BAFOG 307-M, 1300 (36). Benoist, R., 1620 (36). Bernardi, A. L., 1862 (29). Berry, P., 2255 (36 s.l.). Betancur, J. et al., 3756 (1). Binot, P. M., 110 (10). Blanco, C., 278 (38). Boerboom, J., (LBB)9584 (36). Boissier, P. E., 12 (10). Boom, B. M. (et al.), 1722 (37); 1811 (23, type); 5925 (R. sp. nov.?); 8572 (17); 10536 (29). Brant, A. E. & Martinez, 1387 (6, type). Bunting, G. S. & A. Stoddart, 9056 (29). BW, 3204, 5546, 6908 (36). Campbell, D. G. (et al.), 14647 (36); P.21974 (R. sp. nov.?); P.22290 (40). Cardenas, D. et al., 2522 (28, type). Cazalet, P. C. D. & P. D. Pennington, 7629 (Endlicheria metallica). Ceron, C. E. (et al.), 3435 (29); 5170 (36 s.l.). 98 Cid, C. A. (et al.), 981 (17); 4658 (36 s.l.). Clarke, D., 647b (13). Coelho, D. F., (INPA) 3990, (INPA) 4025 (17); (INPA) 46519, (INPA) 46808 (Endlicheria metallica); (INPA) 46810 (21, type). Cogollo, A. (et al.), 290 (28); 5159 (41, type). Cooper, G. P. (et al.), 96 = (Y)10277 type of Ocotea cooperi, 498 = (Y)12116 (29). Cooper, J. J., 10217 (29, syntype of Ocotea stenoneura). Croat, T. B., 19790 (29). Croizat, L., 917 (29). Daly, D. C. (et al.), 3837, 4032 (13); 5358 (29); 5457 (21); 6830, 6898, 7072 (29). Daniel, Bro., 1439 (1). Daniels, A. G. H. & F. P. Jonker, 805 (36). Davidse, G. et al., 25600 (29). Davis, T. A. W., 596 = FD 2716 (38). Delascio, F. & F. Guanchez, 10930 (29). Delgado, L., 741, 1038 (38). Dfaz, C. et al., 4532 (29). Dodson, C. H. (et al.), 2850 (17); 6326, 6339, 6412, 6564, 6658, 13675 (29). Donselaar, J. van, 1891, 2425 (36). Duarte, A. P. (et al.), 263, 5605 (10). Ducke, A., 25674 (19, type); 25681 (21). Dudley, T. R., 10072 (Endlicheria metallica). Elburg, J. P., (LBB)9470 (36). Elburg, R., (LBB)8802 (36). Elcoro, S., 324 (29). Fanshawe, D. B., 130 = FD 2739, 450 = FD 3186 (38); 2863 = FD 6010 (15, type of Nectandra miranda). Ferrnndez, A., 3915 (38); 5476 (29). Ferreira-P.-F.,H., 83-31 (36 s.l.). Field, A., 154 (29). Folli, D. A., 334 (10). Fosberg, F. R., 22333 (1). Franco, P. (et al.), 3956, 4013 (1). Fr6es, R. L. & G. A. Black, 24512 (Paraia bracteata). Fuchs, H. P. & L. Zanella, 22093 (Endlicheria browniana). Gentry, A. (et al.), 21980 (Endlicheria sprucei); 45294, 45342 (29); 55420, 57819, 61109, 61141 (29). Glaziou, A.FRM., 822, 824, 1290, 12119, 12128 (10); 13150 (9, type of Nectandra anomala); 15367, 17742, 18454 (10). G6is, 0. C. & Octavio, 110 (9). Goldstein, I. & L. Salas, 97 (29). G6mez, L. D. & G. Herrera, 23064 (29). Gonggrijp, J. W. & Stahel, 188 (29). Goudot, J., s.n. (1, type of Synandrodaphne laxa). Graham, s.n. (10). Grdndez, C. (et al.), 1611, 1612 (Endlicheria ruforamula). Granville, J.-J. de (et al.), B-4469, B-4667, 4819 (29); 5077, 5077-A [seedling] (37); 8023 (13, type); 12022 (29); 12377 [type], 12383, 12457 (20). Gudinio,E., 143 (31); 533 (29). FLORANEOTROPICA Haber, W. A. & E. Cruz, 6791 (29). Hage, J. L. et al., 2357 (9). Hammel, B., 10494, 10693, 11226 (29). Hartshorn, G. S. (et al.), 1082, 1256, 2630 (29). Hatschbach, G. (et al.), 8936, 18272, 18487, 21269, 53737, 61392 (10). Heringer, E. P. et al., 5081 (Endlicheria paniculata). Hernmndez,L. & N. Dezzeo, 235 (36 s.l.). Herrera, G. (et al.), 2397, 4263 (29). Hodge, W. H. & M. Lescano, 1 (30). Hoffmann, W. A., 178 (10). Holst, B. K. (et al.), 3028 (25); 3287, 3324 (24). Horto-Florestal, (RB)102874, (RB)1 10103 (10). Huashikat, V., 1070 (29). Huber, O., 12557 (25); 12603 (24). Hurtado, F. et al., 1293 (29). Irvine, D., 996 (29). Irwin, H. S., 48584 (33). Jansen-Jacobs,M. J. et al., 3364 (13). Jaramillo, J. (et al.), 8349 (Endlicheria metallica); 13643, 13676 (7). Kawasaki, M. L., 179 (21). Kayap, R., 279 (29); 354 (Endlicheria metallica). Keman, C. & P. Phillips, 1105 (29). Killeen, T. (et al.), 2885, 3530, 3611 (29); 3832 (5); 3836 (29). Killip, E. P., 28400 (18, type). Klug, G., 1258 [type of Endlicheria loretensis], 1506 [type of Ocotea licanioides], 2556 (14). Kroll-Saldafia, B., 134, 326 (Endlicheria metallica); 642 (30). Krukoff, B. A., 4747 (36 st.); 4775, (5, type); 4905 (cf Ocotea); 4932 (Endlicheria metallica, type); 5098 (17); 5171 (36 st.); 5757 (8, type of Nectandra superba); 7996 (Paraia bracteata); 10173 (29); 11286 (Ocotea). Kuhlmann, J. G., 1845? = (RB)20051 (Endlicheria metallica); 20050 (29). Kuhlmann, M. & A. Gehrt, (SP)40043 (10). Kukle, P., 123 (17); 142 (29). Kurz, H., B-9, B-28-14 (40). Lanes, J., 31819 (29). Lanjouw, J. & J. C. Lindeman, 2300 (23). Liene et al., 3970 (10). Liesner, R. L. (et al.), 15655, 16935, 16949 (29). Lindeman, J. C., (LBB)3738, (LBB)4826, (LBB)4841 (36); (LBB)5884, (LBB)5959 (33); (LBB)6031, (LBB)6763 (36). Little, E. L. (et al.), 355, 6646, 21238 (29). Mackenzie, C. A. et al., (INPA/WWF)2108.35 (39). Madrifidn, S., 716 (29). Maguire, B. (et al.), 29830 (29, type of Pleurothyrium cowanianum); 32753, 33017 (25); 35290, 35320 [A], 35320 [B], 35340, 35450 [type of Ocotea celiana], 35450 [A] (24). Malme, G. 0. A., 3309 (Ocotea spixiana). LIST OF EXSICCATAE 99 L. (et al.), 248254 (38); 448-979 (29). Marcano-Berti, Rabelo,B. V. & R. Cardoso,2941 (36). Ramos,J. F & C. F. da Silva, 2834 (40). Marin,E., 805 (36 s.l.). Rangel,0. (et al.), 9008, 9051 (1). Martin,J., s.n. (29). chryso- Renner,S., 991 (17). Matthews,A., 3031 (29, type of Pleurothyrium thyrsus). Renterfa, E. et al., 2821 (Endlicheria metallica). Reynel-R.,C. (et al.), 39, 197 (29); 5115 (Endlicheria Meier,W., 1790 [type], 1797 (12). metallica). Me1inon,M., 1863 [1865 in B] (36, type of Endlicheria Ribeiro,J. E. L. S. (et al.), 943, 1003 (17). grandis). Riedel,L., s.n. [770 fide Mez] (10). Mexia, Y., 4707 (Endlicheria panicilata). Mori, S., 8612 (36); 10669 (10); 14737 (26); 14787, Riera,B., 488 (36); 1710 (13). 14896 (37); 14899 (26, type); 15014 (37, type); Rodrigues,W.A. (et al.), 1469, 1911, 2752, 5369 (17); 5435 [type],5649, 5717, 5763 (27); 6938 (34); 6976 15031, 15044 (33); 15103 (23); 15122, 15334 (33); [type],7009 (40); 7029 (21). 17169(13); 17717 (36); 18052 (Aniba);18147(33); L. & J. A. Dugarte,15884 (3, type). 18234 (29); 19196, 19199 (35); 19744 (17); 20744 Ruiz-Teran, (37); 20774 (35, type);20797 (37); 20850 (33, type); Sabatier,D. (et al.), 573, 1061, 1473, 1870 (36); 1975 20904 (37); 22324, 22670 (29); 23321 (37); 23393 (23); 2193 (36); 3500 (16, type); 3683 (36); 3710, (23); 23416 (16); 23821 (37); 23832 (23); 23872 4014 (23). (37); 23998 (36). Salas,L., TT-205(29). Mutchnick,P, 945 (13); 966 (23). 535 Sandwith,N. Y., 387 [typeof Nectandrapraeclara], (38). Nadeaud,J., s.n. (10). Santos, J. L. dos,(INPA/WWF)1202.5028(36 si.); (21). Nascimento,J. R. et al., (INPA/WWF)1201.288 (INPA/WWF)1202.5767(39). Nascimento,0. C., 735 (19). Santos,T. S. dos, 885 (10). Nee, M. (et al.), 30297 (29); 42516 (17). Saravia,R. S. & I. F. Rego, 1321 (29). Neill, D. A. & W. Palacios,7129 (30, type). Sastre,C., 4551 (20); 5901 (23). Nishimura, A., 93 (Endlicheria 2). aciphylR., 798, 1424 [typeof Aydendron Schomburgk, Nuniez,P. et al., 11424 (29). lum] (29). Occhioni,P., 8580 (10). Schultes,R. E. & I. Cabrera,13687(29); 13805(36 s.l.). Oldeman,R. A. A., B.1119 (29);B.3923 (36); 1448(29). Schulz, J. P., (LBB)7419, (LBB)7521, (LBB)8264, Oliveira,A. R. de, (INPA)59639, (INPA)59887 (Endli(LBB)8455,(LBB)9023(36); (LBB)8178(33). cheria metallica). Schunke-V.,J., 5406 (17); 6726 (5). OrtegaU., A., 005, 211 (29). Sellow,F., s.n. [197 fide Mez] (10, type of Goeppertia? macrocalyx). Palacios,W. (et al.), 1689, 4044 (29); 4454 (32, type); A. S. et. al., 72 (Paraiabracteata);486 (13). Silva, 4936, 5119 (Endlicheria metallica); 6247, 6251, Silva, I. A., 345 (10). 7412 (29). Silva,M. (et al.), 744 (17); 1411 (36 s.l.); 2402 (Paraia Pariona,W. & J. Ruiz, 963 (29). bracteata). R. 26-X (4, type). Pearce, W., Silva, M. G. (et al.), 3604 (36 si.); 4305 (Endlicheria Pennington,R. T. et al., 123 (8). metallica); 7148 (Endlicheria sprucei). Pennington,T. D. (et al.), 10760 (36 s.l.); 14950 (29). Silva, N. T. (et al.), 4672 (Endlicheria bracteolata); Pereira,E. (et al.), 714, 4481, 4520, 4535, 4536 (10). 60720 (36 s.l.). Pereira,0. J.,(et al.), 181, 1205, 2883 (10). A. F., 4757 (29). Skutch, Perry, D., s.n. (29). Phillips, 0. (et al.), 182 (29); 185 (Endlicheriametal- Smith, D. N. (et al.), 2886, 4615, 4720, 4905, 13691, 14354 (29). lica). Pires,J. M. (et al.), 10245, 10949, 13219 (Paraiabracteata); 16962 (29); 51370 (13). Pittier,H., 15274 (29, type of Nectandrameyeriana). Poiteau, P. A., s.n. (29, type of Acrodiclidium kunthianum). Prance,G. T., P. (et al.), 2115 [type] (17); 2181 (21); 3047, 3886, 3919, 4211 (17); 5709, 6432, 7675 (29); 7993 (17); 14013 (Endlicheria ruforamula); 14616 (Endlicheria chrysovelutina); 15860 (21); 17721 (21); 18761-B,(17); 20525 (17); 21642 (40); 21660 (21); 21690 (17); 23447 (21). Prevost,M. F. & D. Sabatier,2920 (23). Quelal,C., 312 (31, type). Smith, S. F. et al., 1503 (Endlicheria metallica). Sobel, G. L. et al., 4923 (29). Sobrevila,C. et al., 1871 (29). Soejarto,D. D. (et al.), 626, 4461, 4502 (29). Sperling,C. R. et al., 5928 (36 s.l.). Stahel,G. & J. W. Gonggrijp,BW 629, BW 670 (36). Stevens,W.D. et al., 24903 (29). Stevenson,P., 126 (30); 500 (29). Steward,W.C. (et al.), P.20098,P.20100,P.20308(17). Steyermark, J. A. (et al.), 59001 (Endlicheria metallica); 60403 (38); 87031, 92862 (29); 93476 [typeof Ocotea steyermarkiana], 93595 (25); 93935 (Ocotea 1); 96512 (29);97853 (25);99866 (29); 109118,109172 (25); 111392(38); 116883(29). 100 FLORA NEOTROPICA Teixeira,L. 0. A. et al., 578 (29). Tillett, S. S. (et al.), 672-83 (30); 45580 (22, type of Ocoteamirecolorata);45660 (29). Tipaz,G. (et al.), 794 (29); 1004 (5). T6mas,Bro., 163 (1). Tresling,J., 343 (29). Triana,J. J., 2040/3 = 1034 [= Schlim, L. J. No.451] (2, type of Nectandravelutina). Trujillo,B., 4122 (29). Ule, E., 9402 (8, type of Nectandradioica). (14); 19409 (36 s.l.); 19421, 19459 (Endlicheria me- tallica); 19472 (36 si.); 19537, 19541 (29). Velloso,H., 382 (10). Vicentini,A. (et al.), 468 (36 s.i.); 628 (21); 653 (39, type); 816 (21); 903 (Endlicheria metallica); 963 (40). Wanderbilt & D. Barros, 223 (Endlicheria paniculata). Weinberg,B., 01-A, 102 [type](11). Werff,H. van der (et al.), 8292 (29); 9815 (Endlicheria chrysovelutina);10188 (14); 11136 (5); 12190 (7, type); 12358 (29); 12954 (23). Williams,LI.(et al.), 114 (29); 1118 (14); 10718, 10728 (29). Wilson-Browne,G., 379 = FD 5793 (29). Valverde,L. & I. Pefia,1138 (29). Vargas,I. et al., 1101 (29). Vargas,W. et al., 1053 (1). Vasquez,E., 119, 120 (29). Young,K., 4072 (29). Vasquez,R. (et al.), 644, 4488, 5296 (14); 5697 (Endlicheria metallica);5950 (29); 6222 (14); 7689 (17); Zak,V. (et al.), 1163 (7); 4452 (29); 4671 (Endlicheria metallica). 8727, 8929 (14); 9564 (17); 10578(14); 11035(29); (34, type);f'12435 (Endlicheriametallica); 14533 Zaruma, J. et al., 453, 810 (Endlicheria metallica). INDEX OF LOCAL NAMES aiju'i-heter77 baaka-apisi86 bala oui 66 broad-leavedsoft silverballi90 buradie90 canau-yek77 canela cedro 48 canela do nativo 49 canelo 77 canelo batallia48 cedre jaune 86 cedre noir 86 86 groot-bladige-zwarte[harde]-pisi gueli[geli]-apisi 86 ishpingo 77 jigua 77 jigua amarilla77 jigua baboso 94 pisie 83 pisie oema 77 quisarraamarilla77 quisarranegra 77 roble 77 roble Amarillo 77 roble playa 77 shirua90 sweetwood 77 tinchi 77 wanchuctar77 watjarang86 yacu ahua yura 77 yalte 40 yana quillu caspi yura 77 yaya 77 yuwich 77 jigua pava 94 lapamaen77 laurel 77 laurel Amarillo 77 laurel grande90 louro bofo 77 louro branco59 louro do baixio 93 louro preto 59 moena 40, 77 moena amarilla79 moena amarillade hoja ancha 77 moena negra 79 muena negra 55 negrillo 77 oliveira-da-praia49 papaja-pisi86 INDEX OF SCIENTIFIC NAMES New names and combinationsin boldface; synonymsin italics. Page numbersin boldface indicateprimary page references;those with an asterix (*) indicate pages with illustrationsor maps. Acrodiclidium5 kunthianum5, 29, 78 Actinodaphne8, 9 Aiouea 3, 4, 5 Aniba 3, 4, 18 juruensis 39 Aydendron5 aciphyllum5, 73 Beilschmiedia 18 Caryodaphnopsis8 Cassytha filiformis 8 INDEX OF SCIENTIFICNAMES Chlorocardium8, 94 Cinnamomum8 Cryptocarya8, 18 Dicypellium 5, 68 Endlicheria3, 4, 5, 6, 7, 8, 9, 18, 19, 22, 23, 24, 25, 26, 29, 31, 55, 92, 93 subgen. Ampelodaphne8, 9, 55 anomala55 bracteolata25, 26, 29, 99 bracteolata25, 26, 29, 99 chrysovelutina25, 27, 28, 100 colombiana21, 40, 42 debilis 59, 60 gracilis 22, 71 grandis 85, 88 juruensis 39 krukovii25 ihotzkyi 25 loretensis 52, 55 metallica 24, 25, 26, 27, 28, 29, 31, 32, 88, 90, 97, 98,99, 100 paniculata48, 98, 100 rubriflora22, 61, 73 ruforamula25, 27, 28, sericea 25, 88 sprucei 57, 59, 98, 99 vinotincta7, 22, 51, 69 Goeppertia6 macrocalvx46 reflectens48 Licaria 3, 5 Mezilaurus3 Mespilodaphne floribunda48 Nectandra3, 4, 5, 6, 10, 18, 24, 88, 95 subgen. Synandrodaphne3, 4, 6, 31 subgen. Eunectandra4, 6 anomala 4, 6, 39, 44, 45 arnottiana4, 73, 78 dioica 4, 6, 24, 43, 44, 88 dominicana4, 6 grandis 5, 6, 7, 24, 44, 36, 88 kunthiana4, 5, 73 laxa 4, 34 meyeriana5, 75 miranda4, 6, 55, 56 paraensis56 praeclara 4, 6, 24, 25, 26, 88, 89 reticulata9 staminea3, 4 superba 4, 6, 24, 43, 44 velutina4, 36, 37 Ocotea complex 5, 18 Ocotea 3, 4, 5, 6, 10, 18, 55, 69, 98, 99 101 celiana 6, 66, 68 cemnuagroup 6 congestifolia 50, 85 cooperi 5, 73, 78 declinata36 deflexa 36 dominicana6 ferruginea69 helicterifoliagroup 5, 6 kunthiana73 licanioides 6, 52 macrocalyx6, 46, 48 mirecolorata6, 64 rotundata69 spixiana45, 98 staminea4, 95 stenoneura5, 78, 98 steyermarkiana6, 68 Oreodaphne4 fragrans 4 Pleurothyrium3, 4, 5, 6, 10, 23, 48, 68, 77 chrysothyrsus4, 73, 77, 78 cowanianum5, 73, 77 glabritepalum78 Paraia bracteata6, 56, 97, 98, 99 Persea 4 Phyllostemonodaphne22 Rhodostemonodaphne1, 2, 3, 4, 6, 7, 8, 9, 10, 11, 12*, 13*, 14, 16, 17, 18, 19, 20, 24, 25, 31, 45, 49, 55, 56, 57, 69, 73, 87, 95 anomala,6, 8, 12*, 14, 17, 18, 19, 20, 21, 33, 37*, 39, 41*, 44, 45, 46, 55 antioquiensis 10, 13*, 14, 17, 18, 20, 22, 32, 58*, 71, 72*, 73 avilensis 7, 8, 9, 14, 17, 18, 19, 20, 21, 23, 32, 37*, 47*, 50, 85 capixabensis6, 7, 10, 13*, 14, 17, 18, 19, 20, 21, 33, 37*, 45, 47*, 48, 49, 50, 68 celiana 6, 7, 10, 12*, 14, 16, 17, 18, 20, 22, 32, 51, 58*, 66, 67*, 68, 69, 95 crenaticupula 7, 12*, 14, 16, 17, 18, 20, 22, 33, 54*, 57, 58*, 59, 60, 66 curicuriariensis 7, 14, 16, 17, 18, 20, 22, 32, 33, 52*, 60*, 61 cyclops 10, 14, 17, 18, 19, 20, 21, 34, 37*, 40, 41*, 42, 43 debilis 7, 14, 17, 18, 20, 22, 33, 59*, 60 dioica 6, 9, 14, 16, 17, 18, 19, 20, 21, 24, 34, 37*, 38, 40, 41*, 43, 44, 64, 88 elephantopus7, 10, 14, 17, 18, 20, 23, 32, 35, 50, 80*, 81*, 84, 85 frontinensis 7, 14, 17, 18, 19, 20, 21, 34, 36, 37*, 40, 41*, 42, 43, 82 grandis 10, 13*, 14, 17, 18, 20, 23, 24, 24, 25, 26, 27, 28, 29, 30, 32, 44, 85*, 86, 87*, 88, 89, 90, 91 grandiscomplex 2, 8, 11, 23, 24, 25, 26, 27, 28, 29, 30, 31, 85*, 87*, 89, 91, 92, 93 grandisgroup 20, 22, 23, 26, 87* juruensis7, 10, 15, 17, 18, 19, 20, 21, 34, 39, 37*, 40* kunthiana4, 6, 7, 8, 9, 10, 11, 13*, 15, 16, 17, 18, 20, 23, 32, 50, 72*, 73, 75*, 77, 78, 79, 80, 82 kunthianagroup 19, 20, 21, 22, 23, 75*, 80* laxa 6, 7, 8, 10, 12*, 15, 17, 18, 19, 20, 21, 22, 31, 34, 35*, 36, 37*, 38, 39, 40, 42, 45 laxa group 6, 19, 20, 21, 22, 23, 37*, 38, 40 leptoclada8, 15, 17, 18, 20, 22, 33, 51, 58*, 61, 63, 67*, 69, 70, 71, 73 licanioides 6, 7, 8, 9, 12*, 15, 16, 17, 18, 20, 21, 22, 33, 51, 52*, 53, 54*, 55, 56 longiflora 15, 17, 18, 19, 20, 21, 34, 35*, 37*, 38, 40 longipetiolata 8, 10, 15, 16, 17, 18, 20, 23, 32, 80*, 81*, 82 macrocalyx 6, 9, 10, 12*, 15, 17, 18, 19, 20, 21, 33, 37*, 45, 46, 47*, 48, 49 102 miranda6, 7, 8, 9, 10, 12*, 15, 16, 17, 18, 20, 21, 32, 51, 52*, 53, 54*, 55, 56 mirecolorata6, 15, 16, 17, 18, 20, 22, 33, 38, 58*, 62*, 64, 65, 66 morii 9, 13*, 15, 17, 18, 20, 23, 33, 80*, 81*, 82, 83, 84 napoensis 15, 16, 178, 18, 20, 22, 23, 33, 72*, 79, 80*, 83 negrensis 12*, 15, 16, 17, 18, 20, 22, 23, 30, 33, 52*, 62*, 63, 64, 80 ovatifolia 15, 17, 18, 20, 21, 30, 34, 35*, 37*, 38, 40 parvifolia 7, 9, 12*, 15, 17, 18, 20, 22, 33, 58*, 67*, 70, 71 penduliflora 3, 8, 9, 10, 15, 17, 18, 20, 24, 32, 68, 80*, 93, 94* FLORA NEOTROPICA peneia 10, 13*, 15, 16, 17, 18, 20, 23, 24, 25, 26, 27, 28, 29, 32, 87*, 91, 92, 93, 94* praeclara 6, 10, 13*, 15, 16, 17, 18, 20, 23, 24, 25, 26, 27, 28, 29, 32, 87*, 88, 89*, 90, 92, 93 recurva6, 9, 10, 15, 16, 17, 18, 20, 23, 24, 25, 26, 32, 87*, 91*, 92 revolutifolia9, 10, 15, 17, 18, 20, 21, 22, 31, 33, 52*, 54*, 56, 57 rufovirgata8, 13*, 15, 16, 17, 18, 20, 22, 33, 58*, 62*, 65, 66 sauilensis13*, 15, 17, 18, 20, 22, 23, 24, 25, 26, 27, 28, 29, 32, 44, 64, 87*, 88*, 89, 91 scandens 7, 8, 9, 10, 12*, 15, 16, 17, 18, 20, 21, 22, 33, 47*, 51, 52*, 53, 55, 56, 69, 80 scandensgroup 20, 21, 22, 23, 52*, 58* sordida 9, 15, 17, 18, 20, 33, 80*, 81*, 83, 84 steyermarkiana6, 7, 12*, 15, 16, 17, 18, 20, 22, 32, 51, 58*, 67*, 68, 69 synandra6, 10, 11, 13*, 15, 16, 17, 18, 20, 23, 32, 72*, 75*, 78, 79 tumucumaquensis 7, 15, 16, 17, 18, 20, 22, 33, 52*, 61, 62*, 63, 70 velutina 6, 7, 15, 17, 18, 19, 20, 21, 34, 35*, 36, 37*, 38, 40, 42 Synandrodaphne3, 4, 6, 31 antillana 3, 4 laxa 3, 4, 34 Urbanodendron6, 68, 95

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Rhodostemonodaphne (Lauraceae)

Rhodostemonodaphne (Lauraceae)

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