. i

The of a Central Amazonian Moist Forest Memoirs of The New York Botanical Garden

ADVISORY BOARD

PATRICIA K. HOLMGREN, Director of the Herbarium The New York Botanical Garden

JAMES L. LUTEYN, Senior Curator SCOTT A. MORI, Senior Curator The New York Botanical Garden The New York Botanical Garden

EDITORIAL BOARD

WILLIAM R. BUCK, Editor The New York Botanical Garden Bronx, New York 10458-5126

WM. WAYT THOMAS, Associate Editor The New York Botanical Garden Bronx, New York 10458-5126

THOMAS F. DANIEL (1991-1996) Department of Botany California Academy of Sciences Golden Gate Park San Francisco, California 94118

The MEMOIRS are published at irregular intervals in volumes of various sizes and are designed to include results of original botanical research by members of The New York Botanical Garden's staff, or by botanists who have collaborated in one or more of The New York Botanical Garden's research programs. Ordinarily only manuscripts of 100 or more typewritten pages will be considered for publication.

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TillS PUBLICATION IS PRINTED ON ACID-FREE PAPER The Lecythidaceae of a Central Amazonian Moist Forest

Scott A. Mori and Nadja Lepsch-Cunha

..~ ..

Cour%ri quionensis

Couralori longipedicel/ala

Lecy/his .rabucajo

Lecylhis porvifruc!o

£. grand/flora

E. brocleo$o £. omozoniciformis Eschwei/ero loev/corpo

The New York Botanical Garden Bronx, New York 10458, U.S.A. Issued: 29 December 1995

MEMOIRS OF THE NEW YORK BOTANICAL GARDEN VOLUME 75 Copyright © 1995

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Mori, Scott A., 1941­ The Lecythidaceae of a central Amazonian moist forest I Scott A. Mori and Nadja Lepsch-Cunha. p. cm.-(Memoirs of the New York Botanical Garden ; v. 75) Includes bibliographical references (p. ) arid index. ISBN 0-89327-396-1 I. Lecythidaceae----Amazon River Region. 2. Rain forest - Amazon River Region. I. Lepsch-Cunha, Nadja, 1962­ II. Title. III. Series QKl.N525 vol. 75 [QK495.L42] 581 s-dc20 [583'.42] 95-37040 CIP MEMOIRS OF THE NEW YORK BOTANICAL GARDEN 75: I-55

The Lecythidaceae of a Central Amazonian Moist Forest

SCOTT A. MORI AND NADIA LEPSCH-CUNHA

Contents

AbstractIResumo . 2 Introduction ...... , 3 Taxonomic Treatment , , 3 Habitat 4 Distribution and Representative Specimens 4 Phenology , 4 Common Names , 4 Uses 5 Comments , 6 Tenninology , 6 Size , , .. , , 6 Tree Bole , 6 Bark...... 6 Stems , , 7 ...... , , 7 " " 8 , .. , .. , .. , 10 Taxonomie Problems , 14 Description of Lecythidaceae Subfamily Lecythidoideae 16 ~of~s., ." .. ,...... 16 Key to Genera, . , 16 Chave para os G€neros , , . , , . , , , .. , 17 Descriptions of Genera and Species 17 I. Allanloma,."",."., . , , 17 I. Allanloma lineafa (Martius ex Berg) Miers 17 2. Bertholletia, , 19 1. Bertholletia exce/sa Humboldt & Bonpland , , 19 3. Carin/ana.... . , " , , .. , , 21 1. decandra Ducke , 21 2. C. Inicrantha Ducke , ...... 22 4, , ...... 22 I. CorythopllOra alta R. Knuth , 24 2, C rirnosa W, Rodrigues subsp. rimosa 24 5. ., , ...... 24 I. Aublet . . . 26 2, C longipedicelfata W. Rodrigues...... 27 3, C mulriflora (1. E. Smith) Eyma ...... 27 4, C stelfata A C. Smith. . . .. 27 5. C. (auari Berg. ' ...... 29 6. ",...... 29 1. Eschweilera OITUU,oniciformis Mori ...... 32 2. E. atropetiolata Mori. . . 32 3. E. bracteosa (Poeppig ex Berg) Miers. . . 33 4. E. carinata Mori ...... 33 5. E. col/ina Eyma...... 33 6. E. coriacea (A. P. de Candolle) Mori . . 34 7. E. cyathiformis Morl . . . . . 34 8. E. grandiflora (Aublet) Sandwith...... 35 9. E. laevicarpa Mori...... 35 10. E. micrantlUJ (Berg) Miers...... 36 2 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN [VOL. 75

11. E. pedicel/ala (Richard) Marl .36 12. E. pseudadecolorans Marl .. .36 13. E. raTlkiniae Man .39 14. E. romeu-cardosoiMori . .39 15. E. tessmannii R. Knuth . .41 16. E. truncata A. C. Smith . .41 17. E. wachenheimii (R. Benoist) Sandwith .. .42 7. ,., . .42 I. Gustavia elliptica Marl . .42 8. Lecyrhis , . ..44 1. barnebyi Marl . . .. 45 2. L. gracieana Marl . .47 3. L. parvifructa Marl . . .. 49 4. L. pisanis CambessMes .. . .. 49 5. L. poiteaui Berg . .. 50 6. L. prance; Marl . . .. 50 7. L. retusa Spruce ex Berg ... 50 8. L. zabucajo Aublet. . . .. 52 9. L. sp. 01. . . .. 53 10. L. sp. 05 .. ..53 Acknowledgments . . . 53 Literature Cited . ..54 Index to Scientific Names .. . .. 55

Abstract

Morl, Scolt A. (Institute of Systematic Botany, The New York Botanieal Garden, Bronx, New York 10458-5126, U.S.A.) and Nadja Lepsch-Cunha (Dept. de Ciencias-Lab. de Sementes, Escola Superior de Agricultura Luiz de Queiroz, 13418-90D-Piracicaba-Sfio Paulo, ). The Lecythidaceae of a central Amazonian moist forest. Mem. New York Bot. Gard. 75: 1-55. 1995. The results of a taxo­ nomic study of the 39 species of the family (Lecythidaceae) found in the lOOO-ha control reserve (Reserve 1501 or "Km 41 ") of the Biological Dynamics of Forest Fragments Project of thc Smithsonian Institution and the Instituto Nacional de Pesquisas da Amaz6nia are reported. Descriptions, keys, habitat preferences, phenology, distribution and representative specimens, com­ mon names (when available), and comments are provided for all species found in the reserve. An illus­ tration of at least one species in each genus is ineluded, and discussions and illustrations of features useful in distinguishing eentral Amazonian species of Lecythidaceae are presented. Three new spccies, Eschweilera rankiniae, E. romeu-cardo.wi. and Lecylhis gracieana, are described. Keys for identification of the genera and species within genera are provided in Portuguesc as well as English.

Resumo

Mori, Scott A. (Institute of Systematic Botany, The New York Botanical Garden, Bronx, New York 10458-5126, U.S.A.) e Nadja Lepsch-Cunha (Dept. de Ciencias-Lab. de Sementes, Esco1a Superior de Agricultura Luiz de Queiroz, 13418-90D-Piracicaba-Sao Paulo, Brasil). The Lecythidaceae of a central Amazonian moist forest. Mem. New York Bot. Gard. 75:1-55. 1995. Os resultados de urn estudo raxon6mico das 39 esp~cies da familia de Castanha-do-Para (Lecythidaceae) que ocorrem numa reserva controle dc 1.000 ha (Reserva 1501 ou "Km 41") do Projeto Dinamica Bio16gica de Fragmentos Florestais sao reportados. Descri90cs, chaves, prcferencias ccol6gicas, distribui9ao e esp~cimes representativos, fenologia, nomes vulgarcs, e comentarios sobre cada esp~cie que se­ encontra na reserva sao aprescntados. Incluem-se comentarios gerais das caractensticas que sao empregadas na identifica9ao de generas e esp~cies dc Lecythidaceae e, pelo menDs, urn desenho de uma esp~cie em cada genero. Tres novas cspl:cies, Eschweilera rankiniae, E. romeu-caniosoi, e Lecythis gracieana sao dcscritas. Chaves para a identifica9iio de generos e esp~cies em Ingles bern como cm Portugues sao apresentadas. 1995J THE LECYTHIDACEAE OF A CENTRAL AMAZONIAN MOIST FOREST 3

Introduction yet to be solved. Mori is convinced that the best way to resolve taxonomic problems in neotropical is OUf central Amazonian study of Lecythidncene is to couple taxonomic and ecological studies. Study of part of the much larger Biological Dynamics of herbarium specimens alone somctimes is not suffi·

Forest Fragments Project CBDFFP) of the Smithson w cient for detecting the nuances of differences among ian Institution and the Instituto Nacional de Pesquisas closely related species of tropical trecs. da Amazonia (INPA). The BDFF Project was previ­ Reserve 1501, also known as Km 41, was selectcd ously known as the Minimum Critical Size of Eco~ for our study because it is the 1OOO-ha control reserve systems Project under the aegis of the World Wildlife of the BDFF Project. The first digit of the rescrve indi­ Fund and INPA. A description of the BDFF Project is cates the ranch upon which the reserve is located (I = provided by Lovejoy and Bierregaard (1990). Fazenda Esteio), the second digit indicates the size The principal goal of the BDFF Project is to assess class of the reserve (5 =control area), and the last two the effects of fragmentation on once continuous trop­ digits indicate the number of replicates of that partic· ieal rain forest. However. as stated in a call for pro­ ular size in the BDFF Project (there is only one con· posals dated 16 October 1990, "Understanding the trol area). Reserve 1501 is also called Km 41 because relative impact of the forest fragmentation processes it is located 41 kilometers along state highway ZF-3 requires as background data detailed inventories, Irom federal highway BR-174 (the Manaus-Boa Vista knowledge of species biology, and undcrstanding of Highway). A map of Reserve 1501, as well as of all the abiotic environment within continuous forest." other reserves in the BDFFP system, is provided by The Lecythidaceae Sub-Project of the BDFF Lovejoy and Bierregaard (1990). Project is an attempt to provide the basic biological Establishment of the Lecythidaceae Sub-Project information needed for understanding the effects of consisted of (I) location of a 100-ha plot and its sub­ forest fragmentation on one of the ecologically most division into hectares, which in turn were divided important families of trces throughout Amazonia and into 20 x 20 m quadrats by a professional surveyor; the Guianas (Mori, 1990; Mori & collaborators, (2) location and tagging of all individuals of 1987; Mori & Pranee, 1990a; Prance & Mori, 1979). Lecythidaceae equal to or greater than 10 em dbh; The specific goals of our work are to (I) rcach an (3) mapping of the individuals of Lecythidaceae; understanding of the of all species of cen­ (4) identification [Q species of the 7791 individuals tral Amazonian Lecythidaceae, (2) determine what of Lecythidaccae; (5) collection of data on topogra­ ecological parameters (topography, past history such phy and soils; and (6) entry of biotic and abiotic data as burning and flooding, edaphic factors, phenology, into a database. intrafamilial interactions, interfamilial interactions, The Lecythidaceae plot is available for use by and interactions with animals) influence the distribu­ other researchers. Studies of Sapotaceae (by T. D. tion of species of Lecythidaccae, (3) determine mini­ Pennington) and palms (by A. Henderson and A. mum population sizes necessary for the continued Scariot) are under way. We encourage others inter­ survival of central Amazonian Lecythidaceae (i.e., ested in the diversity and structure of neotropical where will reservcs have to be placed and how big forcsts to submit research proposals to the BDFF will they have to be), and (4) establish a protocol for Projeet in which the infrastructure of the Lecythida­ sampling Lecythidaceae and, by extrapolation, other ceae plot is utilized. neotropical trees. The purpose of this paper is to provide the basic The Lecythidaceae Sub~Project began in 1987, taxonomie information about Lecythidaceae needed when Peter Becker (ecologist) approached Mori (tax· to promote further systematic and ecological research. onomist) about collaboration. Becker was interested The results of other aspects of our research have been in the dynamics of tropical forests and therefore was (Bassini & Becker, 1990; Mon, 1992; Mon & Beeker, looking for a family of tropical trees that met three 1991) or will be published elsewhere. A paper on the criteria: high diversity, high density, and relatively diversity, density, frequency, and dominance of well·known taxonomy. On the other hand, Mori, who Lecythidaceae in the 100-ha plot is in preparation. had just finished a taxonomic and ecologieal study of Lecythidaceae in central French Guiana (Mod & col­ Taxonomic Treatment laborators, 1987), was looking to expand his studies of Lecythidaceae into Amazonia, especially central Modern taxonomic trcatments of neotropical Le· Amazonia, where diversity of Leqthidaceae is high cythidaceae (Mori & Prance, 1990a; Prance & Mori, and where there are numerous taxonomic problems 1979) are available. Therefore, detailed synonymy, 4 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN [VOL. 75

complete lists of specimens studied, and a list of HABITAT exsiccatae are not provided. However, in order to doe­ Nearly all Lecythidaceae at Reserve 1501 are ument species concepts, we cite several representa­ species of cerrafirme because this is the predominant tive collections for each species and provide complete habitat of the reserve. At least one species ( specimen citations for species described as new to lineata), however, is restricted to the less frequent science in this publication, Annotated specimens doc­ welter habitats along streams. umenting OUf species concepts are available for study at INPA. The New York BOlanical Garden (NY). and. DISTRIBUTION AND REPRESENTATIVE to a lesser extent, the Smithsonian Institution (US). In addition, a list of exsiccatae for Reserve 1501 is SPECIMENS deposited in the library of The New York Botanical A statement of the overall geographic distribution Garden and is also available from the senior author of each species is provided. Distribution maps for all upon request. species known from more than the degree grid square Most of the collections examined for this treatment surrounding Manaus (inclUding Reserve 1501), came from the lOO-ha Lecythidaceae plot at Reserve except newly described species, can be found in 1501, in which 7791 trees of Lecythidaceae ~1O em Prance and Mori (1979) and Mori and Prance dbh are tagged and mapped. In this plot there are 38 (I 990a). A slalemenl aboullhe abundance (Tahle I) of species of Lecythidaceae. One species treated here, each species in the 100-ha Lecythidaeeae plot is pro­ Lecythis poiteaui, has not been recorded from the vided, but more details about ecology will be pre­ 100-ha plot but does occur in Reserve 1501. More­ sented in future publications. over, all species of Lecythidaceae known to occur in Several representative collections are cited for each the reserves of the BDFFP also occur at Reserve species. The collector's name and number, fertility 150I. Consequently, this treatment may be used for state of the specimen (st = sterile, fl = flowers, fr = identifying Lecythidaceae from all of the BDFFP fruits), and tree number in the 100-ha plot are pro­ reserves as well as from terrafirme forests surround­ vided. The specimens are deposited in the herbaria of ing Manaus. It will not be useful for identifying NY, INPA, and, to a lesser extent, US, species of Lecythidaceae found in habitats not found in the BDDFP reserves such as periodically inun­ PHENOLOGY dated forests (vdrzea and igap6). A general statement about the flowering and ­ Descriptions of genera include those features that ing times of each species of Lecythidaceae at Reserve characterize the genera throughout their ranges. 1501 is provided. This information comes from a bi­ However, descriptions of species are based, for the weekly phenological survey between 1988 and 1991 most part, on the features of the species as they occur and from fertile herbarium collections. Because the at Reserve 1501. When complete material for a woody fruits ofLecylhidaceae often persist long after species was not available from Reserve 1501, collec­ the seeds are dispersed, caution has to be used in in­ tions from other areas were utilized to complement terpreting fruiting seasons on the basis of herbarium the descriptions. collections. Additional information about phenology Detailed exploration continues to add to our taxo­ in other areas is included when appropriate. A more nomic knowledge of neotropical trees. Of the 39 spe­ detailed analysis of phenology will be provided in a cies of Lecythidaceae found at Reserve 1501. 13 have future paper. been described since 1978, including three published herein. Two other species of Lecythis in the plot have COMMON NAMES been collected in the sterile condition only and may Common names of Lecythidaceae are most accu­ represent new species. Attaching a name to these two rate at the generic level. The following list provides species awaits the collection of flowers and fruits. the Amazonian names most commonly applied to the Resident collectors at Reserve 1501 have played an genera of Lecythidaceae found at Reserve 1501. important role in helping solve taxonomic problems in the family. Members of the team have worked hard to ALlancoma = ceru collect poorly known species, to gather specimens BerthoLletia = castanha, castanheira representative of all stages of the life cycle of species, Cariniana = tauarf and to provide materials needed for studies of cytol­ Coryrhophora = castanha-jacare. castanha-rana, ogy, anatomy, and chloroplast DNA of Lecythidaceae. ripeiro 1995J THE LECYTHIDACEAE OF A CENTRAL AMAZONIAN MOIST FOREST 5

Table I

Number of individuals of species of Lecythidaceae in the 100-ha plol at Reserve 150 I

Species Code for Species No. of Individuals

Alltmtomll lineata ALL! 5 BertllOlletiu exce/sa BEEX I Carinianu decandra CADE 40 C. micrunlllQ CAMI 29 Coryfhoplwra alta COAL 192 C. rimOSll subsp. rima.'W CORI 208 Courutarj guillnensis CUGU 21 C. longipedicellata CULO 14 C. multiflora CUMU 24 C. stelfura CUST 77 C. IQuu,; CUTA 3 E.n:hweileru umuwnicifurmis ESAF 190 E. lltropetiolura ESAT 571 E. bracteo.w ESBR 105 E. cllrinata ESCA 10 E. collina ESCO 55 E. coriacea ESCR 1539 E. cyutlliformis ESCY 275 E. grandiJJora ESGR 335 E. laevicurpo ESLA 57 E. micTtllltlra ESMI 180 E. pedic:ellata ESPE 100 E. pseudodec%rans ESPS 118 E. rankiniae ESRA 16. E. romeu-c:ardo,w; ESRC 309 E. le.wnannii ESTE 250 E. fruncala ESTR 1321 E. waclrenlzeimii ESWA 926 GU.f/tlvia elliptica GUEL III Lecythi.f barnebyi LEBA 105 L. gracieana LEGR 40 L. parvifructtl LEPA 30 L. risoni:; LEPI 46 L. pojreaa; LEPO o L. prance; LEPR 362 L. remSQ LERE II L. zabucajo LEZA 45 Lecythis 01 LEOI 8 Lecyrhis 05 LE05 51

Couralari = tauari that has appeared on collection labels or in the litera­ Eschweilera = matamatfl., ripeiro ture or (2) the names given [0 the species by woods­ Gustavia =jeniparana, mucurao men associated wilh the BDFF Project. Lecythis = jarana, sapucaia Because of the difficulty in distinguishing species of Lecythidaceae, the application of common names USES to species is mueh more diffieult. This is espeeially Known uses of the species are provided under this tfue in Eschweilera, where even the most experienced heading. The timber of all species and the fibrous woodsmen have difficulty in consistently and accu­ bark (see discussion of bark) of most species have rately distinguishing species. When possible. we have some uses. Therefore, use of wood and bark is not selected common names for species on the basis of recorded under individual speeies unless there is a (I) what we consider to be the most aceurate name specific reference to that species. 6 MEMOIRS OFTHE NEW YORK BOTANICAL GARDEN [VOL. 7S

COMMENTS Table II Statements about (l) pollinalOrs and dispersal Tree size of species of Lecythidaceae at Reserve 1501 agents, (2) features that allow for identification of the species, and (3) indications of taxonomic problems Understory Canopy Emergent remaining to be solved are included in this section. (525 m) (25-35 m) (>35 m) CUTA ALL! BEEX ESBR COAL CADE TERMINOLOGY ESGR CORI CAMI ESPE CULO CUGU In order to facilitate the identification of Lecythi­ ESWA ESAT CUMU daceae in central Amazonia, we provide below short GUEL ESCO CUST descriptions of vegetative, floral, and fruit features of LEBA ESCR ESAF LEOI ESCY ESCA taxonomic importance. For more detailed informa­ ESLA ESMI tion and additional illustrations. the reader is referred ESPS LEPA to Morl and collaborators (1987), Mori and Prance ESRA LEZA (1990a), and Prance and Mori (1979). ESRC LEOS ESTE ESTR LEGR Tree size LEPI Species of Lecythidaceae of central Amazonia can LEPO be arbitrarily divided into understory (to ca. 25 m LEPR LERE tall), canopy (25 to 35 m tall), or emergent (35 to 50 m tall) height classes. Maximum height at maturity is Note: These are subjective estimations of tree size based on used for determining the height classes, Examples are label data and the experience of the senior author. See Table I for species codes. Eschweilera bracteosa, E. grandiflora, E. pedicel­ lata, E. wachenheimii, and Gustavia ellipticQ as understory species; CorythopJzora alta, E. laevicarpa, and as eanopy species; and gent species often, but not always, have some indi­ Cariniana micrantha and Couratari multiflora as cation of buttressing. In central Amazonia, species of emergent species (Table II). Using these subjective Couratari show the greatest tendency toward but­ criteria, we determined that 20.5% of the species of tressing (Table III). Speeies of buttressed Lecythida­ Lecythidaceae at Reserve 1501 are understory, 48.7% ceae appear to be less frequent at Reserve 1501 are canopy, and 30.8% are emergent. A study of tree (23.1 %) than they are in central French Guiana (33%) height versus tree diameter is under way for the (Mori & collaborators. 1987). Lecythidaceae of Reserve I50 I in order to determine if these arbitrary size classes are valid and therefore an indication of forest stratification. Bark The bark of Lecythidaceae is unusually fibrous and is therefore useful in identifying the family in the Tree bole field. The bark of most species peels in long, flexible The presence or absence of buttresses may provide strips that are used or have been used for bark cloth­ some help in the identification of Lecythidaceae ing, caulking boats, cordage for tying together make­ (Table UI). This feature, however. usually has to be shift shelters. making belts for tree climbing, and applied to adults because species with buttresses at even for cigarette paper. The anatomical basis for maturity often do not develop buttresses until they fibrous bark in Lecythidaceae is the stratification of reach larger sizes. Moreover, our placement of a the bark into alternating bands of fibers and softer, species in the nonbuttressed or buttressed category is conducting tissue (Mori. Black & de Zeeuw, 1987). A subjective because buUress size. number of buttresses similar fibrous bark is also found in the Annonaceae, per tree, and number of individuals of a species with and therefore the two families are sometimes con­ buttresses have not been measured. In general. under­ fused in the field. However. the bark of Annonaceae story species of Lecythidaceae lack buttresses, differs from that of the Lecythidaceae by more con­ canopy species mayor may not have them, and emer· spicuous rays in the inner bark (with the exception of 1995] THE LECYTHIDACEAE OF A CENTRAL AMAZONIAN MOIST FOREST 7

Table III rimosa, and most species of LecytJu"s) have distinctly fissured barks; others (e,g" Eschweilera collina and E. Nonbuuressed and buttressed species of romeu-cardosoi) have smooth bark that may possess Lecythidaceae at Reserve 1501 hoop marks and lenticels. Fissured barks in Lecy­ Nonbuttressed Buttressed thidaceae are the result of the accumulation of mul­ tiple periderms, whereas smooth barks in this family ALL! CUGU are probably caused by the periodic sloughing of ring BEEX CULO & CADE CUMO periderms (Mori, Black de Zeeuw, 1987). Another CAMI CUST group of species (e,g" Eschweilera laevicarpa, E, COAL ESCR micrantna, E. pseudodecolorans, and Gustavia ellip­ CORI ESLA lica) have sealloped barks. The scallops are caused by CUTA ESRA irregular peeling of the outer bark because of the for­ ESAF LEPI ESAT LEZA mation of shell periderms (Mori, Blaek & de Zeeuw, ESBR 1987), Finally, several species (e,g., E, coriacea, E. ESCA pedicellata, and E. wachenheimil) have barks that ESCO cannot be classified as fissured or smooth. From a dis­ ESCY ESGR tance they appear smooth, but upon closer examina­ ESMI tion they are very finely eracked and somewhat rough. ESPE ESPS ESRC ESTE Stems ESTR All species of Lecythidaceae possess cortical bun­ ESWA dles. a feature that is useful in deciding if sterile col­ GUEL LEBA lections belong to the Lecythidaeeae because few LEGR other families of tropical trees consistently have cor­ LEPA tical bundles (Mar; & Blaek, 1987). LEPO LEPR LERE LEO! Leaves LEOS descriptions follow the terminology of Hickey Note: These are subjective estimations of tree bunressing (1979). Leaf blade measurements were made on the based on label data and the experience of [he senior author. largest leaf on a speeimen. Intraspecific leaf variation See Table I for species codes. in Lecythidaceae is considerable. Leaves from the top of the crown are usually smaller and thicker than GustQvia), the frequent occurrence of a slight spicy those from the lower part of the erown (Medri & odor in contrast to a slightly fetid odor in Lecythida­ L1eras, 1979). Likewise, leaves associated with inflo­ eeae, the lack of cortical bundles in the stem (see rescences are usually smaller than leaves found on below), and the laek of unilaterally thickened, crystal­ vegetative branches. Lastly, the appearance of the bearing cells in the phloem. leaves changes as the leaves lignify. The newly The slash, a diagonal cut through the bark with a flushed leaves of all species of Leeythidaceae are machete, is used by Amazonian woodsmen to facili­ much more chartaceous in texture than are the older tate species identifieation. In the Lecythidaceac, the more coriaceous leaves. most important slash features are the relative thick­ Some species of Lecythidaceae are deciduous. ness of the inner and outer barks and the color of the Within the deciduous species, some drop their leaves inner bark. For example, in Corythophora, C. rimosa and then tlush new leaves and at more or less subsp. rimosa has very thick outer bark and thinner, the same time (e.g., , L poiteaui, and dark red inner bark, whereas C. alta has thin OUler L. l.abucajo), whereas others (e,g" Couratari guia­ bark and thicker, pinkish inner bark. nensis and C. multiflora) drop their leaves, flower On the basis of external appearance, there are four when leafless, and then flush new leaves, bark types represented at Reserve 1501. Some species The leaves of species of Lecythidaceae are alter­ (Bertholletia excelsa, Corythophora rimosa, subsp. nate, simple, and lack conspicuous stipules, Because 8 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN [VOL. 15

the leaves of many species are morphologically simi­ collaborators. 1987; Mori & Prance, 1990a; Prance & lar, it is often difficult to identify species on the basis MorL 1979), and therefore only the terminology and of leaves alone. Nevertheless, there are a number of features needed for identifying central Amazonian leaf features that are useful for identifying species of speeies of the family will be presented here. Lecythidaceae. The two basie types of flowers of Leeythidaeeae Leaf venation offers some help in identification. are actinomorphic (Allantoma and Gustavia) and The leaves of Allantoma lineata and Cariniana zygomorphic (the remaining genera) (Fig. I). An decandra differ from the leaves of other species by exception is found in Cariniana, in which both being eucamptrodromous throughout and by their aetinomorphic (c. decandra) and zygomorphie (c. strongly percurrent tertiary veins. The leaves of other micrantha) flowers are found. Possession of both species have a greater tendency toward brochidodro­ types of flower suggests that Cariniana is poly­ mous venation and retieulate tertiary veins. The phyletic (see discussion of taxonomic problems). contrast between plane and salient secondary and Zygomorphy and evolutionary trends in the family higher order venation is sometimes taxonomically are most strongly expressed in the androecium. useful. For example, some species have very shiny Therefore, illustrmions of and terminology associated and smooth adaxial leaf surfaces (e.g., Eschweilera with the specialized androecium ofLecythidaceae are amazonicijormis and E. cyathijormis). The most dis­ presented in Figure 2. Variation in androeeial type has tinetive venation is found in E. coLlina, which has been diseussed in detail elsewhere (Mori & collabo­ distinctly salient venation on both surfaces after its rators, 1987; Mori & Prance, 1990a; Prance & Mori, leaves have been dried. 1979). The presence of longitudinal striations on the The androecia of species of Lecythidaceae of cen­ leaves of Couratari stellata and Eschweilera grandi­ tral Amazonia fall into one of the following six types: flora is useful for the identification of these species. (I) actinomorphic with no prolongation of the staminal A whitish cast to the abaxial (lower) leaf surface, ring (Allantoma, Fig. 9; Cariniana decandra and which is caused by the presence of microseopie, Guslavia, Figs. 1,3, 17); (2) very small with the sta­ cuticular papillae, helps to separate Berthollelia minal ring slightly prolonged on one side but not form­ acelsa, Eschweilera atropetiolata, E. carinata, ing a fleshy hood (Cariniana micrantha. Fig. II); (3) Lecythis barnebyi, L. parvijructa, L. poiteaui, and L zygomorphic with a flat androecial hood (e.g., Lecythis retusa from the remaining species. pisonis and L zabucajo, Fig. 3); (4) zygomorphic with Leaf size, even though extremely variable, does the hood appendages markedly curved inward provide some help in identifying some species. The (Bertholletia excelsa, Fig. 10; several species of largest leaves are found in Lecythis barnebyi and the Lecythis, Figs. 19,20); (5) zygomorphic with the hood smallest in Eschweilera rankiniae. Leaf shape, completely coiled inward (Eschweilera spp., Figs. 3, although mostly elliptic, differs in some species of 14,15,16); and (6) zygomorphic with the hood coiled central Amazonian Lecythidaceae. The leaves of E. inward and then outward (Couratari spp., Fig. 13). rankiniae, L. barnebyi, and L parvijructa tend to be Flower size and color are important taxonomic obovate, and those of some other species, e.g., B. features of Lecythidaceae. Large and small flower excelsa and L refusa, tend to be oblong. sizes are species-specific and probably are related to the size of the principal pollinators. The largest flow­ ers are found in the bat-pollinated Lecylhis barnebyi Flowers and L poiteaui. Other large-flowered species, such as Most species of Lecythidaceae possess diurnal Eschweilera grandiflora and E. pedicellata, are polli­ flowers. They usually open their flowers just before nated by robust-bodied species of bee such as EII­ or at daybreak and drop them in the Ime afternoon. laema spp. Some diurnal flowers may open during the early Flowers of Lecythidaceae are generally white, morning hours well before daylight, and others may white with various tinges of pink or red, or yellow. even persist through the night of the day in which Floral parts may be colored differently. A common they opened. However. Lecylhis barnebyi and L color combination in species of Eschweilera is white poiteaui are truly nocturnal, opening their flowers pemls with a yellow amJroecial hood. In other species, after dark and dropping them before daybreak. the hood may be white on the outside but yellow The floral structure of Lecythidaceae has been around the margins and on the inside (e.g., Berthol­ discussed in considerable detail elsewhere (Mori & lelia excelsa and Lecytltis prancei). Exceptions are the [995J THE LECYTH[DACEAE OF A CENTRAL AMAZON[AN MOIST FOREST 9

Gustavia hexapetala Eschweilera pedicel/ata Acti nomorphic Zygomorphic

Fig. 1. AClinomorphic and zygomot]Jhic flowers of Lecythidaceae. Gustavia IIexaperala is a common species in central Amazonia, but it has not yet been collected in the BDFFP reserves.

green petals of Corylhophora rimosa sUbsp. rimosa Locule number and placentation are useful for and Lecythis poiteaui. The color of the anthers of helping distinguish genera. The ovaries of Allantoma Lecythidaceae may differ even within the same are 4-locular with the ovules inserted along the sep­ flower. In general, those species with differentiated tum; those of Bertholletia are 4-locular with the pollen as a pollinator reward have yellow fodder ovules inserted at the base of the septum; those of anthers and while fertile anthers. Yellow seems to be Cariniana and Couratari are 3-locular with the important in directing bee pollinators to the pollinator ovules inserted at the base of the septum; those of reward in the Lecythidaceae. Corylhophora are 2-locular with the ovules inserted There are three types of pollinator reward offered at (he base of the septum or on the floor of the locule; by Lecythidaceae-nondifferentiated pollen, fodder those of Eschweilera are usually 2-locular with the pollen, and nectar. At Reserve 1501, GUSlavia and pre­ ovules inserted on the floor of the locule, but there are sumably Allantoma attract pollinators by offering also 4-locular species; those of Gustavia are 4(-6)­ nondifferentiated pollen as the reward, Le., the pollen locular with the ovules inserted on an expanded collected by the pollinators and used to feed their placenta toward the apex of the septum; and those of larvae is the same as the pollen eventually effccting Lecythis are 4-locular wilh the ovules inserted along fertilization. Other species, such as Corythophora or toward the base of the septum. rimosa subsp. rimosa, Lecythis pisonis, and L. The style also provides some diagnostic features. zabucajo, produce a differentiated fodder pollen in the The styles of species of Allantoma, Cariniana. Cory­ hood that the pollinator collects. As the pollinator thophora. Couratari. Eschweilera, and some species gathers the fodder pollen, fertile pollen is deposited on of Lecythis are usually less than 2 mm long. In con­ its head and back from where it is eventually trans­ trast, the styles of Bertholletia and most species of ferred to the stigmas of subsequent flowers visited. Lecythis are much longcr and obliquely oriented The third type of reward is nectar produced at the apex toward the open end of the flower. Lecythis pisonis, L. of the androecial coil by species of Bertholletia, zabucajo, and some species of Couratari (Fig. 13) Couratari, Eschweilera, and some species of Lecythis. have annular expansions toward the apex of their The pollinator reward of Cariniana is not yet known. style. The calyx-lobes of Lecythidaceae differ mostly in Some species of Lecythidaceae develop a bluish size, degree of imbrication, and thickness. For exam­ green coloration when their flowers are bruised. This ple, the calyx-lobes of Eschweilera grandiflora are coloration may also develop when the leaves are the largest and most imbricate of all species, whereas crushed and the fruits are bruised. Eschweilera laevi­ those of E. collina are much smaller and not imbri­ carpa, Lecythis pisonis, and L zabucajo are the spe­ cate. The calyx-lobes of E. coriacea are thickened cies of Lecythidaceae that possess Ihis feature at into gibbous bases. Reserve 150 I. 10 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN [VOL. 75

Anterior Distal

Distal

---=~-----'-...... -", Staminal Ring Hood Stamina! Ring Proximal A 8 posterior

Hood

Staminal Ring

Ligule

c Staminal Ring

D E

Fig. 2. AClinomorphic and zygomorphic androecia of Lecythidaceae and related terminology. A. Medial section of aclin­ omorphic~f1owered Gustaviu. B. Flower of zygomorphic-flowered Leey/his with petals removed. C. Medial section of nndroeeium of Leey/his. D. Androecium of Lecythis artifieially opened. E. of Lec-ythis with clavate filament. (Reprodueed with permission from Flom Neolropica Monograph 21(11).)

Fruits released (most species) versus drop of fruits to the ground with the seeds inside (Bertholietia ace/sa, L The fruits and seeds of Lecythidaceae at Reserve gracieana, L. patvijructa, and L prancei), (3) pres­ 1501 offer many features useful in classification. ence of a yellow, contort.xJ funicle (only in Gustavia Terminology for typical fruits and seeds is illustrated eltiplica, Fig. 17El, (4) presence of lateral fleshy arils in Figures 4 and 5, and fruit variation among species (Eschweilera spp., Figs. 40, 14F, 161) versus basal of selected genera is shown in Figures 6-8. fleshy arils (Corythophora spp. and most species of Fruit, seed, and embryo features of Lecythidaceae Lecythis, Fig. 4C), (5) presence of membranous, at Reserve 1501 that are especially useful in classi­ winglike arils at the base of the seed (only in fication are (I) indehiscence (Gustavia elliptica, Cariniana, Figs. 4G, 111), (6) presence of membra­ Lecythis gracieana, L. parvijructa, and L. prancei) nous, winglike arils around the seed (only in Coura­ versus dehiscence (most other species: note, however, tari, Figs. 4F, 13G), (7) absence of arils (Bertholietia that the opercular opening of Bertholletia excelsa is excelsa, Fig. IOF, and L. prancei, Fig. 20G), (8) pres­ smaller in diameter than the seeds), (2) retention of ence of plano-convex. cotyledons (only in Gustavia), the fruit in the tree until after the seeds hnvc been (9) presence of leaflike cotyledons (Cariniana and 1995] THE LECYTHIDACEAE OF A CENTRAL AMAZONIAN MOIST FOREST II

fodder fertile

1 em farllle --""--,"'"

stamlnal ring CorYfllopllora Gusfavla lItJJ(fJpsffJ/fJ stamlnal ring r;mosa

aclinQnlor"hlc, 'srille pollen as reward zygomorphlc, fodder pollen as reward

sterUe

1 em

1 em

Lecyfhis zabucajo L 8cyfllis corrugofo zygomorphic, fodder pollen as reward zygomorphic, lodder pollen as reward

1 em

Lecyfllis confertff/orfJ Eschw8/lero zygomorphic, reward unknown pedic8J1affJ

zygomorpl1ic, nectar as reward

Fig. 3. The .androeeiaJ types found in neotropical Lecythidaceae. Note that not all specie~ illustrated in this plate have been collected at Reserve 1501. Nonethel~s, all the androecial types represented here do occur in central Amazonia. [Reproduced with permission from L. B. Holm-Nielsen e[ a1. (eds.), Tropical/oresrs (Academic Press, 1989).]

Couratari. Figs. 50, 13H), and (10) absenee of well­ Fruit size and form provide many features for developed cotyledons (Allantoma, Bertholletia, Cory­ helping to separate species (Figs. 6-8). However. thophora, Eschweilera, and Lecythis). there is considerable fruit variation within species 12 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN [VOL. 75

proximal end c:::.:».~---pedicel scar

infracalycine zone { ~ ~..,..j,--sepal scar supracalycine zone { A ~=="",==.L~--lineof opercular distal end dehiscence

\.,---calycine ring ) \----\--line of opercular opercular opening ( dehiscence -----I;------'~-:::~--I---operc ular ring 8

~aril aril----/ c ~5l-=-funlcle D ~ E

F. G H.

Fig. 4. Fruit, seed, and seedling features of neolropicaI Lecythidaceae. Note that not all species illustrated in this plate have been collected at Reserve 1501. A. Lateral view of Lecytllis. B. View of distal end of fruit of Lecythis. C. Seed of Lecythis showing basal arH surrounding funiele. D. Seed of Eschweilera showing divided lateral ari!. E. Seed of Escllwei/era showing undivided lateral aril. F. Circumferenlially winged seed of Couratari. G. Unilaterally winged seed of Cariniana. H. Lateral germination of seed of Eschweilera tenuifo/ia. I. Apical germination of seed of Eschweilera. (Reproduced with pennission from Flora Neotropica Monograph 21(11).) 1995] THE LECYTHIDACEAE OF A CENTRAL AMAZONIAN MOIST FOREST 13

em 1 em 1 3 em

:\

A .~i\\ . h i ~..•'..I. Couratari :~~:t ;~ stellata Gustavia augusta I j~'

B c o

r/ /

11 em

F E

Bertholletia excelsa

Fig. 5. Seeds and seedlings of neolropical Lecythidaceae. Note [hat not all species illustrated in this plate have been col­ lected at Reserve 1501. (A, B, Gustavia augusta, Nee & Mori 4209. NY; C, D, Courarari stelluta, Mod & Pipoly 15454, NY; E, Bert/wlletia excelsa, Prance blasrogeny coli. no, 4, NY; P, Lecyrhis minor, Prance 2317J, NY.) A. Seed with enl::rrged funicle. B. Seedling with plano-convex cotyledons at base. C. Seed with circumferential wing. D. Seedling with two folia­ ceous cotyledons. E. Seedling without cotyledons. the undifferentiated embryo remains within the lesta. F. Seedling with­ OUl cotyledons, the undifferentiated embryo remains within the testa. (Reprodueed with permission from Flora Neotropica Monograph 21(II).) 14 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN [VOL. 75

Corylhophora rimosa 5ubsp. rimosa Cariniana decandra Couratori longipedicellolo

Couro/ari Courotori guianensis multifloru

Fig. 6. Fruit variation of selected specie.'; of C/lrinilllltl, COIytlwplwra, and Courutari (Cariniunu decandra, Oliveira 355, NY; Coryllwp!lora rimosa subsp. rimo.m, Boom el til. 8706. NY; Courarari glliQ/lensis, Tllomas el al. 5315. NY; C. longj· pedicel/ata, Cunlla et al. J178, NY; C. multiflora, without voucher. redrawn from photo Dt NY).

of Lecythidaceae (Mori & Prance, 1990a; Prance & studies, For example, we have decided that the name Mori, 1979). Eschweilera micranrha should be applied to what we previously called E, apiculata, If we assume that all TAXONOMIC PROBLEMS individuals are correctly identified, this change in names has no ecological significance, at least at the Ecological study depends on the correct identifica­ local level. Another problem is that the two species of tion of the plants and animals being studied-a task Cariniana in the area represent two groups of the that is often very difficult in tropical forests. genus, one of which, C decandra, appears to be more One of the problems in the determination of closely related to ALlantoma than to C micrantha, Lecythidaceae is the difficulty in separating species Upon further study, it may be necessary to transfer on the basis of vegetative features alone. especially the names of some species of Cariniana to Allanroma the leaves. Altempts to determine the names of trees or even to name a new genus. However, a change in without flowers or fruits should consider tree size, the name of a species of Cariniana will have no bear­ growth form, and bark features along with the leaf ing on any ecological study of this species at Reserve characteristics. Consideration of those features will 1501. markedly increase the probability of a correct deter­ A more serious problem is when further study re­ mination. Moreover, every effort should be made to veals that the individuals currently identified as a sin­ gather fertile material of Lecythidaceae in order to gle species tum out to represent more than one species. confirm determinations based on sterile collections, This has happened several times over the course of this Changes in names of species, if the individuals study, For example, individuals currently determined within the species are correctly identified, will have as Escllweilera collina and E. romeu-cardosoi were at no bearing on the conclusions reached in ecological one time considered to represent only E. collina, When 1995] THE LECYTHIDACEAE OF A CENTRAL AMAZONIAN MOIST FOREST 15

£. micron/ho E. brocleosa E. a/ropeliolalo

E: omozonicilormls

~. ~ E corioceo £. grandi/Jora £. col/ina E. oyolhiformis

~ ~ ~ E lessmannii E. wochenl1eimii

Eschweilero Ioev/carpa E truncato

Fig. 7. Fruit variation of selecled species of EscJnvei/era (Eschweilera amazurliciformis, Mor; & Silva 20232, 20580, NY; E. micrantha, Mori & Cardoso 21438. NY; E. arroper;o!ata, Mo,; 2049B, NY; E. bfCIcteosa, Mor; & Silva 19676. NY; E. collina, Pacheco et at. 203a. NY; E. cor;acea. Cunha el af. 882, NY; E. cyatll~r()rtrli.t. Mori 20282, NY; E. grandiflura [bose: Canlla et ai. 918; opex: Freitas 518. bolh NY]; E. laevicarpa, Cunha er al. 883, NY; E. tessmannU, Freitas et al. 716, NY; E, truneata, Canha el af. 886, NY; E. wacl!enlleimii, Cunha et at. 874, NY).

\' #' .. ~ .... 5cm. .\"'~,,.• -, .•.•' ,' ~.'­

Lecylhis parvlfruCla

Lecy'his rabucaja Fig. 8. Fruit variation of selected species of Lecythis (Lecythis parvifrucw, Oliveira et aI. 344, NY; L. wbucaja. Mari et al. 20585, NY). 16 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN [VOL. 75

trees of E. romeu-cardosoi began to flower, it became actinomorphic or zygomorphic, inferior or half­ apparent that two species were represented. Ecological inferior. perfect; calyx entire or of 2-6 triangular to studies in which two species are considered as one broadly ovate lobes; petals 4, 6, 8. or infrequently 12 have little value. Because we have been studying the or 18; arising from symmetrical staminal ring trees in our plot for nearly 6 years. we hope that prob­ in AUamoma, , and Gusravia, the staminal ring lems of this sort have been eliminated. However, one slightly expanded on one side in Cariniana and problem may still exist: that of Lecythis gracieana. We markedly expanded into a straplike ligule with an have not yet been able to adequately understand the expanded hood at apex in remaining genera, the hood vegetative variation of this species, and therefore it appendages with or without anthers, the pollen fertile may tum out that more than onc species is represented in all stamens, or with some stamens with fertile among the 40 individuals of it found in the 1OO-ha plot pollen and others with sterile (fodder) pollen; ovary (see discussion of the species). usually 2-, 4·, or 6-locular, with 2-116 anatropous Ecological study often depends on the accurate ovules per locule, the placentae either axile, then identification of seedlings and saplings. As discussed inserted at apex, base, or throughout lenglh of locule. earlier, there are good fruit and seed charaeters that or inserted on floor of locule. Fruits indehiscent, then can be used to identify individuals to genus and somewhat fleshy and berrylike (Grias and some species. However, seedlings are extremely difficult to species of Gustavia) or with thin, ligneous exocarp identify to species. As a result, every effort should be ( and some species of Gustavia), or dehis­ made to collect the fruits and leaves from the mother cent via circumscissile opereulum, then often large tree to aid in the identification. There is still need for and woody. Seeds winged in Cariniana and Coura­ detailed study of the morphological characteristics of tari, not winged in remaining genera~ funicle well seedlings and saplings of Lecythidaceae. developed and contorted in Gustavia, straight in remaining genera; aril absent or present, when present DESCRIPTION OF LECYTHlDACEAE lateral, basal. or rarely completely surrounding seed; SUBFAMILY LECYTHlDOlDEAE cotyledons leaflike in Cariniana, Couratari, and Couroupira, or thick, plano-convex in Gusravia, or Small, unbranched to large, much-branched emer­ absent in remaining genera. X:; 17. gent trees. Leaves alternate, exstipulate or with minute eaducous stipules, simple, camptrodromous, the mar­ USE OF KEYS gins usually entire, sometimes crenulate to serrate, very large and clustered at branch ends in Grias and The entire lead of the key should be read before some species of Guslavia, medium-sized and not clus­ deeiding what lead to follow. Our keys place the tered in remaining species. terminal, information in the leads in the same order as it occurs axillary, or cauline, of simple or spikes, pan­ in the descriptions, I.e., in the order of vegetative, flo­ iculate arrangements of racemes or spikes, or infre­ ral, fruit, and seed characters. Therefore, the most quently fascicles. Flowers usually diurnal, infre­ diagnostic feature may not be the first to appear in a quently nocturnal (only in several species of Lecyrhis). given lead.

Key to Genera l. Buds globose, the flowers small,:::;;1 cm diam. Seeds with unilateral wing...... 3. Curiniana I. Buds oblong, or if globose, the flowers >1 em diam. Seeds without unilateral wing. 2. Flowers radially symmetrical (actinomorphic). 3. Buds globose; flowers ~2.5 cm diam.; petrus obovate, rounded at apex:; anthers with poricidal dehiscence. Fruits <5 cm long; seeds ovoid, Wilh a contorted, yellow funicle. the aril not developed; cotyledons plano-convex:...... 7. GU.flavia 3. Buds oblong; flowers 0.5-0.7 cm diam.; petals oblong. with inwardly pointing hooks at apex:; anthers with lateral dehi~ccnce. Fruits ~5 cm long; seeds oblong, with a straight. non-yellow funicle, the aril developed; cotyledons not developed (embryo macropodial).. . .. [. Alfuntomu 2. Flowers bilaterally symmetrical (zygomorphic). 4. Androecial hood with extemru flap, stamens <50; ovary 3-locular. Seeds surrounded by chanaceou~ wing; cotyledons foliaeeous...... 5. CfJurufari 4. Androecial hood without ex:ternal flap, stamens 2::50; ovary not 3-locular. Seeds not surrounded by chartaccous wing; cOlyledons nol foliaceous (embryo macropodia\). 5. Calyx 2-1obed at amhcsis. Seeds with thick, hard tcsta...... 2. Bertlwlferia 1995J THE LECYTHIDACEAE OF A CENTRAL AMAZONIAN MOIST FOREST 17

5. Calyx 4-6-1obed at anlhesis. Seeds without thick, hard test.a. 6. All androeeial hood appendages with yellow anthers. Fruits eylindrie or eampanulate...... 4. C()rytlwphora 6. Androecinl hood :lppendages usually without anthers. when present anthers only on proximal appendages and never yellow. Fruits usually globose or turbinate (only E.fcllweilera amllwniciformis with cylindric fruits). 7. Androecial hood with distinct double eoil; OV:lry usually 2~, infrequenrly 4-locular, flat at apex. Seeds usually with Intend nril (only E. wnllzoniciformi.f with basal aril). Bark usually not distinctly fissured.. . . . 6. Eschweilera 7. Androecinl hood without distinct double coil; ovnry 4-loculnr. flnL at npex. Seeds with bnsnl nrit or without nril. Bark usunlly distinclly fissured 8. Lecyrlii.~

Chave para os Generos 1. Gemns florais globosns, flores pequenns, :s; 1 cm diiim. Sementes com aln unilateral. 3. CariniCl1la I. Gemns florais oblongns ou, se globosns, ns flores mniores. >1 cm diiim. Semenles sem nln unilnteral. 2. Flores com simetrin rndindn (nctinomorfns). 3. Gemas f10mis globosa.~; flores ~2.5 em diiim.; peL::tlns obovndns. ::trredondadns no opice; anterns com deiseencin poricidn. Frulos <5 cm eompr.; sementes ov6ides, com funiculo contorcido, dcsenvolvido, nrilo nusenle; cotiledones plano-convexos , 7. Gustav!u 3. Gemns florais oblongns; f10res 0.5-0,7 cm di5.m; petnlns oblongns. ngudns no apice; nnlerns com deiseencin Internl. Frutos 2'5 em compr.; sementes oblongns, funiculo nno eontorcido, pouco dcsenvolvido, 0010 presente; cotiledones nao desenvolvidos...... I. Ai/anton/a 2. Flores com simetria bilntcrnl (zigomorfns). 4. Cnpuz cstnminal com aba externa, estnmes <50; ovario 3-loculnr. Scmentes com nln no redor de lodn n semenlc; cotiledones folioceos...... 5. CourCJtari 4. Cnpuz estnminal sem ab::t exlernn, estnmes ~50; ovorio nao 3-10culnr. Sementes sem nla; coliJedones n1io desenvolvidos. 5. Colice com 2 lncinios nn

DESCRIFTIONS OF GENERA AND SPECIES 1. Allantoma lineata (Martius ex Berg) Miers, Trans. Linn. Soc. London 30(2): 297, t. 65, figs. 4, 5.1874; 1. Allantoma Dueke, Arch. Jard. Bot. Rio de Janeiro 4: 155, t. 16. Canopy trees, unbuuressed. Bark nearly smooth. 1925; Prance in Prance & Mori, FI. Neotrop. Leaves eucamptrodromous, with finely percurrent Monogr. 21: 209-218, figs, 3N. 9, IIS-U. 161-K, tertiary veins. Inflorescences terminal or subterminal, 62-65. 1979; Couratari lineata Martius ex Berg, FI. racemes or once-branched paniculate arrangement of Bras. 14(1): 508, t. 7, 77. 1858. Fig. 9 racemes. Buds oblong. Flowers diurnal, actinomor­ phic; calyx rimlike, sometimes with 5(-6) scarcely Understory to canopy trees, to ca. 25 m tall, unbut­ developed lobes; petals 5, oblong, with inwardly tressed. Bark dark brown, nearly smooth, with scat­ pointing hooks at apex; androecium oblong, without tered vertical lines of lenticels, the ouler bark ca. hood, the stamens inserted at apex and over inside of I mm thick, the inner bark ca. 10 mm thick, white staminal ring, the anthers small, with lateral dehis­ with slight hint of pink, with distinct red boundary cent; ovary 4(-5)-locular, the ovules inserted along between inner and outer barks. Leaves: petioles length of septum. Fruits cylindric. Seeds with basal 12-/8 mm long; blades oblong. 18-19 x 8-9 cm. aril; embryo without cotyledons. glabrous, with 17-25 pairs of secondary veins, the 18 MEMOIRS OF THE NEW YORK BOTANICAL GAROEN [VOL. 75

lem o ~ C

I lem ! i I Ilem G I, E F H/l}7~

Fig. 9. AlIll1ltOlllll !ineala (A-D. Ducke 14/4, NY; E-G, Prance et aJ. 11498, NY). A. Branch with . B. Longitudinal section of flower. C. Cross section of ovary. D. Rower. E. Seed. F. Operculum. G. Fruit (Reproduced with permission from Aora. Neotropica Monograph 21(1).) 1995] THE LECYTHIDACEAE OF A CENTRAL AMAZONIAN MOIST FOREST 19 secondary veins running to near margin, the ter­ without wings or aril, the testa very hard; embryo tiary veins finely percurrent, forming 135° angle with without cOlyledons. midrib; base truncate, infrequently rounded; apex long acuminate; margins entire to finely serrulate. I. Bertholleli. excels. Humboldt & Bonpl.nd, PI. Inflorescences termin

~ \Jr--_-~~_------< ~. ---- ./,

',·f

Fig. to. Bert'wlletia exce.l.m (A-D, Mor; et af. 17503, NY; E, F, Prance et at. 16599, NY; G, Prance blastogeny coil. no. 4, NY). A. Branch with inflorescence. Note how lhc petals are tightly appressed 10 the androecium. B. Medial section of androecium. Note how the hood appendages are swept inward but do not form a complete coil. C. Cross section of ovary. D. Ovary and calyx. Note the 2-parted calyx. sessile ovary, and long, obliquely oriented style. E. Fruit. Note small opercu­ lar opening. F. Seed. G. Seedling. (Reproduced with permisJ::ion from Flora Neolropica Monograph 21 (1).) 1995] THE LECYTHIDACEAE OF A CENTRAL AMAZONIAN MOiST fOREST 21

bark is used to make a dye for staining fish nets and 3. Cariniana may possess medicinal propertIes as it is sometimes sold in the Manaus market in the medicinal sec­ Emergent trees, usually without buttresses. Bark tion. More details about the economic botany of the usually fissured, the inner bark red. Leaves eucamp­ Brazil nut can be found in Man and Prance (199Gb). trodromous, with finely percurrent or reticulate ter­ Comments. This species is characterized by its tiary veins. Inflorescences terminal or in ax.i1s of (1) cylindric trunk and deeply fissured bark; (2) abax­ upper leaves, racemes or spikes or paniculate ar­ ial leaf surface with microscopic, cuticular papillae; rangements of racemes or spikes. Flowers diurnal. (3) calyx with only 2 lobes; and (4) globose fruits that actinomorphic or zygomorphic. small. <10 mm diam; fall to the ground with the seeds inside at maturity. sepals 6; petals 6; androecium without or with Berrholletia excelsa is pollinated by large-bodied slightly developed hood, the anthers with lateral bees belonging to the genera Bombus, Centris, dehiscence; ovary 3-locuJar, the ovules inserted at Eulaema. and Xylocopa. and its seeds are dispersed base of septum. Fruits cylindric or turbinate, the peri­ by rodents, especially agoutis (Dasyprocta spp.) carp woody, thick to very thick. Sceds with unilateral (Mori & Prance, 1990b). wing; embryo with 2 foliaceous cotyledons.

Key to Species of Cariniana

I. Young twigs glabrous. Leaves Wilh entire margins; secondary veint:i without domalia in axils, the tertiary veins finely percurrent; petioles >15 mm long. Flowers actinomorphic; pelals fleshy. cucullate; stamens 10. 5 short and 5 long. Fruits cylindric, the pericarp <5 mm thick. I. C. decandra 1. Young twigs pubescent. Leaves with serrulale margins; secondary veins with domatia in axils, the tertiary veins reticulate; petioles <5 mm long. Flowers zygomorphic; pelals nOl fleshy or cucullate; >10 t:itamens, all more or leI'S the same lenglh. Fruils turbinate. the pericarp > 10 mm thick. . . 2. C. micronlllll Chave para as Especies de Cariniana I. Ramos jovens glabros. Folhas com margent:i inteira..., as nervuras secund5rias scm cavidades pequenas nas aXilat:i (domacia), at:i nervur15 mm compr. Flores aClinomorf5 mm compr. Flores zigomorfas; petalas nno carnosas, planas; estames > I0, LOdos mais ou menos do mesmo comprimento. Frutos turbinados. 0 pericarpo > 10 mm de espeSSura...... 2. C. micranll/Cl

1. Cariniana decandra Ducke, Arch. lard. Bot Rio on longer stamens, the anthers nodding inward, yel­ de Janeiro 4: 153. t 16. 1925; Prance in Prance & low; glabrous; style erect, ca. 0.6 mm Mori, FI. Neolrop. Monogr. 21: 236-237, tigs. 66B, long. Fruils cylindric, 8-1\ x 3-4 em, the exocarp rcd­ 72. 1979. Fig. 6 dish brown. often with longitudinal ribs. Habitat. Terrafirme. The IOO-ha plot contains 40 Emergent tree. to 40 m tall. unbuttrcssed. Bark individuals of this specics ~IO em dbh. slightly fissured. the inner bark red. Leaves: petioles Distribution and representative specimens. Cen­ 15-21 mm long; bladcs elliptic to narrowly elliptic. tral and southwestcrn Amazonia. Oliveira et al. 294 10-25 x 4.5-10 em, glabrous, with 14-17 pairs of sec­ (Fr) (tree 1785); Oliveira et al. 355 (Fr) (tree 1785). ondary veins. without domatia in axils, the tertiary Phenology. Flowers from Dec co Feb throughout veins finely percurrem, forming 13Y' angle with mid­ its range. Flowering trees have not yet been observed rib; base obtuse; apex acuminate; margins entire. In­ at Km 41. florescences with pedicels barely discernible. Flowers Common name. Tauarf. actinomorphic. ca. 5 mm diam.; calyx fused at base Comments. This species is characterized by its into rim 0.6-0.7 mm high, the tobes broadly triangu­ (I) shallowly fissured outer bark with red inner bark. lar; petals glabrous, yellowish white; androecium with (2) percurrent tertiary veins, (3) small flowers, and 10 stamens, 5 long and 5 short, these inserted on mar­ (4) cylindric fruits that are often longitudinally gin of staminal ring, the filaments flattened, especially ribbed. 22 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN [VOL. 15

2. Cariniana micrantha Ducke, Arch. lard. Bot. Rio seeds are heavily preyed upon by capuchin monkeys de Janeiro 5: 175. 1930; Prance in Prnnce & Mori, (Cebus apeLLa). The monkeys knock the nearly ma­ FI. Neolrop. Monogr. 21: 237-238, figs. 17A-C, ture fruits open by banging them against branches. 30, 66A, 73. 1979. Fig. 11 The banging can be heard up to 400 m away from the tree in which the capuchins are harvesting the seeds Emergent trees, to 52 m tall. unbuttressed. Bark (Peres, 1991). with well-developed vertical fissures, the ridges heM Common name. Tauarl. tween the fissures 3-4 em wide, the outer bark 5-10 Uses. The seeds are edible, but they are small and mm thick, the inner bark 10--12 mm thick, distinctly difficult to collect because they are dispersed by the laminated, red. Leaves: petioles 2-5 mm long; blades wind. The empty fruits are used to make toys for elliptic, 9-13 x 4-5 em, pubescent on midrib, espe­ children. cially when young. with 12-15 pairs of secondary Comments. This is one of the largest Lecythida­ veins, with domalia in axils t these lined by hyaline ceae found at Km 41. Even though Prance (Prancc & hairs, the tertiary veins reticulate; base obtuse; apex Mori, 1979) indicated that this species does not have acuminate; margins serrulate, the teeth with scars of domatia, all individuals we have studied at Km 41 caducous hairs. Inflorescences with pedicels short but have them, and they are present in collections at NY discernible. Flowcrs 8-10 mm diam.; caly,,-Iobes determined by Prance as C. micrantha. The margins narrowly ovate to narrowly triangular, 2 x I mm, of the domatia arc fringcd by hyaline hairs. pubescent, green; petals membranous, oblong, 6 x 3 This species is characterized by (I) fissurcd, rela­ mm, pubesccnt. the margins fimbriate, whitc; androe­ tively thick, laminated outer bark; (2) red inner bark; cium with ca. 60 stamens insertcd around staminal (3) serrulate leaf margins~ (4) domatia in the a"i1s of ring and all ovcr interior of hood, the filaments 0.2 the secondary veins; (5) smllll, white flowcrs; and mm long, the anthers versatile; hypanthium globose, (6) turbinate, thick-walled fruils. pubescent; ovary with 4-6 ovules in each loeule, the style 0.7 mm long, thick as opening in staminal ring at base, apicu late at ape", thc lower pan densely 4, Corythophora pubescent. Fruits turbinate, 7-9 x 5.5-6 em, the peri­ Canopy trces, unbuttressed. Bark deeply fissured carp 11-20 mm thick, rcddish to dark brown. or not fissured. Leaves glabrous, brochidodromous, Habitat. Terrafirme. Thc IOO-hn plot contains 29 the tcrtiary vcins reticulate. Inflorescences terminal individuals of this species ?l 0 cm dbh. or in a"ils of upper leaves, simple raccmes or panicu­ Distribution and representative specimens. Cen­ late arrangement of racemes with 2-3 orders of tral and southwestern Amazonia. Freitas eI al. 560 branching. Flowcrs diurnal, zygomorphic; sepals 6; (tl) (tree 6240); Mari et 01. 20246 (Fr) (tree 6834). petals 6; androecium with well-developed bur flat Phenology. Flowcrs mostly from the latter part of hood, the appendages with anthcrs; ovary 2-locular, the dry season into thc early wet season (Oct to Feb) the ovules inserted at base of septum or on floor of and drops seed from the end of the wet season into the Jocule. Fruits cylindric or eampanulate. Seeds with dry season (JulIo Oct) (Prance & Mori, 1979). The basal ari!; embryo without cotyledons.

Key to Species of COlythophora I. Bark not fissured. the outer bark thinner than inner bark, the inner bark pink to red. Leaves with slender petioles, usually ~1.5 mm thick; margins serrulatc 10 less frequently entire. Inflorescences usually <10 cm wide. Flowers with pink to pnrple petals. Fruits cylindric. usually 55 cm diam...... 1. C. a/tel 1. Bark deeply fisl>ured, the ouler bark equal 10 or thicker than inner bMk, the inner bark dark red. Leaves wilh robust petioles. usually >1.5 mm [hick; margins eniire to less frequently selTulate. Inflorescences nsually ~IO cm wide. Flowerl> with green petals. Fruits campanulate, usually >5 cm diam 2. C. rimosa subsp. rimosa

Chave para as Espeeies de Coryrhophora

I. Casca extema nao fissuroda, a casca extema meno~ grossa que a casca iniema em espessura. a casca intema rosa a vermelha. Folhas com pecfolos delgados. geralmente 51,5 mm difun.; margens serrilhada.<; a menos freqi.ientemenlc intciras. Inflorescencias gerolmente <10 em de larguro. Flore!' com petalas rosa.<; aroxas. Frutos cilfndricos, gerolmcnte :55 em dHim...... 1. C. alta 1. Cn.<;ca exlcrna profundamenic fissurada. a casca extema equal ou mais grossa que a ca.<;ca interna em espessuro, a cosca inicma vermelha escura. Folha.<; com pecfolos robustos. gcrolmente > 1.5 mm dinm.; Illargen~ inteira.5 cm di5.m...... 2. C. rill/o.fU subsp. rimosa 1995] THE LECYTHlDACEAE OF A CENTRAL AMAZONIAN MOIST FOREST 23

Fig.H. Carin;ana micranlha (A-E, Mori er at. 20191, NY; H, I, Mod el at. 20675, NY). A. Branch with inflorescences. B. Close-up of inflorescence. C. Lateral view of flower. D. Top view of flower. E. Lateral view of androecium. F. Inferior ovary with all but sepals removed. G. Cross seclion of trilocular OVDry. H. Base of fruit and operculum. I. UnilaternJly winged seeds. 24 MEMOIRS OFTHE NEW YORK BOTANICAL GARDEN [VOL. 75

1. Corythophora alta R. Knuth in Engler, Pflanzenr. 15-25 x 6-10.5 cm, with 10-16 pairs of secondary IV, 219a: 50-51, fig. 10.1939; Mari & Prance, FI. veins; base obtuse to rounded; apex acuminate; mar­ Neatrap. Managr. 21(11): 109-111, figs. 14C, 40, gins entire to serrulate. Inflorescences terminal or in 4IA-B,43. 1990. Fig. 12 axils of uppermost leaves, paniculate arrangements of spikes with 2 orders of branching, usually> 10 em Canopy trees, to 35 m tall, unbuttressed. Bark with wide. Flowers ca. 2.5 em diam.; calyx of 6 broadly distinct, irregular depressions but not fissured, the ovate or triangular lobes, not imbricate at bases, outer bark ca. 1 mm thick, the inner bark ca. 5 rom rounded or acute at apices; petals 6, 5-l3 x 4-8 mm, thiek, pink to red. Leaves: petioles slender, <1.5 rom green; hood of androecium flat, only slightly ex­ diam., io-I8 mm long; blades elliptic, 10-14 x 5-7 panded dorsiventrally, white to yellow, the appen­ em, with 11-14 pairs of secondary veins; base obtuse dages with yellow anthers, the staminal ring with to rounded; apex acuminate; margins serrulate to 50-70 stamens; ovary 2-locular, with 4-7 ovules arrange~ entire. Inflorescences terminal, panicu)ate attached at base of locule, the style not differentiated ment of spikes with 2 orders of branching, usually from summit of ovary. Fruits campanulate, 8-12 x <10 em wide. Flowers 2-3 em diam.; ealyx of 6, 8-11 em. Seeds elongate. 3.5-4 x 1.5-1.7 em. broadly triangular lobes, not imbricate at bases, acute Habitat. Terrafirme. The 100-ha plot contains 208 at apices; petals 6, 8.5-14 x 8-11 mm, red to dark individuals of this species ~l 0 cm dbh. purple; hood of androecium flat, only slightly Distribution and representative specimens. Cen­ expanded dorsiventrally, red to dark purple, the tral Amazonian Brazil. Mori et at. 20034 (fr) (tree appendages with yellow anthers, the ligule with clus­ 2397); Mori el af. 20499 (fl) (tree 558). ter of anther-bearing appendages separated from both Phenology. Flowers most profusely from Mar to the hood appendages and the stamens of the staminal Aug (mid wet to early dry season) and drops seed ring, the staminal ring with 40-90 stamens; ovary 2­ from Aug to Dec (dry to early wet season). locular, with 5-8 ovules inserted on base of locule, Common name. Castanha-jacar6. the style not differentiated from summit of ovary. Comments. This species is characterized by its Fruits cylindric, 7-9.5 x 4--6 cm. Seeds elongate, tri­ (I) thick, fissured outer bark, (2) red inner bark, (3) angular in cross section, 3-4.5 x 1-1.3 cm. green petals, and (4) campanulate fruits. Habitat. Terrafirme. The IOO-ha plot contains 192 Subspecies rubra Mori is found in French Guiana individuals of this species;;:: I0 cm dbh. and Amapa. The two subspecies differ in their petal Distribution and representative specimens. Cen­ color. Subspecies rimosa has green petals; subsp. tral and eastern Amazonian Brazil. Mori et at. 20295 rubra has red petals. The pollinators of subsp. rubra (fr) (tree 7732); van Roosmafen L-32 (fl) (tree 18i8). are robust species of trigonid bees (Mori & Boeke, Phenology. Flowers mainly from Dec to May (wet 1987). The tlowers of subsp. rimosa at first were season) and drops seeds mainly from lui to Sep (dry thought to be nocturnal because of their green col­ season) (Prance & Mari, 1979). oration. However, lhey are now known to be open Common name. Ripeiro vermelho. during the day, but no pollinators have yet been Comments. This species is characterized by its observed visiling them (Mori & Prance, I990a). (1) scalloped bark, (2) relatively thin outer bark and pink to red inner bark, (3) reddish flowers, (4) a group of anther-bearing appendages on the ligule, which is 5. Couratari separated from the staminal ring and from the anther­ bearing haad appendages (Fig. 12C), and (5) cylin­ Emergent trees, the adults usually buttressed. Bark dric fruits. with very fine vertical cracks. never deeply fissured. Leaves brochidodromous. the tertiary veins reticulate 2. Corythophora rimosa W. Rodrigues subsp. or percurrent. Flowers diurnal. zygomorphic; sepals rimosa, Acta Amazonica 4(1): 5-6, fig. I. 1974; 6; petals 6; androecial hood with well-developed Mori & Prance, FI. Neatrap. Managr. 21(11): 111­ inward coil and external tlap. the tlap externally echi­ i3, figs. 5E, 14D, 15B, 40, 41C, 44. 1990. Fig. 6 nate or smooth, the appendages of hood without anthers, the anthers with lateral dehiscence; ovary 3­ Canopy trees, to 30 m tall, unbuttressed. Bark very locular, the ovules inserted on septum toward base of deeply fissured, the outer bark as thick or thicker than locule. Fruits cylindric or campanulate. Seeds com­ inner bark, the inner bark red. Leaves: petioles robust, pletely surrounded by wing; embryo with foliaceous usually >1.5 mm thick, 7-19 mm long; blades elliptic, cotyledons. 1995] THE LECYTHIDACEAE OF A CENTRAL AMAZONIAN MOIST FOREST 25

F 4~A'. " ';4(\;')}'~'

10m

1em

Fig. 12. Corythop!wf(l alta (A. Mod 20266. NY; 8-0. Cun"a et at. 925. NY; E-G, Mori 20290A. NY). A. Branch with inflorescence. B. Top view of flower. C. Medial seclion of androecial hood. D. Cross section of ovary. E. Inferior ovary. F. Base of fruit and operculum. G. Laternl views of seeds. Note arils at top of seeds. 26 MEMOIRS OFTHE NEW YORK BOTANICAL GARDEN [VOL. 75

Key to Species of Couratari

I. Leaves finely crenulale for entire length. the blade oflen with longitudinal .~Iriations. Flowers Wilh leaves present, the androecial hood yellow. with echinate outer surface...... 4. C. sJell(lru I. Leaves entire or crenulate only toward apex, the bJaue without longiluJinal striations. Flowers without leaves, the androecial hood pink to purple, with smoolh outer surface. 2. Petioles long and slender, > 1/4 length of blade, lhe blades < 10 em long, glabrous. Fruits