Endlicheria (Lauraceae)

Organization for Flora Neotropica

Endlicheria (Lauraceae)Author(s): André S. ChanderbaliSource: Flora Neotropica, Vol. 91, Endlicheria (Lauraceae) (Aug. 24, 2004), pp. 1-141Published by: New York Botanical Garden Press on behalf of Organization for Flora NeotropicaStable URL: http://www.jstor.org/stable/4393929 .Accessed: 09/03/2011 09:10

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INTRODUCTION Within this book, Endlicheria and Rhodostemonodaphne are treated as separate mono-

graphs. Traditionally, Endlicheria was placed nearAniba and other genera of the Lauraceae with two-locellate anthers. Rhodostemonodaphne, with four-locellate anthers, was recently segregated from Nectandra. New information, however, suggests that Endlicheria and Rhodostemonodaphne form a monophyletic group of approximately 100 species. Never- theless, the phylogenetic relationships among these species are still unclear and until there is more conclusive evidence we feel this complex group of species should remain within two genera.

ANDRt S. CHANDERBALI AND

SANTIAGO MADRINAN

FLORA NEOTROPICA MONOGRAPH 91

ENDLICHERIA (LAURACEAE)

ANDRE S. CHANDERBALI

; P _ P CANCER

FLORAL NEOTROPICA

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Flora neotropica. Monograph no. 1 New York: Published for Organization for Flora Neotropica by The New York Botanical Garden, 1968-

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Irregular. Each issue has distinctive title. Separately catalogued and classified in LC before monograph no. 40. ISSN 0071-5794 = Flora neotropica.

1. Botany - Latin America- Classification - Collected works. 2. Botany- Tropics - Classification - Collected works. 3. Botany Classification - Collected works. I. Organization for Flora Neotropica. II. New York Botanical Garden.

QK205.F58 581.98'012-dcl9 85-647083

Library of Congress [8508] ISBN 0-89327-454-2

03 04 05 06 07 08 09 10/98765 432 1

ENDLICHERIA (LAURACEAE) ANDRE S. CHANDERBALI

CONTENTS Abstract/Resumen ................................2 Introduction ...............................2 Historical Survey ................................2 Morphology and Anatomy ....3...........................3

Vegetative Morphology ....3...........................3 Inflorescences ...4............................4 Flowers ...............................4 Fruits ...............................5 Wood and Bark Anatomy ...............................5 Floral Anatomy ...5............................5

Embryology ...............................6 Karyology ................................7 Palynology ................................7 Relationships ...7............................7

Systematic Position ....7...........................7 Generic Delimitation and Species Groups ...............................7

Species Concepts ............................... 14 Distribution and Biogeography ............................... 14 Systematic Treatment ............................... 16

Generic Description ............................... 16 Key to the Species ............................... 16 Species Descriptions ............................... 22

Doubtful Names and Excluded Taxa ............................... 123 Acknowledgments ............................... 125 Literature Cited ............................... 125 Numerical List of Taxa ............................... 126 List of Exsiccatae ............................... 127 Index of Local Names ............................... 137 Index of Scientific Names ............................... 138

2 FLORA NEOTROPICA

ABSTRACT Chanderbali, A. S. (Department of Botany, University of Florida, Gainesville, FL

3261 1, USA). Endlicheria (Lauraceae). Flora Neotropica Monograph 91. 1-141. 2004.-A taxonomic monograph of Endlicheria, a South America-centered genus of the Ocotea complex, is presented; and with the benefit of molecular data showing that its members are nested within Rhodostemonodaphne and Ocotea, its delimitation is assessed in morphological detail. Sixty species of Endlicheria are recognized, of which 16 are newly described. Two species are transferred to Rhodostemonodaphne. Nine infrageneric species groups are informally recognized in Endlicheria. Of these, the Endlicheria punctulata species group is closer to species of the Ocotea cernua species group than to its congeners, in both molecular and morphological aspects. Representatives of each of the eight remaining species groups are united with a diverse representation of Rhodostemonodaphne in a well-supported but unresolved clade. The significance of this Endlicheria-Rhodoste- monodaphne alliance is explored from a morphological perspective. It is suggested that the two-locellate anthers that distinguish Endlicheria from Rhodostemonodaphne evolved repeatedly.

RESUMEN Chanderbali, A. S. (Department of Botany, University of Florida, Gainesville, FL

32611, USA). Endlicheria (Lauraceae). Flora Neotropica Monograph 91. 1-141. 2004.- Se presenta una monograffa taxon6mica de Endlicheria, genero perteneciente al complejo de Ocotea con centro de diversidad en America del Sur; y con la ventaja de datos moleculares monstrando que sus miembros estain incluidos dentro de Rhodostemonodaphne y Ocotea, su delimitaci6n generica es evaluada morfologicamente al detalle. Se reconocen 60 especies en Endlicheria, de las cuales 16 se describen como nuevas. Dos especies son transferidas a Rhodostemonodaphne. Nueve grupos infragenericos de especies son infor- malmente reconocidos en Endlicheria. De estos, el grupo de especies de Endlicheria punctulata esta mas cercanamente emparentada al grupo de especies de Ocotea cernua que a las especies congenericas, tanto en los aspectos moleculares como en los morfologicos. Los representantes de los ocho grupos de especies restantes, junto con una representaci6n diversa de Rhodostemonodaphne forman un clado con buen soporte pero sin resolucion. La importancia de la alianza Endlicheria-Rhodostemonodaphne es evaluada desde una perspectiva morfol6gica. Es sugerido que las anteras con dos 16culos que distinguen En- dlicheria de Rhodostemonodaphne podrian haber evolucionado repetidamente.

INTRODUCTION The Lauraceae form a large pantropical family of

about 50 genera and approximately 2500-3000 species of trees, rarely shrubs, and one genus of parasitic vines, Cassytha L. In Endlicheria Nees all species with non-involucrate inflorescences, a dioecious breeding system, and two-locellate anthers are placed. These characters circumscribe a neotropical taxon that differs from sympatric Ocotea Aubl. pro parte and Rhodostemonodaphne Rohwer & Kubitzki mainly in having two rather than four-locellate anthers, a distinction of questionable generic value in Lauraceae (e.g., van der Werff, 1984; Burger, 1988; Rohwer et al., 1991). A recent molecular study of Lauraceae showed that Endlicheria is at least di- phyletic, with more species allied to Rhodostemono-

daphne than to Ocotea (Chanderbali et al., 2001). Here I update the taxonomy of Endlicheria in mon- ographic custom and assess its circumscription in morphological detail while I present additional molecular data relevant to its systematic position.

This treatment is based on almost 2500 collections. In addition to the substantial material available at MO, many of recent collections with several duplicates not yet distributed to other herbaria, loans were received from A, AAU, B, C, COL, E, ECON, F, G, GB, GH, GOET, HBG, HUA, INPA, K, L, LE, MG, NY, P, QAME, R, S, U, UC, US, and VEN.

HISTORICAL SURVEY Nees (1833) described Endlicheria with two spe-

cies. Cryptocarya hirsuta Schott (= Endlicheria pan-

MORPHOLOGY AND ANATOMY 3

iculata) was transferred to Endlicheria, and E. sericea was newly described. In 1836, Nees added three more species and changed the generic epithet to Goeppertia Nees because of the earlier Endlichera Presl in Ru- biaceae. Also in 1836. Endlicheria appeared in the list of Lauraceae genera published in the second edi- tion of John Lindley's Natural System of Botany. Nees supplied this list to Lindley in 1835 and, already aware of Presl's Endlichera, suggested that Schauera Nees be used instead of Endlicheria if Presl's name was to be retained for the rubiaceous genus. Koster- mans (1936) pointed out unsatisfactory consequences with use of either Goeppertia or Schauera in Laura- ceae, and his proposal that Endlicheria be conserved in Lauraceae over Endlichera in Rubiaceae was accepted at the VI Botanical Congress. The generic epithet honors Stephan Ladislaus Endlicher (1804- 1849).

Meissner (1864) treated Endlicheria (as Goepper- tia) in his treatment of Lauraceae for De Candolle's Prodromus. Meissner accepted 14 species in Goep- pertia, four only tentatively, and transferred G. multiflora Miq. to Ampelodaphne Meisn., a newly described genus with two other species. Mez (1889), in his treatment of American Lauraceae, excluded the four species Meissner tentatively described in Goep- pertia and included Ampelodaphne in Endlicheria. Mez apparently used the name Endlicheria instead of Goeppertia because he believed Presl's Endlichera was published without description, i.e., a nomen nudum. Mez recognized 23 species and for the first time provided an infrageneric scheme. Five subgenera were recognized. Subgenera Hemiajouea Mez, An- osphaeria Mez, and Ocoteopsis Mez, were monotypic, accommodating E. paradoxa, E. impressa, and E. anomala, respectively. Subgenus Ampelodaphne (Meisn.) Mez accommodated eight species and subgen. Euendlicheria Mez the remaining 12 species. Kostermans (1937) followed Mez's concept of Endli- cheria in the only subsequent monograph. He included monotypic Huberodaphne Ducke in Endli- cheria and recognized 39 species. In his generic subdivision Kostermans (1937) maintained Mez's monotypic subgen. Ocoteopsis and placed the remaining species in subgen. Euendlicheria Mez. The latter was divided into sections Microlocellata Kos- term. and Macrolocellata Kosterm. on the basis of size and position of locelli in the outer staminal whorls.

Since Kostermans's treatment, several authors have described new species in Endlicheria. Most no- tably, C. K. Allen, in a series of papers published in The New York Botanical Garden's Botany of the Gui-

ana Highlands series, added nine species. Prior to this treatment 68 names were validly published in Endli- cheria.

MORPHOLOGY AND ANATOMY VEGETATIVE MORPHOLOGY

Species of Endlicheria are mostly medium-sized trees less than 25 meters in height, but they can reach up to 40 meters; a few are shrubs. Of the shrubby species, at least E. reflectens and E. vinotincta are consistently reported to be scandent with long slender stems that sprawl across adjacent vegetation. Growth is apparently rhythmic with leaves either produced along the entire length of flushes or, as typical of the Ampelodaphne species group, clustered in close spirals at the tips of each new flush.

Leaves are always simple, entire, and petiolate. Only in Endlicheria dictifarinosa, E. bracteata, E. cocuirey, E. melinonii, and E. verticillata are leaves subsessile. Shape varies from ovate to obovate and is usually constant within species, but in a few (e.g., E. ruforamula and E. paniculata) these extremes are linked by intermediates. Laminae are usually plane, but are strongly bullate in E. bullata and occasionally moderately so in E. glomerata. Leaf apices are usually acuminate, but are caudate in E. longicaudata and E. punctulata, and bluntly rounded to retuse in E. rubra. Leaf bases are most often acute, but vary from attenuate to abruptly contracted into the petiole (i.e., subcordate). Margins are usually curved downward at ca. 900 (recurved), at least in the lower half of the lamina, but they can be plane or recurved throughout.

Venation is mostly pinnate with secondaries alter- nately arranged and evenly spaced along the midrib. Only E. acuminata, E. bracteolata, E. gracilis, and E. krukovii have triplinerved leaves, where one pair of (sub-)opposite secondaries emerges shortly above the base of the midrib, and others (if present) emerge from the midrib around or beyond midlamina. In E. bracteolata and E. sericea quintuplinerved leaves appear when two pairs of (sub-)opposite basal secondaries emerge in the lower lamina. Secondaries range in number from 2 in triplinerved species to over 30 per side in E. bullata, but usually number 5 to 7 pairs. In most species, the secondaries follow arcuate courses and distal pairs form weak loop connections near the leaf apex. Secondaries that form strong loop connections throughout the lamina (i.e., brochidodromous secondaries) are restricted to E. arunciflora, E. bullata, E. dysodantha, and E. longicaudata. Oc- casionally, basalmost secondaries are asymmetric, with one or both merging with the margin near the leaf base or emerging from the junction of midrib and

4 FLORA NEOTROPICA

leaf base. This venation pattern is typical of E. metallica, E. chrysovelutina, and E. klugii, and appears occasionally in E. canescens and E. ruforamula. Ter- tiaries are typically roughly horizontal and parallel, proceeding undivided or once-forked between adjacent secondaries or between midrib and secondaries, but rarely, e.g., in E. reflectens, they reticulate throughout the intercosta. Midribs, secondaries, and tertiaries are usually raised on both sides of the lamina, at least in dried specimens, but one, two, or all vein orders may be immersed above. In E. punctulata all are immersed below.

Variation in color, density, orientation, size, and shape of hairs provides a great array of characters for species identification. Apart from Endlicheria pyriformis, the vegetative surfaces of which would be glabrous were it not for a marginal fringe of hairs on the youngest leaves (in bud), species have conspicuous, and often characteristic, indumentum. Hairs can be straight, crooked, crisped, or tightly crinkled. The latter type of hair is almost restricted to the inner surface of tepals and rarely achieves the density typical of Nectandra Rol. ex Rottb. where, as here, they are termed papillose (Rohwer, 1993a). All other indument terms are used as defined by Steam (1992). Hair orientation, whether erect or closely appressed to the surface, appears to be stable within species, but both extremes are found in the circumscription of E. acuminata, E. anomala, and E. paniculata adopted here. Usually only one type of hair is produced on a surface, but in E. duotincta and E. williamsii a dense tomentose cover that obscures the lower leaf surface is interspersed with longer erect hairs, and in E. dysodantha and E. tomentosa dense tufts of longer more stiffly erect hairs than found elsewhere on the lower leaf surface occupy the axils of secondary veins. In the E. sericea and E. metallica species groups, a dense golden or silvery sericeous indument persists from buds to maturity.

INFLORESCENCES

Inflorescences of Endlicheria are usually panicles with terminal dichasia, i.e., thyrso-paniculate in the sense of Weberling (1989). However, in E. citriodora, E. acuminata, and E. xerampela terminal cymes collapse into pseudo-umbellate fascicles; and in several, terminal botryoids (sensu Weberling, 1989), or deter- minate racemes, are formed via reduction of subterminal lateral cymes to single flowers. The axes of botryoids are occasionally greatly reduced, and flowers are grouped in dense clusters along secondary inflorescence branches (e.g., E. glomerata). Inflorescences are either borne in the axils of new foliage leaves or,

as typical of the Ampelodaphne species group, in the axils of scalelike bracts (cataphylls) that precede leaf crops. Often, the axis of the cataphyll-bearing shoot is greatly reduced and inflorescences are arranged in subterminal clusters above the latest leaf crop. Pistil- late inflorescences are usually shorter, less branched, and fewer flowered than the staminate inflorescences, but can also be equal in these regards.

Inflorescences typically have the indument of twigs and other vegetative surfaces. However, in E. arunciflora, E. arachnocome, and E. verticillata, inflorescences (and outer surface of flowers) are subglabrous although the plants are otherwise densely pubescent. In a few species, inflorescences are more densely pubescent than vegetative organs (e.g., E. formosa and E. metallica).

FLOWERS

Flowers of Endlicheria are organized in the manner typical of Lauraceae, i.e., all are trimerous and consist of two whorls of (sub-)equal tepals and three whorls of stamens surrounding a simple pistil. As is also typical of Lauraceae, stamens are arranged in al- temating cycles, such that those of outermost (whorl I) and innermost whorls (whorl III) stand opposite each other, with those of the intermediate whorl II placed altemately. Rarely, a whorl of staminodes (whorl IV) is produced opposite whorl II inside the innermost staminal whorl (whorl III). A pair of nectariferous glands is usually attached to the base of each whorl III stamen. Locelli are introrse or introrse- latrorse in whorls I and II and extrorse-latrorse in whorl III. With dehiscence, valves that initially covered locelli are raised above anthers with pollen grains attached to their inner surfaces. The pollination of En- dlicheria flowers has not been studied in detail, but as elsewhere in Lauraceae (Kurz, 1982; Kubitzki & Kurz, 1984) it is most likely that short-tongued insects, such as bees and flies, are the pollinating agents. Not surprisingly, the arrangement of stamens and locelli places valves of all three staminal whorls in close proximity over the nectariferous glands that these vis- iting insects seek.

Within the limits of this basic organization, the range of floral variation in Endlicheria is such that a typical flower structure cannot be envisioned. Flower shape ranges from rotate with horizontally spreading tepals, through cyathiform or infundibuliform with erect or ascending tepals, to globose or obconical where tepals open only enough to provide a terminal pore (terminology after Steam, 1992). The androecium is also variable. Stamens of the outer androecial whorls (whorls I and II) are (sub-)equal. Here anthers

MORPHOLOGY AND ANATOMY 5

are two-locellate, and, in most species, the filaments are appreciably narrower than anthers; i.e., stamens are stipitate. In several species however, whorl I and II stamens are sessile with fleshy filaments that are broader than the anthers, and in E. lorastemon filaments equal anthers in width. Also, as noted by Kos- termans (1937), two basic anther types occur in whorl I and II stamens. In species he assigned to sect. Mi- crolocellata, locelli are small and distally placed (Fig. IA-), while in sect. Macrolocellata locelli occupy almost all available space, and sterile tissue, if present, is located mostly above rather than below locelli (Fig. ID-I). These Macrolocellata-type anthers appear in either of two forms. More commonly, they are ovate with apiculate connectives that emerge between and beyond locelli (Fig. ID-F). Here, locelli are often obliquely hemispherical (as though sliced out of a sphere along a diagonal path) with the deeper part adjacent to the main axis of the anther (e.g., E. browniana). Less frequently, Macrolocellata-type anthers are depressed-oblong to elliptic in shape, and connective tissue is either reduced between, level with, or broad above the two locelli (Fig. 1G-I). Here, locelli are suborbicular in shape, or in other words, hemispherical with the deeper part toward the outside of the anther. Such anthers are typical of the Ampelo- daphne species group, and occur also in E. anomala, E. paniculata, and their allies. Stamens of the third androecial whorl (whorl III) are usually sessile with indistinct filaments equal to or broader than anthers. Narrowly stipitate whorl III stamens occur only in E. anomala, E. coriacea, E. punctulata, E. williamsii, and the Ampelodaphne species group. Anthers of whorl III stamens are consistently four-locellate in E. anomala, but occasionally so in E. williamsii. In both cases, the second pair of locelli appears on the adaxial surface of extrorsely bowed anthers. Whorl III anthers are two-locellate in all other species. Nectariferous glands of whorl III stamens are either sessile or raised on short stalks (stipitate) and range in size from filling the space between stamens to inconspicuous. These glands are lacking in E. chrysovelutina, E. longicaudata, E. metallica, and E. mishuyacensis, but are present at the base of all nine stamens in E. vinotincta.

Apart from a slightly deeper receptacle in pistillate flowers, sexual dimorphism is usually limited to ru- dimentary development of male and female organs. The gynoecium is rarely absent in staminate flowers, but pistillate flowers always have staminodes, and these may almost equal their fertile counterparts. In such cases the tri-lobed or reniform stigma that spreads above the staminodes must be noted to deter- mine gender (but see discussion of E. gracilis).

FRUITS

In most species, fruits mature into a blackish drupe subtended by a reddish cupule. In Lauraceae this structure is typical of most neotropical genera and, as elsewhere, the colors are classic omithochorous sig- nals. In Endlicheria, drupes are either ellipsoid, obovoid, ovoid, or spheroid in shape, and cupules range from deeply hemispherical with at least the lower third of drupes included to patelliform with drupes completely exposed. Cupule margins are often entire, but in several species undulate to strongly lobed margins result from a crown of persistent tepals or tepal bases. Pedicels increase during fruit development to varying degrees and range from terete and distinct from cupules to distally swollen and gradually merging with cupules, i.e., claviform.

Cupules are usually glabrous or at least glabrescent when developing from densely pubescent flowers. Only the cupules of E. chalisea have a dense indumentum that completely covers the outer surface at maturity. In this, and several other species however, cupules are densely pubescent inside.

WOOD AND BARK ANATOMY

In terms of wood and bark anatomy Endlicheria belongs to a large group of genera centered around Ocotea, where it appears to be most similar to Aiouea and Pleurothyrium (Richter, 1981). As reported by Richter (1981), the wood of Endlicheria has a specific gravity of 0.5-0.65 g/cm3 and is diffuse-porous with vessel diameters ranging from 100 ,um to 180 gim. Perforation plates are simple, and intervascular pits are 9-13 ,um diameter; fibers are libriform and sep- tate, with simple pits; axial parenchyma is paratra- cheal, and vasicentric to vasicentric-confluent; rays are multiseriate, 2-3 cells wide, 0.4-1.4 mm high, and heterogeneous, but rarely homogeneous; and oil cells are prevalent. Crystals are present in the wood of 50% of the species Richter examined, but the only report of silica in the bark is based on a misidentified representative, Capucho 565 (F), of Aiouea or Cinna- momum.

FLORAL ANATOMY

Species of Endlicheria have not been examined, but from the accounts given for Persea Mill. (Reece, 1939), Laurus L., Umbellularia (Nees) Nutt. (Kasa- pligil, 1951), and Lindera Thunb. (Boyle, 1980), there appears to be a uniform pattem in Lauraceae that should also apply to Endlicheria. As summarized by Rohwer (1993a, 1993b): First, the six median bundles of the tepals branch off the stele of the floral axis either at the same level or slightly different levels. The

6 FLORA NEOTROPICA

-Az-A

4~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~

FIG. 1. Whorl I stamens of Endlicheria. All seen from within except D which is seen from above. A. E. rubriflora (Palacios et al. 4473). B. E. sprucei (Prance et aL 18037). C. E. metallica (Moretti 1443). D. E. browniana (Palacios 13750). E. E. canescens (Tillett & Tillett 45789). F. E. klugii (Jaramillo et aL 1159). G. E. paniculata (Heringer et al. 3062). H. E. arunciflora (Davidse 27495). I. E. anomala (Vdsquez et al. 4778).

remaining vascular tissue provides two carpel bundles (the ventral one supplying the single apical ovule, the dorsal one supplying the ovary wall), style, and stigma. Stamen traces arise from the ventral side of the tepal bundles, and when not anastomosing, it is clear that traces to the staminal glands arise from their respective stamen traces. An increase in the number

of traces in carpels, tepals, and stamens appears to be correlated with the size of these organs.

EMBRYOLOGY Heo et al. (1998) recendly improved the under-

standing of the embryology in Lauraceae, but a rep-

RELATIONSHIPS 7

resentative of Endlicheria is yet to be examined in this respect. Still, the conservative nature of embryol- ogical characters in the family (Heo et al., 1998) suggests that features found in closely related genera also apply to Endlicheria. If so, it is expected that the anther wall development is of the basic type, the tapetum is amoeboid; tapetal cells are 2 or 4 nucleate; cyto- kinesis is successive; microsprore tetrads are tetra- hedral in shape and mature pollen grains are 2-celled. The ovule is anatropous, bitegmic, and crassinu- cellate, the archesporium is 1-celled; megespore tetrads are linear; a single embryo sac is formed via the Polygonum-type pattern; antipodal cells are ephem- eral, the mature embryo sac is elongate and positioned within the nucellus, the nucellar cap is 2-6 cell layers thick, the ovule is pachychalazal with ramified vascular bundles at the chalaza; and endosperm formation is of the nuclear-type.

KARYOLOGY Chromosome counts have not been done for En-

dlicheria, but unless it differs from most genera of Lauraceae that have been investigated (Okada & Ta- naka, 1975), a base number of x = 12, and 2n = 24 chromosomes is expected. In Lauraceae, polyploidy is known only in Cassytha, Laurus, Sassafras J. Presl. (all with 2n = 48), and Neolitsea (Benth.) Merr. (2n = 72). Cassytha has the largest metaphase chromosomes in the family (5-7 gm long) but otherwise these range from 1 Rtm to 3 Rm in length. The cen- tromere may be medial to subterminal, and hetero- chromatin is found only near the ends of the arms.

PALYNOLOGY Six species of Endlicheria-E. canescens (cited

as E. endlicheriopsis), E. glomerata, E. paniculata, E. lhotzkyi (cited as E. sericea), E. tessmannii, and E. verticillata-were examined in a recent palynological study of neotropical Lauraceae (Raj & van der Werff, 1988). As is typical of the family, the pollen grains in Endlicheria are inaperturate and spheroidal with an extremely thin intine but rather thick and stratified extine. In Endlicheria, pollen size ranged from 15 tm to 27 rim, the extine was 0.5-1 ,im, and scattered with <.5-1 ,um long spinules that are 0.5-2 gm apart, and the intine was 1.5-2.5 trn. According to Raj and van der Werff (1988), such pollen is similar to that of other neotropical Lauraceae with two-locellate anthers, but among these, densely spaced processes that surround and hide the basal cushion of spinules are found only in Endlicheria.

RELATIONSHIPS

SYSTEMATIC POSITION

Relationships among Lauraceae have recently been investigated on the basis of wood and bark anatomy (Richter, 1981), inflorescence structure (van der Werff & Richter, 1996), embryology (Heo et al., 1998), and molecular sequence data (Rohwer, 2000; Chanderbali et al., 2001). These studies show that published classifications (i.e., Nees, 1836; Meissner, 1864; Bentham, 1880; Pax, 1889; Mez, 1889; Kos- termans, 1957; Rohwer, 1993b; van der Werff& Rich- ter, 1996), do not wholly reflect phylogenetic relationships. Although differing in level of resolution provided, these studies approach a consensus over the main groups of genera in the family.

Cryptocaryeae (sensu van der Werff & Richter, 1996) receives support from all five studies. This tribe includes all genera with cupules that enclose drupes except for a terminal pore (e.g., Aspidostemon Roh- wer & Richter and Cryptocarya R. Br.), plus noncupulate Beilschmiedia Nees and its allies. Hypodaphnis Stapf, Cassytha, Caryodaphnopsis Airy Shaw, and Neocinnamomum H. Liu, occupy relatively basal and unstable positions. A small clade of neotropical genera including Anaueria Kosterm., Chlorocardium Rohwer et al., Mezilaurus Kuntze ex Taub., Sextonia van der Werff, and Williamodendron Kubitzki & H. G. Richt., lies sister to a large clade that resolves into a noncupulate generic complex centered around Persea, and a cupuliferous clade that separates into two groups corresponding favorably with the tribes Lau- reae of most systems and Kostermans's (1957) Cin- namomeae (Chanderbali et al., 2001). In Cinnamo- meae, Endlicheria belongs to a large generic complex centered around Ocotea where it shares a dioecious lineage with Ocotea s.str. and Rhodostemonodaphne (Chanderbali et al., 2001).

GENERIC DELIMITATION AND SPECIES GROUPS

To better assess generic delimitation, ITS rDNA sequences of 13 species of Endlicheria and 4 of Rho- dostemonodaphne were produced and analyzed with the 94 ITS sequences of higher Lauraceae previously produced by Chanderbali et al. (2001). The molecular techniques and data analyses are briefly summarized here. Total DNA was obtained from silica gel-dried or herbarium leaves using DNeasy (QIAGEN) extrac- tion kits. PCR amplification of the ITS region was usually conducted using the forward primer LAUR 1 designed by Chanderbali et al. (2001) and ITS B from

8 FLORA NEOTROPICA

Blattner (1999), but for some poor quality templates it was necessary to amplify the region in sections (ITS 1 & ITS2) by combining the appropriate univer- sal primers of White et al. (1990) with LAUR 1 and ITS B. The ITS/5.8S sequences produced include all but the first ca. 10 bp of ITS 1 and include the entire ITS 2 and 5.8S regions. PCR products were purified following the protocol provided by QlAquick gel ex- traction kits (QIAGEN) and sequenced using the dye terminator cycle sequencing protocol (Applied Bio- systems). Sequence reactions were analyzed on an ABI 377 automated sequencer (University of Mis- souri-St. Louis, D. E. Lee and Family Sequencing Fa- cility). Both strands of DNA were read and consensus sequences generated using Sequencher ver. 3.1 (Gene Codes Corp., 1998). Sequences were manually aligned using the sequence editing facilities of Seq- pup version 0.6 (D. Gilbert, Indiana University, Bloo- mington, 1996).

Initial Neighbor Joining (NJ) analyses of the 111 taxa ITS data set placed all new Endlicheria and Rho- dostemonodaphne sequences with previously sampled congeners in the dioecious lineage of the Ocotea complex identified by Chanderbali et al. (2001). The data set was therefore reduced to these dioecious taxa and two outgroup lineages for parsimony analyses. Table I provides Genbank information for these accessions. Heuristic searches for most parsimonious trees were conducted with 10 random taxon additions and TBR branch swapping using PAUP* ver. 4.0b4 (Swofford, 1998). Both the MULPARS and COL- LAPSE options were in effect, but the STEEPEST DESCENT option was not employed. Characters were assumed to be unordered (i.e., Fitch parsimony), equally weighted, and gaps were treated as missing data. Parsimony uninformative characters were excluded. Parsimony analyses surpassed memory limi- tations in the first addition replicate and 10,000 equally parsimonious topologies were retained for branch swapping (L = 138, CI = 0.682, RI = 0.849). The majority rule topology with clade support esti- mated from bootstrap analyses (Felsenstein, 1985) with the above heuristic search settings but with MAXTREES set to 100, is shown in Figure 2.

In the dioecious lineage of the Ocotea complex, Rhodostemonodaphne and Ocotea s.str. are techni- cally separated by the arrangement of the four locelli in whorl I and II stamens; locelli are in a shallow arc or horizontal row in Rhodostemonodaphne but in superimposed pairs in Ocotea. Endlicheria is distinguished from dioecious relatives solely by its two- locellate anthers. The limited value of locelli number in generic delimitation has often been noted (e.g., van

der Werff, 1984; Burger, 1988; Rohwer et al., 1991), and the two-locellate condition apparently circum- scribes a polyphyletic assemblage of species in En- dlicheria.

Molecular data place Endlicheria punctulata with Ocotea pauciflora (Nees) Mez, a representative of the Ocotea cernua species group (sensu Rohwer, 1986), in a clade that lies sister to remaining dioecious taxa of the Ocotea complex (Chanderbali et al., 2001; Fig. 2 herein). With members of the 0. cernua species group E. punctulata shares pauciflorous inflorescences with sparsely pubescent flowers and glabrous concolorous leaves with immersed venation below. Endlicheria coriacea differs from E. punctulata mainly in habit and is certainly a second example of a two-locellate species in the Ocotea cernua species group. Rhodostemonodaphne elephantopus Madrifiin most likely belongs here as well. Four locelli arranged in a shallow arc refer this species to Rhodostemono- daphne, but it is isolated there (Madriiinn, 1996a), has the leaf morphology of the 0. cernua species group, and is vegetatively indistinguishable from E. coriacea. The two species of Endlicheria here allied to the Ocotea cernua species group, comprise the En- dlicheria punctulata species group.

Remaining Endlicheria sampled for molecular characters represent all species groups here recognized (see Numerical List of Taxa), and all place with a diverse representation of Rhodostemonodaphne (Fig. 2). Although this Endlicheria-Rhodostemono- daphne alliance is unresolved, it is itself significant. Several species of Endlicheria are morphologically so similar to species of Rhodostemonodaphne that they seem misplaced in their present generic station. With these two genera in a strongly supported clade, morphology-based suspicions (e.g., Rohwer et al., 1991; Madrifinan, 1996a; Madrifiain, 1996b), that separate lineages in Endlicheria are derived from ancestors that would fit the generic characters of Rho- dostemonodaphne, now have molecular support.

Among the species of Endlicheria clearly closer to Rhodostemonodaphne than to congeners are those Kostermans (1937) assigned to section Microlocellata (E. bullata, E. longicaudata, E. rubriflora, E. sprucei), and a new species, E. ferruginosa, here described. These five species comprise the Microlo- cellata species group herein. As in Rhodostemono- daphne, the locelli in these species are distally positioned in the anther (Fig. 1A, B), and flowers are rotate with fleshy tepals. At least E. rubriflora and R. antioquensis Madriiain appear to be sister taxa, and it is likely that the others are each independently derived from Rhodostemonodaphne. As I do not believe

RELATIONSHIPS 9

TABLE I. Sources of plant material. Accessions marked by an asterisk (*) were previously submitted to Genbank by

Chanderbali et al. (2001)

Genbank accession

Taxon Provenance Voucher numbers

Endlicheria anomala (Nees) Mez Brazil, Amazonas Lohmann 343 (MO) AF363371

Endlicheria bracteolata (Meisn.) Brazil, Amazonas Ribeiro 885 (MO) AF363372 C. K. Allen

Endlicheria chalisea Chanderbali Guyana, Iwokrama Chanderbali 252 (MO) AF272271* Reserve

Endlicheria citriodora van der Peru, Iquitos Va'squez 25231 (MO) AF272272* Werff

Endlicheria dysodantha (Ruiz & Bolivia, Santa Cruz Vargas 1098 (MO) AF363373 Pav.) Mez

Endlicheria gracilis Kosterm. Guyana, Iwokrama Chanderbali 250 (MO) AF363374 Reserve

Endlicheria longicaudata (Huber) Brazil, Amazonas Assunqao 366 (MO) AF363375 Kosterm.

Endlicheria metallica Kosterm. Ecuador, Napo Tirado 1010 (MO) AF363376

Endlicheria multiflora (Miq.) Mez Guyana, Karanambo Chanderbali 206 (MO) AF363377

Endlicheria paniculata (Spreng.) Brazil, Parana Silva & Abe 2883 (MO) AF363378 J. F. Macbr.

Endlicheria punctulata (Mez) C. K. French Guiana de Granville 1448 (MO) AF272273* Allen

Endlicheria pyriformis (Nees) Mez Ecuador, Napo Neill 9842 (MO) AF363379

Endlicheria reflectens (Nees) Mez Guyana, Karanambo Chanderbali 208 (MO) AF272274*

Endlicheria rubra Chanderbali Peru, San Martin van der Werff 15436 (MO) AF363380

Endlicheria ruforamula Chander- Peru, San Martin van der Werff 15736 (MO) AF363381 bali

Endlicheria sprucei (Meisn.) Mez Brazil, Amazonas Vicentini 1224 (MO) AF363382

Endlicheria tessmannii O.C. Peru, Loreto Vasquez 25237 (MO) AF363383 Schmidt

Nectandra amazonum Nees Guyana, Iwokrama Chanderbali 217 (MO) AF272289* Reserve

Nectandra cuspidata Nees & Mart. Guyana, Kamarang Chanderbali 279 (MO) AF272291*

Nectandra psammophila Nees & C. Brazil, Sao Paulo Lorea-Hermandez 5595 (MO) AF272292* Mart.

Nectandra turbacensis (HBK) Nees Puerto Rico, Rio Taylor 11746 (MO) AF272295* Grande, El Verde

10 FLORA NEOTROPICA


Chanderbali et al. (2001) (Continued)

Genbank accession


Ocotea botrantha Rohwer U.C. Riverside Scora 99-1 (UCR) AF272297*

Ocotea ceanothifolia (Nees) Mez Guyana, Demerara, Chanderbali (MO) AF272299* Mabura Hill

Ocotea guianensis Aubl. Guyana, Demerara, Chanderbali 232 (MO) AF272302* Mabura Hill

Ocotea helicterifolia (Miesn.) Mexico, Oaxaca Torres 11911 (MO) AF272303* Hemsl.

Ocotea heydeana (Mez & Donn. Honduras, Yoro, Evans 1760 (MO) AF272304* Sm.) Bernardi Pico Pijol

Ocotea nigra Benoist Guyana, Iwokrama Chanderbali 162 (MO) AF272308* Reserve

Ocotea pauciflora (Nees) Mez Guyana, Demerara, Chanderbali 219 (MO) AF2723 10* Mabura Hill

Ocotea percoriacea (Mez) Kosterm. Brazil, Goias Lorea-Hernandez 5584 (MO) AF272311 *

Ocotea pulchella Mart. Brazil, Minas Ger- Lorea-Hernandez 5575 (MO) AF262312* ais

Ocotea schomburgkiana (Nees) Guyana, Iwokrama Chanderbali 286 (MO) AF272315* Mez Reserve

Ocotea spixiana (Nees) Mez Brazil. Minas Ger- Lorea-Hernandez 5574 (MO) AF272316* ais

Ocotea tomentella Sandwith Guyana, Kamarang Chanderbali 284 (MO) AF272317*

Ocotea tristis (Nees & Mart.) Mez Brazil, Minas Ger- Lorea-Hernandez 5577 (MO) AF272318* ais

Pleurothyrium cinereum van der Peru, San Martin van der Werff 15325 (MO) AF272329* Werff

Pleurothyrium insigne van der Ecuador, Napo, Ja- Neill 9033 (MO) AF272330* Werff tun Sacha

Rhodostemonodaphne crenaticupula Guyana, Iwokrama Chanderbali 265 (MO) AF272331* Madrifian Reserve

Rhodostemonodaphne kunthiana Guyana, Iwokrama Chanderbali 257 (MO) AF363384 (Nees) Rohwer Reserve

Rhodostemonodaphne negrensis Brazil, Amazonas Vicentini 628 (MO) AF363385 Madrifian

Rhodostemonodaphne parvifolia Brazil, Amazonas Assun,cao 327 (MO) AF363386

RELATIONSHIPS 11


Chanderbali et al. (2001) (Continued)

Genbank accession


Rhodostemonodaphne peneia Mad- Brazil, Amazonas Ramos 2834 (MO) AF363387 rii~ian

Rhodostemonodaphne praeclara Guyana, Iwokrama Chanderbali 256 (MO) AF272332* (Sandwith) Madrinan Reserve

Rhodostemonodaphne recurva van Brazil, Amazonas Vicentini 653 (MO) AF272333* der Werff

Rhodostemonodaphne scandens Guyana, Iwokrama Chanderbali 271 (MO) AF272334* MadriiaTn Reserve

these species form a monophyletic group, their affinities are discussed individually in the Systematic Treatment, below.

A likely monophyletic group crossing the Endli- cheria-Rhodostemonodaphne boundary is comprised of Endlicheria chrysovelutina, E. metallica, Rhodos- temonodaphne grandis, R. praeclara (Sandw.) Mad- rifia'n, R. peneia MadriiaTn, R. saiulensis Madrifian, and R. recurva van der Werff. All share the peculiar asymmetric basal secondaries described in the leaf venation section above, sericeous indument, and large fruits with robust hemispherical cupules. They assign to different genera on the basis of locelli number, but R. recurva is conceivably intermediate in combining four-locellate anthers in whorl III stamens with two- locellate anthers in whorls I and II. Although flowers are rotate in species assigned to Rhodostemonoda- phne, but campanulate in the two assigned to Endli- cheria, intermediate conditions are found in material that may represent undescribed species. Thus, Stey- ermark 59001 from Bolivar, Venezuela (F, MO) combines campanulate flowers with four-locellate anthers, while in Prance et al. 14616 from Amazonas, Brazil (INPA, NY), and Sdnchez Vega et al. 9345 from San Martin, Peru (MO), two-locellate anthers are found in rotate flowers. The latter two collections may represent additional two-locellate species in this transgeneric species group, but are not described here because anthers are poorly developed and devoid of pollen, possibly indicating hybrid origin. They key out to E. metallica aff. sp. 1 and E. metallica aff. sp. 2, respectively. Pending better material of these entities, E. metallica and E. chrysovelutina are the

only described members of the E. metallica species group.

The Endlicheria browniana species group (species 10-18) may also find its closest relatives in Rhodos- temonodaphne. Here, stamens of all whorls are fleshy and sessile, and anthers are ovate with obliquely hemispherical locelli between and beyond which an apiculate connective protrudes (Fig. ID). In most, the flowers are trumpet-shaped with slender pedicels that widen distally to merge with an infundibuliform receptacle. Especially in E. browniana, E. jefensis, and E. mishuyacensis, flowers are densely papillose inside. Such papillosity is rare in Endlicheria but typical of Rhodostemonodaphne., wherein similarly shaped flowers are found in its type species, R. laxa (Meisn.) Rohwer and, among others, in R. cyclops Madriiain where whorls I and II anthers can be three-locellate, conceivably illustrating the transition from four to two locelli.

Remaining species of Endlicheria can be sorted into groups that appear to be linked by intermediates. Still, rather than constituting a single lineage, these species groups appear to intermingle freely with counterparts in Rhodostemonodaphne.

One close-knit group of species (19-34) includes those previously assigned to Ampelodaphne (Meis- sner, 1864; Mez, 1889), herein, the Ampelodaphne species group. It is characterized by subverticillate leaves and bracteate inflorescences with numerous fragile flowers that have a deep, almost tubular, receptacle below spreading to recurved membranaceous tepals. Here, all stamens have slender distinct filaments, and transversely oblong anthers are dominated

12 FLORA NEOTROPICA

100 Endlidsefl nUIOmIaTI 53 Enck*_ biatodsta .. . _~anma

100 E d EdIdeapanbiata L---Ersdtwi baIs

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clades are indicated.~ ~ ~ ~ ~ ~ ~ ~ ~~~nd

1 10*--- 00p p q . 0. ocd R ills-

FI. 2. Phloeneiffnte of. EndichriaindctdbIT seuenc dt. T..is.... majriymetployo th .00 equa legt tre -L = 138 C- = ' 0.82 RI =F.4)rtie o rac wpigsosE dlceipntaa

paire wit Ocyoteanetuifr affndties oftndiherseies ofindicheianted wlSsqunedta. Rhodstmndpe 50% majrt wellesuppologyeo

but poorly resolved clade. Majority rule percentages are indicated above, and bootstrap values >50% below, branches. Main clades are indicated.

RELATIONSHIPS 13

by suborbicular locelli (Fig. lH). Two subgroups are suggested by venation and leaf texture. In the Endli- cheria multiflora subgroup, also with E. arenosa, E. dictifarinosa, E. levelii, E. macrophylla, and E. melinonii, tertiary and higher-order veins are always immersed above and thus inconspicuous against usually coriaceous laminae. In the E. bracteata subgroup, tertiary and fourth-order veins are prominent above, forming a conspicuous reticulate pattern over usually chartaceous laminae. Other members of the E. bracteata subgroup are E. chalisea, E. cocuirey, E. directonervia, E. glomerata, E. tessmannii, and E. verticillata. Representatives of both subgroups, E. chalisea, E. multiflora, and E. tessmannii, constitute one of the few clades resolved in the Endlicheria- Rhodostemonodaphne alliance by ITS sequence variation (Fig. 2). This clade lies sister to a pair of Rho- dostemonodaphne species while Endlicheria refiec- tens, a species somewhat intermediate between the two subgroups in having the adaxially prominent tertiaries of the E. bracteata subgroup but the rusty tomentose indument typical of the E. multifiora subgroup, appears to share molecular characters with E. sprucei (Fig. 2). This resolution is weakly supported, but it is intriguing from a morphological point of view. The two species of Rhodostemonodaphne, R. crenaticupula Madrinia'n and R. negrensis Madrinian, placed close to core members of the Ampelodaphne species group are morphologically very similar to E. sprucei. Indeed, fruiting specimens of R. crenaticupula are sometimes mistaken for E. sprucei (Madri- nian, 1996b). Further, the sessile outer stamens of E. sprucei (e.g., Fig. 1B) are typical of R. crenaticupula. Therefore, if the molecular signal for close relationship between E. reflectens and E. sprucei has phylogenetic significance, it is conceivable that the Ampe- lodaphne species group relates to members of Rhodostemonodaphne via E. sprucei.

Like members of the Ampelodaphne species group, Endlicheria anomala and E. williamsii have stipitate stamens in all staminal whorls, but their flowers are otherwise quite different. In both, the flowers are rotate with horizontally spreading tepals, and whorl III anthers have four-locellate anthers (incon- sistently so in E. williamsii) although whorls I and II are two-locellate. Rhodostemonodaphne recurva and R. revolutifolia show similar variation in locelli number among androecial whorls, but close relationship with these is not supported by other characters. ITS sequence data places Endlicheria anomala in the En- dlicheria-Rhodostemonodaphne alliance without clearer indications of affinities. Endlicheria anomala

and E. williamsii comprise the E. anomala species group.

In all remaining species of Endlicheria, whorl I and II stamens are stipitate, to a greater or lesser de- gree, but whorl III stamens are broadly stipitate or sessile with filaments equal to or wider than anthers. On the basis of indument and stamen morphology, three groups of similar, and presumably closely related, species can be recognized.

One such group is comprised of Endlicheria canescens, E. citriodora, E. duotincta, E. lorastemon, E. oreocola, E. rubra, E. szyszylowiczii, E. vinotincta, and E. xerampela (species 37-45). Most have a dense reddish tomentose indumentum covering branchlets, and whorls I and II stamens have densely greyish tomentose filaments and ovate anthers with an apically prolonged connective (Fig. lE). These stamens resemble those of Aniba Aubl., and at least flowers of E. citriodora are strikingly similar to those of that genus. However, molecular data unambiguously place E. citriodora (and E. rubra) in the Endlicheria-Rho- dostemonodaphne alliance (Fig. 2) and associate An- iba with Licaria Aubl. (Chanderbali et al., 2001). Endlicheria xerampela is placed here on the basis of overall resemblance to E. canescens, although its whorl I and II anthers are truncate above.

Members of the Endlicheria canescens species group appear to be related to Rhodostemonodaphne species with similar dense reddish indumentum. In particular, E. vinotincta is almost indistinguishable from R. celiana C. K. Allen and R. steyermarkiana (C. K. Allen) van der Werff without counting locelli; in venation and leaf shape E. citriodora and R. kunthiana (Nees) Rohwer are extremely similar; E. canescens shows the asymmetric basal secondaries of R. rufovirgata Madriiain and R. mirecolorata (C. K. Allen) Rohwer; and E. rubra is vegetatively indistinguishable from material, Garc(a-Barriga 13944 from Amazonas, Colombia (NY, US), representing an undescribed species of Rhodostemonodaphne. Relation- ships among these and dioecious species of Ocotea with a similar vestiture, e.g., 0. discrepens C. K. Al- len, 0. endlicheriopsis Mez, 0. indirectinervia C. K. Allen, and 0. rufovestita Ducke, should be investigated further.

The Endlicheria sericea species group (species 46-55) comprises species with stamens similar to those in the E. canescens species group but with a dense sericeous indument that usually persists on the lower leaf surface. Here, leaves are usually ovate and coriaceous with arcuate secondaries that tend to follow asymmetric courses. Close relationship with End-

14 FLORA NEOTROPICA

licheria metallica, E. chrysovelutina, and their allies in Rhodostemonodaphne is suggested by shared sericeous indument, and occasional asymmetric basal secondaries in E. klugii and E. ruforamula, but the stipitate whorl I and II stamens in the E. sericea group speak against it. Instead, a closer link to Rhodoste- monodaphne may be provided by material, e.g., Ac- evedo et al. 1370 (F, MO, US), of an undescribed species. There, the flowers are similar in size and shape to those of E. griseo-sericea but anthers are three-locellate in whorls I and II and four-locellate in whorl III. This unusual androecium is more like that of Rhodostemonodaphne than Endlicheria, but in leaf shape and venation this species appears closer to the E. sericea species group. It is not described for rea- sons given in the discussion of E. griseo-sericea.

Endlicheria acuminata, E. gracilis, and E. krukovii have triplinerved leaves and appear to form a loose association around E. paniculata. All have stipitate whorl I and II stamens, and in E. acuminata, E gracilis, and usually also E. paniculata, anther apices are truncate rather than apiculate. Endlicheria krukovii is close to E. acuminata, although its anther apices are apiculate. Endlicheria nilssonii is poorly known but is placed here on the basis of its stipitate stamens with truncate anther apices.

The above species groups are informal concepts meant to parcel infrageneric heterogeneity into more homogeneous units that can each be considered in more detail. In the case of the Endlicheria punctulata species group (species 1-2), affinities are confidently assumed to lie with Ocotea, while the Microlocellata species group (species 3-7) and E. metallica species group (species 8-9) are clearly best compared to Rho- dostemonodaphne. In the case of the E. browniana species group (species 10-18), counterparts in Rho- dostemonodaphne are conceivably provided by R. laxa and allies. The Ampelodaphne species group (species 19-34) has deviated from ancestral conditions and become very distinctive, but may link to Rhodostemonodaphne via E. sprucei. The E. sericea species group (species 46-55) is also very distinctive but may include a species with the generic characters of Rhodostemonodaphne. The E. canescens species group (species 37-45) is heterogeneous. At least E. vinotincta and E. rubra have clear four-locellate counterparts in Rhodostemonodaphne, but the affinities of remaining members, like the E. anomala and E. paniculata species groups, are unclear.

Given the numerous transgeneric alliances involv- ing species of Endlicheria, Rhodostemonodaphne, and to a lesser extent Ocotea, the current concepts of these three genera warrant careful revision. In Ocotea

all anthers are four-locellate with the locelli arranged in superimposed pairs and species either have bisexual flowers or are dioecious. Rhodostemonodaphne differs from dioecious Ocotea only in having its four locelli arranged in a shallow arc, while Endlicheria differs from Rhodostemonodaphne and dioecious Ocotea only in having two-locellate anthers. Since Endlicheria and Rhodostemonodaphne nest in dioecious Ocotea (Chanderbali et al., 2001; Fig. 2 herein), a rather broad generic concept would conceivably treat this dioecious lineage as one genus; viz. Ocotea. This would make Ocotea monophyletic, provided that its current bisexual members are removed, but whether demoting the delimitation of Endlicheria and Rhodostemonodaphne to infrageneric significance improves the current situation is questionable. Per- haps the Endlicheria-Rhodostemonodaphne alliance alone can be considered for generic status. However, such a group is too heterogeneous to be circumscribed in morphological terms, and might yet be found to be paraphyletic with respect to species of Ocotea. An opportunity to revise generic concepts will arise when relationships among the members of these three genera are more fully elucidated. Until then, this treatment addresses the great practical necessity of providing modem revisions at the species level in Lauraceae, even before all the generic concepts are solidified.

SPECIES CONCEPTS Specific status is here granted to the smallest as-

semblage of individuals (collections) that is morphologically distinguishable from other such assem- blages. Species so circumscribed possess at least one unifying character or character combination that is not found in other species, and may either be internally uniform or variable. Infraspecific categories are avoided because the causes of infraspecific variation are unknown.

The source of species characters varies with species and species groups. Vegetative morphology is almost uniform in the Endlicheria sericea species group but floral variation is important, while flowers are almost identical in the Ampelodaphne and E. punctulata species groups but there is significant vegetative variation. In the other species groups distinguishing characters are provided by both floral and vegetative variation.

DISTRIBUTION AND BIOGEOGRAPHY

Endlicheria is a neotropical genus centered in South America, while reaching Costa Rica in Central

DISTRIBUTION AND BIOGEOGRAPHY 15

America, Guadeloupe in the Caribbean Islands, and its southernmost extreme in the Atlantic coastal forests of SE Brazil (Fig. 3). This distribution is almost paralleled by Rhodostemonodaphne (lacking only in the Caribbean). Ocotea s.str. is similarly distributed, but more widespread in Central America and the Ca- ribbean. Closely related Nectandra s.str. and Pleu- rothyrium are also similarly distributed. The biogeographic significance of the commonality in these generic distributions was partially addressed by Chanderbali et al. (2001).

According to the biogeographic history of Laura- ceae reconstructed there, the lineage to which these genera belong, namely the Ocotea complex, has an ancestral Madrean-Tethyan distribution (sensu Axel- rod, 1975) and radiated in the New World during the Miocene. Furthermore, they constitute a South Amer- ica-centered clade that diverged from a Central Amer- ica-centered sister group, the Ocotea helicterifolia species group, ca. 20 million years ago (Chanderbali et al., 2001). As this divergence time coincides with

increased Andean uplift in the Early Miocene, Chan- derbali et al. (2001) envisaged that the resulting ele- vational barriers to migration isolated the ancestors of Endlicheria and its sympatric relatives in South America. The subsequent biogeographic history of these genera and Endlicheria itself has not been investigated in detail but conceivably involves a complex scenario of diversification along edaphic and ele- vational gradients in South America and dispersal to Central America and the Caribbean Islands.

The species of Endlicheria occur in moist forest habitats from elevations of around sea level to 2500 m in the Andean and Guianian highlands. The limi- tations imposed by low water availability are evident in the absence of Endlicheria from the caatinga vegetation of eastern Brazil and the Chaco region of southern South America (Fig. 3). Species diversity is greatest in the eastern Andean foothills where lowland and lower montane elements intermingle. Central Amazonia and the Guianas are not nearly as rich in diversity, and in the Atlantic coastal forests of SE Bra-

e~~~~~~~I Distibuio o f Endlih;r;a

,'----- ------ ----- ----- ....f -------------- ----- ------- - --- ------- -

~~~~~X----- ------ --------------------- -

30 S.I2 W , k

FIG. 3. Distribution of Endlicheria.

16 FLORA NEOTROPICA

zil, only two species are found. The distribution of E. paniculata almost equals the generic range of Endli- cheria (lacking only in NE South America and the Caribbean Islands) but all other species have narrower ranges. Several species are widespread throughout the lower slopes of the eastern Andes to western, and rarely central, Amazonia, e.g., E. metallica and E. rubriflora. Of these, E. formosa and E. ruforamula range west of the Andes to Costa Rica and Colombian Choco, respectively. Endlicheria bracteolata, E. chalisea, E. pyriformis, and E. szyszylowiczii range from the Guianas through the Amazon basin to lower montane eastem Andean slopes, while E. canescens and E. gracilis range from the Guianas to the eastern An- dean slopes without a central Amazonian presence. Lowland species with wide distributions all appear to be elements of riparian forests, e.g., E. anomala, but narrow distributions for species of terra firme may be collecting artifacts (see Nelson et al., 1989). The possible exceptions, as suggested by relatively well- collected species, are rare. Endlicheria glomerata appears to be endemic to the coastal forests of SE Brazil; E. sericea is essentially an element of the lower montane slopes of the Lesser Antilles and Trinidad but possibly also in adjacent Venezuela; and E. browniana is restricted to the western side of the Andes from Ecuador to Panama.

SYSTEMATIC TREATMENT

GENERIC DESCRIPTION

Endlicheria Nees, Linnaea 8: 37. 1833, nom. cons. Type. Endlicheria hirsuta (Schott) Nees (lectotype, designated by Kostermans, 1937; = Endli- cheria paniculata (Spreng.) J. F. Macbr.). Goeppertia Nees, Syst. laur. 365. 1836. Type. Goepper-

tia hirsuta (Schott) Nees (lectotype here designated; = Endlicheria paniculata (Spreng.) J. F. Macbr.).

Ampelodaphne Meisn., DC. Prodr. 15(1): 81. 1864. Type. Ampelodaphne macrophylla (lectotype here designated).

Huberodaphne Ducke, Arch. Jard. Rio de Janeiro 4: 191. 1925. Type. Huberodaphne longicaudata Ducke (lectotype here designated).

Endlicheria subgen. Ampelodaphne (Meisn.) Mez, Jahrb. Konigl. Bot. Gart. Berlin 5: 125. 1889. Type.

Ampelodaphne macrophylla (lectotype here designated).

Endlicheria subgen. Euendlicheria Mez, Jahrb. Konigl. Bot. Gart. Berlin 5: 114. 1889. Type. Endlicheria browniana Mez (lectotype here designated).

Endlicheria subgen. Hemiajouea Mez, Jahrb. Konigl. Bot. Gart. Berlin 5: 114. 1889. Type. Endlicheria paradoxa Mez (lectotype here designated).

Endlicheria subgen. Ocoteopsis Mez, Jahrb. Konigl. Bot. Gart. Berlin 5: 132. 1889. Type. Endlicheria anomala (Nees) Mez (lectotype here designated).

Endlicheria sect. Microlocellata Kosterm., Recueil Trav. Bot. NMerl. 34: 511. 1937. Type. Endlicheria bullata Ducke (lectotype here designated).

Endlicheria sect. Macrolocellata Kosterm., Recueil Trav. Bot. NMerl. 34: 517. 1937. Type. Endlicheria gracilis Kosterm. (lectotype here designated).

Trees or shrubs. Leaves alternate, widely spaced along branchlets or arranged in close spirals; mostly penninerved, occasionally asymmetrically trinerved, tripli- or quintuplinerved. Inflorescences mostly in axils of foliage leaves, but often in cataphylls, thyrso- paniculate, the terminal dichasia rarely reduced to pseudo-umbels or botryoids. Flowers unisexual, trimerous. Tepals 6, in 2 whorls of 3, usually spreading, rarely inflexed, erect, or reflexed. Stamens 9 in 9 whorls of 9; whorl I and II stamens ? equal, the filaments distinct or undifferentiated, the anthers two- locellate; whorl III stamens usually sessile with in- distinctly differentiated filaments, rarely stipitate, the anthers two-locellate, rarely four-locellate, the basal glands sessile or stipitate, rarely absent; fourth whorl of staminodes occasional; pistillode present or absent. Pistillate flowers usually deeper with smaller sterile stamens; ovary superior; stigma tri-lobed, discoid or reniform. Fruits borne on terete or claviform pedicels; cupules usually hemispherical, occasionally flat, rarely reflexed, the margins entire, undulate or with persistent tepal bases; drupes elliptic, obovoid, ovoid, or spheroid.

The name Schauera was only tentatively offered in place of Endlicheria by Nees (in Lindley, 1836) and is therefore not validly published.

Endlicheria subgen. Anosphaeria Mez is a synonym of Aiouea Aubl.

Key to the species of Endlicheria

1. Lower surface of mature leaves concealed by the indument cover. 2. Indument on lower leaf surface comprised of a dense creamish tomentose cover interspersed with

longer reddish, straight, erect hairs. 3. Leaves alternate, widely distributed along branchlets. Flowers campanulate, densely pubescent

outside. Outer anthers apiculate above. E Andean foothills .......................................................... 44. E. duotincta

SYSTEMATIC TREATMENT 17

3. Leaves clustered at tips of branchlets. Flowers rotate, sparsely pubescent outside. Outer anthers truncate above. W Amazonian lowlands ................................................................. 36. E. williamsii

2. Indument on lower leaf surface comprised of straight appressed hairs, these silvery-grey to golden, uniform in length and orientation. 4. Flowers sparsely pubescent outside, the surface clearly visible.

5. Flowers rotate; tepals spreading horizontally. Anthers of whorl III stamens four-locellate. Flooded Amazonian lowlands .............................................................. 35. E. anomala

5. Flowers obconical or urceolate; tepals inflexed or erect. All anthers two-locellate. Nonflooded forests of E Andean foothills and SW Amazonia ................................................................... 55. E. robusta

4. Flowers densely pubescent outside, the surface not visible. 6. Basalmost secondaries asymmetric, one or both merging with leaf margin near base or emerging from

junction of midrib and leaf base. 7. Tepals broader than long. Whorls I & II stamens sessile, their anthers over topped by a broad

sterile truncate apex. 8. Branchlets sericeous, the hairs closely appressed to the surface, silvery grey. E Andes to central

Amazonia .............................................................. 8. E. metallica 8. Branchlets densely velutinous, the hairs erect, golden to yellowish brown. White sands around

Iquitos, Peru .............................................................. 9. E. chrysovelutina 7. Tepals longer than broad. Whorls I & II stamens stipitate, their anthers ovate with narrowly

apiculate apices. 9. Branchlets sericeous, the hairs appressed. Tepals spreading horizontally; receptacle narrowly

infundibuliform, longer than broad; whorls I & II filaments S-shaped. Cupules shallow, drupes exserted. W Amazonia to E Andean slopes .............................................................. 53. E. klugii

9. Branchlets tomentellose, the hairs ascending. Tepals ascending; receptacle as broad as or broader than long; outer filaments straight. Cupules deeply hemispherical, drupes at least 1/4 included. Central Amazonia to W Andean slopes of Colombia and Ecuador .................... 50. E. ruforamula

6. Basalmost secondaries ? symmetric, emerging from midrib above leaf base. 10. Leaves tripli- or quintuplinerved. The lowermost pair(s) of secondaries subopposite shortly above

the leaf base, arcuate, ascending to beyond midlamina, the others emerging beyond midlamina or absent. 11. Flowers infundibuliform or campanulate, the tepals ascending at anthesis. Guianas,

Amazonia, Pacific coast of Colombia .................................................................... 47. E. bracteolata 11. Flowers rotate, the tepals spreading horizontally at anthesis. Lesser Antilles, NE

Venezuela .................................................................... 46. E. sericea 10. Leaves penninerved. Secondary veins ? uniformly spaced along midrib.

12. Both branchlets and inflorescence axes tomentellose or tomentose, the hairs ascending or sprawling. 13. Hairs on branchlets ca. 0.2 mm long or shorter. Tepals erect to ascending, the inner

surface glabrous. Central Amazonia to W Andean slopes of Colombia and Ecuador .................................................................................................................................. 50. E. ruforamula

13. Hairs on branchlets ca. 0.5 mm long. Tepals spreading, the inner surface pubescent. 14. Leaves ovate. Hairs on branchlets pale to rust red. Tepals chartaceous, sparsely

pilose inside. Mato Grosso, Brazil ....................................................... 52. E. lhotzkyi 14. Leaves obovate. Hairs on branchlets greenish yellow. Tepals fleshy, densely

papillose inside. Amazonas, Brazil ..................................................... E. metallica aff. sp. 1 12. Either or both branchlets and inflorescence axes silvery to rusty sericeous, the hairs closely

appressed to the surface. 15. Receptacle narrowly infundibuliform, longer than broad. Pedicels claviform, indistinct in

flower and fruit. Plants mostly from lowland Amazonia at ca. 100-200 m. 16. Midrib prominent above. Filaments of whorl I & II stamens S-shaped. W

Amazonia to E Andean slopes ......................................................... 53. E. klugii 16. Midrib impressed above. Filaments of whorl I & II stamens straight. W Amazonian

lowlands .................................................... 54. E. argentea 15. Receptacle campanulate or shallowly infundibuliform, broader than long. Pedicels ?

terete, distinct in flower and fruit. Plants mostly from lower montane Andean slopes above 500 m. 17. Leaves drying dark olive-brown above. Receptacle constricted below tepals. Whorl

I and II anthers obovate, the apex broadly truncate to retuse over the locelli. Lower montane Andes .................................................... 49. E. griseo-sericea

18 FLORA NEOTROPICA

17. Leaves drying greenish above. Receptacle gradually merging with tepals. Whorl I and II stamens ovate or depressed-ovate, the apex apiculate between the two locelli. 18. Indument on branchlets, leaves, and flowers golden to rust brown. Anthers of

all whorls depressed-ovate, the locelli ? apical. Cupule margins strongly lobed. E Andes ............................................ 51. E. aurea

18. Indument on branchlets, leaves, and flowers tawny to silvery grey. Anthers of whorls I & II ovate, the locelli introrse-latrorse; anthers of whorl III oblong, locelli extrorse-latrorse. Cupule margins entire or weakly undulate, not strongly lobed. 19. Flowers rotate, the tepals spreading, densely grey-tomentose inside.

Cupules deeply hemispherical, densely sericeous inside. Lesser Antilles, NE Venezuela ........................................ 46. E. sericea

19. Flowers campanulate or infundibuliform, the tepals erect to ascending, glabrous inside. Cupules shallow, glabrous inside. NW South America, Panama ..................................................................... 48. E. tschudyana

1. Lower surface of mature leaves clearly visible through indument cover. 20. Leaves and inflorescences clustered in close spirals at the tips of branchlets. Inflorescence bracts and

bracteoles persistent at anthesis. 21. Tertiary and higher-order veins immersed above, inconspicuous against the lamina.

22. Inflorescence axes and exterior of flowers glabrous. 23. Hairs on branchlets and lower leaf surface reddish to rust brown, erect. Upper Rio

Negro, Brazil, Venezuela ................................................... 19. E. arunciflora 23. Hairs on branchlets and lower leaf surface creamish white, weak, sprawling, silky.

Peruvian Amazonia .......................................................... 20. E. arachnocome 22. Inflorescence axes and exterior of flowers moderately to densely pubescent.

24. Hairs on sides of midrib and secondaries below stiffly straight. 25. Hairs on lower leaf surface very short, ca. 0.2 mm. Well-drained white sand soils.

Central Amazonia ............................................... 21. E. arenosa

25. Hairs on lower leaf surface relatively long, ca. 0.5 mm. Flooded forest, NE South America ............................................... 26. E. melinonii

25. Hairs on sides of midrib and secondaries below curved or crooked, not straight. 26. Leaves ovate to elliptic. Basal glands large, surrounding filaments of the

inner staminal whorl. Central Amazonia ........................................... 22. E. macrophylla 26. Leaves obovate to obovate-elliptic. Basal glands minute, inconspicuous,

leaving the filaments of the inner staminal whorl exposed. 27. Hairs on branchlets, midrib below, and inflorescence axes reddish to rust

brown. Guianas ....................................... 23. E. multiflora 27. Hairs on branchlets, midrib below, and inflorescence axes greyish to

white. 28. Leaves subsessile, the base (sub-)cordate, abruptly contracted into

the petiole. SE Venezuela, Roraima, Brazil ......................... 25. E. dictifarinosa 28. Leaves petiolate, the base acute, gradually tapering into the petiole.

Central to W Amazonia .................................... 24. E. levelii

21. Tertiary and higher-order veins raised above, forming a prominent network over the lamina.

29. Tertiary veins reticulating between secondaries. Shrubs from Guiana shield savanna formations ....................................................... 27. E. reflectens

29. Tertiary veins undivided or once-forked between adjacent secondaries. Trees from wet

forests. 30. Hairs on lower leaf surface appressed. Iquitos, Peru, and vic . .............................. 31. E. tessmannii

30. Hairs on lower leaf surface erect. 31. Branchiets and midrib below tomentose or tomentellose, the hairs ca. 0.5 mm

long or shorter, if longer then sprawling. 32. Leaves subsessile, the base subcordate, abruptly contracted into the petiole.

W Amazonia ........................................... 30. E. cocuirey 32. Leaves petiolate, the base acute or obtuse, gradually tapering into the petiole.

Central Amazonia to E Andean slopes ........................................... 32. E. directonervia 31. Branchlets and midrib below hirsute, the hairs ca. 1 mm or longer, straight, stiff.

33. Hairs on branchlets reddish. Distal branches of inflorescences and exterior of flowers glabrous. Central Amazonia .......................................... 29. E. verticillata


33. Hairs on branchlets greyish white or yellow. Inflorescence axes and exterior of flowers densely pubescent. 34. Indument on vegetative surfaces greyish white. Flowers pedicellate,

distant along terminal cymes. Central Amazonia to E Andes .......... 28. E. bracteata 34. Indument on vegetative surfaces yellowish. Flowers subsessile,

clustered. 35. Laminae plane and cupules densely pubescent outside, the surface

obscured. Guianas, Amazonia, and E Andean slopes ................ 33. E. chalisea 35. Laminae bullate or plane, but cupules always glabrous outside. SE

Brazil ......................................... 34. E. glomerata 20. Leaves and inflorescences arranged evenly along branchlets. Inflorescence bracts and bracteoles

caducous by anthesis. 36. Terminal buds glabrous except for a distal ciliate fringe of hairs. Young leaves and branchlets

completely glabrous. Guianas to E Andes ................................................................... 14 . E. pyriformis 36. Terminal buds densely pubescent, the surface completely covered. Young leaves and/or

branchlets pubescent. 37. Indument on leaves below closely appressed to the surface.

38. Branchlets midway along flush sparsely pubescent, the surface clearly visible. 39. Basalmost secondaries asymmetric, one or both merging with leaf margin near

base or emerging from junction of midrib and leaf base. 40. Flowers campanulate, the tepals erect, glabrous inside. Central Amazonia to

E Andes ................................................ 8. E. metallica 40. Flowers rotate, the tepals spreading horizontally, densely pubescent inside.

San Martin, Peru ................................................ E. metallica aff sp. 2 39. Basalmost secondaries ? symmetric, emerging from midrib above leaf base.

41. Axils of secondary veins below with barbellate tufts of hairs. SW Amazonia and E Andean foothill ................ ..................................... 17. E. dysodantha

41. Axils of secondary veins below without barbellate tufts of hairs. 42. All staminal whorls stipitate, the filaments distinct and narrower than

anthers. Basal glands large, filling the space between stamens. Inner surface of tepals glabrous. 43. Midrib and secondary veins immersed below; tertiary and higher-

order veins immersed above; small trees or shrubs. NE South America ......................................... 1. E. punctulata

43. Midrib and secondary veins prominent below; tertiary and higher- order veins prominent above. Large trees to 30 m. Central Amazonia, Brazil ......... ................................ 2. E. coriacea

42. Outer whorls of stamens subsessile, the filaments inconspicuous, or if obvious as broad as or broader than anthers. Basal glands inconspicuous. Inner surface of tepals pubescent. 44. Flowers depressed-globose, the tepals inflexed, revealing only a

small terminal pore at anthesis. 45. Tertiary veins straight or forked between adjacent secondaries.

Flowers ca. 2 mm diam. Central Amazonia and uplands from Peru to Costa Rica ..................................... 15. E. formosa

45. Tertiary veins reticulating between adjacent secondaries. Flowers 5-7 mm diam. Cajamarca. Peru .......................... 16. E. paradoxa

44. Flowers hypocrateriform, campanulate, or rotate, the tepals ascending, horizontally spreading, or reflexed at anthesis. 46. Anthers of whorl I & II stamens reniform, the apex truncate to

rounded. Locelli suborbicular, minute, just below apex. W Amazonia to E Andes .............. ....................... 4. E. rubriflora

46. Anthers of whorl I & II stamens ovate, the apex apiculate. Locelli obliquely hemispherical, occupying entire anther. 47. Tertiaries laxly reticulating between adjacent secondaries.

48. Indument on branchlets rust brown. W Amazonia .............. ................... 13. E. mishuyacensis

48. Indument silvery grey. Cerro Jefe, Panama .......... 11. E. jefensis

20 FLORA NEOTROPICA

47. Tertiaries straight or once-forked between adjacent secondaries. 49. Receptacle constricted below tepals. Lowland and

lower montane slopes on Pacific side of Andes in Ecuador and Colombia, Panama ..................... 10. E. browniana

49. Receptacle gradually merging with tepals. Antioquia, Colombia ......... ............ 12. E. colombiana

38. Branchlets midway along flush densely pubescent, the surface barely or not visible. 50. Basalmost secondaries asymmetric, one or both merging with leaf margin near

base or emerging from junction of midrib and leaf base. 51. Flowers campanulate, the tepals erect at anthesis, glabrous inside. Stamens

included. Central Amazonia to E Andes ........................................ 8. E. metallica 51. Flowers rotate or hypocrateriform, the tepals spreading at anthesis, densely

pubescent inside. Stamens exserted. 52. All anthers two-locellate; whorls I and II stamens stipitate with S-curved

filaments. W Amazonia to E Andean slopes .................................... 53. E. klugii 52. Whorl III anthers four-locellate; whorls I and II stamens sessile,

petaloid. Manaus, Brazil ..................................... Rhodostemonodaphne recurva 50. Basal secondaries ? symmetric, the basalmost pairs emerging from midrib above

leaf base. 53. Leaves triplinerved. The lowermost pair of secondaries subopposite shortly

above the leaf base, the others emerging after midlamina or absent. 54. Anthers of whorl I & II stamens truncate to emarginate above. W

Amazonia .................................... 58. E. acuminata 54. Anthers of whorl I & II stamens apiculate, the connective prolonged

above. E Andes to W Amazonia ..................................... 59. E. krukovii 53. Leaves penninerved. The secondaries ? uniformly spaced along midrib.

55. Flowers obconical or urceolate, the tepals incurved or erect at anthesis. Stamens included. Cupules seemingly hemispherical when seen from outside but actually shallow, the base thickened. W Amazonia to E Andes ..................................... 55. E. robusta

55. Flowers rotate or hypocrateriform, the tepals patent or reflexed at anthesis. Stamens exserted. Cupules actually deeply hemispherical or if shallow, this not due to a thickened base. 56. Apex of whorl I & II anthers flat, emarginate, or with a hornlike

lobe above each locule. 57. Tepals chartaceous to membranaceous, sparsely pale-pilose

inside. SE Brazil, lower montane Andes to Panama, W Amazonia ............................. 56. E. paniculata

57. Tepals fleshy, densely grey-tomentose inside. E Venezuelan Highlands ............................. 60. E. nilssonii

56. Apex of whorl I & II anthers apiculate, prolonged between locelli. 58. Whorl I & II stamens stipitate, the filaments narrower than

anthers, S-shaped, tomentose. E of Andes in South America ................................................................................................. 53. E. klugii

58. Whorl I & II stamens sessile, the filaments as broad as anthers, densely papillose. W of Andes from Ecuador to Panama ................................................................. 10. E. browniana

37. Indument on leaves below either erect or at least ascending at angle of ca. 300.

59. Indument on branchlets midway along flush sparse and scattered, the surface clearly visible. 60. Leaves triplinerved, the lowermost pair of secondaries subopposite shortly above

the leaf base, the others appearing after midlamina or absent. 61. Tertiary and higher-order veins immersed above. Secondary venation

brochidodromous. Inner surface of tepals and filaments of outer whorls of stamens pubescent. Guianas to E Andes ...................................................... 57. E. gracilis

61. Tertiary and higher-order veins prominent above. Secondary venation eucamptodromous. Inner surface of tepals and filaments of outer whorls of stamens glabrous. W Amazonia ............................. ..................... 58. E. acuminata


60. Leaves penninerved, the secondary veins ? uniformly spaced along midrib. 62. Axils of secondary veins below with barbellate tufts of hairs. Apex of outer

anthers apiculate. Cajamarca, Peru ................................. .............. 18. E. tomentosa 62. Axils of secondary veins below without barbellate tufts of hairs. Apex of

outer anthers rounded, truncate, or emarginate. 63. Leaves densely pubescent below. Outer stamens stipitate, the filaments

distinct; locelli filling anthers. SE Brazil, lower montane Andes to Panama, W Amazonia ............. ............................... 56. E. paniculata

63. Leaves subglabrous below. Outer stamens petaloid, sessile, the filaments as wide as or wider than anthers; locelli minute, occupying only upper half of anthers. 64. Branchlets greenish yellow; leaves oblong with caudate tips.

Secondaries loop-connected. Central to E Amazonia .......... 6. E. longicaudata 64. Branchlets dark brown; leaves obovate to elliptic, tips acuminate.

Basal secondaries ending freely, the distal pairs weakly loop- connected. W Amazonia to E Andes ........................................ 4. E. rubriflora

59. Indument on branchlets midway along flush dense, the surface barely visible or concealed. 65. Lamina narrowly elliptic or lanceolate, ca. 10 times longer than broad, strongly

bullate. Secondaries more than 50. Hairs on all surfaces ca. 2 mm long. SW Amazonia ................................................... 3. E. bullata

65. Lamina ovate to obovate, less than 3 times longer than broad, plane. Secondaries less than 15. Hairs on all surfaces less than 1 mm long. 66. Leaf surface areolate above, the depressions corresponding to the fourth-

order reticulations. All anthers two-locellate. 67. Branchlets reddish tomentose. Inner tepals densely pubescent, the

surface not visible. Tepuis of Venezuela and Colombia ................... 41. E. vinotincta 67. Branchlets greenish yellow tomentose. Inner tepals sparsely pubescent,

the surface clearly visible. Andes of Ecuador and Peru .................... 42. E. oreocola 66. Leaf surface smooth or minutely punctulate above. If with areolate

depressions, then at least whorl III anthers four-locellate. 68. Flowers globose, the tepals erect or inflexed at anthesis. Stamens

included. Amazonia, Guianas, E Andes .................................... 43. E. szyszylowiczii 68. Flowers campanulate to rotate, the tepals ascending to spreading at

anthesis. Stamens exserted. 69. Whorl I and II anthers truncate, rounded or emarginate above;

connectives broad above, level with, or reduced between suborbicular locelli. 70. Outer stamens stipitate, the filaments much narrower than

anthers, locelli large, filling anther. 71. Leaves triplinerved, the lowermost pair of secondaries

subopposite shortly above the leaf base, the others emerging after midlamina. Guianas to E Andes .......... 57. E. gracilis

71. Leaves penninerved, the secondaries ? evenly spaced. 72. Flowers infundibuliform; tepals ascending at

anthesis. Cupule margins entire, the tepals or tepal- bases not persisting. Pacific coast of Colombia

.......................................................................... 38 . E . xeram pela 72. Flowers rotate; tepals spreading at anthesis. Cupule

margins lobed, the tepals or tepal-bases persisting. 73. Whorl III anthers four-locellate. Central to W

Amazonia. Flooded forest ........................... 35. E. anomala 73. All anthers two-locellate. SE Brazil, throughout

Andes to Panama, W Amazonia. Terra firme ................................................................. 56. E. paniculata

70. Outer stamens (sub-)sessile, the filaments barely or not narrower than anthers; locelli small, occupying distal half of anther.

22 FLORA NEOTROPICA

74. Branchlets rusty or grey-tomentose. Tertiaries immersed above. Amazon basin ............... ..................... 7. E. sprucei

74. Branchlets rusty tomentellose. Tertiaries raised above. 75. Laminae plane with flat margins. All anthers two-

locellate. Amazonian Ecuador .......................... 5. E. ferruginosa 75. Laminae with revolute margins. Whorl III anthers

four-locellate. French Guiana .............................................. Rhodostemonodaphne revolutifolia

69. Whorl I and II anthers apiculate above; connectives prolonged between obliquely hemispherical locelli. 76. Tertiaries immersed above. Flowers sparsely pubescent

outside, the surface visible. Guianas, W Amazonia, E Andes .......................................................................................... 37. E. canescens

76. Tertiaries raised above. Flowers densely pubescent outside, the surface barely or not visible. 77. Leaves obovate, the apex blunt, rounded. Indument on

branchlets dark red. W Amazonia and E Andes ............. 40. E. rubra 77. Leaves ovate to elliptic-oblong, acuminate. Indument on

branchlets brown to yellowish. 78. Branchlets tomentellose, the hairs erect. Filaments

of whorl I and II stamens narrower than anthers. W Amazonian lowlands ............................... 39. E. citriodora

78. Branchlets subsericeous, the hairs appressed to ascending. Filaments of whorl I and II stamens equal anthers. W Amazonia and lower montane E Andean slopes ........ 45. E. lorastemon

1. Endlicheria punctulata (Mez) C. K. Allen, Mem. New York Bot. Gard. 15: 68. 1966. Ocotea punctulata Mez, Jahrb. Konigl. Bot. Gart. Berlin. 5. 379. 1889. Type. French Guiana. Without locality and date, Melinon 204 (lectotype, designated by Allen, 1966: P; isolectotype: P).

Treelets or shrubs to 4 m. Branchlets slender, midway along flush 1.5-2.5 mm diam., angular, sparsely strigillose, the surface clearly exposed, reddish brown, the hairs short, to 0.2 mm, straight, appressed, greyish white; terminal buds slender, 3 X 0.6 mm, silvery sericeous. Leaves alternate, widely and evenly spaced along current flush; petioles slender, to 1.5 X

0.1 cm, canaliculate, the indument as on branchlets; laminae stiff-chartaceous to coriaceous, plane to subbullate, obovate, 7-11 X 3-5 cm, the base attenuate, the apex caudate for up to 1.5 cm, the margins minutely recurved throughout; upper surface olive green to light brown, minutely punctulate, the midrib prominent, the higher-order venation immersed; lower surface glabrous, all vein orders immersed, the midrib dark, conspicuous against the lighter lamina; secondary veins 5-6 per side, ? evenly spaced, slightly more distant around midlamina, ascending at 50-60? (more obtusely towards apex), arcuate, the distal pairs loop-connected; tertiaries laxly reticulating between secondaries. Staminate inflorescences evenly spaced

along current flush in the axils of foliage leaves, to 6 cm long with 6 lateral branches, branch orders 2-3, the highest order dichasial, lax, the flowers distant, the axes sparsely grey-strigillose; bracts and bracteoles caducous by anthesis, triangular, with indument as on axes; pedicels terete, to 2 mm long, those supporting secondary flowers slightly shorter. Flowers tubiform, sparsely grey-strigillose outside; receptacle infundibuliform, 0.6 x 1 mm, densely rusty tomentose inside. Tepals chartaceous, triangular, 0.6 X 0.3 mm, erect, surrounding androecium at anthesis, the outer and inner surfaces sparsely grey-strigillose, the margins and apex inside sparsely papillose. Stamens of whorls I and II 0.6 mm tall, stipitate, the anthers ovate, 0.4 x 0.3 mm, glabrous, the apex rounded, the connectives broad above the 2 locelli, these suborbicular, introrse, the filaments laminar, narrower than anthers, sparsely grey-pilose; whorl III stamens stipitate, 0.7 mm tall, the anthers oblong, 0.4 x 0.3 mm, erect, locelli 2, extrorse-latrorse, the filaments narrower than anthers, slender, laminar, sparsely grey- pilose, the basal glands sessile, globose, relatively large, filling the space between filaments of inner and outer whorls; whorl IV wanting; pistillode fusiform. Pistillate inflorescences unknown. Fruits borne on claviform pedicels of up to 1 X 0.3 cm; cupules hemispherical, to 0.5 X 1 cm, glabrous inside and outside, the margins entire; drupes ellipsoid, to 1.5 X 1 cm.


Distribution (Fig. 4) and ecology. Understory treelets or shrubs of lowlands and lower montane forests (200-800 m) in French Guiana, adjacent Suri- name, and the state of Amapd in Brazil. Flowering August and September, fruiting material collected from January to October.

Representative specimens examined. SURINAME. Vic. of Ulemarie R. ca. 150 km upstream from confluence with Litani R., 175 m, 18 Apr 1998 (fr), Hammel etal. 21557 (MO).

FRENCH GUIANA. Saul, Monts La Fumee, 200-400 m, 25 Sep 1982 (fl 6), Mori et al. 14995 (HBG, NY, US); Tumac Humac, Mitiraka, 1 Sep 1972 (fl 6), de Granville 1448 (MO, P); Route de Belizon, Montague Tortue, km 25, 23 Apr 1992 (fr) Acevedo et al. 4834 (F, MO).

BRAZIL. AMAPA: Rio Araguari, Porto Platon, 21 Sep 1961 (fl d), Pires et al. 51168 (B, HBG, K, NY, US); Rio Iaue, first cachoeira, 22 Aug 1960 (fl d), Irwin & Westra 47706 (GH, NY).

Local name. Brazil: louro.

Endlicheria punctulata is distanced from most species of Endlicheria by its concolorous leaves, pauciflorous inflorescences, and distinct filaments in all three staminal whorls. This combination of vegetative and floral features is otherwise only found in E. coriacea, from which E. punctulata is readily distinguished by the minute pin-prick pattern on its upper leaf surface and the immersed venation below. These two species are isolated in Endlicheria and apparently more closely related to species in the Ocotea cernua species group (see Generic Delimitation and Species Groups, above).

Mez (1889) cited five fruiting collections (M6linon 204, 216, 227 & 551, and Jenman 1702) in the protologue of the basionym of this species, Ocotea punctulata. I have seen the Paris (P) duplicates of Me'linon 204, 216, and 551, and the Copenhagen (C) sheet of Melinon 227. All are from French Guiana and are clearly conspecific. However, Jenman 1702, collected in Guyana and seen at BRG, likely deposited also at

--------- - 4- ------

. . . . . . . . .. .....

4"

10 ~ ~ f

\' *'4~~~~~~~~~~~~ 4~~ -- -- -- -- .. .. .. .

~ ~ ~ -. ' ~?- A

A~~~~~~~~~~~~~~~~~~~~~~~~x ~ ~ ~ ~ ~ ~ ~ ~ *~*

FIG. 4. Distribution of Endlicheria punctulata, E. coriacea, E. bullata, and E. rubriflora.

24 FLORA NEOTROPICA

K, belongs to either Ocotea pauciflora or 0. cernua. Both are very similar to E. punctulata, but with smooth rather than punctulate upper leaf surfaces. Al- len's (1966) lectotypification on Melinon 204 pre- vents future confusion.

2. Endlicheria coriacea Chanderbali, sp. nov. Type. Brazil. Amazonas: Manaus, Agropecuairio, Re- serve 1501 (km 41) of the WWF/INPA Biological Dynamics of Forest Fragments Project, 50-150 m, 21 Sep 1989 (fl d), Lepsch da Cunha et al. 391 (holotype: NY; isotypes: NY, MO). Fig. 5

Endlicheriae punctulatae affinis, inflorescentiis et floribus similis, sed statura permajore et foliis subtus nervis prominentibus recedit.

Trees to 30 m. Branchlets slender, midway along flush 2-3 mm diam., distally weakly angular soon terete, sparsely strigillose, the surface clearly exposed, dark brown, the hairs short, to 0.1 mm, straight, appressed, greyish white; terminal buds slender, 3 X 0.6 mm, silvery sericeous. Leaves alternate, widely and evenly spaced along current flush; petioles slender, to 1.5 X 0.1 cm, canaliculate, glabrous; laminae coriaceous, plane, obovate, 4-6 X 8-12 cm, the base attenuate, the apex acute, narrowly acuminate, or caudate for up to 1 cm, the margins minutely recurved throughout; upper surface light to dark brown, waxy, the primary to fourth-order veins raised, their prominence decreasing with rank; lower surface glabrous, all vein orders raised, their prominence decreasing with rank; secondary veins 7-9 per side, ? evenly spaced, slightly more distant around midlamina, patent, diverging at 70-85? (more obtusely towards apex), arching after midcourse, weakly brochidodromous; tertiaries once-forked or laxly reticulating between secondaries. Staminate inflorescences evenly spaced along current flush in the axils of foliage leaves, to 6 cm long with 6 lateral branches, branch orders 2-3, the highest order dichasial, lax, the flowers distant, the axes sparsely grey-strigillose; bracts and bracteoles caducous by anthesis, triangular, the indument as on axes; pedicels terete, to 2 mm long, those supporting secondary flowers slightly shorter. Flowers tubiform, sparsely grey-strigillose outside; receptacle shallowly infundibuliform, 0.6 X 1 mm, densely silvery pilose inside. Tepals membranaceous, ovate, 0.6 X 0.3 mm, ascending, the androecium sur- rounded at anthesis, the outer and inner surfaces sparsely grey-strigillose, the margins sparsely papillose. Stamens of whorls I and 11 0.6 mm tall, stipitate, the anthers depressed-ovate, 0.4 X 0.5 mm, glabrous, the apex rounded, the connectives broad above the 2

locelli, these suborbicular, introrse, the filaments laminar, narrower than anthers, sparsely grey-pilose; whorl III stamens stipitate, 0.7 mm tall, the anthers oblong, 0.4 X 0.3 mm, erect, locelli 2, extrorse- latrorse, the filaments narrower than anthers, slender, laminar, sparsely grey-pilose, the basal glands sessile, globose, relatively large, filling the space between filaments of inner and outer whorls; whorl IV wanting; pistillode filiform. Pistillate inflorescence with indument, color, and branching as in staminate plants, the flowers similar in size and shape; stamens sterile, smaller; ovary glabrous; style slender, distinct from ovary; stigma tri-lobed, 0.3 mm diam. Mature fruits unknown, immature fruits borne on terete pedicels, the cupules glabrous inside and outside, the margins entire.

Distribution (Fig. 4) and ecology. Tall trees known only from lowland forests (50-150 m) on well- drained sand and clay soils around Manaus, Brazil. Flowering from July to September with fruit set ini- tiated by September.

Additional specimens examined. BRAZIL. AmA-

ZONAS: Estrada Manaus-Itacoatiara, km 135, 22 Aug 1973, (fl Y ), Rodrigues et al. 8523 (HBG, INPA, US), 10 Jul 1974 (fl Y), Rodrigues & Loureira 9454 (F, INPA, MO), 13 Sep 1972 (fr), Pires & Coelho 175 (INPA); Reserva 2107 do Projecto Dinamica Biologica dos Fragmentos Florestais, Arv. No. 388, 30 Jul 1982 (fl Y), Mackenzie et al. INPAI WWF 2107.388 (MO).

Local name. Rodrigues et al. 8523 suggest "Louro pirarucu" is the local name, but this may have been misapplied. According to Vicentini et al. (1999), "Louro pirarucu" refers to Licaria cannella (Meisn.) Kosterm. and material of Endlicheria coriacea can be easily mistaken for this species. The strong smell of fish from the inner bark of L. cannella, surely responsible for association with the pirarucu, or ara- paima, is yet to be reported for species of Endlicheria.

Endlicheria coriacea is distinguishable from E. punctulata by its smooth upper leaf surface, prominent venation below, and large stature, but flowers of the two are indistinguishable. Both are apparently two-locellate members of a species group in Ocotea centered around 0. cernua (see Relationships, above). Among the treelets and shrubs typical of this group, E. coriacea and Rhodostemonodaphne elephantopus are remarkable for their arborescence. These two species differ in locelli number and, with E. coriacea known only from central Amazonia and R. elephantopus from French Guiana, they may be allopatric. However, they are vegetatively indistinguishable.

FIG. 5. Endlicheria coriacea (Lepsch da Cunha et al. 391). A. Habit. B. Single leaf showing venation. C. Flower with facing tepal turned down to reveal androecium. D. Flower ls. E. Whorl I stamen seen from within. F. Whorl III stamen seen from without.

26 FLORA NEOTROPICA

3. Endlicheria bullata Ducke, Arch. Jard. Bot. Rio de Janeiro 4: 190. 1925. Type. Brazil. Amazonas: Bom Logar, 25 Jul 1903 (fl d), Hubers.n. R 18359 (holotype: R-n.v.; isotypes: B-n.v., U).

Shrubs to 6 m. Branchlets slender, midway along flush 2-3 mm diam., terete, densely rusty to reddish- hirsute, the surface barely visible through indument cover, the hairs long, to 2.5 mm, straight, rigidly erect; terminal buds densely rusty pubescent, the hairs as on branchlets, ascending. Leaves alternate, widely and evenly spaced along current flush; petioles slender, to 1 X 0.1 cm, terete, the indument as on branchlets; laminae chartaceous, bullate, lanceolate, 15- 35 X 3-4 cm, the base acute to rounded, the apex acute, acuminate for up to 2 cm, the margins minutely recurved throughout; upper surface light green, waxy, the midrib prominulous, the secondaries flat, the tertiaries immersed; lower surface densely rusty hirsute, the hairs as on branchlets, denser on main veins, all vein orders raised, their prominence decreasing with rank; secondary veins numerous, 20-35 per side, +

evenly spaced, slightly more distant around midlamina, diverging at 850, abruptly ascending near margin, brochidodromous; tertiaries laxly reticulating between secondaries. Staminate inflorescences evenly spaced along current flush in the axils of foliage leaves, to 8 cm long with 8 lateral branches, branch orders 3-4, the highest order dichasial, lax, the flowers distant, the axes densely rusty hirsute; bracts and bracteoles caducous by anthesis, lanceolate, the indument as on axes; pedicels terete, to 5 mm long, those supporting secondary flowers slightly shorter. Flowers rotate, 8 mm diam., rusty tomentose outside, the indument sparser towards tepals; receptacle infundibuliform, 2 X 2 mm, rusty velutinous inside. Tepals fleshy, obovate, 3 X 2 mm (the inner whorl slightly narrower), spreading at anthesis, the inner surface glabrous. Stamens of whorls I and II sessile, stout, 1.3 X 1.3 mm, the anthers depressed-ovate, 0.6 X 1 mm, glabrous, the apex rounded, the connectives slightly incurved over the 2 locelli, these suborbicular, just below apex, minute, introrse, the filaments fleshy, broader than anthers, the base rusty tomentose; whorl III stamens stout, sessile, 0.8 mm tall, the anthers depressed-oblong, 0.4 X 1 mm, erect, locelli 2, extrorse-latrorse, the filaments broader than anthers, fleshy, the base rusty tomentose, the basal glands sessile, relatively large, globose-apiculate; whorl IV wanting; pistillode fusiform. Pistillate inflorescence not seen; detached pistillate flowers similar in size and shape to staminate flowers; stamens sterile, smaller; ovary glabrous, ovoid; style stout, weakly distin-

guished from ovary; stigma discoid, 0.8 mm diam. Fruits unknown.

Distribution (Fig. 4) and ecology. Small shrubs from terra firme forest of SW Amazonia in Brazil. Flowering in July and September.

Additional specimens examined. BRAZIL. ACRE: Near mouth Rio Macuahan (tributary of Rio Yaco), 3 Sep 1933 (fl d & Y?), Krukoff 5780 (A, G, K, S, U, US).

Endlicheria bullata is a poorly known but highly distinctive species. Nowhere else in Endlicheria are narrow bullate leaves with numerous secondaries and stiff hirsute vestiture combined with rotate flowers and fleshy sessile stamens. Similar flowers combine with plane laminae in E. ferruginosa and E. rubriflora, while similar leaves and vestiture combine with campanulate flowers and stipitate stamens in E. glomerata. However, outside of Endlicheria these characters all appear in Rhodostemonodaphne scandens Madrinian, albeit with four-locellate anthers.

Although only two collections are cited here (the type and Krukoff 5780), they represent three individuals. Type material is clearly staminate since the protologue is based on a staminate plant, and at least the isotype deposited in Utrecht (U) shows staminate inflorescences attached to twigs. Of Krukoff 5780, sheets at Geneva (G), Stockholm (S), and Utrecht (U) lack inflorescences, while that at the Ar- nold Herbarium (A) shows intact staminate inflorescences, and the Kew (K) sheet has detached staminate flowers. However, the US duplicate of Krukoff 5780 has detached pistillate flowers that appear to be con-specific with staminate ones. Since monoecy is unknown in Lauraceae, the source of these pistillate flowers is unknown unless Krukoff 5780 is a mixed collection.

4. Endlicheria rubriflora Mez, Jahrb. Konigl. Bot. Gart. Berlin 5: 494. 1889. Type. Colombia. With- out locality and date, Triana 1032 (holotype: P). Aniba reticulata A. C. Sm., Bull. Torrey Bot. Club 58:

99. 1931. Type. Peru. Loreto: Alto Amazonas, Yu- rimaguas, lower Rio Huallaga, 138 m, 22 Aug-9 Sep 1929 (fl Y), Killip & Smith 28050 (holotype: NY; isotype: US).

Endlicheria trianae 0. C. Schmidt, Repert. Spec. Nov. Regni Veg. 31: 175. 1933. Type. Colombia. Andes de Antioquia, Triana 2040-1 (holotype: K-n.v.).

Endlicheria wurdackiana C. K. Allen, Mem. New York Bot. Gard. 10(5): 67. 1964. Type. Venezuela. Ama- zonas: Rio Siapa between Raudal Gallineta and Salte Gallineta, 130-150 m, 23 Jul 1959 (fr), Wurdack & Adderley 43592 (holotype: NY; isotypes: GH, U).


Trees to 15 m. Branchlets slender, midway along flush 2-3 mm diam., distally weakly angular, soon terete, sparsely pubescent, the surface clearly exposed, dark to reddish brown, the hairs relatively short, to 0.3 mm, straight, appressed to ascending, yellowish green; terminal buds slender, 2 X 0.6 cm, densely pubescent, the hairs as on branchlets, appressed. Leaves alternate, widely and evenly spaced along current flush; petioles slender, to 1.5 X 0.2 cm, terete, the indument as on branchlets; laminae chartaceous, plane, elliptic to obovate, 7-20 X 3-8 cm, the base obtuse to rounded, briefly attenuate, the apex obtuse to acute, acuminate for up to 2 cm, the margins flat throughout; upper surface greyish green, waxy, the midrib sunken towards petiole, otherwise primary to fourth-order veins raised, their prominence decreasing with rank; lower surface sparsely pubescent, the hairs appressed, ascending, or erect, slightly denser on main veins, all vein orders raised, their prominence decreasing with rank; secondary veins 6- 9 per side, + evenly spaced, slightly more distant around midlamina, patent, diverging at 70-85O (more obtusely towards apex), arcuate, distal pairs loop- connected; tertiaries laxly reticulating between secondaries. Staminate inflorescences evenly spaced along current flush in the axils of foliage leaves, to 10 cm long with 8 lateral branches, branch orders 3-4, the highest order dichasial, lax, the flowers distant, the axes sparsely pubescent, the hairs greenish yellow to grey, ascending, denser on pedicels; bracts and bracteoles caducous by anthesis, lanceolate, densely pubescent, the hairs as on axes, appressed; pedicels terete, to 2 mm long, those supporting secondary flowers slightly shorter. Flowers broadly rotate, 5 mm diam., sparingly rusty strigillose outside; receptacle shallowly infundibuliform, 0.6 X 1.3 mm, densely yellowish tomentose inside. Tepals fleshy, broadly ovate, 2 X 1.6 mm, spreading to recurved at anthesis, the inner surface and margins densely rusty to reddish papillose. Stamens of whorls I and II broadly stipitate, 0.8 m tall, the anthers reniform, 0.6 X 0.8 mm, whorl II slightly narrower, glabrous, the apex rounded, the connectives slightly incurved over the 2 locelli, these suborbicular, just below apex, minute, introrse, the filaments laminar, 0.2 X 0.6 mm, glabrous; whorl III stamens stout, sessile, 0.6 mm tall, the anthers transversely oblong, 0.3 X 0.6 mm, erect, locelli 2, extrorse-latrorse, the filaments as broad as or broader than anthers, columnar, glabrous, the basal glands sessile, globose, relatively large, filling the space between inner and outer whorls of stamens; whorl IV wanting; pistillode filiform. Pistillate inflorescence with indument and color as in staminate plants, but

shorter and with fewer lateral branches, the flowers slightly deeper; stamens sterile, smaller, the anthers auriculate; ovary glabrous, ovoid; style stout, weakly distinguished from ovary; stigma almost sessile, discoid, 0.6 mm diam. Fruits borne on stout claviform pedicels of up to 3 X 0.5 cm; cupules discoid to reflexed, to 4 cm diam., glabrous outside and inside, the margins lobed; drupes ellipsoid, to 3 X 2 cm.

Distribution (Fig. 4) and ecology. Small trees from western Amazonian lowlands and adjacent eastern Andean slopes at 100-1900 m. Found in both flooded and nonflooded soils. Flowering and fruiting throughout the year.

Local names. Colombia: jigua de mierda. Peru: palta amarilla, roble palta, roblecillo puchirin.

Representative specimens examined. COLOMBIA.

ANTiOQUIA: San Luis, Autopista Medellin-Bogota, 790 m, 15 Jan 1983 (fl d), Cogollo & Brand 394 (MO). CAUCA:

El Tambo, Vereda La Romelia, Parque Natural Munchique, 23 Aug 1994 (fr), Jarvenpaa 27 (COL). META: Villavicen- cio, Andes de Bogota, Jan 1856 (fr), Triana 2060 (K). QUIN-

DIO: Genova, 1870 m, 17 Mar 1988 (fl d), Arbelaez et al. 2490 (HBG, MO). TOLIMA: Mariquita, near Honda on Rio

Magdalena, s.d. (fr), Karsten s.n. (LE). VALLE: Sevilla, via Sevilla-La Raquelita, a orilla de la quebrada la Raquelita, 1500 m, 31 Jul 1985 (fr), Devia 1052 (MO).

VENEZUELA. APURE: Reserva Forestal San Camilo, 280-300 m, 27 Mar 1968 (fr), Steyermark et al. 101408 (HBG, NY). BARINAS: Pedraza, El Velador, 14 Dec 1954 (fl Y), Bernardi 1789 (G, NY).

ECUADOR. NAPO: Tena, Estaci6n Biol6gica Jatun Sa- cha, 400 m, 22 Sep 1989 (fl d), Palacios 4473 (G, MO, NY). ZAMORA-CHINCHIPE: Zamora, Jamboe Bajo, E border of Podocarpus National Park, 1100 m, 5 Nov 1996 (fl 6), Clarke et al. 3274 (MO).

PERU. MADRE DE DIOs: Tambopata, Cuzco Amaz6n-

ico, 200 m, 14 Jun 1989 (fl 6), Nuiiez et al. 10804 (MO). PASCO: Oxapampa, Iscozacin, 26 Jun 1986 (fl Y), Pariona & Sebastian 46 (F, MO); Palcazd, 300-600 m, 18 Nov 1985 (fr), Hartshorn et al. 2827 (MO).

BRAZIL. AMAZONAS: Humaitd, Porto Velho, km 30 da estrada Humaita-Labrea, 10 Jun 1982 (fl Y), Filho 82-7 N (HBG); Rio Curuquete, halfway between Cachoeiras Sao Paulo and Republica, 22 Jul 1971 (fl Y), Prance et al. 14498 (HBG, K, MO, NY).

Endlicheria rubriflora has green-drying leaves similar to those of E. longicaudata and E. pyriformis, but its rotate flowers with densely papillose tepals around a central hub of large globose glands and reddish stamens are unmistakable. The cupules of this species are also unmatched by congeners. The cupules appear to start off as shallow patelliform structures, and as fruits mature they flatten to a thick disc and ultimately the margins become recurved, becoming

28 FLORA NEOTROPICA

umbrellalike in aspect. Similar cupules are produced by Rhodostemonodaphne antioquensis (see also Mad- rinain, 1996b), the flowers of which would also be identical but for the four-locellate anthers. Besides the added pair of locelli in R. antioquensis, its ovate leaves with a more coriaceous texture help to separate it from E. rubriflora.

Lowland Amazonian collections, including the type of Aniba reticulata, have slightly smaller flowers, but otherwise Endlicheria rubriflora is morphologically uniform. The type of E. trianae is not available to me, but since Smith's (1933) description does not provide distinguishing characters, I follow Kos- termans's (1937) synonymy under E. rubriflora. The third synonym accepted here, E. wurdackiana, is based on a fruiting specimen that has the cupule and vegetative characters of E. rubrifiora.

5. Endlicheria ferruginosa Chanderbali, sp. nov. Type. Ecuador. Napo: La Joya de los Sachas, Parque Nacional Yasuni, Maxus highway and pipeline under construction, km 27, 230 m, 16 Aug 1993 (fl d), Dik 203 (holotype: MO; isotypes: COL, MO, QCNE). Fig. 6

A speciebus quas Kostermans sectionis Microlocellatae ascripsit ramulis ferrugineo-tomentellis distinguenda.

Trees to 12 m. Branchlets slender, midway along flush 2-3 cm diam., distally weakly angular, soon terete, densely tomentellose, the surface concealed by the indument cover, the hairs short, to 0.1 mm, straight, erect, rust brown; terminal buds plump, 4 X

2.5 mm, rusty sericeous. Leaves alternate, widely and evenly spaced along current flush; petioles slender, to 1.5 X 0.15 cm, semi-terete, the indument as on branchlets; laminae chartaceous, plane, obovate, 12-17 X

3-6 cm, the base acute to cuneate, briefly decurrent, the apex obtuse, acuminate for up to 2 cm, the margins flat; upper surface olive-brown, minutely punctulate, the primary to fourth-order veins raised, their prominence decreasing with rank; lower surface sparsely tomentellose, the hairs as on branchlets, appressed and denser on main veins, all vein orders raised, their prominence decreasing with rank; secondary veins 5-7 per side, ? evenly spaced, slightly more distant around midlamina, ascending at 50-60O

(more obtusely towards apex), arcuate, distal pairs loop-connected; tertiaries roughly horizontal, between secondaries straight to once-forked. Staminate inflorescences evenly spaced along current flush in the axils of foliage leaves, to 6 cm long with 8 lateral

branches, branch orders 3-4, the highest order dichasial, lax, the flowers distant, the axes sparsely pale rusty tomentellose, distalmost branches and pedicels more densely so; bracts and bracteoles caducous by anthesis, lanceolate, rusty sericeous; pedicels terete, to 4 mm long, those supporting secondary flowers slightly shorter. Flowers rotate, 5 mm diam., sparsely rusty strigillose outside; receptacle shallowly infundibuliform, 1 X 2 mm, densely rusty tomentose inside. Tepals fleshy, ligulate, 2 X 2 mm, spreading to recurved, the androecium exserted at anthesis, the outer surface sparsely rusty strigillose, the inner surface more densely so, the margins and apex inside rusty papillose. Stamens of whorls I and 11I1 mm tall, sessile, the anthers transversely oblong, 0.6 X 0.8 mm (whorl II slightly smaller), sparsely minutely papillose, the apex truncate, the connectives level with the 2 locelli, these minute, suborbicular, introrse, the filaments clavate, slightly narrower than anthers, rusty tomentose; whorl III stamens sessile, 1.3 mm tall, the anthers depressed-oblong, erect, locelli 2, extrorse- latrorse, the filaments broader than anthers, columnar, glabrous, the basal glands sessile, globose, apiculate, filling the space between inner and outer whorls of stamens; whorl IV wanting; pistillode wanting. Pis- tillate plants unknown.

Distribution (Fig. 7) and ecology. Known only from the type material taken from a small tree of terra firme lowland forest in Amazonian Ecuador found flowering in August.

The rather large rotate flowers with fleshy tepals and sessile stamens with minutely papillose indument are all characteristic of Rhodostemonodaphne, but two-locellate anthers assign this species to Endli- cheria. Therein, similar flowers are found in E. bullata and E. rubriflora, from both of which E. ferruginosa can be easily distinguished by its rusty tomentellose branchlets. As in those species, the transition to two-locellate anthers from the ancestral four- locellate condition is complete but in other respects the androecium is best matched by members of Rho- dostemonodaphne. In the case of Endlicheria ferruginosa it is Rhodostemonodaphne revolutifolia that comes closest. In staminate flowers of the latter, whorl I and II anthers are two-locellate and broadly petaloid, much like those of Endlicheriaferruginosa, but whorl III anthers are four-locellate, and in pistillate flowers all anthers are four-locellate (Madrifinan, 1996a). Even if pistillate plants of E. ferruginosa turn out to be four-locellate, this species has already advanced further in locelli reduction than Rhodostemonodaphne revolutifolia, and more likely in parallel than from a


co

FIG. 6. Endlicheniaferruginosa (Dik 203). A. Habit. B. Single leaf showing venation. C. Leaf base below. D. Rlower. E. Rlower 15. F. Whorl I stamen seen from within. G. Whorl III stamen seen from without.

30 FLORA NEOTROPICA

- K~~~~~~~~~A

W

W, ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ I"

.......... ---------- ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ , ~ .. . 4. - ~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~ r~~~~~~~~~~~~'~Y

FIG. 7. Distribution of Endlicheriaferruginosa, E longicaudata and E sprucei~~~~~~~~~~~~~~~~~~~.!.'

common ancestor. The indument of R. revolutifolia is much longer and more yellow in color, and the revolute leaves to which its name alludes contrast with the plane laminae of Endlicheriaferruginosa.

6. Endlicheria longicaudata (Ducke) Kosterm., Re- cueil Trav. Bot. Neerl. 34: 515. 1937. Hubero- daphne longicaudata Ducke, Arch. Jard. Bot. Rio de Janeiro 4: 191. 1925. Type. Brazil. Para: Rail- road between Belem and Braganca, near Santa Iz- abel, Jun 1908 (fl cd), Huber s.n. MG 9431 (lectotype, designated by Kostermans, 1937: R-n.v.; isolectotypes: S, U).

Treelets or shrubs to 5 m. Branchlets slender, midway along flush 1-1.5 mm diam., at first sharply angular, soon terete, sparsely pubescent, the surface clearly exposed, light or yellowish green, the hairs short, to 0.2 mm, straight, erect to ascending, pale or yellowish brown; terminal buds slender, 2 X 1 mm,

densely pubescent, the hairs yellowish green, appressed. Leaves alternate, widely and evenly spaced along current flush; petioles slender, to 2 X 0.2 cm, terete, the indument as on branchlets; laminae chartaceous, plane to subbullate, oblong, 10-15 X 4-7 cm, the base acute, attenuate, the apex obtuse, then abruptly caudate for up to 4 cm, the margins minutely recurved throughout; upper surface deep green, waxy, the primary to fourth-order veins raised, their prominence decreasing with rank, creamish yellow, conspicuous against the lamina; lower surface sparsely pubescent, the hairs scattered, erect, persisting only on the midvein, all vein orders raised, their prominence decreasing with rank; secondary veins 4-5 per side, ? evenly spaced, slightly more distant around midlamina, patent, diverging at 70-85' (more acutely towards apex), abruptly ascending after midcourse, brochidodromous; tertiaries laxly reticulating between secondaries. Staminate inflorescences evenly spaced along current flush in the axils of foliage


leaves, to 5 cm long with 4 lateral branches, branch orders 2-3, the highest order dichasial, lax, the flowers distant, the axes sparsely grey-tomentellose, the hairs 0.1 mm long, scattered, erect to ascending; bracts and bracteoles persistent at anthesis, lanceolate, sparsely grey-strigillose, the hairs as on axes, appressed; pedicels gradually increasing in diameter apically, to 2 mm long, those supporting secondary flowers slightly shorter. Flowers rotate, to 5 mm diam., sparsely grey-strigillose outside; receptacle shallow, infundibuliform, 0.5 X 1 mm, densely tawny tomentose inside. Tepals chartaceous, ligulate, 3 X

1.2 mm (the inner whorl slightly broader), horizontally spreading, the inner surface glabrous, the margins and apex inside lightly papillose. Stamens of whorls I and II ligulate, sessile, 1 X 0.6 mm, the anthers minute, transversely oblong, 0.1 X 0.3 mm, sterile (lacking) in whorl I, glabrous, the apex retuse, the connectives reduced between the 2 locelli, these minute, introrse, suborbicular, the filaments ligulate, glabrous; whorl III stamens slender, columnar, 1 X 0.3 mm, the anthers 0.3 X 0.3 mm, depressed-triquetrous, incurved, locelli 2, minute, introrse-latrorse, the filaments as broad as anthers, triquetrous, glabrous, the basal glands absent; whorl IV wanting; pistillode wanting. Pistillate inflorescence with indument, color, and branching as in staminate plants, the flowers similar in size and shape; stamens sterile, smaller; ovary glabrous, ovoid, 0.6 X 0.5 mm; style slender, distinct from ovary, about 0.6 mm long; stigma strongly tri- lobed, 0.7 mm diam., spreading above sterile androecium. Fruits borne on claviform pedicels of up to 2.5 X 0.5 cm; cupules patelliform, to 0.3 X 1 cm, glabrous outside, sparsely grey-strigose inside, the margins lobed, tepals persisting; drupes ellipsoid, to 2.5 X 1 cm.

Distribution (Fig. 7) and ecology. Small treelets or shrubs from terra firme lowland forests (50-200 m) in central to eastern Amazonia. Flowering June to October, fruiting August to June.

Representative specimens examined. BRAZIL. AMAPA: Canteiro de obras de Hidroeletrica de Balbina, Mar 1986 (fr), Cid Ferreira et al. 6713 (INPA, K, MO, NY). AMAZONAS: Vic. of Rio Uatuma, Rio Pitinga, Aug 1979 (fl 6), Cid Ferreira et al. 881 (F, HBG, INPA, MG, NY, R, US); Tapuruquara, Oct 1971 (fl d), Prance et al. 15817 (HBG, INPA, MG, NY). PARA: Belem, Bosque Municipal, Jun 1943 (fl 6), Ducke 1232 (MG, MO, NY, US); lands of Instituto Agron6mico do Norte, Jul 1944 (fl i), Silva 282 (F, K); Tome, Aqu, Rio Acara, Jan 1978 (fr), Nascimento 386 (F, HBG, MG, NY). RONDONIA: Sao Loren,o mines, Nov 1968 (fr), Prance et al. 8892 (HBG, INPA, NY, US);

trail N of Rio Madeira from 2 km below confluence with Rio Abuna, Nov 1968 (fr), Prance et al. 8343 (INPA, NY).

Local names and uses. Called "azywa'yw-pihun" by the Tembe Indians of Brazil. The wood is used to make axe handles and chairs and as a firewood.

Endlicheria longicaudata was described as the only species of Huberodaphne, a genus Ducke (1925) separated from Endlicheria by lack of basal glands in whorl III stamens and shallow cupules with persistent tepals. After Kostermans's (1937) treatment these characters appear in other species of Endlicheria, but the androecium of E. longicaudata is still unmatched therein. Its sessile outer stamens with small, distally positioned locelli are much thinner and more petaloid than others of the Microlocellata species group. In- deed, in some staminate flowers, whorl I stamens can be sterile with empty shallow locelli. Whorl III stamens are also remarkable. These are fleshy, account- ing for most of the androecial tissue, and anthers are incurved and locelli introrse-latrorse. This androecium is reminiscent of Dicypellium and Phylloste- monodaphne. However these genera have bisexual flowers and double-rimmed cupules, and are closely related to Licaria (Chanderbali et al., 2001), while ITS sequence data place Endlicheria longicaudata in the Endlicheria-Rhodostemonodaphne alliance (Fig. 2). Therein, close relationship with Rhodostemono- daphne scandens lacks statistical and morphological support. However, R. parvifolia Madriinan has shallow cupules with persistent tepals and rotate flowers with greenish sessile stamens very similar to those of Endlicheria longicaudata. Interestingly, both species are part of a weakly supported clade (Fig. 2), also with Rhodostemonodaphne scandens, Endlicheria pyriformis, and E. citriodora, which, given the morphological similarities between E. longicaudata and Rho- dostemonodaphne parvifolia, may have phylogenetic significance.

7. Endlicheria sprucei (Meisn.) Mez, Jahrb. Konigl. Bot. Gart. Berlin 5: 125. 1889. Goeppertia sprucei Meisn., DC. Prodr. 15(1): 172. 1864. Type. Brazil. Amazonas: Rio Uaupes, Nov 1852 (fl d), Spruce 2769 (lectotype, designated by Kostermans, 1937: K; isolectotypes: B-n.v., BM-n.v., E, G, K, NY- n.v., OXF-n.v., P).

Small trees or shrubs, 3-7 m. Branchlets slender, midway along flush 2-3 mm diam., distally weakly angular, soon terete, densely greyish green to rusty hirsute, the surface barely visible, the hairs relatively long, to 0.6 mm, straight, erect; terminal buds plump,

32 FLORA NEOTROPICA

1 X 0.6 mm, densely pubescent, the hairs as on branchlets, ascending. Leaves alternate, widely and evenly spaced along current flush; petioles slender, to 1 X 0.3 cm, terete, the indument as on branchlets; laminae chartaceous, plane, elliptic to obovate, 11-23 x 4-8 cm, the base obtuse, or acute, the apex acute, narrowly acuminate for up to 5 cm, the margins minutely recurved throughout; upper surface dull grey, minutely punctulate, the midrib prominulous, the secondaries flat, the tertiaries immersed, inconspicuous against the lamina; lower surface sparsely hirsute, the hairs as on branchlets, slightly denser on main veins, all vein orders raised, their prominence decreasing with rank; secondary veins 4-6 per side, ? evenly spaced, slightly more distant around midlamina, ascending at 50-60o (more obtusely around midlamina), arcuate, brochidodromous; tertiaries laxly reticulating between secondaries. Staminate inflorescences evenly spaced along current flush in the axils of foliage leaves, to 8 cm long with 5 lateral branches, branch orders 2-3, the highest order dichasial, lax, the flowers distant, the axes sparsely rusty to grey-hirsute; bracts and bracteoles caducous by anthesis, lanceolate, densely pubescent, the hairs as on axes, appressed; pedicels terete, to 1.3 mm long, those supporting secondary flowers slightly shorter. Flowers rotate, 3 mm diam., densely grey-pubescent outside, the hairs ascending; receptacle infundibuliform, 0.5 X 0.6 mm, densely grey-tomentose inside. Tepals chartaceous, ligulate, 1.2 X 0.6 mm (the inner whorl slightly broader), the inner surface densely grey- strigose, the indument soon restricted to the base, eventually lost, the margins and apex inside minutely papillose. Stamens of whorls I and II sessile, 0.7 mm tall, the anthers transversely oblong, 0.4 X 0.5 mm, glabrous, the apex truncate, the connectives level with the 2 locelli, these suborbicular, latrorse-introrse, the filaments fleshy, as broad as anthers, the base sparsely grey-pilose; whorl III stamens sessile, 0.8 mm tall, the anthers depressed-oblong, 0.3 X 0.4 mm, slightly bowed towards the outer whorls, locelli 2, extrorse- latrorse, the filaments as broad as anthers, clavate, tapering towards base; sparsely grey-pilose, the basal glands sessile, globose; whorl IV wanting; pistillode wanting. Pistillate inflorescence with indument, color, and branching as in staminate plants, the flowers similar in size and shape; stamens sterile, smaller; ovary glabrous, ovoid; style slender, distinct from ovary; stigma broadly tri-lobed, 1 mm diam. Fruits borne on claviform pedicels of up to 1.5 X 0.1 cm; cupules hemispherical, to 1 X 1.5 cm, glabrous inside and outside, the margins sharply lobed; drupes ellipsoid, to 2 X 1 cm.

Distribution (Fig. 7) and ecology. Small trees or shrubs from nonflooded forests throughout lowland Amazonia (50-300 m). Flowering from May to No- vember with fruits maturing by November and available to July of the following year.

Representative specimens examined. COLOMBIA. CAQUETA: Araracuara, 5 Nov 1991 (fl d), Duivenvoorden 738 (MO); Morelia, 150-300 m, 19 Oct 1941 (fl d), Snei- dern 1191 (COL).

ECUADOR. PASTAZA: Lorocachi, 200 m, 26 May 1980 (fr), Jaramillo et al. 31147 (K, MO, NY). SUCUMBiOS: Lago Agrio, Reserva Faunfstica Cuyabeno, Chiritza, 230 m, 13 Nov 1991 (fl d), Palacios et al. 8845 (MO, NY).

PERU. LORETO: Maynas, Mishuyacu, near Iquitos, 100 m, Oct-Nov 1929 (fl i), Klug 272 (F, NY, US); Estaci6n Biol6gica Rfo Blanco, 150 m, 16 Sep 1985 (fr), Vdsquez et al. 6742 (F, MO, NY); Rio Nanay, Mishana, 80-110 m, 30 Sep 1990 (fl d), Pipoly et al. 12633 (MO).

BRAZIL. AcRE: Cruzeiro do Sul, Estrada do Ale- manha, 4 Nov 1966 (fr), Prance et al. 3008 (HBG, INPA, NY); Rio Jurua & Rio Moa, Serra de Moa Village, 25 Apr 1971 (fr), Prance et al. 12102 (HBG, INPA, NY). AMA- ZONAS: Agropecuario, Fazenda Dimona of WWF/INPA MCS Project, 50-125 m, 23 Oct 1988 (fl d), Boom & Pa- checo 8511 (INPA, MO); Humaita, Estrada Humaita-La- brea, km 70, 15 Jun 1982 (fl Y ), Teixeira et al. 1025 (INPA, K, MG, MO, NY). MATO GROSSO: Novo Aripuana, Rod. do Estanho km 120, 21 Apr 1985 (fr), Cid Ferreira 5682 (INPA, MO, NY, US). PAR": Rio Mapuera, Cach. de Ma- dame e Cache de Ilhas, 15 Aug 1986 (fl 6), Cid Ferreira et al. 7776 (F, INPA, K, MO, NY, US).

Local name. Peru: muena.

Among species of Endlicheria with sessile outer stamens, E. sprucei alone has a dense pin-prick pattern (minutely punctulate) and immersed tertiary venation on the upper surface of greyish green drying leaves. Elsewhere in Endlicheria, the leaves of E. gracilis are similar in color, texture, indument, and surface features, but ovate and triplinerved.

Indument color varies from greyish green to rusty red, but Endlicheria sprucei is otherwise homogeneous. Mez (1889) and Kostermans (1937) both reported that all stamens are provided with a pair of basal glands; however, as in most species of Laura- ceae, and all of Endlicheria except E. vinotincta, only the inner stamens (whorl III) are thus equipped. Fil- aments of whorl I (Fig. 1B) and II stamens are indeed fleshy, but the tissue does not appear to be glandular, nor does it appear to represent a separate structure.

Mez (1889) assigned Endlicheria sprucei to subgen. Ampelodaphne and ITS sequence data suggests close relationship with one of its members, E. reflectens (Fig. 2). As discussed above (Generic Delimita- tion and Species Groups), E. sprucei may occupy an


intermediate position between the Ampelodaphne species group and species of Rhodostemonodaphne, including R. crenaticupula. Fruiting material of the latter has been mistaken for Endlicheria sprucei, but its upper leaf surface is traversed by a reticulate pattern of prominent tertiary veins.

8. Endlicheria metallica Kosterm., Recueil Trav. Bot. Neerl. 34: 543. 1937. Type. Brazil. Ama- zonas: Near mouth of Rio Embira (tributary of Rio Tarauaca), 19 Jun 1933 (fl Y), Krukoff 4932 (holotype: NY; isotypes: A, K-n.v., MO, S, U).

Trees to 35 m. Branchlets stout, midway along flush 3-5 mm diam., striate, angular, silvery sericeous, the surface barely visible to concealed by the indument cover, the hairs short, to 0.3 mm, straight, appressed; terminal buds plump, 3 X 2 mm, silvery sericeous. Leaves alternate, widely and evenly spaced along current flush; petioles slender, to 3 X 0.3 cm, semi-terete, the indument as on branchlets; laminae coriaceous to chartaceous, plane, elliptic to oblong, 11-20 X 4-9 cm, the base acute to obtuse, briefly decurrent, the apex acute to obtuse, acuminate for up to 2 cm, the margins minutely recurved throughout; upper surface greenish grey, waxy, the primary to fourth-order veins raised, their prominence decreasing with rank; lower surface sparsely to moderately pubescent, the hairs appressed, silvery grey, uniformly distributed, all vein orders raised, their prominence decreasing with rank; secondary veins 3-5 per side, ? evenly spaced, slightly more distant around midlamina, ascending at 50-60? (more obtusely around midlamina), arcuate, the lowermost pair asymmetric (one or both merging with margin at base), distal pairs loop-connected, or distal pairs weakly loop-connected; tertiaries close, roughly horizontal to oblique to midrib, between secondaries straight or once-forked. Staminate inflorescences evenly spaced along current flush in the axils of foliage leaves, to 15 cm long with 8 lateral branches, branch orders 3-4, the highest order dichasial, lax, the flowers distant, the axes silvery strigose, distalmost branches and pedicels more densely so; bracts and bracteoles caducous by anthesis, ovate, sericeous; pedicels terete, to 2 mm long, those supporting secondary flowers slightly shorter. Flowers urceolate, up to 2 mm diam., densely silvery sericeous outside; receptacle deeply cyathiform, 1.3 x 1 mm, constricted below tepals, silvery tomentose inside. Tepals chartaceous, depressed-ovate, 0.4 X 0.7 mm, the inner whorl slightly smaller, erect at anthesis, the inner surface sparsely tomentose near base, the margins and apex

inside minutely papillose. Stamens of whorls I and II 0.6 mm tall, sessile, the anthers ovate, 0.3 X 0.4 mm, the apex truncate, the connectives broad above the 2 locelli, these ellipsoid, introrse, the filaments ligulate, as broad as anthers, slightly narrower in whorl II, densely grey-tomentose; whorl III stamens sessile, 0.6 mm tall, the anthers oblong, 0.4 X 0.3 mm, erect, locelli 2, occupying the lower half of the anther, extrorse-latrorse, the filaments broader than anthers, fleshy, the basal glands absent; whorl IV wanting; pistillode filiform. Pistillate inflorescence with indument and color as in staminate plants, but shorter and with fewer lateral branches, the flowers slightly deeper; stamens sterile, smaller; ovary glabrous, ovoid; style stout, weakly distinguished from ovary; stigma minutely tri-lobed, 0.3 mm diam. Fruits borne on stout claviform pedicels of up to 2 X 0.7 cm; cupules heavy walled, hemispherical, to 1.5 X 3 cm, glabrous inside and outside, the margins entire; drupes ovoid, to 4 X 2 cm.

Distribution (Fig. 8) and ecology. Trees from well-drained soils in central to western Amazonia and adjacent eastern Andean foothills at 100-1100 m. Flowering from March to October, with fruits available by August until June of the following year.

Representative specimens examined. COLOMBIA. ANTIOQUIA: San Luis, Quebrada La Cristalina, 550-690 m, 25 May 1987 (fr), Ram(rez & Cdrdenas 1016 (MO); Alto Rico, Rio Claro, 600 m, 9 Oct 1982 (fr), Renterfa et al. 2821 (MO).

ECUADOR. MORONA-SANTIAGO: Pozo petrolero 'Garza' de TENNECO, 35 km al NE de Montalvo, 260 m, 2-12 Jul 1989 (fl Y, fr juv), Zak & Espinoza 4671 (MO). NAPO: Aguarico, Reserva Etnica Huaorani, carretera y oleoducto de Maxus, km 54, 250 m, 26-30 Sep 1993 (fl d), Aulestia & Andi 841 (MO); Estaci6n Experimental INIAP- Napo, Payamino, Reserva Floristica "El Chuncho," 250 m, 5 Apr 1986 (fr), Jaramillo 8349 (B, MO). SANTIAGO-

ZAMORA: Taisha, 500 m, 1 Feb 1962 (fr), Cazalet & Pen- nington 7629 (B, K, US).

PERU. AMAZONAS: Bagua, Imaza-Yamayakat, 400 m, 25 May 1996 (fl d), Vdsquez & Vdsquez 20969 (MO), 450 m, 15 Nov 1997 (fr), Vdsquez et al. 24898 (MO). Cuzco: La Convenci6n, 910 m, 12 Jun 1968 (fr), Dudley 10072 (MO). HUANUCO: Pachitea, Puerto Inca, Llullapichis, 280 m, 16 May 1989 (fl Y), Kroll Saldafia 326 (MO). JUNiN: Juaja, Rio Negro, N of Satipo, 800 m, 20 Aug 1960 (frjuv), Woytkowski 5864 (US). LORETO: Requena, 140 m, 11 Aug 1988 (fl d), van der Werif et al. 10084 (MO); Maynas, Ca- huide, 11 Oct 1984 (fr), Vdsquez & Jaramillo 5697 (MO). MADRE DE Dios: Tambopata, Inca, Puerto Maldonado, carretera Maldonado-Cuzco, km 16, 240 m, 22 Jul 1989 (fr), Vdsquez et al. 12435 (MO). PASCO: Oxapampa, Palcazd, 300-600 m, 26 Mar 1986 (fl d), Hartshorn et al. 2916

34 FLORA NEOTROPICA

~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~T - ..-F- ;

I~~~~~~~~~~~~~~~

0 peX 1 ...S aS C1 ) i X

------F-- -- Dis o of E r m a E c

<~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~CA FIG.8. Dstriutin ofEridicheia etalica,E. hrysveluina,arl E. rownana

(MO). SAN MARTiN: Rioja, Pardo Miguel, 1100 m, 16 Jun 1998 (fl d), Sdnchez Vega et al. 9363 (MO).

BRAZIL. AMAZONAS: Manaus, Igarape do Passarinho, 14 Mar 1956 (fr), Co6lho 3611 (INPA); Rio Purus, Cach- oeira Uby, 22 Jun 1903 (fl 6), Goeldi s.n. MG 3911 (U).

BOLIVIA. LA PAz: Alto Madidi, 370 m, 26 May 1990 (fr), Gentry & Estensoro 70632 (MO); SANTA CRUZ: Es- taci6n del Valle de Sacta, Regi6n del Chapare, 5 May 1989 (fl d3), Moretti 1443 (MO).

Local names. Peru: mautaga, mantaga, moena callhuangia amarilla, moena negra. Bolivia: laurel negro.

Endlicheria metallica shares campanulate flowers and an androecium of nine sessile stamens with E. chrysovelutina, but differs by its closely appressed silvery-grey indument. Rlower shape provides easy contrast with vegetatively similar Rhodostemonoda- phne grandis, R. praeclara, R peneia, and R. saulen- sis, but all fruiting specimens (25 of the 39 known collections) had to be initially identified by locating

two-locellate staminodes remnant on cupule rims. Af- ter this sorting exercise, it became apparent that En- dlicheria metallica is a predominantly western Am- azonian species (reaching the lower slopes of the Andes) that consistently has elliptic or oblong leaves that assume a greenish grey color in the dry state, while these Rhodostemonodaphne species are restricted to northeastern South America and have obovate leaves that dry dark brown (see also Madrifinn, 1996b).

9. Endlicheria chrysovelutina Chanderbali, sp. nov. Type. Peru. Loreto: Maynas, Iquitos, Puerto Al- mendras, 130 m, 27 Jul 1988 (fl d), van der Werff et al. 9815 (holotype: MO; isotypes: F, GH, NY- n.v.). Fig. 9

Endlicheriae metallicae absque dubio proxima, sed ramulis chrysovelutinis et foliis subtus praeclaro pannosis- sericeis differt.

co

__ G *D -- E 0.5mmrHn 1 mm:

FIG. 9. Endlicheria chrysovelutina (AXG, van der Werif et al. 9815; H, Vdsquez & Jaramillo 9183). A. Habit. B. Close-up of lower leaf surface. C. Leaf base below. D. Rlower with facing tepal turned down to reveal androecium. E. Rlower l.s. F. Whorl I stamen seen from within. G. Whorl HII stamen seen from without. H. Fruiting branchlet.

36 FLORA NEOTROPICA

Trees to 20 m. Branchlets stout, midway along flush 4-7 mm diam., striate, angular, densely velutinous, the surface concealed by the indument cover, the hairs relatively long, to 0.6 mm, straight, erect, golden to reddish yellow; terminal buds plump, 2 X 1 cm, densely pubescent, the hairs as on branchlets, ascending. Leaves altemate, widely and evenly spaced along current flush; petioles robust, to 4 X 0.4 cm, semi-terete, striate, the indument as on branchlets; laminae coriaceous, plane, ovate to elliptic, 14-23 X 8-10 cm, the base obtuse to truncate, briefly decurrent, the apex obtuse, acuminate or cus- pidate for up to 1.2 cm, the margins minutely recurved near base or throughout; upper surface light green to olive-brown, waxy, the primary to fourth-order veins raised, their prominence decreasing with rank; lower surface densely golden to yellow pannose-sericeous, the hairs ascending to appressed, reduced on main veins, all vein orders raised, their prominence decreasing with rank; secondary veins 3-5 per side, ? evenly spaced, slightly more distant around midlamina, ascending at 50-60? (more acutely towards apex), arcuate, the lowermost pair asymmetric (one or both merging with margin at base), distal pairs loop- connected; tertiaries roughly horizontal to midrib, closely spaced, between secondaries straight to forked. Staminate inflorescences evenly spaced along current flush in the axils of foliage leaves, to 25 cm long with 12 lateral branches, branch orders 4-5, the highest order dichasial, lax, the flowers distant, the axes golden to yellow velutinous; bracts and bracteoles persistent at anthesis, eventually falling, ovate, sericeous, the hairs as on axes; pedicels terete, to 4 mm long, those supporting secondary flowers slightly shorter. Flowers urceolate, 3 mm diam., densely silvery to yellow tomentellose outside; receptacle cyathiform, 2 X 3 mm, slightly constricted below tepals, grey-velutinous inside. Tepals chartaceous, broadly ovate, 0.5 X 1 cm (the inner whorl slightly narrower), erect, surrounding androecium at anthesis, the outer surface silvery tomentellose, the inner surface glabrous, the margins papillose. Stamens of whorls I and II 0.5 mm tall, sessile, the anthers ovate, 0.3 X 0.4 mm, glabrous, the apex truncate, the connectives broad above the 2 locelli, these ellipsoid, introrse, the filaments ligulate, broader than anthers, densely silvery tomentose; whorl III stamens sessile, the indument and shape as in outer whorls, slightly narrower, locelli 2, introrse-latrorse, the basal glands absent; whorl IV wanting; pistillode filiform. Pistillate inflorescence with indument and color as in staminate plants, but shorter and with fewer lateral branches, the flowers slightly deeper; stamens sterile, smaller;

ovary glabrous; style slender, distinct from ovary; stigma tri-lobed, 0.5 mm diam. Fruits borne on stout claviform pedicels of up to 3 X 0.7 cm; cupules thick walled, hemispherical, to 2 X 3.5 cm, glabrous inside and outside, the margins entire; drupes ovoid, to 3.5 X 2 cm.

Distribution (Fig. 8) and ecology. Medium-sized trees known only from lowland (ca. 120 m) white sand forests around Iquitos, Peru. Flowers have been collected in June, July, and October, and the single fruiting collection was made in June.

Additional specimens examined. PERU. LORETO: Maynas, Puerto Almendras, 122 m, 20 Oct 1982 (fl 9) Vds- quez 3319 (F, MO, NY); Iquitos, carretera Iquitos-Nauta, km 45, 120 m, 12 Jun 1987 (fl d) Vdsquez & Jaramillo 9180 (F, MO), (fr), Vdsquez & Jaramillo 9183 (MO).

Endlicheria chrysovelutina differs from E. metallica by its velutinous branchlets and denser, more felt- like, lower leaf surface indument. Both species most likely belong to a species group in Rhodostemono- daphne centered around R. grandis (see discussion in Relationships, above) wherein an undescribed species, represented by Steyermark 59001 (F, MO) from Mount Roraima, also has the velutinous indument of Endlicheria chrysovelutina. Flowers in Steyermark 59001 are campanulate, as in E. chrysovelutina, but stamens are spathulate and anthers are four-locellate. Were it not for this specimen, E. chrysovelutina would be recognizable in this transgeneric alliance by its indument alone, while flowers, or remnant stamens in fruiting material, are necessary to assign all other members to genus.

10. Endlicheria browniana Mez, Jahrb. Konigl. Bot. Gart. Berlin 5: 115. 1889. New name for Ayden- dron macrophyllum Meisn., DC. Prodr. 15(1): 92. 1864. Type. Panama. Darien: Cape Corrientes, without date (fl 9, fr juv), Seemann 1094 bis (holotype: K).

Trees to 15 m. Branchlets moderately stout, midway along flush 4-5 mm diam., distally weakly angular, soon terete, rusty or silvery strigillose, the surface clearly exposed to concealed by the indument cover, the hairs very short, to 0.1 mm, straight, appressed; terminal buds plump, 3 X 2 mm, sericeous. Leaves alternate, widely and evenly spaced along current flush; petioles slender or robust, to 2.5 X 0.4 cm, semi-terete, the indument as on branchlets; laminae coriaceous to membranaceous, plane, obovate to ovate, 15-35 X 7-15 cm, the base bluntly rounded, or acute and shortly attenuate, the apex broadly obtuse


then abruptly acuminate for up to 2.5 cm, the margins minutely recurved throughout; upper surface dark brown to light or olive green, waxy, the primary to fourth-order veins raised, their prominence decreasing with rank; lower surface sparsely strigillose, the hairs uniformly distributed, all vein orders raised, their prominence decreasing with rank; secondary veins 6-9 per side, ? evenly spaced or closer towards leaf base, ascending at 50-60? (more obtusely towards apex), arcuate, distal pairs loop-connected; tertiaries oblique to midrib, between secondaries once-forked to straight. Staminate inflorescences evenly spaced along current flush in the axils of foliage leaves, to 15 cm long with 8 lateral branches, branch orders 3-4, the highest order dichasial, lax, the flowers distant, the axes strigillose; bracts and bracteoles caducous by anthesis, lanceolate, sericeous; pedicels gradually increasing in diameter apically and merging with receptacle, to 2 mm long, those supporting secondary flowers slightly shorter. Flowers hypocrateriform, 3 mm diam., densely rusty or silvery strigillose outside; receptacle globose, 0.6 X 1 mm, constricted below tepals, densely rusty or silvery tomentose inside. Tepals chartaceous, ovate, 1 X 0.6 mm (the inner whorl slightly broader), spreading at anthesis, the inner surface densely pubescent, the hairs yellowish, tightly crinkled to papillose. Stamens of whorls I and II sessile, 0.6 mm tall, the anthers ovate, 0.4 X 0.5 mm, minutely papillose, the apex apiculate, the connectives prolonged between the 2 locelli, these obliquely hemispherical, introrse-latrorse, the filaments ligulate, as broad as anthers, densely yellowish papillose; whorl III stamens sessile, 0.6 mm tall, the anthers depressed-ovate, 0.3 X 0.5 mm, erect, locelli 2, extrorse-latrorse, the filaments as broad as anthers, ligulate, the indument as in outer whorls, the basal glands sessile, minute, globose; whorl IV wanting; pistillode wanting. Pistillate inflorescence with indument, color, and branching as in staminate plants, the flowers slightly deeper; stamens sterile, smaller; ovary sparsely tomentellose or glabrous, ovoid; style stout, weakly distinguished from ovary; stigma tri- lobed, papillose, 0.5 mm diam. Fruits bome on stout terete pedicels of up to 2 X 0.5 cm; cupules hemispherical, to 1 X 2 cm, glabrous inside and outside, the margins undulate; drupes ovoid, to 3 X 1.5 cm.

Distribution (Fig. 8) and ecology. Small to medium-sized trees from lowlands and lower montane slopes (100-800 m) west of the Andes from Ec- uador to Panama. Flowering material has been collected from September to April, and fruits from October to June.

Additional specimens examined. PANAMA. BOCAS DEL TORO: Chiriquf Lagoon, Fish Creek Mtns., 18 Apr 1941 (fl 6), von Wedel 2257 (A). COCLE: Rd. to Coclesito, 12 mi from Llano Grande, 600 m, 8 Dec 1983 (fr), Churchill et al. 3974 (MO), 9 Dec 1983 (fr), Churchill et al. 3998 (MO). COL6N: Hills just N of Rio Guanche, 100-200 m, 16 Nov 1975 (fl 6), Davidse & D'Arcy 10097 (MO); Rio Boquer6n, 100 m, ca. 14 mi from hwy., 13 Apr 1985 (fr), Hammel 13518 (MO); Rio Guanche, ca. 2.5 km upriver from bridge on rd. to Portobelo, 3 Jun 1975 (fr), Mori et al. 6476 (MO). DARIEN: Cape Corrientes, s.d. (fr), Seemann 1094 (K). PANAMA: El Llano-Carti Rd., 300 m, 6 Sep 1980 (fl d), Sytsma 994 (MO), 21 Mar 1996 (fl 6), Guerra 646b (MO); Gorgas Memorial labs, ca. 25 km NE of Cerro Azul on Rfo Piedras, 550 m, 25 Nov 1974 (fl 2), Mori & Kallunki 3472 (MO). SAN BLAS: El Llano-Carti Rd., km 16.7, trail W to Rio Carti Grande, 250-350 m, 4 Nov 1984 (fl 6), Nevers & Herrera 4186 (MO); El Llano-Carti Rd., km 26.5, along Rio Carti Chico, 200 m, 12 Apr 1985 (fr), Nevers et al. 5339 (MO); Play6n Chico, Rio Grande, 50-100 m, 2 Nov 1991 (fl 2), Herrera et al. 1057 (HBG, MO).

COLOMBIA. CHOC6: Ca. 50 km W of Las Animas, ca. 4 km E of Rfo Pato on Pan American Hwy., 250 m, 11 Jan 1979 (fl 6), Gentry & Renterfa 23983 (HBG, MO), (fr), Gentry & Renteroia 23982 (MO). VALLE: Buenaventura, Quebrada "La Trojita," 30-100 m, 20 Mar 1992 (2 fl & fr), Cogollo et al. 5119 (MO); Rio Calina, 0-5 m, 11 Mar 1944 (fl 6), Cuatrecasas 16855 (F, US).

ECUADOR. CARCHI: San Marcos de los Coaqueres, Chical-Tobar Donoso Trail, 800 m, 8 Feb 1985 (fl 6), 0llgaard et al. 57630 (MO, NY), (fr), 0llgaard et al. 57623A (MO). ESMERALDAS: San Lorenzo, Reserva Etnica Awa, 250 m, 22 Mar 1993 (fl 2), Aulestia & Aulestia 1289 (MO, NY); Eloy Alfaro, Reserva Ecol6gica Cotacachi- Cayapas, Parroquia Luis Vargas Torres, 250 m, 23-27 Oct 1993 (fr), Tirado et al. 586 (MO); Muisne, Sitio San Sal- vador, orillas del Rfo Sucio, 100-150 m, Mar 1995 (fl 6), Palacios 13750 (MO).

Endlicheria browniana is unmistakable in fruit on

account of its hemispherical cupules with undulate margins and stout pedicels. Its hypocrateriform flowers with distally constricted receptacles below horizontally spreading tepals are much like those found in E. jefensis, E. mishuyacensis, and E. pyriformis, but the laxly reticulate tertiaries of these three species are quite unlike the percurrent pattern formed by those of E. browniana.

The concept of Endlicheria browniana adopted here is broader than that suggested by the type material and most of the other collections from the Dar- ien-Choc6 region. These have large obovate coriaceous leaves with rounded bases and stout, seemingly swollen, petioles. Moreover, pistillate flowers have densely pubescent ovaries, a feature otherwise unknown in Endlicheria. These characters seem sufficient to circumscribe a highly distinctive species, but

38 FLORA NEOTROPICA

ovaries are glabrous in a vegetatively indistinguishable collection from quite distant San Blas in Panama (Herrera et al. 1057). Furthermore, in Gentry & Ren- terfa 23982 from the Choco, young drupes are glabrous and leaves are much smaller with rather slender petioles compared to other Darien-Choco material. Given this variation in ovary pubescence and leaf morphology in the Darien-Choco region, and since staminate flowers from there (Cuatrecasas 16855, Gentry & Renteria 13983) are indistinguishable from those collected elsewhere, an expanded concept of E. browniana is unavoidable. Unfortunately, this recourse also increases the range of vegetative variation. Variation in leaf size, shape, and texture appears continuous and can be ignored as a possible source of specific characters, but variation in indument is less readily dismissed. In most collections, including the Darien-Choc6 material discussed above, the indument of branchlets, leaves, inflorescences, and flowers consists of dull rust-colored hairs, but all Ecuadorian material have a shiny silvery-grey indument. How- ever, any apparent correlation between indument color and geographic location is disrupted by two Panamanian collections with silvery-grey indument (von Wedel 2257 and Churchill et al. 3974). Further- more, fruits of silvery-grey forms from Ecuador as well as Panama show the distinctive cupules of Dar- ien-Choc6 material. Within both color morphs, the indument density on branchlets ranges from sparse to subsericeous, and in the latter case provides an aspect reminiscent of the E. sericea group (species 46-55). Separation on the basis of indument color or density alone or in combination would produce either two or four entities, neither of which could be recognized on the basis of other characters.

11. Endlicheria jefensis van der Werff ex Chander- bali, sp. nov. Type. Panama. Panama': Cerro Jefe region, along road to Rio Crist6bal in Chagras drainage, 600 m, 25 Feb 1986 (fl d'), McPherson 8493 (holotype: MO; isotypes: F, HBG, MO, NY, PMA, US). Fig. 10

Endlicheriae brownianae et affinibus similis et nullo dubio his speciebus proxima, sed floribus sub-glabris et foliis ovatis differt.

Trees to 8 m. Branchlets slender, midway along flush 2-3 mm diam., distally weakly angular, soon terete, sparsely strigillose, the surface clearly exposed, dark brown, the hairs short, to 0.1 mm, straight, appressed, greenish grey; terminal buds slender, 2 X 0.6 mm, densely greenish white sericeous.

Leaves alternate, widely and evenly spaced along current flush; petioles slender, to 2 X 0.2 cm, semi-terete, sparsely grey-strigillose above, otherwise glabrous; laminae coriaceous, plane, ovate to elliptic, 9-14 X 3-5 cm, the base acute, attenuate, the apex acute, acuminate for up to 1 cm, the margins minutely recurved throughout; upper surface deep olive-brown, waxy, the primary to fourth-order veins raised, their prominence decreasing with rank; lower surface sparsely strigillose, the hairs as on branchlets, denser on main veins, soon lost, all vein orders raised, their prominence decreasing with rank; secondary veins 4-7 per side, ? evenly spaced, slightly more distant around midlamina, ascending at 50-60 (more obtusely towards apex), arcuate, distal pairs loop-connected; tertiaries laxly reticulate. Staminate inflorescences evenly spaced along current flush in the axils of foliage leaves, to 7 cm long with 12 lateral branches, branch orders 2-3, the highest order dichasial, lax, the flowers distant, the axes sparsely grey-strigillose; bracts and bracteoles caducous by anthesis (none seen); pedicels gradually increasing in diameter apically, to 3 mm long, those supporting secondary flowers slightly shorter. Flowers hypocrateriform, to 4 mm diam., sparsely grey-strigillose, the hairs scattered, soon lost; receptacle deeply infundibuliform, the apex slightly constricted below tepals, 1.8 X 2 mm, densely tawny papillose inside. Tepals chartaceous, broadly ovate, 1.1 X 0.9 mm, spreading, the inner surface rusty papillose. Stamens of whorls I and II sessile, 0.6 mm tall, the anthers broadly ovate, 0.3 X

0.6 mm, glabrous, the apex apiculate, the connectives prolonged between the 2 locelli, these obliquely hemispherical, introrse-latrorse, the filaments broader than anthers, densely rusty papillose; whorl III stamens sessile, 0.6 mm tall, the anthers ovate, 0.4 X 0.4 mm, erect, locelli 2, extrorse-latrorse, the filaments broader than anthers, shortly columnar, rusty yellowish papillose, the basal glands sessile, globose; whorl IV wanting; pistillode wanting. Pistillate plants unknown.

Distribution (Fig. 11) and ecology. Known only from the type material taken from a staminate tree found flowering in February on the forested slopes of Cerro Jefe (Panama).

In the Endlicheria browniana species group, the narrowly ovate to elliptic coriaceous leaves and relatively large subglabrous flowers of E. jefensis are without equal. In Panama, E. browniana has much larger leaves and densely pubescent flowers, and E. formosa has depressed-globose flowers. In adjacent Colombia, E. colombiana has similar indument, but


FIG. 10. Endlicheria jefensis (McPherson 8493). A. Habit. B. Leaf base below. C. Flower. D. Flower ls. E. Whorl Ill stamen seen from without. F. Whorl I stamen seen from within.

40 FLORA NEOTROPICA

7i

-- . - -- -- - - -- --

*~~~~~~~~~~~~~~~~~~~~~~

A ...~~~~~~~~~~~~~J

y %~~~~~~tf4

FIG --- 11 Ditibto of Enlcei eessEclmin n ihycn

broader leaves with canaliculate petioles. Western Amazonian E. mishuyacensis has similar-sized dark- drying leaves, but these are obovate to elliptic, and rust brown rather than greyish hairs appear on the branchlets.

12. Endlicheria colombiana (Meisn.) Mez, Jahrb. Konigl. Bot. Gart. Berlin 5: 117. 1889. Oreoda- phne colombiana Meisn., DC. Prodr. 15(1): 137. 1864. Type. Colombia. Antioquia: Without locality and date (fl 9), Jervise s.n. (lectotype, designated by Kostermans, 1937: K).

Trees to 8 m. Branchlets slender, midway along flush 3-5 mm diam., distally weakly angular and sparsely strigillose, soon terete and glabrous, the surface always clearly exposed, grey to dark brown, the hairs short, to 0.15 mm, straight, appressed, greenish grey; terminal buds plump, 3 X 2.5 mm, densely grey-sericeous. Leaves alternate, widely and evenly spaced along current flush; petioles slender, to 1.5 X

0.3 cm, canaliculate, the indument as on branchlets; laminae chartaceous, plane, elliptic, 11-25 X 5-12 cm, the base obtuse, briefly decurrent, the apex obtuse, acuminate for up to 1 cm, the margins minutely recurved throughout; upper surface light greyish green to olive-brown, waxy, the primary to fourth- order veins strongly raised; lower surface sparsely strigillose, the hairs soon restricted to main veins, eventually lost, the primary to fourth-order veins raised, their prominence decreasing with rank; secondary veins 5-9 per side, ? evenly spaced, slightly more distant around midlamina, ascending at 50 60? (more obtusely towards apex), arcuate, or arching after midcourse, distal pairs loop-connected; tertiaries oblique to midrib, between secondaries straight to once-forked. Staminate inflorescences evenly spaced along current flush in the axils of foliage leaves, to 10 cm long with 9 lateral branches, branch orders 2-3, the highest order dichasial, lax, the flowers distant, the axes sparsely grey-strigillose, distal branches and pedicels more densely so; bracts and bracteoles ca-


ducous by anthesis, none seen; pedicels terete, to 7 mm long, those supporting secondary flowers slightly shorter. Flowers hypocrateriform, 5 mm diam., sparsely grey-strigillose outside; receptacle infundibuliform, 2 X 3 mm, merging with tepals, densely rusty tomentellose inside. Tepals chartaceous, ovate, 2 X 1.3 mm, ascending to spreading at anthesis, the inner surface rusty tomentellose-papillose. Stamens of whorls I and II sessile, 0.7 mm tall, the anthers broadly ovate, 0.6 X 0.7 mm, minutely papillose, the apex apiculate, the connectives bluntly prolonged between the 2 locelli, these obliquely hemispherical, introrse-latrorse, the filaments ligulate, as broad as anthers, rusty tomentellose; whorl III stamens sessile, 1 mm tall, the anthers ovate, 0.6 X 0.5 mm, erect, locelli 2, extrorse-latrorse, the filaments broader than anthers, columnar, rusty tomentellose, the basal glands sessile, globose; whorl IV wanting; pistillode fusiform. Pistillate inflorescence with indument and color as in staminate plants, but shorter and with fewer lateral branches, the flowers slightly smaller; stamens sterile, smaller; ovary glabrous; style slender, distinct from ovary; stigma tri-lobed. Fruits unknown.

Distribution (Fig. 11) and ecology. Small trees from the Andes of Antioquia (Colombia) at ca. 1500 m. The two collections with calendar information were flowering in May and November.

Additional specimens examined. COLOMBIA. AN- TIOQUIA: Medellfn, May 1852 (fl 2), Triana 2040-2 (K); Frontino, Murri, Nutibara-La Blanquita Rd., km 22, 1450 m, 4 Nov 1988 (fl 6), Zarucchi et al. 7120 (INPA, MO). WITHOUT LOCALITY AND DATE: (fl 2), Linden 429 (K); (fl 2), Triana 1033 (P).

Endlicheria colombiana is a poorly known species. The 19th century collections seen by Meissner (1864), Mez (1889), and Kostermans (1937) are all pistillate plants with unfertilized flowers. They have spreading tepals and bear sessile stamens with ovate anthers typical of the E. browniana species group, but their smooth receptacle-tepal transition stands out among the distally constricted receptacle elsewhere in this species group. Vegetatively, these pistillate plants are almost identical and undoubtedly represent the same species, but a staminate plant from near the type locality in Antioquia, Zarucchi et al. 7120, is placed here with hesitation. Flowers of that specimen show the smooth receptacle-tepal transition noted as unique to E. colombiana in the E. browniana species group, but they are much more robust and show a sharper pedicel-receptacle transition than the pistillate flowers. Yet this may be a consequence of disease. Almost all leaves show severe insect damage, several

flowers are infested with fungal mycelia, and there is no sign of pollen in often shallow locelli.

13. Endlicheria mishuyacensis A. C. Sm., Bull. Torrey Bot. Club 58: 102. 1931. Type. Peru. Lor- eto: Mishuyacu, near Iquitos, 100 m, Oct-Nov 1929 (fl d), Klug 204 (holotype: NY; isotypes: F, US).

Trees, 3-20 m. Branchlets slender, midway along flush 2-3 mm diam., angular, sparsely strigillose, the surface clearly exposed, dark brown, the hairs reddish brown, ca. 0.1 mm, appressed; terminal buds slender, 1 X 0.3 mm, tan sericeous. Leaves alternate, widely and evenly spaced along current flush; petioles slender, to 1 X 0.2 cm, canaliculate, the indument as on branchlets; laminae chartaceous, plane, elliptic to obovate, 7-20 X 3-8 cm, the base acute, attenuate, the apex acute, acuminate for up to 1.5 cm, the margins flat; upper surface dark green to blackish brown, waxy, the primary to fourth-order veins raised, their prominence decreasing with rank; lower surface sparsely rusty strigillose, the hairs slightly denser on main veins, all vein orders raised, their prominence decreasing with rank; secondary veins 5-7 per side, ? evenly spaced, slightly more distant around midlamina, patent, diverging at 70-85? (more obtusely around midlamina), arcuate to abruptly ascending after midcourse, distal pairs loop-connected; tertiaries laxly reticulating between secondaries. Staminate inflorescences evenly spaced along current flush in the axils of foliage leaves, to 5 cm long with 3 lateral branches, branch orders 2-3, the highest order dichasial, lax, the flowers distant, the axes rusty strigillose, distal branches and pedicels more densely so; bracts and bracteoles caducous by anthesis, ovate, rusty sericeous; pedicels gradually increasing in diameter apically, to 2 mm long, those supporting secondary flowers slightly shorter. Flowers hypocrateriform, ca. 3 mm diam., densely rusty strigillose outside; receptacle infundibuliform, 0.3 X 0.6 mm, slightly constricted below tepals, rusty papillose inside. Tepals chartaceous, ovate to triangular, 1 X 0.6 mm (the inner whorl slightly narrower), spreading to reflexed at anthesis, the inner surface minutely rusty papillose, the indument reduced to a basal triangular patch in the outer whorl but covering surface of inner whorl. Stamens of whorls I and II 0.5 mm tall, sessile, the anthers ovate, 0.3 X 0.3 mm, minutely rusty papillose near base on the abaxial side, otherwise glabrous, the apex apiculate, the connectives prolonged between the 2 locelli, these obliquely hemispherical, introrse- latrorse, the filaments fleshy, slightly broader than an-

42 FLORA NEOTROPICA

thers, densely rusty papillose; whorl III stamens sessile, 0.6 mm tall, the anthers ovate, equal to those of the outer whorls, erect, locelli 2, extrorse-latrorse, the filaments as broad as anthers, fleshy, columnar, rusty papillose, the basal glands absent; whorl IV wanting; pistillode filiform. Pistillate inflorescence with indument, color, and branching as in staminate plants, the flowers similar in size and shape; stamens sterile, smaller; ovary glabrous, ovoid; style stout, weakly distinguished from ovary; stigma tri-lobed, ca. 0.4 mm diam., minutely papillose. Fruits borne on claviform pedicels of up to 1.5 X 1 cm; cupules shallowly hemispherical, thick walled, to 1 X 2 cm, glabrous inside and outside, the margins entire; drupes ovoid, to 3 X 2 cm.

Distribution (Fig. 11) and ecology. Medium- sized trees already fertile at 3 m occurring in nonflooded lowland (100-400 m) forests in western Ama- zonia. Flowers and fruits have both been collected in all months but January, February, and March.

Representative specimens examined. COLOMBIA. AMAZONAS: Rio Apaporis, entre el Rfo Paco y el Rfo Kan- anari, Soratama, 250 m, 25 Sep 1951 (fl 5d), Schultes & Cabrera 14125 (COL, NY); Rio Miritiparana, Caino Gua- caya, 250 m, 24 Apr 1952 (fr), Schultes & Cabrera 16230 (GH, HBG, NY).

ECUADOR. PASTAZA: Pozo petrolero 'Golondrina' de PETRO-CANADA, 25 km al NW del pueblo de Curaray, 400 m, 23 Jun 1989 (fl Y), Rubio & Gudifio 199 (G, MO); Auca, 115 km al S de Coca, Rfo Tigiiino, 320 m, 29 Apr 1989 (fr), Rubio 115 (G, MO).

PERU. LORETO: Maynas, Mishuyacu, near Iquitos, 100 m, Dec 1929 (fl Y), Klug 703 (F, NY, US); Iquitos, Quis- tococha, 22 Nov 1940 (fl d), Asplund 14682 (G, K, NY, R); Mishana, Estaci6n Biologica Callicebus, 15 Aug 1980 (fr), Foster 4302 (HBG, MO, NY, U).

BRAZIL. AMAZONAS: Basin of Rio Jurua, near mouth of Rio Embira (tributary of Rio Tarauaca), 21 Jun 1933 (fl d5), Krukoff 4959 (A, F, G, K, MO, NY, U); Lago do Cas- tanho, 25 Jun 1973 (fl ), Albuquerque et al. 850 (INPA). ACRE: Cruzeiro do Sul, Rio Jurua & Rio Moa, Serra de Moa village, 28 Apr 1971 (fl d), Prance et al. 12633 (HBG, K, MO, NY, US).

Local names. Colombia: ko-mwa-ko-ree (canoe tree). Peru: muena.

Endlicheria mishuyacensis alone in the E. browniana species group combines rusty pubescence and dark brown leaves with patent secondaries and reticulate tertiaries. Sympatric E. dysodantha is superfi- cially similar with dark leaves of similar size, shape, and color, but has barbellate tufts of hairs in the axils of the secondary veins below and scalariform tertiaries.

14. Endlicheria pyriformis (Nees) Mez, Jahrb. Kon- igl. Bot. Gart. Berlin. 5: 116. 1889. Cryptocarya pyriformis Nees, Syst. laur. 220. 1836. Mespilo- daphne pyriformis (Nees) Meisn., DC. Prodr. 15(1): 108. 1864. Type. French Guiana. Without locality and date (fr), Poiteau s.n. (holotype: B- n.v.; isotypes: G, LE, NY, P).

Endlicheria glaberrima Mez, Bull. Herb. Boiss. 5 (ser. 2): 236. 1905. Type. Peru. Loreto: Yurimaguas, Aug 1902 (fl Y), Ule 6296 (holotype: B-n.v.; isotype: HBG).

Aniba flexuosa A.C. Sm., Phytologia 1: 117. 1935. Type. Brazil. Amazonas: Near mouth of Rfo Embira (tributary of Rio Tarauaca), 21 Jun 1933 (fl ), Kru- koff 5030 (holotype: NY).

Trees to 10 m. Branchlets slender, midway along flush 2-3 mm diam., angular, glabrous, light green; terminal buds slender, 2 X 0.6 mm, sparsely grey- strigillose. Leaves alternate, widely and evenly spaced along current flush; petioles usually slender, usually 1 X 0.1 cm (rarely to 4 X 0.3 cm), canaliculate, glabrous; laminae chartaceous, plane, obovate to elliptic, 10-25 X 3-17 cm, the base cuneate, attenuate, the apex obtuse, acuminate for up to 1.5 cm, the margins flat; upper surface light green, waxy, the primary to fourth-order veins raised, their prominence decreasing with rank, the midrib and secondaries creamish yellow, conspicuous against the greenish lamina; lower surface glabrous, all vein orders raised, their prominence decreasing with rank; secondary veins 6-8 per side, ? evenly spaced, slightly more distant around midlamina, patent, diverging at 70-85? (more obtusely around midlamina), abruptly ascending after midcourse, distal pairs loop-connected; tertiaries laxly reticulating between secondaries. Sta- minate inflorescences evenly spaced along current flush in the axils of foliage leaves, to 15 cm long with 10 lateral branches, branch orders 3-4, the highest order dichasial, lax, the flowers distant, the axes sparsely strigillose, glabrescent; bracts and bracteoles persistent at anthesis, triangular, sparsely strigillose; pedicels gradually increasing in diameter apically, to 5 mm long, those supporting secondary flowers slightly shorter. Flowers hypocrateriform, 3 mm diam., sparsely grey-strigillose outside; receptacle cyathiform, 2 X 1.3 mm, densely pubescent inside, the hairs papillose, crisped, or straight. Tepals chartaceous, ovate, 1 X 0.6 mm, both whorls equal, spreading at anthesis, the inner surface minutely papillose throughout, or the hairs crisped to straight near base. Stamens of whorls I and II broadly stipitate, 1 mm tall, the anthers narrowly ovate, 0.5 x 0.3 mm, glabrous, the apex apiculate, the connectives strongly


prolonged between the 2 locelli, these obliquely hemispherical, introrse-latrorse, the filaments ligulate, almost as broad as anthers, sparsely pubescent, the hairs crisped to papillose; whorl III stamens sessile, 1.2 mm tall, the anthers narrowly ovate 0.6 x 0.3 mm, erect, locelli 2, extrorse-latrorse, the filaments as broad as anthers, columnar, the indument as in outer whorls, the basal glands sessile, minute, globose- apiculate; whorl IV wanting; pistillode filiform. Pis- tillate inflorescence with indument, color, and branching as in staminate plants, the flowers slightly deeper; stamens sterile, smaller; ovary glabrous, ovoid; style slender, distinct from ovary; stigma broadly tri-lobed, 0.6 mm diam. Fruits borne on claviform pedicels of up to 2.5 x 0.5 cm; cupules infundibuliform, to 0.8 X 1.7 cm, glabrous inside and outside, the margins entire; drupes ellipsoid, to 3 x 1.5 cm.

Distribution (Fig. 12) and ecology. Small understory trees of noninundated forests from the Guianas to western Amazonia and along lower An- dean slopes from Peru to Venezuela, at 100-1600 m.

Flowering from April to December. Fruits collected throughout the year.

Representative specimens examined. COLOMBIA. META: Sierra de La Macarena, 630 m, 24 May 1973 (fr), Garcfa et aL 380 (COL); Villavicencio, Bosques de Viena, 28 Apr 1988 (fr), Quiflones 1419 (MO). PLTTuMAYO: Mo- coa, 28 Jul 1990 (fl d), Garcia et al. 103 (MO); Mocoa, 1350-1420 m, 20 Apr-I May 1994 (fr), Ferndndez et al. 11051 (COL, GH).

VENEZUELA. ARAGUA: Henri Pittier National Park, 770 m, 6 Apr 1990 (fl d), Edwards & Roe 423 (MO).

GUYANA. EssEQuiBo: Rupununi Region, Acarai Mtns., 500 m, 5 Mar 1994 (fr), Henkel et al. 4946 (MO), 300-600 m, 2 Nov 1996 (fl 6), Clarke et al. 2865 (MO).

SURINAME. Tafelberg, East Ridge Creek Gorge, 750 m, 29 Aug 1944 (fl d), Maguire 24538 (A, NY, U); Wih- elmmina Mtns., Juliana Top, 1236 m, 13 Aug 1963 (fl 6), Irwin et al. 54744 (A, HBG, MO, U).

FRENCH GUIANA. Montagne Bellevue de 1' Inini, 600 m, 1 Sep 1985 (fl 6), de Granville et al. 7949 (MO, NY, US); Eau Claire, near Saul, 200 m, 11 Aug 1993 (fl 6), van der Werf/et al. 12944 (MO, NY).

4 > --.* . *

f * - t t s- 9 / . g . ;. gS . . ;\rffi * r fi S

FIG 12 D istribution t of Endl hena pynfo. . *

'' ", ';"q.'> \' .. ' ' ' 1; ' '; ii... W 4N ;'\. 4 e ' '',~~ ~~ ~~ ~~ ~~ ~~ ~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~.......

- .7' , >. t T._v.. v

-~~~ .. - - - - - - - f - , .tz-r . :- 7 Az-

FIG. 12. Distribution of Endlicheria pyriformis.

44 FLORA NEOTROPICA

ECUADOR. MORONA SANTIAGO: El Centro Shuar Kankaim, Rfo Kankaim, 500 m, 18 Dec 1985 (fr), Shiki RBAE359 (MO). NAPO: Estaci6n Biol6gica Jatun Sacha, 450 m, 17-21 Nov 1988 (fl 6), Ceron & Iguago 5562 (MO); Reserva Floristica "El Chuncho," 250 m, 7 Oct 1987 (fr), Ceron 2397 (HBG, MO).

PERU. HUANUCO: Rio Cuchara, Villa Ysabel, 21 Sep

1961 (fr), Schunke 5679 (F, K). LORETO: Maynas, Rio Pu- tumayo, Florida, mouth of Rfo Zubineta, 180 m, Oct-Nov 1931 (fl d), Klug 2313 (A, MO); Alto Amazonas, Balsa- puerto, 220 m, Mar 1933 (fl 6), Klug 2958 (GH, K, MO). PUNO: Rio Tavara, 400 m, 19 May 1992 (fl Y), Gentry et al. 76844 (MO); Sandia, between Rio Azata-Colorado, 1100 m, 26 Jun 1986 (fl 6), Niiunez & Munioz 5324 (MO). SAN MARTiN: Lamas, Convento, trail to Tioyacu and Nuevo La- mas, km 68 on Tarapota-Yurimaguas rd., 270 m, 23 Jun 1984 (fl 6), Knapp & Mallet 6531 (F, MO, NY, US); Mar- iscal Caceres, Campanilla, SE of Caserio de Si6n, Rfo Si6n, 3 Oct 1969 (fr), Schunke 3474 (COL, G, GH, NY, US).

BRAZIL. AMAPA: Rio Jari, 400 m, 17 Aug 1993 (fl 9),

de Granville et al. 12329 (MO, US); Colonia do Torrao, 29 Aug 1962 (fl 9), Pires & Cavalcante 52676 (MG, NY, US). AMAZONAS: Manaus, Reserva Florestal Ducke, Manaus- Itacoatiara Rd., km 26, 31 Jul 1997 (fl 6), AssunCdo & Silva 583 (MO); Rio Ituxi, B6ca do Rio Curuquete, 8 Jul 1971 (fl 6), Prance et al. 13987 (HBG, INPA, K, MG, MO, NY). PARA: Altamira-Itaituba Rd., near EMBRAPA station, 30 Oct 1977 (fr), Berg 770 (HBG, MO, NY, U); Oriximind, Rio Trombetas, 17 Jul 1980 (fl 9), Cid Ferreira et al. 1563 (HBG, INPA, MG, MO, NY). RONDONIA: Porto Velho, UHE de Samuel, Rio Jamari, 8 Jun 1986 (fl 6), Cid Ferreira 7387 (F, K, MO, NY).

Endlicheria pyriformis is conspicuous for its lack of indument. Except for a fringe of short appressed hairs on terminal bud scales, vegetative structures are completely glabrous. Sparse hairs are found on inflorescence bracts, bracteoles and floral pedicels, but the inner surface of flowers bears the only noteworthy, and variable, indument. In all material from the Guianas and adjacent Brazil, the tepals and receptacles inside are papillose, as is typical of the E. browniana species group. Such flowers are also found in western Amazonia, e.g., Grandez & Jaramillo 817, but most specimens from western Amazonia and the eastern Andean slopes have a pilose indument of straight erect hairs, e.g., Krukoff 5030 (the type of Anibafiexuosa) and Palacios 3260. Another, conceivably intermediate, indument form is found in Ule 6296 (type of Endlicheria glaberrima), Prance et al. 13987, and Edwards & Roe 423, from Peru, Brazil, and NE Venezuela respectively. In these the hairs are erect, as in material from western Amazonia, but crisped rather than straight. As none of these three forms can be recognized on the basis of other characters, all are here kept together in E. pyriformis.

15. Endlicheria formosa A. C. Sm., Phytologia 1: 118. 1935. Type. Brazil. Amazonas: Basin of Rio Jurua, near mouth of Rio Embira (tributary of Rio Tarauaca), 4 Jul 1933 (fl d), Krukoff 5156 (holotype: NY; isotypes: A, F, G, K, S, U).

Trees to 30 m. Branchlets stout, midway along flush 4-8 mm diam., sharply angular, sparsely rusty strigillose, the surface clearly exposed, dark brown, the hairs very short, to 0.1 mm, straight, appressed; terminal buds slender, 6 X 2 mm, greyish sericeous. Leaves alternate, widely and evenly spaced along current flush; petioles slender, to 4 X 0.3 cm, canaliculate, the indument as on branchlets; laminae chartaceous to coriaceous, plane, obovate, 12-30 X 3-15 cm, the base acute to cuneate, attenuate, the apex obtuse, acuminate for up to 2 cm, the margins minutely recurved throughout; upper surface dark greyish green to olive-brown, waxy, the midrib prominulous throughout or sunken before midcourse, the higher- order venation raised; lower surface sparsely pubescent, the hairs as on branchlets, uniformly distributed, all vein orders raised, their prominence decreasing with rank; secondary veins 8-10 per side, + evenly spaced, slightly more distant around midlamina, ascending at 50-60? (more obtusely around midlamina), arcuate, distal pairs loop-connected; tertiaries oblique to midrib, between secondaries straight to forked. Sta- minate inflorescences evenly spaced along current flush in the axils of foliage leaves, to 10 cm long with 16 lateral branches, branch orders 3-4, the highest order dichasial, lax, the flowers distant, the axes rusty to grey-strigillose, distalmost branches and pedicels more densely so; bracts and bracteoles caducous by anthesis, ovate, grey-sericeous; pedicels terete, to 1 mm long, those supporting secondary flowers slightly shorter. Flowers depressed-globose, 2 mm diam., sparsely to densely rusty strigillose outside; receptacle patelliform, 0.3 X 2 mm, densely rusty papillose inside. Tepals chartaceous, broadly triangular, 0.3 X

0.6 mm (the inner whorl slightly narrower), erect to inflexed at anthesis, the androecium included except for a narrow apical pore through which anther valves protrude, the inner surface densely rusty papillose. Stamens of whorls I and II sessile, unequal, whorl I 0.6 mm long, whorl II half as tall, the anthers ovate, 0.4 X 0.2 in whorl I, 0.3 X 0.3 in whorl II, sparsely papillose, the apex apiculate, the connectives prolonged between the 2 locelli, these obliquely hemispherical, introrse-latrorse, the filaments fleshy, broader than anthers, densely rusty papillose; whorl III stamens sessile, 0.8 mm tall, the anthers narrowly ovate, 0.4 x 0.2 mm, erect, locelli 2, extrorse-latrorse, the filaments broader than anthers, fleshy, densely


rusty papillose, the basal glands globose, sessile; whorl IV wanting; pistillode wanting. Pistillate inflorescence with indument, color, and branching as in staminate plants, the flowers similar in size and shape; stamens sterile, smaller; ovary glabrous, ovoid; style slender, distinct from ovary; stigma tri-lobed, 0.5 mm diam., papillose. Fruits borme on terete pedicels of up to 1 x 0.3 cm; cupules shallowly hemispherical to patelliform, to 2 x 0.3 cm, glabrous inside and outside, the margins entire; drupes ellipsoid to obovoid, to 4 x 2 cm.

Distribution (Fig. 13) and ecology. Medium- sized to tall trees from Amazonian lowlands as well as upland forests from Peru to Costa Rica, at ca. 100- 1700 m. Flowering and fruiting specimens were collected in almost every month of the year.

Representative specimens examined. COSTA RICA. PUNTARENAS: Buenos Aires, Valle de Diquis, 300 m, 24 Nov 1993 (fl 6), Morales et al. 2083 (F, MO, NY); Osa, trail from Palmar Norte to Jalisco, on divide between Que- brada Batambal and Quebrada Benjamin, SW slope of Fila Retinto, 300-400 m, 9 Dec 1988 (fl ), Grayum et al. 9153 (F, MO, NY).

PANAMA. PANAMA: Cerro Jefe, ca. 1.5 mi along Rio Pacora rd. from jct. with Cerro Jefe rd., 750 m, 23 Jan 1986 (fl 6), McPherson & Merello 8131 (HBG, MO).

COLOMBIA. AMAZONAS: Leticia, Tarapaca, Parque Nacional Natural Amacayacu, 100 m, 25 Jun 1991 (fr), Ru- das et al. 2504 (INPA, MO); Rio Caqueta, frente a Villa Azul, 14 Sep 1989 (fl 6), van Andel et al. 293 (K, MO). HUILA: Parque Nacional Natural Cueva de los Guacharos, Rio Suaza, 1710 m, 21 Jun 1979 (fl d), Henao 152 (B). META: Sierra de la Macarena, Rio Guapaya, 500 m, 21 Jan 1950 (fl Y, fr juv), Philipson et al. 2192 (COL, US, NY). PUTUMAYO: Mocoa, 1500-1670 m, 20 Apr-I May 1994 (fr), Bentancur et al. 5404 (COL, HUA, MO); Rfo Caucaya, Parque Nacional La Paya, 240 m, 10-14 May 1993 (fr), Bernal et al. 2039a (MO).

ECUADOR. CARCHI: Maldonado, Parroquia Tobar Donoso, Reserva Etnica Awa, Sabalera, 900 m, 22 Nov 1992 (fr), Aulestia et al. 819 (MO). ESMERALDAS: Eloy Alfaro, Reserva Ecol6gica Cotacachi-Cayapas, Charco Vicente, Rfo San Miguel, 130 m, 15 Jan 1993 (fl 6), Tipaz 2516 (MO), 150 m, 6-9 Sep 1993 (fr), Palacios & Tirado 11165 (MO). NAPO: El Chaco, Rio Quijos, frente a la Estaci6n de Bombeo del Rio Salado, 1400 m, 11 Sep 1990 (fl d), Palacios 5482 (F, MO); Estaci6n Experimental INIAP-Napo, Payamino, Reserva Floristica "El Chuncho," 250 m, 29 Nov 1986 (fr juv), Neill 7514 (HBG, MO, NY). SUCUMBiOS: Lago Agrio, Reserva Faunistica Cuyabeno, 230 m, 15 Nov 1991 (fl 6), Palacios et al. 8963 (F, HBG, MO), 250 m, 30 Jul 1996 (fr), Palacios 13906 (MO). ZAMORA-CHINCHIPE: Nangaritza, Miazi, Rio Nangaritza, 930 m, 26 Oct 1991 (fl 6), Palacios et al. 8644 (F, MO, NY), hill ca. 1 km upstream from Shaime, 900-1100 m, 16 Feb 1994 (fl 6), van der Werf et al. 13147 (F, HBG, MO).

PERU. AMAZONAS: Bagua, Imaza, Comunidad Agu- aruna de Kusd-Listra, Cerro Apag, 600-700 m, 15 Sep 1996 (fl d ), Diaz et al. 8144 (MO). Cuzco: Cuzco, Camisea, Campamento San Martin-C, Camisea Production Unit, 467 m, 12 Jan 1997 (fl Y), Acevedo et al. 8646 (MO); Quispi- canchis, hills around Rfo Araza between Pan de Azucar and Quince Mil Airport, 643 m, 10 Aug 1991 (fr), Nuiniez 13986 (MO). LORETO: Maynas, Iquitos, Nina Rumi (Rio Nanay), 122 m, Jul 1984 (fl d), Vasquez & Jaramillo 5277 (MO, NY); Requena, Reserva Nacional Pacaya, CochaYarina, 125 m, 16 Jul 1985 (fl d), Vdsquez et al. 6665 (MO, NY). MA- DRE DE DIos: Tambopata, Cusco Amaz6nico, 200-220 m, 19 Jun 1989 (fl d), Phillips et al. 468 (F, MO). SAN

MARTiN: Lamas, Alonso de Alvarado, Fundo las "Flores," 800-900 m, 11 May 1973 (fl d), Schunke 6231 (HBG).

BRAZIL. ACRE: Basin of Rio Purus, near mouth of Rio Macauhan (tributary of Rio Yaco), Aug 1933 (fr), Krukoff 5281 (A, B, F, G, MO, NY, U); Manoel Urbano, Rio Purus, Nova Olinda, 24 Nov 1996 (fr), Silveira et al. 1555 (MO). AMAZONAS: Basin of Rio Jurua, near mouth of Rio Embira (tributary of Rio Tarauaca), 8 Jun 1933 (fl ), Krukoff4714 (A, F, G, K, MO, NY, U, US); Limoeiro, Est. Ecol6gica do Juami-Japurd, 26 Apr 1986 (fr), Cid Ferreira et al. 7227 (F, MO, NY, US). MATO GROSSO: Angustura, Machado River region, source of Jatuarana River, Dec 1931 (st), Krukoff 1565 (A, F, G, MO, NY, P, U, US). PARA: Itaituba, estrada Santar6m-Cuiabd, BR 163, km 1221, 21 May 1983 (fl 6, juv), Amaral et al. 1375 (GH, INPA, MO, NY). ROND6NIA:

Ji-Parand, May 1987 (fl ), Cid Ferreira 9037 (INPA); Porto Velho, 23-24 Apr 1987 (fl d), Cid Ferreira 8900 (F, K, MO, NY).

Local names. Costa Rica: aguacat6n. Colombia: yiyiwa-ageyi (Mui). Ecuador: temachi. Peru: cunshi moena, casha moena, moena blanca, moena de altura. Brazil: louro abacate.

Endlicheria formosa differs from most congeners by its depressed-globose flowers with incurved to erect tepals, but belongs in the E. browniana species group on the basis of dense papillosity inside flowers and sessile stamens with ovate anthers. Also, in the E. browniana species group, very similar flowers are found in E. paradoxa. Indeed, without the scalariform to percurrent tertiaries of E. formosa to contrast the reticulate tertiary venation of E. paradoxa, it would be questionable whether larger flower size in the latter is sufficient to separate the two species. Outside of the E. browniana species group, only E. robusta of the E. sericea species group has similar depressed- globose flowers.

Flowers from the Andean uplands and Central

America tend to be larger, more strongly depressed, and more densely pubescent than those from lowland Amazonia. Leaf size and texture is also variable, but as with flowers, variation appears to be continuous.

46 FLORA NEOTROPICA

7---- -~

__ , . . . . .- - . . . .

AV!

. .. . .. .d . . . . .. t. .- xm, ,. ':es y .

FIG. 13. Distribution of Endlicheria formosa and E. paradoxa.

16. Endlicheria paradoxa Mez, Jahrb. Konigl. Bot. Gart. Berlin. 5: 114. 1889. Type. Peru. Cajamarca: Santa Cruz, Jun 1865 (fl d), Pearce s.n. (holotype: K 59; isotype: K 60).

Trees to 18 m. Branchlets stout, midway along flush 5-7 mm diam., angular, sparsely strigillose, soon glabrous, the surface clearly exposed, dark brown, the hairs short, to 0.15 mm, straight, appressed, greyish white; terminal buds plump, 0.8 X 0.5 cm, grey-sericeous. Leaves alternate, widely and evenly spaced along current flush; petioles slender or robust, to 4 cm long and 0.3-0.6 cm diam., semi- terete to canaliculate, the indument as on branchlets; laminae coriaceous, plane, elliptic to obovate, 10-35 X 6-15 cm, the base acute to cuneate, briefly decurrent, the apex rounded or obtuse, acumen lacking or mucronate for up to 0.3 cm, the margins minutely recurved throughout; upper surface olive-brown, waxy, shining, the primary to fourth-order veins prominent, conspicuous against the lamina; lower surface sparsely strigillose, the hairs as on branchlets,

soon lost, all vein orders raised, their prominence decreasing with rank; secondary veins 6-8 per side, + evenly spaced, slightly more distant around midlamina, ascending at 50-60?, arcuate, distal pairs loop- connected; tertiaries reticulating between secondaries. Staminate inflorescences evenly spaced along current flush in the axils of foliage leaves, to 10 cm long with 6 lateral branches, the highest order dichasial, lax, the flowers distant, the axes sparsely strigillose, distal branches and pedicels more densely so; bracts and bracteoles caducous by anthesis, lanceolate, grey-sericeous; pedicels terete, to 7 mm long, those supporting secondary flowers slightly shorter. Flowers depressed-globose, to 7 mm diam., sparsely grey-strigillose outside; receptacle infundibuliform, 1 x 3 mm, rusty papillose inside. Tepals chartaceous, ovate to triangular, 0.6 X 0.6 mm, inflexed at anthesis, the androecium included. Stamens of whorls I and II sessile, 0.6 mm tall, the anthers ovate, 0.5 X 0.5 mm, glabrous, the apex apiculate, the connectives sharply prolonged between the 2 locelli, these obliquely hem-


ispherical, introrse-latrorse, the filaments ligulate, as broad as anthers, rusty papillose; whorl III stamens sessile, 0.6 mm tall, the anthers ovate, 0.5 X 0.5 mm, erect, locelli 2, extrorse-latrorse, the filaments broader than anthers, rusty papillose, the basal glands absent; whorl IV wanting; pistillode wanting. Pistillate inflorescence with indument, color, and branching as in staminate plants, the flowers smaller, slightly deeper; stamens sterile, smaller; ovary glabrous; style stout, weakly distinguished from ovary; stigma discoid, 0.2 mm diam. Mature fruits unknown. Young fruits bome on claviform pedicels of 1 X 0.4 cm; cupules hemispherical, 0.5 X 1.5 cm, glabrous inside and outside, the margins undulate; drupes ellipsoid, to 1.5 X 0.7 cm.

Distribution (Fig. 13) and ecology. Medium- sized trees known only from lower montane forest around 1900-2000 m in Cajamarca, Peru. Flowering material collected in July and with young fruits in October.

Additional specimens examined. PERU. CAJA- MARCA: San Ignacio, Chirinos, vic. of Pacasmayo, 1900- 2000 m, 2 Oct 1997 (fl Y, frjuv), Campos et al. 4480 (MO), (fl Y), Campos et al. 4490 (MO).

Endlicheria paradoxa has been rediscovered almost 150 years after the mid-19th century type material was collected. The additional collections, Cam- pos et al. 4480 and Campos et al. 4490, have smaller leaves than the type, but depressed-globose flowers and reticulate tertiaries refer them to E. paradoxa. Widespread and frequently collected E. formosa has similar depressed-globose flowers, but these are much smaller, and its tertiary veins are straight to once- forked rather than reticulate. Mez's (1889) isolation of E. paradoxa in monotypic subgen. Hemiajouea is based on supposed sterility of whorl III stamens. Sta- minate flowers are known only from the type material, wherein, as Kostermans (1937) also noted, all stamens appear to be fertile with well-developed locelli.

17. Endlicheria dysodantha (Ruiz & Pav.) Mez, Jahrb. Konigl. Bot. Gart. Berlin 5: 118. 1889. Lau- rus dysodantha Ruiz & Pav., Fl. Peruv. 4: plate 355. 1802. Goeppertia dysodantha (Ruiz & Pav.) Nees, Linnaea 21: 514. 1848. Type. Peru. Huan- uco: Macora, without date (fr), Ruiz & Pav6n s.n. (lectotype: MA-n.v., designated by Kostermans, 1937; isolectotypes: B-n.v., BM-n.v.)

Trees to 10 m. Branchlets slender, midway along flush 2-3 mm diam., distally weakly angular, soon terete, sparsely grey-strigillose, the surface clearly ex-

posed, dark brown to black, the hairs very short, to 0.1 mm, appressed; terminal buds slender, 3 X 0.6 mm, densely chalky-white sericeous. Leaves alternate, widely and evenly spaced along current flush; petioles slender, to 1.5 X 0.2 cm, canaliculate, the indument as on branchlets; laminae membranaceous or chartaceous, plane, elliptic-ovate, 10-17 X 3-8 cm, the base obtuse to acute, attenuate, the apex acute, acuminate for up to 2.5 cm, the margins minutely recurved throughout; upper surface blackish green to dark or olive-brown, minutely punctulate, the primary to fourth-order veins raised, their prominence decreasing with rank; lower surface glabrous except for barbellate tufts in the axils of the sparsely strigillose midrib and secondary veins, all vein orders raised, their prominence decreasing with rank; secondary veins 4-6 per side, + evenly spaced, slightly more distant around midlamina, patent, diverging at 70-85? (more obtusely around midlamina), abruptly ascending after midcourse, brochidodromous; tertiaries roughly horizontal, between secondaries straight to once-forked. Staminate inflorescences evenly spaced along current flush in the axils of foliage leaves, to 5 cm long with 5 lateral branches, branch orders 2-3, the highest order dichasial, lax, the flowers distant, the axes sparsely strigillose; bracts and bracteoles caducous by anthesis, narrowly ovate to lanceolate, the indument as on axes; pedicels terete, to 5 mm long, those supporting secondary flowers slightly shorter. Flowers hypocrateriform, 3.5 mm diam., sparsely grey-strigillose outside, the indument denser towards tepals; receptacle infundibuliform, 1 X 1 mm, sparsely grey-pilose inside, the hairs straight, erect. Tepals membranaceous, ovate, 1.2 X 0.8 mm, spreading to recurved at anthesis, the inner surface glabrous except for the minutely papillose tips. Stamens of whorls I and II broadly stipitate, 0.6 mm tall, the anthers broadly ovate, 0.3 X 0.5 mm, glabrous, the apex apiculate, the connectives prolonged between the 2 locelli, these obliquely hemispherical, introrse- latrorse, the filaments laminar, slightly narrower than anthers, glabrous; whorl III stamens columnar, 0.6 mm tall, the anthers ovate, 0.3 X 0.5 mm, erect, locelli 2, extrorse-latrorse, the filaments as broad as anthers, laminar, glabrous, the basal glands sessile, globose, relatively large, filling the space between filaments; whorl IV wanting; pistillode filiform. Pistillate inflorescence with indument, color, and branching as in staminate plants, the flowers similar in size and shape; stamens sterile, smaller; ovary glabrous, ovoid; style slender, distinct from ovary; stigma discoid, 0.3 mm diam. Fruits borne on narrowly claviform pedicels of up to 1.5 X 0.4 cm; cupules infundibuliform, to I x

48 FLORA NEOTROPICA

1.5 cm, glabrous inside and outside, the margins entire; drupes ellipsoid, to 2 X 1.4 cm.

Distribution (Fig. 14) and ecology. Small trees or shrubs from inundated and well-drained soils in SW Amazonia and adjacent eastern Andean foothills. Flowering from June to February, fruits available year round.

Representative specimens examined. ECUADOR. NAPO: Aguarico, Samona Yuturi, Rio Napo, 200 m, 11 Nov 1991 (fl 6), Neill & Rojas 9950 (HBG, MO, NY); Aniangu, Parque Nacional Yasuni, 14 Jul 1982 (fr), Luteyn et al. 8680 (F, MO, NY). PASTAZA: Ceilan, Path from Ceilan to Rfo Conoaco on S side of Rio Curaray, 200 m, 7 Jun 1980 (fr), Brandbyge & Asanza 31785 (MO); Pozo Villana 2 de Arco, 2 km del pueblo de Villano, 400 m, 3 Dec 1991 (fl 9), Tipaz et al. 581 (F, HBG, MO, NY).

PERU. Cuzco: Paucartambo, Pilcopata, Villa Carmen, 720 m, 9 Feb 1975 (fr), Plowman & Davis 5098 (F); Rfo Mapituriani, La Convenci6n, Cordillera Vilcabamba, 670 m, 31 Jul 1968 (fl 6), Dudley 11493 (F, MO, NY). HuANuco: Codo de Pozuzo, 300 m, 22 Oct 1982 (fl 9, fr juv), Foster 9394 (MO); Pachitea, Pucallpa, Sira Mtns., 260-400 m, 31 May 1988 (fr), Wallnofer 17-31588 (MO). JUNiN: Rio Ne- gro to Satipo, 400 m, 17 Aug 1960 (fl 9, fr), Woytkowski 5829 (G, MO). LORETO: Previsto, 420 m, Oct 1962 (fl 6), Woytkowski 7593 (GH, K, MO, US); Rio Santiago, s.d. (fl 6), Tessmann 3999 (F, G, NY). MADRE DE DIos: Manui, Parque Nacional Manui, Cocha Cashu Station, Rio Manu, 300-400 m, 17-24 Aug 1974 (fl 6), Foster 3371 (F, HBG, MO, NY); Tambopata, Explorer's Inn Tourist camp at jct. of Rios La Torre and Tambopata, SW of Puerto Maldonado, Capirona Trail, 600 m, 25 Jan 1989 (fr), Smith et al. 1599 (F, G, K, MO, NY, US). PUNO: Ridge between Rio Candamo and Rio Guacamayo, 40-600 m, 22 May 1992 (fr), Gentry et al. 76951 (MO). SAN MARTiN: Mariscal Caceres, Toca- che Nuevo, Quebrada de Huaquisha, Rio Huallaga, Puerto Pisana, 15 Mar 1971 (fr), Schunke 4766 (F, G, INPA, MO, NY), 400-450 m, 29 Jun 1974 (fl 9, fr), Schunke 7053 (HBG, MO). UCAYALI: Purns, Rio Curanja, circa la comunidad nativa de Colombiana, 250 m, 24 Feb 2000 (fr), Graham & Schunke 1099 (MO).

BRAZIL. AcRE: Caramari Amazonas, Rio Jurua, Lago da Cigana, 150 m, 22 Aug 1986 (fr), Croat 62517 (HBG, MO); Tarauaca, Rio Tarauaca', Seringal Tamandare, Colo-

caqao Santa Maria, 25 Dec 1995 (fr), Ehringhaus et al. 396 (MO). AMAZONAS: Basin of Rio Jurua, near mouth of Rio Embira (tributary of Rio Tarauaca), 12 Jun 1933 (fl 6), Kru- koff4767 (A, F, G, K, MO, NY, U); Rio Purus, Ponto Alegre, 8 Apr 1904 (fr), Huber s.n. (NY).

BOLIVIA. COCHABAMBA: Prov. Carrasco, Valle de Sa- jta, Estacion Piscicola Pirahiba, 215 m, 21 Mar 1995 (fr), Ritter 1706 (MO). LA PAZ: San Antonio, Dec 1907 (fl 6), Buchtien 1986 (US); Prov. Larecaja, Tuiri (near Mapiri, Rio Mapiri), 490-750 m, 12-30 Sep 1939 (st), Krukoff 10787 (A, F, MO); Mapiri, May 1886 (fr), Rusby 2671 (F, NY). SANTA CRUZ: Prov. Ichilo, Parque Nacional Ambor6, 0-2 km SW of El Carmen, 360 m, 9 Nov 1990 (fl 6), Nee 39834

(MO, NY, U), 370-450 m, 17 May 1993 (fr), Vargas et al. 2478 (MO, NY).

Local names. Peru: moenito amarillo, palometa micuna.

Endlicheria dysodantha is immediately recognized by its dark-drying subglabrous leaves with barbellate tufts of hairs in the axils of secondary veins below. Close relationship with E. paniculata and allies is suggested by stipitate whorl I and II stamens and pilose indument inside flowers. This has weak molecular support (Fig. 2), but I prefer to place E. dysodantha close to the E. browniana species group. Like other species therein, in E. dysodantha all anthers are ovate with obliquely hemispherical locelli, and slender claviform pedicels support shallowly hemispherical cupules. Its vestiture of extremely short (0.1 mm) appressed hairs is also characteristic of the E. browniana species group.

18. Endlicheria tomentosa Chanderbali, sp. nov. Type. Peru. Cajamarca, Huarango, El Triunfo (property of Edilberto Delgado), 1500-1800 m, 13 Jul 1996 (fl d), Campos et al. 2938 (holotype:

MO; isotypes: F, G, HBG, MO, NY, U). Fig. 15

A Endlicheriae dysodanthae ramulis et foliis subtus pilis erectis statim diagnoscenda.

Trees to 6 m. Branchlets slender, midway along flush 2-3 mm diam., distally weakly angular, soon terete, densely tomentose, the surface barely visible, the hairs short, to 0.3 mm, straight to crooked, erect to ascending, greenish yellow; terminal buds slender, 3 X 1 mm, densely pubescent, the hairs as on branchlets, ascending. Leaves alternate, widely and evenly spaced along current flush; petioles slender, to 1 X 0.1 cm, terete, the indument as on branchlets; laminae membranaceous or chartaceous, plane, narrow elliptic, 6-10 X 1.5-3 cm, the base acute, briefly decurrent, the apex acute, acuminate for up to 1 cm, the margins minutely recurved throughout; upper surface greyish green to olive-brown, waxy, the primary to fourth-order veins raised, their prominence decreasing with rank; lower surface moderately tomentose, the hairs 0.3 mm long, crookedly erect; denser and forming barbellate tufts in the axils of the midrib and secondaries, all vein orders raised, their prominence decreasing with rank; secondary veins 4-6 per side, ? evenly spaced, slightly more distant around midlamina, ascending at 50-600 (more obtusely around midlamina), arcuate, distal pairs loop-connected; tertiaries laxly reticulating between secondaries. Sta-

SYST'EMATIC TREATMENT 49

- , X bb!; 7 ; X 0 j 9s,v o nt |f 0 r r * S i- k . s 0 T

'LI . .14. Ditibto of .nlcei .yoata E. to etoa ... arnilr,ad..aahoo e

minate inflorescences evenly spaced along current flush in the axils of foliage leaves, to 5 cm long with 3 lateral branches, branch orders 2-3, the highest order dichasial, lax, the flowers distant, the axes sparsely grey-strigillose; bracts and bracteoles caducous by anthesis, lanceolate, the indument as on axes; pedicels terete, to 3 mm long, those supporting secondary flowers slightly shorter. Flowers hypocrateriform, 3.5 mm diam., sparsely grey-strigillose outside; receptacle infundibuliform, 1 X 1 mm, densely pubescent inside, the hairs silvery, straight, rigidly erect. Tepals chartaceous, ovate, 1.2 X 1 mm, spreading to recurved at anthesis, the inner surface sparsely silvery strigillose, the margins and apex inside minutely papillose. Stamens of whorls I and II broadly stipitate, 0.6 mm tall, the anthers broadly ovate, 0.3 X 0.5 mm, glabrous, the apex apiculate, the connectives prolonged between the 2 locelli, these obliquely hemispherical, introrse-latrorse, the filaments laminar, narrower than anthers, glabrous; whorl III stamens sessile, 0.6 mm tall, the anthers

ovate, 0.3 X 0.5 mm, erect, locelli 2, extrorse-latrorse, the filaments broader than anthers, ligulate, glabrous, the basal glands sessile, relatively large, filling the space between filaments; whorl IV wanting; pistillode fusiform. Pistillate plants unknown.

Distribution (Fig. 14) and ecology. Known only from the type material, taken from a small tree found flowering in July in the lower montane forests of Ca- jamarca, Peru.

Flowers of Endlicheria tomentosa are identical to those of E. dysodantha, and in both species the axils of secondary veins on the underside of dark-drying leaves are barbellate. However, in E. tomentosa these barbellate tufts are all but lost amongst a dense erect indumentum, whereas in E. dysodantha they are conspicuous against the background of sparse appressed hairs elsewhere on the lower leaf surface. Laxly reticulate tertiaries in E. tomentosa also contrast with the scalariform pattern formed by the tertiaries of E. dysodantha. If E. tomentosa is correctly assigned to the

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19. Endlicheria arunciflora (Meisn.) Mez, Jahrb. Konigl. Bot. Gart. Berlin 5: 131. 1889. Ampelo- daphne aruncifiora Meisn., DC. Prodr. 15(1): 81. 1864. Type. Venezuela. Amazonas: Rio Negro, San Carlos, 1853-1854 (fl d), Spruce 3061 (lectotype, designated by Kostermans, 1937: K; isolectotypes: B-n.v., BM-n.v., K, NY-n.v., P-n.v.).

Trees to 10 m. Branchlets slender to stout, midway along flush 3-6 mm diam., terete, densely pubescent, the surface barely visible to concealed by the indument cover, the hairs relatively long, to 0.6 mm, straight to crooked, erect, reddish or a few grey; terminal buds plump, 5 X 3 mm, densely rusty pubescent, the hairs as on branchlets, straight, appressed to ascending. Leaves closely spiraled at tips of current flush; petioles robust, to 2 X 0.3 cm, semi-terete, the indument as on branchlets; laminae coriaceous, subbullate, lanceolate to oblong, or elliptic, 10-25 X 2-8 cm, the base obtuse to rounded, the apex obtuse to acute, acuminate for up to 1.5 cm, the margins minutely recurved throughout; upper surface green to olive-brown, minutely punctulate, the midrib prominent, the secondaries impressed, the higher-order venation immersed, inconspicuous against the lamina; lower surface sparsely pubescent, the hairs 0.6 mm long, sprawling to weakly ascending, rusty, denser on main veins, soon lost, all vein orders raised, their prominence decreasing with rank; secondary veins 14-20 per side, ? evenly spaced, slightly more distant around midlamina, patent, diverging at 70-85? (more obtusely around midlamina), abruptly ascending after midcourse, brochidodromous, or strongly loop-connected by midlamina; tertiaries oblique to midrib, between secondaries straight to forked. Sta- minate inflorescences distally clustered in the axils of cataphylls, to 20 cm long with 15 lateral branches, branch orders 3-4, the highest order botryoid, or irregular, lax, the flowers loosely crowded, the axes sparsely rusty tomentose, the hairs restricted to first- and second-order branches, soon lost; bracts and bracteoles persistent at anthesis, lanceolate, sparsely rusty tomentose, glabrescent; pedicels terete, to 2 mm long, those supporting secondary flowers slightly shorter. Flowers campanulate, 2 mm diam., glabrous outside; receptacle cyathiform, 0.6 x 0.6 mm, glabrous inside. Tepals chartaceous, ligulate, 1.3 X 0.6 mm, spreading

to recurved at anthesis, the inner surface papillose near tips, otherwise glabrous. Stamens of whorls I and II stipitate, 0.6 mm tall, the anthers transversely oblong, 0.3 X 0.4 mm, glabrous, the apex truncate or emarginate, the connectives level with or reduced between the 2 locelli, these suborbicular, introrse, the filaments laminar, much narrower than anthers, sparsely grey-villose; whorl III stamens stipitate, 0.6 mm tall, the anthers equal to outer whorls, erect, locelli 2, extrorse-latrorse, the filaments narrower than anthers, laminar, sparsely grey-villose, the basal glands sessile, globose; whorl IV wanting; pistillode filiform. Pistillate inflorescence unknown, old pistillate flowers glabrous, stamens sterile, smaller; ovary glabrous, ovoid; style stout, weakly distinguished from ovary; stigma broadly tri-lobed, 0.6 mm diam. Fruits borne on terete pedicels of up to 5 X 3 mm; cupules hemispherical, to 0.7 X 1.5 cm, glabrous inside and outside, the margins entire; drupes ellipsoid, to 2 x 1 cm.

Distribution (Fig. 14) and ecology. Medium- sized trees in flooded forests of the upper Rio Negro area in Brazil and adjacent Venezuela at 100-200 m. Flowering July to October, fruits available from No- vember to March.

Additional specimens examined. VENEZUELA. AMAZONAS: Rio Negro, Cerro Cucuy, 2 Mar 1944 (fr), Baldwin 3211 (A, MO, US); Rio Casiquiare, Cafno Pimichin, 128 m, 6-19 Jul 1969 (fl 6), Bunting et al. 4079 (MO, U); lower Rio Baria, 80 m, 22-23 Jul 1984 (fl 6), Davidse 27595 (MO, NY, U); San Carlos de Rio Negro, 120 m, 16 May 1979 (fr), Liesner 7466 (MO, VEN); Atabapo, Nov 1989 (fr), Marin 438 (MO); Rio Yatua, 2 Dec 1984 (fl Y, fr juv), Stergios & Aymard 7524 (MO, NY); Rio Orinoco, Cano Tama-Tama, 125-150 m, 23 Jun 1959 (fr), Wurdack & Adderley 43153 (NY); Rio Pacimoni-RioYatua, along Rio Yatua between mouth and Laja Catipan, 110-120 m, 13 Jul 1959 (fl d), Wurdack & Adderley 43435 (F, K, NY, U, US).

BRAZIL. AMAZONAS: Rio Negro, Rio Curicuriary, 7 Sep 1979 (fl d), Kubitzki et al. 79-172 (G, HBG, INPA), 18 May 1973 (fr), Silva et al. 1707 (HBG, INPA); Caman- aus, Rio Miua, 30 Oct 1978 (fl 6), Nascimento 826 (HBG, NY); Rio Curicuriari, 13 Jul 1979 (fl d), Poole 1972 (HBG, K, MG, MO, NY).

The subglabrous inflorescences of Endlicheria arunciflora occur in only two other species of Endli- cheria, both also of the Ampelodaphne species group. Of the two, Endlicheria verticillata has prominent tertiaries above, i.e., it belongs to the E. bracteata subgroup, while E. arachnocome has unique arachnoid indument and never shows the brochidodromy typical of E. arunciflora. Brochidodromy appears to be unstable in E. arunciflora since the basalmost second-

52 FLORA NEOTROPICA

aries sometimes lack loop connections (e.g., Poole 1972). Even so, E. arunciflora is easily distinguished from E. arachnocome by its rusty tomentose branchlets.

20. Endlicheria arachnocome Chanderbali, sp. nov. Type. Peru. Loreto: Maynas, Rio Nanay, Pun- chana, 90 m, 21 Apr 1993 (fl d), Rimachi 10532 (holotype: MO; isotype: NY) Fig. 16

Species habitu cum Endlicheriae arunciflorae optime congruens, sed differt ramuliis et foliis subtus pilis arach- noides praeditis.

Trees to 12 m. Branchlets slender to stout, midway along flush 3-6 mm diam., terete, densely pubescent, the surface barely visible to concealed by the indument cover, the hairs relatively long, to 0.6 mm, crooked, sprawling, greyish white; terminal buds plump, 5 X 3 mm, densely pubescent, the hairs straight, appressed to ascending, white. Leaves closely spiraled at tips of current flush; petioles robust, to 3 X 0.4 cm, semi-terete, the indument as on branchlets; laminae chartaceous, plane, broadly elliptic, 15-30 X 5-11 cm, the base obtuse to acute, the apex acute, acuminate for up to 2 cm, the margins minutely recurved throughout; upper surface light greyish green to olive-brown, minutely punctulate, the midrib prominent, the secondaries prominulous, the higher-order venation immersed, inconspicuous against the lamina; lower surface sparsely pubescent, the hairs 0.6 mm long, sprawling to weakly ascending, greyish white, denser on main veins, all vein orders raised, their prominence decreasing with rank; secondary veins 9-11 per side, ? evenly spaced, slightly more distant around midlamina, ascending at 50-60? (more obtusely around midlamina), arcuate to arching after midcourse, distal pairs loop-connected; tertiaries oblique to midrib, between secondaries straight to arched. Staminate inflorescences distally clustered in the axils of cataphylls, to 35 cm long with 10 lateral branches, branch orders 3-4, the highest order botryoid, or irregular, the flowers loosely crowded, the axes moderately grey-tomentose, the indument thinning distally, lost by third-order branches; bracts and bracteoles persistent at anthesis, lanceolate, the indument as on axes; pedicels terete, to 2 mm long, those supporting secondary flowers slightly shorter. Flowers campanulate, 2 mm diam., glabrous outside; receptacle cyathiform, 0.6 X 0.6 mm, glabrous inside. Tepals chartaceous, ligulate, 1.3 X 0.6 mm, spreading to recurved at anthesis, the inner surface glabrous, the margins minutely papillose near

apex. Stamens of whorls I and II stipitate, 0.6 mm tall, the anthers transversely oblong, 0.3 X 0.4 mm, glabrous, the apex truncate or emarginate, the connectives level with or reduced between the 2 locelli, these suborbicular, introrse, the filaments laminar, much narrower than anthers, sparsely grey-villose; whorl III stamens stipitate, 0.6 mm tall, the anthers equal to outer whorls, erect, locelli 2, extrorse- latrorse, the filaments narrower than anthers, laminar, sparsely grey-villose, the basal glands sessile, globose; whorl IV wanting; pistillode filiform. Pistillate inflorescence with indument, color, and branching as in staminate plants, the flowers slightly deeper; stamens sterile, smaller; ovary glabrous, ovoid; style slender, distinct from ovary; stigma broadly tri-lobed, reniform, 0.6 mm diam. Fruits borne on terete pedicels of up to 6 X 4 mm; cupules hemispherical, to 0.5 X 1.3 cm, glabrous inside and outside, the margins entire; drupes ellipsoid, to 2 X 1 cm.

Distribution (Fig. 14) and ecology. Small trees from seasonally flooded (tahuampa) lowland forests near Iquitos, Peru, at ca. 100 m. Flowering February to October, fruiting November to March.

Additional specimens examined. PERU. Loreto: May- nas, Mishana, Santa Maria de Nanay, 90 m, 2 Oct 1990 (fl 6), Pipoly et al. 12727 (MO); Rfo Nanay, 130 m, 26 Jul 1984 (fl 6), Vasquez et al. 5434 (F, MO, NY), 140 m, 18 Aug 1988 (fl Y), van der Werif et al. 10207 (HBG, MO, NY), Morona Cocha, 120 m, 21 Mar 1977 (fr), Gentry et al. 18525 (F, MO); Iquitos-Nauta, km 10, 90 m, 3 Apr 1989 (fl 6), Rimachi 9113 (MO); Mishuyacu, near Iquitos, 100 m, Apr 1930 (fl 6), Klug 1264 (F, NY, US), May-Jun 1930 (fl 6), Klug 1403 (F, NY, US); Iquitos, RIo Itaya, Buena Suerte, 122 m, 16 Nov 1986 (fr), Vasquez & Jaramillo 8414 (F, MO, NY); Rfo Momon, 95 m, 7 Feb 1985 (fr), Rimachi 7739 (US); Puerto Almendras, 130 m, 22 Feb 1979 (st), Gentry et al. 24873 (HBG, MO), 122 m, 16 Jul 1984 (fl 6), Vdsquez & Jaramillo 5253 (MO, NY), 14 Aug 1984 (fl 6), Vasquez & Jaramillo 5457 (MO), 2 Jun 1986 (fl 6), Vdsquez & Jaramillo 7620 (MO), 4 Jun 1986 (fl 6), Vdsquez & Jar- amillo 7647 (MO), 20 Jul 1982 (fl 6), Vdsquez & Jaramillo 3157 (MO); Punchana, 90 m, 25 Feb 1993 (fl 6), Rimachi 10402 (MO); Alto Nanay, 5 Mar 1968 (fr), Simpson 782 (G, US); Requena, Sapuena, Bagazan-Rfo Ucayali, 130 m, 13 Jan 1987 (fr), Vdsquez & Jaramillo 8766 (MO).

Material of Endlicheria arachnocome was previously distributed as E. arunciflora. The two species share subglabrous inflorescences and immersed tertiary venation above, and without noting the authen- ticity of the arachnoid indument of E. arachnocome its representatives could be thought diseased E. arunciflora. However, sprawling greyish white hairs and eucamptodromous secondaries consistently distinguish this species from rusty tomentose and brochi-


3.~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~o

0.5 mm

0.3 -m

FIG. 16. Endlicheria arachnocome (Rimachi 10532). A. Habit. B. Leaf base below. C. Flower. D. Flower l.s. E.

Whorl I stamen seen from within. F. Whorl HI stamen seen from without.

54 FLORA NEOTROPICA

dodromous E. arunciflora. Further, with E. arachnocome known only from Peruvian tributaries of the Rio Amazonas and E. arunciflora only from the upper Rio Negro and Rio Orinoco around the Brazil-Venezuela border, these two riparian species appear to be allopatric in different drainage systems.

21. Endlicheria arenosa Chanderbali, sp. nov. Type. Brazil. Amazonas: Mun. Manaus, Estrada Ma- naus-Caracaraf, km 130, Igarape Lages, 11 May 1974 (fl d), Nelson & Lima P21108 (holotype: MO; isotypes: F, GH, HBG, INPA, K, NY, US).

Fig. 17

Endlicheriae macrophyllae accedens, ob pilis perbrevi- bus ab eo removendum.

Trees to 5m. Branchlets usually stout, midway along flush (3-)5-8 mm diam., terete, densely tomentose, the surface concealed by the indument cover, the hairs relatively short, to 0.2 mm, erect, rust brown to dark red; terminal buds plump, 1 X 0.6 mm, rusty tomentose, the hairs as on branchlets, erect, crisped to crooked. Leaves closely spiraled at tips of current flush; petioles robust, to 3 X 0.4 cm, semi-terete, the indument as on branchlets; laminae stiff chartaceous, plane to subbullate, obovate to elliptic, 10-27 X 3- 10 cm, the base acute to obtuse, the apex acute, acuminate for up to 2 cm, the margins minutely recurved throughout; upper surface olive to reddish brown, waxy, the midrib and secondaries prominulous, the higher-order veins immersed; lower surface sparsely pubescent, the hairs on the upper surface of the midrib and secondaries crooked to crisped, dark red, soon falling, those on the sides of the main veins and lamina light brown to translucent, 0.2 mm long, straight, erect, persistent, denser on main veins, all vein orders raised, their prominence decreasing with rank; secondary veins 8-11 per side, ? evenly spaced, slightly more distant around midlamina, patent, diverging at 70-85? (more acutely towards apex), arching after midcourse, distal pairs loop-connected; tertiaries oblique to midrib, between secondaries once-forked to straight. Staminate inflorescences distally clustered in the axils of cataphylls, to 25 cm long with 12 lateral branches, branch orders 2-4, the highest order botryoid, lax, the flowers distant, the axes rusty tomentose, the indument moderately dense, thinning distally; bracts and bracteoles persistent at anthesis, lanceolate, the indument as on axes; pedicels slender, terete, to 1 mm long, reduced to lacking below secondary flowers. Flowers campanulate, to 1.5 mm diam., sparsely pubescent outside; receptacle cyathi-

form, 0.7 X 0.6 mm, slightly constricted below tepals, glabrous inside. Tepals membranaceous, ovate, 0.6 X 0.5 mm, spreading to recurved at anthesis, the inner surface glabrous except for sparsely papillose tips and distal margins. Stamens of whorls I and II stipitate, 0.5 mm tall, the anthers transversely oblong, 0.25 x 0.3 mm, glabrous, the apex truncate to emarginate, the connectives level with or reduced between the 2 locelli, these suborbicular, introrse, the filaments laminar, narrower than anthers, glabrous; whorl III stamens stipitate, 0.5 mm tall, the anthers depressed- oblong, 0.2 X 0.3 mm, bowed towards the outer whorls, locelli 2, extrorse-latrorse, the filaments narrower than anthers, laminar, silvery strigose on the adaxial side, the basal glands stipitate, globose, relatively large, filling the space between filaments of outer and inner whorls; whorl IV wanting; pistillode fusiform. Pistillate inflorescence with indument and color as in staminate plants, but shorter and with fewer lateral branches, the flowers similar in size and shape; stamens sterile, smaller; ovary ellipsoid, glabrous; style slender, distinct from ovary, 0.3 mm long; stigma tri-lobed, papillose, 0.25 mm diam. Fruits bome on short terete pedicels of up to 2 mm, rarely claviform and 1 cm long; cupules hemispherical, to 0.6 x 1 cm, glabrous inside and outside, the margins entire; drupes ovoid, to 1 X 0.8 cm.

Distribution (Fig. 18) and ecology. Small trees or shrubs from white sand (campina) forests in lowland Amazonia and savanna formations of north- western South America at 100-350 m. Flowering specimens have been collected throughout the year, and fruiting material from September to April.

Additional specimens examined. COLOMBIA. AMA- ZONAS: Corregimiento Puerto Santander, Rio Caqueta, 100- 350 m, 13-21 Feb 1997 (fl d ), Arbeldtez & Sueroke 711 (U), (st), Arbelaez & Sueroke 715 (U). VAUPES: Rio Kuduyari, Cerro Yapoboda, 450 m, 5-6 Oct 1951 (fl d), Schultes & Cabrera 14356 (US), quarzite savannah near headwaters, Apr 1953 (fr), Schultes & Cabrera 20003 (US).

VENEZUELA. AMAZONAS: Atabapo, 120 m, 8 Dec 1978 (fl i, fr), Huber & Tillett 2975 (MO, NY); Rio Ori- noco, SerraYapacana base, 100 m, 1 Jan 1951 (fr), Maguire et al. 30601 (NY); Sabana de Basil, Cano Chimoni, 22 Feb 1989 (fr), Stergios et al. 13295 (HBG, NY); sabana de arena blanca al N de San Juan de Ucata, 70-80 m, 20 Oct 1989 (fl 5), Romero & Melgueiro 2129 (GH, MO).

BRAZIL. AMAZONAS: Parque Nacional do Jad, Cam- pina do Pataud, 30 Aug 1998 (fl 5), Vicentini et al. 1317A (INPA, MO), 2 Sep 1998 (fl Y), Vicentini et al. 1359 (INPA, MO), 6 Sep 1998 (fl d), Vicentini et al. 1394 (INPA, MO); Itapiranga, Rio Uatuma, 13 Aug 1979 (fl d), Cid Ferreira et al. 266 (HBG, INPA, MO, NY, US), Igarap6 Catitu, 20 Aug 1979 (fl d), Cid Ferreira 582 (HBG, MO, NY); Estrada


,4~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~4.

FIG. 17. Endlicheria arenosa (Nelson & Lima P21108). A. Habit. B. Inflorescence. C. Leaf base below. D. Flower E. Flower ls. F. Pistillode. G. Whorl I stamen seen from within. H. Whorl III stamen seen from without.

56 FLORA NEOTROPICA

Mr Al~~~~~~~~~~~

17~~~~~~~~~:yk

A~~~~~~~~~~~..x

4 .......... ...............~

-IA1 DstibtinofEnliheiamarphll ad aen,

da BR 17, km 20, 27 Jul 1961 (fl T), Rodrigues & Coe'lho 3033 (NY); Estrada da Praia Dourada, Tarumizinho, 29 Jun 1976 (fl Y), Monteiro 1246 (INPA); Estrada do Ponta Ne- gra-Tarumi, 24 Oct 1966 (fr), Allen et al. 323 (HBG, INPA, NY); Estrada Manaus-Caracaraf, km 11, near Igarape Leao, 25 Aug 1959 (fl 6), Rodrigues & Chagas 1259 (NY), 10 May 1961 (fl 6), Rodrigues & Coilho 2564 (INPA, NY), 17 Mar 1967 (fr), Prance et al. 4678 (HBG, MG, NY, R), km 130, 15 Feb 1974 (fr), Loureiro et al. s.n. INPA 47996 (INPA), 18 Feb 1974 (fr), Steward etaaL P20357(GH, HBG, INPA, K, MO, NY, R, S, US), km 125, 19 Feb 1979 (fr), Coelho et al. 896 (INPA); Estrada Manaus-Itacoatiara, kmn 181, 21 Dec 1966 (fr), Prance et al. 3817 (HBG, MG, NY); Igarap6 da Cachoeira Alta do Tarum&, 6 Jun 1961 (fl 6), Rodrigues & Chagas 2743 (INPA, NY), 12 Jun 1961 (fl d), Rodrigues & Chagas 2783 (INPA, NY), 30 Jul 1962 (fl d), Rodrigues & Chagas 4558A (INPA, NY); Igarapd da Cachoeira Grande, 28 Aug 1962 (fl d), Rodrigues & Chagas 4611 (NY); Igarap6 de Bolivia, 28 Dec 1955 (fr), Cotlho s.n. INPA 3206 (INPA, MG, NY); Igarape de Riacho Grande, 14 Aug 1957 (fl 6), Rodrigues 508 (INPA); Parque 10, 5 Jun 1956 (fl,2), Coilho s.n. INPA 3895 (INPA, MO, NY);

Ponta Negra, 12 May 1961 (fl 3), Rodrigues & Coelho 2588 (INPA, NY), 24 Jan 1962 (fr), Rodrigues & Lima 4124 (G, INPA, NY), 2 Dec 1960 (fr), Rodrigues et al. 1980 (INPA, MO, NY), 24 May 1972 (fr), Coilho 160 (INPA); Reserva Florestal Ducke, campinarana Acari, 21 Aug 1979 (st), Ku- bitzki & Poppendieck 79-29 (HBG, NY); Rio Negro, May 1910 (fl 3 ), Ule 8848 (G, L, NY, MG, US); Nova Prainha, Rio Aripuana, Projeto RADAM/BRAZIL, SB-20-ZB, Ponto 02, 10 Aug 1976 (fl 6), Mota & Monteiro s.n. INPA 61281 (INPA), Ponto 10, 9 Jul 1976 (fl 6), Mota s.n. INPA 60608 (INPA), Ponto 15, 15 Sep 1976 (fl 6), Ramos etal. s.n. INPA 62154 (INPA); Presidente Figueiredo, Rio Uatuma, UHE Balbina, 26 Sep 1988 (fl 6), Lopes et al. 124 (INPA); Rio Aripuana, 53 km W along Trans-Amazon Hwy., 27 Jun 1979 (fl 6), Calderon et al. 2704 (MPA, NY, MO, US); Serra Araca, Rio Jauari, 29 Feb 1984 (fl juv), Rodrigues et al. 10494 (INPA, K, MO, NY). PARA: Campos do E de Faro, Aug 1907 (fl 6), Ducke 8429 (MG); Campos do Mariapiry, Jul 1912 (fl 6), Ducke s.n. R 19970 (HBG, U); Campos do Tigre, Dec 1919 (fl 6), Ducke s.n. R 11373 (U); Estrada de T. Santa, km 65, 16 Sep 1988 (fl 6), Soares 462 (INPA); Oriximinu, Rio Mapuera, Campina das Tres Ilhas, 19 Aug


1986 (fl d), Cid Ferreira et al. 7803 (F, GH, NY, MO, US), 25 Sep 1987 (fr), Farney et al. 2012 (MO), 25 Nov 1987 (fr), Cid Ferreira 9676 (INPA, MO).


Endlicheria arenosa is conspicuous in the Ampe- lodaphne species group for its indument of stiffly erect straight hairs only ca. 0.2 mm in length. Such short hairs are occasionally found in E. directonervia but E. arenosa is best compared with species with which it shares immersed tertiary venation above and rusty tomentose branchlets and inflorescences. Of these, E. macrophylla has similar flowers, but hairs at least four times as long provide a soft pilose indument on the leaves below. Further, E. macrophylla appears to be a species of flooded forests while E. arenosa has been found only on well-drained sands.

Specimens from open savannas and scrub vegetation in Brazil (Rodrigues et al. 10494), Colombia (Arbeldez & Sueroke 711, 715) and Venezuela (Huber & Tillett 2975, Romero & Melgueiro 2129, Stergios et al. 13295) depart from the typical campina forest material by their denser, darker red indument. They also maintain crooked hairs on the veins below beyond leaf maturity, but the short erect hairs that persist on older leaves are typical of E. arenosa.

22. Endlicheria macrophylla (Meisn.) Mez, Jahrb. Konigl. Bot. Gart. Berlin. 5: 128. Ampelodaphne macrophylla Meisn., DC. Prodr. 15(1): 81. 1864. Type. Brazil. Amazonas: Manaus, Apr 1851 (fl d ), Spruce 1453 (lectotype, designated by Kos- termans, 1937: K; isolectotypes: B-n.v., BM-n.v., C-n.v., E, G, GH-n.v., L.-n.v., NY-n.v., P, W- n.v.).

Trees to 15 m. Branchlets slender to stout, midway along flush 3-6 mm diam., terete, densely rusty tomentose, the surface barely visible to concealed by the indument cover, the hairs relatively short, to 0.5 mm, erect, crooked, matted; terminal buds plump, 3 X 2 mm, the indument as on branchlets. Leaves closely spiraled at tips of current flush; petioles robust, to 3.5 X 0.5 cm, semi-terete, the indument as on branchlets; laminae stiff chartaceous, plane, ovate to ovate-elliptic, 15-25 X 5-10 cm, the base rounded to acute, the apex acute, acuminate for up to 1.5 cm, the margins minutely recurved throughout; upper surface greyish green to olive-brown, minutely punctulate, the midrib and secondaries prominulous, the higher-order venation immersed, inconspicuous against the lamina; lower surface densely pubescent,

the hairs 1 mm long, erect, straight or curved near the tip, uniformly distributed, all vein orders raised, their prominence decreasing with rank; secondary veins 8- 11 per side, ? evenly spaced, slightly more distant around midlamina, ascending at 50 60? (more obtusely around midlamina), arcuate, the lowermost pair occasionally asymmetric (one or both merging with margin at base), distal pairs loop-connected; tertiaries oblique to midrib, between secondaries straight to forked. Staminate inflorescences distally clustered in the axils of cataphylls, to 30 cm long with 12 lateral branches, branch orders 3-4, the highest order botryoid, or irregular, the flowers loosely crowded, the axes densely rusty villose, the hairs 1 mm long, crooked; bracts and bracteoles persistent at anthesis, lanceolate, greyish white villose; pedicels terete, to 1 mm long, those supporting secondary flowers slightly shorter. Flowers campanulate, 2 mm diam., sparsely greyish white villose outside; receptacle narrowly cyathiform, 1 X 0.3 mm, slightly constricted below tepals, glabrous inside. Tepals membranaceous, ligulate, 1 X 0.6 mm (the inner whorl slightly broader), ascending to recurved at anthesis, the inner surface lightly papillose near the apex, otherwise glabrous. Stamens of whorls I and II stipitate, 0.6 mm tall, the anthers transversely oblong, 0.3 X 0.4 mm, glabrous, the apex truncate to emarginate, the connectives level with or reduced between the 2 locelli, these suborbicular, introrse, the filaments laminar, much narrower than anthers, sparsely grey-villose; whorl III stamens stipitate, 0.6 mm tall, the anthers equal to outer whorls, erect, locelli 2, extrorse-latrorse, the filaments narrower than anthers, laminar, sparsely grey-villose, the basal glands sessile, globose; whorl IV wanting; pistillode filiform. Pistillate inflorescence with indument and color as in staminate plants, but shorter and with fewer lateral branches, the flowers similar in size and shape; stamens sterile, smaller; ovary glabrous, ovoid; style slender, distinct from ovary; stigma tri- lobed, 0.4 mm diam. Fruits borne on short terete pedicels of up to 1.5 X 1 mm, often subsessile and clustered; cupules hemispherical, to 1 x 1.5 cm, glabrous inside and outside, the margins entire; drupes ellipsoid, to 2 X 1.2 cm.

Distribution (Fig. 18) and ecology. Medium- sized riparian trees of central and western Amazonian lowlands ca. 100 m. Flowering from September to June, fruits available by December until the following August.

Representative specimens examined. COLOMBIA. CAQUETA: Rio Caqueta, Vega del Cafno Paufil, 9 km NE de Araracuara, 1 Dec 1988 (fr), Sanchez et al. 1870 (MO).

58 FLORA NEOTROPICA

BRAZIL. AMAZONAS: Manaus, Igarape da Cachoeira Grande, 28 May 1936 (fl d), Ducke 441 (A, F, K, MO, NY, R, U, US); Rio Cuieiras, margin of Rfo Branchinho, 12 Sep 1973 (fl d), Prance et al. 17785 (HBG, INPA, K, MO, NY, R, US). PARA: Oriximina, Rio Trombetas, 17 Jul 1980 (fl d ), Cid Ferreira et al. 1601 (HBG, INPA, MO, NY); San- tarem, Rio Curuauna, Prainha, 19 Jan 1979 (fr), Santos 563 (INPA, MG). ROND6NIA: Bananeiras, Madeira-Mamor6 Railway, 13 Sep 1963 (fl d), Maguire et al. 56624 (HBG, MO); Porto Velho, UHE de Samuel, Rio Jamari, 14 Jun 1986 (fl d), Thomas et al. 5089 (F, INPA, K, MO, NY).


Endlicheria macrophylla is the only member of the Ampelodaphne species group with ovate leaves. Such leaves are prevalent in available material, but occasional ovate-elliptic forms reduce the diagnostic value of leaf shape for this species. Of close relatives, E. multiflora shows similar soft pilose indument on the leaves below, and also has rusty tomentose branchlets, but its minute and stipitate basal glands are inconspicuous compared to the sessile globose glands that surround the whorl III filaments of E. macrophylla. Further, E. multifiora appears to be restricted to the Guianas while central Amazonian E. macrophylla can best be compared to sympatric E. arenosa. The latter bears a much shorter indument and occurs in well-drained white sand soils, whereas E. macrophylla is a species of flooded forests.

Flowers from Rondonia are less densely pubescent and have more reflexed tepals than those from north- em Amazonia.

23. Endlicheria multiflora (Miq.) Mez, Jahrb. K6n- igl. Bot. Gart. Berlin 5: 130. 1889. Goeppertia multifiora Miq., Stirp. Surin. Sel. 203. 1851. Type. Suriname. Without locality and date (fl d), Host- mann & Kappler 1163 (holotype: U; isotypes: B- n.v., BM-n.v., G, K-n.v., MO, OXF-n.v., P, S, W- n.v.)

Endlicheria villosa Mez, Jahrb. Konigl. Bot. Gart. Berlin 5: 129. 1889. Type. Jamaica? Without date (fl d), March s.n.? (syntypes: GOET-2 sheets).

Trees to 15 m. Branchlets slender to stout, midway along flush 3-6 mm diam., terete, densely rusty tomentose, the surface barely visible to concealed by the indument cover, the hairs relatively short, to 0.3 mm, crooked, erect, matted; terminal buds plump, 3 X 2 mm, the indument as on branchlets. Leaves closely spiraled at tips of current flush; petioles robust, to 2 X 0.4 cm, semi-terete, the indument as on branchlets; laminae coriaceous to stiff chartaceous,

plane to subbullate, obovate to narrowly elliptic, 12- 20 X 4-10 cm, the base acute to rounded, the apex obtuse to acute, acuminate for up to 1.5 cm, the margins minutely recurved throughout; upper surface dark green to olive-brown, minutely punctulate, the midrib and secondaries prominulous, the higher-order venation immersed, inconspicuous against the lamina; lower surface densely pubescent, the hairs of the midrib and secondaries ca. 0.6 mm long, crooked, densely matted, rusty red, those on the lamina greyish to translucent, 1 mm long, erect, straight or curved near tip, uniformly distributed, all vein orders raised, their prominence decreasing with rank; secondary veins 7- 9 per side, ? evenly spaced, slightly more distant around midlamina, patent, diverging at 70-85O (more obtusely around midlamina), arching after midcourse, distal pairs loop-connected; tertiaries oblique to midrib, between secondaries straight to forked. Staminate inflorescences distally clustered in the axils of cataphylls, to 25 cm long with 14 lateral branches, branch orders 3-4, the highest order botryoid, or irregular, the flowers clustered, the axes densely rusty villose, the hairs 1 mm long, crooked; bracts and bracteoles persistent at anthesis, lanceolate, greyish white villose; pedicels terete, to 1 mm long, those supporting secondary flowers slightly shorter. Flowers campanulate, 2 mm diam., densely greyish white villose outside; receptacle narrowly cyathiform, 1 X 0.3 mm, slightly constricted below tepals, glabrous inside. Te- pals membranaceous, ligulate, 1 X 0.6 mm (the inner whorl slightly broader), spreading to recurved at anthesis, the inner surface lightly papillose near the apex, otherwise glabrous. Stamens of whorls I and II stipitate, 0.6 mm tall, the anthers transversely oblong, 0.3 X 0.4 mm, glabrous, the apex truncate to emarginate, the connectives level with or reduced between the 2 locelli, these suborbicular, introrse, the filaments laminar, much narrower than anthers, sparsely grey- villose; whorl III stamens stipitate, 0.6 mm tall, the anthers equal to outer whorls, erect, locelli 2, extrorse-latrorse, the filaments narrower than anthers, laminar, sparsely grey-villose, the basal glands minutely stipitate, globose; whorl IV staminodial or absent, columnar; pistillode filiform. Pistillate inflorescence with indument and color as in staminate plants, but shorter and with fewer lateral branches, the flowers similar in size and shape; stamens sterile, smaller; ovary glabrous, ovoid; style slender, distinct from ovary; stigma tri-lobed, 0.4 mm diam. Fruits borue on short, terete pedicels of up to 5 X 3 mm, often subsessile and clustered; cupules hemispherical, to 1 X 1.5 cm, glabrous inside and outside, the margins entire; drupes ellipsoid, to 1.7 X 1.2 cm.


Distribution (Fig. 19) and ecology. Medium- sized trees of lowland riparian forests (5-100 m) in Guyana, Suriname, and bordering areas of Brazil and Venezuela. Flowering specimens collected from June through November, fruits collected in all months but September and January.

Representative specimens examined. VENEZUELA. BOLiVAR: Rio Venamo, 90 m, 31 Jul 1981 (fl d), Aymard et al. 372 (MO, NY); Rfo Uiri-yuk, 16 Aug 1962 (fl 6), Maguire et al. 46713 (NY).

GUYANA. BERBICE: Berbice River Kuruduni Ck., 50 m, 30 Apr 1995 (fr), Mutchnick 1243 (MO), Melissa Falls to the Gate, 120 m, 3 May 1995 (fr), Mutchnick 1303 (MO). DEMERARA: Demerara River, Sep 1889 (fl 6), Jenman 4127 (K, US), 19 Mar 1923 (fl 9, fr), Persaud 59 (F, INPA, K, MO, NY, U). ESSEQUIBO: Cuyuni-Mazaruni Region, Ma- zaruni River, Jun 1889 (fl 6), Jenman 5321 (K, NY, US); Potaro-Siparuni Region, Iwokrama Rainforest Reserve, Es- sequibo River, 80 m, 13-27 Jul 1996 (fl 6), Chanderbali et al. 153 (BRG, MO, US); Rupununi Region, Torobaroe Ck., 145 m, 21 Nov 1987 (fr juv), Jansen-Jacobs et al. 1118 (B, K, NY, U).

SURINAME. Nickerie R., Blanche Marie Falls, 20 Jun 1965 (fl d), Maas & Tawjoeran s.n. LLB. 10928 (U); Re- chter Coppename R., 14 May 1963 (fr), Wessels Boer 1381 (F, NY, U).

BRAZIL. PARU: Oriximind, Rio Trombetas, 29 Nov 1907 (fl 9), Ducke 8920 (U); Rio Mapuera, 12 Aug 1986 (fl d), Cid Ferreira et al. 7652 (F, INPA, K, MO, NY).

Local names. Guyana: bastard silverballi. Suri- name: siroewaballi oenilebobandikoro (Arawak)

In the Ampelodaphne species group, Endlicheria multiflora shares densely rusty tomentose branchlets and inflorescence axes with E. arenosa, E. macrophylla, E. melinonii, and E. reflectens, but the minute basal glands associated with its whorl IH stamens suggest affinity with E. dictifarinosa and E. levelii. Grey- ish tomentose branchlets and inflorescence axes, and subsessile leaves with subcordate bases, distinguish E. dictifarinosa from E. multiflora, but delimitation from E. levelii is not as straightforward. In the Suri- namese type of E. multiflora, most collections from

R EU

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60 FLORA NEOTROPICA

Guyana, and two collections from adjacent Bolivar in Venezuela, branchlets and inflorescence axes bear a dense rusty tomentose cover of ca. 0.6 mm long hairs whilst material from the upper Rfo Negro-Rio Ori- noco type locality of E. levelii have a dense greyish setose-villose indumentum of 1.2 mm long hairs. On the basis therefore of indument color and length, most specimens can be assigned to either E. multiflora or E. levelii without hesitation. Conceivably intermediate conditions are found only in Cid Ferreira et al. 7652 and Ducke 8920 (both from northern Para in Brazil), Maas & Tawjoeran s.n. (from Suriname), and Maguire & Fanshawe 23481 and Hoffinann et al. 2029 (both from Guyana). In these specimens, new leaves have sparse sprawling rusty hairs resembling those in the several sparsely pubescent specimens of E. levelii. Yet these 0.6 mm long rust-red hairs are a magnitude of order shorter than the 1.2 mm greyish hairs of the latter. Since these specimens differ from E. levelii in both indument length and color, but from typical E. multiflora only in indument density, they are more parsimoniously assigned to the latter species. With this recourse adopted, E. multiflora and E. levelii appear allopatric with the former centered in the Guianas and the latter in central Amazonia.

Ampelodaphne dasyantha Meisn., listed as a synonym of Endlicheria multiflora by Mez (1889) and Kostermans (1937), is an illegitimate name based on, among others, Hostmann & Kappler 1163, the type of Goeppertia multiflora. The only synonym of En- dlicheria multiflora accepted here, E. villosa, was based on two unlabeled collections found by Grise- bach among W. T. March's collections from Jamaica. As Endlicheria does not occur in Jamaica and the Am- pelodaphne species group is entirely South American, Grisebach's annotations and Mez's (1889) protologue justifiably express uncertainty about collector and locality. These sheets were certainly misplaced among March's Jamaican collections but their source is still unknown. If strongest resemblance with Cid Ferreira et al. 7652 and Ducke 8920 is any clue, northern Para in Brazil seems a likely locality.

24. Endlicheria levelii C. K. Allen, Mem. New York Bot. Gard. 10 (5): 62. 1964. Type. Venezuela. Amazonas: Rio Orinoco 10 km above mouth of Rio Atabapo, Canio Yagual, 150 m, 30 May 1954 (fl 1), Level 127 (holotype: NY).

Trees to 20 m. Branchlets slender to stout, midway along flush 3-6 mm diam., terete, densely pubescent, the surface barely visible, the hairs relatively long, to 1.2 mm, straight, or crooked, erect, greyish white; ter-

minal buds plump, 3 X 3 mm, densely pubescent, the hairs as on branchiets, straight, ascending. Leaves closely spiraled at tips of current flush; petioles slender, to 3 X 0.3 cm, semi-terete, the indument as on branchlets; laminae coriaceous to chartaceous, plane, obovate to elliptic, 10-25 X 3-9 cm, the base acute, the apex obtuse to acute, acuminate for up to 1.5 cm, the margins minutely recurved throughout; upper surface greyish green to olive-brown, minutely punctulate, the midrib and secondaries prominulous, the higher-order venation immersed, inconspicuous against the lamina; lower surface densely pubescent, the hairs 1.2 mm long, rigid, ascending, greyish, soon lost on lamina, persistent and denser on main veins, all vein orders raised, their prominence decreasing with rank; secondary veins 7-11 per side, + evenly spaced, slightly more distant around midlamina, patent, diverging at 70-850 (more obtusely around midlamina), arcuate, distal pairs loop-connected; tertiaries oblique to midrib, between secondaries once-forked to straight. Staminate inflorescences distally clustered in the axils of cataphylls, to 20 cm long with 18 lateral branches, branch orders 3-4, the highest order botryoid, or irregular, the flowers loosely crowded, the axes densely greyish white villose, the hairs 1 mm long, straight to crooked; bracts and bracteoles persistent at anthesis, lanceolate, the indument as on axes; pedicels terete, to 2 mm long, those supporting secondary flowers slightly shorter. Flowers campanulate, 2 mm diam., densely greyish white villose outside; receptacle cyathiform, 0.6 X 0.4 mm, slightly constricted below tepals, glabrous inside. Te- pals membranaceous, ligulate, 0.6 X 0.5 mm (the inner whorl slightly broader), spreading to recurved at anthesis, the inner surface lightly papillose near the apex, otherwise glabrous. Stamens of whorls I and II stipitate, 0.6 mm tall, the anthers transversely oblong, 0.3 X 0.4 mm, glabrous, the apex truncate to emarginate, the connectives level with or reduced between the 2 locelli, these suborbicular, introrse, the filaments laminar, much narrower than anthers, sparsely grey- villose; whorl III stamens stipitate, 0.6 mm tall, the anthers equal to outer whorls, erect, locelli 2, extrorse-latrorse, the filaments narrower than anthers, laminar, sparsely grey-villose, the basal glands minutely stipitate, globose; whorl IV staminodial or absent, columnar; pistillode filiform. Pistillate inflorescence with indument and color as in staminate plants, but shorter and with fewer lateral branches, the flowers similar in size and shape; stamens sterile, smaller; ovary glabrous, ovoid; style slender, distinct from ovary; stigma tri-lobed, 0.4 mm diam. Fruits borne on short, terete pedicels of up to 5 X 3 mm, often


subsessile and clustered; cupules hemispherical, to 1 X 1.5 cm, glabrous inside and outside, the margins entire; drupes ellipsoid, to 1.7 X 1.2 cm.

Distribution (Fig. 19) and ecology. Medium- sized riparian trees from central and western Ama- zonia at 80-250 m. Flowering from April to October, fruits available year round.

Representative specimens examined. COLOMBIA. VAUPES: Rio Apaporis, Soratama, 250 m, 1 Sep 1951 (fl 6), Schultes & Cabrera 13849 (GH, U, US).

VENEZUELA. AMAZONAS: Atabapo, Rio Cunucun- uma, 180-210 m, 28-30 Jan & 6-8 Feb 1982 (fr), Steyer- mark et al. 126162 (HBG, MO, NY); Atures, 43 km NE de Santa Barbara del Orinoco, 220 m, Apr 1990 (fr), Marin 981 (MO); Rio Negro, Rfo Baria, 80 m, 21 Jul 1984 (fl 6), Dav- idse 27575 (MO, NY, US). BARINAS: Rfo Zulia, Santa Bar- bara de Barinas, 21 Sep 1988 (fl 6), Valverde & Penia (MO).

BRAZIL. AMAZONAS: Rio Curicuriari, Igarape Cariua, 12 Jul 1979 (fl 6), Maia et al. 562 (HBG, INPA, K, MO, NY); Rio Negro, between Manaus and Sao Gabriel, Ilha Acaburu, 4 Apr 1979 (fl 6), Maia et al. 430 (HBG, MO, NY, US); Rio Solimoes, Sao Paulo de Olivenqa, near Pal- mares, 1 1 Sep-26 Oct 1936 (fr), Krukoff 8096 (A, F, G, K, MO, NY, U); Rio Tea, affluente do Rio Negro, 12 Jun 1976 (fl Y), Coelho 460 (INPA); Serra Araga, Rio Jauari, 28 Jul 1985 (fr), Silva 181 (GH, K, MO, NY). MATO GROSSO: Nucleo Pioneiro de Humboldt, 28 Oct 1973 (fr), Berg & Steward P19932 (HBG, INPA, K, MO, NY); Serra do Ron- cador, Rio Vau, 11 Oct 1964 (fr), Prance & Silva 59385 (GH, MO, NY, US).

BOLIVIA. SANTA CRUZ: Prov. Velasco, Reserva Biol- 6gica El Refugio, 210 m, 23 May 1995 (fl 6), Guilln & Chore 3814 (MO), ibid., 27 Jan 1997 (fr), Carrion et al. 531

(MO).

Local name. Venezuela: laurel de babilla.

Endlicheria levelii is distinguished from E. multiflora primarily by the greyish white indument of its branchlets and inflorescences (see discussion of the latter), and from E. dictifarinosa by its acute rather than subcordate leaf bases. The type specimen and material from Bolivia and adjacent Mato Grosso in Brazil have a dense indumentum on the leaves below, but hairs are restricted to midribs below in all other specimens. These less pubescent specimens are reminiscent of E. bracteata wherein hairs can be equally long, or longer, but leaves tend to dry blackish green and always show prominent tertiaries above.

25. Endlicheria dictifarinosa C. K. Allen, Mem. New York Bot. Gard. 12 (3): 108. 1965. Type. Ven- ezuela. Bolivar: Maihia Rapids along Rio Paragua, 500-510 m, 1 Jan 1962 (fl 6), Steyermark 90535

(holotype: NY; isotypes: G, U).

Trees, 5-20 m. Branchlets slender to stout, midway along flush 3-6 mm diam., terete, densely tomentose, the surface barely visible to concealed by the indument cover, the hairs relatively short, to 0.3 mm, crooked, erect, matted, rusty to greyish white; terminal buds plump, 3 X 2 mm, the hairs as on branchlets, ascending. Leaves closely spiraled at tips of current flush; petioles robust, to 1 X 0.4 cm, semi- terete, the indument as on branchlets; laminae coriaceous to stiff chartaceous, plane, obovate, 11-25 X

5-13 cm, the base abruptly rounded to cordate, the apex obtuse, acuminate for up to 1.5 cm, the margins minutely recurved throughout; upper surface light greyish green to reddish brown, minutely punctulate, the midrib and secondaries prominulous, the higher- order venation immersed; lower surface moderately pubescent, the hairs 0.7 mm long, crooked on the midrib and secondaries, on the lamina straight and erect, uniformly distributed, all vein orders raised, their prominence decreasing with rank; secondary veins 7- 11 per side, ? evenly spaced, slightly more distant around midlamina, patent, diverging at 70-85? (more obtusely around midlamina), arching after midcourse, distal pairs loop-connected; tertiaries oblique to midrib, between secondaries straight to forked. Staminate inflorescences distally clustered in the axils of cataphylls, to 25 cm long with 12 lateral branches, branch orders 3-4, the highest order botryoid to irregular, the flowers loosely crowded, the axes densely villose, the hairs 1 mm long, crooked, greyish white throughout, a few reddish, or distally rusty; bracts and bracteoles persistent at anthesis, lanceolate, densely greyish whitevillose; pedicels terete, to 1 mm long, those supporting secondary flowers slightly shorter. Flowers campanulate, 2 mm diam., densely greyish white or rusty villose outside; receptacle narrowly cyathiform, 1 X 0.3 mm, slightly constricted below tepals, glabrous inside. Tepals membranaceous, ligulate, 1 X 0.6 mm (the inner whorl slightly broader), spreading to recurved at anthesis, the inner surface lightly papillose near the apex, otherwise glabrous. Stamens of whorls I and II stipitate, 0.6 mm tall, the anthers transversely oblong, 0.3 X 0.4 mm, glabrous, the apex truncate to emarginate, the connectives level with or reduced between the 2 locelli, these suborbicular, introrse, the filaments laminar, much narrower than anthers, sparsely grey-villose; whorl III stamens stipitate, 0.6 mm tall, the anthers equal to outer whorls, erect, locelli 2, extrorse-latrodrse, the filaments narrower than anthers, laminar, sparsely grey-villose, the basal glands minutely stipitate, globose; whorl IV staminodial or absent, columnar; pistillode filiform. Pistillate inflorescence unknown. Fruits borne on

62 FLORA NEOTROPICA

short, terete pedicels of up to 5 X 3 mm, often subsessile and clustered; cupules hemispherical, to 1 X 1.5 cm, glabrous inside and outside, the margins entire; drupes ellipsoid, to 1.7 X 1.2 cm.

Distribution (Fig. 19) and ecology. Small to medium-sized riparian trees from SE Venezuela and the adjacent state of Roraima in Brazil at ca. 100-800 m. Flowering material collected in January, May, and from July to October, and fruiting specimens from November to May.

Representative specimens examined. VENEZUELA. AMAZONAS: Rio Orinoco, Tama-Tama, 125-150 m, 28 Jul 1959 (fl d), Wurdack & Adderley 43647 (B, COL, F, G, NY, W); Sierra Parima, vecindades de Simarawochi, Rio Mata- cuni, 795-830 m, 18 Apr-23 May 1973 (fr), Steyernark 107391 (G, HBG, NY). BOLiVAR: Cedenlo, Icutd, cuenca del Rio Nachare, 150 m, 1 Aug 1995 (fl d), Knab-Vispo & Rodriguez 479 (MO, VEN); Uriman, Rio Garoni, 480 m, 31 Aug 1954 (fl d), Bernardi 1626 (COL, G, NY).

BRAZIL. RORAIMA: Boa Vista, Estaqao Ecol6gica de Maraca, 19 May 1987 (fr), Lima 783 (E, INPA, K, MO, NY); Rio Mucajai, river island, 17 Mar 1971 (fr), Prance et al. 11069 (HBG, K, MO, NY, US).

Local names and uses. Venezuela: laurel hormiguero, peroua-yek (Arecuna), wakamei (Ye'kwana), Brazil: moroman (Uaica-Mucajai). Fruits edible.

Endlicheria dictifarinosa shares greyish tomentose branchlets and inflorescences with E. levelii but is distinguished by its subsessile leaves with subcordate bases. The hairs of the two also differ in length, with those of E. dictifarinosa half as long as the ca. 1.2 mm hairs in E. levelii. Occasionally, distal inflorescence branches, pedicels, and exterual flower parts are rusty tomentose (e.g., Bernardi 1626), or rust colored hairs are interspersed throughout the dominant greyish indumentum (e.g., Knab-Vispo & Rodriguez 479). These specimens appear marginal between E. dictifarinosa and E. multiflora on the basis of indument color but do not show the soft pilose indument that typically persists on lower leaf surfaces in the latter. Their peculiar subsessile leaves are also unmatched in E. multiflora. Leaves of similar shape combine with adaxially prominent third- and fourth- order veins in E. bracteata, E. cocuirey, and E. verticillata.

26. Endlicheria melinonii Benoist, Bull. Soc. Bot. France 75: 977. 1928 (nomen nudum), validated in Arch. Botan. V: 63. 1931. Type. French Guiana.

Cayenne, without date (fl d), Melinon s.n. (holotype: P; isotype: P).

Trees to 30 m. Branchlets slender to stout, midway along flush 3-6 mm diam., terete, densely rusty tomentose, the surface barely visible to concealed by the indument cover, the hairs relatively short, to 0.3 mm, crooked, erect, matted; terminal buds plump, 3 X 2 mm, densely rusty tomentose. Leaves closely spiraled at tips of current flush; petioles robust, to 1 X 0.3 cm, semi-terete, the indument as on branchlets; laminae chartaceous to coriaceous, plane, obovate, 8- 15 X 3-7 cm, the base acute, the apex obtuse, acuminate for up to 1 cm, the margins minutely recurved throughout; upper surface dark green to olive-brown, minutely punctulate, the midrib and secondaries prominulous, the higher-order venation immersed, inconspicuous against the lamina; lower surface densely rusty pubescent, the hairs ca. 0.5 mm long, erect, straight, uniformly distributed, all vein orders raised, their prominence decreasing with rank; secondary veins 5-8 per side, ? evenly spaced, slightly more distant around midlamina, patent, diverging at 70-85? (more obtuse around midlamina), arching after midcourse, distal pairs loop-connected; tertiaries oblique to midrib, between secondaries straight to forked. Sta- minate inflorescences distally clustered in the axils of cataphylls, to 11 cm long with 8 lateral branches, branch orders 2-3, the highest order condensed, the flowers clustered, the axes densely rusty villose, the hairs 0.7 mm long, crooked; bracts and bracteoles persistent at anthesis, ovate, the indument as on axes; pedicels wanting. Flowers campanulate, 2 mm diam., densely rusty villose outside, the hairs sparser on tepals; receptacle cyathiform, 1 X 0.4 mm, slightly constricted below tepals, sericeous inside. Tepals membranaceous, ovate, 1 X 0.6 mm (the inner whorl slightly broader), erect at anthesis, the inner surface glabrous, the margins minutely papillose. Stamens of whorls I and II stipitate, 0.6 mm tall, the anthers transversely oblong, 0.3 X 0.4 mm, glabrous, the apex truncate to emarginate, the connectives level with or reduced between the 2 locelli, these suborbicular, introrse, the filaments laminar, much narrower than anthers, sparsely grey-villose; whorl III stamens stipitate, 0.6 mm tall, the anthers equal to outer whorls, erect, locelli 2, extrorse-latrorse, the filaments narrower than anthers, laminar, sparsely grey-villose, the basal glands sessile, globose; whorl IV wanting; pistillode fusiform. Pistillate inflorescence with indument and color as in staminate plants, but shorter and with fewer lateral branches, the flowers similar in size and shape; stamens sterile, smaller; ovary glabrous,


ovoid; style slender, distinct from ovary; stigma tri- lobed, 0.3 mm diam. Cupules hemispherical, sessile, to 8 x 5 mm, glabrous inside and outside, the margins undulate, drupes ellipsoid, to 1.5 X 1 cm.

Distribution (Fig. 20) and ecology. Tall trees from flooded and nonflooded moist forests of NE South America at ca. 200-450 m. Flowering material collected in August, September, and January, and fruits in February and September.

Additional specimens examined. VENEZUELA. DELTA AMACURO: El Palmar, Rio Grande, 19 Aug-7 Sep 1964 (fl 6), Marcano-Berti 361 (F, G, HBG, SPA, MO, NY).

FRENCH GUIANA. Cayenne, s.d. (fl d), Patris s.n. (G), Saul, La Fum6e Mtn. Trail, 200-300 m, 16 Sep 1984 (fr juv), Mori et aL 20927 (MO, NY), 220 m, 21 Jun 1988 (st), Gentry et al. 62990 (NY); Bassin du Sinnary, 22 Feb 1989 (fr), Sabatier 2365 (MO, NY); Crique Passoura, 26 Sep 1992 (fr juv), Sabatier & Prevost 4073 (B, MO, NY); Mont Atachi Bacca, Region de l'Inini, 420 m, 10 Jan 1989

(fr juv), Granville 10538 (B, MO, NY, U, US); Pistie de Saint-Elie, 26 Sep 1991 (fl 9), Sabatier & Prevost 3819 (MO, NY, U).

SURINAME. Brokopondo, 21 Jan 1965 (st), van donselaar 2037 (MO); Rikanau prope Moengo, 2 Jun 1954 (st), Lindeman 5943 (U), (st), Lindeman 6056 (U); Suriname River, Wilhelminagebergte-Mapane Ck., 1 Oct 1953 (st), Lindeman 4787 (NY, U); s.loc., 2 Mar 1957 (st), Heyligers 485 (U).

BRAZIL. AMAPA: Reserva INCR, Rio Falsino, 22-26 Aug 1983 (fl dc), Campbell et al. 14650 (NY).

Local names. Venezuela: laurel negro. Suriname: harige pisie, pisie.

Kostermans (1937) placed Endlicheria melinonii in synonymy under E. multiflora. The two share rusty tomentose indument and obovate leaves but E. melinonii is easily distinguished by its sessile flowers with erect tepals. Further, stiff straight ca. 0.5 mm long hairs on the lower leaf surface permit recognition on vegetative grounds alone. Sterile specimens thus as-

I ~~~~~~~~~~~~~~~~~~~~~~~7

.-^tsx55rl X*ir;tY; f EL M\- F.

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FIG. 20. Distribution of Endlicheria melinonii E. reflectens, and E. bracteata.

64 FLORA NEOTROPICA

signed establish its presence in Suriname and reduce the severity of an otherwise striking disjunction between French Guiana-NE Brazil and NE Venezuela. Although the lack of material from intervening Guy- ana may be a collecting artifact, similar disjunctions between Suriname and Venezuela are found in the range of widespread E. bracteolata and E. canescens. As both have been found in Guyana only around the upper Cuyuni-Mazaruni Region that borders Vene- zuela, perhaps here too is where E. melinonii occurs in Guyana. Nevertheless, if real, the absence of these three species from relatively well-collected central Guyana is enigmatic.

27. Endlicheria reflectens (Nees) Mez, Jahrb. Kon- igl. Gart. Berlin 5: 126. 1889. Goeppertia reflectens Nees, Linnaea 21: 514. 1848. Type. Guyana. Without locality and date (fl d), Schomburgk 475/ 801 (holotype: B-n.v.; isotypes: E, K-n.v.).

Shrubs to 4 m. Branchlets slender, midway along flush 2-3 mm diam., distally weakly angular, soon terete, densely rusty tomentose, the surface concealed by the indument cover or barely visible, the hairs moderately long, to 0.5 mm, straight to crooked, erect, loosely matted; terminal buds slender, 3 X 1 mm, densely pubescent, the hairs as on branchlets, ascending. Leaves altermate, widely and evenly spaced along current flush or loosely spiraled at tips of branchlets; petioles slender, to 1 X 0.2 cm, terete, the indument as on branchlets; laminae chartaceous to coriaceous, plane, obovate to elliptic, 5-12 X 2-5 cm, the base acute, briefly decurrent, the apex obtuse to acute, acuminate for up to 1 cm, the margins minutely recurved throughout; upper surface dull green to olive-brown, waxy, the primary to fourth-order veins raised, their prominence decreasing with rank; lower surface densely pilose, the hairs translucent, 0.5 mm long, erect, straight or curved near tips, uniformly distributed, all vein orders raised, their prominence decreasing with rank; secondary veins 4-6 per side, ? evenly spaced, slightly more distant around midlamina, ascending at 50-60o (more obtuse around midlamina), arcuate, distal pairs loop-connected; tertiaries reticulating between secondaries. Staminate inflorescences distally clustered in the axils of cataphylls, to 12 cm long with 8 lateral branches, branch orders 2-3, the highest order irregular to botryoid, lax, the flowers distant, the axes sparsely pubescent, the hairs light brown, 0.5 mm long, soft, erect, straight to crooked; bracts and bracteoles persistent at anthesis, lanceolate, densely pubescent, the hairs as on axes, appressed to ascending; pedicels terete, to 2 mm long, those sup-

porting secondary flowers slightly shorter. Flowers campanulate, 3 mm diam., sparsely tawny pilose outside, the hairs ascending; receptacle cyathiform, 0.6 X 0.5 mm, slightly constricted below tepals, glabrous inside. Tepals membranaceous, ovate, 1 X 0.6 mm (the inner whorl slightly broader), spreading to recurved at anthesis, the inner surface glabrous, the margins minutely papillose. Stamens of whorls I and II narrowly stipitate, 0.6 mm tall, the anthers transversely oblong, 0.3 X 0.5 mm, glabrous, the apex truncate to emarginate, the connectives level with or reduced between the 2 locelli, these suborbicular, introrse, the filaments laminar, much narrower than anthers, glabrous; whorl III stamens broadly stipitate, 0.6 mm tall, the anthers depressed-elliptic, 0.3 X 0.5 mm, slightly bowed towards the outer whorls, locelli 2, extrorse-latrorse, the filaments slightly narrower than anthers, laminar; opaque pilose near base, the basal glands sessile, globose, relatively large, filling the space between filaments of inner and outer whorls; whorl IV wanting; pistillode wanting. Pistillate inflorescence with indument, color, and branching as in staminate plants, the flowers similar in size and shape; stamens sterile, smaller; ovary glabrous, ovoid; style stout, weakly distinguished from ovary; stigma broadly tri-lobed, 0.5 mm diam. Fruits borne on slender terete pedicels of up to 1 X 0.3 cm; cupules hemispherical, to 1 X 1.5 cm, glabrous inside and outside, the margins entire; drupes ellipsoid, to 2 X 1 cm.

Distribution (Fig. 20) and ecology. Shrubs from savanna formations of the Guiana shield at ca. 100- 500 m. Flowering from March to November, and fruits available from January to June.

Representative specimens examined. COLOMBIA. VICHADA: Puerto Ayacucho, laja outcrops behind Casuar- ito, 4 Apr 1984 (fr), Gentry & Stein 46329 (MO, NY).

VENEZUELA. BOLfVAR: Igneous outcrops at Agua Amena, 100 m, 6 Sep 1985 (fl 6), Steyermark et al. 131425

(MO, NY). GUYANA. ESSEQUIBO: Rupununi Region, Rupununi

R., Karanambo, 1 Sep 1988 (fl 6), Maas et al. 7136 (K, MO, NY, U); Dadanawa, 120-150 m, 25 Jun 1989 (fl 6), Gillespie et al. 1691 (MO, NY, U, US).

BRAZIL. AMAZONAS: Rio Branco, Aug 1913 (fl 6), Kuhlmann 3376 (U). RORAIMA: Rio Branco, Surumu, Jul 1909 (fl 6'), Ule 8125 (G, K); SEMA Ecological Reserve, Ilha de Maraca, 11 Jul 1952 (fl d), Milliken et al. 439 (E, MO, NY).

Endlicheria reflectens alone in the Ampelodaphne species group has reticulate tertiaries. Here too basal glands of whorl III stamens extend between filaments of whorls I and II to an extent unmatched in this species group. Elsewhere in the genus, similarly large


glands are found in E. acuminata and E. gracilis, both with triplinerved leaves. Its obovate to elliptic leaves with pilose lower surfaces and rusty tomentose branchlets and inflorescence are reminiscent only of E. multiflora. Ranges of the two species overlap in SW Guyana where, as elsewhere, E. multiflora are single-stemmed trees of riparian forests and E. reflectens are shrubs of savanna formations with several slender stems that sprawl across adjacent vegetation. The habit and habitat of E. reflectens is shared with E. arenosa, but nonreticulate tertiaries immersed above and short stiffly erect hairs below leaves of the latter provide easy vegetative contrast.

28. Endlicheria bracteata Mez, Repert. Spec. Nov. Regni Veg. 16: 306. 1920. Type. Peru. San Martin: Moyabamba, without date (fl d), Weberbauer 4680 (holotype: B-n.v.; fragment, F; photo [neg. 3813 ex B], F, G, GH, NY).

Trees to 10 m. Branchlets slender to stout, midway along flush 3-7 mm diam., terete, densely hirsute, the surface barely visible, the hairs very long, to 2 mm, straight, erect, grey to rusty red; terminal buds slender, 4 X 2 mm, densely pubescent, the hairs as on branchlets, ascending. Leaves closely spiraled at tips of current flush; petioles short, robust, to 0.5-1 X 0.4 cm, semi-terete, the indument as on branchlets; laminae chartaceous, plane, obovate (often spathulate), 15-40 X 6-15 cm, the base acute or abruptly contracted into the petiole, the apex obtuse, acuminate for up to 3 cm, the margins minutely recurved throughout; upper surface dark green, the primary to fourth-order veins raised, their prominence decreasing with rank, or primary and secondary veins sunken; lower surface moderately pubescent, the hairs as on branchlets, rigidly erect, greyish green, denser on main veins, all vein orders raised, their prominence decreasing with rank; secondary veins 13-20 per side, gradually further apart towards apex, patent, diverging at 70-85? (more obtuse towards apex), arcuate to arching after midcourse, distal pairs loop-connected; tertiaries oblique to midrib, between secondaries once-forked to straight. Staminate inflorescences distally clustered in the axils of cataphylls, to 20 cm long with 10 lateral branches, branch orders 3-4, the highest order botryoid to irregular, lax, the flowers distant, the axes densely greyish hirsute, the hairs 1.5 mm long, straight, erect; bracts and bracteoles persistent at anthesis, lanceolate, the indument as on axes, ascending; pedicels terete, to 2 mm long, those supporting secondary flowers slightly shorter. Flowers campanulate, 3 mm diam., sparsely grey-villose out-

side; receptacle globose, 0.8 X 0.6 mm, constricted below tepals, glabrous inside. Tepals chartaceous, ovate, 1 X 0.8 mm, spreading to recurved at anthesis, the inner surface minutely papillose towards apex, otherwise glabrous, the margins papillose. Stamens of whorls I and II stipitate, relatively tall, up to 1 mm, the anthers transversely oblong, 0.3 X 0.5 mm, glabrous, the apex truncate to emarginate, the connectives level with or reduced between the 2 locelli, these suborbicular, introrse, the filaments laminar, narrower than anthers, glabrous; whorl III stamens stipitate, equal to outer whorls, the anthers depressed-oblong, erect, locelli 2, extrorse-latrorse, the filaments narrower than anthers, laminar, glabrous, the basal glands broadly stipitate, globose, apiculate; whorl IV staminodial or absent, columnar; pistillode fusiform. Pis- tillate inflorescence with indument and color as in staminate plants, but shorter and with fewer lateral branches, the flowers slightly deeper; stamens sterile, smaller; ovary glabrous, ovoid; style slender, distinct from ovary; stigma broadly tri-lobed, 0.5 mm diam. Fruits borue on terete pedicels of up to 7 X 3 mm; cupules hemispherical, to 1 X 2 cm, initially rusty or grey-velutinous outside, the indument thinning with age, glabrous inside, the margins entire; drupes ellipsoid, to 2 X 1.5 cm.

Distribution (Fig. 20) and ecology. Small understory trees of nonflooded lowland Amazonia to ca. 1200 m on the adjacent eastem Andean slopes. Flow- ering material collected from January through Octo- ber, fruits in March, July, August, and December.

Representative specimens examined. ECUADOR. NAPO: Tena, Estaci6n Biol6gica Jatun Sacha, 400 m, 26 Jun 1991 (fl 3), Palacios 7500 (MO).

PERU. HuANUCO: Leoncio Prado, Rupa Rupa, Tingo Marfa, 672 m, 21 Oct 1971 (fr), Schunke 5060 (F, G, GH, MO, US); Pachitea, Sira Mtns., 800 m, 12 Aug 1988 (fl d

juv), Morawetz & Wallnofer 11-12888 (MO). LORETO:

Maynas, Iquitos, Puerto Almendras, 122 m, 16 Jun 1988 (fl i), Vasquez & Soto 12350 (GH, HBG, MO); Pumayacu, between Balsapuerto and Moyobamba, 600-1200 m, Aug- Sep 1933 (fl d), Klug 3187 (F, G, GH, K, MO, NY, US). MADRE DE DIos: Tambopata, Zone Reserva de Tambopata, Rio Tambopata, 260 m, 23 Aug 1990 (fl d), Alexiades & Nicole 1042 (K, MO), 280 m, 19 Aug 1990 (fl d ), Reynel et al. 5244 (MO). SAN MARTiN: Jepelacio, near Moyob-

amba, 1100 m, Jul 1934 (fl d), Klug 3745 (F, GH, MO, NY, US); Mariscal Caceres, Tocache Nuevo, 400 m, 29 Jun 1978 (fl Y), Schunke 10304 (F, G, HBG, MO, NY). UCAYALI:

Coronel Portillo, Pucallpa, Estaci6n Experimental Alexan- der von Humboldt, Arboreto "A. Solagar Cavero," 23 Jul 1980 (fl d), Sousa 76 (INPA).

BRAZIL. ACRE: Tarauaca, Vit6ria Velho, 28 Sep 1994 (fl d), Silveira et al. 943 (MO). AMAZONAS: Manaus, Re-

66 FLORA NEOTROPICA

serva Florestal Ducke, Manaus-Itacoatiara Rd., km 26, 10 Aug 1965 (fl d), Rodrigues 7028 (INPA, NY), 4 Dec 1997 (fr), Souza & AssunVdo 491 (MO). PARA: Oriximina, Rio Trombetas, Porto Trombetas, 29 Aug 1980 (fl 6), Cid Fer- reira et al. 1884 (HBG, INPA, K, MO, NY). ROND6NIA: Santa Barbara, Rod. BR 364, km 120, 29 May 1982 (fl 5), Teixeira et al. 852 (F, MO, NY, US).

Local names. Peru: moena, shisho moena, shisho

moenita. Brazil: louro.

Endlicheria bracteata is conspicuous amongst species of the Ampelodaphne species group with prominent tertiary and fourth-order veins above (i.e., the E. bracteata subgroup) for its dense indumentum of stiff greyish 2 mm long hairs on its branchlets, lower leaf surfaces, and inflorescence axes. A globose receptacle with strongly constricted apex below spreading tepals and definitely stipitate basal glands associated with whorl III stamens are also distinctive. Leaf bases usually taper gradually into the petiole, but occasional subcordate forms (e.g., Klug 3187) resemble those of typical of E. cocuirey, a species with a much shorter indumentum of ca. 0.6 mm long hairs. The variation of leaf shape in E. bracteata is paralleled by E. verticillata. The two coincide in central Amazonia where, according to label data, E. verticillata is found in flooded and E. bracteata in well- drained habitats. Further, the hairs tend to be reddish rather than greyish in E. verticillata, distal inflorescence axes and external flower parts are glabrous, receptacles are campanulate without constricted apices, and basal glands are definitely sessile.

As young cupules can be densely pubescent outside (e.g., Schunke 7245 & 8069), fruiting specimens of Endlicheria bracteata can be confused with E. chalisea, a vegetatively similar species where densely pubescent cupules are typical. However, in E. bracteata these hairs soon fall and mature cupules are either glabrous or only sparsely pubescent. Further- more, cupules here are always glabrous inside, while in E. chalisea, cupules remain densely pubescent inside even if the hairs outside are sometimes lost.

29. Endlicheria verticillata Mez, Bull. Herb. Boiss. 5 (ser. 2): 235. 1905. Type. Brazil. Amazonas: Rio Jurua, Jun 1901 (fl d), Ule 5584 (holotype: B- n.v.; isotypes: G, HBG, K).

Trees, 3-20 m. Branchlets slender to stout, midway along flush 4-8 mm diam., terete, densely rusty hirsute, the surface barely visible, the hairs long, to 2 mm, straight, erect; terminal buds slender, 4 X 2 mm, densely pubescent, the hairs as on branchlets, ascending. Leaves closely spiraled at tips of current flush;

petioles relatively short, robust, to 0.5-1 X 0.4 cm, semi-terete, the indument as on branchlets; laminae chartaceous, plane, obovate, 15-35 X 5-12 cm, the base acute or abruptly contracted into the petiole, the apex obtuse, acuminate for up to 3 cm, the margins minutely recurved throughout; upper surface olive- brown, the primary to fourth-order veins raised, their prominence decreasing with rank, or midrib and secondaries impressed, the higher-order venation prominent; lower surface moderately pubescent, the hairs as on branchlets, rigidly erect, reddish, denser on main veins, all vein orders raised, their prominence decreasing with rank; secondary veins 1 1-15 per side, gradually further apart towards apex, patent, diverging at 70-85? (more obtuse towards apex), arcuate to arching after midcourse, distal pairs loop-connected; tertiaries oblique to midrib, between secondaries once-forked to straight. Staminate inflorescences distally clustered in the axils of cataphylls, to 25 cm long with 12 lateral branches, branch orders 3-4, the highest order botryoid to irregular, lax, the flowers distant, the axes glabrous except for the sparsely rusty hirsute peduncle; bracts and bracteoles persistent at anthesis, lanceolate, sparsely rusty hirsute, glabrous after second-order branches; pedicels terete, to 2 mm long, those supporting secondary flowers slightly shorter. Flowers campanulate, 3 mm diam., glabrous outside; receptacle cyathiform, 0.8 X 0.6 mm, glabrous inside. Tepals chartaceous, ovate, 1 X 0.8 mm, spreading at anthesis, the inner surface glabrous, the margins papillose. Stamens of whorls I and II stipitate, 0.6 mm tall, the anthers transversely oblong, 0.4 X 0.5 mm, glabrous, the apex truncate to emarginate, the connectives level with or reduced between the 2 locelli, these suborbicular, introrse, the filaments laminar, narrower than anthers, sparsely grey-tomentose; whorl III stamens stipitate, equal to outer whorls, the anthers depressed-oblong, erect, locelli 2, extrorse- latrorse, the filaments narrower than anthers, laminar, sparsely grey-tomentose, the basal glands sessile, globose; whorl IV wanting; pistillode fusiform. Pistillate inflorescence with indument and color as in staminate plants, but shorter and with fewer lateral branches, the flowers slightly deeper; stamens sterile, smaller; ovary glabrous, ovoid; style slender, distinct from ovary; stigma broadly tri-lobed, 0.6 mm diam., the lobes minutely papillose. Fruits borne on terete pedicels of up to 6 X 4 mm; cupules shallowly hemispherical to patelliform, to 0.3 X 1 cm, glabrous inside and outside, the margins entire; drupes ellipsoid, to 3.5 X 1.5 cm.

Distribution (Fig. 21) and ecology. Small to medium-sized trees of flooded lowland forests throughout central Amazonia at ca. 90-200 m. Flow-

SYST'EMATIC TREATMENT 6

~~~~~~~~~~~~~~~~~~~~~~~r.~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~J

. ..... .......

FIG. 21. Distribution of Endlicheria verticillata, E. cocuirey, and E. tessmannii

ering March to October, fruits available from August to the following June.

Representative specimens examined. COLOMBIA. AMAZONAS: Leticia, Tarapaca, Parque Nacional Natural Amacayacu, 100 m, 21 Jun 1991 (fl 6), Rudas et al. 2290 (MO), 125 m, 12 Jun 1992 (fr), Rueda 445 (MO).

PERU. LORETO: Maynas, Rio Amazonas, Yanamono Explorama Tourist Camp, halfway between Indiana and mouth of Rio Napo, 120 m, 15 Jun 1986 (fl d), Gentry et al. 54550 (F, MO, NY); Sapuena, Arboreto Jenaro Herrera, 140 m, 10 Aug 1988 (fr), van der Werif et aL 10053 (F, HBG, MO, NY). UCAYALI: Pucallpa, Pau-Cocha, 200 m, 6 May 1961 (fl c), Woytkowski 6304 (F, GH, K, MO, NY).

BRAZIL. AcRE: Sena Madureira, trail to Rio laco from kn 7 along Sena Madureira to Rio Branco rd., 1 Oct 1968 (fr), Prance et al. 7744 (HBG, INPA, MG, MO, NY, R); Traumaturgo, Rio Alto Jurua, Reserva Extractivista do Alto Jurua, 4 Apr 1993 (fr), Silveira et al. 468 (NY). AMAZONAS: Manaus, Igap6 no Parana do Careiro, 21 Oct 1947 (fl d), Ducke s.n. R 19966 (HBG, MO, U); Livramento, Tres Casas, bacia do Madeira, 19 Sep 1962 (fr), Duarte 7368 (HBG, INPA, MO); Parana do Autaz Mirim, 18 Apr 1966 (fr), Rod- rigues & Mello 7760 (INPA, NY); Rio Negro, 23 Sep 1975 (fl ; ma), Kubitzki 75-34 (B, HBG, INPA). PARk: Rio Ja-

pure, 14 Oct 1904 (fl 6), Ducke 6764 (G); Rio Tapaj6s, Missao Cururu, 17 Jul 1959 (fl 6), Elger 914 (HBG). RoND-

6NA: Rio Abuna, 18 Jul 1968 (fl S), Prance et al. 6182 (HBG, INPA, MO, NY, R).

Local names. Peru: moena, muena amarilla, pucacuro caspi (ant tree). Brazil: louro abacate, louro cedro, louro jambo.

Endlicheria verticillata shares glabrous flowers with E. arachnocome and E. arunciflora, but on account of its long hirsute vestiture and subsessile leaves with prominent tertiary and fourth-order veins above is best compared to E. bracteata, a species with densely pubescent inflorescences and rather different flowers (see above).

30. Endlicheria cocuirey Kosterm., Recueil Trav. Bot. N6erl. 34: 522. 1937. Type. Peru. Loreto: Florida, Rio Putumayo at mouth of Rio Zubineta, 180 m, May-Jul 1931 (fl d), Kiug 2253 (holotype: F; isotypes: A, B-n.v., G, GH, K, MO, NY, S, US).

68 FLORA NEOTROPICA

Trees, 4-15 m. Branchlets stout, midway along flush 4-6 mm diam., terete, densely rusty to grey- tomentose, the surface barely visible to concealed by the indument cover, the hairs relatively short, to 0.6 mm, straight to crooked, erect; terminal buds plump, 4 X 3 mm, densely pubescent, the hairs as on branchlets, ascending. Leaves closely spiraled at tips of current flush; petioles robust, to 8 X 4 mm, semi- terete, the indument as on branchlets; laminae chartaceous, plane, obovate, 15-30 X 6-11 cm, the base rounded to cordate, the apex obtuse, acuminate for up to 1 cm, the margins minutely recurved throughout; upper surface dark green to olive-brown, waxy, the primary to fourth-order veins raised, their prominence decreasing with rank; lower surface sparsely grey- tomentose, the hairs as on branchlets, scattered, erect to ascending, slightly denser on main veins, all vein orders raised, their prominence decreasing with rank; secondary veins 11-16 per side, ? evenly spaced, slightly more distant around midlamina, patent, diverging at 70-85? (more obtuse around midlamina), arcuate to arching after midcourse, distal pairs loop- connected; tertiaries oblique to midrib, between secondaries once-forked to straight. Staminate inflorescences distally clustered in the axils of cataphylls, to 20 cm long with 15 lateral branches, branch orders 3- 4, the highest order botryoid to irregular, the flowers clustered, the axes densely rusty to grey-tomentose; bracts and bracteoles persistent at anthesis, ovate, the indument as on branchlets; pedicels terete, to 1 mm long, reduced to lacking below secondary flowers. Flowers campanulate 2 mm diam., sparsely grey- tomentose outside, the hairs thinning distally, lost by tepals; receptacle cyathiform, 0.6 X 0.5 mm, glabrous inside. Tepals membranaceous, elliptic, 0.8 X 0.6 mm, spreading to recurved at anthesis, the inner surface glabrous, the margins minutely papillose near apex. Stamens of whorls I and II stipitate, 0.6 mm tall, the anthers depressed-ovate, 0.3 X 0.4 mm, glabrous, the apex truncate to emarginate, the connectives level with or reduced between the 2 locelli, these suborbicular, introrse, the filaments laminar, narrower than anthers, sparsely grey-tomentose outside; whorl III stamens stipitate, 0.6 mm tall, the anthers oblong, 0.3 X 0.2 mm, erect, locelli 2, extrorse-latrorse, the filaments narrower than anthers, laminar, densely grey-tomentose inside, the basal glands sessile, globose; whorl IV staminodial, columnar; pistillode filiform. Pistillate plants unknown.

Distribution (Fig. 21) and ecology. Small trees of nonflooded forest in western Amazonia at ca. 100- 200 m. Flowering specimens collected from March to June. Fruits unknown.

Additional specimens examined. PERU. LORETO: Requena, Sapuena, Arboreto Jenaro Herrera, 125 m, 2 Mar 1989 (fl juv), Freitas 53 (MO).

BRAZIL. AMAZONAS: Rio Jutahy, Riosinho Juruema, 2 Jun 1945 (fl d), de Lemos Fr6es 21022 (F, NY).

Local names. Peru: hioma cocuir-ey (Huitoto), puspo moena.

Endlicheria cocuirey is a poorly known species that shares subsessile leaves with contracted or subcordate bases and adaxially prominent tertiaries with E. bracteata and E. verticillata. The latter has glabrous flowers, and E. cocuirey is distinguished from E. bracteata by its short tomentose rather than stiff hirsute indument. Similar indument and leaves, but with immersed tertiaries above, are otherwise found only in E. dictifarinosa.

31. Endlicheria tessmannii 0. C. Schmidt, Notizbl. Bot. Gart. Berlin-Dahlem 10: 227. 1928. Type. Peru. Loreto: Iquitos, without date (fl d), Tess- mann 5146 (holotype: B-n.v.; isotype: NY).

Trees to 22 m. Branchlets stout, midway along flush 4-8 mm diam., angular, densely pubescent, the surface concealed by a dense tomentellose cover of grey to yellowish hairs (0.1 mm or less) interspersed with straight to crookedly sprawling rusty hairs of 0.7-1 mm; terminal buds plump, 5 X 4 mm, densely pubescent, the hairs as branchlets, ascending to appressed. Leaves closely spiraled at tips of current flush; petioles robust, to 7 X 0.5 cm, semi-terete, the indument as on branchlets; laminae chartaceous to membranaceous, plane, obovate to elliptic, 20-45 X 8-15 cm, the base acute, the apex obtuse, acuminate for up to 4 cm, the margins minutely recurved throughout; upper surface dull greyish green to olive- brown, the primary to fourth-order veins raised, their prominence decreasing with rank; lower surface sparsely pubescent, the hairs appressed to ascending, 0.1 mm long, denser on main veins, interspersed with sprawling early caducous hairs of 0.7 mm on the midrib, all vein orders raised, their prominence decreasing with rank; secondary veins 10-15 per side, ? evenly spaced, slightly more distant around midlamina, patent, diverging at 70-850 (more obtuse around midlamina), arcuate to arching after midcourse, distal pairs loop-connected; tertiaries oblique to midrib, between secondaries forked. Staminate inflorescences distally clustered in the axils of cataphylls, to 30 cm long with 15 lateral branches, branch orders 3-4, the highest order botryoid to irregular, the flowers loosely crowded, the axes densely pubescent, the indument as


on branchlets; bracts and bracteoles persistent at anthesis, ovate, densely pubescent, the hairs appressed to ascending, silvery grey; pedicels of terminal flowers terete, to 0.5 mm long, lacking below penultimate flowers. Flowers campanulate, 2.5 mm diam., densely pubescent outside, the hairs yellowish to grey, 0.3 mm long, straight, ascending, sparser towards tepals; receptacle cyathiform, 0.6 X 0.6 mm, glabrous inside. Tepals membranaceous, elliptic, 1.2 x 0.7 mm, spreading to recurved at anthesis, the inner surface glabrous, the margins and apex inside papillose. Sta- mens of whorls I and II stipitate, 0.6 mm tall, the anthers transversely oblong, 0.25 X 0.3 mm, glabrous, the apex truncate to emarginate, the connectives level with or reduced between the 2 locelli, these suborbicular, introrse, the filaments laminar, narrower than anthers, the base sparsely grey-pilose; whorl III stamens stipitate, ca. 0.6 mm tall, the anthers oblong, 0.3 X 0.2 mm, erect, locelli 2, extrorse-latrorse, the filaments narrower than anthers, laminar, the base sparsely grey-pilose, the basal glands sessile, globose; whorl IV wanting; pistillode filiform. Pistillate inflorescence with indument and color as in staminate plants, but shorter and with fewer lateral branches, the flowers slightly deeper; stamens sterile, smaller; ovary glabrous, ovoid; style slender, distinct from ovary; stigma tri-lobed, 0.4 mm diam. Fruits borne on stout terete pedicels of up to 1 X 0.4 cm; cupules hemispherical, to 1 x 2 cm, sparsely brown tomentellose outside, the indument thinning with age, densely yellowish strigose inside, the margins entire; drupes ellipsoid, to 2 X 1.5 cm.

Distribution (Fig. 21) and ecology. Trees from nonflooded forests around Iquitos, Peru, at ca. 100 m. Flowering September to November, fruits January to July.

Representative specimens examined. PERU. LOR- ETO: Maynas, Iquitos, carretera Iquitos-Nauta, km 21.5, trocha de penetraci6n hasta el caserfo de Yarana, 120 m, 1 Feb 1995 (fr), Rimachi 11305 (MO); 7 km SW of Iquitos, 31 Jul 1972 (fr), Croat 18609 (MO); Estaci6n Experimental IIAP, Allpahuayo, 106 m, 1 Jun 1990 (fr), Vdsquez & Jaramillo 13945 (MO); Las Amazonas, Explornapo Camp, 100-140 m, Feb 1991 (fr), Pipoly etal. 13554 (MO); Mishuyacu, near Iquitos, 100 m, Oct-Nov 1929 (fl d), Klug 161 (F, NY, US); Punchana, trocha de la Comunidad de San Antonio, 120 m, 30 Jun 1998 (fr), Rimachi 12263 (MO); Rio Amazonas, trail from Caserfo Santa Maria de Ojeal, 30 Sep 1977 (fl d), Rimachi 3226 (MO); Yanamono Explorama Tourist Camp, halfway between Indiana and mouth of Rio Napo, 130 m, 23 Jan 1983 (fr), Gentry et al. 39734 (F, G, MO); Rio Mo- m6n, 4 Sep 1972 (fl ; ma), Croat 19982 (MO); Estaci6n Biologica Callicebus, 130, 26 Oct 1980 (fl d), Vasquez et

al. 651 (F, G, MO, NY); Varadera de Mazan, 27 Sep 1972 (fl d ), Croat 20771 (F, G, MO).

Local names. Peru: muena, moena amarilla.

Endlicheria tessmannii has unique indument in the Ampelodaphne species group. Vegetative surfaces of no other species therein show the extremely short (ca. 0.1 mm) hairs found in E. tessmannii. Moreover, only here are hairs on the laminae and veins below appressed or weakly ascending. In all other species of the Ampelodaphne species group, the hairs are erect on the lower leaf surface. Further, the flowers of E. tessmannii bear an indument of short (ca. 0.2 mm) straight hairs on the outer surface that is quite unlike the villose indumentum typically found in this species group. These characters circumscribe a narrowly distributed species that is morphologically close to the widespread and highly variable E. directonervia.

32. Endlicheria directonervia C. K. Allen, Mem. New York Bot. Gard. 10(5): 60. 1964. Type. Ven- ezuela. Amazonas: Cerro Sipapo, 125 m, 17 Jan 1949 (fl d), Maguire & Politi 28428 (holotype: NY; isotypes: GH, US).

Trees to 30 m. Branchlets slender to stout, midway along flush 4-8 mm diam., terete, densely greyish or rusty tomentose, the surface concealed by the indument cover, the hairs relatively long, to 0.7 mm, straight, erect, sometimes interspersed with longer crookedly erect hairs of 1 mm; terminal buds plump, 6 X 4 mm, densely pubescent, the hairs as on branchlets, erect to ascending. Leaves closely spiraled at tips of current flush; petioles robust, to 7 X 0.5 cm, semi- terete, the indument as on branchlets; laminae chartaceous to membranaceous, plane, obovate to elliptic, 15-45 X 6-18 cm, the base acute, the apex obtuse, acuminate for up to 4 cm, the margins minutely recurved throughout; upper surface green to olive- brown, sparsely tomentose, the primary to fourth- order veins raised, their prominence decreasing with rank; lower surface moderately pubescent, the hairs of mixed length, 0.3-1 mm, erect to ascending, straight to crooked, longer hairs denser, persisting, all vein orders raised, their prominence decreasing with rank; secondary veins 12-18 per side, ? evenly spaced, slightly more distant around midlamina, patent, diverging at 70-85o (in lower lamina often perpendicular to midrib, more acutely towards apex), arcuate, or arching after midcourse, distal pairs loop-connected; tertiaries oblique to midrib, between secondaries forked. Staminate inflorescences distally clustered in the axils of cataphylls, to 20 cm long with

70 FLORA NEOTROPICA

10 lateral branches, branch orders 3-4, the highest order botryoid to irregular, or condensed, the flowers clustered, the axes rusty to grey-tomentose; bracts and bracteoles persistent at anthesis, ovate, densely silvery grey-villose; pedicels terete, to 2 mm long, those supporting secondary flowers slightly shorter. Flow- ers campanulate, 3 mm diam., sparsely grey or densely yellowish pubescent outside, the hairs 0.7 mm long, crooked; receptacle cyathiform, 0.6 x 0.6 mm, glabrous inside. Tepals membranaceous, ovate, 1.2 X 0.7 mm, spreading to recurved at anthesis, the inner surface glabrous, the margins and apex inside papillose. Stamens of whorls I and II stipitate, 0.6 mm tall, the anthers transversely oblong, 0.25 X 0.3 mm, glabrous, the apex truncate to emarginate, the connectives level with or reduced between the 2 locelli, these suborbicular, introrse, the filaments laminar, narrower than anthers, the base sparsely grey-pilose; whorl III stamens stipitate, ca. 0.6 mm tall, the anthers oblong, 0.3 X 0.2 mm, erect, locelli 2, extrorse- latrorse, the filaments narrower than anthers, laminar, the base sparsely grey-pilose, the basal glands sessile, globose; whorl IV wanting; pistillode filiform. Pistil- late inflorescence with indument and color as in staminate plants, but shorter and with fewer lateral branches, the flowers slightly deeper; stamens sterile, smaller; ovary glabrous, ovoid; style slender, distinct from ovary; stigma tri-lobed, 0.4 mm diam. Fruits borne on terete pedicels of up to 1 X 0.3 cm; cupules hemispherical, to 1 X 2 cm, sparsely brown tomentose outside, the indument thinning with age, densely yellowish strigose inside, the margins entire; drupes ellipsoid, to 2 X 1.5 cm.

Distribution (Fig. 22) and ecology. Small to large trees of both flooded and nonflooded forests in Ama- zonia and adjacent lower montane eastern Andean slopes, to ca. 1500 m. Flowering June through following January, fruits available throughout the year.

Representative specimens examined. COLOMBIA. AMAZONAS: Leticia, near El Marco, 200 m, 7 Sep 1963 (fl d), Soejarto 580 (GH, K, US); Rio Apaporis, Cachivera de Jirijirimo y alrededores, 250 m, 12 Jun 1951 (fr), Schultes & Cabrera 12423 (GH, NY, U, US); Soratama, 250 m, 21 Jun 1951 (fr), Schultes & Cabrera 12735 (GH, NY). CA- QUETA: Cordillera Oriental, vertiente oriental, bosques entre Sucre y La Portada, 1200-1350 m, 5 Apr 1940 (fl d), Cua- trecasas 9123 (COL, US); Florencia, carretera Florencia- Suaza, km 28, 1000-1400 m, 10 Nov 1993 (fl d), Ramirez et al. 4886 (MO). META: Sierra de la Macarena, 1500 m, 30 Dec 1949 (fr juv), Philipson & Idrobo 2013 (COL, NY).

VENEZUELA. AMAZONAS: Yavita, Rio Guiania, Ma- roa, 127 m, 14 Feb 1942 (fr), Williams 14370 (A, G, F); Rio Negro, Cano 12 km NE of San Carlos, 120 m, 15 Apr 1979 (fr), Liesner 6657 (MO).

ECUADOR. MORONA-SANTIAGO: Pumpuentza, 250 m, 29 Jun 1980 (fr), Brandebyge & Asanza 32418 (MO); Tukupi, 250 m, Jun 1980 (fr), Brandebyge & Asanza 32271 (MO). NAPO: Aguarico, Reserva Ethnica Huaorani, carretera y oleoducto de Maxus, km 119-120, 235 m, Mar 1995 (fr), Aulestia et al. 3593 (MO); Yasuni Forest Reserve, E of Pontificia Universidad Cat6lica del Ecuador Scientific Sta- tion, 225 m, 20 Jun 1995 (fr), Acevedo & Cedenlo 7416 (MO); Archidona, carretera Hollin-Loreto, Rfo Huataraco, 800-1000 m, 23-30 Aug 1989 (fr), Cer6n & Factos 7649 (MO); Loreto, faldas del Volcan Sumaco, 690 m, 2 Mar 1996 (fl Y, fr juv), Vargas & Grefa 764 (MO); Tena, Estaci6n Biol6gica Jatun Sacha, 450 m, 12 Nov 1987 (fr), Cer6n 2679 (HBG, MO). PASTAZA: Puerto Sarayacu, 3 Oct 1974 (fl 6), Lugo 3906 (GB); Pozo petrolero Villan6n 2 de ARCO, 400 m, 1-18 Dec 1991 (fl 6), Hurtado 2979 (MO). SANTIAGO- ZAMORA: Taisha, 1500 ft, 7 Feb 1962 (fr), Cazalet & Pen- nington 7716 (NY). SucuMBios: Cascales, Parroquia El Dorado, 250 m, 5 May 1997 (fr), Freire et al. 2226 (MO).

PERU. AMAZONAS: Bagua, Imaza, Comunidad de Ya- mayakat, 500 m, 17 Oct 1996 (fl d), Vdsquez & Jaramillo 20300 (MO); Rio Cenepa, vic. of Huampami, 200-250 m, 8 Aug 1978 (fr), Ancuash 1343 (F, G, HBG). HUANUCO: Pachitea, Pucallpa, 1080 m, 22 Dec 1987 (fl 6), Wallnofer 14-221287 (MO). LORETO: Requena, Sapuena, Arboreto Jenaro Herrera, 170 m, 15 Sep 1987 (fl juv), Vdsquez & Jaramillo 9606 (F, MO, NY), 3 Nov 1980 (fl Y), Castillo 65 (F). PAsco: Oxapampa, Iscozacfn, 16 Jun 1986 (fl 6), Pariona & Sebastian 25 (F, MO, NY); Palcazu, 300-600 m, 4 Jun 1984 (fl 6), Hartshorn et al. 2600 (MO, NY). PUNO: Carabaya, cabeceras del Rfo Candamo, 800-850 m, 14 Nov 1996 (fr), Cornejo & Balarezo 2701 (MO).

BRAZIL. ACRE: Mancio Lima, Mancio Lima Ramal do Banho, 8 Nov 1991 (fr), Cid Ferreira et al. 10642 (MO): AMAZONAS: Humaita, on plateau between Rio Livramento and Rio Ipixuna, Cipoal, 7-18 Nov 1934 (fr), Krukoff 7206 (A, NY); Estrada Manaus-Caracarai, km 130, 13 Nov 1973 (frjuv), Berg et al. P19527 (HBG, INPA, MO, NY); Reserva Florestal Ducke, Manaus-Itacoatiara Rd., km 26, Bosque de Palmeiras P-1068, Jul 1963 (fl 6), Rodrigues 5389 (F, HBG, INPA, NY); Presidente Figueiredo, Rio Uatuma, UHE Bal- bina, 15 Aug 1986 (fl 6), Freitas et al. 239 (INPA); Rio Ituxi, Labrea airstrip, 28 Jun 1971 (fl 9 ), Prance et al. 13888 (HBG, INPA, MO, NY, US); Rio Negro, right side of Ilha Tamandua (locally Ilha Maraj6), 19 Oct 1987 (fr), Maas et al. 6798 (K, MO, NY, U, US); Sao Gabriel de Cachoeira, Camanaus, 19 Oct 1978 (fl 6), Nascimento 685 (HBG, MG, NY).

Local names. Colombia: ha-ro, las brisas, laurel.

Venezuela: laurel hediondo. Ecuador: ocatoe (Huaor- ani). Peru: muena, palta moena, roble anis amarillo, tinci, tikis, uchitinchi, und yuwich (Huambisa), wau

yuwich (Huambisa). Brazil: louro, louro cururti.

As circumscribed here Endlicheria directonervia is a highly variable species. Type material is conspicuous for rusty tomentose indumentum and secondary


; < - e; ~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~. . .. ....o;. -v . .

FIG. 22. Distibution of Endlicheria directonervia.~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~ .. .. .. ..

AMr.

...4 ~~~~~~~~~~~~ A "'k ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ A~-.-

.,.~~~~~, ~~ ~ ~

Wk AVjA *~R~

FIG. 22. Distribution of Endlicheria directonervia.

veins directed ? perpendicular to midrib, but only Rodrigues 5389, Berg et al. P19527 (both near Ma- naus, Brazil), Soejarto et al. 580 (Leticia, Colombia), and Wallnofer 14-221287 (Huanuco, Peru) match these characters. All other specimens here assigned to E. directonervia depart from typical material by one or more characters.

In Williams 14370, from near the type locality in Venezuela, and several collections from Ecuador (e.g., Lugo 3925 and Hurtado 2979), the perpendicularly directed secondaries of the type combine with sparse sprawling greyish hairs. Provided that these can be assigned to E. directonervia, specimens with similar indument (e.g., Ceron & Factos 7649) or with a mixture of tomentose and longer sprawling greyish hairs (e.g., Vdsquez & Jaramillo 20300), but without perpendicularly directed secondaries, become acceptable as well. Similarly, another variant combines dense greyish tomentose indument with (e.g., Freitas et al. 239 and Ram(rez et aL 4886) and without (e.g., Cua- trecasas 9123) laterally directed secondaries. The re-

maining specimens, Acevedo & Cede*io 7416, Aules- tia et al. 3593 (Napo, Ecuador), Schultes & Cabrera 12423 & 12735 (Amazonas, Colombia), and Vdsquez & Jaramillo 9606 and Vdsquez et aL 12391 (Loreto, Peru) have non-perpendicular secondaries and relatively short erect hairs of ca. 0.3 mm on the leaves below interspersed, on the midrib, with sprawling 0.5-0.7 mm hairs. These specimens approach E. tessmannii, but their hairs are still an order of magnitude longer (0.1 mm) and erect rather than appressed.

The variation in E. directonervia can be arranged so that specimens depart from each other by only one character, either indument or venation. Furthermore, no geographic or altitutudinal correlation can be dis- cerned. Therefore, separation into distinct species seems unwarranted.

33. Endlicheria chalisea Chanderbali, Novon 6: 329. 1996. Type. Guyana. Essequibo: Potaro- Siparuni Region, Kato and vicinity, dry forests N

72 FLORA NEOTROPICA

of town, 750 m, 19 Mar 1989 (fl d), Hahn et al. 5795 (holotype: MO; isotypes: F, NY, U, US).

Trees to 25 m. Branchlets stout, midway along flush 5-8 mm diam., terete, densely hirsute, the surface concealed by the indument cover, the hairs long, to 1.2 mm, straight, rigidly erect, yellowish; terminal buds plump, 2 X 1.5 mm, densely pubescent, the hairs as on branchlets, ascending. Leaves loosely spiraled at tips of current flush; petioles robust, to 4 X 0.5 cm, semi-terete, the indument as on branchlets; laminae chartaceous to coriaceous, plane to subbullate, broadly elliptic to obovate, 15-30 X 7-16 cm, the base acute, the apex obtuse, acuminate for up to 2.5 cm, the margins minutely recurved throughout; upper surface green to olive-brown, waxy, the midrib and secondaries immersed to sunken, the higher-order venation raised; lower surface densely yellowish hirsute, the hairs erect, up to 0.8-1.2 mm long, denser on main veins, all vein orders raised, their prominence decreasing with rank; secondary veins 6-14 per side, + evenly spaced, slightly more distant around midlamina, patent, diverging at 70-85o (more acutely towards apex), arching after midcourse, distal pairs loop- connected; tertiaries oblique to midrib, between secondaries once-forked to straight. Staminate inflorescences distally clustered in the axils of cataphylls or foliage leaves, to 7 cm long with 8 lateral branches, branch orders 2-3, the highest order condensed, the flowers clustered, the axes densely yellow to reddish hirsute; bracts and bracteoles persistent at anthesis, ovate, the indument as on axes; pedicels terete, short, to 1 mm long, reduced to lacking below secondary flowers. Flowers campanulate, up to 3 mm diam., densely pubescent, the hairs yellowish, straight to crooked, sparser towards tepals; receptacle cyathiform, 1 X 1 mm, slightly constricted below tepals, glabrous inside. Tepals membranaceous, elliptic, 1.3 X 1 mm, the tips recurved at anthesis, otherwise erect, the inner surface glabrous, the margins and apex inside lightly papillose. Stamens of whorls I and II 0.5 mm tall, stipitate, the anthers transversely oblong, 0.25 X 0.5 mm, glabrous, the apex truncate to emarginate, the connectives level with or reduced between the 2 locelli, these suborbicular, introrse, the filaments laminar, much narrower than anthers, glabrous; whorl III stamens stipitate, 0.6 mm tall, the anthers depressed-oblong, 0.3 X 0.4 mm, erect, locelli 2, extrorse-latrorse, the filaments narrower than anthers, laminar, glabrous, the basal glands sessile, globose; whorl IV staminodial, stipitate, the apex elliptic- ovate; pistillode fusiform. Pistillate inflorescence with indument, color, and branching as in staminate plants, the flowers sessile, slightly deeper; stamens sterile,

smaller; ovary glabrous, ellipsoid, 0.5 X 0.3 mm; style slender, distinct from ovary; stigma tri-lobed, 0.3 mm diam. Fruits bome on short claviform pedicels of up to 0.7 X 0.5 cm; cupules hemispherical, to 2 X 2 cm, densely velutinous outside, the indument thinning with age, sericeous inside, the margins entire; drupes ellipsoid, to 4 X 1.5 cm.

Distribution (Fig. 23) and ecology. Medium- sized trees of lowland Amazonia and upland forests of the Guiana highlands and easterm Andean slopes to ca. 2500 m. Flowering specimens collected from December through August, fruits available from Feb- ruary through November.

Representative specimens examined. VENEZUELA. AMAZONAS: Atabapo, Alto Rfo Casiquiare, 160 m, 6 Mar 1990 (fr), Aymard & Delgado 8483 (MO, NY); Rfo Mata- cuni, 1 Feb 1990 (fr), Stergios & Velasco 14495 (HBG, INPA, NY, US).

GUYANA. EsSEQUIBO: Potaro-Siparuni, Iwokrama Rainforest Reserve, 13-27 Jul 1996 (fl d), Chanderbali et al. 159 (MO); N Pakaraimas, Ciong Valley, Manawarrai Mtn., 2000 m, 1 Jun 1995 (fr), Mutchnick 1464 (MO); Mt. Ayangana, 1140 m, 16 Aug 1960 (fr), Tillett et al. 45163 (K, MO, NY, US); Mabura region, Kurupukari main, km 6, 23 Mar 1994 (fr), Ek & Hammond 1033 (MO); Upper Takatu- Upper Essequibo, Rupununi, rd. from Lethem to 25 km past Surama Village entrance, 90-110 m, 28 Feb 1990 (fr), Ac- evedo 3432 (MO, NY, US).

FRENCH GUIANA. Riviere Grand Inini-Bassin du Maroni, 11 Jul 1990 (st), Sabatier & Prevost 3195 (MO).

PERU. LORETO: Requena, Sapuena, Arboreto Jenaro Herrera, 8 Jul 1986 (fl d), Valcdrcel & Chota 1/98 (MO), 9 Dec 1980 (fl d), Castillo 87 (F). UCAYALI: Coronel Portillo, Cordillera Azul, 1200 m, 4 Jun 1981 (fr), Young & Sullivan 663 (MO).

BRAZIL. AMAZONAS: Manaus, Reserva Florestal

Ducke, Manaus-Itacoatiara Rd., km 26, 27 Jun 1964 (fl d), Rodrigues & Loureiro 5926 (F, HBG, NY); Presidente Fi- gueiredo, Rio Uatuma, UHE Balbina, 15 Jul 1990 (fl d), Cid Ferreira et al. 7582 (K, MO, NY). PARA: Oriximina, Rio Trombetas, terra firme adjacent to lago Moura, 25 Aug 1980 (fl d), Cid Ferreira et al. 1832 (HBG, INPA, MO).

BOLIVIA. LA PAZ: Prov. Sud Yungas, 7 km de Huan- cane en carretera a San Isidro, 2300 m, 13 Dec 1989 (fl 9), Smith et al. 13908 (MO); ibid., Huancane, 7.5 km hacia el sud sombre el camino nuevo, 2410 m, 9 Mar 1980 (fr), Beck 3192 (MO).

Local names. Venezuela: laurel carutillo, laurel babosa. Guyana: yellow silverballi (locally usually applied to Aniba spp.), wild pear. Peru: moena de hoja grande. Brazil: louro de folha larga, louro imbauba.

Endlicheria chalisea is immediately recognized in fruit by its densely pubescent cupules. However, it resembles E. glomerata in the long-hirsute yellowish


> . j > e - z ^ ; ~* V .'Wr

i//A

F%IG. 23. Distribution of Endlicheria chalisea and E. glomerata.

indumentum and (sub-)sessile flowers in dense clusters. The two species are sometimes vegetatively indistinguishable but the densely pubescent cupules of E. chalisea have never been found in the SE Brazilian range of E. glomerata. There, cupules, whether young or mature, are glabrous.

34. Endlicheria glomerata Mez, Jahrb. Konigl. Bot. Gart. Berlin 5: 127. 1889. Type. Brazil. Rio de Janeiro: Maua, 15 November 1874 (fl 9), Glaziou 7781 (lectotype, designated by Kostermans, 1937: P; isolectotypes: B-n.v., K).

Trees to 8 m. Branchlets slender to stout, midway along flush 4-8 mm diam., terete, densely pubescent, the surface concealed by the indument cover, the hairs yellowish, 0.4-1 mm long, straight to crooked, erect; terminal buds plump, 1 X 0.5 cm, densely pubescent, the hairs as on branchlets, ascending. Leaves spiraled at tips of current flush; petioles robust, to 3.5 X 0.4 cm, semi-terete, the indument as on branchlets; lam-

inae chartaceous, bullate (rarely plane), obovate to elliptic, 17-32 X 4-13 cm, the base obtuse, the apex obtuse to acute, acuminate for up to 2 cm, the margins minutely recurved throughout; upper surface light green to olive-brown, the midrib and secondaries con- vex between bullae (prominulous when laminae plane), the higher-order venation raised; lower surface moderately to densely pubescent, the hairs as on branchlets, slightly denser on main veins, all vein orders raised, their prominence decreasing with rank; secondary veins 10-14 per side, ? evenly spaced, slightly more distant around midlamina, patent, diverging at 70-850 (more obtuse around midlamina), arching after midcourse, distal pairs loop-connected; tertiaries oblique to midrib, between secondaries straight to forked. Staminate inflorescences distally clustered in the axils of cataphylls, to 20 cm long with 12 lateral branches, branch orders 3-4, the highest order condensed, the flowers clustered, the axes densely yellowish to rusty pubescent, the hairs as on branchlets; bracts and bracteoles persistent at anthe-

74 FLORA NEOTROPICA

sis, ovate, the indument as on axes; pedicels terete, to 0.5 mm long, reduced to lacking below secondary flowers. Flowers campanulate, 2.5 mm diam., grey- villose to glabrous outside; receptacle cyathiform, 0.5 x 0.6 mm, constricted below tepals, silvery sericeous inside. Tepals chartaceous, broadly ovate, 1 X 0.8 mm, erect to spreading at anthesis, the inner surface glabrous, the margins papillose. Stamens of whorls I and II stipitate, 0.6 mm tall, the anthers depressed- ovate, 0.3 x 0.4 mm, glabrous, the apex truncate to emarginate, the connectives level with or reduced between the 2 locelli, these suborbicular, introrse, the filaments laminar, much narrower than anthers, glabrous; whorl III stamens stipitate, equal to outer whorls, the anthers depressed-ovate, 0.3 X 0.4 mm, erect, locelli 2, extrorse-latrorse, the filaments narrower than anthers, laminar, glabrous, the basal glands sessile, globose; whorl IV wanting; pistillode fusiform. Pistillate inflorescence with indument and color as in staminate plants, but shorter and with fewer lateral branches, the flowers slightly deeper; stamens sterile, smaller; ovary glabrous, ovoid; style slender, distinct from ovary; stigma tri-lobed, 0.3 mm diam. Fruits borne on short terete pedicels of up to 5 X 3 mm, often sessile; cupules hemispherical, to 0.5 X 1 cm, glabrous inside and outside, the margins entire; drupes ellipsoid, to 2.5 X 1.5 cm.

Distribution (Fig. 23) and ecology. Small trees from the lower montane Atlantic coastal forests of SE Brazil at 200-700 m. Flowering specimens collected in May, July, September, October, and November. Fruits collected in April, June, July, August, and No- vember.

Additional specimens examined. BRAZIL. GoIAs: Niquelandia, Fazenda Engenho, 580 m, 27 Jun 1997 (fr), Oliveira et al. 770 (MO). MINAs GERAIS: Caratinga, Esta- cvo Biol8gica de Caratinga, 6 Aug 1984 (fr), Strier 992 (MO, NY); Caratinga, Fazenda Maced6nea/Cenibra, Ipaba Trilho do Triangulo (T2-37), 4 Nov 1991 (fr), Braga et al. s.n. BHCB 27250 (MO); Ilheu, Fazenda da Tabunha, Capi- chava, 220 m, 31 Aug 1930 (fr), Mexia 5025 (G, GH, NY, US); Leopoldina, Rod. BR-1 16, Rio Pombos, 14 Jul 1988 (fl 5), Hatschbach & Silva 52169 (HBG, US), 10 Oct 1992 (fl 5), Hatschbach et al. 57930 (HBG, MO, NY, S); Mar- lieria, Parque Estadual do Rio Doce, 400 m, 18 Sep 1975 (fl d), Heringer & Eiten 15083 (HBG, US), 22 Sep 1975 (fl 5), Heringer & Eiten 15154 (MO, US), 23 Mar 1976 (fl Y), Heringer 16026 (HBG, K, U), 200 m, 30 Aug 1973 (fr), Martinelli et al. 23 (MO), 29 Jun 1993 (fr), Costa s.n. BHCB 32676 (MO), 13 Sep 1997 (fl 5), Lombardi 1987 (MO); Tim6teo, Parque Estadual do Rio Doce, 20 Oct 1982 (fl d), Heringer 18530 (HBG), 22 Oct 1982 (fl d), Heringer 18580 (HBG); Vi,oca, oxcart rd. to Sao Miguel, ca. km 4, 685 m, 23 Sep 1930 (fr), Mexia 5091 (F, G, GH, MO, NY, S, U,

US), 3 Oct 1930 (fl d), Mexia 5138 (F, G, GH, MO, NY, P, S, U, US). RiO DE JANEIRO: Governador Portela, Monte Sinai, 1935 (fl 9, fr), Machado Nunes 318 (HBG, MO, U); Maud, Nov 1890 (fl d), Schwacke s.n. R 30973 (R), 26 Jun 1901 (fr) Hemmendorif 457 (R, S); Morro de Viraqa6, Praia Grande, 3 Apr 1891 (fl 9, fr), Glaziou 18451 (F, L, K, NY, P); Capelinha de St. Antonio, 22 Aug 1894 (st), Glaziou s.n. (E, NY); Rio das Flores, Fazenda Santa Genoveva, 500-600 m, 7 Oct 1971 (fl 5), Sucre 7779 (HBG, MO); s.loc., 13 May 1878 (fr), Glaziou 8093 (K, P).

Local names. Brazil: canela, canelao.

Endlicheria glomerata alone of the Ampelodaphne species group inhabits the Atlantic coastal forests of SE Brazil. In most flowering specimens, a long hirsute vestiture of stiffly erect yellowish 1 mm hairs combines with bullate laminae and densely clustered subsessile pubescent flowers. This character combination is unique to E. glomerata but appears to be unstable. Laminae range from strongly bullate in Mexia 5138, through moderately so in the type material, to plane in Glaziou 18451 and Oliveira et al. 770 without accompanying vegetative or floral differences.

Two other indument variants show parallel insta- bility in bullae formation. The specimens from Parque Estadual do Rio Doce (Costa s.n. BHCB 32676, Her- inger 18530, 18580, and 16026, Heringer & Eiten 15083 and 15154, Lombardi 1987, and Martinelli et

al. 23), have a much shorter indumentum of 0.4 mm hairs, while in Hatschbach et al. 57930, Hatschbach & Silva 52169, and Sucre 7799 vegetative indument

is as in typical material but distal inflorescence branches and flowers are glabrous. Attaching specific importance to the single character that distingishes both variants from typical material would result in three very similar, and sympatric, species that cannot be confused with others in the genus, but probably with each other-in my opinion an unnecessary in- flation of species diversity.

35. Endlicheria anomala (Nees) Mez, Jahrb. Kon- igl. Bot. Gart. Berlin 5: 133. 1889. Goeppertia anomala Nees, Syst. laur. 370. 1836. Type. Brazil. Amazonas: Rio Amazonas, near Ega, Sep 1831 (fl 5), Poeppig 2552 (lectotype, designated by Kos- termans, 1937: W-n.v.; isolectotypes: B-n.v., BM-n.v., G-n.v., KIEL-n.v., LE-n.v., LZ-n.v., NY, OXF-n.v., P-n.v.).

Goeppertia polyantha Meisn. DC. Prodr. 15(1): 175. 1864. Syntypes. Brazil. Amazonas: Rio Negro, Barra, Jul 1851 (fl 5), Spruce 1648 (B-n.v., BM- n.v., G-n.v., K-n.v., LE-n.v., NY-n.v., OXF-n.v., P-


n.v., US-n.v.); ibid., Apr 1851 (fl d juv), Spruce 1433 = 'Nectandra (6)' (B-n.v., BM-n.v., E, G-n.v., GH-n.v., K-n.v., LE-n.v., M, NY-n.v., OXF-n.v., P-n.v.). Guyana. Without locality and date (fl Y), Schomburgk 784 (BM-n.v., B-n.v., F-n.v., G-n.v., K, MO, OXF-n.v., P, U, US, W-n.v.).

Ocotea simulans C. K. Allen, Mem. New York Bot. Gard. 10(5): 99. 1964. Type. Venezuela. Amazonas: Rio Casiquiare, just above Capihuara, 100-130 m, 24 Jun 1959 (fl 6), Wurdack & Adderly 43169A (holotype: NY; isotype: US).

Trees to 15 m or shrubs. Branchlets slender, midway along flush 2-3 mm diam., distally weakly angular, soon terete, densely pubescent, the surface concealed by the indument cover to barely visible, the hairs short, to 0.5 mm, straight, appressed, ascending, or erect, silvery-grey to rust-brown; terminal buds slender, 3 X 1 mm, densely pubescent, the hairs as on branchlets. Leaves alternate, widely and evenly spaced along current flush; petioles slender, to 1 X 0.2 cm, semi-terete, the indument as on branchlets; laminae chartaceous, plane, ovate, 8-17 X 2-4 cm, the base obtuse to rounded, briefly decurrent, the apex acute, acuminate for up to 2 cm, the margins minutely recurved throughout; upper surface dull greyish green to dark olive-brown, waxy, the midrib sunken, secondary and higher-order venation raised, their prominence decreasing with rank; lower surface densely sericeous or tomentose, the hairs appressed, ascending, or erect, uniformly distributed, all vein orders raised, their prominence decreasing with rank; secondary veins 4-6 per side, + evenly spaced, slightly more distant around midlamina, ascending at 50-60? (more obtuse around midlamina), arcuate, distal pairs loop-connected; tertiaries roughly horizontal to oblique to midrib, between secondaries straight to forked. Staminate inflorescences evenly spaced along current flush in the axils of foliage leaves, to 10 cm long with 8 lateral branches, branch orders 3-4, the highest order dichasial, lax, the flowers distant, the axes sparsely rusty to grey-strigillose; bracts and bracteoles caducous by anthesis, lanceolate, the indument as on axes, denser; pedicels terete, to 3 mm long, those supporting secondary flowers slightly shorter. Flowers rotate, 5 mm diam., sparsely rusty strigillose outside; receptacle patelliform, 0.5 X 1 mm, densely rusty tomentose inside. Tepals chartaceous, obovate, 2 X 1 mm, spreading, the androecium exserted at anthesis, the outer surface sparsely rusty to grey-strigillose, the inner surface sparsely rusty to grey-strigillose near base, otherwise minutely papillose throughout, the margins papillose. Stamens of whorls I and II stipitate, 0.7 mm tall, the anthers transversely oblong to obovate, 0.3 X 0.4 mm, glabrous,

the apex truncate, the connectives level with or broad above the 2 locelli, these suborbicular, the filaments laminar, narrower than anthers, slender, glabrous; whorl III stamens stipitate, ca. 1 mm tall, the anthers oblong, 0.5 X 0.3 mm, bowed towards the outer whorls, locelli 4, upper pair introrse-latrorse, lower pair extrorse-latrorse, the filaments narrower than anthers, laminar, glabrous, the basal glands sessile, globose; whorl IV wanting; pistillode fusiform. Pistillate inflorescence with indument and color as in staminate plants, but shorter and with fewer lateral branches, the flowers similar in size and shape; stamens sterile, smaller; ovary glabrous; style slender, distinct from ovary; stigma tri-lobed, 0.4 mm diam. Fruits borne on claviform pedicels of up to 1 X 0.5 cm; cupules scarcely broader, patelliform, to 0.6 cm diam., strigose inside, the margins lobed, tepal bases persisting; drupes ellipsoid, to 1.2 X 0.6 cm.

Distribution (Fig. 24) and ecology. Small to medium-sized trees or shrubs frequent in inundated lowland forests (ca. 50-300 m) throughout the Am- azon basin. Flowering and fruiting throughout the year.

Representative specimens examined. COLOMBIA. AMAZONAS: Puerto Narifio, Rio Loretoyacu, Trapecio Amaz6nico, 100 m, Jun 1973 (fl d), Soejarto et al. 4198 (NY); Rio Yari, Quebrada El Mochilero, 120-200 m, 21 Apr 1986 (fr), Galeano et al. 1024 (MO). META: Puerto L6pez, 240 m, 27 Aug 1944 (fl Y), Little & Little 8259 (COL). VAUPES: Rio Vaupes, Mitd, 9 Jul 1976 (fl d), Zarucchi et al. 1830 (COL, MO); 2 mi upriver from Mitui, 1 Nov 1976 (fl 6), Davis 216 (COL, K). VICHADA: Parque Nacional Natural "El Tuparro," La Linea Roja, just S of Rio Tomo, 100 m, 12 Mar 1985 (fr), Zarucchi and Barbosa 3679 (MO).

VENEZUELA. AMAZONAS: Atabapo, Rio Asisa, 100 m, Oct 1989 (fl Y), Delgado 868 (MO); Atures, Puerto Aya- cucho, 37 m, 23 Jul 1981 (fl 6), Castillo 1282 (MO, NY). ApuRE: Pedro Camejo, Laguna la Guacharaca, 70 m, 24 Feb 1979 (fl d), Davidse & Gonzales 15701 (MO); Parque Na- cional "Santos Luzardo," 5 Apr (fr), G6mez et al. 649 (MO). BOLiVAR: Cedefno, Rio Parguaza, 30 m, 25-28 Jan 1989 (fr), Cuello 637 (MO, NY); Rio Pargueni, 1-10 km above mouth, 90 m, 10 Dec 1955 (fl 6, most diseased), Wurdack & Mon- achino 39775 (NY); Sierra Ichdn, cercanias del Salto Maria Espuma (Salto Ichdn) del Rio Ichdn, base de la sierra Ichdn, tributary del Rio Paragua, 500 m, 28 Dec 1961 (fr), Stey- ermark 90315 (NY).

ECUADOR. NAPO: Aguarico, Parque Nacional Yasuni, 200 m, 22 Sep 1988 (fl Y, fr juv), Ceron & Gallo 4898 (MO, NY); Rio Lagarta Cocha, 6 Aug 1992 (fl 6), Delprete et al. 6141 (MO). SUCUMBiOS: Lago Agrio, Reserva Faun- istica Cuyabeno, 230 m, 4 Oct 1991 (fl d), Palacios et al. 8144 (MO).

PERU. LORETO: Loreto, Nauta, Reserva Nacional Pacaya-Samiria, Rio Yanayacu, campamento Palizada, 90 m,

76 FLORA NEOTROPICA

. | . . .|.f. ~~~~~~~~~~~ ', _ ̂-S O r | 0 r i | i | w . | . . . X T f~~~~~~~~~~~~~~~~. .. .. . .

7 4 - -

FIG. 24. Distribution of Endlicheria anomala.

3 Nov 1992 (fl 6), Del Carpio & Ruiz 1620 (MO); Maynas, Iquitos, 100 m, 3-11 Aug 1929 (fl c), Killip & Smith 27192 (NY); Requena, Avispa Cocha, Rfo Tapiche, 180 m, 9 Jan 1984 (fl 6), Vdsquez et aL 4778 (MO). UCAYALI: Coronel Portillo, Yarinacocha, Cafno de Alijandriam vecindad de Lago Yarinacocha, 210 m, 22 Nov 1997 (fl Y, fr), Graham & Schunke 365 (MO).

BRAZIL. ACRE: Cruzeiro do Sul, Rio Jurua, Igarape Preto, 29 Oct 1966 (fl juv), Prance et aL 2972 (HBG, NY, R). AMAZONAS: Barcelos, Rio Araca, 29 Jul 1985 (fl 6), Cordeiro 309 (INPA, MO); Humaita, Rio Madeira, lago do Purusinho, 15 May 1985 (fl 6), Henderson etal. 458 (INPA, K, MO, NY); Manaus, 28 Mar 1937 (fl Y, fr), Ducke 440 (K, MO, NY, R, U); Bom Jardim, near Flutuante do INPA, 7 Apr 1971 (fl 6), Prance et al. 11737 (COL, INPA, MO, NY, R, U); Rio Negro, Ilha da Costa Arirarra, 28 June 1979 (fl 6), Poole 1717 (INPA, MO); Ilha das Onqas, 24 Jun 1992 (fl 6), Mori & Gracie 22473 (INPA, MO, NY); Tefe, lago de Tefe, 27 Feb 1972 (fl 6), Albuquerque et al. 570 (INPA). PARA: Oriximink, Rio Trombetas, 9 Jul 1980 (fl J), Cid Ferreira et al. 1388 (INPA, MO, NY); Rio Paru do Oeste, Lago Ara,c, 8 Sep 1980 (fl 6), Cid Ferreira et al. 2321 (INPA, MO, NY). ROND6NIA: Rio Jaciparand, 4 km above

Jaciparana, 28 Jun 1968 (fl d), Prance et al. 5295 (MO, NY); Rio Pacaas Novos, 8-25 km above mouth, 6 Aug 1968 (fl d), Prance et al. 6834 (MO, NY).

BOLIVIA. BEM: Guayaramerin, Rio Mamor6, 15 Feb 1978 (fl 9, fr), Anderson 12052 (NY). LA PAZ: Prov. Itur- ralde, Luisita, 180 m, 29 Feb 1984 (fl 9, fr), Beck & Haase 10129 (MO). PANDO: Manupiri, Rio Manupiri, 278 m, Sep 1996 (fr), Paniagua & Foster 690 (MO). SANTA CRUZ:

Prov. Velasco, Reserva Ecologfia El Refugio, 160 m, 24 Apr 1995 (fl d), GuillUn & Chore 3292 (MO); Campos de San Ram6n, 8-10 Aug 1995 (fl d), Halloy et aL 4319 (MO).

Local names. Colombia: ko-ma-nee-nee-ko (Makuna), chaviaco negro, pee-shee'. Venezuela: laurel blanco, laurel blanco de orilla del rio, laurel de revalta, laurel del rio, laurel fino, laurel rebalsero, laurel oriyera, laurelito. Peru: canela, isma moena, moena de bajo, muena blanca, sanango, yacu muena. Brazil: louro canela, louro do igap6, louro flor, louro tambaqui.

Endlicheria anomala is the only species of the genus in which whorl HI stamens are consistently pro-


vided with four-locellate anthers. The rotate flowers with an androecium in which all stamens have distinct filaments and whorl III stamens are closely grouped with their anthers extrorsely bowed, are also unmistakable. The claviform pedicels and shallow platelike cupules with persistent tepal bases resemble those of E. lorastemon but are much smaller in size.

Most specimens, including the type material and the type of Ocotea simulans, bear an appressed indument of silvery hairs, the denser forms of which are reminiscent of the Endlicheria sericea species group, but several specimens, including the type material of Goeppertia polyantha, have a rusty tomentose vestiture much like that typical of Endlicheria paniculata. However, ascending hairs of intermediate orientation also appear (e.g., Cavalcante & Silva 1782 and Mori & Gracie 22473), and flowers and fruits are uniform irrespective of indument orientation.

36. Endlicheria williamsii 0. C. Schmidt, Repert. Spec. Nov. Regni Veg. 31: 177. 1933. Type. Peru. Loreto: Maynas, Rio Nanay, Maquisapa, 5 Jul 1929 (fl d), Williams 1193 (holotype: F; isotype: NY).

Trees to 20 m. Branchlets relatively stout, midway along flush 4-5 mm diam., distally weakly angular, soon terete, densely tomentose, the surface concealed by the indument cover, the hairs relatively short, to 0.3 mm, straight to crisped, erect, rusty to reddish brown; terminal buds plump, 4 X 3 mm, densely pubescent, the hairs as on branchlets, ascending. Leaves closely spiraled at tips of current flush; petioles robust, to 2.5 X 0.3 cm, terete, slightly flattened above, the indument as on branchlets; laminae chartaceous, plane, broadly elliptic, 12-20 X 6-12 cm, the base obtuse, briefly decurrent, the apex obtuse, abruptly acuminate for up to 1.5 cm, the margins minutely recurved throughout; upper surface greyish to olive- brown, minutely punctulate, the midrib and secondaries prominent, tertiary and higher-order venation immersed; lower surface obscured by a short tomentose cover of 0.3 mm long creamish hairs interspersed with straight erect reddish hairs of 0.7 mm, all vein orders raised, their prominence decreasing with rank; secondary veins 5-7 per side, ? evenly spaced, slightly more distant around midlamina, ascending at 50-60O (more acutely towards apex), the lowermost pair sometimes asymmetric (one or both merging with margin at base), distal pairs loop- connected; tertiaries roughly horizontal, between secondaries straight or forked. Staminate inflorescences distally clustered in the axils of cataphylls, to 20 cm

long with 14 lateral branches, branch orders 3-4, the highest order dichasial, lax, the flowers distant, the axes densely reddish to rusty tomentose; bracts and bracteoles caducous by anthesis, lanceolate, densely rusty strigose; pedicels terete, to 2 mm long, those supporting secondary flowers slightly shorter. Flow- ers hypocrateriform, 5 mm diam., sparsely rusty strigose outside; receptacle infundibuliform, 1 X 1 mm, densely grey-tomentose inside. Tepals chartaceous, narrowly elliptic or ligulate, 2 X 1 mm (the inner whorl slightly broader), ascending to spreading at anthesis, the inner surface and margins sparsely rusty papillose, otherwise glabrous. Stamens of whorls I and II stipitate, 1 mm tall, the anthers transversely oblong, 0.3 X 0.4 mm (slightly narrower in whorl II), the apex truncate, the connectives broad above the 2 locelli, these suborbicular, introrse, the filaments laminar, narrower than anthers, sparsely whitish grey- pilose, the hairs straight, erect; whorl III stamens broadly stipitate, 1 mm tall, pressed together, the anthers oblong, bowed towards the outer whorls, locelli 2, extrorse-latrorse, occasionally a second introrse- latrorse pair above, the filaments almost as broad as anthers, wider towards base, the indument as in outer whorls, the basal glands sessile, globose; whorl IV wanting; pistillode fusiform. Pistillate inflorescence with indument and color as in staminate plants, but shorter and with fewer lateral branches, the flowers slightly deeper; stamens sterile, smaller; ovary glabrous, ovoid; style slender, distinct from ovary; stigma tri-lobed, 0.4 mm diam. Fruits borme on short claviform pedicels of up to 0.5 X 0.3 cm; cupules hemispherical, to 1 X 1.5 cm, glabrous outside, tawny sericeous inside, the margins entire; drupes ovoid, to 2 x 1.2 cm.

Distribution (Fig. 25) and ecology. Medium- sized trees of seasonally flooded forest in western Amazonian at ca. 100-200 m. Flowering specimens collected in June and July, fruits in January, March, April, and September.

Representative specimens examined. COLOMBIA. AMAZONAS: Rio Yari, Quebrada El Mochilero, 120-200 m, 21 Apr 1986 (fr), Galeano et al. 1065 (MO).

PERU. LORETO: Maynas, Rio Ampiyacu, between Esti- r6n and Tierra Firme, 24 Apr 1977 (fr), Plowman et al. 7013 (F, GH, MO, NY); Rio Nanay, Iquitos, Almendro, 17 Jun 1976 (fl d), McDaniel & Rimachi 20744 (MO); Mishana, 120 m, 11 Jan 1976 (fr), Gentry et al. 15865 (F, G, MO, NY); QuebradaYarana, 130 m, 10 Jul 1988 (fl d), Vdsquez et al. 10939 (MO); Timbuchi, Jun-Jul 1929 (fl juv), Williams 1002 (F), (fl d), Williams 1003 (F, G), (fl Y, fr), Williams 1004 (F, G); Rio Ampiaco, 24 Sep 1972 (fr juv), Croat 20680 (HBG, MO, NY).

78 FLORA NEOTROPICA

. . . . . . . . . .~~~.t .

-7-

4~~~~~~~~~~~~~ $~~~~~~~~~~~~

FIG. 25. Distribution of Endlicheria williamsii E canescens, and E. xerampela.

Local names. Colombia: jigua. Peru: isma muena, pampa muena.

Endlicheria williamsii is immediately recognized by the mixture of short creamish tomentose indument that completely obscures the lower surface of the ter- minally clustered leaves. Similar indument combines with widely spaced leaves in E. duotincta, and on account of its phyllotaxy E. williamsii may be mistaken for a member of the Ampelodaphne species group, but its flowers suggest affinity with E. anomala. The receptacle is deeper in E. williamsii but the androecium is very similar, even occasionally showing the four-locellate whorl m anthers typical of E. anomala. (e.g., Vdsquez et al. 10939).

37. Endlicheria canescens Chanderbali, Novon 6: 328. 1996. Type. Suriname. Saramacca River Toe- koemoetoe Creek, 5 Oct 1994 (fl di), Maguire 24898a (holotype: MO; isotypes: A, F, G, K, NY, U, US, W).

Trees to 35 m. Branchlets slender, midway along flush 2-3 mm diam., angular, densely tomentose, the surface concealed by the indument cover, the hairs relatively short, to 0.3 mm, straight to crisped, erect, reddish to rust brown; terminal buds plump, 3 X 2 mm, densely rusty tomentose. Leaves alternate, widely and evenly spaced along current flush; petioles slender, to 2.5 X 0.3 cm, semi-terete, the indument as on branchlets; laminae stiff chartaceous, plane, broadly elliptic, 10-20 X 5-8 cm, the base acute, often asymmetric, briefly decurrent, the apex acute, acuminate for up to 1.5 cm, the margins minutely recurved throughout; upper surface reddish to olive- brown, minutely punctulate, the midrib and secondaries depressed, the higher-order venation immersed; lower surface densely tomentose, the hairs erect, straight, tan to light brown, slightly denser on main veins, all vein orders raised, their prominence decreasing with rank; secondary veins 4-7 per side, + evenly spaced, slightly more distant around midlamina, ascending at 50-60o (more acutely towards apex),


arcuate, the lowermost pair often asymmetric (one or both merging with margin at base), distal pairs loop- connected; tertiaries oblique to midrib, between secondaries once-forked or straight. Staminate inflorescences evenly spaced along current flush in the axils of foliage leaves, to 15 cm long with 8 lateral branches, branch orders 2-3, the highest order dichasial or irregular, lax, the flowers distant, the axes rusty tomentose; bracts and bracteoles caducous by anthesis, lanceolate, rusty tomentose; pedicels terete, to 3 mm long, those supporting secondary flowers slightly shorter. Flowers broadly rotate, to 5 mm diam., sparsely rusty to grey-strigillose outside; receptacle broadly infundibuliform, 2 X 3 mm, slightly constricted below tepals, densely grey-velutinous inside. Tepals fleshy, obovate, 1.9 X 1.3 cm, spreading at anthesis, the inner surface densely greyish white tomentose, the margins densely reddish brown papillose. Stamens of whorls I and II broadly stipitate, relatively short, 0.6 mm, the anthers depressed-ovate, 0.3 X 0.5 mm (slightly narrower in whorl II), glabrous, the apex sharply apiculate, the connectives prolonged between the 2 locelli, these obliquely hemispherical, introrse-latrorse, the filaments clavate, tapering towards base, densely grey-tomentellose outside; whorl III stamens sessile, 0.6 mm tall, the anthers depressed-oblong, 0.3 X 0.6 mm, erect, locelli 2, extrorse-latrorse, the filaments as broad as anthers, columnar; densely grey-tomentellose inside, the basal glands sessile, globose, apiculate; whorl IV wanting; pistillode filiform. Pistillate inflorescence with indument and color as in staminate plants, but shorter and with fewer lateral branches, the flowers slightly deeper; stamens sterile, smaller; ovary glabrous, ellipsoid, 1 X 0.6 mm; style stout, weakly distinguished from ovary; stigma minutely tri-lobed, 0.3 mm diam. Fruits borne on claviform pedicels of up to 1.5 X 0.7 cm; cupules hemispherical, to 1 X 1.5 cm, glabrous outside, rusty sericeous inside, the margins entire; drupes ellipsoid, to 1.3 X 0.9 cm.

Distribution (Fig. 25) and ecology. Large trees of terra firme forests from the Guianas, western Ama- zonia, and eastern Andean slopes at ca. 100-1700 m. Flowers collected in May, June, September, October, November, and January, and fruits in October, No- vember, April, May, and June. If not a sampling artifact, it is conceivable the May-June flowering sea- son results in October-November fruits, while the September-January flowering period provides fruits from April to June.

Representative specimens examined. COLOMBIA. AMAZONAS: Leticia, Tarapaca, Parque Nacional Natural

Amacayacu, Cabania Pamate, Rfo Cotuhe, 100 m, 29 Jun 1991 (fl d), Rudas et al. 2642 (MO). PUTUMAYO: Mocoa, San Antonio, Alto Campucaca, La Mariposa, 1350 m, 20 April-I May 1994 (fr), Ferndndez et al. 11101 (MO).

VENEZUELA. BOLiVAR: Between San Ignacio de Ya- runi and San Francisco de Yaruni, 1200 m, 4 Jan 1975 (fl Y), Steyermark 111387 (NY).

GUYANA. ESSEQUIBO: Upper Mazaruni River Basin,

Kamarang R., 24 Oct 1960 (fl d), Tillett & Tillett 45789 (F, K, NY, US).

SURINAME. Natuurpark Brownsberg, 16 Jun 1917 (fl

d ), Suriname Forest Bureau 2936 (A, MO, U), 23 Jun 1925 (fl d), Suriname Forest Bureau 6884 (MO, US).

ECUADOR. NAPO: Aguarico, Reserva Etnica Huaor- ani, 247 m, 10-14 Nov 1993 (fl 6), Dik 767 (MO); Reserva Faunistica Cuyabeno, Zancudo, 230 m, 5 Oct 1991 (fl Y), Palacios et al. 8164 (MO); Orellana, Parque Nacional Ya- sunf, 230 m, 6 Oct 1993 (fr), Dik 665 (QCNE-n.v., MO). PASTAZA: Pastaza, Pozo petrolero "Danta 2" de UNOCAL, 50 km SSE of Cararay, 365 m, 1-20 Oct 1990 (fl 6), Es- pinoza & Coba 378 (MO). ZAMORA-CHINCHIPE: Nangar- itza, Pachicutza, Camino al Hito, Cordillera del C6ndor, 1200-1300 m, 20 Oct 1991 (fl Y), Palacios et al. 8399

(MO). PERU. CAJAMARCA: San Ignacio, Ricardo Palma, 1650

m, 20 May 1998 (fl 6), Campos & L6pez 4930 (MO); Ta- baconas, La Bermeja, 1600-1700 m, 19 Nov 1997 (fr), Cam- pos & Cano 4695 (MO). LORETO: Alto Amazonas, Cerros Campanquiz, Pongo de Manseriche, 300-550 m, 19-21 Oct 1962 (fl 6), Wurdack 2362 (F, GH, NY, UC, US).

BRAZIL. AMAZONAS: Benjamin Constant, s.d. (st), Brees 40 (INPA, MO).

Local names. Colombia: amarillo. Brazil: louro comum.

This species was first brought to my attention as Endlicheria endlicheriopsis (Mez) Kosterm., a combination based on Ocotea endlicheriopsis Mez. How- ever, Kostermans had overlooked the four-locellate anthers in the pistillate type material of the basionym, and Endlicheria canescens was described to accom- modate the two-locellate plants formerly associated

with Ocotea endlicheriopsis (Chanderbali, 1996). From the several congeners with similar rusty to-

mentose branchlets Endlicheria canescens is easily recognized by its rotate flowers where fleshy obovate

tepals have densely greyish white tomentose inner

surfaces. Immersed tertiaries above and a tendency to have asymmetric leaf bases with a pair of secondaries emerging from the juncture of leaf base and petiole are also useful for identification. Flower shape and subsessile whorl I and II stamens with rather small locelli are reminiscent of species with Microlocellata- type anthers, of which E. ferruginosa has similar in-

dument, but in E. canescens whorl I and II anthers are

80 FLORA NEOTROPICA

provided with sharply apiculate rather than truncate apices.

38. Endlicheria xerampela Chanderbali, sp. nov. Type. Colombia. Choco: Halfway between Certe- gui and Las Animas, 100 m, 17 Aug 1976 (fl d), Gentry & Fallen 17803 (holotype: MO; isotypes: F, G, HBG, NY). Fig. 26

Endlicheriae canescentis aemulans, floribus infundibularibus et antheris sex externis late oblongis connectivo nec supra loculos producto differt.

Trees to 18 m. Branchlets slender, midway along flush 3-4 mm diam., angular, densely reddish brown tomentose, the surface concealed by the indument cover, the hairs relatively short, to 0.3 mm, straight, erect; terminal buds plump, 3 X 2 mm, densely pubescent, the hairs as on branchlets, ascending. Leaves alternate, widely and evenly spaced along current flush; petioles slender, to 1.5 X 0.15 cm, semi-terete, the indument as on branchlets; laminae chartaceous, plane, ovate, 8-15 X 3-7 cm, the base obtuse, the apex acute, acuminate for up to 1 cm, the margins minutely recurved throughout; upper surface rusty to yellowish green, sparsely rusty strigose, waxy, the midrib and secondaries immersed, the higher-order venation prominulous; lower surface sparsely rusty tomentose, the hairs as on branchlets, scattered on lamina, denser on main veins, all vein orders raised, their prominence decreasing with rank; secondary veins 4-6 per side, ? evenly spaced, slightly more distant around midlamina, ascending at 50-60o (more obtuse around midlamina), arcuate, distal pairs loop- connected; tertiaries roughly horizontal, between secondaries straight to forked. Staminate inflorescences evenly spaced along current flush in the axils of foliage leaves, to 7 cm long with 8 lateral branches, branch orders 3-4, the highest order essentially dichasial, but the internodes reduced and flowers arranged in pseudo-umbellate fascicles, the axes sparsely rusty tomentose; bracts and bracteoles caducous by anthesis, ovate, the indument as on axes; pedicels terete, to 2 mm long, those supporting secondary flowers slightly shorter. Flowers infundibuliform, 2.5 mm diam., sparsely rusty strigose outside; receptacle broadly infundibuliform, 0.5 X 1 mm, densely rusty pilose inside. Tepals chartaceous, ovate, I x 0.6 mm (the inner whorl slightly narrower), ascending at anthesis, the inner surface sparsely grey- tomentose, the margins and tips papillose. Stamens of whorls I and II stipitate, 0.4 mm tall, the anthers depressed-oblong, 0.3 x 0.4 mm, glabrous, the apex

truncate to emarginate, the connectives level with or reduced between the 2 locelli, these suborbicular, introrse, the filaments laminar, narrower than anthers, sparsely rusty pilose; whorl III stamens sessile, 0.6 m tall, the anthers depressed-oblong, 0.3 X 0.4 mm, erect, locelli 2, extrorse-latrorse, the filaments broader than anthers, ligulate, sparsely rusty pilose near base, the basal glands sessile, globose, apiculate; whorl IV wanting; pistillode minute, filiform. Pistillate inflorescence with indument and color as in staminate plants, but shorter and with fewer lateral branches. Mature pistillate flowers unknown. Fruits borue on claviform pedicels of up to 7 X 4 mm; cupules hemispherical, to 1 X 1.5 cm, glabrous outside, rusty sericeous inside, the margins entire; drupes ovoid, to 1.5 X 1 cm.

Distribution (Fig. 25) and ecology. Medium- sized trees from the lowland (0-150 m) forests of the Pacific coast of Colombia. Flowering specimens collected in April, August, and October, fruits in March and April.

Additional specimens examined. COLOMBIA. CAUCA: Bajo Calima, Concesi6n Pulapel/Buenaventura, 100 m, Oct 1987 (fl 9, juv), Monsalve 1975 (MO); Lopez, Rio Naya, "El Carmen," 50-150 m, Apr 1992 (fr), Cogollo et al. 5167 (MO). VALLE: Buenaventura, Concesi6n de Pul- papel, 0-50 m, Apr 1992 (st), Cogollo et al. 5190 (MO), (fl , juv), Cogollo et al. 5195 (MO); Corregimento San Isidro,

Quebrada Ordofiez, 40 m, Mar 1989 (fr), Devia & Prado 2669 (MO); Rio Cajambre, "El Chorro," 0-50 m, Apr 1992 (fr), Cogollo et al. 5182 (MO).

Endlicheria xerampela is most similar to E. canescens. The two share a dense reddish tomentose vestiture and deeply hemispherical cupules with a rusty sericeous indument inside. However, the upper leaf surface of E. xerampela is smooth rather than minutely punctulate, the flowers are infundibuliform rather than rotate, and apices of whorl I and II anthers are truncate rather than apiculate. None of the other species of Endlicheria known from the Choco region are remotely similar.

39. Endlicheria citriodora van der Werff, Ann. Mis- souri Bot. Gard. 78: 415. 1991. Type. Peru. Lor- eto: Requena, Sapuena, Arboreto Jenaro Herrera, 140 m, 7 Aug 1988 (fl d), van der Werf et al.

9991 (holotype: MO; isotypes: AMAZ-n.v., F, G, HBG-n.v., MO, US).

Trees to 25 m. Branchlets stout, midway along flush 4-6 mm diam., angular, striate, densely rusty tomentellose, the surface concealed by the indument


FIG. 26. Endlicheria xeramipela (A-E, Gentry & Fallen 17803; F, Cogollo et aL 5167). A. Habit. B. Flower. C. Flower l.s. D. Whorl III stamen seen from without. E. Whorl I stamen seen from within. F. Fruits.

82 FLORA NEOTROPICA

cover, the hairs short, to 0.2 mm, straight, erect; terminal buds plump, 3 X 1.5 mm, densely brown tomentellose. Leaves alternate, widely and evenly spaced along current flush; petioles striate, robust, to 2 X 0.6 cm, semi-terete, the indument as on branchlets; laminae chartaceous, plane, elliptic-ovate, 15- 30 X 7-14 cm, the base obtuse to truncate, briefly decurrent, the apex acute, acuminate for up to 1.5 cm, the margins minutely recurved throughout, often revolute near leaf base; upper surface olive-brown, the midrib and secondaries flat to immersed, the higher- order veins prominulous; lower surface sparsely pubescent, the hairs 0.2 mm long, curved, weakly ascending to appressed, uniformly distributed, all vein orders raised, their prominence decreasing with rank; secondary veins 7-10 per side, ? evenly spaced, slightly more distant around midlamina, ascending at 50-60O (more acutely towards apex), arching after midcourse, distal pairs loop-connected; tertiaries oblique to midrib, between secondaries once-forked to straight. Staminate inflorescences evenly spaced along current flush in the axils of foliage leaves, to 15 cm long with 14 lateral branches, branch orders 2- 3, the highest order essentially dichasial, but the in- termodes reduced and flowers arranged in pseudo- umbellate fascicles, the axes densely brown tomentellose; bracts and bracteoles caducous by anthesis, lanceolate, the indument as on axes; pedicels terete, to 2 mm long, those supporting secondary flowers slightly shorter. Flowers hypocrateriform, 2 mm diam., densely rusty strigillose outside; receptacle deeply infundibuliform, 0.7 X 0.5 mm, slightly constricted below tepals, rusty tomentose inside. Te- pals chartaceous, ovate, 1 X 0.6 mm, both whorls equal, spreading at anthesis, the tips incurved, the inner surface densely rusty tomentellose. Stamens of whorls I and II stipitate, 0.8 mm tall, the anthers ovate, 0.5 X 0.3 mm, glabrous, the apex apiculate, the connectives prolonged between the 2 locelli, these obliquely hemispherical, introrse-latrorse, the filaments clavate, narrower than anthers, densely grey- tomentose; whorl III stamens broadly stipitate, 0.6 mm tall, the anthers ovate, 0.3 X 0.3 mm, erect, locelli 2, extrorse-latrorse, the filaments as broad as anthers, wider towards base, ligulate, densely grey-tomentose inside, the basal glands absent; whorl IV wanting; pistillode wanting. Pistillate inflorescence with indument, color, and branching as in staminate plants, the flowers slightly deeper; stamens sterile, smaller; tepals erect; ovary glabrous, ovoid; style stout, weakly distinguished from ovary; stigma discoid, 0.5 mm diam. Fruits borne on claviform pedicels of up to 1.3 x 0.3 cm; cupules hemispherical, to 1 x 2 cm, gla-

brous inside and outside, the margins entire; drupes ellipsoid, to 3 X 2 cm.

Distribution (Fig. 27) and ecology. Medium- sized trees from lowland forests of western Amazonia at ca. 130-215 m. Flowering material known from June, August, and September, and fruits from January.

Representative specimens examined. COLOMBIA. VAUPES: Rio Paca, Wacaricuara and vic., 215 m, 1-3 Jun 1953 (fl d), Schultes & Cabrera 19540 (US).

PERU. LORETO: Maynas, Iquitos, Estaci6n Experimen- tal IIAP, Allpahuayo, 130 m, 24 Aug 1988 (fl d), van der Werf et al. 10242 (F, G, MO); Rfo Nanay, Mishana, 140 m, 10 Jan 1983 (fr), Gentry et al. 39301 (G, MO), Aug 1988 (fl Y), van der Werif et al. 10187 (G, MO); Requena, Sa- puena, Arboreto Jenaro Herrera, 130 m, 18 Sep 1980 (fl 6), Castillo 34 (F, MO), 170 m, 15 Sep 1987 (fl 6), Vdsquez & Jaramillo 9593 (G, MO, US).

Local names. Peru: anis moena, limon moena.

Endlicheria citriodora has ovate-apiculate whorl I and II anthers typical of the E. canescens species group (species 37-45), but the very short reddish hairs that cover its striate branchlets and petioles are without equal therein. Further, the truncate leaf base is immediately diagnostic whenever evident. As noted in the protologue (van der Werff, 1991), the tendency for ultimate cymes to collapse into pseudo-umbels is unusual. Elsewhere in Endlicheria, pseudo-umbels appear in relatively pauciflorous inflorescences in E. acuminata and E. xerampela, species further differing from E. citriodora by truncate to emarginate anther apices.

Collectors of this species consistently report a strong lemonlike scent and glaucous lower leaf surfaces, but the odor disappears in herbarium specimens and the glaucous cast is seldom retained (e.g., Castillo 34).

40. Endlicheria rubra Chanderbali, sp. nov. Type. Peru. San Martin: Naranjillo, along rd. between Rioja and Pedro Ruiz, 950 m, 22 Mar 1998 (fl d), van der Werif et al. 15436 (holotype: MO; isotypes: F, G, GH, HBG, MO, NY, QRS, U, US).

Fig. 28

Ex affinitate Endlicheriae canescentis et specierum affinium foliis obtusissimis distinguenda.

Trees to 25 m. Branchlets relatively stout, midway along flush 3-5 mm diam., angular, densely tomentose, the surface barely visible, reddish brown, the hairs short, to 0.3 mm, straight to crisped, erect, dark red; terminal buds slender, 2 x 0.6 mm, densely pu-

SYSTENMATIC TREATMENT 83

~~~~~~~~~. I~~~~~~~~~~~~~~~~~~~~.

V

AR~~~~

4 - ---------- 4, Irc

W, 7

T.~~~~~~~~~~~~~ lv ~

FI.7.Dsriuin.fEdl.e.a....oaE..r Evnticaan recl

bescent, the hairs as on branchlets, ascending. Leaves alternate, evenly spaced along current flush; petioles slender, to 1 x 0.2 cm, semi-terete to canaliculate, the indument as on branchlets; laminae stiff chartaceous, plane, obovate, 8-15 X 2-5 cm, the base acute, the apex blunt to rounded, the margins minutely recurved throughout; upper surface olive to reddish brown, the primary to fourth-order veins raised, their prominence decreasing with rank, but the midrib sunken towards petiole; lower surface sparsely tomentose, the hairs erect, straight to curved, dark red, denser on main veins, all vein orders raised, their prominence decreasing with rank; secondary veins 5-8 per side, ? evenly spaced, slightly more distant around midlamina, ascending at 50 60? (more obtuse around midlamina), arcuate, distal pairs loop-connected; tertiaries oblique to midrib, between secondaries straight to forked. Staminate inflorescences evenly spaced along current flush in the axils of foliage leaves, to 12 cm long with 10 lateral branches, branch orders 3-4, the highest order dichasial, lax, the flowers distant, the

axes densely red tomentellose; bracts and bracteoles caducous by anthesis, lanceolate, the indument as on axes; pedicels gradually increasing in diameter apically, to 1.5 mm long, those supporting secondary flowers slightly shorter. Flowers hypocrateriform, to 2 mm diam., densely rusty tomentellose outside; receptacle cyathiform, 1 X 1 mm, slightly constricted below tepals, densely greyish tomentose inside. Te- pals fleshy, broadly ovate, 0.8 X 0.6 mm (the inner whorl slightly broader), spreading, the inner surface densely greyish rusty tomentellose, the hairs erect, crisped, the margins and apex inside rusty papillose. Stamens of whorls I and II broadly stipitate, ca. 0.6 mm tall, the anthers ovate, 0.4 X 0.3 mm (slightly narrower in whorl II), glabrous, the apex apiculate, the connectives prolonged between the 2 locelli, these obliquely hemispherical, introrse-latrorse, the filaments fleshy, slightly narrower than anthers, densely grey-tomentellose; whorl Ill stamens sessile, 0.6 mm tall, the anthers narrowly ovate to lanceolate, 0.3 X 0.15 mm, erect, locelli 2, narrow, slitlike, extrorse-

84 FLORA NEOTROPICA

FIG. 28. Endlicheria rubra (van der Weriffet al. 15436). A. Habit. B. Leaf base below. C. Flower. D. Flower I.s. E. Whorl I stamen seen from within. F. Whorl III stamen seen from without.


latrorse, the filaments apically as broad as anthers, widening towards base, fleshy, densely grey- tomentellose, the basal glands sessile, laminar, apiculate; whorl IV wanting; pistillode wanting. Pistillate inflorescence with indument and color as in staminate plants, but shorter and with fewer lateral branches, the flowers slightly deeper; stamens sterile, smaller; ovary glabrous, ellipsoid; style stout, weakly distinguished from ovary, 0.2 mm long; stigma tri-lobed, 0.5 mm diam. Fruits unknown.

Distribution (Fig. 27) and ecology. Medium- sized trees known from poorly drained forests in western Amazonia and eastern Andean slopes at ca. 800 m. Flowering at least in March, May and July.

Additional specimens examined. BRAZIL. ACRE:

Cruzeiro do Sul, Rio Jurua and Rio Moa, vic. of Porangaba, 17 May 1971 (fl juv), Maas et al. P13058 (HBG, MG, NY, U). AMAZONAS: Rfo Ituxi, vic. of B6ca do Curuquete, 12 Jul 1971 (fl 2), Prance et al. 14150 (HBG, K, MO, NY, US).

Among species with dense reddish vestiture En- dlicheria rubra is immediately recognized by its obovate leaves with blunt apices and recurved margins. The type material is made even more conspicuous by a glaucous cast on the leaves below. This feature does not appear in the Brazilian specimens, possibly lost through preservation in alcohol, but otherwise they provide good vegetative matches and the narrowly ovate anthers of the type also appear in the pistillate flowers of Prance et al. 14150.

The unusual leaf morphology of Endlicheria rubra, complete with glaucous cast below, appears in a fruiting collection from Amazonian Colombia, Garcfa-Barriga 13944 (NY, US), but four locelli arranged in a shallow arc in whorl I and II staminodes assign this specimen to Rhodostemonodaphne. Garcia-Barriga 13944, apparently representing an undescribed species, is so similar to Endlicheria rubra that my initial impression was that it provided the fruits of this new species. These are immature, but as they already show deeply hemispherical cupules, this suggests deep floral receptacles in pistillate flowers, as found in E. rubra. Given these similarities, it would be surprising if the two were not sister species.

41. Endlicheria vinotincta C. K. Allen, Mem. New York Bot. Gard. 10(5): 63. 1964. Type. Venezuela. Amazonas: Rio Yatua, Cerro Neblina, Cafion Grande, below Cumbre Camp, 1650 m, 26 Nov 1957 (fl ), Maguire et al. 42244 (holotype: NY).

Scandent shrubs to 3 m. Branchlets relatively slender, midway along flush 2-4 mm diam., distally weakly angular, soon terete, densely tomentose, the surface concealed by the indument cover, the hairs short, to 0.5 mm, crisped, erect, dark red; terminal buds plump, 5 X 3 mm, reddish tomentose-pannose. Leaves alternate, widely and evenly spaced along current flush; petioles robust, to 1.5 X 0.4 cm, semi- terete, the indument as on branchlets; laminae coriaceous, plane to subbullate, ovate, 8-2 X 3-8 cm, the base rounded, briefly decurrent, the apex acute, acuminate for up to 4.5 cm, the margins minutely recurved throughout; upper surface olive-brown, waxy, areolate, the primary to fourth-order veins raised, their prominence decreasing with rank; lower surface sparsely tomentose, the hairs as on branchlets but pale brown, denser on main veins, all vein orders raised, their prominence decreasing with rank; secondary veins 3-4 per side, ? evenly spaced, slightly more distant around midlamina, patent, diverging at 70-85? (more acutely towards apex), arcuate, distal pairs loop-connected; tertiaries laxly reticulating between secondaries. Staminate inflorescences evenly spaced along current flush in the axils of foliage leaves, to 15 cm long with 9 lateral branches, branch orders 3-4, the highest order dichasial, lax, the flowers distant, the axes reddish tomentose; bracts and bracteoles caducous by anthesis, lanceolate, the indument as on axes; pedicels terete, to 3 mm long, those supporting secondary flowers slightly shorter. Flowers hypocrateriform, 5 mm diam., densely reddish tomentose throughout; receptacle infundibuliform, 0.6 X 1 mm. Tepals chartaceous, elliptic to obovate, 1.6 X 1.3 mm (the inner whorl slightly narrower), spreading to recurved at anthesis. Stamens of whorls I and II broadly stipitate, 0.8 mm tall, the anthers ovate, 0.3 X 0.6 mm, densely reddish tomentose outside, the apex glabrous, truncate to emarginate, the connectives level with or slightly reduced between the 2 locelli, these obliquely hemispherical, introrse, the filaments stout, clavate, gradually tapering below anthers, rusty tomentose, provided with a pair of minute sessile basal glands; whorl III stamens sessile, 0.8 mm tall, the anthers ovate, 0.3 X 0.6 mm, erect, locelli 2, extrorse-latrorse, the filaments as broad as anthers, ligulate, fleshy, rusty tomentose, the basal glands minute sessile; whorl IV wanting; pistillode wanting. Pistillate inflorescence with indument, color, and branching as in staminate plants, the flowers similar in size and shape; stamens sterile, smaller; ovary glabrous, ovoid; style stout, indistinct from ovary; stigma discoid, 0.5 mm diam. Fruits borne on claviform pedicels of 0.4 X 0.3 cm; cupules infundibuliform, to 0.7 X 0.4 cm, glabrous

86 FLORA NEOTROPICA

inside and outside, the margins strongly lobed, tepals persisting; drupes ellipsoid, to 1.3 X 0.8 cm.

Distribution (Fig. 27) and ecology. Scandent shrubs from sandstone formations of the Guiana Highlands at ca. 350-1650 m. Flowering specimens have been collected in April, October, and November, and the only fruiting specimen in April.

Additional specimens examined. COLOMBIA. GUAINIA: Maimachi, Serranfa del Naqu6n. Cerro Minas, alrededores del Helipuerto-15 y camino hasta la cima del Cerro, 900 m, 7 Apr 1993 (fl 5), Madrinidn & Barbosa 936 (COL), (fr), Madrinidn & Barbosa 948 (MO), (fl 5), Mad- rinan & Barbosa 953 (MO).

VENEZUELA. AMAZONAS: Atabapo, Cerro Duida, cuesta de arenisca de granda fino al pie del Cerro Duida, 360 m, 10 Nov 1982 (fl 5), Gudnchez 2154 (MO); meseta de arenisca grande ubicada al Sur del Rio Matakuni, 1050 m, 19 Nov 1991 (fl 5), Huber 13261 (MO); Cerro de la Neblina, slope N of Rio Mawarinuma, above "Pto. Chimo," 500 m, 25 Apr 1984 (fl 5), Stein & Gentry 1660 (F, MO, US); Rio Negro, Cerro Aracamuni, summit, 1550 m, 16 Oct 1987 (fl i), Liesner & Delascio 22008 (MO), 1415 m, 16- 18 Oct 1987 (fl d), Delascio & Liesner 13508 (MO, VEN), 600 m, 21 Oct 1987 (fl 5), Liesner & Carnevali 22292 (MO), 1400 m, 25 Oct 1987 (fl 5), Liesner & Carnevali 22414 (MO).

In Endlicheria, E. vinotincta alone inhabits the summits and upper slopes of the sandstone mountains (tepuis) of the Guiana highlands. In its scandent habit, the rich dark red vestiture that covers its branchlets and inflorescences, including inner floral surfaces, and stamens all provided with basal glands, this is a very distinctive species of Endlicheria. Yet all these characters are matched by the sympatric Rhodoste- monodaphne celiana. Leaves of Endlicheria vinotincta have prominent tertiaries above and pale brown indument while in Rhodostemonodaphne celiana the tertiaries are immersed above and a dark red indument covers the lower leaf surface, but flowers of the two differ mainly in locelli number. Another sympatric species of Rhodostemonodaphne, R. steyermarkiana, is vegetatively similar, but appears to be an erect shrub (Madrinian, 1996b), and, like R. celiana, has dark red indument on the leaves below.

The faint glaucous cast persisting on lower leaf surfaces in Gudnchez 2154 and Huber 13261 hints at a useful field character for this species.

42. Endlicheria oreocola Chanderbali, sp. nov. Type. Peru. Cajamarca: San Ignacio, Romerillo, base of Cordillera de Romerillo, 1570 m, 15 Nov

1997 (fl d), Campos & Nuniez 4669 (holotype: MO; isotypes: F, HBG, NY). Fig. 29

Nova species Endlicheriae vinotinctae proxima, fo- liorum texturae et colori similis, sed floribus indumento gris- eis differt.

Treelets or shrubs to 8 m. Branchlets stout, midway along flush 4-6 mm diam., angular, densely greenish yellow tomentose, the surface concealed by the indument cover, the hairs relatively short, to 0.2 mm, straight to crooked, erect; terminal buds plump, 5 X 3 mm, densely pubescent, the hairs as on branchlets, ascending. Leaves alternate, widely and evenly spaced along current flush; petioles slender, to 2 X

0.2 cm, semi-terete, distally furrowed, the indument as on branchlets; laminae coriaceous, subbullate to plane, ovate, 9-15 X 3-7 cm, the base acute, briefly decurrent, the apex acute, briefly acuminate for up to 0.5 cm, the margins minutely recurved throughout; upper surface yellowish green to reddish brown, waxy, areolate, all vein orders prominulous; lower surface sparsely tomentose, the hairs as on branchlets, scattered on lamina, denser on main veins, all vein orders raised, their prominence decreasing with rank; secondary veins 4-6 per side, ? evenly spaced, slightly more distant around midlamina, ascending at 50-60O (more obtuse around midlamina), arcuate, distal pairs loop-connected; tertiaries roughly horizontal, between secondaries once-forked to straight. Stami- nate inflorescences evenly spaced along current flush in the axils of foliage leaves, to 3 cm long with 3 lateral branches, branch orders 2-3, the highest order dichasial, the flowers distant, the axes densely yellowish green tomentose; bracts and bracteoles caducous by anthesis, ovate, the indument as on axes; pedicels terete, to 1 mm long, those supporting secondary flowers slightly shorter. Flowers infundibuliform, 3 mm diam., densely greyish green tomentose outside; receptacle broadly infundibuliform, 0.6 X 1 mm, densely grey-tomentose inside. Tepals fleshy, ovate, 1 X 0.6 mm, ascending at anthesis, the inner surface sparsely grey-tomentose. Stamens of whorls I and II broadly stipitate, 0.4 mm tall, the anthers ovate, 0.3 X 0.4 mm, glabrous, the apex apiculate, the connectives prolonged between the 2 locelli, these obliquely hemispherical, introrse-latrorse, the filaments stout, clavate, slightly narrower than anthers, densely grey- tomentose; whorl III stamens stout, sessile, 0.5 m tall, the anthers oblong, 0.3 X 0.2 mm, erect, locelli 2, extrorse-latrorse, the filaments broader than anthers, columnar, densely grey-tomentose, the basal glands sessile, globose; whorl IV wanting; pistillode wanting. Pistillate inflorescence with indument, color, and

FIG. 29. Endlicheria oreocola (Campos & Nuntez 4889). A. Habit. B. Leaf base below. C. Flower. D. Flower 1.s. E. Whorl I stamen seen from within. F. Whorl III stamen seen from without.

88 FLORA NEOTROPICA

branching as in staminate plants, the flowers similar in size and shape; stamens sterile, smaller; ovary glabrous, ovoid; style slender, distinct from ovary; stigma discoid, 0.3 mm diam. Fruits borne on claviform pedicels of up to 1 X 0.4 cm; cupules shallowly infundibuliform, to 0.3 X 1 cm, glabrous inside and outside, the margins entire; drupes obovoid, to 2 X 1.5 cm.

Distribution (Fig. 27) and ecology. Small to medium-sized trees or shrubs of pre-montane cloud forest vegetation from ca. 1500-2500 m on the eastern Andean slopes of Peru and Ecuador. Flowers known from November and January, fruits from Jan- uary and April.

Additional specimens examined. ECUADOR. ZAMORA-CHINCHIPE: Zamora, Parque Nacional Podocar- pus, carretera Loja-Zamora, Estaci6n Cientifica San Fran- cisco, 2250 m, Jan 1995 (fl 6), Palacios & Tirado 13425 (QCNE, MO), (fl Y), Palacios & Tirado 13439 (QCNE, MO), (fl Y, fr), Palacios & Tirado 13459 (QCNE, MO), 2100 m, 20 Apr 2000 (fr), Neill et al. 12616 (QCNE, MO).

The dense greenish yellow tomentose indument found on branchlets of the type material of Endli- cheria oreocola is unmatched in the genus. A glaucous cast appearing on the underside of younger leaves offers another striking feature but appears to be lost with age, and older leaves attach to reddish brown tomentose branches. Older leaves of E. oreocola resemble those of E. vinotincta in their coriaceous texture, ovate shape, and areolate upper surfaces. Yet, this is certainly not its closest relative. Flowers of E. oreocola are greyish green rather than reddish tomentose outside, and although the shape and size of the stamens in the two species are much alike, in E. oreocola only whorl III stamens are provided with basal glands. Still, as no other species of Endlicheria is remotely similar it seems best to com- pare E. oreocola with E. vinotincta, if only to em- phasize its peculiarities.

43. Endlicheria szyszylowiczii Mez, Jahrb. Konigl. Bot. Gart. Berlin. 5: 121. 1889. Type. Peru. Ca- jamarca: Tambillo, without date (fl d), Jelski 165 (lectotype, designated by Kostermans, 1937: B- n.v.; isolectotypes: L, MO, S, US).

Trees to 15 m. Branchlets stout, midway along flush 4-6 mm diam., distally weakly angular, soon terete, densely rusty to reddish tomentose, the surface concealed by the indument cover, the hairs relatively long, to 0.6, straight to crisped, erect to ascending; terminal buds plump, 2 x 2 mm, densely pubescent,

the hairs rusty to red, ascending. Leaves alternate, widely and evenly spaced along current flush; petioles slender, to 2.5 X 0.3 cm, semi-terete, the indument as on branchlets; laminae coriaceous to chartaceous, plane to subbullate, ovate to elliptic, 9-25 X 4-10 cm, the base acute to rounded, briefly decurrent, the apex acute to obtuse, acuminate for up to 2.5 cm, the margins minutely recurved throughout; upper surface dull greyish green to olive-brown, minutely punctulate, the primary to fourth-order veins raised, their prominence decreasing with rank, or tertiary and higher-order veins immersed; lower surface densely pilose, the hairs soft, erect, up to 1 mm long, rust brown, slightly shorter and denser on main veins, all vein orders raised, their prominence decreasing with rank; secondary veins 4-6 per side, ? evenly spaced, slightly more distant around midlamina, ascending at 50-60? (more obtuse around midlamina), arcuate, distal pairs loop-connected; tertiaries roughly horizontal, between secondaries straight, or forked. Staminate inflorescences evenly spaced along current flush in the axils of foliage leaves or cataphylls, to 25 cm long with 14 lateral branches, branch orders 3-4, the highest order dichasial, lax, the flowers distant, the axes densely rusty tomentose; bracts and bracteoles caducous by anthesis, narrow ovate to lanceolate, the indument as on axes; pedicels terete, to 2 mm long, those supporting secondary flowers slightly shorter. Flowers depressed-globose to urceolate, 1.5 to 3 mm diam., sparsely rusty to grey-strigose outside; receptacle deeply cyathiform, 1 X 2, densely rusty red velutinous inside. Tepals chartaceous, broadly ovate, 1 X 1.5 mm (the inner whorl slightly narrower), incurved to erect at anthesis, the inner surface minutely papillose. Stamens of whorls I and II broadly stipitate, 0.6-1 mm tall, the anthers ovate, 0.3-0.5 X 0.3-0.6 mm, glabrous, the apex apiculate, the connectives prolonged between the 2 locelli, these obliquely hemispherical, introrse-latrorse, the filaments ligulate, almost as broad as anthers, densely grey-tomentose; whorl III stamens sessile, 0.6-1 mm tall, the anthers depressed-oblong, 0.2-0.3 X 0.3-0.5 mm, erect, locelli 2, extrorse-latrorse, the filaments as broad as anthers, wider towards base, the indument as in outer whorls, the basal glands sessile, globose; whorl IV wanting; pistillode filiform. Pistillate inflorescence with indument, color, and branching as in staminate plants, the flowers similar in size and shape; stamens sterile, smaller; ovary glabrous, ovoid; style slender, distinct from ovary; stigma minutely tri-lobed, ca. 0.2 mm diam., emerging beyond incurved tepals. Fruits borne on short claviform pedicels of up to 5 X 5 mm; cupules hemispherical, to 0.5 x 1 cm, glabrous inside


and outside, the margins undulate, tepal bases persisting; drupes ellipsoid to obovoid, to 1.5 X 1 cm.

Distribution (Fig. 30) and ecology. Medium- sized trees ranging throughout the Amazonian lowlands and reaching the surrounding highlands of the Guiana shield to the north, the Brazilian shield to the south, and lower montane Andes to the west at 200- 2200 m. Flowering material collected from February through November, fruits in July, and September through November.

Representative specimens examined. COLOMBIA. ANTIoQJIA: San Antonio de Prado, 2200 m, Nov 1989 (fl d ), de Escobar et al. 8851 (HUA).

VENEZUELA. AMAZONAS: Rfo Negro, Canyon Grande de La Neblina, 5 Feb 1984 (fld ), Funk 6102 (MO).

GUYANA. EssEQuIBo: Wassarai Mtns., 14 km S of Kassikaitu R., 1135 m, 10 Sep 1999 (fl Y), Clarke et al. 8397 (MO).

PERU. AMAzONAS: Mendoza, 1600 m, 16 Aug 1963 (fl d ), Woytkowski 8323 (GH, MO). CAJAMARCA: Tambillo, s.d. (fl juv), Jelski 196 (L). LORETO: Maynas, Bosque Na-

cional Alexander von Humboldt, 300 m, 25 Nov 1977 (fl ), Froehner 67 (HBG, MO). MADRE DE DIos: Tambopata,

Lago Sandoval, 13 km NE of Puerto Maldonado, 200 m, 25 Jul 1989 (fr), Nuiiez 11195 (HBG, MO); Parque Nacional "Bahuaja-Sonere," Ex Santuario Nacional Pampas del Heath, 10 Jul 1997 (fl S), Diaz & Pereira 9020 (MO). PAsco: Oxapampa, 1800 m, 4 Mar 1986 (fl d), van der Werif et al. 8340 (HBG, MO, NY). SAN MARTiN: Rioja, Pardo Miguel, Venceremos, Caserfo El Afluente, Bosque de Protecci6n de Alto Mayo, 1440-1520 m, 14 Nov 1996 (fr), Sdnchez & Dillon 8687 (MO).

BRAZIL. AcRE: Brasileia, Reserva Extrativista Chico Mendes, Seringal Porongaba, 15 Apr 1992 (fl d), Saraiva & de Lima 1523 (MO); Cruzeiro do Sul, BR 364, km 42, ramal 4 do Projeto Santa Luzia (INCRA), Sep 1985 (fr), Rosas et al. 216 (MO, NY). AMAPA: Rio Jari, Sep 1968 (fl d6), Silva 968 (HBG, NY). AMAZONAS: Novo Aripuana, BR 230, Rod. Transamaz8nica 400 km de Humaita, Projeto INCRA-Rfo Juma, vicinal dos Gadchos, 4 May 1985 (fl 6), Cid Ferreira et aL 6033 (F, GH, INPA, K, MO, NY, US); Presidente Figueiredo, Rio Uatuma, UHE Balbina, 7 Jul 1986 (fl Y), Thomas et aL 5370 (F, INPA, K, MO, NY, US). MATO GRosso: Sinop, 25 Sep 1985 (fr), Thomas et al.

It

A~~~~~~~~~~~~~~~~~~~~~~~~~

4,~~~~~~~~~~~~~~~~~

FIG~~~~~~~~~~~~~~~~~~~~~~ 30DsrbtoFfEdihrasyzlwcuEdoicaadElrseo

90 FLORA NEOTROPICA

4059 (HBG, INPA, MO, NY). PARA: Belem, 27 Jul 1944 (fl Y), Silva 309 (NY, UC, US); Rio Mapua, 18 Jul 1950 (fl 6), Black et al. 50-9809 (INPA, NY, US). ROND6NIA: Gua- jara Mirim, 175 m, 9 Apr 1987 (fl Y), Nee 34692 (F, INPA, GH, K, MO, NY, US).

BOLIVIA. LA PAZ: Prov. Franz Tamayo, 1600 m, 23 May 1992 (fl Y), Perry 999 (MO); Prov. Larecaja, Copa- cabana, 10 km S of Mapiri, 850-950 m, 8 Oct-15 Nov 1939 (fr), Krukoff11103 (A, F, G, K, MO, NY, S, U, US). SANTA

CRUZ: Prov. Caballero, Parque Nacional Ambor6, 31 Mar 1996 (fl 6), Jardim et al. 2576 (NY).

Local names. Peru: moena, roble fusi. Brazil: louro de f61ha peluda, louro vermelho.

Endlicheria szyszylowiczii is readily distinguished from other species with dense reddish tomentose branchlets by its depressed-globose to campanulate flowers with tepals incurved to erect at anthesis. In vegetatively similar species, ascending tepals may yield infundibuliform flowers, but horizontally spreading tepals predominate. At first glance the higher-elevation material of E. szyszylowiczii (e.g., van der Werif et al. 8340) may be mistaken for E. duotincta because the pale lower leaf surface contrasting with the erect reddish hairs produce an aspect similar to that bestowed by the mixed indument of the latter. Despite a wide geographic and altitudinal range, the only noteworthy internal variation appears as slightly reduced floral size and paler pubescence in extra-Andean collections.

44. Endlicheria duotincta Chanderbali, sp. nov. Type. Peru. Cajamarca: San Ignacio, Huarango, Nuevo Mundo, road between Caserio Gosen and Nuevo Progress, 1350-1400 m, 23 Jul 1997 (fl d), Rodrigues & Reyes 1807 (holotype: MO; isotypes: F, G, HBG, NY, US, U). Fig. 31

Foliis subtus indumento cum Endlicheriae williamsii optime congruens, sed floribus campaniformis et foliis alternis differt.

Trees to 15 m. Branchlets stout, midway along flush 4-6 mm diam., angular, densely rusty red tomentose, the surface concealed by the indument cover, the hairs moderately long, to 0.5 mm, crooked to crisped, erect; terminal buds plump, 2 X 2 mm, densely pubescent, the hairs as on branchlets, appressed. Leaves alternate, widely and evenly spaced along current flush; petioles robust, to 3.5 X 0.4 cm, striate, the indument as on branchlets; laminae stiff chartaceous, plane to subbullate, ovate to elliptic, 10- 30 X 5-13 cm, the base acute to rounded, the apex acute, acuminate for up to 1.5 cm, the margins flat

throughout; upper surface light green to olive-brown, minutely punctulate, the midrib prominulous, the higher-order venation immersed; lower surface obscured by a short creamish tomentose indument interspersed with straight erect reddish hairs of 0.7 mm, all vein orders raised, their prominence decreasing with rank; secondary veins 6-8 per side, ? evenly spaced, slightly more distant around midlamina, ascending at 50-60o (more acutely towards apex), arcuate, distal pairs loop-connected; tertiaries roughly horizontal, between secondaries straight or forked. Staminate inflorescences evenly spaced along leafless flushes in the axils of cataphylls, to 20 cm long with 14 lateral branches, branch orders 2-3, the highest order dichasial, lax, the flowers distant, the axes densely rusty tomentose; bracts and bracteoles persistent at anthesis, lanceolate, the hairs as on axes, appressed; pedicels terete, to 2 mm long, those supporting secondary flowers slightly shorter. Flowers campanulate, to 2.5 mm diam., densely pubescent outside, the hairs rusty red, appressed to ascending, reduced to a basal triangular patch in the inner whorl of tepals; receptacle cyathiform, 2 X 1.5 mm, rusty pilose inside. Tepals chartaceous, ovate, 1.3 X 0.6 mm (the inner whorl slightly narrower), ascending at anthesis, the inner surface sparsely papillose towards the margins and apex, otherwise glabrous. Stamens of whorls I and II broadly stipitate, 1 mm tall, the anthers ovate, 0.5 X 0.4 mm, glabrous, the apex apiculate, the connectives prolonged between the 2 locelli, these suborbicular, introrse, the filaments laminar, hardly narrower than anthers, densely grey-tomentose; whorl III stamens broadly stipitate, 1 mm tall, the anthers ovate, 0.5 X 0.4 mm, erect, locelli 2, extrorse-latrorse, the filaments almost as broad as anthers, laminar, the indument as in outer whorls, the basal glands sessile, globose; whorl IV wanting; pistillode filiform. Pistil- late inflorescence with indument and color as in staminate plants, but shorter and with fewer lateral branches, the flowers slightly deeper; stamens sterile, smaller, sessile, the filaments as broad as anthers; ovary glabrous, ellipsoid; style slender, distinct from ovary, 1 mm long; stigma minutely tri-lobed, 0.2 mm diam. Fruits borne on claviform pedicels of up to 5 mm long; cupules hemispherical, to 1 X 1.5 cm, glabrous outside, densely rusty strigose inside, the margins sharply lobed, tepal bases persisting; drupes ellipsoid, to 2 x 1 cm.

Distribution (Fig. 30) and ecology. Medium- sized trees of pre-montane forests from ca. 1300- 1900 m on the eastern Andean slopes of Ecuador and Peru. Flowering material collected in February, July,

4~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~-

ttI t~ ~ ~ ~ ~ ~ ~ ~~~V

FIG. 31. Endlicheria duotincta (Rodr(gues & Reyes 1807). A. Habit. B. Leaf base below. C. Flower. D. Flower with facing tepal turned down to reveal androecium. E. Flower 1.s. F Whorl I stamen seen from within. G. Whorl HI stamen seen from without.

92 FLORA NEOTROPICA

August, and September, and fruiting material in June and August.

Additional specimens examined. ECUADOR. ZAMORA-CHINCHIPE: Quebrada Las Pavas, 1800 m, 21 Aug 1975 (fl Y), Samaniego & Vivar 90 (QAME, US); Que- brada de Los Monos, cerca de Sabanilla, 1600 m, 4 Sep 1975 (fl Y, fr juv), Little et al. 221 (COL, QAME, US).

PERU. SAN MARTiN: Rioja, Pedro Ruiz-Moyobamba rd., km 390, Venceremos, 1800-1900 m, 29-31 Jul 1993 (fl d), Smith 4491 (MO), 1790 m, 9-10 Aug 1983 (fr), Smith & Vdsquez 4775 (MO); Rioja-Pomacochas rd., below Ven- ceremos, ca. 20 km NW of Rioja near Restaurant El Amigo, 1600 m, 8 Feb 1984 (fl ?), Gentry & Smith 45169 (MO); vic. of Puente Serranoyacu, 1590-1840 m, Aug 1983 (fr), Luna 112 (MO).

Local name. Ecuador: laurel blanco.

Endlicheria duotincta is the second species in the genus in which tomentose indument conceals the lower leaf surface. This remarkable vestiture is also found in E. williamsii, but may have originated independently there since its flowers are comparable to those of E. anomala, while the campanulate flowers of E. duotincta closely resemble those of E. ruforamula of the E. sericea species group. Although a position in the latter is suggested on floral grounds, the rusty tomentose branchlets of E. duotincta would be anomalous there, and a place among those with similar indument, only less dense on leaves below-i.e., the E. canescens species group-is preferred until its affinities become more clear.

45. Endlicheria lorastemon Chanderbali, sp. nov. Type. Ecuador. Zamora-Chinchipe: Nangaritza, Rio Nangaritza, Pachicutza, 1000-1200 m, 6 Dec 1990 (fl J), Palacios & Neill 6577 (holotype: MO; isotypes: F, GH, HBG, NY, QCNE, U, US).

Fig. 32

Species ramulis sub-sericeis foliis subtus pilosis a con- generis diversa.

Trees, 5-40 m. Branchlets slender, midway along flush 2-3 mm diam., distally weakly angular, soon terete, densely pubescent, the surface barely visible to concealed by the indument cover, the hairs relatively short, to 0.3 mm, straight, appressed to ascending, yellowish to rust brown; terminal buds slender, 3 X 1 mm, densely rusty sericeous. Leaves altemate, widely and evenly spaced along current flush; petioles slender, to 2.5 X 0.15 cm, semi-terete, the indument as on branchlets; laminae chartaceous, plane, ovate, 10-25 X 4-9 cm, the base acute, briefly decurrent, the apex acute, acuminate for up to 4 cm, the margins

flat throughout; upper surface olive-brown, minutely punctulate, the primary to fourth-order veins raised, their prominence decreasing with rank; lower surface sparsely pubescent, the hairs as on branchlets, but erect, denser on main veins, all vein orders raised, their prominence decreasing with rank; secondary veins 3-5 per side, + evenly spaced, slightly more distant around midlamina, ascending at 50-60? (more obtuse towards apex), arcuate, distal pairs loop- connected; tertiaries roughly horizontal, between secondaries forked to straight. Staminate inflorescences evenly spaced along current flush in the axils of foliage leaves, to 10 cm long with 8 lateral branches, branch orders 2-3, the highest order dichasial, lax, the flowers distant, the axes densely pubescent, the hairs reddish to light brown, appressed to ascending; bracts and bracteoles caducous by anthesis, lanceolate, rusty sericeous; pedicels terete, to 2 mm long, those supporting secondary flowers slightly shorter. Flowers hypocrateriform, 3 mm diam., densely tawny to rusty pubescent outside, the hairs ascending; receptacle infundibuliform, 0.6 X 1 mm, silvery velutinous inside. Tepals chartaceous, ligulate, 1 X 0.5 mm (the inner whorl slightly narrower), ascending to spreading at anthesis, the inner surface sparsely silvery tomentose near base, otherwise glabrous, the margins minutely papillose. Stamens of whorls I and II broadly stipitate, 0.5 mm tall, the anthers depressed-ovate, 0.2 X 0.3 mm, glabrous, the apex broadly apiculate above the 2 locelli, these suborbicular, introrse, the filaments ligulate, equaling or slightly narrower than anthers, densely silvery-grey tomentellose; whorl III stamens columnar, 0.6 mm tall, the anthers oblong, 0.3 X 0.2 mm, erect, locelli 2, extrorse-latrorse, the filaments slightly broader than anthers, wider towards base, densely silvery-grey tomentellose, the basal glands stipitate, minute, globose; whorl IV wanting; pistillode filiform. Pistillate inflorescence with indument, color, and branching as in staminate plants, the flowers similar in size and shape; stamens sterile, smaller; ovary glabrous, ovoid; style stout, weakly distinguished from ovary; stigma minutely tri-lobed, 0.3 mm diam. Fruits borue on stout, claviform pedicels of up to 1 X 0.5 cm; cupules patelliform, to 0.3 X 1 cm, glabrous inside and outside, the margins sharply lobed, tepal bases persisting; drupes spheroid to obovoid, to 1 X 1 cm.

Distribution (Fig. 30) and ecology. Small to large trees from western Amazonia and adjacent lower montane eastern Andean slopes, at ca. 120-1200 m. Flowering specimens have been collected continu- ously from August through January and fruits from January to June.


3 cm ~~~~~~~~~~

FIG. 32. Endlicheria lorastemon (Palacios & Neill 6577). A. Habit. B. Leaf base below. C. Flower. D. Flower 1.s. E. Whorl I stamen seen from within. F. Whorl mI stamen seen from without.

94 FLORA NEOTROPICA

Additional specimens examined. COLOMBIA. AMA-

ZONAS: Rio Apaporis, Soratama, near mouth of Rio Kan- anari, 250 m, 14 Dec 1951 (fl 9), Schultes & Cabrera 14903 (COL, U); Rio Caqueta, Sep 1989 (fl juv), Urrego et al. 1012 (MO). CAQUETA: Araracuara, 22 Dec 1993 (fl 3), Vester & Roman 865 (COL, MO).

ECUADOR. NAPO: Aguarico, Reserva Etnica Huaor- ani, carretera y oleoducto de Maxus, km 108, 235 m, 18 Jan 1995 (fr), Aulestia & Omehuat 3236 (MO), km 72, 270 m, 23-31 Jan 1994 (fr), Dik & Andi 1001 (MO, QCNE-n.v.), km 75-76, entre el Rio Tivacuno y Rio Yasuni, 17-20 Feb 1994 (fr), Aulestia & Gonti 1755 (MO); Orellana, Parque Nactional Yasuni, 230 m, 9-13 Jan 1987 (fl d), Ceron & Coello 3257 (MO); Tena, Estaci6n Biol6gica Jatun Sacha, 450 m, 17 Aug 1993 (fl d), Palacios & Tipaz 11074 (MO). PASTAZA: Rio Chullana, ca. 15 km N of Puerto Sarayacu, 16 Oct 1974 (fl 9), Lugo 4188 (GB); Rio Curiacu, ca. 8 km W of Puerto Sarayacu, 19 Oct 1974 (fl 9), Lugo 4241 (GB); Rio Bobonaza, Caimito, between Puerto Sarayacu and Pa- cayacu, 25 Oct 1974 (fl 9), Lugo 4302 (GB), ca. 4 km E of Pacayacu, 31 Oct 1974 (fl 9), Lugo 4394 (GB), between Pacayacu and Canelos, 3 Nov 1974 (fl 9), Lugo 4450 (GB); Via Auca, 115 km al S de Coca, cerca del Rio Tigiiino, 320 m, 29 Apr 1989 (fr), Rubio 8 (MO). SANTIAGO-ZAMORA:

Cordillera Cutucd, 1600 m, 17 Nov-5 Dec 1944 (fl d), Camp E-1165 (NY). ZAMORA-CHINCHIPE: Nangaritza, Rio Nangaritza, Miazi, 1000 m, 9 Dec 1990 (fl 3), Neill & Pa- lacios 9618 (MO), (fl d), Palacios & Neill 6669 (MO), (fl d), Palacios & Neill 6688 (MO), 20 Oct 1991 (fl 3), Pa- lacios et al. 8492 (COL, MO), Pachicutza, 6 Dec 1990 (fl 3), Palacios & Neill 6572 (MO); Zamora, Parque Nacional

Podocarpus, Guarderia Rio Bombuscaro, sendero al Mira- dor, 1100 m, Jan 1995 (fl d), Palacios & Tirado 13310 (MO).

PERU. LORETO: Maynas, Iquitos, Puerto Almendras, UNAP, 122 m, 17 Jan 1993 (fl 3), Grdndez et al. 5512 (MO).

BRAZIL. ACRE: Cruzeiro do Sul, sub-base do Projeto RADAM/BRASIL, 8 Mar 1976 (fr), Ramos & Mota 350 (INPA); Mancio Lima, 12 Oct 1989 (fl 9), Cid Ferreira et al. 10031 (INPA, NY); Serro do Moa, Local Central, 30 Sep 1984 (fl 3), Cid Ferreira et al. 5094 (F, INPA, K, MO, NY, US); Rio Moa, 8 km above Cachoeira Grande, 27 Apr 1971 (fr), Prance et al. 12563 (HBG, MO, NY). AMAZONAS: Rio Solimoes, Tefe, entrada do Lago Tefe, Santa Missoes, 16 Oct 1982 (fl d), Amaral et al. 109 (INPA, MO); Parand de Tefe, 16 Oct 1982 (fl d), Cid Ferreira & Lima 3270 (INPA, K, MO, R); Fonte Boa, sub-base do Projeto RADAMI BRASIL, Carta SA-l9-XD-PT? 06, 4 Jun 1976 (fr), Ramos 450 (INPA).

Local names. Ecuador: ocatoe, yahuemomo (both are Huaorani names).

Endlicheria lorastemon typically has straplike whorl I and II stamens where filaments equal anthers

in width, but occasional stipitate stamens (e.g., Amaral et al. 109 and Cid Ferreira & Lima 3270) reduce the diagnostic value of an otherwise unique

feature. Still, no other species of Endlicheria combines erect hairs on the leaves below with a dense subsericeous cover of appressed to ascending hairs on the branchlets.

As for Endlicheria duotincta, affinities are unclear since the subsericeous branchlets of E. lorastemon recall the E. sericea species group while the claviform pedicels below shallow platelike cupules resemble those of E. anomala.

46. Endlicheria sericea Nees, Linnaea 8: 38. 1833. Goeppertia sericea (Nees) Nees, Syst. laur. 369. 1836. Type. Trinidad. Without locality and date (fl), Sieber 175 (holotype: B-n.v.; isotypes: BM- n.v., E, G, GH, L, LZ-n.v., MO, NY-n.v., P, W- n.v.).

Goeppertia sericea (Nees) Meisn. var. opaca Meisn. DC. Prodr. 15 (1): 174. 1864. Type. Dominica. Without locality and date (fl), Imray 457 (holotype: K-n.v.).

Endlicheria guadaloupensis Mez, Jahrb. Konigl. Bot. Gart. Berlin 5: 124. 1889. Type. Guadeloupe. With- out locality and date (fr), Duchassaing s.n. (holotype: B-n.v. isotype: W-n.v.).

Trees to 20 m. Branchlets relatively stout, midway along flush 3-5 mm diam., distally weakly angular, soon terete, silvery to golden sericeous, the surface concealed by the indument cover, the hairs short, to 0.2 mm, straight, appressed; terminal buds plump, 3 X 2.5 mm, sericeous. Leaves alternate, widely and evenly spaced along current flush; petioles slender to robust, to 2 X 0.3 cm, semi-terete, the indument as on branchlets; laminae coriaceous to chartaceous, plane, ovate, 10-20 X 2-10 cm, the base obtuse, briefly decurrent, the apex acute, acuminate for up to 2 cm, the margins minutely recurved throughout; upper surface light olive-green, waxy, the midrib flat, sunken towards petiole, drying dark, conspicuous against the lamina, the secondaries immersed; tertiaries prominulous; lower surface densely sericeous, the hairs as on branchlets, uniformly distributed, all vein orders raised, their prominence decreasing with rank; secondary veins 4-5 per side, ? evenly spaced, slightly more distant around midlamina, or basal pairs closer, subopposite, all ascending at 50-60? (more obtuse around midlamina), arcuate, distal pairs loop- connected; tertiaries roughly horizontal, between secondaries once-forked to straight. Staminate inflorescences evenly spaced along current flush in the axils of foliage leaves or cataphylls, to 20 cm long with 15 lateral branches, branch orders 3-4, the highest order dichasial, lax, the flowers distant, the axes silvery to golden sericeous; bracts and bracteoles ca-


ducous by anthesis, lanceolate, the indument as on axes; pedicels terete, to 2 mm long, those supporting secondary flowers slightly shorter. Flowers rotate, to 4.5 mm diam., densely silvery sericeous outside; receptacle shallowly cyathiform, 2 X 4 mm, silvery pilose inside. Tepals fleshy, ovate, 1.6 X 1 mm, spreading, the androecium exserted at anthesis, the inner surface densely grey-tomentose, the margins and apex inside rusty papillose. Stamens of whorls I and II broadly stipitate, 0.6 mm tall, the anthers ovate, 0.4 X 0.3 mm, glabrous, the apex apiculate or truncate to emarginate, the connectives prolonged between, broad above, or level with the 2 locelli, these suborbiculate, introrse-latrorse, the filaments laminar, slightly narrower than anthers, basally grey-pilose; whorl III stamens sessile, 0.7 mm tall, the anthers oblong, 0.4 X 0.3 mm, erect, locelli 2, extrorse- latrorse, the filaments as broad as anthers, columnar, grey-pilose, the basal glands sessile, globose; whorl IV wanting; pistillode wanting. Pistillate inflorescence with indument and color as in staminate plants, but shorter and with fewer lateral branches, the flowers slightly deeper; stamens sterile, smaller; ovary glabrous; style slender, distinct from ovary; stigma tri-lobed, 0.3 mm diam. Fruits borne on stout claviform pedicels of up to 2 X 0.5 cm; cupules hemispherical, to 1 X 1.3 cm, glabrous outside, sericeous inside, the margins entire to undulate; drupes ellipsoid, to 2 x 1.3 cm.

Distribution (Fig. 33) and ecology. Medium- sized trees from lower montane forests in the Lesser Antilles, Trinidad, and northern Andes in Venezuela at ca. 300-1350 m. Flowering throughout the year, fruits collected from February through April and in October.

Representative specimens examined. DOMINICA. Laudat, Jul 1881 (fl 6), Eggers 403 (G, GH, HBG, P); Castle Bruce Trail, 400 m, 25 Feb 1946 (fl ?, fr juv), Beard 628 (A, MO, NY, U).

GUADELOUPE. Basse-Terre near Duclos, Petit Bourg, 200 m, 30 Mar 1956 (fr), Smith 10356 (K, NY); Pidgeon, 21 Aug 1973 (fl ?), Sastre & Sastre 2046 (NY, P).

MARTINIQUE. Trace des Jesuites NW of Fond St.- Denis, entrance E of Morne Rouge-Fond St. Denis rd., 450 m, 22 Jul 1987 (fl Y), Daly 5293 (MO, NY); Mamne Rose, 600 m, 28 Sep 1984 (fl 2), Rollet 1672 (A).

ST. LUCIA. Savanne Edmund district, SE of Piton Troumass6e, 1800-2000 ft, 20 Nov 1960 (fi 2, fr), Procter 21585 (A); Zeno Top Soufriere, 3 Oct 1986 (fr), Pierre & Slane 357 (A).

ST. VINCENT. Silver Spoon, above Three Rivers, 1-7 Apr 1960 (fl 2), Howard 11145 (GH, NY); Mt. St. Andrews summit, Feb 1972 (fr) Howard & Howard 18005 (A, NY).

VENEZUELA. MIRANDA: Cerros del Bachiller, 200-

690 m, 21, 27, & 28 Mar 1978 (fr), Steyermark & Davidse 116957 (G, MO); Los Guayabitos (arriba de Baruta), 1350 m, Mar 1958 (fr), Aristeguieta 3012 (MO, NY). TACHIRA:

Cerro Las Minas, 17 km SE of Santa Ana, 1150-1250 m, Nov 1979 (fr), Steyermark et al. 119895 (G, MO, VEN).

Local names. Dominica: bois marble, laurier bord de mer, laurier p6te. St. Lucia: laurier gris, laurier gwa gwen, laurier marbre, sweetwood. St. Vincent: sweetwood.

Endlicheria sericea is distinguished from all other species where silvery to golden sericeous indument obscures the lower leaf surface by its rotate flowers with fleshy spreading tepals that bear a dense greyish tomentose cover on the inner surfaces. Stout claviform pedicels below hemispherical cupules and a sunken midrib in the lower lamina are also diagnostic.

This species has been collected most frequently in the Lesser Antilles and the type is from Trinidad, but presence on the South American mainland is suggested by three collections from the Venezuelan An- des (viz., Steyermark & Davidse 116957, Steyermark et al. 119895, and Aristeguieta 3012). All are fruiting specimens that are vegetatively indistinguishable from insular material and, moreover, show truncate anther apices in whorl I and II staminodes. Except for E. griseo-sericea with elliptic to obovate leaves and much larger cupules, and E. bracteolata with triplinerved leaves, all other species with densely sericeous lower leaf surfaces have definitely apiculate anthers.

Endlicheria guadaloupensis is based on fruiting material from Guadeloupe (Duchassaing s.n.) that could not be located. From Mez's (1889) descriptions and key to species it is apparent that he distinguished this species from E. sericea by ellipsoid rather than ovoid fruits and nonpersistence of tepals on cupule margins. Without floral correlates the specific value of these differences is questionable, and Kostermans (1937), who examined the type of E. guadaloupensis, considered it conspecific with that of E. sericea. His synonymy is maintained not only because the differences in fruit shape are rather slight, but also because tepal persistence appears to be temporary in E. sericea.

The type material of E. sericea appears to be a mixed collection since the E, GH, and P duplicates bear staminate inflorescences, the MO duplicate pistillate inflorescences, and the G duplicate diseased inflorescences.

47. Endlicheria bracteolata (Meisn.) C. K. Allen, Mem. New York Bot. Gard. 10(5): 64. 1964. Goeppertia sericea (Nees) Nees var. bracteolata

96 FLORA NEOTROPICA

- . . e-- . . . . .-- * * - -- -.-

.,,~ ~ ~ ~ ~~.

FIG 33 Distribution of Endlicheria serice and E bracteota. I

FIG. 33. Distribution of Endlicheria sericea arld E. bracteolata.~ .7'

Meisn. DC. Prodr. 15(1): 174. Type. Venezuela. Amazonas: San Carlos, Rio Negro, 1853-4 (fl c, juv), Spruce 3092 (syntypes: B-n.v., BM-n.v., E, G, GH, L, NY, OXF-n.v., P-n.v., U, W).

Trees to 15 m. Branchlets slender, midway along flush 3-4 mm diam., distally weakly angular, soon terete, silvery to golden sericeous, the surface concealed by the indument cover, the hairs rather short, to 0.15 mm, straight, closely appressed to the surface; terminal buds plump, 3 X 3 mm, sericeous. Leaves alternate, widely and evenly spaced along current flush; petioles slender, to 1.5 X 0.3 cm, semi-terete, the indument as on branchlets; laminae chartaceous to coriaceous, plane, ovate, 10-20 X 5-8 cm, the base obtuse to rounded, briefly decurrent, the apex acute, acuminate for up to 2 cm, the margins minutely recurved throughout; upper surface light green to olive- brown, the midrib immersed, secondary and tertiary veins prominulous; lower surface densely silvery to golden sericeous, the hairs uniformly distributed, all vein orders raised, their prominence decreasing with

rank; secondary veins 2-4 per side, the lowermost pair(s) subopposite shortly above the leaf base, ascending at 5060O, arcuate, distal pairs loop- connected; tertiaries roughly horizontal, between secondaries straight or forked. Staminate inflorescences all evenly spaced along leafless flushes in the axils of cataphylls, or the distal ones in the axes of foliage leaves, to 10 cm long with 10 lateral branches, branch orders 3-4, the highest order dichasial, lax, the flowers distant, the axes golden sericeous; bracts and bracteoles caducous by anthesis, lanceolate, sericeous; pedicels terete, to 1 mm long, those supporting secondary flowers slightly shorter. Flowers infundibuliform, 2 mm diam., golden sericeous outside; receptacle infundibuliform, 1 X 1 mm, densely silvery pilose inside. Tepals chartaceous, ovate to ligulate, 1 x 0.5 mm (the inner whorl slightly broader), ascending, surrounding androecium at anthesis, the inner surface moderately silvery tomentose, the margins minutely papillose. Stamens of whorls I and II stipitate, 0.8 mm tall, the anthers depressed-elliptic to


ovate, 0.5 X 0.5 mm, glabrous, the apex rounded to truncate, the connectives broad above the 2 locelli, these suborbiculate, introrse-latrorse, the filaments laminar, narrower than anthers, densely silvery pilose; whorl III stamens sessile, 0.8 mm tall, the anthers oblong, 0.4 x 0.3 mm, erect, locelli 2, extrorse- latrorse, the filaments broader than anthers, laminar, densely silvery pilose, the basal glands sessile, globose, apiculate; whorl IV staminodial; pistillode filiform. Pistillate inflorescence with indument and color as in staminate plants, but shorter and with fewer lateral branches, the flowers campanulate; stamens sterile, smaller; ovary glabrous; style slender, distinct from ovary; stigma tri-lobed, 0.3 mm diam. Fruits borne on narrowly claviform pedicels of up to 0.5 X

0.2 cm; cupules hemispherical, to 0.5 X 0.7 cm, glabrous outside, sericeous inside, the margins entire; drupes ellipsoid, to 1 X 0.5 cm.

Distribution (Fig. 33) and ecology. Medium- sized trees from seasonally inundated lowland and lower montane forests (50-1000 m) throughout northern South America. Flowers and fruits available year round.

Representative specimens examined. COLOMBIA. AMAZONAS: Rio Apaporis, Jirijirimo, 250 m, 25-26 Nov 1951 (fl d), Garcfa-Barriga 13703 (COL, NY, US), 27 Nov 1951 (fl d), Schultes & Cabrera 14606 (GH, U). ANTIO-

QUiA: Urrao, Parque Nacional Natural "Las Orquideas," 1300-1500 m, 3 Apr 1992 (fl 6), Cdrdenas & Alvarez 3267 (MO), Sector Calles, margen derecha del Rfo Calles y de la quebrada "El Guaguo," 12 Feb 1989 (fl Y, fr juv), Cogollo et al. 3928 (MO), camino de Venados arriba hacia Calles, 30 Jul 1988 (fl ?), Cogollo et at. 3638 (MO). VALLE: Barco, Rio Cajambre, Apr 1944 (fr), Cuatrecasas 17216 (F, US); Buenaventura, Caserfo "San Isidro," CONIF, 100-150 m, 23 Mar 1992 (fr), Cogollo et al. 5127 (INPA, MO), 25 Mar 1992 (fl juv), Cogollo et al. 5129 (MO).

VENEZUELA. AMAZONAS: Atabapo, Rio Asisa, 100 m, Oct 1989 (fl d), Delgado 835 (MO); Atures, Puerto Aya- cucho, Rio Cataniapo, 100-110 m, 11 Nov 1980 (fl d), Gua'nchez 388 (HBG, MO, VEN). BOLiVAR: Cedeno, 20 km E of Turiba, 6-11 Dec 1970 (fl d), Marcano-Berti 2581 (G, MO); Rio Las Ahallas, 14 Mar 1985 (fr), Liesner 18665

(MO, NY). GUYANA. ESSEQUIBO: Cuyuni-Mazaruni Region, Pa-

ruima Falls to Paruima Mission, Kamarang-Wenamu Trail, 600 m, 8 Aug 1951 (fl Y), Maguire & Fanshawe 32463 (GH, MO, NY, U, US); Kamarang R., Utschi River mouth, 500 m, 20 Oct 1960 (fl Y), Tillett & Tillett 45705 (F, K, NY, US); Kurupung R., Makreba Falls, 24 Feb 1939 (fr), Pinkus 263 (F, G, GH, MO, NY, U, US).

SURINAME. Tafelberg, 610 m, 3 Sep 1944 (fl 6), Ma- guire 24712 (A, K, MO, NY, U, US); Marowijne R., Wane Ck., Feb 1918 (fr), Gonggrijp 3692 (U).

FRENCH GUIANA. Fleuve Approuague, Grand Can-

ori and Crique Sapokay, 26 Jul 1968 (fl Y), Oldeman T-35 (MO, P), entre le Saut et la crique Couata, 31 Jul 1968 (fl Y), Oldeman T-55 (NY, P, U).

PERU. LORETO: Maynas, Rfo Nanay, Puerto Almen- dras, 122 m, 4 Jun 1986 (fl d), Vdsquez & Jaramillo 7644 (MO); Upper Rio Mazan, near Base Araguana, 8 Jul 1976 (fl d), Gentry & Revilla 16539 (F, HBG, MO).

BRAZIL. AMAPA: Colonia do Torrao, 28 Aug 1962 (fl d ), Pires & Cavalcante 52649 (NY, US); Rio Oiapoque, Rio Ingarari, 15 Sep 1960 (fl Y), Irwin et al. 48290 (F, NY, U). AMAZONAS: Jauarete, Vaupes, Rio Negro, 22 Oct 1945 (fl d), Froes 21243 (F, MO, NY); Manaus, Cachoeira baixa do Taruma, 12 Sep 1966 (fl Y), Prance et al. 2269 (F, GH, HBG, INPA, K, MG, NY, R). PARA: Porto Trombetas, 12 Aug 1986 (fl d), Soares 189 (INPA).

Local names. Colombia: jigua. Venezuela: dimukuima (Ye'kwana), laurel blanco, laurel plateado, laurel rebalsero. French Guiana: cedre. Brazil: louro cedinha.

Endlicheria bracteolata is easily distinguished from other species with sericeous indument by its triplinerved leaves. Occasional production of a second pair of basal secondaries shortly above the leaf base results in quintuplinervation similar to that frequent in E. sericea. Indeed, Meissner (1864) considered this species a mere variety of E. sericea, but rotate flowers in the latter contrasting infundibuliform flowers with only ascending tepals in E. bracteolata supports Al- len's (1964) elevation to specific status. The inflorescence bracteoles to which the specific epithet alludes are present on the type material and other specimens with immature inflorescences, but fall by the onset of anthesis. Specimens from Valle and Antioquia in Co- lombia have slightly darker indument and tend to have larger leaves but otherwise cannot be distinguished from material east of the Andes.

48. Endlicheria tschudyana (Lasser) Kosterm., Bol. Tecn. Inst. Agron. N. No. 28, Addenda post p. 75. 1953. Aniba tschudyana Lasser, Bol. Soc. Venez. Ci. Nat. 11(72): 181. 1948. Type. Venezuela. Ar- agua: Parque Nacional de Rancho Grande, 14 Feb 1946 (fl Y), Lasser 2036 (holotype: VEN; isotypes: MO, VEN).

Trees to 12 m. Branchlets stout, midway along flush 5-7 mm diam., angular, silvery to pale yellow sericeous, the surface concealed by the indument cover, the hairs short, to 0.1 mm, straight, appressed; terminal buds plump, 4 X 3 mm, sericeous. Leaves alternate, widely and evenly spaced along current flush; petioles robust, to 4 X 0.4 cm, semi-terete, the indument as on branchlets; laminae coriaceous, or

98 FLORA NEOTROPICA

chartaceous, plane, ovate to elliptic, 13-35 X 8-15 cm, the base obtuse to subcordate, briefly decurrent, the apex acute, acuminate for up to 1.5 cm, the margins minutely recurved throughout; upper surface greyish green to olive-brown, the primary to fourth- order veins raised, their prominence decreasing with rank; lower surface densely sericeous, the hairs as on branchlets, uniformly distributed, all vein orders raised, their prominence decreasing with rank; secondary veins 5-8 per side, + evenly spaced, slightly more distant around midlamina, ascending at 50-60' (more acutely towards apex), arcuate, the lowermost pairs sometimes patent, diverging at 70-85?, and abruptly ascending after midcourse, distal pairs loop- connected; tertiaries roughly horizontal, between secondaries once-forked to straight. Staminate inflorescences evenly spaced along current flush in the axils of foliage leaves or cataphylls, to 12 cm long with 8 lateral branches, branch orders 3-4, the highest order dichasial, lax, the flowers distant, the axes rusty to grey-sericeous; bracts and bracteoles caducous by anthesis, ovate, the indument as on axes; pedicels terete, to 3 mm long, those supporting secondary flowers slightly shorter. Flowers cyathiform to infundibuliform, to 3 mm diam., silvery-grey to rusty sericeous outside; receptacle cyathiform or infundibuliform, 1.5 X 2.5 mm, grey-pilose inside. Tepals chartaceous, elliptic, 0.8 X 0.5, erect to ascending, surrounding androecium at anthesis, the outer surface densely silvery or rusty sericeous, the indument reduced to a basal triangular patch in the inner whorl, the inner surface glabrous or sparsely grey-strigillose. Stamens of whorls I and II narrowly stipitate, 0.5 mm tall, the anthers ovate, 0.3 X 0.3 mm, glabrous, the apex apiculate, the connectives prolonged between the 2 locelli, these suborbicular, introrse-latrorse, the filaments laminar, narrower than anthers, grey-pilose or tomentose; whorl III stamens columnar, 0.6 mm tall, the anthers depressed-oblong, 0.2 X 0.3 mm, erect, locelli 2, extrorse-latrorse, the filaments as broad as anthers, columnar, the indument as in outer whorls, the basal glands sessile, globose; whorl IV wanting; pistillode fusiform. Pistillate inflorescence with indument and color as in staminate plants, but shorter and with fewer lateral branches, the flowers slightly deeper; stamens sterile, smaller; ovary glabrous; style stout, weakly distinguished from ovary; stigma discoid, 0.3 mm diam. Fruits borue on stout terete pedicels of up to 0.8 X 0.3 cm; cupules shallowly hemispherical, to 0.8 X 1.5 cm, glabrous outside and inside, the margins undulate to lobed, tepal bases persisting; drupes ellipsoid, to 1.7 X 1.2 cm.

Distribution (Fig. 34) and ecology. Small trees of lower montane and cloud forests from NW South America to Panama at ca. 300-1600 m. Flowering and fruiting specimens collected throughout the year.

Additional specimens examined. PANAMA. BOCAS DEL TORO: Along rd. to Chiriquf Grande, 10 road-mi from continental divide, 300 m, 9 Feb 1987 (fr), McPherson 10444 (MO). COCLE: Near sawmill 16.7 km N of turnoff to Coclesito from Llano Grande, 700 ft, 7 Mar 1978 (fl 5), Hammel 1858 (MO). DARIEN: Alturas de Nique, 850-1100 m, 24 Aug 1987 (fl 5), McPherson 11582 (HBG, MO); Trocha de Rancho Frio, 900 m, 23 Sep 1989 (fl Y), Aranda et al. 978 (MO). PANAMA: Cerro Jefe, 650 m, 27 Aug 1986 (fl Y), McPherson 10000 (MO), 750-850 m, 19 Jul 1987 (fl Y), McPherson 11310 (MO), 800 m, 13 Jan 1988 (fl 9 juv, fr), McPherson 11936 (MO), 900 m, 27 Dec 1986 (fl), Val- despino et al. 278 (MO).

COLOMBIA. ANTiOQuIA: San Francisco, carretera a Aquitania, 500-900 m, 8 Apr 1990 (fl 5, juv), Ca'rdenas et al. 2645 (MO); San Luis, carretera hacia Aquitania, a 12 km de la Autopista Medellin-Bogota, 850 m, 24 Nov 1988 (fl 5), Cogollo et al. 3735 (MO).

VENEZUELA. ARAGUA: Parque Nacional Henri Pit- tier, 4 Aug 1991 (fl 5, juv), Cardozo et al. 1822 (MO), Fila de Paraiso, above Portachuelo, 1500 m, 5 Jul 1963 (fr), Stey- ermark 91524 (K, NY), Periquito Trail, 18 Jun 1962 (fr), Allen 20 (NY), Mar 1959 (fr), Aristeguieta 3845 (NY), Pico Periquito, 12 Dec 1979 (fr), Manara s.n. VEN 172708 (VEN), 1250-1600 m, 4 Sep 1960 (fl 5, juv), Steyermark & Agostini 7 (F, NY, US), Rancho Grande, Aug 1963 (fr), Aristeguieta 5107 (VEN), 15 Jun 1962 (fr), Lasser & Allen 13 (NY), 29 May 1975 (fr), Ferrari & Benitez de Rojas 1456 (F). CARABOBO: Aut6nomo Mora, cuenca hidrografica del Rio Moran, 700-1100 m, 3-5 May 1991 (fr), Diaz & Ninio 289 (MO); Rio San Gian, arriba de La Toma, 750-850 m, 30 Mar 1966 (fl 5, juv), Steyermark & Steyermark 95347 (G, NY, VEN). YARACUY: Cerro La Chapa, 1200-1400 m, 9-10 Nov 1967 (fl 9), Steyermark et al. 100298 (HBG, NY), 21 Oct 1982 (fl 5, juv), Davidse et al. 20840 (MO); San Felipe, Serrania Santa Maria-Cerro La Chapa, 1150-1250 m, 12 Aug 1992 (fr), Meier & Walter-Weisbeck 2586 (MO).

ECUADOR. NAPO: Estaci6n Biol6gica Jatun Sacha, Rio Napo, 8 km al E de Mishualli, 450 m, 4 Sep 1987 (fr), Cer6n et al. 2090 (MO), 3-4 Jun 1988 (fr), Neill et al. 8475 (MO); Parque Nacional Yasuni, Rio Tiputini, 200-300 m, 5 Oct 1996 (fl 5), Romoleroux et al. 2567 (MO). SUCUMBiOS:

Lago Agrio, Reserva Faunistica Cuyabeno, Tarapoa- Tipishca, cruce del Rio Cuyabeno, 230 m, 14 Nov 1991 (fl 5), Palacios et al. 8938 (MO, NY, US), 8940 (MO).

PERU. HUANUCO: Pucallpa, Sira Mtns., 800 m, 9 Feb 1988 (fl 5), Morawetz & Wallnofer 13-9288 (MO).

Local names. Colombia: laurel bobo, laurel bongo.

The Venezuelan type material of Endlicheria tschudyana was taken from a pistillate plant with pat-


..

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W' 'At~ .. ~ - -

'2 ~ -J ~ ~ ~ ~ ~.

FIG 3 Disributon o Endlchera tscudyaa andE grseo srice

ent, almost perpendicular, secondary veins in the lower lamina of cordiform leaves. Such leaves are found again in Aristeguieta 3845, but all other specimens from the type locality have ovate to elliptic leaves with more ascending secondaries. Since flowers from the type locality are either pistillate or immature, the concept of E. tschudyana adopted here is largely based on plants from elsewhere. Although this recourse increases its geographical range considerably, it is not without justification. In the type material, the floral receptacle is infundibuliform and incurved tepals close around the stigma. These flowers are unusual but are matched by pistillate plants from Pan- ama (e.g., McPherson 10000). It therefore becomes plausible that vegetatively indistinguishable staminate plants from Panama, Hammel 1858 and Mc- Pherson 11582, provide mature staminate flowers of E. tschudyana. However, the two offer slightly different manifestations of this, with Hammel 1858 showing cyathiform, and McPherson 11582 infundibuli-

form, flowers. As the most advanced stage of floral development found in the type locality, Steyermark & Agostini 7 shows an infundibuliform receptacle, it is likely that McPherson 11582 provides the more typical representation. Furthermore, with infundibuliform staminate flowers also in Cogollo et al. 3735 from Colombia, the disjunction between Venezuela and Panama is considerably reduced.

Yet, it is possible that flower shape is unstable in E. tschudyana. An apparent correlation between cyathiform flowers and robust branchlets with coriaceous leaves can be disregarded since three collections from Ecuador, Palacios et al. 8938 and 8940, and Romoleroux et al. 2567, combine cyathiform flowers with slender branchlets and relatively thin leaves. Furthermore, branchlets and leaves also vary in the type locality. With both floral shapes accommodated in E. tschudyana, a staminate plant identical to Hammel 1858 in both vegetative and floral terns, Morawetz & Wallnofer 13-9288, extends the range of

100 FLORA NEOTROPICA

the species to Peru. With these specimens all included, E. tschudyana accommodates all of the E. sericea species group material with silvery sericeous branchlets, pinnate venation, campanulate or infundibuliform flowers, with erect to ascending, internally glabrous tepals, ovate anthers with apiculate apices, and shallowly hemispherical cupules.

49. Endlicheria griseo-sericea Chanderbali, sp. nov. Type. Ecuador. Napo: Archidona, Volcain Sumaco, km 50 along road between Hollin and Loreto, Guagua Sumaco Community, 1300 m, 5 Feb 1992 (fl 6), Rubio & Alvarado 2395 (holotype: MO; isotypes: F, G, GH, HBG, K, MO, NY, P, QCNE, U, US). Fig. 35

Ex affinitate Endlicheriae sericeae et specierum affinium floribus urceolatis et antheris obovatis distinguenda.

Trees to 30 m. Branchlets stout, midway along flush 5-7 mm diam., angular, densely grey to yellowish sericeous, the surface concealed by the indument cover, the hairs short, to 0.2 mm, straight, appressed to ascending; terminal buds plump, 0.4 X 0.4 mm, densely sericeous. Leaves alternate, widely and evenly spaced along current flush; petioles robust, to 4 X 0.3 cm, semi-terete, the indument as on branchlets; laminae chartaceous, or membranaceous, plane, elliptic to obovate, 15-30 X 5-13 cm, the base obtuse, or acute, briefly decurrent, the apex acute, or obtuse, acuminate for up to 2 cm, the margins minutely recurved throughout; upper surface dark olive- brown, the primary to fourth-order veins raised, their prominence decreasing with rank; lower surface densely grey to yellowish sericeous, the hairs as on branchlets, sparser on main veins, all vein orders raised, their prominence decreasing with rank; secondary veins 7-9 per side, ? evenly spaced, slightly more distant around midlamina, ascending at 50-60? (more obtuse towards apex), arcuate, distal pairs loop- connected; tertiaries roughly horizontal, between secondaries straight or forked. Staminate inflorescences evenly spaced along leafless flushes in the axils of cataphylls, to 18 cm long with 12 lateral branches, branch orders 3-4, the highest order dichasial, the flowers loosely crowded, the axes densely pubescent, the indument as on branchlets; bracts and bracteoles persistent at anthesis, ovate, sericeous; pedicels terete, to 3 mm long, those supporting secondary flowers slightly shorter. Flowers urceolate,.to 2.5 mm diam., densely yellowish tomentellose; receptacle cyathiform, 2 x 3 mm, constricted below tepals, densely

grey-sericeous inside. Tepals chartaceous, narrowly ovate to triangular, 1.5 X 0.5 mm (the inner whorl slightly narrower), erect to spreading at anthesis, the outer surface densely pubescent, the hairs as on receptacle, reduced to a basal triangular patch in the inner whorl, the inner surface grey-tomentose, the margins and apex inside minutely papillose. Stamens of whorls I and II stipitate, 0.6 mm tall, the anthers obovate, 0.3 X 0.3 mm, glabrous, the apex truncate, the connectives broad above the 2 locelli, these suborbicular, introrse, the filaments laminar, narrower than anthers, glabrous; whorl III stamens sessile, 0.6 mm tall, the anthers oblong to obovate, 0.4 X 0.3 mm, erect, locelli 2, extrorse-latrorse, the filaments as broad as anthers, columnar, glabrous, the basal glands sessile, globose; whorl IV wanting; pistillode wanting. Pistillate inflorescence with indument and color as in staminate plants, but shorter and with fewer lateral branches, the flowers similar in size and shape; stamens sterile, smaller, glabrous or the filaments grey-tomentose; ovary glabrous; style slender, distinct from ovary; stigma minutely tri-lobed, 0.3 mm diam. Fruits bome on stotit claviform pedicels of up to 1.5 X 0.4 mm; cupules hemispherical, to 1.5 X 2.5 cm, glabrous outside, sericeous inside, the margins entire; drupes ellipsoid, to 4.5 X 2.5 cm.

Distribution (Fig. 34) and ecology. Medium- sized to large trees of lower montane Andean forests at ca. 800-1500 m. Flowering specimens collected in August, November, and February, fruits in April, May, June, October, November, and December.

Additional specimens examined. ECUADOR. Mo- RONA-SANTIAGO: Cordillera de Cutucu, Mendez-Morona Rd., 800 m, 4 Feb 1989 (fl Y), van der Werif & Palacios 10385 (MO). NAPO: Archidona, faldas al sur del Volcan Sumaco, carretera Hollin-Loreto, km 45, 1100 m, 10 Nov 1989 (fl Y), Palacios 4747 (G, K, MO, NY), km 50, Gua- gua Sumaco, 1000 m, 29 Apr-2 May 1989 (fr), Cer6n & Hurtado 6642 (MO), 1100 m, I May 1993 (fr), Neill et al. 8953 (G, MO, NY), km 31, Comuna Challua Yacu, 1200 m, 4 Oct 1989 (fr), Palacios & Iguago 4541 (MO); carretera Hollin-Loreto-Coca, km 40, 1200 m, 11 Dec 1987 (fr), Neill et al. 8094 (HBG, K, MO, NY, US), km 25, 1230 m, 10-19 Nov 1988 (fr), Hurtado & Alvarado 963 (MO); Or- ellana, Parque Nacional Yasunf, carretera y Oleoducto de Maxus en construcci6n, km 53-54, 250 m(?), 13-16 Sep 1993 (fr), Dik 405 (MO). PASTAZA: Pastaza, via Puyo- Macas, Pitirishca, 800 m, Aug 1993 (fl 6, juv), Palacios 11001 (F, MO).

PERU. SAN MARTiN: Rioja, km 399 of carretera Mar- ginal, trail to Quebrada Venceremos and Rio Serranoyacu, 67 km E of Pomacochas, 8 km W of bridge over Rio Ser- ranoyacu, 1400-1500 m, 13 Jun 1985 (fr), Knapp & Alcorn 7778 (MO).

I0~~~~~~~~~

FIG. 35. Endlicheria gniseo-sericea (Rubio & Alvarado 2395). A. Habit. B. Flower with facing tepal turned down to show androecium. C. Flower I.s. D. Whorl I stamen seen from within. E. Whorl HI stamen seen from without.


Local name. Ecuador: sacha palta (Quichua).

Endlicheria griseo-sericea is a remarkable new species in the E. sericea species group, differing from all other members by its urceolate flowers with the receptacle strongly constricted below spreading tepals, and obovate anthers with broadly truncate to retuse apices in whorl I and II stamens. The characteristic dark olive-brown color assumed by the upper leaf surface in the dry state is a useful diagnostic character, and in plants with a shiny greyish sericeous vestiture the contrast is especially striking.

Co-occurring with Endlicheria griseo-sericea in Ecuador but reaching further north to Colombia is a vegetatively similar entity with flowers of similar size and shape, but in which whorls I and II anthers are three-locellate and whorl III anthers are four-locellate. I have refrained from describing this material, prefer- ring instead to annotate it as cf. E. griseo-sericea, because in the only staminate collections, Acevedo et al. 1370 (F, MO, US), Henao 9 (COL), and van der Wertf & Palacios 9246 (HBG, MO, NY), the anthers are devoid of pollen and the unusual androecium may be a consequence of disease or hybridization. Several pistillate plants are known-Boom & Beardsley 8449 (MO, NY), Campos & Campos 3092 (MO) Cogollo et al. 2531, 6556, 7061, & 7062 (MO), Henao 21 & 281 (COL), Jorge & Torres 1270 (COL), Neill et al. 12614 & 12615 (MO), Pipoly et al. 17315 & 17453 (MO) Vdsquez & Rojas 21920 (MO)-and whether in flower or fruit, locelli number is as in staminate plants, but I cannot rule out the possibility that pollen donors were two-locellate.

50. Endlicheria ruforamula Chanderbali, sp. nov. Type. Peru. San Martin: Rioja, along road between Rioja and Pedro Ruiz, 850-1050 m, 26 Mar 1998 (fl d), van der Werif et al. 15736 (holotype: MO; isotypes: AMAZ, F, G, GH, HBG, K, MO, NY, P, QCNE, QRS, U, US). Fig. 36

Ex affinitate Endlicheriae sericeae et specierum affinium floribus campanulatis et ramulis tomentellis distinguenda.

Trees to 40 m. Branchlets stout, midway along flush 5-6 mm diam., angular, densely tomentellose, the surface concealed by the indument cover, the hairs short, to 0.15 mm, straight, ascending, reddish brown; terminal buds plump, 5 X 4 mm, densely pubescent, the hairs as on branchlets. Leaves alternate, widely and evenly spaced along current flush; petioles robust, to 5 X 0.4 cm, semi-terete, striate, the indument as on branchlets; laminae coriaceous, plane, ovate to ob-

ovate, 15-30 X 5-15 cm, the base obtuse to acute, briefly decurrent, the apex acute to obtuse, acuminate for up to 2 cm, the margins minutely recurved throughout; upper surface greyish green to olive- brown, waxy, the primary to fourth-order veins raised, their prominence decreasing with rank; lower surface densely pubescent, the hairs appressed, concealing surface, rusty red to golden yellow, reduced on main veins, all vein orders raised, their prominence decreasing with rank; secondary veins 5-7 per side, ? evenly spaced, slightly more distant around midlamina, ascending at 50-60o (more obtuse around midlamina), arcuate, the lowermost pair sometimes asymmetric (one or both merging with margin at base), distal pairs loop-connected; tertiaries roughly horizontal, between secondaries forked. Staminate inflorescences distally clustered in the axils of cataphylls, or evenly spaced along current flush in the axils of foliage leaves or cataphylls, to 25 cm long with 30 lateral branches, branch orders 3-4, the highest order dichasial, lax, the flowers distant, the axes rusty tomentellose; bracts and bracteoles persistent or caducous by anthesis, triangular, the indument as on axes; pedicels terete, to 4 mm long, those supporting secondary flowers slightly shorter. Flowers campanulate, to 3 mm diam., rusty to greyish tomentellose outside; receptacle cyathiform, 2 X 2 mm, rusty pilose inside. Tepals chartaceous, ovate, 1 X 0.7 mm (the inner whorl slightly narrower), erect to ascending at anthesis, the outer surface rusty to greyish tomentellose-sericeous, this reduced to a basal triangular patch in the inner whorl, the inner surface glabrous. Stamens of whorls I and II stipitate, 1 mm tall, the anthers ovate, 0.5 X 0.5 mm, glabrous, the apex broadly apiculate, the connectives bluntly prolonged between the 2 locelli, these suborbiculate, introrse- latrorse, the filaments laminar, narrower than anthers, glabrous or basally grey-tomentose; whorl III stamens sessile, ca. 1 mm tall, the anthers depressed-oblong, 0.3 X 0.5 mm, erect, locelli 2, extrorse-latrorse, the filaments as broad as anthers, ligulate, glabrous or sparsely grey-tomentose near base, the basal glands sessile, globose; whorl IV wanting; pistillode well- developed or wanting. Pistillate inflorescence with indument and color as in staminate plants, but shorter and with fewer lateral branches, the flowers slightly deeper; stamens sterile, smaller or equaling those of staminate flowers; ovary glabrous; style slender, distinct from ovary; stigma weakly tri-lobed, 0.2 mm diam. Fruits borue on stout terete pedicels of up to 1 x 0.4 cm; cupules hemispherical, to 1.5 x 2.5 cm, glabrous outside, rusty sericeous inside, the margins entire; drupes ellipsoid, to 4.5 X 2.5 cm.

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FIG. 36. Endlicheria ruforamula (A-G, van der Werff et al 15736; H, Vdsquez et al 11954). A. Habit. B. Leaf base below. C. Flower. D. Flower with facing tepal turned down to reveal androecium. E. Flower l.s. F. Whorl I stamen seen from within. G. Whorl III stamen seen from without. H. Fruits.


Distribution (Fig. 37) and ecology. Medium- sized to large trees of flooded and nonflooded Ama- zonian lowlands, eastern lower montane slopes of the Andes, and Pacific coast of Colombia and Ecuador at ca. 100-1200 m. Flowers and fruits available throughout the year.

Additional specimens examined. COLOMBIA. AMA- ZONAS: Leticia, Tarapacd, Parque Nacional Natural Ama- cayacu, Rio Cotuhe, 100 m, 19 Jun 1991 (fl d), Rudas et al. 2116 (MO), Cabafia Lorena (Inderena) y Caino Lorena, 100 m, 21 Jun 1991 (fl d), Rudas et aL 2262 (NPA, MO), 100 m, 21 Jun 1991 (fl d), Rudas et al. 2288 (MO), Cabania Pamat6; Canio Pamate, 100 m, 27 Jun 1991 (fl d), Rudas et al. 2588 (MO). CHOC6: Entre Curunde y San Jose de Pal- mar, 500 m, 30 Aug 1976 (fl 6), Forero et al. 2379 (COL, MO). VALLE: Cordillera Occidental, Rio Anchicaya, 230- 260 m, 13 Oct 1943 (fr), Cuatrecasas 15292 (F, U, US); Buenaventura, Caserfo "San Isidro," Reserva Forestal de CONIF, 30-100 m, 23 Mar 1992 (fr), Cogollo et al. 5128 (GH, MO).

ECUADOR. ESMERALDAS: San Lorenzo, Parroquia Ri- caurte, Centro Pambilar, 500 m, 21 Jan 1993 (fl d), Aulestia

& Aulestia 975 (MO, NY); Quininde, Fundaci6n Paraiso de Papagayos, Centro de Rescate de Aves y Mamiferos, 6-8 Jul 1996 (fr), Clark et al. 2784 (MO). NAPo: Aguarico, Reserva Etnica Huaorani, carretera y oleoducto de Maxus, km 92-96, 250 m, 20 Mar 1994 (fl 9, fr juv), Aulestia & Gonti 1986 (MO); Estaci6n Experimental INIAP-Napo, Payamino, Reserva Floristica "El Chuncho," 250 m, 5 Apr 1986 (fr), Jaramillo 8372 (MO), 250 m, 10-11 Sep 1986 (fr), Palacios & Neill 1289 (HBG, MO, NY); Estaci6n Cien- tffica Yasuni, 200-300 m, 15 Oct 1998 (fl 9), Romoleroux et al. 3502 (MO); La Joya de los Sachas, Pompeya, carretera de Maxus, km 1-5, 220 m, 23-29 Nov 1992 (fr), Grijalva et al. 243 (MO); Aiiangu, Parque Nacional Yasunf, 260-350 m, Apr 1986 (fr), SEF 8740 (NY); Tena, Estaci6n Biol6gica Jatun Sacha, 450 m, 8 Nov 1987 (fl 6), Cero$n 2600 (HBG, K, MO, NY), 450 m, 17-24 Feb 1988 (fr), Cer6n 3704 (MO), 400 m, 27 Nov 1992 (fr), Zuleta 42 & 50 (MO); Via Payamino-Loreto, 250 m, 13 Sep 1986 (fl d, juv), Zaruma 695 (MO). PASTAZA: Pastaza, 300 m, 21-28 Feb 1990 (fl cI), Zak & Espinoza 4835 (MO, NY), Curaray, 290 m, 13- 30 Nov 1990 (fr), Espinoza & Coba 588 (MO); Arajuno, 700 m, 3-14 Sep 1998 (fl c3), Freire & Santi 3294 (MO).

PERU. HuANuco: Pachitea, Dantas, trochas de estudio

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UNA La Molina, km 42, 22 Jan 1987 (fr), Diaz & Baldeon 2304 (MO, NY). JUNiN: Satipo, Gran Pajonal, E of Che- quitavo on trial to Kotampaz, 1200 m, Apr 1984 (fl Y), Smith 6764 (MO). LORETO: Alto Amazonas, Rio Huallaga, Shucushuyacu, 250 m, 13 Sep 1981 (fr), Vdsquez & Jar- amillo 2441 (F, MO, NY); Maynas, Caserfo Gamitana, Re- serva del Rfo Mazan, 116 m, 21 Jun 1990 (fr), Grdndez et al. 1611 (MO), (fl ?), Grdndez et al. 1612 (MO); Iquitos, Estaci6n Experimental IIAP, Allpahuayo, 130 m, 24 Aug 1988 (fl 6, juv), van der Weriffet al. 10237 (MO); carretera Iquitos-Nauta, km 44, 150 m, 5 Apr 1989 (fr), Vdsquez et al. 11954 (MO); Punchana, Rfo Mom6n, 2 Jul 1998 (fl 6), Rimachi 12301 (MO); Rfo Tigre-Rio Corrientes, Cocha Be- lem, 116 m, 20 May 1987 (fl 6), Grandez & Chiquispama 982 (MO); Sargento Lores, Constancia Norte, 116 m, 17 Apr 1997 (fr), Vdsquez et al. 23419 (MO). MADRE DE DIos: Tambopata, Cusco Amaz6nico, 200 m, 8 Oct 1991 (fr), Ti- mana & Jaramillo 2476 (MO); Zona Reserve de Tambopata, 280 m, 13 Aug 1990 (fr), Reynel & Meneses 5089 (MO), 14 Aug 1990 (fl 6), Reynel & Meneses 5125 (MO), Hermosa Chica, Nativa de Infierno, 260 m, 19 Dec 1990 (fl 6, juv), Pesha 68 (K, MO).

BRAZIL. ACRE: Mancio Lima, Parque Nacional Serra do Divisor, Rio Azul, 11 May 1996 (fl 6), Daly et al. 9024 (MO), (fr), Daly et al. 9025 (MO); Manoel Urbano, Rio Purus, margem esquerda, seringal Nova Olinda, Colocaqao Nova Olinda, 24 Nov 1966 (fr), Silveira et al. 1565 (MO); Marechal Taumaturgo, Rio Jurua, Reserva Extrativista de Alto Jurua, 200 m, 31 Mar 1993 (fr), Daly 7656 (MO); Peix- oto, 13 Aug 1989 (fr), Santos et al. 39 (INPA); Rio Macau- han (tributary of Rio Yacu), 4 Aug 1933 (fr), Krukoff 5279 (F, G, MO, NY, U, US); Sena Madureira, Rio Macaua, Fa- zenda Sao Jose, 30 Mar 1994 (fr), Daly et al. 8104 (MO), Seringal Capital, 7 Oct 1978 (fr), Lima & Sousa 230 (INPA). AMAZONAS: Coari, Rio Urucu, 7 Dec 1993 (fr), Aguiar et al. RUC-122 (INPA); ibid., Rio Urucu-Petrobras, Santo An- tonio, Porto Helio, 2 May 1988 (fl 6), Matos et al. 207 (INPA); Humaita, estrada Humaita-Ldbrea, km 80, Igarape do Riozinho, 4 Jun 1982 (fl 6), Filho 82-2N & 82-3N (HBG), near Tres Casas, 14 Sep-il Oct 1934 (fr), Krukoff 6116 & 6406 (A, F, G, K, MO, NY, U, US); Itapiranga, Rio Pitinga, 24 Aug 1979 (fr), Cid Ferreira et al. 690 (F, HBG, INPA, K, MO, NY, US); Manaus, Cachoeirinha, 1929 (fl 6 ), Ducke 2480 (U); Novo Japura, entre Tamandare e Man- guari, Rio Japura, 12 Nov 1982 (fr), Cid Ferreira & Lima 3624 (F, MO, NY, US); Presidente Figueiredo, Rio Uatuma, UHE Balbina, 23 Apr 1987 (fr), Bilby et al. 230 (MO); Rio Cunhua, Deni Indian Village, 29 Nov 1971 (fr), Prance et al. 16527 (HBG, INPA, NY, US); Rio Ituxi, 8 Jul 1971 (fl Y, fr juv), Prance et al. 14013 (F, HBG, INPA, NY, US); Sao Paulo de Oliven,a, 13 May 1945 (fl Y, frjuv), de Lemos Froes 20899 (NY, US). PARA: Rio Branco de 6bidos, Barro Vermelho, 27 Dec 1913 (fr), Ducke s.n. M.G. 15252 (NY); Rio Tapaj6s, 31 May 1923 (fl 6), Ducke s.n. R 17542 (U); Rio Itapacaru, 4 Apr 1924 (fl Y, fr), Kulhmann s.n. R 18360 (U). RONDONIA: Ji-Parana, Gleba G, km 3, 29 Mar 1983 (fl 6), Silva 6059 (MO, NY), km 2, 29 Mar 1983 (fl 6), Filho 83-30 (HBG), 31 Mar 1983 (fr), Filho 83-49 (HBG), km 3,

Mar 1983 (fr), Silva 6078 (MG); Porto Velho, 18 Jun 1986 (fr), Cid Ferreira 7485 (K, MO, NY).

BOLIVIA. PANDO: Madre de Dios, 165 m, 21 May 1991 (fl Y, fr), Killeen 3880 (MO); Manuripi, 35 km al N de Puerto America, 200 m, 19 May 1994 (fr), Jardim 764 (MO); Triunfo, 54 km SW Cobija, 250 m, 3 Aug 1988 (fl Y), Pennington et al. 80 (F, MO).

Local names. Ecuador: ocatue (Huaorani), honcatohue (Huaorani), urcuayua (Quichua), plata gigante. Peru: inchaquito, moena hoja ancha. Brazil: louro branco, louro de folha cinzenta, louro pichuri, louro seda.

Endlicheria ruforamula is distinguished from others of the E. sericea species group by its campanulate flowers and rusty tomentellose branchlets and inflorescences. Further, its large hemispherical cupules with entire margins dry a characteristic brick-red color that is unmistakable among the dark brown to blackish cupules of similar size and shape produced by E. griseo-sericea, E. sericea, and E. tschudyana.

Still, the species circumscribed by these three peculiarities is a variable one. In flowers of the type material and other staminate plants from lower An- dean montane regions (e.g., Cer6n 2600 and Forero et al. 2379), inflorescence bracts and bracteoles persist at anthesis. Furthermore, the pistillode is remarkably well-developed and provided with a clearly differentiated, albeit small, stigma. However, in most staminate plants from lowland Amazonia, inflorescence bracts and bracteoles are lost at anthesis and the pistillode is absent from flowers. The latter are also slightly smaller and have greyish rather than reddish irndument. Yet, a narrower species concept than adopted here would provide two entities of which only staminate plants may be distinguished. A third group of specimens (e.g., Vdsquez et al. 11954), mostly in fruit, have asymmetric basal secondaries that originate from the junction of the midrib and leaf base in a manner reminiscent of E. metallica and members of Rhodostemonodaphne in the R. grandis species group (sensu Madrifinan, 1996b). This venation is very distinctive and may be sufficient to circumscribe an easily recognizable species. However, in the only staminate plant with this venation (Freire & Santi 3294) inflorescence bracts and bracteoles persist at anthesis, and flowers, complete with pistillode, are indistinguishable from typical material.

Representatives of Endlicheria ruforamula were previously assigned to E. lhotzkyi, a species that maintains an indument of much longer hairs, has rotate flowers, and has smaller fruits in shallow patelliform cupules with lobed margins.


51. Endlicheria aurea Chanderbali, sp. nov. Type. Bolivia. La Paz: Prov. Nor Yungas, 13.7 km NW of San Pedro, 1500 m, 15-16 Jan 1983 (fl d), Solomon 9255 (holotype: MO; isotypes: K, NY, U, US). Fig. 38

Ex affinitate Endlicheriae sericeae et specierum affinium locellis antherarum sub-apicalibus distinguenda.

Trees, 4-25 m. Branchlets slender, midway along flush 3-4 mm diam., angular, densely golden to reddish sericeous, the surface concealed by the indument cover, the hairs short, to 0.2 mm, straight, appressed; terminal buds plump, 3 X 3 mm, the indument as on branchlets. Leaves alternate, widely and evenly spaced along current flush; petioles slender, to 1.5 X 0.2 cm, semi-terete, striate above, the indument as on branchlets; laminae chartaceous to coriaceous, plane, ovate to lanceolate, 9-28 X 3-10 cm, the base obtuse to acute, briefly decurrent, the apex acute, acuminate for up to 2 cm, the margins minutely recurved throughout; upper surface dull greyish green to olive- brown, waxy, shining, the primary to fourth-order veins raised, their prominence decreasing with rank; lower surface densely sericeous, the hairs as on branchlets, uniformly distributed, all vein orders raised, their prominence decreasing with rank; secondary veins 4-5 per side, ? evenly spaced, slightly more distant around midlamina, ascending at 50-60? (more obtuse around midlamina), arcuate, distal pairs loop- connected; tertiaries roughly horizontal, between secondaries straight to once-forked. Staminate inflorescences evenly spaced along current flush in the axils of foliage leaves, to 10 cm long with 10 lateral branches, branch orders 3-4, the highest order dichasial, lax, the flowers distant, the axes densely rusty sericeous; bracts and bracteoles caducous by anthesis, lanceolate, rusty sericeous; pedicels terete, to 1 mm long, those supporting secondary flowers slightly shorter. Flowers infundibuliform, to 3 mm diam., rusty sericeous outside; receptacle shallowly infundibuliform, 0.5 X 1 mm, densely rusty pilose inside. Tepals chartaceous, ovate, 1 X 0.6 mm (the inner whorl slightly narrower), erect to ascending at anthesis, the tips slightly incurved over stamens, the outer surface densely rusty sericeous (distal margins and apex of whorl II glabrous), the inner surface densely grey-tomentose (distal margins and apex of whorl I glabrous). Stamens of whorls I and II stipitate, 0.5 mm tall, the anthers ovate, 0.3 X 0.4 mm (whorl II slightly smaller), glabrous, the apex apiculate, the connectives slightly prolonged between the 2 locelli, these obliquely hemispherical, subapical, the filaments laminar, narrower than anthers, grey-

tomentose; whorl III stamens columnar, 0.5 mm tall, the anthers ovate, erect, dimensions as in outer whorls, locelli 2, subapical, the filaments as broad as anthers, columnar, densely grey-tomentose, the basal glands sessile, globose; whorl IV wanting; pistillode wanting. Pistillate inflorescence with indument and color as in staminate plants, but shorter and with fewer lateral branches, the flowers similar in size and shape; stamens sterile, smaller; ovary glabrous; style slender, distinct from ovary; stigma tri-lobed, 0.3 mm diam. Fruits borne on slender terete pedicels of up to 0.7 X 0.2 cm; cupules shallowly hemispherical, 0.3 x 1 cm; glabrous outside and inside, the margins undulate; drupes ellipsoid, 1.5 X 1 cm.

Distribution (Fig. 39) and ecology. Small to medium-sized trees of eastern Andean forests from Colombia to Bolivia at ca. 850-1530 m. Flowering from October through January, fruits collected in Jan- uary and late April to early May.

Additional specimens examined. COLOMBIA. Pu- TUMAYO: Mocoa, San Antonio, Alto Campucaca, La Mar- iposa, 1350 m, 20 Apr-1 May 1994 (fr), Pilar Franco et al. 5499 (MO).

ECUADOR. ZAMORA-CHINCHIPE: Nangaritza, Miazi, Rio Nangaritza, 1200 m, 10 Dec 1990 (fl d), Palacios 6732 (MO, NY); Zamora, Parque Nacional Podocarpus, 1400 m, Jan 1995 (fl Y, fr juv), Palacios & Tirado 13342 (MO).

PERU. PUNO: Moro, Jan 1866 (fl d), Pearce s.n. (K 52, MO 1611827).

BOLIVIA. LA PAZ: Prov. Larecaja, Cocacabana (about 10 km south of Mapiri), 850-950 m, 8 Oct-15 Nov 1939 (fl 5, juv), Krukoff 11262 (A, F, G, K, MO, NY, U, US); Prov. Nor Yungas, 4 km NE above Incahuara, 13.5 km above San Pedro, 1500-1530 m, 23 Jan 1984 (fl d), Gentry & Solomon 44524 (MO); Mapiri, San Carlos, 850 m, 11 Dec 1926 (fl d), Buchtien 745 (HBG, NY, US), 9 Jan 1927 (fl d), Buch- tien 746 (GH), 31 Jan 1927 (fl 5), Buchtien 748 (F, NY, US).

Endlicheria aurea has a most unusual androecium

where, in stamens of all whorls, depressed-ovate anthers with almost apically positioned locelli sit atop densely pubescent filaments. Whorl I and II anthers, though never whorl III anthers, may be ovate in similarly sericeous species, but locelli are nevertheless more introrse than apical. The position of the locelli and the densely pubescent filaments are reminiscent of stamens in Aniba, and as is typical of that genus, in E. aurea the tepals surround the stamens at anthesis. Advantages of having locelli placed closer to the tops of stamens in such flowers seem obvious, but of the several species of Endlicheria with erect tepals only E. aurea has such anthers.

~~o' 000 fA, X I H~~~~~~~~~~~~~~~3c

0.5mm0

FIG. 38. Endlicheria aurea (Solomon 9255). A. Habit. B. Single leaf showing venation. C. Leaf base below. D. Close up of lower leaf suface. E. Flower from above. F. Flower with facing tepal turned down to reveal androecium. G. Flower l.s. H. Whorl III stamen seen from without. I. Whorl I stamen seen from within.


*1~~~~~~~~~~~~~A

........ ... - \ --- --- ---- 4---

FIG 39 Distribution of Endlicheria aurea E lhorzkyi and E klugu

r~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~M

FIG. 39. Distribution of Endlicheria aurea, E. Ihotzkyi, and E. klugEi.~~~~~~~~~~~~~.. .. .. . . . .

Kostermans (1937) noted denser indument inside flowers of Buchtien's collections but assigned them (and Pearce's) to Endlicheria lhotzkyi, a species of Brazilian cerrado with rotate flowers, a conventional androecium, and an indumentum of much longer, more ascending hairs.

52. Endlicheria Ihotzkyi (Nees) Mez, Jahrb. K6nigl. Bot. Gart. Berlin 5: 122. 1889. Ocotea Ihotzkyi Nees, Syst. laur. 475. 1836. Strychnodaphne lhotzkyi (Nees) Meisn., D.C. Prodr. 15(l): 143. 1864. Type. Brazil. Mato Grosso: Cujaba, Nov (fl 9), Manso & Lhotzky 84 (lectotype, designated by Kostermans, 1937: B-n.v.; isolectotypes: G-n.v., GZU-n.v.; fragments, F, NY; photos [neg. 3813 ex B], F, GH, NY).

Trees to 15 m. Branchlets slender, midway along flush 3-5 mm diam., distally weakly angular, soon terete, pubescent, the surface concealed by the indument cover, the hairs relatively long, to 0.5 mm,

straight to crooked, weakly appressed to ascending, yellowish to rusty red; terminal buds plump, 4 X 5 mm, the indument as on branchlets. Leaves alternate, widely and evenly spaced along current flush; petioles slender, to 2 X 0.3 cm, semi-terete, the indument as on branchlets; laminae coriaceous, plane, ovate, 9-25 X 4-10 cm, the base obtuse to acute, briefly decurrent, the apex acute, acuminate for up to 1 cm, the margins minutely recurved throughout; upper surface dull greyish green to olive-brown, waxy, the primary to fourth-order veins raised, their prominence decreasing with rank; lower surface densely golden sericeous, the hairs as on branchlets, uniformly distributed, all vein orders raised, their prominence decreasing with rank; secondary veins 5-6 per side, ? evenly spaced, slightly more distant around midlamina, ascending at 50 60? (more obtuse around midlamina), arcuate, distal pairs loop-connected; tertiaries roughly horizontal, between secondaries forked. Staminate inflorescences evenly spaced along current flush in the axils of foliage leaves or cata-


phylls, to 15 cm long with 10 lateral branches, branch orders 3-4, the highest order dichasial, lax, the flowers distant, the axes densely pubescent, the indument as on branchlets; bracts and bracteoles caducous by anthesis, lanceolate, sericeous; pedicels gradually increasing in diameter apically, to 1.3 mm long, those supporting secondary flowers slightly shorter. Flow- ers rotate, rusty sericeous outside, the indument thinning distally; receptacle shallowly infundibuliform, 0.5 x 1 mm, densely grey-pilose inside. Tepals chartaceous, ovate, 1.3 x 0.8 mm, spreading at anthesis, the androecium exserted, the outer surface sparsely rusty sericeous, the inner surface sparsely grey-pilose near base, the margins and apex minutely papillose, otherwise, glabrous. Stamens of whorls I and II broadly stipitate, 1 mm tall, the anthers ovate, 0.5 x 0.5 mm, glabrous, the apex rounded to truncate, the connectives hardly prolonged between the 2 locelli, these suborbicular, introrse-latrorse, the filaments laminar, slightly narrower than anthers, densely grey- pilose; whorl III stamens broadly stipitate, 1 mm tall, the anthers oblong 0.5 x 0.4 mm, erect, locelli 2, extrorse-latrorse, the filaments slightly narrower than anthers, laminar, densely grey-pilose, the basal glands sessile, globose; whorl IV wanting; pistillode wanting. Pistillate inflorescence with indument, color, and branching as in staminate plants, the flowers similar in size and shape; stamens sterile, smaller; ovary glabrous; style stout, indistinct from ovary; stigma tri- lobed, 0.3 mm diam. Fruits bome on short claviform pedicels of up to 5 x 3 mm; cupules shallowly infundibuliform, 0.3 X 1 cm, glabrous inside and outside, the margins sharply lobed, tepal bases persisting; drupes ellipsoid, to 1.5 X 1 cm.

Distribution (Fig. 39) and ecology. Small trees of gallery forests apparently restricted to the cerrado vegetation found in the states of Mato Grosso and Goia's, Brazil, at ca. 400-500 m. Flowers collected in May, June, and November, fruits from May to Octo- ber.

Representative specimens examined. BRAZIL. GoLas: Araguaina, 500 ft, 11 Aug 1963 (fr), Maguire et al. 56091 (HBG, MO, NY). MATO GROSSO: Barra do Gargas- Xavantina Rd., 77 km from Barra do Garqas, 400 m, 15 Jun 1966 (fr), Hunt & Ramos 6004 (NY); ca. 270 km N of Xav- antina, 4 May 1968 (fl d), Ratter et al. 1278 (E, K, MO, NY, U); close to Xavantina-Sao Felix Rd., 6 Apr 1968 (fl d), Ratter et al. 840 (E, K, NY); Corrego do Surucucu, 10 Oct 1986 (fr), Sidney & Onishi 1308 (NY); Rio Araguaia, Xavantina, 400 m, 8 Jun 1966 (fr), Irwin et al. 16777 (HBG, MG, MO, NY).

Local names. Brazil: louro amarello, louro dorado, louro dourado, louro roxo.

Unlike most members of the Endlicheria sericea species group, E. Ihotzkyi maintains a vestiture of ascending rather than closely appressed hairs. These hairs provide a shaggy aspect unmistakable among the smoothly sericeous branchlets typical of vegetatively similar species. Hairs are also ascending in E. ruforamula but are more reddish in color and less than half the length found in E. lhotzkyi. Flowers of E. lhotzkyi resemble those of Antillean E. sericea in being rotate with horizontally spreading tepals, but the tepals are chartaceous rather than fleshy with inner surface indument restricted to the base and consisting of straight rather than crooked or crinkled hairs. Fruits of E. lhotzkyi are conspicuous for the shallow cupules with strongly lobed margins. Of vegetatively similar species, these cupules are matched only by the An- dean E. aurea and densely sericeous forms of E. anomala, both with very different flowers. Goeppertia chrysophylla Meisn., cited in synonymy by Mez (1889) and Kostermans (1937), is an illegitimate name based on a duplicate of the type material of E. lhotzkyi deposited in DeCandolle's herbarium, now in Geneva (G).

53. Endlicheria klugii 0. C. Schmidt, Repert. Spec. Nov. Regni Veg. 31: 173. 1933. Type. Colombia. Putumayo: Umbria, 325 m, Dec 1930 (fl d), Klug 1904 (lectotype, designated by Kostermans, 1937: NY; isolectotypes: A, B-n.v., F-n.v., GH, K-n.v., MO, S, US).

Trees to 15 m. Branchlets relatively stout, midway along flush 4-5 mm diam., angular, silvery to tawny sericeous, the surface concealed by the indument cover, the hairs short, to 0.15 mm, straight, appressed; terminal buds relatively plump, 1 X 0.6 mm, rusty sericeous. Leaves alternate, widely and evenly spaced along current flush; petioles long and slender, to 7 X

0.4 cm, semi-terete, the indument as on branchlets; laminae chartaceous, plane, broadly ovate to elliptic, 12-35 X 6-18 cm, the base obtuse to rounded, rarely acute, briefly decurrent, the apex broadly acute, acuminate for up to 6 cm, the margins minutely recurved throughout; upper surface dull slate grey to olive- brown, minutely punctulate, the midrib and secondaries prominent, the higher-order venation prominulous; lower surface sparsely or densely pubescent, the hairs silvery to rust brown, appressed, uniformly distributed, the epidermis clearly visible to obscured, all vein orders raised, their prominence decreasing with rank; secondary veins 4-6 per side, ? evenly spaced, slightly more distant around midlamina, ascending at 50-60o (more obtuse around midlamina), arcuate, the


lowermost pair occasionally asymmetric (one or both merging with margin at base), distal pairs loop- connected; tertiaries oblique to midrib, between secondaries straight to once-forked. Staminate inflorescences evenly spaced along current flush in the axils of foliage leaves, to 25 cm long with 9 lateral branches, branch orders 4-5, the highest order dichasial, lax, the flowers distant, the axes rusty to tawny sericeous; bracts and bracteoles caducous by anthesis, lanceolate, the indument as on axes; pedicels terete, to 2 mm long, those supporting secondary flowers slightly shorter. Flowers hypocrateriform, 2.5 mm diam., densely pubescent outside, the hairs appressed, reddish to brown; receptacle narrowly infundibuliform, 2 X 1 mm, grey-velutinous inside. Tepals chartaceous, ligulate, 1 X 0.6 mm (the inner whorl slightly narrower), spreading at anthesis, the inner surface grey-tomentose near base, the margins and apex inside sparsely papillose, otherwise glabrous. Stamens of whorls I and II stipitate, 0.6 mm tall, the anthers narrowly ovate, 0.3 X 0.25 mm, glabrous, the apex apiculate, the connectives prolonged between the 2 locelli, these suborbiculate, introrse-latrorse, the filaments S-shaped, narrower than anthers, densely grey- tomentose; whorl III stamens broadly stipitate, 1 mm tall, the anthers depressed-obovate, 0.3 X 0.3 mm, erect, locelli 2, extrorse-latrorse, the filaments slightly narrower than anthers, ligulate, densely grey- tomentose, the basal glands sessile, minute, globose; whorl IV wanting; pistillode fusiform. Pistillate inflorescence with indument and color as in staminate plants, but shorter and with fewer lateral branches, the flowers slightly deeper; stamens sterile, smaller; ovary glabrous, ellipsoid; style slender, distinct from ovary, 0.6 mm long; stigma broadly tri-lobed to discoid, 0.5 mm diam. Fruits borne on stout, claviform pedicels of up to 2 X 0.8 cm; cupules shallowly infundibuliform to patelliform, to 0.4 X 1.3 cm, glabrous outside, sparsely rusty strigose inside, the margins undulate; drupes ellipsoid to obovoid, to 3 X 1.4 cm.

Distribution (Fig. 39) and ecology. Small trees from the nonflooded forests of western Amazonia and adjacent eastern Andean foothills at ca. 150-1300 m. Flowering September to following March, fruits available year round.

Representative specimens examined. COLOMBIA. PUTUMAYO: Umbrfa, 325 m, Dec 1930 (fl d), Klug 1884

(G, GH, K, MO, NY, S). ECUADOR. NAPO: Estaci6n Experimental INIAP-

Napo, Payamino, Reserva Florfstica "El Chuncho," 250 m, 16-26 Feb 1986 (fl Y), Neill 7152 (MO, NY, US); Tena,

Estaci6n Biol6gica Jatun Sacha, 400 m, 2 Jan 1998 (fl d), Neill 11028 (MO).

PERU. AMAZONAS: Bagua, Imaza, Comunidad de Ya- mayakat, 320 m, 14 Feb 1996 (fl d), Jaramillo et al. 1159 (MO); Rfo Santiago, Caterpiza, 180 m, Sep 1979 (fr), Hu- ashikat 436 (HBG). HUANUCO: Pachitea, Pucallpa, Sira Mtns., 680 m, 8 Apr 1988 (fr), Wallnofer 111-8488 (MO). LORETO: Alto Amazonas, Rio Huallaga, Shucushuyacu, 250 m, 12 Sep 1981 (fl d), Vdsquez & Jaramillo 2425 (F, MO, NY); Maynas, Iquitos, Estaci6n Experimental IIAP, Allpahuayo, 150 m, 24 Mar 1992 (fl d), Vdsquez et al. 18095 (MO). MADRE DE Dios: ManU, Atalaya, 500-700 m, 8 Dec 1983 (fl Y, fr), Foster & Wachter 7286 (G, MO). PAsco: Oxapampa, Palcazu, 380 m, 24 Jan 1984 (fl Y), Foster 9483 (G, MO, NY), 300-600 m, 13 Nov 1985 (fr), Hartshorn et al. 2824 (MO).

BRAZIL. AcRE: Bujari, Floresta Estadual do Antimari, 10 Mar 1997 (fr), Daly et al. 9425 (MO). AMAZONAS: Rio Ia, 24 Feb 1977 (fl d), Mori et al. 9097 (F, HBG, INPA, MO, NY, U, US); Rio Javari, behind Palemeiras Army Post, 31 Jul 1973 (fr), Lleras et al. P16960 (HBG, K, MO, NY, R, US).

Local name. Peru: yuwich (Huambisa).

The S-curved whorl I and II filaments of Endli- cheria klugii are unique in the genus. These curious filaments combined with the ovate anthers confer on stamens an aspect that resembles a cobra raised in strike position (Fig. IF). The flowers are themselves remarkable for their slender receptacles below horizontally spreading tepals.

The long petioles supporting broadly ovate leaves with arcuate secondaries are sometimes matched by E. ruforamula, from which E. klugii can be distinguished, even without flowers, by its closely appressed indument on branchlets and shallow cupules with clavate pedicels.

54. Endlicheria argentea Chanderbali, sp. nov. Type. Peru. Loreto: Requena, Sapuena, Arboreto Jenaro Herrera, 130 m, 14 Jul 1989 (fl 6), Vdsquez & Jong 12397 (holotype: MO; isotypes: AMAZ, HBG, MO). Fig. 40

Inter Endlicheriae sericeae et specierum affinium floribus infundibularibus ad E. klugii accedens sed foliis supra costa impressis et staminibus rectis differt.

Trees to 5 m. Branchlets slender, midway along flush 2 mm diam., distally weakly angular, soon terete, sericeous, the surface concealed by the indument cover, the hairs short, to 0.3 mm, straight, appressed, silvery to straw-colored, shining; terminal buds slender, 2 X 0.6 mm, sericeous. Leaves altemate, widely and evenly spaced along current flush; petioles slen-

-5

1mm~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~1

FIG. 40. Endlicheria argentea (Vdsquez & Jong 12397). A. Habit. B. Single leaf showing venation. C. Leaf base below. D. Rlower. E. Rlower with tepals removed. F. Whorl I stamen seen from within. G. Whorl III stamen seen from without.


der, to 1.5 X 0.2 cm, terete, the indument as on branchlets; laminae chartaceous, plane, narrowly ovate to lanceolate, 14-17 X 3-5 cm, the base acute, briefly decurrent, the apex acute, acuminate for up to 4 cm, the margins flat throughout; upper surface olive- brown, minutely punctulate, the midrib impressed, the higher-order veins prominulous; lower surface sericeous, the hairs silvery, uniformly distributed, concealing the epidermis, all vein orders raised, their prominence decreasing with rank; secondary veins 3- 4 per side, closer near base and apex, widely spaced around midlamina, ascending at 50-60? (more acutely towards apex), arcuate, distal pairs loop-connected; tertiaries roughly horizontal, between secondaries once-forked to straight. Staminate inflorescences evenly spaced along current flush in the axils of foliage leaves, to 10 cm long with 8 lateral branches, branch orders 2-3, the highest order dichasial, lax, the flowers distant, the axes silvery sericeous; bracts and bracteoles caducous by anthesis, narrowly lanceolate, sericeous; pedicels terete, to 2 mm long, those supporting secondary flowers slightly shorter. Flowers tubiform, to 5 mm diam., sericeous outside; receptacle infundibuliforn, 2 X 1.5 mm, densely silvery pilose inside, tepals chartaceous, ovate, 1 X 0.6 mm (the inner whorl slightly narrower), ascending at anthesis, surrounding androecium, the inner surface densely pubescent, the hairs as in receptacle, ascending to erect, the margins and apex inside papillose. Stamens of whorls I and II stipitate, 1 mm tall, the anthers broadly ovate, 0.3 X 0.5 mm, glabrous, the apex apiculate, the connectives prolonged between the 2 locelli, these obliquely hemispherical, introrse, the filaments laminar, narrower than anthers, densely silvery pilose, the hairs as in receptacle; whorl III stamens sessile, 1 tall, the anthers depressed-obovate, 0.3 X 0.3 mm, erect, locelli 2, extrorse-latrorse, the filaments as broad as anthers, laminar, the indument as in outer whorls, the basal glands sessile, ovate, the apex apiculate; whorl IV wanting; pistillode filiform. Pistillate plants unknown.

Distribution (Fig. 41) and ecology. Known only from the type (but see discussion), a small plant of seasonally inundated lowland (ca. 130 m) forest in Peru, found flowering in July.

Endlicheria argentea differs from similarly sericeous species by its tubiform flowers with tall slender stamens, but the impressed midrib above is also distinctive. Perhaps its closest relationship lies with E. klugii, with which it shares narrowly infundibuliform receptacles and depressed-obovate whorl III anthers. However, the flowers of E. argentea are externally

silvery sericeous rather than reddish or brown and filaments of whorl I and II stamens are straight rather than curved. Endlicheria argentea also appears to be a less robust species with slender branchlets and petioles less than half the dimensions found in E. klugii.

Urrego et al. 1218 (MO), from Rio Caquetai in Amazonian Colombia, is an exact vegetative match and likely belongs here but its sterile condition pre- vents definite identification.

55. Endlicheria robusta (A. C. Sm.) Kosterm., Re- cueil Trav. Bot. Neerl. 34: 556. Cryptocarya robusta A. C. Sm., Bull. Torrey Bot. Club 58: 97. 1931. Type. Peru. Junin: San Nicholas, Pichis Trail, 1100 m, 4-5 Jul 1929 (frjuv), Killip & Smith 26077 (holotype: NY; isotypes: F, US).

Trees to 20 m. Branchlets relatively slender, midway along flush 2-3 mm diam., weakly angular, densely strigillose, the surface barely visible, dark brown, the hairs very short, to 0.1 mm, appressed, tawny or light brown; terminal buds slender, 2 X 0.2 mm, reddish sericeous. Leaves alternate, widely and evenly spaced along current flush; petioles slender, to 2 X 0.2 cm, semi-terete to broadly canaliculate, the indument as on branchlets; laminae chartaceous, plane, ovate to elliptic, 8-20 X 3-10 cm, the base acute, briefly decurrent, the apex acute, acuminate for up to 3 cm, the margins flat or weakly recurved in lower half; upper surface olive-brown to light green, minutely punctulate, the primary to fourth-order veins raised, their prominence decreasing with rank; lower surface sericeous to sparsely strigillose, the hairs tawny to light brown, dull to lustrous, uniformly distributed on lamina, sparser on midrib and secondaries, all vein orders raised, their prominence decreasing with rank, the midrib often darker and crooked after midcourse; secondary veins 5-7 per side, + evenly spaced, slightly more distant around midlamina, ascending at 50-60O (more obtuse towards apex), arcuate, distal pairs loop-connected; tertiaries oblique to midrib, between secondaries straight or forked. Staminate inflorescences evenly spaced along current flush in the axils of foliage leaves, to 10 cm long with 8 lateral branches, branch orders 2-3, the highest order dichasial, lax, the flowers distant, the axes grey- strigillose, distalmost branches and pedicels more densely so; bracts and bracteoles persistent at anthesis, lanceolate, silvery sericeous; pedicels terete, to 3 mm long, those supporting secondary flowers slightly shorter. Flowers obconical, or urceolate, 2 mm diam., sparsely grey-strigillose outside; receptacle deeply cyathiform, the base abruptly truncate, 1.3 x 2 mm,


L /~~~~~~~~~~~~~~~~~~~~~~~

.Api ' ' '';

F Db o c r a r

FIG. 41. Distnbution of Endllicheria argentea and E. robusta.

densely grey-pilose inside. Tepals chartaceous, broadly ovate, 0.6 x 1 mm (the inner whorl slightly narrower), inflexed to erect at anthesis, the androecium included, the inner surface sparsely grey- strigillose, the margins and apex inside sparsely papillose. Stamens of whorls I and II stipitate, 0.6 mm tall, broadly anthers ovate, 0.5 X 0.5 mm, glabrous, the apex sharply apiculate, the connectives narrowly prolonged between the 2 locelli, these suborbiculate, introrse-latrorse, the filaments laminar, slightly narrower than anthers, sparsely grey-tomentellose; whorl III stamens sessile, 0.7 mm tall, the anthers narrowly oblong, 0.4 X 0.2 mm, erect, locelli 2, extrorse- latrorse, the filaments broader than anthers, the indument as in outer whorls, the basal glands sessile, globose; whorl IV wanting; pistillode fusiform. Pistillate inflorescence with indument and color as in staminate plants, but shorter and with fewer lateral branches, the flowers similar in size and shape; stamens sterile, smaller; ovary glabrous, ovoid; style slender, distinct from ovary; stigma minutely tri-

lobed, 0.5 mm diam. Fruits borne on claviform pedicels of up to 1 X 0.3 cm; cupules shallowly hemispherical, to 0.5 X 1 cm, glabrous inside and outside, the margins entire; drupes ovoid, to 1 X 0.8 cm.

Distribution (Fig. 41) and ecology. Small to medium-sized trees from western Amazonia and adjacent lower montane slopes of the eastern Andes at ca. 100-1800 m. Flowering specimens collected from March through July and from November through De- cember, fruits from April through July and from Oc- tober through January. If not a sampling artifact, collections of this species suggest two discrete reproductive cycles per year, with an early flowering period from March to July providing fruits from Oc- tober to January, and a late flowering period in No- vember and December responsible for fruit availability from April to July.

Representative specimens examined. ECUADOR. NAPO: Huaorani, Rio Shiripuno, Quehueiri-ono, 300 m, 14 May 1995 (fl 6), Miller et al. 638 (MO). PASTAZA: Auca,


Rio Tiguino, 115 km S de Coca, 320 m, 29 Apr 1989 (fl d),

Rubio 12 (MO); Lorocachi, 200 m, 25 May 1980 (fl d), Brandbyge & Asanza 30885 (AAU). SUCUMBiOS: Lago Agrio, Reserva Faunistica Cuyabeno, 265 m, 11 Mar-13 May 1990 (fl d3), Balslev et al. 97016 (MO). ZAMORA- CHINCHIPE: Nangaritza, faldas de las Cordillera del C6n- dor, 1600-1800 m, 5 Dec 1990 (fl d), Palacios & Neill 6522 & 6531 (MO).

PERU. AMAZONAS: Condorcanqui, Cenepa, Comuni- dad de Tutino, 500 m, 18 Jul 1997 (fl Y), Vdsquez et al. 24341 (MO), 24 Jul 1997 (fr), Rojas et al. 172 (MO). JUNiN:

San Nicolis, Pichis Trail, 1100 m, 4-5 Jul 1929 (fr), Killip & Smith 25975 (NY). LORETO: Maynas, Iquitos, carretera Iquitos-Nauta, km 44, 150 m, 14 Dec 1988 (fr), Vasquez & Jaramillo 11442 (GH, HBG, MO); Rio Napo, San Pedro, Isla de Mangua, below caserfo Juancho Playa, 6 Apr 1979 (fl d), Rimachi 4397 (MO). PASCO: PEPP Forest Reserve, Puerto Mairo, 300-600 m, 5 Jun 1986 (fl J, juv), Hartshorn et al. 2971 (MO); Santiago Soto, Buenos Aires, 300-600 m, 18 May 1986 (fl 6, juv), Hartshorn & Quijano 2940 (MO).

BRAZIL. AcRE: Cruzeiro do Sul, Rio Jurua & Rio Moa, Serra de Moa village, 22 Apr 1971 (fr), Prance et al. 12233 (HBG, MG, MO, NY, U). AMAZONAS: Labrea, sub- base do Projeto RADAM/BRAZIL, SG-20-YA, Ponto 02, 20 Jun 1976 (fl 6), Mota s.n. INPA 60364 (INPA); Vila Bittencourt, Rio Japura, Serrinha, 16 Nov 1982 (fr), Amaral et al. 488 (INPA, MG, MO).

Local names. Peru: moena blanca, moenilla, tikis, tindu, tunchi, tinchi.

Endlicheria robusta is alone among species with dense sericeous indument in having obconical or urceolate flowers with incurved or erect tepals that provide only a narrow terminal pore at anthesis. In the E. browniana species group, E. formosa and E. paradoxa have similar flowers as well as appressed indument; but although E. robusta can be subsericeous, its leaves are never obovate as in E. formosa, and the tertiaries never reticulate as in E. paradoxa. Similar flowers combine with rusty tomentose vegetative vestiture in E. szyszylowiczii.

A few specimens from Jenaro Herrera are placed here with hesitation. In Spichiger & Loizeau 4161 and 4108 (A) and Valcarcel & Chota 6/619 (MO), staminate flowers are more densely pubescent than ever encountered in E. robusta, and the fruiting specimen, Encarnaci6n 26150 (G, MO, US), shows rather small

shallow cupules with spheroid drupes. Such fruits are also found in Gentry et al. 37246 (F, G, MO) from Junfn, Peru, and Berg & Steward P19913 (HBG, K, MO, NY, US) from Mato Grosso, Brazil, extending considerably the geographic range of this variant. Similar cupules and densely pubescent flowers are found in E. tschudyana but the shape of the flowers and sharply apiculate whorl I and II anthers point to E. robusta.

56. Endlicheria paniculata (Spreng.) J. F. Macbr., Publ. Field Mus. Nat. Hist., Bot. Ser. 13 (2, 3): 850. 1938. Citrosma paniculata Spreng. Syst. Veg. 2: 545. 1825. Type. Brazil. Without locality, Sel- low s.n. (holotype: B [fide Kostermans, 1937]- n.v.).

Cryptocarya hirsuta Schott, Syst. Veg. 4(2): 405. 1827. Endlicheria hirsuta (Schott) Nees, Linnaea 8: 38. 1833. Goeppertia hirsuta (Schott) Nees, Syst. laur. 366. 1836. Type. Brazil. Rio de Janeiro: San Cris- tovao, Pohl 5611 (lectotype, designated by Koster- mans, 1937: W-n.v.; isolectotype: U).

Nectandra lucida Nees, Linnaea 8: 47. 1833. Type. Bra- zil. Without locality, Sellow s.n. (holotype: B [fide Kostermans, 1937]-n.v.; fragment, GZU [fide Roh- wer, 1993a]-n.v.).

Goeppertia longifolia Nees, Syst. laur. 368. 1836. En- dlicheria longifolia (Nees) Mez, Jahrb. Konigl. Bot. Gart. Berlin 5: 119. 1889. Type. Peru. Cuchero, Nov 1821 (fl 5), Poeppig 1520 (syntypes: B-n.v., BM- n.v., G-n.v., GOET-n.v., KIEL-n.v., LE-n.v., LZ- n.v., NY-n.v., OXF-n.v., P-n.v., W-n.v.; fragment, U).

Goeppertia panicularis Nees, Syst. laur. 368. 1836. En- dlicheria panicularis (Nees) Mez, Jahrb. Konigl. Bot. Gart. Berlin 5: 128. 1889. Type. Brazil. Rio de Janeiro: Without locality, Oct-Dec (fl 5), Lhotzky 43 (holotype: B [fide Kostermans, 1937]-n.v.).

Goeppertia cantagallana Meisn., DC. Prodr. 15(1): 173. 1864. Type. Brazil. Minas Gerais: Canta Gallo, 1859 (fl 5), Peckholt 115 (holotype: M-n.v.; isotype: U).

Goeppertia hirsuta (Schott) Nees, var. coriacea Meisn., DC. Prodr. 15(1): 172. 1864. Syntypes. Brazil. Minas Gerais, Sellow s.n. (G-n.v., M-n.v.); Weddell s.n. (G-n.v., M-n.v.); Claussen 454 (G-n.v., M-n.v.); Aug 1840 (fl juv), Claussen 2454 [cited as 1454 by Meissner] (G-n.v., K, M-n.v.).

Goeppertia hirsuta (Schott) Nees, var. hirsutior Meisn., DC. Prodr. 15(1): 172. 1864. Endlicheria poeppigii Kosterm., Recueil Trav. Bot. Neerl. 34: 555. 1937. Type. Peru. Loreto: Maynas, Yurimaguas, Jun 1831 (fl 5), Poeppig 2298 (holotype: W-n.v.; isotypes: B- n.v., G-n.v., LE, LZ-n.v., OXF-n.v., U).

Goeppertia hirsuta (Schott) Nees, var. latifolia Meisn., DC. Prodr. 15(1): 172. 1864. Type. Brazil. Near Vi- t6ria, without date (fl 5), Sellow 433 (holotype: B- n.v.; isotype: K).

Endlicheria hirsuta (Schott) Nees, var. glabrata Glaziou, Bull. Soc. Bot. France 12 (3): 590. 1912. Syntypes. Brazil. Rio de Janeiro: Tijuca and Corcovado, 19 Oct 1868 (fl 5), Glaziou 3092 (B-n.v., K-n.v., P); road to Macaco, near Vista Chineza, 26 Dec 1886 (fl 5), Glaziou 16315 (B-n.v., K-n.v., P).

Endlicheria hirsuta (Schott) Nees, var. robusta Glaziou, Bull. Soc. Bot. France 12(3): 590. 1912. Syntypes. Brazil. Minas Gerais: Biribiry, 28 Mar 1892 (fr), Glaziou 19795 (B-n.v., K-n.v., P)


Endlicheria boliviensis Kosterm., Recueil Trav. Bot. Neerl. 34: 553. 1937. Type. Bolivia. Santa Cruz: Buena Vista, 500 m, 14 Mar 1925 (fl d), Steinbach 6985 (holotype: B-n.v.; isotypes: GH, K, MO, NY, S, U).

Endlicheria racemosa Lasser, Bol. Tecn. Minist. Agric. 3: 8. 1942. Type. Venezuela. Carabobo: Hospital near San Joaquin, 1000 m, 15 Aug 1918 (fl d), Pittier 8017 (holotype: VEN-n.v.; isotypes: GH, MO).

Trees to 10 m (rarely 20 m). Branchlets slender to stout, midway along flush 3-6 mm diam., angular, densely pubescent, the surface barely visible to concealed by the indument cover, the hairs relatively short to long, (0.3-)0.6(-1.5) mm, straight, erect to appressed, rusty red to grey; terminal buds plump, 3 X 2 mm, densely pubescent, the hairs as on branchlets, ascending. Leaves alternate, widely and evenly spaced along current flush; petioles slender, to 2 X

0.3 cm, terete, the indument as on branchlets; laminae chartaceous to membranaceous, plane, ovate to obovate, 7-30 X 2-12 cm, the base acute to obtuse, the apex obtuse to acute, acuminate for up to 1.5 cm, the margins minutely recurved throughout; upper surface reddish to olive-brown, waxy, the midrib and secondaries sunken, the higher-order venation raised or immersed; lower surface densely pubescent, the hairs as on branchlets, uniformly distributed or denser on main veins, all vein orders raised, their prominence decreasing with rank; secondary veins 4-6 per side, ? evenly spaced, slightly more distant around midlamina, ascending at 50-60O (more obtuse around midlamina), arcuate, distal pairs loop-connected; tertiaries roughly horizontal, between secondaries straight to forked. Staminate inflorescences evenly spaced along current flush in the axils of foliage leaves, to 20 cm long with 14 lateral branches, branch orders 3-4, the highest order dichasial, lax, the flowers distant, the axes densely pubescent, the indument as on branchlets; bracts and bracteoles caducous by anthesis, rarely persisting, ovate to lanceolate, densely pubescent, the hairs as on axes, ascending; pedicels terete, to 4 mm long, those supporting secondary flowers slightly shorter. Flowers rotate, 2-4 mm diam., sparsely to densely silvery or rusty strigose outside; receptacle infundibuliform, 0.5 X 1 mm, densely rusty or grey-pilose inside. Tepals chartaceous to membranaceous, ovate to ligulate, 1.5 X 0.7 mm, spreading at anthesis, the inner surface densely silvery grey-strigose, the margins minutely papillose. Stamens of whorls I and II stipitate, 1 mm tall, the anthers ovate, 0.6 x 0.4 mm, glabrous, the connectives broad above, reduced between, or homlike over, the 2 locelli, these suborbicular, introrse-latrorse, the filaments laminar, narrower than anthers, sparsely

grey-pilose; whorl III stamens broadly stipitate, 1 mm tall, the anthers oblong, 0.5 X 0.4 mm, erect, locelli 2, extrorse-latrorse, the filaments equal to or slightly narrower than anthers, ligulate, the indument as in outer whorls, the basal glands sessile, globose; whorl IV wanting; pistillode fusiform. Pistillate inflorescence with indument and color as in staminate plants, but shorter and with fewer lateral branches, the flowers similar in size and shape; stamens sterile, smaller; ovary glabrous, ovoid; style stout, indistinct from ovary; stigma broadly tri-lobed, 0.6 mm diam. Fruits borne on claviform pedicels of up to 2 X 0.5 cm; cupules hemispherical, to 1 X 1.5 cm, glabrous outside, strigose inside, the margins entire, or tepal bases persisting; drupes ovoid, to 2.5 x 1.2 cm.

Distribution (Fig. 42) and ecology. Small to medium-sized trees distributed from the Atlantic coastal forests of SE Brazil, through the lower slopes of the Andes in tropical South America to Panama in Central America. Occurring at 50-1000 m throughout its geographic range and reaching over 2000 m in the Andes. Flowers and fruits available throughout the year.

Representative specimens examined. PANAMA. PANAMA: El Llano-Carti Rd., 8-11 km from Inter- American Hwy., 300-400 m, 13 Aug 1975 (fl d), Mori 7708 (MO); Serranfa de Maje, 5-6 hrs walk from Choc6 Village, 650-800 m, 31 Mar 1982 (fr), Knapp et al. 4496 (MO). SAN

BLAS: El Llano-Carti Rd., Pacific side, 350 m, 13 Feb 1983 (fr), Hamilton & Stockwell 2905 (MO); Pemasky, carretera Nusagandi-Cartf, 5 Jun 1994 (fl 9, juv), Galdames et al. 1362 (MO).

COLOMBIA. CUNDINAMARCA: Laguna de Pedro Palo, 4 km from rd. Bogota to La Mesa, 2070 m, Oct 1990 (fr), Wijninga 580 (U). HUILA: Rio Negro, 30 km ESE of Baraya, ca. 2300 m, Oct 1944 (fl 9), Little 8878 (A), 30 Oct 1944 (fl 9), Little 8886 (A). MAGDALENA: Sierra Nevada de Santa Marta, 1700-1900 m, 5 Oct 1972 (fr), Kirkbride 2387 (US). META: La Macarena, Parque Nacional Natural Tini- guas, s.d. (fr), Polanco 222 (MO).

VENEZUELA. LARA: Distrito Moran, Quebrada Los Cedros, 1500 m, 8 Jul 1974 (fl d), Steyermark & Espinoza 110275 (MO, NY, VEN); Distrito Yaritagua, near border with Distrito Urachiche of Estado Yaracuy, 1450 m, 28 Mar 1975 (fl 6), Steyermark et al. 111746 (F, G, NY). TACHIRA:

Mata Mula, N of Delicias on rd. to Bramon, 1750 m, 26 Jul 1979 (fl d), Steyermark & Liesner 118687 (MO). YARA-

cuy: Cocorote, carretera Cocorote-Fila Las Cumaraguas, 1500-1600 m, 13 May 1994 (fr), Benftez de Rojas et al. 5096 (MO); Nirgua, al NW de Montalban, entre torres de relevo (TV), y Cumbre de Capotillo, 1350-1500 m, 19 Mar 1999 (fr), Meier & Kunert 4632 (MO).

ECUADOR. NAPO: Estaci6n Experimental INIAP- Payamino, 250 m, 18-26 Feb 1986 (fr), Palacios et al. 1030 (MO); Rio Wai-si-aya, 300 m, 28 Aug 1981 (fl d), Brand-


| ._ -................ ; ..... -- -- -- - - ------- ---

M.,4 1........ .- .-- - - -- , -,P{

I f-ioB''!?t-i 4;257.

...................... --:- - --- - - - ---e- i-

* f

FIG. 42. Distribution of Endlicheria paniculata.

byge et al. 36206 (AAU). SucuMBios: San Pablo de Kan- tesiya, Rfo Aguarico, 250 m, 23 May 1992 (fl d), Pfrommer 6 (MO).

PERU. AMAZONAS: Above mouth of Quebrada Cikan- inci, N of Rio Cenepa, 30 Dec 1972 (fl S), Berlin 753 (F, MO); Vista Alegre, puente sobre el Rio Salas, 1525 m, 30 Jun 1998 (fl d), Sdnchez et al. 9575 (MO). HuANUco: Pachitea, Honoria, Bosque Nacional de Iparia, 300-400 m, 30 Jan 1967 (fl &), Schunke 1587 (GH, INPA, K); Puerto Inca, Llullapichis, 270 m, 1 Dec 1989 (fl juv), Kroll Saldania 795 (G, MO). LORETO: Alto Amazonas, above Pongo de Manseriche, Rio Santiago, 200 m, 23 Dec 1931 (fl &), Mexia 6329 (F, GH, K, MO, NY); Yurimaguas, 140 m, 7 Jan 1933 (fl J), Klug 2833 (GH, K, MO). MADRE DE Dios: Mand, Parque Nacional Mand, Rfo Manu, 350 m, 10 Sep 1986 (fr), Foster 11339 (INPA, NY); Tambopata, Comuni- dad Nativa de Infierno, 260 m, 17 Jul 1989 (fr), Alexiades & Diaz 849 (MO). PASCO: Oxapampa, Chantabamba, 1890 m, 28 May 1982 (fl f), Smith et al. 1748 (MO); Palmazu, 2200 m, 2 Oct 1984 (fl 9), Smith et al 8670 (MO). SAN MARTiN: Mariscal Caceres, Tocache Nuevo, Quebrada de Canuto, 250 m, 2 Jan 1979 (fl 9), Schunke 10657 (MO).UCAYALI: Coronel Portillo, Padre Abad, La Divisoria

cerca a Rio Chino, 1400-1600 m, Jun 1976 (fr), Schunke 9242 (MO); Bosque Nacional Alexander von Humboldt; km 86 Pucallpa-Tingo Maria Rd., 270 m, Jan 1978 (fl d), Froehner 171 (HBG, MO).

BRAZIL. BAHIA: Rod. Porto Seguro-Eunapolis, km 31, 26 Nov 1970 (fl 6), Filho & Emmerich 2934 (R). DisTiuTo FEDERAL: Brasflia, bacia do Rio Saio Bartolomeu, 24 Jan 1980 (fl 6), Heringer et al. 3187 (MO); C6frego Quilombo, 18 Sep 1980 (fl 6), Heringer et al. 5513 (MO). EsPiRfRo SANTO: Concei,cao do Castelo, Rod. BR-262, Dec 1984 (fl dc), Hatschbach & Silva 48651 (MO); Santa Teresa, Reserva Biologica de Nova Lombardia, 700-750 m, 4 Feb 1985 (fl d), Peixoto et aL 3466 (MO). GoIAs: Serra do Caiap6, ca. 12 km S of Caiap6nia, 840 m, 2 May 1973 (fl d), Anderson 9604 (NY); Campinacu, 400 m, 10 Oct 1991 (fr), Cavalcanti et al. 947 (MO). MATO GRosso: Caba. do Rio Taquaraussu (Chapada), Mar 1911 (fl Y), Hoehne 3517 (R); Santa Anna da Chapada, Aug 1902 (fr), Robert 4856 (K). MATO GRosso Do SuL: Ponta Pora, Ponto Alto, 12 Feb 1983 (fl c), Hatschbach 46151 (MO); Rod. BR-163, 1 km L de Mundo Novo, 7 Nov 1993 (fl d), Hatschbach et al. 58562 (MO). MINAS GERAIS: Serro do Espinhaqo, 1100 m, 12 Feb 1969 (fl Y), Irwin et al. 23151 (MO); Rio Acima, Feb 1994


(fl 6), Costa s.n. BHCB 26334 (MO). PARANA: Cerro Azul, Caeceira do Ribeirao do Tigre, 14 Apr 1987 (fl 6), Hatsch- bach et al. 51226 (MO); Palotina, Perola Independente, 13 Mar 1985 (fl Y), Hatschbach & Silva 48978 (MO). Rio DE JANEIRO: Angra dos Reis, Reserva Biologica Est. da Praia do Sul, 2 Mar 1985 (fl 6), Araujo & de Oliveira 601 (MO); Santa Maria Madalena, Pedra Dubois, 900-1195 m, 22 Feb 1983 (fl 9), Plowman & de Lima 12895 (K, MO). Rio GRANDE DO SUL: Itacolumi, prope Gravataf, 11 Jan 1950 (fl 6), Rambo 45269 (K, MO); Porto Alegre, Morro da Pol- icia (Caseata), 9 Dec 1901 (fl 6), Malme 755 (R). SANTA CATARINA: Ararangua, Passo do Sertao, 10 m, Feb 1952 (fl 6), Reitz 4423 (MO); Horto Florestal I.N.P., Ihirama, 350 m, 2 Nov 1953 (fr), Reitz & Klein 1140 (MO). SAO PAULO: Paraguacu Paulista, Fazenda Sao Jose, 425-450 m, 8 Feb 1865 (fl 9), Eiten et al. 5899 (K, MO); Morro das Pedras, Iguape, Dec 1922 (fl 6), Brade 8195 (R).

BOLIVIA. BENI: Prov. Ballivian, Serrania del Pil6n La- jas, 850-900 m, 19-26 Feb 1990 (fl 9), Smith et al. 14052 (MO), 16 Feb 1992 (fl 9), Killeen & Smith 3634 (MO). LA PAZ: Prov. Itturalde, Buena-Vista Tacana, 17 Feb 1997 (fl 6), Serato 305 (MO); Prov. Murillo, Zongo valley, Cahua hydroelectric plant, 1200-1400 m, 23 Dec 1984 (fl 9), Sol- omon 12948 (MO). PANDO: Puerto Nuevo, Rio Ortan, Feb 1924 (fr), Meyer 146 (MO). SANTA CRUZ: Prov. Ichilo, Buena Vista, 300 m, 22 Mar 1995 (fl 9), Abbott & Isaacs 16484 (MO); Prov. Sara, Rio Surutd, 400 m, 2 Oct 1925 (fr), Steinbach 7264 (GH, K, MO, U).

PARAGUAY. ALTO PARANA: Centro Forestal alto Pa- rand, 12 km 0 de Puerto Presidente Strossner, 1 Feb 1984 (fl 6), Little 40105 (MO); Reserva Biol6gica Itabo, 9 Oct 1990 (fr), Schinini & Marmori 27009 (MO). AMAMBAY: Parque Nacional Cerro Cord, 300 m, 20 Feb 1982 (fl 6), Solomon et al. 7140 (MO); Torin, Aug 1985 (fr), Hatsch- bach & Cervi 48903 (MO). CAAGUAZU: S.loc., 12 Nov 1874 (fr), Balansa 2026 (K); s.loc., 5 Feb 1905 (fl 6), Hass- ler 8935 (K). CAAZAPA: Linea del Bosque Parque hacia Ao Ita y Ao Jakuy, 20 Jul 1986 (fr), Molas 771 (MO); Tavai, Prop. Trosiuk, 17 Mar 1989 (fl 6, juv), Soria 3419 (MO). CANENDIYUJ: Mbaracayu Natural Reserve, 13 Jan 1998 (fl 9), Zardini & Guerrero 47737 (MO); Lagunita, Sendero Arroyo Moroti, 13 Mar 1997 (fl 9), Jimenez et al. 1809 (MO). SAN PEDRO: Colonia 8 de Diciembre, 12 km al SE de Chore, 30 Sep 1987 (fr), Zardini & Ben(tez 3182 (MO); Colonia Naciente, 8 km al SE de Chore, 2 Oct 1987 (fr), Zardini & Benftez 3429 (MO).

Local names. Ecuador: shiringochy (Siona). Peru: moena rosada, moenilla, roble, sacha muena, tinchi. Brazil: canela amarella, canela frade, canela garuva, canela paluda. Bolivia: laurel. Paraguay: laurel aguacate, laurel moroti.

As circumscribed here, Endlicheria paniculata accommodates considerable variation in leaf size and

shape, but more disturbing to any sense of intraspe- cific uniformity are the different manifestations of vestiture. The type material and most collections of

this species are from the Atlantic coastal forests of SE Brazil where one usually finds rusty tomentose indument covering the lower surfaces of narrowly ovate leaves. However, this indument can appear with broadly ovate (e.g., Araujo 6776), or even obovate leaves (e.g., Heringer et al. 3187). Similarly irrespective of leaf size and shape, a subsericeous vestiture of pale appressed hairs, e.g., Sellow 433 (type of E. hirsuta var. latifolia) and Glaziou 3092 (type of E. hirsuta var. glabrata), or a hirsute cover of longer, more stiffly erect, dark red hairs (e.g., Rambo 45269 and Brade 8195), are both found in SE Brazil.

These vegetative instabilities are repeated throughout the range of E. paniculata. The typical form, with rusty tomentose ovate leaves, is widespread, ranging from SE Brazil through lower montane Andean slopes from Bolivia (e.g., Solomon 12948) to Venezuela (e.g., Steyermark & Espinoza 110275). The other variants are also widespread. Sub- sericeous indument reappears with ovate leaves in NW South America, including the type of E. racemosa, and ranges into Panama, while in western Ama- zonia it combines with obovate leaves in the type of E. longifolia. Likewise, hirsute indument reappears in western Amazonia with either ovate (e.g., Schunke 1617) or obovate leaves (e.g., Poeppig 2298, the type of E. poeppigii). However, as in SE Brazil, tomentose forms that are otherwise indistinguishable from these subsericeous and hirsute forms are found in sympatry. Flowers also vary in E. paniculata, but only slightly and not in correlation with indument. All are rotate with horizontally spreading tepals, stipitate whorl I and II stamens, and broadly stipitate whorl III stamens with relatively large globose glands, but the anther apices in whorl I and II may be truncate or emarginate. The latter condition may combine with hornlike lobes persisting above each locule, as was indicated as diagnostic for E. boliviensis (Kostermans, 1937), and appears throughout the range of E. paniculata. In Mexia 6329, staminate flowers are provided with four whorls of stamens (i.e., whorl IV members are fertile). This remarkable situation appears to be an abnormal- ity never repeated in the species, nor elsewhere in Endlicheria.

Despite the internal variation, E. paniculata is easily recognized by its pinnate venation since other species with rotate flowers and stipitate stamens with truncate anther apices, E. acuminata and E. gracilis, have triplinerved leaves.

57. Endlicheria gracilis Kosterm., Recueil Trav. Bot. NMerl. 34: 528. 1937. Type. Colombia. San-


tander: Vicinity of Barranca Bermeja, Magdalena Valley, between Sogamoso and Colorado Rivers, 100-500 m, 14 Nov 1934 (fl Y), Haught 1414 (holotype: U; isotypes: A, G).

Endlicheria grisea Kosterm., Reinwardtia 6: 285. 1962. Type. Guyana. Essequibo: Wabuwak, Kanuku Mtns., ca. 650 m, Oct 1948 (fl Y), Wilson-Browne 454 (holotype: K; isotype: NY).

Trees, or shrubs, to 7 m. Branchlets slender, midway along flush 2-3 mm diam., distally weakly angular, soon terete, rusty tomentose, the surface barely visible or exposed, the hairs short, to 0.3 mm, straight to crooked, erect to ascending; terminal buds slender, 1 X 0.6 mm, densely pubescent, the hairs as on branchlets, ascending. Leaves alternate, widely and evenly spaced along current flush; petioles slender, to 1.3 x 0.2 cm, terete, the indument as on branchlets; laminae chartaceous, plane to subbullate, narrowly to broadly ovate, 5-10 X 1.5-5 cm, the base obtuse to acute, briefly decurrent, the apex acute, acuminate for up to 1.5 cm, the margins minutely recurved throughout; upper surface dull grey, minutely punctulate, the midrib and secondaries prominulous, the higher-order venation immersed; lower surface sparsely pubescent, the hairs erect to ascending, slightly denser on main veins, all vein orders raised, their prominence decreasing with rank; secondary veins 3-(4) per side, the lowermost pair subopposite shortly above the leaf base, ascending at 50-60?, arcuate, upper pairs more obtuse, emerging around or beyond midlamina, abruptly ascending after midcourse, loop-connected; tertiaries roughly horizontal, between secondaries straight or forked. Staminate inflorescences evenly spaced along current flush in the axils of foliage leaves, to 3 cm long with 4 lateral branches, branch orders 2-3, the highest order dichasial, lax, the flowers distant, the axes sparsely pubescent, the hairs as on branchlets; bracts and bracteoles caducous by anthesis, lanceolate, densely rusty pannose; pedicels terete, to 2 mm long, those supporting secondary flowers slightly shorter. Flowers rotate, 3 mm diam., sparsely pubescent outside, the hairs appressed to ascending, silvery grey; receptacle broadly infundibuliform, 0.3 X 1.3 mm, densely rusty tomentose inside. Tepals chartaceous, ovate, 0.6 X 0.5 mm, spreading to recurved at anthesis, the inner surface sparsely silvery grey-strigose. Stamens of whorls I and II stipitate, 0.5 mm tall, the anthers transversely oblong, 0.3 x 0.5 mm, glabrous, the apex truncate to emarginate, the connectives level with or reduced between the 2 locelli, these suborbicular, introrse, the filaments laminar, much narrower than anthers, sparsely grey- strigillose; whorl III stamens broadly stipitate, 0.5

mm tall, the anthers depressed-oblong, 0.3 X 0.5 mm, erect, locelli 2, extrorse-latrorse, the filaments slightly narrower than anthers, clavate, densely grey- strigillose inside, the basal glands sessile, globose, relatively large, filling the space between filaments of the inner and outer whorls; whorl IV wanting; pistillode wanting. Pistillate inflorescence with indument, color, and branching as in staminate plants, the flowers similar in size and shape; stamens sterile?, dimensions as in staminate flowers; ovary glabrous, ovoid; style stout, weakly distinguished from ovary; stigma tri-lobed, 0.3 mm diam. Fruits borne on slender clav- ifonn pedicels of up to 1.5 X 0.4 cm; cupules shallowly hemispherical to patelliform, to 7 mm diam., glabrous, the margins undulate; drupes ovoid, to 2 X 1.3 cm.

Distribution (Fig. 43) and ecology. Small trees or shrubs from western Amazonia and adjacent lower montane slopes from Peru to Venezuela and east to the Guianas and NE Brazil at ca. 100-1100 m. Ap- parently absent in the Amazon basin although known to occur in both flooded and well-drained soils. Most flowering specimens collected from September through November, but also in January and May. Fruits available year round.

Representative specimens examined. COLOMBIA. AMAZONAS: Leticia, Parque Nacional Natural Amacayacu, 120 m, 30 Mar 1992 (st), Rudas et al. 4057 (MO). ANTiO- QUIA: San Luis, Vereda Berlin, 400-530 m, 26 Oct 1989 (fr), Cogollo & Ram(rez 4361 (MO).

VENEZUELA. AMAZONAS: Atabapo, camino entre Culebra y la falda del extremo norte del Cerro Duida, 220 m, 9 Feb 1982 (fr), Steyermark et al. 126285 (HBG, MO, NY); Culebra Savanna, 150 m, 14 Nov 1950 (fl Y), Maguire et al. 29436 (F, G, GH, MO, VEN). BOLiVAR: Quebrada 0- paru-ma, 1065-1220 m, 20-21 Nov 1944 (fl i, fr), Stey- ermark 60428 (F); Selva de Oparuma, Kavanay6n, 30 May 1946 (fl 6), Lasser 1881 (F, US).

GUYANA. ESSEQUIBO: Rupununi, Kuyuwini Landing, 250-350 m, 25 Oct 1992 (fl Y), Jansen-Jacobs et al. 3085 (B, MO, NY, U); Potaro-Siparuni, Iwokrama Rainforest Re- serve, 8-16 Jul 1997 (st), Chanderbali et al. 250 (MO).

SURINAME. Lucie R., small granitic islands in river near confluence of Oost R., 225 m, 12 Sep 1963 (fl Y), Irwin et al. 55631 (F, NY, S, U, US).

ECUADOR. NAPO: Tena, Estaci6n Biol6gica Jatun Sa- cha, 400 m, 3-6 Sep 1989 (fr), Palacios 4386 (G, MO, NY). PASTAZA: UNOCAL petroleum exploration well site "Ma- zaramu," 390 m, Apr 1990 (fr), Beck et al. 1054 (MO). Suc. UMBiOS: Gonzalo Pizarro, 30 km NE of Lago Agrio, 1050 m, 23 Mar 1990 (fr), Cer6n et al. 9226 (MO); Lago Agrio, Reserva Faunistica Cuyabeno, 230 m, 13 Nov 1991 (fl Y), Palacios et al. 8840 (HBG, MO, NY). ZAMORA-

CHINCHIPE: Nangaritze, Miazi, 900-1000 m, 21 Oct 1991 (fl ?, fr), Palacios et al. 8585 (MO, NY); Zamora, Parque


I

.7

C mV. .,,. ...-.w,s.+W ^i ;E'--, --5W1z, r.

FIG. 43. Distribution of Endlichenia gracilis anld E. acuminata.

Nacional Podocarpus, Guarderna Rio Bombuscaro, sendero al Mirador, 1100 m, Jan 1995 (fl 9), Palacios & lirado 13298 (MO).

PERU. AMAZONAS: Condorcanqui, El Cenepa, Comu- nidad de Tutino, 500 m, 20 Jul 1997 (fr), Rojas et al. 91 (MO). LoRETo: Alto Amazonas, Rfo Marant6n, Pongo de Manseriche, Cerros Campanquiz, 500-550 m, 19-21 Oct 1962 (fl d), Wurdack 2339 (F, G, GH, K, NY, S, UC); Rfo Huallaga, Shucushuyacu, 250 m, 14 Sep 1981 (fr), Vasques & Jaramillo 2501 (F, MO, NY). MADRE DE Dios: Manu, Parque Nacional Manut, Cocha Cashu Station, Rio Manu, 250 m, 3 Sep 1989 (fl 9), Foster & Beltrdn 13050 (MO).

BRAZIL. ACRE: Brasileia, Seringal Porongaba, Colo- cagko Sao Jose, 30 May 1991 (fl 9, fr), Daly et al. 6795 (MO, NY). AMAPA: Rio Agauaria, 9 Oct 1961 (fl d), Pires et al. 51599 (COL, G, GH, NY, US); Mun. Oiapoque, BR 156 between Cal,oene and Oiapoque, 17 km SSE of Oia- poque, 3 Dec 1984 (fr), Mori et al. 17170 (MO).MARANHAo: Mun. Monqao, basin of Rfo Tariaqu, Ka'apor Indian Reserve, 7 km from Urutawy, 5 Jun 1985 (fr), BalMe 988 (MO).

Local names and uses. Venezuela: muneu-yek. Brazil: mahamira, aiju'ywahu. According to W. L.

Balee, used as a hunting charm by the Ka'apor Indi- ans of NE Brazil. The leaves are rubbed on the head of a hunter to help him kill maha deer.

Endlicheria gracilis is immediately recognized by its rotate flowers and the minutely punctulate upper surface of triplinerved ovate leaves that assume a slate grey color in the dried state. As noted by Kostermans (1937), the androecium is remarkably well-developed in pistillate flowers with stamens as large as those in staminate flowers and anther valves often dehisced. Further, in at least one collection, Jansen-Jacobs 3085, pollen is present inside locelli although the pistil is clearly fertilized and developing toward fruit. As staminate flowers lack a pistillode altogether, it seems likely that E. gracilis is androdioecious rather than dioecious. Still, an ITS sequence taken from Chan- derbali et al. 250 assigns this species to the Endli- cheria-Rhodostemonodaphne clade (Fig. 2), wherein, as so far known, other species are dioecious. Therein its affinities are unclear. The androecium of E. gracilis is similar to that of E. paniculata and its allies, but


the slender claviform pedicels and shallow cupules with undulate margins are reminiscent of E. sprucei and E. longicaudata, species here compared with members of Rhodostemonodaphne. The appearance of remarkably similar leaves and fruits in R. parvifolia also seems to support affinities with Rhodostemono- daphne. Were it not for coriaceous leaves with smooth upper surface in R. parvifolia, the two species would be separable only in flower, themselves so similar in external appearance that it would be necessary to note differences in stamen shape and locelli number.

58. Endlicheria acuminata Kosterm., Recueil Trav. Bot. Neerl. 34: 553. 1937. Type. Brazil. Ama- zonas: Lago Jurua Miry, Jun 1901 (fl d), Ule 5581 (holotype: L; isotypes: G, HBG, K).

Trees to 15 m. Branchlets slender, midway along flush 2-3 mm diam., angular, sparsely pubescent, the surface barely visible to clearly exposed, the hairs silvery-grey to pale brown, short, to 0.3 mm, straight to crooked, erect or appressed; terminal buds slender, 3 X 1 mm, densely rusty pubescent, the hairs as on branchlets, ascending. Leaves alternate, widely and evenly spaced along current flush; petioles slender, to 1 X 0.2 cm, semi-terete, the indument as on branchlets; laminae membranaceous to chartaceous, plane, ovate to ovate-elliptic, 5-15 X 2-4 cm, the base obtuse to acute, briefly decurrent, the apex acute, acuminate for up to 1.5 cm, the margins minutely recurved throughout, or flat after midlamina; upper surface dull greyish green, waxy, the midrib and secondaries immersed to sunken, the higher-order venation raised; lower surface sparsely pubescent, the hairs erect or appressed, uniformly distributed, all vein orders raised, their prominence decreasing with rank; secondary veins 2-4 per side, the lowermost pair subopposite shortly above the leaf base, ascending at 50-60?, almost straight to arcuate, other pairs emerging around or beyond midlamina, more obtuse, arcuate, weakly loop-connected; tertiaries roughly horizontal, between secondaries once-forked to straight. Staminate inflorescences evenly spaced along current flush in the axils of foliage leaves, to 5 cm long with 3 lateral branches, branch orders 2-3, lateral second-order branches often reduced to a single flower, internodes of terminal cymes reduced, the flowers clustered in pseudo-umbels, the axes sparsely pubescent, the hairs 0.3 mm long, greyish, appressed to ascending; bracts and bracteoles caducous by anthesis, lanceolate, the indument as on axes; pedicels terete, to 2 mm long, those supporting secondary

flowers slightly shorter. Flowers rotate, 3 mm diam., sparsely grey-strigillose outside; receptacle cyathiform, 0.5 X 1 mm, densely silvery grey-tomentose inside. Tepals membranaceous to chartaceous, ovate, 1 X 0.7 mm, spreading at anthesis, the inner surface glabrous. Stamens of whorls I and II stipitate, 0.5 mm tall, the anthers transversely oblong, 0.3 X 0.5 mm, glabrous, the apex truncate, the connectives level with the 2 locelli, these suborbicular, introrse, the filaments laminar, much narrower than anthers, glabrous; whorl III stamens sessile, 0.6 mm tall, the anthers depressed- oblong, 0.3 X 0.4 mm, erect, locelli 2, extrorse- latrorse, the filaments broader than anthers, widening further towards base, laminar, glabrous, the basal glands sessile, globose, relatively large, filling the space between filaments; whorl IV wanting; pistillode wanting. Pistillate inflorescence with indument, color, and branching as in staminate plants, the flowers similar in size and shape; stamens sterile, smaller; ovary glabrous, ovoid; style stout, weakly distinguished from ovary; stigma broadly tri-lobed, 0.5 mm diam. Fruits borne on slender cylindrical pedicels of up to 1.3 X 0.3 cm; cupules patelliform or shallowly hemispherical, to 0.5 X 1 cm, glabrous outside, sericeous inside, the margins entire; drupes ellipsoid, to 3.5 x 1.5 cm.

Distribution (Fig. 43) and ecology. Small trees of flooded forests in western Amazonia at ca. 100- 300 m. Flowering from April to August, fruits available year round.

Representative specimens examined. COLOMBIA. VAUPES: San Jose del Guaviare, La Libertad, cerca a Re- serva Indfgena "Asunci6n," Canio Grande, 300 m, 10 Aug 1989 (fl d), Marulanda & Mdrquez 1108 (HUA 89052), Cano Nare, 280 m, 17 Aug 1989 (fl ?, fr), Marulanda & Ma'rquez 1536 (HUA).

ECUADOR. NAPO: Aguarico, Reserva Faunfstica Cuy- abeno, Laguna Zancudo Cocha (Iripari), 230 m, 28 Sep 1991 (fl 5), Palacios et al. 7804 (MO); Lagunas de Cuyabeno, 300 m, 21 Aug 1981 (fl 5, juv), Brandbyge et al. 33861 (AAU). SUCUMBiOS: Lago Agrio, Reserva Faunfstica Cuy- abeno, 230 m, 30 Sep 1991 (fl Y), Palacios et al. 7884

(MO). PERU. LORETO: Maynas, Rfo Amazonas, Yanamono

Explorama Tourist Camp, halfway between Indiana and mouth of Rfo Napo, 140 m, 26 Jul 1991 (fl 5), Vdsquez & Grdndez 17475 (MO); Requena, Sapuena, Arboreto Jenaro Herrera, Canio Lobito, 160 m, 17 Aug 1994 (fl 5), Ortiz et al. 124 (MO). MADRE DE Dios: Puerto Maldonado, Rfo Madre de Dios, 250 m, 22 Apr 1977 (fl Y), Gentry et al. 19636 (MO, NY); Tambopata, Cuzco Amaz6nico, Fundo Concepci6n, Rfo Madre de Dios, 200 m, 19 May 1989 (fl 5 ), Phillips & Ntiuiez 52 (MO).


BRAZIL. ACRE: Cruzeiro do Sul, Rio Jurua & Rio Moa, near Uruburetama, 25 May 1971 (fl d), Maas et al. P13307 (HBG, INPA, NY); Rio Muru, Seringal Vit6ria Velha, Colocagao Barro Vermelho, 20 Jun 1995 (fl d), Fi- gueiredo et al. 908 (MO); Senador Guiomard, basin of Rio Purus, Rio Iquiri, 6 Mar 1997 (fr), Daly 9293 (MO). AMA-

ZONAS: Rio PurUs, between Campina and Tambaqui, Jun 1971 (fl 3), Prance et al. 13391 (HBG, INPA, NY); Lago Preto, 2 km N of Labrea, 25 Jun 1971 (fl Y), Prance et al. 13695 (HBG, INPA, K, MO, NY, US).

BOLIVIA. BENI: Marban, parque Isiboro-Secure, de Puerto San Lorenzo, 200 m, 25 May 1992 (fr), Seidel et al. 6557 (MO); Yacuma, E of San Bora, Bosques de Chimanes, near Rfo Mozeruma, 250 m, Nov 1988 (fr), Foster et al. 12483 (MO).

Local names. Peru: cunchi moena, muena. Brazil: louro, louro do igapo, louro preto do igap6.

Endlicheria acuminata shares triplinerved ovate leaves with E. gracilis. In both as well, the flowers are rotate, stipitate whorl I and II stamens have transversely oblong anthers with truncate apices, and large basal glands completely fill the space between filaments of all staminal whorls. Yet the two are not easily confused because the upper leaf surface in E. acuminata is smooth and waxy rather than minutely punctulate, and tertiaries are raised rather than immersed.

As in Endlicheria anomala and E. paniculata, either erect or appressed hairs constitute the vestiture of branchlets and lower leaf surfaces of E. acuminata without accompanying differences. Type material and a few other specimens are tomentose, but in most the hairs are definitely appressed. This subsericeous form of E. acuminata is vegetatively indistinguishable from E. krukovii, where, however, the flowers are infundibuliform, all anthers are ovate with acuminate apices, and basal glands are minute and inconspicuous. As cupules in the two are also indistinguishable, both being shallowly hemispherical and densely sericeous inside, only staminodes remnant on cupule rims can assign fruiting specimens to species.

59. Endlicheria krukovii (A. C. Sm.) Kosterm., Re- cueil Trav. Bot. Neerl. 34: 531. 1937. Aniba krukovii A. C. Sm., Phytologia 1(3): 117. 1935. Type. Brazil. Amazonas: Near mouth of Rio Embira (tributary of Rio Taracau), 26 Jun 1933 (fl Y), Krukoff 5023 (holotype: NY; isotypes: A-n.v., G, K).

Trees, 3-22 m. Branchlets slender, midway along flush 3-4 mm diam., angular, densely strigose, the

surface obscured or barely visible, the hairs short, to 0.3 mm, straight, appressed, grey to light brown; terminal buds slender, 3 X 0.6 mm, densely pubescent, the hairs as on branchlets, ascending. Leaves alternate, widely and evenly spaced along current flush; petioles slender, to 2 x 0.2 cm, semi-terete, the indument as on branchlets; laminae chartaceous to membranaceous, plane, ovate to obovate, 10-25 X 5- 10 cm, the base acute, briefly decurrent, the apex acute, acuminate for up to 3 cm, the margins minutely recurved throughout, or flat beyond midlamina; upper surface dull greyish green to olive-brown, waxy, the midrib and secondaries sunken to immersed, the higher-order venation raised; lower surface sparsely pubescent, the hairs as on branchlets, appressed, uniformly distributed, all vein orders raised, their prominence decreasing with rank; secondary veins 2-4 per side, the lowermost pair subopposite shortly above the leaf base, ascending at 50-600, arcuate to almost straight, other pairs emerging around or beyond midlamina, more obtuse, arcuate, weakly loop-connected; tertiaries roughly horizontal, between secondaries once-forked to straight. Staminate inflorescences evenly spaced along current flush in the axils of foliage leaves, to 20 cm long with 12 lateral branches, branch orders 3-4, the highest order dichasial, lax, the flowers distant, the axes densely pubescent, the hairs appressed, increasing in density and obscuring epidermis distally; bracts and bracteoles caducous by anthesis, lanceolate, the indument as on axes; pedicels terete, to 1 mm long, those supporting secondary flowers slightly shorter. Flowers infundibuliform, 2 mm diam., densely silvery-grey pubescent outside, the hairs appressed; receptacle infundibuliform, 0.6 X 0.6 mm, glabrous inside. Tepals membranaceous to chartaceous, broadly ovate, 0.6 x 0.6 mm, ascending at anthesis, the inner surface glabrous, the margins and apex inside minutely papillose. Stamens of whorls I and II stipitate, 0.6 mm tall, the anthers ovate, 0.4 x 0.3 mm, glabrous, the apex apiculate, the connectives prolonged between the 2 locelli, these obliquely hemispherical, introrse-latrorse, the filaments laminar, much narrower than anthers, glabrous; whorl III stamens sessile, 0.6 mm tall, the anthers ovate, 0.4 x 0.3 mm, erect, locelli 2, extrorse-latrorse, the filaments broader than anthers, wider towards base, glabrous, the basal glands sessile, minute, globose-apiculate; whorl IV wanting; pistillode wanting. Pistillate inflorescence with indument and color as in staminate plants, but shorter and with fewer lateral branches, the flowers slightly deeper; stamens sterile, smaller; ovary glabrous, ovoid; style stout,


weakly distinguished from ovary; stigma strongly tri- lobed, 0.6 mm diam. Fruits borne on slender cylindrical pedicels of up to 1 X 0.3 cm; cupules patelliform to shallowly hemispherical, to 0.5 X 1 cm, glabrous outside, sericeous inside, the margins entire; drupes ellipsoid, to 2.5 x 1.3 cm.

Distribution (Fig. 44) and ecology. Small to medium-sized trees in nonflooded forests of the eastern Andean foothills but also in seasonally inundated lowlands in western Amazonia at ca. 200-1000 m. Flowering specimens collected from February through May, and in October and November; fruits available year round.

Representative specimens examined. ECUADOR. NAPO: Estaci6n Experimental INIAP-Napo, Payamino, Re- serva Floristica "El Chuncho," 280 m, 17 May 1986 (fr), Baker 7017 (QAME-n.v., QCNE-n.v., MO, NY); Santa Ce- cilia, 340 m, 30 Mar 1972 (fl d, juv), Dwyer & Simmons 9748 (MO). PASTAZA: Pastaza, 365 m, Oct 1-20 1990 (fl d), Espinoza & Coba 369 (MO). SucuMsios: Gonzalo Pi- zarro, Bosque Protector Los Cedros, cuenca del Rio Tigre,

1000 m, 17 Mar 1992 (fl c), lipaz et al. 709 (MO); Lago Agrio, Reserva Faunfstica Cuyabeno, 230 m, 20 Oct 1991 (fl 9), Palacios et al. 8027 (MO).

PERU. AMAZONAS: Bagua, Imaza, Comunidad de Ya- mayakat, 600 m, 9 Jun 1997 (fl 6), Vdsquez et al. 23948 (MO); Rfo Santiago, Quebrada Kusu', Bagua, 250 m, 30-31 Oct 1962 (fl 6), Wurdack 2493 (K, NY, S, UC, US). Cuzco: Rio Mapituriani, La Convenci6n, 14 Sep 1976 (fr), Was- shausen & Encarnaci6n 649 (K, MO, NY, US). HukNUco: Pachitea, Honoria, Quebrada de Macuya al oeste del Rio Pachitea, 300-400 m, 6 May 1968 (fl d), Schunke 2570 (MO). LORETO: Alto Amazonas, Puerto Melendez, below Pongo de Manseriche, s.d. (fl 6), Tessmann 4734 (NY); Maynas, Iquitos, km 44 carretera Iquitos-Nauta, terreneos del Comite de Reforestaci6n Iquitos (CRI), 150 m, 14 Mar 1989 (fr), Vdsquez et aL 11928 (HBG, MO). MADRE DE

DIos: Manu, Parque Nacional Manu', Cocha Cashu Station, Rio Manu, 350 m, 1 Feb 1981 (fr), Foster & Wright 8254 (MO); Tambopata, Cuzco Amaz6nico, trail to Lago San- doval across Rio Madre de Dios, ca. 12 km E of Puerto Maldonado, 200 m, 21 Feb 1990 (fl T, juv), Gentry & Nufnez 69392 (MO). PAsco: Oxapampa, Palcazt, 300-600 m, 7 Dec 1984 (fr), Hartshorn et al. 2689 (MO, NY, U). SAN

4-a ~ ~~~~~~~-

Air

FIG. 44. Distribution of Endlichenia krukovii and E. nilssonii.

DOUBTFUL NAMES AND EXCLUDED TAXA 123

MARTiN: Huinguillo, 500-600 m, 21 Mar 1962 (fl d, juv), Woytkowski 7179 (HBG, MO); San Martin, 5-15 km E of Shapaja on rd. to Chazuta, 200-300 m, 9 Apr 1986 (fl d), Knapp & Mallet 7024 (F, MO, NY, US). UCAYALI: Coronel Portillo, Yarinacocha, Nueva Esperanza de Panaillo, 148 m, 1 Apr 1988 (fl d, juv), Vdsquez & Jaramillo 10458 (F, MO, NY).

BRAZIL. ACRE: Tarauaca, 26 May 1977 (fl d), Mir- anda 20 (INPA); Xapuri, Rio Acre, 10 Nov 1991 (fr), Daly et al. 7282 (MO, NY).

BOLIVIA. PANDO: Manupiri, 35 km al N de Puerto America, 200 m, 30 Apr 1994 (fl d), Jardim 594 (MO).

Local names. Ecuador: ajua. Peru: moenilla, tinchi, yuwich (Huambisa). Brazil: louro.

Among species with triplinerved leaves, Endli- cheria krukovii is conspicuous for its relatively mul- tiflorous inflorescence with densely pubescent infundibuliform flowers within which all stamens have ovate anthers. Yet, from representatives of E. acuminata with similar appressed indument, only floral characters are distinguishing. Although E. acuminata has been found only in flooded habitat, and most collectors have indicated terra firme conditions for E. krukovii, any ecological distinction seems weak since occasional reports, e.g., Vdsquez et al. 23948, suggest that E. krukovii also tolerates inundation.

60. Endlicheria nilssonii C. K. Allen, Mem. New York Bot. Gard. 10(5): 66. 1964. Type. Venezuela. Bolivar: Wooded ridge, La Danta, along provi- sional road from km 125 to 127 between Leupa and Cerro Venamo, 1200 m, 17 Apr 1960 (fr), Steyermark & Nilsson 255 (holotype: NY).

Trees to 30 m. Branchlets stout, midway along flush 3-5 cm diam., angular, sericeous, the surface concealed by the indument cover, trichomes short, to 0.1 mm, straight, appressed, light yellowish green; terminal buds plump, 4 X 3 mm, sericeous. Leaves alternate, widely and evenly spaced along current flush; petioles robust, to 1.5 X 0.3 cm, striate, the indument as on branchlets; laminae coriaceous, plane, ovate to elliptic, 10-15 X 4-6 cm, the base acute, briefly decurrent, the apex acute, acuminate for up to 0.5 cm, the margins minutely recurved throughout; upper surface deep green, waxy, the midrib and secondary veins immersed, the tertiaries prominulous; lower surface sparsely pubescent, the hairs as on branchlets, appressed, uniformly distributed, all vein orders raised, their prominence decreasing with rank; secondary veins 3-4 per side, ? evenly spaced, slightly more distant around midlamina, ascending at 45-60? (more acutely towards apex), arcuate, distal

pairs loop-connected, the tertiaries roughly horizontal, between secondaries straight to forked. Staminate and pistillate inflorescences unknown. Fruits borne on claviform pedicels of up to 1 X 0.4 cm; cupules hemispherical, to 0.5 X 1 cm, glabrous outside and inside, the margins entire; drupes ellipsoid, to 1 X 0.7.

Distribution (Fig. 44) and ecology. Known only from the type material, taken from a tree found fruiting in April at ca. 1200 m in the highlands of eastern Venezuela.

Endlicheria nilssonii is a poorly understood yet distinctive species. The combination of densely sericeous branchlets but only sparse appressed hairs on the leaves below is otherwise found only occasionally in E. metallica. Flowers are not known, but one staminode found persisting in fruit shows a distinct and densely pubescent filament and an ovate anther with a truncate apex. However, a tepal, also found on the cupule margin, provides a better appreciation of the flowers of E. nilssonii. This tepal is rather large and fleshy but, more important, densely pubescent with papillose hairs covering the inner surface. As such tepals only occur with rotate flowers (e.g., E. sericea), it is most likely that flowers of E. nilssonii are rotate. Even if not the case, the combination of pubescent fleshy tepals, densely sericeous branchlets, and sparse appressed hairs on the leaves below is unique in En- dlicheria.

In the truncate anther apices, pinnate venation, and sparse appressed indument on leaves below, this species approaches to subsericeous forms of E. paniculata, with which it is placed, with hesitation. Better material may later more clearly indicate its affinities.

DOUBTFUL NAMES AND EXCLUDED TAXA

Endlicheria balsamea Vattimo, Rodriguesia 33 (46): 23. 1978. Type. Brazil. Rio de Janeiro: 14 Jan 1932 (fr), Paulino & Victorio s.n. R 148881 (holotype: R; isotypes: R [7 sheets]).

The protologue describes a representative of Aiouea, since it states that nine two-locellate stamens and well-developed whorl IV staminodes occur together with fruits. However, I could not find stamens or staminodes in the ample type material before me, and I therefore refrain from proposing a new combination.

Endlicheria debilis Kosterm., Recueil Trav. Bot. Neerl. 34: 555. 1937. Type. Peru. Loreto: Alto


Amazonas, Balsa Puerto (lower Rio Huallaga basin), 150-350 m, 28-30 Aug 1929 (fr), Killip & Smith 28400 (holotype: NY; isotypes: B-n.v., F, US) = Rhodostemonodaphne debilis (Kosterm.) Chanderbali comb. nov.

Kostermans apparently overlooked the four- locellate staminodes on the isotype deposited at F, and found before me by V. Koch on the isotype at US. Those from androecial whorls I and II are narrowly stipitate with obovate anthers with the locelli arranged in a very shallow, almost horizontal, arc, and the whorl III staminode is sessile with four locelli in two superimposed pairs. These staminodes assign Killip & Smith 28400 to Rhodostemonodaphne, wherein R. crenaticupula is an excellent vegetative match. The latter differs in that whorl I and II stamens in both staminate and pistillate plants are larger and more sessile and cupule margins more strongly lobed.

Endlicheria endlicheriopsis (Mez) Kosterm., Meded. Bot. Mus. Herb. Rijks Univ. Utrecht 25: 43. 1936. Type. French Guiana, Melinon 605 (holotype: P-n.v.) = Ocotea endlicheriopsis Mez.

Endlicheria goeldiana Vattimo, An. XV Congr. Soc. Bot. Bras., 169. 1964. Type. Brazil. Amazonas: Rio Purus, 20 Jun 1903 (fl), Goeldi s.n. HAMP 3902 (holotype: R).

The type material available to me consists of detached leaves. As there is no sign of the flowers described in the protologue, I cannot confirm or deny that this material belongs to Endlicheria.

Endlicheria grandis Mez, Jahrb. Konigl. Bot. Gart. Berlin 5: 124. 1889. Type. French Guiana. Meli- non s.n. (holotype: B-n.v.; isotypes: NY-n.v. P- n.v.) = Rhodostemonodaphne grandis (Mez) Roh- wer.

Endlicheria impressa (Meisn.) Mez, Jahrb. Konigl. Bot. Gart. Berlin 5: 132. 1889. Phoebe impressa Meisn., in DC., Prodr. 15(1): 33. 1864. Type. French Guiana. Patris s.n. (holotype: G-n.v.) =

Aiouea impressa (Meisn.) Kosterm.

Endlicheria juruensis (A. C. Sm.) Kosterm., Recueil Trav. Bot. Neerl. 34: 542. 1937. Aniba juruensis A. C. Sm., Phytologia 1(3): 116. 1935. Type. Bra-

zil. Amazonas: Near mouth of Rio Embira (tributary of Rio Tarauaca), Jun 1933 (fl 9, fr juv), Krukoff 4775 (holotype: NY; isotypes: A, F, G, K, MO, U) = Rhodostemonodaphnejuruensis (A. C. Sm.) Chanderbali comb. nov.

Staminodes in the type material are four-locellate with the locelli arranged in the shallow apical arch typical of Rhodostemonodaphne. Other characters as- sociating this species with Rhodostemonodaphne include sessile stamens and dense papillosity in the inner surface of horizontally spreading tepals. After the type collection, several more pistillate representatives have been found (listed below), but staminate plants are still unknown.

Additional specimens examined. ECUADOR. CAR-

CHI: Tulcan, Parroqui Chical, Reserva Indigena Awa, 1200 m, 23-27 May 1992 (fl 9), Tipaz et al. 1104 (MO), 900 m, May 1992 (fl Y), Quelal et al. 704 (MO). ESMERALDAS:

San Lorenzo, carretera Lita-San Lorenzo, 500 m, 9 Jul 1990 (fr), Rubio et al. 454 (MO). MORONA-SANTIAGO: Along new rd. Mendez-Morona, 650 m, 16 Aug 1989 (fl Y), van der Werff & Gudinio 11136 (MO).

PERU. SAN MARTiN: Mariscal Caceres, Tocache Nuevo, Camino a Santa Rosa, Rfo Mishollo, 350-370 m, 5 Aug 1973 (fr), Schunke 6726 (F, MO, NY).

BOLIVIA. LA PAZ: Prov. Franz Tamayo, Serranfa de Chepite, 700 m, 3-8 Apr 1992 (fl Y, fr), Killeen 3832 (MO).

Endlicheria loretensis 0. Schmidt, Repert. Spec. Nov. Regni. Veg. 31. 178. 1933. Type. Peru. Lor- eto: Mishuyacu, near Iquitos, 100 m, Apr 1930 (fl J), Klug 1258 (holotype: F-n.v.; photo neg. 40486 ex F) = Ocotea or Rhodostemonodaphne.

Endlicheria maguireana C. K. Allen, Mem. New York Bot. Gard. 10(5): 68. 1964. Type. Venezuela. Amazonas: RioYatua, 1100-1150 m, 26 Dec 1957 (fl), Maguire et al. 42521 (holotype: NY; isotypes: K-n.v., MO, S, U, US-n.v.) = Aiouea maguireana (C. K. Allen) Renner.

Endlicheria tomentella Mez, Jahrb. Konigl. Bot. Gart. Berlin 5: 115. 1889. Type. Peru. Ancach: Near Moro, Pearce s.n. (holotype: K-n.v.) =

Aiouea tomentella (Mez) Renner.

Endlicheria zapoteoides Lundell, Wrightia 1: 145. Type. Mexico. Chiapas: Cascada near Siltepec, 1600 m, 1 Mar 1945 (fl, fr), Matuda 5153 (holo-

LITERATURE CITED 125

type: TEX-n.v.; isotypes: MO, US-n.v.) =

Beilschmiedia zapoteoides (Lundell) Kosterm.

Goeppertia argentea (Griseb.) Meisn., in DC., Prodr. 15(1). 174. 1864. Aydendron argenteum Griseb., Fl. Brit. W. Ind. Isl. 1: 285. 1860. Type. Dominica, Imray 365 (holotype: GOET-n.v.) = Aniba bracteata (Nees) Mez.

Goeppertia caudata Meisn., in DC., Prodr. 15(1). 175. 1864. Syntypes. Brazil. Amazonas: Prov. Rio Negro, prope Panure ad Rio Vaupes, Spruce 2638 (syntypes: K-n.v., M-n.v.) = Ocotea debilis Mez.

Goeppertia geminiflora Meisn., in DC. Prodr. 15(1). 175. 1864. = Systemonodaphne geminiflora (Meisn.) Mez, provided Martin s.n. from French Guiana is cited as the type. = Acrodiclidium geminiflorum (Meisn.) Mez, provided Guillemin 231 from Rio de Janeiro, Brazil, is cited as the type.

ACKNOWLEDGMENTS This study was undertaken as part of my doctoral

dissertation research at the University of Missouri-St. Louis and the Missouri Botanical Garden. I am in-

debted to these two institutions for the opportunity to pursue graduate studies in their excellent facilities. For providing unlimited access to his expertise in the Lauraceae, my dissertation advisor, Henk van der Werff, is specially thanked. Grants from the Mallinck- rodt Foundation administered by the International Center for Tropical Ecology, University of Missouri- St. Louis, and from the Smithsonian Institution's Bi- ological Diversity of the Guianas Program, are grate- fully acknowledged for their generous support of the laboratory and field components of this study, respectively.

I thank Jim Solomon, Manager of the Missouri Botanical Garden Herbarium, for requesting loans of Endlicheria and for providing a place in the herbarium where they could be studied in relative luxury. I also thank the curators of institutions that provided loans, sometimes searching for specific specimens among their indets, and allowing me to keep them for the five-year duration of this study.

Barbara Alongi prepared the fine illustrations of the new species of Endlicheria. Mike Vieth helped to take the SEM photographs of stamens. Henk van der Werff corrected the Latin diagnoses and Rosa Ortiz corrected the Spanish abstract. Susanne Renner, Henk van der Werff, and the three reviewers, Mike Nee, William Burger, and Santiago Madriinan, provided useful comments on earlier drafts of the manu- script.

LITERATURE CITED

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Axelrod, D. I. 1975. Evolution and biogeography of Madrean-Tethyan sclerophyll vegetation. Ann. Mis- souri Bot. Gard. 62: 280-334.

Bentham, G. 1880. Laurinae. Pp. 146-165 in G. Ben- tham & J. Hooker (eds.), Genera plantarum. Vol. 3. Reeve & Co., London.

Blattner, F. 1999. Direct amplification of the entire ITS region from poorly preserved plant material using re- combinant PCR. Biotechniques 29: 1180-1186.

Boyle, E. M. 1980. Vascular anatomy of the flower, seed, and fruit of Lindera benzoin. Bull. Torrey Bot. Club. 107: 409-417.

Burger, W. C. 1988. A new genus of Lauraceae from Costa Rica, with comments on problems of generic and specific delimitation within the family. Brittonia 40: 275-282.

Chanderbali, A. S. 1996. Novelties in Guianian Endli- cheria (Lauraceae). Novon 6: 328-334.

, H. van der Werff & S. S. Renner. 2001. Phy-

logeny and historical biogeography of Lauraceae: evidence from the chloroplast and nuclear genomes. Ann. Missouri Bot. Gard. 81: 104-134.

Ducke, A. 1925. Plantes nouvelles ou peu connues de la region amazonienne (Ille Partie). Arch. Jard. Bot. Rio de Janeiro 4: 1-210.

Felsenstein, J. 1985. Confidence limits on phylogenies: an approach using the bootstrap. Evolution 39: 783- 791.

Heo, K., H. van der Werff & H. Tobe. 1998. Embry- ology and relationships of Lauraceae (Laurales). Bot J. Linn. Soc. 126: 295-322.

Kasapligil, B. 1951. Morphological and ontogenetic studies on Umbellularia californica Nutt. and Laurus nobilis L. Univ. Calif. Publ. Bot. 25: 115-240.

Kostermans, A. J. G. H. 1936. Studies in South Amer- ican Malpighiaceae, Lauraceae and Hemandiaceae, especially of Suriname. Meded. Bot. Mus. Utrecht 25: 1-70.

. 1937. Revision of Lauraceae II. The genera En- dlicheria, Cryptocarya (American species) and Li- caria. Recueil Trav. Bot. Neerl. 34: 500-609.

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Kurz, H. 1982. Fortpflanzungsbiologie einiger Gattun- gen neotropischer Lauraceen und Revision der Gat- tung Licaria (Lauraceae). Thesis, University of Ham- burg, Hamburg.

Kuzoff, R., J. Sweere, D. Soltis, P. Soltis & E. Zimmer. 1998. The phylogenetic potential of entire 26S rDNA sequences in plants. Molec. Biol. Evol. 15: 251-263.

Lindley, J. 1836. A natural system of botany. Longman, Rees, Orne, Brown & Green, London.

Madrifiain, S. 1996a. New species of Rhodostemono- daphne (Lauraceae) from northeastern South Amer- ica. Brittonia 48: 45-66.

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NUMERICAL LIST OF TAXA

E. punctulata species group

1. E. punctulata (Mez) C. K. Allen 2. E. coriacea Chanderbali

Microlocellata species group

3. E. bullata Ducke 4. E. rubriflora Mez

5. E. ferruginosa Chanderbali 6. E. longicaudata (Ducke) Kosterm. 7. E. sprucei (Meisn.) Mez

E. metallica species group

8. E. metallica Kosterm. 9. E. chrysovelutina Chanderbali

LIST OF EXSICCATAE 127

E. browniana species group

10. E. browniana Mez 11. E. jefensis van der Werff ex Chanderbali 12. E. colombiana (Meisn.) Mez 13. E. mishuyacensis A. C. Sm. 14. E. pyriformis (Nees) Mez 15. E. formosa A. C. Sm. 16. E. paradoxa Mez 17. E. dysodantha (Ruiz & Pav.) Mez 18. E. tomentosa Chanderbali

Ampelodaphne species group

19. E. aruncifiora (Meisn.) Mez 20. E. arachnocome Chanderbali 21. E. arenosa Chanderbali 22. E. macrophylla (Meisn.) Mez 23. E. multiflora (Miq.) Mez 24. E. levelii C. K. Allen 25. E. dictifarinosa C. K. Allen 26. E. melinonii Benoist 27. E. reflectens (Nees) Mez 28. E. bracteata Mez 29. E. verticillata Mez 30. E. cocuirey Kosterm. 31. E. tessmannii 0. C. Schmidt 32. E. directonervia C. K. Allen 33. E. chalisea Chanderbali 34. E. glomerata Mez

E. anomala species group

35. E. anomala (Nees) Mez 36. E. williamsii 0. C. Schmidt

E. canescens species group

37. E. canescens Chanderbali 38. E. xerampela Chanderbali 39. E. citriodora van der Werif 40. E. rubra Chanderbali 41. E. vinotincta C. K. Allen 42. E. oreocola Chanderbali 43. E. szyszylowiczii Mez 44. E. duotincta Chanderbali 45. E. lorastemon Chanderbali

E. sericea species group

46. E. sericea Nees 47. E. bracteolata (Meisn.) C. K. Allen 48. E. tschudyana (Lasser) Kosterm. 49. E. griseo-sericea Chanderbali 50. E. ruforamula Chanderbali 51. E. aurea Chanderbali 52. E. lhotzkyi (Nees) Mez 53. E. klugii 0. C. Schmidt 54. E. argentea Chanderbali 55. E. robusta (A. C. Sm.) Kosterm.

E. paniculata species group

56. E. paniculata (Spreng.) J. F. Macbr. 57. E. gracilis Kosterm. 58. E. acuminata Kosterm. 59. E. krukovii (A. C. Sm.) Kosterm. 60. E. nilssonii C. K. Allen

LIST OF EXSICCATAE In addition to all collections of Endlicheria examined, this index includes several specimens that were

previously annotated as Endlicheria but belong to other genera.

Abbott, J. R. & L. C. Isaacs, 16484 (56). Acevedo, P., 3432 (33). Acevedo, P. & Cedeino, 7416 (32). Acevedo, P. & F. Ramfrez, 9967 (cf. 59). Acevedo, P. et al., 1370 (cf. 49); 3484 (27); 4834 (1);

4930, 6035 (14); 8646 (15). Adalardo-Oliveira, A., 2643 (35); 2738 (22). Agiular, M. & D. Castro, 1119 (58). Aguiar, I. J. A. et al., RUC-122 (50). Aguirre-Galviz, L. E., 1117 (13). Albuquerque, B., 135, 136 (35). Albuquerque, B. & L. Coelho, 415 (35). Albuquerque, B. & Elias, 67-22 (43). Albuquerque, B. & J. Lima, 240 (35). Albuquerque, B. et al., 570,742,765,778 (35); 850 (13);

955, 1050 (35). Alexiades, M. N. & C. Diaz, 849 (56). Alexiades, M. N. & G. Nicole, 1042 (28).

Alexiades, M. N. et al., 437 (17). Allen, C. K., 20 (48). Allen, C. K. et al., 323 (21). Aluisio, J., 267 (43). Alvarez, E. et al., 482 [Ocotea cernua (Nees) Mez]; 1150

(7). Amaral, I. L. et al., 109 (45); 157 (24); 448 (15); 488

(55); 1375 (15). Ancuash, E., 133 (8); 179 (56); 283 (8); 404 (55); 1343

(32). Andel, T. van et al., 293 (15). Anderson, A., 1521, s.n. (46). Anderson, W. R., 7759, 9604 (56); 12052 (35); 35171

(56). Angeli, C., 587 (56). Appun, C., 377 (23); 2006 (27). Aquilar, M. & D. Castro, 566 (43). Aranda, J. E. et al., 978 (48).


Araujo, D., 7321, s.n. R. 30974 (56). Araujo, D. & R. de Oliveira, 6779 (56). Araujo, D. et al., 7918 (56). Arbelaez, G. et al., 2490 (4). Arbelaez, M. V. & F. Sueroke, 711, 715 (21). Argent, G. C. G. in P. W. Richards, 6712 (52). Argent, G. C. G. et al., 6577 (52). Aristeguieta, L., 3012 (46); 3845, 5107 (48). Asplund, E., 14682 (13). Assunc,o, P. A. C. L., 128 (6). Assun,co, P. A. C. L. & C. F. da Silva, 553 (6); 583 (14). Assun,co, P. A. C. L. et al., 366 (6); 531 (33). Aulestia, C. & M. Aulestia, 975 (50); 1289 (10). Aulestia, M., 3640 (8). Aulestia, M. & J. Andi, 513 (56); Andi 516 (15); 841

(8). Aulestia, M. & 0. Gonti, 1755 (45); 1986 (50). Aulestia, M. & A. Omehuat, 3236 (45). Aulestia, M. et al., 819, 1433 (15); 3593 (32). Ayala, F., 817, 6781, 2954, 3026, 3039 (35); 3136 (cf.

59); 3491 (15); 3587 (35). Aymard, G. & L. Delgado, 6646 (47); 6752 (25); 8483

(33). Aymard, G. et al., 372 (23); 5707, 7381 (35).

Bahia, R. P., 77 (6). Baitello, J. B. & 0. T. Aguilar, s.n. SPSF-8148 (56). Baker, 90 (6). Baker, M. A., 7017 (59). Balansa, 2026 (56). Baldwin, J. T. Jr., 3211 (19). Balee, W. L., 988 (57). Balee, W. L. & B. G. Ribeiro, 1579 (6). Balslev, H. et al., 97016 (55); 97178 (15); 97282 (cf. 59);

97283 (Ocotea cernua); 97440, 97447 (55). Bamps, P., 5145, 5326 (6). Bang, A. M., 1676 (17); 1691 (43). Barbosa, M., 866, 870 (27). Barbour, P. J., 5742 (17). Barclay, A. S. et al., 596 (35). Barreto, M., 3344 (56). Barrier, S., 2361, 2370, 2457, 3256 (46). Barros, W. D. de, 223 (56). Beard, J. S., 189, 228, 628 (46). Beard, P., 1438 (46). Beck, G., 3192 (33); 8176 (56). Beck, G. & R. Foster, 13983 (4). Beck, G. & R. Haase, 10129, 10146 (35). Beck, G. et al., 19650 (cf. 59). Beck, H. T. et al., 1054 (57). Belanger, 161, 979 (46). Benitez de Rojas, C. et al., 5096 (56). Bennett, B., et al., 4306 (57). Benson, W., UNICAMP 10847 (56). Bentancur, J., 5404, 5405 (15); 5418 (14). Berg, C. C., BG 770 (14). Berg, C. C. & W. C. Steward, P 19913 (cf. 55); P19932

(24). Berg, C. C. et al., P19527 (32); P19760 (35).

Berlin, B., 753 (56); 1781 (55); 1783 (32). Bernal, R., et al., 2039a (15). Bernardi, L., 1626 (25); 1789 (4); 18223, 18223b (56). Bertoni, B. S., 916, 917 (56). Bilby, R. et al., 230 (50). Billiet, F. & B. Jadin, 5966 (17). Black, G. A., 48-2639 (24). Black, G. A. et al., 50-9809 (43). Blanco, C., 874 (25); 1138 (24); 1280 (47). Bonifaz, C., 16 (14). Boom, B. & D. Beardsley, 8449 (cf. 49). Boom, B. & M. Pacheco, 8511 (7). Boone, W., 1038 (56). Bourdy, G., 164 (56). Brade, A. C., 7941, 8195, 9062 (56). Braga, P. I. S. et al., s.n. BHCB 27250 (34). Brandbyge, J. & E. Asanza, 30651, 30885 (55); 31785

(17); 32271 32418 (32). Brandbyge, J. et al., 33861 (58); 36206 (56). Brees, E. M., 40 (37). Breteler, F. J., 4859, 4869 (35). Brina, A. E., s.n. BHCB 39260 (56). Brunner, D. R. et al., 917 (56). Buchtien, O., 736 (43); 745, 747, 748 (51); 1986 (17). Bunting, G. S. et al., 4079 (19).

Calder6n, C. E. et al., 2704 (21). Camp, W. H., E-1165 (45). Campbell, D. G. et al., 435 (47); 14650 (26); P21953,

P21960 (7); P21974 (22). Campos, J. & L. Campos, 3092 (cf. 49). Campos, J. & 0. Cano, 4695 (37). Campos, J. & P. Lopez, 4930 (37). Campos, J. & S. Nuniez, 4669 (42). Campos, J. et al., 2938 (18); 4480, 4490 (16). Capucho, P., 565 (Aiouea or Cinnamomum) Carauta, J. P. P., 6251 (56). Carauta, J. P. P. & E. Kautsky, 3284 (56). Carauta, J. P. P. et al., 5175 (56). Cardenas, D. & E. Alvarez, 3267 (47). Cardenas, D. et al., 2645 (48). Cardona, F., 2784 (25). Cardozo, A., et al., 1822 (48). Carnevali, G. & I. Ramirez, 2706, 2707 (35). Carpio, Del C., 2236 (29). Carpio, Del C. & J. Ruiz, 1620 (35). Carri6n, A. et al., 531 (24). Castanieda, R. R., 4068 (cf. 59); 5178 (cf. 44). Castillo, A., 13, 34 (39); 65 (32); 87 (33); 168 (Ocotea

sp.); 1281, 1282, 1300 (35); 1433, 1538 (47); 1632 (35); 2183, 2225, 2309 (47); 2437, 3653 (35).

Cavalcante, P. & M. Silva, 1782 (35). Cavalcanti, T. B. et al., 947 (56). Cazalet, P. C. D. & T. D. Pennington, 7629 (8); 7716

(32). Cerner, 116, 117 (35). Cer6n, C. E., 882 (32); 1986,2397,2411 (14); 2679 (32);

3704 (50); 7875 (17).


Cer6n, C. E. & M. Cer6n, 2600 (50); 4520 (53); Cer6n 4626 (57).

Cer6n, C. E. & F. Coello, 3257 (45). Ceron, C. E. & M. Factos, 7649 (32). Cer6n, C. E. & N. Gallo, 4898, 4973 (35). Cer6n, C. E. & F. Hurtado, 3882 (37); 6642 (49). Cer6n, C. E. & C. Iguago, 5562 (14). Cer6n, C. E. et al., 2090 (48); 8781 (32); 9226 (57). Chagas, J., s.n. INPA 1313 (47). Chagas, J. & D. F. Coelho, s.n. INPA 3695 (7). Chanderbali, A. S. et al., 14 (33); 147, 151, 152, 153

(23); 159 (33); 206 (23); 208, 216 (27); 250 (57); 252 (33); 269 (23).

Chavez, E., 92 (53). Churchill, H. W. et al., 3974, 3998 (10). Cid Ferreira, C. A., 676 (21); 3936 (35); 5563 (22); 5682

(7); 7387 (14); 7485 (50); 8900, 9037 (15). Cid Ferreira, C. A. & J. Lima, 3270 (45); 3590, 3615

(15); 3624 (50). Cid Ferreira, C. A. et al., 203 (35); 266 (21); 291 (22);

582 (21); 690 (50); 881 (6); 1388 (35); 1563 (14); 1601 (22); 1832 (33); 1884 (28); 1888 (Ocotea sp.); 2200, 2321 (35); 5094 (45); 6033 (43); 6713 (6); 7124 (22); 7227 (15); 7257 (35); 7582 (33); 7652 (23); 7776 (7); 7803 (21); 10031 (45); 10487 (43); 10642 (32).

Clark, H. L., 7587 (35). Clark, H. L. & G. Gomez, 8047 (24). Clark, H. L. & P. Maquirino, 7910 (35); 8324 (33). Clark, J. L., 2784 (50). Clark, J. L., et al., 3274 (4). Clarke, D., 2865 (14); 3695 (27); 4414 (57); 6795 (27);

7342, 7516, 7623, 7975, 8070 (14); 8397 (43). Claussen, P., 9, 2093, 2454, 18412 (56). Coelho, D., 29 (22); 160 (21); 3611 (8); s.n. INPA 3206,

s.n. INPA 3895 (21); s.n. INPA 43578 (43). Coelho, D. & L. Coelho, 26 (7). Coelho, D. et al., 896 (21). Coelho, L., 65 (22); 152 (47); 460 (24). Coelho, L. & D. Coelho, 39 (35). Coelho. L. & A. Miranda, s.n. INPA 36033 (6). Cogollo, A. & J. Brand, 380, 394, 415 (4). Cogollo, A. & C. C. Estrada, 290 (Rhodostemonodaphne

antioquensis). Cogollo, A. & J. G. Ramfrez, 4361 (57). Cogollo, A. et al., 2531 (cf. 49); 3638 (47); 3735 (48);

3928 (47); 5119 (10); 5127 (47); 5128 (50); 5129, 5141 (47); 5167, 5182, 5190, 5195 (38); 6119, 6206 (47); 6556, 7061, 7062 (cf. 49).

Collella, M. et al., 1845 (35). Command, R., s.n. G 8341/153 (46). Cordeiro, I., 309 (35). Cornejo, F. V. & A. Balarezo, 2701 (32). Costa, L. V., s.n. BHCB 22314, s.n. BHCB 26334 (56);

s.n. BHCB 32676 (34). Cowan, R. S., 1624 (46). Cowan, R. S. & J. J. Wurdack, 31529 (47). Cramer, 81, (23). Cremers, C., 3982 (14); 5724 (1); 6404 (14).

Croat, T. B., 18609 (31); 18846 (29); 18851 (35); 19156, 19680 (29); 19982 (31); 20680 (36); 20771 (31); 58625 (14); 62517 (17); 62737 (cf. 59); 74133 (14).

Croizat, L., 1052 (25); 919B (57). Cuatrecasas, J. A., 9123 (32); 15292 (50); 16855 (10);

17216 (47). Cuello, N., 637 (35). Cunha, O., et al., 206 (24); 216 (6).

Daly, D. C., 5293 (46). Daly, D. C. et al., 4050 (14); 5086, 5630 (15); 6194 (28);

6795 (57); 7282 (59); 7656, 8104, 9024, 9025 (50); 9293 (58); 9425 (53); 9488 (29).

Damiao, C., 2590 (35). Daniels, A. G. H. & F. P. Jonker, 847, 990 (14). Davidse, G., 27575 (24); 27595 (19); 27893 (35). Davidse, G. & W. D'Arcy, 10097 (10). Davidse, G. & A. Gonzales, 12971, 14663, 15701 (35). Davidse, G. et al., 20840 (48). Davidson, C. & G. Martinelli, 10041 (43). Davis, E. W., 216 (35). Delascio, F. & R. Liesner, 13508 (41). Delgado, L., 582 (35); 835 (47); 868 (35). Delprete, P. et al., 6141 (35). Devia, W., 1052 (4). Devia, W. & F. Prado, 2669 (38). Diaz, C. & M. Alexiades, 3657 (17); 3661 (56). Diaz, C. & S. Balde6n, 2304 (50). Diaz, C. & J. Pereira, 9020 (43). Diaz, C. et al., 783, 8144 (15); 1079 (31); 7710 (59);

8469 (55). Diaz, W. & M. Niiio, 289 (48). Dick, C., 6 (7). Dik, A., 203 (5); 207 (15); 405 (49), 481 (14); 665, 767

(37). Dik, A. & J. Andi, 1001 (45). Dodson, C. et al., 15112 (32). Donselaar, J. van, 1700, 1789, 2465, 2719 (23); 2037

(26). Duarte, A. P., 7368 (29). Ducke, A., 248 (22); 440 (35); 441 (22); 1232, 1728 (6);

1733 (47); 2083 (35); 2478 (13); 2479 (22); 6764 (29); 7185 (35); 8429 (21); 8920 (23); s.n. MG 19238 (22); s.n. MG 15252 (50); s.n. R 2480 (50); s.n. R 11373 (21); s.n. R 17542 (50); s.n. R 18656 (Licaria sp.); s.n. R 19961, s.n. R 19962 (6); s.n. R 19966 (29); s.n. R 19970 (21); s.n. R 60261 (6).

Ducke, A. & M. Bandeira, s.n. R 162 (56). Dudley, T. R., 10072 (8); 11493 (17). Duivenvoorden, 738 (7). Dus6n, P., 12129 (56). Duss, P. 226, 575, 2217, 3624, 4093 (46). Dutra, J., s.n. R. 30975 (56). Dwyer, J. D. & J. E. Simmons, 9748, 9758A (59).

Edwards, K. S. & T. Roe, 423 (14). Eggers, H. F. A. von, 332, 403, 740 (46). Ehringhaus, C. et al., 396 (17). Eiten, G. et al., 5899 (56). Ek, R. C. & D. Hammond, 1033 (33).


Elger, W. A., 914 (29). Encarnaci6n, F., 872 (47); 941 (28); 1241 (35); 1261

(15); 26150 (cf. 55). Ernst, W. R., 1867 (46). Escobar, L. de et al., 8851 (43). Espinoza, S. & T. Coba, 369 (59); 378 (37); 588 (50).

Fanshawe, D. B., 1501 (33). Farney, C. et al., 1744 (35); 2012 (21). Fernandez, A. 6550 (25); 6887, 7415 (47); 7846 (24);

7968 (47). Fernandez, J. C. & Susanna, 8414, 8460 (35). Fernmndez, J. L., 11051 (14). Fernandez, J. L. et al., 11101 (37). Fernandez-Casas, J. & J. Molero, 3818, 4205, 5638 (56). Ferrari, G. & C. E. Benitez de Rojas, 1456 (48). Ferreira, A. R., 236 (22). Ferreira, L. V., 237 (22). Ferreyra, R., 19452 (59). Feuillet, C., 10187 (1). Figlioulo, R. & T. Lima, 125 (35). Figueiredo, C. & I. Riveiro, 553 (cf. 59). Figueiredo, C. et al., 908 (58). Filho, H. F. P., 82-114 (22); 82-2 N, 82-3 N, 83-30 N,

83-49 N (50); 82-51 N, 82-55 N (29); 82-7 N (4). Filho, H. F. P. & L. M. da Silva, 47 (33). Florschutz, P. A. & P. J. M. Maas, 3069 (14). Focke, 101 (23). Folli, D. A., 1546 (56). Fontella, P. J. et al., 476 (56). For. Dept. Brit. Guiana, 2183 (27); G233 (57); 2931,

2949 (23). Forero, E. et al., 2379 (50). Foster, R. B., 4302 (13); 5406, 5475, 9394, 9688 (17);

5949 (59); 9483 (53); 11339 (56). Foster, R. B. & B. d' Achille, 11937 (cf. 59). Foster, R. B. & J. Arce, 11052, 11066 (17). Foster, R. B. & S. Baldeon, 12815 (59). Foster, R. B. & H. Beltran, 13050 (57). Foster, R. B. & C. Janson, 6249 (17). Foster, R. B. & J. Terborgh, 5093, 6154 (17). Foster, R. B. & T. Wachter, 7286 (53). Foster, R. B. & P. Wright, 8254 (59); 8368 (17). Foster, R. B. et al., 3371, 7140 (17); 8925 (55); 11605,

11783 (56); 12483 (58). Fournet, 4205 (46). Francisco, s.n. MG. 21114 (47). Freire, E. & L. Santi, 3294 (50). Freire, E. et al., 2226 (32). Freitas, 52 (58); 53 (30). Freitas & Mota, 467 (15). Freitas, C. A. A. et al., 239 (32). Froehner, C., 67 (43); 171 (56). Fr6es, R. L. de, 20899 (50); 20960 (35); 21022 (30);

21211 (7); 21243 (47); 22620 (24); 23875 (29); 23966, 26368 (35); 28878 (6); 34834 (35).

Funk, V., 6102 (43).

Galdames, C. et al., 1362 (56). Galeano, G. et al., 1001, 1019, 1024 (35); 1065 (36);

1120 (35). Garcia, L. et al., 72, 103, 380 (14). Garcia-Barriga, H., 13703 (47); 13944 (Rhodostemono-

daphne sp.). Gardner, 349, 5595 (56). Gentry, A. H., 28964 (35); 43479 (17); 48990,49120 (6). Gentry, A. H. & J. Aronson, 25184 (36); 25188 (15). Gentry, A. H. & S. Estensoro, 70632 (8). Gentry, A. H. & M. Fallen, 17803 (38). Gentry, A. H. & N. Jaramillo, 58022 (15); 65576 (cf. 59). Gentry, A. H. & P. Nuiiez, 69332, 69392 (59). Gentry, A. H. & R. Ortiz, 74245 (29). Gentry, A. H. & E. Renterfa, 23982, 23983 (10). Gentry, A. H. & J. Revilla, 16514 (35); 16539 (47);

16678 (35); 20368, 20507 (15); 20841 (35); 69177 (28).

Gentry, A. H. & D. N. Smith, 45169 (44). Gentry, A. H. & J. Solomon, 44524 (51). Gentry, A. H. & B. Stein, 46329 (27). Gentry, A. H. & K. Young, 31842 (15). Gentry, A. H. et al., 2533 (15); 7695 (17); 15865 (36);

18525 (20); 19636 (58); 21944 (55); 21980 (7); 22365 (31); 24873 (20); 25797 (35); 25983 (39); 26850 (17); 27088 (59); 27145 (17); 28884 (13); 28964 (35); 31118 (Ocotea sp.); 31649 (15); 37107 (39); 37246 (cf. 55); 38124 (15); 39301 (39); 39734 (31); 42743, 42763, 42779, 42925 (15); 43074 (cf. 8); 43779, 45597, 45598, 45764, 45776, 51521 (15); 54550 (29); 56205 (cf. 59); 56281 (7); 59300,59347, 59377 (56); 62990 (26); 68712 (cf. 59); 68974 (59); 76844 (14); 78108 (17).

Gillespie, L. & H. Persaud, 1557 (23). Gillespie, L. et al., 1691, 1851 (27). Giovianni, F., 32-A (35). Glasgo, A. & V. Slane, 21 (46). Glaziou, A., 3092 (56); 7781, 8093 (34); 12120 (56);

14210 (35); 14212 (21); 16315 (56); 18451 (34); 19795, 20459, 22056, 97811 (56); s.n. (34).

Gleason, H. A., 356 (23). Goeldi, A., s.n. HAMP 3902 (type of Endlicheria goel-

diana); s.n. MG 3911 (8). G6mez, R. et al., 649 (35). Gonggrijp, 3692 (47). Gonggrijp & Stahel, 5607 (14). Goodland, R., 4504 (56). Goulding, M., 41 (35); 1379 (22). Graham, J. & J. V. Schunke, 365 (35); 1099 (17). Grandez, C., 502 (29). Grandez, C. & A. Chiquispama, 894 (35); 912 (15); 919

(29); 982 (50); 996 (35). Grandez, C. & G. Criollo, 1582 (cf. 59). Grandez, C. & N. Jaramillo, 794 (13); 817 (14). Grandez, C. et al., 537 (15); 1526 (58); 1611, 1612 (50);

1921 (7); 4228, 4233 (13); 5512 (45). Granville, J.-J. de et al., 384 (14); 1106 (1); 1331 (14);


1448 (1); 1451, 1462, 2133, 3145, 3723, 3739, 3979, 4949, 5547, 7406, 7675, 7949, 8380, 9005, 9476 (14); 10538 (26); 10794 (14); 10816 (1); 12329 (14); B.3713, B.4356, B.5496 (1).

Grayum, M. et al., 9153 (15). Grenand, 2078 (14); 2870 (1). Grenand & M. F. Pr6vost, 2089 (14). Grewal et al., 384 (23). Grijalva, A. et al., 243 (50); 645 (15). Guanchez, F., 102, 166, 388, 988 (47); 1016 (25); 2154

(41). Gudnchez, F. & A. Brown, 2753 (47). Gudiino, E. & N. Andi, 2064 (17). Gudifio, E. & S. Papa, 1902 (17). Gudiiio, E. et al., 871, 1041 (55). Guedes, M., s.n. R 20065, s.n. R 20081 (43). Guerra, C., 646b (10). Guilding, L., s.n. K H1/45/97-199 (46). Guillaumet, J. L. et al., 5707, 5716 (22). Guillen, R. & R. Chore, 2918, 3283, 3292 (35); 3814

(24). Guillen, R. & C. Medina, 3781 (35). Guillen, R. & V. Roca, 3313 (35). Guillen, R. et al., 3115, 4074 (35); 4454 (24). Gurgel, 15140 (56).

Hahn, L., 1068 (46). Hahn, M., 168, 983, 3184 (46). Hahn, W., 2088, 2591 (56). Hahn, W. et al., 1347 (56); 5795 (33). Halloy, S. et al., 4319 (35). Hamilton, C. & H. Stockwell, 2905 (56). Hammel, B., 1858 (48); 13518 (10). Hammel, B. et al., 21557 (1). Hartshorn, G. & J. Quijano, 2940 (55). Hartshorn, G. et al., 2600 (32); 2606 (4); 2689 (59); 2824

(53); 2827 (4); 2916 (8); 2971 (55). Hassler, E., 8935 (56). Hatschbach, G., 14075, 14366, 39144, 42573, 46151

(56). Hatschbach, G. & E. Barbosa, 59364 (56). Hatschbach, G. & A. C. Cervi, 48903 (56). Hatschbach, G. & 0. Guimaraes, 14678 (56). Hatschbach, G. & R. Kummrow, 52332 (56). Hatschbach, G. & J. M. Silva, 48651, 48978 (56); 52169

(34); 61556 (56). Hatschbach, G. et al., 13246, 51226 (56); 57930 (34);

58562, 61580 (56). Haught, O., 1414 (57). Helme, N., 351 (8). Hemmendorff, E., 457 (34). Henao, J. E., 9, 21 (cf. 49); 152 (15); 281 (cf. 49). Henderson, A. et al., 342 (7); 458 (35). Henkel, T., 4946 (14). Henkel, T. & M. Chin, 601 (23). Henkel, T. & R. Williams, 687, 2091 (23). Henkel, T. et al., 1771 (23); 3430, 3713 (27); 5087 (23).

Heringer, E. P., 16026 (34); 18203 (56); 18530, 18580 (34).

Heringer, E. P. & G. Eiten, 15083, 15154 (34). Heringer, E. P. et al., 1942, 3059, 3062, 3187, 3210,

3211, 3233, 5081, 5184, 5513, 6211, 6245, 6404, 7174 (56).

Herrera, G., 4967, 5031 (15). Herrera, H. et al., 1057 (10). Hetschko, s.n. R. 30983 (56). Heyligers, P. C., 485 (26). Hill, S. R., 13066 (35). Hill, S. R., 24166 (46). Hodge, W. H., 400, 1123, 1317, 2252, 3843 (46). Hodge, W. H. & B. T. Hodge, 2760 (46). Hoehne, F. C., 3517 (56). Hoffman, B. & D. Gopaul, 365 (27). Hoffman, B. & H. Jacobs, 1072 (27). Hoffman, B. et al., 992 (27); 1358, 2029, 2382, 2400

(23). Hohenkerk, L. S., 837 (23). Holm-Nielsen, L. et al., 19908, 19914, 20029 (35). Hopkins, M. J. G., 1595 (6). Hopkins, M. J. G. et al., 633 (27); 1592 (32). Homer, C. et al., 233 (25). Hostmann, F. W. & A. Kappler, 1163 (23). Howard, R. A., 11145 (46). Howard, R. A. & B. R. Howard, 18005 (46). Howard, R. A. & E. S. Howard, 19541 (46). Hoyos, S. E. & J. J. Hemrnndez, 292 (Rhodostemonoda-

phne antioquensis). Huashikat, V. 46 (59); 76, 436 (53); 443, 781 (32); 836

(53); 1475 (cf. 59); 1488 (32). Huber, J., s.n. MG 513, MG 9431 (6); s.n. R 18359 (3). Huber, O., 13261 (41). Huber, 0. & S. Tillett, 2975 (21). Hunt, D. R. & J. F. Ramos, 6004 (52). Hurtado, F., 2654 (17); 2979 (32). Hurtado, F. & A. Alvarado, 963 (49). Hurtado, F. & D. Neill, 1467, 1477, 1509 (55).

Idrobo, J. & R. E. Schultes, 944 (14); 1187 (33). Imray, J., 185, 194, 199, 334 (46). Irwin, H. S., 2688 (56). Irwin, H. S. & L. Y. Th. Westra, 47706 (1). Irwin, H. S. et al., 9097, 13077, 23151 (56); 16777,

16993, 17122, 17291 (52); 47273, 47504, 47933 (1); 48290 (47); 54744, 54747, 54946, 54975, 55132 (14); 55631 (57).

Itaipu Binacional, 129, 924 (56).

Jansen-Jacobs, M. et al., 171, 261 (27); 1118 (23); 2699 (27); 3085 (57); 3311, 3394 (27); 4413 (23); 4575 (57); 4599 (23); 5101 (14); 5374 (27).

Jaramillo, J., 8349 (8); 8372 (50). Jaramillo, J. & E. Grijalva, 11582 (cf. 59). Jaramillo, J. et al., 31147 (7). Jaramillo, N. & S. Katip, 790 (53). Jaramillo, N. et al., 894 (55); 1084 (59); 1159 (53).


Jaramillo, R. et al., 7103 (cf. 45). Jardim, A. et al., 594 (59); 764 (50); 2576 (43). Jarvenpaa, T., 27 (4). Jelski, 165, 196 (43). Jenman, G. S. 1702 (Ocotea sp.); 4127, 5321, 5823 (23). Jervise, s. n. K H842/98-8 (11). Jimenez, B. & G. Manrn, 1331 (56). Jimenez, B. et al., 1809 (56). Jones, J. & C. Davidson, 9639 (58). Jorge, H & R. Torres, 1270 (cf. 49). J0rgensen, P. M., 4462 (56). Jouvin, P. P., 472 (56).

Karsten, s.n. (4). Kayap, R., 354 (8); 1374 (55). Killeen, T., 3832 (Rhodostemonodaphne juruensis);

3880 (50). Killeen, T. & K. Smith, 3634 (56). Killip, E. P. & A. C. Smith, 25975, 26077 (55); 27192

(35); 28050 (4); 28291 (14); 28400 (Rhodostemon- odaphne debilis); 29870 (13).

Kirkbride, J. H. Jr., 2387 (56). Klein, R., 7, 21, 571 (56). Klug, G., 26, 161 (31); 204 (13); 272, 273 (7); 411, 621,

703 (13); 728 (35); 1264, 1403 (20); 1884, 1904 (53); 2253 (30); 2313 (14); 2833 (56); 2958 (14); 3187, 3745 (28).

Knab-Vispo, C. & G. Rodriguez, 479 (25). Knapp, S. & P. Alcom, 7778 (49). Knapp, S. & J. Mallet, 653 (14); 2829 (27); 7024 (59);

7110, 8557 (14). Knapp, S. et al., 4496 (56). Kroll Saldania, B., 134, 326 (8); 560, 689, 795 (56). Krukoff, B. A., 1565 (15); 1933 (14); 4714 (15); 4717,

4767 (17); 4775 (Rhodostemonodaphne juruensis); 4855 (17); 4932 (8); 4943 (53); 4959 (13); 5023 (59); 5030 (14); 5156 (15); 5204 (17); 5279 (50); 5281 (15); 5780 (3); 6116 (50); 6300 (58); 6406 (50); 7206 (32); 7265 (22); 8096 (24); 8293 (29); 10787 (17); 11103 (43); 11262 (51).

Kubitzki, K., 75-34 (29); 87-22 (22). Kubitzki, K. & H. Poppendieck, 79-29 (21). Kubitzki, K. et al., 79-172 (19). Kuhlmann, J. G., 171 (35); 3376 (27); s.n. R 11966, R

136580 (56); s.n. R 18360 (50); s.n. R 20028 (53); s.n. R 20036 (29).

Kurtz, B. C. et al., 80, 87 (56). Kvist, L. P. & L. Freitas A. 706, 736, 808, 921 (35).

Labroy, s.n. (35). Lanjuow, J. & J. Lindeman, 2367 (14). Larpin, D., 851 (14). Lasser, T., 1881 (57); 2036 (48). Lasser, T. & C. K. Allen, 13 (48). Leng, H., 128 (23). Lepsch da Cunha, N. M. et al., 391 (2). Level, S., 127 (24). Lewis, G. P., 1488 (25). Liesner, R. L., 3961, 4183, 6237 (35); 6657 (32); 7137

(35); 7466 (19); 8479, 8640, 8665 (35); 18657, 18665, 18671, 19092 (47); 24576 (25).

Liesner, R. L. & G. Carnevali, 22292, 22414 (41). Liesner, R. L. & H. Clark, 8963 (47). Liesner, R. L. & F. Delascio, 22008 (41). Liesner, R. L. & B. Holst, 21854 (47). Liesner, R. L. & G. Morillo, 13929 (25). Lima, J., 783 (25). Lima, J. & S. Sousa, 230 (50). Lindeman, J. C., 379 (47); 2741 (14); 4509 (47); 4590

(23); 4787 (26); 5272 (23); 5381 (cf. 32); 5601 (23); 5943, 6056 (26); 6651 (23).

Lindeman, J. C. & J. H. de Haas, 747 (56). Linden, 429 (11). Lisb6a, P. & 0. P. Monteiro, 954 (7). Little, E. L. Jr., 8878, 8886, 40105, 40141 (56). Little, E. L. Jr. & R. G. Dixon, 21035 (15). Little, E. L. Jr. & R. R. Little, 8259, 8308, 9565 (35). Little, E. L. Jr. et al., 221 (44). LLB (Landsbosbeheer, Suriname), 12570 (37). Lleras, E. et al., P16960 (53). Lohmann, L. G., 110, 343 (35). Lombardi, J. A., 1987 (34). Lopes, M. et al., 124 (21). L6pez, A. et al., 8635 (35). Lorea, F. G., 5581 (56). Lorea, F. G. & M. Gorgulho, 5579, 5580 (56). Loureiro, A. et al., 742, 1320 (35); 5739 (47); s.n. INPA

35784 (7); s.n. INPA 37822, s.n. INPA 37846 (22); s.n. INPA 37949 (7); s.n. INPA 38000, s.n. INPA 38870, s.n. INPA 38878 (35); s.n. INPA 39526 (22); s.n. INPA 47996 (21, 22); s.n. INPA 48137 (47).

Lugo, H. S., 3906, 3925, 4125 (32); 4188, 4241, 4302, 4394, 4450 (45).

Luna, M. L., 112 (44). Luteyn, J. L. et al., 8541 (59); 8586 (cf. 59); 8680, 9004,

9019 (17).

Maas, P. J. M. & J. A. Tawjoeran, s.n. LLB 10928 (23). Maas, P. J. M. & L. Y. Th. Westra, 3815 (23). Maas, P. J. M. et al., 4599, 6637 (35); 6798 (32); 7136,

7166, 7215, 7308 (27); P13058 (40); P13307 (58). Machado Nunes, G., 318 (34). Maciel, U. N. & C. S. Rosairio, 1559 (24). Mackenzie, C. A. et al., INPA/WWF 2107.388 (2). Madriiain, S., 710 (35). Madrifian, S. & C. Barbosa, 936, 948, 953 (41). Maguire, B., 24538 (14); 24712 (37, 47); 24898a (37);

40775, 40790 (14). Maguire, B. & D. B. Fanshawe, 23481 (23); 32463 (47). Maguire, B. & L. Politi, 27384, 28159, 28380 (47);

28428 (32). Maguire, B. & J. J. Wurdack, 34727 (25); 34859 (35). Maguire, B. et al., 29436 (57); 30601 (21); 36314 (47);

42244 (41); 42521 (Aiouea maguireana); 46713 (23); 53519 (47); 56091 (52); 56624 (22); 60121 (47).

Maia, L. A. et al., 208 (35); 344 (22); 430, 438, 562 (24). Mallorguim, R., 16 (56). Malme, G., 755 (56).


Manara, B., s.n. VEN 172708 (48). Manso & Lhotzky, 84 (52). Marcano-Berti, L., 361 (26); 2528 (25); 2581 (47). March, W. T.?, s.n. (23). Marin, E., 438 (19); 981 (24). Marfn, G. & B. Jimenez, 322 (56). Martinelli, G., 151 (56). Martinelli, G. & D. Sucre, 200 (56). Martinelli, G. et al., 23 (34). Marulanda, 0. & S. Marquez, 1108-HUA 88921 (35);

1108-HUA 89052, 1536 (58). Mathias, M. E. & D. Taylor, 5514 (29); 5532 (58); 5957

(17). Matos, F. et al., 207 (50). Matuda, E., 5153 (Beilschmiedia zapoteoides). McDaniel, F. & L. Santiago, 2539 (58). McDaniel, F. & T. Wilkes, 2483 (29). McDaniel, S. & M. Rimachi, 18838 (58); 20744 (36). McDowell, T., 2663 (47). McDowell, T. & S. Tiwari, 1861 (27). McPherson, G., 8493 (12); 10000, 10444, 11310, 11582,

11936 (48). McPherson, G. & M. Merello, 8131 (15). Mecenas, V. V. & F. Q. Leite, 92 (56). Meier, W. & 0. Kunert, 4632 (56). Meier, W. & K. Walter-Weisbeck, 2586 (48). Meier, W. et al., 2756 (25). Melinon, 204, 216, 227, 551 (1). Mello, F., 24 (35); s.n. INPA 78 (47); s.n. INPA 3526

(6). Mello, F. & L. F. Coelho, s.n. INPA 4167 (47). Mello, F. & M. Emmerich, 2934 (56). Mello, F. & J. Ribamar, s.n. INPA 58316 (43). Mendez, P. et al., 22 (15).

Mendonqa, R. C. & F. C. Silva, 61 (56). Mennaga, A. M. W., 86 (14). Mexia,Y., 5025, 5091, 5138 (34); 6268a (53); 6329 (56). Meyer, G., 146 (56). Miers, J., 4270 (56). Miller, J. S. & G. Davidse, 1649 (24). Miller, J. S. et al., 638 (55). Miller, R., 524 (35). Milliken, W. & S. Bowles, 304 (25). Milliken, W. et al., 142, 439 (27); 524 (35). Miralha, J. M. S., 78 (24). Miralha, J. M. S. et al., s.n., Araca Inventory BO-6-611

(24). Miranda, S. I., 20 (59); 1000 (27). Molas, L., 771, 820 (56). Molino, J.-F., 1663 (14). Monsalve, M., 1975 (38). Monteiro, 0. P., 1246 (21); 1252 (32). Monteiro, 0. P. & J. Lima, 145 (7). Moore, S., 518 (52). Moraes, P. L. R. de, 122, 123, 403, 561 (56). Morales, J. F. et al., 2083 (15). Morawetz, W. & B. Wallnofer, 11-12888 (28); 13-9288

(48). Moretti, C., 1196 (14); 1443 (8).

Mori, S., 7708 (56); 23935 (14). Mori, S. & C. Gracie, 21867, 21962, 22473, 22492 (35). Mori, S. & J. Kallunki, 3472 (10). Mori, S. & T. Pennington, 18041 (1). Mori, S. et al., 6476 (10); 9097 (53); 9178 (Ocotea sp.);

14910, 14995 (1); 15059, 15649 (14); 17170 (57); 20614 (7); 20927 (26); 22268 (1); 22979 (14).

Morillo, G. & R. Liesner, 8818, 8936 (25). Moscheta, I. & D. dos Saldovino, 3 (56). Mota, C. D. A., 685 (28); s.n. INPA 60364 (55); s.n.

INPA 60608 (21). Mota, C. D. A. & 0. P. Monteiro, s.n. INPA 61281 (21). Muinera, M. L. E., 13 (cf. 49). Mutchnick, P., 687, 1243, 1303 (23); 1464 (33).

Nadruz, M. et al., 573 (56). Nascimento J. R. & E. C. da Pereira, 606 (cf. 53). Nascimento, 0. C., 386 (6); 685 (32); 826 (19). Nee, M., 34692 (43); 39040 (56); 39834 (17). Nee, M. & L. Bohs, 49533 (17). Neill, D., 7152 (53); 7164 (cf. 59); 7266 (14); 7514 (15);

8730 (53); 9842 (14); 9886 (32); 10095 (17); 11028 (53).

Neill, D. & F. Hurtado, 8807 (55). Neill, D. & W. Palacios, 9618 (45). Neill, D. & W. Rojas, 9950 (17). Neill, D. et al., 8094 (49); 8475 (48); 8953 (49); 10168

(15); 12614, 12615 (cf. 49); 12616 (42). Nelson, B. W. & J. F. Lima, P21108 (21). Nevers, G. de & D. Cavagnaro, 4795 (56). Nevers, G. de & H. Herrera, 4186 (10). Nevers, G. de et al., 5339 (10). Nicholls, H. A., s.n. (46). Nicholson, D. H., 4237 (46). Nunhez, P., 5907, 6202, 6260 (17); 11195 (43); 13986

(15). Nunfez, P. & C. Munioz, 5324 (14). Nufiez, P. et al., 8017 (17); 10804 (4); 14261 (17); 14373

(cf. 59); 14429 (56).

Oldeman, R. A. A., 79, 2448, B. 2993, B.3580, T-35, T- 55 (47); 1251, 2866, B. 1881, B. 817, B. 3996, B 4034 (14); T-23 (1).

Oldenburger et al., 1221 (cf. 33). Oliveira, F. C. A. et al., 770 (34). Oliveira, P. I., 256, 926 (56). 0llgaard, B. et al., 34616 (17); 57630, 57623A (10). Ortiz, R. & J. Vargas, 206 (53). Ortiz, R. et al., 122, 124 (58).

Palacios, W., 1005 (14); 1370 (53); 1423 (59); 3260 (14); 3496, 3496 (55); 4044 (15); 4212, 4386 (57); 4397 (28); 4413, 4473 (4); 4747 (49); 5482 (15); 5597, 5663 (53); 6732 (51); 7500 (28); 10294 (33); 11001 (49); 11268 (17); 13750 (10); 13897, 13906 (15); 13936 (33).

Palacios, W. & E. Freire, 5119 (8). Palacios, W. & C. Iguago, 4541 (49); 4696 (8). Palacios, W. & D. Neill, 1289 (50); 1516 (53); 6522,

6531 (55); 6572, 6577, 6669, 6688 (45).


Palacios, W. & G. Tipaz, 11074 (45). Palacios, W. & M. Tirado, 11165 (15); 13298, 13302

(57); 13310 (45); 13342 (51); 13425, 13439, 13459 (42).

Palacios, W. et al., 1030 (56); 1183 (53); 4936, 7726 (8); 7804, 7884 (58); 7879 (15); 8027, 8042 (59); 8142, 8144 (35); 8164, 8165, 8399 (37); 8492 (45); 8585 (57); 8644 (15); 8840 (57); 8845 (7); 8938, 8940 (48); 8963, 9492 (15).

Paniagua, N. & R. Foster, 690 (35). Pariona, W., 399 (32). Pariona, W. & J. Ruiz, 974 (32). Pariona, W. & A. Sebastian, 25 (32); 46 (4). Patris, s.n. G8341107, G8341109, G8341110,

G8341111, G8341112, G8341113, G8341117 (26). Paulino R. & F. Victorio, s.n. R 2322 (type of Endlicheria

balsamea). Pavon, J., 506 (17); s.n. G 8341 (17). Pearce, R., s.n. K 52, MO 1611827 (51); s.n. K 59, K 60

(16). Peckolt, 115 (56). Peixoto, A. et al., 3466, 3504 (56). Pennington, R. T. & J. Ramos, P22768 (43). Pennington, R. T. et al., 80 (50). Pereira, E., 5356 (56). Perry, A., 999 (43). Persaud, A. C., 49, 59 (23). Pesha, V., 68 (50). Pfrommer, A., 6 (56). Philipson, W. R. & J. M. Idrobo, 2013 (32). Philipson, W. R. et al., 2192 (15). Phillips, 0. & P. Nuinez, 52 (58). Phillips, 0. et al., 468 (15). Pierre, L. L. Jr. & V. Slane, 357 (46). Pilar Franco, R. et al., 5337 (37); 5499 (51). Pinkus, 263 (47). Pinto, F. & P. Q. Bernal, 1624, 1642 (cf. 59). Pipoly, J. J., 10316 (47). Pipoly, J. J. & G. Samuels, 6868 (24). Pipoly, J. J. et al., 10607 (33); 12202 (13); 12633 (7);

12727 (20); 13554 (31); 14321 (28); 16280 (15); 16545, 16568, 16718, 17080 (47); 17315, 17453 (cf. 49).

Pires, J. M. & R. P. Belem, 12405 (14). Pires, J. M. & G. A. Black, 112 (6); 1055 (13); 1229 (24). Pires J. M. & P. B. Cavalcante, 52649 (47); 52676 (14). Pires, J. M. & N. T. Silva, 10647, 10658 (6). Pires, J. M. et al., 9235 (56); 13887 (35); 16821 (25);

50825 (14); 51168 (1); 51599 (57). Pires, 0. & D. Coelho, 175 (2). Pittier, H., 8017 (56). Pizziolo, W., 280 (56). Plotkin, M., 159 (47). Plowman, T. & E. W. Davis, 5098 (17). Plowman, T. & H. C. de Lima, 12895 (56). Plowman, T. & J. V. Schunke, 11575 (14). Plowman, T. et al., 6693, 7013 (36). Poeppig, 1520, 2298 (56); 2552 (35). Pohl, 5611 (56).

Poiteau, M., s.n. G 122, G 123, G 124, G 128, LE 2811, NY 99497, P 162 (14).

Polanco, R., 222 (56). Pollard, 79 (27). Poncy, O., 89 (14). Poole, J. M., 1717 (35); 1972 (19). Prance, G. T., 30162 (22). Prance, G .T. & D. F. Coelho, 17581 (7). Prance, G. T. & T. D. Pennington, 2004 (cf. Rhodoste-

monodaphne tumucumaquensis Madriniain). Prance, G. T. & A. E. Prance, 14774 (35). Prance, G. T. & N. T. Silva, 59385 (24). Prance, G. T. et al., 1343 (Ocotea diffusa van der Werff);

1563 (14); 2269, 2688 (47); 2972 (35); 3008 (7); 3180 (47); 3563 (29); 3817 (21); 3886 (6); 3919 (7); 4211 (27); 4282 (25); 4678 (21); 5290, 5295 (35); 5327 (22); 6182 (29); 6834 (35); 7744 (29); 7810 (17); 8343, 8892 (6); 9961 (Ocotea sp.); 11069 (25); 11511 (24); 11568, 11596, 11737 (35); 12102 (7); 12161 (cf. 59); 12214 (17); 12233 (55); 12436 (7); 12501 (17); 12514 (cf. 59); 12563 (45); 12633 (13); 13391, 13442, 13485, 13695 (58); 13888 (32); 13987 (14); 14013 (50); 14150 (40); 14498 (4); 14616 (E. metallica aff sp. 1); 14671 (24); 15035, 15725 (47); 15817 (6); 16527 (50); 16695 (58); 17742 (7); 17785 (22); 18032, 18037 (7); 24626 (35); 29998 (22).

Prevost, M. F. & Grenand, 1958 (14). Proctor, G. R., 18205, 21575, 21585 (46). Proenqa, C., 863 (56). Pulle, A., 223 (14).

Qazarin, P. R., 11 (29). Quelal, C. et al., 704 (Rhodostemonodaphne juruensis). Quentin, R. P., 1091 (46). Quifiones, L., 1419 (14).

Raimondi, 4098 (43). Ramage, G. A., s.n. (46). Rambo, B., 43408, 45269, 63615 (56). Ramfrez, J. G. & D. Cardenas, 1016 (8). Ramfrez, J. G. et al., 4886 (32). Ramirez, R. C., 1068 (29). Ramos, J., 450 (45); s.n. INPA 54118 (29); s.n. INPA

62184 (47). Ramos, J. & D. Coelho, 693 (32). Ramos, J. & G. Mota, 192, 332 (7); 350 (45). Ramos, J. & R. Sousa, 68 (52). Ramos, J. et al., 684 (Ocotea sp.); s.n. INPA 62154 (21). Ratter, J. A. et al., 840, 1278, 1944, 5590, 5634, 5713

(25). Ravedutti, C. et al., 7 (56). Regnell, A. F., s.n. R. 30958, R. 30976 (56). Reif, 1785 (56). Reitz, P. R., 3098, 4423, C153 (56). Reitz, P. R. & R. Klein, 286, 931, 1140, 1676, 1833 (56). Renterfa, E. et al., 2821 (8). Restrepo, D. & A. Matapi, 376, 466 (7). Revilla, J., 210 (29); 410 (35); 460 (29); 625 (35); 636

(13); 675, 695 (35); 798 (29); 885 (35); 889 (29); 945


(7); 1153 (29); 1224 (28); 2111 (35); 3627 (47); 3772 (29).

Revilla, J. & J. Forero, 4175, INPA 82158 (22). Revilla, J. et al., 2472 (35). Reynel, C. & E. Meneses, 5089 (50); 5115 (8); 5125

(50). Reynel, C. et al., 5244 (28). Ribeira, R. & I. Silva, 2061 (56). Ribeiro, J. E. L. S. et al., 885 (47). Ribeiro, R., 544 (56). Ribeiro, R. & W. L. Araujo, 130 (56). Ribeiro, R. et al., 1021 (cf. 56). Richards, P. W., 6871 (52). Riedel, L., 1142 (56); 1374, 1451 (35). Rimachi, M. Y, 2533 (29); 2590 (58); 3226 (31); 3506

(36); 4286 (35); 4397 (55); 4644 (7); 7739, 9113 (20); 9906 (29); 10391 (28); 10402, 10532 (20); 11199 (13); 11305, 12263 (31); 12301 (50); 12302 (29).

Ritter, N., 1706 (17). Robert, A., 4856 (56). Robertson, K. R. & D. F. Austin, 90, 100 (35). Rodrfgues, E. & P. M. Reyes, 1807 (44). Rodrigues, H., 2545 (24). Rodrigues, R. S., s.n. MG 8265, s.n. MG 8811 (43). Rodrigues, W., 374 (47); 508 (21); 552 (53); 574 (47);

1064 (32); 1596 (35); 1972 (22); 5389 (32); 6715 (22); 7028 (28); 7977 (cf. Ocotea rufovestita); 8780 (6); 8781 (7); 8835 (35).

Rodrigues, W. & J. Chagas, 1259 (21); 2350 (47). Rodrigues, W. & D. Coelho, 461 (21); 2473 (22); 2588,

2743, 2783, 3033, 4558A (21); 5278 (35). Rodrigues, W. & L. Coelho, 2564 (21); 5306 (7). Rodrigues, W. & J. Lima, 2605 (35); 4124 (21). Rodrigues, W. & A. Loureiro, 5926 (33); 7075 (47);

9454 (2). Rodrigues, W. & F. Mello, 5355 (35); 7760 (29). Rodrigues, W. et al., 114 (56); 941 (55); 1980 (21); 8523

(2); 10494 (21). Rojas, R. et al., 91 (57); 172 (55); 506 (32). Rollet, B., 1672 (46). Romero, G. A. & E. Melgueiro, 2010 (35); 2129 (21). Romoleroux, K., 2122 (cf. 59). Romoleroux, K. & R. Foster, 1908 (15). Romoleroux, K. et al., 2567 (48); 3502 (50). Rosales, J. et al., 1245 (25). Rosario, C. S. et al., 1032 (6). Rosas, A. et al., 216 (43). Rubio, D., 8 (45); 11, 12, 40 (55); 115 (13); 306 (14). Rubio, D. & A. Alvarado, 2395 (49). Rubio, D. & E. Gudiiio, 199 (13). Rubio, D. et al., 454 (Rhodostemonodaphne juruensis);

920 (15). Rudas, A. et al., 2116 (50); 2152 (35); 2262, 2288 (50);

2290 (29); 2504 (15); 2588 (50); 2610, 2642, 2648 (37); 3601 (15); 4057 (57); 5817 (15).

Rueda, R., 445 (29); 1116 (57). Rueda, R. & J. C. Ruiz, 679 (35). Ruiz, H. & J. Pav6n, s.n. (17).

Rufz, J. C., 1436, 1466a, 1515 (35). Ruiz, J. C. & H. Murphy, 228 (13); 230 (31). Ruiz, J. C. et al., 6285 (29). Rusby, H. H., 572, 2671 (17). Ruzz, J., 1409 (15).

Sabatier, D. G., 1087 (47); 2365 (26); 3501 (14). Sabatier, D. G. & M. F. Prevost, 3195 (33); 3819, 4073

(26). Salino, A., 3274 (56). Samaniego, A. V. & A. C. Vivar, 90 (44). Sampaio, A., 2223 (56). Sanchez, I. V. & M. Dillon, 8687 (43); 8916, 8925 (28). Sanchez, I. V. et al., 9345 (E. metallica aff. sp. 2); 9575

(56); 29363 (8). Sanchez, M. et al., 1870 (22). Sanoja, E., 2522 (25). Santos et al., 39 (50). Santos, M. R., 563 (22). Saraiva, R. S. & L. de Lima, 1523 (43). Sastre, C ., 1488 (1); 1619 (14); 1644 (1); 1651 (14). Sastre, C. & F. Sastre, 2046 (46). Schinini, A. & G. C. Marmori, 27009 (56). Schomburgk, R., 171(240) (23); 475/801 (27); 784 (35). Schrainer, s.n. R 61160 (56). Schultes, R. E., 3337 (cf. 59); 7149 (35). Schultes, R. E. & I. Cabrera, 12423 (32); 12658, 12664

(35); 12735 (32); 13067, 13073, 13076, 13236, 13241 (35); 13566 (13); 13849 (24); 14125 (13); 14356 (21); 14606 (47); 14903 (45); 15566 (35); 16230 (13); 17033, 17898, 17898, 18421 (35); 19540 (39); 20003 (21); 22578, 24200 (35).

Schultes, R. E. & F. L6pez, 9954 (24). Schultes, R. et al., 24174, 24200 (35). Schunke, J. V., 1587, 1604, 1617 (56); 2570 (59); 2924

(56); 3329 (17); 3474, 5679 (14); 4532, 4766 (17); 5060 (28); 6231 (15); 6726 (Rhodostemonodaphne juruensis); 6957 (28); 7053 (17); 7245, 8069 (28); 9242 (56); 10304 (28); 10657 (56); 12481 (17).

Schwacke, C. A. W., 95 (56); 351 (22); 617 (35); s.n. R 30973 (34); s.n. R 61142 (56).

Seemann, B. C., 1094 (10). SEF, 8740 (50); 9247 (32). Seidel, R. et al., 6557 (58). Sellow, 433, 1166, 2244, B 383, c 418 (56). Serato, A., 305 (56). Setz, E., F779 (43). Shiki, D., RBAE359 (14). Shillingford, C. A., 517 (46). Sidney, 1308 (52). Sieber, F. W., 175 (46). Silva, A., 135, 282 (6); 309 (43). Silva, A. S. L. da et al., 635 (14). Silva, E. S., 185 (24). Silva, J. A., 181 (24). Silva, J. M. & L. M. Abe, 2883 (56). Silva, J. M. & E. Barbosa, 2435 (56). Silva, J. M. et al., s.n. INPA/WWF 2303.4840 (28). Silva, M. F., 788, 823, 824 (35); 960, 976 (22).


Silva, M. F. & R. Souza, 2546 (Ocotea nigrescens Vi- centini); 2630 (Ocotea diffusa van der Werff).

Silva, M. F. et al., 381, 680 (35); 808 (47); 1707 (19); 1752 (35).

Silva, M. G., 1042 (6); 6059, 6078 (50); 7148 (7). Silva, N. T. da, 38 (6); 968 (43); 57857 (6). Silveira, M. et al., 468 (29); 943 (28); 1499 (cf. 59); 1555

(15); 1565 (50). Simonis, J. E. et al., 121 (56). Simpson, D. R., 782 (20). Slane, V. & P. Acevedo, 287 (46). Slane, V. & B. Rollet, 514 (46). Smith, A. C., 2221, 3129, 3376 (27); 10356 (46). Smith, D. N., 4491 (44); 5318 (56); 6764 (50). Smith, D. N. & S. Vasquez, 4775 (44). Smith, D. N. et al., 1748, 8670 (56); 13908 (33); 14052

(56). Smith, H. H. & G. W. Smith, 240, 323, 353, 1840 (46). Smith, S. F. et al., 1503 (8); 1599 (17). Sneidern, 1191 (7). Snethlage, E.(?), s.n. MG 9411 = R18362 (6). Soares, E., 189 (47); 462 (21). Soejarto, D., 580 (32). Soejarto, D. et al., 4198 (35). Solomon, J. C., 9255 (51); 12948 (56). Solomon, J. C. et al., 7140 (56). Soria, N., 3419 (56). Sothers, C. A., 727 (6). Sousa, J., 76 (28). Souza, M. A. D. & P. A. C .L. Assunqao, 491 (28). Souza, M. A. D. et al., 396 (28). Sperling, C. R. et al., 6157 (14). Spichiger, R. & F. Encarnaci6n, 1043, 1122 (35). Spichiger, R. & P. A. Loizeau, 4108, 4161 (cf. 55); 4758,

4759 (28). Spruce, R., 1453, 1453 bis (22); 1757 (35); 2769 (7);

3061 (19); 3092 (47); Nectandra 6 (35). Stahel, G., 43 (14); 236 (23). StehIl, H., 387, 919, 1168 (46). Stehle, H. & M. StehIl, 6337 (46). Stein, B. & A. Gentry, 1660 (41). Steinbach, J., 3274, 5291, 5369, 6985, 7264 (56); 7264

(35). Steinbach, R. F., 765 (56). Stergios, B. & G. Aymard, 4171, 7371, 7386, 7400 (35);

7524 (19); 9006, 9104 (35). Stergios, B. & P. Stergios, 11427 (47). Stergios, B. & J. Velazco, 14495 (33). Stergios, B. et al., 8187, 8280, 9607, 9610, 9706, 9839,

13259 (35); 13295 (21). Steward, W. C. et al., P20357 (21). Steyermark, J. A., 57935 (47); 59001 (cf. Rhodostemon-

odaphne grandis); 60428 (57); 75564 (47); 90315 (35); 90316 (47); 90535 (25); 91524 (48); 97760 (47); 107391 (25); 107466 (47); 111387 (37).

Steyermark, J. A. & G. Agostini, 7 (48). Steyermark, J. A. & G. Davidse, 116876, 116957 (46). Steyermark, J. A. & V. C. Espinoza, 110275 (56). Steyermark, J. A. & R. Liesner, 118687 (56).

Steyermark, J. A. & S. Nilsson, 255 (60). Steyermark, J. A. & P. Redmond, 117081 (35). Steyermark, J. A. & C. Steyermark, 95347 (48). Steyermark, J. A. et al., 100298 (48); 101408 (4); 111746

(56); 119895 (46); 126162 (24); 126285 (57); 131425 (27); 131500 (35).

Strier, K. B., 992 (34). Sturrock, 583 (46). Stutz de Ortega, L. C., 1238, 2175, 2223 (56). Sucre, D, & P. I. S. Braga 1404, 1825 (56). Suriname Forest Bureau, 2936, 3197 (37); 3494 (23);

5884, 6884 (37). Sytsma, K., 994 (10).

Talbot, H. F., s.n. (56). Tameirao Neto, E., 87, 2653 (56). Tameirao Neto, E. & M. S. Werneck, 1244, 1245 (56). Teixeira, E. M. & A. E. Brina, s.n. BHCB 36260 (56). Teixeira, L. 0. A. et al., 852 (28); 1025, 1163 (7); 1221

(22). Tello, 560 (56). Tessmann, G., 3439 (35); 3058a, 3999, 4248 (17); 4734

(59); 4736 (14); 5126 (35); 5146 (31); 5288 (29). Thomas, W. et al., 4059, 5370 (43); 5089 (22); 6775 (29). Thomsen, K., 595 (15). Tillett, S. S. & C. L. Tillett, 45705 (47); 45789 (37). Tillett, S. S. et al., 43990, 45163 (33). Timana, M. & N. Jaramillo, 2466 (59); 2476 (50); 2538

(59). Timana, M. & 0. Phillips, 1868, 1878, 1882 (15). Tipaz, G., 2516, 2632 (15); 2730 (17). Tipaz, G. et al., 581 (17); 709 (59); 1104 (Rhodostemon-

odaphne juruensis). Tirado, M., 1010 (8). Tirado, M. et al., 462 (15); 586 (10). Toledo, F. R. N. & J. A. Lombardi, s.n. BHCB 46022

(56). Torres, J., 156, 300 (35); 3092 (cf. 59). Triana, J. 1032 (4); 1033 (11); 1059 (4); 2040-2 (11);

2060 (4). Tunqui, S., 215 (32); 316, 589, 649 (cf. 59).

Ule, E., 5 (56); 5581 (58); 5584 (29); 5710 (17); 6296 (14); 8125 (27); 8848 (21); s.n. R. 61128 (56).

Urrego, L. E. et al., 1012 (45); 1218 (cf. 51). Usteri, P. A., 3026 (56).

Valcarcel, J. H., 383-I/F (29). Valcarcel, J. H. & M. Chota, 6/619 (cf. 55); 7/75 (28); I/

98 (33). Valdespino, I. A. et al., 278 (48). Valle, M. H., 5 (56). Valverde, L. & I. Penia, 1094 (24). Varejao, de Jesus M., s.n. INPA 140522 (22). Vargas, H., 1152 (59). Vargas, H. & P. Grefa, 764 (32). Vargas, I. G. et al., 120 (15); 1098, 1137, 2478 (17). Vargas, R., 2 (56); 1096 (35); 3319 (9); 4193 (32); 6611,

10989 (35); 12287 (58); 12296 (29). Vasquez, R. & G. Criollo, 1785 (13).

INDEX OF LOCAL NAMES 137

Vasquez, R. & C. Grandez, 17475 (58); 17487 (15). Vasquez, R. & N. Jaramillo, 40 (cf. 59); 203 (56); 1256

(35); 2425 (53); 2441 (50); 2501 (57); 3157 (20); 3165 (47); 5253 (20); 5258 (35); 5277 (15); 5457 (20); 5697 (8); 6085, 7174 (15); 7620 (20); 7633, 7644 (47); 7647 (20); 8114, 8215 (31); 8270 (15); 8398 (35); 8414, 8766 (20); 9123, 9125 (29); 9180, 9183 (9); 9593 (39); 9606 (32); 10228 (15); 10458 (59); 10749 (13); 11442 (55); 11467 (cf. 59); 11577 (35); 13609 (31); 13637 (28); 13711, 13945 (31); 14922 (39); 20300 (32).

Vasquez, R. & R. Jong, 12397 (54). Vasquez, R. & R. Rojas, 21920 (cf. 49). Vasquez, R. & T. Soto, 12350 (28). Vasquez, R. & H. Valderrama, 1363 (31). Vasquez, R. & A. Vdsquez, 20969 (8). Vasquez, R. et al., 111 (35); 648 (13); 651 (31); 4778

(35); 5307 (28); 5434 (20); 6393 (58); 6665 (15); 6666 (29); 6675 (15); 6742 (7); 10934 (15); 10939 (36); 11928 (59); 11954 (50); 12391 (32); 12435 (8); 14393 (31); 17706 (39); 18095 (53); 19459 (8); 21426, 21528, 21535 (53); 23419 (50); 23530 (13); 23873 (32); 23906 (8); 23948, 24025 (59); 24341 (55); 24898 (8); 25231 (39); 25237 (31).

Velazco, J., 587 (35); 859 (47); 1032 (33). Velloso, H., 170, 205, 209, 398 (56). Vester, H. & R. Roman, 865 (45). Vicentini, A. & E. da C. Pereira, 903 (8). Vicentini, A. & E. M. Venticinque, 1000 (32). Vicentini, A. et al., 635 (cf. 53); 993 (cf. 55); 1224 (7);

1317A, 1359, 1394 (21). Vidal, J., 1342, 1710, 2020 (56).

Wallnofer, B., 17-31588 (17); 111-8488 (53); 14- 221287 (32).

Walter, B. M. T. et al., 3517 (56). Wasshausen, D. C. & F. Encarnaci6n, 649 (59). Weberbauer, 4680 (28). Wedel, H. von, 2257 (10). Werff, H. van der, 12953, 12964 (14).

Werff, H. van der & E. Gudinlo, 11136 (Rhodostemono- daphne juruensis).

Werff, H. van der & W. Palacios, 9246 (cf. 49); 10385 (49).

Werff, H. van der et al., 8340, 8342 (43); 9776 (39); 9796 (13); 9815 (9); 9991 (39); 10045 (28); 10048 (58); 10053 (29); 10084 (8); 10187 (39); 10207 (20); 10237 (50); 10242 (39); 13116, 13147 (15); 15436 (40); 15736 (50); 15738 (8).

Wessels Boer, J. G., 1043 (14); 1381 (23). Whitefoord, C., 3533, 4267, 4490, 5349 (46). Wijninga, V., 580 (56). Wilde, L., 6-88 (47); 14-88 (22). Williams, L., 609 (35); 1002, 1003, 1004, 1193, 1203

(36); 1494, 1877 (35); 11348 (47); 14370 (32); 14788 (47); 15179 (25); 15367 (35).

Wilson-Browne, Fr. G., 104, 178 (27); 454 (57); 470 (27).

Wolfe, 12194 (17). Worbes, M., 1012 (22). Woytkowski, F., 5829 (17); 5864 (8); 6304 (29); 6315

(35); 7179 (59); 7593 (17); 8323 (43). Wurdack, J. J., 2339 (57); 2362 (37); 2493 (59). Wurdack, J. J. & L. S. Adderley, 43064 (35); 43153 (19);

43169A, 43169B (35); 43435 (19); 43592 (4); 43647 (25).

Wurdack, J. J. & J. V. Monachino, 39775 (35).

Young, K. & G. Sullivan, 663 (33).

Zak, V., 4093 (55). Zak, V. & S. Espinoza, 4671 (8); 4835 (50). Zardini, E. M., 7710, 7845, 7911 (56). Zardini, E. M. & C. Benftez, 3159, 3182, 3182, 3205,

3205, 3236, 3429 (56). Zardini, E. M. & L. Guerrero, 47737, 47746 (56). Zarucchi, J. L., 3031, 3054 (35). Zarucchi, J. L. & C. E. Barbosa, 3679 (35). Zarucchi, J. L. et al., 1830 (35); 3093 (22); 7120 (11). Zaruma, J., 695 (50); 810 (8). Zaruma, J. & A. Arguello, 453 (8). Zuleta, J., 42 (50); 45 (57); 50 (50); 142 (59).

INDEX OF LOCAL NAMES

aguacat6n, 45 aiju'ywahu, 119 ajua, 123 amarillo, 79 anis moena, 82 ant tree, 67 azywa'yw-pihun, 31

bastard silverballi, 59 bois marble, 95

canela, 74, 76 canela amarella, 117 canela frade, 117 canela garuva, 117 canela paluda, 117 canelao, 74 canoe tree, 42 casha moena, 45 cedre, 97 chaviaco negro, 76

cunchi moena, 121 cunshi moena, 45

dimukuima, 97

harige pisie, 63 ha-ro, 70 hioma cocuir-ey, 68 honcatohue, 105


inchaquito, 105 isma moena, 76 isma muena, 78 jigua, 78, 97 jigua de mierda, 27

ko-ma-nee-nee-ko, 76 ko-mwa-kb-ree, 42

las brisas, 70 laurel, 70, 117 laurel aguacate, 117 laurel babosa, 72 laurel blanco, 76, 92, 97 laurel blanco de orilla del rfo, 76 laurel bobo, 98 laurel bongo, 98 laurel carutillo, 72 laurel de babilla, 61 laurel de revalta, 76 laurel del rio, 76 laurel fino, 76 laurel hediondo, 70 laurel hormiguero, 62 laurel moroti, 117 laurel negro, 34, 63 laurel oriyera, 76 laurel plateado, 97 laurel rebalsero, 76, 97 laurelito, 76 laurier bord de mer, 95 laurier gris, 95 laurier gwa gwen, 95 laurier marbre, 95 laurier pete, 95 limon moena, 82 louro, 23, 57, 58, 66, 70, 121, 123 louro abacate, 45, 67 louro amarello, 109 louro branco, 105 louro canela, 76 louro cedinha, 97 louro cedro, 67 louro comum, 79

louro cururu, 70 louro de folha cinzenta, 105 louro de f6lha larga, 72 louro de f6lha peluda, 90 louro do igap6, 76, 121 louro dorado, 109 louro dourado, 109 louro flMr, 76 louro imbauba, 72 louro jambo, 67 louro pichuri, 105 Louro pirarucu, 24 louro preto do igap6, 121 louro roxo, 109 louro seda, 105 louro tambaqui, 76 louro vermelho, 90

mahamira, 119 mantaga, 34 mautaga, 34 moena, 66, 67, 90 moena amarilla, 69 moena blanca, 45, 114 moena callhuangia amarilla, 34 moena de altura, 45 moena de bajo, 76 moena de hoja grande, 72 moena hoja ancha, 105 moena negra, 34 moena rosada, 117 moenilla, 114, 117, 123 moenito amarillo, 48 moroman, 62 muena, 32,.42, 69, 70 muena amarilla, 67 muena blanca, 76 muneu-yek, 119

ocatoe, 70, 94 ocatue, 105

palometa micuna, 48 palta amarilla, 27

palta moena, 70 pampa muena, 78 pee-shee', 76 peroua-yek, 62 pisie, 63 plata gigante, 105 pucacuro caspi, 67 puspo moena, 68

roble, 117 roble anis amarillo, 70 roble fusi, 90 roble palta, 27 roblecillo puchirfn, 27

sacha muena, 117 sacha palta, 102 sanango, 76 shiringochy, 117 shisho moena, 66 shisho moenita, 66 siroewaballi oenilebobandikoro, 59 sweetwood, 95

temachi, 45 tikis, 70, 114 tinchi, 114, 117, 123 tinci, 70 tindu, 114 tunchi, 114

uchitinchi, 70 und yuwich, 70 urcuayua, 105

wakamei, 62 wau yuwich, 70 wild pear, 72

yacu muena, 76 yahuemomo, 94 yellow silverballi, 72 yiyiwa-ageyi, 45 yuwich, 110, 123

INDEX OF SCIENTIFIC NAMES

New names and combinations are in boldface and synonyms are in italics. Page numbers in boldface indicate primary page references.

Acrodiclidium geminiflorum, 125

Aiouea, 5, 16 impressa, 124 maguireana, 124, 132 tomentella, 124

Ampelodaphne, 3, 11, 16 arunciflora, 51 dasyantha, 60 macrophylla, 16, 57

Anaueria, 7 Aniba, 13, 72, 106

INDEX OF SCIENTIFIC NAMES 139

Aniba (continued) bracteata, 125 flexuosa, 42, 44 juruensis, 124 krukovii, 121 reticulata, 26, 28 tschudyana, 97,98

Aspidostemon, 7 Aydendron,

argenteum, 125 macrophyllum, 36

Beilschmiedia, 7 zapoteoides, 125, 133

Caryodaphnopsis, 7 Cassytha, 2, 7 Chlorocardium, 7 Cinnamomeae, 7 Cinnamomum, 5, Citrosma

paniculata, 184 Cryptocarya, 7,

hirsuta, 2, 114 pyriformis, 42 robusta, 112

Cryptocaryeae, 7

Dicypellium, 31

Endlichera, 3, Endlicheria

sect. Macrolocellata, 3, 5, 16 sect. Microlocellata, 3, 5, 8, 16, subgen. Ampelodaphne, 3,16, 32, subgen. Anosphaeria, 3, 16 subgen. Euendlicheria, 3, 16 subgen. Hemiajouea, 3, 16, 47 subgen. Ocoteopsis, 3, 16 acuminata, 3, 4, 14, 20, 65, 82, 117, 119*, 120, 121,

123, 127 Ampelodaphne species group, 3, 4, 5, 11, 13, 14, 33, 51,

57, 58, 59, 60, 64, 66, 69, 74, 78, 127 bracteata sub-group, 13, 51, 66 multiflora sub-group, 13

anomala, 3, 4, 5, 6. 13, 16, 17, 21, 74, 76*, 78, 92, 94, 109, 121, 127

anomala species group, 13, 14, 127 arachnocome, 4, 18, 49*, 51, 52, 53*, 54, 67, 127 arenosa, 13, 18, 54, 55*, 56*, 57, 67, 127 argentea, 17, 110, 111*, 112, 113*, 127 arunciflora, 3, 4, 6, 18, 49*, 51, 52, 54, 67, 127 aurea, 18, 106, 107*, 108*, 109, 127 balsamea, 123, 134 boliviensis, 114,117 bracteata, 3, 19, 61, 62, 63*, 65, 66, 67, 68, 127 bracteolata, 3, 16, 17, 64, 95, 96*, 97, 127 browniana, 5, 6, 11, 16, 20, 34*, 36, 37, 38, 127 browniana species group, 11, 14, 41, 45, 48, 51, 114,

127

Endlicheria (continued) bullata, 3, 8, 16, 21, 23*, 26, 28, 127 canescens, 4, 6, 7, 13, 16, 22, 67, 78*, 79, 80, 127 canescens species group, 13, 14, 82, 92, 127 chalisea, 5, 13, 16, 19, 66, 71, 72, 73*, 127 chrysovelutina, 4, 5, 11, 14, 17, 34*, 35*, 36, 126 citriodora, 4, 13, 22, 31, 80, 82, 83*, 127 cocuirey, 3, 13, 18, 62, 66, 67*, 68, 127 colombiana, 20, 38, 40*, 41, 127 coriacea, 5, 8, 19, 23*, 24, 25*, 126 debilis, 123 dictifarinosa, 3, 13, 18, 59*, 61, 62, 68, 127 directonervia, 13, 18, 57, 69, 70, 71*, 127 duotincta, 4, 13, 16, 78, 89*, 90, 91*, 92, 94, 127 dysodantha, 3, 4, 19, 42, 47, 48, 49*, 127 endlicheriopsis, 7, 79, 124 ferruginosa, 8, 22, 26, 28, 29*, 30*, 79, 126 formosa, 4, 16, 19, 33, 44, 45, 46*, 114, 127 glaberrima, 42, 44 glomerata, 3, 4, 7, 13, 16, 19, 26, 72, 73*, 74 goeldiana, 124, 130 gracilis, 3, 5, 14, 16, 20, 21, 32, 65, 117, 119*, 121,

127 grandis, 124 grisea, 118 griseo-sericea, 14, 17, 95, 99*, 100, 101*, 102, 127 guadaloupensis, 94, 95 hirsuta, 16, 114 impressa, 3, 124 jefensis, 11, 19, 37, 38, 39*, 40*, 127 juruensis, 124 klugii, 4, 6, 14, 17, 20, 108*, 109, 110, 112, 127 krukovii, 3, 14, 20, 121, 122*, 123, 127 levelii, 13, 18, 59*, 60, 61, 62, 127 lhotzkyi, 7, 17, 105, 108*, 109, 127 longicaudata, 3, 5, 8, 21, 27, 30*, 31, 119, 126 longifolia, 114, 117 lorastemon, 5, 13, 22, 77, 89*, 92, 93*, 94, 127 loretensis, 124 macrophylla, 13, 18, 56*, 57, 58, 59, 127 maguireana, 124 melinonii, 3, 13, 18, 59, 62, 63*, 64, 127 metallica, 4, 5, 6, 11, 14, 16, 17, 19, 20, 33, 34*, 36.

105, 123, 126 metallica aff. sp. 1, 11, 17, 134 metallica aff sp. 2, 11, 19, 135 metallica species group, 4, 11, 14, 126 Microlocellata species group, 8, 14, 31, 126 mishuyacensis, 5, 11, 19, 37, 40*, 41, 42, 127 multiflora, 18, 58, 59*, 60, 62, 63, 65, 127 nilssonii, 14, 20, 122*, 123, 127 oreocola, 13, 21, 83*, 86, 87*, 88, 127 panicularis, 114 paniculata, 2, 3, 4, 5, 6, 7, 14,16 20, 21, 48, 77, 114,

116*, 117, 119, 121, 123, 127 paniculata species group, 14, 127 paradoxa, 3, 16, 19, 45, 46*, 114, 127 poeppigii, 114, 117 punctulata, 3, 4, 5, 8, 19, 22, 23*, 24, 126 punctulata species group, 2, 8, 14, 126


Endlicheria (continued) pyriformis, 4, 16, 19, 27, 31, 37, 42, 43*, 44, 127 racemosa, 114, 117 reflectens, 3, 4, 13, 18, 32, 59, 63*, 64, 65, 127 robusta, 17, 20, 45, 112, 113*, 114, 127 rubra, 3, 13, 14, 22, 82, 83*, 84*, 127 rubriflora, 6, 8, 16, 19, 21, 23*, 26, 27, 28, 126 ruforamula, 3, 4, 14, 16, 17, 92, 102, 103*, 104*,

105, 109, 110, 127 sericea, 3, 7, 16, 17, 18, 94, 95, 96*, 97, 105, 109,

123, 127 sericea species group, 4, 13, 14, 38, 45, 77, 92, 94,

100, 102, 105, 109, 127 sprucei, 8, 13, 14, 22, 30*, 31, 32, 119, 126 szyszylowiczii, 13, 16, 21, 88, 89*, 90, 127 tessmannii, 7, 13, 18, 67*, 68, 69, 71, 127 tomentella, 124 tomentosa, 4, 21, 48, 49*, 50*, 127 trianae, 26, 28 tschudyana, 18, 97, 99*, 100, 105, 114, 127 verticillata, 3, 4, 7, 13, 18, 51, 62, 66, 67*, 68, 127 villosa, 58, 60 vinotincta, 3, 5, 13, 14, 21, 32, 83*, 85, 86, 88, 127 williamsii, 4, 5, 13, 17, 77, 78*, 90, 92, 127 wurdackiana, 26, 28 xerampela, 4, 13, 21, 78*, 80, 81*, 127 zapoteoides, 124

Goeppertia, 3, 16, anomala, 74 argentea, 125 cantagallana,1 14 caudata, 125 chrysophylla, 109 dysodantha, 47 geminiflora, 125 hirsuta, 16

var. coriacea, 114 var. glabrata, 114 var. hirsutior, 114 var. latifolia, 114 var. robusta, 114

longifolia, 114 multiflora, 3, 58, 60 panicularis, 114 polyantha, 74, 77 reflectens, 64 sericea, 94

var. bracteolata, 95 sprucei, 31

Huberodaphne, 3, 16, 31 longicaudata, 16, 30

Hypodaphnis, 7

Laureae, 7 Laurus, 5, 7

dysodantha, 47

Licaria, 13, 31, 128 cannella, 24

Lindera, 5

Mespilodaphne pyriformis, 42

Mezilaurus, 7

Nectandra, 4, 15 lucida, 114

Neocinnamomum, 7 Neolitsea, 7

Ocotea, 2, 5, 7, 8, 13, 14, 24, 124, 127, 132, 133, 134 cernua, 24, 128 cernua species group, 8, 23 debilis, 125 diffusa, 134, 136 discrepens, 13 endlicheriopsis, 13, 79, 124 helicterifolia species group, 15 indirectinervia, 13 lhotzkyi, 108 nigrescens, 136 pauciflora, 8, 24 punctulata, 22, 23 rufovestita, 13, 135 simulans, 74, 77

Ocotea complex, 8 15 Ocotea s.str, 7, 8, 15 Oreodaphne

colombiana, 40

Persea, 5, 7 Phoebe

impressa, 124 Pleurothyrium, 5, 15 Phyllostemonodaphne, 31

Rhodostemonodaphne, 2, 7, 8, 11, 13, 14, 15, 28, 33, 34, 36, 51, 84, 86, 105, 119, 124, 130

antioquensis, 8, 28, 129, 131 celiana, 13, 86 crenaticupula, 13, 33, 124 cyclops, 11 debilis, 124, 132 elephantopus, 8, 24 grandis, 11, 34, 36, 105, 124, 136 juruensis, 124, 132, 134, 136, 137 kunthiana, 13 laxa, 11, 14, 51 mirecolorata, 13 negrensis, 21 parvifolia, 31, 119 peneia, 11, 34 praeclara, 11, 34 recurva, 11, 13, 20 revolutifolia, 13, 22, 28, 30 rufovirgata, 13 sauilensis, 11, 34

INDEX OF SCIENTIFIC NAMES 141

Rhodostemonodaphne (continued) scandens, 26, 31 steyermarkiana, 13, 86 tumucumaquensis, 134

Rubiaceae, 3

Sassafras, 7 Schauera, 3, 16 Sextonia, 7

Strychnodaphne lhotzkyi, 108

Systemonodaphne geminiflora, 125

Umbellularia, 5

Williamodendron, 7

Endlicheria (Lauraceae)

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