~ A Taxonomic Treatment. of the Palm Subtribe Attaleinae (Tribe Cocoeae) SIDNEY F. GLASSMAN ILLINOIS BIOLOGICAL MONOGRAPHS 59 UNIVERSITY OF ILLINOIS LIBRARY AT URBANA-CHAMPAIGN BIOLOGY 9 § 200 Te Ly bd le © -) Pay : a “ Pe toe Ayia . = 4 Ya j i ts : 5 Alene j 4 i ’ ™ Digitized by the Internet Archive in 2011 with funding from University of Illinois Urbana-Champaign http://www.archive.org/details/taxonomictreatme59glas A Taxonomic Treatment of the Palm Subtribe Attaleinae (Tribe Cocoeae) SIDNEY F. GLASSMAN ILLINOIS BIOLOGICAL MONOGRAPHS 59 UNIVERSITY OF ILLINOIS PRESS _ Urbana and Chicago Illinois Biological Monographs Committee David S. Seigler, chair Daniel B. Blake Joseph V. Maddox Lawrence M. Page © 1999 by the Board of Trustees of the University of Illinois Manufactured in the United States of America Pi 5 A 32 1 This edition was digitally printed. This book is printed on acid-free paper. Library of Congress Cataloging-in-Publication Data Glassman, Sidney F. A taxonomic treatment of the palm subtribe Attaleinae (tribe Cocoeae) / Sidney F. Glassman. cm. — (Illinois biological monographs ; 59) Includes bibliographical references (p. _) and index. ISBN 0-252-06786-X (pbk. : acid-free paper) 1. Palms—Classification. 2. Palms—Classification—South America. lL. Title. II. Series. QK495.P17G548 1999 584'.5—dc21 98-58105 CIP Tee vial SF Abstract Detailed taxonomic treatments of all genera in the subtribe Aftaleinae are included in the present study. Attalea contains 21 species; 2 interspecific hybrids, A. x piassabossu and A. X voeksiz; and | intergeneric hybrid, x Attabignya minarum. Orbignya has 11 species; 1 interspecific hybrid, O. x teixerrana; 2 intergeneric hybrids, x Attabignya minarumand Xx Maximbignya dahlgreniana; | putative intergeneric hybrid, Ynesa colenda; and 1 unde- scribed putative intergeneric hybrid. Scheelea contains 31 species and 2 undescribed putative intergeneric hybrids. Maximiliana contains | species with 1 intergeneric hybrid, x Maximbignya dahigreniana, and | undescribed putative intergeneric hybrid. A total of 13 new species are described here (Attalea, 3, Orbignya, 1, and Scheelea, 9), as well as 3 new combinations and name transfers. The main emphasis of this treatment is taxonomic, but other system- atic aspects have been considered, such as leaf anatomy and chemotax- onomy as well as geographical and ecological distribution. Evolutionary relationships among all four genera, as well as alliances among species within each genus (including a tentative division into infrageneric catego- ries), are followed by a section on geographical distribution of species within each genus, a description of each genus, and keys to their species. Taxonomic treatments of all four genera and their individual species in- clude original as well as other pertinent publications, complete synonymy, designation of types, a detailed morphological description, a list of speci- mens examined, distribution and habitat, vernacular names, economic importance, a discussion of evolutionary relationships, and nomenclatural problems. A total of 10 tables, 201 illustrations, and 17 distribution maps are also included. Illustration and map numbers are listed as Figs. in bold type tor each species. Lists of doubtful and excluded species for each genus are also incorpo- rated into the text, as are acknowledgements, a list of references, and an alphabetical index to names of all taxa in the subtribe Attaleznae. Cladistic studies are not included in the present treatment due to in- complete diagnostic morphological data for a number of taxa. At this juncture I should point out that this work does not represent a conven- tional monograph or revision. However, the total amount of information contained in it, and found nowhere else, forms a basis for future study that should lead to a more complete monographic treatment. Therefore, the main purpose of this study is to lay the groundwork to stimulate further research (including the collection of additional specimens) of this com- plex group of palms. Acknowledgments I am indebted to the National Science Foundation for three grants (GB6899, GB3737, and BMS7509779) and to the Research Board and the Department of Biological Sciences at the University of Illinois at Chicago for additional assistance. These grants enabled me to take field trips to Brazil; visit herbaria in Europe, South America, and the United States; and study cultivated palms of various genera in the subtribe Attaleinae at Fairchild Tropical Garden in Miami, Florida. I would like to thank J. B. Harborne and C. Williams, both at the University of Reading, England, for carrying out chemotaxonomic studies on flavonoids; the late Timothy Plowman of the Field Museum for numerous valuable suggestions; and the late Rolf Singer of the Field Museum for revising the Latin descrip- tions of new species. Special thanks go to Larry Noblick of Montgomery Botanical Center for important field information on the distribution of Attalea in Bahia; to Margaret Kleist and Joan Cureton of UIC for prepara- tion of the original manuscript; to my son Bob for his computer exper- tise and excellent editorial assistance; to my wife, Ida, for continuous sup- port in the preparation of this manuscript; to Marlene Werner of the Field Museum and Janice Rajecki of UIC for preparation of the illustrations; to several people at the Instituto Botanica, Sao Paulo, especially A. Milanez and D. Vital, for making available to me transportation, guides, drivers, and herbarium facilities during my trip to Brazil in 1976; to the late Pedrito Silva of CEPLAC in Salvador for facilitating the shipment of palm speci- mens to the United States and for useful information on the disposition of Gregorio Bondar’s palm collections; to Leticia Faria of the Federal University of Salvador, Edna Olivera of EMBRAPA in Salvador, Luis Silva of CEPEC in Itabuna, and Luciano Lima of CEPLAC in Petrolina for making available transportation, drivers, and herbarium facilities (all of which was coordinated by Larry Noblick) during our trip to Bahia, Bra- zil, in 1988. I am also grateful to the curators of the herbaria cited for the privilege of studying their specimens. dasa: hiaewed Topsail Eee AEA Wadi thy «esd macypl ie 0 ee » o> wf oh |. ee 4 +A A > —— lar ‘tl al re a oe ‘ar ae ‘ ree rag vert: dol vee ats | te tc ig] siti a ot) brut Pnaodl onsen hi oat (eR IOS OE Dew eves % Oyr i ty loniiDlo Wire ens? migotold Tae _ & ogr_blen sie) et feldnns aig seo orn seins Lets i ants bik oot ANH alt bie nairtiA thuve scorn pues io ee sends Ati nd Mrofeg eUoiey to ented bode : ‘ daaga mn) til cited | ebitolt Veil ty) ceobrged ts “gone fren realy te wsienass Wint) aio ox Wed 2 ew Pw pew ial way sual ah shim riya ay eothoata don ar casein’ atively bres RO MSg we ahtalti ar erin coun wit watse bist eel a1 ayPrenb ine! seh) grein earsew M bight ody te yee Hest & “ remeymolt te dodo el ot mq aatunits gage woivnze onli Ye wolarstinell: on) co veMigniobrliphehi Jemtednd 18 yan? hea ateuutep iy WA DIV ier eotossh) rts of bus raicilaregeM 0) ahi o— Magu iiugines ad wi dig ate gen ot pqiinaiinet Bogie ort aque tinouniisived co us atier yen whan ieiat LaMibs inelinaes® hituhid meds ey verte scout aneen theplavedann cuts zis to filet al ‘ersitaingl{) alt lo coder it A te ine dine! bee onl warreh #. «lutoasey pene evi aii vO ? 4 on HA ‘ = - fy y ‘a ) ‘ ‘at (ta.4 ke’ Cie ie Va hy n is ig Ve teagt ip Pio _ ie PW ts be i “| hat tes Li. yn via ure ‘ * si ' n ¥ . Ts Th te " M anciedened Piy ween a ee .* 7 Tatilg 5 : Jdor vedio a A es | ey 4 wie vi | Hy) (PN ie ri ut" 7, rd ” R hes ty og ie ay a rh 4 mark] " A : a “ Wi D \ *, A a hy a 4 ayes! | 7 | ules Ak Introduction The purpose of this study is to correlate all available information pertain- ing to the systematics (mainly taxonomy) of the subtribe Attaleinae. It is my hope that the present treatment will stimulate further research on this group of palms as it has in some of my other taxonomic treatments (Butia in 1979, and Syagrus in 1987). Moore (1973) classified palms into 15 different groups, which in turn were divided into alliances and units. The four genera being treated here (Attalea, Orbignya, Scheelea, and Maximiliana) were placed in the cocosoid group in the Cocos alliance within the Attalea unit. In a more recent for- mal classification of the family Palmae (Arecaceae) by Dransfield and Uhl (1986), the above genera were included in the subtribe Attaleinae within the tribe Cocoeaeand subfamily Arecoideae. Attaleinae differs from the other subtribes in Cocoeae by having two kinds of inflorescences, usually on the same plant: a staminate, usually with staminate flowers only; and an an- drogynous inflorescence, in which a pair of staminate flowers forms a series of triads with one pistillate flower on the proximal part of each rachilla, and on the distal portion of the same rachilla there are only stami- nate flowers. All other subtribes in Cocoeae (Beccariophoenicinae, Butiinae, Elaeidinae, Bactridinae) usually have only androgynous inflorescences. The four genera in subtribe Attaleinae can be differentiated by the fol- lowing key: 1. Anthers coiled, petals of staminate flowers divided into 4 groups of spe- cies: (1) 3 in number and spatulate in shape; (2) 1-8 by fusion and broad below with notched or coarsely toothed tips, frequently with ad- ditional lanceolate petals, rarely with only lanceolate petals (up to 8 in number); (3) 1 species with 3 petals with fleshy hooked tips; (4) 1 spe- cles with curved lanceolate petallss x spor cles quieren hyoe oration we aqeyerchs Ze A Taxonomic Treatment of the Palm Subtribe Attaleinae (Tribe Cocoeae) 1. Anthers usually straight, not coiled, petals of staminate flowers usually 3 in number, lanceolate or more or less linear in shape, with entire tips 2. Petals of staminate flowers, more or less linear, somewhat fleshy, usu- ally convex outside and grooved inside or more or less terete and SIMIAN fete oa 214 3 & eke eee ee ee Scheelea 2. Petals of staminate flowers lanceolate, usually flattened, not fleshy 3. Petals usually much longer than stamens ............ Attalea 3. Petals usually much shorter than stamens, which are consistently Gaim number 2. icc 6 eet tre ee eee ae ee Maximiliana Henderson and Balick (1991), Henderson (1994, 1995), and Hender son, Galeano, and Bernal (1995) have concluded that subtribe Attaleinae consists of only one genus, Aftalea, because several intermediate forms obscure supposed differences between genera. In the latter article, all published names of the 4 genera in Attaleinae are lumped together; 29 recognized species of Attaleaand 4 hybrids are listed but at least 17 of these names are considered to be synonyms or species dubia by me. On the other hand, in the subtribe Attaleinae I recognize a total of 66 species (includ- ing 13 new species) and several interspecific and intergeneric hybrids within 4 different genera. I disagree with this lumping concept. As long as I can recognize 4 distinct genera based mainly on differences in the staminate flowers, I cannot accept 1 genus to cover all taxa in the Attaleinae. Most intermediate forms can be treated as either (1) intergeneric hybrids (e.g., x Maximbignya dahlgreniana, x Attabignya minarum, and Ynesa colenda); (2) ancestral types that are differentiating at the extremes of their ranges (currently, I cannot cite any examples of this possibility); or (3) represen- tatives of subgeneric types within a genus (e.g., O. luetzelburgzi, O. polysticha, O. phalerata, and O. crassispatha). All four species of Orbignya are differ- entiated by their petals in the staminate flowers (see “Key to Species of Orbignya”), but are held together in one genus by the coiled stamens. I believe that the subtribe Attaleinae is presently a highly evolving group of palms with many diverse characteristics. Further research, such as more collections and DNA analysis, may lead to a less controversial classification. Lumping all of the genera into one genus and the wholesale reduction of a large number of species to synonomy ignores the problem and results in a simplistic classification. Introduction 3 History of Subtribe Attaleinae Attalea Attalea (named after Attalus I, king of Pergamum, Greece, 241-197 B.c.) was first distinguished as a genus by Kunth in Humboldt, Bonpland, and Kunth (1816), in which he described one species, A. amygdalina. Close- ly related genera were later described by Martius in 1826 (Maximiliana) and 1837 (Orbignya), by Karsten in 1857 (Scheelea), by Dugand in 1940 (Parascheelea) , by Cook in 1942 ( Ynesa), and by Bondar in 1957 (Markleya). Since its original description, Attalea has been treated taxonomically by Martius (1824, 1826, 1844, 1845, 1853), Karsten (1857), Drude (1881), Barbosa Rodrigues (1903b), Burret (1929a), Bondar (1942a, 1942b, 1964), Dugand (1953, 1954), Wessels Boer (1965, 1972, 1988), and me (1977a). Drude (1881) divided the genus into three sections: (1) Attalea verae (in- cluding six species with staminate flowers having flattened petals), (2) Cylindrostachys (containing only A. nucifera, characterized by having its staminate flowers arranged all around the rachilla) and (3) Pseudoscheelea (containing A. princeps and A. phalerata, which later were transferred to the genus Scheelea). Burret (1929a) also divided Attalea into three sections: Euattalea (including 15 species split into two groups based on size of the trunk) and two other sections differentiated by the arrangement of stami- nate flowers on the rachillae, size of trunk, and amount of fibers in the endocarp. Bondar (1942a, 1942b, 1964) described several new species and constructed keys to 16 species found in Brazil. Wessels Boer recognized 7 species from Suriname (1965) and 14 species from Venezuela (1972, 1988), but within the genus Aftalea he included other closely related gen- era, Maximiliana, Orbignya, Scheelea, Markleya, and Parascheelea. In 1973, Moore considered all of these except Markleya as separate genera within the Aitalea unit and listed 30 species for the genus Attalea; In my 1977a article, I did a preliminary taxonomic study of the genus Attalea in which I tentatively recognized 21 taxa; Balslevand Henderson (1987) transferred Ynesa colenda to genus Attalea. Orbignya Martius established the genus Orbignya in 1837a (named after A. d’Orbigny, a French collector in the 1820s and 1830s), but no species were described in this article. The first two taxa (O. phalerata and O. humilis) were delineated in 1844 by Martius. Other species were described or trans- ferred from other genera (mainly Attalea) by Drude (1881), Barbosa 4 A Taxonomic Treatment of the Palm Subtribe Aftaleinae (Tribe Cocoeae) Rodrigues (1879, 1888, 1891b, 1898, 1903b), Burret (1929a, 1930, 1932, 1940), and Bondar (1954a). Burret (1929a) did a comprehensive study of Orbignya, recognizing 19 species. The genus was divided into three sections: Distichanthus Burret, Pleiostichanthus Burret, and Spirostachys Burret. In the first two sections the staminate flowers are arranged along one side of the rachilla (and these in turn are differentiated by abundance or scarcity of fiber clusters in the endocarp), while the arrangement of staminate flowers is spiral in section Spirostachys. In addition, a partial key to sections and species was included. Of 13 species partly keyed out in the first section, 8 are listed as unknown or doubtful by Burret; of 4 listed in the second section, 2 are considered doubtful; while in the last section the 2 species are not keyed out, but 1 of these is doubtful. Bondar (1964) listed 14 species in his treatment of Brazilian Orbignya, but no keys were included; Wessels Boer (1965, 1972, 1988) submerged all species of Orbignya (as well as other genera in the subtribe Attaleinae) from Suriname and Venezuela under the genus Aftalea, sensu lata; in a preliminary taxonomic treatment of Orbignya, I included keys and cited specimens of 21 recognized taxa (1977b); and Anderson and Balick (1988) did a taxonomic study of the Babassi complex in which two species and hybrids of Orbignya were discussed. Scheelea Scheelea (named after C. W. Scheele, a Swedish pharmacist and chem- ist, 1742-86) was established as a new genus by Karsten (1857). In this ar- ticle he described four new species and transferred three others from Attaleaand Maximiliana. Karsten (1861, 1866) also published illustrations of the species he described. During the next several decades, other taxa of Scheelea were described or transferred from other genera (mainly Attalea) to Scheelea by Barbosa Rodrigues (1891a, 1891b, 1894, 1898, 1899, 1903a, 1907), Hooker (1897), Beccari (1916), Burret (1929a, 1934b, 1940), Bailey (1933, 1947), Bartlett (1935), and Dugand (1959). Undoubtedly, the most comprehensive study of Scheelea was done by Burret (1929a). In the same article he also treated the genera Attalea, Orbignya, and Maximiliana. Besides describing 13 new species of Scheeleaand transferring 8 others to the genus from Attaleaand Maximiliana, he divided Scheelea into two sections, Synalphocaryum Burret and Dialphocaryum Burret. In the first section (characterized by having very large endocarp fibers dis- Introduction 5 tributed in dense clusters and 1-5 closely arranged pistillate flowers on each androgynous rachilla) Burret included 23 species, and in section Dialpho- caryum (differentiated by having endocarp fibers about one-half as small and distributed in smaller clusters and 5-many loosely arranged pistillate flowers per androgynous rachilla) 15 taxa were listed. Burret also con- structed a partial key to the species within the first section and a more or less complete key to those in the second section. In the same article Burret emphasized the pitfalls in undertaking a comprehensive study of Scheelea because a number of taxa are based on incomplete descriptions as well as inadequate material. Wessels Boer (1965, 1972, 1988) treated all species of Scheelea, as well as other closely related genera (Attalea, Orbignya, Maxi- miliana, Parascheelea, and Markleya), as part of the genus Aittalea, sensu lata. In my 1977c article, I published preliminary studies of Scheelea; and in my 1977a article, I discussed all six genera in the Attalea complex. One important study not discussed in my 1977c article is the FAO Oil- seed Mission for Venezuela (Claassen, Jenkins, and Markley, 1949). A to- tal of five species of Scheelea (S. macrocarpa, S. macrolepis, S. maracaibensis, S. humboldtiana, and S. passargei) along with distribution maps were men- tioned as possible oil producers. It was estimated that one of these palms (S. macrocarpa) was represented by almost 6 million trees. Unfortunately, there is no record of where, if any, herbarium specimens were deposited. Therefore, identifications and distribution records of these species can- not be accurately verified. Maximiliana The genus Maximiliana (named after Maximilian Joseph I, king of Ba- varia, 1756-1825), was established by Martius (1824), but no species were described until 1826 (M. regiaand M. insignis) and 1844 (M. crassispatha). Other species of Maximiliana were described or transferred from other related genera to Maximiliana by Karsten (1857), Grisebach (1864), Spruce (1871), Barbosa Rodrigues (1875, 1891b), Wendland (1878), Drude (1881), and Burret (1929a). Kuntze (1891) erected the genus Englerophoenix as a new name for Maximiliana Martius 1824, which was published as a homonym for Maximilianea Martius 1819 (in the family Cochlospermaceae). Maximiliana Martius 1824 was later given conserved name status (see Voss 1983). Some other authors (Barbosa Rodrigues, 1903b and Bondar, 1938, n.d.) consid- ered Englerophoenix as a separate, distinct genus. 6 A Taxonomic Treatment of the Palm Subtribe Altaleinae (Tribe Cocoeae) Drude (1881) divided Maximiliana into section Eumaximiliana (includ- ing M. maripaand M. regia) and section Scheelea (including M. insignis and M. tetrasticha) based on whether the stamens were longer than the petals in the staminate flowers. Barbosa Rodrigues (1903b) recognized two sec- tions (Inayaand Jnayay) under Englerophoenix, primarily based on the pres- ence or absence of trunks. The most comprehensive study of Maximiliana was done by Burret (1929a), in which he also treated Attalea, Orbignya, and Scheelea. He keyed out nine species of Maximiliana in two sections, Exanthera and Cryptanthera, distinguished by having anthers extending a considerable distance beyond the petals rather than anthers included in the petals or scarcely longer than the petals. He also excluded the two sections identified by Barbosa Rodrigues. Burret (1953) distinguished Maximiliana from its closest relatives, Attalea, Scheelea, and Orbignya, by the following: (1) the endocarp is thin rather than thick at the location of the embryo pore, (2) the embryo pore is covered with an operculum instead of devoid of an operculum, and (3) the en- docarp lacks fibers rather than includes fibers. Wessels Boer (1965, 1972) submerged all species of Maximiliana, as well as its three closely related gen- era, under the genus Attalea, sensu lata, and I (1978a, 1978b) published preliminary studies of the genus Maximiliana but only one species was even- tually recognized. Henderson (1995) and Henderson, Galeano, and Bernal (1995) reduced M. maripa to synonomy under A. manpa. Systematic Studies of Subtribe Attaleinae As with other groups of palms, individual genera and species may show a range of morphological variation depending on the age and kind of habi- tat of individual palms. Examples: Some normally acaulescent palms may exhibit a short trunk when older (e.g., A. humilis); some palms that are arborescent when mature may remain in an acaulescent juvenile state for a long period of time due to growing in adverse habitats (e.g., A. funifera). The leaves are pinnate, and the middle series pinnae are either in clus- ters of 2-several or regularly arranged and usually 1-several cm wide. Ju- venile palms often exhibit longer petioles and leaves and sometimes wider pinnae than mature plants. Petioles vary less in length in adult plants and are consistently absent or very short in some species. Mature palms grow- ing in wetter habitats may have leaf and inflorescence parts with larger dimensions than those growing under optimal conditions. Sterile bracts are usually several mm to several cm thick and sulcate on the outer sur- Introduction 7 face. Individuals usually have separate staminate and androgynous inflor- escences on the same plant. Sometimes individuals may have only stami- nate inflorescences or predominately pistillate inflorescences on the same plant. Staminate rachillae and flowers are usually longer on the lower parts of the rachis and rachillae than on the upper parts. Basically, four distinct kinds of staminate flowers exist, based on the shape of their petals and anthers (see the key above). The size of individual staminate flowers within each genus does not vary much; however, stamens vary in number (6-75) in species of Attaleaand Orbignya, but are constant (6) in species of Scheelea and Maximiliana. Anthers usually vary within a few mm in individual speci- mens; these are coiled in species of Orbignya, but are usually straight in the other three genera. The petals are usually three in number, but some species of Orbignya have as many as eight. The shape of the petals in each genus is delineated in the key above. Sepals are usually much shorter than petals and three in number. The length of pistillate rachillae is fairly con- stant in individual species, and pistillate flowers do not vary much in size. The number of stigmas ranges from three to eight. Fruits vary in size and shape based on their degree of development. Some species are consistently one-seeded, others have as many as eight or more seeds. Younger fruits are usually narrower and smaller, and fruits of many species turn orange or yellow at maturity. Except for some chemotaxonomic studies in the subtribe Attaleznae, previous systematic investigations based on other than gross morphology have not yielded much information for phylogenetic studies. Very few chromosome counts have been made for this subtribe. Darling- ton and Wylie (1945) reported the following: A. cohune, A. spectabilis, and O. lydiae are n= 16, but all three taxa belong to genus Orbignya; and Read and Moore (1967) report n= 16 for A. alleniz. Very little or no informa- tion has been published about polyploidy, aneuploidy, breeding experi- ments, or population studies. Comprehensive palm pollen studies have been carried out by Punt and Wessels Boer (1966), who surveyed 20 species in the subtribe Attaleznae, and by Thanikaimoni (1971), Sowunmi (1972), and me (unpublished data), in which I treated a number of taxa in this subtribe as part of a broader survey of palms. In all cases, the results were similar. Due to the lack of distinct differences in pollen structure among taxa, palynology appears to be a poor tool for studying evolutionary relationships among genera and among species of each genus in the subtribe Altaleinae. I reached the same conclusions in my unpublished palynological survey of 8 A Taxonomic Treatment of the Palm Subtribe Attaleinae (Tribe Cocoeae) genera in the Syagrus group. Chemotaxonomic surveys of genera in the subtribe Attaleinae were carried out by Glassman et al. (1981), who stud- ied the leaf flavonoids and lipids, and Williams, Harborne, and Glassman (1983, 1985), who surveyed the leaf flavonoids of various palms in the tribe Cocoeae. No major differences were found among species of Attalea and Orbignya. Chemotaxonomic analyses of flavonoids and lipids by Glassman et al. (1981) showed some differences among the three species of Scheelea tested; however, later flavonoid studies by Williams, Harborne, and Glassman (1983, 1985) of several species of Scheelea were inconclusive and somewhat puzzling. Five of the seven taxa studied are confined to Central America; and three of these, S. rostrata, S. preussii, and S. zonensis, were reduced to one species (S. rostrata) because I could not find any major morphologi- cal differences among them. Results showed that S. rostrata differed from S. preussui and S. zonensis by three and four flavonoid compounds, respec- tively, whereas it differed from S. lundelliiand S. liebmannii, the other dis- tinct Central American species, by only one and two compounds, respec- tively. Results of major significance were found in the genus Maximiliana. Chemotaxonomic studies of four different collections (labeled different species) of Maximiliana showed the presence of flavonoid sulphates in two of the four collections (Williams, Harborne, and Glassman; 1983). All four collections also revealed the presence of flavonoid C-glycosides and tricin. This combination of characteristics separated these specimens (one spe- cies) from all other species of Orbignya, Scheelea, and Attalea treated in the survey. I carried out a leaf anatomy survey of more than 100 collecting num- bers in Attalea and related genera. Cross sections of middle series pinnae were prepared in the same manner as described for Syagrus and Butia (subtribe Butiinae) in my 1972b, 1979, and 1987 publications. It was not possible to construct a key to all genera and species in the subtribe Attaleinae as I did in the subtribe Butiinae, based on leaf anatomy alone. Only a handful of species in each genus could be keyed out. No clear pattern between related groups of species could be established. When a key was attempted for all taxa, very few differences could be found to dis- tinguish a large percentage of the species. The relatively few species that could be keyed out were frequently those in different genera occurring in the same couplet or distantly related species in the same genus. The usefulness of the leaf anatomy survey, therefore, lies in the distinct char- Introduction 9 acteristics of certain isolated species of Attalea, Orbignya, and Scheelea, which adds to the morphological information. Specifically, in a survey of 20 different collections of Orbignya and its hybrids, most of the species showed similar or the same leaf anatomy pat- terns, whereas some others had distinct patterns (e.g., O. cwatrecasana, O. luetzelburgu, and x Maximbignya dahligreniana). Even though no related groups of species could be distinguished, the survey was useful because it showed that x Maximbignya dahlgreniana and O. luetzelburgii appear to be distinct and that other isolated species could be differentiated by their leaf anatomy. Cross sections of middle pinnae of 10 specimens of Maximiliana (identified as several different species) did not reveal any unique combi- nation of characteristics that could distinguish them from other genera in the subtribe. There is a certain amount of variability in the specimens observed, but most of them have elongated midribs, divided expansion cell tissue, common or very common distribution of adaxial nonvascular fibers, and frequent or common distribution of abaxial nonvascular fibers. Fossil fruits of cocoid palms have been reported from the Eocene from several different localities by Moore (1973); however, none of these can be definitely attributed to any genera in the Attaleinae. Unfortunately (as stated in the abstract), a cladistic analysis of the subtribe Attaleinae was not undertaken because of incomplete diagnostic morphological data for certain taxa. It is my hope that future research and new collections will fill in these morphological gaps and eventually lead to a more complete monographic treatment of this group. a | Ae! ee Ve Tew VET f 4 ‘ - 2.) ” 2 E YP Wee ; ‘i ‘ Pan 4 on ic = . oy oan peti yay . ¥s) hel i vin nie % Vay Sct al av ibe ie coreg oe as, ites a sneth ee ii ath fad ate ila a ae er. Eee mae 1) id i Nadal fet on fuged i Te A, 7 sation! Lagi sty, Pas A PAPUTE : oc fags + rane’ site ¥ 7 i . sid a o pair x dst sagt ys byse any wd - eiey is Das giclee Ky HSER: te id a ee 34 Pele Loe 14s dean Lange es He sil) ie 9 inc ssvesraiont Tals h ic) Md ro i eed SORE rhs Sha) 274 at 19 Be ae 1 ee 4 ia Jae sea ies} ia ‘tow as ne LaLa 7) fey ci / e " at: ‘9 ‘bioters a hates io satel oi) th garg 21 Peet Fay < , > Pans F a ‘ ae eee qc am Nes yeicery ied joker ats bei TO ely “abate ; ary bya : 7 , Sait | i pA eta y PItsiAt F iy b sey: ff ‘ af ae oh is aa 1a) P teh Loc ‘7 uy Gite Lp : Vie + WBS: Hite a mets; a Ts y eh x) ‘inns is hee hy “at ity? fiw rfon Gre: Trost: A K mE ee eae - a mY REE seen Tid, leggy ee el ai ke nypwere’; % a v ; Pew 7” rom 5 i é :- >y Frao } feet ber : BAP G)ict jin * Fe Por dmg eye ase > vhs at . § lee Ee Mahan) ove en pif ty * t F r ; Lh ey ry a j O24), . . & wey Ase: Y (ghags' dx ’ ‘ 4 ail ve ogg) i, t : 4 ‘Tas’ se ‘i i ei . te | Pe aiegss 1. 08>, cpl ai ied aie od laf wre ae Pee |, ni eve eae rets iene “8 te! y wD ¥ OM od TM a ea Rie Ae ; S| Carr ; ' m yw age) é ieee Til % ee ee ee A d , - tat Magen ast * bev? tedeidaeert ht ied tdgere ity | loo hea 24 PAS ; an % th ie ce j i : z ‘ : 7 P Wk , ne 4 y bY 5 | At ) enn Genus Attalea Geographic Distribution of Genus Attalea The genus Aftalea is almost entirely South American in distribution (see table 1). Its center of distribution appears to be eastern Brazil, specifically in the state of Bahia. Of the 24 taxa (21 species, 2 interspecific hybrids, and | intergeneric hybrid) recognized in this treatment, 17 are native to Brazil, 5 to Colombia, 2 each to Peru and Panama, and | each to Venezu- ela and Paraguay. The greatest concentration of Brazilian species occurs in Bahia (10); others are in Minas Gerais (4); Goias and Sao Paulo (3 each); Espirito Santo, Rio de Janeiro, Mato Grosso, Alagoas, and Sergipe (2 each); and Santa Catarina, Paraiba, and Pernambuco (1 each). Only 3 taxa are found in the Amazon region of Colombia, Peru, and Venezuela. In contrast to genus Syagrus (Glassman, 1987), most members of Attalea inhabit mesic inland or coastal forests (Brazil: 12, Colombia: 3, Peru: 2, Panama: 2), whereas several others are found in drier habitats such as cerrados, campos, roadsides, savannas, and rocky hillsides with limestone (Brazil: 4, Colombia: 2, Venezuela: 1, Paraguay: 1). Some of the species have limited or restricted distributions, and some of these are known from a relatively few collections. At least three species (A. burretiana, A. funifera, and A. humilis) are found mainly in the Atlantic coastal forest of Bahia, where they are fairly common, with the latter species extending into ad- jacent Espirito Santo, Rio de Janeiro, and Sao Paulo. Some species have a limited distribution in parts of central and western Bahia but are common in these small areas (A. pindobassu, A. seabrensis, and A. barreirensis); oth- ers from Goias, A. brasiliensis and A. exigua (which are less common), were collected from relatively few areas. With further exploration, however, some of these species eventually may have their known ranges extended. Perhaps the most common and widespread species in Brazil is A. geraensis, which forms dense stands in cerrados and disturbed areas in parts of Minas 12 A Taxonomic Treatment of the Palm Subtribe Altaleinae (Tribe Cocoeae) Gerais, Sao Paulo, Goias, Bahia, and Mato Grosso. Another fairly common species is A. apoda (A. camposportoana), which has been observed in sev- eral localities in Minas Gerais. The most common species of Attalea out- side of Brazil are A. allenii, collected from several localities in Panama and a number of provinces in Colombia, and A. ferruginea, known from a num- ber of localities in the Venezuelan Amazon region and from Colombia and Peru. A. nucifera also seems to be fairly common in several provinces in Colombia; but A. septuagenata is apparently known mainly from its type locality in Colombia. Table 1. Geographic Distribution of Attalea Species Country and State Habitat A. allenii Panama: Colon, Canal Zone, Primary and secondary forests, Comarca de San Blas, Bocas along streams, hills, and ravines del Toro; Colombia: Valle, Antioquia, Bolivar, Narino, Choc6é A. amygdalina Colombia: Valle, Caldas, Forests along streams in mountain Quindio, Antioquia, Risaralda areas; most of former habitats converted to coffee plantations A. apoda Brazil: southern Minas Gerais Common in primary and second- from Serra de Mantiqueira to ary forests, areas along hillsides, Copelinho and Serro de Palacio and in cerrados A. barreirensis Brazil: western Bahia Cerrados A. brasiliensis Brazil: Goias, Fer Cal area Remnant of climax deciduous of Brasilia upland forest in limestone derived soils A. burretiana Brazil: eastern Bahia and Remnants of Atlantic coastal Sergipe forest, transitional forest, and Mata Cipo A. compta Brazil: western Minas Gerais, Forested areas mostly west of Rio Sao Francisco A. dubia Brazil: Espirito Santo, Rio Atlantic coastal forest de Janeiro, Sao Paulo, Parana, and Santa Catarina along the coast A. exigua Brazil: Goias, Minas Gerais, Campos and roadsides Bahia, Mato Grosso, Maranhao? A. ferruginea Venezuela: Amazonas, Bolivar; Savannas and open mesic forests Colombia: Amazonas, Caqueta, in sandy soils Guainia, Vaupes; Peru: Loreto; Brazil: Amazonas Genus Atialea Table 1. Cont. Species A. funifera A. geraensts A. humilis A. iguadummat A. nucifera A. oleifera A. X piassabossu A. pindobassu A. salvadorensis A. seabrensis A. septuagenata A. tessmannii Country and State Brazil: coastal Bahia, Alagoas, Sergipe Brazil: Minas Gerais, Sao Paulo, Mato Grosso, Goias, southwestern Bahia; Paraguay: Cordillera Brazil: eastern Bahia, Espirito Santo, Rio de Janeiro, Sao Paulo Panama: Colon, Comarca de San Blas Colombia: Santander, Narino, Bolivar, Norte de Santander, Tolima, Cundinamarca Brazil: Paraiba, Pernambuco, Alagoas Brazil: eastern Bahia Brazil: central Bahia Brazil: eastern Bahia, near Amelia Rodrigues Brazil: central Bahia Colombia: Amazonas, endemic to Rio Miritiparana area Peru: Loreto, Madre de Dios, Ucayali; Brazil: Acre 13 Habitat Sand dunes and Atlantic coastal forest Cerrados (fine sandy fertile soils) and roadsides Coastal forest, wet restingas Extremely wet primary forests, along streams, and on ridges and slopes Dense forests Atlantic coastal forest Atlantic coastal forest Transitional mesophytic forest Secondary forest Transitional mesophytic forest Forested areas along rivers Flood-free and seasonally inundated forests Lleras, Giacometti, and Coradin (fig. 3, 1983) illustrated a distribution map of 34 species of Attalea extracted primarily from the literature. A to- tal of 12 of these taxa have been treated by me either as synonyms or uncertain species or have been transferred to other genera. Relationships within Genus Attalea Alliances or relationships between taxa cannot always be accurately deter- mined because descriptions of several species are incomplete; i.e., either the androgynous inflorescence with its pistillate flowers or the staminate inflorescence has not been collected or described in the field. In addition to this, some species are known from only one or two collections. As ex- 14 A Taxonomic Treatment of the Palm Subtribe Attaleinae (Tribe Cocoeae) plained earlier, this lack of diagnostic data is the main reason for not undertaking a cladistic analysis. It is possible that a few of these taxa are actually narrow endemics, but it is likely that with further exploration the ranges of these as well as other species may be extended and morphologi- cal gaps will be filled in. Hence, it is important that additional collections be made before the habitats where these palms grow are completely de- stroyed because of agricultural and lumbering practices. A number of botanists are currently making extensive palm collections throughout the state of Bahia as well as in other Brazilian states, which should be impor- tant in clarifying relationships between certain poorly known species. Although leaf anatomy proved to be an important systematic tool in my other studies of Syagrus and allied genera (1972b, 1978c, 1987) and Butia (1979), for Attalea it has not provided much additional data that can be correlated with the morphological characteristics. In the following outline, I have tentatively divided the genus into infrageneric categories. The larg- est subgeneric division is based on clustering of pinnae. Seven species (A. exigua, A. barreirensis, A. allenii, A. funifera, A. tessmannii, A. seabrensis, and A. dubia) have their middle series pinnae arranged in clusters of 2-5, whereas middle series pinnae of the remaining 14 are regularly arranged. In A. seabrensis and A. X piassabossu, however, middle series pinnae are mostly regularly arranged, but only the lower one-third to one-half of the rachis has clustered pinnae. Of the taxa with clustered pinnae, three are acaulescent, while the others are arborescent. A. barreirensis may be more closely allied to A. allenii than to A. exigua from central Brazil. A. exigua differs from the other two in having much longer pistillate rachillae and a distinct leaf anatomy pattern (abaxial nonvascular fiber bundles [NVF]) attached to the base of veins instead of alternating with the veins (see fig. 75). On the other hand, A. alleniiis distinct from the other two species by having petals of staminate flowers with punctate glands and is geographi- cally disjunct. Concerning the arborescent taxa, A. funiferais one of the most distinct, apparently with few close relatives, because of its unusually long petiolar fibers and grooved fruits with a very thick mesocarp and endocarp with- out visible fibers; A. tessmannii is probably without close relatives because it is the only species of Attalea with clustered pinnae in which the stami- nate flowers completely encircle the staminate rachilla instead of being inserted on only one side of the rachilla. One other species of Altalea, A. iguadummat, has spirally arranged staminate flowers, but does not appear to be closely related to A. tessmannii. A. x piassabossu is a hybrid between Genus Aittalea 15 A. funifera and A. burretiana; and A. x voeksi, between A. funifera and A. humilis; A. seabrensis seems to be closely related to A. pindobassu. Taxa with regularly arranged pinnae are more difficult to characterize; nevertheless, there seem to be several distinct species or groups of related species. As with the subdivision having clustered pinnae, these members can also be divided into palms with trunks and palms without trunks. The acaulescent species are separated into two groups based on their number of stamens. The two taxa with more stamens (12-52) are A. amygdalina and A. ferruginea, but each has several distinct characteristics. Concern- ing the group of four taxa with fewer stamens (6-10), A. iguadummat and A. nucifera appear to be the most distinct because of their rather special- ized features, such as spirally arranged staminate flowers and staminate flowers with reddish glandular hairs, respectively. The remaining taxa, A. humilis and A. geraensis, appear to be allied to one another, although A. humilis has more distinct characteristics than A. geraensis. Regarding the eight remaining arborescent members, A. septuagenata is easily differentiated from the others by an unusually high stamen num- ber in each flower (60-75). A. pindobassu, A. burretiana, and A. seabrensis are apparently closely related because they have fairly wide middle series pinnae, relatively long staminate rachillae, and staminate flowers similar in size. They differ primarily in the arrangement of the lower pinnae along the rachis. A. oleifera and A. compta are probably closely related because they have pinnae of similar width and staminate rachillae, staminate flowers, and fruits of similar size. They differ mainly in the number of seeds in their fruit. A. salvadorensis may be a hybrid between A. humilis and A. burretiana. Finally, A. apoda and A. brasiliensis seem to be closely allied because of their similar appearance. They differ mainly in the number of seeds in their fruit and the nervation of their staminate petals. Evolution within the genus Attalea, as well as within other genera in the subtribe Attaleinae, probably has been taking place since the Eocene period. Fossil cocoid fruits of mostly indeterminate genera from this period have been reported from several localities (see Moore, 1973). In some ways, the evolution of Aftalea parallels that of Syagrus and other genera in the subtribe Butunae. Both Attalea and Syagrus are almost entirely South American in distribution with a large number of acaulescent species occupying similar habitats, although more members of Attalea are found in mesic rather than dry habitats. In addition to this, Attaleais not easily divided into infrageneric categories, but part of the problem may be attributed to the genus being incompletely known or poorly understood. The disparity between species 16 A Taxonomic Treatment of the Palm Subtribe Attaleinae (Tribe Cocoeae) or groups of species may be due to its long evolutionary history, and per haps the well-differentiated, less closely related taxa have been in existence a much longer period of time than the more closely related species. It is unfortunate that the fossil record is not complete enough to determine more accurately when some of these evolutionary events took place. Outline of Tentative Division of Aittalea into Infrageneric Categories Subgenus I. All middle series pinnae in clusters of 2-6. Section 1. Plants acaulescent for the most part, leaf rachis 1.0-—3.5 m long. A. exigua, A. barreirensis, A. allenii. Section 2. Plants arborescent for the most part, leaf rachis 6-18 m long. Subsection A. Staminate flowers completely encircling the staminate rachillae. A. tessmanniz. Subsection B. Staminate flowers on one side of rachillae, sheath and petiole fibers up to 3 m long, endocarp of fruit without visible fibers. A. funifera. Subsection C. Staminate flowers on one side, fibers up to several cm long, endocarp with at least some visible fibers. A. dubia. Subgenus II. Middle series pinnae regularly arranged except for one spe- cies with partially clustered middle series pinnae. Section 3. Plants mostly acaulescent. Subsection D. Stamens 12—52 in number. A. ferruginea, A. amygdalina. Subsection E. Stamens 6-10 in number. Series a. Staminate flowers completely encircling rachillae. A. iguadummat. Series b. Staminate flowers arranged on one side of rachillae Subseries 1. Petals of staminate flowers with reddish glandu- lar hairs. A. nucifera. Subseries 2. Petals of staminate flowers without glandular hairs. A. humilis, A. geraensis. Section 4. Plants mostly arborescent. Subsection F. Stamens 60-75 in number, mesocarp of fruit with con- spicuous air pockets. A. septuagenata. Subsection G. Stamens 6-12, mesocarp without air pockets. Series c. Middle series pinnae 6-8 cm wide, anthers 9-14 mm long. A. seabrensis, A. pindobassu, A. burretiana. Series d. Middle series pinnae 3.5—5.5 cm wide, anthers 6-8 mm long. A. oleifera, A. compta, A. apoda, A. brasiliensis. Genus Attalea 17) Taxonomic Treatment of Genus Attalea In the present paper I am recognizing 21 species and 2 hybrids of genus Attalea (out of 68 published taxa), including the descriptions of 4 new species. All published names pertaining to Attaleaas a genus are included here. Following the taxonomic treatment isa list and discussion of doubtful and uncertain species of Attalea. A list of excluded species of Attalea will be found after the discussion of hybrids of Maximiliana. Subfamily Arecoideae Tribe Cocoeae Martius in Endlicher, Gen. Plant. 254. 1837. (“Cocoineae”). Subtribe Attaleinae Drude in Engler and Prantl, Natural. Planzenfam. 2, 3:27, 78. 1887. (“Attaleae”). TYPE: Attalea. Attalea Kunth, zn Humboldt, Bonpland, and Kunth, Nov. Gen. Sp. 1:folio edition 248. quarto edition 309. 1816; Martius, 1824, 1826, 1844, 1845, 1853; Drude, 1881; Barbosa Rodrigues, 1903b; Burret, 1929a, 1938; Bondar, 1942a, 1942b; Dugand, 1953, 1954; Wessels Boer, 1965, 1972, 1988; Glassman, 1977a. TYPE: Attalea amygdalina Kunth, in Humboldt, Bonpland, and Kunth, Nov. Gen. Sp. 1:309. 1816; section Attalea Verae Drude, Martius FI. Bras. 3:1881; section Euattalea Burret, Notizbl. Bot. Gart. Berlin-Dahlem 10:1929a; section Cylindrostachys Drude, Martius Fl. Bras. 3:1881; section Chaunostachys Burret, Notizbl. Bot. Gart. Berlin-Dahlem 10:1929a; section Dasystachys Burret, Notizbl. Bot. Gart. Berlin Dahlem 10:1929a. Pindarea Barb. Rodr., Pl. Jard. Rio de Janeiro 5:17. 1895. 1903b. TYPE: P. concinna Barb. Rodr., Pl. Jard. Rio de Janeiro 5:17, t. 4C. 1895. (= Attalea dubia [Martius] Burret). Lithocarpos Targioni-Tozzetti, Mem. Mat. Fis. Soc. Ital. Sci. Modena 20:312. 1833. (non Lithocarpos Blume 1825-26). TYPE: L. cocciformis Targioni-Tozzetti, Mem. Mat. Fis. Soc. Ital. Sci. Modena 20:312. 1833 (illegitimate name). Sarinia Cook, Nat. Hort. Mag. 21:78. 1942. TYPE: S. funifera (Martius) Cook, Nat. Hort. Mag. 21:78. 1942. (= Attalea funifera Martius). Trees (in 12 species), from 4—6 m tall to 25-30 m tall and 13-46 cm in diam, trunks usually single, sometimes caespitose, at first with prominent scars caused by abscission of leaves, later becoming obscured through weathering; without emergent stem or acaulescent (in 8 species), but with 18 A Taxonomic Treatment of the Palm Subtribe Attaleinae (Tribe Cocoeae) subterranean stem and occasionally with short aereal trunk; leaves borne at apex of stem in a dense crown or cluster, with youngest ones in the cen- ter; sheathing leaf base 20-130 cm long, usually consisting of a mass of relatively thin or coarse interwoven fibers, individual fibers sometimes up to 3.6 m long and of commercial value, surfaces sometimes with brown- ish lepidote indument; petiole absent or short (6-34 cm long) in some taxa to 2.0-3.6 m long in others, and 2.5—-17.0 cm wide, margins fibrous, becoming smooth with age, surfaces glabrous, ferrugineous lepidote or tomentose, sometimes with dark ferrugineous indument consisting of sharp, narrow, irregularly curved scales; leaf rachis 1-10 m long, surface glabrous or glaucous in several species, rarely soft blackish pubescent or harsh blackish pubescent, ferrugineous lepidote in others or whitish brown lepidote, becoming glabrescent or green with age; blades pinnately compound, pinnae reduplicate in vernation (inverted V-shaped in cross section), 50-200 in number on each side of rachis, uppermost and low- ermost series usually much shorter and narrower than middle series pin- nae, which are 40-43 cm long and 1.5—2.5 cm wide in some species and up to 130-40 cm long and 6-8 cm wide in others, usually regularly (sin- gly) arranged or in clusters of 2-6 (in several species), regularly or irregu- larly spaced, usually lie flat, but divergent in a few taxa, intervals between pinnae or clusters 0.5-9.0 cm apart, mostly with acuminate or aristate asymmetrical tips, surfaces usually glabrous or glaucous, but ferrugineous or brownish lepidote or tomentose on margins and tips in several species, or with small patches of brown ramenta along midrib grooves or abun- dant reddish punctate dots on abaxial side in some others, cross veinlets sometimes prominent; inflorescences located within leaf crown among older leaf bases, several per tree and usually of two kinds: (1) androgynous with both pistillate and staminate flowers and (2) staminate with only staminate flowers, for the most part; each inflorescence consists of a basal prophyll (usually concealed by a leaf sheath), one or more peduncular bracts, and a much larger and wider rameal bract or sterile bract that at maturity (anthesis) splits longitudinally along a midventral line, exposing part or most of the rachis and its flowers; expanded part of staminate ster- ile bract 26 cm long and 4-6 cm wide in some species to 235 cm long and 30 cm wide in others (including beak or rostrum of 4-27 cm long) and up to 4 cm thick, sulcate, depths of grooves variable, grooves often irregu- larly spaced, woody-fibrous, sometimes brittle, glabrous, glaucous or with brownish indument, frequently becoming greenish or glabrous with age, peduncular or narrowed basal part of sterile bract 16—60 cm long; rachis Genus Attalea 19 of staminate inflorescence 20—24 cm long in several species to 200 cm long in a few others, peduncle 10-125 cm long, rachillae numerous, 3—7 cm long in several species to 38-41 cm long in a few others, glabrous or whit- ish or brownish pubescent; staminate flowers numerous, mostly arranged in two rows along one side of the rachilla, twisting of rachillae sometimes gives the appearance of a third row of flowers, spirally arranged around rachilla in two species, occasionally with one basal pistillate flower on some rachillae, staminate flowers 10-15 mm long and 2-3 mm wide in several species to 20-30 mm long and 6—7 mm wide in a few others, petals mostly 3 (rarely 4), always flattened, usually with acuminate tips and denticulate margins, obscurely or prominently veined, usually glabrous and free of glands, but margins sometimes with patches of brownish white pubes- cence, or with tiny punctate glands covered with brownish pubescence, becoming glabrous with age exposing glands, or with patches of reddish glands or short reddish glandular hairs, or dark brown in color with white lepidote or farinose indument along margins, sepals usually 3, very short (mostly 0.5-2.0 mm), triangular, with free, acuminate or acute tips and fused bases, stamens vary in number from consistently 6 to 6-10, 10-15, 17-21, 15-52, and 60-75, separate, included in the corolla, anthers dorsifixed, thecae usually straight, 3-6 mm long (in several species) to 12- 14 mm long in a few others, filaments 1—4 mm long, inserted on recep- tacle; expanded part of androgynous sterile bract from 29-34 cm long to 158-63 cm long (usually with prominent beak 6—33 cm long), 4—6 cm wide in some species to 40 cm wide in others and up to 4 cm thick, mostly deeply sulcate, grooves usually irregularly spaced, woody-fibrous in texture, some- times brittle, surfaces glabrous, glaucous, or brownish pubescent pedun- cular or narrowed basal part 20-80 cm long; rachis of androgynous inflorescence 10-140 cm long, brownish or whitish tomentose or lepidote, or glabrous, usually with many rachillae that are very short or sessile in several species, and 1-18 cm long in others, with 1-8 pistillate flowers on lower half of rachilla, each usually accompanied by two adjacent staminate flowers that fall off early in anthesis, each androgynous rachilla with a nar- rower staminate rachilla extension (very brittle and easily broken off) con- taining one or two rows of staminate flowers only, 6-18 cm long; (in spe- cies descriptions of all four genera, the androgynous rachillae will be referred to as lower pistillate portion and upper narrower staminate ex- tension); pistillate flowers 2—5 cm long and 1-3 cm in diam, petals 3, usu- ally same size as the 3 sepals, sometimes slightly longer or shorter, texture of both more or less smooth, obscurely or distinctly nerved, petals rarely 20 A Taxonomic Treatment of the Palm Subtribe Attaleinae (Tribe Cocoeae) rugose and glandular dotted, pistil 2-4 cm long, ovary sometimes glabrous, more commonly with whitish or brownish indument, rarely farinose, style very short or obscure, stigmas usually 3 in number, occasionally 4 or 6, each 6-12 mm long, staminodial ring 0.5—1.5 cm high, usually glabrous and with smooth margins, but rarely brownish tomentose and with ciliate margins; staminate flowers (from rachillae of pistillate flowers or extended part of rachilla with staminate flowers only) often the same size as those from stami- nate inflorescence, but frequently abnormal or sterile, rarely with petals up to 4cm long, stamens usually the same number as in staminate flowers from staminate inflorescence, but anthers sometimes much smaller and probably sterile; pubescent and glaucous androgynous sterile bracts and inflorescences gradually become glabrous and greenish or brownish, and rachillae usually become thickened during fruit development; fruit from 4-5 cm long and 2.5 cm in diam to 15 cm long and 9 cm in diam, mostly ovoid in shape, beak usually 0.5-2.5 cm long, sometimes with distinct lon- gitudinal grooves or irregularly angled (probably due to crowding of fruits in infructescence), persistent perianth 1.5—5.0 cm high, staminodial ring connate, 0.8—1.5 cm high, margins usually smooth, but others with brown ciliate margins; epicarp usually greenish and glabrous, sometimes yellow, ferrugineous tomentose at maturity in several species, mostly fibrous, 1—2 mm thick, but woody-fibrous and 4—6 mm thick in a few others; mesocarp soft, 0.5-2.0 mm thick (in several species) 4—6 mm thick (in several oth- ers, sometimes divided into two parts) and rarely up to 13 mm thick, oily in anumber of species, sometimes with conspicuous air pockets; endocarp usually woody or bony in texture 0.5—2.0 cm thick; fiber clusters usually scattered and inconspicuous, sometimes completely free of visible fibers, but in several species with prominent irregular patches of fibers or with prominent patches arranged in a circular pattern; seeds more commonly 1-2, less commonly 3-5, rarely 8 in number, 2-5 cm long and 1.0—2.5 cm in diam, yielding an edible oil in a number of species. Key to Species of Attalea 1. Middle series pinnae in clusters of 2—6 or only lower one-third to one- half of rachis with clustered pinnae in some arborescent species. . . Joie ah SUNT il eee (A. seabrensis and A. x piassabossu). 2. Plants acaulescent, leaf rachis 1.0—3.5 m long. 3. Staminate rachillae 10-15 cm long, staminate flowers 20-24 mm long, stamens about one-third as long as petals, leaf sheath fibers Genus Attalea 1 very long)/(upito 3 m), angled toward) base yy :).)5 (04/2 +) Pai sar bP Cae e eae, wee cf) 2 4. A. funifera (Brazil: Bahia). 3. Staminate rachillae 3-8 cm long, staminate flowers 12-16 mm long, stamens one-half to two-thirds as long as petals, leaf sheath fibers much shorter, angled upward. 4. Pistillate portion of androgynous rachilla up to 9 cm long, middle series pinnae 1.5—2.0 cm wide, endocarp of fruit with conspicuous xing, of ber, ChASteTs).. 05.0: 465 i Pyseoalevaq ee .... 1. A. exigua (Brazil: Goias, Minas Gerais, Mato Grosso). 4. Pistillate portion of androgynous rachilla very short (0.5-1.0 cm), middle series pinnae 3.0—5.5 cm wide, endocarp of fruit with inconspicuous clusters of fibers. 5. Petals of staminate flowers without punctate glands covered with brownish lepidote indument, anthers 9-11 mm long, ups ofpistillate petals trdewtate). scald paieps.s oye een fide ie A laicds te werner vb ieg 5 md 2. A. barreirensis (Brazil: Bahia). 5. Petals of staminate flowers with distinct punctate glands covered with brownish lepidote indument, anthers 5—7 mm long; tipsjof pisullate*petals aculte 4... 2.2 i124 )eeyei Slenen petiaags hier: 2 Rn Bk seh k 3. A. allenii (Panama, Colombia). 2. Plants arborescent, leaf rachis 6-18 m long. 6. Middle series pinnae mostly regularly arranged, only lower one-third to almost one-half of rachis with clustered pinnae. 7. Lower staminate rachillae 15-20 cm long, pistillate portion of androgynous rachillae 1-3 cm long, stigmas 3—6 in number. . bier ep ty ot egepey ch eh prea dent 14. A. seabrensis (Brazil: Bahia). 7. Lower staminate rachillae 25-29 cm long, pistillate portion of androgynous rachillae 12-18 cm long, stigmas 3 in number. . afd byes tog pres, OS ERLE epee A. X piassabossu (Brazil: Bahia). 6. Pinnae clustered throughout except for extreme upper pinnae. 8. Staminate flowers completely encircling the staminate rachillae AV east ately pe wis eee ahcphscihia Aad 5. A. tessmannii (Peru). 8. Staminate flowers in two rows on one side of the staminate rachillae. 9. Staminate flowers 21-24 mm long, petals 3-5 mm wide and valvate, fruits 10-15 cm long, mesocarp up to 13 mm thick, individual sheath and petiole fibers up to 3.5 m long, mostly projected downwards toward base................2005: eR A Ee tes ap a eure Ma 4. A. funifera (Brazil: Bahia). 22 A Taxonomic Treatment of the Palm Subtribe Altaleinae (Tribe Cocoeae) 9: Staminate flowers 10-20 mm long, petals 1.5-2.5 mm wide and imbricate, fruits 6.0-8.5 cm long, mesocarp 3—4 mm thick, individual sheath and petiole fibers up to several cm long, projected upward away from the base.................. .. 6. A. dubia (Brazil: Espirito Santo to Santa Catarina). 1. Middle series pinnae regularly arranged along rachis, sometimes lower one-tenth of rachis with loose, widely spaced pairs of pinnae (A. pindobassu). 10. Plants acaulescent. 11. Stamens 12-52 in number, staminate petals without patches Lis of reddish glandular hairs, and staminate flowers not spirally arranged around rachilla. 12. 12: Middle series pinnae 80-95 cm long and 3-4 cm wide, margins and apices ferrugineous-lepidote, staminate flowers 2.5—3.0 mm wide, petals with patches of brown- ish white lepidote indument, stamens 15-52 in number, petals of pistillate flowers rugose and glandular dotted, fruits 6.5-9.0 cm long, mesocarp about 1 mm thick. . . 8. A. ferruginea (Venezuela, Colombia, Peru, Brazil). Middle series pinnae 115-25 cm long and 6-8 cm wide, margins and apices more or less glabrous, staminate flowers 4-6 mm wide, petals mostly glabrous, stamens 12-21 in number, petals of pistillate flowers not rugose or glandular dotted, fruits 10-12 cm long, mesocarp 2— 6 mmtthicksus52.. ..% 7. A. amygdalina (Colombia). Stamens 6-15 in number, staminate petals either with reddish glandular hairs or staminate flowers spirally arranged in A. nucifera and A. iguadummat. nS. 13. Middle series pinnae 100-148 cm long and 6—7 cm wide; staminate rachillae 13-19 cm long. 14. Petals of staminate flowers with scattered patches of reddish glandular hairs at anthesis, staminate flowers arranged on one side of rachillae ............. Pe es ee 9. A. nucifera (Colombia). 14. Petals of staminate flowers without reddish glandu- lar hairs, staminate flowers spirally arranged around rachillae’ 275.4% 10. A. iguadummat (Panama). Middle series pinnae 46-95 cm long and 2.5-5.5 cm Genus Attalea 23, 15: 1D. wide, staminate rachillae 5-11 cm long, staminate petals without reddish glandular hairs, flowers arranged on one side of rachilla. Leaf rachis 2.5—5.3 m long, rachis of staminate inflorescence 34— 44 cm long, stamens consistently 6 in number, middle series pin- nae 3.5-5.5 cm wide, anthers 8-12 mm long....... 11. A. hu- milis (Brazil: Bahia, Espirito Santo, Rio de Janeiro, Sao Paulo). Leaf rachis 1.5—2.2 m long, rachis of staminate inflorescence 20-— 30 cm long, stamens 6-10 in number, middle series pinnae 2.5— 3Oicmewide, anthers 7-9’ mmilon@. (3) Ae. 12. A. geraen- sis (Brazil: Minas Gerais, Goias, Bahia, Sao Paulo, Mato Grosso). 10. Plants arborescent. 16. Petiole and leaf rachis covered with indument of dark, sharp, 16. curved scales, stamens 60-75 in number, petals of staminate flowers with white lepidote or farinose margins, mesocarp of fruit MAL CONS PICUOWS dik POCKElS * hoi Ae hohe oe sw ee eee PO VHISLAE stcios Steen: SaaS 13. A. septuagenata (Colombia). Petiole and leaf rachis not covered with indument of sharp, curved scales, stamens 6—12 in number, petals of staminate flowers more or less glabrous, mesocarp without air pockets. 17. Staminate rachillae 15-31 cm long, rachis of staminate inflorescence 85-150 cm long. 18. Middle series pinnae 6-8 cm wide, anthers 9-14 mm long. 19. Lower pinnae with several widely spaced loose clus- ters of 2-3, petiole 1.2—2.0 m long, stigmas 6 in num- ber, pistillate portion of androgynous rachilla 2.0— 4.5 cm long, epicarp of fruit 3-5 mm thick ..... SL CER ihe 8 ie Laren 15. A. pindobassu (Brazil: Bahia). 19. Lower pinnae not in widely spaced loose clusters, petiole very short or absent, stigmas 3 in number, pistillate portion of androgynous rachillae either less than | cm, 6-7 cm, or 13-14 cm long, epicarp of fruit P5220 main THick Wht ie Yee etal ee A eer s ARC eee d 9504 MD 16. A. burretiana (Brazil: Bahia). 18. Middle series pinnae 3.5—5.5 cm wide, anthers 6-8 mm long. 20. Petiole 0.6—1.5 m long, staminate rachillae 18-22 cm long, staminate flowers 12-15 mm long, stamens 6— 24 A Taxonomic Treatment of the Palm Subtribe Attaleinae (Tribe Cocoeae) 9 in number, fruits.1—2 seeded: .... ..2. 62. Jaa Se ene oe 17. A. oleifera (Brazil: Pernambuco). 20. Petiole 0.1—0.2 m long, staminate rachillae 15-18 cm long, staminate flowers 15-18 mm long, stamens 10- 12 in number, fruits with 3—4 seeds............ bia sits ous Saat. 18. A. compta (Brazil: Minas Gerais). 17. Staminate rachillae 6-14 cm long, rachis of staminate inflorescence 35-45 cm or 60-80 cm long. 21. Staminate rachillae 6-11 cm long, usually acaulescent, but some- times with trunk up to) Aim) talliys. 93,0 0)2. 38 5 ee sjactd sala chee siateee cube eat Nee pop tn ie 11. A. humilis (Brazil: Bahia). 21. Staminate rachillae 12—14 cm long, trees up to 30 m tall. 22. Rachis of staminate inflorescence 60-80 cm long, staminate rachillae 12-14 cm long, staminate flowers 17-20 mm long, petals strongly nerved, mesocarp 1 mm thick, fruits 1 seeded oa an Gt ieee chant. i 19. A. apoda (Brazil: Minas Gerais). 22. Rachis of staminate inflorescence 35—45 cm long, staminate rachillae 9-11 cm long, staminate flowers 13-16 mm long, mesocarp 2.5-3.0 mm thick. 23. Staminate flowers mostly with four petals, each strongly nerved, fruits about 9 cm long, | seeded, fiber clusters in endocarp nob prominenti.<, gods. 4)3))- ae yds Paani deputy ab 20. A. salvadorensis (Brazil: Bahia). 23. Staminate flowers with three petals, each obscurely nerved, fruits about 6 cm long, 2 seeded, fiber clusters in. endocarp prominent s.j4n0. 201.0 *): ee if a eye i el ea 21. A. brasiliensis (Brazil: Goias). Taxonomic Treatment of Species of Attalea 1. Attalea exigua Drude, Martius FI. Bras. 3:439. t. 100, fig. 1. 1881; Burret, 1929a; Glassman, 1977a; Henderson, Galeano, and Bernal, 1995. TYPE: Brazil, between Goias and Cuyaba, 1844, Weddell 2965 (lec- totype, P!, only 1 of 2 sheets, flower of sheet 2 belongs to Orbignya; isotype F!). Figs. 37, 75, 78. Acaulescent; sheathing leaf base and petiole not measured; leaves about 1 m long (fide Drude); 50-60 pinnae on each side of rachis, those from middle series in clusters of 2-3, 50-54 cm long, 1.5-2.0 cm wide, with asymmetrical tips; expanded part of staminate sterile bract about 28 cm Genus Attalea 25 long and 3.5 cm wide (immature?), peduncular part about 12 cm long; rachis of staminate inflorescence about 19 cm long, peduncle 10 cm long, rachillae 30—40 in number, 6-7 cm long; staminate flowers mostly in single row on one side of the rachilla, 12-15 mm long, petals flat, 2-3 mm wide, distinctly nerved, with acuminate tips, margins somewhat denticulate, sepals 1-2 mm long, stamens 6—7 in number (9, fide Drude), anthers 5— 6 mm long, filaments 2-3 mm long; androgynous inflorescence not mea- sured; androgynous rachilla (from Weddell 2022 [B]) with stalk of 9cm and flowering part of 9 cm, with 7 pistillate flower scars, extended staminate rachilla about 3 cm long (incomplete); fruit (from Weddell 2022 [B]) 6 cm long and 3.5—4.0 cm in diam (including beak of 8 mm long), epicarp and mesocarp not readily distinguishable, fibrous, about 2 mm thick, endocarp hard, 6-8 mm thick, with conspicuous ring of fiber clusters confined to middle of endocarp, seed cavities 2, seeds not seen. Distribution and Habitat. See table 1. Vernacular Name. /ndaza rasteira. Representative Specimens Examined. BRAZIL. Goias: near Salinas, 1844, Weddell 2022 (B, P); near Goias, campo, July 1895, Glaziou 22267 (P). Minas Gerais: Riacho das Varas, Campo, Mar. 1893, Glaziou 20022 (P). Specimens Tentatively Included. BRAZIL: Maranhao: Snethlage 735 (B), 737 (B). As previously mentioned, this taxon is most similar to A. barrecrensis from Bahia and A. allenii from Panama and Colombia; however, its present dis- tribution is uncertain because the last known collection was made in 1895. 2. Attalea barreirensis Glassman. sp. nov. TYPE: Brazil, Bahia, munic. Barreiras, 28 km west of Barreiras, cerrado, Aug. 1976, Glassman 13048 (holotype, F!; isotype, SP!). Figs. 31-32, 69, 78. Palma acaulis, pinnis mediis aggregatis, 35-46 cm longis, 3.0—4.6 cm latis, bractea mascula 36-46 cm longa, 9-13 cm lata, fusco-ferruginea, inflorescentia mascula multiramosa, pedunculo 21—25 cm longo, rachide 16-27 cm longa, rachillis 4-8 cm longis, flores masculi 14-16 mm longi, stamina 6 numero, antherae 9-11 mm longae, fructus 5.0—5.5 cm longus, 3.0—4.5 cm diametro, exocarpio fibroso, 1 mm crasso, mesocarpio pulposo, 1 mm crasso, endocarpio osseo, 5 mm crasso, semina I, 2.2 cm longa, 2 cm lata. Acaulescent, plants solitary or caespitose; sheathing leaf base not mea- sured, petiole 14—45 cm long, leaf rachis 0.8—1.8 m long; 60-112 pinnae on each side of rachis, those from middle series in clusters of 2-4, 35- 46 cm long, 3.0-4.6 cm wide, with asymmetrical tips; expanded part of 26 A Taxonomic Treatment of the Palm Subtribe Attaleinae (Tribe Cocoeae) staminate sterile bract 36—46 cm long (including beak of 4 cm long), 9- 13 cm wide and 2—3 mm thick, deeply sulcate, covered with dense brown- ish tomentum, peduncular part about 20 cm long; rachis of staminate inflorescence 16-27 cm long, peduncle 21—25 cm long, rachillae 35-79 in number, 4—8 cm long; staminate flowers arranged in two rows along one side of the rachilla, 14-16 mm long and 4—5 mm wide, petals ob- scurely nerved, mostly with acute tips, sepals 1-2 mm long, stamens 6 in number, anthers 9-11 mm long, filaments about 2 mm long; androgynous sterile bract 21-30 cm long and 6-9 cm wide, rachis of androgynous inflorescence 8—11 cm long, peduncle 8-10 cm long; pistillate rachillae very short or absent, pistillate flowers with apices of petals tridentate, 3.0— 3.5 cm long and 1.5—2.0 cm in diam, stigmas 3; expanded part of fruit- ing sterile bract about 29 cm long (including beak of 4 cm long) and 8 cm wide, deeply sulcate, with brownish tomentose indument, becoming glabrous with age; rachis of infructescence 16 cm long, with about 32 fruits per rachis, fruiting rachillae short, 0.5-1.0 cm long, with only 1 fruit per rachilla; fruit yellow to reddish-orange, turning purple when mature (fide Noblick, 1991), 5.0-5.5 cm long and 3.0—4.5 cm in diam, persistent perianth 2-4 cm high, staminodial ring about | cm high, epicarp fibrous, about 1 mm thick, mesocarp soft, about 1 mm thick, endocarp hard, about 5 mm thick, with scattered fiber clusters mostly arranged in a circle, seed | in number, about 2.2 cm long and 2 cm in diam, with central cav- ity about 1 cm long. Distribution and Habitat. Brazil, common in western Bahia, in cerrados at elevations of 600-800 m. According to Noblick (1991), this species was also observed at the old airport in Barreiras, between Cocos and Coribe in the south and various points between, e.g., Baianopolis. Noblick ob- served large populations from Sao Desiderio and Roda Velho to Barreiras and north to Sao Marcelo; he believed that this species eventually will be found in southern Piaui, Goias, and Minas Gerais. Vernacular Name. Catolé (fide Noblick, 1991). Representative Specimens Examined. BRAZIL. Bahia: munic. Barreiras, 28 km west of Barreiras, cerrado, Aug. 1976, Glassman 13047 (F, SP); Baianopolis, 12-13 km sul do povoado de Cocos (Rio Sao Desiderio) na estrada Varzeas-Correntina, sandy soil, cerrado, 700-800 m, Feb. 1986, Noblick and Lobo 4528 (HUEFS); Coribe, 7-8 km sul da cidade na BR172 (Coribe/Cocos), common, poor sandy soil, in cerrado, 650-700 m, Jan. 1986, Noblick and Lobo 4522 (HUEFS). This taxon appears to be most similar to A. exigua from Mato Grosso and Genus Attalea 97 A. allen from Panama and Colombia in being acaulescent, having clustered middle series pinnae, and having similar-sized staminate flowers (12-16 mm long). The new species can be differentiated from A. exigua by longer an- thers (9-11 vs. 5—6 mm) and 1-seeded fruits with inconspicuous patches of endocarp fibers rather than 2-seeded fruits with a conspicuous ring of en- docarp fibers. A. barreirensis has the same number of stamens and very short androgynous rachillae as in A. allenii, but has longer anthers (9-11 vs. 5—7 mm), narrower middle series pinnae (3.0 cm vs. 4.0—5.5 cm), and smooth staminate petals rather than petals with punctate glands. The fruits of A. barreirensis are edible, and the sweet seeds are used lo- cally in making candy. 3. Attalea allenii H. E. Moore, Gentes Herb. 8:191, fig. 82. 1949; Dugand, 1953; Glassman, 1977a; de Nevers, 1987; Henderson, Galeano, and Bernal, 1995. TYPE: Panama (Allen 4103, holotype, MO!; isotype, BH!). Figs. 42, 65, 82. Acaulescent or with very short trunk; sheathing leaf base about 73 cm long, petiole 0.6—3.6 m long, densely ferrugineous-lepidote on convex (adaxial) side, 2.5 cm in diam at base; leaf rachis 3.3-3.6 m long, densely ferrugineous-lepidote on flat (abaxial) side, glabrous adaxially; 85-114 pinnae on each side of rachis, those from middle series in clusters of 3-6, intervals of 5-9 cm between clusters, 75—95 cm long, 4.0—5.6 cm wide, with more or Jess prominent cross veinlets, with acuminate asymmetrical tips, margins of pinnae covered with ferrugineous lepidote indument chiefly near tips; expanded part of staminate sterile bract 30—38 cm long (includ- ing beak of 6 cm long) and 4—6 cm wide, deeply sulcate, peduncular part 19 cm long; rachis of staminate inflorescence 22-39 cm long, peduncle about 28 cm long, rachillae 30-50 in number, 3.0-—4.5 cm long, covered with a dark brown lepidote indument; about 12—15 staminate flowers per rachilla, arranged in two rows along one side of the rachilla, 10-15 mm long, petals flat, 2-4 mm wide, with acuminate tips and denticulate mar- gins, obscurely or more or less prominently nerved, with tiny punctate glands covered with brownish lepidote indument that exposes the glands more clearly when the indument falls off, sepals 1-2 mm long, stamens 6 in number, anthers 5-7 mm long, filaments 1-2 mm long; expanded part of androgynous sterile bract about 40 cm long (including beak of 6.5 cm long), about 6 cm wide, deeply sulcate, grooves irregularly arranged, cov- ered with dense brownish lepidote indument, peduncular part 38-40 cm long; androgynous inflorescence mostly unbranched, rachis 10-11 cm 28 A Taxonomic Treatment of the Palm Subtribe Attaleinae (Tribe Cocoeae) long, peduncle 34-35 cm long, 25-35 pistillate flower scars, pistillate flowers sessile or on short rachillae up to 0.5 cm long, staminate exten- sion of each rachilla about 8 cm long; pistillate flowers 2.5-2.7 cm long and 1.5—2.0 cm in diam, sepals 1.5—1.7 cm long, shorter than petals, pis- til about 2 cm long and 1 cm in diam, ovary covered with dense dark brown indument, stigmas 3, about 1 cm long, staminodial ring about 0.7 cm high; staminate flowers (from rachillae of pistillate flowers) 13-16 mm long, petals about 5 mm wide, with acute tips, more or less nerved, with scat- tered patches of brown lepidote indument, sepals 1-4 mm long, stamens 6 in number, anthers about 6 mm long, filaments 3 mm long; complete fruiting sterile bract not seen; infructescence unbranched, rachis 13-14 cm long, peduncle about 34 cm long; fruit 6.0—7.5 cm long (including beak of 5-8 mm long), 3.5—4.0 cm in diam, persistent perianth 2—3 cm high, staminodial ring 1.5—-2.0 cm high and 2.5—3.0 cm in diam, epicarp fibrous, 1.0-1.5 mm thick, mesocarp soft, about 0.5 mm thick, endocarp hard, up to 1 cm thick, fiber clusters obscure, seeds 1—3 in number, about 3 cm long and 1.5 cm in diam. Distribution and Habitat. See table 1. Vernacular Names. Bangue, mangue, igua (Panama); mangue, taparo, tapa- ro chiquito, taparin, corozo chiquito (Colombia). Representative Specimens Examined. PANAMA. Colon: Salud Hills, Lao and Holdridge 197 (MO); Canal Zone: Coco solo, Sept. 1972, Gentry 6298 (MO, R); Comarca de San Blas: El] Llano-Carti road, km 16.7 west to Rio Carti Grande, Nov. 1984, de Nevers and Herrera 4152 (NY). COLOMBIA. Valle: vicinity of Buenaventura, forests in concession of Carton Colombia, Feb. 1967, Moore et al. 9460 (BH); Buenaventura, May 1926, Cook 64 (US); slopes and ravines of Aqua Dulce, an island in Buenaventura Bay, Feb. 1967, Moore et al. 9468 (BH); Bahia de Buenaventura, Quebra de San Joaquin, Feb. 1946, Cuatrecasas 19948 (COL, F); Buenaventura, Camp Cartin, Mar. 1967, D'Alessandro and Gutimes 19 (BH); R. Salaqui, Hydro Camp 14, May 1967, Duke 11377 (BH); km. 15 Carretera Buenaventura— Buga, Finca de Moises-Caicedo, May 1958, Patino 203 (COL); munic. Istmina, Correg. de Togroma, May 1958, Patino 205 (COL); Rio Calima, La Troijita, Feb. 1944, Cuatrecasas 16397 (COL); Corcovada region, upper Rio San Juan, dense forest, Apr. 1939, Killip 35311 (US); Dindo area, Bajo Calima, pluvial forest, July 1984, Gentry and Monsalve 48429 (NY); Antio- quia: entre Dabeiba y Llanitos, Feb. 1942, Cuatrecasas and Metcalf 30184 (COL); Bolivar: Bajo Magdalena, Orillas del Brazo de Mompos, Aug. 1940, Najar 11 (COL); Narino: Tumaco, May 1958, Patino CL-1 (COL), CL-2 Genus Attalea 29 (COL), CL-3 (COL); Choco: environs of Quibido, secondary forest, mixed with Jessenia and Bactris, July 1984, King et al. 548 (NY); same locality, par- tially disturbed forest, June 1985, King et al. 671 (NY); environs of Las Animas, roadside, entrance to Pan American Highway, secondary forest, King et al. 664 (NY). This taxon seems to be most closely related to A. exigua from Goias and Minas Gerais because both species are acaulescent and have clustered mid- dle series pinnae. A. allenii differs from A. exiguain its mostly unbranched rather than branched androgynous inflorescences and its petals of stami- nate flowers with punctate glands. According to Dugand (1961), A. allenii as well as other species of Attalea from Colombia (A. nucifera, A. septuagenata, A. amygdalina) have edible seeds that yield a fine edible oil. This product has not been exploited com- mercially for lack of efficient machines to break the hard endocarp with- out damaging the seeds. Endosperm of seed is used as a food and drink by local inhabitants (fide King et al. 664). 4. Attalea funifera Martius ex Sprengel, Syst. Veg. 2:624. 1825; Martius, Hist. Natur. Palm. 2:136, ts. 95-96, fig. 4. 1826; Bondar, figs. 4-10. 1942b; Noblick, 1991; Henderson, Galeano, and Bernal, 1995. LECTO- TYPE: (Glassman, 1977a) Brazil, Espirito Santo, Porto Seguro and Bahia (Martius, ts. 95-96, fig. 4. 1826). Figs. 5-8, 43, 68, 80. Sarinia funifera (Martius) O. F. Cook, Nat. Hort. Mag. 21:78. 1942. A. acaulis Burret, Repert. Spec. Nov. Regni Veg. 32:103. 1933. TYPE: Brazil, Bahia, Werdermann 3182 (holotype, B). A. funiferavar. acaulis Burret, in Bondar, Bol. Inst. Centr. Fom. Econ. Bahia 13:fig. 8. 1942b. pro syn. Acaulescent or trees 1.5-15.0 m tall and 25-30 cm in diam; sheathing leaf base about 1.3 m long, individual fibers up to 3.6 m long; petiole 0.4— 2.0 m long, margins with long rigid fibers up to 3.5 m long; leaf rachis 3.5— 6.0 m long; 130-60 pinnae on each side of rachis, those from middle se- ries in clusters of 3-4 (2-9), divergent, often arranged at different angles in separate interrupted groups, 60-120 cm long (up to 150 cm in culti- vated specimens) and 4—7 cm wide; staminate sterile bract 75-135 cm long (including beak of 13-27 cm long), 14 cm wide, and 3-4 cm thick, deeply sulcate; rachis of staminate inflorescence 30-40 or 60-85 cm long, pe- duncle 35-70 cm long, rachillae 70-80 or 90-145 in number, 10-15 or 18- 22 cm long on basal part; staminate flowers in two rows along one side of the rachilla, 21-24 (30) mm long, petals flat, 3-5 mm wide, with acute or 30 A Taxonomic Treatment of the Palm Subtribe Attaleinae (Tribe Cocoeae) acuminate tips, distinctly nerved, sepals 0.5-1.0 mm long, stamens 6 in number, anthers 4-7 mm long, filaments 3—4 mm long; androgynous ster- ile bract 80-165 cm long (including beak of 16-37 cm long), about 12 cm wide and 4mm thick, deeply sulcate; rachis of androgynous inflorescence 35-55 cm long, peduncle 40 cm long, with many rachillae, pistillate por- tion sessile, 2.0-2.5 or 4-6 cm long, with 1-4 pistillate flowers, each 3.0— 3.5 cm long, stigmas 3 in number; fruiting sterile bract 60-93 cm long (including beak of 16 cm long) and 9 cm wide, deeply sulcate, rachis of infructescence 47—72 cm long, with many rachillae; mature fruit 10-15 cm long (including beak of 2.0-2.5 cm long) and 5.5—9.0 cm in diam, surface marked by 3 longitudinal grooves being sutures between carpels which are not completely closed (fide Cook, 1942), persistent perianth about 3 cm high, staminodial ring about 4 cm across and 1.4 cm high, epicarp fibrous, 2.0—2.5 mm thick, mesocarp soft, oily, 3-6 mm thick (13 mm thick, fide Burret, 1929a), endocarp hard, without fiber clusters, 1.5— 1.8 cm thick, seeds 1-3 in number, about 4 cm long and 1.5-1.8 cm in diam. Distribution and Habitat. Martius ex Sprengel (1825) cites the follow- ing: “Southern Brazil, mainly along strand, between 10—20 degrees So. lat., in coastal forests near ocean in Espirito Santo and Bahia: Porto Seguro and IIheus, entering Minas Gerais.” According to Bondar (1942b), how- ever, piassava is not native in the states of Espirito Santo and Minas Gerais; it is distributed in littoral soils of Tertiary origin in Bahia from the mu- nicipality of Jiquirica, south of Salvador, to Rio Corumbahy, in the munici- pality of Prado. He also saw dense native forests of an acaulescent stage of piassava (which he called A. acaulis) from Reconcavo da Bahia to Rio Real near the border of Sergipe. Bondar speculated that these particular stands are trunkless and have a poor grade of fiber because they grow on poor soils with less than normal precipitation. Other fairly large popula- tions of mostly acaulescent piassava (probably several thousand plants) were observed in August 1988 by Larry Noblick and me in the sand dunes area of Itapoan, in the environs of Salvador. The acaulescent plants oc- cur mainly on the slopes, whereas in the ravines, the plants are mostly arborescent (6—10 m tall). Palm associates include Allagoptera arenana, two species of Bactris, and Syagrus coronata. According to Medeiros-Costa (1985), A. funifera occurs in the states of Espirito Santo, Bahia, Sergipe, and Alagoas (mainly in sand dunes and remnants of the Atlantic coastal forest); however, Voeks (1987) reports that the southern limit of this species is near Caraiva, in southern Bahia; and Voeks (1985, 1988) discusses reproductive ecology of this species. Genus Attalea 31 In the northernmost part of its range (i.e., Maceio, Alagoas), the plants are unusually small (fide Noblick, 1991). One of the densest populations observed by Larry Noblick and me was in Bahia, on the island of Itaparica, along highway BR101, from Colonia de Ferias to Nazaré. Perhaps one hundred thousand trees were growing in the region, mostly on red, sandy soils, in remnants of the Atlantic coastal forest, associated with Syagrus botryophora, Bactris sp., and Polyandrococos caudescens. In a few localities near Salvador, A. funifera hybridizes with A. burretiana to produce A. x piassa- bossu; A. funifera also crosses with A. humilis, resulting in A. x voeksi, known only from one station. These taxa will be treated in more detail under “Hybrids of Attalea.” Vernacular Names. Piassava, piacaba, piracaba, coquilho, coquilla (fruits), piassaveira (fibers). Representative Specimens Examined. BRAZIL. Bahia: Bondar 21 (F- 404620), Bondar s.n. (F-619760) ; Salvador, sand dunes, Nov. 1935, Dahlgren s.n. (F-614747); 9 km east of Nazaré, along BA245, remnant of Atlantic coastal forest, stand of 750—1,000 trees, Aug. 1988, Noblick and Glassman 4578 (BAH, F), 4579 (BAH, F), 4580 (BAH, F), 4581 (BAH, F); munic. Aguas Claras, 15 km west of Salvador, along BR324, 500 m north of access to USIBA, remnant of Atlantic coastal forest, in low area, above stream associated with Syagrus botryophora, Bactris, and Polyandrococos, about 300 trees in area, near FRIMASA meat processing plant, Aug. 1988, Noblick and Glassman 4586 (BAH, F). CULTIVATED. BRAZIL. Jard. Bot. Rio Jan., July 1886, Glaziou 16483 (BR, C, MO, P); Bailey and Bailey 484 (BH); Dahlgren s.n. (F-611639). As previously mentioned, this taxon is probably most closely related to A. dubia because both are mostly arborescent with clustered middle series pinnae. They differ mainly in the size of their staminate rachillae and flowers, the number of stamens per flower, the size of their fruits, and the very long fibers from the leaf base in A. funifera. According to Cook (1942), A. funifera belongs in a separate genus be- cause of the unusual fibrous structure of the sheathing leaf bases and exceptionally thick and uniform texture of the endocarp (i.e., not rein- forced by an indurated layer of mesocarp fibers that are fused and in- corporated into the endocarp). He described a new genus (Sarinia) and subsequently made a new combination, Sarinia funifera; however, the genus was originally thought to be invalid because it lacked a formal Latin description. Cook compared this taxon with other genera de- scribed by him having similar characteristics, Ynesa, Bornoa, Temenia, and Ethnora, but apparently most (except Ynesa) are invalid because they were 32 A Taxonomic Treatment of the Palm Subtribe Attaleinae (Tribe Cocoeae) published without Latin diagnoses after 1935. On page 84, Cook men- tioned that the diagnostic characters of Sarinia—long fibers, interrupted pinnae, and uniform endocarp—are scarcely different in Latin: fibris longis, pinnis interruptis, endocarpiis uniforms. Therefore, according to the rules of botanical nomenclature, this genus is treated as having a valid description. These characteristics, however, are insufficient to delineate Sarinia funifera as a separate genus because other members of Attalea have simi- lar specialized characteristics, such as leaf sheaths with very long fibers, interrupted pinnae, homogeneous (nonfibrous) endocarp, and grooved or angled fruits. A. funiferais known as the original source of piassava, a coarse rigid fiber used extensively in Europe and in the United States for brushes, brooms, and street cleaning machinery (Cook, 1942). Similar fibers have been ob- tained from Leopoldinia piassaba and other palms not related to Attalea. According to Bondar (1942b), piassava fiber is extracted for commercial use almost entirely from native trees in coastal forests in southern Bahia. No known cultivated piassava plantations existed at that time. Fonseca (1927) reported that piassava is “spontaneously” cultivated for fiber and oil production all along the coast of Bahia. At the time, the species was in danger of extermination because of the method of fiber extraction, i.e., some plants were cut down to pull the leaf fibers, whereas others were uprooted to search for fibers on the roots in addition to the leaf bases. Fruits yield oil equal to that from babassti. Heyne (1927) said oil from seeds of piassava was used for oiling watches and other fine instruments, and seeds were used to carve buttons and rosaries. Booth (1889) reported that piassava fibers were used for cables, ropes, blankets, hats, and fences in Brazil, but in Europe the fiber was used mostly for brushes and brooms by itself or mixed with other fibers. The main types of fiber-producing plants are bananeira, young trunkless plants that yield a fresh and supple fiber, and coquiera, mature plants that yield two or three kinds of fibers. One of these, from the younger leaves, is similar to fibers of bananeira. In the late 1880s, about 7,000 tons of piassava fiber was exported to Eu- rope from Brazil. According to Bondar (1942b) fruits and seeds had commercial value as a vegetable ivory in the manufacture of buttons and carved objects, the oil could be used as a fuel in gasoline engines, and the fruits were an ex- cellent source for the production of charcoal. Genus Aitalea oa 5. Attalea tessmannii Burret, Notizbl. Bot. Gart. Berlin-Dahlem 10:538. 1929a. 12:155. 1934; Glassman, 1977a; Henderson, 1995; Hender- son, Galeano, and Bernal, 1995. TYPE: East Peru, Upper Amazon region, lower Itaya, Soledad, in flood-free forests, June 1925, Tessmann 5167 (holotype, B!; isotypes, G!, F!, NY!). Figs. 52, 82. Trees 15-30 m tall and 25—46 cm in diam; sheathing leaf base and peti- ole not measured; leaves about 18 m long (fide Gentry et al. 29032); middle series pinnae in clusters of 2-5, 82 cm long and 4—6 cm wide; staminate sterile bract 2.4 m long (fide Burret); staminate inflorescence not mea- sured, with numerous rachillae, 25-28 cm long; staminate flowers ar- ranged in pairs, completely encircling the rachilla, about 286 staminate flowers per rachilla, each 14-16 mm long, petals flat, 2-3 mm wide, dis- tinctly nerved, with acuminate tips and denticulate and short ciliate mar- gins on upper third of petal, stamens 6—9 in number (12, fide Burret), anthers 4—5 mm long, filaments 2.0—2.5 mm long; androgynous inflor- escence not measured; pistillate portion of androgynous rachilla 13-18 cm long, mostly with 1-3 pistillate flowers, each about 2.5 cm long, sepals shorter than petals, about 1 cm long, stigmas 3, extended staminate rachilla 7-9 cm long, with sterile staminate flowers; rachillae of infruc- tescence 9-13 cm long, mostly 3 fruits per rachilla; fruit 11-15 cm long and 4.5-5.5 cm in diam (including beak of 1.3 cm long), persistent peri- anth about 4 cm high, staminodial ring about 1.5 cm high, epicarp fibrous, 1.0-1.5 mm thick, mesocarp soft, 0.5 mm thick, endocarp 10-15 mm thick, with abundant, scattered, and conspicuous fiber clusters, but obscured by dark brown deposits of tannin, seed cavities 1-3, surrounded by a circu- lar fiber-free area. Distribution and Habitat. See table 1. Vernacular Names. Conta, chonta. Representative Specimens Examined. PERU. Loreto: prov. Maynas, Yanamono Explorama Tourist Camp on Rio Amazonas, between Indiana and mouth of Rio Napo, July 1980, Gentry et al. 29032 (MO); same local- ity, June 1983, Gentry and Vasquez 42322 (MO). Specimens Tentatively Included. PERU. Madre de Dios: prov. Manu, Parque Nacional Manu, Rio Manu, Rio Sotileja, steep forested hills, alt. 400-500 m, Oct. 1986, Foster and d’Achille 11554 (F). The spiral arrangement of staminate flowers around the rachillae is unusual for Attalea; however, a recently described species from Panama, 34 A Taxonomic Treatment of the Palm Subtribe Altaleinae (Tribe Cocoeae) A. iguadummat, also has this arrangement. All other species in the genus have staminate flowers in 1-2 rows along one side of the rachilla. Burret (1929a) classified A. tessmanniiin a separate section, Dasystachys. It would seem that A. lessmannii and A. iguadummat are closely allied because of their unusual flower arrangement, but there are some major differences. The first species is a tall tree with clustered middle series pinnae, stami- nate rachillae 25-28 cm long, and fruits 11-15 cm long, whereas A. iguadummat is acaulescent with regularly arranged pinnae, staminate rachillae 13-18 cm long, and fruits 7-10 cm long. According to Henderson and Balick (1991), staminate petals of this species are 3—5 in number, linear in shape, and 1.5 mm long and the flowers have 11-14 stamens; however, my observations of type material show that A. tessmannii has 3 flat petals, 2-3 mm wide and 14—16 mm long, and 6-9 (12) stamens. 6. Attalea dubia (Martius) Burret, Notizbl. Bot. Gart. Berlin-Dahlem 10:537. 1929a; Dahlgren, pls. 27—28, 32. 1959; Reitz, 118-24, figs. 52-55. 1974; Glassman, 1977a; Henderson, Galeano, and Bernal, 1995. Figs. 57, 74, 78. Orbignya dubia Martius, Hist. Natur. Palm 3:304, t. 169, fig. 6. 1845. LECTOTYPE: (Glassman, 1977a) Brazil, Rio de Janeiro, prov. Sebastionopolitana (Martius, Hist. Natur. Palm. 3:t. 169, fig. 6. 1845). Pindarea dubia (Martius) Hawkes, Arq. Bot. Est. S. Paulo 2:185. 1952. Attalea indaya Drude, Martius FI. Bras. 3:437, t. 100, fig. 2. 1881. LECTOTYPE: (Dahlgren, pl. 27, 1959) Brazil, Rio de Janeiro, Corcovado, Glaziou 8070 (C!; isolectotypes, BR!, F!, P!). Pindarea fastuosa Barb. Rodr., Pl. Jard. Rio de Janeiro 5:23, t. 5A. 1895. t. 59B. 1903b. LECTOTYPE: (Glassman, 1972a) Culti- vated. Brazil, Rio de Janeiro (Barb. Rodr., t. 5A, 1895). Attalea concinna (Barb. Rodr.) Burret, Notizbl. Bot. Gart. Berlin- Dahlem 10:537. 1929a; Glassman, 1977a. Fig. 73. Pindarea concinna Barb. Rodr., P|. Jard. Bot. Rio Jan. 5:17, t. 4c. 1895; Sert. Palm. Bras. 1:t. 59A. 1903b. LECTOTYPE: (Glassman, 1977a) Brazil, origin of specific locality not indicated; cultivated, Jard. Bot. Rio Jan. (Barb. Rodr., Sert. Palm. Bras. 1:t. 59A. 1903b). Trees 5-25 m tall and 20-40 cm in diam; complete sheathing leaf base not seen; petiole 1-2 m long, 15 cm wide near base and 6 cm thick, mar- gins fibrous; leaf rachis 6-7 m long; 114 pinnae on each side of rachis, Genus Attalea 35 those from middle series in distinct clusters of 2—4, 90-150 cm long and 4.5-6.5 cm wide, with acute or acuminate asymmetrical tips; expanded part of staminate sterile bract 65-78 cm long (including beak of 10-15 cm long), about 6 mm thick, and 20-30 cm wide, deeply sulcate except for smooth apex, peduncular part 30-60 cm long; rachis of staminate inflorescence 40-55 cm long, peduncle about 22 cm long, with many rachillae, lower ones 21—30 cm long, upper ones 15—20 cm long; stami- nate flowers in two rows on one side of the rachilla, 10-20 mm long, pet- als flat, 1.5—2.5 mm wide, with acuminate tips, distinctly nerved, sepals 1— 2 mm long, stamens 6-10 in number, anthers 5-8 mm long, filaments 1.0—2.5 mm long; expanded part of androgynous sterile bract about 1 m long (including beak of 20 cm long), 30-45 cm wide and 5 mm thick, deeply sulcate, peduncular part (incomplete) 20 cm long, with brownish lepidote indument; rachis of androgynous inflorescence 60—140 cm long (fide Barbosa Rodrigues, 1895), peduncle 1.0—1.2 m long (fide Barbosa Rodrigues, 1895), with many rachillae, pistillate portion of each rachilla 8-11 cm long, with 3-5 pistillate flowers, extended staminate rachilla 6— 15 cm long, staminate flowers (adjacent to individual pistillate flowers) up to 20 mm long, mostly sterile with tiny stamens, staminate flowers (on staminate part of rachilla) 10-15 mm long, mostly sterile with tiny unde- veloped stamens; pistillate flowers 2.5—4.0 cm long and 1.5-1.7 cm in diam, sepals and petals about same size, pistil 2.0—2.5 cm long and about 1 cm in diam, mostly light brown lepidote, stigmas 3 in number, 0.5—1.0 cm long, style very short or absent, staminodial ring dark brown, about 0.8 cm high; rachis of infructescence 1 m long, peduncle about 60 cm long; fruit 6.0-8.5 cm long (including beak of 1.5 cm long) and 3-4 cm in diam, persistent perianth 2-3 cm high, staminodial ring 1 cm high, with ciliate margins, epicarp fibrous, 1-2 mm thick, mesocarp soft, 3-4 mm thick, endocarp hard, 4—6 mm thick, fiber clusters scattered and incon- spicuous, seeds usually 1 in number, rarely 2, about 2.5 cm long and 1 cm in diam. Distribution and Habitat. See table 1. Reitz (1974) says that the distribu- tion of this palm in Santa Catarina is irregular and discontinuous, but is more frequent in coastal forests to the north of the state of Santa Catarina, principally in the municipalities of Garuva and Joinville, and is rare or ab- sent in the southern half of the state. According to Burret (1938), this spe- cies was frequently seen in forests in the vicinity of Rio de Janeiro and in the streets near the Jardim Botanico and was very common in forests ap- proaching the Haupstadt station. It was also seen in the state of Rio de 36 A Taxonomic Treatment of the Palm Subtribe Altaleinae (Tribe Cocoeae) Janeiro on a trip to Serra de Tijuca and in Serra Estrella and Serra do Mar in Itataya. No specimens were cited by Burret to verify these observations. Vernacular Names. Camarinha, palmito do chao, indaya, coqueiro indaia, indaya-assu, inaya, naia, perina, anaja. Representative Specimens Examined. BRAZIL. Sao Paulo: 225 km south of Sao Paulo, Feb. 1973, Krapovickas et al. 21354 (F, MO). CULTIVATED. BRAZIL. Jard. Bot. Rio Jan., Glaziou 17341 (F, US); Oct. 1893, Glaziou 20534 (B, MO, NY, P); Glaziou 14365 (C, F, MO, NY, P, US); Glaziou 14456 (B); Glaziou 16480 (C, F, G, MO, P); Burret 207 (B). Specimens Tentatively Included. BRAZIL. Sao Paulo: Caragauatatuba, swampy forest behind beach, Jan. 1985, Gentry and Zardini 49360 (F, MO). CULTIVATED. BRAZIL. Jard. Bot. Rio Jan., Bailey and Bailey 517 (BH); Jard. Bot. Sao Paulo, Toledo and Pereira s.n. (SP-50433); Jard. Bot. Lagoa de Freitas, Glaziou 16489 (F). No specimens were cited by Martius (1845) for A. dubia, nor could any be found. Therefore, his illustration was chosen as lectotype. The same situation applies to Pindarea fastuosa by Barbosa Rodrigues (1895, 1903b). For A. indaya, Drude (1881) cited both Glaziou 303 and 8070with no her- barium indicated. Glaziou 8070 (C) was chosen as lectotype because it was illustrated in Dahlgren (1959) and is represented by ample material. Speci- mens of Glaziou 303 could not be found, but photographs of an infructes- cence were seen at C. A. dubiais probably most closely allied to A. funifera because both have an arborescent habit and clustered middle series pinnae. It differs from A. funifera in the much smaller 1 seeded rather than 1-3 seeded fruits, thickness of mesocarp, and length of petiole fibers. 7. Attalea amygdalina Kunth in Humboldt, Bondpland, and Kunth, Nov. Gen. Sp. 1:310, ts. 95-96. 1816; Dugand, 1940, 1953; 1954; Glass- man, 1977a; Henderson, Galeano, and Bernal, 1995. TYPE: Colom- bia, dept. Choco, near Zitara, Humboldt and Bonpland s.n. (holo- type, P!). Figs. 1-2. Attalea victoriana Dugand, Mutisia 18:9. 1953. 20:4. 1954; Glassman, 1977a. TYPE: Colombia, dept. Valle, Correg. Galicia and Ceilan, east of Bugalagrande, sitio “E] Almendronal,” primeras estriba- ciones de la Cordillera Central, vertiente occidental, June 1945, Patino s.n. (holotype, COL 29718!). Figs. 29, 40, 81. Attalea uberrima Dugand, Mutisia 18:4. 1953. 20:5. 1954; Glassman, 1977a. TYPE: Colombia, dept. Caldas, munic. Palestina, Jan. 1942, Jaramillo Mejia 199 (holotype, COL!). Figs. 28, 82. Genus Aittalea 37 Acaulescent; leaves 7.0—7.5 m long; sheathing leaf base not measured; petiole about 55 cm long and 40 cm wide at base; middle series pinnae about 120 on each side of rachis, regularly arranged, about 5—6 cm apart, 90-125 cm long and 6-8 cm wide, with asymmetrical tips; expanded part of staminate sterile bract 2.35 m long (including beak of 19 cm long), deeply sulcate; rachis of staminate inflorescence 50—68 cm long, peduncle 55-110 cm long, rachillae numerous, 15-18 cm long, covered with whit- ish lepidote indument; staminate flowers in two rows on one side of rachilla (in Patino 213a few lower rachillae have one basal pistillate flower), 12-20 mm long, petals 2-6 mm wide, distinctly nerved, with acute or acuminate tips, sepals 1-2 mm long and 1 mm wide, stamens 12-21 in number, anthers 4.5—7.0 mm long, filaments 2—4 mm long; expanded part of androgynous sterile bract 1.5 m long and 18 cm wide, deeply sulcate; rachis of androgynous inflorescence 1.2—1.4 m long, peduncle 0.7—1.2 m long, androgynous rachillae numerous, pistillate portion 2—7 cm long, each with 2-3 pistillate flowers, 3.0-3.5 cm long and 2.5—3.0 cm in diam, petals shorter than sepals, pistil 2.5-3.0 cm long, densely ferrugineous pubescent on upper half, stigmas 3, about 1 cm long, staminodial ring about 1 cm high, extended staminate rachilla 10-30 cm long, covered with whitish lepidote indument, staminate flowers anomalous, 2—3 cm long, petals 5-7 mm wide, sepals 3-8 mm long, stamens 12-21 in number, in one row on one side of the rachilla; fruit 8-12 cm long, 4.0-6.3 cm in diam (including beak of 1.0—-1.5 cm long), persistent perianth 2.5—3.2 cm long, epicarp fibrous, about 1 mm thick, mesocarp soft and fibrous, 2-6 mm thick, endocarp hard, 4.5—5.0 cm thick, seeds 2—4 in number, 4—5 cm long and 1-2 cm in diam. Distribution and Habitat. See table 1. According to Bernal (1989), most of this species’ former habitat is now converted into coffee plantations, and it is considered to be endangered. Vernacular Names. Almendro, palma almendron, palma de cueso, taparo. Representative Specimens Examined. COLOMBIA. Valle: Patino s.n. (COL-29721 through 29729); Patino s.n. (COL-1177, includes sheet of photos); El] Almendronal, 12 km de Bugalagrande, 1945, Patino I (F); Caldas: Agudas, E] Boqueron (Pito), Aug. 1959, Patino 213 (COL); Antioquia: munic. Bolivar, Finca La Palmera, Aug. 1959, Patino 215 (COL). In 1977a, I designated Humboldt and Bonpland s.n. (P) as the lectotype of A. amygdalina because no specimens were cited in the original article. Since no other specimens of this species collected by Humboldt and Bonpland have been found, I am satisfied that the collection from P is the holotype. Dugand (1953) discussed the type locality with reference to comments 38 A Taxonomic Treatment of the Palm Subtribe Attaleinae (Tribe Cocoeae) by Karsten (1857:257). In addition to the locality listed above, Humboldt and Bonpland said that A. amygdalina was cultivated in orchards near Cartago and Guaduas. Dugand believed that the species seen by these two men in Guaduas was not the same one they saw in Cartago. They passed through Guaduas in April 1801 but traveled to Cartago in October dur- ing the time this palm usually flowers. He also thought that the plant seen in flower there was the basis for Humboldt and Bonpland’s description of A. amygdalina. In Cartago, they probably were informed that this spe- cies originated in the province of Choco, near Zitara, because it seems that the two explorers did not visit the Choco region themselves. This is the type species of Attalea because it is the first one described in the genus (Glassman, 1977a). The morphology of the staminate flowers definitely place it in the genus Attalea; however, there is insufficient her- barium material (four staminate rachillae, one pistillate rachilla, a packet of staminate flowers, an incomplete description, and an uncertain type locality) to delineate it as a clear-cut species. Therefore, I previously (1977a) designated A. amygdalinaas a species dubium. On the other hand, Henderson, Galeano, and Bernal (1995) recognize A. amygdalinaas a good species and place A. uberrima and A. victoriana in synonymy under it. Af- ter careful consideration, I have decided to follow Henderson, Galeano, and Bernal (1995) and treat A. amygdalina as a good species. This taxon seems to be most closely related to A. ferruginea, as both are acaulescent with regularly arranged pinnae and with similar-sized stami- nate flowers, anthers, and pistillate flowers. A. amygdalinais distinguished by having fewer stamens per flower (12—21 vs. 15-52), longer and broader middle series pinnae with glabrous rather than ferrugineous lepidote margins, and smooth rather than rugose and glandular pistillate flowers. De Nevers (1987) considered A. iguadummat to be allied to A. amygdalina (A. victoriana) because of the regularly arranged pinnae and acaulescent habit. They differ, however, in the arrangement of staminate flowers on the rachilla, size of androgynous rachillae, number of stamens, and size of anthers. 8. Attalea ferruginea Burret, Notizbl. Bot. Gart. Berlin-Dahlem 11:1044. 1934a; Glassman, 1977a. TYPE: Venezuela-Brazil border, Rio Negro, Lako s.n. comm. G. Huebner 166 (holotype, B!). Figs. 46, 59, 82. Acaulescent; sheathing leaf base 70-80 cm long, petiole 2.0—2.4 m long; leaf rachis 4.0—4.5 m long, densely ferrugineous lepidote on convex side, becoming glabrescent with age; pinnae rigid, 100-130 on each side of Genus Atialea 39 rachis, those from middle series regularly arranged, intervals of 3.5-10.0 cm between pinnae, 80-94 cm long and 3-4 cm wide, with acuminate, asymmetrical tips, margins, and especially apical 10-15 cm of pinnae usu- ally covered with ferrugineous lepidote indument on adaxial side, adaxial midrib usually lined with patches of brownish ramenta; expanded part of staminate sterile bract 30-50 cm long (including beak of 7-10 cm long), 5-7 cm wide, deeply sulcate, densely ferrugineous lepidote, peduncular part about 55 cm long; rachis of staminate inflorescence 25-35 cm long, peduncle about 55 cm long, rachillae 35-55 in number, 9-11 cm long, from 10-25 flowers per rachilla, arranged in one row along one side of the rachilla; staminate flowers 17—22 mm long, petals flat, slightly unequal in length, about 3-4 mm wide, with acute or acuminate tips, more or less nerved below, smooth above, margins with patches of brownish-white lepi- dote indument, sepals 1-2 mm long, stamens 16-52 in number (fide Wessels Boer, 1988), anthers 3—5 mm long, filaments 2.5-3.0 mm long; expanded part of androgynous sterile bract 40-57 cm long and 5-9 cm wide, deeply sulcate, covered with ferrugineous lepidote indument, pe- duncular part about 67 cm long; rachis of androgynous inflorescence 24— 35 cm long, peduncle 0.57-1.5 m long, with 20-30 rachillae, pistillate portion of each rachilla 2—4 cm long, with 1-2 pistillate flowers, staminate extension of each rachilla about 4 cm long with 8 staminate flowers; pis- tillate flowers 2.8—3.2 cm long and 1.8—2.5 cm in diam (3.5—4.5 cm x 3 cm in young fruit), sepals shorter than petals, about 2.5 cm long, ferrugine- ous lepidote mostly on apical part, petals usually rugose and covered with tiny glandular dots on narrowed apical portion, more or less nerved on basal part, usually with ferrugineous indument, becoming glabrescent with age, pistil 2.0—2.5 cm long and 1.3—1.5 cm in diam, usually dark brown- ish in color, stigmas 3 in number, 0.5—1.0 cm long, upper part of ovary covered with whitish tomentose indument, staminodial ring 0.8—-1.0 cm high, and about 2.5 cm in diam, dark brown in color, but usually covered with a dense brownish tomentose indument and tiny glandular dots, margins usually ciliate; complete infructescence not seen; fruit 6.5—9.0 cm long (including beak of 1-2 cm long), 4—5 cm in diam, usually dark brown in color and covered with a ferrugineous indument, persistent perianth 3.5—5.0 cm high, staminodial ring 1.0—1.5 cm high and 3—4 cm across, with brown ciliate margins, epicarp fibrous, 1.0—1.5 mm thick, mesocarp soft, about 1 mm thick, endocarp hard, 0.8—1.5 cm thick, fiber clusters con- spicuous in some fruits but obscured by dark brownish tannins in others, seeds 1—3 in number, about 3.5 cm long and 0.8—1.3 cm in diam. 40 A Taxonomic Treatment of the Palm Subtribe Attaleinae (Tribe Cocoeae) Distribution and Habitat. See table 1. According to Wessels Boer (1988), in Venezuela this palm occurs in relatively open forests on granitic table- lands and white sandy soils that are poorly drained and usually inundated during part of the year. Vernacular Name. Curua. Representative Specimens Examined. COLOMBIA. Vaupes: Rio Negro, San Felipe, and vicinity, Oct. 1952, Schultes et al. 18040 (BH, US). VEN- EZUELA. Amazonas: near Sanariapo, poorly drained sandy soil July 1967, Wessels Boer 1894 (U); Puerto Ayacucho, in well-drained sandy clay, July 1967, Wessels Boer 1906 (U); near San Carlos de Rio Negro, white sandy soil, Jan. 1968, Wessels Boer 2322 (U); dept. Atabapo, bosque ribereno in Trapichote, 15 km rio abajo de Santa Barbara, July 1982, Huber et al. 6440 (NY); Bolivar: from Puerto Ayacucho to Puerto Paez, savanna and poorly developed forest, along creeks, Aug. 1967, Wessels Boer 1945 (U). PERU. Loreto: prov. Maynas, Mishana, Rio Nanay, noninundated lowland forest, stamens 34—36 in number, May 1978, Gentry et al. 22347 (F, MO); between Rios Nanay and Itaya, on white sand, Nov. 1977, Gentry et al. 21010 (MO). Specimens Tentatively Included. PERU. Loreto: prov. Maynas, Mishana, Rio Nanay, upland forest on white sand, Mar. 1982, Gentry et al. 36535 (MO); July 1980, Gentry et al. 28878 (MO). VENEZUELA. Amazonas: dept. Atures, 23 km northeast of Puerto Ayacucho, evergreen lowland forest, Apr. 1978, Davidse and Huber 15357 (MO); dept. Atabapo, Cucurital del Caname, south bank, Middle Cano Caname, evergreen forest, Apr. 1977, Davidse et al. 17025 (NY). COLOMBIA. Santander, Curare Opon, June 1979, Renteria et al. 1499 (NY). This species appears to be closely allied to A. amygdalina from Colom- bia, as both are acaulescent with regularly arranged pinnae, have similar- sized staminate flowers and anthers, and have similar-sized pistillate flowers. A. ferruginea is differentiated by having a larger number of stamens per flower (16-52 vs. 12-21), shorter and narrower middle series pinnae whose margins and apices are ferrugineous lepidote instead of glabrous, and pistillate flowers with rugose and glandular dotted petals rather than smooth petals. A. ferruginea is also distinguished from most of the other species of Attalea by having almost all of its parts covered with a ferrugineous indument instead of being glabrous or having other types of indument. As I noted in 1977a, Wessels Boer equated A. racemosa Spruce with A. ferruginea probably because pinnae of the holotype (Spruce 54K) have ferrugineous margins. In fact, Henderson, Galeano, and Bernal (1995) Genus Attalea 4] reduce A. ferruginea to synonymy under A. racemosa. However, I am treat- ing A. racemosa as a species incerta because staminate flowers were not de- scribed or collected and because it was transferred to Orbignya by Drude in 1881. A more detailed discussion of A. racemosa can be found under “Doubtful and Uncertain Species of Attalea.” 9. Attalea nucifera Karsten, Linnaea 28:255. 1857. t. 68. 1861; Drude, t. 101. 1881; Burret, 1929a; Dugand, 1940, 1953, 1954; Henderson, Galeano, and Bernal, 1995. LECTOTYPE: (Glassman, 1977a). Co- lombia (Nova Granada): Guaduas (Karsten, Florae Columbianae 1:t. 68. 1861). Figs. 30, 55a, 55b, 81. Acaulescent; leaves 4.6—7.3 m long (fide Karsten), sheathing leaf base and petiole 1.5—2.5 m long; leaf rachis 5.0—5.5 m long; 90-146 pinnae on each side of rachis, those from middle series regularly arranged, 100-130 cm long and 5.5-6.7 cm wide, with asymmetrical tips; staminate inflorescence incomplete, rachillae numerous (6 cm long, fide t. 68, Karsten), 15-19 cm long (Castaneda 5339), covered with whitish lepidote indument; staminate flowers arranged in two rows along one side of rachilla, 14-17 mm long, petals flat, 2-3 mm wide, with acuminate tips, nerves more or less distinct at anthesis, becoming more prominent with age, with scattered or dense patches of reddish glands or short reddish glandular hairs at anthesis, becoming caducous and almost completely disappearing with age, sepals 1.5—-2.0 mm long, stamens 6-7 in number, anthers 5-6 mm long, filaments about 2 mm long; expanded part of an- drogynous sterile bract 31 cm long (including beak of 6 cm long) and 10 cm wide, peduncular part about 21 cm long, deeply sulcate; androgynous inflorescence mostly unbranched, rachis 15 cm long, peduncle about 23 cm long, about 50 pistillate flowers per inflorescence; pistillate flowers almost sessile, 2.0—2.5 cm long and 1.5 cm in diam, sepals shorter than petals, petals with strongly fluted margins, both distinctly nerved and with reddish glandular hairs; pistil about 2 cm long, ovary about 1 cm long and 1 cm in diam, densely brownish pubescent; stigmas 3, about 1 cm long, staminodial ring 0.5—-1.0 cm high, densely brownish pubescent; staminate flowers (from androgynous inflorescence) 12-14 mm long, with abundant reddish glandular hairs; fruiting sterile bract not measured; infructescence about 18 cm long, peduncle about 25 cm long, rachillae very short, about 0.5 cm long, each fruiting scar subtended by one long caudate bract about 6 cm long and two shorter bracts; fruit 6.5-10.0 cm long, (including beak of 1 cm long), and 4.5-5.0 cm in diam, persistent perianth about 2 cm 42 A Taxonomic Treatment of the Palm Subtribe Attaleinae (Tribe Cocoeae) high, staminodial ring about 3 cm across and 2 cm high, epicarp fibrous, about | mm thick, mesocarp soft, about 1 mm thick, endocarp hard, 5-8 mm thick, seed cavities 3 in number, 3.0-3.2 cm long and 1.0-1.5 cm in diam. Distribution and Habitat. See table 1. Vernacular Names. Palma real (Narino), taparo calimeno (Choc6), almendron (Santander). Representative Specimens Examined. COLOMBIA. Santander: east of Pt. Berrio, selvas compactas, May 1949, Scolnik 198016 (US); munic. Puerto Wilches, entre Gomez and 80 km del Atlantico R.R., Apr. 1960, Romero Castaneda 8392 (COL). Narino: Tumaco, near Pinal Dulce, Oct. 1955, Romero Castaneda 5339 (COL). Specimens Tentatively Included. COLOMBIA. Choc6: munic. Baudo, entre Cola del Barco and Canalete, May 1958, Patirio 206 (COL). No specimens were cited by Karsten (1857) nor could any be found; therefore, t. 68 (1861) was chosen as lectotype. Karsten (1857) described the staminate flowers as completely surround- ing the rachillae. As a result, both Drude (1881) and Burret (1929a) in- cluded A. nucifera in a separate section of Attalea because of this unusual flower arrangement for Attalea. A close look at t. 68 of Karsten (1861), however, reveals that the staminate flowers are actually in two rows along one side of the rachilla instead of spirally arranged. Collections purported to be this species (see specimens examined) also have the staminate flowers arranged in two rows on one side of the rachilla. Plate 101-I of Drude (1881) illustrates a pinna and staminate inflorescence based on Trail 212 (K) from Brazil (near Barreiras de Mutum), which is erroneously labeled A. nucifera. The staminate flowers completely encircle the rachilla, but the specimen is undoubtedly a species of Orbignya. A. nucifera differs from other Colombian species of Attalea as follows: Both A. nucifera and A. allenii are acaulescent and have a similar number of stamens, but A. nucifera has regular rather than clustered middle series pinnae. The remaining taxa are distinguished mainly by the number of stamens per flower. In A. nucifera there are 6—7, whereas A septuagenata has 60-75, A. amygdalina, 12-21, and A. ferruginea, 16-52. All of the above taxa have regular middle series pinnae and all except A. septuagenata are acaulescent. Even though certain discrepancies exist between Karsten’s original description (1857) and the specimens cited here, these can be resolved if one interprets the staminate flowers of the staminate rachilla in Karsten’s Genus Attalea 43 t. 68 (1861) to be in two rows on one side instead of spirally arranged. The other differences could be included within the range of morphological variation considered due to incomplete or broken off parts, e.g., length of staminate rachillae about 7 cm long in Karsten’s t. 68 and 15-19 cm long in Castaneda 5339. On a more positive note, staminate flowers of Castaneda 5339 and Scolnik et al. 19S016 match Karsten’s illustration closely, especially the glandular hairs on the petals. Therefore, in view of the above evidence, I am reasonably certain that the specimens cited here (except for Patino 206, which lacks staminate flowers, but has similar pinnae and an androgynous inflorescence with sessile pistillate flowers) are represen- tative of A. nucifera. With reference to Dugand’s (1953) cited specimens as examples of A. nucifera (e.g., Najar 11 [COL] and Rangel-Calindo s.n. [COL]), I have con- cluded that the first one is most probably A. alleni mainly because of the clustered pinnae, and the other specimen has insufficient material for positive determination. 10. Attalea iguadummat de Nevers, Ann. Missouri Bot. Gard. 74:506, fig. 2. 1987; Henderson, Galeano, and Bernal, 1995. TYPE: Panama, Colon, Santa Rita Ridge, Feb. 1986, de Nevers 7197 (holotype, CAS!; isotypes, K, MO, PMA). Acaulescent or with short trunk; petiole 55-80 cm long, 6-7 cm wide; leaf rachis 6.75—7.25 m long; 104—9 pinnae on each side of rachis, those from middle series regularly arranged, 144-48 cm long and 6.5-7.0 cm wide; staminate sterile bract 115-200 cm long and 30 cm wide; rachis of staminate inflorescence 35-45 cm long, peduncle 90-100 cm long, rachillae 40-50 in number, 13-18 cm long; staminate flowers spirally ar- ranged around rachillae, each 13-17 mm long, petals about 1 mm wide, flat, sepals 1.0-1.5 mm long, stamens 8-10, anthers 2-3 mm long, straight, filaments 2—3 mm long; androgynous sterile bract as in staminate inflores- cence, rachis of androgynous inflorescence 35-45 cm long, peduncle 60— 70 cm long; pistillate flowers on short pedicels or sessile, sepals 2.5—3.5 cm long, petals 4.0-4.5 cm long, stigmas 3, staminodial ring 1—2 cm high, with 15-20 lobes; infructescence with 45-65 fruits, 7-10 cm long and 4.5— 6.3 cm in diam, epicarp fibrous, 1-2 mm thick, mesocarp fibrous, 3-5 mm thick, endocarp 2-8 mm thick, without evident fiber clusters, seeds 1—3 in number. Distribution and Habitat. See table 1. Vernacular Name. /gua dummat (Kuna Indians of San Blas). 44 A Taxonomic Treatment of the Palm Subtribe Altaleinae (Tribe Cocoeae) Representative Specimens. De Nevers cites about six specimens from Panama, but I have seen only some of the type material. This is the second species of Attalea described from Panama. The other species, A. allenii, can be easily distinguished from A. iguadummat by its clustered rather than regular pinnae, its one-sided rather than spiral stami- nate flower arrangement, and its petals of staminate flower with punctate glands. Both species are acaulescent. A. iguadummat probably has few close relatives because of its unusual staminate flower arrangement. The only other species in the genus with spirally arranged staminate flowers is A. tessmannii, but it is arborescent with clustered middle series pinnae. According to Henderson and Balick (1987), the staminate petals of this species are similar to those of Scheelea. The petals are narrow (about 1 mm wide), but they are flat, like other members of the genus Aitalea. 11. Attalea humilis Martius ex Sprengel, Syst. Veg. 2:624. 1825; Martius, p. 121. 1844. t. 168, fig. 1. 1845; Burret, 1929a; Glassman, 1977a; Henderson, 1995. LECTOTYPE: (Dahlgren, pl. 29, 1959) Brazil, plures provincias, Princ. M. Neovidensis s.n. (M!). Figs. 15-17, 53, 60, 77. A. compta var. acaulis Martius, Hist. Natur. Palm. 2:t. 75. 1826. LEC- TOTYPE: (Glassman, 1972a) Brazil (Martius, t. 75, 1826). A. borgesiana Bondar ex Dahlgren, Trop. Woods 77:42. 1944; Bondar, Bol. Inst. Centr. Fom. Econ. Bahia 13:67, figs. 20-22. 1942b (sine descr. lat.). LECTOTYPE: (Glassman, 1977a) Brazil, Bahia, on hills, 100 m, in silicious soils of tertiary rock, Reconcavo de Bahia, Fazenda Carmo, munic. Sao Sebastiao, no date listed, Bondar s.n. (F-619755!). A. borgesiana Hawkes, Arq. Bot. Est. S. Paulo 2:176. 1952. Superflwous name. Acaulescent or with short trunk up to | m tall; sheathing leaf base about 50 cm long; petiole 50-80 cm long (70-150, fide Noblick, 1991) and 6 cm wide at base, margins fibrous; leaf rachis 2.5-5.3 m long; 85-130 pinnae on each side of rachis, those from middle series regularly arranged, about 1.5-3.0 cm apart, 65-95 cm long, and 3.5-5.5 cm wide, with acute, asym- metrical tips; expanded part of staminate sterile bract 0.9-1.8 cm long (including beak of 10-20 cm long), 8-13 cm wide, and 4 mm thick, deeply sulcate; rachis of staminate inflorescence 34-44 cm long, brownish or whitish tomentose, peduncle about 65 cm long, rachillae numerous, whit- ish tomentose, 6-11 cm long; staminate flowers arranged in two rows along Genus Attalea 45 one side of the rachilla, each 16-21 mm long, petals flat, somewhat un- equal, 2-3 mm wide, with acute or acuminate tips and denticulate mar- gins, distinctly nerved, sepals 1.0-1.5 mm long, stamens 6 in number, anthers 8-12 mm long, filaments 2—3 mm long; androgynous sterile bract 56-80 cm long (including beak of up to 16 cm long), 11-14 cm wide, deeply sulcate; rachis of androgynous inflorescence 28-36 cm long, pe- duncle 20-26 cm long, individual pistillate rachillae very short (about 1 cm long) or sessile; pistillate flowers 3—4 cm long, 1.0—1.5 cm in diam, sepals shorter than petals, both strongly nerved, pistil about 2 cm long, ovary whitish brown pubescent, style short, stigmas 3 in number, about 1 cm long, staminodial ring about 5 mm high; extended staminate rachilla 2—5 cm long, with 10-20 staminate flower scars arranged on one side, staminate flowers anomalous, 2—4 cm long, petals flat, 3-5 mm wide, with acuminate tips and denticulate margins, distinctly nerved, sepals 0.5—2.5 mm long, stamens 6 in number, anthers 7-11 mm long (or sometimes undeveloped—only 2 mm long), filaments about 3 mm long; fruiting ster- ile bract about 60 cm long and 17 cm wide, covered with brownish indument; rachis of infructescence about 28 cm long, peduncle about 31 cm long; fruits numerous, about 100 or more on each rachis; fruit 6-9 cm long and 4-8 cm in diam (including beak of 1 cm long), persistent peri- anth 3.5-4.0 cm high, staminodial ring 4-5 mm high, epicarp 1-2 mm thick, mesocarp soft and fibrous, 2—3 mm thick, endocarp hard, 8-12 mm thick, with inconspicuous scattered fiber clusters, seeds 1-3 in number, 1.5-2.5 cm long, 1.2—1.8 cm in diam, oily, with acid taste. Distribution and Habitat. See table 1. According to Martius (1844), A. humilis occurs in “eastern Brazil, prov. Sebastianopolis, Espirito Santo, Porto Seguro, Minas Gerais, Bahia and Pernambuco,” but no specimens have been seen from Minas Gerais or Pernambuco. According to Burret (1938), A. humilis was seen on dry hills and mountains in the vicinity of Rio de Janeiro, and it was also common between Rio de Janeiro and Serra Estrela, near Pilar and on the way to Serra do Mar. No specimens were collected to verify these observations. This palm reaches its northernmost limit near Sao Sebastiao do Passé in Bahia. Vernacular Names. Caiolé, pindova, palmerim, palmerinha, coco de pindoba (Burret, 1929a); pendoba mirim, pindoba brava, anaja mirim. Representative Specimens Examined. BRAZIL. Sao Paulo: Socavao, perto de Bananal Mandioca Palma, Feb. 1883, Herb. Saldanha 7106 (R); Espirito Santo: 1946, Bondar 20 (F-404620); Bahia: Bondar 22 (F-404620), on coast road between Alcobaca and Prado, 7 km northwest of Alcobaca 46 A Taxonomic Treatment of the Palm Subtribe Attaleinae (Tribe Cocoeae) and | km north along road from Rio Itanheninga, restinga, Jan. 1977, Harley et al. 17968 (F); munic. Alcobaca, Rod. BAO001, 5 km south of Alcobaca, restinga in solo arenoso, Mar. 1978, Mori et al. 9611 (NY); munic. Candeias, 50 km northwest of Salvador, 4—5 km south of turnoff to Sao Sebastiao, grassy hillsides, Aug. 1976, about 1,000 plants seen on two hill- sides, Glassman 13011 (F, SP), 13012 (F, SP), 13013 (F, SP), 13014 (F, SP); Amelio Rodrigues, remnant of Atlantic coastal forest, mata secundario, adjacent to bamboo forest, abundant on hillside, alt. 150 m, associated with Syagrus botryophora, Polyandrococos, Bactris, and Attalea burretiana, Aug. 1988, Noblick and Glassman 4575 (BAH, F); Rio de Janeiro: estrada do Porto da Estrella, Aug. 1925, Kuhlmann 19164 (RB); without locality, Glaziou 2756 (BH, BR, C, P). CULTIVATED. Jard. Bot. Rio Jan., Dahlgren s.n. (F-611620). Bahia: Fazenda Camurucu, Amelio Rodrigues, tree about 25 years old, trunk about | m tall, Aug. 1988, Noblick and Glassman 4577 (BAH, F). No specimens were cited by Martius (1825, 1844, 1845) for A. humilis or for A. compta var. acaulis (1826); lectotypes were chosen for each. Type specimens of all new species described by Bondar in his 1942a and 1942b articles were listed as isotypes deposited in F. Hence, I assumed that the holotypes were deposited in another herbarium in Brazil. Since the holotypes were never located, I designated all isotypes of Attalea in the above articles as lectotypes (1977a). According to the late Pedrito Silva of CEPLAC in Salvador, all of Bondar’s original palm collections (including type specimens) were discarded after he died in 1954. Fortunately, B. E. Dahlgren of the Field Museum of Natural History in Chicago previously had received a duplicate set of most of these specimens. Originally, I considered A. borgesianaas a distinct species, mainly because its habitat (grassy hillsides) is different from that of A. humilis (wet forested areas). The size differences of the flowering parts could be attributed to the fact that the grassy hillsides where A. borgesiana grows are subjected to continued burning and cutting. In fact, in August 1976, I counted about 1,000 plants of this species on two hillsides in the Sao Sebastiao area; but when I revisited the same area in August 1988, only a handful of palms were still present there. This particular region was probably originally Atlantic coastal forest but has been completely destroyed along with the palms by extensive burning and cutting for farming and grazing. A. humilis appears to be most closely related to A. geraensis, which is also acaulescent and has regularly arranged middle series pinnae. It differs from this species in the wider middle series pinnae, fewer stamens, longer anthers, fewer stigmas, and longer fruits with fewer seeds; A. humilzs is Genus Atialea 47 found mainly in restingas and coastal forests, whereas A. geraensis is mostly a cerrado palm. Apparently, A. humilis crosses with A. funifera to produce A. x voeksiiand with A. burretiana resulting in the putative hybrid A. salvadorensis. Both hybrids are rare, each known from one locality. 12. Attalea geraensis Barb. Rodr., Pl. Jard. Bot. Rio Jan. 6:22, t. 7. 1898 (“ceraensis”); Sert. Palm. Bras. 1:66, t. 56. 1903b; Glassman, fig. 10, 1967. 1977a; Noblick, 1991. LECTOTYPE: (Glassman, 1972a) Bra- zil, Minas Gerais, Alfenas (Barb. Rodr., t. 7. 1898). Figs. 4, 56, 63, 80. ? Attalea monogyna Burret, Notizbl. Bot. Gart. Berlin-Dahlem 10:534. 1929a; Glassman, 1977a. TYPE: Brazil, Goias, Serra dos Pyreneos, common in forest, July 1894, Glaziow 22269 (holotype, B, de- stroyed; isotypes, BH!, BR!, C!, K!, MO!, P!, US!). Attalea guaranitica Barb. Rodr., Palm. Paraguay. 27:t. 4d. 1899. t. 57B. 1903b. TYPE: Paraguay, ad Cordilla dos Altos, prope Pueblo Valenzuela, ad ripas Rio Y-Aka (Barbosa Rodrigues, 1899). LEC- TOTYPE: (Glassman, 1977a) Paraguay, Cordillera de Piribebuy, campo rupestre sicco, Hassler 1860 (G). Figs. 48, 64, 81. Acaulescent; sheathing leaf base about 20 cm long, petiole about 10 cm long (15-80 cm long, fide Noblick, 1991; probably includes petiole size of juvenile leaves) and 4 cm wide at base, both with long, fine fibrous margins and brownish lepidote indument; leaf rachis 1.4—2.2 m long, brownish lepidote on axis; 75—79 pinnae on each side of rachis, those from middle series regularly arranged, 46-71 cm long and 2.2—3.0 cm wide, with acuminate or aristate asymmetrical tips; expanded part of staminate ster- ile bract 35-50 cm long (including beak of 9 cm long), 7-10 cm wide, and 5 mm thick, sulcate except for apex, grooves 1.0—1.5 mm apart, pedun- cular part about 16 cm long; rachis of staminate inflorescence 20-30 cm long, peduncle 15 cm long (60 cm and 40 cm, fide Barbosa Rodrigues), with many rachillae, 3-8 cm long, about 20 flowers per rachilla, arranged in two rows along one side of the rachilla; staminate flowers 13-17 (21) mm long, petals flat, 2-5 mm wide, with acute or acuminate tips, distinctly nerved, margins denticulate, sepals 1-2 (3) mm long, stamens 6—10 in number, anthers 5-9 mm long, filaments 2-3 mm long; expanded part of androgynous sterile bract 29-34 cm long (including beak of 6 cm long), 4-6 cm wide, and 5 mm thick, deeply sulcate except for apex, peduncu- lar part about 20 cm long; rachis of androgynous inflorescence 15-27 cm 48 A Taxonomic Treatment of the Palm Subtribe Attaleinae (Tribe Cocoeae) long, peduncle about 20 cm long, with many rachillae, pistillate portion of each rachilla 2.5—4.0 cm long, with 1-3 pistillate flowers, staminate extension of each rachilla 2.5—3.0 cm long; pistillate flowers 2.5-3.5 cm long and 1.8—2.5 cm in diam, petals and sepals more or less equal in size, sepals sometimes shorter, pistil 1.8—2.0 cm long and 0.8-1.0 cm in diam, whitish brown lepidote on proximal half, stigmas mostly 3 in number (4— 6 in Bahia, fide Noblick, 1991), 6-10 mm long, style short or absent, staminodial ring 0.8-1.0 cm high and 1 cm across; expanded part of fruit- ing sterile bract 26-35 cm long (including beak of 10-14 cm long), 8-10 cm wide and 5 mm thick, sulcate, grooves about 1 mm apart, peduncular part about 19 cm long; rachis of infructescence 15-17 cm long, peduncle about 20 cm long, with many rachillae, each 1.5—3.5 cm long; mature fruit yellow, covered with a rusty velvet tomentum (Noblick, 1991) and very aromatic (fide Fonseca, 1927), 5-7 cm long and 3.0—4.5 cm in diam (in- cluding beak of 1-2 cm long), persistent perianth 2.7-3.5 cm high, staminodial ring about 1 cm high and 2 cm across, epicarp fibrous, 1.0— 1.5 mm thick, mesocarp soft, 1.5—4.0 mm thick, endocarp hard, 0.2—1.0 cm thick, fiber clusters arranged in a circle and mostly prominent, seeds 2—4 (6) in number, about 2 cm long and 1.5 cm in diam. Distribution and Habitat. See table 1. A. geraensis is probably the most widespread and one of the most common species of Aitalea in Brazil and extends into adjacent Paraguay. Vernacular Names. /ndaya do campo, catolé (Brazil); mbocaya guazu, coco de la cordillera, bocaia guazul (Paraguay). Representative Specimens Examined. BRAZIL. Sao Paulo: campos das Sete Lagoas, Fazenda Campininha, 3.7 km northwest of Padua Sales, Apr. 1960, Eiten and Eiten 1902 (BH); 4.6 km northwest of Padua Sales, cerrado, Sept. 1960, Exten and Eiten 2212 (BH); degraded campo cerrado, Exten and Eiten 2220 (BH); 4 km north of Padua Sales, cerrado, Aug. 1960, Ezten and Eiten 2208 (BH); Sertao de Itirapina, 12 km northwest of city, burned-over savanna, July 1965, Glassman and Gomes 8014 (F, SP), 8017 (F, SP); 5 km south of Emas, 7 km north of Pirassununga, in cerrado, common, July 1969, Glassman 8744 (F, SP), 8745 (F, SP). Minas Gerais: munic. Curvelo, 54 km north of Sete Lagoas, near entrance to road to Curvelo, highway BR7, cerrado remnant, July 1965, Glassman and Gomes 8106 (F, SP). Mato Grosso: 40 km south of Rondopolis, along BR163, common in cerrado, Sept. 1976, Glassman 13085 (F, SP), 13087 (F, SP). Goias: 26 km northeast of Rio Verde, dense cerrado, along roadside and plowed fields, about 75,000 plants seen in area, Aug. 1976, Glassman 13065 (F, SP), 13066 (F, Genus Attalea 49 SP), 13068 (F, SP), 13073 (F, SP), 13074 (F, SP). Bahia: munic. Baianopolis, 7 km south of Rio Angico on road to Inhumas and Correntina, cerrado, sandy soil, alt. 700—720 m, Oct. 1988, Noblick and Lima 4628 (BAH, CEPEC, CPATSA, F). Noblick (1991) also cites specimens from munic. Coribe and munic. Cocos, in Bahia. PARAGUAY. Cordilleras de Valenzuela, Nov. 1878, Balansa 3316 (P); Valle de l’y-acan guazu, near Valenzuela, Jan. 1884, Balansa 4775 (P); 5 km east of Caacupe, Cordillera de Altos, Apr. 1978, Schinini 14805 (F). Specimens Tentatively Included. BRAZIL. Minas Gerais: vicinity of Mendanha, Sept. 1936, Archer 4074 (US); without locality, May 1881, Glaziou 13296 (C, P). Cultivated. Jard. Bet. Rio de Janeiro, Jan. 1924, Bailey and Bailey 601 (BH), 601a (BH). A. geraensis is probably most closely allied to A. humilis. It differs mainly in its number of stamens, width of pinnae, and size of anthers. Burret (1929a) cites Glaziou 8069and 20022 for A. geraensis. The first is doubtful and probably out of range (coastal Rio de Janeiro), and 20022 has been identified as A. exigua. In 1977a, I treated A. monogynaas a syn- onym of A. oleifera because Burret said they were probably the same. The type specimens of A. monogyna (Glaziou 22269) appear to be closer to A. geraensis in total characteristics, and hence are tentatively included under that species here. No specimens were cited in the original article of A. guaranitica (Barbosa Rodrigues, 1899), but the locality was listed as Cor- dillera dos Altos, prope pueblo Valenzuela, ad ripas Rio Y-aka. However, Barbosa Rodrigues later cited Hassler 1860 (1903b), hence it was chosen as the lectotype. This palm is often used as an indicator of good soils and the seeds are edible. 13. Attalea septuagenata Dugand, Mutisia 18:3. 1953. 20:4. 1954; Caldasia 7:145. 1955; Glassman, 1977a; Henderson, 1995; Henderson, Galeano, and Bernal, 1995. TYPE: Colombia, Amazonas, Rio Miriti- Parana, Mar. 1952, Schultes and Cabrera 15796 (holotype, COL!; isotype, BH!). Figs. 27, 39, 66, 81. Trees 18—20 m tall; leaves 5.4—6.7 m long; sheathing leaf base and peti- ole not measured; petiole and leaf rachis glabrous on upper flat surface, densely dark ferrugineous lepidote, consisting of sharp, narrowly oblong, curved irregular scales, 1.0—1.5 cm long on lower convex surface; middle series pinnae regularly arranged, distributed at 4—5 cm intervals along rachis, 90-103 cm long and 3.5—4.5 cm wide, with asymmetrical tips; ex- 50 A Taxonomic Treatment of the Palm Subtribe Attaleinae (Tribe Cocoeae) panded part of staminate sterile bract 59 cm long (incomplete), 14 cm wide, and about 6 mm thick, deeply and irregularly sulcate, covered with a densely ferrugineous lepidote indument, peduncular part 39 cm long (incomplete) ; rachis of staminate inflorescence 1.0—1.2 m long, peduncle 1.2 m long, rachillae numerous, 13-19 cm long, whitish in color; stami- nate flowers on one side of the rachilla, mostly in two rows, 15-18 mm long, petals flat, with acute or acuminate tips, obscurely nerved, 3-4 mm wide, dark brown with white lepidote or farinose indument along margins, sepals about 1 mm long, stamens 60-75 in number (fide Dugand), anthers 4—5 mm long, very narrow, filaments 2.0-2.5 mm long; androgynous inflorescence not measured; androgynous rachillae about 9 cm long with 2 pistillate flower scars, pistillate flowers not seen; fruit 10.0-11.5 cm long and 5.5—6.0 cm in diam (including beak of 1.0—1.3 cm long), persistent perianth about 4 cm high, staminodial ring about 1.5 cm high, epicarp fibrous, 1.0-1.5 mm thick, mesocarp soft, about 3 mm thick, with conspicu- ous air pockets in cross section, endocarp hard, 1.0—1.5 cm thick, seeds 2-3 in number, 3.5—4.5 cm long and 1.3 cm in diam. Distribution and Habitat. See table 1. Only known from the Miriti- Parana area. Vernacular Names. Karjona, makuna, tanikuma, yukuna, mirana, kujita, kurua. REPRESENTATIVE SPECIMENS EXAMINED. Only type material was examined. Henderson (1995) cites Galeano et al. 2078 in which the stamens are only one-half the number reported by Dugand (1953). This palm differs from all of the other species of Attalea by the petiole and leaf rachis covered with dark, sharp, curved scales and the unusually large number of stamens (60-75). The only species that has stamens ap- proaching this number is A. ferruginea, which has 16-52 stamens. A. septuagenata also differs from this taxon mainly in the smaller staminate flowers (15-18 mm vs. 17-22 mm) and larger fruit (10.0-11.5 cm vs. 6.5— 8.0 cm) with conspicuous air pockets in the mesocarp. 14. Attalea seabrensis Glassman sp. nov. TYPE: Brazil, Bahia, munic. Seabra, Veredinha, gallery forest, Aug. 1976, Glassman 13034 (ho- lotype, F!; isotype, SP!). Figs. 23-26, 35, 45, 62, 80. Caudex 15-20 m altus, folium regulariter pinnatum ex parte, pinnis mediis 0.8—1.1 m longis, 4—6 cm latis, inflorescentia mascula multiramosa, pedunculo 40-50 cm longo, rachide 1.0-1.8 m longa, rachillis 15-20 cm longis, flores masculi 15-20 mm longi, stamina 9-13, antherae 5-9 mm Genus Attalea 51 longae, bractea androgyna 0.8—1.0 m longa, 15-30 cm lata, inflorescentia androgyna multiramosa, pedunculo 50-70 cm longo, rachidi 70-80 cm longi, rachillis 1-3 cm longis, flores feminei 4-5 cm longi, 2-3 cm diametro, fructus 6.5-9.0 cm longus, 5.5—7.0 cm diametro, epicarpio lignoso et fibroso, 2-6 mm crasso, mesocarpio pulposo, 1-3 mm crasso, endocarpio osseo, 1— 2 cm crasso, fibris conspicuis, semina 3-4, 3 cm longa, 1.2—1.5 cm lata. Trees 15-20 m tall and 25-40 cm in diam; petiole 0.6—2.0 m long; leaf rachis 4.0—6.3 m long; 150-200 pinnae on each side of rachis, only lower one-third to one-half of rachis with pinnae in clusters of 2—4, and lying in divergent planes, remaining pinnae regularly arranged, those from middle series 0.8-1.1 m long and 4—6 cm wide; staminate sterile bract 1—2 m long, 15-30 cm wide; rachis of staminate inflorescence 1.0—1.8 m long, peduncle 40-50 cm long, rachillae 250-90 in number, lower ones 15—20 cm long; staminate flowers arranged in two rows along one side of the rachilla, those on lower part 15—20 mm long, those on upper part 13-15 mm long, pet- als flat, 4-5 (8) mm wide, with acute to acuminate tips and denticulate margins, faintly or distinctly nerved, sepals 1.5—3.0 mm long, stamens 9— 13 (17) in number, anthers 5-9 mm long, filaments 1-2 mm long; ex- panded part of androgynous sterile bract 0.8—1.0 m long and 15-30 cm wide (including beak of 9-30 cm long), peduncular part 50-60 cm long; rachis of androgynous inflorescence 70-80 cm long, peduncle 50-70 cm long, rachillae numerous (250-300), pistillate portion of each 1-3 cm long, staminate extension 7—12 cm long, one pistillate flower per rachilla, each 4—5 cm long, 2-3 cm in diam, pistil 3 cm long and 1.5 cm in diam, stigmas 3-6 in number, 1.0—1.5 cm long, staminodial ring about | cm high; fruit 150-220 in number, 6.5—9.0 cm long and 5.5-7.0 cm in diam, red- dish brown in color, persistent perianth 3—4 cm high, staminodial ring 1.0— 1.5 cm high, epicarp woody and fibrous, 2-6 mm thick, mesocarp soft, yellow or orange when fresh, 1-3 mm thick, endocarp bony, 1—2 cm thick, with more or less conspicuous irregular fiber clusters, seeds 3—4 (6) in number, each 3 cm long and 1.2—1.5 cm in diam. Distribution and Habitat. Brazil, central Bahia, in transitional forest remnants, on hills and in valleys, associated with streams, mostly at eleva- tions of 500-1,200 m. In August 1988, Larry Noblick and I observed large populations of this palm from Barra do Mendes south to Seabra, Palmeras, Boninal, Piata, Jussiape, Ituacu, and Contendas do Sincora. Actually, Serra do Tombador is the dividing line between populations of A. seabrensis on the western slopes, which extend farther south, and A. pindobassu on the eastern slopes, which extend farther north along this mountain range. 52 A Taxonomic Treatment of the Palm Subtribe Altaleinae (Tribe Cocoeae) Vernacular Names. Palmerao, palmeira, catolé, acu. Representative Specimens Examined. BRAZIL. Bahia: munic. Seabra, Veredinha, about 291 km west of Salvador, depressed semi-moist lowland area, mostly in pure stands, in gallery forest of about 50 trees, Aug. 1976, Glassman 13035 (F, SP), 13036 (F, SP); Serra do Sincora, Caraibinha, 15- 25 km from Contendas do Sincora, 500-600 m, transitional forest rem- nant, about 2,000 trees in region, Aug. 1988, Noblick et al. 4590 (CEPEC, F); Serra do Sincora, Fazenda Andrai, munic. Ituacu, 10-15 km south of Barra da Estiva, transitional forest remnant, 940 m, about 100 trees seen in area, Aug. 1988, Noblick et al. 4591 (CEPEC, F); Fazenda Cocho, munic. Piata, Riacho Cochéo, 31 km south of Boninal, along stream, 1,060 m, about 300 trees seen, Aug. 1988, Noblick et al. 4597 (ALCB, F), 4598 (ALCB, F); Fazenda Lagoa, munic. Barra do Mendes, povoado of Areias, 25 km from Barra do Mendes, alt. 920 m, stand of about 50 trees, Aug. 1988, Noblick et al. 4599 (ALCB, F). The new species seems to be most closely related to A. pindobassu be- cause both are arborescent, have staminate flowers of about the same length with a similar number of stamens, have 6 stigmas in the pistillate flowers, and have similar-sized fruits with an exceptionally hard and thick (4-6 mm) epicarp. The lower one-third to one-half of the rachis in A. seabrensis have clustered pinnae and short anthers (5-9 mm); in A. pindobassu the lower pinnae are arranged in several widely spaced loose clusters of 2-3 and the anthers are 10-12 mm long. A. seabrensis is also probably closely related to A. burretiana. Differences are those mentioned above for A. pindobassu plus the much longer peti- oles (up to 2 m) and 3-6 stigmas. In A. burretiana, the pinnae are regu- larly arranged throughout; anthers are 9-10 mm long; petioles are very short or absent; epicarp of the fruit is only 1.5—-2.0 mm thick; and pistil- late flowers have 3 stigmas. Leaves of this palm are used to cover roofs; leaf rachises are used in construction of doors and walls; fruits are sweet tasting and seeds are uti- lized as food and as a cooking oil. 15. Attalea pindobassu Bondar, Field Mus. Natur. Hist. Bot. 22:462. 1942a; p. 62, figs. 15-16. 1942b; Henderson, Galeano, and Bernal, 1995. LECTOTYPE: (Glassman, 1977a) Brazil, Bahia, Serra de Ouro, 1940, Bondar s.n. (F-619761!). Figs. 18-22, 80. Trees up to 15 m tall and 35-50 cm in diam; sheathing leaf base about 1.5 m long; petiole 1.2—2.0 m long, leaf rachis 5-7 m long, abaxial side of Genus Attalea 53 rachis and petiole with ferrugineous or whitish indument, 160—90 pinnae on each side of rachis, lower pinnae with several widely spaced loose clus- ters of 2-3, those from middle series regularly arranged, 80-135 cm long and 5-7 cm wide with asymmetrical tips; staminate sterile bract 1.3—2.0 m long and 21-25 cm wide (including beak of 25-60 cm long); rachis of staminate inflorescence 0.8—1.5 m long, peduncle 0.4—1.0 m long, with ferrugineous indument, rachillae 220-470 in number, lower ones 18-31 cm long, whitish tomentose; staminate flowers arranged on one side of rachilla, mostly in two rows, lower ones 20-22 mm long, upper ones 15— 17 mm long, petals with acute tips and denticulate margins, 3-7 mm wide, stamens 8-12 in number, anthers 9-12 mm long; androgynous sterile bract similar in size to staminate bract, rachis of androgynous inflorescence | m long, peduncle 50-60 cm long, rachillae about 270 in number, pistil- late portion of rachilla 2.0—4.5 cm long, 1-2 pistillate flowers per rachilla, each 4.5 cm long and 2.5 cm in diam, petals and sepals 3.5—4.0 cm long, pistil about 4 cm long and 2 cm in diam, farinose on upper half, stigmas 6, whitish when fresh, turning black when dried, about 6 mm long, staminodial ring about 1.5 cm high; extended staminate rachilla about 19 cm long, staminate flowers arranged along one side of rachilla, mostly in one row, 9-12 mm long, petals flat with acute tips about 4 mm wide, some irregular in shape, sepals 2.0-3.5 mm long, stamens 11-12 in number, anthers only about 1 mm long and probably sterile, filaments 0.5 mm long; fruit 8—10 cm long and 5.0—6.5 cm in diam, yellowish brown in color, persistent perianth about 3-4 cm high, staminodial ring 1.0—1.3 cm high, epicarp woody-fibrous, 3-5 mm thick, mesocarp yellow or orange, oily, edible, and sweet in some fruits, flavorless in others, 1-3 (2-6) mm thick, endocarp bony, 8-10 mm thick, with inconspicuous fiber clusters, seeds usually 3—4 in number. Distribution and Habitat. Bondar (1942b) lists the following localities: Bahia, center of state, munic. de Campo Formoso, Saude, Jacobina, Miguel Calmon. Serra de Ouro, munic. Djalma Dutra (old name for Miguel Calmon) is on the caption under fig. 15. In August 1988, Larry Noblick and I observed several hundred thousand trees of this species in transitional forest areas between Pindobacu on the north and Tapirimuta on the south. No palms were seen in Pindobacu itself, but in the neighboring povoado of Tapicuru, thousands of pindobassu trees were seen along the hillsides. Between Pindobacu and Saude, about 1,000 more trees were observed; and between Saude and Jacobina, several thousand additional trees were seen, mostly after the village of Caem was 54 A Taxonomic Treatment of the Palm Subtribe Altaleinae (Tribe Cocoeae) passed. The greatest concentration of A. pindobassu was observed between Miguel Calmon and Itapura, in the Serra do Sapucaia (Serra do Ouro) for a distance of 50 km, at elevations of 450-600 m. We estimated there are probably several hundred thousand trees along hillsides and in valleys, and this region may possibly be the center of distribution for the species. This palm is usually found on the lower, wetter mountain slopes or in wet valleys at elevations of 350-900 m. Vernacular Names. Pindobassu (fide Bondar), palmeira, coco palmeira, babacu, coco babacu (fide Noblick). Representative Specimens Examined. BRAZIL. Bahia: Fazenda Santo Cristo, munic. Piritiba, near Posto Feliz, 12 km north of Tapirimuta, rem- nant of transitional forest, alt. 710 m, about 600 trees in area of 1,000 hectares, Aug. 1988, Noblick et al. 4600 (ALCB, F); Noblick et al. 4601 (ALCB, F); Fazenda Riachao, along Rio Mangabera, povoado of Tapicuru, turn- off north of town of Pindobacu, transitional forest, alt. 410 m, several thousand trees in area, Aug. 1988, Noblick et al. 4602 (ALCB, F); Noblick et al. 4603 (ALCB, F); Serra do Ouro, Fazendinho, munic. Jacobina, about 10 km south of Jacobina, dense transitional forest, dissected by streams, alt. 450 m, about 5,000 trees in area, Aug. 1988, Noblick et al. 4604 (ALCB, F); Fazenda Estreita Campo Grande, 2 km west of Miringaba, transitional forest remnant, alt. 730 m, a few hundred trees in area, Aug. 1988, Noblick et al. 4605 (ALCB, F). As previously mentioned, A. pindobassu seems to be closely allied to A. burretiana. It differs from that species mainly in having 6 instead of 3 stig- mas, a much longer petiole (0.5-1.5 m long), and lower pinnae with sev- eral widely spaced loose clusters of 2—3 rather than lower pinnae not clus- tered in this manner. This species is probably more closely related to A. seabrensis, which has clustered pinnae on the lower one-third to one-half of its rachis, whereas in A. pindobassu, the lower pinnae are in several widely spaced loose clusters of 2—3. They also differ in size of anthers and rela- tive prominence of fiber clusters in the endocarp. Both species are fairly tall trees with similar-sized staminate flowers and fruits with an exception- ally thick epicarp. According to Bondar (1942b), an edible oil extracted from the seeds of this palm is used in the diet of local inhabitants; a certain number of fruits are shipped to Salvador (Bahia) for use in the oil industry. Larry Noblick relates a story of a young boy (from a family of 14 children) from Serra de Tobira (Jussiape), a remote area in Bahia with no roads. The staple food of the family for a number of years was fruits of pindobassu: the Genus Attalea 55 yellow mesocarp (dende) and seeds were the main parts eaten. Leaves are used in dwellings, and the apical meristem is edible. 16. Attalea burretiana Bondar, Field Mus. Natur. Hist. Bot. 22:460. 1942a; Bol. Inst. Centr. Fom. Econ. Bahia 13:63-64, figs. 17-19. 1942b. LECTOTYPE: (Glassman, 1977a) Brazil, Bahia, Arata, Bondar 24 (F-619754!). Figs. 9-11, 47, 71, 78. Attalea concentrista Bondar, Field Mus. Natur. Hist. Bot. 22:461. 1942a, 1942b, 1964. LECTOTYPE: (Glassman, 1977a) Brazil, Bahia, munic. S. Antonio de Jesus, Castro Alves, Amargosa, Aeria, S. Ines, alt. 200—400 m, on granitic or gneissic terrains, Bondar s.n. (F-619759!). Trees 10-30 m tall and 35—40 cm in diam; sheathing leaf base about 0.65 m long, petiole absent, or 1-6 cm long and 17-30 cm wide near base, mostly with dark brown or grayish white indument, margins with relatively long stiff fibers 11-30 cm long, leaf rachis 5.5-8.5 (10) m long; 140-200 pinnae on each side of rachis, those from middle series regularly arranged, 95-140 cm long and 5-7 (8) cm wide, with acute, asymmetrical tips, par- tially covered with brownish lepidote indument,; staminate sterile bract 1.2-1.9 m long (including beak of 15-40 cm long) and 10-20 cm wide, deeply sulcate; rachis of staminate inflorescence 80-125 cm long, peduncle 45-62 cm long, rachillae about 300 in number, 22—26 cm long on lower part, 16-20 cm long on upper part; staminate flowers in two rows on one side of the rachilla, 18-25 mm long on lower part, 14-15 mm long on upper part, petals usually 3 in number (rarely 4, in Noblick 4722), flat, 3— 4 mm wide, with acuminate tips and denticulate margins, prominently nerved, sepals 1-2 mm long, stamens 6—9 in number, anthers 9-10 (14) mm long, filaments 2-3 mm long; androgynous sterile bract 2.3—2.5 m long (including beak of 25—40 cm long) and 25-45 cm wide, deeply sul- cate, rachis of androgynous inflorescence 0.85—1.2 m long, peduncle about 0.8 m long, rachillae numerous, pistillate portion of each rachilla either less than 1 cm, 6-7 cm, or 13-14 cm long, 1-2, 2-4, or 5-7 pistillate flowers per rachilla, staminate extension of each rachilla 15-16 cm long; in Noblick and Glassman 4584, rachillae on lower one-third of androgynous rachis with staminate flowers only, but rachillae on upper two-thirds of rachis with the normal arrangement of pistillate and staminate flowers; pistillate flowers 3.5—4.0 cm long and 2 cm in diam, sepals much shorter than pet- als, one 0.8 cm long, the other two fused, about 1.5 cm long, pistil 2.5 cm long, stigmas 3 in number, subsessile, about 1.2 cm long, staminodial ring 56 A Taxonomic Treatment of the Palm Subtribe Attaleinae (Tribe Cocoeae) about 5 mm high; fruit 8-11 cm long (including beak of 1 cm long) and 4.5-6.0 cm in diam, covered with a rusty brown indument, persistent pe- rianth 3-5 cm high, staminodial ring 1.0—2.5 cm high, 2.5—3.0 cm across, dark banded on outer and inner margins, epicarp fibrous, 1.5—2.0 mm thick, mesocarp soft, 3-6 mm thick, endocarp hard, 1-2 cm thick, with more or less conspicuous fiber clusters, seeds 1—3 in number, 3—4 cm long and 1.0—1.5 cm in diam. Distribution and Habitat. Brazil, Bahia, municipalities adjacent to Sal- vador, alt. 10-100 m, associated with A. x piassabossuand in Agua Comprida (Bondar, 1942b); confined to eastern Bahia in the Atlantic coastal forest and its remnants and in the transitional forest (mata cipo), from Serra da Pioneira, near Amargosa in the north, to Cravolandia and Gandu in the south, and extending southwest to several km north of Itororo. In August 1988, Larry Noblick and I observed dense populations of many thousands of trees on inaccessible, steep mountain slopes and in valleys on the road to Amargosa along highway BA046; this area may be the center of distri- bution for A. burretiana. Thousands of trees were also seen both south and north of Salvador, especially about 15 km north of Salvador on BR324; along the old airport road; and near Valeria, on the road to Feira Santana, in remnants of the Atlantic coastal forest, about 350 m in elevation. Other large populations of this palm were observed along BR101 both north and south of Gandu for about 50 km or more. According to Noblick (1991), populations of A. burretiana were also observed within the borders of Sergipe on the north and Espirito Santo on the south, and it appears to thrive in areas where the original forest was cleared. Vernacular Names. Palmeira, andaia, catolé, pindoba-grauda. Representative Specimens Examined. BRAZIL. Bahia: Aratu, Bondar s.n. (F-619753); without definite locality, 1940, Bondar s.n. (F-619758); munic. Cravolandia, Fazenda Ponto Novo, 25 km east of Santa Ines, mata primario, about 1,000 trees seen in area, Aug. 1976, Glassman 13008 (F, SP), 13009 (F, SP); vicinity of Arata, 25 km west of Salvador, dense mata, about 75-100 trees seen in area, Aug. 1976, Glassman 13010 (F, SP); Dom Macedo Costa, 2 km leste da cidade, mata hygrofilica, yellow latosol red- dish clay, only a few trees seen, rare, May 1985, Noblick and Lemos 3810 (F, HUEFS); Fazenda Serra Branca, Amelio Rodrigues, remnant of Atlantic coastal forest, mata secundario, stand of about 100 trees, Aug. 1988, Noblick and Glassman 4574 (BAH, F); Noblick and Glassman 4576 (BAH, F); city limits of Salvador, Avenida Parallela, circa 1 km from center de Adminis- tracao da Bahia going toward airport, Atlantic coastal forest, about 750 trees in area, Aug. 1988, Noblick and Glassman 4582 (BAH, F); Noblick and Genus Attalea 57 Glassman 4584 (BAH, F); 20 km north of Gandu, remnant of transitional forest, along BR101, a few thousand trees seen in area along hillsides, Aug. 1988, Noblick et al. 4588 (CEPEC, F); Alto Seco, 20 km from Laje, alt. 350 m, in mata cipo, several hundred trees seen in valleys and on steep slopes in region, Aug. 1988, Noblick et al. 4589 (CEPEC, F); on Fazenda, 46 km northeast of Itororo, alt. 300 m, along BR101, on grassy hillsides and in remnants of transitional forest with Syagrus romanzoffiana as one of its as- sociates, a few hundred trees seen in region, Aug. 1988, Noblick et al. 4594 (CEPEC, F); 10-15 km from Palmira, along rocky dirt road, remnant of transitional forest, about 50 trees in area on steep, inaccessible slopes, Aug. 1988, Noblick et al. 4595 (CEPEC, F); Noblick et al. 4596 (CEPEC, F); munic. Jandaira, 14-18 km east of Jandaira, Fazenda Lagoa Escura, disturbed coastal rain forest, Dec. 1988, Noblick 4722 (F). A. burretiana seems to be most closely related to A. pindobassu and A. seabrensis. Similarities and differences between these three species have been discussed previously. Medeiros-Costa (1985) said that A. burretiana may be conspecific with A. camposportoana (now transferred to A. apoda). According to my investi- gations, they are distinct species and differ as follows: A. burretianais found primarily in eastern Bahia and has staminate rachilla 15-17 cm long, an- thers 9-10 mm long, 3 stigmas, fruits 8-11 cm long with 1-3 seeds, and mesocarp 3-6 mm thick; A. camposportoana (A. apoda) has been collected mainly in Minas Gerais and has staminate rachillae 10-15 cm long, anthers 7-8 mm long, 4 stigmas, fruits 5-7 cm long with one seed, and mesocarp about | mm thick. A. burretiana crosses with A. funifera to produce A. x piassabossu in a few localities in the vicinity of Salvador. This taxon will be treated in more detail under “Hybrids of Attalea.” Henderson, Galeano, and Bernal (1995) reduce this species to synonymy under A. oleifera Barb. Rodr. Leaves of A. burretiana are used as a covering for houses and to shade vegetable crops. Fruits and seeds are edible, and seeds yield a fine cook- ing oil. 17. Attalea oleifera Barb. Rodr., Nov. Rev. Bras. Rio Jan. 7:123. 1881; Sert. Palm. Bras. 1:69, t. 58. 1903b; Burret, Notizbl. Bot. Gart. Berlin- Dahlem 10:534. 1929a; Henderson, Galeano, and Bernal, 1995. LECTOTYPE: (Glassman, 1977a) Brazil, common in stands in arid or dry lands along the basin of Rio Sao Francisco, prov. Alagoas, Pernambuco, and Ceara (Barb. Rodr., Sert. Palm. Bras. 1:t. 58, 1903b). Figs. 41, 67, 77. 58 A Taxonomic Treatment of the Palm Subtribe Altaleinae (Tribe Cocoeae) Trees 7—25 m tall and 30-45 cm in diam (fide Barb. Rodr.); sheathing leaf base not measured; leaves 9 m long, petiole 0.6—1.5 m long, leaf ra- chis 4.0—7.5 m long, both with ferrugineous indument,; 180-200 pinnae on each side of rachis, intervals of 3-9 cm between some pinnae, those from middle series regularly arranged, 80-115 cm long and 3.0—5.5 cm wide, with asymmetrical tips; staminate sterile bract 0.5-2.6 m long (fide Barb. Rodr.) (including beak of 30 cm long), 19—26 cm wide, covered with a ferrugineous tomentum, deeply sulcate; rachis of staminate inflorescence 1.0-1.4 m long, peduncle about 1.25 m long, both covered with a whitish tomentum, rachillae numerous, 18-22 cm long, covered with a brownish white lepidote indument; staminate flowers arranged in two rows along one side of the rachilla, 1.5—2.5 cm long, petals 2-3 mm wide, distinctly nerved, with acuminate tips and denticulate margins, sepals 1-2 mm long, stamens 6—9 in number, anthers 6—8 mm long, filaments |.5—2.0 mm long; androgynous sterile bract 0.65—2.65 m long (fide Barb. Rodr.) (including beak of 30 cm long), and 24—40 cm wide, covered with a ferrugineous tomentum, deeply sulcate; rachis of androgynous inflorescence 1.0—1.3 m long, peduncle about 1.25 m long, both covered with a whitish tomentum, androgynous rachillae numerous, pistillate portion 10—11 cm long (25 cm long, fide Barb. Rodr.), covered with a brownish white lepidote indument, with 6-8 pistillate flowers (2-4, fide Lyra-Lemos, 1987), each 2.5-3.5 cm long and 1.5—2.0 cm in diam, pistil 2.5 cm long and 1.2 cm in diam, cov- ered with brownish lepidote indument, stigmas 3, about 6 mm long, staminodial ring about 7 mm high; extended staminate rachilla 8—11 cm long; fruit 7-9 cm long and 4.0—4.5 cm in diam, ferrugineous tomentose, epicarp fibrous, about 1 mm thick, covered with ferrugineous tomentum, mesocarp soft, about 3 mm thick, watery-oily, endocarp hard, 7-10 mm thick, dark brown, with scattered fiber clusters, seeds | in number, some- times 2, 3.0-3.5 cm long, endosperm solid, oily. Distribution and Habitat. See table 1. Medeiros-Costa (1982) reported this species from Paraiba, Alagoas, Sergipe, Bahia, and Minas Gerais, but cites specimens only from Pernambuco; and Lyra-Lemos (1987) cited specimens from Alagoas. Noblick (1991) did not list this palm from Ba- hia. The reports of A. oleifera from Rio Sao Francisco and Minas Gerais by Barbosa Rodrigues probably should refer to A. compta. Vernacular Names. Catolé, palmeira, pindoba (Medeiros-Costa, 1982). Representative Specimens Examined. BRAZIL. Cultivated. Rio de Janeiro: Jardim Publico de Campos, Rio de Janeiro, 1882, 1885, Glaziou 15556 (B, C, G, MO, P). Genus Attalea 59 A. oleiferaseems to be closely allied to A. compta because both are arbores- cent with regularly arranged pinnae and have similar-sized anthers. It dif- fers from this taxon mainly in the longer staminate rachillae, the shorter staminate flowers, and the smaller l-seeded fruits. Fruit is economically important: the mesocarp and endosperm of seed are oily and are used by native inhabitants as a source of cooking and il- luminating oil; leaves are used as a thatching material. 18. Attalea compta Martius, Hist. Natur. Palm. 237. & 41. 97. lel: Bondar, figs. 1-3, 11. 1942b; Dahlgren, pl. 26. 1959; Balick, Ander- son, and Medeiros-Costa, 1987. LECTOTYPE: (Glassman, 1977a) Brazil, in plures provincias, Princ. M. Neovidensis s.n. (M!). Figs. 50, 79. Trees 10-15 m tall, about 15 cm in diam (2.5—7.5 m x 20-32 cm, fide Balick, Anderson, and Medeiros-Costa, 1987); sheathing leaf base 0.4—1.3 m long, petiole 10-20 cm long, margins coarsely fibrous; leaf rachis 4.0— 6.5 m long; 132-91 pinnae on each side of rachis, those from middle se- ries regularly arranged about 1.5—1.8 cm apart, 90-110 cm long, and 3.5— 4.0 cm wide, with abundant reddish punctate dots on abaxial side (observed in lectotype, but not mentioned by Balick, Anderson, and Madeiros-Costa, 1987 in their description), more or less prominent cross veinlets on adaxial side, with acute asymmetrical tips, margins of pinnae covered with ferrugi- neous lepidote indument chiefly near tips; expanded part of staminate sterile bract 0.75-1.60 m long, 11-30 cm wide, and 1 cm thick, deeply sulcate and plicate as well, grooves irregularly distributed; rachis of stami- nate inflorescence 0.8—1.0 m long, peduncle 0.4—1.3 m long, rachillae numerous, 15—18 cm long; staminate flowers 15-18 mm long, petals flat, 4—5 mm wide, with acute to acuminate tips, distinctly nerved, sepals 2mm long, stamens 10-12 in number, anthers 7-8 mm long, filaments about 1 mm long; androgynous inflorescence and pistillate flowers not recorded; fruit 8.0—9.5 cm long and 5.0—-5.5 cm in diam, persistent perianth 4.0-4.5 cm high, staminodial ring 0.5-1.0 cm high and 2.5 cm across, epicarp fibrous, 2—3 mm thick, mesocarp soft, 2-3 mm thick, oily, endocarp woody, about 1 cm thick, fiber clusters inconspicuous, seeds 3—4 in number, about 3 cm long and 1 cm in diam. Distribution and Habitat. Martius (1826) listed A. compta from the fol- lowing nine different Brazilian states: along the coast in forests of Bahia (Ilheus), Piaui, Pernambuco, and Maranhao and in Espirito Santo (Porto Seguro), Rio de Janeiro, Sao Paulo, Minas Gerais, and Goias. Bondar 60 A Taxonomic Treatment of the Palm Subtribe Attaleinae (Tribe Cocoeae) (1942b) said that this species forms dense forests in the north of Espirito Santo and in all of the coastal area of southern Bahia up to Reconcavo in the municipalities of Pocoes, Boa Nova, Jequie, and Rio Novo. According to Balick, Anderson, and Medeiros-Costa (1987), A. compta occurs in the state of Minas Gerais in the municipalities of Uberlandia and Santa Fé, where it hybridizes with O. oleifera to produce X Attabignya minarum. This taxon will be discussed in more detail under “Hybrids of Attalea.” Noblick (pers. comm.) collected specimens of A. compta from Vazante, Minas Gerais, near the headwaters of Rio Sao Francisco; however, no Orbignya palms were observed in this location. From Martius’s description of A. compta occurring “in plures provincias,” it would be impossible to determine the exact type locality. The particu- lar specimen was chosen as lectotype because it contains information simi- lar to that in the original article and was also illustrated by Dahlgren (1959). Several other specimens collected by Princ. M. Neovidensis deter- mined as A. compta are also deposited at M. Some have no localities listed, but two others are inscribed with Rio Sao Francisco. The main course of this river runs through Bahia and into Minas Gerais for some distance until it branches into several smaller tributaries above Pirapora. Balick, Ander- son, and Medeiros-Costa (1987) reported A. compta from Uberlandia, which is more than 200 km west of these tributaries, and from Santa Fé, which is considerably closer to Rio Sao Francisco. Before trying to estab- lish the probable type locality, it will be necessary to make more collec- tions to get a better idea of the distributional range of this palm. As it now stands, the only precise localities are those cited in Balick, Anderson, and Medeiros-Costa (1987) and recent collections of Noblick from Minas Gerais. I have seen the following specimens determined as A. compta by Bondar from Bahia and Espirito Santo, without specific localities: Bondar s.n. (F-619757) and Bondar 18 (F-404618); the specimens of the first one do not match the original description closely, and the specimen from Espirito Santo consists of leaf parts only. The localities mentioned above for the distribution of A. compta probably refer to other species of Attalea that occur in eastern Bahia and Espirito Santo, such as A. funifera, A. humilis, and A. burretiana. Recent collections of the last three taxa by Larry Noblick and me have enabled us to pinpoint the morphological charac- teristics and distribution patterns more accurately. It is not likely that A. compta is found in all the nine states mentioned by Martius (1826), nor is it likely that it occurs in both eastern Bahia and central Minas Gerais, since there is such a wide disjunction between these two places. There are Genus Aitalea 61 sufficient differences between A. comptaand the above three species from eastern Bahia to conclude that the native distribution of A. comptais prob- ably confined to Minas Gerais, mainly west of Rio Sao Francisco. According to a survey done by Tenorio (1982), A. compta was observed in about 50 different localities in Minas Gerais. It is uncertain how many collections were made to verify its identity, but if most of these observa- tions are accurate, then A. compta must be considered a common or abun- dant species, probably with populations of several hundred thousand trees. Vernacular Names. Pindoba-ussu, babassu (Espirito Santo); naza, catolé (Bahia); andaja, ndaja (Princ. M. Neovidensis); indaia, andaia, palmeira (Minas Gerais, fide M. Balick); bandarra, baguagu (Minas Gerais). It should be noted that the names listed above for Espirito Santo and Bahia prob- ably belong to other species of Attalea. Representative Specimens Examined. BRAZIL. No localities listed on two sheets, but Rio Sao Francisco listed on two others, Princ. M. Neovidensis s.n. (M). This species appears to be closely allied to A. brasiliensis because their staminate flowers are similar in size with a similar number of stamens and their anthers are similar in size. A. compta differs chiefly in the longer stami- nate rachillae, distinctly nerved rather than obscurely nerved staminate petals, and larger fruits with inconspicuous rather than prominent fiber clusters in the endocarp. It also seems to be closely related to A. oleifera, differing principally in the narrower pinnae, shorter staminate rachillae, longer staminate flowers, and longer fruits with several seeds rather than one seed. Henderson, Galeano, and Bernal (1995) reduce this species to synonymy under A. oleifera Barb. Rodr. 19. Attalea apoda Burret, Repert. Spec. Nov. Regni Veg. 32:105. 1933. TYPE: Brazil, Minas Gerais, Porto de Rio Paracatt and Piquiero, sandy cerrado, Sept. 1895, Glaziou 22266 (holotype, P!; isotypes, BR!, C!, G!, MO!). Attalea camposportoana Burret, Notizbl. Bot. Gart. Berlin-Dahlem 14:257. 1938; Glassman, 1977a. TYPE: Minas Gerais, Serra de Mantiqueira, on road between Juiz da Fora and Barbacena, by Palmyra, Dec. 1937, Burret 17 (holotype, B!; isotype, RB!). Figs. 2586). 79: Trees 8-13 m tall; sheathing leaf base not measured; petiole (incom- plete) about 34 cm long and 8 cm wide; leaf rachis about 6.7 m long (fide Burret), more or less covered with a brownish indument; middle series pinnae regularly arranged along rachis, with brownish indument along 62 A Taxonomic Treatment of the Palm Subtribe Attaleinae (Tribe Cocoeae) edges, 96-110 cm long and 3.5—4.7 cm wide (7 cm, fide Burret), mostly with acuminate, asymmetrical tips; staminate sterile bract 2 m long (fide Burret) (including beak of about 27 cm long), deeply sulcate; rachis of staminate inflorescence 60-80 cm long, peduncle 18 cm long, rachillae numerous, 12—14 cm long; staminate flowers arranged in two rows along one side of the rachilla, 17-20 mm long, 3-5 mm wide, petals strongly nerved, sepals 1 mm long, stamens 8—9, anthers 6-8 mm long, filaments 2 mm long; complete androgynous sterile bract not measured, deeply sulcate, beak about 15 cm long; rachis of androgynous inflorescence 30 cm long (incomplete), peduncle 24-32 cm long, rachillae numerous, pistillate portion of each rachilla 1-3 cm long, 2-4 pistillate flowers per rachilla, staminate extension of each rachilla 5-7 cm long, with many staminate flowers; pistillate flowers 1.8—2.0 cm long, pistil about 1.5 cm long, stigmas 4 in number, staminodial ring about 0.6 cm high; staminate flowers (from staminate extension) 15-22 mm long, petals flat, about 5 mm wide, with acuminate tips and denticulate margins, more or less prominently nerved, sepals 2 mm long, stamens 9 in number, anthers 6- 7 mm long, filaments about 2 mm long; fruit 5—7 cm long and 3.0-3.5 cm in diam, persistent perianth 3—4 cm high, epicarp about 1 mm thick, mesocarp | mm thick, endocarp bony, 5-8 mm thick, with scattered, in- conspicuous fiber clusters, seeds ] in number, 2.3 cm long and 1.9 cm in diam. Distribution and Habitat. See table 1. Vernacular Name. Catolé. Representative Specimens Examined. BRAZIL. Minas Gerais: Fazenda Catolé, 27 km east of Copelinha, along Catolé correqeio, along hillside and in Manihot plantation, about 1,000 young stemless plants and mature trees seen, Aug. 1976, Glassman 13004 (F, SP), 13005 (F, SP), 13006 (F, SP), 13007 (F, SP); munic. Juiz da Fora, 7 km south of Eubang de Camara, 10 trees seen on fazenda grazing area, also common for 20 km north of Juiz da Fora, Aug. 1976, Glassman 13000 (F, SP). Specimens Tentatively Included. BRAZIL. Minas Gerais: Serro de Palacio, part of stand of thousands of trees, July 1965, Glassman and Gomes 8043 (F, SP). Goias: Meia Ponte, Rio Vagabine, virgin forest, July 1894, Glaziou 22270 (BR, C, K, MO, P). This taxon differs from A. compta, which also occurs in Minas Gerais, mainly in the fewer number of stamens (8-9 vs. 10-12), smaller fruits (5— 7 cm vs. 8.0-9.5 cm) and fewer seeds (1 vs. 3-4). A. apoda seems to be closely related to A. oleifera. Both are arborescent, have regularly arranged Genus Attalea 63 pinnae, and have l-seeded fruits. They differ primarily in the size of their fruits, the size of their staminate and pistillate rachillae, and the number of stigmas. This species also appears to be closely related to A. brasiliensis; comparisons will be discussed under that species. Figure 9, illustrated in my 1967 article, shows a stand of Attalea from Serro de Palacio that may possibly be this species. Unfortunately, only fruits were collected from these trees. A. apoda Burret (1933) was originally thought to be a synonym of A. geraensis Barb. Rodr., but after reexamining the type specimens, I discov- ered that its characteristics matched those of A. camposportoana more closely than those of A. geraensis, especially since the pistillate flowers have four stigmas. Other features, such as the size of staminate flowers, num- ber of stamens, and size of anthers also match those of A. camposportoana closely. When this change was made, it became apparent that A. apoda is now the “correct” name because it was described by Burret at an earlier date (1933) than A. camposportoana (1938) was. Henderson, Galeano, and Bernal (1995) reduce this taxon to synonymy under A. speciosa Martius, which I treat as a synonym of O. phalerata Martius. 20. Attalea salvadorensis Glassman, sp. nov. TYPE: Brazil, Bahia, 78 km north of Salvador (vicinity of Amelia Rodrigues), near turnoff for Itapetingui, along highway BR324, mata secundario, in Atlantic coastal forest remnant, associated with Elaeisand Bactris, about 200 Attalea trees seen in area, Aug. 1976, Glassman 13016 (holotype, F!; isotype, SP!). Figs. 51, 72, 77. Caudex ca 30 m altus, folium regulariter pinnatum, pinnis mediis 106— 18 cm longis, 3.5—4.2 cm latis; bractea mascula 61 cm longa, inflorescentia mascula multiramosa, pedunculo 67 cm longo, rachide 40—45 cm longa, rachillis 10-11 cm longis, flores masculi 13-15 mm longi, petalis 3-4, stamina 6-9, antherae 7-9 mm longae; fructus 9 cm longus, 5 cm diamet- ro, epicarpio 1.5—2.0 mm crasso, mesocarpio 2.5—3.0 mm crasso, endocar- pio osseo, 10-15 mm crasso, fibris inconspicuis, semen 1. Trees up to 30 m tall; petiole 80-95 cm long; leaf rachis 6.0—7.5 m long; pinnae regularly arranged along rachis, those from middle series 106-18 cm long and 3.5—4.2 cm wide; expanded part of staminate sterile bract about 61 cm long and 10 cm wide, peduncular part about 72 cm long; rachis of staminate inflorescence 40-45 cm long, peduncle 67 cm long, rachillae numerous, 10-11 cm long; staminate flowers arranged in two rows along one side of the rachilla, each 13-15 mm long, mostly with 3- 64 A Taxonomic Treatment of the Palm Subtribe Attaleinae (Tribe Cocoeae) 4 petals, each about 2 mm wide, strongly nerved with acuminate tips and denticulate margins, sepals 3-4 in number, about 1 mm long, stamens 6-— 9 in number, anthers 7—9 mm long, filaments about 2 mm long; complete androgynous inflorescence not seen; fruit about 9 cm long and 5 cm in diam (including beak of 1.5 cm long), persistent perianth 3—4 cm high, staminodial ring about 1 cm high, 2.5 cm across, epicarp fibrous, 1.5—2.0 mm thick, mesocarp soft, 2.5—-3.0 mm thick, endocarp hard, 10-15 mm thick, fiber clusters scattered, not conspicuous, seeds 1 in number, off- center, about 1.5 cm in diam. Distribution and Habitat. See table 1. Vernacular Names. None recorded. Representative Specimens Examined. Only material from the type lo- cality has been collected. Superficially, this newly described species seems to be most closely allied to A. brasiliensis from Goias because both have regularly arranged pinnae, are arborescent, have relatively short staminate rachillae, and have stami- nate flowers similar in size. It differs primarily in the staminate flowers with 3-4 petals instead of 3, which are strongly, rather than obscurely, nerved and longer fruits with inconspicuous fiber clusters. Staminate flowers (from the staminate inflorescence) with 4 petals are unusual in the genus Aftalea. Therefore, I am not certain whether the 4 petals in this particular speci- men is teratological or if it is an inherited trait. The flowers, however, do not appear to be infested with insects or attacked by a bacterial, viral, or fungal disease. Also, the single seed is off-center, indicating that one or more other seeds may have been suppressed from developing. A more plausible explanation for the origin of A. salvadorensis may be that it is a hybrid between A. burretianaand A. humilis, the two most com- mon species of Attalea in the general area where it was collected. Table 2 shows a comparison of the three taxa. The data in the table reveal that only one characteristic, pinnae length, is more or less intermediate between the two taxa. The following features of A. salvadorensis are closer to those of A. burretiana: size of plant, rachis length, and stamen number; the length of petiole, pinnae width, length of stami- nate rachillae, microscopic cross sections of pinnae, and size of fruit more closely resemble those of A. humilis. Its flower length is smaller than those in either putative parent, and the 3-4 petals are unlike those in either one. Analysis of table 2 does not strongly indicate that A. salvadorensis is a hybrid between A. burretianaand A. humilis; however, Glassman 13016 may Genus Atialea 65 Table 2. Comparison of A. salvadorensis with Its Possible Hybrid Parents, A. burretiana and A. humilis Plant size Petiole length Rachis length Pinnae length Pinnae width Staminate rachillae length Staminate flower length Stamen number Petal number Fruit size Seed number Microscopic cross section of pinnae Geographic distribution A. burretiana trees 10-30 m tall sessile or 1-6 cm 5.5-7.8 m 95-140 cm 5-7 cm 22-26 cm 18-25 mm 6-9 3 8-11 x 4.5-6.0 cm 1-3 expansion cell tissue (ECT) not divided, adaxial nonvascular fibers (NVF) mostly in short oval clusters, abaxial NVF com- mon in separate clusters and at- tached to some veins eastern Bahia, in Atlantic coastal forest and rem- nants A. salvadorensis up to 30 m tall 80-95 cm 6.0-7.5 m 106-18 cm 3.5-4.2 cm 10-11 cm 13-15 mm 6-9 3-4 9x5cm 1, off-center ECT divided, adaxial NVF in small rounded clus- ters, abaxial NVF common in separate clusters and attached to most veins eastern Bahia, only known for one local- ity in Atlantic coastal forest remnant A. humilis acaulescent, trunk up to 1 m tall 50-80 cm 5.5-8.5 m 65-95 cm 3.5-5.5 cm 6-11 cm 16-21 mm 6 3 6-9 cm x 4-8 cm 1-3 ECT divided, adaxial NVF in small rounded clusters, abaxial NVF common in separate clusters and attached to most veins eastern Bahia, Espirito Santo, and Rio de Janeiro, in At- lantic coastal forest and remnants represent a backcross with one of these species, probably A. humilis (even though most of the palms in the stand appear to be A. burretiana) , because it seems to share more characteristics with this taxon than with A. burre- ttana. Certainly, more collections are necessary before a final determina- tion of the status of this taxon can be made. 21. Attalea brasiliensis Glassman sp. nov. TYPE: Brazil, Goias, 23.5 km north of central Brasilia, Fer Cal area, remnant of climax decidu- ous upland forest, Aug. 1976, Glassman and Eiten 13057 (holotype, F!; isotype, SP!). Figs. 33-34, 70, 79. 66 A Taxonomic Treatment of the Palm Subtribe Attaleinae (Tribe Cocoeae) Caudex 4-6 m altus, folium regulariter pinnatum, pinnis mediis 76-82 cm longis, 3.5 cm latis; bractea mascula 69 cm longa, inflorescentia mascula multiramosa, pedunculo 53 cm longo, rachide 43 cm longa, rachillis 8-11 cm longis, flores masculi 14—16 mm longi, stamina 10-11, antherae 6—7 mm longae, bractea androgyna 124 cm longa 15 cm lata, inflorescentia androgyna multiramosa, pedunculo ca 25 cm longo, rachidi 74 cm longi, rachillis feminis ca 2.5 cm longis, rachillis masculis 7-10 cm longis, fructus 6 cm longus, 4.8—5.2 cm diametro, epicarpio 1 mm crasso, mesocarpio 3 mm crasso, endocarpio osseo 10-13 mm crasso, fibris con- spicuous, semina 2, 2.2 cm longa 1.2—1.5 cm lata. Trees 4—6 m tall; sheathing leaf base not measured; petiole (incom- plete) 13 cm wide near base, margins with fibers 8—15 cm long, leaf ra- chis not measured; pinnae regularly arranged along rachis, those from middle series 76-82 cm long and 3.5 cm wide with acuminate, asymmetri- cal tips; staminate sterile bract deeply sulcate, expanded part 69 cm long (including beak of 16 cm long); rachis of staminate inflorescence 43 cm long, peduncle 53 cm long, rachillae numerous, 8—11 cm long; staminate flowers arranged in two rows along one side of the rachilla, each 14-16 mm long, petals flat, 4-5 mm wide, with acuminate tips and smooth margins, obscurely nerved, sepals 1.5 mm long, stamens 10-11 in num- ber, anthers 6—7 mm long, filaments about 1.5 mm long; expanded part of androgynous sterile bract deeply sulcate, grooves unequally spaced, about 78 cm long (including beak of 15 cm long) and 15 cm wide, pe- duncular part about 46 cm long; rachis of androgynous inflorescence about 74 cm long, peduncle about 25 cm long, rachillae numerous, pis- tillate portion of each rachilla about 2.5 cm long, 1 pistillate flower scar per rachilla, staminate extension of each rachilla 7-10 cm long; pistillate flowers not seen; fruit 6 cm long and 4.8—5.2 cm in diam, persistent pe- rianth 3 cm high, staminodial ring 0.7 cm high and 3 cm across, epicarp about 1 mm thick, mesocarp about 3 mm thick, endocarp 10-13 mm thick, dotted with prominent fiber clusters, seed cavities 2, 2.2 cm long and 1.2—1.5 cm in diam. Distribution and Habitat. See table 1. Representative Specimens Examined. BRAZIL. Goias: 23.5 km north of central Brasilia, Fer Cal area, remnant original low trees in climax de- ciduous upland forest area, in limestone derived soil, Aug. 1976, Glassman and Eiten 13058 (F, SP), 13059 (F, SP), 13060 (F, SP). The new species seems to be closely allied to A. apoda from Minas Gerais because both are arborescent, have regularly arranged pinnae, have rela- Genus Attalea 67 tively short staminate and androgynous rachillae, and have relatively small fruits. A. brasiliensis differs in having staminate flowers with obscure rather than strongly nerved petals, a thicker mesocarp, and 2-seeded rather than l1-seeded fruits with prominent instead of inconspicuous clusters of fibers. Doubtful and Uncertain Species of Attalea Attalea agrestis Barb. Rodr., Enum. Palm. 42. 1875; Sert. Palm. Bras. 1:t. 55. 1903b. LECTOTYPE: (Glassman, 1972a) Brazil, Rio Uauincha (Barb. Rodr., t. 55, 1903b). Orbignya agrestis (Barb. Rodr.) Burret, 1929a. Acaulescent; petiole 20-25 cm long, leaf rachis 1 m long, tomentose; middle series pinnae regularly arranged, 30 cm long and 2.5 cm wide; inflorescence 30-35 cm long; fruit 4 cm long and 2.8 cm in diam, with 3 seeds. Distribution. Amazon region of Brazil. Vernacular Name. Curua-y (Barb. Rodr.). Representative Specimens. No authentic specimens seen. Burret (1929a) cited Huebner 4b (B), but this specimen is not diagnostic (Glass- man, 1977b). In Barbosa Rodrigues’s original article (1875), Barbosa Rodrigues 324 was cited, but this specimen could not be found. Therefore, the above illus- tration was chosen as lectotype. The genus to which this taxon belongs is uncertain because staminate flowers were not described or illustrated in t. 55. Burret (1929a) transferred this species to Orbignya because of its resemblance to O. sabulosa Barb. Rodr. Henderson, Galeano, and Bernal (1995) treat this taxon as a synonym of A. microcarpa Martius, which I treat as a doubtful species. Attalea blepharopus Martius, Palmet. Orbign. 116, t. 5, fig. 2, t. 31C. 1844; Hist. Natur. Palm. 3:t. 167. 1845. TYPE: Bolivia, d’Orbigny 34 (ho- lotype, P, destroyed?). Scheelea blepharopus (Martius) Burret, 1929a. Burret (1929a) transferred this species to Scheelea without explanation, but probably did so because staminate flowers were described as fleshy by Martius (1844). Staminate flowers illustrated by Martius (t. 167), however, appear to have flattened petals. Since the description of the staminate flowers is questionable and no other specimens (besides the type, which 68 A Taxonomic Treatment of the Palm Subtribe Attaleinae (Tribe Cocoeae) was probably destroyed) have been collected, this taxon should be con- sidered an uncertain species. No other species of Attalea have been reported from Bolivia; however, S. princeps is the only species of Scheelea collected from that country. At any rate, I cannot equate S. blepharopus with S. princeps because of a lack of available specimens for examination and a questionable description. Henderson (1995) and Henderson, Galeano, and Bernal (1995) treat this taxon as synonymous with A. phalerata Martius. Attalea excelsa Martius ex Sprengel, Syst. Veg. 2:624. 1825; Martius, t. 96, fig. III, 1-2. 1826. t. 169, fig. 3. 1845. TYPE: Brazil, Maranhao and Para (no specimens cited). LECTOTYPE: (Henderson, 1995) Martius, Hist. Natur. Palm. 2:t. 96, figs. 1-3. 1826. Scheelea martiana Burret, Notizbl. Bot. Gart. Berlin-Dahlem 10:661. 1929a. Burret (1929a) transferred this species to Scheelea, but was obliged to give ita new name because S. excelsa Karsten (1857), based on a different type, was already published. Staminate flowers were not mentioned in Martius’s original description; however, Burret justified his transfer to Scheelea on the presence of fiber clusters in the endocarp of the fruit. This is a question- able distinction because a number of species of Attalea also have clusters of fibers in the endocarp. On the other hand, there is nothing in the descrip- tion to place it in Aftalea either. Therefore, this binomial is designated as a species dubium. Henderson (1995) and Henderson, Galeano, and Bernal (1995) treat this taxon as a synonym of A. phalerata Martius. Attalea goeldiana Huber, Bull. Herb. Boiss. ser. 2, 6:268. 1906. TYPE: Bra- zil, Rio Acre (no specimens cited). Type not designated. Scheelea goeldiana (Huber) Burret, 1929a. I am considering this species as doubtful because Huber’s description is inadequate and lacks illustrations. Burret probably transferred it to Scheelea because Huber placed it in section Pseudoscheelea Drude of the genus Attalea. Henderson (1995) and Henderson, Galeano, and Bernal (1995) treat this binomial as a synonym of A. insignis (Martius) Drude. Attalea gomphococca Martius, Hist. Natur. Palm. 3:301, t. 167, fig. 6. 1845. LECTOTYPE: (Glassman, 1972a) Central America (t. 167, fig. 6). Scheelea gomphococca (Martius) Burret, p. 666. 1929a. Since Martius’s original description and illustration is based mainly on the fruit and since the exact locality is in doubt, it is difficult to delimit Genus Attalea 69 this taxon with any degree of confidence. Apparently, Burret had no ba- sis for transferring it to Scheelea because the staminate flowers were not described. Henderson, Galeano, and Bernal (1995) treat this binomial as a synonym of A. butyracea (Mutis ex L.f.) Wessels Boer. Attalea hoehnei Burret, Notizbl. Bot. Gart. Berlin-Dahlem 10:522. 1929a. TYPE: Brazil, Mato Grosso or Acre, Agua Limpa, Campo, Hoehne 2196 (holotype, B, destroyed?). The entire description of Burret indicates this taxon is probably based on depauperate specimens. Size of pinnae, staminate inflorescence, rachillae, and staminate flowers are all very small for this genus and hence are difficult to compare to those of other species. Henderson (1995) and Henderson, Galeano, and Bernal (1995) treat this binomial as a synonym of A. phalerata Martius. Attalea lapidea (Gaertner) Burret, Notizbl. Bot. Gart. Berlin-Dahlem 10:533. 1929a; Henderson, Galeano, and Bernal, 1995. Cocos lapidea Gaertner, Fruct. Sem. Pl. 1:16, t. 6, fig. 1. 1788. LEC- TOTYPE: (Glassman, 1972a) Brazil, Bahia (Gaertner, t. 6, fig. HS L7S8)e Burret placed this palm close to A. funifera based on the similarity of the fruits; however, it should be considered a doubtful species because there is virtually no information on the leaves, inflorescences, flowers, or size of the plant. Attalea microcarpa Martius, Palm. Orbign. 125, 1844. t. 168, fig. 2. 1845. t. Z16, fig. 5. 1849. TYPE: Brazil, Para, Martius s.n. (holotype, M, de- stroyed?). LECTOTYPE (Henderson, 1995) Martius, Hist. Natur. Palm. 3:t. 168, fig. 2. 1845. Orbignya microcarpa (Martius) Burret, 1929a. Size of plant, leaves, rachis, pinnae, and androgynous and staminate inflorescences and flowers not measured; fruit about 3.7 cm long and 2.5 cm in diam. The description and illustrations are insufficient for determination (neither the leaves nor the staminate flowers are mentioned), but Burret thought it may belong to Orbignya. No specimens were cited by Martius; however, according to Burret, the infructescence illustrated by Martius (t. 168) was preserved in the Munich collections. Nevertheless, this name should be treated as a species dubium because even the genus to which it 70 A Taxonomic Treatment of the Palm Subtribe Attaleinae (Tribe Cocoeae) belongs is uncertain. On the other hand, Henderson (1995) and Henderson, Galeano, and Bernal (1995) treat A. microcarpaasa good spe- cies. They also place O. sagotiiand O. polystichain synonymy, but I consider both as bona fide species. Aitalea monosperma Barb. Rodr., Enum. Palm. Nov. 42, 1875. t. 57A. 1903b. LECTOTYPE: (Glassman, 1972a) Brazil, Para (t. 57A, 1903b). A. spectabilis var. monosperma (Barb. Rodr.) Drude, Martius Flora Bras. 3:440. 1881; Burret, 1929a. Burret (1929a) said this palm may belong to Orbignya, but it is differ- ent from O. spectabilis. Staminate flowers were neither described nor illus- trated by Barbosa Rodrigues or Drude, hence determination of the genus is doubtful. No specimens were cited by Barbosa Rodrigues, but Drude listed Sagot 601 and 831 from French Guiana. The latter specimen is part of the type collection of O. sagoti. Henderson (1995) and Henderson, Galeano, and Bernal (1995) treat this taxon as a synonym of A. spectabilis. Attalea pixuna Barb. Rodr., Enum. Palm. Nov. 43. 1875. LECTOTYPE (Glassman, 1972a). Brazil: Para, calcareous soil of Igarape Bom Jardim, Villa de Itaituba, Rio Tapajos (t. 49, 1903b). Orbignya pixuna (Barb. Rodr.) Barb. Rodr., Prot. App. 49. 1879. t. 49. 1903b; Glassman, 1977b. See Orbignya pixuna for discussion. Attalea pycnocarpa Wessels Boer, Pittieria 17:299. 1988. TYPE: Venezuela, Terr. Amazonas, Puerto Ayacucho, mesophytic forest on well- drained sandy clay, July 28, 1967, Wessels Boer 1910 (holotype, U!). Since staminate flowers were not described, it is difficult to determine the genus of this taxon. Wessels Boer surmised that it may belong to Orbignya, but there is insufficient evidence to draw this conclusion. Henderson (1995) and Henderson, Galeano, and Bernal (1995) treat this species as a synonym of A. butyracea. Attalea racemosa Spruce, Journ. Linn. Soc. 11:166. 1871; Wessels Boer, 1972, 1988. TYPE: Venezuela, Amazonas, Rio Negro, Spruce 54 (holotype, K!; isotype, P!). Orbignya racemosa (Spruce) Drude, 1881. This taxon will be discussed under O. racemosa. Genus Attalea 7) Attalea rhynchocarpa Burret, Notizbl. Bot. Gart. Berlin-Dahlem 12:617. 1935; Dugand, 1940, 1954; Henderson, Galeano, and Bernal, 1995. TYPE: Colombia, Rio Frio, bei Salonique, Dryander s.n. (holotype, B, destroyed?). The description for this taxon is incomplete. Neither staminate or pis- tillate inflorescences or flowers nor length and width of pinnae are de- scribed. Attalea spectabilis Martius, Hist. Natur. Palm 2:136, t. 96, figs. 1-2. 1826; Wessels Boer, 1965, 1972, 1988. Orbignya spectabilis (Martius) Burret, 1929a. This taxon will be discussed under O. spectabilis. Attalea wallisii Huber, Bull. Herb. Boiss., ser. 2, 6:267. 1906. TYPE: Brazil, Amazonas, Rio Purus (no specimens cited). LECTOTYPE (Glass- man, 1977a) Brazil: Rio Purus or Rio Acre (Huebner 163-B!). Scheelea wallisi (Huber) Burret. The original description by Huber is mainly a comparison of charac- ters with A. humboldtiana, which it most closely resembles. Burret probably transferred the species to Scheelea because of its close resemblance to A. humboldtiana. In 1934 he gave S. wallisiia rather lengthy description, except for the flowers. Since neither staminate nor pistillate flowers are known, this species is doubtful. Henderson (1995) and Henderson, Galeano, and Bernal (1995) treat this taxon as a synonym of A. butyracea. y ey: r sar i - ea od) See f Sot Tiel Braet 2) - 4 . ‘ ae mo ’ oI sS ‘ > ; By Kee ont ich sheila Ae tras soil at PUR Bidig? (peer aye EE ebee | funldB0r Bhd ape , 40 yonder? £ Ah de ps ‘3 , 5 rt ee | Sheer “IN, Sadidluane ergh sss ohn el IN. pga ab 416; Pettey ceed Won lara Pagereh syn er Dant r n 7 heater ex’) 6 tet eel 4 Gas S—I Ant! ae le Poe ieee pei Js teva Ln ia Fell Bet ai 0M | CEE te ne ldabliga cgi ; a Ta) CS ah Hata ” hae MALOSTL! ) Pi ven ek be bg} on Shee ert Ly aie iT he ed nel, thin fH fu sith. te hed TEER pet sermons. ores: 36). 14 ik VD VIR) eile Ay NED OR. yi " age o (yale 7 shew ae iy es reali ae Wy we ys: ta 5 7 f i "6 eee Airs uliirl (web lee Sitka ae oe CP INRA Bao tego stat hating 7 ‘ate /] ‘7, Jered pare, a0 Wivac: THA). eraraba tag ied x . Apaeztn! TVA aps beech, aun ek vy NT te aT | 90h “) eve eye a C1 NOCTENG “OTN te 8) Oh ye Sah ah KEOs ae a nee TONEY Ag Mah AS WM; § eee | PERE MT wei ma 9 . AOE 6 45-14 Se ie RPOAED WES > heer toares ey sm ‘ by » "Vy ’ > ~ ; ik ly Ghee) ™ 7 (wy ph ide ‘ eal } Mm a i; LP Sapa. Wi ater o ‘ , ' é Pree Med GT RR te , i tt Te my th i rae) 01 uy) ariel pier: al ANC : ar nm = 7 a A ‘ BS We Laie heen f Arie hs iad’, NO } ued We bom WAR We age . Healing, Frag atven py Rss parapet,’ ae ee i 1) Barat ye ; ; (hi Te. Tye tetamey aI) ie hie aud OF, ena J Ps : WEAN, 7 i wn, ee i Aol. ; Genus Orbignya Geographic Distribution of Genus Orbignya The genus Orbignya (11 species) is essentially South and Central Ameri- can in distribution (see table 3). Its center of distribution appears to be wet forested areas of northern Brazil. A total of 7 of the 9 South Ameri- can species as well as 3 hybrids, O. x teixeirana, x Maximbignya dahlgreniana, and x Attabignya minarum, are found in Brazil. Other South American taxa occur in Suriname (3, including x Maximbignya dahlgreniana); Colombia (3, including Ynesa colenda); French Guiana, Guyana, and Venezuela (2 each); and Bolivia, Peru, and Ecuador (1 each). In Brazil 4 taxa each are found in Amazonas and Maranhao; 3 each in Goias, Bahia, and Piaui; 2 each in Para and Minas Gerais; and one each in Ceara and Mato Grosso. In Central America and the Caribbean | species is found in each of the following: Mexico, Belize, El Salvador, Guatemala, Honduras, Nicaragua, and Haiti. In contrast to genera like Syagrus and Attalea, which have a number of taxa growing in xerophytic regions, and much like the genus Scheelea, most species of Orbignya occupy mesophytic or wet forested areas. Some per- sist in drier disturbed habitats after the original vegetation has been cut. In spite of deforestation, several species of Orbignya are still fairly com- mon or abundant today: O. phalerata is common or abundant in several states of Brazil in mesophytic or wet forests; x Maximbignya dahlgreniana forms large uniform populations associated with O. phaleratain Suriname; O. cohuneis common in several provinces along the Atlantic coast in Gua- temala; O. guacuyule is apparently still fairly common in forests in several states of Mexico; O. polysticha appears to be common in forests in the Amazon regions of Peru, Venezuela, and Brazil; and judging from recent collections, O. sagotii seems to be holding its own in wet forests of Suri- name. On the other hand, certain other species may not be as common Table 3. Geographic Distribution of Orbignya Species O. brejinhoensis O. cohune O. crassispatha O. cuatrecasana O. eichlen O. guacuyule O. luetzelburgit O. oleifera O. phalerata O. polysticha O. sagotit Hybrids: O. X teixeirana x Maximbignya dahlgreniana x Altabignya minarum Country and State Brazil: Bahia, east of Rio Sao Francisco Belize; El] Salvador; Honduras: Atlantida; Guatemala: Izabel, Alta Verapaz, Petén; Nicaragua: Zelaya Haiti: western part of southern peninsula and eastern part of Massif de la Hotte Colombia: Valle Brazil: Goias, Bahia, Maranhao, Piaui Mexico: Oaxaca, Guerrero, Michoacan, Colima, Jalisco, Nayarit Brazil and Venezuela: Amazonas; Colombia: Vaupes Brazil: western Minas Gerais, Munic. de Pirapora, Santa Fé; Bahia: west of Rio Sao Francisco, Munic. de Barra, Bianapolis, Cocos, Sao Desiderio Bolivia: departments of Santa Cruz and Beni; Brazil: Amazonas, Para, Maranhao, Ceara, Piaui, Mato Grosso, Goias; Guyana and Suriname Amazon regions of Peru, Venezuela, Brazil; French Guiana French Guiana; Suriname; Guyana; Brazil: Amazonas Brazil: Maranhao (Cod6, Caxias), Goias (Tocatinopolis), Piaui (Terezina) Brazil: Para, Maranhao; Suriname Brazil: Minas Gerais Habitat Mesophytic forests in stream valleys at 400-900 m alt. Climax forests, Atlantic lowland and mountain forests, in well- drained soils Scattered stands, rocky areas Rain forest, along streams Forests and secondary growth areas Forests and deciduous forests Rain forests, sandy savannas, sandy soil between forest and open savanna Forests, along river margins, adjacent to cerrados or caatingas Wet forests, sandy soils, subhydrophytic forest Forests, rain forests on white sandy soil Wet, well-drained soil in undergrowth of savanna-like forest on sandy soil often among granite in dense shade Cerrados (soils of low water and low fertility during dry season) Matas, mesophytic forests, in clay or sandy loam soils Forests, along streams Genus Orbignya 75 Table 3. Cont. Species Country and State Habitat Ynesa colenda Ecuador: Los Rios, Manabi, Dense wet forests on northern Esmeraldas, Guayas; coastal plain; more abundant in Colombia: Narino drier forests of western and southern coastal plain; much destruction of vegetation, but persists in pastures on well- drained and hilly ground as O. phalerata; O. brejinhoensis, which was known only from several stands in Bahia, in the municipality of Oliveira dos Brejinhos, recently has been reported from a number of other localities in Bahia; O. eschlen is known from a number of collections in Goias, Piaui, and Maranhao, and more recently from Bahia; O. olezferais known from limited areas in Minas Gerais and several localities in Bahia; O. cuatrecasana has a limited distribution in the department of Valle in Colombia; and O. luetzelburgii is apparently local in the Amazon regions of Brazil and Venezuela and in the province of Vaupes in Colombia. According to Markley (1971), the genus Orbignya (especially O. phale- rata) originated in the interior high plateau of Goias. With few exceptions, babassu areas in Brazil are connected with watersheds of the highlands of Goias. These highlands contain Archaezoic rocks that have undergone long periods of erosion, resulting in complex river systems. The shoreline habitats provided a means of migration and dispersal for many genera of palms that probably evolved during the Cretaceous period. Currently, O. phalerata occurs in greatest density and pure stands on deltas in Maranhao, in flood plains of the Paraguay River drainage basin, and in broad river valleys of the interior. In these areas, there is little or no competition from dicotyledonous trees. Relationships within Genus Orbignya In contrast to Attaleaand Scheelea, which are divided into subgenera based on clustering of middle series pinnae, Orbignya seems to fall into four natural groups or subgenera based on morphology of the staminate flower petals. In the first group (comprising four species), petals are flattened and 1-8 in number resulting from fusion of 2-several petals and additional petals resembling anthers; petals are usually of two shapes, lanceolate with acute tips and broad with laciniate or coarsely toothed tips. The second 76 A Taxonomic Treatment of the Palm Subtribe Attaleinae (Tribe Cocoeae) subgenus, with five species, has 3 flattened spatulate petals (very wide near the middle and narrow at the base) with acute or obtuse tips; in the third group petals are curved and lanceolate; and the last group has fleshy pet- als with hooked, acute tips. The first subgenus can be split into two dis- tinct units or sections based on the size of plants, the number of stamens per flower, and clustering of middle series pinnae (see the “Key to Spe- cies and Hybrids of Orbignya”). O. eichleri is the only member of the first grouping. It is distinct and does not seem to be closely related to species of the second unit; however, it does hybridize and backcross with O. phalerata to produce O. x teixeirana. The second section, characterized by arborescent plants, regularly arranged pinnae, and flowers with 24-57 stamens, comprises three species that seem to form a close alliance. O. phalerata has a much broader geographic range than the other two spe- cies, O. oleifera and O. brejinhoensis. The second subgeneric group is divided into three acaulescent and two arborescent species. The latter two palms from Central America appear to be closely allied, differing mainly in the size of their staminate rachillae and staminate flowers, size of pistillate rachillae, and number of stamens. O. guacuyule may have been derived from O. cohune, which has a more extensive range. Apparently, the acaulescent taxa do not show very close relationships. O. cuatrecasana differs from the others in having pinnae with glandular dots, a large number of stamens (20-24), and very large fruits (11-14 cm); and O. polysticha is differentiated from O. sagotii chiefly in having staminate flowers completely encircling the staminate rachilla in- stead of being on one side of the rachilla. In the third subgeneric group, O. crassispatha can be easily separated from the other subgenera by its curved, flattened petals and clustered instead of regularly arranged middle series pinnae. It seems to be with- out close relatives in Orbignya. The fourth subgenus is represented by one species, O. luetzelburgii, whose staminate flowers are so distinct that it was transferred to a separate ge- nus (Parascheelea) by Dugand (1941). In contrast to Syagrus, the genus Orbignya seems to have a higher per- centage of closely related species. In Orbignya, only about 5 of 11 taxa are isolated, whereas in Syagrus more than 50 percent of the species appear to be without very close relatives, and most of these represent sections containing a single species. Scheelea seems to be more of a homogeneous genus than Orbignya because a higher percentage of its species are closely Genus Orbignya 77 allied, mainly because its staminate flowers are very much alike, i.e., all have fleshy petals mostly of the same shape (more or less terete) and con- sistently with 6 stamens. On the other hand, in Orbignya the petals are of four distinct shapes and have stamens ranging from 6 to 57 in number. In Attalea, petals are fairly similar in shape (consistently lanceolate), but like Orbignya its species have evolved high stamen numbers (up to 64). It is not certain when Orbignya first appeared in the geological record, but as previously mentioned, its cocoid ancestors were present in the Eocene period. Three of the four genera in the subtribe Attaleinae (Orbignya, Scheelea, and Attalea) have a similar distribution, being found in both South and Central America. The ranges of both Orbignya and Scheelea extend as far north as Mexico, but Attalea reaches only Panama in Central America. Maximiliana, the fourth genus, with only one species, is distributed in northern South America. Outline of Tentative Division of Orbignya into Infrageneric Categories Subgenus I. Staminate flowers with 1-8 petals, usually of two distinct shapes, one as a result of fusion of 2 or more petals with coarsely toothed tips, the other lanceolate with acute tips. Staminate flowers arranged on one side of the rachillae. Section 1. Acaulescent, middle series pinnae in clusters of 2—4. O. ezchleri. Section 2. Arborescent, middle series pinnae regularly arranged. O. brejinhoensis, O. oleifera, O. phalerata. Subgenus II. Staminate flowers with 3 petals, usually separate, spatulate, and flattened. Staminate flowers mostly spirally arranged around rachillae. Section 3. Acaulescent. Subsection A. Staminate flowers with 20-24 stamens. O. cuatrecasana. Subsection B. Staminate flowers with 11-15 stamens. O. polysticha. Subsection C. Staminate flowers with 8—12 stamens, arranged on one side of rachillae. O. sagotzz. Section 4. Arborescent. O. cohune, O. guacuyule. Subgenus III. Staminate flowers with 3 curved, flattened lanceolate pet- als. Middle series pinnae in clusters of 2—4. Arborescent. O. crassispatha. Subgenus IV. Staminate flowers with 3 narrow, fleshy, curved petals with hooked tips, and connate at base. Middle series pinnae regularly ar- ranged. Acaulescent. O. luetzelburgit. 78 A Taxonomic Treatment of the Palm Subtribe Attaleinae (Tribe Cocoeae) Taxonomic Treatment of Genus Orbignya In the present paper, I am recognizing only 11 species of Orbignya out of 30 that have been included in this genus. This number includes 1 newly described species, O. brejinhoensis, and 1 new combination, O. crassispatha. The remaining 19 names have been designated species incerta or species dubia (9) or nomen nudum (1), or reduced to synonymy (9). In addition to 11 species of Orbignya, Iam recognizing | interspecific hybrid, O. x teixeirana, 2 intergeneric hybrids, x Maximbignya dahlgreniana (described as new) and x Attabignya minarum; and 1 putative hybrid, Ynesa colenda. In the taxonomic treatment of Orbignya the same format as that used for Attalea is followed. ORBIGNYA Martius ex Endlicher, Gen. Plant. 257. 1837a (conserved name); Burret, Notizbl. Bot. Gart. Berlin-Dahlem 10:493-543. 1929a; Glassman, Phytologia 36:89-115. 1977b; not Orbignya Bertero, Mercurio Chil. 16:737. 1829 (Euphorbiaceae). TYPE: Orbignya phalerata Martius. Section Distichanthus Burret, Notizbl. Bot. Gart. Berlin-Dahlem 10:498. 1929a. TYPE: none listed. Section Pleiostichanthus Burret, Notizbl. Bot. Gart. Berlin-Dahlem 10:499. 1929a. TYPE: none listed. Section Spirostachys Burret, Notizbl. Bot. Gart. Berlin-Dahlem 10:499. 1929a. TYPE: none listed. Parascheelea Dugand, Caldasia 1:10. 1940. TYPE: P. anchistropetala Dugand. = Orbignya luetzelburgii Burret. Trees (in 6 species) 6-38 m tall (up to 70 m in one species, fide Ryder, 1978) and 30-50 cm in diam; acaulescent in 5 species. Sheathing leaf base 0.5-2.0 m long; petiole absent or 0.15—2.5 m long, whitish tomentose and greenish punctate in at least 1 species, petiole margins with fibers 10-32 or more cm long and 3 mm thick; leaf rachis 2-18 m long, frequently covered with a ferrugineous indument; 50-290 pinnae on each side, those from middle series regularly arranged (in 9 species) or in clusters of 2—4 (in 2 species and | hybrid), with intervals of from 2 cm between pinnae or clusters in several species to 5—7 cm apart in several others, pinnae from middle series 60-150 cm long and 2.5-9.0 cm wide, glaucous abaxially in some species, rarely with scattered, tiny brownish glandular dots adaxially, margins near tips frequently with ferrugineous indument, pinnae tips acute or acuminate and asymmetrical; expanded part of staminate ster- Genus Orbignya 79 ile bract from 35 cm long and 4—5 cm wide to 2.5 m long and 60 cm wide (including beak of 10-40 cm long), deeply sulcate, frequently covered with a brownish or ferrugineous indument, peduncular part of bract (fre- quently included in size of whole bract) 0.35—1.0 m long; rachis of stami- nate inflorescence 15-60 cm long, peduncle 0.2—3.0 m long, with numer- ous rachillae (30-600), usually covered with a whitish indument or sometimes with a brownish farinose indument, individual rachillae 7—30 cm long, staminate flowers in pairs and completely encircling rachilla or single or in pairs on one side of rachilla, each 7-15 mm long, petals of four types: (1) consistently 3, usually of one shape (spatulate and flattened), about 1 mm wide at base and 3-8 mm wide near middle; (2) petals 1-8 in number, resulting from complete fusion of 2-several petals, and usually with one or more additional separate petals, usually of two shapes, one irregular and mostly broad throughout, with notched and coarsely toothed tips, the other lanceolate, with acute tips, rarely with only lanceolate petals; (3) 3 fleshy petals with hooked, acute tips; and (4) 3 flat, curved lanceolate petals; sepals 3 in number, usually 1-2 (5) mm long, stamens 6—57 in number, anthers 1-2 mm long, spirally twisted and coiled, filaments 1.5—3.5 mm long; androgynous sterile bract 0.35—1.9 m long, 5.5-60.0 cm wide and 2-3 cm thick (including beak of up to 32 cm long), peduncular part 23-50 cm long, deeply and irregularly sulcate, frequently brownish pubescent; rachis of androgynous inflorescence 0.2—1.0 m long, peduncle 0.15-1.5 m long, frequently with brownish indument, androgy- nous rachillae numerous (up to 300 in some species) , comprising a basal pistillate portion with each pistillate flower accompanied by two staminate flowers that fall off early and a narrower terminal portion with staminate flowers only (staminate rachilla extension); basal portion from almost sessile to 10-15 cm long, frequently with scattered whitish lepidote indument, pistillate flowers 1-11 per rachilla, each 2.0—4.5 cm long and 1.5-4.0 cm in diam, sepals 3 and petals 3, either equal in size or sepals somewhat shorter than petals and more or less nerved, pistil 2—4 cm long, ovary glabrous or brownish lepidote, style very short or sessile, stigmas 3— 8 in number, each 0.5-1.5 cm long, staminodial ring 0.5-1.5 cm high; transitional pistillate flowers about 15 mm long, sepals 3, broad and im- bricate, about 9 mm long, petals 3, narrow, lanceolate, black in color, about 15 mm iong, stigmas black, pistil about 8 mm long, staminate flowers from pistillate rachilla frequently sterile and transitional, 5-8 mm long and 2 mm wide, petals black, flat, somewhat fleshy and curved, stamens mostly 6 in number, anthers 1.0—1.5 mm long, coiled and twisted, pistillode about 80 A Taxonomic Treatment of the Palm Subtribe Attaleinae (Tribe Cocoeae) 4 mm long; staminate extension of androgynous rachilla usually un- branched and frequently broken off, 11-21 (28) cm long, sometimes with 8-14 branches; fruit usually orange when ripe, 3.5—-14.0 cm long and 2.0- 9.5 cm in diam, persistent perianth 3-6 cm high, staminodial ring 1.0- 1.5 cm high, epicarp fibrous, 1.5—3.5 cm thick, mesocarp soft, 1-6 mm thick, endocarp hard, 0.5—2.5 cm thick, fiber clusters usually inconspicu- ous, but occasionally with conspicuous fibers, seeds 1-7 in number, each 2.5-5.2 cm long and 1.0—1.5 cm in diam. Key to Species and Hybrids of Orbignya 1. Staminate flowers with 1-8 petals, which are usually of two distinct shapes: one resulting from complete fusion of 2 or more petals and mostly broad throughout, with notched or coarsely toothed tips, the other lanceolate with acute tips; staminate flowers arranged on one side of rachilla, pistillate flowers with 3-8 stigmas. 2. Acaulescent or up to 8 m tall (in O. x teixeirana), staminate flowers with 12—24 stamens, middle series pinnae in clusters of 2-5. 3. Middle series pinnae 45—95 cm long and 2.5—3.6 cm wide, stami- nate sterile bract 35-65 cm long and 4—5 cm wide, staminate ra- chis 18-35 cm long, rachillae of staminate inflorescence up to 15 CMON EINES OL BE PES 2 1. O. ezchlen (Brazil: Goias). 3. Middle series pinnae up to 110 cm long and up to 4 cm wide, staminate sterile bract 85-130 cm long and 10-13 cm wide, stami- nate rachis 50-100 cm long, rachillae of staminate inflorescence up to 20cm Tong 02s Se ee O. x teixeirana (Brazil: Goias). 2. Trees 6-38 m tall, staminate flowers with up to 57 stamens, middle series pinnae regularly arranged. 4. Stamens 8-9 or 11-20 in number, anthers 5-10 mm long, usu- ally twisted, sometimes partially coiled or straight. 5. Stamens 8-9, anthers 5-6 mm long, usually twisted, sometimes straight, not‘coiled, fruit l-seeded |... .5 2. Ae OO walt ABSA OR Ynesa colenda (Ecuador and Colombia). 5. Stamens 11-20, anthers 8-10 mm long, usually twisted, some- times partially coiled, fruit. 2—4 seeded». 0.0.5 h204 rN A See x Attabignya minarum (Brazil: Minas Gerais). 4. Stamens up to 57 in number, anthers 1-2 mm long, distinctly coiled. 6. Petiole 80-100 cm long, pistillate portion of androgynous Genus Orbignya 81 rachilla up to 1 cm long, stigmas mostly 6-8 in number, fruits 8-10 emrlong 2.4 J) fas 2. O. brejinhoensis (Brazil: Bahia). 6. Petiole absent or up to 50 cm long, pistillate portion of an- drogynous rachilla 2-15 cm long, stigmas 3—6 in number, fruits 10-14 cm long. 7. Petiole absent or up to 10 cm long, pistillate portion of an- drosynous rachillaP—Jem ION Gis pists airs. Hele Bo is faye huis: oy 3. O. oleifera (Brazil: Minas Gerais, Bahia). 7. Petiole 15-50 cm long, pistillate portion of androgynous RACHA TONS CmAONS: ps2;? stasis pey-nrsies ies ays ee # as o: Sy ars Si 2) Rae 4. O. phalerata (Brazil, Bolivia, Guianas). 1. Staminate flowers consistently with 3 petals, usually separate, spatulate, and flattened (about 1 mm wide at base and 3-8 mm wide near middle, usually with acute or obtuse tips); or sometimes lanceolate, flat, and curved with acuminate tips; or narrow, fleshy, connate for about one- third of corolla base and with curved, hooked, acute tips; staminate flowers mostly completely encircling the rachilla (except O. sagotii), pistillate flowers mostly with 3—4 stigmas. 8. Staminate flowers usually with spatulate or curved, flattened lan- ceolate petals. 9: 4) Middle series pinnae in clusters of 2—4, staminate flowers with eluved petals. ...0i) aja

bP bb bb val macrolepis (Burret) Wessels Boer. = Scheelea macrolepis Burret. macropetala (Burret) Wessels Boer. = Maximiliana maripa (Correa) Drude. . maracaibensis Martius. = Scheelea maracaibensis (Martius) Burret. . maripa (Correa) Martius. = Maximiliana maripa (Correa) Drude. . osmantha (Barb. Rodr.) Wessels Boer. = Scheelea osmantha Barb. Rodr. parviflora Barb. Rodr. = Scheelea anisitsiana Barb. Rodr. phalerata Martius ex Sprengel. = Scheelea phalerata (Martius ex Sprengel) Burret. . pixuna Barb. Rodr. = Orbignya (species dubia). . polysticha (Burret) Wessels Boer. = Orbignya polysticha Burret. princeps Martius. = Scheelea princeps (Martius) Karsten. regia (Martius) Wessels Boer. = Maximiliana maripa (Correa) Drude. rostrata Oersted. = Scheelea rostrata (Oersted) Burret. sagoti (Trail ex Im Thurn) Wessels Boer. = Orbignya sagotii Trail ex Im Thurn. speciosa Martius. = Orbignya phalerata Barb. Rodr. . Spectabilis var. polyandra Drude. See Orbignya pixuna (Barb. Rodr.) Barb. Rodr. under “Doubtful and Uncertain Species of Orbignya.” transitiva Barb. Rodr., Prot. App. 49, 1879; Les Palmiers 29, 1882. Superfluous name. This is a superfluous name for A. attaleoides (which has staminate flowers like Scheelea and has been previously discussed). After describing M. attaleoides as a new species in 1875, Barbosa Rodrigues (1879, 1882) de- cided that it was a transitional species between this genus and Attalea. Therefore, he renamed it A. transitiva. Excluded Species of Maximiliana M. attaleoides Barb. Rodr. = Scheelea sp. (species confusum). M. crassispatha Martius. = Orbignya crassispatha (Martius) Glassman. M. inajai Spruce, Journ. Linn. Soc. 11:163. 1871. = Syagrus inajai (Spruce) Beccari, Agric. Colon. 10:467. 1916. TYPE: Brazil, Amazonas, Rio Negro, Spruce 83 (holotype, K!). M. insignis Martius. = Scheelea insignis (Martius) Karsten. M. tetrasticha Drude. = Scheelea (species dubia) . M. venatorum (Poeppig ex Martius) H. Wendland in Kerchove, Palm. 251. 1878. Cocos venatorum Poeppig ex Martius, Hist. Natur. Palm. 3:325. 1853. = Oenocarpus or Euterpe? TYPE: Peru, Tocache, Maynas Alto, Poeppig 1998 (holotype, W, destroyed; isotype, P!). ‘ cy ae A tava Frcaln lta ” Fahy Suubovthe 4. wench edb (ats pul Wien the a Sebiiaten ime i ML A. ehderahaticds Fe eet) Vries hs Mee. ‘By Teor Nw | ' Oo eae = A. Hreradnideiree Bist, > Sel eee amaaiiendeune te i Coton) Nanas Sel aang ~~ orem abate le pamdinfat ola eagle era ati oe eae or) same Bit. Bodie wf = rohine sattbiceiatas ley larete: KS _obmenrealynegy, » A, nay a the re ‘ on aeaaiey tial ag) ne avenged? sj ppt ‘ea een Mage Marie cree ania ‘he nid irol, Pu, . UT SRS ans oes si Nal hye 7 ie fret tat te bien i . moh EH. ninegouen) & trices Martius, tighten pean My bitin} carte Rete Weis (eseltvst) ahegtawii athena = | A. Fiore (Mar oe Viewge!- Jeon Peden ates 3 Lm Mae (¢ ah t ; eR ssc waite) } oiled, . oY 4, POS eEBA De Sed, oh SN an Naas vic sete: i4 Meeaeetceet NO teat ay 9 LAL ithe yl atl ‘pt ina TRS hT (eset ee os » + ‘ of Pri ine i y } od : iy . by § op er , etal EEE het an pg eT juried xs Wetpo*t ware aexT Be be ae i. ae AR ty, apts hee <9 iY IOL “hereutesh Wi orpdaled) OO OU) SRE cathode Rohe tit iia tak: wae, Polaron | Heh See Oy yh Pe i Lee anders A Donated tam ty ey eh Bae: (Mitte. Po fi ery ty { Pai, TUGR Wappcr th rm os ; Px. | ae hie BS UEP RTA a (tee c ws ' - 19 tee ‘Wivis “al +h hee tite : al edlinw act Janu 3) e'e yiekeedy alee dave) 7”), oF on | a : As | nw i% Ww 4) ae Ges t re, Bu: ™ ‘ Roving iee9 é 17 * 2) i wars 4 6A wna wre ih bes Peak tS | Kean | mm yy mea “a revi Ki ia “ul r , ork, ? | ne A a)? 2 iv 7 r i er ) ty Z the ' fay! 7 el ii Ae Cra’) ee ae ao (ey e NT ah aNd : pred M 5 * yivr e Tid fh Pr? ' Pie Ae ee ya® “ ¥ - ‘ \ j nl » ' _ i-# ny « Glossary Morphological Terms Used in Text Acaulescent: Without a vertical aboveground trunk. Androgynous inflorescence: Containing both pistillate and staminate flow- Ets. Arborescent: With a vertical aboveground trunk. Bract or sterile bract: Usually broad, leaf-like organ that subtends the in- florescence; woody in the Attaleinae; formerly called a spathe. Ciliate: Bearing a fringe of hairs on an organ (e.g., staminodial ring). Connate: Fused or united parts. Endocarp: Innermost layer of pericarp. Epicarp (also exocarp): Outer layer of pericarp. Farinose: Covered with a mealy coating or indument. Indument: Any covering such as hairs (pubescent) or scales (lepidote). Inflorescence: Structure bearing flowers; formerly called spadix. Lamina: Blade of pinna or leaflet. Lanceolate: Relatively narrow and tapering at both ends. Lepidote: Indument consisting of scales (i.e., Lepidoptera = scale wings). Mesocarp: Middle layer of pericarp. Pedicel: Stalk of an individual flower. Peduncle: Stalk of an inflorescence. Perianth: Collective term for sepals (calyx) and petals (corolla). - Pericarp: Wall of fruit, usually comprising three layers: epicarp, mesocarp, and endocarp. Persistent perianth: Sepals and petals remaining on base of fruit during its development. Petiole: Stalk of whole leaf. Pinnae: Divisions of a pinnate leaf blade, either separate or in clusters. Pistillode: Sterile or undeveloped pistil of a staminate flower. Rachilla(e): Branch(es) that bear the flowers. Rachis: Axis of a leaf blade bearing pinnae or axis of an inflorescence. 210 A Taxonomic Treatment of the Palm Subtribe Altaleinae (Tribe Cocoeae) Rugose: Densely wrinkled surface. Sessile: Lacking a stalk (e.g., lacking a petiole). Sheath or sheathing base: Base of leaf that clasps the stem. Staminode: Sterile or undeveloped stamen of a pistillate flower. Staminodial ring: Structure located on inside base of pistillate flower and becoming larger on inside base of persistent perianth of fruit during its development. Tomentose: Covered with dense wooly hairs. Ecological Terms Used in Text For further information on this subject, consult Eiten (1972, 1974, 1978, 1982, 1984). Atlantic coastal forest: Tropical mesophytic forest of evergreen or semideciduous trees along eastern coast of Brazil (from Rio Grande do Norte to Rio Grande do Sul), usually in mountainous terrain. Much of the region has been cleared for crops, pastures, and Euca- lyptus plantations. Fortunately, a number of areas have been spared. In northeastern Bahia, palm vegetation (especially comprising sev- eral species of Attalea) is a prominent part of the landscape with hundreds of thousands of individual trees. Caatinga: Unfortunately, there are two different examples of this type of vegetation: (1) Amazon region: closed canopy forest of low or tall trees on white sandy soil; (2) northeastern Brazil: semidesert vegeta- tion on shallow soils that dries out completely during the dry season consisting of arborescent and acaulescent palms, spiny shrubs, low trees, cacti, and terrestrial bromeliads. Campo limpo: Predominantly grassy with many herbs, semishrubs, and ground vines. Campo sujo: Short grass or tall grass with very scattered low shrubs. Cerrado region: Central Brazil. Moderate rainfall, dry season of five months, sterile, very deep soils. Vegetation is xeric, semideciduous low arboreal woodland (open canopy) or low forest with closed canopy. Both arborescent and acaulescent palms are prevalent. Dominance is usually shared by several species. Consult Eiten (1972, 1978) for fur- ther information. Dominant: Tallest and most abundant species in a population; palms ex- hibit dominance in many plant communities. Gallery forest: Forest that closely follows a stream that is a constant source of water, even during the dry season. Frequently, pure stands of palms (e.g., O. phalerata) comprise these forests. Glossary 211 Llanos: Large areas of grassland at low altitudes with readily available water from rainfall or high water table found in Colombia, Venezuela, and the Guianas. Mata cipo: Forest with many lianas or vines, mostly in disturbed areas (e.g., in northeastern Bahia). Restinga vegetation: On podzolic sandy soil of a narrow coastal plain, from Pernambuco to Santa Catarina, Brazil. Low broadleaf forest with many palms. On sites from highly drained sand dunes to low, flat swampy ground. Several species of palms are found in this type of vegetation. Savanna: Broad term usually referring to a grassy area with scattered trees and shrubs. Consult Eiten (1982) for more detailed information. Secondary growth: Vegetation growing in disturbed areas cleared or par- tially cleared of original vegetation by burning or agricultural prac- tices. The new vegetation that replaces the original is usually entirely different, e.g., a bamboo forest coming into a partially cleared palm forest of A. burretiana in central Bahia. Shared dominance or codominance: Two species in a population more or less equally common and equally distributed. Occasionally the two species hybridize and backcross in a palm forest community. Several examples are given in the text. Transitional forest: Disturbed areas where secondary growth vegetation is mixed with Atlantic coastal forest vegetation in northeastern Bahia. Certain species of palms are found here. ed Nae ve nit ot i mi — ied 7 i aN a) ; vol eaiaaa om sine oSteatl yw alee Arh fpaal bbs we teat! A et yp ies ‘Se cee ssa ict tokio eine fev Pt ‘iy Sn tame ncontligh Cath WV CtRL® Lewitt Toth abel Sate ¥ 4s ‘i overt ava an ATH MD WS SH ALOR EIC.Y TH YO iftive ee a ee wend Tinie RaniRRO sexist ot oer DwemReaden ie? Daler are et RE | eee | ® ‘ty sift ni bargi i} rita arn! acy ao ese lie +5" a a — at i ios pit fh Wr 8 mg IE ony iti in MD ates De a Saawariya Aoi rh PGR BSL RE ek) gpnbiad ve ad astt't yin a tanta wei Apa iciraald 1 BWOTy MONTE yt cH NTE, TITUS S aM “He ‘rent ee pena beretyeoy Mey ie o eebh th sete aE meg abl Saige sind biestih i} en Hig meg boethdsaty eee He yi watts hopesnars qe tee ” + Novte oo Rin Covet lg elaieahent Ht ina ea roid MOM MgO beh i wag CRE a Se ey Re rey | MOR TR LU un ae Sa shal eETU NR Uh ¥ icant insite Wes'y ae Tr 2h NENTS Oink lid gi! So aie Re by HM: se a wh ye VE USCA DSAT A fiery bith WM al rie Pyigey| noe v4 Wat ieee dren it aration! Hohe L ciewigny Av iF « beplibide 4 AeeaphteP hh basil vis wy vy Mises Wy ba SF pei hed A hay * i f iba Hil ag ob huts tii vis’ tat tee? i. SES ers cw! yr ity ‘ais ey wopabeey Yh yo a shy Ae rh i! Ae isd Mig Cons Pa Ao Ahi Tat aires At RY ne font day ia crm ' i el i oy i ee HOH ne yyy ipalvive mie p digs nvr, a " hth ie vy reatt wale nats ve pernty if, BAY. ey Vib , vtcraen, PYipapesbenaie j rv nua ii Atl Aen freee, miorls wk . rn bs } Lge WHA ICa) vee Cerne Cae Une MPa ii wey 6 cutie | py pa Coed Brawl. Mele ie ig al unity ALR, vty weap is 9 if, Venrert Leen werk, is The 2) ee A | Vepre (Tur rn er bee finreegt witty a DUK 2 he deel a id SCA) she ae Oy eth , russe wy Dah dtioahe Aa les, i! sprue A ay ai fe ha? Wad ori ~ . ge bia ewe ve pcidiiais, tat ht ” in r pope phe aie we ny Hage coor vtdnihes, i i here gorest; Rocca thang p folk.» ws i yeersa charlene water een 4 Seu Yarra tun ali tee. G. plaleran)) Cont dhrac Gowen ae References Anderson, A. B., and S. Anderson. 1985. A “tree of life” grows in Brazil. Natur. Hist. 94:40-47. Anderson, A. B., and M. J. Balick. 1988. Taxonomy of the babassi com- plex (Orbignya spp: Palmae). Systematic Botany 13:32-50. Anderson, A. B., M. J. Balick, and C. Pinheiro. 1985. O que e babacu? 36th National Botanical Congress of Brazil. Curitiba. Jan. 20-26, 1985. Bailey, L. H. 1933. Certain palms of Panama. Gentes Herb. 3:33-116. . 1939. Certain palms of the Greater Antilles. 2. Article 19. The great Carossier. Gentes Herb. 4:263-65, figs. 167—70. . 1947. Palmae indigenae Trinitenses et Tobagenses. Gentes Herb. 7:353—445. Balick, M. J. 1985. Current status of Amazonian oil palms. Jn C. Pesce, Oil Palms and Other Oilseeds of the Amazon. pp. 172-91. Balick, M. J., A. B. Anderson, and J. T. M. de Medeiros-Costa. 1987. Hy- bridization in the babassu palm complex. 2. Attalea compta x Orbignya oleifera (Palmae). Brittonia 39:26—36. Balick, M. J., C. U. B. Pinheiro, and A. B. Anderson. 1987. Hybridization in the babassu palm complex: Orbignya phalerata x O. eichler. Am. J. Bot. 74:1013-32. Balslev, H., and A. Henderson. 1987. The identity of Ynesa colenda (Palmae). Brittonia 39:1-6. Barbosa Rodrigues, J. 1875. Enumeratio palmarum novarum quas Valle Fluminis Amazonum. Rio de Janeiro. pp. 1-43. . 1879. Protesto-Appendice ao Enumeratio Palmarum Novarum. Rio de Janeiro. pp. 1-48, 2 pls. . 1881. Attalea oleifera. Nova Revista Brazileira Rio de Janeiro 7:123. . 1882. Les Palmiers: Observations sur la monographie de cette famille dans la flora Brasiliensis, pp. 1-53, 4 pls. Rio de Janeiro. . 1888. Palmae Amazonenses novae. Vellosia 1:33-56. . 1891a. Plantas novas cultivadas no Jardim Botanico do Rio de Janeiro 1:17-19, 30-32. 214 A Taxonomic Treatment of the Palm Subtribe Attaleinae (Tribe Cocoeae) . 1891b. Palmae Amazonenses novae. Vellosia 2:91—112. . 1894. Plantas novas cultivadas no Jardim Botanico do Rio de Janeiro 4:24. . 1895 (“1896”). Plantas novas cultivadas no Jardim Botanico do Rio de Janeiro 5:16—23, t. 4c. . 1898. Palmae Mattogrossenses novae vel minus cognitae. Rio de Janeiro. pp. 1-88. . 1899. Palmae novae Paraguayenses quas descripsit et iconibus illustrativit. Rio de Janeiro. pp. 1-66. .1903a. Palmae, in R. Chodat and E. Hassler, Plantae Hasslerianae. Bull. Herbar. Boiss. 3:622—26. . 1903b. Sertum palm. Brasil., ou relation des palmiers nouveaux du Bresil 1:1-140. 91 pls.; 2:1-114, 83 pls. Bruxelles. . 1907. Supplementum ad sertum palm. Brasil. Contr. Jard. Bot. Rio Jan. 4:105—-23. Bartlett, H. H. 1935. Botany of the Maya area. 3. Scheelea lundellii, a new “Corozo” palm from the department of Petén, Guatemala. pp. 45— 47, 5 pls. Beccari, O. 1888. Le palme incluse nel genere Cocos Linn. Malpighia 2:150. . 1916. Il genere Cocos Linn. e le palme affine. Agric. Colon. 10:617. Bernal, R. 1989. Endangerment of Colombian Palms. Principes 33:113- 28. Bondar, G. 1938. n.d. Palmeiras Bahianas do genero Attalea. Seu aproveitamento economico e descripcao de quatro especies novas. Unpublished ms. . 1942a. New palms of Bahia. Field Mus. Natur. Hist. Bot. 22:457— G3; . 1942b. A Piassaveira e outras palmeiras Attaleaneas na Bahia. Inst. Centro Fom. Econ. Bahia Bol. 13:22—30, 59-70. . 1954a. Nova especie de Orbignya, produtora do oleo de babacu. Arq. Jard. Bot. Rio Jan. 13:57-59. . 1954b. O babacu e outras palmeiras produtoras de amendoas oleaginosas no Brasil. Ministeria Agricultura, Servico de Informacao Agric. Rio de Janeiro. 40 figs. 64 pp. . 1957. Novo genero e nova especie de palmeiras da Tribo Attaleine. Arq. Jard. Bot. Rio Jan. 16:49-55, fig. 1, photos 1-3. . 1964. Palmeiras do Brasil. Instituto de Botanica. Sao Paulo. figs. 1-57, pp. 5-159. Booth, W. S. 1889. Kew Bull. add. ser. 2:250, 237-42. Braun, A. 1968. Cultivated palms of Venezuela. Principes 12:39-103, 111- 36, figs. 63-65. References 215 Burret, M. 1929a. Die Palmengattungen Orbignya, Attalea, Scheelea, und Maximiliana. Notizbl. Bot. Gart. Berlin-Dahlem 10:493—543, 651-701. . 1929b. Palmae Cubenses et Domingenses a Cl. E. L. Ekman 1914-— 1928 lectae. Kong]. Svensk. Vetensk. ser. 3, 6:3—28, ts. 8-11. . 1930. Palmae novae Luetzelburgianae. Notizbl. Bot. Gart. Berlin- Dahlem 10:1013-26. . 1932. Attalea cohune Mart. wirklich eine Orbignya. Notizbl. Bot. Gart. Berlin-Dahlem 11:688-90. . 1933. Palmae neogeae III. Fedde Repert. Spec. Nov. Regni Veget. 32:102-5. . 1934a. Palmae neogeae V. Notizbl. Bot. Gart. Berlin-Dahlem 11:1037-50. . 1934b. Palmae neogeae VII. Notizbl. Bot. Gart. Berlin-Dahlem 12:151-59. . 1937. Uber die bisher unbekannte Heimat einer Alten Gartenpal- me, Scheelea osmantha Barb. Rodr. Notizbl. Bot. Gart. Berlin-Dahlem 13:476-77. . 1938. Palmae Brasiliensis. Notizbl. Bot. Gart. Berlin-Dahlem 14:231-60. . 1940. Palmae neogeae 12. Notizbl. Bot. Gart. Berlin-Dahlem 15:99-108. . 1953. Systematische Ubersicht die Gruppen der Palmen. Willdeno- wia 1:57-—74. Campos Porto, J. 1936. Plantas indigenas e exoticas provenientes da Amazonia, cultivadas no Jardim Botanico do Rio de Janeiro. Rodriguesia 11:99. Claassen, C. E., D. W. Jenkins, and K. S. Markley. 1949. Report of the FAO Oilseed Mission in Venezuela. Food and Agricultural Organization of the United Nations, Washington. figs. 13-20, pp. 35-44. Cook, O. F. 1939. Bornoa, an endemic palm of Haiti. Natl. Hort. Mag. 18:245-80. . 1940. Aublet the botanist, a pioneer against slavery, with a memo- rial genus of palms. Washington Acad. Sci. 30:294-99. . 1942. A new commercial oil palm in Ecuador. Natl. Hort. Mag. 21:70-85, figs. 1-6. Cuatrecasas, J. 1947. Vistazo a la vegetacion natural del Bajo Calima. Rev. Acad. Colomb. Ciencia 7:306-12. Dahlgren, B. E. 1936. Index of American Palms. Field Mus. Natur. Hist. Bot. 14:1—438. . 1959. Index of American palms. Plates. Field Mus. Natur. Hist. Bot. 14:pls. 1-412. 216 A Taxonomic Treatment of the Palm Subtribe Attaleinae (Tribe Cocoeae) Damme r, U. 1902. Plantae novae Americanae imprimis Glaziovianae. Bot. Jahrb. Syst. 31, Beiblatt 70:22—23. Darlington, C. D., and A. P. Wylie. 1945. Chromosome atlas of flowering plants. George Allen and Unwin, London. p. 397. de Nevers, G. 1987. The genus Attalea (Palmae) in Panama. Ann. Missouri Bot. Gard. 74:505-10. Dransfield, J., and N. W. Uhl. 1986. An outline of a classification of palms. Principes 30:3-11. Drude, O. 1881. Palmae in Martius, Flora Brasiliensis 3:254—460. . 1887. Palmae in Engler and Prantl, Natur. Pflanzenf. 2. 3:1—-93. Dugand, A. 1940. Un genero cinco especies nuevas de palmas. Caldasia 1:10-19. . 1941. Notas sobre palmas Colombianas y una de Brasil. Caldasia 1:17-29. . 1943. Noticias botanicas Colombianas. II. Especies nuevas y criticas. Caldasia 2:285-93. . 1944. Palmas nuevas o criticas Colombianas II. Caldasia 2:451, 454, 4 figs. . 1953. Notas sobre el genero Attalea (Palmae) en Colombia. Mutisia 18:1-10. . 1954. Notas adicionales sobre el genero Attalea en Colombia. Mutisia 20:3-5. . 1955. Palmas nuevas y notables de Colombia 2. Caldasia 7:129- 57. . 1959. Una palma nueva Scheelea del Bajo Magdalena, Colombia. Mutisia 26:1—6. . 1961. Palms of Colombia. Principes 5:135—44. Eiten, G. 1972. The cerrado vegetation of Brazil. Bot. Rev. 38:201-341. . 1974. An outline of the vegetation of South America. Symp. 5th Cong. Int'l. Primat. Soc. pp. 529-45, fig. 1. . 1978. Delimitation of the cerrado concept. Vegetatio 36(3):169- 78. . 1982. Brazilian “savannas.” Ecological Studies. Ecology of Tropi- cal Savannas 42:25-47, figs. 1-13. Springer-Verlag. Berlin. . 1984. Vegetation of Brasilia. Phytocoenologia 12(2-3):271-92, figs. 1-17. Gebriider Borntraeger, Berlin. Fonseca, E. Teixeira da. 1924. O babassti. Ministeria Agricultura Rio de Janeiro. 44 pp. . 1927. Oleos vegetaes Brasileiros, ed. 2. Rio de Janeiro. pp. 97-101, 261. Froes Abreu, S. 1929. O coco babassti e o problema do combustiva. Ministeria Agricultura Rio de Janeiro. References Daley Furley, P. A. 1975. The significance of Orbignya cohune on the nature and in the development of the soil profile. Biotropica 7:32-36. Galeano, G., and R. Bernal. 1989. La identidad de Scheelea insignis (Palmae). Caldasia 16:10-13. Glassman, S. F. 1967. Collecting palms in Brazil. Principes 11:41-52. . 1972a. A revision of B. E. Dahlgren’s Index of American Palms. J. Cramer, Lehre, Germany. 294 pp. . 1972b. Systematic studies in the leaf anatomy of palm genus Syagrus. Am. J. Bot. 59:775-88, figs. 1-14, ts. 1-3. . 1977a. Preliminary taxonomic studies in the palm genus Aftalea H. B. K. Fieldiana Bot. 38:31-61. . 1977b. Preliminary taxonomic studies in the palm genus Orbignya Mart. Phytologia 36:89-115. . 1977c. Preliminary taxonomic studies in the palm genus Scheelea Karsten. Phytologia 37:219-50. . 1978a. Preliminary taxonomic studies in the palm genus Maximiliana Mart. Phytologia 38:161-—72. . 1978b. Corrections and changes in recent palm articles published in Phytologia. 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H. 1876. Descriptions of new species and varieties of palms collected in the Valley of the Amazon in north Brasil, in 1874. J. Bot. 14(n:s. vol. 5):323—33, 353-59, t. 183. . 1884. In E. F. Im Thurn, Memoranda on the palms of British Guiana. Timehari, J. Royal Agric. Commiss. Soc. of British Guiana 3:219-76. Demerara. Uhl, N. W., and J. Dransfield. 1987. Genera palmarum. A classification of palms based on the work of Harold E. Moore Jr. Bailey Hortorium and International Palm Society. 610 pp., 200 figs. Uhl, N. W., and H. E. Moore. 1973. The protection of pollen and ovules in palms. Principes 17:111-49, figs. 1-29. Voeks, R. A. 1985. Preliminary observations on the reproductive ecology of the piassava palm (Attalea funifera Mart.) An. Acad. Bras. Cienc. 57:524—25. . 1987. A biogeography of the piassava fiber palm (Attalea funifera Mart.) of Bahia, Brazil. Ph.D. diss. University of California, Berkeley. 288 pp. . 1988. Changing sexual expression of a Brazilian rain forest palm (Attalea funifera Mart.) Biotropica 20:107-13. Voss, E. G. (ed.). 1983. International code of botanical nomenclature as adopted by the thirteenth International Botanical Congress, Sydney. Aug. 1981. Junk, The Hague/Boston. Wallace, A. R. 1853. Palm trees of the Amazon and their uses. London. 48 pls., 129 pp. Wendland, H. A. 1878. In O. Kerchove de Denterghem, Les Palmiers— Histoire Iconographique, etc. Paris. 348 pp. 222 A Taxonomic Treatment of the Palm Subtribe Attaleinae (Tribe Cocoeae) Wessels Boer, J. G. 1965. The indigenous palms of Suriname. Leiden. 172 PP: . 1971. Bactris X moore, a hybrid in palms. Acta Bot. Neerland. 20:167-72. . 1972. Clave descriptiva de las palmas de Venezuela. Acta Bot. Venez. 6:299-362. . 1988. Palmas indigenas de Venezuela. Pittieria 17:9-332. (Span- ish trans. of unpublished ms. with additions.) _n.d. Palms of Venezuela. Unpublished ms. Williams, C., J. B. Harborne, and S. F. Glassman. 1983. Flavonoids as taxo- nomic markers in some cocosoid palms. Plant System. Evol. 142:157- 69. Williams, C., J. B. Harborne, and S. F. Glassman. 1985. Further flavonoid studies on Attalea species and some related cocosoid palms. Plant System. Evol. 149:233-39. Attalea Figures 224 A Taxonomic Treatment of the Palm Subtribe Attaleinae (Tribe Cocoeae) Se i : gre a AN EFL cet >. > 1. A. amygdalina. From Humboldt, Bonpland, and Kunth t. 95-96. 1816. Androgy- nous inflorescence with mostly pistillate flowers below and staminate flowers above. 7p Aa) Attalea Figures "WN JO SMITA “742 moog pup tayuao mwonog “KYeAQ 7481 doy, ‘siamoy areururerg -497UA9 doy, “e[[TYIe1 UO pue aseq Ie SIOMOTJ dIeUTUTEIS payoene YIM 19MOLJ 9P][DStd Ya] Oy “9181 “96-46 3 uNny pue ‘purjduog ‘pjoquiny wo1y ‘puyvpstun “VY A Taxonomic Treatment of the Palm Subtribe Attaleinae (Tribe Cocoeae) 226 ‘98uv1 UIe]JUNOU SIy) INOYsnoIY) JURpUNGY ‘oO1lNeL[eg Op eLIIg ‘sIRIayg seul ‘vpodv ‘y “¢ 227 Attalea Figures "S2ARI] UIIMI9q aseq WOIJ UdyR) WNIJ SuIpjoy sauioy “yD ‘[‘opess99 ul ‘O[aAIND ‘sisuavsad ‘y “F 228 A Taxonomic Treatment of the Palm Subtribe Attaleinae (Tribe Cocoeae) 5. A. funifera. Municipality of Salvador. Itapoan sand dunes. Dense stand in background. Note large number of cut palms. 229 Attalea Figures ‘repuog Aq o0Yg ‘UMOUY 10U AjI]LIO] eX eiyeg “‘nafiunf y °9 A Taxonomic Treatment of the Palm Subtribe Attaleinae (Tribe Cocoeae) 230 7. A. funifera. Bahia. Bolandeira. Staminate inflorescence. Photo by Bondar. Attalea Figures 8. A. funifera. Bahia. Bales of piassava fiber in warehouse. Photo by Vosylius. 232 A Taxonomic Treatment of the Palm Subtribe Attaleinae (Tribe Cocoeae) x4 GH S) ~ n o / & o n = o ss) 8 ° Zz E bo = ig eg vo > bo & = re) = o 50 i 2) o So) ae) 5 SS ~ = a) 3 Zz vi ep i= 8 m4 1S E 8 a is:} —Q 8 = ‘S g 3 SS < or) se | S) I bp ~ 12) S o 42 8 = = S & 3 SS 233 Attalea Figures (A) 68Gh ‘19 12 YOUQON’ ‘2dUadsa}INAFU ‘esOSIeUTY “VIYLY “‘VUDYALING "Y “OT A Taxonomic Treatment of the Palm Subtribe Altaleinae (Tribe Cocoeae) 234 ‘snijAso, Aq O10Yg ‘UONaSs sso19 UI diedosaut yI1Y} A19A pue s1aqy jo siaysnjo snondidsuod ajON “WN JO MATA [RUIIIX9 PUL “UONIIS SsO1D ‘eulpnysuo 7 -7ysu 07 Ya] “vuniyjasing *Y *[ | % 2, Ole, ot, ait, ae, (ak Oh te x eo | iA oe. cate tt pe oS i ie Ld delluakes ‘DUDIALING ‘VW JO ISI1OJ UL 99.1) pLIqdy 0} IX9U YIIGON pur vuey sopeares eiyeg ‘nssoqvssnd x “y Fall Nn 7 = S ~~ = x A Taxonomic Treatment of the Palm Subtribe Attaleinae (Tribe Cocoeae) < a Sd, pone 4 » ; nN in) wt ~~ 8 8 ss ~~ Ss z 0 Ss S 5 = oO. be 2) = | ae) iS CS = = ~_ vo oO. jot e) s= = ° & n my SS & jot) B= = ie} << a4 = a) ° Zz = fe) 52) Ss = is} Y Ss << iss] —Q = & 5 & os x < a) re 237 Attalea Figures "(H) E8SP UDUUSSDID pun YHGON “227 ‘eiyeg ‘nssogussnid x ‘VY “PT puqgdy 0} 1xauU saduadsa10yUT BUIPjOY YIGON pue 381095 ‘AITeIO] JuIeG IOpRATeS 238 A Taxonomic Treatment of the Palm Subtribe Attaleinae (Tribe Cocoeae) the iy e WL Lue : > ~ nn oO res & ie} ie} 2 = CS 2 cS a = © eS oD = iss} 6) S fe) rs) o n uo) S is") 42) S 2) fo) o jot) © a & ‘5 2 S & or — S vo 12) n Ko = fon] cS) < 72) 5 QO := 8 4 HS] E S & iss] -Q «| ™ iS 3 = 27. A. septuagenata. Holotype. Schultes and Cabrera 15796 (COL). Note dense dark indument consisting of sharp curved scales on three rachises on right and regu- larly arranged pinnae on left. Attalea Figures 251 Kw 199 « daramillo-die jis Lor completa. fr Ae ee ence team : 4 in “ot t BO. Nam HERBARIOC NACIONAL COLOMBIANO Attelea uberrims Dugand Bp. NOV. oren pelme acaule. Circunferencia 2.63 m. gn le base. Diémetro 0,76 m. Largo de le | aoje 9,15 m. Ancho del pecfolo en le bas 3,40 me Poliolas beseles 0,90 m. ¥oliolas| nedies 1,15 m. Foliolas extremes 0,50 m, NEGATIVE NO. SIASFZ COLCUBIA pepto. de Caldse; Municipio de Palestins. Heciends "La Pleats”, unos 1200 m sit, Ne. 199 R JAR AMILLS MEIIA wor? Bnero,l 28. A. uberrima. Holotype. Jaramillo Mejia 199 (COL). Individual androgynous rachillae each with two pistillate flowers and very long attached staminate rachillae (up to 30 cm).= A. amygdalina. 252 A Taxonomic Treatment of the Palm Subtribe Attaleinae (Tribe Cocoeae) ed EB Notes de Victor li. Patiiio 12 a 16 hojas de 7.10 8 7255 me. de largo por 2 8 2.15 m. de anchura; pecfolo 0.55 m. heste las prineres leci{nias. spate masculine 2.35 me; inflorescencis 1.75 m. Repata Cemenine 1.50 m y es ice 1.40 m. de loo cusles 0.70 m. estén constitu- {dos por el ped@nculo. entre 4O y 50 rru- tos por recimo adulto. HERBARIO NACIONAL COLOMBIANO Attelea victoriens Dugend Gp. Nov. “TYPUS" SOLOMBLA yepto. del Valle; Corregimientos de Golici: y Ceilén, 01 &. de Bugelagranie, sitio "s) Alunendronel"; imeres estribeciones de le sordillere Centrel. No g/n. Victor Me Petifo Junio 1945. sch a aha TCHICAGO NATURAL | HISTORY MUSEUM |...» 99. A. victoriana. Holotype. Patino s.n. (COL). Note large pistillate flowers and relatively short attached staminate rachillae. = A. amygdalina. Attalea Figures MA REM Dee BB Eos — alia eae” Sail * ‘ ‘ 'StON OF ATTALEA HBK. ALLIANCE fe Hersario NaAcionat CortomeBiano SOLOMBIA R. Romero Castaheda 30. A. nucifera. Castaneda 5339 (COL). Part of staminate inflorescence showing long staminate rachillae (up to 19 cm) and staminate flower scars arranged in two rows along one side of rachilla. Original description by Karsten (1857) erroneously described short staminate rachillae (6 cm) and staminate flowers completely sur- rounding rachilla. 253 acne 254 A Taxonomic Treatment of the Palm Subtribe Attaleinae (Tribe Cocoeae) F Tuy as Creat VERS ry ;¢ ; teche °F ILtrNors, [FIELD MUSEUM OF NATURAL HISTORY wenken HOG 31. A. barreirensis. Glassman 13048 (SP). Left to right: Individual staminate flowers. Staminate inflorescences showing relatively short staminate rachillae with one- sided staminate flower scars. Attalea Figures 255 ORTVERSITY GF iLLiNcis CIRCLE 39. A. barreirensis. Glassman 13047 (F). Part of leaf showing clustered middle se- ries pinnae. 256 A Taxonomic Treatment of the Palm Subtribe Attaleinae (Tribe Cocoeae) 33. A. brasiliensis. Glassman and Eiten 13057 (SP). Top left: Part of sterile bract with long beak. Bottom left: Separate staminate flowers. Right: Staminate inflorescences. 257 Attalea Figures (FlELe OF |NATURAL HISTORY) wegative nO | | g2593 34. A. brasiliensis. Glassman and Eiten 13059 (F). Portions of leaves showing regu- larly arranged pinnae. A Taxonomic Treatment of the Palm Subtribe Attaleinae (Tribe Cocoeae) 258 FIELD MUSEUM OF NATURAL HISTORY Bae, Diy ie : | | LJ cence with stami- 35. A. seabrensis. Glassman 13034 (SP). Part of staminate inflores nate flowers on one side of rachillae. i } y i : , ' ; fj y ; ‘es Abs. bee" i SE DAEs a me ; : } ie Scie ate? vel te toxeoesnT wines A HR. ay as . : m =| AS a + wa ss dali _ ' ' j wn Mi { : | A ee 4 t rene | " a Pid a . é 4 a Lith | ; } AS any, i i hg i ; i I oi Misex; ny ‘ ' ‘ a ; q ) Tan) & i H Ye i; ; fee ' Medd | Me my f vA . , ‘ " 7 : a x Uy Fe a r i . aa =i ia ag ‘i sf" bic a on ee... erseniieaiiianet iS See eet A pact» adore yowntt omen? al gif sania’ AM. Wil? is Bae wie aby ai maiisla vay te 7 me Pe i. 6, (2) TY / : ee. pe Pre weemee ct Spry ae ih sani Ali WOT, Hah aU re penal atta, = he i . i ie MeN Demcey! LB ai r, Ut ies'+43 yi ae A) a rye lai ae) a oa ae “4 gens i | Ore ae | pdb (yey (ait xi Waa suns: ne fp sien portal iigrnd shapers gl i oo (rama wp how S yatta ASonk Niwnnt WO arth dish webs red bine Maite te arial totiwoitic mo wok saben ere gees sik ih Beasg Wi site. - nae a 2 inh oe ; a x os + er ; i +" = ’ es _ - 1 | * Dh p oe ; ery at Hi ie iy 6 rt ; Us verte am pats ental veh ‘om ; Ay a a 1 : a ' , i » x y 7 . Poy ; wt < . nin ; a ; ae, i res F ap 5 ® gist in 1. = % - : ‘ he nay he =e ‘s We ~ y 1 aca i j os 260 A Taxonomic Treatment of the Palm Subtribe Attaleinae (Tribe Cocoeae) 36. A. brasiliensis. 37. A. exigua. 38. A. pindobassu. 39. A. septuagenata. Top left: Staminate flower. Top right: Stamen. Glassman 13057 (F). Bottom: Cross section of fruit showing conspicuous fiber clusters in endocarp and two seed cavities. Glassman 13058 (F). Left: Staminate flower. Weddell 2965 (P). Right: Cross section of fruit showing narrow epicarp and mesocarp and conspicu- ous fiber clusters in endocarp. Weddell 2022 (B). Far left: Pistillate flower with perianth. Next left: Pistil with staminodial ring. Center to nght: Various views of staminate flowers showing nerved petals. Far nght: Stamen. Bondar s.n. (F-619761). Left to nght: Longitudinal section of fruit showing 2 seeds, large irregular dark areas of tannin. Cross section of fruit showing 2 seeds, air spaces in mesocarp, clusters of fibers and irregular dark areas of tannin in endocarp. Stamen. Staminate flower showing clusters of glands on inner mar- gins of petals. Schultes and Cabrera 15796 (BH). 261 Attalea Figures 262 A Taxonomic Treatment of the Palm Subtribe Altaleinae (Tribe Cocoeae) 40. A. victoriana. 4]. A. oleifera. 42. A. alleniti. 43. A. funifera. Top row, left to nght: Staminate flower. Stamen. Bottom row, left to right: Pistillate flower with perianth. Pistil with staminodial ring. Patino 215 (COL), Patino s.n. (COL-39725). =A. amyg- dalina. Top: Staminate flower showing partially nerved petals. Bot- tom row, left to ght: Pubescent pistil. Pistillate flower. Glaziou 15556 (G). Top center: Stamen. Cook 64 (US). Bottom row, left to right: Lon- gitudinal and cross sections of fruit. Moore et al. 9460 (BH). Apical portion of leaf showing asymmetrical tip and dark pu- bescent margin. Moore et al. 9460 (BH). Staminate flower showing punctate glands and nerved petals. Cook 64 (US). Left to right: Cross section of fruit with seed, inconspicuous fibers in endocarp, and conspicuous line of fibers on both sides of mesocarp. Dahlgren s.n. (F-611639). Staminate flower. Stamen. Glaziou 16483 (MO). Attalea Figures 263 264 A Taxonomic Treatment of the Palm Subtribe Attaleinae (Tribe Cocoeae) 44. A. barreirensis. 45. A. seabrensis. 46. A. ferruginea. 47. A. burretiana. Left to ight: Longitudinal section of fruit showing single seed. Staminate flower. Stamen. Glassman 13048 (F). Left to right: Cross section of fruit showing 2 seeds and con- spicuous fibers in endocarp. Glassman 13036 (F). Staminate flower. Stamen. Glassman 13034 (F). Top row, left to right: Cross section of fruit showing 3 seeds and fiber clusters. Balick et al. 1000 (F). Staminate flower show- ing floccose patches of indument on lower half of petals. Schultes et al. 18040 (US). Second row: Stamen. Third row, left to right: Pistil. Pistillate petal with nerves. Bottom row, left to right: Pistillate flower. Top view of staminodial ring of fruit showing ciliate margins. Schultes et al. 18040 (BH). Top row, left to nght: Longitudinal and cross sections of fruit showing thick mesocarp. Bondar s.n. (F-619753). Bottom row, left to nght: Staminate flower showing petals with dentate mar- gins. Variations of shapes of stamens. Bondar s.n. (F-619754, F-619759). Apical portion of leaf showing asymmetrical tip and dark lepidote margin. Glassman 13008 (F). Attalea Figures 265 266 48. A. 49. A. 50. A. 52 A. A Taxonomic Treatment of the Palm Subtribe Attaleinae (Tribe Cocoeae) guaranitica. uberrima. compta. . salvadorensis. tessmannit. Left to right: Pistillate flower. Staminate flower with partially nerved petals. Stamen. Schinini 14805 (F). =A. geraensis. Left to right: Pistillate flower. Pistil. Staminate flower showing nerved petals. Stamen. Jaramillo Mejia 199 (COL). = A. amygdalina. Top row: Staminate flower with petals and stamens. Bottom: Sepals and petals of pistillate flower and staminate rachilla with staminate flowers and one pistillate flower at base. Martius, Hist. Natur. Palm. 2. t. 97. 1824. Left to right: Staminate flowers showing four petals with strong nerves, acuminate tips, and denticulate margins. Sta- men with fairly long anther. Glassman 13016 (F). Left: Staminate flowers with nerved petals. Top center: Stamen. Tessmann 5167 (NY). Bottom right: Cross section of fruit show- ing conspicuous fiber clusters and ring around seeds. Tessmann s.n. (B). 267 Attalea Figures 268 A Taxonomic Treatment of the Palm Subtribe Attaleinae (Tribe Cocoeae) 53. A. humilis. 54. A. X piassabossu. 55a. A. nucifera. Far left and center: Longitudinal and cross sections of fruit showing relatively narrow mesocarp. Bondar s.n. (F-619755). Top right: Stamen. Bottom right: Staminate flower with nerved petals and denticulate margins. Glassman 13014 (F). Left to right: Cross section of two-seeded fruit showing epicarp and two mesocarp layers (outer one is dotted). Pistillate flower with attached staminate rachilla. Bondar s.n. (F- 619732). Pistil. Staminate flower. Stamens. Bondar. s.n. (F- 619762). Left to right: Longitudinal and cross sections of fruit showing scattered fibers. Staminate flowers showing patches of dark glands. Skolnik et al. 195016 (US). Stamens. Romero Castaneda 5339 (COL). Attalea Figures 269 270 A Taxonomic Treatment of the Palm Subtribe Attaleinae (Tribe Cocoeae) 55b. A. nucifera. 56. A. geraensis. 57. A. dubia. Top row, left to right: Pistillate flower. Pistillate petal showing strong nerves and fluted margins. Bottom row, left to right: Pistillate flower. Pistil. Pistillate petals showing strong nerves and fluted margins. Romero Castaneda 8392 (COL). Top row, left to ight: External view of fruit showing rugose ex- terior. Cross section of fruit with three seeds. Glassman 13087 (F). Pistillate flower with attached staminate rachilla. Pistil. Eiten and Eiten 2212, 2220 (BH). Bottom row, left to ght: Stami- nate flower. Stamen. Glassman 8744 (F). Top row, left to right: Pistillate flower. Pistil showing stami- nodial ring. Glaziou 17341 (P). Bottom row, left to right: Stami- nate flower. Stamen. Krapovickas 23154 (F). i) 1 \ 1cm ala) Nyy ( av2 A Taxonomic Treatment of the Palm Subtribe Attaleinae (Tribe Cocoeae) 58. A. apoda. Left to nght: Longitudinal and cross sections of fruit showing single seed. Glassman 13000 (F). Staminate flower. Stamen. Glassman 13006 (F). Attalea Figures 273 . 1 7 k & ts f ; | ieee - , et: f { ) BE ABeatidvorpte ucae scr of vie Palin Bubseette ial Nase vay ; my 2 X ‘ ; »] y , a : ; a a i ; wk : a 7 ; (eo © i ; F « i 1 = *. , P s ' a t r ” f : ay . 7 ee en a Se a 4 ‘ . 7 ; - { iY rs aun «> P i . y } ee ‘ . . A eee AAI) alee He aR EA Ait ee Wit eS Vk r% aaah \ + c a ® - A) * af 3 i i / - é = ei ‘ i + { ru Attalea Pinnae Cross Sections All illustrations are diagrams. Lamina (on left) shows upper and lower epi- dermis (double lines), adaxial nonvascular fiber bundles (NVF—upper solid patches), abaxial NVF (lower solid patches), and veins of three dif- ferent sizes (empty circles). Midrib (on right) shows main vascular bundle (MVB) and expansion cell tissue (ECT). Sometimes, fibers are present in either or both (solid patches). Voucher specimens for each cross section are also listed. Most sections are magnified approximately 90x. 276 A Taxonomic Treatment of the Palm Subtribe Attaleinae (Tribe Cocoeae) 59. A. ferruginea. Wessels Boer 1945 (U). 60. A. humilis. Glaziou 8069 (C). Note irregularly shaped adaxial NVF and divided ECT. 61. A. camposportoana. Glassman 13006 (F). =A. apoda. 62. A. seabrensis. Glassman 13034 (F). Attalea Pinnae Cross Sections 59 60 61 62 OOO GOL OS 7 277 278 A Taxonomic Treatment of the Palm Subtribe Attaleinae (Tribe Cocoeae) 63. A. geraensis. Glassman 13065 (F). 64. A. guaranitica. Hassler 1860 (G). = A. geraensis. 65. A. allenii. O. F Cook 64 (US). 66. A. septuagenata. Schultes and Cabrera 15796 (BH). Note irregular size and dis- tribution of NVF. 280 A Taxonomic Treatment of the Palm Subtribe Attaleinae (Tribe Cocoeae) 67. A. oleifera. Glaziou 15556 (G). 68. A. funifera. Dahlgren s.n. (F-16483). Note broad midrib, sharply pointed MVB, divided ECT, and triangular large vein. 69. A. barreirensis. Glassman 13047 (F). 70. A. brasiliensis. Glassman 13060 (F). Attalea Pinnae Cross Sections 281 4459000 a Bee ee ba, U aa (% e@eoeeer”? © (90000200 FO 6 . olor O Se eS 282 A Taxonomic Treatment of the Palm Subtribe Attaleinae (Tribe Cocoeae) 71. A. burretiana. Glassman 13008 (F). 72. A. salvadorensis. Glassman 13016 (F). 73. A. concinna. Bailey and Bailey 517 (BH). = A. dubia. Attalea Pinnae Cross Sections a Se eee rie dd Wl el 73 940.00 0 0-0 0 s an 283 284 A Taxonomic Treatment of the Palm Subtribe Attaleinae (Tribe Cocoeae) 74. A. dubia. Glaziou 8070 (F). Note differences from figs. 59, 60. 75. A. exigua. Weddell 2965 (F). 76. A. X piassabossu. Bondar s.n. (F-619762). Attalea Pinnae Cross Sections 285 ag Use sees () =I TIT be 9.00 9.09 9.0 Po | ) WE A Vand Seber pafinet ot he Pike Sishior ie . \ if f, 4 , Me | Nema yee ¥ 1, dahlias, (Eloi BOF. CE), Tee diene rc in A eaten, Weil f 3 pil eas iT Aj aie POEL Ff rs ia ‘ . , P / = tr 7 Attalea Distribution Maps 288 A Taxonomic Treatment of the Palm Subtribe Attaleinae (Tribe Cocoeae) @ A humilis O A. salvadorensis sk A. x piassabossu + A. oleifera 77. A. humilis, A. salvadorensis, A. x piassabossu, A. oleifera. 289 Attalea Distribution Maps @ A. exigua © A. burretiana A. dubia + A. barreirensis 78. A. exigua, A. burretiana, A. dubia, A. barreirensis. 290 A Taxonomic Treatment of the Palm Subtribe Attaleinae (Tribe Cocoeae) @ A. compta O xAttabignya minarum *& A. brasiliensis + A. camposportoana 79. A. compta, x Attabignya minarum, A. brasiliensis, A. camposportoana (= A. apoda). Attalea Distribution Maps 291 @ A. geraensis O A. seabrensis * A. pindobassu + A. funifera 80. A. geraensis, A. seabrensis, A. pindobassu, A. funifera. 292 A Taxonomic Treatment of the Palm Subtribe Attaleinae (Tribe Cocoeae) @ A. victoriana O A. guaranitica * A. nucifera + A. septuagenata A A.iguadummat 81. A. victoriana (= A. amygdalina), A. guaranitica (= A. geraensis), A. nucifera, A. septuagenata, A. iguadummat. Attalea Distribution Maps 293 A. allenii A. uberrima A. ferruginea A. tessmannii >+* O @ Ynesa colenda 82. A. allenii, A. uberrima (= A. amygdalina), A. ferruginea, A. tessmannit, Ynesa colenda. Mie staal - j ME iy Teaeintinie lente eyy (i lar tity Hietia \ + i anita A "erty ety A amy pipe ae fee eer’ s af i} -. Ret - ; rl s aah a . Aes : ATs me : © 8 : . i A - i A iy a | prenatal amie 1 yy a 7 Wy ip ae) eee Orbignya Figures 296 A Taxonomic Treatment of the Palm Subtribe Attaleinae (Tribe Cocoeae) 83. O. phalerata. Near Terezina. Piaui, Brazil. Natural stand of trees showing shape. Photo by B. E. Dahlgren. Orbignya Figures 297 84. O. phalerata. Near Terezina. Piaui, Brazil. Tree showing several infructescences with subtending sterile bracts. Photo by B. E. Dahlgren. 298 A Taxonomic Treatment of the Palm Subtribe Attaleinae (Tribe Cocoeae) f, Fy et” We 85. O. phalerata. Near Terezina. Piaui, Brazil. Boy holding relatively large stami- nate inflorescence and sterile bract. Photo by B. E. Dahlgren. 299 Orbignya Figures Aq 010Y ‘spaas ajoym jo adeys pur spaas Q-Z SUIMOYS SINJJ JO SUONIAS SSOITD * ‘uai3[YeQg “Ag [Izeag ‘Inelg ‘eulzalay Iwan ‘vyp1gvyd ‘CE ‘98 300 A Taxonomic Treatment of the Palm Subtribe Attaleinae (Tribe Cocoeae) i 87. O. phalerata. Far left: Androgynous inflorescence. Center: Various views of fruits and pistillate and staminate flowers. Far right: Staminate inflorescences. Lower right: Stamens with coiled antlers. Martius, Hist. Natur. Palm. 3:t. 170. 1845. 7: iy Oh AIN &) ; ~ ' \ ; j \ My i = 4) i i , no & ( ms) j ’ 7 ’ | — , 4 he ‘ bs ‘ ‘ i ni a pomp pay nth a4 Hi) or ts gong Cac ¥) Fubherleriait A 7 ap cae Wignninngs % h-siaacnss decreed tio Se as ———— 0 = Ps : } r 4 yA 1 { a, heh ; Vy Fi AR iwi | e De aald iy ‘ 1% " Nj " 4 7. By) ia et ye we “eRAM ae) = Tike mat MN - A *: 7a raid netlairealeeetiietiiaanatiae Unt asad” melanie ina ren natal , * * ay La i , aaa rm i. ’ f Crees Lx J fa Ni te ; : , { . indy % 1 i i ee dt aaa) ky t “ hy "i wat (Sih ate iq Ah Ve ia iW NY AT . ar Pit ere ye ‘ \ veal NOL beak a . mY in 7 : i is F ve vi 302 A Taxonomic Treatment of the Palm Subtribe Attaleinae (Tribe Cocoeae) 88. O. cohune. Pistillate and staminate flowers and fruit. Barbosa Rodrigues. Sert. Palm. 1:t. 54. 1903. 89. O. X teixeirana. Staminate flowers. Balick et al. (1987b). 90. x Attabignya minarum. Staminate flowers. Balick et al. (1987b). Orbignya Figures 303 A Taxonomic Treatment of the Palm Subtribe Attaleinae (Tribe Cocoeae) 304 ‘(ZI808-da) “u's snpuocg "Maqyrta ‘O “16 Orbignya Figures 305 92. O. guacuyule. Hodge, Principes 19:133. 1975. 306 A Taxonomic Treatment of the Palm Subtribe Attaleinae (Tribe Cocoeae) pl. 2, fig. 2.1947. 93. O. cuatrecasana. Cuatrecasas. Rev. Acad. Colomb. Cienc. 7 Orbignya Figures 307 Zs i ee (172% Bas 94. O. crassispatha. Plumier, Nov. Pl. Amer. Gen. t. 1. 1703. 308 A Taxonomic Treatment of the Palm Subtribe Attaleinae (Tribe Cocoeae) HERBARIO NACIONAL COLOMBIANO CHICAGO NATURAL HISTORY MUSEUM NEGATIVE WO S34 87 wenes_ s Centimeters a i er te, HERBARIO NACIONAL COLOMBIANO Dugan Bene & BPe NOVe acaule o tallo corto; hojus de 5-7 met. Reve “curda"s rro de Circasia (Voupés), 00-900 mealtd | No, 71729. Castrecanns, coleet10-K-1939. yh 95. O. luetzelburgii. Cuatrecasas 7172 (COL). Type specimen of Parascheelea anchistropetala. Orbignya Figures 96. O. luetzelburgii. Staminate inflorescence at base of leaves. Schultes, Principes 18:6, fig. 1. 1974. 310 A Taxonomic Treatment of the Palm Subtribe Attaleinae (Tribe Cocoeae) 97. Orbignya sp. Drude, Martius Fl. Bras. 3:t. 101. 1881. Far left and far right: Pin- nae. Center: Infloresence. Staminate sterile bract. This is probably the source of error of A. nucifera, followed by Burret, in which the staminate flowers are spirally arranged around the rachilla. Upper right corner: Flowers belong to Attalea sp. ee MUR a ty FOL Zou ts, oe ores ee i a if D4 mt Wsaivenneait rusian A Sit mula ; “y va \ ie | ‘Saigua oi grteayliins (REMY Harb r dg © mat wd oe r ripen e fu mein } feviaiieoe DA. theese Si house fateecba: Paice ly peice hier TA, pen em AD i shit tics Hew abide iar Wy 4 Magpies: \" Anne ant eis) ogee Wigs te) lnpigg “ausvae a a eth thy and tet furs ra hewingg: sabe dilie dixie UOTE deal" un Aertate Prenat he YB Re DS piemaprrettl) geste Jab angylnceie ‘on a msi) Wr enciont andes Lepin bgbetbeetiaredt Det 4 totnenres) a! ste don) brie been egerie puinesne iGtyisni ee be 8 aritventte Toit on \ pn pagal ft) Oe — AEM) Feed wale le penile / ag ir , e Saban sega eres wparlierls ak y yee: Resa Lagi tpg | thing anlage i | oY it gf ay ; uj ' ay (1 ven a OS Gee a ae ee _ 312 A Taxonomic Treatment of the Palm Subtribe Attaleinae (Tribe Cocoeae) 98. O. brejinhoensis. 99. O. cuatrecasana. 100. O. cohune. Left: Cross section of fruit showing relatively thick mesocarp, seeds, and scattered fibers in endocarp. Glassman 13042 (F). Center: Pistil showing 6 stigmas and staminodial ring. Top night: Stamen with coiled anther. Bottom right: Staminate flower showing petals with irregular margins. Glassman 13041 (F). Left to nght: Staminate flower showing spatulate petals fused at base. Cook 81 (US). Apical portion of pinna showing asym- metrical tip and pubescent margin. Schultes et al. 7373 (GH). Pistillate flower with attached staminate flower at base. Pis- til showing 4 stigmas, pubescent style, and pubescent staminodial ring. Cuatrecasas 16389 (COL). Far left: Longitudinal and cross sections of fruit showing rela- tively thin mesocarp, single seed, and lack of fibers. Yuncker 4970 (F). Right of center: Pistil showing 3 stigmas and short staminodial ring. Glaziou 16468 (MO). Top rnght: Stamen showing coiled anther. Bottom right: Staminate flower show- ing spatulate petals and coiled anthers. Glaziou 16468 (B). Orbignya Figures 313 314 A Taxonomic Treatment of the Palm Subtribe Attaleinae (Tribe Cocoeae) 101. O. eichleri. 102. O. guacuyule. 103a. O. phalerata. Top left: Cross section of fruit showing 5 seeds. Bottom left: Staminate flower showing fused petals with irregular lacini- ate tips and one lanceolate petal. Middle, top to bottom: Pistil- late flower with 6 stigmas. Pistil with 7 stigmas. Pistillate petals with apical projections. Right: Stamen showing coiled anther. Pistil with staminodial ring removed showing 4 stig- mas and pubescent ovary. Bondar s.n. (F). Left to right: Cross and longitudinal sections of fruit showing ring of fibers in endocarp (cross section) and single seed. Moore and Valiente 6199 (BH). Staminate flower showing nerved spatulate petals. Stamen showing coiled anthers stretched out from drying. Moore and Valiente 6199 (BH). Pistil showing 3 stigmas, pubescent ovary, and staminodial ring. Moore and Cetto 6405 (BH). Top row, left to right: Staminate flowers showing various de- grees of fusion of petals and irregular laciniate tips. Flower on far right with partially nerved petals. Bottom: Coiled an- thers. Kuntze s.n. (NY). Orbignya Figures 315 316 A Taxonomic Treatment of the Palm Subtribe Attaleinae (Tribe Cocoeae) 103b. O. phalerata. 104. O. polysticha. 105. O. sagotit. Top row, left to right: Staminate flower with partially nerved petals. Sanders 1762 (FTG). Staminate petals showing strong nerves and irregular tips (five views). Balick et al. 1359 (NY). Anthers (left). Martius, t. 32 (1844). Anthers (ght). Burret, t. 9 (1929a). Second row, left to right: Pistillate flower with 4 stigmas. Pistil with 6 stigmas and pubescent ovary. Kunize s.n. (NY). Cross sections of fruit showing relatively thick en- docarp and seeds. Dahlgren s.n. (F-614714). Bottom row, left to right: Staminate flowers showing partially fused and sepa- rate petals. Frontal view of calyx with scars. Coiled anthers. Orbigny 20 (M). Longitudinal section of fruit showing rela- tively thick endocarp and seeds. Dahlgren s.n. (F-614714). Two views of staminate flowers showing spatulate petals. Prance et al. 2155 (NY), Bunting et al. 3571 (F). Left to right: Staminate flower with spatulate petals. Sagot 601 (P). Stamens with coiled anthers. Wessels Boer 276 (U). Pis- tillate flower. Pistil. Sagot 831 (P). Orbignya Figures oad, 318 A Taxonomic Treatment of the Palm Subtribe Aétaleinae (Tribe Cocoeae) 106. O. crassispatha. 107. O. luetzelburgit. 108. x Maximbignya dahlgreniana. 109. O. X teixeirana. Left to right: Transitional flower showing outer imbricate pistillate sepals, a pistil with black style and stigmas, and 3 black, narrow fleshy staminate petals. Figueras and Louis 2785 (F). Pistillate flower with imbricate sepals and pet- als. Pistil showing black style and stigma with constricted staminodial ring. Ekman 7164 (US). Left to right: Staminate flowers showing hooked and curved fleshy petals (two views). Wessels Boer 2374 (F); Luetzelburg 21969 (B). Pistillate flower. Pistil. Wessels Boer 2374 (F). Left to right: Staminate flower showing flat- tened sickle-shaped petals and stamens with twisted anthers. Wessels Boer 805 (U). Enlarged stamen with twisted anther. Staminate flower with flattened petals. Bondar s.n. (F). Cross sec- tion of fruit showing very thick mesocarp and 3 seeds. Bondar s.n. (F). Left to right: Pistillate flower showing 4 stigmas and nerved sepals. Pistil with staminodial ring. Separate petal with fluted edges and darkened tip. Balick et al. 1312 (NY). Orbignya Figures 319 320 A Taxonomic Treatment of the Palm Subtribe Attaleinae (Tribe Cocoeae) 110a-d. Ynesa colenda. All figures are of staminate flowers from staminate inflor- escences. Figure a represents a collection from several trees; but b and c each represent flowers collected from a single tree showing two kinds of petals and two kinds of anthers. Figure a. Little 6518 (BH). Figure b. Dodson 5753 (BH). Figure c. Dodson and Duke 7742 (BH). Figure d. Staminate flowers. After Balslev and Henderson, fig 8. 1987. 322 A Taxonomic Treatment of the Palm Subtribe Attaleinae (Tribe Cocoeae) 111. Hybrid between Orbignya sp. and Scheelea sp. or Attalea sp. Sanders 1735 (FTG). 112. O. oleifera. 113. O. oleifera. Left to nght: Staminate flower with flat narrow petals. Coiled anthers. Cross section of fruit with 3 seeds and clusters of fibers in en- docarp. Staminate flowers. Balick, Anderson, and Medeiros Costa (1987). Noblick and Lima 4645 (F). Left: Cross section of fruit showing seeds and conspicuous clus- ters of fibers. Right: Variation of staminate flowers from one collection. Orbignya Figures 323 dle She os) i wig > : ~ tba c se ) 9 ~ 4 rn - f he ee twn 2 tad 9 it ) , ae ~ * “ Feat tess “ yas oes a a a ff - 7. 2 Cn ¥ pve 7” it oh b if “ ‘a aha eee ih aT a 1 ae is 7k We Li Ri " fins f. oF ¢ eth er . : > ain Orr ? Nit Sere . : N teed OF Saher 2g seer 0c kL Orbignya Pinnae Cross Sections All illustrations are diagrams. Lamina (on left) shows upper and lower epi- dermis (double lines), adaxial nonvascular fiber bundles (NVF—upper solid patches), abaxial NVF (lower solid patches), and veins of three dif- ferent sizes (empty circles). Midrib (on right) shows main vascular bundle (MVB) and expansion cell tissue (ECT). Sometimes, fibers are present in either or both (solid patches). Voucher specimens for each cross section are also listed. All sections are magnified approximately 90x. 326 A Taxonomic Treatment of the Palm Subtribe Attaleinae (Tribe Cocoeae) 114. O. brejinhoensis. 115. O. phalerata. 116. O. luetzelburgit. 117. O. polysticha. Glassman 13041 (F). Note similarity in shape and distribution of NVF to that in O. phalerata. Glassman 10418 (F). Note divided expansion cell tissue (ECT), abundant adaxial NVF mostly regular in shape, and common abaxial NVF. Luetzelburg 21969 (M). Note abundant, irregu- larly arranged and shaped adaxial NVF and common, large, and irregularly shaped abax- ial clusters attached to base of small and me- dium veins. Wessels Boer 2409 (U). Note abundant, adaxial NVF, scattered abaxial NVF, and common fiber clusters in ECT. Orbignya Pinnae Cross Sections 327 Se ee ee ae ae 414 °2080 cd r 206 ee e0°6 O° gp *grveeaes § 60.900,.0 240 117 . PA907 G4 08 On00.0 0000, fMeaa™ 328 A Taxonomic Treatment of the Palm Subtribe Aitaleinae (Tribe Cocoeae) 118. O. sagotiz. 119. O. huebneni. 120. x Maximbignya dahlgreniana. Bondar s.n. (F). 121. O. cohune. Huebner 100 (B). Note abundant, regularly shaped adaxial NVF, fairly common abaxial NVF, and common fiber clusters in midrib and ECT. Huebner 64 (B). Note abundant and irregu- larly shaped adaxial NVF and uncommon abaxial NVF. Note uncommon, small adaxial and abaxial NVF. Steyermark 45693 (F). Note abundant, large, irregularly arranged clusters of adaxial NVF and scattered abaxial NVF. Orbignya Pinnae Cross Sections 329 118 | ° SLOT arr aR Ea afrone cD O; Oo Ore, 0, (Oo 6 Se eee Pee Bee yO 0 0 oO. < ELSES pee aaron DAEs” SOFT P TF ep osgtGrgys 121 G0" OF Se7e--O .0 0 ee ee 330 A Taxonomic Treatment of the Palm Subtribe Attaleinae (Tribe Cocoeae) 122. O. crassispatha. 123. O. guacuyule. 124. O. cuatrecasana. Ekman H7164 (NY). Note abundant, irregu- larly shaped adaxial NVF, fairly common abaxial NVF, and divided ECT with few fiber clusters. Bennington 9488 (NY). Note abundant, very small clusters of adaxial NVF, fairly common abaxial NVF, and divided ECT with few fiber clusters. Schultes et al. 7373 (GH). Note abundant, ir- regularly shaped adaxial NVF, fairly common abaxial NVF, and divided ECT with abundant fiber clusters. Orbignya Pinnae Cross Sections yay ———_ omy a me Gils eee ibe. ALT eae l ' ‘2.1? Naar F YI , we I ts - +; i or, rm | ; wy. et Be LD ue ered oe wy ail i al mt, ee ie iv oe re i iq) “ b. A besa j me At a. ae AS Danencintite erent xo at Pliny SaeaeOn + le a gis \ ns fi us Ct d ; 4 shee me Orbignya Distribution Maps A Taxonomic Treatment of the Palm Subtribe Aittaleinae (Tribe Cocoeae) 334 O. phalerata O. brejinhoensis O. sagotii O. luetzelburgii ¥x+@ 125. O. phalerata, O. brejinhoensis, O. sagotii, O. luetzelburgit Orbignya Distribution Maps 335 O. eichleri O. oleifera O. polysticha x Maximbignya dahlgreniana oxx+@6 O. x teixeirana 126. O. eichlen, O. oleifera, O. polysticha, x Maximbignya dahlgreniana, O. X teixeirana 336 A Taxonomic Treatment of the Palm Subtribe Attaleinae (Tribe Cocoeae) + x Maximbignya dahigreniana x O. luetzelburgii * O. cuatrecasana 127. x Maximbignya dahlgreniana, O. luetzelburgii, O. cuatrecasana Orbignya Distribution Maps 337 + O. phalerata x O. polysticha @ o. sagotii 128. O. phalerata, O. polysticha, O. sagoti ft RAPS 1 Tide gran e Yrwouernecet 06 fhe Miter Sa cuwaa D + ie ong O 4 Hoyts @s Scheelea Figures Scheelea Figures 341 129. S. regia Lectotype of Scheelea Karsten, Fl. Columb. 2:t. 176. 1866. = S. butyracea. 342 A Taxonomic Treatment of the Palm Subtribe Attaleinae (Tribe Cocoeae) 130. S. attaleoides Top row, left to nght: Fruit. Staminate flower. Stamen. Bottom: Part of androgynous inflorescence. Karsten, Fl. Columb.1:t. 67. 1861. =S. insignis. 131. S. attaleoides Left to right: Part of staminate inflorescence. Part of leaf show- ing base of pinnae. Pistillate flower. Pistils with and without staminodial ring. Karsten, Fl. Columb.1:t. 67. 1861. = S. insignis. 343 Scheelea Figures 344 A Taxonomic Treatment of the Palm Subtribe Attaleinae (Tribe Cocoeae) 132. S. kewensis. Fruits and staminate and pistillate flowers. J. Hooker, Curtis Bot. Mag. t. 7552, 7553. 1897. 133. S. insignis. Left and bottom: Part of leaf showing clustered pinnae, Cen- ter: Pistillate and staminate flowers. Far right: Staminate inflorescence. Martius, Hist. Natur. Palm. 2:t. 94. 1826. Scheelea Figures 345 346 A Taxonomic Treatment of the Palm Subtribe Attaleinae (Tribe Cocoeae) CHICAGO NATURAL HISTORY MUSEUM inches. NEGATIVE NO centimeters 2 53495 || HERBARIO NACIONAL COLOMBIANO Scice hes rn da nick Dies. tet. Ne Aomeroecn ots in mala Indo. CLES Jel Frits bay re; fentoe ventha, athet? scx 134. S. magdalenica. Romero Castaneda 8225 (COL). Several androgynous rachillae showing pistillate flowers below and rather short naked extension of staminate portion of rachilla. Scheelea Figures 347 EAN HERBARIC NACIONAL COLOM ae bes Comisarvia de? Caquera: crtre Florencia ¥ Yeeee 4 om alt RECASAS 3 QUAT | CHICAGO mare | A | | TET ie MUSEUM ' 2 NEGSTIVE 0 om SRZH FB HEVISION OF ATTALEA HBK. ALLIANCE & GLASSMAN 135. S. butyracea. Cuatrecasas 8945 (COL). Several androgynous rachillae showing pistillate flowers below and rather long naked extension of staminate portion of rachilla. 348 A Taxonomic Treatment of the Palm Subtribe Attaleinae (Tribe Cocoeae) (cnicaco narurAL| | TT yt UT | HISTORY MUSEUM |... bi lsoovo Lt 136. S. rostrata. Moore 6450 (BH). Portions of androgynous inflorescence showing androgynous rachillae with pistillate flowers below and rather short- to medium- sized extensions of staminate portion of rachillae. 349 Scheelea Figures 3S OF NEGATIVE NO. itabeiebiercn ce Te Tt ARAL IA NANT sy LE HISTORY MUSEUM nctes | CHICAGO NATURAL 137. S. rostrata. Moore 6450 (BH). Portion of staminate inflorescence showing rather long staminate rachillae with staminate flowers. 350 A Taxonomic Treatment of the Palm Subtribe Attaleinae (Tribe Cocoeae) 138. S. osmantha. Photo by A. C. Langlois from Trinidad. 351 Scheelea Figures pur seypryoes woys AJaaneja1 Surmoys SIIUDISIIO[JUI JIVUIUILIS *(AYD) ‘SIOMOYy AIeUTUTeIS pasuese ATTe11ds L80L LOUvLD ap ‘sisuaunins -¢ GE] 352 A Taxonomic Treatment of the Palm Subtribe Attaleinae (Tribe Cocoeae) FIELD MUSEU NATURAL HISTOY NEGATIVE No. REVISION OF ATTALEA HBK. ALLIANCE Veheoloa heoranyebler bhi 62369 ere Di | Moy 19PL y By: Geek 2 140. S. degranvillei. de Granville 5566 (CAY). Whitish staminate rachillae showing spiral staminate flowers with whitish anthers. Scheelea Figures 558 141. S. fwndellii. Base of petiole showing long fibers. Bartlett (1935). 354 A Taxonomic Treatment of the Palm Subtribe Attaleinae (Tribe Cocoeae) 142. S. lundellii. Staminate inflorescence. Bartlett (1935). Scheelea Figures 355 Mh c aye. IM, TOU Mk 4 pat : Me ELRD 4) 1 1 He \ 143. S. liebmannii. Various fruits, seeds, and staminate and pistillate flowers. Hernandez Xolocotzi (1947). 356 A Taxonomic Treatment of the Palm Subtribe Attaleinae (Tribe Cocoeae) 144. S. liebmannii. Androgynous inflorescence. Hernandez Xolocotzi (1947). Si Scheelea Figures ‘(LP6[) IZ2OD0[0X ZapueUIO} SUOTIIIS JINIF PUL SPIIS SNOLILA “WUUDWUGI) “S “CHT 358 A Taxonomic Treatment of the Palm Subtribe Attaleinae (Tribe Cocoeae) Museum botanicum Berolinense. -, ° i Shi rkin? Ovinvespan Mth « Ante * Nee . mae hratilien hay rian na yebib oy Gap War as Whe Fhe Sant ye Pile ak Vag > 4 We vie «200, eg ee ange tae eae om 146. S. microspadix Burret. Holotype. Hopp 3010 (B). = S. phalerata. Scheelea Figures 359 , dated Ad Tune : A ‘Barb.Rod des.dap.nat. 147. S. princeps var. corumbaensis Barb. Rodr., Palm. Matt. t. 21A. 1898. =S. phalerata. 360 A Taxonomic Treatment of the Palm Subtribe Aftaleinae (Tribe Cocoeae) a8, Pe see pia by Bee > os eae ptt ge Xe See 148. S. guianensis. Staminate inflorescence at base of acaulescent plant. de Granville 7087 (CAY). cD) is} =) a joe 9) o on =| io} ba be 9°] tee he Ba =) iota) o be c E & jot) 5 ie) a ~n =) S es Ou. r) i= oO 12) Nn a =) i) 1S) < “3 a = LY ~ = 7) D > Loot 4 N 09 Ss N Ss S 3 Scheelea Figures 362 A Taxonomic Treatment of the Palm Subtribe Attaleinae (Tribe Cocoeae) 150. S. fairchildensis. 151. S. butyracea. 152. S. amylacea. 153. S. huebnerni. 154. S. lundellii. 155. S. wesselsboerii. Left to nght: Staminate flower. Pistillate flower. Read 900 (BH). Left to nght: Staminate flower showing partially nerved pet- als. Dryander 12 (B). Pistil showing 3 stigmas and large staminodial ring. Cuatrecasas 8945 (COL). Left to right: Staminate flower. Pistillate flower showing 4 stig- mas. Pistil showing staminodial ring. Glaziou 16489 (P). Cross section of fruit showing 4 seeds and conspicuous fiber clusters in endocarp. Dahlgren and Millar s.n. (F-611601). Left to nght: Staminate flower. Cross section of fruit showing 5 seeds. Krukoff 1615 (F). Left to nght: Staminate flower. Stamen. Sieyermark 45718a (F). Left to nght: Staminate flower. Wessels Boer 1990 (F). Cross sec- tion of fruit showing 3 seeds. Schulz and Rodrigues 420 (U). Scheelea Figures y 5mm 363 364 A Taxonomic Treatment of the Palm Subtribe Attaleinae (Tribe Cocoeae) 156. S. bassleriana. 157. S. insignis. 158. S. huebneri. Top: Pistillate flower showing staminate extension of rachilla. Middle left: Pistil with 3 stigmas and pubescent ovary. Tessmann 5237 (NY). Middle right: Staminate flower. Tessmann 5490 (NY). Bottom: Cross section of fruit with seeds and con- spicuous Clusters of fibers. Tessmann 5490 (B). Top row, left to right: Pistillate flower. Pistil with 4 stigmas and pubescent ovary. Triana 731 (P). Bottom row, left to nght: Stami- nate flower. Stamen. Dugand and Jaramillo 2915 (COL). Cross section of fruit showing relatively conspicuous clusters of fibers. Killip 34270 (US). Top row, left to right: Cross section of fruit with one seed. Pis- til without and with staminodial ring, both showing pubes- cent ovary. Pistillate flower with petals covering stigmas. Pistillode showing long narrow stigmas and pubescent ovary. Far left: Normal staminate flower. Stamen. Bottom row, left to right: Series of five staminate flowers from androgynous rachilla showing normal unisexual in 1| and 2, transitional unisexual in 3 and 4, and transitional bisexual in 5. Krukoff 5618 (NY). Scheelea Figures v 1om on Vio) 365 366 A Taxonomic Treatment of the Palm Subtribe Attaleinae (Tribe Cocoeae) 159. S. kewensis. 160. S. Lebmannii. 161. S. macrocarpa. 162. S. leandroana. 163. S. magdalenica. Left to nght: Staminate flower. Royal Bot. Gard. Kew s.n. (K). Cross section of fruit with conspicuous fiber clusters and 3 seed cavities. Nuss s.n. (BH). Left to right: Stamen. Staminate flower. Moore et al. 9482 (BH). Pistillate flower with finely nerved sepals and petals. Pistil. Moore 6121 (BH). Far left: Cross section of fruit showing prominent fibers in endocarp and 2 seeds. Bailey 252 (BH). Top center: Pistillate flower with striated sepals and dark constricted tips. Top nght: Staminate flower. Bottom center: Pistillate flower showing se- pals removed and fluted margins of petals. Bottom right: Pis- til. Steyermark and Davidse 116494 (MO.) Left to right: Cross section of fruit with relatively thick meso- carp, conspicuous fibers in endocarp, and 3 seeds. Dahlgren s.n. (F-611640). Pistillate flower. Pistil showing 4 stigmas and pubescent ovary. Bailey 462 (BH). Staminate flower showing unequal petals. Dahlgren s.n. (F-611651). Left to right: Cross section of fruit. Dugand 558 (COL). Pistil- late flower. Najar 4a (COL). Pistil. Romero Castaneda 8225 (COL). Staminate flower. H. Smith 2639 (NY). Scheelea Figures 367 368 A Taxonomic Treatment of the Palm Subtribe Attaleinae (Tribe Cocoeae) 164. S. lauromuelleniana. 165. S. maracaibensis. 166. S. macrolepis. 167. S. moorei. 168 S. plowmanii. 169. S. osmantha. Top to bottom: Cross section of fruit with thick mesocarp, fiber clusters in endocarp, and 3 seed cavities. Dahlgren s.n. (F-611607). Staminate flower. Dahlgren s.n. (F- 404661). Top row, left to right: Staminate flower showing nerved petals. Wessels Boer 2463 (U). Pistillate flower showing nerved sepals. Bottom: Cross section of fruit showing conspicuous fibers in endocarp and 2 seeds. Wessels Boer 2007 (F). Top to bottom: Cross section of fruit with ring of fibers in endocarp and 2 seeds. Staminate flower. L. Williams DIOL). Left to right: Cross section of fruit with relatively thick mesocarp, conspicuous fiber clusters, and 3 seeds. Moore et al. 8392 (BH). Pistillate flower showing 3 stigmas and distinctly nerved apical parts of perianth. Moore et al. 8539 (BH). Left to nght: Cross section of fruit with conspicuous clus- ters of fibers and 3 seeds. Moore et al. 10214 (BH). Stami- nate flower showing nerved petals. Plowman et al. 6778 (BH). Left: Cross section of fruit with relatively thick mesocarp, scattered fibers, and 2 seeds. Bottom center: Pistillate flower. Bottom right: Pistil with dense band of pubes- cence. Dahlgren s.n. (F-611598). Top right: Staminate flower. Glaziou 16487 (F). Scheelea Figures 369 370 A Taxonomic Treatment of the Palm Subtribe Attaleinae (Tribe Cocoeae) 170. S. phalerata. 171. S. princeps. 172. S. rostrata. 173. S. antsitsiana. 174. S. weberbaueri. 175. S. salazarit. Left to nght: Pistillate flower showing nerved perianth and 4 stigmas. Pistil showing dark staminodial ring. Weddell 2030 (P). Staminate flower with partially nerved petals. Glaziou 22268 (P). Cross section of fruit with conspicuous clusters of fibers and 4 seeds. Glassman 13091 (F). Top: Cross section of fruit showing clusters of fibers and 4 seeds. Schinini et al. 31543 (F). Bottom left: Pistillate flower. Bottom nght: Staminate flower. Schinini et al. 32178 (F). Top row: Staminate flower. Long 154 (F). Bottom row, left to right: Cross section of fruit showing scattered fibers and single seed. Holm and Iltis 809 (BH). Pistillate flower. Pistil (staminodial ring removed) with pubescent ovary and 3 stig- mas. Moore 6540 (BH). Top: Cross section of fruit with fiber clusters and 4 seeds. Schinini et al. 20316 (F). Bottom: Staminate flower. Hassler 7165 (G). Top: Staminate flower. Killip and Smith 25141 (NY). Bottom: Cross section of fruit showing scattered fibers and 2 seeds. Weberbauer 1848 (B). Top: Cross section of fruit with scattered fibers and 2 seeds. Moore et al. 8478 (BH). Bottom: Staminate flower with partially nerved petals. Moore et al. 8481 (BH). Scheelea Figures 371 Sf2 A Taxonomic Treatment of the Palm Subtribe Attaleinae (Tribe Cocoeae) 176. S. guianensis. 177. S. camopiensis. 178. S. degranvillei. 179. S. maripensis. 180. S. maripensis. 181. S. maripensis. 182. Hybrid between Scheelea sp. and Maximiliana maripa. de Granville 7087 (CAY). Staminate flower showing finely nerved fleshy petals. de Granville 2087 (CAY). Staminate flower showing smooth fleshy petals. de Granville 9094 (CAY). Staminate flowers showing rather thick, irregularly nerved, and wrinkled fleshy pet- als. de Granville 10174 (CAY). Staminate flower showing rela- tively long, irregularly nerved, and wrinkled fleshy pet- als. Sist 172 (CAY). Staminate flower showing relatively long, irregularly nerved, and wrinkled fleshy petals. Sist 171 (CAY). Pistillate flower and young pistil. Staminate flower showing narrow fleshy petals with hooked tips and very long anthers. Wessels Boer 1330 (F). = SNS g-2 Zan = i| Ra Le Ni ase f h Teiaeracniy' i ‘Wepinjent ay th ain se had ‘. : le] / vt Seg \) v Py i night + ‘ital 47 nig va ioe seth Pun Vapiomunwe Tye | heyhey Af HW silts =f pa . plete a) f : En fas veh yep? J #4 vy on Vredae Uahs ot - “enti ‘ Pi er i Pet Nae a dehy "PL ate } y } Vi ae i ir v t q ‘ P tert . vet oer $= oF } r + 1 ; 3 i ] Scheelea Pinnae Cross Sections All illustrations are diagrams. Lamina (on left) shows upper and lower epi- dermis (double lines), adaxial nonvascular fiber bundles (NVF—upper solid patches), abaxial NVF (lower solid patches), and veins of three dif- ferent sizes (empty circles). Midrib (on right) shows main vascular bundle (MVB) and expansion cell tissue (ECT). Sometimes, fibers are present in either or both (solid patches). Voucher specimens for each cross section are also listed. All sections are magnified approximately 90x. 376 A Taxonomic Treatment of the Palm Subtribe Altaleinae (Tribe Cocoeae) 183. S. bassleriana. Tessmann 5490 (NY). Note abundant, evenly sized adaxial NVF, scattered abaxial NVF, and fibers in ECT. 184. S. huebnen. Krukoff 5572 (NY). Note uncommon adaxial NVF and relatively large ECT with fiber clusters. 185. S. kewensis. Furtado s.n. (BH). Note abundant, irregularly shaped adaxial NVF, rare adaxial NVF, and divided ECT. 186. S. insignis. Triana 731 (P). Note relatively large ECT with fiber clusters and midrib with fiber clusters. Scheelea Pinnae Cross Sections 377 o9 © @ee ee Se : a Pe &®& ee moO 0 © 0.4 0,00) ia TAF teeth by 640 2, 9@°e 186 378 A Taxonomic Treatment of the Palm Subtribe Attaleinae (Tribe Cocoeae) 187. S. osmantha. Glaziou 16487 (F). Note abundant, irregularly shaped adaxial NVFEF, scattered abaxial NVF, and fiber clusters in ECT. 188. S. macrocarpa. Wessels Boer 2106 (U). Note abundant, irregularly shaped adaxial NVF and common abaxial NVF. 189. S. phalerata. Glassman 13063 (F). Note abundant adaxial NVF and uncommon abaxial NVF. 190. S. anisitsiana. Hassler 7165 (G). Note common adaxial NVF and rare abaxial NVF. Scheelea Pinnae Cross Sections 379 —a—a—S—S=—==_=_™™™\™X—_»_—__ a Be Oe 7 ee BF 6l).0.0,0uPubibe diate a 380 A Taxonomic Treatment of the Palm Subtribe Altaleinae (Tribe Cocoeae) 191. S. amylacea. Dahlgren s.n. (F-611601). Note mostly horizontal irregular adaxial NVF, uncommon abaxial NVF, and divided ECT. 192. S. liebmannii. Moore and Cetto 6229 (BH). Note abundant adaxial NVF and common abaxial NVF mostly attached to base of veins. 193. S. lundellu. Lundeli 3752 (MICH). Note abundant adaxial NVF, scattered abaxial NVF, and fiber clusters in ECT. 194. S. rostrata. Bailey and Bailey s.n. (BH). Note abundant adaxial NVF, scattered abaxial NVF, and fiber clusters in ECT. Scheelea Pinnae Cross Sections 381 192 009894 pO pe yD er 6 04 - ROTI es 4° O89 % *@ eb be ge? ° oe @ 382 A Taxonomic Treatment of the Palm Subtribe Attaleinae (Tribe Cocoeae) 195. S. maracaibensis. Wessels Boer 2007 (US). Note common adaxial NVF and scat- tered abaxial NVF. 196. S. wesselsboerii. Wessels Boer 1990 (U). Note abundant irregularly shaped and distributed adaxial NVF and scattered abaxial NVF. 197. S. lauromuelleriana. Dahlgren s.n. (F-611607). Note abundant irregularly shaped and distributed adaxial NVF, common abaxial NVF, and divided ECT. Scheelea Pinnae Cross Sections — Fee™s Pe .0.0.600. 6 0. ONIN a 6 o ORK) | Re ccc- gee” * 383 195 196 197 384 A Taxonomic Treatment of the Palm Subtribe Attaleinae (Tribe Cocoeae) 198. Hybrid between Scheelea sp. and Maximiliana maripa. Wessels Boer 1330 (U). Note short clusters of adaxial NVF, common abaxial NVF, and divided ECT. Scheelea Pinnae Cross Sections 385 2 0 @@e@ ee = 7 Bp eis e Coe ® - Ore ere oS eee ' i 4 . us : Py i a se ns 4 Pit 7 A * i 1. Whe i ALP be aro ine hu i Yalan Ha ae ns, ‘a “it lng a) } mm Bi mown pone * saa! roto ’ A 1 ¥ d turn i hy x ft DRY so 4 x \ “ P a : a ; wt , j _ f ; , / i 7 <4 Pau £ t ; » ee | ; ‘ “ ) } ’ = : - = a 6 j Z P al { = ae ; +s } i a. m= i i \ Bia : = a ‘ 7 = ! r ws ns . Sng ees Ae ans et et tk ty a a — rieemcyubindllspos iieveanie ——m ai I ' . ia! ange re ee fs " ee A we are ra « ¢ , ae ¢ 9 % ) » 6 o e aa igav ¢ I ie I a aN a — ae. ee : i ( i : am agra : ; { ny yo" — : Ny fi — \ Wa Ieee sti if gh WRIA ohm Le Wald atin ei ‘Aus g 5 hes vat aM TT ATT ar ay | i \ ~ ‘ — | | | | | 7 ——— © ase fa ee OE | RE a = ENN A Oe eR ‘ W r _ 4 ‘ { 4 f ¥ po 7 z pr f f ™~ ‘ a f 5 i 7 ' Se ih) Se eee a) Scheelea Distribution Maps 388 A Taxonomic Treatment of the Palm Subtribe Attaleinae (Tribe Cocoeae) @ S. huebneri * S. phalerata 199. S. huebnen, S. phalerata Scheelea Distribution Maps 389 . moorei . tessmannii . Macrocarpa . magdalenica . maracaibensis onHoOo«x+®6 NNNHH ODN . parviflora 200. S. moore, S. tessmannii, S. macrocarpa, S. magdalenica, S. maracaibensis, S. parviflora (= S. anisitsiana) 390 A Taxonomic Treatment of the Palm Subtribe Attaleinae (Tribe Cocoeae) @ S. osmantha + S. bassleriana XS. salazarii O S. butyracea OS. weberbaueri 201. S. osmantha, S. bassleriana, S. salazarii, S. butyracea, S. weberbaueri Scheelea Distribution Maps 39] S. insignis S. wesselsboerii S. princeps S. macrolepis S. cephalotes oOx xX + ®@ S. plowmanii 202. S. insignis, S. wesselsboerii, S. princeps, S. macrolepis, S. cephalotes, S. plowmani 392 A Taxonomic Treatment of the Palm Subtribe Attaleinae (Tribe Cocoeae) S. guianensis S. camopiensis S. degranvillei * X + @ S. maripensis 203. S. guianensis, S. camopiensis, S. degranvillei, S. maripensis Maximiliana Figures A Taxonomic Treatment of the Palm Subtribe Attaleinae (Tribe Cocoeae) 394 ‘9681 G6 3 ‘SHAE ‘spedas pur sjejad yoys A139 pure suaureys paiiasxa Surmoys siaMoy ayeuTUE)s [eNpIA “Ipuy -woyog “xed |eisip uo siamoy ayeurues pue qed jewrxoid uo siamoy ayeyusid Surmoys eyryses snouAsoipuy :doy, “40z Maximiliana Figures \ 205. Staminate rachilla with staminate flowers spi- rally arranged. Martius, t. 93. 1826. 395 396 A Taxonomic Treatment of the Palm Subtribe Attaleinae (Tribe Cocoeae) 206. Left to right: External view of fruit showing persistent perianth. Longitudinal section of fruit showing persistent perianth and two seed cavities. Drude, t. 104. 1881. Maximiliana Figures 397 207. Relatively young tree showing staminate (eft) and androgynous (right) inflorescences subtended by sterile bracts with long beaks. Georgetown Botanic Garden. Guyana. Dahlgren and Persaud s.n. (F-610752). 398 A Taxonomic Treatment of the Palm Subtribe Attaleinae (Tribe Cocoeae) PR ovr Sth EXPEDITION TO BRAZILIAN AMAZONTS BABIN OF RIO epee Tey a 208. Part of leaf showing clustered pinnae and part of staminate inflorescence showing numerous staminate rachillae surrounding rachis. Krukoff 7067 (NY). why fh Mesn wo, ssaiivcute p's oh a FN Sue i ; ; a a uy " | ” - +8) ‘ . fad ” et hts sbi tore ine a basins = aap gl | | ; awn adi m Tt vig i Sues ase pea €s ie ri y , by ale ppt ones | ier an ae : a egitie ta tora, arti: oT" iat “i ‘ie | . | os | a , . , n , “s r a ne 7 7 li aT Pin: ¥ 7 j Ph met Sn abated meh le i yi Wd : ‘Semen 7 Aue the 400 A Taxonomic Treatment of the Palm Subtribe Altaleinae (Tribe Cocoeae) 209. Staminate flower showing short petals and sepals and exserted stamens. N. L. Britton 494 (NY). 210. Staminate flower showing short petals and sepals and exserted stamens. Rusby and Squires 413 (F). 211. Staminate flower. L. Williams 12816 (F). 212. Pistillate flower showing 3 stigmas. Martius s.n. (G). 213. Pistil showing staminodial ring. Martius s.n. (G). 214. Left and center: Longitudinal and cross sections of fruit showing single seed, relatively thin epicarp, and mesocarp. Right: Top view of staminodial ring show- ing ciliate margins. Wessels Boer 1920 (U). Maximiliana Figures ——— 1 209 Z 12 Ae: 1cm 0.5 cm 0.5 cm rea 401 : : f te al ¥ ' ‘; } poe "yy Lee 20D. 4, Syenmeinie, Fors 2 a ad, ’ uf tg ale ‘ . 4 ‘y ; d (? ae oa | * es byt a! — =) i af L a = -~ - hi A A af ~ #} ws S TT; Sewnaa ToWwes Bag uni ger re | eel i ali ss dec > Fi! note 41 ie Ca) ae Ph see a tes 1 Meee be! p We a Roe z Maximiliana Pinnae Cross Section Lamina (on left) shows upper and lower epidermis (double lines), adaxial nonvascular fiber bundles (NVF—upper solid patches), abaxial NVF (lower solid patches), and veins of three different sizes (empty circles). Midrib (on right) shows main vascular bundle (MVB) and expansion cell tissue (ECT). Fibers are present in both (solid patches). A voucher specimen for the cross section is listed. The section is magnified approximately 90x. 404 A Taxonomic Treatment of the Palm Subtribe Attaleinae (Tribe Cocoeae) 215. M. maripa. Wessels Boer 1920 (U). Note small rounded common clusters of adaxial NVF, abaxial NVF alternating with veins, and divided ECT with fiber clus- ters. Maximiliana Pinnae Cross Section 405 215 ) a A ee We} C076 r) 10s) Os. O1 0 202, of « > Fw . bat a) ¥ . : a” « } i j ‘ | The a é A, . ee Taneitnrenie Prearivenpt of the Yala , rma ‘a ed, cP0) ee ry - ‘ Ae \ i ‘ ’ } df i A, t ; oh < \ x bh | r ‘ i — * , aie ca i i = / z Se ; ; \ ki a = : = \ , z . A Mw a a x - \ CT de J a ; | , . — 215 UN) aorta Phas ore TABI) MG MrT tren idedh My r x . 4 ‘ ~ " +a,'& ¥. et UNyPaleinating with veins, oral divided tao - , { a id r prdpes Vere e -~ f aoe PO au, meine tanner anne Ss Te + “ i = : Muathoy, Tr : ; ; . vy ‘ by =. = —/ A, 1 : - é ie = i ; .. = é —_— I . 1 M “3 t ‘ x a m< , - » me ‘ { if pee ue 2 A ' _ 4 7 er ; ‘2-8 (a i E - i" oh f a on = : Piet. a ee Vv ® “*) * 2 i - Py ac 7 LY j rat a i i vt eas "Fy os 4 eh “s , ih ee raip ah | 4 i Ay ‘iW if ii Li By! a i! yh