11197 FIELDIANA: BOTANY A Continuation of the BOTANICAL SERIES of FIELD MUSEUM OF NATURAL HISTORY VOLUME 32 FIELD MUSEUM OF NATURAL HISTORY CHICAGO, U.S.A. 1. A New Guatemalan Spige/i'o. By Dorothy N. Gibson 1 2. Three New Nicaraguan Epidendrvuns. By Alfonso H. Heller 7 3. Syagrus oleracea ( Mart. ) Becc. and Closely Related Taxa. By S. F. Classman 13 4. Tropical American Plants, X. By Louis O. Williams 35 5. Two New Guatemalan Tournefortias. By Dorothy N. Gibson 65 6. A New Member of Morganella. By Patricio Ponce de Leon 69 7. A New Odontoglossum from Nicaragua. By Alfonso H. Heller 73 8. Studies in the Palm Genus Syagrus Mart. II. By S. F. Glassman 77 9. Revision of the Genus Vascellum (Ly coper daceae). By Patricio Ponce de Leon 109 10. A Conspectus of the Palm Genus Butia Becc. By S. F. Glassman 127 11. Studies in American Plants, II. By Dorothy N. Gibson 173 12. Tropical American Plants, XI. By Louis O. Williams 179 13. The Juglandaceae of Guatemala. By Louis O. Williams and Antonio Molina R 207 14. An Overlooked Genus of the Scrophulariaceae. By Louis O. Williams 211 15. A Synopsis of the Palm Genus Syagrus Mart. By S. F. Glassman 215 16. A New Hybrid in the Palm Genus Syagrus Mart. By S. F. Glassman 241 THE JUGLANDACEAE OF GUATEMALA LOUIS 0. WILLIAMS AND ANTONIO MOLINA R. AN OVERLOOKED GENUS OF THE SCROPHULARIACEAE LOUIS 0. WILLIAMS A SYNOPSIS OF THE PALM GENUS STAGRUS MART. S. F. GLASSMAN A NEW HYBRID IN THE PALM GENUS STAGRUS MART. S. F. GLASSMAN FIELDIANA: BOTANY VOLUME 32, NUMBERS 13, 14, 15, 16 Published by FIELD MUSEUM OF NATURAL HISTORY JULY 24, 1970 The Library of the THE JUGLANDACEAE OF GUATEMALA LOUIS 0. WILLIAMS Chief Curator, Botany, Field Museum of Natural History AND ANTONIO MOLINA R. Field Associate AN OVERLOOKED GENUS OF THE SCROPHULARIACEAE LOUIS 0. WILLIAMS Chief Curator, Botany, Field Museum of Natural History A SYNOPSIS OF THE PALM GENUS STAGRUS MART. S. F. GLASSMAN Research Associate, Palms, Field Museum of Natural History University of Illinois at Chicago Circle A NEW HYBRID IN THE PALM GENUS SYAGRUS MART. S. F. GLASSMAN Research Associate, Palms, Field Museum of Natural History University of Illinois at Chicago Circle FIELDIANA: BOTANY VOLUME 32, NUMBERS 13, 14, 15, 16 Published by FIELD MUSEUM OF NATURAL HISTORY JULY 24, 1970 CONTENTS PAGE The Juglandaceae of Guatemala by Louis 0. Williams and Antonio Molina R. 207 An Overlooked Genus of the Scrophulareaceae by Louis 0. Williams . . . .211 A Synopsis of the Palm Genus Syagrus Mart, by S. F. Glassman 215 A New Hybrid in the Palm Genus Syagrus Mart, by S. F. Glassman . . . .241 FIELDIANA . BOTANY Volume 32, No. 15 July 24, 1970 Publication 1103 A Synopsis of the Palm Genus Syagrus Mart.1'2 S. F. GLASSMAN RESEARCH ASSOCIATE, PALMS FIELD MUSEUM OF NATURAL HISTORY AND UNIVERSITY OF ILLINOIS AT CHICAGO CIRCLE Syagrus as a genus was originally described by Martius in 1824, and in subsequent papers he established a number of species under this name (1826, 1844, 1845). Other species of Syagrus were de- scribed by Barbosa Rodrigues (1875, 1879, 1888), Beccari (1916) Burret (1932, 1933, 1937, 1940), Wessels Boer (1964), and Glassman (1967, 1968a, 1968c). Burret (1937) divided the genus into two sec- tions, Glaziova (containing S. weddelliana, S. insignis, and S. hoehnei), and Orthosyagrus, which included all the other species of Syagrus known to him at the time. Apparently, no one else has attempted to further subdivide the genus into sections or subgenera. Section Glaziova was later taken out of Syagrus and transformed into the new genus Lytocaryum by Toledo (1944). Subsequently, Burret and Potztal (1956) described Microcoelum as a segregate genus of Lyto- caryum. Drude (1881) included Syagrus as one of five sections of the genus Cocos. Except for Eucocos, consisting of C. nucifera, the other sec- tions (Syagrus, Langsdorffia, Arecastrum, and Diplothemiopsis) con- tain species which I presently consider as part of the genus Syagrus. In a later paper, Drude (1887) elevated the section names to sub- generic status; but combined the latter two names under subgenus Arecastrum, incorporated Langsdorffia into subgenus Syagrus, and reduced Glaziova from a separate genus to a subgenus of Cocos. Bec- cari (1887, 1888) divided Cocos into four genera, Barbosa, Rhyticocos, Syagrus, and Cocos. Syagrus included only S. cocoides, and under the genus Cocos, he considered four subgenera: Eucocos, Glaziova, Are- castrum, and Butia. In 1916, Beccari elevated Arecastrum and Butia ^ ! Slnce jhi? pa?ei; h'as gone to Press' a "umber of changes have been made in the text and the phylogenetic tree. These changes will appear as parts of future articles in this journal. 2 This research has been supported by N. S. F. grant no. GB 6899. Library of Congress Catalog Card Number: 70-124572 215 216 FIELDIANA: BOTANY, VOLUME 32 to generic status, and retained the previous generic separation of Barbosa, Rhyticocos, and Arikury (= Arikuryroba Barbosa Rodrigues, 1891) from Cocos. The remaining species of Cocos, except C. nucifera, were transferred to the genus Syagrus. Others who considered Sy- agrus to be part of the genus Cocos are Barbosa Rodrigues (1899, 1900, 1901, 1903, 1907) and Bondar (1939, 1941, 1942, 1964). In the latter article, Bondar subdivided the genus Cocos in Brazil into five groups: Butia, Barbosa, Arikury, Arecastrum, and Syagrus. In previous articles pertaining to Syagrus, I discussed the limits of the genus (1965), described several new taxa (1967, 1968a, 1968c), made some new combinations and reduced a number of species to synonymy (1963, 1968b, 1969, 1970), and transferred several closely related genera to Syagrus (1968b, 1970). The purpose of the present paper is to show probable phylogenetic relationships of the various species of Syagrus by dividing the genus into subgenera, sections, and groups. Due to lack of fossil material and meager evidence from other fields of study, this information is based primarily on gross morphology. Results of this work are de- picted in the form of a key, a phylogenetic tree, and descriptions of various subdivisions of the genus (subgenera, sections, and groups) with a discussion and a list of species under each. A complete re- vision of Syagrus, including descriptions of the genus and all of its species, will appear at a future date. The genus Syagrus is essentially South American in distribution, with only one species (S. amara) occurring in the West Indies. Of the South American taxa (45 species, two varieties, and four hy- brids), 39 of these (including all four hybrids) occur in Brazil, eight in Paraguay, three in Colombia, three in Argentina, two in Peru, two in Bolivia, two in Surinam, two in Uruguay, one in Ecuador, one in British Guiana, and one is known only from cultivation. In contrast to the artificial key to species of Syagrus proposed by me in 1965, the following key is constructed along phylogenetic lines. Closely related species are placed in the same section or group and are keyed out together. The order of appearance in the key, how- ever, is not intended to show a hierarchy of primitive and advanced species. KEY TO THE GENUS SYAGRUS 1. Middle pinnae mostly in tight or loose cluster of 2-4 along rachis of leaf. Subgenus A. Syagrus. 2. Middle pinnae mostly in loose clusters of 2-4, occasionally single. GLASSMAN: SYNOPSIS OF SYAGRUS 217 3. Stem largely subterranean, upright portion very short or absent, middle pinnae up to 1 cm. wide, branched part of spadix 8-25 cm. long. Section I. CAMPYLOSPATHA. 4. Pinnae up to 43 cm. long, mostly with acuminate tips Group a. 5. Expanded part of spathe up to 18 cm. long and 8 cm. wide, branched part of spadix up to 9 cm. long, individual branches gnarled and twisted, 1.0-3.5 cm. long 1. S. pleioclada. 5. Expanded part of spathe up to 30 cm. long and 3.2 cm. wide, branched part of spadix up to 21 cm. long, individual branches straight, up to 16 cm. long 2. S. mendanhensis. 4. Pinnae up to 24 cm. long, mostly with oblique tips Group b. 6. Expanded part of spathe 11-18 cm. long and 2-4 cm. wide, branched part of spadix 8-13 cm. long, branches 4-6 in number, each up to 8 cm. long; female flowers 10-14 mm. long 3. S. loefgrenii. 6. Expanded part of spathe up to 35 cm. long and 4-6 cm. wide, branched part of spadix up to 25 cm. long, branches up to 20 in number, each up to 15 cm. long, female flowers 6-8 mm. long. 4. S. campylospatha. 3. Stem upright, slender, up to 20 m. tall, middle pinnae 0.75-4.2 cm. wide, branched part of spadix 27-70 cm. long Section II. SYAGRUS. 7. Flowering spadix branches relatively few, 9-15 in number. Group c. 8. Female flowers 12-22 mm. long, fruit subglobose, exocarp striated 5. S. stratinicola. 8. Female flowers 7-11 mm. long, fruit ovate, exocarp not stri- ated. 9. Middle pinnae up to 2.8 cm. wide, male flowers 10-19 mm. long, fruit up to 5 cm. long 6. S. cocoides. 9. Middle pinnae up to 0.75 cm. wide, male flowers 5-9 mm. long, fruit up to 2.2 cm. long 7. S. cardenasii. 1. Flowering spadix branches numerous, 25-50 or more in number. 10. Pinnae up to 117 cm. long, expanded part of spathe up to 104 cm. long and 21 cm. wide, endosperm of seed rumi- nate, female flowers 10-13 mm. long. Group d. 8. S. amara. 10. Pinnae up to 56 cm. long, expanded part of spathe up to 62 cm. long and 9 cm. wide, endosperm of seed homogene- ous, female flowers 6-9 mm. long Group e. 11. Fruit obovate, non-beaked, up to 2.5 cm. long and 2.2 cm. in diameter 9. S. allenii. 11. Fruit ovate, short beaked, up to 4.0 cm. long and 2.2 cm. in diameter 10. S. orinocensis. 2. Middle pinnae mostly in tight clusters of 2-4. 12. Petiole margins with long, spine-like projections Section III. CORONATA. 13. Female flowers 9-13 mm. long, pinnae 3-4 cm. wide 11. S. coronata. 13. Female flowers 6-7 mm. long, pinnae 2-3 cm. wide 12. S. treubiana. 12. Petioles with fibrous or smooth margins. 218 FIELDIANA: BOTANY, VOLUME 32 14. Endocarp cavity more or less triangular in cross- section, seeds 3-lobed with truncate ends. Section IV. INAJAI. 15. Endosperm ruminate, female flowers 12 mm. long, 9-10 mm. wide. .Grcup f. 13. S.smithii. 15. Endosperm homogeneous, female flowers 5-7 mm. long, 5-7 mm. wide Group g. 14. S. inajai. 14. Endocarp cavity rounded or irregular in cross- section, but not triangular, seeds round or irreg- ular in cross-section, but not 3-lobed. 16. Female flowers 5-9 mm. long. 17. Female flowers mostly strongly nerved, with obtuse tips. Section V. SANCONA. 18. Female flowers 5-6 mm. long. 15. S. ecuadorensis. 18. Female flowers 7-9 mm. long. 16. S. sancona. 17. Female flowers mostly smooth, with acute tips. Section VI. ARECASTRUM. 19. Endocarp cavity very irregular in shape, seed gibbous-uncinate, pin- nae mostly with acuminate tips. Group h. 17. S. romanzoffiana. 19. Endocarp cavity mostly regular in shape, seed not gibbous-uncinate, pinnae mostly with oblique tips. Group i. 18. S. comosa. 16. Female flowers 12-35 mm. long. 20. Endosperm of seed ruminate, fe- male flowers with obtuse, keeled tips Section VII. BARBOSA 19. S. pseudococos. 20. Endosperm of seed homogeneous, female flowers with acuminate or acute tips. Section VIII. OLERACEA. 21. Acaulescent or small trees up to 4.5 m. tall, middle pinnae 13- 36 cm. long, 0.5-2.5 cm. wide, mostly with oblique tips. Group j. 22. Middle pinnae 0.5-1.6 cm. wide. 23. Acaulescent, pinnae up to 13 cm. long, female flowers up to 13 mm. long. 20. S. rachidii. 23. Small trees up to 4.5 m. tall, pinnae up to 36 cm. long, female flowers up to 22 mm. long. 21. S. flexuosa. GLASSMAN: SYNOPSIS OF SYAGRUS 219 22. Middle pinnae 2.0-2.5 cm. wide. 24. Acaulescent, spadix branches 5-8 in number, up to 22 cm. long, fruit turbinate, endocarp 6-8 mm. thick. 22. S. duartei. 24. Small trees up to 3 m. tall, spadix branches 12- 17 in number, up to 12 cm. long, fruit ovoid, endocarp 2-4 mm. thick along sides. 23. S. glaucescens. 21. Taller trees 10-36 m. tall, mid- dle pinnae 42-80 cm. long, 2.4- 4.0 cm. wide, mostly with acu- minate tips Group k. 25. Female flowers 25-35 mm. long, scars in- flated, up to 10 mm. in diameter. 24. S. macrocarpa. 25. Female flowers 12-22 mm. long, scars not in- flated, up to 6 mm. in diameter. 26. Tree up to 36 m. tall, leaf rachis up to 120 cm. long, pin- nae up to 100 pairs per leaf, each up to 42 cm. long. 25. S. gomesii. 26. Tree 10-20 m. tall, leaf rachis 175-263 cm. long, pinnae 134-152 pairs, each up to 80 cm. long. 6. S. oleracea. 1. Middle pinnae unclustered for the most part, rarely in loose clusters of 2. Subgenus B. Syctgropsis. 27. Spadices simple, unbranched Section IX. DIPLOTHEMIOPSIS. 28. Spadices extending beyond tips of spathes for several cm., fruits 1-2 chambered, 1-2 seeded Group 1. 29. Spathes more or less woody in texture, expanded part up to 19 cm. long, flowering part of spadix 10-12 cm. long, male flowers 5-7 and 8-10 mm. long, fruit ovate, up to 2 cm. long 27. S. campicola. 29. Spathes papery in texture, expanded part 4-5 cm. long, flowering part of spadix 3.0-5.5 cm. long, male flowers 3-5 mm. long, fruit rounded, 1.0-1.4 cm. long 28. S. leptospatha. 28. Spadices completely enclosed by spathes, fruits 1-chambered and 1-seeded. Group m. 30. Flowering part of spadix up to 18 cm. long, female flowers 6-7 mm. long, with broad, acute or obtuse tips 29. S. acaulis. 220 FIELDIANA: BOTANY, VOLUME 32 30. Flowering part of spadix up to 12 cm. long, female flowers 7-14 mm. long, with acute or acuminate tips. 31. Female flowers 10-14 mm. long. 32. Pinnae up to 7 mm. wide, with acuminate or oblique tips, un- clustered throughput, female flowers 4-5 mm. wide, mostly nerved, tips acuminate 30. S. petraea. 32. Pinnae up to 12 mm. wide, mostly with oblique tips, usually single, but in occasional clusters of 2, female flowers 5-8 mm. wide, mostly smooth, tips acute 31. S. glazioviana. 31. Female flowers 7-10 mm. long 32. S. glazioviana var. alpina. 27. Spadices with 2-many branches. 33. Petiole margins spiny, pinnae mostly with oblique tips. Section X. BUTIA. 34. Spathes deeply plicate-sulcate for the most part, fruits 1-chambered and 1-seeded Group n. 35. Acaulescent, middle pinnae in loose clusters of 2 or single, 2.0-2.5 cm. wide, endosperm of seed homogeneous. 33. S. vagans. 35. Trees up to 4 m. tall, middle pinnae unclustered through- out, 3.0-3.5 cm. wide, endosperm of seed ruminate. 34. S. schizophylla. 34. Spathes shallowly plicate-sulcate, appearing smooth, fruits 1-3-chambered and 1-3-seeded Group o. 36. Female flowers 10-16 mm. long, 6-10 mm. wide, ma- ture fruit 3.0-4.2 cm. long. 37. Acaulescent or with trunk 1-2 m. tall, 10-20 cm. in diameter, middle pinnae 0.8-1.3 cm. wide, up to 50 cm. long, rachis of leaf up to 76 cm. long, up to 42 pairs of pinnae, spathe up to 43 cm. long and 5 cm. wide 35. S. paraguayensis. 37. Trees 8-10 m. tall, up to 40 cm. in diameter, middle pinnae 2.0-2.4 cm. wide, up to 81 cm. long, rachis of leaf up to 170 cm. long, up to 72 pairs of pinnae, spathe up to 125 cm. long and 12 cm. wide. 36. S. yatay. 36. Female flowers 3-8 mm. long, 3.5-5.0 mm. wide, ma- ture fruit 1.8-2.6 cm. long. 38. Acaulescent or with very short trunk, middle pin- nae 0.7-1.1 cm. wide, up to 40 cm. long, spathe up to 33 cm. long and 3 cm. wide, spadix with up to 22 branches, petiolar spines short, usually not ex- ceeding 2 mm. in length 37. S. arenicola. 38. Trees 3-6 m. tall, middle pinnae 1.8-2.7 mm. wide, 60-70 cm. long, spathe up to 135 cm. long, 4-16 cm. wide, spadix with 40-60 branches, petiolar spines frequently only a few mm. long, but in older specimens up to 11 cm. long. 39. Spathe up to 16 cm. wide, densely brownish tomentose on outside, female flowers 3-5 (6) mm. long 38. S. eriospatha. 39. Spathe up to 8.5 cm. wide, glaucous on outside, female flowers 4-8 mm. long. . .39. S. capitala. 33. Petiole margins fibrous or becoming smooth with age, pinnae with oblique or acuminate tips. GLASSMAN: SYNOPSIS OF SYAGRUS 221 40. Fruit splitting in 3 sections at maturity, endo- carp thin and papery, pinnae silvery-glaucous or brownish pubescent. - Section XL GLAZIOVA. 41. Middle pinnae up to 40 cm. long and 2.0 cm. wide, usually brownish pubescent, male and female flowers 6-8 mm. long, endo- sperm of seed ruminate. . . .40. S. hoehnei. 41. Middle pinnae up to 23 cm. long and 1.5 cm. wide, usually silvery glaucous, male and female flowers 2.5-5.0 mm. long, endo- sperm of seed homogeneous. 42. Middle pinnae 1.0-1.5 cm. wide, exterior of spathe usually covered with long blackish-brown hairs, spadix branches up to 20 cm. long 41. S. insignis. 42. Middle pinnae 0.5-0.8 (1.2) cm. wide, exterior of spathe usually covered with short chestnut-brown hairs, spadix branches up to 13 cm. long. 42. S. weddelliana. 40. Fruit not splitting into 3 sections at maturity, endocarp woody or bony, pinnae neither pu- bescent nor silvery-glaucous, for most part. 43. Trees up to 18 m. tall, leaf rachis up to 160 cm. long, 50-80 pairs of pinnae, each up to 3.5 cm. wide, mostly with oblique tips, spathes 65-110 cm. long, female flowers 4-7 mm. long. Section XII. SYAGROPSIS. 44. Middle pinnae up to 2.7 cm. wide, spathe up to 20 cm. wide, male flow- ers 7-13 mm. long, mature fruit mostly ovate, up to 4.5 cm. long. 43. S. botryophora. 44. Middle pinnae up to 3.5 cm. wide, spathe up to 6 cm. wide, male flow- ers 3-6 mm. long, mature fruit mostly rounded, up to 2.5 cm. long. 44. S. ruschiana. 43. Acaulescent, leaf rachis 25-70 cm. long, 10-30 pairs of pinnae, each up to 1.0 cm. wide, mostly with acuminate tips, spathes 7-40 cm. long, female flowers 4-13 mm. long. Section XIII. GRAMINIFOLIA. 45. Expanded part of spathe 6-12 cm. long, brownish tomentose, branched part of spadix 4-13 cm. long, fruits 1-chambered. Group p. 46. Branched part of spadix up to 4 cm. long, up to 3 branches in number, each branch up to 3 cm. long. . .45. S. lilliputiana. 46. Branched part of spadix 10-13 cm. long, 13-15 branches in 222 FIELDIANA: BOTANY, VOLUME 32 number, each branch 6-11 cm. long 46. S. hatschbachii. 45. Expanded part of spathe 24-39 cm. long, glaucous or brownish pilose, branched part of spadix 20-30 cm. long, fruits 1-2-cham- bered Group q. 47. Spadix branches 19-31 in number, spathes mostly glau- cous, female flowers 5-7 mm. long, mostly with obtuse tips. 47. S. archeri. 47. Spadix branches 2-7 in num- ber, spathes brownish pilose, becoming glabrous with age, female flowers 7-13 mm. long, with acute tips 48. Female flowers 10-13 mm. long, spathes 1-2 cm. wide, spadix branches 4-7 in. number. 48. S. graminifolia. 48. Female flowers 7-10 mm. long, spathes 0.8-1.2 cm. wide, spadix branches 2-4 in number .... 49. S. gra- minifolia var. nana. The genus Syagrus seems to fall into two natural divisions, sub- genus Syagrus, with clustered pinnae, and subgenus Syagropsis with unclustered pinnae. In the phylogenetic tree (fig. 1), I have shown these as two major lines of development. Subgenus Syagrus is divided into two branches, one with tightly clustered pinnae and the other with loosely clustered pinnae. I have not given formal names to these subdivisions because the latter branch (including sections Syagrus and Campylospatha) appears to represent a transitional stage of development between clustered and unclustered pinnae, and hence may not be a natural grouping. Sec- tion Syagrus is separated from Campylospatha chiefly on its larger dimensions and by being arborescent rather than acaulescent. It is split into three groups based on whether the spadix branches are few or many, size of female flowers and pinnae, and the nature of the endosperm. Some people may wonder why S. amara, previously in- cluded under genus Rhyticocos, is placed in section Syagrus. If one disregards the ruminate endosperm, which I believe evolved inde- pendently in several different sections of the genus Syagrus, e.g., Syagrus, Barbosa, Inajai, Butia, and Glaziova, then the relation- ships with section Syagrus become apparent. The loosely clustered GLASSMAN: SYNOPSIS OF SYAGRUS 223 pinnae, the numerous spadix branches, the triangular, acute-tipped female flowers, and the ovoid, short-beaked fruits would warrant placing S. amara in this section, probably near S. orinocensis and S. allenii. Section Campylospatha is split into two groups based on the length of the pinnae and whether their apices are oblique or acumi- nate. The first group of S. pleioclada and S. mendanhensis especially seems to be a natural one because of the remarkable resemblance of the two species in the vegetative stage. The other group (S. campy- lospatha and S. loefgrenii) does not show such a close relationship. Concerning the sections with tightly clustered pinnae, Oleracea is probably ancestral to all of the others in this grouping because it is the least specialized. This section appears to be a natural unit be- cause the female flowers are relatively large and usually have acumi- nate or acute tips. In general, other sections in this grouping (except Barbosa) have smaller female flowers with acute or obtuse tips. Sec- tion Oleracea is differentiated into two groups, one comprising acau- lescent plants or small trees mostly with oblique-tipped pinnae, and another group of taller trees having pinnae with acuminate tips. Syagrus flexuosa may be intermediate between the two groups be- cause it has pinnae with both oblique and acuminate tips. Appar- ently, each group represents a reduction series, with a decrease in size of the stem in S. duartei and S. rachidii and a decrease in the size of female flowers and width of middle pinnae in S. rachidii, on the one hand; and the reduction in size of female flowers in S. oleracea and S. gomesii, and reduction in the size of the leaf rachis, and number and length of pinnae in S. gomesii, on the other hand. Among the more specialized sections of subgenus Syagrus, sec- tion Coronata is the only one with spiny petioles. This character seems to have arisen independently because it is also found in sec- tion Butia of subgenus Syagropsis. The broad, spine-like projections on the petiole of S. coronata shows a strong similarity to those of S. capitata, S. eriospatha, and S. yatay, but the resemblance is prob- ably only superficial. Section Inajai is characterized by a triangular endocarp cavity in cross-section and a three-lobed seed with truncate ends. Such char- acteristics are so highly specialized that they are found nowhere else in the genus Syagrus. Therefore, it seems highly unlikely that these features could have developed independently. Syagrus smithii was originally described under the genus Chrysallidosperma by Moore (1963) ; and even though the endosperm is ruminate, the resemblance x camposportoana ' x te ^xeiri -x. matafome ecuadorensis /\ 'sancon^ ARECASTRUM SANCONA. smithi^ Common Ancestor FIG. 1. Phylogenetic tree showing probable relationships of various species of Si/ 224 arenicola eripspatha capitata var. alpin /acaulis glaziovian Jtraea Hybrids x camposportoana (Bondar) Classman 3. S. x teixeiriana Classman . coronata x S. romanz off iana) (S. oleracea x S. romanz off iana) x matafome (Bondar) Classman 4. S. x tostana (Bondar) Classman . coronata x S. varans) (S. coronata x S. sohizophvlla) 225 226 FIELDIANA: BOTANY, VOLUME 32 of the fruit and seed of this species and S. inajai is so strong that it could not logically be considered coincidental. In my 1965 paper, I keyed out S. inajai with other species having spiny petiole margins but this is incorrect because the margins are actually fibrous. The strongly nerved female flowers with obtuse tips of section Sancona are apparently a unique combination of characters within Syagrus. Section Barbosa also has blunt-tipped female flowers and may be remotely related to Sancona; but its female flowers are smooth and unusually large (up to 32 mm. long), the fruit is promi- nently beaked and the endosperm is ruminate. Syagrus pseudococos was formerly treated under the genus Barbosa, and although it has no very close relatives within Syagrus, it appears to have the general characteristics of the genus as a whole. The final section in the subgenus Syagrus is Arecastrum, charac- terized by having small female flowers (about 5-7 mm. long and 4- 7 mm. wide) with acute tips and small, ovate fruits, 2-3 cm. long and 1.2-1.7 in diameter. It consists of two groups, differentiated by the nature of the endocarp and seed, and tips of the pinnae. For a long time S. romanzoffianum was treated under the separate genus Arecastrum because of the irregular endocarp cavity and gibbous- uncinate seed, but its affinities with the genus Syagrus are now un- mistakable. Besides the close similarity to S. comosa, it is known to hybridize with at least two different species of Syagrus and is also thought to cross with several others (Classman, 1968b). As previously mentioned, the other major division of Syagrus is the subgenus Syagropsis, characterized by having unclustered pinnae. Section Syagropsis seems to be the least specialized section in this grouping. It differs from section Graminifolia, to which it is most closely related, in its larger dimensions, i.e., tall trees rather than acaulescent, much longer leaf rachis, wider and more numerous pin- nae, and much longer spathes. Graminifolia is divided into two spe- cialized groups, one with 1-2-chambered fruits and 1-2 seeds (S. ar- cheri and S. graminifolia) which is also characteristic of parts of the sections Diplothemiopsis and Butia. Evidently, multiple-chambered fruits with more than one seed arose independently in the three sec- tions mentioned above because members having these features are not closely related. The other group in the section Graminifolia, which includes S. hatschbachii and S. lilliputiana, is characterized by having both its vegetative and reproductive structures extremely re- duced in size. Syagrus lilliputiana is perhaps the smallest species in GLASSMAN: SYNOPSIS OF SYAGRUS 227 the genus with a leaf rachis measuring only 25 cm. long and a spadix up to 5 cm. in length. Section Diplothemiopsis is apparently derived from Graminifolia by the complete loss of spadix branches. Of the two groups in this section, the one with S. campicola and S. leptospatha is more special- ized because its spathes are shorter than the spadices and the fruits are 1-2-chambered with 1-2 seeds. Syagrus leptospatha, particularly, shows extreme specialization in the spathe texture which is papery rather than woody, and, in addition to this, the spathes, spadices, and male flowers are considerably reduced in size. The other group in Diplothemiopsis is less specialized, but it seems to show a trend toward reduction in size of the female flowers (from 10-14 mm. long to 6-7 mm. long) and change in their shape from acute or acuminate to broadly acute or obtuse tips. The remaining two sections, Butia and Glaziova, are among the more highly specialized within the subgenus Syagropsis. They do not appear to have close relatives in any of the other sections. Butia is described here as having spiny petioles and pinnae with oblique tips. It has two well-defined groups which are distinguished chiefly by the depth of grooving in the spathes and the number of chambers and seeds in the fruit. Syagrus schizophylla was formerly treated under a separate genus, Arikuryroba because its seeds have ruminate endosperm; however, it appears to be closely related to S. vagans with homogeneous endosperm (Glassman, 1968b) . Both species have male and female flowers which are almost identical in size and shape. An interesting sidelight is that each is known to hybridize with the same species, S. coronata (also with spiny petiole margins), of the subgenus Syagrus. The other group of species in this section was previously included under the genus Butia, but I gave reasons for transferring them to Syagrus in a recent paper (Glassman, 1970). These closely related species appear to represent one reduction series of larger female flowers (10-16 mm. long and 6-10 mm. wide) and larger fruits (3.0-4.2 cm. long) in S. yatay and S. paraguayensis, to smaller, female flowers (3-8 mm. long and 4-5 mm. wide) and smaller fruits (1.5-2.6 cm. long) in S. capitata, S. arenicola, and S. eriospatha) ; and another series in the attainment of the acaulescent habit in S. paraguayensis and S. arenicola from arborescent ancestors. The last section to be discussed here, Glaziova, consists of three species. Until recently it was thought to be comprised of two differ- ent genera, Lytocaryum (S. hoehnei) and Microcoelum (S. insignis and S. weddelliana) . Syagrus hoehnei was split off from the other two 228 FIELDIANA: BOTANY, VOLUME 32 species because of the ruminate endosperm, however, the three spe- cies make up a closely knit group characterized by fruits which de- hisce into three parts at maturity and by a papery-thin endocarp. The alliances are unmistakably with Syagrus because the leaves, spathes, spadices, and male and female flowers fit the general char- acteristics of the genus. Section Glaziova seems to show a trend toward decrease in length and width of the middle pinnae and a re- duction in size of both male and female flowers, and possibly a loss of ruminate endosperm in the seed. Under this interpretation, S. weddelliana would be the most highly evolved species in the section. The following subgenera, sections, groups, and species are listed according to their appearance in the key. Descriptions and synon- ymy of subgenera and sections are given along with their type spe- cies. Each species under the sections is merely listed with its known geographical distribution, and synonyms are given only when a new combination is made. Complete synonymy and descriptions will appear in a subsequent paper. SYAGRUS subgenus A. Syagrus. Cocos, subgen. Syagrus Drude, Engler & Prantl Natur. Pflanzenf. II, 3: 81. 1887; in part. Acaulescent or trees up to 36 m. tall. Petiole margins mostly fibrous or smooth, seldom spiny, middle pinnae mostly in tight or loose clusters of 2-4, occasionally single, with acuminate or oblique tips; spathes deeply plica te-sulcate; spadices branched, branches 4-many in number; female flowers mostly with acute or acu- minate tips, occasionally obtuse, 5-35 mm. long; mature fruit with a single locule and single seed ; endocarp cavity more or less circular in cross-section, occasionally triangular or irregularly lobed; seed mostly conforming to shape of endocarp cavity, endosperm usually homogeneous, occasionally ruminate. Type species: Syagrus cocoides Mart, which is also the type spe- cies of the genus. This subgenus contains more than one-half of the described spe- cies of Syagrus. I consider it to be representative of the genus as a whole because the sections show a closer relationship to each other and seem to be less specialized than the subgenus Syagropsis. Section I. CAMPYLOSPATHA Classman, sect. nov. Palma acaulis. Piniis in gregibus laxis dispositis ca. 1 cm. latis; spadix pars ramosa 8-25 cm. longa; endospermium non ruminatum. Acaulescent. Petiole margins fibrous; middle pinnae mostly in loose clusters of 2-4, occasionally single, 15-43 cm. long, up to 1.0 cm. wide, with oblique or acuminate tips; expanded part of spathe 18-35 cm. long and 2-8 cm. wide; branched part of spadix 8-25 cm. long, branches 4-20 in number, each branch GLASSMAN: SYNOPSIS OF SYAGRUS 229 1-15 cm. long; male flowers 4-10 mm. long; female flowers 6-14 mm. long, 4-8 mm. wide; fruit ovate, 1.7-3.0 cm. long and 1.0-2.0 cm. in diameter; endosperm of seed homogeneous. Type species: Syagrus campylospatha (Barb. Rodr.) Becc. Section Campylospatha consists of two groups distinguished by the length of the pinnae and the nature of its apices. Group a. Pinnae up to 43 cm. long, mostly with acuminate tips. 1. S. pleioclada Burret — Brazil (Minas Gerais). 2. S. mendanhensis Glassman — Brazil (Minas Gerais). Group b. Pinnae up to 24 cm. long, mostly with oblique tips. 3. S. loefgrenii Glassman — Brazil (Sao Paulo) . 4. S. campylospatha (Barb. Rodr.) Becc.— -Paraguay. Section II. SYAGRUS. Cocos, sect. Syagrus Drude, Mart. Fl. Bras. 3: 401. 1881, in part. Rhyticocos Beccari, Malpighia 1 : 353. 1887, in part (as genus). Syagrus sect. Orthosyagrus Burret, Notizbl. 13:679. 1937. Trees up to 20 m. tall. Petiole margins fibrous; middle pinnae mostly in loose clusters of 2-4, occasionally single, 30-117 cm. long, 0.75-4.2 cm. wide, with oblique or acuminate tips; expanded part of spathe 38-104 cm. long and 3.3- 21 cm. wide; branched part of spadix 27-70 cm. long, branches 9-50 in number, each branch 22-56 cm. long; male flowers 5-19 mm. long; female flowers 6-18 mm. Icng, 4-11 mm. wide; fruit subglobose or ovoid, 2.2-5.8 cm. long and 1.6-3.7 cm. in diameter; endosperm of seed homogeneous, sometimes ruminate. Type species: Syagrus cocoides Mart. Drude (1881) listed the following species under Cocos, section Syagrus: S. botryophora, S. romanzoffiana (in part), S. inajai, S. co- coides, S. comosa, S. graminifolia, S. flexuosa, and S. macrocarpa. I have recognized only S. cocoides as belonging to section Syagrus; the remainder are placed in several other sections of the genus. This section is divided into three groups based mainly on the number of spadix branches, size of pinnae, spathes and female flowers, and nature of the endosperm. Group c. Flowering spadix branches 9-15 in number, endosperm homogeneous. 5. S. stratinicola Wessels Boer — Surinam. 6. S. cocoides Mart. — Brazil (Para, Amapa, Bahia, Piaui, Ceara, Mato Grosso and Goias) ; British Guiana. 7. S. cardenasii Glassman — Bolivia. 230 FIELDIANA: BOTANY, VOLUME 32 Group d. Flowering spadix branches up to 50 in number, pinnae up to 117 cm. long, female flowers 10-13 mm. long, endosperm ruminate. 8. S. amara (Jacq.) Mart. — Lesser Antilles (Martinique, Do- minica, Guadeloupe and St. Lucia). Group e. Flowering spadix branches up to 40 in number, pinnae up to 56 cm. long, female flowers 6-9 mm. long, endo- sperm homogeneous. 9. S. allenii Glassman— Colombia. 10. S. orinocensis (Spruce) Burret — Colombia, Venezuela. Section III. CORONATA Glassman, sect. nov. Palma 9 m. alta. Petiolus cum marginus spinosus; piniis in gregibus strictis dispositis 2.0-4.2 cm. latis. Trees up to 9 m. tall. Petiole margins armed with long, spine-like projections for most of its length; middle pinnae in tight clusters of 2-4, 40-72 cm. long, 2.0- 4.2 cm. wide, mostly with acuminate tips; expanded part of spathe 54-75 cm. long, 8-17 cm. wide; branched part of spadix 38-54 cm. long, branches 42-48 in number each branch up to 30 cm. long; male flowers 6-16 mm. long; female flowers 6- 13 mm. long, 4-7 mm. wide; fruit 2.5-2.7 cm. long, 1.5-2.0 cm. in diameter; endo- sperm of seed homogeneous. Type species: Syagrus coronata (Mart.) Becc. This section consists of two closely related species, therefore no subdivision into groups is necessary. 11. S. coronata (Mart.) Becc.- — Brazil (Bahia). 12. S. treubiana (Becc.) Becc. — Only known from cultivation. Section IV. INAJAI Glassman, sect. nov. Chrysallidosperma H. E. Moore, Principes 7: 109. 1963, in part (as genus). Palma 12 m. alta. Piniis in gregibus strictis dispositis 3-4 cm. latis; fructus cavus endocarpus triquetras; semine trilobo cum termini truncati endospermium ruminatum aut non ruminatum. Trees up to 12 m. tall. Petiole margins fibrous; middle pinnae in tight clusters of 2-4, 66-77 cm. long and 3-4 cm. wide, mostly with acute or acuminate tips; expanded part of spathe 80-85 cm. long, 10-14 cm. wide; branched part of spadix 38-50 cm. long, branches 35-64 in number, each branch 31-40 cm. long; male flowers 5-13 mm. long; female flowers 5-12 mm. long, 5-10 mm. wide; fruit ovoid, with short beak, 4-7 cm. long, 3-4 cm. in diameter, endocarp cavity irregularly triangular in cross-section, trivittate within; seed 2.0-3.5 cm. Icng, 1.2-1.8 cm. in diameter, three-lobed or three-angled in cross-section, truncate at each end, with sculptured impressed sides, endosperm homogeneous or ruminate. GLASSMAN: SYNOPSIS OF SYAGRUS 231 Type species: Syagrus inajai (Spruce) Becc. I have included S. inajai and S. smithii in the same section be- cause of the unique characteristics of the fruit and seed, however, I have placed them in separate groups based on the differences in the endosperm and size of the female flowers. Moore (1963) originally described S. smithii in another genus, Chrysallidosperma, as having affinities with Barbosa and Rhyticocos. In the present paper, I have included the latter two genera in other sections of the subgenus SYAGRUS. Group f. Endosperm of seed ruminate, female flowers 12 mm. long, 9-10 mm. wide. 13. S. smithii (H. E. Moore) Glassman, comb. nov. — Peru. Chrysallidosperma smithii H. E. Moore, Principes 7: 110. 1963. Group g. Endosperm of seed homogeneous, female flowers 5-7 mm. long, 5-7 mm. wide. 14. S. inajai (Spruce) Becc.— Brazil (Para and Amazonas); Surinam, French Guiana. Section V. SANCONA Glassman, sect. nov. Palma 20 m. alta. Piniis in gregibus strictis dispositis 3.5-4.7 cm. latis; flores feminei 5-9 mm. alti cum venas firmas et apices obtuses. Trees up to 20 m. tall. Petiole margins fibrous or smooth; middle pinnae in tight clusters of 2-4, 56-100 cm. long and 3.5-4.7 cm. wide, with acute or acumi- nate tips; expanded part of spathe up to 100 cm. long, and 21 cm. wide; branched part of spadix 76-109 cm. long, branches 100-138 in number, each branch 42- 63 cm. long; male flowers 6-12 mm. long; female flowers mostly strongly nerved, with obtuse tips, 5-9 mm. long and 4-7 mm. wide; fruit ovoid, 3.0-3.4 cm. long, 1.5-2.0 cm. in diameter, with short beak. Type species: Syagrus sancona Karsten. This section contains two closely related species, hence it is not divided into groups. In fact, they may even be conspecific, but due to lack of complete collections of S. ecuadorensis I am reluctant to combine the two species. 15. S. ecuadorensis Becc. — Ecuador. 16. S. sancona Karsten — Colombia, Peru, and Venezuela. Section VI. ARECASTRUM (Drude) Glassman, comb. nov. Cocos, sect. Arecastrum Drude, Mart. Fl. Bras. 3: 402. 1881. Cocos, 232 FIELDIANA: BOTANY, VOLUME 32 subgen. Arecastrum Beccari, Malpighia 1: 351. 1887; Drude, Engler & Prantl, Natur. Pflanzenf. II, 3: 81. 1887. Arecastrum Beccari, L'Agric. Colon. 10: 446. 1916 (as genus). Trees 7-15 m. tall. Petiole margins fibrous or smooth; middle pinnae in tight clusters of 2-4, 41-85 cm. long and 2.0-3.2 cm. wide, with acuminate or oblique tips; expanded part of spathe 72-150 cm. long, 7.5-14 cm. wide; branched part of spadix 32-125 cm. long, branches 19-80 in number, each branch 23-62 cm. long; male flowers 6-16 mm. long; female flowers smooth in texture, with acute tips, 4.5-7.0 mm. long, 4-7 mm. wide; fruit ovoid, 2.0-3.0 cm. long and 1.2-1.7 cm. in diameter, slightly beaked, endocarp cavity regular or very irregular in cross- section; seed ovoid and smooth or gibbous-uncinate. Type species: Syagrus romanzoffiana (Chamisso) Classman. Besides S. romanzoffiana, Drude (1881) included the following species in section Arecastrum: S. coronata, S. oleracea, S. eriospatha, S. schizophylla, S. yatay, and S. capitata. When he elevated Are- castrum to subgeneric status in 1887, Drude also added S. petraea and S. acaulis. In the present paper I have placed the above species in several different sections of the genus Syagrus. On the other hand, Beccari (1916) considered only S. romanzoffiianum under the genus Arecastrum. Section Arecastrum is divided into two groups, represented by one species each. Group h. Endocarp cavity very irregular, seed gibbous-uncinate, pinnae mostly with acuminate tips; length of pinnae, size of spathe, and length and branches of spadices of much larger dimensions than the next group. 17. S. romanzoffianum (Chamisso) Classman. — Brazil (Bahia, Minas Gerais, Goias, Mato Grosso, Sao Paulo, Guanabara, Parana, Santa Catarina and Rio Grande do Sul); Para- guay, Bolivia?, Argentina, and Uruguay?. Group i. Endocarp cavity mostly regular in shape, seed not gib- bous-uncinate, pinnae mostly with oblique tips. 18. S. comosa (Mart.) Mart. — Brazil (Minas Gerais, Bahia?, Ceara, Piaui, Maranhao, Goias and Mato Grosso). Section VII. BARBOSA (Beccari) Classman, comb. nov. Bar- bosa Beccari, Malpighia 1: 349. 1887; L'Agric. Colon. 10: 439. 1916 (as genus). Cocos, sect. Langsdorffia Drude, Mart. Fl. Bras. 3: 401. 1881. GLASSMAN: SYNOPSIS OF SYAGRUS 233 Trees 10-15 m. tall. Petiole margins fibrous; middle pinnae in tight clusters of 2-4, up to 85 cm. long and 4 cm. wide, with acuminate or oblique tips; spadix branches up to 34 cm. long; male flowers 14-22 mm. long; female flowers smooth in texture, 20-32 mm. long, 12-13 mm. wide, with obtuse, keeled tips; mature fruit 6.0-6.8 cm. long, 3.8-4.0 cm. in diameter, with distinct beak; endosperm of seed ruminate. Type species: Syagrus pseudococos (Raddi) Glassman. Although the name Langsdorffia has priority over Barbosa, I have avoided its use because it was originally applied as a generic name in the family Balanophoraceae by Martius in 1818. It was first used in the Palmae by Raddi in 1820 as Langsdorffia pseudococos, thus be- coming a homonym. In 1887, Drude incorporated section Lans- dorffia into the subgenus Syagrus. Beccari (1887, 1916) recognized only one species under the genus Barbosa and in the present paper I am transferring this species to the genus Syagrus under section Barbosa. 19. Syagrus pseudococos (Raddi) Glassman, comb, nov.— Brazil (Guanabara, Sao Paulo, Bahia?) . Langsdorffia pseudococos Raddi, Mem. Soc. Ital. Sci. Modena 18: 345. 1820. Bar- bosa pseudococos (Raddi) Beccari, Malpighia 1 : 349. 1887. Section VIII. OLERACEA Glassman sect. nov. Palma acaulis aut 36 m. alta. Pinniis in gregibus strictis dispositis 0.5-4 cm. latis; flores feminei 10-35 mm. alti cum apices acuminates aut acutates; endo- spermium non ruminatum. Acaulescent or trees up to 36 m. tall. Petiole margins fibrous or smooth; middle pinnae in tight clusters of 2-4, 13-80 cm. long, 0.5-4 cm. wide, with acumi- nate or oblique tips; expanded part of spathe 16-100 cm. long, 3.5-14 cm. wide; branched part of spadix 15-68 cm. long, branches 8-50 in number, each branch 8.5-45 cm. long; male flowers 8-22 mm. long; female flowers 10-35 mm. long, 6-19 mm. wide, with acuminate or acute tips; fruit 2-9 cm. long, 1.0-4.5 cm. in diameter; endosperm of seed homogeneous. Type species: Syagrus oleracea (Mart.) Becc. This section is divided into two groups based chiefly on the size of the plants and size and tips of the pinnae. Group j. Acaulescent or small trees up to 4.5 m. tall, middle pinnae 13-36 cm. long, 0.5-2.5 cm. wide, mostly with oblique tips. 20. S. rachidii Glassman — Brazil (Sao Paulo). 21. S. flexuosa (Mart.) Becc. — Brazil (Minas Gerais, Sao Paulo, Mato Grosso, Goias, and Bahia). 234 FIELDIANA: BOTANY, VOLUME 32 22. S. duartei Glassman — Brazil (Minas Gerais). 23. S. glaucescens Glaz. ex Becc. — Brazil (Minas Gerais). Group k. Taller trees 10-36 m. tall, middle pinnae 42-80 cm. long, 2.4-4.0 cm. wide, mostly with acuminate tips. 24. S. macrocarpa Barb. Rodr. — Brazil (Guanabara and Minas Gerais) . 25. S. gomesii Glassman — Brazil (Sao Paulo). 26. S. oleracea (Mart.) Becc. — Brazil (Parana, Sao Paulo, Gua- nabara, Espirito Santo, Minas Gerais, Goias, Bahia, Per- nambuco, Paraiba and Ceara) ; Paraguay. SYAGRUS subgenus B. Syagropsis Glassman, subgenus nov. Palma acaulis aut 18 m. alta. Petiolus cum marginus fibrosus aut spinosus; pinniis aequaliter pinnatisecta; fructus loculus 1-3; semine 1-3 endospermium non ruminatum interdum ruminatum. Acaulescent or trees up to 18 m. tall. Petiole margins fibrous; smooth or spiny; middle pinnae unclustered for the most part, rarely in loose clusters of 2, with oblique or acuminate tips; spathes deeply or less frequently shallowly plicate- sulcate; spadices usually branched, sometimes unbranched, branches 2-many in number; female flowers mostly with acute tips, occasionally acuminate or obtuse, 2.5-16 mm. long; mature fruit with 1-3 locules and 1-3 seeds; endocarp cavity (with single locule) more or less circular in cross-section, seeds conforming to shape of endocarp cavity or locules, endosperm usually homogeneous, occasionally ruminate. Type species: Syagrus botryophora (Mart.) Mart. Neither the genus Syagrus nor Cocos was previously divided into subgenera based on clustering of pinnae, hence it became necessary to establish a new name for the grouping with unclustered pinnae. As mentioned before, subgenus Syagropsis contains sections which seem to be more specialized than subgenus Syagrus. Although sec- tions Syagropsis, Graminifolia and Diplothermiopsis show fairly close affinities, Butia and Glaziova appear to be more distantly related to the other three sections and to each other. Section IX. DIPLOTHEMIOPSIS (Drude) Glassman, comb, nov. Cocos, sect. Diplothemiopsis Drude, Mart. Fl. Bras. 3: 404. 1881, in part. Acaulescent. Petiole margins fibrous or smooth; middle pinnae unclustered for the most part, 26-38 cm. long, 0.3-1.3 cm. wide, with acuminate or oblique tips; expanded part of spathe usually woody in texture, occasionally papery, 4-23 cm. long, 0.5-3.0 cm. wide; spadix unbranched, flowering part 3.0-18 cm. long; male GLASSMAN: SYNOPSIS OF SYAGRUS 235 flowers 3-20 mm. long; female flowers with acuminate, acute or obtuse tips, 4.5- 14 mm. long, 4-8 mm. wide; fruit 1.0-2.8 cm. long, 0.8-1.7 cm. in diameter, with 1-2 chambers; seeds 1-2, endosperm homogeneous. Type species: Syagrus petraea (Mart.) Becc. Drude (1881) included only S. acaulis and S. petraea in this sec- tion, but in a later paper (1887) he incorporated these species into the subgenus Arecastrum. Beccari (1887) did not think that section Diplothemiopsis was a natural one, and subsequently (1888) listed S. acaulis with subgenus Arecastrum. In the present paper, I am recognizing Diplothemiopsis as a distinct section, probably derived from Graminifolia by loss of its spadix branches. The section is dif- ferentiated into two apparently well-defined groups. Group 1. Spadices extending beyond tips of spathes for several cm., fruits 1-2-chambered, 1-2-seeded. 27. S. campicola (Barb. Rodr.) Becc. — Paraguay. 28. S. leptospatha Burret — Brazil (Mato Grosso). Group m. Spadices completely enclosed by spathes, fruits 1 -cham- bered and 1-seeded. 29. S. acaulis (Drude) Becc. — Brazil (Goias). 30. S. petraea (Martius) Becc.- — Bolivia; Brazil (Mato Grosso and Piaui?). 31. S. glazioviana (Dammer) Becc. — Brazil (Goias). 32. S. glazioviana var. alpina (Drude) Glassman — Brazil (Goias). Section X. BUTIA (Beccari) Glassman, comb. nov. Cocos, sub- gen. Butia Becc., Malpighia 1 : 352. 1887. Butia Beccari, L'Agric. Colon. 10: 471. 1916 (as genus). Arikuryroba Barb. Rodr., PL Nov. Cult. 1 : 5. 1891; Arikury Beccari, I.e. 445. 1916; in part. Acaulescent or trees up to 10 m. tall. Petiole margins mostly armed with short spines, occasionally with coarse spines up to 11 cm. long on basal part; middle pin- nae unclustered for the most part, 40-81 cm. long, 0.7-3.5 cm. wide, mostly with oblique tips; expanded part of spathe usually shallowly plicate-sulcate, sometimes deeply plicate-sulcate, 30-135 cm. long, branches 22-100 in number, each branch 18-62 cm. long; male flowers 3-11 mm. long; female flowers 3-16 mm. long, 3.5- 10 mm. wide; fruit 1.8-4.2 cm. long, 1.0-2.8 cm. in diameter, with 1-3 chambers; seeds 1-3, endosperm mostly homogeneous, sometimes ruminate. Type species: Syagrus capitata (Mart.) Glassman. Beccari (1887) included S. schizophylla under subgenus Butia, but later (1916) he transferred this species to the genus Arikury (= Ariku- ryroba) when he elevated Butia to generic rank. In the present paper 236 FIELDIANA: BOTANY, VOLUME 32 I am placing both genera plus a few other species of Syagrus under section Butia which is split into the following groups. Group n. Spathes deeply plicate-sulcate, fruits 1-chambered and 1-seeded. 33. S. vagans (Bondar) Hawkes— Brazil (Bahia). 34. S. schizophylla (Mart.) Classman— Brazil (Bahia). Group o. Spathes shallowly plicate-sulcate, appearing smooth, fruits 1-3-chambered and 1-3-seeded. 35. S. paraguayensis (Barb. Rodr.) Glassman— Paraguay and Argentina. 36. S. yatay (Mart.) Glassman — Argentina, Paraguay, Uruguay and Brazil (Rio Grande do Sul). 37. S. arenicola (Barb. Rodr.) Frambach ex Dahlgr.— Paraguay and Brazil (Mato Grosso and Minas Gerais)?. 38. S. eriospatha (Mart, ex Drude) Glassman — Brazil (Rio Grande do Sul and Santa Catarina) . 39. S. capitata (Mart.) Glassman — Brazil (Minas Gerais, Goias, Sao Paulo, Parana, Santa Catarina and Rio Grande do Sul) and Uruguay. Section XI. GLAZIOVA (Mart, ex Drude) Burret, Notizbl. 13: 677. 1937. Glaziova Mart, ex Drude, Mart. Fl. Bras. 3: 395. 1881 (as genus). Cocos subgen. Glaziova (Mart, ex Drude) Beccari, Mal- pighia 1 : 443. 1887; Drude, Engler & Prantl Naturl. Pflanzenf. II, 3: 81. 1887. Lytocaryum Toledo, Arq. Bot. Est. Sao Paulo 2: 6. 1944; Burret & Potztal, Willdenowia 1 : 387. 1956. Lytocaryum sect. Glazi- ova (Mart, ex Drude) Toledo, and sect. Maxburretia Toledo, I.e. 7. Microcoelum Burret & Potztal, I.e. 387. Trees up to 5 m. tall. Petiole margins smooth or fibrous; middle pinnae un- clustered, silvery-glaucous or brownish pubescent, 21-40 cm. long, 0.5-2.0 cm. wide, mostly with oblique tips; expanded part of spathe brownish or blackish tomentose on outside, 40-56 cm. long, 2.5-7.0 cm. wide; branched part of spadix 36-40 cm. long, branches 38-62 in number, each branch 8-20 cm. long; male flowers 2.5-8.0 mm. long; female flowers 2.5-8.0 mm. long, 2.5-6.0 mm. wide; fruit 1.7-3.3 cm. long, 1.0-2.3 cm. in diameter, usually dehiscing along 2 or 3 seams at maturity, endocarp thin and papery, 0.5-1.0 mm. thick; endosperm of seed homogeneous or ruminate. Type species: Syagrus weddelliana (Wendl.) Becc. Glaziova was originally described as a genus of palms by Drude (1881) in which he included G. insignis and G. martiana (= S. weddel- GLASSMAN: SYNOPSIS OF SYAGRUS 237 liana). Beccari (1887) reduced Glaziova to a subgenus under Cocos, and in 1910 he described a new species, G. treubiana (now included in section Coronata), under the genus Glaziova. Later, Beccari (1916) incorporated the three species listed above into the genus Syragrus, chiefly because it was discovered that the name Glaziova previously was published as a genus in the family Bignoniaceae in 1868 by Bureau. Since this section apparently consists of closely related species, I have not chosen to divide it into groups. 40. S. hoehnei Burret— Brazil (Sao Paulo). 41. S. insignis (Hort. ex Drude) Becc.- — Brazil (Guanabara and Espirito Santo?) . 42. S. weddelliana (Wendl.) Becc. — Brazil (Guanabara). Section XII. SYAGROPSIS Trees up to 18 m. tall. Petiole margins fibrous or smooth; middle pinnae un- clustered, 53-62 cm. long, 2.7-3.5 cm. wide, mostly with oblique tips; expanded part of spathe 65-110 cm. long, 6-20 cm. wide; branched part of spadix 50-98 cm. long, branches 34-100 in number, each branch 34-55 cm. long; male flowers 3- 13 mm. long; female flowers 4-7 mm. long, 3-7 mm. wide; fruit 2.5-4.5 cm. long, 2.0-2.5 cm. in diameter; endosperm of seed homogeneous. Type species: Syagrus botryophora (Mart.) Mart. As previously stated, this section seems to be the least specialized in the subgenus Syagropsis, and is probably ancestral to section Graminifolia. Syagrus botryophora was formerly considered to be a variety of Arecastrum romanzoffianum by Beccari (1916), and S. ruschiana was included in the genus Arikuryroba by Toledo (1944) . Both species show a close affinity, therefore this section is not broken down into groups. 43. S. botryophora (Mart.) Mart.— Brazil (Bahia). 44. S. ruschiana (Bondar) Glassman — Brazil (Espirito Santo). Section XIII. GRAMINIFOLIA Glassman, sect. nov. Palma acaulis. Folia rachis 25-70 cm. longa; pinniis utrinque 10-30 aequaliter pinnatisecta 0.4-1.0 cm. latis cum apices acuminates; spatha 7-40 cm. longa; flores feminei 4-13 mm. alti. Acaulescent or with short trunk up to 1 m. high. Petiole margins fibrous or smooth; middle pinnae unclustered, 26-55 cm. long, 0.4-1.0 cm. wide, mostly with acuminate tips; expanded part of spathe brownish pubescent or glaucous, 6.5- 39 cm. long, 1-3 cm. wide; branched part of spadix 5-30 cm. long, branches 2-31 in number, each branch 3-18 cm. long; male flowers 3-10 mm. long; female flowers 238 FIELDIANA: BOTANY, VOLUME 32 4-13 mm. long, 3.5-7.0 mm. wide; fruit 1.2-2.0 cm. long, 0.7-1.4 cm. in diameter, with 1-2 chambers; seeds 1-2, endosperm homogeneous. Type species: Syagrus graminifolia (Drude) Becc. Section Graminifolia is divided into the following two groups. Group p. Expanded part of spa the 6-12 cm. long, brownish to- mentose, branched part of spadix 4-13 cm. long, fruits 1-chambered. 45. S. lilliputiana (Barb. Rodr.) Becc. — Paraguay. 46. S. hatschbachii Glassman- — Brazil (Parana). Group q. Expanded part of spathe 24-39 cm. long, glaucous or brownish pilose, branched part of spadix 20-30 cm. long, fruits 1-2-chambered. 47. S. archeri Glassman — Brazil (Minas Gerais, Sao Paulo, and Goias) . 48. S. graminifolia (Drude) Becc.— Brazil (Piaui, Goias and Mato Grosso?). 49. S. graminifolia var. nana (Drude) Becc. — Brazil (Goias) . It should be pointed out here that superficially, S. archeri resem- bles S. arenicola of section Butia more closely than S. graminifolia. The first two species have shallowly plicate-sulcate spathes which may become deeply grooved with age, middle pinnae about the same length and width, and male and female flowers approximately the same size. Syagrus archeri, however, differs from S. arenicola pri- marily in having fibrous rather than spiny petiole margins and pinnae with acuminate rather than oblique tips; hence it would appear to be more closely related to species in section GRAMINIFOLIA. Neither mature fruits nor seeds of S. arenicola have been seen by me, therefore, comparisons of these characteristics with S. archeri can- not be made. REFERENCES BARBOSA RODRIGUES, J. 1875. Enumeratio palmarum novarum quas Valle Fluminis Amazonum in- ventas et ad sertum Palmarum, p. 40. Rio de Janeiro. 1879. Protestp-Appendice ao Enumeratio Palmarum Novarum, pp. 45-46. Rio de Janeiro. 1888. Palmae Amazonenses novae. Vellosia 1 : pp. 33-56. 1891. Plantas novas cultivadas no Jardin Botanico do Rio de Janeiro 1: pp. 10-14. 1899. Palmae novae Paraguayenses, pp. 7-13. Rio de Janeiro. 1900. Palmae Hasslerianae novae, pp. 5-13. Rio de Janeiro. GLASSMAN: SYNOPSIS OF SYAGRUS 239 1901 Palmae Uruguayenses novae vel minus cognitae. Contr. Jard. Bot. Rio de Janeiro 2: pp. 23-43. 1903. Sertum Palmarum Brasiliensium, ou relacion des palmiers nouveau du Bresil, decouverts, decrits et dessines d'apres nature 1: pp. 86-102. 1907. Supplementarum ad Sertum Palmarum Brasiliensium. Contr. Jard. Bot. Rio de Janeiro 4: pp. 105-123. BECCARI, O. 1887. Le palme incluse nel genere Cocos Linn. Malpighia 1: pp. 343-354; 441-454. 1888. Le palme incluse nel genere Cocos Linn. Malpighia 2: pp. 85 95; 147- 156. 1910. Glaziova Treubiana, nouvelle espece de Cocoinee, avec observations sur le genre Cocos. Ann. Jard. Bot. Buitenzorg ser. 2, suppl. 3: pp. 791-806. 1916. II genere Cocos Linn, e le palmae affine. L'Agricoltura Coloniale 10: pp. 435-471; 489-532; 585-623. BONDAR, G. 1939. Palmeiras da Bahia. Inst. Central Fomento Econ. Bahia Bol. 6: pp. 3-22. 1941. Palmeiras do Genero Cocos e descricao de duas especies novas., I.e. Bol. 9: pp. 13-53. 1942. New palms of Bahia. Field Mus. Nat. Hist. Bot. 22: pp. 457-463. 1964. Palmeiras do Brasil. Pp. 58-89. Institute de Botanica. Sao Paulo. BUREAU, M. E. 1868. Sur quelques Bignoniacees nouvelles. Adansonia 8: pp. 379-380. BURRET, M. 1932. Palmae neogeae I. Notizbl. 2: pp. 313-317. 1933. Palmae neogeae III. Fedde Rep. 32: pp. 102-115. 1937. Die palmengattung Syagrus Mart. Notizbl. 13: pp. 677-696. 1940. Palmae neogeae XII. Notizbl. 15: pp. 99-108. BURRET, M. and POTZTAL, E. 1956. Microcoelum, eine neue palmengattung (Cocoideae). Willdenowia 1: pp. 386-388. DRUDE, 0. 1881. Martius Flora Brasiliensis 3: pp. 398-428. 1887. Palmae. In Engler and Prantl, Die naturlichen Pflanzenfamilien II. 3: p. 81. GLASSMAN, S. F. 1963. Nomenclatural changes in the Family Palmae. Rhodora 65: pp. 259- 261. 1965. Preliminary studies in the palm genus Syragrus Mart. Fieldiana: Bot. 31: pp. 285-299. 1967. New species in the palm genus Syagrus Mart. Fieldiana: Bot. 31: pp. 235-245. 1968a. New species in the palm genus Syagrus Mart. II. Fieldiana: Bot. 31: pp. 285-299. 1968b. Studies in the palm genus Syagrus Mart. Fieldiana: Bot. 31: pp. 363- 397. 1968c. Syagrus oleracea (Mart.) Becc. and closely related taxa. Fieldiana: Bot. 32: pp. 13-33. 240 FIELDIANA: BOTANY, VOLUME 32 1969. Studies in the palm genus Syagrus Mart. Fieldiana: Bot. 32: 77-1030. 1970. A conspectus of the palm genus Butia Becc. Fieldiana: Bot. 32: 127- 172. MARTIUS, C. F. P. VON 1818. Eschweg. Journ. Bras. 2: p. 178 (Langsdorffia). 1824. Palmarum familia ejusque genera denus illustrata. P. 18. Munich. 1826. Hist. Nat. Palm. 2: pp. 118, 130. 1844. Palmetum Orbignyanum. In d'Orbigny, Voyage dans L'Amerique meridionale 7: pp. 1-140. 1845. Hist. Nat. Palm. 3: p. 292. MOORE, H. E. 1963. Two new palms from Peru. Principes 7: pp. 107-115. TOLEDO, J. F. 1944. Estudos sobre algumas Palmeiras do Brasil. I. Um novo Genero da Tribu Cocoeae. Arq. Bot. S. Paulo II: pp. 3-9. WESSELS BOER, J. G. 1964. The indigenous palms of Suriname, pp. 170-172. Leiden. Publications 1101, 1102, 1103, 1104