BIOLOGY MR 1 9 C The person charging this material is re- sponsible for its return to the library from which it was withdrawn on or before the Latest Date stamped below. Theft, mutilation, and underlining of books are reasons for disciplinary action and may result in dismissal from the University. To renew call Telephone Center, 333-8400 UNIVERSITY OF ILLINOIS LIBRARY AT URBANA-CHAMPAIGN L161— O-1096 (J I FIELDIANA: BOTANY A Continuation of the BOTANICAL SERIES of FIELD MUSEUM OF NATURAL HISTORY VOLUME 31 The Library of the JAN 1 8 1979 FIELD MUSEUM OF NATURAL HISTORY CHICAGO, U. S. A. 520.5 -FB vi 31 TABLE OF CONTENTS PAGE 1. Two New Species of Palms from Nicaragua. By S. F. Glassman ... 1 2. Tropical American Plants, VI. By Louis 0. Williams 11 3. Agriculture, Tehuacan Valley. By C. Earle Smith, Jr 49 4. Flora, Tehuacan Valley. By C. Earle Smith, Jr 101 5. Preliminary Studies in the Palm Genus Syagrus Mart, and Its Allies. By S. F. Glassman 145 6. Tropical American Plants, VII. By Louis O. Williams 165 7. Supplement to Orchids of Guatemala. By Donovan S. Correll .... 175 8. Preliminary Notes on Scrophulariaceae of Peru. By Gabriel Edwin . . 223 9. New Species in the Palm Genus Syagrus Mart. By S. F. Glassman . . 233 10. Tropical American Plants, VIII. By Louis O. Williams 247 11. Notes on the Flora of Costa Rica, I. By William C. Burger 273 12. A New Eurystyles from Nicaragua. By Alfonso H. Heller 279 13. New Species in the Palm Genus Syagrus Mart. By S. F. Glassman . . 285 14. A Revision of the Family Geastraceae. By Patricio Ponce de Leon . . 303 15. Studies in American Plants. By Dorothy N. Gibson 353 16. Two New Nicaraguan Juglandaceae. By Antonio Molino R 357 17. Studies in the Palm Genus Syagrus Mart. By S. F. Glassman .... 363 18. Tropical American Plants, IX. By Louis O. Williams 401 STUDIES IN AMERICAN PLANTS DOROTHY N. GIBSON TWO NEW NICARAGUAN JUGLANDACEAE ANTONIO MOLINA R. STUDIES IN THE PALM GENUS SYAGRUS MART. S. F. GLASSMAN TROPICAL AMERICAN PLANTS, IX LOUIS 0. WILLIAMS FIELDIANA: BOTANY VOLUME 31 NUMBERS 15, 16, 17, 18 Published by FIELD MUSEUM OF NATURAL HISTORY MAY 23, 1968 tfufmtf if Hfims JULH1968 .. _ >ni BimtNi MATI STUDIES IN THE PALM GENUS SYAGRUS MART. SIDNEY F. CLASSMAN Research Associate, Palms FIELDIANA: BOTANY VOLUME 31 NUMBER 17 Published by FIELD MUSEUM OF NATURAL HISTORY MAY 23, 1968 PUBLICATION 1047 Library of Congress Catalog Card Number: 68-26375 PRINTED IN THE UNITED STATES OF AMERICA BY FIELD MUSEUM PRESS Studies in the Palm Genus Syagrus Mart. Since the publication of my paper dealing with preliminary studies of Syagrus (Glassman, 1965), I have accumulated additional infor- mation pertinent to the genus. This has resulted from a collecting trip to Brazil, a visit to several European herbaria, and an oppor- tunity to examine herbarium specimens not previously seen. Re- cently, I finished two articles in which eight new species of Syagrus were described (Glassman, 1967, 1968) . The present paper, however, deals primarily with nomenclatural changes and alignments, emendations of descriptions, and evaluation of type specimens of certain taxa. Several new combinations are made, two species are transferred to hybrid status, and several spe- cies are reduced to synonymy. This work has been supported by National Science Foundation grant no. GB 3737. Syagrus cocoides Martius, Palmet. Orbign. 134. 1844. Cocos weddellii Drude, Mart. Fl. Bras. 3: 411. 1881. C. drudei Becc., Mal- pighia 1 : 445. 1886. S. drudei (Becc.) Becc., L'Agric. Colon. 10: 466. 1916. Cocos weddellii was considered by Beccari to be a homonym when published in 1881 because Cocos weddelliana Wendl., an entirely dif- ferent species, was described previously in 1871. In 1886, Beccari gave the former taxon a new name, Cocos drudei; later he transferred the specific epithet from Cocos to Syagrus. Recently, I examined the holotype of Cocos weddelii (Weddell 2690, P) which consists of various leaf parts, spathe, and spadix, and male and female flowers. These specimens match the type ma- terial and other specimens of S. cocoides very closely, hence C. wed- dellii becomes a synonym of this species. S. cocoides is distributed in Brazil in the states of Pard, Amazonas, Maranhao, Bahia, Ceara, Piauhy, and Goyaz. S. hatschbachii Glassman, Fieldiana: Botj 31: 240. 1967. This species is based on Hatschbach 11668 (F, holotype) and Hatschbach 8091 (RB, paratype), both from the state of Parand Brazil. 363 364 FIELDIANA: BOTANY, VOLUME 31 After describing this species as new, I found a specimen labelled Butia microspadix Burret with the following data: BRAZIL: Rio Grande do Sul, det. M. Burret, Luederwaldt 12267 (SP). Even though this plant is immature, it shows a striking resemblance to S. hatschbachii. I then looked up the original article of Butia micro- spadix (Notizblatt 10: 1050. 1930) and found that Burret cited the following: BRAZIL: without locality, probably Sao Paulo state, Sellow s.n.; campos, without locality, J. Keller s.n. In addition to these, Burret listed another specimen "very near to this species," BRAZIL: Sao Paulo, Raiz da Serra, Luederwaldt— com. F. C. Hoehne 12267. I have not been able to locate the first two cited palms. They prob- ably were destroyed at the Berlin-Dahlem herbarium during World War II. The other specimen bears the same number as the one I saw (SP), but the locality data is different, i.e., one says state of Sao Paulo, the other says state of Rio Grande Do Sul. Luederwaldt 12267 (SP) is similar to the holotype and paratype of S. hatschbachii in that the spathe is covered with a brownish tomen- tum and the leaf has 20 pairs of pinnae. The leaf and floral parts, however, are obviously immature and hence it is risky to put a final determination on the specimen. The description of B. microspadix given by Burret is similar to that of S. hatschbachii only in the brownish tomentose spathes and the size of the male flowers. It differs from the latter species in the larger pinnae (37 cm. long and 0.7 cm. wide, rather than 27 cm. long and 0.4 cm. wide) and in the branched part of the spadix which is shorter (6 cm., rather than 9-13 cm.) and has more branches (25, rather than 13-15). Size of the female flowers is not mentioned in the description. Until I see other specimens of Butia microspadix, I am forced to designate it as species incerta and at the same time recognize S. hatschbachii as a valid species. Syagrus loefgrenii Glassman, Fieldiana: Bot. 31: 240. 1967 (corrected from lofgrenii). The above species is based on A. Lofgren 573, BRAZIL: Sao Paulo state, Rio Claro, campo, June 5, 1888 (SP) ; therefore, the spelling of the specific epithet should be changed from lofgrenii to loefgrenii. Since this species was described I have seen other specimens which can be included here. These were also collected in Sao Paulo FIG. 1. Syagrus campylospatha. Holotype of Cocos campylospatha. Hassler 1733 (G). LftHTftE PHRftGUftRiENSES KHK 365 366 FIELDIANA: BOTANY, VOLUME 31 state, but near Itirapina, about 40 km. N.W. of the type locality, in a burned-over savanna, July 2, 1965, Classman & Gomes 8011, 8012, 8013, 8015, 8016 (CHI). The plants cited above (1-4 specimens were collected for each number) grew in an area apparently subjected to periodic fires, so that measurements for certain organs may vary somewhat from the original description. The overall appearance and special character- istics, however, approach that of the holotype. The following de- scription of S. loefgrenii is emended to include the range of variation in this particular population. Acaulescent palm, 74-80 cm. tall. Petiole up to 22 cm. long and 6 mm. wide, sheathing base up to 22 cm. long; rachis of leaf blade 35-44 cm. long; pinnae 26-34 pairs, in loose clusters of 2-3 or single, middle ones 15-20 cm. long and 0.5-1.0 cm. wide, mostly with oblique tips; expanded part of spathe 11-18 cm. long and 2-4 cm. wide; branched part of spadix 8-12 (13) cm. long, branches 4-6 in number, each branch 6-7 (8) cm. long; male flowers 7-10 mm. long on lower part, 4-7 mm. long on upper part; female flowers mostly 10-14 mm. long (occasionally 8-9 mm. long) and 4-5 mm. wide. Fruit (immature) 1.7 cm. long and 0.9 cm. in diam- eter, beaked. Syagrus campylospatha (Barb. Rodr.) Becc., L'Agric. Colon. 10: 465. 1916. Figs. 1-2. Cocos campylospatha Barb. Rodr., Palm. Hassl. Nov. 9. 1900. ?C. acaulis ssp. glauca Drude ex Lindman, Bih. Sv. Vet. Akad. Handl. 26(3) : 23. 1900. Figure 3. ?C. apaensis Barb. Rodr., Bull. Herb. Boiss. 2(3) : 625. 1903. ?S. apaensis (Barb. Rodr.) Becc., L'Agric. Colon. 10: 465. 1916. After comparing type material of S. campylospatha and S. apaen- sis, I noticed a striking similarity in the indument of the pinnae. The upper surface is covered with a dense, appressed white-brownish pubescence while the lower surface is glaucous. Further investiga- tion of leaves, spadices, and flowers of other specimens identified as S. campylospatha reveals that S. apaensis falls within the range of variation of that species. Although the male flowers of the two taxa are about the same size, the sepals which are normally less than 2 mm. long, are very irregular in size (up to 3.5 mm. long) and shape in S. apaensis. This condition, however, may be teratological. Cocos acaulis ssp. glauca was originally considered to be synony- mous with Butia pungens Becc. of Argentina by Beccari (1916). Petioles of the latter taxon, however, are armed with short spines whereas C. acaulis ssp. glauca has unarmed petioles. Pinnae of the holotype shows a remarkable resemblance to those of S. campylo- spatha, but unfortunately there is no flower material for comparison. FIG. 2. S. campylospatha. Part of holotype of C. campylospatha. 367 .,.,» ia«rfm«*!ii;i !' n»"*f w txiw*«' ' r**1!!1' -• v%/«. 368 CLASSMAN: STUDIES IN PALM GENUS SYAGRUS MART 369 Therefore, I am provisionally including this subspecies as a synonym under S. campylospatha. The following is an emended description of S. campylospatha. Acaulescent palm. Patiole up to 19 cm. long and 0.7 cm. wide, sheathing base up to 20 cm. long; rachis of leaf up to 70 cm. long; pinnae up to 38 pairs, mostly in clusters of 2, middle ones up to 24 cm. long and 0.7-1.0 cm. wide, mostly with oblique tips, these sometimes bidentate, upper surface mostly appressed white- brownish pubescent, lower surface mostly glaucous; 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 branch up to 15 cm. long; male flowers 7-8 mm. long on lower part, 5-6 mm. long on upper part; female flowers 6-8 mm. long, 4-6 mm. wide; fruit (immature?) 2 cm. long and 1.2 cm. in diameter. Specimens examined : PARAGUAY: In campis Cordillera de Altos, Hassler 1733 (G, holotype of Cocos campylospatha; K, NY, P, G); Plaine de Pirayu, campos, June 1877, Balansa 2711 (P) ; Arroyo Tre- mentino, in region of Rio Apa, Hassler 7688 (G, holotype of Cocos apaensis); in campo aprico arenoso, "Yatai," Aug. 26, 1893, Regnell A. 1879 (S, holotype of Cocos acaulis ssp. glauca). Syagrus petraea (Martius) Becc., L'Agric. Colon. 10: 467. 1916. Cocos petraea Martius, Palmet. Orbign. 100, t. 9, fig. 2. 1844. In my 1965 article, I considered S. glazioviana (Dammer) Becc. as synonymous with S. petraea. After examining the holotype and other specimens of these two taxa, I concluded that they are distinct. The following key points out the main differences: Pinnae up to 6 mm. wide, mostly with acuminate tips; unclustered throughout; female flowers 10-14 mm. long, 4-5 mm. wide, mostly nerved, tips acuminate. S. petraea Pinnae up to 12 mm. wide, mostly with oblique tips, usually single, but in occa- sional clusters of 2; female flowers 10-13 mm. long, 5-8 mm. wide, mostly smooth in texture, tips acute S. glazioviana Cited specimens of S. petraea: BOLIVIA: Prov. Chiquitos, Sierra de Santiago, D'Orbigny 21 (P, holotype; M). Doubtful specimens: BRAZIL: Matto Grosso, Sept. 1839, Gardner 2979 (K); Piauhy, 1841, Gardner 2970 (K). In 1881, Drude cited a number of specimens under S. petraea and its varieties, but did not specify which specimens belong to which varieties. Gardner 2970 and 2979 (among several specimens cited in this grouping), resemble the holotype of S. petraea in the size, shape, FIG. 3. S. campylospathal Holotype of Cocos acaulis ssp. glauca. Regnell A, 1879 (S). 370 FIELDIANA: BOTANY, VOLUME 31 and texture of the female flowers, but the pinnae, although they are only 6-7 mm. wide, have oblique tips like S. glazioviana. In this case, it is difficult to come to any definite conclusions because so little material of S. petraea has been seen. Syagrus glazioviana (Dammer) Becc., L'Agric. Colon. 10: 466. 1916. Figure 4. Cocos glazioviana Dammer, Engl. Bot. Jahrb. 31 : 21. 1902. Cocos petraea var. genuina Drude, Mart. Fl. Bras. 3: 426. 1881. Cited specimens: BRAZIL: Goyaz, Morro do Frota, prope Meia Ponte, Sept. 1894, Glaziou 22251,. (C, lectotype; FI, G, K, P) ; Glaziou 22255 (G, K, P, US) ; Goyaz, Copelinhe de S. Antonio, Feb. 23, 1894, Glaziou 22256 (F, K, P); Goyaz, Amarelas, Anapolis, July 21, 1952, Macedo 3557 (S, SP); Goyaz, entre Patrocinho & Goyaz, Campos, 1844, Weddell 2536 (P, holotype of Cocos petraea var. genuina Drude). I have designated Glaziou 22254 from Copenhagen as the lecto- type of C. glazioviana because it is the most complete collection (leaf, spathe and spadix, flowers and fruit). There are four sheets from Paris with this number, but two of these sheets have branched spa- dices. All of the other isotypes listed here, including the lectotype, have simple spadices. Therefore, I am excluding the two sheets with branched spadices from the type material. Glaziou 22253 is the type of Cocos graminifolia var. glazioviana Dammer (which has branched spadices), so it is very likely that some of this material was inad- vertently mixed with Glaziou 22254- Nevertheless, in his description of C. glazioviana, Dammer states that the spadix has 3-4 branches. The following is an emended description of S. glazioviana. Besides correcting certain errors, it also includes additional information not in the original description. Acaulescent palm. Petiole up to 5 cm. long, sheathing base about 10 cm. long; rachis of leaf up to 55 cm. long; pinnae up to 13 pairs, usually unclustered but occa- sionally in two's, mostly irregularly distributed on rachis, middle ones up to 38 cm. long and 8-12 mm. wide, mostly with oblique tips; expanded part of spathe up to 18 cm. long and 2-3 em. wide; spadix simple, up to 12 cm. long; male flowers 8- 14 mm. long on upper part, 15-20 mm. long on lower part; female flowers 10-13 mm. long, 5-8 mm. wide; fruit (mature?) 2.8 cm. long, 1.7 cm. in diameter, slightly beaked, endocarp bony, 0.5-1 mm. thick, cavity smooth, trivittate; seed shrivel- led, 1.4 cm. long, 1.1 cm. in diameter. Syagrus glazioviana var. alpina (Drude) Glassman, comb. nov. Cocos petraea var. alpina Drude, Mart. Fl. Bras. 3: 426. 1881. S. pe- traea var. alpina (Drude) Becc., L'Agric. Colon. 10: 467. 1916. FIG. 4. Syagrus glazioviana. Holotype of Cocos glazioviana. Glaziou 2225 1+ (C). 371 372 FIELDIANA: BOTANY, VOLUME 31 Cited specimens: BRAZIL: Goyaz, Sertao Amaroleite, Sept. -Oct. 1844, Weddell 2775 (P, holotype). Doubtful specimens: Goyaz, be- tween Cabeceira do Rio Sambambaia & Ponzo do Barbatimao, Sept. 19, 1894, Glaziou 22257 (£, FI, G, K, NY, P). The main reason for transferring var. alpina to S. glazioviana is that the female flowers are similar in texture and shape. The two taxa differ in the following: Female flowers 7-9 mm. long, male flowers up to 10 mm. long, fruit 1.7 cm. long, 1.3 cm. in diameter, pinnae with acuminate or oblique tips. S. glazioviana var. alpina Female flowers 10-13 mm. long, male flowers up to 20 mm. long, fruit 2.8 cm. long, 1.7 cm. in diameter, pinnae mostly with oblique tips. S. glazioviana var. glazioviana The holotype material of var. alpina consists of three sheets. On two of the sheets, the pinnae are only 5 mm. wide and have acumi- nate tips, but the pinnae are up to 10 mm. wide with oblique tips on the third sheet. On all three sheets, however, the female flowers are about the same size, shape, and texture. I have included Glaziou 22257 here tentatively because the female flowers are 7-10 mm. long, but the pinnae are consistently 9-11 mm. wide and mostly with oblique tips. Unfortunately, I cannot account for the differences between the leaves of the three holotype sheets, unless the specimens with the narrow, acuminate pinnae are immature or underdeveloped. SYAGRUS PETRAEA var. PLATYPHYLLA (Drude) Becc., L'Agric. Colon. 10: 467. 1916. varietas confusa. Cocos petraea var. platy- phylla Drude, Mart. Fl. Bras. 3: 426. 1881. BRAZIL: Between Goyaz & Cuyaba, 1844, Weddell 2952 (P, holo- type). Holotype material consists of two sheets; both have leaves, but only one sheet has floral parts. The pinnae are up to 9 mm. wide with oblique tips and resemble the leaves of S. glazioviana, but they are not of the same texture. The spathe is shallowly grooved as in the genus Butia, the spadix is simple, and both the male and female flowers appear to be immature. Weddell 2952 does not fall into any recognizable species of Syagrus; but since the flowers are immature, and the leaves and floral parts may have come from different plants, I am designating var. platyphylla as varietas confusa. FIG. 5. Syagrus graminifolia. Holotype of Cocos graminifolia. Burchell 5956 (K). 373 374 FIELDIANA: BOTANY, VOLUME 31 Weddell 2795 (P) was cited by Drude (1881) under varieties of Cocos petraea without specifically designating to which variety it be- longed. This number comprises two sheets. Both have leaves that resemble S. glazioviana, but one sheet has a simple spadix and the other a branched spadix. The former is probably S. glazioviana whereas the latter may be S. graminifolia. The type collection of Cocos graminifolia var. nana Drude is Weddell 2995, so it is conceiv- able that some of the specimens of this number could have been mixed with Weddell 2795. Cocos RUPESTRIS Barb. Rodr., Prot. App. 45, 1879; Les Palm. 29, 1882. Nomen incertum. Barbosa Rodrigues gave the following locality data for this name. BRAZIL: Minas Gerais, ad margines Rio Sapucahy, montibus Serra da Tromba; inter saxalapideus, inter gramineas. Drude (1881) lists this as a synonym of Cocos petraea Martius. No specimens are cited and there are no illustrations published in either article. The de- scription is too brief to determine where it might be placed, therefore I have designated Cocos ruprestris as nomen incertum. Syagrus graminifolia (Drude) Becc., L'Agric. Colon. 10: 466. 1916. Figures 5-6, 18. Cocos graminifolia Drude, Mart. Fl. Bras. 3: 415, t. 91, 1881. S. graminifolia var. glazioviana (Dammer) Becc., L'Agric. Colon. 10 : 466. 1916. C. graminifolia var. glazioviana Dam- mer, Engl. Bot. Jahrb. 31 : 23. 1902. Cited specimens: BRAZIL: Piauhy near Paranahyba R., between S. Domingos and Borda do Mato, Burchell 5956 (K, holotype; P); Goyaz, Catalao, 1834, /. Riedel and B. Luschnatt s.n. (LE, paratype of Cocos graminifolia); Goyaz, Chapado de Catalao, cerrados da regiao do Paranaiba, July, 1892, Uhle 794 (R) ; Brazil, without local- ity, Glaziou 22252 (C, PI, G, K, P) ; Goyaz, inter Lagoa do Piquiero and Buraco da Onca in campis, Sept., 1895, Glaziou 22253 (C, holo- type of Cocos graminifolia var. glazioviana; F, FI, G, K, P). Both the holotype and isotype of C. graminifolia have fruits with one or two chambers (locules), but no seeds are present. The parti- tion separating the locules is more or less firm. Fruits of the genus Syagrus are typically one-chambered. The genus Butia Beccari (1916) characteristically has a 3-seeded fruit (or is 1-2-seeded by abortion), but the petioles are conspicuously spiny on the margins and the exterior of the spathes is not grooved (plicate-sulcate) . S. graminifolia however, has unarmed petioles and grooved spathes. In the genus Syagrus the spathes are always grooved, but sometimes FIG. 6. S. graminifolia. Isotype of Cocos graminifolia var. glazioviana. Glaziou 22253. 375 376 FIELDIANA: BOTANY, VOLUME 31 the grooves are obscured by a waxy coating or by a tomentum. Some species may have armed petioles (e.g., S. coronata (Martius) Becc., S. vagans (Bondar) Hawkes, S. camposportoana (Bondar) Glassman, S. x matafome (Bondar) Glassman), but none of these has more than one-chambered fruits. The two-chambered fruit is also characteristic of several other species of Syagrus (S. archeri Glassman, S. campicola (Barb. Rodr.) Becc., and S. leptospatha Burret) which will be discussed in another part of this article. Perhaps these species should be placed in a sep- arate section of the genus, but S. graminifolia and S. archeri have branched spadices whereas the other two species (S. campicola and S. leptospatha) have simple spadices. In his original description of Cocos graminifolia var. glazioviana, Dammer stated that the spadices were not branched. All of the type specimens cited above (Glaziou 22253) have spadices with 3-5 branches, so Dammer was in error. As previously suggested, type material of Cocos glazioviana (Glaziou 22254) was probably mixed up with this number. To incorporate additional information I am emending the orig- inal description of S. graminifolia as follows: Acaulescent palm. Petiole up to 15 cm. long, 4 mm. wide, sheathing base up to 10 cm. long; rachis of leaf up to 70 cm. long; pinnae mostly unclustered, middle ones 35-40 (55) cm. long and 4-7 mm. wide, mostly with long acuminate tips; ex- panded part of spathe up to 30 (39) cm. long and 2 cm. wide; branched part of spadix up to 30 cm. long, branches 4-7 in number, each branch up to 18 cm. long; male flowers 4-7 mm. long on upper part, 8-10 mm. long on lower part; female flowers mostly 10-13 mm. long, 4-7 mm. wide; fruit (mature) up to 2 cm. long and 1.3 cm. in diameter, slightly beaked, endocarp about 1 mm. thick, cavity with 1-2 locules, separated by a firm partition (when 2 locules present); seeds not seen. Syagrus graminifolia var. nana (Drude) Becc., L'Agric. Colon. 10: 466. 1916. Cocos graminifolia var. nana Drude, Mart. Fl. Bras. 3:415, t. 91. 1881. Cited specimens: BRAZIL: entre Goyaz & Cuyaba, 1844, Weddett, 2995 (P, holotype; F); Goyaz, campos entre Patrocinho & Goyaz 1844, Weddell 2541 (P, paratype). At present, I am not certain whether var. nana is distinct from var. graminifolia because so few specimens have been seen. Until I see further material of these two taxa, I am separating them by the following: Female flowers 7-10 mm. long, male flowers up to 7 mm. long; spathes 0.8-1.2 cm. wide, spadix branches 2-4 in number var. nana CLASSMAN: STUDIES IN PALM GENUS SYAGRUS MART 377 Female flowers 10-13 mm. long, male flowers up to 10 mm. long, spathes 1-2 cm. wide, spadix branches 4-7 in number ... . var. graminifolia Syagrus archeri Glassman, Fieldiana: Bot. 31 : 235. 1967. Fig- ure 17. Since the publication of this taxon I have examined a number of additional specimens. BRAZIL: Minas Gerais, near Lavras, burned- over pasture, Sept. 20, 1936, Archer WW (A, holotype; BH, MO, NY, US) ; 16 km. N. of Lavras, campo natural, July 9, 1965, Glass- man & Gomes 8018, 8019, 8020, 8021, 8022, 8023, 8024- (CHI). We collected other specimens of this species near Paraopeba, Cur- velo and Diamantina state of Minas Gerais, but these plants were lost in transit. The specimens collected 16 km. north of Lavras re- vealed some interesting features not mentioned in the original article. Many of the fruits are two-chambered with flattened seeds, whereas others have a single chamber with globose seeds. Apparently, the two-seeded fruits are the fully mature ones. Many of the specimens collected also showed a wide range of variation in measurements, therefore I am emending the original description to include this new information. Acaulescent palm, or with very short trunk. Petiole up to 8 cm. long and 1 cm. wide, sheathing base up to 15 cm. long; rachis of leaf up to 69 cm. long; pinnae up to 28 pairs, unclustered, glabrous on both surfaces, middle ones up to 41 cm. long and 1 cm. wide, mostly with acuminate tips; expanded part of spathe up to 30 cm. long and 3 cm. wide, glaucous on outside, becoming eglaucous with age; branched part of spadix up to 20 cm. long (23 cm.— fruiting), branches up to 19 in number (31 branches— fruiting), each branch up to 10 cm. long; male flowers 3-5 mm. long on upper part, 6-7 mm. long on lower part; female flowers 5-7 mm. long, 4-6 mm. wide; fruit (mature) 1.8-1.9 cm. long, 1.2-1.4 cm. in diameter, short beaked, endocarp 1-1.5 mm. thick, cavity with 1-2 locules separated by firm parti- tions; seed (when single), irregularly globose, 7.5-9.0 mm. long and 7-9 mm. in diameter, cavity 2-2.5 mm. in diameter; seed (when two are present) flattened laterally, 10 mm. long, 6 mm. in diameter. Syagrus campicola (Barb. Rodr.) Becc., L'Agric. Colon. 10: 465. 1916. Figures 7-8, 19. Cocos campicola Barb. Rodr., Palm. Hassl. Nov. 6, 1900; Sert. Palm. Bras. l:t. 71. 1903. Cited specimens: PARAGUAY: In campis Ip£ hu, Sierra de Mara- cayu, Hassler 5057 (G, holotype; BM); in Valle fluminis Y-aca in arenosis pr. Piribebuy, Hassler 6871 (G). According to Barbosa Rodrigues (1903), t. 71 shows a single- chambered fruit with a trivittate cavity. Hassler 6871 has one fruit 2 cm. long, 1.1 cm. in diameter with a beak 5 mm. long, but there are Dr. L HASSkER, Plantae Para^uarienses. 1900 Ar<)ui. o mttrctt d.m Hlcrt. Geocce, i«S* FIG. 7. Syagrus campicola. Hassler 6871 (G). 378 FIG. 8. Syagrus campicola. Hassler 6871 (G). 379 380 FIELDIANA: BOTANY, VOLUME 31 two distinct chambers. The petioles are unarmed and the spathes are definitely grooved, so this species does not fall into the genus Butia. It appears to be more closely related to the following species. Syagrus leptospatha Burret, Notizbl. Konigl. Bot. Gart. Mus. Berlin 15: 105. 1940. Figures 9 and 20. Cited specimens: BRAZIL: Mattogrosso, Boliche Seco, Campo Grande, Sept. 6, 1936, Archer & Gehrt 36429 (SP, holotype; US). Type specimens contain several fruits which are either single- or double-chambered, mostly with seeds. Although the spathes are papery, they are still grooved on the lower part, and the petioles are unarmed. This taxon most closely resembles S. campicola in being acaulescent, having unclustered pinnae which are long and narrow, simple spadices which extend beyond the tips of the spathes for sev- eral centimeters, and two-chambered fruits. The two taxa differ, however, in the following: Spathes more or less woody in texture, inflated part 10-20 cm. long, 1.0-1.3 cm. wide, spadix (excluding peduncle) 10-12 cm. long, male flowers 5-10 mm. long; fruit ovate, up to 2 cm. long and 1 cm. in diameter S. campicola Spathes papery in texture, inflated part 4-5 cm. long, 0.5-1.0 cm. wide, spadix 3-6 cm. long, male flowers 3-5 mm. long; fruit rounded, 1.0-1.4 cm. long, 0.8-1.2 cm. in diameter S. leptospatha Syagrus aff. coronata (Martius) Becc. L'Agric. Colon. 10: 466. 1916. Cocos botryophora var. ensifolia Drude, Mart. Fl. Bras. 3: 409. 1881. Arecastrum romanzoffianum var. ensifolium (Drude) Becc. L'Agric. Colon. 10: 459. 1916. There are enough authentic specimens of Cocos botryophora var. ensifolia (Glaziou 8058— P, holotype; C, F, FI, G, LE, R) available for study to definitely state that it does not belong to Syagrus botryo- phora (Martius) Martius. S. botryophora has unclustered pinnae and female flowers 5-6 mm. long and 6-7 mm. wide, whereas var. ensi- folia has pinnae in clusters of 3-4 and female flowers 9-11 mm. long and 7-8 mm. wide. In a previous paper (Glassman, 1965), I stated that I had not seen sufficient type material of Cocos botryophora Martius to pin it down definitely. Since then, I have seen additional leaf material from an oversized sheet (Martius s.n., M, holotype) which shows unclustered pinnae. In 1916, Beccari transferred the varietal epithet, ensifolia to Are- castrum romanzoffianum. The latter species has female flowers 4.5- 6 mm. long and 4-6 mm. wide and clustered pinnae, which are mostly 2-3 cm. wide, rather than 3-4 cm. wide as in the type material of var. ensifolia Drude. FIG. 9. Syagrus leplospatha. Holotype. Arc/zer c£ Geftrt 55^^S (SP). 381 382 FIELDIANA: BOTANY, VOLUME 31 Unfortunately, specimens of Glaziou 8058 lack certain diagnostic structures to conclusively place them under S. coronata. For in- stance, there are no petioles (in S. coronata they are armed with flat spine-like margins), no fruits, and the spadices are incomplete. The size and shape of the female flowers, the width and clustering of the pinnae, and the length of the individual rachillae, however, compare very favorably with specimens of S. coronata. Syagrus romanzoffiana (Chamisso) Glassman, comb. nov. Fig- ures 10 and 14. Cocos romanzoffiana Chamisso, Choris Voyage Pitt. 5-6, t. 5-6. 1822. Arecastrum romanzoffianum (Chamisso) Becc., L'Agric. Colon. 10: 447. 1916. A. romanzoffianum var. genuinum Becc., I.e. 455. t. 1-2, 3, fig. 7. 1916. A. romanzoffianum var. genui- num minus Becc., I.e. 456, t. 3, fig. 8. 1916. A. romanzoffianum var. australe (Martius) Becc., I.e. 459, t. 3, figs. 2-6. 1916. C. australis Martius, Palmet. Orbign. 95, t. 1, fig. 2, t. 30C. 1844. A. romanzof- fianum var. micropindo Becc., I.e. 462, t. 3, fig. 1. 1916. C. arechava- letana Barb. Rodr., Contr. Jard. Bot. Rio de Jan. 2: 43. 1901. C. acrocomioides Drude, Mart. Fl. Bras. 3: 409, t. 87, fig. 3. 1881. 1C. martiana Drude & Glaziou ex Drude, Mart. FL Bras. 3: 418, t. 88-89. 1881. C. plumosa Hooker, Bot. Mag. 86: t. 5180. 1860. C. geriba Barb. Rodr., Prot. App. 43. 1879. C. datil Griseb. & Drude, Symbol. Flor. Argent. 283. 1879. In a recent article (Glassman, 1965), I questioned the wisdom of segregating genera in the Syagrus alliance on the basis of fruit and seed characters alone. The genus Arecastrum Becc. is based on the very irregular endocarp cavity and irregular, gibbous-uncinate seed. It is true that these are distinctive characteristics, but the fruits and seeds within the genus Syagrus are quite variable. S. inajai (Spruce) Becc. has a fruit with an irregularly triangular endocarp cavity and seed three lobed in cross-section; and four species, S. archeri Glass- man, S. graminifolia (Drude) Becc., S. campicola (Barb. Rodr.) Becc., and S. leptospatha Burret, have fruits which are 1-2-chambered with as many seeds. If the fruits and seeds are used as a basis for splitting up Syagrus into separate genera, then other characteristics also could be used. About two-thirds of the species have clustered pinnae, the other third unclustered pinnae. Unclustered pinnae is one of the characteristics of the genus Butia Becc. Several species, e.g., S. vagans (Bondar) Hawkes, S. camposportoana (Bondar) Glassman, FIG. 10. Syagrus romanzoffiana. Minas Gerais, Fazenda Pecegueiro, North of Braganga. Photo by Glassman. 383 384 FIELDIANA: BOTANY, VOLUME 31 S. coronata (Martius) Becc., S. tostana (Bondar) Glassman and Ari- kuryroba schizophylla (Martius) Bailey, have petioles with spiny or spine-like margins which is also characteristic of the genus Butia. A number of species are acaulescent, but a majority of the taxa have definite trunks. Most of the species have branched spadices, but some, e.g., S. campicola, S. leptospatha, S. petraea, S. glazioviana, and S. acaulis (Drude) Becc. have simple, unbranched spadices. The acaulescent habit and unbranched spadices are also found in the genus Allagoptera Nees. At the present time, a more reasonable approach would be to divide Syagrus into various subgenera and sections and use several characteristics to tie them all together. This has been done to a limited extent by others, especially Drude (1881); however, I plan to make further changes and subdivisions before I complete my re- vision of the genus Syagrus. In Brazil, there is good evidence to show that S. romanzoffiana hybridizes with S. coronata in the state of Bahia and with S. oleracea (Martius) Becc. in the state of Sao Paulo. In addition to this, hy- brids between S. romanzoffiana and various species of Butia have been reported by Barbosa Rodrigues (1903), Beccari (1916), Bailey (1936), and Bondar (1964). Division of S. romanzoffiana into varieties by Beccari (1916) is not very satisfactory. Even though this species has a wide range of distribution (northern Brazil to Uruguay), differences in size and shape of the fruit and size of the tree is too variable to consider clear- cut varieties. In general, fruits are 2.0-2.6 cm. long and 1.2-1.7 cm. in diameter, but many of the fruits collected from cultivated plants are longer (up to 3.0 cm. long) and broader (up to 2.4 cm. in diameter). A. romanzoffianum var. botryophorum (Martius) Becc. has been transferred to Syragrus botryophora (Glassman, 1965) and var. ensi- folium has been placed provisionally under S. coronata, as previously mentioned. The holotype of Cocos plumosa (Kew Gardens 22-44164, K) has fruits 2.7 cm. X 2.1 cm., but the larger size may be due to cultiva- tion. Some specimens in the type collection of Cocos martiana (Gla- ziou 8056, cult. Rio de Janeiro, G) have female flowers 6-7 mm. long and pinnae up to 3.8 cm. wide, both larger than average. Illustra- tions of fruits by Drude (1881, t. 89) resemble S. romanzoffiana closely. In their description, Drude and Glaziou state that the fruits are 2.5-3.0 cm. X 1.5 cm., and the tree is up to 20 m. tall. Here again, cultivation may account for the larger than normal size. £ O .3" •S m 385 386 FIELDIANA: BOTANY, VOLUME 31 Other synonyms given above, C. arechevaletana, C. acrocomioides, C. geriba, and C. datil, seem to fit the general description of S. ro- manzoffiana by their type specimens or published descriptions. The following is an emended description of S. romanzoffiana. It should be noted that all of the dimensions are frequently larger in cultivated specimens. Palm up to 15 m. tall, trunk up to 40 cm. or more in diameter. Naked petiole up to 45 cm. long and 4 cm. wide, sheathing base 50-84 cm. long; rachis of leaf up to 320 cm. long, pinnae up to 150 pairs, mostly in clusters of 2-5, middle ones up to 85 cm. long, about 2-3 cm. wide, mostly with acuminate tips; expanded part of spathe up to 150 cm. long and 14 cm. wide; branched part of spadix up to 125 cm. long, branches up to 80 or more in number, each branch up to 62 cm. long; male flowers 7-10 mm. long on upper part, 11-16 mm. long on lower part; female flowers averaging 4.5-6.0 mm. long and 4-6 mm. wide; fruit 2.0-2.6 cm. long and 1.2- 1.7 cm. in diameter, slightly beaked, endocarp up to 0.5 cm. thick, very irregular in shape; seed gibbous-uncinate, 0.8-1.2 cm. long and 0.6 cm. in diameter, cavity very small. Syagrus schizophylla (Martius) Glassman, comb. nov. Fig- ure 11. Cocos schizophylla Martius, Fl. Bras. 2: 119. t. 85. 1826. Arikuryroba capanemae (Barb. Rodr., Plant. Nov. Cult. 1: 6. 1891. C. capanemae (Barb. Rodr.) Drude in Engler & Prantl, Pflanzenfam. Nachtr. 1 : 57. 1897. C. arikuryroba Barb. Rodr., Palm. Mattogross. Nov. 25. 1899. Arikury schizophylla (Martius) Becc., L'Agric. Colon. 10: 445. 1916. ("Aricuri")- Arikuryroba schizophylla (Martius) Bailey, Gentes Herb. 2: 196. 1930. In a key to the Syagrus alliance, Moore (1963) separated Arikury- roba Barb. Rodr. from the other genera as having spinose-dentate petioles, pinnae with oblique or toothed tips, and seeds with ruminate endosperm. As previously mentioned, several species of Syagrus have spiny petioles; and pinnae with oblique or toothed tips are found in a number of members of Syagrus, e.g., S. campylospatha (Barb. Rodr.) Becc., S. vagans (Bondar) Hawkes, S. loefgrenii Glassman, S. glazioviana (Dammer) Becc., S. glaucescens (Glaziou) Becc., S. rachidii Glassman, and S. duartei Glassman. S. schizophylla differs from all other species of Syagrus mainly in the ruminate endosperm of the seed. It is probably most closely related to S. vagans (fig. 12), an acaulescent palm, because both have mostly unclustered pinnae with oblique tips, spiny petioles, spadix and spadix branches and male and female flowers about the same size, FIG. 12. Syagrus vagans. Dense stand in foreground. Brazil, Bahia, Santa Teresinha. Photo by Bondar. 387 FIG. 13. Syagrus x camposportoana. Brazil, Bahia, Pogoes. After Bondar (1964). 388 GLASSMAN: STUDIES IN PALM GENUS SYAGRUS MART 389 and fruits similar in size and shape. Although S. schizophylla is usu- ally a tree 2-4 m. tall, it is sometimes acaulescent (Bondar, 1964). FIG. 14. Top row: longitudinal and cross-sections of fruits of Syagrus roman- zoffiana (\eft)—Dahlgren 610582 (F) and right-— Bondar 619773 (F). Middle and bottom rows: longitudinal and cross-sections of fruit with seed enclosed (left), with seed removed (middle), and external views of seeds (right) of S. x camposportoana —Bondar 619765 (F). This species apparently crosses with S. coronata to produce S. tostana, which will be discussed later. For the above reasons, S. schizophylla seems to be closely allied to species of Syagrus and therefore the ruminate endosperm alone is insufficient grounds for keeping it in a separate genus. FIG. 15. Syagrus coronate. Brazil, Bahia, near Salvador. Photo by Bondar. 390 FIG. 16. Syagrus x tostana. Brazil, Bahia, outskirts of Salvador. Photo by Bondar. 391 392 FIELDIANA: BOTANY, VOLUME 31 Syagrus x camposportoana (Bondar) Glassman, stat. nov. Figures 13-14. Cocos camposportoana Bondar, Field Mus. Nat. Hist. Bot. 22: 460. 1942. Arecastrum camposportoanum (Bondar) Hawkes, Arq. Bot. S. Paulo II : 175. 1952. S. camposportoana (Bondar) Glass- man, Rhodora 65: 260. 1963. Bondar (1964) considered this taxon as a natural hybrid between Cocos romanzoffiana and C. coronata, but he did not formally change its status. It is known only from one locality. BRAZIL: Bahia, Fa- zenda Gangugy, municipio Pocoes, planted from seedlings coming from Itabuna, 1940, Bondar 619765 (F, lectotype); 610582, 619766, 61 9767 '(F). S. romanzoffiana S. x camposportoana S. coronata Grouping of leaves on trunk No special arrangement Grouped into spiral rows Grouped into spiral rows Petiole margins Mostly smooth Armed with long, flat, spine-like projections Armed with long, flat, spine-like projections Rachis length Up to 320 cm. Up to 300 cm. (fide Bondar) Up to 216 cm. No. of pinnae per leaf Up to 150 pairs 130 pairs (incomplete) Up to 85 pairs Average number of pinnae per cluster 2-5 3-5 3-4 Middle pinnae width 2-3 cm. 3-4 cm. 3.0-3.5(4) cm. Middle pinnae length Up to 85 cm. Up to 105cm. 66 cm. Spadix branched part Up to 125 cm. long Up to 120 cm. long Up to 51 cm. long Length of each branch Up to 62 cm. Up to 57 cm. 29cm. Length of male flowers 7-10 mm. and 11-16 mm. 10-11 mm. and 13-15 mm. 7-11 mm. and 12-16 mm. Length and width of female flowers 4.5-6 X 4-6 mm. 6-7 X 4-5 mm. 9-13 X 5-7 mm. Fruit length and diameter 2.0-2.6 X 1.2-1.7 cm. 3 X 2 cm. Up to 2.6 X 1.8-2.0 cm. Endocarp cavity Very irregular in outline, grooved within Less irregular in outline, grooved within Trivittate, smooth within, slightly truncate at one end Seed length and diameter 0.8-1.2 X 0.6cm. 1.1 X 1.0cm. 1.4-1.6 X 1cm. Seed shape Gibbous-uncinate Gibbous-uncinate Ovate Seed cavity Very small 4-5 mm.[in diameter 4-5 mm/in diameter GLASSMAN: STUDIES IN PALM GENUS SYAGRUS MART 393 The foregoing table shows the relationship between the three taxa involved. Inspection of this chart shows that S. x camposportoana has the following characters in common with S. romanzoffiana: rachis, spadix, spadix branches, and female flowers are about the same length, and both have gibbous-imcinate seeds; and it is similar to S. coronata (fig. 15) in the grouping of leaves on trunk, the flat, spine- like petiole margins, the width of the pinnae and the relatively large seed cavity. It is intermediate between the two species in the shape of endocarp cavity; and its dimensions are larger than either species in the longer pinnae and slightly longer fruit. In some cultivated specimens of S. romanzoffiana, however, the fruits may be up to 3 cm. long. Syagrus x tostana (Bondar) Glassman, stat. nov. Figure 16. Cocos tostana Bondar, Field Mus. Nat. Hist. Bot. 22: 458. 1942. Arikuryroba tostana (Bondar) Hawkes, Arq. Bot. S. Paulo II: 175. 1952. S. tostana (Bondar) Glassman, Rhodora 65: 261. 1963. Bondar (1964) said that this taxon is undoubtedly a spontaneous hybrid between C. coronata and C. schizophylla, but he made no formal transfer at the time. All three taxa occur together in the vicinity of Salvador, in the state of Bahia, Brazil. S. x tostana is known only from the type locality: Pituba, Salvador, 1940, Bondar 619775 (F, lectotype); 619735, 619776(F). The following chart shows similarities and differences between the three taxa. S. coronata S. x tostana S. schizophylla Grouping of Grouped into No special No special arrange- leaves on trunk 5 spiral rows arrangement ment Petiole margins Armed with long, Armed with long, Armed with short, flat, spine-like spine-like curved spines for projections for projections for most of length most of length more than Yi its length, dentate beyond this point and gradually becoming spineless for last 10cm. of length Rachis length Up to 2 16 cm. Up to 214 cm. Up to 145 cm. No. of pinnae Up to 85 pairs Up to 65 pairs Up to 41 pairs per leaf No. of pinnae 3-4 Unclustered Unclustered per cluster Tips of pinnae Mostly acuminate Mostly acuminate Mostly oblique FIG. 17. Syagrus archeri. Longitudinal section of single-chambered fruit (upper left), and longitudinal and cross-sections of double-chambered fruit (lower left and center left). Glassman & Gomes 8025 (CHI). Longitudinal and cross-sections of single-chambered fruits and seeds (lower right and upper right). Glassman & Gomes 8021 (CHI). 394 GLASSMAN: STUDIES IN PALM GENUS SYAGRUS MART 395 f FIG. 18. Syagrus graminifolia. Cross-sections of double and single-chambered fruits. Burchell 5956 (K) Chart showing similarities and differences between the three taxa (continued}. S. coronata S. x tostana S. schizophylla Middle pinnae width 3.0-3.5(4) cm. Up to 4 cm. 3.0-3. 5 cm. Middle pinnae length Up to 66 cm. Up to 72 cm. Up to 59 cm. Spadix branched part Up to 51 cm. Up to 73 cm. Up to 60 cm. long Length of each branch Up to 29 cm. Up to 45 cm. Up to 28 cm. Length of male flowers 7-11 mm. and 12-16 mm. 8-11 mm. and 11-14 mm. 3-5 mm. and 5-7 mm. Length and width 9-13x5-7 mm. 10-11.5 X 6-7 mm. 5-8 X 4-6 mm. of female flowers Fruit length and Up to 2.6 X 1.8- Up to 4 X 2.3 cm. Up to 3 X 2.5 cm. diameter 2.0 cm. Seed length and 1.4-1.6 X 1 cm. 2.1 X 1.3 cm. diameter Endosperm Homogeneous Homogeneous 1.7 X 1.4cm. Ruminate Analysis of the above chart shows that S. x tostana is similar to S. coronata in the rachis being about the same size, acuminate pinnae, size of male and female flowers, and homogeneous endosperm of seed, FIG. 19. Syagrus campicola. Cross-sections of double-chambered fruits show- ing one seed in each. Hassler 6871 (G) FIG. 20. Syagrus leptospatha. Cross-section of single-chambered fruit with seed (upper row), and cross-sections of double-chambered fruits with seeds (lower row). Archer & Gehrt 36^29 (SP) 396 GLASSMAN: STUDIES IN PALM GENUS SYAGRUS MART 397 and is similar to S. schizophylla in the grouping of the leaves on the trunk and the mostly unclustered pinnae. It is intermediate be- tween the two species in the arrangement of the spines on the petiole, and the number of pinnae per leaf; and its measurements are larger than either species in the slightly longer pinnae, longer spadix and spadix branches, and longer fruits and seeds. REFERENCES BAILEY, L. H. 1936. The genus Butia. Gentes Herb., 4: 49-50. BARBOSA RODRIGUES, J. 1903. Sertum Palmarum Brasiliensium ou Relacion des Palmiers Nouveaux du Bresil, Decouverts, Decrits et Dessines d'apres Nature, 1: 116. BECCARI, O. 1916. II Genere Cocos Linn, e le Palmae Affine. L'Agricoltura Coloniale 10: 435-471; 489-532; 585-623. BONDAR, G. 1964. Palmeiras do Brasil. 58-89, figs. 7-30. Institute de Botanica. Sao Paulo. DRUDE. 1881. Martius Flora Brasiliensis 3: 398-428. GLASSMAN, S. F. 1965. Preliminary studies in the palm genus Syagrus Mart, and its allies. Fieldiana: Bot., 31 : 147-164, figs. 44-50. 1967. New species in the palm genus Syagrus Mart. Fieldiana: Bot., 31: 235- 245, figs. 1-7. 1968. New species in the palm genus Syagrus Mart. II. Fieldiana: Bot., 31: 285-299, figs. 1-15. MOORE, H. E. 1963. Two new palms from Peru. Principes, 7: 107-115. Publications 1045, 1046, 1047, and 1048 UNO, UNIVERSITY OF ILLINOIS-URBANA