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.
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240 FIELDIANA: BOTANY, VOLUME 32
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Publications 1101, 1102, 1103, 1104