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FLORA OF PERU 

PART I, NO. 2 
BY 

J. FRANCIS MACBRIDE 

CURATOR, PERUVIAN BOTANY 




BOTANICAL SERIES 

FIELD MUSEUM OF NATURAL HISTORY 

VOLUME XIII, PART I, NUMBER 2 

AUGUST 17. 1960 



E UKttn OF THE 

\UG 2 9 I960 PUBLICATION 895 

OF ILLINOIS 



FLORA OF PERU 

PART I, NO. 2 
BY 

J. FRANCIS MACBRIDE 

CURATOR, PERUVIAN BOTANY 




BOTANICAL SERIES 

FIELD MUSEUM OF NATURAL HISTORY 

VOLUME XIII, PART I, NUMBER 2 

AUGUST 17, 1960 

PUBLICATION 895 



Library of Congress Catalog Card Number: 36-101+26 



PRINTED IN THE UNITED STATES OF AMERICA 
BY CHICAGO NATURAL HISTORY MUSEUM PRESS 



& 



2- 

FLORA OF PERU 



J. FRANCIS MACBRIDE 



16. PALMAE. Palm Family. 

Ligneous, smooth or more or less rudely spiny or prickly, acaules- 
cent or caulescent, sometimes scandent or caudices low or tall, clus- 
tered or more often solitary, slender or columnar, nearly always strict, 
not rarely magnificent trees conveying the impression of great nobil- 
ity, each crowned as it were with a beautifully proportioned cluster 
of several to many usually palmate or pinnate frond-like leaves, these 
again, especially among low or smaller species, variously divided or 
even entire, often few and gracefully ornamental. Petioles in greater 
or less degree sheath-like, spadix (inflorescence) commonly with 2- 
several spathes, the lower much longer, the upper split in anthesis, 
coriaceous or ligneous. Not unlike Cyclanthaceae, but fruit rarely 
a syncarp; leaves at least tardily deciduous, the small (rarely about 
1 cm. long), soon often chaff -like flowers briefly open, single or ter- 
nate, the central ordinarily female, its perianth usually persisting. 
Ovary commonly 3-celled, usually 1 ovule developing. Fruit more 
or less drupiform, variously fleshy, even plum-like, not infrequently 
fibrous or (and) coriaceous, rarely papery, the seed frequently in- 
cluded in a mesocarp that is extremely indurated and covered with 
the often intricately branched raphe; the fruit within soft or even 
early liquid; endosperm finally hard or firm, ruminated or homo- 
genous; embryo basal, lateral or apical, sometimes at an intermediate 
position. Few families are so generally recognized as this one, but 
it is so diverse in all characters that it can be and has been (see 
most standard reference works) described in great detail, although 
this is not necessary here. 

Notwithstanding the relatively few who have proposed and de- 
veloped the taxonomy, there has been considerable disparity regard- 
ing generic relationships and even more concerning generic lines. By 
and large the former as presented in recent works has been generally 
accepted but the latter in many cases still often expresses disagree- 
ments. In equivocal genera my own desire would be to give prefer- 
ence to unique or at least apparently stable characters, and to re- 
strict the use of recurring or variable ones to species or groups 
(sections) within the genus. 

321 



322 FIELD MUSEUM OP NATURAL HISTORY BOTANY, VOL. XIII 

In the case of the palms the contemporary who has advanced 
their study most intelligently is M. Burret, whose thoughtful work, 
cited on many of the following pages, is acknowledged with appre- 
ciation. The notable bibliographical contribution to the study of 
palms is that of my friend B. E. Dahlgren (Field Mus. Bot. 14: 1-456. 
1936). Compendiums like this lead without loss of time to all the 
literature available. In this family the citations to Weberbauer re- 
fer to the Spanish edition (El Mundo Veg. Andes Peru, 1945). The 
generic key, traditionally usable only by those whose vocation is the 
study of palms, has been devised with some sacrifice of precision for 
the benefit of those whose interest in them is a hobby; the natural 
but impractical division of the family, however, on character of fruit, 
has, regrettably, been retained, with the addition of one or two sup- 
plementary characteristics. 

The usefulness of palms, especially in primitive societies, has been 
described in numerous articles and books. Now, however, many 
think of palms chiefly as a source of coconuts (Cocos nucifera L.) 
and dates (Phoenix dactylifera L.) or are entranced by their noble or 
graceful beauty. 

Many contemporaries have been mentioned since the first num- 
ber of Part I of Field Museum's Flora of Peru appeared; to this 
number I would add a single name, Aven Nelson, my student men- 
tor, who enjoyed fame because his interest in others exceeded that 
in himself. 

Leaves pinnate; male flowers in dense catkins; fruits capitately 

crowded, strongly tubercled 1. Phytelephas. 

Leaves flabellate-palmate-parted; fruits never as above. 
Fruits tessellate; petioles not ligulate, leaves subconcolor; male 

flowers (at least) amentaceous. 

Spadix branches long; flowers seriately spiralled; seeds esulcate. 

2. Mauritia. 

Spadix branches short; flowers often distichous; seeds sulcate. 

3. Lepidocaryum. 

Fruits corky roughened or smooth; petioles ligulate; leaves silvery 
beneath. 

Epicarp corky; leaves subbasally parted 4. Chelyocarpus. 

Epicarp smooth; leaves biparted, each half incised. 

5. Tessmanniophoenix. 

Leaves various but never palmate, often pinnatifid, pinnate or fur- 
cate, rarely simple; fruits various. 



FLORA OF PERU 323 

Endocarp (nut) without obvious pores, the carpels (usually 3) dis- 
tinct or partly coalescent, in any case not finally an osseous 
shell; plants smooth or (and) with caudex root props; female 
petals sometimes imbricate. 
Leaf divisions or leaf more or less expanded apically, cuneate- 

based; caudex root-supported. 
Spadix branches simple; stamens 9-many; spathes often 

3-many. 
Spadix bisexual; leaf parts often much wider at apex. 

9. Iriartea. 
Spadix unisexual; leaf parts (Peru) oblanceolate. 

10. Catoblastus. 

Spadix often simple, axis thick, sexes separate; stamens 12-16; 
fruit villous 11. Wettinia. 

Leaves more or less pinnate, rarely entire, often bifid or segments 
oblique, sometimes ensiform-acuminate. 

Caudices slender, low or tall. 

Peduncle-spathes or bracts often 3-many; flowers rarely 
monoecious in diverse spadices, these finally simply 
scarred. 

Flower groups long-seriate (always?) . . 13. Wendlandiella. 
Flower groups not or not clearly seriate (always?). 
All spadices same sex on a plant ... 14. Chamaeodorea. 

Spadices of each sex on a plant 15. Morenia. 

Peduncle-spathes 2 or 3; flowers monoecious in diverse spa- 
dices; plants arundinaceous 16. Hyospathe. 

Caudices stout or (and) spathes only 2. 
Spathes 5; leaf segments rigid, ensiform; caudex robust, 

waxy; monoecious 12. Ceroxylon. 

Spathes 2; characters otherwise than above. 

Leaves unevenly pinnate or (and) flowers immersed. 

Flowers sessile, male petals sublanceolate, female im- 
bricate 16. Hyospathe. 

Flowers immersed, calyx segments imbricate as con- 
nate female petals; leaves various as in Chamaeo- 
dorea. 

Anther cells united, sagittate-cordate, filaments cupu- 
lately united 6. Calyptronoma. 



324 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Anther cells free, lax from cupulate tube. 

7. Geonoma. 

Anther cells free but porrect, filaments usually con- 
nate 8. Taenianthera. 

Leaves evenly pinnate, segments acuminate, divaricate; 

female petals imbricate. 

Flowers sessile, spadix branches pendulous, sheath fusi- 
form, split; male sepals valvate. 

17. Jessenia, 18. Oenocarpus. 

Flowers more or less immersed, spadix elongate, erect, 
sheath cylindric, entire; male sepals imbricate. 

19. Euterpe. 

Endocarp (nut) with usually 3 obvious pores, the 3 or more carpels 
coalescent; typified by the pores and osseous shell of the culti- 
vated coconut, but fruit (as Aiphanes) may be drupe-like and 
endocarp may be thin (as Desmoncus). 

Plants aculeate (more or less, rarely not, Bactris) ; sepals as petals 

valvate or connate (always?). 

Upper pinnae reduced, spiniform; endocarp thin or firm; plants 
dependent, sprawling, clinging 28. Desmoncus. 

Upper pinnae developed normally to rachis tip; endocarp osse- 
ous; plants independent (erect unless at base). 

Pinnae erose; male flowers usually immersed, congested, 
female corollas urceolate; spathes cymbiform; peri- 
carp often dry, rough, even aculeate. .26. Astrocaryum. 

Pinnae same, male flowers sessile, often scattered; petals 
fleshy, all basally connate, usually valvate; pericarp 
subfleshy 25. Aiphanes. 

Pinnae at least terminal, attenuate; flowers sessile, vari- 
ously disposed; pericarp more or less fleshy. 

27. Bactris. 

Plants smooth, more or less fleshy (lower pinnae rarely spini- 
form); perianth parts con volute-imbricate; leaf segments 
attenuate apically; fruit 1-6-seeded. 
Stamens 6; petals subcoriaceous; fruit 1-3-seeded. 

Petals minute; stamens subexserted or exserted; seeds 1. 

24. Maximilliana. 

Petals elongate, thick; stamens included; seeds 1-3. 

22. Syagrus, 23. Scheelea. 



FLORA OF PERU 325 

Stamens 6-many, anthers direct, included; petals lanceolate; 

seeds 2-6 21. Attalea. 

Stamens 12-24, anthers not direct, at least sinuous; petals 

ovate; seeds 2-6 20. Orbignya. 

1. PHYTELEPHAS R. & P. 

Elephantusia Willd. Sp. PL 4, pt. 2: 175b. 1805. Yarina Cook, 
Journ. Wash. Acad. Sci. 17: 223. 1927. Polandra Cook, I.e. 228. 

Reference: Cook, Journ. Wash. Acad. Sci. 17: 218-230. 1927. 

Smooth, with erect or inclined and rooting, moderately to very 
stout trunks, the species in Peru low but with many or few tall (elon- 
gate) terminal pinnate leaves. Flowers large, dioecious, the basal 
(Peru) spadices and peduncles squamate, the male rather amenta- 
ceous, more or less pendulous, simple, more or less scurvy, elongate, 
the female capitate, compact, erect. Stamens many, clustered, an- 
thers basifixed. Sepals 3, imbricate, as the 5-10 rather fleshy narrow 
acuminate petals. Staminodia many; ovary 4-9-sulcate, -celled, each 
cell with a single erect ovule, style terminal, stigma long-filiform. 
Fruits more or less pressed together forming a large or small syn- 
carp, 1-celled, more or less succulent or lignescent, shortly broadly 
and acutely tubercled, the hard nutlets loosely fibrous, embryo sunken 
near hilum in plane ivory-like albumen. The several segregate gen- 
era proposed by Cook have doubtful merit unless the diagnostic 
characters are accepted also in several other genera as defining sep- 
arate groups. Many illustrations are available, but some at least 
scarcely refer to the two original species unless, as possible, the sev- 
eral others proposed, as Panama species, are the same or merely vari- 
ants; consequently, only one or two drawings and a photograph are 
listed which represent at least the genus as it occurs in Peru. It 
stands apart in the family on account of the 4-9 united carpels, 
usually maturing several large long seeds. 

Vegetable ivory is supplied by the fruits of this extraordinary 
palm, described well by the authors: Early the fruits are filled with 
a liquid as clear as water, in place of which it is sometimes used; after 
some days it turns milky and acid; later it changes into a sweet and 
pleasant emulsion and successively acquires greater consistency, solid- 
ity and weight until it becomes as hard as ivory; small objects, as 
buttons, may be turned in a lathe with more ease than ivory ones 
because it is neither splintery nor fibrous; furthermore, it remains 
white for many years. The leaves are commonly used for thatch. 



326 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Besides the following, two or three species have been illustrated under 
other names without diagnosis. 

Trunks as petioles none or short, stout; leaf segments approximate, 
opposite; fruit lignescent P. macrocarpa. 

Trunks as petioles more or less developed; leaf segments distant; 
fruit succulent P. microcarpa. 

Phytelephas macrocarpa R. & P. Syst. Veg. 301. 1798; 224. 

Trunk none or short and inclined; leaves equally pinnate, crowded, 
the rachis basally incrassate, sessile or nearly, segments about 100 
pairs, mostly exactly opposite, basally reduplicate, linear-lanceolate, 
attenuate, the medial about 12 dm. long, 3.5 cm. wide, the upper 
scarcely 8 dm. long; spadices axillary, male ascending, fleshy, spici- 
form, 10 dm. long or slightly longer, peduncle compressed, 2.5 dm. 
long, 1.5 cm. thick, rachis nearly 8 dm. long, flowers sessile or sub- 
sessile, a cluster of many (150-280) stamens, filaments 7 mm. long, 
anthers 4 mm. long; spathes 2, glabrate, the outer nearly 2 dm. long, 
the inner 1 dm. long, 5 cm. across, linear-fusiform; peduncle and 
female spadix longer than male, the 12-20 sessile clusters apically 
compressed in an apical oval receptacle; bracteolate ovaries many, 
coalescent, the few central ones fertile, style filiform, 5 cm. long, stig- 
mas 5 or 6, 5 cm. long; fruits in a globose mass (12-20) 2.5 dm. in 
diameter or longer, the mature by mutual pressure 4-6-angulate- 
obpyramidate, apically muricate-tubercled, nutlets about 4 cm. long, 
2.5 cm. broad, perianth fragile. After Spruce, Journ. Linn. Soc. 11: 
179. 1871, who described P. aequitorialis of Ecuador (Polandra Cook), 
trunk 5 meters or taller, leaves unequally pinnate, the segments 
aggregate, male capitula stalked or racemed on a pendulous spadix, 
each capitulum with a thousand or more stamens; may occur in 
adjacent Peru. Illustrated, Huber, Bull. Herb. Boiss. se"r. 2, 6: pi. 11. 

San Martin: Vitoc, (Ruiz & Pawn). Tarapoto, (Spruce). Hua- 
nuco: Chanchamayo, San Antonio de Playa, Macora, (Ruiz & Pawn, 
type). Loreto: Headwaters of Rio Purus, (Chandless) . Rio Acre: 
Toward origin, (Huber}. "Pulu-puntu," "polo-ponto," "humiro." 

Phytelephas microcarpa R. & P. Syst. Veg. 302. 1798. Yarina 
microcarpa (R. & P.) Cook, Journ. Wash. Acad. Sci. 17: 223. 1927. 

Fruiting sometimes with no stem at all, finally with inclined stem 
about 1 dm. thick, spirally areolate with the deep leaf-scars; petiole 
elongate, leaf segments equal (Spruce), slender, with 6-7 leaves, seg- 
ments widely spaced, spreading, retrorse below, the lower much 



FLORA OF PERU 327 

shorter, the petioles perhaps 3-6 dm. long (Cook). The thick muri- 
cated rind of the fruit is reddish within, tough but may be eaten, 
flavor of melon or milky cheese; the albumen is drunk while still 
watery, eaten when fleshy, suggesting immature coconut, when ma- 
ture ivory-like (Spruce). Otherwise apparently unknown. Illus- 
trated, Pflanzenfam. 2, Abt. 3: 87 (flowers, fruits, at least in part); 
88 (plant). 

Hudnuco: Pozuzo, Pampa Hermosa, (Ruiz & Pavdn, type). 
Loreto: Near mouth of Rio Napo, (Spruce). Rio Pastaza, (Spruce). 
Ecuador? "Yarina," "rullipunta," "anon de palma." 

2. MAURITIA L. f. 

Mauritietta Burret, Notizbl. Bot. Gart. Berlin 12: 609-611. 1935. 
Reference: Burret, I.e. 605-609. 1935. 

Trees, the columnar trunk with a dense crown of deeply or fla- 
bellately divided semicircular leaves, the often large inflorescences 
pendent from their axils. Branches of spadix simple, at least male 
amentaceous, sessile, exserted from the spathe, the flowers densely 
spiralled, the internodes vaginate. Calyx 3-lobed, corolla 3-fid, male 
approximate, female few on short branches, staminodia connate with 
petals. Ovary (absent in male flowers) 3-celled, stigma sessile, ovule 
basal, erect. Fruit ellipsoid or globose, 1-seeded, tessellate with re- 
trorse lustrous scales (and leaves). Small prickly-stemmed species 
have been separated as a distinct genus by Burret, also by Hawkes; 
cf. M. peruviana Becc. The question of the correct name for those 
who wish to use the segregate genus is arguable, and perhaps conser- 
vation could here be useful. The segregate has both sexes in aments. 

Stems, leaves smooth, these concolor. 

Rachis strongly flexuose; fruit depressed M. flexuosa. 

Rachis slightly angled; fruit acute M. vinifera. 

Stems, leaves prickly, these pale-scaly beneath M. peruviana. 

Mauritia flexuosa L. f. Suppl. 454. 1781; 290. 

Smooth, the trunk to 7 dm. thick, 50 meters tall, medially slightly 
tumid, the leaves deeply radiately flabellate; stout petioles subterete, 
canaliculate above, subequaling the blades, these 8-12 dm. long, seg- 
ments 2-4 cm. wide; spadices 2-3 meters long, shortly peduncled, the 
male branches flexuose, elongate-cylindric, about 30, many 1 meter 
long, flexuose between the floriferous branchlets, these 5 cm. long; 



328 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

bracts scabrous-ciliate; flowers densely fuscous-velutinous or flaves- 
cent, calyx (male) turbinate, corolla 3-parted from stipiform base, 
segments lanceolate, acute, petals much smaller, stamens 6, anthers 
erect, oblique-cordate; calyx (female) urceolate, shortly lobed, corolla 
3-lobed; fruit 4 cm. high and thick, depressed-globose. After Drude; 
Ule found it on the Rio Jurua Mirim nearly at the Peruvian bound- 
ary, and Weberbauer saw it frequently cultivated but also in a natural 
state. Possibly occurring is the similar M. minor Burret, Notizbl. 
Bot. Gart. Berlin 11: 1. 1930, from adjacent Colombia, smaller in all 
its parts. Illustrated, Pflanzenf am. 2, Abt. 3 : 42, fig. 33 (habit) ; 
Drude, Mart. Fl. Bras. 3, pt. 2: pis. 62, fig. 2 (fruit), 63, (habit), 65, 
fig. 1, 67, fig. 2; Weberbauer, 584, pi. 33. 

San Martin rMoyobamba, ( Weberbauer) . Loreto: Tessmann, fide 
Dahlgren. To the Guianas. "Aguaje," "achual," "buritisol" (Ule, 
Brazil). 

Mauri tia peruviana Becc. Ann. Bot. Gard. Calcutta 12, pt. 2: 
225. 1918. Mauritiella peruviana (Becc.) Burret, Notizbl. Bot. Gart. 
Berlin 12: 609. 1935. Lepidococcus peruvianus (Becc.) A. D. Hawkes, 
Arquiv. Bot. Sao Paulo 2: 2. 1952. 

Type about 20 meters tall, the leaves deeply parted into many 
narrow long-acuminate segments, white waxy puberulent above, 
minutely and sparsely brown squamate beneath, the scales hair-like, 
the margins and midrib smooth; largest segments 7.5-8 dm. long, 
2.5 cm. wide; male spadix 1.5 meter long, spathes of the spike bear- 
'ing branches broadly infundibuliform, 6-7 cm. long, about as wide 
at opening; larger spikes 1 cm. long, the 18-20 male flowers (female 
unknown) oblong, obtuse, not distinctly biseriate, 6 mm. long, 3 mm. 
broad ; stamens 6. It seems rather closely related to M. armata Mart. 
For the basic purpose of this work identification the traditional 
designation of this little palm, with apparently solitary trunk (author), 
and its allies as a subgroup of Mauritia seems to be the useful classi- 
fication. Francia Chisaki kindly copied for me Beccardi's description 
from the University of California library at Berkeley. 

San Martin: In woody savannah, Moyobamba to Rioja, (Weber- 
bauer 4717, type). 

Mauritia vinifera Mart. Hist. Nat. Palm. 2: 42, pis. 38, 39. 
1824; 291. 

Resembles M . flexuosa; cylindric caudex early annulate; male spa- 
dix branches about half as long, less bent or angled between the stouter 



FLORA OF PERU 329 

branchlets; bracts and flowers finally glabrescent; fruit ovoid-ellip- 
soid, acute, 5 cm. long, 3.5-4 cm. thick. May extend into south- 
eastern Peru from Bolivia; listed by Dahlgren I.e. 418. Illustrated, 
Drude in Mart. Fl. Bras. 3, pt. 2: pis. 62, 67. 
Peru (see above). Bolivia; Brazil. 

3. LEPIDOCARYUM Mart. 

Slender smooth erect or flexuose stemmed with flabelliform pal- 
mately parted leaves, slender petioles, elongate inflated convolute 
sheaths. Floriferous branches of the spadices short, often distichous. 
Ovule horizontal. Allied to Mauritia, to which it has been referred. 

Male spadix branches many; flowers of both sexes shorter than 8 mm. 

L. tenue. 
Male spadix branches 6 or 7; flowers at least 8 mm. long. 

L. Tessmannii. 

Lepidocaryum tenue Mart. Hist. Nat. Palm. 2: 51, pi. 7. 1823. 
L. enneaphyllum Barb. Rodr. Enum. Palm. Nov. 19. 1875, fide Trail 
and Drude. 

Caudex slender, to several meters tall; fronds 12-15 or more, to 
2 meters long, glabrous except ciliate-spinulose margins of pinnae and 
midnerves, green, lustrous; petioles basally vaginate-lanceolate, ante- 
riorly compressed-terete, to three times longer than the flabellate- 
pinnatifid blade, this medially biparted even to base, mostly with 4. 
lanceolate acute pinnae parted to base (pinnae constantly 9, the 2 
ultimate binate, fide Rodriguez) or joined below, medially at least 
7.5 cm. wide, primary nerves acutely prominent above; male spadices 
not seen, the hermaphrodite to 1 meter long, peduncles subcompressed, 
spathes membranous, linear-lanceolate, obliquely truncate, glabrate, 
the rachis extension stout, glabrous or evanescently and finely tomen- 
tose, the 3-4 amentiferous branches 2.5 cm. distant, exserted from a 
tomentose spathe; spathes of branches 15-20, cyathiform, base of 
each black-annulate; scales suborbicular, subamplexicaul with rachis, 
lustrous; flowers solitary in each little spathe, pale rose color, the 
calyx campanulate, 3-denticulate, the corolla two times longer, petals 
erect, lanceolate, acute; stamens 6; anthers linear-oblong, 9, ovate 
(in male flower observed in hermaphrodite spadix); ovary ovoid, 
attenuate at both ends, minutely squamulate; stigma pyramidally 
connate, twice as long as ovary; fruit oblong-cylindric, tessellate- 
squamate, lower margin of scales membranous, obscurely ciliolate, 



330 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

apically black-fuscous; inner seed coat soft, very thin, reddish-gray 
with fuscous veins and adhering to scales; seed ellipsoid, testa casta- 
neous, lustrous; raphe developed entire length; albumen solid, osse- 
ous, embryo softer, conical, lateral. Fruit size of a walnut (Martius), 
presumably typical in character for the genus. Francia Chisaki gen- 
erously copied for me the detailed original description, here much 
reduced; also that of the apparent synonym, nearly without signifi- 
cant characterization. Since types of both plants are from Brazil 
near Rios Negro and Trombetas, the occurrence in Peru may be 
questioned. F.M. Negs. 18587, 18588. 

Peru (fide Dahlgren). Brazil. "Irapay." 

Lepidocaryum Tessmannii Burret, Notizbl. Bot. Gart. Berlin 
10: 771. 1929. 

Original plants a few cm. to 2 meters tall; petioles (as known) in- 
cluding sheathed portion about 1 meter long, stout, subglabrous, 6 mm. 
broad below the leaf, this rigid, 7 dm. long or longer, concolored, 
4-parted, the lateral incised 2 cm. above the rachis, the subequal 
lobes 6.5-7.5 cm. wide, the 5-6 primary nerves spinulose above as 
the inner margins of the inner lobes and the outer of the outer lobes; 
primary nerves 2, prominent only above, intermediate secondary few, 
transverse, curved above; male spadix with peduncle this 4.5 dm. 
long nearly 8.5 dm. long, branches 6 or 7, lower 13-17 cm. long; 
primary spathes narrow, the free part 11 cm. long, 4-5 cm. wide, sec- 
ondary funnelform, fuscous furfuraceous, caducous; flowering branch- 
lets little exserted, without flowers for 8 mm., the male flowers in 
about 7 rows, before anthesis nearly 1 cm. long; calyx turbinate, 
4 mm. long, clearly and acutely dentate, densely striate as the linear 
petals, these 2 mm. wide; female spadix nearly 5.5 dm. long, branched 
part 1.5 dm. long, the 4 branches 7-11 cm. long; flowers (before an- 
thesis) at least 8 mm. long, calyx half as long; fruit (after Tessmann 
drawing) scarlet, obovoid, 1.6 cm. thick, 3 cm. long with perianth 
and beak, this rather slender, acute, nearly 3 mm. long. Flowers 
rather white, seen on plants a few cm. high; fruit eaten after it has 
been softened in water. Related to L. tenue Mart, with more numer- 
ous male spadix branches, flowers of both sexes smaller; less closely to 
L. gracile Mart, (author). 

Loreto : Below Rio Morona, (Tessmann 4906, type) . "Irapay." 

4. CHELYOCARPUS Dammer 
Reference: Burret, Notizbl. Bot. Gart. Berlin 10: 394-397. 1928. 



FLORA OF PERU 331 

Trunk annulate, smooth as the flabellate leaves and ligulate peti- 
oles, these transversely rhomboid in cross section, obtusely margined, 
lightly bicanaliculate beneath and above, the ventral ligule triangu- 
late, the dorsal shorter. Leaves basally multi-parted, the elongate- 
cuneate apically laciniate, the laciniae minutely bidentate at tips. 
Fruiting spadix with only 2 spathe scars. Perianth segments 4, free 
or nearly, coriaceous beneath the globose fruit, this with pseudo- 
suberose tessellate epicarp similar to Pholidocarpus. Seed globose, 
broad raphe with few branches nearly to apex, embryo medially lateral. 

Chelyocarpus Ulei Damm. Notizbl. Bot. Gart. Berlin 7: 395 
(51). 1920. 

Usually 3-4, rarely 6 meters tall, the remotely annulate stem 7 or 
8 cm. thick; sheath unknown; petioles to 7.5 dm. long, only 5-7 mm. 
across, 3 mm. thick, the angles obtuse; ligule at base of leaf 3-angled, 
1 cm. wide, 8 mm. long, extended as a 2 mm. wide wing on the petiole, 
that at the leaf base beneath 2 mm. long; leaves green above, more 
or less white silvery beneath, semicircular, 7-13-parted to base, the 
divisions 3.5^4.5 dm. long, 4-10 cm. wide, 3-5-laciniate, these lobes 
5-10 cm. long with apical teeth 1-2 mm. long; transverse nerves 
between the longitudinal many; spadix in fruit 6 dm. long, branched 
part 2.5 dm. long, early white-floccose, peduncle as rachis strongly 
compressed; scars 13 mm. distant, upper about 8 mm. from lower 
branch, lower nearly the same distance above the base; branches 
mostly simple, about 1.5 dm. long, 2-3 mm. thick in herbarium, cari- 
nate-pulvinate below fruit, this globose, about 2.5 cm. in diameter, 
spiralled, solitary, with a solitary short ovate bract. Spadix descrip- 
tion after Burret, I.e., who also considers the possibility that C. Wal- 
lisii (Wendl.) Burret, the seed (described fully by him) with uniform 
albumen, probably from Colombia or Ecuador, may belong to this 
genus or Tessmanniophoenix, the ripe seed of the latter unknown. 
Type of C. Ulei (Ule 5885} being from Rio Jurua Mirim at the little 
community of Bele"m, nearly on the boundary with Peru, the species 
grows also no doubt within Peru, if environment is favorable. Illus- 
trated, Burret, I.e. 12: 152 (photograph). 

Loreto (surely). Adjacent Brazil. 

5. TESSMANNIOPHOENIX Burret 

Stems cylindric, smooth as the biconvex petioles, these with rounded 
margins, ligulate, the ligule protracted below the leaf-blade. Leaves 
palmate-flabellate, equally biparted nearly to base, the 2 divisions 



332 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

also divided. Spadix simply branched, the well-developed peduncle 
apparently with few deciduous spathes, none between the branches, 
these long-bracteate. Flowers solitary, hermaphrodite, sessile, 1- 
bracted. Perianth erect, closed (anthers exserted), segments seem- 
ingly 4, also irregularly 5 or 6. Stamens 7, 6, or 8, free. Filaments 
dilated, ovate, contracted above, anthers oblong, dorsally affixed 
near sagittate base, shortly incised at apex. Perianth little accres- 
cent beneath globose fruit, the obscure stigmas apical. Pericarp 
thin, epicarp smooth, mesocarp probably fleshy, endocarp thin but sub- 
osseous. Nearly Chelyocarpus Damm. which it resembles in petioles 
and division of the leaves and probably also in flowers, but well- 
marked by the smooth pericarp according to the author, who com- 
ments (I.e. 11: 500) on the validity of the genus as shown by T. dianeura 
Burret, from Colombia, also with short stout style, infundibuliform 
stigma as in Thrinaceae but that with calyx and corolla united and 
ordinarily only 1 carpel developed. The author (I.e. 12: 152-155) 
has distinguished the similar T. chuco (Mart.) Burret of Brazil, pos- 
sibly to be found in Peru, by the leaves, which are basally divided 
into halves, while those of T. longibracteata resemble those of Chelyo- 
carpus Ulei; photographs by H. A. Johnstone presented by Burret 
show clearly this difference. 

The noted ethnographer (Guenther Tessmann) made botanical 
collections with excellent data as noted elsewhere in this work; this 
generic name could be given greater euphony by omitting one con- 
necting (repetitive) vowel, regardless of tradition. Tessmanniodoxa 
Burret (one of the repetitive consonants also not deleted), Notizbl. 
Bot. Gart. Berlin 15: 336. 1941, defining generically Thrinax chuco 
Mart., formerly included here by Burret (I.e. 10: 400), is unlikely to 
be found in Peru. 

Tessmanniophoenix longibracteata Burret, Notizbl. Bot. 
Gart. Berlin 10: 398. 1928; 11: 315. 1932. 

Known to attain 5 meters, the stem 7 cm. in diameter, the 10-15 
leaves silvery beneath (except major nerves) with a dense or floccose 
puberulence as the younger spadices; petioles glabrous, rounded, stri- 
ate (dried) above, 2 meters long or longer, 6 mm. thick, below apex 
1 cm.; ligule fleshy coriaceous, triangular, smooth above, densely sul- 
cate beneath, protracted below and free from the blade, this a meter 
long, membranous, divided to 3 cm. above the base, the two parts 
semiflabellate, each cuneately 6-7-lobed nearly to base, the 2 medial 
lobes the widest, scarcely medially incised into broad segments, these 
shortly 2 dentate; laciniae of lower lobes acuminate; longitudinal nerves 



FLORA OF PERU 333 

many, transverse serpentine ones more obvious above; spathes 3, the 
lowest 19 cm. long, linear, 3 cm. wide below, 3.5 cm. above, the second 
extending about 2 dm. over the former, the third 2.5 dm. over the 
second, all rather strong but still soft, rather densely but evanescently 
floccose; spadix 8.5 dm. long, peduncle as rachis strongly compressed, 
6-7 cm. thick, little attenuate, annular scars 2 (3), about 1.5 dm. 
distant, branches 1-1.5 dm. long, scarcely divergent, basal bract lin- 
ear, 7 cm. (type) to 2 dm. long, the upper shorter; perianth erect, 
subglobose, closed, anthers entirely exserted; perianth mostly irreg- 
ular, segments nearly free, 4, also 6, rarely 5, glabrous; stamens 
mostly 7 (6 or 8), filaments free, subfiliform, anthers oblong, black 
punctate; carpels 2, free, slightly contracted apically, stigma dilated, 
rounded; fruits on umbonate pulvinae from solitary spiralled flowers 
but rather irregularly and thus pseudoverticillate, the single elongate 
dorsal bract uncinate-curved; fruit depressed globose, 8 mm. in diam- 
eter (immature); stigma obscure, scarcely excentric, sterile carpel 
present; perianth 7 mm. across, segments apparently 4, the 2 free 
sepals ovate, equaled by the alternating petals; mesocarp thin, endo- 
carp hard, seed basal. After author, who noted that he had little 
doubt of the identity of the flowering (Weberbauer) and fruiting 
specimens. 

Huanuco: Rios Pozuzo, Palcaza, (Weberbauer 6765, type); Tess- 
mann 4830, fruit. Junin : Capuahuanas, Rio Pichis, (Killip & Smith 
26717). "Uchapanga." 

6. CALYPTRONOMA Griseb. 
Reference: L. H. Bailey, Gentes Herb. 4: 153-172. 1938. 

Resembles Geonoma in general and floral character, the filaments 
cupulate and tubularly connate below, but the sagittate-cordate par- 
allel anther cells united to the connective, and the ovary, at least 
early, 3-celled. Leaf segments long-acuminate, often narrowly, even 
filiform-caudate. Peduncles of the laterally branched spadices elon- 
gate, the lower apically parted spathe much shorter. Included by 
Burret in Calyptrogyne Wendl. but distinguished by Drude; Bailey 
(I.e. 156) emphasizes in contrast the arboreal habit, uniformly pin- 
nate leaves, axillary rather than terminal spadix, 1-2-branched from 
a central axis, male and female flowers normally in same cavity any- 
where on axis, style central, mesocarp firm-ligneous, marked with 
stout interlocking fibers or ribs; perhaps these characters, impres- 
sive as they seem, will be found to be similar developments, varying 
in degree. 



334 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Spadix branches brownish-tomentose C. synanthera. 

Spadix branches white pruinose C. Weberbaueri. 

Calyptronoma synanthera (Mart.) Bailey, Gentes Herb. 4: 
166. 1938. Geonoma synanthera Mart. Hist. Nat. Palm. 2: 13, pi. 18. 
1823. Calyptrogyne synanthera (Mart.) Burret, Bot. Jahrb. 30: 137. 
1930. 

Leaves 1-1.5 meters long, pinnatifid, the lanceolate acuminate 
pinnae subfalcate; male spadices branched, 3 dm. long or longer, 
flowers approximate, the corollas little exceeding calyx. Otherwise 
unknown, but Burret (I.e.) observed that the vertical bract-enclosed 
ranks or lines of the flower pits are less marked, more delicate in con- 
trast to the same characteristic, especially in C. robusta Trail of the 
upper Amazon and in C. Kalbreyeri (Burret) Bailey of Colombia, 
and, also, in contrast to the latter, the pits are less crowded; the obo- 
void fruits, 9 X 12 mm., are said by Burret to resemble those of the 
latter in size. C. robusta Trail (Journ. Bot. 330. pi. 183, fig. 3. 1876; 
Mart. Fl. Bras. pi. 122} as to type had solitary erect stem to 5 meters 
tall, densely annulate, widely divaricate leaves 2-2.5 meters long, 
5-7 pairs of long-acuminate pinnae, nerves many, costa beneath fur- 
furaceous, spadix 9-12 dm. long, spike 1.5 dm. long, 7-17-branched, 
pits 8-10-ranked, lip reflexed, flowers unknown, fruit ligneous, ellip- 
soid, 2 cm. long, more than 1 cm. thick, smooth, the mesocarp coarsely 
fibrous. Type of C. robusta from Rio Jauary, Brazil, and quite pos- 
sibly is the little known C. synanthera or, in any case, it will be found 
in Peru. F.M. Neg. 18531. 

Hudnuco: Chicoplaya, (Ruiz & Pav6n, type). Brazil? 

Calyptronoma Weberbaueri (Burret) Bailey, Gentes Herb. 4: 
166. 1938. Calyptrogyne Weberbaueri Burret, Bot. Jahrb. 30: 139. 
1930. 

Type about 4 meters tall; petioles 6 dm. long, sheath short, early 
tomentose below (where 5 mm. broad), toward apex acutely carinate- 
canaliculate, finally gradually applanate, 4 mm. broad; leaf-segments 
laterally 7, filiform caudate, rather regularly disposed, the largest 
apical ones 4.5 cm. wide, 9.5-10.5 cm. long, 8- or 9-nerved, the nerves 
acute especially above, beneath (in herb.) brownish-paleaceous; up- 
per spathe (incomplete in type) 22 cm. long, brown within, paler 
without, deciduously ferrugineous tomentose as peduncle below, this 
compressed, 3- nearly 4 dm. long, at insertion of lower spathe 1.5 cm. 
broad; spadix about 8 dm. long, the 10-13 simple branches white- 



FLORA OF PERU 335 

pruinose, 6 mm. thick, 2-2.5 dm. long, basal bract broadly ovate, re- 
flexed, sterile base short; flower pits in 8 ranks, these 6 mm. distant, 
but along 2 elevated lines rather well separated; pit bracts (lip) in 
fruit reflexed, 2.5 mm. broad, lanceolate sepals then 3.5 mm. long; 
fruit pruinose, obovoid, 11-14 mm. long, 8 mm. thick, rounded api- 
cally, subincurved to nearly stiped base, epicarp branched-fibrous, 
strongly 14-nerved toward base. 

Puno: Chunchusmayo, 900 meters, (Weberbauer 1234, type). 

7. GEONOMA Willd. 

Reference: Burret, Bot. Jahrb. 63: 144-266. 1930; second page 
references refer to descriptions by Drude in Mart. Fl. Bras. 3, pt. 2. 
1882. 

Smooth, slender, the trunk annulate, the terminal or alternate 
leaves entire, apically bifid or pinnatisect with acuminate segments, 
tubular sheath. Spadices variable in development, the lower of the 
2 spathes (these usually caducous) incomplete, truncate, concave, 
the upper compressed or fusiform. Flowers glumaceous, unisexual, 
immersed in spirally or vertically disposed pits (1-3 in each), the 
central (if present) female developed after the male; both sexes with 
sepals slightly imbricate below, petals ciliate, connate toward base. 
Male flowers deciduous, filaments united into a truncate, entire, den- 
tate or laciniate tube, the 6 (typically) anthers with free cells. Ovary 
1-celled, ovule ascending, long style lateral, the 3 stigmas subulate. 
Fruit with fibrous usually thin pericarp; endosperm uniform. G. 
multiflora Mart., 261, G. laxiflora Mart., 239, are scarcely to be ex- 
pected within Peru, their known distribution being out-of-line with 
that of other Brazilian species extending into Peru; cf. also G. myri- 
antha Damm. 

It is certainly fortunate for the users of this work that Burret's 
key (I.e. 145-161) has been available; also, he has clearly presented 
his definition of the generic boundaries and his interpretation of the 
relative importance of specific characters. With laudable humility, 
he expressed his conviction that his key (and his conclusions) are 
pioneer studies. 

KEY TO GEONOMA 
(After Burret, inserts by me) 

Staminodal tube cylindric, truncate or obscurely dentate (subgenus 
Eugeonoma Spruce). 



336 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Spadix simple, rarely bifurcate (branches 3, G. Tessmannii). 
Peduncles well-developed, subequaling or longer than spikes (see 

G. Tessmannii). 

Alveoli oval, obliquely truncate, upper edge not produced; 
spathes two-thirds as long as spadix. 

Leaf segments 3 pairs G. granditrijuga. 

Leaf segments 5-8 pairs G. gracilipes. 

Leaf segments mostly 6 pairs G. multisecta. 

Alveoli bilabiate, upper edge acute, extended; spathes often 

three-fourths as long as spadix. 
Spathes one-half as long as spadices. 

Leaf segments oblong-subquadrate G. Brongniartii. 

Leaf segments cuneate-based G. cuneifolia. 

Spathes two-thirds (at least) as long as spadices, equaling 

or exceeding pedicel G. adscendens. 

Peduncles short, exceeded by spikes only a little (G. Tessmannii, 

spike small). 
Leaves simple. 

Spadix normally 3-branched, slender G. Tessmannii. 

Spadix simple or branches stout. 
Spikes 2-10 cm. long. 

Alveoli lips (lower) 7-8 mm. distant; spikes rather stout. 

G. Uleana. 

Alveoli lips 4-5 mm. distant; spikes stouter. 

G. pycnostachys. 

Spikes 2-2.5 dm. long G. piscicauda. 

Leaves pinnate G. Raimondii. 

Spadix branches 3 (2) -many. 
Alveoli bilabiate, spiralled or verticillate, upper edge somewhat 

produced. 

Spadix branches stout or alveoli spiralled. 
Branches rather stout; flowers as alveoli mostly rather large. 
Spathes narrow, often long-tubular, thin, the upper often 
high above lower. 

Leaf segments 2-3 pairs G. andina. 

Leaf segments 7 pairs G. floccosa. 

Spathes broad, firm to coriaceous, subequal, little remote; 
leaf segments many. 



FLORA OF PERU 337 

Alveoli subapproximate; some spadix branches branched. 

Spadix axis 7-9 mm. thick G. Weberbaueri. 

Spadix axis 4.5-5.5 mm. thick G. helminthoclada. 

Alveoli congested; spadix branches simple. 

G. congestissima. 

Branches slender; flowers as alveoli small. 
Spadix branches 3, slender, only to 6 cm. long; leaves 

simple G. Tessmannii. 

Spadix larger or characters not as above. 
Alveoli approximate. 
Spathes 1.5-3.5 dm. long; branches usually simple. 

G. interrupta. 
Spathes as noted shorter; spadix branches long, 2-3- 

branched G. myriantha. 

Alveoli laxly spiralled; leaves simple, oblanceolate. 

G. leptospadix. 
Spadix branches slender, alveoli alternate in verticils. 

G. myriantha. 
Alveoli uniformly edged; spathes more or less dilated, firm or 

inner membranous. 
Branches stout; alveoli mostly subregularly in verticils. 

G. megalospatha, G. andicola. 
Branches slender; alveoli spiralled. 

Leaves unevenly pinnate . . . . G. Poeppigiana, G. interrupta. 

Leaves evenly pinnate, segments 3 (4) pairs . . . G. Killipii. 

Staminodal tube divided into 6 linear teeth (subgenus Astrandroe- 

cium Spruce; cf. Taenianthera Burret, staminodal tube dentate 

or lobate). 

Alveoli in 5-6 series little inclined toward axis; leaf segments small. 

Leaf segments various, 1-7.5 cm. wide G. Spruceana. 

Leaf segments (apical) rhombic, 1.5 cm. wide, 2.5 dm. long. 

G. camptoneura. 
Alveoli in 7-8 series; leaf segments larger G. longisecta. 

Geonoma adscendens Damm. ex Burret, Bot. Jahrb. 63: 175. 
1930. 

Finally erect, to 1 meter tall, 8 mm. in diameter above; sheath 
amplexicaul, early membranous, 8 cm. long, petioles about 2 dm. 
long, hardly 2 mm. thick, leaves 28 cm. long to the cleft (lobes nearly 



338 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

1.5 dm. long), segments rather regularly 4 pairs, the broader upper 
slightly falcate, acuminate, 8 cm. wide at rachis, the lowest 1.5-2 cm. 
wide, primary nerves 26 or 27 in each lateral lobe, rather acutely 
prominent beneath, secondary nearly as high; spadices simple, flower- 
ing in axils of withered leaves, slender, about 2-3.5 dm. long, spathes 
narrow, apparently including the peduncles, these 12-25 cm. long; 
floral pits laxly separated, bilabiate, upper lip shorter, acute, lower 
emarginate, axis nearly 2.5 mm. thick, glabrous but minutely granu- 
late; sepals (male) about 3.5 mm. long, petals two-thirds connate, 
lobes ovate, staminal cup rather robust, female calyx (also before 
anthesis) nearly 3 mm. long, petals about as male, staminodal tube 
about a third longer, truncate. Perhaps comparable to G. arundi- 
nacea Mart., 180, Amazonian, the flowers mostly laxly spiralled in 

3 series, the short peduncle conspicuously exceeded by the spike, the 
nerves about 20 (Burret). 

Cuzco: Steep wooded outcrop, 1,700 meters, St. Anna above Haci- 
enda Idma, (Weberbauer 5033, type). "Cuculli." 

Geonoma andicola Damm. ex Burret, Bot. Jahrb. 63: 218. 1930. 

Leaves narrowly pinnate, the linear leaflets not at all falcate, the 
first broadest and only about 1 cm. wide, the petioles apieally 7 mm. 
broad, rounded beneath, early leprose; primary nerve one, rather 
well-developed above, fuscous beneath with crowded narrow scales 
between the primary and secondary nerves (the latter marginal), the 
tertiary more or less developed, the larger minutely lepidote, others 
sparsely; spadices duplicately branched, glabrous, outer spathe sub- 
coriaceous, 3.5 cm. broad, 23 cm. long, the inner reddish furfuraceous 
or glabrate; peduncle to 5 cm. long, at base of lowest branch 7 mm. 
thick; rachis to base of top branch 8.5 cm. long, primary branches 8, 
acutely angled, broadly bracted at base, the lower bifurcate, 2.5- 
3.5 cm. pedunculate, the rest simple, the spikes floriferous to rounded 
tips, 11.5-14.5 cm. long, axis 5 mm. thick; alveoli in 6 nearly vertical 
series, 6 or 7 mm. distant, the upper lip none, even fruiting margins 
rounded, the lower long-extended, always deeply emarginate; sepals 
4.5 mm. long, oblong; petals 4.5 mm. long, these two-thirds connate, 
staminodal tube scarcely incised, subtruncate, exceeded 1 mm. by 
stigmas; fruit at least 1 cm. long, 6 mm. broad, obscurely mammil- 
late, densely elevated-lineolate; male flowers fallen. Type noted as 

4 meters tall. 

Puno: Chunchusmayo, 1,500-1,800 meters, (Weberbauer 1345, 
type; 573). 



FLORA OF PERU 339 

Geonoma andina Burret, Bot. Jahrb. 63: 188. 1930. 

Leaves (type) to 47 cm. long, broadly elliptic, primary nerves on 
each side 18 or 19, segments 3 pairs, strongly scattered, lightly fal- 
cate, acuminate, the basal nearly 4 dm. long, the apical on interior 
margin nearly 2 dm. long, 3.3-4.2 cm. wide; primary nerves very 
prominent above, somewhat slenderer than the secondary beneath, 
toward the rachis furfuraceous, the tertiary many, obvious both sides, 
clearly shortly appressed-pubescent beneath, the primary nerves me- 
dial, segments of adult leaves 7 mm. distant; spadix over 4 dm. long, 
the branched part 1.5 dm. long, the rachis 3 cm. long, the spathes 
equaling three-fourths of the peduncle, the lower 14 cm. long, appla- 
nate below, 1 cm. broad, the upper 1 dm. above the lower; peduncle 
furfuraceous toward base, bracted above; branches 5, basally bracted, 
axis in flower nearly 2.5 mm. thick, 10-12 cm. long or longer, granu- 
lose, extended into a slender spine; alveoli rather laxly spiralled, 
upper lip somewhat produced, lower incised; male flowers not open. 

Cajamarca: Tambillo, Prov. Cutervo, (Raimondi, type). "Frutilla." 

Geonoma Brongniartii Mart. Palm. Orbign. 24. pi. 12, fig. 1, 
pi. 23C, figs. 3, 5, 6. 1847; 174. 

A most attractive subacaulescent palm, the overall height only 
1 meter (Ule), the 6 or 7 long leaves reddish beneath, clustered on a 
very short caudex this with a few supporting roots the few rufes- 
cent spadices erect; leaves about a meter long, the petioles shorter 
than the 2-3 pairs of oblong subquadrate acute segments, these to 
1.5 dm. wide; primary nerves about 24 each side, more prominent 
beneath, the alternating secondary impressed above, nearly as incras- 
sate as the primary below and scattered paleaceous, the slender ter- 
tiary many (7-9); spathes membranous, finally fibrous, outer 1.5, 
inner 2.5 dm. long; peduncles about 4 dm. long, the rachis about as 
long, cylindric, attenuate to acute mucro 12-15 mm. long; early 
flowers female, soon (as usual) with 2 male in each remote pit, this 
with a broad emarginate bract or lip; calyx 2 mm. long, lobes lanceo- 
late, carinate, ciliolate, subequaling the ovate-oblong, subobtuse, 
marginally membranous sublacerate corolla segments; stamen tube 
urceolate; female rachis at maturity incrassate, red, the globose black 
fruits 8 mm. in diameter. Burret (I.e.) corrected Martius' descrip- 
tion, noting the spadix as both male and female and that fig. 6 is in- 
correct as to scales; but cf. G. cuneifolia Burret. F.M. Neg. 38642. 

Loreto (no doubt, as Ule 5593 nearly on the Brazilian boundary). 
Adjacent Brazil; Bolivia. 



340 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Geonoma camptoneura Burret, Notizbl. Bot. Gart. Berlin 11: 
201. 1931. 

Rachis fuscous furfuraceous, the apical portion seen (19 cm. long) 
with widely divaricate membranous segments, 2.5 dm. long on upper 
margins, 14 or 15 cm. wide, nearly rhombic, the outer margin sub- 
parallel to rachis; primary nerves 14 or 15, sigmoid, strongly curved 
to margins, to 1.5 cm. distant, secondary beneath nearly twice wider 
than primary, the many tertiary prominent both sides; spadix (frag- 
ment) 16.5 cm. long, the 8 branchlets 10.5-13 cm. long, minutely 
pilosulous but green, apically aculeate, the spine short, the male 
flowers immersed, axis nearly 3 mm. thick, granulose; alveoli in 6 
series little inclined, 4-5 mm. distant; lower lip densely costate, some- 
what protracted, subobtuse, the upper slightly but obviously extended 
from the axis; flowers mostly emerged at anthesis, the faded perianth 
3.5 mm. long, somewhat shorter than corolla; stamens long-exserted; 
staminode teeth 6, linear. Reminds one of G. latisecta Burret with 
quite different leaf-segments. 

Loreto: Yurimaguas, Williams 7836, type. 

Geonoma congestissima Burret, Bot. Jahrb. 63: 224. 1930. 

Arundinaceous, 4 meters tall (type) ; petioles 3 mm. broad at apex, 
2 dm. long or longer without sheath, this apically auriculate, early 
tomentose, the many nerves nearly vertical; leaf -rachis beneath as 
petioles scurfy, 3.8 dm. long, not at all extended; primary nerves 
31 or 32 each side, prominent beneath, minutely lepidulous, tertiary 
above obsolete, beneath 7 between secondary and primary, finely 
appressed setulose; leaves membranous, subelliptic, to about 28 cm. 
broad, to apices 53 cm. long, to bifurcation 1.5 dm. long, the 3 pairs 
of distinct segments falcate, narrowly acuminate, 5-10 cm. wide, to 
about 2.5 dm. long, interval between basal and medial pair nearly 
7.5 cm. long, primary nerves 8-10 except shortly lobed terminal part 
11-12-nerved; spadix about 2.5 dm. long, branches simple, 6-10, 
rachis 2.5-10 cm. long, spathes subligneous, early fulvous tomentose, 
1 dm. long, 2 cm. broad at apex, peduncle 8-16 cm. long, spikes very 
dense, about 7-11 cm. long, before anthesis 8-10, in fruit 15-17 cm. 
in diameter, axis alveolate to rounded apex; alveoli approximate in 
7 series, the parts 3-3.5 mm. distant; upper as lower lip short, the 
latter rounded, finally often reflexed, entire; male sepals before an- 
thesis nearly 4 mm. long, the outer spathulate, 1 mm. wide, petals 
to 3.25 mm. long, to one-half connate, staminal cup (without stipe) 
1 mm. long, as filament to connective, this to 0.75 mm. long, black, 



FLORA OF PERU 341 

anthers 2 mm. long; female sepals oblong, 4 mm. long, as petals, these 
to one-half connate, the staminodal tube little shorter, scarcely in- 
cised; fruits congested, subglobose, 7 mm. in diameter, 6 mm. long, 
broadly rounded and submammillate apically, broadly protracted 
basally, minutely tuberculate, epicarp membranous, mesocarp finely 
fibrous, endocarp cartilaginous. 

San Martin: Among shrubs, 1,300 meters, Moyobamba, (Weber- 
bauer 4560, type). 

Geonoma cuneifolia Burret, Notizbl. Bot. Gart. Berlin 11: 199. 
1931. 

Simulates G. Brongniartii but the caudex developed, leaves about 
2 meters long or longer, segments definitely cuneate at base, the 
lower 5-7 cm. wide and with many more primary nerves; peduncle 
to 5.5 dm. long; spadix 6-8 dm. long, the spike 2 dm. long, 3 mm. 
thick. The much larger leaves with basally long-cuneate segments 
may distinguish this species from the one to which the author origi- 
nally referred the type. 

Loreto: Rio Ucayali, (Tessmann 3317, type). 

Geonoma floccosa Damm. ex Burret, Bot. Jahrb. 63: 203. 1930. 

Type a meter tall, petioles 4.5 dm. long, at apex 3 mm. thick, 
canaliculate above, basally vaginate; rachis nearly 3.5 dm. long; pri- 
mary nerves 22 or 23; segments on each side about 7, apical broadest 
(primary nerves about 7), 14-20 cm. long, mostly 3-3.5 cm. wide, 
but sometimes narrower, basal primary nerves about 7, basal seg- 
ments nearly 2-2.5 cm. wide, all falcate; nerves subequal except 
tertiary, these obscure above, rather many beneath, sparsely and 
minutely pubescent; spadix 5 dm. long, simply branched; spathe 
narrow, to 34 cm. long, subequaling or little longer than peduncle, 
this as branches densely long-floccose-tomentose, somewhat unevenly 
deciduous; upper bracts oblong; rachis not flexuose at union with the 
7 or 8 divergent branches; alveoli rather densely spiralled in 5 series 
little inclined toward the axis and after anthesis equally 5 or 6 mm. 
distant, the upper lip clearly extended, the lower often cleft; male 
flowers emerged; fruit subglobose, apiculate at base, rounded at apex, 
1 cm. long, half as broad, densely and coarsely verruculose, endocarp 
little exceeding one-third mm., the seed 7 mm. in diameter. Flowers 
brownish-purple (collector) . 

Junin: Low moist shrub-wood, west of Huacapistana, (Weber- 
bauer 2277, type). 



342 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Geonoma gracilipes Damm. ex Burret, Bot. Jahrb. 63: 173. 
1930. 

Original plant recorded as only 1 dm. tall, the fuscous-leprose 
petioles and leaf-rachis each about 3.8 dm. long; segments (each side) 
5-8, mostly 1.5-2.5 cm. apart, all or nearly all with many primary 
nerves, falcate, long-acuminate, the broader apical 15-17 cm. long, 
4-5 cm. wide; primary nerves each side about 28, to 7 mm. distant, 
subequaling the secondary, the tertiary conspicuous beneath; spike 
7.5-11 cm. long, including the more or less spinescent tip, the axis 
to 4 mm. thick, alveoli densely in 6-7 series; sepals 2.25-2.5 mm. 
long, ovate-oblong, the petals 3 mm. long, staminodal tube subentire; 
fruit subglobose, minutely granular, 6 mm. in diameter, the pericarp 
one-third mm. thick; male flowers unknown. G. Jussieuana Mart. 
(Palm. Orbign. 24. pi 12, fig. 2, pi. 23 A. 1847), to which Burret (172) 
with doubt referred the following, may prove to be the same but, as 
he noted, the species of Martius ex char, (mostly from description 
and drawing of D'Orbigny) and from illustrations has petiole much 
shorter than leaf-rachis, peduncle and spadix subequal, in contrast 
to the specimen of Weberbauer with petioles and leaf-rachis subequal, 
peduncle and spadix very unequal; also, if the same species, and vari- 
ety, it would be more expected in southern Peru as the type is from 
northeast of Cochabamba, Bolivia. 

San Martin: Shrub-wood, 1,100 meters, Moyobamba, (Weber- 
bauer 4557, type). 

Geonoma granditrijuga Burret, Bot. Jahrb. 63: 171. 1930. 

Low (type 2 meters), the leaves, apparently, regularly with 3 pairs 
of subequal and subequally disposed segments, all glabrous, the ter- 
minal one 14 cm. decurrent, nearly 10 dm. long to the bifurcation, 
this to apex of lobes 3.7 dm., upper margins direct, narrowly acumi- 
nate, apex little falcate, the medial to 9 cm. wide; primary nerves 
finally arcuate, 34 or 35 on each side, strongly prominent only above, 
the secondary half as high, the tertiary fine, especially beneath; spa- 
dix simple, peduncle and spike subequal, 21 cm. long or longer, 2.5- 
3 mm. thick below spike, lower spathe 12-14 mm. long, narrow, 
robust, spike apex caudiform-spiniform, 1 cm. long, 3 mm. thick, 
leprose; alveoli in 6 exactly vertical lines, the parts of each equally 
5 mm. distant; sepals of male flowers 2.25 mm. long, the outer ovate, 
the rest ovate-oblong; petals two- thirds connate; staminal cup stout, 
aJithers nearly 1.5 mm. long to tip of connective; sepals of female 
(young) ovate-oblong, acute, staminodal tube obscurely dentate. 



FLORA OF PERU 343 

Spadix of 3672, male flowers emerging, nearly 7.5 dm. long, lower 
spathe 3.2 dm. long, linear, subligneous, to 16 mm. broad, upper 
scarcely longer, peduncle nearly 4.5 dm. long, about 12 cm. larger 
than spathe, spike to spine-like tip 2.85 dm. long. This inflorescence 
doubtless belongs here (author); so, why not include the measure- 
ments above? 

Huanuco: Sparse woods, 700 meters, Monzon to the Huallaga, 
(Weberbauer 3673, type; 3672). 

Geonoma helminthoclada Burret, Bot. Jahrb. 63: 222. 1930. 

Leaf-rachis dorsally plane, glabrous, narrowly triangular toward 
the filiform extension; segments irregularly aggregate, at base very 
narrowly reduplicate, 1.5 cm. wide, midrib above more or less prom- 
inent, beneath the one strong nerve besides the marginal, the tertiary 
obsolete; spadix (fragment) twice branched or more, peduncle 6.5 cm. 
long, extending into 4 branches 3-3.5 dm. long, 4.5-5.5 mm. thick, 
minutely tuberculate, rounded (espinose) apically; alveoli in 3 alter- 
nating verticils, upper lip moderately but conspicuously and thinly 
produced, the lower split, the series 8-9 mm. distant; female flowers 
under anthesis 1 cm. long, staminodal tube equaling petals, truncate, 
scarcely incised. Recalls the much more northern G. undata Klotzsch, 
213, but the leaves, from the single apical fragment seen, appear to 
be distinctly different (author). 

Amazonas: Chachapoyas, (Raimondi 509, type). 

Geonoma interrupta (R. & P.) Mart. Hist. Nat. Palm. 2: 8, 
pi. 7. 1823; 249. Martinezia interrupta R. & P. Syst. Veg. 296. 1798. 

Type a few meters tall, leaves unequally and interruptedly pin- 
nate, segments linear-oblong, falcate-acuminate, multinerved, 4 dm. 
long or longer; inner spathe coriaceous, cuspidate, 2.5 cm. broad, 
linear-cuspidate, unevenly grayish tomentose; spadices branched be- 
low, the rarely bifid branches 1.5 dm. long or longer, shortly attenu- 
ate, sparsely puberulent; flowers more or less approximate, 5-seriate, 
about 2 mm. long, calyx 3 times shorter than or equaling (?) corolla, 
lobes strongly concave, subserrulate; staminodal cup (female flowers) 
about as long as corolla; fruit globose; acute, about 4 mm. in length. 
After Martius who seems not to have seen the earlier developed 
male flowers as he notes distinct female and male spadices. G. eu- 
spatha Burret (Notizbl. Bot. Gart. Berlin 11: 10. 1930) from adjacent 
Colombia has larger flowers, the male 2.5 mm. long, spathes much 



344 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

longer; it is one of several Colombian species proposed in the same 
paper that may be expected in Peru. F.M. Neg. 18509. 

Huanuco: Cuchero, Pozuzo, (Ruiz & Pav6n, type). "Cuyol," 
"siasia," (Ruiz & Pavon), "chica-chica." 

Geonoma Killipii Burret, Notizbl. Bot. Gart. Berlin 11: 320. 
1932. 

Stems arundinaceous, the small leaves with 3 pairs (rarely a fourth 
reduced pair) of leaflets on a rachis 23 cm. long, the slender petiole 
concave above; leaf -blade 33 cm. long, paler beneath, primary nerves 
22 each side; leaflets falcate, the apical twice as wide as the medial, 
these longer, the basal nearly directly divaricate, sigmoid, half as wide 
as the medial, all very narrowly long-acuminate; secondary and pri- 
mary nerves equally strong beneath; peduncle 4 cm. long, rachis 5 
cm. long, verrucose as the 8 branches, the 2 lower of these furcate, 
all slender, at most 1.5 mm. thick at internodes; alveoli deep, lower 
margin produced, inflated, truncate, the upper obscurely extended, 
rarely spiralled, often decussate, 3-4 mm. distant; fruit (dried) black, 
subglobose, at least 5 mm. in diameter. The detached leaf (26532} 
scabrous, petiole (above sheath) 2.5 dm. long, rachis 3 dm. long, upper- 
most leaf segments 13.5 cm., decurrent, upper margin 1 dm. long, 
medial 6-7 cm., length of rachis 13 cm. on margin, at most 6-6.5 cm. 
wide, lowest segments 1.7-2 cm. long (rachis), 1.5 dm. on upper mar- 
gin, at most 3.5 cm. wide; spadix 19 cm. long. Similar to G. panicu- 
ligera Mart, but with smaller slender leaf-segments; the alveoli are 
not spiralled or 3-whorled, but placed crosswise (author). 

Junin : Puerto Bermudez, (Killip & Smith 26594, type; also 26532} . 

Geonoma leptospadix Trail, Journ. Bot. 14: 327, pi. 183, fig. 2. 
1876; 230. 

Caudex reed-like, 1.5-2 meters tall, fulvous, glabrous; leaves 10-12 
(petioles 4-5 dm. long, sulcate above), the beaks about 1 dm. long, 
long-decurrent, simple, bifurcate, the deltoid segments about 4 cm. 
wide; primary nerves faint, 27 pairs; spadices 3-14, in flower and fruit 
on same plant, reddish, 2.5-4 dm. long, peduncle to 2 dm. long, spa- 
dices finally often inflexed, 3-5-branched, flower pits scattered, ob- 
scurely 5-ranked, bract 2-lobed or emarginate; male sepals and petals 
subequally, medially connate; sterile staminal tube of female flowers 
subentire or minutely 6-denticulate; fruit globose, 6 mm. in diameter, 
black-purple. Stem usually partly buried; in one plant I found 14 
spadices, the lower ten in fruit (author). Recorded, besides lower 



FLORA OF PERU 345 

Amazon (type locality), at Jurua Mirim, Brazil, near Peruvian 
boundary (Ule 5516, 5516b, det. Burret). F.M. Neg. 38651. 
Peru (no doubt). Amazonian Brazil. "Ubim." 

Geonoma longisecta Burret, Bot. Jahrb. 63: 257. 1930. 

Type 2 meters tall with 12 very rigid leaves each with 2 pairs of 
approximate (2-2.5 cm. distant) leaflets, these little falcate, the lower 
8 dm. long, about medially 7 cm. wide, apical about 7 dm. long or 
longer on upper margin, 1 dm. wide, the apices not seen; rachis 
filiform-produced, fuscous scurfy beneath; primary nerves of larger 
segments nearly direct, very prominent above, obscure, scaly be- 
neath, the secondary impressed above, very prominent beneath, the 
tertiary there more obvious than above; peduncle 4.5 cm. long, 6 mm. 
thick at apex, the partly twice branched spadix 2.5 dm. long, nearly 
3 dm. wide; rachis 1 dm. long, branches 12, the upper 6 simple, in 
flower 9.5-11.5 cm. long, axis 2.5 mm. long, pilosulous-granulose, 
alveolate to spineless tip; alveoli in 7 series, upper lip thin-margined, 
lower produced, often emarginate; stamens 2.5 mm. long, petals not 
exserted; female flowers (early) scarcely 2 mm. long, staminodal tube 
exserted, teeth linear; fruit oval, rounded both ends, about 13 mm. 
long, 11 mm. broad, densely and minutely granular, pericarp at least 
1 mm. thick, seed 1 cm. long, 9 mm. broad. Resembles G. Spruceana 
Trail with much shorter and narrower leaf segments and with the 
spadix rachis marked by the terminal flowers overreaching the spine- 
less point. 

Loreto: Flood-free wood, Iquitos, (Tessmann 5087, type). 

Geonoma megalospatha Burret, Bot. Jahrb. 63: 218. 1930. 

A larger counterpart of G. andicola Damm., the type 6 meters tall, 
petioles apically 9 mm. broad, filiform rachis extension stouter, about 
5 mm. long, leaflets nearly 12 mm. wide, tertiary nerves 7-9; spadices 
deciduously furfuraceous, outer spathe 36 cm. long, at base 5.5, 
medially 6.5-7 cm. broad, coriaceous; lowest branch spadix to apex 
37 cm., rachis to base of ultimate branches 23 cm. long, primary 
branches 13, upper 7 simple, lowest to 7 cm. pedunculate, spikes 
16-17.5 cm., axis 7 mm. thick; sepals nearly 5 mm. long, petals 4.5 
mm. long; fruit stoutly stiped (stipe 2-3 mm. long), rounded at base, 
obscurely apiculate, including stipe 13 mm. long, 9 mm. broad, lightly 
rugulose, yellowish, pericarp nearly 1.25 mm. thick. In all parts, 
notably spadix, larger and more robust; the spathes are nearly twice 
as long and wide, spadix branches thicker, longer, pinnae wider 



346 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

(author). To an ecologist this vigorous condition resulting in greater 
size may suggest maximum natural conditions, to a horticulturist, a 
personal achievement in providing them. Grows with Ceroxylum 
crispum Burret at the altitudinal boundary for Peruvian species of 
palms. 

Huanuco: Pozuzo, (Weberbauer 6800, type). 

Geonoma multisecta Burret, Notizbl. Bot. Gart. Berlin 12: 
155. 1934. Taenianthera multisecta Burret, I.e. 11: 13. 1930. T. 
Weberbaueri Burret, Bot. Jahrb. 63: 269. 1930, fide Burret, I.e., 1934. 

Type like Taenianthera acaulis in habit; petioles (triangular) as 
leaf-blade to base of uppermost furcate segment about 7 cm. long; re- 
maining segments mostly 6 pairs, long-cuneate to base, upper to 6.5, 
rest mostly 4-6 cm. wide, all falcate, narrowly acuminate; primary 
nerves many; spadix simple, about 14 cm. long, peduncle 12 cm. long 
or longer; spathes slender, narrowly vaginate, not half as long as pe- 
duncles; spike 16.5-21.5 cm. long, 8 mm. broad including bracts, the 
nearly concealed axis 5 mm. broad; pits densely congested even in 
fruit, in 7 series, little inclined; lower lips strongly extended, deeply 
incised, in each series 4-5 mm. distant; sepals 3.25 mm. long, oblong- 
lanceolate, subequaling petals, to two-thirds connate; fruit oval, 9 
mm. long, 6 mm. thick, densely tuberculate, little protracted at 
base. After seeing more material with very irregular segments I 
believe that T. Weberbaueri is not distinct (author). 

Huanuco: In shrub-wood, Monzon, 900 meters, (Weberbauer 3450, 
type of T. Weberbaueri). Adjacent Colombia. 

Geonoma myriantha Damm. Verb. Bot. Ver. Brandenb. 48: 
1. 1906 (1907); 236. G. paniculigera Mart. var. papyracea Trail, 
Journ. Bot. 16: 326. 1876; fide Burret, with query. 

Type 4 meters high, with several broadly oval interrupted-pinnate 
(3 pairs) leaves to 4.5 dm. broad; petioles to 7 dm. long, the juvenile 
bright brown; rachis about 5 dm. long, nerves (each side) about 28; 
leaf divisions 6-8 cm. distant, the lower 7, medial 12, uppermost 18 
cm. wide; spadices lax, lower branch much branched, all filiform to 
3 dm. long, 2 mm. thick, the lower pedicels closely brown-red verru- 
culose; alveoli 2-3 mm. distant, 5-ranked; male flowers reddish, 
scarcely 2 mm. long, outer segments carinate, inner ovate-lanceolate; 
filaments basally connate, anther cells linear. G. Heinrichsiae Burret 
(Notizbl. Bot. Gart. Berlin 12: 43. 1934) of Ecuador has laxly dis- 
posed alveoli. Type, Ule 5882, from nearly at the boundary, as the 



FLORA OF PERU 347 

variety of Martius' species; the latter, known from southern Colombia 
not very far from the Rio Putumayo, is distinguished apparently 
chiefly by thin, often split lower lip of the flower pits; the species 
may be found to merge. G. laxi flora Mart., 239, from lower Rio 
Purus, etc., is marked by the simple or little separated 2 pairs of leaf 
segments, and especially by the laxly disposed verticils, these with 
intervals 2-3 times longer than the alveoli themselves. 
Loreto (certainly). Adjacent Brazil; Bolivia. 

Geonoma piscicauda Damm. Verh. Bot. Ver. Brandenb. 48: 
122. 1906 (1907); 182. G. Wittiana Damm. I.e. as to spadix (Burret); 
cf. Hyospathe brevipedunculata Damm. 

Low (1-2 meters), apparently branched at base, with simple 
leaves, the sheath at least 1 dm. long, truncate, petioles 16 cm. long, 
3 mm. thick, leaf-blade elongate-obovate, apically bifid, 5 dm. long, 
21 cm. broad, rachis 5 dm. long, primary nerves 30 each side; outer 
spathe 7 cm. long, spadix simple, cylindrical, (peduncle 7-8 [13] cm. 
long), about 2.5 dm. long, 7-10 mm. in diameter, alveoli 7-ranked, 
7-10 mm. distant; male calyx lobes conchiform, carinate, ciliate, 

3 (4) mm. long, corolla lobes ovate, acute, 3.5 (4) mm. long; filaments 
5 mm. long, anther cells 1.5 mm. long; female calyx lobes ovate, acute, 
5 mm. long, corolla tube subglobose, 2 mm. long, lanceolate lobes 

4 mm. long; style trifid, 5 mm. long; fruit subglobose, 6 mm. in 
diameter, 7-8 mm. long, attenuate toward base. Type from Jurua 
Mirim (Ule 5520), nearly at the Peruvian boundary; G. Wittiana 
Damm. I.e. 124 (at least very near, Burret) from neighboring Seringal 
Betem (not Para). 

Peru (no doubt). Brazil. 

Geonoma Poeppigiana Mart. Palm. Orbign. 35. 1847; 263. 

Leaves unevenly pinnate, younger leaves divided into few seg- 
ments, 7.5-10 cm. long, adult a meter long or longer with 6-8 seg- 
ments, these linear-oblong, 4 dm. long or longer, 12-25 mm. wide, 
opposite or subalternate, attenuate into an acutely falcate acumen; 
petioles convex below (as rachis), concave above, sparsely reddish 
tomentose as younger leaves beneath, the peduncles (to 5 dm. long) 
and spadices in part densely so; primary nerves acutely prominent 
only above, the alternating secondary nerves there impressed, stout 
below, in age hard, pale, glabrous; outer spathes linear, 2 cm. wide, 
subcoriaceous; male spadix simply few-branched (rarely entire), the 
acutely mucronate (mucro 4-20 mm. long) branches 3-4 times shorter 



348 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

than the long peduncle, the naked part reddish verrucose-squamate; 
bractlets acute, binate or ternate toward apex of peduncle and at 
base of branches, these 2-5, always simple, each erect; peduncles of 
branches 6-10 mm. long; rachis with the crowded immersed flowers 
cylindric, the series mostly 8, obliquely spiralled, the lip of the 
alveoli entire; male calyx subequaling corolla. Size and color of 
flowers of G. interrupta, calyx segments suboblong, corolla lobes ovate, 
acute, same color and texture, the staminodal cup shortly dentate, 
but unlike G. interrupta in the membranous inner spathe (Martius) ; 
Burret (Notizbl. Bot. Gart. Berlin 11: 321. 1931) applied this name, 
obscurely understood, to the following collections but without new 
characterization. 

Huanuco: Cuchero, (Poeppig, type). Loreto: Yurimaguas, (Kil- 
lip & Smith 28026). Santa Rosa, (Killip & Smith 28729). 

Geonoma pycnostachys Mart. Hist. Nat. Palm. 2: 16, pis. 17, 
21, fig. 1. 1823; 181; 503 (Drude). 

A meter or two high, the remotely annulate stem less than a cm. 
thick; leaves 10-12, simple, spreading, narrowly elliptic or oblong- 
spathulate, one-fifth to one-sixth bilobed, apical divisions broadly 
ovate, inflexed-acuminate, the entire leaves 3-4 dm. long, 10-12 
(-17) cm. wide, petioles 5-8 cm. long; primary nerves each side about 
30; spadices erect-spreading, 12-16 cm. long, ferrugineous lepidote, 
the short peduncle gradually incrassate into the 9-12 cm. long rachis, 
this in flower more than 1 cm. thick, acute; alveoli exserted, approxi- 
mate in 8-10 series, the long-produced lip orbicular; flowers reddish, 
male corolla semitrifid, staminal cup short-funnelform, female scarcely 
half as long as trifid calyx, long exceeded by the cylindric staminodal 
cup, this 6-denticulate; fruit ovoid, obtuse, 6 mm. long, olive-black. 
Male flowers carmine red, subpersisting (collector). F.M. Neg. 18523. 

Loreto: Flood-free terrain, mouth of the Santiago, (Tessmann 
4446). Amazonian Brazil. 

Geonoma Raimondii Burret, Bot. Jahrb. 63: 182. 1930. 

Leaves (in fragment seen) pinnate, the segments rigid, dark green 
above, 4.5-6 dm. long, 8.5 cm. wide; primary nerves 4-9, direct, 1 cm. 
distant, prominent above, beneath much slenderer than the second- 
ary, the intermediate tertiary 12-14, minutely appressed setulose, 
densely pale puncticulate; upper spathe 2.5 cm. above the lower; 
peduncle above shortly ovate-bracted, tuberculate, little shorter than 
the stout spike, this with fertile female flowers to 12 mm. in diameter; 



FLORA OF PERU 349 

alveoli in 7-8 series little inclined, 5 more inclined, lower lobe broadly 
rounded, entire; flowers more than half emerged, sepals and petals 
finally subequal and exceeded by the narrow staminodal tube, this 
obscurely 6-dentate; fruit ovoid, faintly striate, 11 mm. long, 5 mm. 
broad, pericarp thin, endocarp within fuscous, the seed 7X4 mm., 
male flowers unknown. G. Trauniana Damm., 183, from the lower 
Jurud is similar but has many leaf-segments, all except the apical 
with only 2-3 primary nerves; G. Trailii Burret, 183, upper Ama- 
zonian, has leaves with 3-4 pairs of sigmoid leaf -segments, 22 primary 
nerves on each side. 

Amazonas: (Raimondi 978, type). "Palmide." 

Geonoma Spruceana Trail, Journ. Bot. 14: 328. 1876; 256. 
G. juruana Damm. Verh. Bot. Ver. Brandenb. 48: 119. 1907, fide 
Burret. 

Stems 3 or 4 to 5 meters tall (type), 2.5 cm. thick; leaves 3-7 cm. 
long (with petiole), the 2-21 pinnae lanceolate-rhombic, ligulate or 
grass-like, subfalcate, 3-6 (7.5) dm. long, 1-7.5 cm. wide, sub- 
coriaceous, concolor, primary veins 11-15 pairs, the sublinear blades 
always 3-nerved; spadices 2-3 (4) dm. long, reddish or green, branches 
4-8, simple or 2-8-branched below; peduncle to 1 dm. long, com- 
pressed; spathes lanceolate, obtuse, 7 cm. to nearly 2 dm. long, 
coriaceous; flower pits 5-7-ranked; male calyx membranous, seg- 
ments 3 mm. long; staminal tube 3-crenate but splitting into 6 
teeth or 6-lobed from the first; fruit oval or subglobose, 6 cm. X 1.2 
cm. After Trail, who designated several Brazilian variants, with 
doubt as to their taxonomic validity. Burret adopted two: var. 
intermedia Trail, leaves multisected, smaller, segments denser, and 
var. compta Trail, leaves larger, segments laxer. He cites Trail 177 
from Tabatinga, at the Brazilian boundary with Loreto, as typical, 
but the collection is not listed by Trail, I.e., unless it is an error for 
170, 176 or 174; type of Dammer's species (Ule 5744), however, was 
from mouth of Rio Jurud Miry (Mirim) nearly at the Peruvian 
boundary. G. Spixiana Mart., 263, little known, as to Maynas 
(Yurimaguas) specimen by Poeppig may be this; Martius himself 
suggested he might be mistaken in considering it the same as a 
species of Bahia, Brazil. 

Peru (surely; see note above). Amazonian Brazil. 

Geonoma Tessmannii Burret, Bot. Jahrb. 63: 181. 1930. 
Stems several from the same rhizome, 9 mm. thick, lustrous, the 
upper internodes 1 dm. long; sheaths 1 cm. long; petioles 12 cm. long; 



350 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

leaves simple, often laterally cleft, never disjoined, obovate-lance- 
olate, shortly cuneate at base, upper to 17.5 cm. wide, 4.5 dm. long 
to apex, about one-fifth bifurcate, upper margin 14.5 cm. long, lobes 
shortly acuminate, membranous, paler beneath; primary nerves 26 
both sides, slightly curved near rachis, the secondary subequal be- 
neath, the tertiary obscure above, about 8 visible beneath; spadix 1 
dm. long, branches 3; upper spathe (fragment) 7 mm. long; pe- 
duncles about 5 cm. long (2 broadly rounded, nearly truncate bracts), 
at apex 2 mm. thick, as branches densely granulose, these 5-6 cm. 
long, spikes 4.2-4.6 cm. long, dense, pits in 5 or 6 vertical series, 
bilabiate, lower lip inflated, emarginate; male flowers more than 
one-half emerged, 3 mm. long, stamens exserted 4 mm.; staminodal 
tube of female flowers denticulate. 

Loreto: Non-inundated woods, mouth of Rio Santiago, (Tessmann 
4225, type). 

Geonoma Uleana Damm. Verh. Bot. Ver. Brandenb. 48: 122. 
1906; 181. 

Stems scarcely 5 mm. thick, the annuli 1.5-3.5 cm. distant; 
sheaths cylindric, fibrous, oblique, 6 cm. long; petioles about 12 cm. 
long, 2 mm. thick, leaf -rachis 18 cm. long, the oblong leaf attenuate 
to base, apically bifid, the lateral margins quite parallel, 2-2.5 dm. 
long, 7.5-10.5 cm. wide; primary nerves 18-20, early, as rachis, 
floccose-pilose; spadices in the older or lower axils, the included 
simple peduncles 3 cm. long, spathe 5-6 cm. long; spadix 7-11 cm. 
long, 3 mm. thick, alveoli deep, 5 mm. distant, 4-ranked; fruits 
apiculate. Type, Ule 5521, mouth of Rio Tejo, near Peruvian 
boundary. 

Loreto: Iquitos, (Ule 6879). Adjacent Brazil. 

Geonoma Weberbaueri Damm. ex Burret, Bot. Jahrb. 63: 221. 
1930. 

Type 6 meters tall, the caudex about 5 cm. in diameter at the 
great spadix, this a meter and a half long, cernuous, duplicate- 
branched, early furfuraceous; leaves 16 dm. long with petiole, this 
4 dm. long, 12 mm. broad at apex, at apex of sheath 16 cm. broad; 
leaf-rachis above as petioles early broadly canaliculate, finally sub- 
plane, apex not at all produced; segments 1-nerved, linear, attenuate 
but not at all falcate, middle 10-12 mm. wide, diminished upward; 
primary nerve prominent above, acute beneath, marginal secondary 
broader beneath than primary, tertiary partly obvious, intervals 



FLORA OF PERU 351 

beneath minutely white puncticulate; peduncle about 2.5 dm. long, 
somewhat shorter than branched part, rachis 57 cm. long, branches 
often 20, the 12 lower branched, the lowest with about 10 simple 
branches; spikes floriferous to summit, not spine- tipped, to 3 dm. 
long, axis about 8 mm. thick; alveoli (as commonly) 3-verticillate in 
6 lines, in young fruit about 5 mm. distant, upper lip conspicuously 
thin, finally often split unevenly, lower deeply emarginate; sepals 
broadly oblong, 5 mm. long, as also petals, these one-half connate, 
equaled by the staminodal tube, this scarcely incised, subtruncate; 
fruit (young) oblong, mammillate-cusped, pericarp thick; male flow- 
ers fallen. 

Hudnuco: Among evergreens, mostly shrubs, southwest of Mon- 
zon, 2,200 meters, (Weberbauer 3552, type). 

8. TAENIANTHERA Burret 
Reference: Burret, Bot. Jahrb. 63: 267-270. 1930. 

Simulates Geonoma but the free anther cells after anthesis are 
porrect, being, without any angle, in a direct line with the filament, 
and the stigma (known) is applanate. Sepals lanceolate or narrow 
(female flower). Filaments connate only toward base. Spadices 
in known species simple or furcate, pits dense, spathes 2, narrowly 
vaginate, slender. Indubitably the basic character of the species 
assigned by Burret to a distinct group is very real, being clearly stabi- 
lized, probably ancient; in monographic work, to emphasize the char- 
acter, generic status may be preferred, but in floristic, especially when 
the plants concerned possess human interest, a more generally useful 
taxonomy, when possible, is desirable, and subgeneric rank may be 
more feasible. The author himself (267) remarked (trans.): I was 
in doubt if it would not be better to treat this as a subgenus but I 
have found no intermediate development of the anthers in the many 
species of Geonoma examined. In other words, for students whose 
interest prompts them to emphasize specific relationships by many 
generic names this is a "good" genus; others, if they have the pleasure 
of recognizing one of the following closely allied plants, especially 
when not in flower, will refer to it as Geonoma, whether in cultiva- 
tion or native. 

Leaves simple or laterally cleft, basally long-cuneate, nerves ascend- 
ing, straight T. tamandua, T. macrostachys. 

Leaves somewhat pinnatisected, rounded or shortly cuneate at base, 
nerves more spreading, curved. 



352 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Floral pits 5-7-ranked, somewhat inclined. 

Leaf-segments 2-3.5 dm. X 3-9 cm T. tapajotensis. 

Leaf-segments about 2 dm. X 2-5 cm T. oligosticha. 

Floral pits in 10-15 verticils or little inclined series. 
Leaf-segments 3-6 pairs, upper multinerved. 

Leaf -segments distant T. acaulis. 

Leaf-segments partly united T. Lagesiana. 

Leaf -segments many, 1-nerved T. camana. 

Taenianthera acaulis (Mart.) Burret, Bot. Jahrb. 63: 267. 1930. 
Geonoma acaulis Mart. Hist. Nat. Palm. 2: 18, pis. 4, 9. 1823. 

Leaves pinnate, erect or suberect, rosulately congested (with spa- 
dix) on caudex, sometimes as many as 12, a meter long or longer 
including the long slender petiole, the younger fuscous-leprose; seg- 
ments 3-6, the more or less divaricate subequal lateral oblong- 
lanceolate, (1) 2-10 cm. wide, (2) 3-4 dm. long, few- to many-nerved, 
the furcate terminal one much wider; peduncle about 6 dm. long; 
spadix striate, 6-9 dm. long, simple, cylindric; lower spathe scarcely 
1 dm. long, upper 2-2.5 dm. long; spike incrassate-cylindric, 8-13 cm. 
long, 7-10 cm. thick, densely flowered, the deep pits 10-12-ranked, 
the lip emarginate; flowers 4-8 mm. long, male corolla at least twice 
longer than calyx, deeply 3-parted, short or campanulate stamen 
tube extended into broadly liguliform filaments, the tube in female 
flowers urceolate, dentate. Burret noted that the anthers were 
longer than shown in the otherwise good illustration by Drude (Mart. 
Fl. Bras. 3, pt. 2: pi. 121, fig. 1. F.M. Neg. 18501. 

Peru (probably, as at Jurua Mirim, nearly on the Brazilian bound- 
ary). To Colombia; Amazonian Brazil. "Palmilla" (Dahlgren). 

Taenianthera camana (Trail) Burret, Bot. Jahrb. 63: 270. 1930. 
Geonoma camana Trail, Journ. Bot. 14: 324. 1876. 

Caudex (type) 1-1.5 meters tall, 2.5-4 cm. in diameter, densely 
annulate, erect or flexuose; leaves pinnate, glaucous green, 1.5-2.5 
meters long (petiole 6-12 dm. long, sulcate), the 17-21 (31) segments 
subfalcate-linear, 1-2-nerved, or lanceolate-rhombic, the upper sev- 
eral-nerved; spadix 4.5-6 dm. long (peduncle to about 1 dm. long), 
spathes 2-3 dm. long, linear-tubular, spike 1.5-2 dm. long, slender, 
obtuse, the densely congested spiralled pits in about 15 ranks; fruit 
ellipsoid, scabrous, black, 9 mm. long, 6 mm. thick. Type from 



FLORA OF PERU 353 

inundated areas on the Rfos Gavary and Jutabi on the Brazilian 
border. F.M. Neg. 38643. 

Loreto (no doubt; cf. note above). Adjacent Brazil. "Juriti- 
ubim" (Trail). 

Taenianthera Lagesiana (Damm.) Burret, Bot. Jahrb. 63: 268. 
1930. Geonoma Lagesiana Damm. Verh. Bot. Ver. Brandenb. 48: 121. 
1906. 

Type 2 meters tall with ample dark green long-oval more or less 
3-pinnate leaves, the petioles at least 3.5 dm. long, the rachis about 
7.5 dm. long, nerves (each side) about 35; segments 3-8 cm. distant, 
to 4.5 dm. long, long-acuminate; spathe obliquely open, peduncle 
4.5 dm. long or longer, stout, the 2 branches about 2.5 dm. long, 
densely flowered, 1 cm. in diameter; male flowers 3.5 mm. long, calyx 
lobes spathulate, ciliate, 3 mm. long, corolla 3.5 mm. long, lobes 
oblong; staminal tube short, anther cells clavate, 1.5 mm. long. 
Distinct from G. acaulis Mart, by the longer leaves with partly united 
segments (Burret), a character that may prove variable; type, Ule 
5745 from Jurua Mirim, nearly on the Peruvian boundary. 

Peru (certainly). Adjacent Brazil. 

Taenianthera macrostachys (Mart.) Burret, Bot. Jahrb. 63: 
268. 1930. Geonoma macrostachys Mart. Hist. Nat. Palm. 2: 19, 
pi. 20. 1823. 

Acaulescent, the thick caudex bearing 10 or more crowded simple 
or subsimple leaves about 1.5 to nearly 2 dm. long, the younger reddish 
tomentose or furfuraceous, soon glabrous, coriaceous, long-decurrent 
into petiole at least about 3 dm. long, triangular-lanceolate, falcate, 
furcate, the apical segment at least 2.5 dm. long, each part 1 dm. 
wide, primary nerves about 30; peduncles tomentose, about 8 dm. 
long, spadix erect, 1 meter long, upper spathe narrowly tubular, 
about 3 dm. long; spike 1.5-2 dm. long, 8-9 mm. thick, attenuate 
and mostly extended into a filiform sterile portion; pits deep, approx- 
imate, the short lip rounded, bilobate or deeply emarginate; flowers 
3 mm. long, calyx and corolla subequal, male petals free, female 
connate medially, staminal tube lageniform in male flower, deeply 
6-dentate in female flower; fruit dark purple, lustrous, ellipsoid, 7 or 
8 mm. long, perhaps larger (Burret). Burret also observes (I.e.) that 
Drude's plate (for G. acaulis), not that of Martius, depicts correctly 
the similar stamen-character. Since it has been collected nearly 
at the eastern boundary of Peru (Rio Tejo, Ule) it must extend 



354 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

into Loreto or Madre de Dios; however, Burret noted (I.e.) that the 
record of Poeppig from Cuchero does not pertain to this palm. F.M. 
Negs. 18514; 29875. 

Loreto (no doubt; see note above). Brazil; Bolivia; Colombia. 

Taenianthera oligosticha Burret, Notizbl. Bot. Gart. Berlin 11 : 
201. 1931. 

Acaulescent, leaves small, primary nerves about 20, rachis to 
2 dm. long, deciduously fuscous scurfy as peduncles, segments 4-5 
pairs, concolor, narrowly long-acuminate, apical broader than the 
rest, about 5 cm. wide, 2 dm. long (upper margin), primary nerves 8, 
others about 2-2.5 cm. wide, primary nerves 3-5, little narrowed to 
base; secondary nerves little narrower beneath than primary, the 
tertiary more obvious above than below; spadix simple, slender, the 
peduncle much longer than the spike, this 13-17 cm. long, nearly 6 
mm. thick (with flowers), apically (type) spinose; pits approximate, 
in 7 scarcely declined series, 2 mm. distant, lower lip emarginate, 
upper not at all or obscurely produced but forming a very acute 
margin; female sepals somewhat emerging, 2.5 mm. long, petals 
shorter, densely and acutely nervose, staminodal tube 6-lobed. 
Peduncle sometimes longer than 2 meters, petiole 6 dm. long, sheath 
7 cm. long, rachis 28 cm. long (Klug). Near T. acaulis but series of 
flower groups and nerves fewer, spadix smaller (author). 

Loreto: Rio Nanay, Williams 787, type; 740. Mishuyacu, 
Klug 448. 

Taenianthera tamandua (Trail) Burret, Bot. Jahrb. 63: 268. 
1930. Geonoma tamandua Trail, Journ. Bot. 14: 323. 1876. 

Near T. macrostachys; type with leaves 1.5-2 meters long, dark 
green, strongly plicate, primary nerves 25-27 each side, nearly 
straight, petiole 1.5 dm. long, deeply canaliculate; spike white tomen- 
tose, 2.5-3.3 dm. long, to 11 mm. thick, obtuse, the pits 10-11-ranked, 
the lower lip bifid, tomentose. Incompletely known, but, as Burret 
remarks, spike definitely longer, thicker; type from non-inundated 
area at the Rio Javary. F.M. Neg. 38665. 

Peru (probably). Brazil. "Tamandua-pecu" or "-ubim" (Trail). 

Taenianthera tapajotensis (Trail) Burret, Bot. Jahrb. 63: 269. 
1930. Geonoma acaulis Mart, subsp. tapajotensis Trail, Journ. Bot. 
14: 342. 1876. Geonoma tapajotensis (Trail) Drude in Mart. Fl. Bras. 
3, pt. 2: 508. 1882. 



FLORA OF PERU 355 

Habit of T. acaulis but smaller in all parts; petiole, with sheath, 
4.5-5.5 dm. long; leaves scarcely 1 meter long, segments 3-4 pairs, 
broadly ovate-lanceolate, long-acuminate, 2-3.5 dm. long, 3-9 cm. 
wide; spadix 5-6 dm. long, spathes firm-membranous, the lower 5, 
the upper about 13 cm. long; spike 5-7.5 cm. long, 3^4 mm. thick, 
with a mucronate sterile extension 1 cm. long, pits 6-7-ranked; 
flowers (Trail) as in T. acaulis; fruit globose, about as large as a pea 
but immature. Ordinarily 3 pairs of leaf-segments, but sometimes 
1 or 2 narrow segments also, both sides. 

Loreto: Mouth of Rio Santiago, flood-free wood, (Tessmann 4715, 
det. Burret). Amazonian Brazil. 

9. IRIARTEA R. & P. 

Socratea Karst. Linnaea 28: 263. 1856. Iriartella Wendl. Bon- 
plandia 8: 103. 1860. 

Handsome palm marked by partly exposed more or less aculeate 
roots supporting and seemingly bracing the tall stem, this bearing 
a coma of long pinnate leaves with many deltoid unevenly dentate 
or laciniate segments (nerves prominent beneath) and 2-several 
spathes; flowers with both sexes in the same spadix, all the younger 
(corniform) or the intermediate of each three, female, spiralled. Male 
flowers subsymmetric, sepals orbicular, concave, broadly imbricate. 
Ovary 3-celled, stigma early apical or central. Fruit 1-seeded, ovoid 
or globose, stigma position various. Staminodia always present 
(Burret). As to original species see remarks about Socratea sta- 
mens are 12-20, stigma terminal or subterminal, embryo sub-basal, 
slightly below middle of seed, raphe distinctly anastomosed; the 
leaves of /. exorrhiza are merely bifid; in Iriartella the fruit has 
stigma at base, embryo subterminal. Socratea is like Iriartea but, 
especially, stamens many (to 25), stigma in fruit (as embryo) sub- 
terminal, leaves, at least as to original species, equally pinnate, 
spadices solitary, spathes 4-8. Drude treated Socratea as a section 
(1882), later as a subgenus (1889). Bentham and Hooker (1880) 
wrote that Socratea, Iriartella, Dictyocaryum could all be treated as 
subgenera or sections. Recently Burret accepted them. The tax- 
onomy most generally useful would be that of Drude, a conclusion 
reached by Spruce (Journ. Linn. Soc. 11: 133. 1871), who remarked 
that similar differences exist in single genera of exogens as Cordia; 
all these palms (Socratea, Iriartella, Iriartea) are so alike in habit that 
the Indians unhesitatingly give them the same generic name (Pax- 
iuba); and a closer examination reveals so many essential resem- 



356 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

blances that the botanists will probably endorse the opinion of the 
Indians and continue to unite them under one title. To date, verily, 
this is a vain hope. 

Stigmas apical or subapical; stems cylindric; fruit subglobose. 

Embryo apical; stamens 20-25 I. exorrhiza. 

Embryo sub-basal; stamens (known) 15. 

Flowers, fruits in 7 series 7. Weberbaueri. 

Flowers, fruits in fewer series 7. deltoidea. 

Stigmas excentric; embryo lateral. 

Stems ventricose; fruit globose 7. ventricosa. 

Stems cylindric; fruit oblong-linear 7. stenocarpa. 

Iriartea deltoidea R. & P. Syst. Veg. 298. 1798. Ceroxylon 
deltoideum (R. & P.) HBK. Nov. Gen. & Sp. 1: 308. 1815. 

Caudex 20 meters tall or taller; earliest leaves bifid, apical seg- 
ments of adult deeply bilobed or bifid, all glabrous, size of 7. exorrhiza; 
spadix 6 dm. long or longer, branches incrassate, excavate-scrobicu- 
late; male calyx glabrous; spathes 10-12, deciduous; stamens 15; 
fruits subglobose, (stigma remains subapical), yellowish-brown, with 
their flesh mucilaginous (Martius), raphe branches ascending, retic- 
ulately anastomose, embryo sub-basal. Illustrated, R. & P. Prodr. 
pi. 32. 1794; Mart. Hist. Nat. Palm. 1: pi Z5, figs. 3, 7; Weberbauer, 
594, pi. 35. F.M. Negs. 18532; 18532A. 

Huanuco: Pozuzo, (Ruiz & Pavdn, type). Junin: Chanchamayo 
Valley, (Raimondi 2641; 10405). La Merced, (Weberbauer 1839; 
1850). Puno: San Govan, (Lechler 1725; 10405). "Huacra-pona," 
"camona," "morona." 

Iriartea exorrhiza Mart. Hist. Nat. Palm. 2: 36, pis. 33-34. 
1824; 538. Socratea exorrhiza (Mart.) Wendl. Bonplandia 7: 103. 
1860. 

Tall, sometimes 20 meters or taller, cylindric but with many 
partly branched aerial roots at base, at least the exposed portion 
prickly, and crowned with a dense cluster (10-15) of oblong leaves 
4-6 meters long with 15-20 pairs of deltoid or oblique-rhombic seg- 
ments finally 5 dm. long, 1.5-2 dm. wide, shorter but mostly as wide 
toward apex, all radiately about 10-dentate (or entire); spadices 
1-4 below the coma, stoutly peduncled, about 5 dm. long, the 5 or 6 
spathes ventricose-inflated base to apex, before anthesis horn-like, 
4 dm. long, finally caducous, branches nutant, slender, densely flow- 



FLORA OF PERU 357 

ered; flowers yellowish, the larger male nearly 1 cm. long; stamens 
20-25; fruit olive-yellowish, perhaps reddish, 3 cm. long, more than 
2 cm. thick, exocarp indurate, seed 2.5 cm. long, 18 mm. in diameter, 
reticulately rugulose-lineate, stigmatic remains scarcely discernible; 
embryo at depressed apex of seed, barely within the hard horny 
albumen; raphe of many flat filaments radiating from base to apex, 
subramose, anastomosing (Spruce). By Drude (I.e.) as by Trail 
(Journ. Bot. 6 (15): 130. 1870) regarded as a variable species in- 
cluding 7. exorrhiza Mart. var. elegans (Karst.) Drude (539), leaf- 
segments about 5, incised-dentate, spathes mostly 4, fruit finally 
yellow, somewhat smaller, and var. Orbignyana (Orbigniana) (Mart.) 
Drude (540), leaf-segments many, radiately dentate, fruit orange, 
2 cm. long, scarcely 1.5 cm. thick. Both of these have been re- 
corded as varieties or species from adjacent Colombia or (and) 
Brazil so no doubt they occur within Peru. Trail (I.e.) observed 
"a complete transition" between 7. exorrhiza and 7. Orbignyana but 
fide Martius the seed of the latter is only a third as large. The 
curious roots that support or brace the beautiful stems near the base 
are illustrated by Drude in Pflanzenfam. 2, 6: 14, and the flowers 
and fruit in Mart. Fl. Bras. 3, pt. 2: pi. 126. F.M. Neg. 18533. 

San Martin: Tocache, (Poeppig, fide Drude). Colombia; Brazil 
to Guianas. "Huacra," "huacra-pona." 

Iriartea stenocarpa (Burret) Macbr., comb. nov. Iriartella 
stenocarpa Burret, Notizbl. Bot. Gart. Berlin 11: 233. 1931. 

Type 3 meters tall, slender, the sheath (apical part seen) hirsute 
velutinous, the longer trichomes yellowish, retrorse, the shorter whit- 
ish, dense; petioles about 2.5 dm. long (probably longer), terete but 
sulcate-costate, rather closely and retrorsely white-pubescent, the 
rachis densely so with short and long trichomes; segments rhombic- 
cuneiform, decurrent at base 6-6.5 cm., free 15-16.5 cm., pubescent 
as midrib above, more or less glabrescent, conspicuously soft beneath; 
immature fruiting spadix 36 cm. long; peduncle 29 cm. long; spathes 
4, laxly vaginate, shortly and softly pubescent, nearly attaining 
the lower branches; rachis 2.5 cm. long; branches 4-6, simple, curved, 
glabrate, 21 cm. long; fruit laxly spiralled, linear, curved, 10-11 
mm. long, 3 mm. thick (young); perianth cupulate, somewhat en- 
closing fruit-base, 2 mm. high, 3.5 mm. across; sepals erect, broadly 
rounded as the twice as long petals. Distinct from 7. setigera Mart, 
by the larger perianth and the very slender fruit (author); the 
Martius' species probably occurs on the upper branches of Rio Japura 



358 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

within Peru; the exposed roots are sparsely muricate, the slender 
stem 2-3 meters high and prominently tomentose-setose above, the 
leaves tomentulose beneath, the apical segments entire, the lateral 
erose-dentate. 

Loreto: Rios Napo and Amazonas, (Hopp 110, type). 

Iriartea ventricosa Mart. Hist. Nat. Palm. 2: 37, pis. 85, 36. 
1824; 537. 

Caudex 20-35 meters, ventricose medially or lower, annulate 
above at intervals of 2-3 dm. ; roots sparsely aculeate, at least about 
2 meters high; leaves usually 7 or fewer, to 6 mm. long, petioles 
subterete, 1 dm. across, sheath scarcely 3 dm. long; rachis acutely 
trigonous, grayish tomentose; pinnae many, cuneate-flabellate, soon 
glabrate, veins and plicae 10, finally splitting nearly to base, the 
lower laciniae much longer and wider (nearly 2 meters long), the 
upper gradually reduced to hardly 6 dm. long, all laciniae apically 
broadly sinuate-dentate and erose; spadices often binate from the 
same annulus, about 5 dm. long, spreading or subpendulous in fruit; 
peduncle 7 cm. long, abruptly dilated at base; branches at most 3 
dm. long, 8 mm. thick, simple or furcate at base, deeply alveolate 
entire length, spathes many, upper before anthesis, lower tardily 
deciduous; stamens 12-15; fruit globose, minutely umbonate; stigmas 
on fruit clearly excentric; embryo lateral just above middle of seed, 
this with fragile testa, filaments of raphe clearly anastomose. In 
part after Spruce, who noted it as frequent between 700 and 1,200 
meters in the eastern Andean valleys, along with /. exorrhiza Mart, 
and I. deltoidea R. & P. F.M. Neg. 13535. 

San Martin: Tarapoto, (Spruce). Loreto: Yurimaguas, (Poep- 
pig~). "Tarapoto" (Spruce), "huacra-pona," "conduma." Amazonian 
Brazil. 

Iriartea Weberbaueri Burret, Notizbl. Bot. Gart. Berlin 10: 
921. 1930. 

Fragment of rachis (probably upper portion) broadly triangular, 
laterally widely excavated, dorsally nearly plane, opaque, obscurely 
but densely and minutely setulose; segments (apical) about 5 dm. 
long, cuneate, yellowish; sheath strongly puncticulate, dark brown 
within, heavy-ligneous; fruiting spadix pendent, in entirety nearly 
2 meters long; peduncle 3.5 dm. long, 4 cm. thick at apex, terete, 
scars 12, approximate at base, remotely annulate above, the ultimate 
toward apex three-fourths encircling the peduncle; primary branches 



FLORA OF PERU 359 

crowded, 27, the lower 9 furcate, the rest simple, the basal coarsely 
callose within, the upper applanate-rounded; branchlets as seen about 
9 dm. long, below 8 mm. thick, flowers (per scars) ternate, 2 male; 
alveoli 5 mm. long, 2.75 mm. broad, in 7 vertical series, equally 
distant 1.5 cm.; fruit (known) globose, 2.8 mm. in diameter, scarcely 
excentric at minutely mammillate apex, lustrous, light yellowish- 
brown; perianth (in fruit) to 1.5 cm. across; pericarp fleshy squamose 
within; seed about 1.5 cm. in diameter. Distinguished from /. 
deltoidea R. & P. by the much larger fruits, thicker spadix rachis, 
the flowers and fruits in more series; from /. stenocarpa Mart, sim- 
ilarly by the size of fruit, further by the cylindric stem (author). 

Puno: Chunchusmayo, Prov. Sandia, (Weberbauer 1279, type). 
"Morona." 

10. CATOBLASTUS Wendl. 

Acrostigma Cook & Doyle, Contr. U. S. Nat. Herb. 16: 228. 1913. 
Catostigma Cook & Doyle, I.e. 230. 

Reference: Burret, Notizbl. Bot. Gart. Berlin 10: 932. 1930. 

Aerial roots short, stem slender. Flowers remote, all male or 
female in each simple or simply branched spadix; spathes 5 or more, 
the upper longer. Petals free or basally imbricate. Stigma rostrate, 
borne on a columnar style or sessile at the base of the single fertile 
carpel (Catostigma). Stamens 9-15, staminodes none or filiform. 
Taxonomically significant as evidence of relationship, it would be 
more useful to treat the floristic character of Catostigma as sectional 
in definition; the differences in development, too, may be tenuous 
or merely specific; for instance, Acrostigma was based primarily on 
having 3 equal carpels at anthesis, while Catostigma shares with 
Catoblastus the diagnostic character of a single enlarged carpel at 
anthesis. Spadices often several from the same leaf annulus. The 
group (sens, lat.), in view of the variation in distribution of male 
and female flowers, could be referred to Iriartea as a subgenus. 

Catoblastus Drudei Cook & Doyle, Contr. U. S. Nat. Herb. 16: 
233, fig. 41- 1913. Iriartea pubescens Karst. var. krinocarpa Trail, 
Journ. Bot. 5 (15) : 332. 1876. C. pubescens (Karst.) Wendl. Bon- 
plandia 8: 104. 1860, var. krinocarpa Trail ex Drude in Mart. Fl. 
Bras. 3, pt. 2: 543, pi. 127, fig. 2. 1882, not C. pubescens Drude, 
Pflanzenfam. 2, Abt. 3: fig. 48, Gl, G2. 1887, nor Wendland, fide 
Cook & Doyle. Catostigma Drudei (Cook & Doyle) Burret, Notizbl. 
Bot. Gart. Berlin 10: 932. 1930. 



360 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Caudex 3-5 meters tall, 2-3.5 cm. in diameter; leaves pubescent 
below, 13-15 dm. long (sheath puberulent-tomentose, 3-5 dm. long), 
segments oblanceolate, 10-11 pairs, basal 1.75 dm. long, nearly 2 cm. 
wide, medial 3-4 dm. long, 5-7 cm. wide, apical deltoid (connate), 
2 dm. long, 12 cm. wide; spadices 3-5 from one axis with 5 simple 
caudate branches to 5 cm. long, nutant, in fruit pendulous; female 
flowers about 5 mm. long, the tumid fertile ovary equaled by the 
ovate-acute petals, the sterile thick gibbous style little longer; sterile 
anthers (or anther) present, mucronate; fruit about 2 cm. long, 
12 cm. in diameter, stellate puberulent, style rudiment persisting. 
Habit of /. setigera Mart, (see under /. ventricosa) ; when not in 
flower most readily distinguished by the narrower usually more 
numerous pinnae, pubescent below. 

This is a species quite different from Iriartea pubescens Karst. of 
Colombia as beautifully illustrated by Karsten (PL Col. 1, pi. 81); 
type locality, Rio Yavari on the boundary with Brazil, on non- 
inundated terrain. 

Loreto (no doubt). Amazonian Brazil. "Ponilla." 

11. WETTINIA Poepp. 

Wettinella Cook & Doyle, Contr. U. S. Nat. Herb. 16: 235. 1913, 
fide Burret. 

Reference: Burret, Notizbl. Bot. Gart. Berlin 10: 939-942. 1930. 

Aspect of Iriartea but as in Catoblastus the flowers of one sex in 
a spadix, this short, club-like so compactly clothed with the pubescent 
fruits. Petals of male flowers subulate, free. Fertile ovary 1, often 
with 1 or 2 sterile, the much longer style from the base with elongate 
stigmas. Embryo basal. Spadices simple (8-15 for each node) or 
4-5-branched, 4 of the 5 inflorescences maturing (Wettinella), this 
subgroup also with sepals broader than petals (authors). The pollen 
tuberculate, not smooth, as, by error, in lit. (Burret). 

Leaf-segments about 20 pairs. 
Stamens about half as long as petals, sepals (female) at most half 

as long W. augusta. 

Stamens at least as long as petals, sepals (female) at least a third 

as long W. Weberbaueri. 

Leaf-segments about 40 pairs W. maynensis. 

Wettinia augusta Poepp. & Endl. Nov. Gen. & Sp. 2: 39, 
pis. 153, 154. 1838; 939. 



FLORA OF PERU 361 

Similar to W. maynensis, aerial roots interposed; leaf -pinnae 18-20 
pairs; spathes several, yellowish pilosulous; spadices simple, 8-15, 
verticillate; stamens about half as long as petals; sepals of female 
flowers a third to half as long as petals; style and 3 stigmas rather 
slender; raphe fleshy. The finally incomplete spathes remain on 
peduncle as coriaceous sheaths. Kunth (Enum. PI. 3: 109. 1841) 
proposed the species name Poeppigii, perhaps not aware that the 
author of the genus (published in 1836) had added the species later. 
Illustrated, Contr. U. S. Nat. Herb. 16. pi 63. F.M. Neg. 29887. 

San Martin: Tocache, (Poeppig 2058, type). 

Wettinia maynensis Spruce, Journ. Linn. Soc. 3: 194. 1859. 
Catoblastus maynensis (Spruce) Drude in Mart. Fl. Bras. 3, pt. 2: 
544. 1882. Wettinella maynensis (Spruce) Cook & Doyle, Contr. 
U. S. Nat. Herb. 16: 235. 1913. 

Aerial roots strict, subaculeate; stem to 12 meters tall, 3-4 dm. 
in diameter, smooth, annulate; leaves 5 or 6, 4 mm. long, equally 
pinnate, pinnae 38-40 pairs, the lowest reduced, the medial 1 meter 
long, 7.5 cm. wide, finally incised, veins many, nearly contiguous at 
semivertical reduplicate base; spadices 3 (6) in each verticil, 1 female, 
2 male, mature fruit usually on fourth ring below the leaves; spathes 
6, fusiform, appressed pubescent, the incomplete outer 1 dm. long, 
the 3 complete about 3 times longer; male branches 6, 1.5 dm. long, 
densely flowered and straight, in anthesis sepals scale-like, rigid, 
castaneous, free; petals 14 mm. long, subulate; stamens 13 (12-16); 
female branches 5-7, crowded, 2 dm. long, in fruit about half as 
thick, sepals 4-6 mm. long, petals 10-12 mm. long; ovaries 3, united 
and with style, this central, 14 mm. long, villous, the abortive 
ovaries rarely deciduous; stigmas erect, 6 mm. long; fruit dry, gray 
villous, endocarp membranous, seed 22 mm. long, half as thick, 
albumen uniform. Not infrequent both north and south of the Rio 
Mayo, 1,000-1,500 meters, associated with Euterpe and Iriartea ven- 
tricosa according to Spruce, who, I.e. 190 (by lapsus, as shown in 
index) wrote "W. illaqueans." 

San Martin: Tarapoto to Moyobamba, (Spruce, type). Ecuador. 
"Shulla-chonta," "pullo-coroto," "cullo-coroto." 

Wettinia Weberbaueri Burret, Notizbl. Bot. Gart. Berlin 10: 
939. 1930. 

Original tree 10 meters tall, petioles apparently subterete and 
1.5 dm. long; rachis triangled above, furfuraceous, segments seen 



362 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

disposed regularly, narrow, maximum seen 5.4 dm. long, upper some- 
what reduced, strongly drawn together at base, subconcolor both 
sides; male spadix to upper spathe tip 23 cm. long, inner spathes 
finally laciniate, mostly several cm. long, 1.5-3 cm. wide, fourth 
and fifth early complete, thickly inflated-fusiform, tardily split; pe- 
duncle shortly ferrugineous, velvety, finally annulate by the rudi- 
ments of the spathes, these early reddish sericeous strigillose; spadix 
fusiform, in flower 7 mm. thick at base, attenuate, the male flowers 
approximate, 1 cm. long; sepals as petals linear, the former 1, the 
latter 8-9 mm. long, acuminate, the 12 stamens at least as long; 
female spadix similar; sepals 6-10 mm. long, 4-5 mm. broad at base, 
acute, petals to a third longer; fertile carpel yellowish villous; stigmas 
at least 5 mm. long, 1.75 mm. thick. Differs from W. augusta Poepp. 
& Endl. in subequal stamens and petals, stouter stigmas, probably 
also in reddish indument and leaves (author). 

Puno: Prov. Sandfa, Chunchusmayo, 900 meters, (Weberbauer, 
no number, type). 

12. CEROXYLON Humb. & Bonpl. 

Reference: Burret, Notizbl. Bot. Gart. Berlin 10: 841-854. 1929. 

Tall, slender or robust, the columnar stem more or less wax- 
whitened, annulate, crowned with many multipinnate leaves their 
rigid fleshy or coriaceous segments ensiform, and at their nodes elon- 
gate, much-branched spadices, the polygamous or monoecious flowers 
on different branches. Spathes 3 or more, the upper complete, cori- 
aceous. Pedicels short, often arcuate or bent. Calyx minute, petals 
ovate- or subulate-lanceolate, acuminate, more or less adnate; male 
flowers with 9-15 stamens; female with 9-12 staminodes, 1 sterile 
stamen. Fruit with 1 or 3 fertile cells, residual style at base, pericarp 
smooth or granulate; seed erect, raphe ascending from base, branched 
reticulate, albumen uniform. Seedlings develop rootstocks. 

The type species is the Colombian C. andicola HBK., the most 
famous wax palm of the Andes. The fruit of C. Schultzii Burret of 
Colombia, as probably that of other palms, is sought by parrots 
(Arnold Schultze). 

KEY (after Burret) 

Calyx teeth at least half as long as corolla tube; leaf -segments regu- 
larly disposed. 

Spadix large, robustly 3-much-branched C. latisectum. 

Spadix small, the branches most simple C. Weberbaueri. 



FLORA OF PERU 363 

Calyx teeth minute, many times shorter than corolla; leaf -segments 

aggregate. 
Leaves to 6 dm. long; branched part of spadix 2.5 dm. long. 

C. crispum. 
Leaves 2 meters long; spadix ample C. verruculosum. 

Ceroxylon crispum Burret, Notizbl. Bot. Gart. Berlin 10: 849. 
1929. 

A small species with, in general, about the same characteristics 
and measurements (except the floral) as C. verruculosum except as 
follows: petioles 2-2.5 dm. long, 1 cm. wide at apex, nearly plane 
above, convex beneath; segments 40-50, mostly 2-3-aggregate, 
crisped (direction various), lowest 15-18 cm. long, apical about 12 
cm. long, maximal medial 21 cm. long, little wider than 1.5 cm., all 
attenuate to oblique apex, very rigid, the midnerve above and sur- 
face beneath furfuraceous; peduncle spathes 5, lower 3 cm. broad; 
branches about 30, branchlets 10, early sparsely furfuraceous; young 
fruits densely, distinctly verruculose; calyx tube 0.5 mm. long, teeth 
minute, obscure; corolla tube at least 1.5 mm. long, teeth acuminate, 
striate, 3 (-4) mm. long. These palms, together with Geonoma mega- 
lospatha Burret, grow at the highest boundary for Peruvian species 
(Burret). 

Huanuco: Chinchao, Acomayo, 2,800-2,850 meters, (Weberbauer 
6825, type). 

Ceroxylon latisectum Burret, Notizbl. Bot. Gart. Berlin 10: 
844. 1929. 

Trunk known to attain 11 or 12 meters, 2.5 dm. in diameter a 
meter above the terrain, petioles 8 dm.; leaves 1.75 meters, the seg- 
ments disposed evenly in one plane; rachis of apical part of leaf 
acute above, plane or nearly beneath, fuscous furfuraceous as the 
segments finely beneath, all yellowish-green above, the latter at apex 
13 cm. long, 1 cm. wide, 24 cm. below, about 4 cm. wide, those of 
the upper third subequal, broader, abruptly contracted, the basal 
more approximate, 3.3 dm. long, nearly 5 mm. wide; peduncle stout, 
13 dm. long, branched part 7.5 dm. long, the divaricate branches 
3-many times branched; spathes many, lowest ligneous, applanate, 
at base 8.5 cm. broad, 2.5 dm. above the base 10.5 cm. broad; lowest 
primary branches nearly 4 dm. long, 5 cm. pedunculate, intermediate 
branches shorter, slenderer, flexuose; fruit globose, smooth, 2 cm. in 
diameter without perianth; calyx teeth and tube each 0.75 mm. long, 



364 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

equaling corolla tube, its lanceolate acuminate teeth about 3 mm. 
long; seed globose, 13 mm. in diameter. Fruit at maturity fiery red 
(collector). Illustrated, Weberbauer, page 504, pi. 19. 

Amazonas: Congon to Conila, 2,950 meters, (Weberbauer 7161, 
type). 

Ceroxylon verruculosum Burret, Notizbl. Bot. Gart. Berlin 10: 
850. 1929. 

Trunk 7 meters tall, 2 dm. in diameter, smooth, green but thinly 
waxed; sheaths 12 dm. long, petioles 5 dm. long, leaves 2 meters long, 
the segments aggregate, yellowish-green above, very finely white fur- 
furaceous beneath as the spathes and spadix-rachis, the former com- 
pressed basally, 8.5 cm. broad, at least 18 cm. long, probably longer; 
peduncle long, stout, transversely oval, spathes many; branches 
divaricate, twice branched, the primary nearly 5 dm. long, the many 
slender branchlets flexuose, the rachis at base 2 cm. thick; fruits laxly 
spiralled, globose, without perianth 1.8 cm. in diameter, minutely but 
distinctly and densely verruculose; perianth (in fruit) with low calyx, 
the teeth obscure, the corolla tube cylindric, 2 mm. high, teeth 
acutely acuminate, 4 mm. long; pericarp thin, one third mm. thick; 
seed globose, 12 mm. in diameter. Some leaves have segments only 
about 1 cm. wide, others, larger, one with terminal, regularly disposed 
segments; some segments 38 cm. long are 2.5 cm. wide; there are also 
color differences and some segments, nearly 7 dm. long, 3.5 cm. wide, 
are bidentate and in groups of 2 or 3 (author) ; these are all probably 
normal variations and belong to the species. 

Junin: Huacapistana, 2,700 meters, (Weberbauer 2284, type). 

Ceroxylon Weberbaueri Burret, Notizbl. Bot. Gart. Berlin 10: 
848. 1929. 

Type 5 meters high; sheaths lacerate, the short petioles canalic- 
ulate above, thin-margined, 1.5 cm. broad at apex; rachis margins 
excavated, acute above, pale furfuraceous beneath as segments, these 
rather subevenly disposed, the lower very narrow, shorter, the apical 
2 dm. long, about 1 cm. wide, the medial 5 dm. long, 8 mm. wide; 
spadix 11 dm. long, branched part 2.7 dm. long, visible spathes 4, 
the lower dilated at base, ligneous, the upper thin; peduncles com- 
pressed above to 7 mm. broad at apex; branches 40-50, only the lower 
sparsely branched, all glabrous; fruit obovoid, with perianth at least 
13 mm. long, 10 mm. thick, glabrous, rugose but not all verruculose; 
calyx cylindric, tube 0.5 mm. long with the small teeth to one-half 



FLORA OF PERU 365 

as long or subequaling corolla tube, this with triangular or sub- 
acuminate petals. 

Puno: Prov. Sandia, Yuncacoya, 1,800 meters, (Weberbauer 1157, 
type). 

13. WENDLANDIELLA Damm. 

Sheath cylindric, obliquely open, leaves simple or paripinnate, 
petiole triangular, pinnae 2-3 each side, lower subopposite, the upper 
4-, remaining 3-nerved. Inflorescence subdigitate, interfoliaceous, 
spathes 2, cylindric, obliquely open, nearly covering peduncle, 
branches filiform, densely flowered. Calyx of female flowers 3-fid, 
the orbicular lobes gibbous as the similar corolla segments, these 
twice as large. Staminodes 3, minute, subulate. Ovary sessile, sub- 
globose, 3-celled, stigmas 3 (-4) -reflexed, 1-2 cells sterile, 1 pendu- 
lous ovule in each cell. Probably related to Chamaeodorea Willd. but 
distinctive in flower segments and branched inflorescence (author) ; 
flowers glomerulate in long series (Burret). 

Honors worthily the Director of the Garden at Herrenhausen, 
Hannover, great admirer and student of palms. 

Leaves simple W. simplicifrons. 

Leaves pinnate. 

Segments 2 each side W. polyclada. 

Segments 3-4 each side W. gracilis. 

Wendlandiella gracilis Damm. Bot. Jahrb. 36: Beibl. 80: 32. 
1905. 

Stem (type) only 4 cm. thick, to 1.5 meters tall, leaves remote, 
sheath 5 cm. long, petiole to 6 cm. long, lower pinnae 16-17 cm. long, 
1.6 cm. wide, the medial 18 cm. long, 2 cm. wide, the uppermost 2 
dm. long, 2.5-3 cm. broad, elongate-obovate-lanceolate; lower spathe 
8 cm. long, upper tubular, peduncle canaliculate above, 13 cm. long, 
apically ventricose, subdigitate, the crowded flowers scarcely 1 mm. 
across; male calyx as corolla lobes 3, connate at base; stamens 6, 
anthers affixed dorsally, introrse; stigmas 3; fruit orange-red, oval, 
8 mm. long, nearly 6 mm. thick, pericarp thin, fibers none, the 
2 larger raphe branches 3-furcate, calyx (fruiting) 1.5 mm. across, 
lobes broadly rounded, petals ovate; basal sterile carpels 2; stigmas 
small. Description of male flowers and fruits after Burret from 
Williams and Killip and Smith specimens. 



366 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Loreto: Huimbaio, Pampa de Sacramento, between Rio Ucayali 
and Rio Huallaga, (Huber 1541, type). Lower Huallaga, Williams 
4566. Puerto Arturo, (Killip & Smith 27775). 

Wendlandiella polyclada Burret, Notizbl. Bot. Gart. Berlin 11: 
203. 1931. 

Caespitose, dioecious, the stem of type 12 dm. high, 5 cm. in 
diameter, the leaves about 6; sheath green, even the margins, efi- 
brose, obliquely truncate, glabrous, long-striate; petioles 14 cm. long, 
rachis 4-8 cm. long, segments 2 pairs, concolor, thin-membranous, 
narrowly acuminate, the larger lower leaves 27 cm. long, 7-8 mm. 
wide at base, widening to 3 cm., acuminate, the apical about 3 dm. 
long, at least 1.5 cm. wide at base, 4.5-5 cm. at upper third; spadix 
above sheath 3 dm. long, duplicate-branched; visible spathes 2, largely 
covering peduncle, little dilated; branches 4, these with 4 attenuate 
branchlets 4.5-9 cm. long; male flowers in about 3 vertical series, 
above often 2, at most 3 mm. across; calyx nearly 1.5 mm. across, 
the 3 lobes suborbicular; filaments connate at dilated base; pistil 
with 3 stigmas; fruits solitary, spiralled, oblong (juvenile); stigmas 
elongate, sulcate above, at base of fruit the sterile cells 2; calyx 
fleshy in fruit, lobes rounded, corolla larger, lobes lacerate. Fruiting 
spadix (same collection) detached. 

Loreto: Rio Itaya, Soledad, (Tessmann 5240, type). 

Wendlandiella simplicifrons Burret, Notizbl. Bot. Gart. Berlin 
11:316.1932. 

Arundinaceous, 2-3 meters tall, unique at least in Peru by the 
simple leaves, these glabrous (as densely striate, obliquely truncate 
unevenly dentate sheath), nearly 4 dm. long (petioles 7-8.5 cm. long), 
thin, dark green, subconcolor, not at all produced at base, shortly 
acute, about 14 cm. wide at the bifurcation; rachis 17 cm. long, lobes 
23 cm. long, inner margin nearly direct, outer curved to apex, ob- 
scurely, remotely appressed denticulate; spadix solitary from leaf- 
sheath, in fruit 29 cm. long, simply branched; peduncle 21 cm. long, 
robust, without trace of spathes, medially 4 mm. thick; rachis 6 cm. 
long, branches 7, spiralled, upper spreading, lower reflexed, to 5.5 
cm. long, acute; flowers vertically disposed; fruit red, suboval, 1 cm. 
long, 6 mm. broad, fleshy, obtusely rounded, subacute at base, finely 
rugulose, efibrous; calyx (fruit) 1.5 mm. across, obscurely lobed, stig- 
mata basal, seed obovoid-oblong, 8.5 mm. long, 5.5 mm. thick; 



FLORA OF PERU 367 

raphe branches 3 or 4, bifurcately few-branched; embryo medial- 
lateral. 

Junin: Puerto Bermudez, (Killip & Smith 26515, type). 

14. CHAMAEODOREA Willd. 
Nunnezharia R. & P. Prodr. 137. pi 81. 1794. 
Reference: Burret, Notizbl. Bot. Gart. Berlin 11: 724-767. 1933. 

Stems solitary or several, slender, annulate, the leaves simple and 
bifid or variously pinnatisect. Sheath tubular. Spadices in or below 
the leaves, simple or branched; spathes 3 or more, enclosing peduncle, 
cleft, more or less persistent. Bracts, bractlets none. Flowers dioe- 
cious, sometimes more or less immersed; calyx cupulate, entire, 3- 
lobed or -parted, petals free to somewhat connate, valvate or (female 
flower) sometimes imbricate; stamens 6, anthers included. Ovary 
3-celled, stigmas minute, recurved, ovules solitary, basal, erect. 
Staminodes sometimes lacking. Fruit of 1 (2-3) globose or oblong 
carpels, stigmas basal, pericarp coriaceous or fleshy, scarcely fibrous; 
raphe branches obscure, endosperm cartilaginous, uniform. The 
dioecious flowers and absence of scars after they (as fruit) have fallen 
apparently mark the group, sens. lat. Name conserved; Kuntze's 
transfer (Rev. Gen. 2: 729-730. 1891) of names, including Morenia 
species, to the earlier cognomen of Ruiz and Pavon, "corrected" by 
him to Nunnezharoa as after Nunez de Haro, Archbishop of Mexico, 
being without further interest is not listed. 

The small flowers of C. fragrans (R. & P.) Mart, ("sangapilla") 
exhale a wonderful fragrance which spreads a long distance in the 
mon tafia (Ruiz and Pavon); for this it is cultivated (Weberbauer). 
The shoots, called palmitos, and spadices as in similar groups are 
eaten cooked or in salads and may be excellent. Sap of ripe fruits is 
highly irritating. 

Flowers not or little immersed; fruits on slight not deep depressions; 

perianth usually ecostate. 
Leaves 6-multi-pinnate. 

Leaf segments many pairs, 1-2.5 cm. wide. 

Lateral segments 1-nerved C. angustisecta. 

Lateral segments 2-3-nerved C. pauciflora. 

Leaf segments 6-8 pairs, 7 cm. wide C. boliviensis. 

Leaves simple or segments 2, rarely 5. 



368 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Leaves often with 2 segments C. integrifolia. 

Leaves merely deeply bifid C. Pavoniana, C. fragrans. 

Flowers (unless C. depauperata) well-sunken, as fruit; perianth, at 

least female, costate. 
Leaves pinnate. 
Segments 2-3 pairs. 

Male calyx 1 mm. long, 3 mm. across C. depauperata. 

Male calyx minute, annulate C. lanceolata. 

Segments about 6 pairs C. Herrerae. 

Leaves subsimple, deeply bifid C. geonomoides. 

Chamaeodorea angustisecta Burret, Notizbl. Bot. Gart. Ber- 
lin 11: 318. 1932. 

Caudex 1 meter high, 2.5 cm. in diameter, internodes 5 mm. long; 
sheath glabrous, long-costate-striate; petioles robust, 1 meter long; 
leaves 1.5 meter long, segments many, evenly disposed, all except 
the apical 1-nerved, narrowly linear, somewhat falcate, narrowly 
acuminate, concolor, thin, the apical 2-3 times wider (almost 2 cm.), 
the primary nerves 2-3, maximum seen 2.5 cm. wide, 3.5 dm. long, 
the smaller nerves few, very fine, conspicuous both sides; male spa- 
dix 8 dm. long; peduncle nearly enclosed in spathes, these 6, diameter 
of sheath but 4-7 mm. in diameter, upper the longest; branches about 
12, simple, to 1.5 dm. long, slender; male flowers solitary, laxly 
spiralled, calyx high-cupulate, the 3 lobes enervose as the valvate 
petals; stamens 6; fruiting spadix orange-red, 6 dm. long, pendent, 
peduncle 4 dm. long, spathe scars 8, lower approximate, rachis 1 dm. 
long, branches to 15, porrect, 1-1.5 dm. long, fruits solitary, laxly 
disposed (perianth 4 mm. across), oblong, black, fleshy, 16 mm. long, 
half as broad, epicarp and mesocarp fragile, endocarp fibers rather 
firm in one plane. Leaves resemble those of Morenia linearis R. & P. 

Junin: Colonia Perene", (Killip & Smith 25091, type; also 24926; 
25103). Ayacucho: Kimpitiriki, (Killip & Smith 22864). 

Chamaeodorea boliviensis Damm. Notizbl. Bot. Gart. Berlin 
6: 262. 1915; 756. 

Slender trunk nearly 1 cm. in diameter, sheaths cylindric, petioles 
9-12 cm. long, leaf-rachis 4 dm. long, 2-3 mm. thick, segments re- 
mote, 6-8 pairs, the apical confluent, the lateral sigmoid, attenuate 
to base, long-acuminate, about 2.5 dm. long, 7-7.5 cm. wide, primary 
nerves 5, secondary binate between two of these; peduncle of female 



FLORA OF PERU 369 

spadix at least 3.5 dm. long, 3-5 mm. thick, closely bracted, these 
cylindric, apically 4-7-branched, the branches densely floriferous 
nearly to base, 13-16 cm. long; fruit oblong, black, 7 mm. long, 5 mm. 
thick; embryo slightly supramedial. Somewhat resembles C. lance- 
olata (R. & P.) Kunth with fewer leaf-segments, subequally nerved, 
merely acuminate, the petiole and rachis stouter (author). Type 
from Alto Rio Acre, vicinity of Lojija, (Ule 115b), near boundary. 
Madre de Dios (surely). Adjacent Bolivia. 

Ghamaeodorea depauperata Damm. Notizbl. Bot. Gart. Berlin 
6: 263. 1915. 

Type at most a meter tall, petioles 1.5 dm. long, rachis slender, 
nearly 3 dm. long, lateral segments remote, subopposite, 2-3 each 
side, lanceolate-falcate, long-acuminate, 2-2.5 dm. long, 22-33 mm. 
wide, primary nerves 4-5, the secondary as many, apical segments 
broadly elongate-lanceolate, at base 11, at middle 6 cm. wide, 32 cm. 
long, primary nerves 8-9, secondary as many; male peduncle bracted 
to apex, the 7 slender orange branches 17-20 cm. long, the rather 
densely disposed flowers a little immersed; sepals 3, 1 mm. long, 
3 mm. broad, rounded as the 3 petals, these 2 mm. long and broad; 
female flowers unknown. The label data, Alto Acre, Sao Francisco, 
(Ule 9155b), may be incorrect (author); if not, the locality is at the 
Brazilian boundary. 

Madre de Dios (probably) . Adjacent Brazil or Bolivia. 

Chamaeodorea fragrans (R. & P.) Mart. Hist. Nat. Palm. 2: 
4, pi. 3, figs. 1, 2. 1849; 730. Nunnezharia fragrans R. & P. Syst. 
Veg. 1:294. 1798. 

Stems scarcely 1.5 cm. thick, flexuose, inclined, rarely erect; 
leaves at most 6, glabrous, simple, deeply bifurcate, the divisions 
3.5 dm. long, not over 3.5 cm. wide, linear-rhombic, outer margin 
apically crenate-incised, nerves 12 each side, acute; spadices some- 
what pendulous, solitary, 2-4 dm. long, very slender, the usually 
2-6 branches about 2 dm. long; spathes 3, membranous; calyx trifid; 
corolla 3 mm. long, fleshy, female 3-parted; 2 aborted ovaries in- 
durate at base of fertile; anthers adnate below; fruit black, lustrous, 
12 mm. long, mesocarp insipid, seed black, raphe laxly reticulate 
(Spruce). Forms large beds under the tall trees and perfumes the 
forest far and wide with its orange-colored male flowers, especially 
in August (Spruce). Illustrated, Oerst. L'Amer. Central 14. pi. 5, 
figs. 18-20. F.M. Neg. 18539. 



370 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

San Martin: Tarapoto, 600-1,000 meters, (Spruce 65}. Rio Mayo, 
Moyobamba, (Weberbauer 4554; 585). Huanuco: Pozuzo, Cuchero, 
(Ruiz & Pavdn, type). Loreto: Shapaya on the Huallaga, (Vie 6850). 
Bolivia. "Chutasllium" (Ruiz & Pavon), "sangapilla" (Spruce). 

Ghamaeodorea geonomoides (Spruce) Drude in Mart. PI. Bras. 
3, pt. 2: 531. 1882; 757. Nunnezharia geonomoides Spruce, Journ. 
Linn. Soc. 11: 122. 1871. 

A meter or two tall with extremely slender, glabrous stem; peti- 
oles long-sheathed; leaves cuneate-obovate, deeply bifid, the furcate 
portions subfalcate, scarcely acuminate; veins 10 pairs; spadices 
nearly 5 dm. long, simply branched; peduncles arcuate-pendulous, 
2.5 dm. long; branches slender, flexuose, about 3 dm. long, alveolate, 
the 1-flowered pits oblong-cymbiform, more or less remote; spathes 
2, longer than peduncle at anthesis, narrowly fusiform before open- 
ing, finally fibrous, deciduous; calyx semi-immersed, shortly 3-lobate, 
transversely rugulose, lowest lobe slightly higher, persisting, the 3 
suborbicular petals 3 times as long; anthers erect, turgid, deeply 
emarginate both ends. Because of the alveolate rachis and the lack 
of female flowers in the type the author questioned the genus. 

San Martin: Mount Campana, 1,000-1,300 meters, (Spruce 67, 
type). Moyobamba, (Weberbauer 4628; 585). Junin: Chancha- 
mayo, (Raimondi). Colonia Perene", (Killip & Smith 24929). 

Chamaeodorea Herrerae Burret, Notizbl. Bot. Gart. Berlin 11 : 
748. 1933. 

Leaves small, the blades 4-4.5 dm. long, thin, the segments about 
6 pairs, regularly disposed, the lower more approximate, all lance- 
olate-sigmoid, acuminate; apical segments nearly two times wider 
than the rest, the maximum seen about 2.3 dm. long, 3.5 cm. wide; 
larger nerves in each segment (apical excepted) 5; fruiting spadix 
high, 2.5 dm. or more pedunculate; spathes narrowly cylindric; 
branches about 6, a dm. long; fruit insertions laxly spiralled, oblong 
or narrow, somewhat excavated, perianth closely costate, the short 
calyx 3-lobed, the outer petals imbricate; fruit obovoid, 9 mm. long; 
embryo dorsally at slightly above the middle of the seed; male 
spadix peduncle slender, branches 10, 1 dm. long, flowers rather 
densely spiralled, immersed, subglobose, the lower smaller; petals 
laterally free; anthers oblong, cells connate. Distinguished from re- 
lated species by the slender finely divided leaves; nevertheless, per- 



FLORA OF PERU 371 

haps closest to C. geonomoides (Spruce) Drude with simple leaves 
(author). 

Cuzco: Valle de Santa Ana, (Herrera 3641, male and female 
spadices, type). "Cuyuli." 

Chamaeodorea integrifolia (Trail) Damm. Verh. Bot. Ver- 
Brandenb. 48: 125. 1906; 735. Morenia integrifolia Trail, Journ. 
Bot. 14. 331. 1876; var. nigricans Trail, I.e. M. Lechleriana Wendl. 
ex Damm. Card. Chron. ser. 3, 36: 246. 1904. 

To about 1.5 meters high, the simple submedially bifid leaves 
sometimes as long (petioles 3-5 dm. long, sheath 1.5 dm. long), nerves 
12-20 pairs, the two (rarely 3-5) segments broadly lanceolate, acumi- 
nate, to 1 dm. wide at base, the outer margin remotely denticulate; 
spadices (male) single, verticillate in the axils, slender, 5-6 dm. long, 
peduncles 3 dm. long, spathes 4-5, imbricate; flowers remote, solitary, 
spiralled; calyx cupulate, petals orbicular, deep green; filaments 
slender, anthers free. Burret noted that in spite of the difference 
in the number of primary nerves ex char, be believes M. Lechleriana 
to be the same and that a mixture of material may have occurred. 
Wood soft; flowers dark orange below, greenish above, violet-scented 
(Tessmann). Type of a variety with 3-5-parted leaves (dried dark) 
from Tabatinga at the boundary with Peru; also from Jurua Mirim 
and mouth of Rio Tejo, nearly at the boundary (Ule). 

Loreto : Mouth of Santiago, (Tessmann 4558) . Puno : San Govan, 
(Lechler, type, M. Lechleriana). To Colombia and Amazonian Brazil. 
"Sangapilla." 

Chamaeodorea lanceolata (R. & P.) Kunth, Enum. PI. 3: 172. 
1841; 755. Martinezia lanceolata R. & P. Syst. Veg. 297. 1798. 

Graceful, the stem often a little flexuose, a meter or two tall, 2-3 
cm. thick, the few (4-6) equally (mostly 5) pinnate leaves 1 meter 
long borne laxly; leaf -segments lanceolate, sometimes narrowly, 
the lateral usually 2-3 dm. long, 2-4 cm. wide, subequally multi- 
nerved, the terminal ovate-lanceolate, 7-9 cm. wide, all acuminate; 
spathes 4 or 5, long-tubular; peduncles at least 2.5 dm. long, spadices 
3-5 dm. long, the many branches of the male 1.5-2 dm. long, nutant- 
arcuate, the fewer female suberect; male flowers about 4 mm. across, 
calyx minute, annulate, petals cohering apically, the adnate stamens 
with erect anthers; female flowers depressed, obtuse; berry maturing 
black in the yellowish branches, 12-15 mm. long, ellipsoid. Illus- 



372 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

trated, Mart. Palm. Orbign. pi. 16 A and Drude in Mart. Fl. Bras. 3, 
pt. 2: pi. 125. F.M. Neg. 18542. 

Huanuco: Chinchao, Cuchero, (Ruiz & Pavdn, type). El Valle 
de Monzon, 700-900 meters, (Weberbauer, 593; 594). Rio Acre: 
Seringal Auristella, (Ule 9152). Bolivia; Brazil. 

Chamaeodorea pauciflora Mart. Hist. Nat. Palm. 2: 5, pi. 3, 
fig. 3. 1823; 735. Morenia (?) pauciflora Drude in Mart. Fl. Bras. 3, 
pt. 2: 526. 1882. C. amazonica Damm. Notizbl. Bot. Gart. Berlin 6: 
263. 1915, fide Burret. 

Low, the petioles 2 dm. long, leaf-rachis 4.5 dm. long, segments 
of the ovate leaves opposite or subopposite, 9-10 pairs, 2-4 cm. 
distant, not decurrent, linear-lanceolate, sigmoid, about 2 dm. long, 
1-2 cm. wide, lateral 3-, apical 4-6-nerved; peduncle of the simple 
female spadix 3 dm. long, spathe cylindric, obliquely open, rachis 
about 3-5 dm. long, gradually attenuate toward apex, flowers 3- 
4 mm. distant, slightly immersed, calyx as corolla lobes ovate, 
acute, 1.5 mm. long, 2 mm. broad, the former 3, dorsally puber- 
ulent, the latter 2, glabrous; staminodes minute, ovary subglobose, 
stigmas sessile. After Dammer; Drude (I.e. 527), who also knew 
only the female flowers, suggested that the species apparently ex- 
hibits the characters of both Morenia and Chamaeodorea. The de- 
scription of the Brazilian type agrees essentially : spadix rachis 3 mm. 
thick, 2.5 dm. long, laxly scrobiculate, calyx urceolate, trifid, petals 
suborbicular, connivent, fruit ellipsoid, olive-black, 10 or 11 mm. 
long, 7-8 mm. in diameter, sarcocarp thin, embryo sublateral, in- 
clined toward base. Type of Dammer's species from Jurua Mirim 
(Ule 5595), nearly at the Peruvian boundary. F.M. Neg. 18545. 

Loreto (no doubt). Amazonian Brazil. 

Chamaeodorea Pavoniana Wendl. ex Damm. Gard. Chron. 
ser. 3, 36: 246. 1904; 731. C. cataractarum Hort. not Liebm. ex 
Mart., 1849. C. Ruizii Wendl. I.e.? 

Stems proliferous, 3-4 meters high, scarcely 2 cm. thick; petioles 
1-4 cm. long; leaves 5-8, simple, cuneate-obovate, more than medi- 
ally furcate, 4-4.5 dm. long (or 3-3.5, fide Burret), the divergent 
divisions broadly lanceolate, acuminate, the inner margin straight, 
the apex slightly serrate-crenate; primary nerves 10-11 pairs; spa- 
dices few-branched; fruit rounded-oblong, the seed rounded-ovate, 
9 mm. long, 8 mm. broad (Dammer). Near C. fragrans ex char, 
and perhaps doubtfully distinct. However, Burret (I.e.) noted that 



FLORA OF PERU 373 

C. Ruizii Wendl. is not identical with C. fragrans but extraordinarily 
near C. Pawniana; the leaves are little larger and the primary nerves 
15-16; only known from sterile material. 
Peru (fide Linden). 

15. MORENIA R. & P. 

Reference: Burret, Notizbl. Bot. Gart. Berlin 13: 332-339. 1936. 

Character of Chamaeodorea Willd. but calyx trilobed, the seg- 
ments spreading in the male flowers, the corolla stellate, the lobes 
nearly free. Stamens erect (Drude). Included in the former group 
by Bentham and Hooker and Dammer, but this is the earlier name. 
In view of the floral variations now known within the genus, sens, 
lat., only sectional divisions appear to be feasible. Drude's char- 
acter is questionable, as the anthers may be erect (if included) in 
species accepted by him as Chamaeodorea. Dr. D. Gabriel Moreno 
was a physician of Lima, well-informed in botany and physical 
science (authors). Martinezia R. & P. (Prodr. Fl. Peruv. 148. pi. 82. 
1794) was named in honor of an archbishop of Colombia; Burret 
(Repert. Sp. Nov. 34: 169-170. 1933) showed, apparently correctly, 
that the name was based on a mixture of materials, including, 
among other groups, Morenia; in any case, it is expedient to follow 
Burret's decision and drop the name. The Willdenow name could 
be conserved. 

Leaf segments lanceolate M. Poeppigiana, M. fragrans. 

Leaf segments linear M. macrocarpa, M. linearis. 

Morenia fragrans R. & P. Prodr. Fl. Peruv. 140, pi 23. 1794. 

A slender graceful species, the type observed as 2 meters tall with 
4 or 5 multipinnate leaves a meter long or longer; stem arundinaceous, 
slightly flexuose, more or less conspicuously annulate, scarcely 2.5 
cm. thick; pinnae lanceolate, divaricate, acute, 1.5 dm. long, medial 
wider than 5 cm., midnerve prominent beneath, secondary 4-6, pale; 
spadices clustered, the female simply branched, to 5 dm. long, the 
male nutant. Fruit, fide Orbigny, ellipsoid, black, 6-8 mm. long. 
According to Martius in note under his M. Poeppigiana the leaf 
pinnae are somewhat broader, 4 dm. long, nearly 5 cm. wide, a 
little more unequal and falcate-acuminate, spathes of the male spadix 
larger, and fragrant flowers a little longer, yellowish, and, finally, 
the fruit ovoid. Illustrated, Mart. Palm. Orbign. pi. 3, fig. 1 (plant) ; 
16c (stem, fruit, leaf); Bot. Mag. pi. 5492 (fide Hooker). 



374 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Amazonas: Chachapoyas, (Mathews 31082}. Huanuco: Pozuzo, 
Muna, (Ruiz & Pavdn, type). Bolivia. "Siasia" (Ruiz & Pavon), 
"san capilla" (Martius). 

Morenia linearis (R. & P.) Burret, Notizbl. Bot. Gart. Berlin 11: 
316. 1932. Chamaeodorea linearis (R. & P.) Mart. Hist. Nat. Palm. 2: 
5. 1823; 3: 307. 1849. Martinezia linearis R. & P. Syst. Veg. 297. 
1798. 

Leaves abruptly pinnate, the linear leaflets very acute; spadices 
compound, branches recurved and incurved, dioecious. Apparently 
not re-collected unless as M. macrocarpa Burret. F.M. Neg. 29564. 

Huanuco: Pozuzo, (Ruiz & Pavdn, type). 

Morenia macrocarpa Burret, Notizbl. Bot. Gart. Berlin 13: 
333. 1936. 

Dioecious, 6 meters high, the rachis of the rather robust leaves 
glabrous, these with linear direct and gradually acuminate segments 
paler beneath, many nerves little more slender than the midnerve; 
fruiting spadix 4.5 dm. long, ascending at base, finally recurved; 
peduncle 2.5 dm. long, spathe-scars 5; rachis 12 cm. long, the 20 
branches cernuous; fruits spiralled, orange, 20-22 mm. long, 13 mm. 
in diameter; calyx scutellate, subtrilobed; seed obovate-oblong, 14 
mm. long, 1 cm. in diameter; raphe branches 2, the lower curved, 
the embryo a little above the middle; male spadix 2.5 dm. long, 
spathes 5, the upper moderately inflated; peduncle 13 cm. long, rachis 
1 dm. long, branches 20-25, spiralled, slender, to 7 cm. long; flowers 
rarely solitary, usually 2 or many together; calyx rounded-triangular; 
petals subovate, 2.5 mm. long, equaled by the 6 stamens, these with 
oblong sagittate apically incised anthers. 

Huanuco: Between Monzon and the Huallaga, (Weberbauer 3600, 
type; also 8597, male). 

Morenia Poeppigiana Mart. Hist. Nat. Palm. 3: 161. 1837; 309, 
pis. ll>0-m. 1849. 

Seems to have no character distinct from M. fragrans as remarked 
by Hooker (Bot. Mag. pi. 5492. 1865), except that the male flowers 
are described as inodorous; leaf-pinnae about 3-4 cm. wide, lanceo- 
late-acuminate; lower spathe 2.5 cm. long, upper 5 cm. long. Plate 
140 is a landscape of the type locality with several small palms in 
the right foreground; plate 141, however, is a drawing of spadices in 
flower and fruit, and detailed analysis; Martius' description is com- 



FLORA OF PERU 375 

plete but seems, ex char., to be without contrasting definitions. 
F.M. Negs. 29901; 29902. 

Huanuco: Near Rio Chinchao, (Poeppig, type). 

16. HYOSPATHE Mart. 

Reference: Burret, Notizbl. Bot. Gart. Berlin: 854-859. 1929. 

Small, arundinaceous, the leaves pinnate or pinnatisect, the 
branched spadices below or between them, the small flowers monoe- 
cious, normally male. Spathes 2, long-fusiform, 3 together, not, in 
anthesis, immersed. Sepals of male flowers connate, petals narrow, 
valvate, both imbricate in female flowers, these with 3-celled ovary; 
staminodes present, but sometimes minute. Fruit slightly fleshy; 
embryo basal. 

KEY (after Burret) 

Leaf segments (known) with 1 primary nerve, 2.5 cm. wide. 

H. gracilis. 

Leaves nearly simple or segments, at least mostly, several-nerved. 
Spadices about 1.5-2 dm. long, lowest branches 5-13 cm. long. 

Male flowers subsessile, only 2 mm. long H. Ulei. 

Male flowers pedicellate, 3 mm. long H. Tessmannii. 

Spadices larger, lower branches 2-4 dm. long. 
Lower branches scarcely 2 dm. long; fruit 1.5 cm. long. 

H. elegans. 
Lower branches 2.5-4 dm. long. 

Male flowers sessile, about 3 mm. long H. micropetala. 

Male flowers pedicellate. 

Leaves basally long-cuneate; calyces and pedicels subequal. 

H. brevipedunculata. 

Leaves not long-cuneate at base; pedicels longer than calyx. 

H. Weberbaueri. 

Hyospathe brevipedunculata Damm. Verh. Bot. Ver. Bran- 
denb. 48: 126. 1907. Geonoma Wittiana Damm. I.e. 124 as to leaves 
fide Burret, Notizbl. Bot. Gart. Berlin 11: 235. 1931. 

Leaves, in outline, obovate, the lower of the two pairs of leaflets 
sigmoid, lanceolate, 4 dm. long, 5.5 cm. wide, long-acuminate, 6- 
nerved, the upper, placed 3.5-4 cm. distant, similar (unless as to 
apex, unknown), nearly 4 dm. long, 9 cm. wide, 9-11-nerved; spadix 



376 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

rachis about 4 dm. long, peduncle 9 cm. long, branches many, 1-1.5 
cm. pedicellate, 3-3.5 dm. long, flower glomerules about 5 mm. 
distant; male calyx cupulate, 3-denticulate, corolla lobes lanceolate, 
3 mm. long, a third as wide; filaments unequal, the 3 longer 2 mm. 
long, the shorter 1 mm. long as the oval anthers; female calyx min- 
utely denticulate, 1 mm. long, the broadly lanceolate corolla lobes 
twice as long; staminodes minute, linear. Petioles 2.5 cm. long, 
leaf-nerves 11-12 (the synonym from nearby Seringal Bele'm, not 
Para!). Type, Ule 5881, from Rio Jurua Mirim, in adjacent Brazil; 
also collected at Seringal Bele'm on the same river. 
Loreto (no doubt). Amazonian Brazil. 

Hyospathe elegans Mart. Hist. Nat. Palm. 2: 1, pi. 2. 1823; 
856; 521 (Drude). 

Trunk about 2 meters tall, to 3 cm. thick; leaves 5 or more, lax 
or arcuate, at least a meter long, sheath 2-3 dm. long, truncate, 
petioles 1.5-2 dm. long, rachis 6-9 dm. long; segments many, various 
and mostly remote, nearly 5 dm. long, 1-nerved, linear-acuminate, 
1.5-nearly 2 cm. long, the 6-8-nerved intermixed, lanceolate, 6-8 cm. 
wide, the secondary nerves unequally interposed; spathe before an- 
thesis subequaling spadix; peduncle and rachis of spadices subequal, 
the 12-24 divaricate branches mostly 2 (rarely 3.5) dm. long, near 
base 2 mm. thick, much longer than rachis, rigid, attenuate; male 
flowers at least 3.4 mm. long, the cupulate calyx segments much 
shorter than the broadly lanceolate petals, these exceeded by the 
stamens; female calyx 1.5 mm. long, corolla about twice longer; fruit 
about 1.5 cm. long, 8 mm. in diameter. Original material in part 
belongs to two or more species of Geonoma (Burret). Known from 
adjacent Brazil. Illustrated, Mart. I.e. pi. Z8, figs. 5, 6. F.M. Negs. 
18527; 18528; 18528a (in part, H. filiformis Wendl.). 

Loreto (probably). Amazonian Brazil. 

Hyospathe gracilis [Poeppig] Wendl. ex Drude in Mart. Fl. 
Bras. 3, pt. 2: 523. 1883 (err. sub nom. filiformis). 

Slender, a meter or two high, the nearly equally pinnatisect leaves 
with linear-lanceolate falcate-acuminate more or less strongly veined 
and nerved segments, the solitary uppermost 3.5 dm. long, 2.5-3 cm. 
wide, the upper pair about 2.5 dm. long, 8 cm. wide, broadly connate; 
peduncles 2-3 cm. long, spadices slender, 2 dm. long at base, rachis 
incrassate, branches about 30, crowded, basal to 1.8 dm. long, apical 
to 1 dm. long, strict; glomerules 3-flowered, approximate, male early 



FLORA OF PERU 377 

3-4 mm. long, at anthesis broadly stellate, sepals all 1.5 mm. long, 
petals basally connate, finally linear-lanceolate, long-acuminate, 3 
mm. long, stamens coherent toward base; female flowers ovoid, 1.5 
mm. long, sepals scarcely imbricate, the ovate acute petals twice 
longer. F.M. Neg. 29888. 

San Martin: Rocky island, Rio Tocache, (Poeppig 2057, type). 

Hyospathe micropetala Burret, Notizbl. Bot. Gart. Berlin 10: 
857. 1929. 

Type little more than 3 meters tall, the upper stem only 6-8 mm. 
in diameter, internodes 2.3-3 cm. long; sheath 12.5 cm. long, truncate 
at apex, longitudinally nerved, nearly glabrous, sparsely scurfy; peti- 
oles 17 cm. long, rounded except lightly canaliculate above; leaves 
thin, subconcolored both sides, elliptic or obovate-elliptic, nearly 
entire, apically bilobed or more or less unevenly 2-3 segmented 
each side and with 20 primary nerves, the rachis 24-29 cm. long, 
the segments falcate, acuminate, the medial on upper margin about 
26 cm. long; spadix 33.5 cm. long, glabrous, smooth; branches about 
16, slender, 22-25.5 cm. long, very shortly bracteate at base; male 
flowers sessile, before anthesis 3 mm. long, calyx 1 mm. long, petals 
oblong, 2.5 mm. long; anthers oval, scarcely 0.5 mm. long. Flowers 
orange, stamens bright brown, pleasantly scented (Tessmann). 

Loreto: Inundated woods, San Antonio, mouth of Rio Pastaza, 
(Tessmann 4935, type). 

Hyospathe Tessmannii Burret, Notizbl. Bot. Gart. Berlin 10: 
856. 1929. 

Stem 2.5 meters high, 8 mm. in diameter at the spadix, the 
internodes 2.5-3 cm. long; sheath closely costate, sparsely fuscous 
scurfy as the petioles, these 1.5 dm. long, canaliculate above, rounded- 
triangular beneath; leaves subelliptic or possibly obovate-elliptic, 
about 5.5 dm. long, rather thin, drying dark, nearly concolor, primary 
nerves 19; rachis 3.5 dm. long; segments 3 pairs, falcate, narrowly 
acuminate, subequal or the basal slightly narrower, the medial on 
upper margin 3 dm., the apical about 25 cm. long, 5-6.5 cm. wide; 
primary nerves at most 11 mm. distant, the intermediary well- 
defined on both sides, numerous, the oblique transverse veins more 
obvious above; spadix in flower 15-19 cm. long, branches 7-9, spathes 
17 cm. long, about 1.7 cm. across, glabrous and nearly smooth, ob- 
scurely costate; peduncle 3 cm. long; lower branches 9.5-13 cm. 
long, little flexuose to apex, basal bracts acute as the floral, these 



378 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

7 mm. long; pedicels rather robust, with calyx 3 mm. long, teeth 
rather subfiliform, acute; petals 4.5 mm. long, linear, at most 1 mm. 
wide, densely striate; filaments filiform, 3 mm. long, anthers linear, 
at least 1 mm. long. Petals scarlet, anthers white, leaves used for 
roofs (collector). 

Loreto: Non-inundated woods, mouth of Rio Santiago, (Tess- 
mann 3980, type). "Palmiche." 

Hyospathe Ulei Damm. Verh. Bot. Ver. Brandenb. 48: 127. 1907. 

A low (1-3 meter) slender palm with irregularly pinnatifid leaves 
segments now 1-nerved, again many-nerved and small (1 dm. 
long) spadices; petioles about 1.5 dm. long, minutely brown-scaly 
as the 3-angled rachis; leaves ovate in outline with 4-5 uneven 
segments on each side, the lowest about 22, the uppermost 16 cm. 
long, the width 1.5-7 cm.; lateral nerves 21 in all, each segment more 
or less sigmoid, long-acuminate; secondary nerves 2 between the 
primaries, 3-4 tertiaries between the secondaries; peduncle 2.5 cm. 
long, 6 mm. thick at base, rachis 1 dm. long, type with 10 lateral 
branches 4-7 cm. long, 3-7 mm. distant; lower flowers ternate, the 
2 male 4.5 mm. long; calyx 2 mm. long, with 1 mm. long tip; stamens 
unequal, anthers linear, 1.5 mm. long; female calyx 1 mm. long, 
corolla twice as long; staminodes filiform. Unique in unevenly pin- 
nate leaves and small spadix (author) . 

Loreto: Cerro de Escalero, (Ule 4?P). 

Hyospathe Weberbaueri Damm. ex Burret, Notizbl. Bot. Gart. 
Berlin 10: 858. 1929. 

Three to 5 meters tall, the trunk about 17 mm. in diameter 
beneath the spadix; petioles (known) 26.5 cm. long, minutely and 
sparsely furfuraceous, rounded-triangular, the margins subacute; 
rachis 6.6 dm. long, extended into a filum above the fork of the rigid 
leaf-blade, this paler beneath and with 26 primary nerves to 1.5 cm. 
distant; segments 3 on each side, the apical pair broader, all falcate, 
rather abruptly acuminate; secondary nerves conspicuous both sides, 
the obliquely transverse veins approximate; spadix of male flowers 
before an thesis 38.5 cm. long, peduncle about 5 cm. long; branches 
about 15, the lower 3 dm. long, the upper 2.5 dm. long, slender, 
flexuose; floral bracts shortly triangular; pedicels slender, with calyx 
2 mm. long, the former longer; petals linear, 4 mm. long; filaments 
2-5 mm. long, anthers linear; spadix with fruit (Weberbauer 3657} 
48 cm. long, peduncle 9.5 cm. long, rachis blood-red, the 12 stout 



FLORA OF PERU 379 

branches to 3.2 dm. long, toward apices strongly flexuous; mature 
fruit green, with perianth 12 mm. long, without, 10 mm. long, 
apically rounded, apiculate; perianth 3 mm. high, calyx 2 mm. long, 
teeth short, broad, petals 3-angled. 

Huanuco: Near Rio Monzon toward Rio Huallaga, 600-700 me- 
ters, (Weberbauer 8650, type; also 3657}. 

17. JESSENIA Karst. 

Reference: Burret, Notizbl. Bot. Gart. Berlin 10: 301-312. 1928; 
839-840. 1929. 

In general character similar to Oenocarpus, at least in Peru, but 
minute calyx segments (all) imbricate, broadly cordate. Stamens 
12-20, anthers versatile on apically straight (type) or inflexed fila- 
ments, the anther connective produced. Male petals valvate, female 
convolute-imbricate. Fruit fibrous or also slightly succulent, the 
single seed with ruminate albumen, grooved within or irregular. 
Burret separates this from Oenocarpus chiefly on the basis of many 
(instead of 6) stamens and the ruminate albumen; both characters 
are sometimes variable in other groups (as Bactris, Euterpe) probably 
also the anther connective. When honored by Karsten, Carl Jessen 
was professor of botany at Eldena, Prussia. 

Fruit 3-3.5 cm. long, entire spadix 1-2 meters long J. bataua. 

Fruit 4 cm. long, spadix branches 1 meter long or longer. 

J. Weberbaueri. 

Jessenia bataua (Mart.) Burret, Notizbl. Bot. Gart. Berlin 10: 
300, 302. 1928; 839. Oenocarpus bataua Mart. Hist. Nat. Palm. 2: 
23, pis. 24, 25. 1823. 

Only in age the columnar stems smooth when 20 meters high or 
higher, 2 dm. in diameter, in youth more or less marked by or 
enclosed in the spiniform remnants of the leaf-sheaths; leaves 8-10, 
crowded, erect-spreading, 10 meters long or longer, the equally dis- 
tant segments linear-lanceolate, mostly about 2 meters long, 1 dm. 
wide; spadices few, 1-2 meters long, the many branches fastigiate, 
strict, incrassate above; lower spathe half as long as upper, extended 
into a fuscous tomentose mucro; male petals ovate-oblong, subacute; 
fruit violet-purplish, cylindric-ellipsoid (stigmas nearly on the rounded 
apex), 3-3.5 cm. long, 2-2.25 cm. in diameter, the oblong seed 
acute at both ends. Known from Rios Japura and lea, adjacent 
Brazil; no doubt occurs in Peru. J. polycarpa Karst. (111., Fl. 



380 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Colomb. 1. pi. 98), listed by Dahlgren as in Peru, and certainly 
similar, is apparently known only from Venezuela and Colombia; 
the "Maranon" mentioned by Karsten cannot be Rio Maranon of 
Peru. Burret (Notizbl. Bot. Gart. Berlin 10: 839. 1929) observed 
that this species actually has rings of thin white fibers on the upper 
part of the stem and is distinct from J. bataua. Illustrated, Wallace, 
Palm Trees Amazon, pis. 10, 11. 

The thin mesocarp, triturated with water, forms a creamy sweet- 
ish beverage similar to that from fruits of Euterpe (Spruce) . 

Peru (probably). Brazil to Guiana. "Ungurauy," "sacumana." 

Jessenia Weberbaueri Burret, Notizbl. Bot. Gart. Berlin 10: 
840. 1929. 

Caudex 12 meters tall, nearly 4 dm. in diameter, leaves 10 meters 
long, rachis 1.5 dm. thick (collector), upper fragment seen 3.5 cm. 
high, 2.5 cm. broad, ashy furfuraceous, the regularly disposed seg- 
ments 4-5 cm. distant, linear, 5-6 cm. wide, apex abruptly con- 
tracted, bifid, paler beneath, sigmoid or falcate, perhaps plicate 
above along the longer nerves; fruiting spadix lustrous; peduncle 
very stout, below the lower spathe 12.5 cm. thick, below the upper 
11.5 cm., above that and at the lower branches 8.5 cm. in diameter, 
the space between the spathe scars 9 cm., between the upper of 
these and the lower branches 6.2 cm.; branches stout, minutely 
bracted at base, about 1 meter long or longer, the peduncles 1-2 
meters longer, these apically 7 mm. thick, medially 8 mm. thick; 
vestigial flowers mostly ternate, the 2 lateral male; fruit oblong, 
4 cm. long (3.8 without perianth), 2.3 cm. broad, beak short, slightly 
excentric; perianth to 1.5 cm. high, segments rhombic; seed 2.5 cm. 
long, 1.5 cm. in diameter. Unique in large fruit and long, very 
stout branches; fruit of J. poly car pa Karst. without perianth is 
3.3 cm. long (Burret). 

San Martin: Moyobamba, (Weberbauer 4561, type). "Sinami." 

18. OENOCARPUS Mart. 

Reference: Burret, Notizbl. Bot. Gart. Berlin 10: 291-301. 1928. 

Slender, tall, the unarmed trunks often in clusters, the leaves 
pinnatisect, the spadices infrafoliar in base of dense coma, simple or 
divided. Upper spathe exceeding lower, often acuminate. Flowers 
monoecious, not immersed, the lower or all 3-glomerate, the inter- 
mediate usually female. Stamens 6, anthers versatile; sepals as 



FLORA OF PERU 381 

petals valvate, unless at base. Female flowers much smaller, seg- 
ments imbricate, staminodes commonly obsolete. Fruit 1-seeded, 
drupiform, more or less fibrous or succulent, stigma terminal or sub- 
terminal (type). Seed with plane albumen, embryo basal. The 
name, "wine fruit," applies best perhaps to the species "Pataua" or 
"Bataua," now, probably not necessarily, separated as part of a 
distinct group, Jessenia. For convenience the two species of the 
latter are included in this key. 

Leaf-segments long-acuminate, mostly aggregate. 

Trunks solitary 0. bacaba. 

Trunks clustered 0. multicaulis. 

Leaf -segments abruptly acuminate, laxly distant; trunks solitary. 

Segments 4-5 dm. long 0. minor. 

Segments 1-2 meters long (see Jessenia for descriptions). 

Fruit 3-3.5 cm. long J. bataua. 

Fruit 4 cm. long J. Weberbaueri. 

Oenocarpus bacaba Mart. Hist. Nat. Palm. 2: 24, pi. 26, 
figs. 1-2. 1823; 469 (Drude). 

Stem solitary, to 20 meters tall, 1.5-2 dm. in diameter, the 
marcescent sheaths persisting below the dense spreading coma of 8 
or more leaves, these about 5 meters long with many crowded (in 
clusters of 2-6) linear-lanceolate acuminate segments a meter or 
more long, 4-5 cm. wide; spathes lignescent, caducous before anthesis; 
spadices usually many, flowering and fruiting dependent beneath the 
coma; upper spathe twice longer than lower; spadix branches 5 dm. 
long or longer, strict, basally pedicelliform; male flowers 7 mm. long, 
petals ovate-oblong, acute, short calyx cupulate, stamens early con- 
nivent, anthers bifid; female flowers depressed globose, 5 mm. high, 
the large sepals (enclosing petals) con volute-imbricate; fruit pur- 
plish, pruinose, globose-ellipsoid, plum-like, acute, 2.5 cm. long, 2 cm. 
thick, the stigmas apical; seed 16 X 12 mm. Type from Rio Negro; 
collected also on Rio Solimoes. Illustrated, Wallace, Palm Trees 
Amazon, pi. 9; Drude, Mart. Fl. Bras. 3, pt. 2: pi. 108, fig. 3 (fruit). 

San Martin: Tocache, (Poeppig, fide Drude). 

Oenocarpus minor Mart. Hist. Nat. Palm. 2: 25, pi. 27 (part). 
1823; 471 (Drude). 

Stem solitary, to 10 meters high with a diameter of 7.5 cm. or less, 
annulate; leaves many in a dense coma, often 2-3 meters long, equally 



382 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

pinnate, the 50-60 segments 4-5 dm. long, 4-5 cm. wide, lanceolate, 
abruptly attenuate-acuminate apex, strongly 5-plicate, glaucous or 
puberulent beneath; spadices below leaves, about 3 dm. long, the 
25-35 simple branches 2.5-3 dm. long, early flexuose toward base; 
spathes fuscous tomentose, deciduous after anthesis, the upper ros- 
trate; male petals acute, 3 mm. long; segments of female flowers 
all reniform-orbicular; fruit lustrous, black, ellipsoid, cusped apically, 
1.5 cm. long, 1 cm. in diameter. The fusiform sheaths are more than 
double the diameter of the stem, open with a single slit, split up into 
soft fibers, soon fall away (Spruce). Said to occur in all the wooded 
valleys of the Rio Amazon and listed by Dahlgren (222) as in Peru. 
Illustrated, F.M. Negs. 18853; 18853a; Drude, I.e. pi 108 (fruit). 

Peru (probably). Brazil; Venezuela. "Ciamba," "manaqui." 

Oenocarpus multicaulis Spruce, Journ. Linn. Soc. 11: 142. 
1871; 299 (Burret); 470 (Drude). 

Stems clustered (6-10), 3-10 meters high, to 1.5 dm. thick, re- 
motely annulate; leaves broadly lanceolate, 3-4 meters long; sheath 
9.5 dm. long, ventricose, splitting down the middle into fibers and 
finally deciduous with the leaf; leaf-segments about 60 each side, 
linear-lanceolate, acute, about 1 meter long, 7-8 cm. wide, the lower 
and apical equidistant, the rest 2-3-aggregate, all glaucous or whitish- 
pulverulent beneath; spadix shortly (7 cm.) pedunculate, scapiform, 
deciduous spathe tomentose, the lower 2.5 dm. long, the upper 6 dm. 
long, elongate-fusiform, obtusely rostrate; rachis 1 meter long, 
branches many, nearly 5 dm. long; flowers shortly bracteolate at pit, 
male flowers yellowish, petals ovate-lanceolate, little exceeding sta- 
mens, female flowers whitish, sepals and petals finally orbicular; 
fruit ellipsoid, cusped, stigma apical, 2.5 cm. long, 2 cm. in diameter, 
black-purple, mesocarp thin, sweet, the seed oblong. Fruit edible 
(Tessmann). 

San Martin: Tarapoto, (Spruce 63, type). Loreto: Mouth of 
Rio San Isidro, (Tessmann 4995). "Ciamba," "ciama," "sinami," 
"sinamillo." 

19. EUTERPE Gaertner 

Reference: Burret, Bot. Jahrb. 63: 49-76. 1929. 

Slender, often tall, monoecious, the trunk smooth but annulate, 
solitary (Peru), the petiolar sheaths of the pinnate leaves forming a 
conspicuous crown-shaft. Spadices inter- or infrafoliar, the short 



FLORA OF PERU 383 

outer spathe open apically, the inner elongate, often rostrate; 
branches slender, few to many, spreading or pendent. Flowers in 
groups of 3, sometimes partly or deeply immersed; stamens 6, petals 
valvate, sepals imbricate as also petals of female flowers; staminodes 
ordinarily obvious. Fruit small, subglobose, drupe-like, 1-celled, the 
stigmatic scar lateral or subapical, early somewhat succulent, soon 
hard; seed with plane or ruminate albumen, the embryo lateral 
(or essentially apical; Bailey). The contradiction in characters ac- 
cepted within this group points up the tenuous lines of generic demar- 
cation in the family. Bailey (Gentes Herb. 7: 417-419. 1947) dis- 
cussed the validity of the name and suggested that the presence of 
clearly homogeneous and completely ruminate albumen and super- 
ficial or pitted flowers may indicate two genera. 

Early leaves simple, bilobed; filaments apically geniculate; petals 
obtuse; female flowers superficial; albumen ruminate. 

Peduncle and rachis subequal E. ensiformis. 

Peduncle somewhat shorter than rachis E. megalochlamys. 

Early leaves flabellate-pinnate; filaments straight; petals obtuse; 
female flowers somewhat immersed E. precatoria. 

Euterpe ensiformis (R. & P.) Mart. Hist. Nat. Palm. 2: 32, 
pi. 31. 1824; 58. Martinezia ensiformis R. & P. Syst. Veg. 1: 297. 
1798. 

Caudex erect, cylindric, 12 meters tall; leaf pinnae subensiform, 
very long-acuminate; rachis laxly flowered; sepals of male flowers 
broadly triangular, 4 times shorter than ovate petals, roseate within, 
female whitish; fruit globose, about 6 or 7 mm. in diameter, dull 
blue-black, the stigma scar excentric, scarcely succulent (Kunth). 
Trunk slender, curved, leaves about 8, pinnae horizontal, spadix 
rachis and long peduncle subequal (Tessmann); 5-12 meters high, 
leaf (with petiole, this 6-10 dm. long) 3-4 meters long, spadix rachis 
rose-red, fruit olive-green (Weberbauer). F.M. Neg. 18555. 

Huanuco: Pozuzo, (Ruiz & Pavdn, type; Tessmann, fide Burret). 
Near Rio Monzon, 600 meters, (Weberbauer 3585}. Monzon, 900 
meters, (Weberbauer 3438}. "Cuyol," "coyol." 

Euterpe megalochlamys Burret, Bot. Jahrb. 63: 59. 1929. 

Type with caudex 7 meters tall, leaves 12 dm. long the rachis 
early fuscous tomentose the nutant fruiting spadices glabrous, 
about 7.5 dm. long, the peduncle somewhat shorter; leaf-segments 



384 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

subregular, about 40 on each side, the lower 27 cm. long, to 1 cm. 
wide, the medial to 32 cm. long, 2 cm. wide, the upper 11 cm. long, 
1 cm. wide, all flexible, paler beneath, mid-nerve prominent above 
as the 3 secondary nerves (besides marginal) both sides, abruptly 
acute, often subcucullate or bifid; bracts at base of the many rachis 
branches (these 2-3.5 dm. long), oblong, acute, 2.5 cm. long; fruits 
densely spiralled, 10 or 11 mm. in diameter, light brown, smooth, 
the stigma residuum above the middle; perianth (in fruit) 8 mm. 
across, calyx a third shorter than corolla, both smooth; seed 7 mm. 
long, 8 mm. broad, albumen deeply ruminate. 

La Libertad: Valle Rio Mixiollo, 1,900 meters, (Weberbauer 7070, 
type). 

Euterpe precatoria Mart. Palm. Orbign. 10. pis. 8, 18. 1847; 70. 

Caudex strict, 12-20 meters tall, 1.5 dm. in diameter at base, 
7 cm. at apex; sheaths cylindric, 3 times shorter than the 12-15 
spreading leaves, these to 3 meters long with approximate lax or 
pendent linear long-acuminate pinnae 6 dm. long or longer; spadices 
infrafoliar, the sheaths reddish tomentose, the branches 4.5 dm. long, 
flowers crowded, flavescent; petals of the male broadly lanceolate, 
acute; sepals and petals of the female suborbicular; fruit globose, 
8-10 mm. diameter, residual stigmas excentric; albumen plane. 
Dahlgren, 185, lists locality Maynas (i.e. the area of Yurimaguas) ; 
Huber found it common on the Rio Solimoes and it is also known 
from adjacent Colombia and Bolivia (the type). 

Peru (probably). Colombia to Bolivia. "Yuyu-chonta" (Dahl- 
gren). 

20. ORBIGNYA Mart. 
Reference: Burret, Notizbl. Bot. Gart. Berlin 10: 493-516. 1929. 

Character in general resembles that of Attalea HBK. but the 
free anther cells of the 12-24 stamens are elongate and contorted or 
sinuous, petals ovate or ovate-lanceolate, plane. Fruit 2-6-seeded. 
It is curious that, as independent developments, similar anther 
contortion occurs in some exogens, as Bombacaceae, Cucurbitaceae 
and Columelliaceae. Burret (I.e. 493^497) has presented well the 
relative characters of this and related genera; as he remarks, male 
flowers are necessary to recognize this genus and it may be sug- 
gested that at least in floristic work expediency should take prece- 
dence over emphasis of a single technical characteristic so that 



FLORA OF PERU 385 

determination may be more practical, at least rarely if ever impos- 
sible. Indicative of the closely uniform development of these groups 
may be the occurrence in each of similarly distinctive characters, as 
Burret makes clear. 

The distinguished naturalist, Alcide D'Orbigny, of the Natural 
History Museum of Paris, collected in southern Peru in 1830 and 1834. 

Orbignya polysticha Burret, Notizbl. Bot. Gart. Berlin 11: 324. 
1932. 

Acaulescent (or shortly caulescent) ; leaves 4 meters long, rachis 
(fragment, probably medial) laterally excavated, fuscous furfuraceous 
as segments beneath, especially apically, these (few seen) apparently 
regularly disposed, narrowly reduplicate at base, 8 dm. long, 3.2 cm. 
wide, concolor, larger nerves few, densely and finely nervose-striate, 
the transverse veins prominent both sides, many short, others 6 or 
7 mm. long, narrowly produced; male spadix (part) 3 dm. long, gla- 
brous rachis 22 cm. long, branches simple, upper flowers strongly 
congested; lower branches 7-9 cm. long, sterile for 2-4 cm., sparsely 
ovate-acuminate bracteate; spikes 3-4 mm. thick or with flowers 
nearly 2 cm. in diameter, these spirally in 8 or 9 series, the impressed 
binate flowers about 7 mm. long, conspicuously bracteate, bracteoles 
minute, petals strongly involute apically; calyx 1 mm. high; petals 
spathulate, linear at base, suborbicular at tip; stamens 15, filaments 
filiform, subglobose anthers more or less contorted. Only 0. Dam- 
meriana Barb. Rodr., origin unknown, has spiralled flowers and it is 
not closely related, according to Burret, who referred here with little 
doubt the second collection, this however with stem 4 dm. high, leaf 
segments 6 dm. long, 1.8 cm. wide, edible fruit dark orange. 

Loreto: Mishuyacu near Iquitos, (Klug 205, type). Santa Rosa, 
below Yurimaguas, (Killip & Smith 28814). "Catirina." 

21. ATTALEA HBK. 

Reference: Burret, Notizbl. Bot. Gart. Berlin 10: 517-543. 1929. 

Tall or low (rarely stemless), smooth, the leaves pinnate, the 
flowers monoecious in each spadix (rarely in each tree), this simple 
or simply branched. Stamens 6-many, filaments free, anther par- 
allel, often exserted, the calyx segments small, free or connate at 
base, the petals valvate. Female flowers much the larger, sepals as 
petals imbricate, coriaceous. Fruit ovoid or oblong-ellipsoid, cusped 
or pointed, 1-3 (-5) -seeded, fibrous, 3 (-5) -pored at base. 



386 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Cocos (C. nucifera L., coconut or cocotero), cultivated for the well- 
known subtrigonal-ovoid fruit, also 3-pored at the base, is now re- 
garded as a monotype genus; other palms besides Attalea HBK. that 
are native to Peru and in general similar include particularly Syagrus 
Mart., Scheelea Karsten, Orbignya Mart, and Maximilliana Mart. 
See Burret' s thoughtful discussion (I.e. 493-497). 

Attalea Tessmannii Burret, Notizbl. Bot. Gart. Berlin 10: 538. 
1929. 

Vegetatively incompletely known but ex drawing by collector 
petioles laterally fibrous, leaf-segments at least the lower irregular, 
perhaps disposed in groups; spathe about 24 dm. long, relatively thin, 
extended rather suddenly into a long small beak; female spadix 
branches many, 2 (seen) 5 dm. long, virgate, 5 smaller 13-18 cm., 
all certainly from same plant, basal bracts linear, acute, rigid, 2.2- 
3.5 cm. long, glabrous, one flower in shallow pit 5-7 cm. above the 
base; basal bract about 2.5 cm. long, floral bracts broadly ovate; 
sepals oblong, longer than petals; stigmas 3; lower sterile male flowers 
above female remotely, then rather densely spiralled, the bracts 
mostly 3 (lower 5) mm. long; sepals and petals linear or sublanceo- 
late, to 1.5 cm. long, stamens 12, a third as long (anthers sterile); 
fruits (with perianth and beak) 12-12.5 cm. long, 5 cm. across or 
broader, fuscous furfuraceous, more or less abruptly apiculate with 
beak 2-2.5 cm. long, style with stigmas conically protracted; perianth 
in fruit at least one-third as long as the latter, segments ovate to 
broadly ovate, broadly apiculate, petals somewhat longer; mesocarp 
3 mm. thick, mostly fibrous, the multifibrose endocarp dark-colored; 
male spadix branches 25-27 cm. long, rigidly bracted, the flowers 
more or less spiralled in 5 little inclined, 3 strongly inclined series, 

2 in juxtaposition in axils of very rigid bracts, the smaller bractlets 
ovate-oblong; flowers 1.5 cm. long, sepals oblong, 1.33 mm. long, 
petals acute, costate-nerved, denticulate, minutely scabrous; stamens 
12, anthers sagittate, linear, about 5 mm. long; pistil reduced, the 

3 subulate styles 1 mm. long. No other unquestionably dioecious 
species is known to me from the four related genera; the species is 
further especially noteworthy by the conspicuously bracted relatively 
thick spikes of spirally disposed male flowers, and by the abundant 
fibrous tissue of the dark-colored endocarp (author). 

Cultivated at Pard and collected by Huber, probably on the Rio 
Ucayali (Burret, I.e. 12: 155. 1934). 



FLORA OF PERU 387 

Loreto: Flood-free woods, Soledad, lower Rio Itaya, (Tessmann 
5167, type; 5167a, male). Yarina-cocha, middle Ucayali, (Tess- 
mann 5395, male). "Conta." 

22. SYAGRUS Mart. 

Resembles Scheelea but the putamen of the acute 1-seeded fruit 
rugose within except for 3 smooth bands. Female flowers smaller 
than male, disk prominent. Stamens 6, exceeded by the lanceolate 
petals. Bentham and Hooker followed Drude in regarding this as 
a subgenus of Cocos L.; however, compare Scheelea Karst. and Atta- 
ka HBK. 

Syagrus Tessmannii Burret, Repert. Sp. Nov. 32: 106. 1933. 

Caudex to 20 meters tall, in type 3.5 dm. in diameter at base, 
1.5 dm. at the coma of 17 leaves; petioles 1.5 meters long, dilated 
toward base, marginally tomentose; sheath fleshy coriaceous, semi- 
circular at base dissolving into an intermixed mass of fine and very 
fine fibers; leaf-rachis 3.1 meters long; segments about 160, base to 
apex mostly in distinct groups of 3-4, the basal very narrow and 
shorter, the upper about 2.5 dm. long (rarely 1 meter long, 5 cm. 
wide), all acuminate, glabrous, slender nerves few above, many be- 
neath, obvious, transverse veins obsolete; spadices large, interfoliar; 
upper spathe ventricose, 1 meter long, deeply sulcate, deciduously 
scurfy tomentose; peduncle at apex nearly 3 cm. broad, rudimentary 
spathes or bracts linear or lanceolate, 1-2; branches straight, many, 
rather densely spiralled on the long rachis, to about 5 dm. long, all 
flowers male (no rudimentary female), scrobiculi shallow, 2-flowered 
except toward apex, bracts most minute; flowers ellipsoid-rotund, to 
12 mm. long, calyx 2 mm. long, sepals oblong-lanceolate, carinate, 
petals oblong-subobtuse; filaments 2.5 mm. long, anthers linear, sagit- 
tate, to 7 mm. long; fruit (after Tessmann drawing) yellowish, oval, 
erostrate, with perianth 33 mm. long, 24 mm. in diameter; female 
spadix larger and each flower with 2 sterile male flowers on the larger 
part of the many branches; fruiting bracts minute, broadly rotund; 
flowers (known) before anthesis 8 mm. high, about 6 mm. broad, 
rounded-ovoid; sepals as shorter petals slightly imbricate in bud, not 
seen in anthesis; sterile flowers 6-10 mm. long, calyx 1-2 mm. long, 
petals oblong-elliptic. The collector's painting shows an individual 
tree with male flowers and fruit. From the Brazilian S. inajai 
(Spruce) Becc. and S. ecuadoriensis Becc. this species is distinct by 
the thinner leaves without transverse veins; notable is the cushion 



388 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

of fibers of the young sheaths; it may be illustrated by Huber (Bull. 
Herb. Boiss. se>. 2, 6: pi. 13, as a species of Cocos) under a nomen 
nudum (tall, the spadix much-branched), which, of course, must be- 
come a nomen dilendum as suggested by Burret; the transfer of 
Huber's name without description also gave it no standing. 

Fleshy fruit eaten by cattle; stem, used for construction, harbors 
edible larvae, the wood serves for lances, etc. (collector). 

Loreto: Mouth of Apaga, (Tessmann 4811, type). Middle Ucay- 
ali, Rio Ayuaytia, (Tessmann 3286, female spadix). "Inchaui," 
"intshawui." 



23. SGHEELEA Karst. 
Reference: Burret, Notizbl. Bot. Gart. Berlin 10: 651-689. 1929. 

Resembles Orbignya Mart, and Attalea HBK. but the petals of 
the male flowers are fleshy, elongate-clavate or cylindrical, and the 
stamens are straight as in the latter but only 6. Fruit 1-3-seeded, 
the seeds ellipsoid, albumen plane. Cymba (spathe-valve) heavy, 
ligneous as in Attalea. S. Wallisii (Huber) Burret (I.e. 657) from 
the Rio Ucayali is essentially a nomen nudum. Huber (Bull. Herb. 
Boiss. se>. 2, 6: 267. 1907) described the drupes as ellipsoid-cylindric; 
only original material will enable identification (Burret). Only one 
species in the following key after Burret is acaulescent. 

Stigmas (as cells) mostly more than 3; female flowers in mostly 1-5 

branches; fruiting bracts large; endocarp fibers large (section 

Syncalphocaryum Burret). 

Acaulescent S. Weberbaueri. 

Caulescent. 

Leaf-segments regularly disposed S. Bassleriana. 

Leaf-segments 2-4-aggregate S. cephalotes. 

Stigmas mostly 1 (-3); female flowers lax on 5 or more branches; 

bracts as fibers small (section Dialphocaryum Burret). 
Fruit rounded, the beak slender, the perianth to 2 cm. high. 

Perianth one-third as large as fruit; flowers (male) 1.5 cm. long, 
yellow S. stenorhyncha. 

Perianth one-fourth as large as fruit; flowers (male) 1 cm. long, 

yellowish S. brachyclada. 

Fruit extended into thick beak; flowers yellowish, female 2.5 cm. 
long, male 1.5 cm. long S. Tessmannii. 



FLORA OF PERU 389 

Scheelea Bassleriana Burret, Notizbl. Bot. Gart. Berlin 10: 
655. 1929. 

Type 12 meters tall, fragment (apical) of leaf seen about 1 meter 
long, its narrowly linear rachis fuscous furfuraceous only beneath, 
the acutely angled segments gradually decreasing in size, regularly 
disposed, narrowly reduplicate at base, linear, long-acuminate, shortly 
and unequally bifid, subpruinose beneath, green above, midnerve 
very prominent, secondary nerves obscure, transverse conspicuous, 
obsolete beneath; male spadix (fragment) branches about 2-2.5 dm. 
long, bracts scarcely 2 mm. long, abruptly contracted, the flowers 
densely spiralled; petals 14 mm. long, stamens one-third as long, 
anthers oblong, 3 mm. long; fruit oval, 9.5 cm. (with perianth) long, 
5-5.5 cm. in diameter, slender acumen of rounded apex 1 cm. long; 
perianth cupulate, 2.4 cm. high, sepals ovate, subacuminate, 2.8 mm. 
long, little longer than the broader petals; mesocarp (dried) entirely 
fibrous, 4 mm. thick, endocarp 3-celled, 8 mm. thick, densely and 
rather regularly spotted, the many fibers mostly 2-4-aggregate, small 
fibers nearly none. Trunk to 4 dm. in diameter, flowers yellowish, 
oily flesh of fruit edible (Tessmann) ; name commemorates supporter 
of botanical work. 

Loreto: Yarina-cocha, Rio Ucayali, (Tessmann 5490, type; also 
3266}. "Shevao," "shebon." 

Scheelea brachyclada Burret, Notizbl. Bot. Gart. Berlin 10: 
680. 1929. 

Apical part of leaf seen 9 dm. long, glabrous, the rachis triangu- 
lar, segments reduced upward, nearly regularly disposed, spreading 
in one plane (lower probably aggregate?), narrowly linear, reduplicate 
at base, rounded but bidentate at apex, medial nerve prominent 
above as 3 lateral nerves each side, the numerous transverse ones 
obvious only above; female branchlets many (broad-based bract 
pungently acuminate), 1-1.5 dm. long, upper fourth to half bearing 
sterile male flowers, lower part 4-5 (rarely 3) secund or subdistichous 
female, the cupulae of bracts 1 cm. across, about 2.5 cm. long; sepals 
long- triangular, longer than petals, equaling stigmas; fruits with 
perianth 7 cm. long, 3 cm. in diameter, oblong-cylindric, the sub- 
rotund apex with beak-like style 7 or 8 mm. long, transverse section 
1-seeded (2 cells aborted), mesocarp 2.5 mm. in diameter, entirely 
fibrous, endocarp 8 mm. thick, fibers brown, often few aggregate; 
perianth in lobes 2 cm. high, sepals and petals ovate, the latter very 
broadly, abruptly contracted; male spadix branches small, slender 



390 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

(basal bracts 1.5-2 cm. long), 10-12.5 cm. long, floriferous to 1.5 cm. 
of base, rather crowded flowers finally subsecund; floral bracts minute, 
suborbicular; flowers 8 or 9 mm. long, petals subulate, about twice 
as long as stamens (anthers 2.5 mm. long), sepals oblong, 0.5 mm. 
long. Description of leaves and male spadix from 5493, which col- 
lection was accompanied by careful drawing of fruit so that the 
identity is scarcely to be doubted (Burret) ; fruit, from the aquarelle, 
8.3 cm. long, 3.5 cm. in diameter, 9 mm. rostrate from pulvinate- 
conic style, perianth lobes 2.5 cm. high. Differs as S. Tessmannii 
from the obscure S. kewensis Hook. f. (Bot. Mag. pis. 7552, 7553. 
1897), cultivated, by the not exserted stigmas. 

Loreto: Soledad, Rio Itaya, (Tessmann 5237, type). Yarina- 
cocha, middle Ucayali, (Tessmann 5493}. 

Scheelea cephalotes (Poepp.) Karst. Linnaea 28: 269. 1856; 660. 
Attalea cephalotes Poepp. ex Mart. Palm. Orbign. 10: 119. 1847. 

Type with caudex several meters tall, smooth below, residual peti- 
oles toward apex among the leaves, these to 3 meters long including 
the petioles, fibrillose toward base and to 1 meter long; leaf-rachis 
lepidote-tomentose, pinnae reduplicate (2-4-aggregate), about 1- 
2.5 cm. distant, linear, acuminate, to 1 meter long, 16-18 mm. 
wide, pale green, punctate beneath, the midrib prominent above, 
acute, secondary fewer than 8, branches of male spadices about 
1 dm. long, densely fastigiate, subterete, flexuose below; bracts 
minute, flowers 8-10 mm. long, calyx minute, ovate-suborbicular 
segments apiculate, petals linear, subfleshy, shortly acute, anthers 
subsessile; female spadix bracteate at base, coriaceous bractlets 2, 
flowers unknown in type; fruit nearly 1 dm. long, about 3.5 cm. in 
diameter, fibrose without, fleshy within, the residual stigmas apical; 
seed solitary, linear-oblong, subtrigonous, albumen cartilaginous. 
Weberbauer noted stem 7 meters high, fruiting spadix without pe- 
duncle 6 dm. long, 3.5 dm. across. Illustrated, Mart. Hist. Nat. 
Palm. 3. pi. 169. F.M. Neg. 31307. 

Huanuco: Mouth of Rio Pozuzo, (Weberbauer 6762}. Loreto: 
Yurimaguas area, (Maynas), Poeppig, type. "Shapaja." 

Scheelea stenorhyncha Burret, Notizbl. Bot. Gart. Berlin 10: 
675. 1929. 

Type with stem 4 dm. in diameter, 25 meters high, bearing 15- 
18 leaves; male spadix branches 2.5-3.5 dm. long, bract 7 mm. long, 



FLORA OF PERU 391 

farinose puberulent, flowers spiralled, attenuate to nearly acute apex; 
floral bracts ovate, pungent, minute; flowers 17 mm. long, yellow, 
sepals oblong, petals scarcely broader than 1 mm.; stamens one- 
fourth as long as flowers, anthers nearly 3 mm. long; fruiting branch 
female below, fruits spiralled; bracts forming a cup about 1 cm. 
across; perianth in fruit 1.8-2 cm. high, to 2.5 cm. in diameter; 
sepals, petals subequal, about ovate, abruptly acute; fruit cylindric, 
6 cm. long (with perianth and beak), 2.3-2.5 cm. in diameter, beaks 
5-7 mm. long, transverse section 1-seeded (2 cells obliterated), meso- 
carp thin, entirely fibrous, endocarp cinnamon-colored, 6 mm. thick, 
inner fibers none. Related to S. regia Karst. of Colombia with male 
flowers only 8 mm. long, stamens half as long. 

Loreto: Soledad, Rio Itaya, (Tessmann 5256, type). "Shapaja." 

Scheelea Tessmannii Burret, Notizbl. Bot. Gart. Berlin 10: 
682. 1929. 

Trunk 12 meters tall, 5 dm. in diameter, the crown with about 
15 leaves; spadix about 4.5 dm. long, apically attenuate, the male 
flowers finally in 5 laxly spiralled series, 15-17 mm. long, sepals 
scarcely 1 mm. long, petals cylindric, 1 mm. thick, stamens shorter 
than 5 mm.; lower part of branchlets with 12-17 slightly secund 
female flowers, before anthesis oblong, 2.7 cm. long; sepals oblong- 
acuminate, stigmas included; fruit 8 cm. long, cylindric-terete, 3 cm. 
in diameter, rather gradually extended about two-thirds its length 
into a stout beak; mesocarp 2-4 mm. in diameter, endocarp 6-7 mm. 
thick, minutely fibrous; perianth nearly 2.5 cm. high, sepals and 
petals subequal, the former ovate-oblong, the latter broadly rounded, 
both acute. Fruit recalls that of S. kewensis Hook. f. (cf. S. brachy- 
clada) but more slender and stigmas included; also, male flowers 
spiralled, petals much longer (author). Photograph shows a strong 
but slender trunk, smooth but annulate, leaf-rachis little arched, 
leaf-segments on same plane, apparently regularly disposed, the in- 
terfoliar spadices about 10. 

Loreto: Inundated and non-inundated woods, Iquitos, (Tess- 
mann 5085, type). "Shapuja," "chapaja." 

Scheelea Weberbaueri Burret, Notizbl. Bot. Gart. Berlin 10: 
659. 1929. 

Completely stemless, the leaves 9-10 meters long, the rachis (frag- 
ment, probably medial) 3 cm. broad, acutely carinate above, rounded 
beneath, only dorsally fuscous furfuraceous; segments aggregate, di- 



392 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

rection diverse, 4-5 cm. wide, green above, paler beneath, the mid- 
nerve stout, longitudinal and transverse nerves obscure or faint both 
sides; male spadix (flowers unknown) 6 dm. long, basal part 14 cm. 
long, 2 cm. in diameter at apex, rachis glabrous, much attenuate, 
branchlets slender, very numerous, spiralled, lower flowers in 5-6 
series on upper third of rachis approximate, floral bracts minute; 
fruiting spadix 8 dm. long (base 2.5 dm. long), upper bracts large; 
apices of branchlets with sterile male flowers; fruits mostly abnormal, 
more or less compressed obovoid, 7-9 cm. long (with perianth and 
beak this 7 mm. long), about 4-4.5 cm. in diameter, rounded api- 
cally; perianth mostly lacerate, 2.5 cm. high, segments rather ovate; 
mesocarp 2-2.5 mm. thick, outer part finely fibrous, endocarp densely 
sulcate, 1-2-celled (transversely). 

Junin: La Merced, (Weberbauer 1848, type). "Shapaja." 

24. MAXIMILLIANA Mart. 
Reference: Burret, Notizbl. Bot. Gart. Berlin 10: 689-701. 1929. 

Resembles Scheelea in appearance vegetatively but the approx- 
imate (in circles) male flowers with minute petals equaled or far 
exceeded by the 6 filaments, the linear anthers affixed at bifid base, 
pendulous. Female flowers few, much larger, sepals coriaceous, 
broadly imbricate, petals little longer. Fruit 1-seeded, pointed or 
one of the rounded lower halves free from fibers (Drude). 

Sepals of male flowers about a sixth as long as petals . . M. venatorum. 
Sepals of male flowers a third as long as petals M. stenocarpa. 

Maximilliana stenocarpa Burret, Notizbl. Bot. Gart. Berlin 
10: 696. 1929. 

In general similar to M. venatorum but leaf-segments probably 
16 dm. long, 5 cm. wide, little narrowed below, narrowly bidentate 
apically, midnerve stout above, secondary nerves 6; apex of spathe 
seen strongly compressed, margins rounded, minutely and decidu- 
ously furfuraceous; male spadix glabrous (only fragment seen), prom- 
inently carinate, bracts 5 mm. long, often dentate at base, acute, 
perianth 2 mm. long, sepals 0.75 mm. high, ovate, filaments longer 
than petals, anthers 8 mm. long; female spadix branches about 2 dm. 
long, bracts 1 cm. long, flowers 7-9 inserted on lower two-thirds, 
elliptic, 1.6 cm. long, sepals ovate, the little longer petals acute, den- 
tate; upper third of branches sterile male flowers, these with sub- 
oblong petals, sepals 1 mm. long, anthers 5 mm. long; fruiting 



FLORA OF PERU 393 

branches 2; cup of perianth bracts 7 mm. across; fruit 4.5 cm. long, 
18 mm. in diameter, deciduously tomentose, sepals ovate, acute, 
petals rotund, apiculate; mesocarp fleshy, finely fibrous, endocarp 
smooth, beaked, 4 X 1.6 cm., the pores about 5 mm. above the base. 
Contrasts with M. venatorum, as the calyx of male flowers in com- 
parison with petals is several times longer, and the heads of the 
branches as well as the male flowers are notably longer (author). 

Loreto: Iquitos, (Tessmann 5081, type; also 5078, male) . "Ynay- 
uca," "inayuca." 

Maximilliana venatorum (Poepp.) Wendl. Kerch. Palm. 251. 
1878; 698. Cocos venatorum Poepp. ex Mart. Hist. Nat. Palm. 3: 
325. 1850. Attalea venatorum (Poepp.) Mart. I.e. 

Caudex less than 8 meters high, leaves with long petioles, the 
4-aggregate pinnae in part erect, all lepidote at base, linear, long- 
acuminate; spathe pubescent, longitudinally sulcate, long-cuspidate; 
spadix about 8 dm. long, branches of female erect, glaucous-pruinose; 
stamens 6, exserted, 4 times longer than the perianth; fruit oval. 
Burret described the Tessmann specimen about as follows: apical 
leaf fragment with triangular rachis minutely fuscous scurfy beneath; 
segments aggregate nearly to apex, the lower variously directed, all 
rigid, concolor, secondary nerves about 3, the ulterior obscure above, 
many very fine beneath, the transverse more prominent above; lower 
segments seen 3-aggregate, 7 dm. long, 2 cm. wide, the upper mostly 
binate, the uppermost very narrow; branches male spadix fragment, 
incrassate at base, 14-16 cm. long, sparsely bracteate to about one- 
third, finally densely flowered, about 2 mm. thick; floral bracts sub- 
rotund, minutely extended, oblong, nearly 7 cm. long; perianth 
2.75 mm. high, sepals scarcely 0.5 mm. long, petals narrowly oblong, 
filaments as long, anthers linear, 7 mm. long. F.M. Neg. 31308. 

Indians use petioles as blow-pipes for the arrow poison Urari 
(Poeppig). 

San Martin: Tocache, (Poeppig). Loreto: Yurimaguas, (Poep- 
pig). Yarina-cocha, (Tessmann 5492). "Ynaynca," "inayuca." 

25. AIPHANES Willd. 

Martinezia R. & P. Prodr. 148, pi. 32. 1793 (per Dombey, litt.); 
1794 (per title) ; Syst. Veg. 295. 1798, a mixture fide Burret, Repert. 
Sp. Nov. 34: 169-170. 1933. 

Reference: Burret, Notizbl. Bot. Gart. Berlin 11: 557-577. 1932. 



394 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Spiniferous nearly all over, and thus resembling many species of 
Bactris but the spadices elongate and sooner or later free from the 
slender or thin, not cymba-like spathe. Pinnae usually cuneate- 
based, sometimes strongly oblique at the truncate or erose apex. 
Flowers variously disposed as to sex but ordinarily 1 female with 
2 male, now and then the spadix mostly female, not infrequently 
entirely below. Stamens 6, linear or suborbicular, the male flowers 
not immersed. Ovary 3-celled. Fruit a small, 1-seeded, firm or 
fleshy depressed-globose drupe, the seed rugose or pitted, albumen 
plane, solid. A. caryotifolia (HBK.) Wendl., ascribed to Peru by 
Martius, is probably an error (cf. Burret, I.e. 560); the species, at 
least as to typical state, is restricted, perhaps, to Colombia or area 
north of Peru. 

Flowers scarcely immersed, male higher than basal, acute; female 
staminodal cup teeth 6, obvious (subgenus Macroanthera Burret). 

Leaf-segments 3-5-aggregate A. Ernesti. 

Leaf-segments distinct A. praemorsa. 

Flowers, especially female, more or less immersed; male often broader 
than high, rounded; staminodal teeth obscure or little obvious 
(subgenus Brachyanthera Burret). 
Caulescent. 
Leaf pairs 5-12, cuneate or deltoid leaflets. 

Leaflets 5 pairs, small A. Ulei. 

Leaflets 12 pairs, ample A. deltoidea. 

Leaf pairs many, narrow. 

Leaflets all subequal A. gracilis. 

Leaflets unequal A. Weberbaueri. 

Acaulescent A. Tessmannii. 

Aiphanes deltoidea Burret, Notizbl. Bot. Gart. Berlin 11: 568. 
1932. 

Cespitose, the larger stems to 2 meters long, regularly spiny as 
the petioles, these 3.5 dm. long, densely fuscous scurfy, not setose, the 
spines 3 cm. long, black; rachis aculeate, setose and pubescent, be- 
neath furfuraceous; leaves 10 or 12, about 14 dm. long, oblong, the 
segments around 12 pairs, very unevenly disposed, 2-3-aggregate, 
widely spaced, the lower above the base mostly 1-aculeate, 4-4.5 cm. 
long, all above smooth and glabrous, subglabrous beneath, broadly 
elongate-deltoid-cuneate, the apical decurrent 2-2.5 cm., inner mar- 



FLORA OF PERU 395 

gin 2.6 dm. long, 8-11 cm. wide, the apex praemorsely obliquely 
lobulate and caudately produced 6 cm., lower segments shorter but 
often relatively wider; spadices interfoliar, about 1 meter long, the 
branched part 47 cm. long; lower spathe linear, about 3 dm. long, 
to 12 mm. wide, furfuraceous, free part 7 dm. long, linear, not at all 
acuminate; peduncle as rachis and many divaricate branches these 
slender, flexuose, the lower 3.5, the upper about 1 dm. long furfura- 
ceous, setose and setulose; glomerules mostly 2 males, upper flowers 
all male, the latter broader than high, sepals oblong, petals broadly 
ovate, costate, anthers oval; female sepals rotund, petals connate, 
lobes triangular, staminodal cup obscurely dentate. Male flowers 
orange, female more greenish, all odorless; fruit eaten, sweet. 

Loreto: Mouth of Rio Santiago, (Tessmann 4709, type) . "Chiqui- 
chiqui," "chica-chica," "shica-shica." 

Aiphanes Ernesti Burret, Notizbl. Bot. Gart. Berlin 11: 560. 
1932. Martinezia Ulei Damm. I.e. 6: 266. 1915, not 1907. 

Petioles, leaf -rachis and spathes whitish-tomentose, brown-squa- 
mate and dark brown aculeate with spines to 3.5 cm. long; leaf- 
segments 3-5-aggregate, elongate-cuneate, to 3.5 dm. long, 1 dm. 
wide, apically erosely crenate-dentate and tricuspidate, glabrous 
above, pubescent beneath, the upper of the sparsely setose margins 
long-produced; outer spathe membranous toward apex, finally only 
fibrous, the inner elongate-lanceolate, subligneous, sparsely short- 
aculeate, to 6 dm. long, 3 cm. wide; spadix long-peduncled, branches 
many to 26 cm. long, 3 mm. thick at base, tortuous, densely flowered; 
flowers solitary toward apex; male calyx subhyaline, lobes long- 
lanceolate, 1.5 mm. long, corolla fleshy, petals ovate, acuminate, 
3 mm. long, 1 mm. wide, those of female cordate, 1.5 mm. long; 
fruit red. Type, Alto Acre (Ule 116B, type). 

Peru (probably). Adjacent Brazil. 

Aiphanes gracilis Burret, Notizbl. Bot. Gart. Berlin 11: 566. 
1932. 

Similar to A. Weberbaueri, but slenderer and smaller; type half 
meter tall; petioles about 3 dm. long, densely black-setose and -acule- 
ate, the larger spines nearly 4.5 cm. long, these as setae but smaller 
on the lower part of the leaf -rachis and the leaf-segments beneath; 
leaf-blades to 9.5 dm. long, the 17-20 pairs of segments plicate in the 
nerves, rigid, the apical 2 subequal or narrower, to about 14 cm. long, 
the longest about medial, 1.5 dm. long on midnerve, upper margin 



396 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

produced 3-5 cm., to 3.5 cm. wide, upper segments decreasing in size; 
spadix slender, about 13 dm. long, sparsely and minutely black- 
setose, the branched part 6 dm. long, the branches 12-15, scattered; 
upper male flowers 2 or solitary. Except that the terminal leaf- 
segments are little if at all wider than the rest, the differences appear 
to be relative developments. 

Amazonas: Rio Mayo north of Moyobamba, (Weberbauer 4665, 
type; 585). 

Aiphanes praemorsa (Poepp.) Burret, Notizbl. Bot. Gart. Ber- 
lin 11 : 575. 1932. Bactris praemorsa Poepp. ex Mart. Palm. Orbign. 
10: 66. 1847. 

Low caudex and basally vaginate petiole aculeate, the smooth 
leaf-rachis lepidote-tomentose, the few deltoid leaflets apically erose- 
denticulate, marginally aculeolate, early lepidote beneath, finally 
green; drupe turbinate-globose. Seems to differ from A. caryotifolia 
(HBK.) Wendl. in that the indument and the leaf-rachis are not 
aculeate, and the segments are larger (Martius) . No fruit at Munich, 
but the leaves most resemble A. caryotifolia; the rachis, however, is 
quite without spines; noteworthy, also, is the white tomentum; in 
A. Ernesti the lower leaf segments are strongly drawn together 
(Burret). 

San Martin: River woods near Tocache, (Poeppig, type). 

Aiphanes Tessmannii Burret, Notizbl. Bot. Gart. Berlin 11: 
564. 1932. 

Acaulescent; petioles 3 dm. long; spines black, few, to 5 cm. long, 
setae many, yellowish-brown, these also on both sides of leaves and 
on spadices; leaves long-elliptic, 12 dm. long, rachis not at all acule- 
ate, extended as a filum; apical lobes 7-nerved, 11-12.5 cm. across, 
the inner margin 2 dm. long, the apical unevenly lobulate-dentate; 
remaining segments 1-nerved, 19-20 on each side, rather regularly 
spaced, mostly the medial larger, these on midrib about 28 cm. long, 
on upper caudate margin 32 cm. long, all sublinear, 2.5-3 cm. wide, 
thin, yellowish-green with long rigid trichomes; branched part of 
spadix 4.5 dm. long; peduncle elongate; lower spathe apparently 
black-setose, the upper very long, thin, narrow, yellowish-setose; 
branches not aculeate, the setae minute; glomerules rarely with 2 
male flowers, the upper with flowers nearly always solitary; petals 
reflexed, ovate-rounded, nervose, calyx not produced basally, sepals 



FLORA OF PERU 397 

suboblong, anthers oval; female flowers with broadly rounded sepals, 
staminodal teeth obscure. An outstanding species (author). 
Loreto: Mouth of Rio Santiago, (Tessmann 4281, type). 

Aiphanes Ulei (Damm.) Burret, Notizbl. Bot. Gart. Berlin 11: 
568. 1932. Martinezia Ulei Damm. Verh. Bot. Ver. Brandenb. 48: 
127. 1907. 

Slender, 1-4 meters tall, the pinnate leaves with 5 pairs of cuneate- 
truncate, setose-ciliate leaflets about 1 cm. distant and a single pair 
6-8 cm. from these, the 3-angled rachis somewhat scaly and strongly 
black-spiny; leaflets (known) to 6 cm. long, 2 mm. wide at tip, the 
marginal setae 1 mm. long, 2-4 mm. distant; spadix 3 dm. long or 
longer; peduncle sparsely aculeate; rachis pilosulous or glabrous, 
elongate branches strict, female flowers on lower third or half, male 
approximate in 2 series above. 

Loreto: Cerro de Escalero, (Ule 6880, type). 

Aiphanes Weberbaueri Burret, Notizbl. Bot. Gart. Berlin 11: 
565. 1932. 

Ascending stems to a meter high, the younger portion long- 
aculeate with dark brown spines, these black on the petioles (frag- 
ment seen 3 dm. long), many, to 6 cm. long; rachis aculeate, yellowish 
setulae toward apex, produced into a filum above the bifurcation, 
longer than 1 meter; apical segments both 5-nerved, the inner margin 
7.5 cm. long, at the apex 3.5-6.5 cm. wide, praemorsely denticulate, 
the remaining 18-20 all 1-nerved, very unevenly disposed in clusters 
at long intervals, those below the apical somewhat narrower, the 
larger medial segments about 21 cm. long on the midnerve, 24 cm. 
on the upper margin, to 4.5 cm. wide above, thin, not at all plicate, 
green both sides, often beneath above base 1-aculeate, both sides 
between and on veins short- and long-brownish-setulose, the upper 
margin (not lower) caudate, the apex unevenly lobulate; spadices 
rather slender, branched part 42 cm. long, branches about 24 (pe- 
duncle sparsely aculeate, setose, 4 dm. long), the lower 12 remote, 
largest seen about 3.5 dm. long, rigid, subflexuose, densely brown 
setose; lower glomerules laxly spiralled, upper and those of upper 
branchlets male, mostly solitary; petals broadly ovate, obtuse, an- 
thers small, suborbicular; fruits oval, setose, with perianth 12 mm. 
long, 1 cm. thick, sepals rotund, petals connate below, cup obscurely 
dentate. 

Huanuco: Rio Pozuzo, (Weberbauer 6775, type). 



398 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

26. ASTROCARYUM G. F. W. Meyer 
Reference: Burret, Repert. Sp. Nov. 35: 114-158. 1934. 
Spinescent and vegetatively similar to Aiphanes rarely acaules- 
cent, often robust, and, especially, the spathe-valves cymba-like, the 
peduncles short, extended as the rachis of the reduced spadix, the male 
immersed in the crowded, usually simple branches. Fruit 1 (-2) 
-seeded, the pericarp slightly fleshy but soon dry, fibrous within, the 
somewhat accrescent perianth persisting, the 3 subapical, often radi- 
ately ornamental pores regularly at the same height (Burret) ; albu- 
men grooved. Entirely mature fruit with perianth is necessary to 
determine the species (Burret). 

Spadix branches with 2-many female flowers laxly disposed above 

base; fruit not long-cuneate or applanate, not aculeate or setose 

(subgenus Pleiogynanthus Burret). 
Fruit early scurfy and setulose; staminodal cup elongate. 

A. chambira. 
Fruit glabrous; staminodal cup short. 

Leaf -segments 3-4-aggregate A. tucuma. 

Leaf-segments equidistant A. jauari. 

Spadix branches with 1 female flower, basal or subbasal; endocarp 

normally cuneate below, often laterally somewhat prismatically 

applanate, mostly aculeate or setose (subgenus Monogynanthus 

Burret). 
Calyx and corolla of female flowers subequal. 

Plants acaulescent A. huicungo. 

Plants caulescent A. murumura. 

Calyx shorter than corolla in female flowers. 

Fruit finely setulose; female calyx setose A. macrocalyx. 

Fruit smooth; female calyx esetulose A. chonta. 

Astrocaryum chambira Burret, Repert. Sp. Nov. 35: 122. 1934. 

Caudex solitary, 3.5 dm. in diameter, annulate, the internodal 
spines to 9 cm. long; leaves 16, about 6 meters long, petioles deeply 
canaliculate, 1 dm. wide, spines to 1 dm. long or longer, rachis acu- 
leate and setose; segments many, more or less approximate and 
unevenly disposed, the linear basal not at all reduplicate, the apical 
rather abruptly contracted, bifid, mostly subobtuse, the remaining 
similar but gradually attenuate, all lustrous above, smooth, margin- 
ally rather coarsely setose (-ciliate), opaque (probably early silvery) 



FLORA OF PERU 399 

beneath, the midrib prominent on both sides, the secondary nerves 
obscure above, more obvious beneath; upper spathe (fragmentary) 
a meter long, furfuraceous, aculei slender, 2-2.5 cm. long, not cov- 
ered by the dense setosity; peduncle 12 cm. long, shortly aculeate, 
early tomentose and sparsely aculeate, the many branches spiralled, 
to 4.5 dm. long, female flowers bright brown, 2-3 on the lower half 
of the branch, the upper part a thick, densely white-villous bracted 
spike of male chocolate-colored blossoms; female with 2 rudimentary 
male, calyx glabrate, cupulate, little or not longer than the similar 
but contracted corolla, this white-furfuraceous and black-setulose, 
nearly equaled by the staminodal cup; male calyx lobes narrowly 
lanceolate, acuminate, the corolla base connately stiped, the oblong 
lobes nearly as long; anthers linear, 3 mm. long; fruit elongate- or 
obovoid-globose, to 7 cm. long including beak (1 cm. long) and cup- 
ulate perianth (13 mm. long), 4 cm. in diameter or more; epicarp 
early scurfy and black-setulose, finally opaque, densely gibbous, 
mesocarp fibrous, endocarp obovoid, notably produced at base, the 
calyx more or less damaged, the corolla nearly crenulate. Distin- 
guished from A. tucuma Mart, by the coniform fruiting perianth 
and the very high staminodal cup of the female flowers; A. macro- 
carpum Huber (Bull. Herb. Boiss. se>. 2, 6: 271, pi. 13. 1906), of 
adjacent Brazil, scarcely described, has about 6 leaves, rather weak 
spines, lower internodes at least twice longer than broad, fruit larger. 

Tessmann noted usual uses for similar palms, including the fiber 
and fruits, and observed this species frequently in areas rarely 
flooded near Iquitos, on the Ucayali, Tamaya, Yarina-cocha, mouth 
of the Santiago, and Rio Chambira. 

This may be the species referred to by Spruce (Journ. Linn. Soc. 
11: 157. 1871) as A. vulgare Mart, under the same native name, a 
species not, apparently, within the range; this native name has 
been listed, as by Dahlgren, also for A. tucuma Mart.; Drude's illus- 
tration of the fruit of the latter could be this species (Burret). 

San Martin: Tarapoto, (Raimondi). Loreto: Flood-free area 
near Iquitos, (Tessmann 5079, type). Brazil. "Chambira." 

Astrocaryum chonta Orbigny ex Mart. Palm. Orbign. 10: 84, 
pi 4, figs. 1, 2; pi. 29C. 1847; 149. 

Caudex at least above long-aculeate, to 10 meters tall; leaves 
9-12, finally divaricate, 6 or 7 meters long, 2-5 meters wide, the 
linear-acuminate subfalcate pinnae 1 meter long or longer, to 7.5 cm. 
wide, regularly subopposite, about 2.5 cm. distant, not at all plicate, 



400 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

pale or silvery-tomentose beneath, the rachis (this tomentose) and 
margins black setulose, the spines about 1-2.5 cm. long; midnerve 
prominent above, the secondary many, very fine, approximate; 
spathe fusiform, 9 dm. long, violet-tinged, closely black aculeate 
especially toward the apex; spadix branches many, 7-10 dm. long; 
female flowers subsessile, at base of each branch, calyx as corolla 
cupulate, the latter 2 times longer, esetulose; male flowers shortly 
pedicellate, calyx segments acute, carinate, the corolla divisions 4 
times higher, oblong-linear, obtuse, the filaments 6, basally cohering; 
fruit yellow, long-obovoid, smooth, 6-7 cm. long, scarcely 2.5 cm. 
thick above, the residual stigmas apical, the pores subapical; black 
seed radiately veined. After Martius (I.e. 85), who referred here a 
fruiting specimen (Herb. Parker Webb) labeled as collected by Pavon, 
not seen by Burret; however, as it is known from the upper Rio Purus 
it well may occur in flooded woods and shores in Madre de Dios. 
Plate 29 (I.e.) is incorrect in irregular pinnae; fruit perianth sche- 
matic (Burret). 

Wood hard but fragile; putamen very sweet (but not eaten) fruit 
used as a fire-starter (D'Orbigny). 

Peru (perhaps). Bolivia; Amazonian Brazil. "Chonta." 

Astrocaryum huicungo Damm. ex Burret, Repert. Sp. Nov. 
35: 146. 1934. 

Acaulescent; leaf rachis medially 3-angled, early furfuraceous be- 
neath, densely aculeate; segments many, regularly disposed, long- 
linear, abruptly contracted at base, about 3 cm. wide, smooth, 
glabrous, blue-green (glaucous) beneath, marginally spinulose toward 
apex; spadix (branched part) nearly 2.5 dm. long, the many branches 
as the rachis aculeate, the male spikes rather long; fruits compactly 
crowded, thus laterally somewhat prismatic, cuneate below, mostly 
oblong-obovoid, verruculose and short-setulose, 5-8 cm. long, rounded 
to the short cusp; fruiting perianth 2-2.5 cm. high, calyx to three- 
fourths, staminodal cup to two-thirds as high as corolla, both calyx 
and corolla long-setulose-hirsute; mesocarp apparently not at all 
fleshy, dry. Resembles A.jauarense Trail ex Drude (Mart. Fl. Bras. 
3, pt. 2: 372. 1881), with long bracts at the female flowers (no trace 
of these found on this fruiting specimen) ; also the setae on the fruits 
of Trail's species are 1 cm. long (Burret); A. Ulei Burret (I.e. 147) 
from Rio Acre is also acaulescent but the glabrous female calyx is 
much shorter than the corolla and also suggests the caulescent 
A. murumura with subequal calyx and corolla, the latter setulose. 



FLORA OF PERU 401 

San Martin: Moyobamba, 800 meters, (Weberbauer, type; 147). 
"Huicungo." 

Astrocaryum jauari Mart. Hist. Nat. Palm. 2: 76, pis. 52, 65, 
fig. 1. 1823 (Burret), 1824 (Dahlgren); 125. 

Sometimes 14 meters tall, the spiny zones 3-5 cm. long; leaves 
about 3 meters long, the short petioles and costa below densely 
rufous tomentose and spinescent; segments linear, long-acuminate, 
equidistant, arcuate-pendulous from conduplicate base, 7-8 dm. long, 
remotely aculeate marginally, silvery beneath; spadix nearly 1 meter 
long, hidden among the leaves; peduncle white tomentose, erect, 
4 dm. long, branches many, 3^t dm. long, rachis tomentulose; spathe 
ventricose, acuminate, fuscous tomentose and aculeate; scrobiculi 
of 3-6 flowers 2-3 cm. distant, a single female (glabrous, ciliate as 
the subequal corolla) with 2 often rudimentary male, this lower part 
finally equaled by the thick upper spike (1.5 dm. long) of cuspidately 
bracted immersed male flowers; stamens included, disk fleshy, calyx 
as the one-third to one-half longer corolla, 3-parted; fruit insipid, 
glabrous, greenish-yellow, obovoid, shortly acuminate, 4-5 cm. long, 
2.5-3 cm. in diameter, putamen 3-3.5 cm. long, 2-2.5 cm. in diam- 
eter, the slender black fibers subparallel. Inundated areas, forming 
the "jauary woods" (Huber). Illustrated, Wallace, Palms Amazon, 
pi 42. 

Loreto: Soledad, lower Rio Itaya, (Tessmann 5210). Amazonian 
Brazil. "Huirirma," "jauari" (Brazil). 

Astrocaryum macrocalyx Burret, Repert. Sp. Nov. 35: 150. 
1934. 

Low, perhaps to 9 meters tall, the caudex 2.5 dm. in diameter 
with 15 leaves (Tessmann); leaf-rachis narrowly 3-angled, setulose 
and aculeate beneath, the segments regularly disposed, not redupli- 
cate, the lower (as seen) 6 dm. long, 5 cm. wide, the upper gradually 
wider, all with secondary nerves not at all plicate, fragile, smooth 
except marginally spinulose, lustrous above, paler but scarcely sil- 
very beneath; upper spathe (apical part) 6 dm. long, fuscous setose- 
velutinous and aculeate; peduncle spadix to 2 cm. long, aculeate, 
pilose, the bracts oblong or lanceolate, the branched part 4 dm. long, 
aculei or setae lacking, branches many; female flowers at base of each 
branch 1, sessile, the bract little produced, covered by the trichomes 
and setae of the truncate calyx, the longer corolla narrowed above, 
furfuraceous and densely black-setulose as the pistil, staminodal an- 



402 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

mil us present; male flowers all fertile (spikes pedunculate, yellowish- 
pilose), in 6 vertical series, calyx small, narrowly 3-denticulate, petals 
obovate, cucullate; fruits (by mutual pressure) prismatic below, obo- 
void-oblong, 6.5-8 cm. long (beak 5 mm. long), finely setulose; peri- 
anth 1.8 cm. high, calyx half as long as corolla, both setulose, 
mesocarp more or less reticulate fibrous. Resembles A. murumura 
Mart, with calyx and corolla of female flowers subequal. A. uro- 
stachys Burret (I.e. 151), of Rio Napo, Ecuador, so probably also in 
Peru, has peduncles without aculei or setae, and rudimentary flowers 
at the ends of the male spikes. 

Loreto: On the Maran6n, (Hoppll26, type). Iquitos, (Tessmann 
5117, probably; Burret). "Huicungo" (Tessmann). 

Astrocaryum murumura Mart. Hist. Nat. Palm. 2: 70, pis. 
58, 59. 1823; 148. 

Caudex rarely attains 6 meters, 2.5 dm, in diameter, the coma 
globose with a mixture of leaves, sheath remains and spines often 
several to 10 dm. long; petioles aculeate, leaves 3-4 meters long, 
segments 30-40 or more both sides, unequally distant, linear-lanceo- 
late, falcate-acuminate, about 5 dm. long, half as wide, mostly un- 
evenly floccose-tomentose below, secondary nerves mostly 3, very 
prominent; spadix about 1 meter long, the long peduncle tomentose 
and aculeate, the upper spathe fuscous setose and aculeate, rachis 
1 dm. long, branches 200 or so, 1-1.5 dm. long, pedicels 3-^4 cm. long; 
male flowers 3 mm. long, half emerged, turbinate-obovoid; female on 
dilated base of branches, 1 cm. high, 6 mm. across, calyx glabrescent, 
about equaled by the campanulate setose corolla; stigmas elongate, 
exserted; fruit 5-6 cm. long, 4 cm. in diameter above the middle, 
sparsely setulose, putamen turbinate-obovoid, arcuately lined and 
coarsely stellate a little below the acute apex. After Drude. Said 
by Spruce (Journ. Linn. Soc. 11 : 157. 1871) to grow along the Hual- 
laga to the foot of the Andes, and by Huber (Bull. Herb. Boiss. se>. 2, 
6: 267. 1906) to be found in nearly all the Amazonian region, includ- 
ing the upper Purus, at the edge of flooded zones; not included in 
Peru, however, by Burret. F.M. Negs. 18572; 185720. 

Peru (cf. above). Amazonian Brazil. "Huicunga," "uicungu," 
"murumura" (Martius). 

Astrocaryum tucuma Mart. Hist. Nat. Palm. 2: 77, pi. 65, 
fig. 2. 1823; 119. A. princeps Barb. Rodr. Enum. Palm. 22. 1875, 
fide Burret. 



FLORA OF PERU 403 

Caudex 15 meters tall or taller, the spreading spines 1 dm. long; 
spathes at insertion strongly tumid, spinescent, the 8-12 (or more?) 
leaves 2-3 meters long, segments 3-4-aggregate, costa aculeate, 
whitish-tomentose beneath, setae ciliate, linear-acuminate, 6-8 dm. 
long, 3-4 cm. wide; spadix 1.5 meters long, spathe smooth below, 
aculeate toward long-acuminate tip, rachis smooth, branches slender, 
finely tomentulose, 400-500, erect, 2.5-3 dm. long; female flowers at 
nearly 1 dm. from base, the lageniform setulose calyx deeply parted, 
subequaling the glabrous corolla, scarcely 1 cm. long, the staminodal 
cup less than half as long; male flowers immersed, minutely bracteo- 
late, calyx minute, 4-5 times shorter than corolla; fruit glabrous, 
putamen acute at base, obovoid, rostrate. Included by Dahlgren 
in his list but possibly replaced entirely within Peru by similar spe- 
cies as A. chambira Burret, which compare, but may be known as 
near as Rios Solimoes and Japurd (Burret). 

Peru (perhaps). Amazonian Brazil. 



27. BAGTRIS Jacquin 

Guilielma Mart. Palm. Fam. 21. 1824. Pyrenoglyphis Karst. Fl. 
2: 141. 1866. Martinezia R. & P. Prodr. Fl. Peruv. 148, pi. 32. 1794, 
in part. Amylocarpus Barb. Rodr. Contr. Journ. Bot. Rio Janiero 3: 
69. 1902, fide Burret, I.e. 168. Yuba (Barb. Rodr.) Bailey, Gentes 
Herb. 7: 416. 1947, not Amylocarpus Currey, 1857. 

Reference: Burret, Repert. Sp. Nov. 34: 167-253. 1934. 

Strikingly to inconspicuously spinescent (rarely not at all), usu- 
ally clustered (cespitose), the straight annulate commonly slender 
trunks rather rarely very tall. Leaves sometimes in part subtermi- 
nal, simple, bifurcate or pinnate, the often many segments attenuate- 
acuminate, characteristically more or less ciliate, at least at tip. 
Spathes cymbiform, ordinarily firm-chartaceous (sometimes lignes- 
cent), the inner exceeding the entire spadix, this interfoliar, simple 
or once-branched, its peduncle often soon divaricate or pendulous. 
Flowers sessile, monoecious in same spadix, typically on the female 
scattered or the sexes intermixed or the lower ternate, the central 
female; stamens 6, 9 or 12, filaments subulate and long (known), the 
erect bifid anthers affixed at bifid base, calyx annular, urceolate or 
3-parted; female calyx similarly varies, about half to nearly com- 
pletely enclosing the 3-denticulate corolla, the staminodia obsolete 
or free or infrequently (subgenus Pyrenoglyphis') forming a ring; 



404 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

ovary 3-celled. Fruit 1-celled, 1-seeded, ovoid-subglobose, sometimes 
suboblong, stigmas sessile, terminal, pericarp slightly to very suc- 
culent, endocarp osseous, more or less subapically 3-pored, seed 
pendulous, raphe reticulate, albumen uniform. Subgenus Guilielma 
has no staminal cup, and the leaf-segments are usually in more than 
one plane (Bailey), but extraordinarily characteristic are the flat 
band-like fibers radiating from the pores of the endocarp (Burret) ; 
this is usually uniformly covered with fibrous or other tissues (as 
prickles). The fruiting perianth may be somewhat accrescent, in- 
durated, forming a double merely crenate or entire cup but, especially 
in Pyrenoglyphis, it is scalloped and shallow (Bailey). As to Pyreno- 
glyphis, fide Burret, no intermediates as regards the development of 
the staminodal ring were observed and he therefore retains the genus; 
practically, this single difference may be regarded as defining a sub- 
genus and still indicate the relationships. Finally, there is the segre- 
gate Amylocarpus (Yuba). Bailey (I.e.) implied but did not add basic 
information to justify acceptance of Burret's considered judgment 
that the group more correctly is treated as a subgenus, having no 
distinctive fruit character, but (as redefined by Burret) including 
species with lower flowers 3-glomerulate, the upper only male, no 
female; this able and thoughtful student noted that the floral position 
is too variable within the genus to define, in itself, generic lines. 

Burret, who devised no key himself (I.e.), remarked that Drude's 
key to the Brazilian species is quite unnatural; the following, com- 
piled from literature only, is entirely expedient. 

Staminodial ring more or less developed (subgenus Pyrenoglyphis). 

Spines pale; leaf-segments to 6 dm. long; spadix branched. 

B. Brongnartii. 

Spines black; leaf-segments to 4 dm. long; spadix simple. 

B. concinna. 

Staminodial ring obsolete, staminodia free (Bactris). 
Endocarp fibers radiate in flat bands from pores (Burret) ; leaves 
strongly crisped; fruit colored, edible (subgenus Guilielma). 

Upper spathe sparsely brownish aculeate; spadix branches 10-12. 

B. gasipaes. 

Upper spathe densely long black aculeate; spadix branches few. 

B. ciliata. 

Endocarp (known) not fibrous as above; leaves and (or) fruits also 
different (apparently, ex char.). 



FLORA OF PERU 405 

Flowers of lower rachis or its branches typically or usually all 
ternate, 1 male, upper all single, male (subgenus Amylo- 
carpus; see also B. Lakoi). 

Calyx (female) as subincluded glabrous corolla cupulate or 
subcylindric, in fruit subequal; spathes smooth, spadix 
small, not setose. 
Primary nerves 9 or 10; spadix simple. 

Leaves deeply bifurcate, glabrous B. Killipii. 

Leaves submedially bifurcate, apex setulose. . .B. naevia. 

Primary nerves 14; spadix bifurcate B. amoena. 

Calyx (also fruiting) low, 3-lobed, corolla much longer, pilose, 
cylindric; upper spathes setose or aculeate. 

Leaves shortly bifid B. hirta. 

Leaves pinnate. 
Leaf-segments 10 pairs or fewer, setose-ciliate. 

Segments narrowly lanceolate B, cuspidata. 

Segments oblong-lanceolate B. mitis. 

Leaf -segments many B. Lakoi. 

Flowers not so regularly disposed by sex, usually 2 male with 
1 female, and single females also scattered; sometimes (as 
B. Lakoi) single, or several male may occur between the 
ternate groups (Bactris sens. str.). 
Leaf -segments few or leaves merely parted. 

Spadix simple; fruit glabrous B. angustifolia. 

Spadix few-branched. 

Petioles 3 dm. long, long-spinescent; fruit glabrous. 

B. fissifrons. 

Petioles 1-5 cm. long, aculeate; fruit setose. 

B. trichospatha. 

Leaf-segments many. 

Segments linear, about 12 mm. wide B. Lakoi. 

Segments 2-7 cm. wide. 

Many segments 3-4 cm. wide or wider. 
Leaves beneath, spathes tomentose . . . B. trichospatha. 
Leaves green, margins setulose. 

Petioles to 5 dm. long, setulose, spiny. 

B. chaetochlamys. 



406 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Petioles 6-9 cm. long, finally smooth. 

B. actinoneura. 
Many segments 2-2.5 cm. wide or narrower, if linear, not 

narrowly. 
Spines black; leaves glabrous above, bidentate. 

B. riparia. 

Spines yellowish; leaves green both sides, acuminate. 

B. chloracantha. 

Bactris actinoneura Drude & Trail, ex Drude in Mart. Fl. Bras. 
3, pt. 2: 344, pi 76. 1882; 211. 

Caudices 2 or 3, a meter or two (-3) high, 2-3 cm. thick, annulate, 
the spines black, compressed (as on sheaths), 2-3 cm. long; leaves 
about 8, 2.5-3 meters long, equally pinnate, sheaths 4-5 dm. long, 
petioles slender, 6-9 cm. long and, as costa, finally smooth; segments 
20-25 each side, unevenly disposed, sometimes binate, sessile, broadly 
ovate-arcuate, strongly falcate-acuminate, rather abruptly caudate, 
2-3 dm. long, 4-7 cm. wide, smooth, multinerved, one prominent 
above, the 14-16 secondary radiating from base, many confluent at 
tip, obscurely or not setulose on margins; spadix nutant, about 2 dm. 
long, spathe shorter, densely black-aculeate; peduncles scarcely setu- 
lose, rachis 2 cm. long, the 6-12 branches slender, 8-12 cm. long, 
female flowers to above the middle, approximate, esetulose, after 
anthesis 5 mm. long, calyx 3-lobed, glabrous, corolla twice as long, 
urceolate, 3-dentate; male flowers and fruit unknown. F.M. Neg. 
18601. 

San Martin: Tocache, (Poeppig). Loreto: Yurimaguas, (Poep- 
pig 2072). Amazonian Brazil. 

Bactris amoena Burret, Repert. Sp. Nov. 34: 180. 1933. 

Type solitary, 1 cm. in diameter, internodes 11-14 cm. long; peti- 
oles 2.5 dm. long, rather stout, quite smooth as the sheath this 
fuscous furfuraceous, apex broadly ovate-produced and the simple 
leaf-blade, not medially bifurcate; leaves obovate, cuneate to base, 
primary nerves 14, slender, glabrous both sides, about 5.5 dm. long, 
3.5 dm. wide, the lobes subovate, shortly incurved-acuminate, 27 cm. 
long (on upper margin) ; spadix bifurcate, smooth as the fragmentary 
spathe; free part of fruiting peduncle 2 cm. long, recurved, branches 
5-6 cm. long, 3-flowered glomerules to apices, not alveolate, densely 
spiralled; fruit smooth, ovoid, robustly rostrate, 7 mm. wide, 12 mm. 



FLORA OF PERU 407 

high, with perianth, this cupuliform, 3 mm. high, the 3-lobed calyx 
two-thirds as long as the 3-lobed corolla. Suggests B. simplex Burret 
(I.e. 179), with 10 primary nerves and aculeate petioles 13 cm. long; 
less, B. integrifolia Wallace, abundantly spinescent (author) ; species 
range of variation is of course unknown; it also simulates B. bifida 
Mart. (Pyrenoglyphis, fide Burret). Type from Rio Putumayo (Rio 
lea) on the boundary (Lako 4)> 

Peru (no doubt). Adjacent Colombia or Brazil. 

Bactris angustifolia Damm. Verh. Bot. Ver. Brandenb. 48: 128. 
1906 (-7); 201. 

Type a meter or two high, the simple leaves with spiny petioles 
10-13 cm. long (spines few, to 6 cm. long, brown); sheath fibrous; 
rachis to 22 cm. long, blade medially bifid, 48 cm. long, 7 cm. wide, 
cuneate, 8-nerved each side, the apical, little divaricate lanceolate 
lobes 2.5 dm. long, 4 cm. wide, attenuate to base, marginally aculeate; 
lower spathe fusiform, the appressed aculei brown, 2-3 mm. long; 
peduncle 7-8 cm. long, 2 apical branches 3.5-6 cm.; flowers approxi- 
mate (injured) ; fruits black. B. sphaerocarpa Trail, 201, with leaves 
more than medially bifid or biparted, the lobes basally linear-lanceo- 
late, 6-8-nerved, and widely known rather near Peru, may occur 
there; type of Dammer's species from headwaters of the Rio Jurua, 
near the boundary, may be a variant. 

Peru (undoubtedly). Adjacent Brazil. 

Bactris Brongnartii Mart. Palm. Orbign. 59. pi. 7, figs. 2, 28 A. 
1847 (fig. 2 under name B. maraja) ; 251. Pyrenoglyphis Brongnartii 
(Mart.) Burret, Repert. Sp. Nov. 35: 251. 1934. B. maraja Mart. 
Hist. Nat. Palm. 2: 93, pi. 71, fig. 1. 1823 as to fruiting spadix, name 
probably based on a mixture (Burret). B. rivularis Barb. Rodr. 
Enum. Palm. 36. 1875. 

Stems usually clustered, rarely 10 meters tall, cylindric or con- 
stricted at the nodes, the internodes stouter above, unequal in length 
and in the upper part densely aculeate, the spines pale, divaricate; 
petioles, spathes and leaf -rachis also whitish-aculeate; leaves crowded, 
9-12, somewhat crisped, adult spreading, 3-4 meters long, segments 
to nearly 6 dm. long, 3 cm. or so wide, lanceolate, acute, marginally 
rather long-aculeate, duplicate; spadix branched, early included in 
the fusiform spathe, 3 dm. long, densely and shortly aculeate; calyx 
of female flowers shorter than 5 mm., the oblong-cylindric corolla 



408 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

included, minutely pubescent; fruit turbinate or obovate-globose, 
shortly cusped, 14 mm. in diameter, violet-black, fleshy, acid; puta- 
men depressed globose or lenticular, 3-pored. Illustrated, Barb. 
Rodr. Sert. Palm. Bras. 2: pis. 30, 31 (B. rivularis). 

Loreto: Inundated terrain, Cashiba Playa, (Tessmann 3267). 
Edge of Rio Itaya near Iquitos, (Tessmann 5086). Bolivia; Brazil. 
"Chontilla," "nieja," "niejilla," "tana," (all Tessmann). 

Bactris chaetochlamys Burret, Repert. Sp. Nov. 34: 208. 1934. 

Caudex (type) to 5 cm. in diameter; sheaths as petioles these 
to 5 dm. long fuscous furfuraceous, minutely black setulose and 
aculeate with rather dilated spines, to 4 cm. long, nearly subulate, 
to 7 cm. long on the petioles; rachis 11 dm. long, furfuraceous be- 
neath, minutely more or less darker setulose all over; segments 20-22, 
only the upper rather regularly disposed, a few of the remaining aggre- 
gate, the larger 3.5 dm. long, 4 cm. wide, all thin, lightly but clearly 
sigmoid, green both sides, glabrous but marginally setose-ciliate, ob- 
scurely bidentate, the upper part narrowly caudate, the lower obscure; 
peduncle compressed, 1.5 dm. long, velvety setulose as the 2.5 cm. 
long rachis; branches about 8; male flowers crowded, disposed all 
over, as the scattered female; fruit glabrous, to 2 cm. long, 17 mm. 
in diameter, cusp 1.5 mm. long, perianth 3 mm. high, calyx striate, 
fuscous setose, as the little longer corolla (about 2 mm. long), at last 
medially multilobed; putamen subglobose; staminodal cup none. 
No doubt the upper spathe has the indument of the peduncle; notable 
are the long thick serpent-like setae of the perianth (author). Type 
from upper Rio Napo near the boundary with Peru. 

Loreto (surely). Amazonian Ecuador. 

Bactris chloracantha Poeppig ex Mart. Hist. Nat. Palm. 2: 
145. 1837; 234; 346 (Drude). 

Clustered (cespitose) stems about 2 meters tall, 8 cm. in diameter, 
more or less aculeate with straw-colored subterete spines, especially 
abundant on the sheaths and petioles, the latter also white-floccose, 
3-5 cm. long; leaves equally pinnate, 6-9 dm. long, the segments 
2-5-aggregate, many, linear-oblanceolate, long-acuminate, caudate, 
2-2.5 dm. long, 2-2.5 cm. wide, green both sides, margins setulose; 
spadix 2 dm. long or longer, slender (upper spathe aculeate, Martius), 
peduncle compressed, 2-3 cm. long, branches about 12, a dm. or so 
long, floriferous to base; female flowers few, disposed to above the 
middle of the branches, 5-6 mm. long, the urceolate corolla at least 



FLORA OF PERU 409 

twice longer than calyx, glabrous; fruit ovoid, acute, glabrous, dark 
violet. F.M. Neg. 18606. 

Loreto: Yurimaguas, (Poeppig 2107, type). Brazil. "Cachi- 
rana," "caxirama." 

Bactris ciliata (R. & P.) Mart. Hist. Nat. Palm. 2: 95, pi 71. 
1823. Martinezia ciliata R. & P. Syst. Veg. 295. 1798. Guilielma 
ciliata (R. & P.) Wendl. Kerch. Palm. 246. 1878; 240. 

Tall, the trunks and leaf-rachises aculeate and hirsute with spread- 
ing white trichomes; leaf -segments equidistant, linear-lanceolate, 
long-acuminate, to 4 dm. long, about 3 cm. wide, marginally and 
midnerves above setose-aculeate, hirtellous below, the midnerve 
prominent above, not decurrent, the secondary and tertiary strongly 
approximate, glabrate only above; spathe 4 dm. long or longer (terete 
acumen about 12 mm. long), early fusiform, lignescent, completely 
covered with setiform aculei 12 mm. long, more robust on the ante- 
rior part and imbricate, like an animal pelt, but toward apex of spathe 
the covering spreading and reversed; spadix branched; fruit obovoid- 
oblong, putamen obovoid (Martius). This palm has the trunk cov- 
ered with long black sharp spines; its wood is black on the exterior, 
solid, and very hard; nevertheless, it is easy to work lengthwise so 
that the Indians use it for bows, arrow points, canes and blow-pipes, 
beautifully polished; shoots are tasty, raw or cooked (Ruiz & Pavon). 
The upper spathe is covered with long black prickles; the spadix with 
young flowers is very small and has only a few branches (Burret, 
after the type). Burret adds that the species is rather near G. gasi- 
paes (HBK.) Bailey. 

Huanuco: Pozuzo, Cuchero, (Ruiz & Pavdn, type). Junin: 
Chanchamayo, Huabal, (Ruiz & Pavdn). "Chonta." 



Bactris concinna Mart. Hist. Nat. Palm. 2: 99, pi. 72, figs. 
1823; 242; 335 (Drude). Pyrenoglyphis concinna (Mart.) Burret, 
Repert. Sp. Nov. 34: 242. 1934. 

Gregarious, several meters tall, the stem about 2 cm. in diameter, 
remotely annulate, aculeate above (as petioles) with subulate-com- 
pressed black spines 2-3 cm. long; leaves 1.5-2 mm. long, petioles 
2-3 dm. long, rachis less spiny, toward apex smooth; segments 30-40 
each side, mostly 3 cm. distant, often alternate, linear-lanceolate, 
the longer lower 3-4 dm. long, to 2 dm. long toward apex, 1.5-2 cm. 
wide, acute or the lower margin caudate, 1-nerved, the margins black 



410 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

ciliate; spadix stout, simple, finally nutant, long peduncle compressed, 
the lower spathe membranous, ferrugineous, 5 cm. long, the upper 
cylindric-clavate acuminate, nearly 2 dm. long, finely black setose 
and aculeate; rachis shorter than peduncle, male flowers congested, 
surrounded by female on lower half, these 3-4 mm. long, urceolate 
setulose calyx little longer than the hirsutulous corolla, staminodal 
cup present; fruits many, lustrous, dark violet, ellipsoid-obovoid, by 
pressure sometimes angled, 2-3 cm. long, half as thick, putamen 
scarcely 1 mm. thick, 3 stellate pores medial or a little above the 
middle. B. aristata Mart. (P. aristata Burret, 242), perhaps occur- 
ring, has fewer leaf-segments, rachis of spadix mostly biparted, setose 
corolla 2-3 times longer than calyx (female), fruit glabrous. Another 
upper Amazon species is B. gaviona Trail (P. gaviona (Trail) Burret, 
246), the 30-35 segments equal, the peduncle stout with 4 long 
branches, the female corolla 2-3 times longer than calyx, inflated, 
the fruit setose-scabrous. Spruce (Journ. Linn. Soc. 11: 147. 1871) 
treated B. concinna Mart, as a subgroup (Endochlamys). Illustrated, 
Mart. Fl. Bras. 3, pt. 2: pi 78, fig. 1; Barb. Rodr. Sert. Palm. Bras. 2: 
pi. BE. Wood used in making weapons (Tessmann). F.M. Negs. 
18609; 18610. 

Loreto: Among shrubs of shores and flooded terrain, (Tessmann 
3263). To Colombia and Amazonian Brazil. "Niejilla." 

Bactris cuspidata Mart. Hist. Nat. Palm. 2: 101, pi 73 B, 
figs. 1, 2. 1823; 185. 

Stem scarcely 5 mm. thick, smooth, glabrous below, fuscous to- 
mentose above; leaves 4 or 5, mostly 6-7 dm. long, sheath and petiole 
aculeate (spines dark, 5-10 mm. long), costa smooth, 3 dm. long, the 
segments 5-8 pairs, narrowly falcate-lanceolate, caudately acumi- 
nate, 1-1.5 dm. long, 2.5-3 cm. wide, 1-nerved, secondary nerves 
many, prominent, glabrous except for the sparsely setose margins 
and nerves or rarely setulose beneath; spathe sparsely long-setose; 
spadix below the leaves, erect, 6 cm. long, peduncle hidden in sheath 
and spathe, rachis (about 3 dm. long) divided into 2 or 3 branches; 
upper spathe about 6 cm. long, fusiform, coriaceous, tomentose and 
early capillaceously setulose; female calyx annulate, 3-lenticulate, 
corolla 3 times longer, hirsutulous. According to Burret the var. 
mitis of Drude (not Mart.) is B. ucupensis Spruce (Journ. Linn. Soc. 
15: 150. 1871), calyx higher, spathe not setulose; perhaps, as sug- 
gested by Burret, is a variant; like B. cuspidata and B. mitis Mart, 
known from western Amazonian Brazil so may well occur in Peru. 



FLORA OF PERU 411 

Type from Rios Yapura and Solimoes, so probably, as listed by 
Dahlgren, in Peru. F.M. Neg. 18611. 
Peru (cf. note). Brazil. 

Bactris fissifrons Mart. Hist. Nat. Palm. 2: 103, pi. 73B, figs. 
3, 4- 1823; 202; 327 (Drude). 

Stem unarmed, 2-3 meters tall, 1-2 cm. in diameter, vaginate 
above; petioles slender, about 3 dm. long, spinescent above with 
black teeth, aculei 1-1.5 cm. long, leaf rachis 4 dm. long, smooth as 
the 2-4 segments, the upper broadest, multinerved, marginally black- 
setose; peduncle nutant, upper spathe fusiform-acuminate, aculei 

1 cm. long, appressed, black, finally deciduous; spadix 2 dm. long or 
longer, branches 2-4, fastigiate (rachis obsolete), spreading, densely 
flowered to base; female flowers (male unknown) 4-5 mm. long, calyx 
cupulate, 2 mm. long, corolla appressed setulose, twice as long; fruit 
ellipsoid, mucronate-rostrate (5 mm. long), glabrous, 2.5 cm. long, 
1.5 cm. in diameter, putamen depressed gibbous-turbinate. B. sphae- 
rocarpa Trail (Journ. Bot. 15: 8, pi. 184- 1877; Fl. Bras. 325), widely 
distributed near Peru, has sheath and petioles aculeate or smooth, 
leaves simple and usually deeply bifid or with 1 opposite pair of seg- 
ments, spadix simple, short-cylindric, calyx as longer corolla glabrous 
or ciliate, in fruit lacerate; said by the author to be variable; Dam- 
mer proposed another, possibly distinct, as B. angustifolia (Verh. Bot. 
Ver. Brandenb. 48: 128. 1906 [-7). F.M. Neg. 18612. 

Peru (no doubt). Adjacent Brazil; Colombia. 

Bactris gasipaes HBK. Nov. Gen. & Sp. 1 : 302. 1815. Guilielma 
gasipaes (HBK.) Bailey, Gentes Herb. 2: 187. 1930; 237. G. speciosa 
Mart. Hist. Nat. Palm. 2: 81, pis. 66, 67. 1823. G. microcarpa Huber, 
Bol. Mus. Goeldi 4: 476. 1904; 239, essentially a nomen nudum. 

Trunks usually cespitose, 20 (-30) meters tall, lightly annulate, 
long (3 cm.) aculeate, with a coma of incurved crisped leaves about 

2 meters long; segments approximate, many, linear-lanceolate, long- 
acuminate, about 6 dm. long, 3 cm. wide, midnerve prominent above, 
the 3 or 4 secondaries both sides, margins remotely setulose or mi- 
nutely aculeate; upper spathe sparsely brownish aculeate; spadix 
early yellowish tomentose (peduncle aculeate), branches 2-3 dm. 
long; male flowers crowded, 5-6 mm. long, calyx 1.5 mm. long, sub- 
patelliform, corolla as subconnate disk fleshy, turbinate globose; 
female flowers 7-8 mm. long, calyx 1 mm. long; fruit mostly 4 cm. 
long, 3 cm. in diameter, applanate from base or concave-ovoid or 



412 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

globose-rounded, glabrous, as many as 70 or 80 to a spadix, the puta- 
men (often aborted) central, acutely turbinate from base, 2 cm. long, 
half as thick. 

Burret (I.e. 237-239) gave a re'sume' of the evidence relating to 
the probable origin of the species and suggested that G. microcarpa 
Huber (I.e.), fruit scarcely 1.5 cm. thick, may be the native state; 
this seems to be borne out by Spruce (Journ. Linn. Soc. 11: 81. 1871), 
who, observing that even in the Andes the large-fruited examples are 
around habitations, expressed the opinion that a native small-fruited 
form might still be found; he noted a small seedless variety (Wallace, 
I.e., found the seed usually aborted). Further, Spruce gave a prac- 
tical description: the clustered stems grow to 30 meters high and are 
thickly armed with long prickles; the numerous curling and droop- 
ing leaves rarely exceed 2 meters and have 50-60 leaflets on each 
side, aggregate by threes and fours and pointing in all directions; the 
fruits are massed into large pendulous corymbs; and if from their size 
and vivid colors of yellow and red they may be likened to a well- 
ripened peach (hence the name Peach Palm) in shape they more re- 
semble a hen's egg, although usually rather more conical (!) ; the thick 
firm flesh is mealy when cooked, something between chestnut and 
potato in flavor and superior to either. Tessmann, as recorded by 
Burret, 237, described the popular "masato" of Loreto, made as 
"chiche" or, less conventionally, by the use of sugar; after four or 
five days of fermentation, the drink was "very potent." Illustrated, 
HBK. I.e. 7, pi. 700; Wallace, Palm, pi 3; Contr. U. S. Nat. Herb. 16: 
opposite 308, pis. 55, 56; Weberbauer, 148, pi. 2 A. 

Loreto: Pampa de Sacramento, (Huber, type, G. microcarpa). 
Yarina-cocha, middle Ucayali (Tessmann, perhaps, fide Burret). 
Typical state apparently only in cultivation. "Pijuanyo," "pifuayo" 
or "sara-pifuayo" (Tessmann), "pisho-guayo" (bird fruit, Quechua), 
"chonta ruru," "chonta dura." 

Bactris hirta Mart. Hist. Nat. Palm. 2: 104, pis. 60, 74, figs. 1-3. 
1826; 184. B. mollis Damm. Verh. Bot. Ver. Brandenb. 48: 129. 1907, 
fide Burret. 

Stems a meter or two high, a half cm. or so thick, not at all spiny 
but fuscous tomentose above and the petioles (5 cm. long) as leaf 
rachis densely but softly black-setose; leaves cuneate-obovate, shortly 
bifid, more or less lanuginose beneath, pilose above, lobes triangular, 
scarcely acute, nerves about 12, width below sinus 9-13 cm., length 
2.5 dm.; spadix infrafoliar, bifid, erect-spreading in fruit, 5-10 cm. 



FLORA OF PERU 413 

long, outer spathe reddish-tomentose, inner coriaceous, fuscous-gla- 
brous; peduncle compressed, setose, 2-4 dm., the 2 branches half 
as long, male flowers approximate above; female corolla setulose, 
2.5 mm. long, long-exceeding the calyx; fruit olive-black, acutely 
cusped, 1 cm. long, 8 mm. in diameter, sparsely setulose. Leaves of 
B. mollis as to type a little deeper parted, the fruit somewhat smaller, 
differences not specific (Burret). F.M. Neg. 18614. 

Loreto: Leticia, Ule 6221 (type, B. mollis). Adjacent Colombia; 
Brazil. 

Bactris Killipii Burret, Repert. Sp. Nov. 34: 175. 1933. 

Type (about 2 meters tall) smooth but the sheaths (12 cm. long) 
and inner spathes fuscous furfuraceous, the peduncles early arachnoid 
tomentose, larger than the spike (4 cm. long) ; petioles 3dm. long, at 
first fuscous leprose; leaves simple, deeply bifurcate, glabrous, 4.6 dm. 
long, each lobe with 9 direct primary nerves; lobes to 3.5 cm. wide 
at base, strongly dilated above, rather abruptly contracted and nar- 
rowed in the upper third, the inner margin straight, 4 dm. long; 
spadix simple, 12 cm. long; dilated part of spathe about 6.5 cm. long, 
acute, slightly produced; glomerules 3-flowered, dense, not alveolate, 
male flowers solitary toward apex; female calyx tubular, 3-dentate 
as included corolla, pistil glabrous, staminodal tube none. Resem- 
bles B. Luetzelbergii Burret, 174, and B. Huberiana Burret, I.e., in the 
directly disposed nerves and inner margins but the lobes are much 
wider and strongly dilated; in the latter the rachis is 12 cm. long, 
lobes 26, 2 cm. long, and as it is known from adjacent Colombia and 
Brazil it may be found in Peru. 

Loreto: Iquitos, (Killip & Smith 27305, type). 

Bactris Lakoi Burret, Repert. Sp. Nov. 34: 187. 1934. 

Slender, more or less setose-hirsute, the stems nearly 7 mm. thick 
with about 10 oblong pinnate leaves 6.5 dm. long or longer (petioles 
to 2.5 dm. long or longer) ; spathes as petioles densely black-setose, 
the slender setae whitish at the tumid base; segments hirsute both 
sides, 20-25, mostly aggregate, narrowly linear, falcate toward the 
acuminate apex, the larger intermediate to 1.5 dm. long, about 
12 mm. wide, the apical shorter, wider, all on nerves and veins both 
sides rather long-setulose, flexibly hirsute and margins ciliate; upper 
spathe 6.5 cm. long, densely black-setose; spadix branches 2 (-3), in 
fruit recurved, 5-6 cm. long, peduncles densely setose, shorter; 



414 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

branches about 3 cm. long, the 3-flowered glomerules regularly dis- 
posed (type), laxly spiralled; fruiting perianth with broadly 3-lobate 
calyx, the fuscous setulose corolla many times longer, staminodal 
cup none; fruit mammillate, striate and transversely lineolate. 
The Hopp specimen has 3 spadix branches; one, also near the base, 
is mostly male, the others have male flowers scattered between the 
usual 3-glomerate ones; allied to B. hylophila Spruce, and relations 
(more northern), but the foliage is much smaller, more delicate 
(author) . 

San Martin: (Hopp 1114}- Rio Putumayo on the Brazil bound- 
ary, (Carl Lako 1, type). Brazil. 

Bactris mitis Mart. Hist. Nat. Palm. 2: 102. 1823; 185. B. cus- 
pidata Mart. var. mitis Drude in Mart. Fl. Bras. 3, pt. 2: 329. 1881 
in part, fide Burret. 

Resembles B. cuspidata Mart, but leaves interruptedly pinnate, 
(3) 6-10 pairs, oblong-oblanceolate, about 2 dm. long, 1.5-4 cm. 
wide, the lower 2-nerved, the medial 1-nerved, the broader apical 
2-3-nerved, all glabrous or margins setose-ciliate; spadix branches 
2-3, spathes appressed fuscous setose. Type from the Rio Japura 
but similar species, probably variants, have been recorded from the 
Rios Purus and Jauary. F.M. Neg. 18623. 

Peru (probably). Adjacent Brazil. 

Bactris naevia Poepp. ex Burret, Repert. Sp. Nov. 34: 179. 1933. 

Stems arundinaceous, about 8 mm. thick, the upper internodes 
4.5-5 cm. long, smooth except the leaf-lobes toward apex black- 
setulose, otherwise glabrous or early furfuraceous as the sheaths, 
these ovate below the petiole, the latter (seen) 16.5 cm. long, the 
leaf-rachis 1.5 dm. long and the upper spathe (free part) 6.5 cm. 
long; leaves nearly obovate, rather shortly contracted, scarcely to 
the middle bifurcate, about 27 cm. long, 16.5 cm. wide, primary 
nerves 10, lobes short, subovate, abruptly acuminate, 14 cm. long; 
spadix simple, to 7.5 cm. long, broken sheath 4.5 cm. long, spike 
glabrous, 3 cm. long, the glomerules 3-flowered to apex, spiralled, 
lightly alveolate; perianth in fruit glabrous, calyx 3-lobed as corolla, 
subequal, nearly 5 mm. across; staminodal cup none; fruit small 
(pea size), glabrous, globose, 6-7 mm. in diameter. Confused with 
B. simplicifrons Mart., 176, of the lower Amazon, typically at least 
with deeply parted leaves, the lobes sigmoid. B. brevifolia Spruce, 
178, the known range more northern, has rachis 1 dm. long, leaf lobes 



FLORA OF PERU 415 

(as to original) about 18 cm. long and 8 cm. wide (author) ; of course 
the stability of these observed differences and thus their genetic sig- 
nificance are as yet unknown. Trail, Journ. Bot. 15 (6): 1. 1877, 
treated the Spruce plant as a variety of that of Martius, in which 
Drude, I.e. 321, included it. F.M. Neg. 31315. 

Loreto: Drier woods, Yurimaguas, (Poeppig 2475, type, Herb. 
Munich; at Vienna, Poeppig 2476). Brazil (Rio Madeira). 

Bactris riparia Mart. Hist. Nat. Palm. 2: 97, pi. 71, fig. 4- 1823; 
220; 353 (Drude). 

Crowded, stoloniferous, spiny at the nodes, a few to 15 meters 
tall, 5-10 cm. in diameter, the spines black, 1-7 cm. long; leaves 
1.5-2 meters long, the 40-50 segments 3-5-aggregate, linear-acumi- 
nate, the apex bidentate, the upper part 4-5 dm. long, 2 cm. wide, 
glabrous above, the shorter lower part minutely pilose or setulose, 
the petioles and leaf-rachis sparsely aculeate; spadix 3-4 dm. long, 
peduncle minutely aculeate, elongate, the shorter (1 dm. long) rachis 
with 20-30 branches; upper spathe fusiform, long-acuminate, seti- 
form-aculeate especially toward tip, the scarcely pungent setae 
spreading, black; male flowers ochroleucous, calyx minute, corolla 
3-4 times longer, female flowers rarely scattered on the middle of 
the branches, calyx 1 mm. long, corolla cylindric, many times longer, 
white tomentose and appressed setulose; fruit spheroid, acutely 
cusped, glabrous, 1.5-2 cm. in diameter. The similar B. longifrons 
Mart. (I.e. 106), of the lower Amazon, listed from Peru by Dahlgren, 
is probably a lapsus; spathe tomentose between the brown spines, 
these 2.5 cm. long, terete. F.M. Neg. 18628 (B. longipes, 18618). 

Peru (perhaps). Adjacent Brazil. 

Bactris trichospatha Trail, Journ. Bot. 15: 41. 1877; 204; 339 
(Drude). 

Stems clustered, rhizomatous, rarely 3 meters tall, 1.5-2.5 cm. 
thick, usually aculeate as the sheath, petioles (1-5 cm. long) and 
ordinarily leaf rachis; leaves 1.5-2.5 meters long, segments many, 
one variety 4-6, broadly lanceolate, falcate, long-acuminate, mostly 
1.5-2.5 dm. long, 3-5 cm. wide, reduced toward apex, secondary 
nerves 5-7 (9-12), glabrous above, velutinous beneath, all except 
apical, free; spadix erect, 2-3 dm. long or longer, upper spathe lig- 
neous, reddish tomentose, branches 6-10, fastigiate; male flowers 
white, 5 mm. long, calyx segments subulate, petals ovate, female 



416 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

setose, usually twice longer than calyx, few toward the apex of the 
branches; fruit dark violet, subglobose, setose, 1.5 cm. in diameter. 
Peduncle and inner spathe covered with short brown bristles re- 
sembling velvet (Trail). Trail lists Tabatinga at the boundary with 
Peru as one of several localities. B. longisecta Burret (I.e. 205), re- 
lated, has linear-lanceolate segments 5.5-6 dm. long, 4.5 cm. wide, 
the upper much smaller. F.M. Neg. 18635. 

Loreto (undoubtedly). Adjacent Brazil. 



28. DESMONCUS Mart. 
Reference: Burret, Repert. Sp. Nov. 36: 197-221. 1934. 

Supported by the modified spine-like segments of the whip-like 
rachis end of the pinnate leaves or scandent, the stem in time much 
elongated, remaining slender, the leaf-rachis with straight or more 
or less aculeate (Peru), mostly (or entirely) recurved prickles. Upper 
spathe often recurved-aculeate from a conico-lagenif orm base (Peru) . 
Stamens 6 or sometimes 9. Otherwise like Bactris, and as various 
in floral characters. Name conserved. 

Rachis scarcely filiform, the branches probably many. 

Leaf-segments long-acuminate both ends D. prunifer. 

Leaf -segments acute at base D. longifolius. 

Rachis subfiliform as the few (if any) branches. 

Spadix branches 3-4 D. leptospadix. 

Spadix perhaps simple D. vacivus. 

Desmoncus leptospadix Mart. Palm. Orbign. 10: 52. 1847; 216; 
308 (Drude). Atitara leptospadix (Mart.) Ktze. Rev. Gen. 727. 1891. 
D. leptoclonos Drude ex Damm. Verh. Bot. Ver. Brandenb. 48: 129. 
1907, fide Burret. 

Stem diameter 5 mm.; sheath elongate, membranous, smooth or 
shortly setose-aculeate; leaves about 1 meter long, rachis shortly and 
sparsely aculeate, segments exactly opposite, foliose 3-4 pairs, the 
larger lanceolate or elliptic, acuminate, 17 cm. long, 4 cm. wide, the 
narrower 2 dm. long, 2.5 cm. wide, lacking toward base, toward apex 
spiniform (3-4 pairs), trigonous-conical, the lower 1.5-2.5 cm. long, 
spreading or separated; spathe not aculeate as spadix, the latter 
2-3 dm. long, long-included, the rachis filiform, the 3-4 very slender 
branches lax, 4-10 cm. long; fruit oblong-ellipsoid, acute both ends; 



FLORA OF PERU 417 

putamen about 8 mm. long, 5 mm. in diameter. D. mitis Mart. 
Hist. Nat. Palm. 90. 1823; 217; 308 (Drude) has 15-18 pairs of lan- 
ceolate pinnae (6-8 pairs subulate, lanceolate) ; D. setosus Mart. I.e., 
315 (Drude) 9-11 pairs (4-5 subulate), linear leaf segments only 
1.5-2 dm. long, 1.5 cm. wide; both have been recorded at more than 
one station not far from eastern Peru. Liana 2.5 meters tall, 5 mm. 
thick (Tessmann). 

Loreto: Yurimaguas, (Poeppig, type). Soledad, lower Rio Itaya, 
(Tessmann 5236). Amazonian Brazil. 

Desmoncus longifolius Mart. Palm. Orbign. 10: 52. 1847; 
211. 

Sheath 4 dm. long or longer, rather densely aculeate as leaf-rachis 
to tip with straight subulate prickles and shorter hamate ones, 2-8 mm. 
long; leaf -segments 7-8 pairs, lanceolate, long-acuminate both ends, 
3 dm. long, 2.5-4 cm. wide, 2-3 pairs subulate, hamate, straight, 
strongly enlarged at nodose base, 3.5-4 cm. long. Burret (I.e.) has 
added the following information from Weberbauer specimen (spathe 
lacking) : upper part of peduncle aculeate with small straight prickles, 
black above; rachis 16 cm. long, branches about 20, scattered, basal 
bract obvious, oblong, subacuminate; flowers mostly 3-glomerulate; 
perianth (fruiting) cupulate, nearly 3 mm. high, calyx low, minutely 
3-denticulate, corolla more than twice as high; fruit ellipsoid, over- 
all 19 mm. long, 14 mm. in diameter, rather rugose, sublustrous, apex 
not at all mammillate, base shortly contracted, pulp yellow, sweet. 

Huanuco: Pozuzo, (Ruiz & Pavdn, type, Herb. Webb). Junin: 
Open wood near La Merced, (Weberbauer 1906}. "Cachapurina." 

Desmoncus prunifer Poepp. ex Mart. Hist. Nat. Palm. 2: 148. 
1837; 216. Atitara prunifer (Poepp.) Ktze. Rev. Gen. 727. 1891. 

Sheath, petiole, inner spathe and peduncle black-setose and acule- 
ate, spines conical, straight or especially on the leaf-rachis hamate, 
very dense on the lower spathe; leaf -segments 10-15 pairs, oblong, 
acute at base, long-acuminate, the 4-5 spiniform pairs subulate; fruit 
oval, 2.5 cm. long. Ascends to 10 meters; leaves (with excurrent 
rachis) 14 dm. long; spadices subaxillary, solitary, branched, longer 
than 4 dm., the peduncle nearly 2 dm. long; drupe orange, succulent, 
acid-sweet; albumen edible (Martius). Unfortunately, no other data. 
F.M. Neg. 31323. 

Loreto: Yurimaguas, (Poeppig, type). 



418 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Desmoncus vacivus Bailey, Gentes Herb. 8: 186. 1949. 

Weak, scandent, only minutely aculeate; sheaths apically retic- 
ulate; leaves about 5 pairs, opposite or nearly, lanceolate-elliptic, 
acuminate, 10-12 cm. long, 2-3.5 cm. wide, lateral nerves promi- 
nent; petioles and rachis sparsely aculeate with hamate prickles 
1-2 mm. long; terminal spiniform leaflets about 4 pairs, to 2 cm. 
long, very slender, reflexed; spadix perhaps simple; fruit ellipsoid, 
2-2.5 cm. long, 2 cm. thick, minutely cusped, smooth; seed strongly 
reticulate. No comparison made by author; it will doubtless be 
found, whatever its correct name, in adjacent Peru, as type, R. E. 
Schultes 3941, is from Rio Igaraparana near the boundary. 

Loreto (no doubt). Colombia. 



It may be appropriate to cite here, in connection with the generic 
problems referred to above, the work of P. B. Tomlinson. Tomlin- 
son's summary of systematic anatomy of the palms will appear in 
the forthcoming series, edited by C. R. Metcalfe, Anatomy of the 
Monocotyledons, published by the Clarendon Press, Oxford. 




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