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Full text of "Flora of Peru"

no. % 

FLORA OF PERU 





BOTANICAL SERIES 

FIELD MUSEUM OF NATURAL HISTORY 

VOLUME XIII, PART IV, NUMBER 2 

NOVEMBER 28, 1958 



PUBLICATION 861 



THE LIBRARY OF. THE 

DEC 1 9 1953 
UNIVERSITY OF ILLINOIS 



To renew call Telephone Center, 333-840O 




FLORA OF PERU 



BY 

ROGERS MCVAUGH 

CURATOR OP VASCULAR PLANTS, UNIVERSITY OF MICHIGAN 




BOTANICAL SERIES 

FIELD MUSEUM OF NATURAL HISTORY 

VOLUME XIII, PART IV, NUMBER 2 

NOVEMBER 28, 1958 

PUBLICATION 861 



Library of Congress Catalog Card Number 36-10426 



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



FLORA OF PERU 



ROGERS MCVAUGH 

MYRTACEAE. Myrtle Family 

Shrubs or trees or rarely subherbaceous. Leaves simple, oppo- 
site (except in some introduced genera), exstipulate, entire or rarely 
crenate, punctate with resinous or pellucid glands, usually pinnately 
veined. Mid vein usually elevated and prominent on the lower sur- 
face. Principal lateral veins usually uniting distally into a "marginal 
vein" which extends nearly the length of the blade and more or less 
parallel to the margin but somewhat separated from it. Flowers 
borne on axillary (or rarely terminal) branches, solitary or in spe- 
cialized bracteate inflorescences with opposite branching, these mod- 
ified in various ways, e.g., by elongation of the axis and reduction of 
the lateral axes to one flower each ("racemes"); by suppression of 
the axis and reduction of the lateral axes to one flower each (flowers 
in "glomerules" or "umbelliform clusters"); by reduction of the 
lateral axes to one pair, these arising just below the flower which 
terminates the central axis ("dichasium") ; by potentially indefinite 
elongation of both central and lateral axes, this resulting in a "pan- 
icle" with proximal branches elongate and a transition from these 
to short simple branches and terminal triads of flowers. Flowers 
regular or essentially so, hermaphrodite or rarely by abortion uni- 
sexual. Ovary inferior, the hypanthium adnate to the ovary its 
whole length or prolonged beyond it so that the stamens, petals and 
calyx-lobes appear to arise from the distal margin of a short tube 
surrounding the summit of the ovary. Calyx-lobes usually 4 or 5, 
distinct and imbricate, or the calyx calyptrate and circumscissile, or 
rupturing irregularly in anthesis. Petals usually 4 or 5 (sometimes 
reduced in number or size, or wanting). Stamens usually indefi- 
nitely many, in one-many series about the margin of the usually 
thickened calycine disk, usually inflexed in the bud. Filaments 
usually filiform and distinct in Peruvian species. Anthers usually 
short, versatile or basifixed, bilocular, opening (at least in Peruvian 
genera) by longitudinal slits. Style simple, elongate, with small 
capitate or peltate stigma. Ovary 2- to many-locular, the placentae 
affixed to the axis or parietal and coalesced into a central axis, the 
ovules 2 or more. Fruit fleshy or capsular. Embryo various. 

569 



570 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

The genera now referred to Myrtaceae were divided by DeCan- 
dolle among three tribes, namely, Chamaelaucieae (with dry uni- 
locular and usually indehiscent fruit), Leptospermeae (with dry 
multilocular and usually dehiscent fruit), and Myrteae (with fleshy 
multilocular fruit). This system was set forth in detail in the 
Prodromus 3: 207-288. 1828. Essentially the same arrangement 
was followed by Bentham, in Benth. & Hook. f. Gen. PI. 1: 690- 
720. 1865. Niedenzu, in Engler & Prantl, Nattirl. Pflanzenfam. Ill 
(7): 57-105. 1893, erected two subfamilies; the first, Myrtoideae, 
comprised the one tribe Myrteae, and the second, Leptospermoi- 
deae, included the two tribes Leptospermeae and Chamaelaucieae. 

The subfamily Leptospermoideae is especially developed in the 
Australian region and to a lesser extent elsewhere in the southwest 
Pacific. The total number of species approaches 1000, including 
according to some authors nearly 500 species of the vast Australian 
genus Eucalyptus. The only American member of the subfamily is 
the endemic Chilean species Tepualia stipularis (Barn.) Griseb. All 
the native Peruvian Myrtaceae are members of the tribe Myrteae, 
but several species of Eucalyptus have been introduced for shade and 
for ornament, and at least one has become widespread. 

Key to the Tribes (Peruvian representatives only) 

Fruit dry, capsular, consisting of the capsule immersed in the hard- 
ened hypanthium, the valves sometimes projecting beyond the 
rim of the hypanthium; petals and calyx-lobes united into an 
operculum which is dehiscent at anthesis; flowers usually in 
pedunculate axillary umbels; adult leaves glabrous, lanceolate 
and long-petiolate, and mostly alternate, the juvenile ones often 
broad, subsessile and more or less opposite. 

Tribe I. Leptospermeae 

Fruit fleshy (a few- or many-seeded "berry") ; petals and calyx-lobes 
free, or in a few genera united into an operculum; flowers vari- 
ously arranged, never in pedunculate axillary umbels; leaves 
opposite, rarely markedly long-petiolate .... Tribe II. Myrteae 



Tribe I. LEPTOSPERMEAE DC. 
1. EUCALYPTUS L'Her. 

Glabrous trees or shrubs, the leaves alternate, leathery, usually 
elongate, lanceolate and 8-12 cm. long or more, markedly petiolate 
and often hanging vertically; juvenile foliage (of seedlings or shoots 



FLORA OF PERU 571 

from felled trees) often broad, subsessile and more or less opposite, 
rarely hairy; flowers usually in pedunculate axillary umbels, some- 
times forming panicles; bracts and bracteoles deciduous so early as 
to be seldom seen; ovary usually 3- to 4-locular, immersed in and 
surrounded by the fleshy hypanthium which hardens in fruit and is 
prolonged beyond the summit of the ovary into a rim which bears 
the numerous stamens; petals and calyx-lobes united into an oper- 
culum which is continuous with the rim of the hypanthium in bud 
and circumscissile at anthesis; stamens widely spreading in anthesis 
and forming the showy part of the flower; style about as long as the 
operculum; ovules and seeds numerous, but only a few in each locule 
fertile. 

A large and almost exclusively Australian genus at one time sup- 
posed to include nearly 500 species; some recent authors have sug- 
gested that the actual number is somewhat smaller. A recent account 
of the Northern Australian species, by S. T. Blake in Austral. Jour. 
Bot. 1: 185-352, pi. 1-36. 1953, includes 50 species in this part of the 
continent; J. M. Black, in Fl. South Austral, ed. 2, 612-632. 1952, 
lists 52 species. 

Numerous species have been introduced into the tropical and 
warm-temperate regions of America for ornament, for purposes of 
reforestation, for wood and for lumber; only the following seems to 
have been entirely successful. 

Eucalyptus globulus Labill. Voy. 1: 153, t. 13. 1799. 

A large tree, said to reach a height of 75-90 meters, with pale 
deciduous bark and yellowish green angled branchlets; adult leaves 
alternate, lanceolate and often falcate to narrowly ovate, 2-3 (-6) 
cm. wide at base, 12-25 cm. long, (3-) 5-8 times as long as wide, 
attenuate from base to the slenderly pointed apex, the base often 
obliquely unequal-sided, abruptly rounded to the flexuous petiole 
2-4 cm. long; mid vein pale, flat or concave above, convex beneath; 
leaf-margins bordered by heavy cartilaginous veins about equal to 
the midvein but compressed at right angles to the plane of the leaf 
and often standing somewhat above and below it; lateral veins deli- 
cate and inconspicuous, joining an equally slender and nearly straight 
submarginal vein just within the cartilaginous border; foliage with 
numerous small dark glands on both surfaces; juvenile shoots and 
leaves conspicuously whitened and waxy-glaucous, their leaves oppo- 
site, sessile, ovate to oblong, cordate, abruptly short-acuminate at 
tip, 4-5 cm. wide, 7-15 cm. long; flowers large, subsessile, solitary or 



572 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

rarely 3 in an axil, on a massive, broadly 2-angled peduncle up to 
5 mm. long; buds 1.7-2.5 cm. long, conspicuously whitened by a 
heavy waxy-farinose coating; hypanthium about 1 cm. long, trun- 
cate at base, strongly 4-angled, obpyramidal, irregularly and coarsely 
warty-roughened especially on the angles and the thickened margin; 
calyptra dome-like, roughened like the hypanthium, usually with a 
broad knoblike or acute central beak; stamens 1.5 cm. long (the 
flower when expanded 3-4 cm. across), borne on the inner edge of 
the disk which projects about 3 mm. beyond the thickened margin 
of the hypanthium; style 8-10 mm. long; fruit 2-2.5 cm. broad and 
high, flat-topped or the surface convex, the 4-5 valves not exserted 
but nearly plane with the surface; seeds 1-3 mm. long, very numer- 
ous, prismatic, irregularly several-angled. 

A native originally of Victoria, New South Wales, and Tasmania, 
this species is now extensively planted and naturalized from Cali- 
fornia to Argentina and Chile, especially in high semi-arid regions; 
it is a valuable timber tree of rapid growth, and now forms a charac- 
teristic feature of the landscape in many parts of Latin America. 
According to Acosta Solis, "El eucalipto en el Ecuador," in Flora 
(Quito) 15-16: 149-194. 1945, E. globulus was introduced into Ecua- 
dor in 1865 and has become a valuable resource in the inter-Andine 
region of that country. It has likewise become abundant in Peru, 
as in Cuzco, where, according to Herrera in Contr. Fl. Depto. Cuzco, 
ed. 2, 148-149. 1921, it was introduced about 1880 and subsequently 
became general in all the provinces of that Department. 

Amazonas: Chachapoyas, 2,700 meters, Williams 7562. Junin: 
Tarma, 3,000-3,200 meters, Killip & Smith 21870. "Eucalipto." 

Another species, Eucalyptus camaldulensis Dehnh., Cat. PI. Hort. 
Camald. ed. 2: 20. 1832 (E. rostrata Schlecht., 1847, non Cav., 1797), 
has been collected near Yucay, Cuzco, by Soukup (no. 840). The 
inflorescence is a 6- to 10-flowered axillary umbel, on a peduncle 1-2 
(-3) cm. long, the pedicels 3-8 mm. long; buds ovoid, 5-10 mm. long, 
the calyptra longer than the base (up to 3 times as long), abruptly 
narrowed to a stout straight beak 1-6 mm. long; fruit nearly hemi- 
spheric, 4-6 mm. high, 5-8 mm. broad, the 3-4 deltoid or narrowly 
pointed valves projecting 1.5-3 mm. beyond the equator. 

Tribe II. MYRTEAE DC. 

Classification of the subtribes of the Myrteae has been based 
principally upon characters of the mature embryo. Such a classifi- 






FLORA OF PERU 573 

cation appears to be in the main a natural one, but practically it 
presents many difficulties. Members of this family are usually col- 
lected when in flower, at a time when it is impossible to ascertain the 
structure of the embryo. Flowers and fruits are rarely found on the 
same plant at the same time, so that any usable system of classifi- 
cation must utilize characters of flower and inflorescence in addition 
to those which may be found in the mature fruit. The subtribes 
accepted in the present treatment are those proposed by Berg, and 
the arrangement of genera follows in the main that of his "Revisio 
Myrtacearum Americae," in Linnaea 27: 1-472. 1855-56. The sub- 
tribes are characterized as follows: 

1. Subtribe Myrciinae Berg. Cotyledons foliaceous, contortu- 
plicate. Radicle elongate. 

2. Subtribe Eugeniinae Berg. Cotyledons fleshy, distinct, or 
somewhat or completely fused, or conferruminate; radicle very short. 

3. Subtribe Pimentinae Berg. Embryo spiral, subspiral or un- 
cinate-curved. Radicle elongate; cotyledons very short. 

Key to Flowering Material 

Inflorescence compound, usually many-flowered (flowers often 30- 
200 or more), with branches compound and opposite near the 
base of the panicle, becoming irregularly ternate or solitary near 
tips; calyx-lobes, if developed, usually 5; bracts and bracteoles 
usually deciduous at anthesis or before. 

Calyx-lobes evident in bud and in anthesis, usually with thin dis- 
tal and lateral margins which are imbricate or at least con- 
tiguous at base; central axis of the inflorescence well developed 
and about as long as the primary lateral branches .... Myrcia 

Calyx closed in bud or merely the tips of the lobes free, either cir- 
cumscissile or splitting irregularly between the lobes; central 
axis of the inflorescence often aborted at the node where the 
lowest lateral branches emerge, and the panicle as a whole 
seeming to consist of two nearly equal parts. 

Calyx closed in bud, calyptrate, circumscissile; petals minute or 
often wanting; malpighiaceous hairs often present and con- 
spicuous; bracts mostly deciduous Calyptranthes 

Calyx closed in bud or the tips of the lobes free, the buds open- 
ing by irregular longitudinal splitting between the calyx- 
lobes nearly or quite to the summit of the ovary; petals 



574 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

small, usually present; malpighiaceous hairs usually not 

apparent; bracts sometimes persistent Marlierea 

Inflorescence not as above, 1- to several-flowered (flowers rarely as 
many as 30), racemose, regularly dichotomous or the flowers 
solitary, sub-umbellate or glomerate; calyx various, the lobes 
usually 4 and distinct if the flowers are more than 7; bracteoles 
often persistent. 

Cymes lateral on old wood, thrice dichotomous, 2-3 cm. long, with 
persistent paired bracts and a central sessile flower in each 
fork; calyx-lobes 4; flowers red; stamens 4-8, 1.5-2 cm. long. 

Myrrhinium 

Inflorescence racemose or dichasially branched, or the flowers 
glomerate or solitary; cymes, if occasionally twice or thrice 
dichotomous, irregularly branched and in leaf-axils near the 
tips of twigs; flowers mostly white or cream-color, rarely pink 
or red; stamens usually shorter and far more numerous. 

Calyx closed or essentially so in the bud, opening at anthesis by 
irregular longitudinal slits. 

Flowers about 4, nearly sessile, glomerate (in a much abbrevi- 
ated raceme), the clusters appearing involucrate because 
of the relatively large persistent bracts; calyx-lobes 4; 
ovary bilocular, the locules 2-ovulate Plinia 

Flowers 1, or 3-7 in a dichasium, pedicellate or the dichasia 
pedunculate, not involucrate; calyx-segments usually 5; 
ovary 2- to 7-locular, the locules many-ovulate . . Psidium 

Calyx-lobes normally developed even in the bud, the calyx at 
anthesis not or scarcely splitting beyond the bases of the 
lobes. 

Calyx-lobes 5. 

Small or prostrate shrubs or subshrubs with coriaceous vac- 
cinioid leaves 1-2 cm. long or less; flowers solitary; 
bracteoles foliaceous and persistent; Andean paramos. 

Anthers sagittate, the connective dilated Ugni 

Anthers oblong or subrotund, not sagittate, the filaments 

filiform Myrteola 

Shrubs or trees with larger leaves; flowers solitary or in 
small dichasia; bracteoles deciduous; middle and low 
elevations. 



FLORA OF PERU 575 

Leaves coriaceous, subsessile, suborbicular, 1.5-7.5 cm. 
long and wide; dichasium stout, 3- to 7-flowered; 
Lima, on Pacific slopes .... Myrcianthes. quinqueloba 

Leaves membranaceous or chartaceous, petiolate or elon- 
gate or both, 1.5-3.5 times as long as wide; flowers 
solitary or in slender 3 (-7) -flowered dichasia; culti- 
vated and escaped, otherwise mostly Amazonian 
lowland. 

Flowers usually 4, in two decussate pairs at the lowest 
nodes of an axillary branch; calyx-lobes broadly 
rounded, 3-6 mm. long, spreading after anthesis; 
mature petioles 10-15 mm. long; 3-4 lowest pairs 
of veins in the leaf closely grouped, the succeeding 
ones increasingly distant Campomanesia 

Flowers 1-7 in an axil; calyx-lobes, if developed, 3 mm. 
long or less (in one species with narrow terminal 
appendages 6-14 mm. long); mature petioles 10 
mm. long or usually less; veins of leaf uniformly 
spaced, the lower ones not markedly aggregated. 

Psidium 
Calyx-lobes 4. 

Stamens 25-50, the filaments 15-22 mm. long, red; style 
15-22 mm. long, about three times as long as the 

petals; plants heavily villose or tomentose Acca 

Stamens much more numerous, or much shorter, or white; 
style rarely more than 15 mm. long (usually much 
less), usually less than twice as long as the petals; 
plants various. 

Inflorescence a raceme, the flowers in opposite decussate 
bracteate pairs; central axis of raceme abortive at 
tip (the terminal flower wanting) and the axis often 
so much abbreviated that the flowers appear glom- 
erate or umbellate. 

Calyx-tube much prolonged above the ovary, circum- 
scissile at base after anthesis; flowers usually 4, 
small (buds in our species 5 mm. long or less) and 
nearly sessile, with conspicuous paired persistent 
involucre-like bracteoles; ovules 2 or 4 in each 
locule Myrciaria 



576 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Calyx-tube little or not at all prolonged, not circum- 
scissile, persistent with the lobes in flower and 
fruit; inflorescence various; ovules 2 or many. 
Flowers glomerate (in clusters of about 4-6), mostly 
pubescent, medium-sized (buds in ours 5-12 
mm. long), the clusters appearing involucrate 
because of the relatively large persistent bracts; 
calyx in the bud very shortly 4-lobed at sum- 
mit, and later splitting irregularly; ovules 2. 

Plinia 

Flowers glomerate to racemose, not involucrate; 
calyx-lobes free their whole length, or loosely 
coherent at base; ovules mostly numerous. 

Eugenia 

Flowers solitary or in simple or compound dichasia, the 
primary axis of the inflorescence if once-forked with 
a sessile or nearly sessile flower in the fork. 
Small or prostrate shrubs or subshrubs with coriaceous 
vaccinioid leaves 1 cm. long or less; flower soli- 
tary; bracteoles foliaceous, persistent; paramos in 

the Andes Myrteola 

Shrubs or trees with larger leaves; flowers various; 

middle and low elevations. 

Flowers 3-7, in a dichasium; calyx-lobes 1.5 mm. 
long, ciliate, deciduous at anthesis with the 
bracts and bracteoles; southern Ecuador. 

Blepharocalyx salicifolius 
Inflorescence various; calyx-lobes not ciliate and 

deciduous. 

[Stigma peltate?; placentation parietal, the pla- 
centae introrse, bilamellate?] ; flowers 1 or 3, 
large, long-pedunculate, the calyx-lobes 7-9 
mm. long; Amazonian lowlands. .Psidium sp. 
Stigma small and simple, hardly broader than the 
style; placentae simple, axillary; flowers and 
inflorescence various. 

Flowers, if more than one, in simple or com- 
pound dichasia; if solitary only, then oc- 
curring in many or most of the mature leaf- 
axils, not confined to the basal nodes of a 
new branch Myrcianthes 



FLORA OF PERU 577 

Flowers, if more than one, in elongate or 
much abbreviated racemes; if solitary only, 
then mostly confined to the 1-2 lowest 
nodes of a new branch, and often in the 
axils of much reduced bracts Eugenia 

Key to Fruiting Material 

Cotyledons foliaceous, contortuplicate; radicle elongate; testa mostly 
membranaceous, fragile; calyx-lobes, if present, usually 5; in- 
florescence compound, usually many-flowered, with branches 
opposite near base and irregularly ternate or solitary near tips; 
bracts and bracteoles rarely present (i.e., deciduous about the 
time of anthesis). 

Calyx-lobes normally developed and present on the fruit, their 
basal margins contiguous; central axis of the inflorescence 
well developed and about as long as the primary lateral 
branches Myrcia 

Calyx wanting (the tip of the fruit umbilicate) or represented by 
a shrunken calyptra attached at one side, or the calyx longi- 
tudinally ruptured below the base of the lobes and the latter 
irregularly margined proximally in fruit; central axis of the 
inflorescence often aborted at the node where the lowest lat- 
eral branches emerge. 

Calyx closed in bud, calyptrate, circumscissile, the calyptra 
sometimes persistent in fruit or, if deciduous, the fruit um- 
bilicate; [malpighiaceous hairs often present and conspic- 
uous; bracts mostly deciduous] Calyptranthes 

Calyx closed in bud or the tips of the lobes free; lobes at ma- 
turity separated by irregular longitudinal ruptures extend- 
ing nearly or quite to the summit of the ovary; fruit 
crowned by the unevenly margined (usually 5) often re- 
flexed lobes, or these irregularly deciduous; [malpighiaceous 
hairs usually not apparent; bracts of the inflorescence often 
persistent on the branches after the flowers fall] . . Marlierea 
Cotyledons not as above, but small and fleshy, or large and plano- 
convex, or the embryo undivided; inflorescence 1- to several- 
flowered (flowers rarely more than 30), not branched as above; 
calyx-lobes usually 4 and distinct if the flowers are more than 7; 
bracteoles often persistent. 



578 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Cymes lateral on old wood, thrice dichotomous, 2-3 cm. long, with 
persistent paired bracts and a central sessile flower in each 
fork; calyx-lobes 4; embryo arcuate, the cotyledons hardly 
distinct, the testa hard, bony Myrrhinium 

Inflorescence racemose or apparently the flowers glomerate, or 
dichasially branched, or the flowers solitary; cymes, if occa- 
sionally twice or thrice dichotomous, terminal and irregular. 

Inflorescence a raceme, the flowers in opposite decussate brac- 
teate pairs; central axis of raceme abortive at tip (the 
terminal flower wanting) and the axis itself often so much 
abbreviated that the flowers appear glomerate or umbel- 
late in the axils; calyx-lobes usually 4. 

Seeds small, numerous, with curved embryo, long radicle and 
short cotyledons of Subtribe Pimentinae Acca 

Seeds few, one or two filling the entire fruit; embryo undi- 
vided or the cotyledons large, free, plano-convex. 

Cotyledons free, plano-convex; flowers glomerate, mostly 
pubescent, the clusters appearing involucrate because 
of the relatively large persistent bracts; calyx splitting 
irregularly longitudinally, the lobes persistent . . Plinia 

Embryo undivided or the cotyledons partly separated; 
flowers, if glomerate, not involucrate; calyx various, 
if closed in bud then not splitting irregularly. 

Calyx-tube prolonged above the ovary, circumscissile at 
base after anthesis and leaving a circular scar on the 
fruit; flowers small and nearly sessile, usually 4. 

Myrciaria 

Calyx-tube not circumscissile, scarcely or not at all pro- 
longed, the lobes persistent; flowers glomerate or in 
racemes Eugenia 

Flowers solitary or in simple or compound dichasia, the primary 
axis of the inflorescence if once-forked with a sessile or 
nearly sessile flower in the fork; calyx-lobes 4 or 5, or the 
calyx irregularly longitudinally dehiscent. 

Seeds one or two; embryo undivided or the cotyledons plano- 
convex and distinct, longer than the radicle; testa thin; 
calyx-lobes 4, usually distinct and persistent. 
Embryo undivided; flowers solitary, or racemose in some 
axils Eugenia 



FLORA OF PERU 579 

Cotyledons fleshy, distinct, plano-convex; flowers in dicha- 

sia, or partly or all solitary Myrcianthes 

Seeds several or many; embryo arcuate, uncinate or spiral, 

with long radicle, very short cotyledons and usually bony 

testa; calyx-lobes 4 or 5, or the calyx splitting irregularly. 

Calyx-lobes 5, subequal, spreading; flowers solitary at leafy 

or leafless nodes. 

Shrubs with stiff vaccinioid leaves mostly 2 cm. long or 

less; flowers borne at leafy nodes; bracteoles folia- 

ceous, persistent; fruit 1 cm. in diameter or less; 

seeds with arcuate embryo and bony testa. 

Leaves less than 1 cm. long, narrowly sagittate with 

inrolled margins and then apparently 1.5 mm. 

wide; peduncle 4-5 mm. long; filaments filiform, 

the anthers not sagittate Myrteola acerosa 

Leaves 1-2 cm. long, elliptic, 3-8 mm. wide; peduncle 
9-12 mm. long; filaments dilated and flattened, 

the anthers sagittate Ugni 

Tree with thin leaves often 10 cm. long or more; flowers 
often at leafless nodes; bracteoles deciduous at an- 
thesis; fruit 4-5 cm. in diameter; seeds with spirally 
involute embryo, tough membranaceous and gland- 
ular- verrucose testa Campomanesia 

Calyx splitting irregularly from summit to base or the lobes 
4 and distinct (if the lobes 5, the flowers 3-7 or the 
embryo merely curved or the testa bony). 
Calyx-lobes 4 and distinct (if, rarely, 5, the leaves 2 cm. 
long or less and the bracteoles foliaceous and per- 
sistent) . 

Fruit about 5 mm. long and wide; flowers solitary; 
small or prostrate shrubs or subshrubs with coria- 
ceous vaccinioid leaves 1 cm. long or less . Myrteola 
Fruit (when known) larger; upright shrubs or trees 

with larger leaves. 

Leaves membranaceous, glabrous; flowers 3 to 7, in 
a dichasium; calyx-lobes 1.5 mm. long, ciliate 
and deciduous with the bracts and bracteoles 

at an thesis; Ecuador Blepharocalyx 

Leaves coriaceous, heavily pubescent to tomentose; 
flowers solitary or in 3's or short axillary ra- 



580 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

cemes; calyx-lobes 2.5-9 mm. long, persistent; 

fruit ellipsoid, 8-15 mm. long Acca 

Calyx splitting irregularly or with 5 lobes Psidium 

Subtribe MYRCIINAE Berg 

Trees or shrubs, with the inflorescence in Peruvian species cy- 
mose-paniculate, the peduncles rarely few-flowered. Calyx-lobes if 
distinct usually 5. Cotyledons foliaceous, contortuplicate, the rad- 
icle elongate. Ovary 2-3 (rarely -4) -locular, the ovules 2 in each 
locule, collateral, affixed to the central axis. 

1. MARLIEREA Camb. 

Reference: Berg, Linnaea 27: 12-18. 1855; and in Mart. Fl. Bras. 
14, pt. 1:31-38. 1857. 

Calyx closed in the bud (or the tips of the lobes free), splitting 
irregularly in anthesis into 5 (-4) longitudinal lobes which are de- 
ciduous from, or subpersistent on, the fruit. Petals 5 (-4), small and 
inconspicuous, or none. Inflorescence-axis usually abortive above 
the first node, and the panicles consequently appearing paired. 
Bracts of the inflorescence sometimes persistent through anthesis. 

A genus of more than 60 described species, mostly confined to 
tropical South America east of the Andes. The distinction between 
Myrcia and Marlierea is not a clear one, and it is probable that the 
latter represents a phylogenetically diverse group of species which 
have been somewhat arbitrarily assigned to the genus because of the 
irregularly splitting calyx. 

Branchlets prominently 2-winged, the wings 0.5-3 mm. high, extend- 
ing from just between the axillary buds at a node to the keeled 
or angled base of the petiole at the node above; mid vein im- 
pressed above; buds glabrous, turbinate, 2 mm. long, nearly 
closed; calyx-lobes in fruit 4, persistent, 1.3 mm. long and wide, 

somewhat separated at base M. bipennis 

Branchlets terete or compressed, not winged; midvein, buds and 

calyx various. 

Midvein flat or convex on the upper surface of the leaf, up to 
1-1.5 mm. broad at base, if sulcate this at base of blade only. 
Inflorescence, including the hypanthium, strongly velutinous or 
at least with numerous loosely ascending reddish or reddish- 
yellow silky hairs up to 1 mm. long. 



FLORA OF PERU 581 

Inflorescence strongly velutinous; flowers large, the buds 
7 mm. long; calyx- tips free; leaves acute or acuminate, 
13-18 cm. long M. velutina 

Inflorescence loosely silky-hairy; flowers small, the buds 2.5 
mm. long, closed, apiculate; leaves caudate-acuminate, 
4.5-7 cm. long M. caudata 

Inflorescence glabrous or essentially so; at least the tips of the 
calyx-lobes evident in the bud. 

Leaves 9-14 cm. long, 2-3.5 times as long as wide; lateral 
veins 12-15 pairs, the transverse veins obscurely retic- 
ulate; leaves finely and obscurely dark dotted; calyx- 
lobes in bud minute, ciliate M. scytophylla 

Leaves 7.5 cm. long or less, 1.7-2.2 times as long as wide; 
lateral veins 6-8, the transverse veins prominently and 
coarsely reticulate; leaves with 1-3 large translucent dots 
per square mm.; calyx-lobes in bud distinct, the inner 
broadly scarious-margined, 2.5 mm. wide. . . . M. areolata 

Midvein sharply and narrowly impressed on the upper surface, or 
in one species broad but concave or broadly sulcate. 

Inflorescence, including buds, with numerous ascending lustrous 
yellowish-white hairs up to 1 mm. long; buds 4-5 mm. long, 
mostly concealed by the hairs; midvein concave or sulcate; 
receptacle tomentose within M. spruceana 

Inflorescence pubescent, often sparsely so, with short, pale or 
reddish hairs 0.5 mm. long or less; buds glabrous or essen- 
tially so, 1.5-3.5 mm. long; midvein sharply and narrowly 
impressed; receptacle glabrous within. 

Leaves cordate-auriculate, nearly sessile, the petioles 3-4 mm. 
long; inflorescence finely hispidulous with minute stiff 
erect hairs; buds closed with a prominent narrow apic- 
ulum; staminal ring short-hairy M. subulata 

Leaves acute to cuneate or somewhat rounded at base, on 
petioles 4-10 mm. long; inflorescence pubescent with ap- 
pressed or ascending hairs; buds closed or the calyx-lobes 
distinct; staminal ring glabrous. 

Petioles transversely rimose, the reddish-brown or whitish 
outer papery layers separating but persistent; calyx in 
bud with 4 very small deltoid separate tips; lower 
branches of the panicle straight and much elon- 



582 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

gated, spikelike with numerous sessile flowers and 

short squarrose bracts M. umbraticola 

Petioles smooth and with unbroken surface, usually dark; 
buds closed or with distinct calyx-lobes; inflorescence 
various. 

Inflorescence thinly pubescent with pale hairs; bracts 
and bracteoles deciduous before an thesis; lateral 
veins of the leaves not impressed above; buds 3- 

3.5 mm. long, closed at apex M. imperfecta 

Inflorescence pubescent with lustrous rufous hairs; bracts 
and bracteoles persistent, squarrose; lateral veins 
impressed above; buds 1.5-2 mm. long with 5 dis- 
tinct calyx-lobes M. squarrosa 

Marlierea areolata McVaugh, Fieldiana Bot. 29: 175. 1956. 

A shrub or tree, glabrous except the vegetative buds thinly stri- 
gose, the inflorescence with a few appressed hairs near base, the 
calyx-segments strigose on both surfaces near tips; leaves broadly 
elliptic, (2-) 3-4.5 cm. wide, (4.5-) 6-7.5 cm. long, mostly 1.7-2.2 
times as long as wide, bluntly acuminate, rounded or acute at base, 
the margins decurrent on the petiole 1 mm. thick, 6-7 mm. long; 
midvein plane or convex above (the whole somewhat impressed), 
prominent beneath; lateral veins 6-8 pairs, somewhat elevated 
above, prominent beneath, the small transverse veins prominently 
reticulate and forming irregularly angular areoles on the order of 
1 mm. across; marginal vein scarcely distinct, 2-4 mm. from margin, 
formed of the looped and somewhat diminished tips of the laterals, 
with a second, smaller and irregular submarginal vein, and numer- 
ous areoles beyond it; blades lustrous on both sides, darker and 
smoother above, the veins more prominent beneath; leaves with 
large translucent glandular dots, these 1-3 per square mm., con- 
spicuous on both surfaces or obscure in mature leaves; inflorescence 
a many-flowered broad panicle, 5-7 cm. long and wide, probably 
always axillary but often appearing terminal, 3 times compound, 
usually branching 5 mm. from base or less, the axis terete or com- 
pressed, 1.5-3 mm. thick; flowers mostly solitary or in 3's on very 
short lateral branchlets from the secondary branches; bracts and 
bracteoles deciduous before anthesis; buds 3 mm. long and almost 
as wide, gland-dotted, darkening in drying, the body subglobose, 
the hypanthium narrowed at base into an ill-defined pseudostalk 
0.5 mm. long; calyx-lobes 4, in bud distinct, strongly imbricated, 



FLORA OF PERU 583 

the outer pair rounded or bluntly triangular, 1.5 mm. long, 2 mm. 
wide, somewhat inclosing the much larger inner pair, which are 
broadly scarious-margined and irregularly erose, truncate at apex, 
about 1.7 mm. long, 2.5 mm. wide; calyx at anthesis explanate, the 
disk becoming shallowly bowl-shaped, 2.5 mm. wide, the calyx split- 
ting slightly between the lobes or the inner lobes tearing free at base; 
style about 4 mm. long; stamens 75-100, about as long as the style, 
the anthers about 0.4 mm. long; petals 4, unequal, the larger broad 
and short, 2.5 mm. wide. A distinctive species of uncertain generic 
position, suggesting its affinity to Myrcia (Aulomyrcia) in the large 
and distinct calyx-lobes, and the deciduous bracts and bracteoles. 
It is better referred to Marlierea because of the tendency of the re- 
ceptacular disk to flatten after anthesis, and the accompanying dis- 
tortion and splitting of the calyx, which even though slight is unlike 
any species of Aulomyrcia known to me. As a minor character may 
be mentioned the terminal or falsely terminal inflorescence in this 
species; this character recurs often throughout the genera Calyp- 
tranthes and Marlierea, whereas in Myrcia the panicles are more often 
from the lower axils. F. M. Neg. 23474. 

Loreto: Stromgebiet des Ucayali von 10 S. bis zur Miindung, 
G. Tessmann 3264, anno 1923 (G, type). 

Marlierea bipennis (Berg) McVaugh, Mem. N. Y. Bot. Card. 
10: 79. 1958. Myrciaria bipennis Berg, Linnaea 31: 259. 71862. 
Myrcia bipennis (Berg) McVaugh, Fieldiana Bot. 29: 189. 1956. 

A shrub or small tree to at least 2.5-3 meters high, with promi- 
nently 2-winged branchlets, the wings 0.5-3 mm. high, running from 
just above each axil and between the leaf-bases to the base of the 
petiole of the next node; plants nearly glabrous, closely strigose on 
the vegetative buds (densely) and the inflorescence and youngest 
shoots (thinly) with appressed lustrous rufous, narrowly fusiform, 
sessile and partly dibrachiate hairs up to 0.5 mm. long; scattered 
pale appressed fusiform hairs persistent on the lower leaf-surface; 
leaves elliptic to lanceolate or ovate, 3-5 cm. wide, 7-15 cm. long, 
(2-) 2.5-3.5 times as long as wide, narrowed from the middle or 
below to the shortly and often obscurely acuminate or merely acute 
tip, the base rounded or acute, the margins cuneately decurrent on 
the petiole 1-1.5 mm. thick, 3-4 mm. long; midvein impressed above, 
prominent beneath, keeled near base and the keel passing gradually 
into the wing of the branchlet; lateral veins very slender, close and 
parallel, about 20-25 pairs, somewhat larger than the nearly equal 
intermediate veins, all obscure and seen as fine lines on both surfaces; 



584 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

marginal vein about equaling the laterals, nearly straight, 1 (-2) mm. 
from margin; blades dull olive green, very smooth but impressed - 
puncticulate above, markedly paler and finely dark-dotted beneath; 
inflorescence axillary, very short and slender, the axis less than 1 mm. 
long, apparently abortive, the flowering branches paired (from the 
lowest nodes) or with an additional pair from the next and approxi- 
mate node, each branch up to 3 cm. long; rachis half as long as the 
branch or more, 1 mm. thick or less, gland-dotted, terete or nearly so; 
flowers 5-9, solitary or in 3's at the tips of the short branches up to 
5 mm. long; bracteoles linear, 0.6 mm. long, deciduous before anthe- 
sis; buds (immature) narrowly obconic, 2 mm. long; flowers not seen; 
hypanthium (in fruit) 2.5 mm. across, glabrous within, with well- 
marked rim and depressed center with straight sides; stamens prob- 
ably 75-100; calyx-lobes (in fruit) rounded, scarcely separated at 
base, about 1.3 mm. long and wide; fruit globose, 8-10 mm. in diam- 
eter, dark purple or black, very finely verruculose. U. of Mich. 
Neg. 482. 

Peru (probably). Amazonian Brazil (Rio Negro) and Venezuela 
(Rio Casiquiare). This is the type-species of Sect. Myrciopsis 
McVaugh, a well-marked subgeneric taxon which is here referred 
to Marlierea rather than to Myrcia because of the nearly closed 
buds which split longitudinally between the calyx-lobes at an- 
thesis; the presence of dibrachiate hairs; the tendency of the in- 
florescence to abort at the lowest node with the production of 
paired lateral panicles. The fruiting calyx is strongly suggestive of 
species of Myrcia. The species is readily recognized by the charac- 
teristic winged branchlets. 

Marlierea caudata McVaugh, Fieldiana Bot. 29: 176. 1956. 

A tree to 7 meters high, the compressed branchlets, vegetative 
buds, and inflorescence shaggy with numerous loosely ascending red- 
dish or reddish-yellow silky hairs up to 1 mm. long on the buds and 
hypanthium; hairs on branchlets often intermixed with many short 
erect reddish hairs; mature leaves with a few hairs beneath, ovate or 
elliptic, 1.7-2.5 cm. wide, 4.5-7 cm. long, 2-3 times as long (includ- 
ing the acumen) as wide, about equally narrowed to both ends, the 
apex then prolonged into a prominent narrow acumen 3 mm. wide 
at base, 1-2 cm. long, the base acute to rounded, the margins de- 
current on the stout petiole 1 mm. thick, 2-4 mm. long; mid vein 
smooth and convex above, prominent but nearly flat beneath; lat- 
eral veins 12-15 pairs with numerous intermediate ones, all very 
slender, obscure on both surfaces; marginal vein about equaling the 



FLORA OF PERU 585 

laterals, less than 1 mm. from margin; both surfaces dull, obscurely 
and sparingly gland-dotted, the upper surface darker, and somewhat 
impressed-puncticulate at least when young; inflorescence axillary 
or terminal, the axis either abortive and 1-2 mm. long or elongate 
and leafy, the 1-4 flowering branches 3-5 cm. long, the slender axis 
somewhat compressed, about 1 mm. wide below the first node; 
flowers few, sessile, the flowering nodes 1 or 2 below the terminal 
triad, the flowers 1 or 3 on each short lateral branch; bracteoles 
linear, glabrous on the backs, 1.5 mm. long, appressed to the bud, 
deciduous after anthesis; buds about 2.5 mm. long, prominently and 
abruptly apiculate, obconic with long hirsute narrow base, glabrous 
on the hemispheric distal half, opening to the level of the stamens 
or a little below in 4 irregularly oblong lobes; hypanthium deeply 
cup-shaped, glabrous within, sparingly hairy among the bases of the 
stamens; style glabrous, 4.5 mm. long; stamens about 100, white, 
the longest 4-5 mm. long, the anthers 0.3 mm. long; petals small, 
obovate, white, about 0.7 mm. wide, 1.3 mm. long; fruit subglobose, 
8-11 mm. in diameter; seeds reniform, 7-9 mm. long. 

Loreto: Mishuyacu, near Iquitos, alt. 100 meters, forest, G. Klug 
235, Oct.-Nov., 1929 (F, type); forest between [lower] Rio Nanay 
and Rio Napo, June 6, 1929, L. Williams 718. 

Marlierea imperfecta McVaugh, Fieldiana Bot. 29: 176. 1956. 

Tree 6 meters high, the inflorescence thinly pubescent with pale 
or reddish-based appressed or ascending hairs 0.2 mm. long, or longer 
on the vegetative buds; leaves elliptic, 4.5-7 cm. wide and 12 cm. 
long, or up to 9.5 cm. wide, 25 cm. long, about 2.5 times as long as 
wide, acuminate (often narrowly so), rounded to acute at base, the 
margins decurrent on the dark channeled petiole 1.5-3 mm. thick, 
5-8 mm. long; mid vein impressed above, prominent beneath; lateral 
veins 10-15 pairs and some intermediate ones, somewhat raised on 
both sides, more prominent (as also the reticulum formed by the 
small transverse veins) beneath; marginal vein about equaling the 
laterals and arched between them, 2-5 mm. from margin, with a 
faint submarginal vein beyond it; blade dark, smooth and obscurely 
impressed-puncticulate above, the lower surface dull, coppery brown, 
finely dark-dotted; inflorescence falsely terminal, the several branches 
apparently arising from the lowest nodes of an abortive terminal 
axis, each branch forming a broad panicle 8-14 cm. long and almost 
as broad, 3 times compound, the peduncle 2.5-4 cm. long, flattened, 
1.5-2 mm. wide below the first node; branches about 5 pairs, mostly 
opposite; flowers sessile or nearly so, solitary or in 3's near the tips 



586 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

of the branches; bracts not seen, apparently deciduous before an- 
thesis; buds 3-3.5 mm. long, glabrous except a small apical tuft, 
obovate, closed and rounded or scarcely apiculate at tip, the base 
rather broadly obconic; calyx rupturing at anthesis and splitting 
into 4 irregular lobes about 1.5 mm. long and up to 2.5 mm. wide; 
disk glabrous, concave, about 3 mm. across; style 4-4.5 mm. long; 
stamens about 100, about equaling the style; petals 3-4, suborbic- 
ular, ciliate, often very broad at base but the attachment narrow, 
about 2.5 mm. long and broad. 

Loreto: Mishuyacu, near Iquitos, alt. 100 meters, forest, Jan. 
1930, G. King 787 (US, type). 

Marlierea scytophylla Diels, Verh. Bot. Ver. Brandenb. 48: 
187. 1907. 

A shrub or tree up to 8 meters high, glabrous except that the 
vegetative buds are finely strigose, the divisions of the calyx are cili- 
ate and strigose within, and a few hairs sometimes persist about the 
base of the inflorescence; leaves elliptic, 3-5.5 cm. wide, 9-14 cm. 
long, 2-3.5 times as long as wide, the tips acuminate, the base cune- 
ate, acute, or somewhat rounded above the very base where the 
margins are cuneately decurrent on the stout petiole 1.5 mm. thick, 
5-10 mm. long; midvein convex above, 1-1.5 mm. wide at base, 
prominent beneath; lateral veins 12-15 pairs with some interme- 
diate ones nearly the same size, inconspicuous above, slightly ele- 
vated beneath; marginal vein about equaling the laterals and arched 
between them, 1.5-2.5 mm. from the margin; blades smooth and 
rather dull green above, with numerous but often obscure glandular 
dots, the lower surface yellow-green, finely and sparingly dark-dot- 
ted; inflorescence a many-flowered axillary or falsely terminal pan- 
icle 5-8 cm. long, mostly twice-compound, the central axis longer 
than the lateral branches, or abortive and the lateral panicles paired; 
flowers mostly sessile along the spikelike branches, the longest of 
which are 3 cm. long and about 10-flowered; bracts broad-based, 
acute, 1-2.5 mm. long, at least in part persistent through anthesis 
and evident along the spikes as abortive flowers fall; buds 2-2.5 mm. 
long, obconic, open at the apex, the lobes small and ciliate; calyx 
opening by four irregular splits; disk glabrous, about 2.3 mm. wide; 
style 5-5.5 mm. long; stamens probably 75-100, 5 mm. long (Diels) ; 
fruit globose to oblate, finely glandular- verruculose, 1-1.5 cm. in 
diameter. A little-known species of the Amazon Basin, readily dis- 
tinguished even in fruit from similar species of Calyptranthes by the 
convex rather than impressed midvein. 



FLORA OF PERU 587 

An imperfect specimen with immature fruit, which was col- 
lected on the Rio Nanay, Loreto (Williams 767), appears to be a 
Marlierea and may be the present species, but the leaves tend toward 
ovate rather than elliptic and the inflorescence is somewhat strigose. 
F. M. Neg. 23411. According to Diels' original description of M. 
scytophylla the leaves are deeply sulcate above, but, as plainly shown 
in the photograph, the midveins are convex rather than impressed 
in Ule's no. 6044, the type specimen. 

Peru (probably). Amazonian Brazil and Venezuela. 

Marlierea spruceana Berg, in Mart. Fl. Bras. 14, pt. 1: 34. 
1857. M. spruceana a latifolia Berg, I.e. 515. 1859. M. spruceana 
/3 angustifolia Berg, I.e. 

A tree up to 9 meters high, strigose on the branchlets, inflores- 
cence and the lower surfaces of at least the young leaves with lus- 
trous yellowish-white hairs, these up to 1 mm. long (on the buds) or 
often shorter, those on the leaves short, often with a few dibrachiate 
hairs, up to 0.5 mm. long, interspersed; leaves ovate-elliptic, often 
broadest a little below the middle, 6-7 cm. wide, 15-18 cm. long (or 
those subtending the inflorescence smaller, 2.5-4 cm. wide, 8-13 cm. 
long), 2-3 times as long as wide, gradually narrowed to an acuminate 
tip, rounded or subcuneate at base, the margins decurrent on the 
very stout petiole 1.5-2 mm. thick, 6-12 mm. long; midvein on the 
upper surface concave, sulcate at least near the base, prominent 
beneath; lateral veins 15-20 pairs, slender, slightly raised on both 
surfaces; marginal veins about equaling the laterals, slightly arched 
between them, 1.5-3 mm. from margin and often with a weak sec- 
ondary marginal vein; leaves sometimes blackening above in drying, 
both surfaces dull, the glands obscure, somewhat apparent above; 
inflorescence terminal, forming a broad panicle with up to 200 flow- 
ers, 10-12 cm. long, 4 to 5 times compound, or the central axis 
abortive and the lower branches 5-8 cm. long; principal branches 
usually with 3-4 nodes, the peduncle 2-4 cm. long below the lowest 
node, compressed and 2-2.5 mm. wide below that node; flowers ses- 
sile, mostly in terminal clusters of 3; bracts and bracteoles ovate, 
acute, 1-3 mm. long, deciduous before anthesis; buds closed, obo- 
void, 4-5 mm. long, with some dark glands partly concealed among 
the hairs, opening irregularly into about 4 lobes which are glabrous 
on the inner surface; style glabrous, 7 mm. long; stamens 100-125, 
about 5 mm. long, the anthers 0.3 mm. long; petals 4, ovate, trun- 
cate at base, ciliate, 3 mm. wide, 4 mm. long; disk 5 mm. across in 
anthesis, the hypanthium deeply concave, tomentose within; fruit 



588 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

globose, 1-1.5 cm. in diameter, tomentose. Probably conspecific is 
M. uaupensis Berg, in Mart. Fl. Bras. 14, pt. 1: 516. 1859, of Co- 
lombia, which has the leaves broader (the terminal 6 cm. wide, 12.5 
cm. long), short-petiolate and subcordate, the buds broadly ovoid, 
and the pubescence of longer and whiter hairs. F. M. Neg. 19879. 
Loreto: Mishuyacu, Klug H.11. Amazonian Brazil and Colombia. 

Marlierea squarrosa McVaugh, Fieldiana Bot. 29: 177. 1956. 

A shrub 2 meters high, densely pubescent on vegetative buds and 
sparingly on young branchlets, petioles, and inflorescence, with ap- 
pressed or ascending flexuous lustrous rufous hairs up to 0.5 mm. 
long; leaves broadly elliptic or somewhat ovate, 3.5-6 cm. wide, 9- 
13 cm. long, about 2.5 times as long as wide, rather abruptly nar- 
rowed to a slenderly acuminate tip, rounded at base, the margins 
decurrent on the petiole 1 mm. thick, 5-7 mm. long; mid vein and 
10-15 pairs of lateral veins impressed above, prominent and raised 
beneath; marginal vein about equaling the laterals and arched be- 
tween them, not impressed above, 2-3 mm. from margin; small veins 
not elevated nor prominently reticulate above; leaves drying brown, 
dull, the upper surface darker and nearly eglandular, the lower cop- 
pery and very sparingly dotted; inflorescence a broad many-flowered 
axillary panicle 7-10 cm. long, 3 to 4 times compound, with up to 
10 pairs of branches, the lowest up to 4 cm. long, the peduncle 2 cm. 
long, compressed and up to 1.5 mm. wide below the lowest node; 
flowers subspicate, mostly toward the tips of the branches, often sub- 
opposite, sessile, solitary or in short-peduncled clusters of 3; bracts 
ovate, divaricate, hairy, acute, 1-1.5 mm. long, broad-based and 
rounded on the backs, persistent through an thesis; buds glabrous, 
drying black, broadly obovoid, 1.5-2 mm. long, with numerous 
raised glandular dots, contracted at base to a very short pseudo- 
stalk; calyx-lobes 5, rounded, red-ciliate and appressed-pubescent 
within, strongly unequal, the two outer much smaller, about 0.7 mm. 
long and wide, the three inner up to 2 mm. wide and 1 mm. long, 
the calyx as a whole splitting irregularly from the bases of the lobes 
to the summit of the ovary; disk 1.7-2 mm. wide, glabrous, flat after 
anthesis; style 3.5 mm. long; stamens about 50, 3 mm. long; anthers 
0.6 mm. long; petals 3, suborbicular, about 2.5 mm. long; fruit not 
seen. This species, like M. areolata, seems to cross the supposed 
generic lines between Marlierea and Myrcia (Aulomyrcia) . It has 
the irregularly splitting calyx, persistent bracts and explanate disk 
of Marlierea, but the coppery color and free calyx-lobes of some 
species of "Aulomyrcia." It might conceivably be a hybrid involv- 



FLORA OF PERU 589 

ing Marlierea umbraticola, which also has impressed veins in the 
leaves. 

Loreto: Mishuyacu, Klug 169 (US, type). 

Marlierea subulata McVaugh, Fieldiana Bot. 29: 177. 1956. 

Tree 5 meters high, the branchlets, inflorescence and petioles 
finely hispidulous with yellowish erect hairs about 0.2 mm. long, or 
those on the vegetative buds, base of inflorescence and base of hy- 
panthium slightly longer and appressed; leaves ovate or lanceolate, 
nearly sessile, 4.5-7 cm. wide, 13-18 cm. long, 2.5-3.5 times as long 
as wide, narrowed toward the apex from the middle or below, and 
gradually acuminate, rounded and cordate-auriculate at base, the 
margins passing abruptly into the ventral angles of the nearly flat, 
hispidulous and somewhat expanded summit of the petiole 1.5-2 mm. 
thick, 3-4 mm. long; mid vein impressed above, prominent beneath; 
lateral veins about 10 pairs, impressed above, prominent and scalari- 
form beneath, the intermediate ones weak; marginal vein about 
equaling the laterals and markedly arched between them, 2.5-7 mm. 
from margin, with an indistinct submarginal vein beyond it; blades 
smooth, dark and lustrous above, with no glands apparent, the lower 
surface paler, dull, with occasional dark glands; inflorescence a rather 
few-flowered narrow panicle, 2 to 3 times compound, 7-15 cm. long, 
and up to 6 cm. wide at base; peduncle less than half as long as the 
whole panicle, distally flattened, 2-2.5 mm. wide; flowering nodes 
5-6, the branches opposite or subopposite, the flowers solitary or in 
3's on pedicels up to 5 mm. long; bracts divaricate, glabrous, subu- 
late, up to about 3 mm. long, more or less persistent through an thesis; 
bracteoles similar, about 1 mm. long, somewhat appressed to the 
pseudostalk 0.5-1 mm. long; buds obovate, about 3 mm. long in- 
cluding the prominent narrow apiculum, glabrous except near base 
and sometimes for a tuft of small hairs crowning the apiculum, the 
hypanthium gradually narrowed toward the base and then abruptly 
rounded to the pseudostalk; calyx in bud completely closed, in an- 
thesis splitting irregularly into 4 lobes, these up to 2 mm. long and 
wide, glabrous except at apex; disk cup-shaped, 2.5 mm. across, the 
staminal ring short-hairy; style 6 mm. long or more; stamens 75-100, 
about equaling the style, the anthers about 0.3 mm. long; petals 4, 
obovate or subrotund, 1 mm. wide, 1-1.5 mm. long, silky-villous 
without; fruit globose to oblate, up to 1 cm. long and 1.5 cm. across. 

Loreto: Mishuyacu, near Iquitos, alt. 100 meters, forest, May, 
June, 1930, G. Klug 1341 (F, type). 



590 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Marlierea umbra ticola (HBK.) Berg, Linnaea 27: 17. 1855. 
Myrtus umbraticola HBK. Nov. Gen. & Sp. 7: 258 (folio ed. p. 199). 
1825. Marlierea insculpta Diels, Verb. Bot. Ver. Brandenb. 48: 188. 
1907. 

A shrub or small tree 5-8 meters high, the inflorescence thickly 
and finely pubescent with appressed or ascending lustrous rufous or 
silver-tipped hairs up to 0.3 mm. long; lower leaf-surface with a few 
appressed pale hairs with darker bases; leaves lanceolate or elliptic, 
2.5-5.5 cm. wide, 8.5-17 cm. long, (2.8-) 3-4.5 times as long as wide, 
the tip gradually or abruptly and often conspicuously acuminate, 
the base acute to cuneate or somewhat rounded, the margins decur- 
rent on the stout, transversely rimose petiole 1.5-2 mm. thick, 4-10 
mm. long; petiole at maturity with the outer layers smooth and 
reddish brown, cracking transversely and at length longitudinally, 
separating and persisting; midvein impressed above, elevated nearly 
its own thickness beneath; lateral veins 20-25 pairs, never promi- 
nent but the major ones slightly impressed as delicate lines above, 
slightly raised beneath; marginal vein about equaling the laterals or 
slightly weaker, arched between them, 1-2.5 mm. from margin; 
blades dark and lustrous but without apparent glands above, dull 
green or brown, and finely dark-dotted beneath; inflorescence a 
many-flowered axillary panicle 7-10 cm. long, mostly twice com- 
pound with the flowers sessile or nearly so along the slender divari- 
cate branches, the lowest of which may be as long as the central 
branch or even longer; peduncle 1-2 cm. long, terete or slightly com- 
pressed, usually rimose like the petioles, 1-2.5 mm. thick; branches 
straight, slender, up to 10 cm. long and with more than 40 flowers 
in bracteate, opposite or sometimes alternate pairs; bracts persist- 
ent, squarrose, ovate, broad-based and somewhat surrounding the 
buds, acute, 1 mm. long or less; buds 2-2.5 mm. long, dark or black 
in drying, glabrous except the ciliate tips, broadly obovoid, the hy- 
panthium obconic, the tip rounded; calyx-lobes 4, approximate in 
the bud, very small, with deltoid ciliate tips, the calyx splitting 
irregularly at anthesis between the lobes into irregular glabrous divi- 
sions about 2.5 mm. long and 2 mm. wide; disk glabrous; style 4-5 
mm. long; stamens 60-75, about 4 mm. long, the anthers 0.3 mm. 
long; fruit globose, smooth or minutely verruculose, 5-10 mm. in 
diameter. A commonly collected species of the basins of the Orinoco 
and Casiquiare rivers in Colombia and Venezuela, and known also 
from various stations in Amazonian Brazil; the type of M. insculpta 
came from Marary on the upper Rio Jurud, Ule 5080, 5081. F. M. 
Negs. 23407, 36908. 



FLORA OF PERU 591 

Peru (probably). Amazonian Bolivia and Brazil to Venezuela, 
and eastern Colombia. 

Marlierea velutina McVaugh, Fieldiana Bot. 29: 178. 1956. 

A shrub or tree, densely velutinous on the branchlets, inflores- 
cence, buds, and petioles with coarse sharp erect yellowish brown 
hairs up to 1 mm. long; leaves on both surfaces thinly hirsutulous 
with similar hairs; leaves elliptic, oblong, or lanceolate, 4-6 cm. wide, 
13-18 cm. long, about 3 times as long as wide, acute or shortly and 
slenderly acuminate at tip, narrowly rounded at base, the margins 
decurrent on the stout petiole 1.5-2 mm. thick, 5-7 mm. long; mid- 
vein flat above or sulcate near base, velutinous, prominent and ele- 
vated its own diameter beneath; lateral veins about 20 pairs, slightly 
elevated above, prominulous beneath; marginal vein about equaling 
the laterals, somewhat arched between them, 1.5-3 mm. from mar- 
gin; blades at maturity dull, dark and obscurely punctate above, 
yellow green or brown and hairy beneath, without apparent glands; 
inflorescence of many-flowered, 2 or 3 times compound, axillary or 
falsely terminal panicles 6-10 cm. long, the flowers solitary or in 3's 
near the tips of the branches; peduncle 3.5-5 cm. long, compressed, 
2.5 mm. wide below the first branches; primary branches 1.5-2.5 cm. 
long, the lateral pedicels 3-5 mm. long, the central flowers sessile or 
nearly so; bracts and bracteoles deciduous before an thesis, linear, 
2-3 mm. long; buds probably about 7 mm. long, not seen; hypan- 
thium 3-4 mm. long, broadly ellipsoid or urceolate to subglobose, 
with about 8 prominent longitudinal ridges; calyx-lobes in bud united, 
with free, bluntly deltoid tips about 1 mm. long, 1.5 mm. wide, the 
calyx in anthesis splitting irregularly into 4 somewhat elliptic lobes 
2-3 mm. wide, 3-4 mm. long, the split extending into the edge of the 
staminal disk and the margin of the disk recurved with the lobes; 
disk about 4 mm. wide, deeply depressed at center, velutinous; style 
7 mm. long, hairy two-thirds of its length; stamens very numerous, 
probably about 200; petals obovate-cuneate, 2 mm. wide, 3 mm. 
long; ovary bilocular, with 2 ovules in each locule. Known only 
from the type, Rusby 2683, collected at the falls of the Rio Madeira, 
Brazil. 

Peru (probably). Amazonian Brazil. 

2. CALYPTRANTHES Swartz 

Reference: Berg, Linnaea 27: 18-33. 1855; and in Mart. Fl. Bras. 
14, pt. 1:38-55. 1857. 



592 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Calyx completely closed in the bud, circumscissile, the operculum 
usually attached at one side in anthesis, finally completely dehiscent. 
Petals none, or in a few species 2-3 (-5), small and inconspicuous. 
Inflorescence-axis usually abortive above the first node, with the 
paired panicles arising from opposite axils at the lowest nodes of the 
axis. Pubescence usually at least in part of dibrachiate hairs. 

A distinctive genus of perhaps 50-75 species, ranging from Florida 
through the West Indies and eastern South America to Uruguay. 
Early reports of this genus from Asia were based primarily on species 
now referred to Syzygium Gaertn., a genus of the Eugeniinae. 

Flowers very large for the genus, the buds 7-8 mm. long, convex or 
nearly flat at the apex, lacking a narrow apiculum; inflorescence 
pale-scurfy and also appressed-puberulent with minute brown- 
ish dibrachiate hairs 0.1 mm. long; leaves 25-39 cm. long, nar- 
rowly elliptic, nearly sessile, appearing cordate-auriculate at 
base, the stout petiole 4 mm. long C. maxima 

Flowers smaller, the buds if 5 mm. long or more fusiform and apicu- 
late, or noticeably hirsute or velutinous; inflorescence glabrous 
to hairy; leaves usually smaller and slender-petiolate, if sessile 
or essentially so the buds not as above. 

Leaves sessile, with veins impressed on the upper surface; blades 
cordate at base or the margins much produced and plicate. 

Leaves ovate, cordate, 9-12 cm. long; inflorescence glabrous; 
buds fusiform, 6-7 mm. long C. sessilis 

Leaves obovate, with the margins near base produced into 
puckered folds, the blades 30-38 cm. long; inflorescence 
appressed-hirsutulous; buds 3.5 mm. long, obovoid. 

C. plicata 

Leaves petiolate, cuneate to acute or somewhat rounded at base, 
the veins usually not impressed. 

Inflorescence of paired spikes, the individual flowers sessile along 
the axis or the lowest in sessile or very short-peduncled 
groups of three. 

Buds glabrous; leaves 2.5-6 cm. long, often obovate with 
rounded or sometimes short-acuminate tip; flowers 
mostly 3 or 5 (-11) in each spike C. pulchella 

Buds strigose or hirsute; leaves 7-16 cm. long, elliptic to 
ovate, usually prominently and often narrowly acumi- 
nate; flowers more numerous. 



FLORA OF PERU 593 

Branchlets and inflorescence, including the buds, thickly 
rufous-hirsute; buds broadly obovoid to nearly glo- 
bose, 5-6 mm. long, concealed by the hairs; flowers 
8-13 in each spike C. krugioides 

Branchlets and inflorescence with appressed yellow or 
brown dibrachiate hairs; buds obovoid or broadly 
fusiform, 2-2.5 mm. long, rather sparingly hairy. 

Spikes mostly 8-12 cm. long, the numerous flowers in 
several sessile clusters of 10-20 flowers each; leaves 
short-acuminate, the lateral veins not impressed 
above, slender and closely parallel C. densiflora 

Spikes 3-5.5 cm. long, the flowers 25 or fewer on each, 
in small sessile clusters of 1-3 each; leaves with nar- 
row acumen 1.5-2 cm. long; lateral veins impressed 
above, the principal ones prominent beneath and 
contrasting with the less conspicuous intermediates. 

C. brevispicata 

Inflorescence of paired panicles or compound dichasia, the basal 
branches elongated and again branched, or occasionally 
with solitary terminal flowers. 

Branches of the inflorescence uniformly but sometimes thinly 
beset with appressed or erect hairs; hypanthium vari- 
ously strigose to tomentose, except in C. multiflora. 

Hypanthium glabrous; inflorescence loosely pubescent with 
numerous erect or somewhat appressed soft pale rufous 
hairs; midvein convex above; buds 2-2.5 mm. long. 

C. multiflora 

Hypanthium strigose or variously velutinous or tomentose, 
if nearly glabrous the midvein sulcate or narrowly im- 
pressed above. 

Inflorescence-branch an umbelliform cyme 2-5 cm. long 
with 15 flowers or fewer, the ovate boat-shaped 
bracts subfoliaceous, persistent; inflorescence, in- 
cluding the flowers, thickly hirsute with coarse rufous 
sessile dibrachiate hairs up to 1.5-2 mm. long and 

attached near one end C. longifolia 

Inflorescence-branch paniculate or by reduction race- 
mose, often 3 to 4 times compound and many- 
flowered (if short and few-flowered not hirsute as 
above); bracts completely deciduous before an the- 



594 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

sis or occasionally a few (especially the basal ones) 
persisting; hairs of the inflorescence various, mostly 
sessile and less than 1 mm. long. 

Hairs of the inflorescence golden-yellow, dibrachiate, 
up to more than 1 mm. long, the basal stalk of the 
hair erect and often as long as the spreading or 
ascending branches; leaves 6 cm. long or less, 
rounded to obscurely acuminate at tip; branch- 
lets 2-winged; flowers mostly 10 or fewer on each 

branch C. tridymantha 

Hairs of the inflorescence sessile and somewhat ap- 
pressed and mostly less than 0.5 mm. long, or with 
very short hairs intermixed, or the inflorescence 
velutinous or tomentose, usually with red or rusty 
hairs. 

Leaves large, mostly more than 15 cm. long (often 
20-30 cm. long), with 20-35 pairs of lateral 
veins; buds obovoid or obconic, scarcely apic- 
ulate, 3-5 mm. long; inflorescence with abun- 
dant rufous pubescence. 

Lateral and marginal veins scarcely apparent on 
lower leaf-surface, the surface covered with 
very numerous, closely appressed pale hairs 
up to 0.2 mm. long; blades tapering from the 
middle or below to a slender apex; inflores- 
cence 5 cm. long or less, few-flowered. 

C. macrophylla 

Lateral and marginal veins forming a conspicuous 
gridiron pattern on the lower leaf-surface, 
which is glabrous or sparingly appressed- 
pubescent at maturity; blades abruptly and 
narrowly acuminate; inflorescence 6-10 cm. 

long, many-flowered C. speciosa 

Leaves of moderate size, usually less than 20 cm. 
long or, if longer, the lateral veins 15 pairs or 
fewer, or the buds 2-2.5 mm. long; buds and 
pubescence various. 

Flowers small, the buds 2-2.5 mm. long, obovoid, 
the apex rounded or shortly apiculate; pan- 
icles mostly 3 times compound, many-flow- 
ered, the branches sparingly covered with 






FLORA OF PERU 595 

appressed pale or sometimes reddish hairs, 
the hypanthium strigose, sometimes very 
sparingly so; paired panicles from an abor- 
tive flattened axis 10 mm. long or less. 
Lower leaf-surfaces with few dark hairs and 
usually with rather numerous persistent, 
nearly colorless appressed hairs; leaves 
elliptic, broadest at the middle, 5-10 cm. 
long with 12-15 lateral veins on each side; 

style 4-4.5 mm. long C. ruiziana 

Lower leaf-surfaces glabrous except for a few 
dark hairs; leaves ovate or lanceolate, usu- 
ally widest somewhat below the middle, 
9-15 (-25) cm. long, with 20-25 pairs of 
veins on each side; style 5-6 mm. long. 

C. simulata 

Flowers larger, the buds 3-6 mm. long, variously 
shaped; panicle compound, or by reduction 
racemoid, its branches and the hypanthium 
uniformly and usually conspicuously ap- 
pressed-hairy, velutinous or tomentose with 
ferruginous or dark reddish hairs; panicles 
various. 

Inflorescence-branch a spike with all flowers 
sessile, or the lower branches 1-2 cm. long, 
1- or 3-flowered; buds 5-6 mm. long, abun- 
dantly hirsute, the hypanthium hairy 

within C. krugioides 

Inflorescence-branch a panicle, usually many- 
flowered and 3 times compound; buds 3-4 
mm. long, appressed-hairy to tomentose, 
the hypanthium glabrous within. 
Midvein impressed above; hypanthium ap- 
pressed-hairy, the hairs of the inflores- 
cence rusty-brown; buds fusiform, 3-3.5 
mm. long; panicle narrow, the lowest 
branches about 1.5 cm. long. 

C. tessmannii 

Midvein convex above (and then sometimes 
sulcate) or raised in a narrow ridge; hy- 
panthium loosely velutinous or tomen- 



596 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

tose; buds obovoid or ellipsoid; lower 

branches of the panicle relatively long. 

Leaves 15-21 cm. long, about 4 times as 

long as wide, the straight marginal 

vein and the 10-15 short lateral veins 

prominent beneath; petiole very stout, 

3 mm. thick, 7 mm. long; buds broadly 

ellipsoid, heavily tomentose with dark 

red hairs, the hypanthium urceolate 

in anthesis C. rufotomentosa 

Leaves 10-16 cm. long, about 2.5 times as 
long as wide, the marginal vein con- 
sisting of a series of loops or arches 
between the 8-12 pairs of laterals; 
petiole 1 mm. thick, 8-10 mm. long; 
buds obovoid, sparingly or rather 
densely velutinous with rusty-brown 
or reddish hairs, the hypanthium prob- 
ably turbinate in anthesis. 

C. cuspidata 

Branches of the inflorescence (including the hypanthium) 
completely glabrous or with a very few scattered hairs 
about the base and the nodes of the panicle. 

Panicles with 20 flowers or fewer, sometimes reduced and 
raceme-like or spikelike; peduncle and rachis filiform 
or very slender, often terete and nearly straight, usually 
less than 1 mm. thick; branchlets narrowly 2-winged. 

Flowers mostly sessile on the axis, only the lower clusters 
pedunculate; wings of the branchlets often 0.5 mm. 
high; buds 4-7 mm. long, obtuse or obscurely 
apiculate C. pulchella 

Flowers mostly on very long slender pedicels; wings of 
the branchlets scarcely higher than thick; buds 2-3 
mm. long, narrowly and conspicuously apiculate. 

C. bipennis 

Panicles many-flowered, 3 to 4 times compound, the pe- 
duncle usually somewhat angular near summit and 
1-1.5 mm. thick, the rachis often irregularly enlarged 
and zigzag; wings, if produced on branchlets, not per- 
sisting through the growing season. 



FLORA OF PERU 597 

Lower leaf-surface obscurely gland-dotted, the dots about 
15 per square mm.; leaves relatively narrow, mostly 
2.5 times as long as broad, or longer; panicle- 
branches irregularly alternate C. paniculata 

Lower leaf-surface prominently dark-dotted, the dots 
more than 50 per square mm. ; leaves broader, mostly 
2.3 times as long as broad, or less; panicle-branches, 
both large and small, often verticillate or fasciculate. 

C. crebra 

Calyptranthes bipennis Berg, Linnaea 31: 248. 71862. 

A shrub or tree up to 15 meters high, glabrous except for a few 
hairs on the young terminal vegetative buds; young leafy branchlets 
2-winged, the wings up to about 0.2 mm. high, arising just above the 
axil at one node and terminating in stipule-like tips up to 1 mm. 
long between the petiole bases at the node above; leaves elliptic, 
lanceolate or ovate, (1-) 2-5 cm. wide, 3-7 (-11) cm. long, 2-3 (-4) 
times as long as wide, acuminate (often slenderly so), the base cune- 
ate, or rounded and finally subcuneate, the margins decurrent on 
the petiole 2-3 mm. long and up to 1 mm. thick; midvein prominent 
beneath, on the upper surface broadly impressed near base in a 
rounded channel, plane near the tip; lateral veins slender, slightly 
raised on both surfaces or but obscurely so above, about 10-12 prin- 
cipal pairs but with relatively strong intermediate veins, the blade 
thus often seeming to have numerous close parallel veins; marginal 
vein continuous, 0.5-1 (-2) mm. from margin, little arched between 
the laterals; leaves dull and pale green in drying, or the lower sur- 
face pale brown; upper surface smooth, usually with at least a few 
impressed dots; lower surface more or less prominently brown dotted; 
inflorescence axillary, the axis either abortive and 0.5-2 mm. long, 
or elongate and leafy; flowering branches 2, opposite, arising from 
the lowest nodes of the axis, filiform, 2-5 (-9) cm. long, 0.2-0.7 mm. 
thick below the nodes, 1 to 3 times compound; peduncle 9-22 mm. 
long; nodes of the flowering axis 1-3, the pedicels filiform, 2-7 mm. 
long, 1-flowered or with 2 or 3 nearly sessile flowers at the tip; bract 
at base of flowering branch linear, 0.7 mm. wide, 1.7 mm. long; other 
bracts and bracteoles not seen; buds obovoid, prominently gland- 
dotted, attenuate at base at least when young, 2-3 mm. long, nar- 
rowly apiculate, the apiculum often conspicuous; calyptra about one- 
third as long as the unopened bud; style 3-5 mm. long; petals none; 
stamens about 75, up to 4 mm. long; fruit globose, 5-7 mm. in 



598 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

diameter. The several collections which I should refer to this species 
show unusual variation in size of leaves, but are otherwise so similar 
that there is little doubt that they are conspecific. The type speci- 
men, which is in young bud only, bears small and narrow leaves up to 
1 cm. wide and 4 cm. long. The inflorescences are poorly developed, 
each branch bearing three pedicellate flowers only. Superficially this 
specimen bears little resemblance to such large-leaved examples as 
those in Killip & Smith's no. 27702 and Krukoff 's nos. 5203 and 5205, 
but in characters other than leaf-size there is good agreement. The 
development of the inflorescence and the presence of 1-flowered or 
3-flowered pedicels evidently vary from plant to plant, and under 
conditions which affect vigor and development; in some cases both 
1-flowered and 3-flowered pedicels are found on the same plant, and 
I regard as taxonomically unimportant, in this case, the difference 
between the 3-flowered cymes of the type specimen, and the approxi- 
mately 20-flowered panicle-like branches of Killip & Smith's no. 
27702. The type specimen of C. bipennis apparently is merely a 
small-leaved extreme, collected at an early stage in development, of 
a species population which is widespread and relatively uniform in 
Brazilian Amazonia and adjacent Peru. Univ. of Mich. Neg. 430. 

San Martin: Tarapoto, Williams 6584- Loreto: Yurimaguas, 
Killip & Smith 27702. Fl. Huallaga ad cataractas, July, 1856, 
Spruce 4596, type. Amazonian Brazil. 

Calyptranthes brevispicata McVaugh, Fieldiana Bot. 29: 181. 
1956. 

A tree 4 meters high, the branchlets, inflorescence, petioles and 
lower leaf-surfaces sparingly covered with appressed fusiform yellow- 
ish brown dibrachiate hairs up to 0.5 mm. long (or to 1 mm. long 
on the midvein); leaves elliptic-lanceolate, 3.5-5 cm. wide, 11-15 cm. 
long, about 3 times as long as wide, narrowed toward the apex from 
about the middle, gradually or rather abruptly long-acuminate, the 
tip 3 mm. wide at base, 1.5-2 cm. long; base of blade acute and 
finally cuneate, the margins decurrent on the stout petiole 1.5 mm. 
thick, 8 mm. long; midvein sharply sulcate and impressed above, 
prominently elevated beneath; lateral veins 20 or more pairs, im- 
pressed above, prominent beneath and contrasting with the smaller 
and less conspicuous intermediate veins; marginal vein nearly 
straight, about equaling the laterals, 2-3 mm. from margin, slightly 
impressed above and prominently elevated beneath, a smaller but 
distinct submarginal vein beyond it; blades somewhat lustrous, green 
and impressed-puncticulate above, somewhat yellowish beneath, 



FLORA OF PERU 599 

with small dark dots and numerous appressed hairs; inflorescence 
axillary, the axis very short and flat, about 2 mm. long and wide, 
the flowering branches 2, spicate, 3-5.5 cm. long, from the lowest 
nodes of the axis, the peduncle short, about 1 cm. long, nearly terete, 
1-1.3 mm. thick below the first node; flowers 25 or fewer on each 
branch, 1-3 in small sessile clusters at the 3-4 principal nodes and 
the tip; bracts deciduous before an thesis; buds about 2.5 mm. long, 
broadly fusiform, brown-hairy and somewhat arachnoid-pubescent, 
broadly and sometimes obscurely apiculate; calyptra domelike; hy- 
panthium after dehiscence broadly campanulate, 1.5 mm. high, about 
2 mm. across the rim; style 6 mm. long; stamens about 50, white, to 
6 mm. long, the anthers 0.3 mm. long. Univ. of Mich. Neg. 454. 

Loreto: Florida, Rio Putumayo, at mouth of Rio Zubineta, alt. 
200 meters, forest, March-April, 1931, G. King 2040 (US, type). 

Calyptranthes crebra McVaugh, Fieldiana Bot. 29: 181. 1956. 

Tree 4-10 meters high, completely glabrous or the vegetative 
buds strigose with reddish fusiform dibrachiate hairs up to 0.5 mm. 
long, and a few similar hairs persistent at the base and about the 
nodes of the inflorescence; leaves elliptic to ovate, 4-5 cm. wide, 
8.5-10.5 cm. long, about 1.8-2.3 times as long as wide, short-acumi- 
nate, the base cuneate, the margins decurrent on the stout petiole 
1-1.5 mm. thick, 4-5 mm. long; midvein broadly sulcate above, 
prominent beneath; lateral veins 12-15 pairs in addition to the 
scarcely less conspicuous intermediates, slightly elevated on both 
surfaces but obscure on both; marginal vein about equaling the lat- 
erals, nearly straight, 1.5-2 mm. from margin; upper surface olive 
green to brown in drying, impressed-puncticulate, the lower pale or 
dark tan, with abundant (more than 50 per square mm.) dark prom- 
inent glands; inflorescence axillary or falsely terminal, the axis about 
1 mm. long, the 2 flowering branches from the lowest nodes, 5-11 cm. 
long, mostly 3 times compound, many-flowered, the peduncle often 
half as long as the entire inflorescence, terete, 1 mm. thick or a little 
more; lowest branches of the panicle sub-verticillate, three larger 
and as many smaller arising nearly together; upper nodes mostly 
alternate and enlarged, the branchlets often fasciculate, the rachis 
zigzag and flattened; flowers yellow-green (Klug), near tips in 3's or 
in clusters up to 10; bracts deciduous before anthesis; buds 2 mm. 
long or less, obovate, somewhat expanded above the middle, shortly 
apiculate; hypanthium after dehiscence broadly campanulate, about 
1.3 mm. long and 1.5 mm. wide; calyptra concave; style 5 mm. long; 
stamens 60-75, about as long as the style; anthers about 0.2 mm. 



600 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

long; fruit globose, about 5 mm. in diameter, finely glandular- ver- 
ruculose, surmounted by the very short flaring neck of the hypan- 
thium. Univ. of Mich. Neg. 470. 

Loreto: Mishuyacu, near Iquitos, forest, alt. 100 meters, Octo- 
ber-November, 1929, G. Klug 77, 78 (F, type), 607. 

Calyptranthes cuspidata DC. Prodr. 3: 258. 1828. 

Probably a tree, the vegetative buds, petioles, branchlets and 
inflorescence velutinous with rusty brown or light reddish brown 
partly dibrachiate hairs up to 0.5 mm. long; leaves glabrous both 
sides, elliptic or oblanceolate, 4-5 cm. wide, 10-16 cm. long, about 
2.5 times as long as wide, at apex cuspidate-acuminate with acumen 
1-1.5 cm. long, the base acute, the margins decurrent on the chan- 
neled petiole 1 mm. thick, 8-10 mm. long; midvein convex or nar- 
rowly sulcate above, prominent beneath; lateral veins 8-12 pairs, 
sometimes with nearly similar intermediate veins, slightly convex 
on both sides of the blade; marginal vein about equaling the laterals 
and arched between them, or scarcely forming a distinct vein, 2-4 
mm. from margin, often with smaller veins evident beyond it; blade 
smooth and featureless above, paler and dull beneath with pale in- 
conspicuous glandular dots; inflorescence to 10 cm. long, 3 times 
compound, many-flowered; flowers sessile, clustered near the tips of 
the branches; buds 3.2-4 mm. long, obovoid, the calyptra rounded 
and domelike, punctate, and (in the type) glabrate; hypanthium 
markedly produced beyond the ovary, the orifice about 1 mm. across; 
stamens probably 50-75. The type, which I saw in Munich in 1954, 
bears both buds and opening flowers; it is labeled "Rio dos Enganos, 
in flumen Japura, Provinciae Rio Negro," Martins (F. M. Neg. 
19884). Krukoff's no. 6221 (NY, US), from the State of Ama- 
zonas, Mun. Humayta, near Tres Casas, is referred to this species 
with some doubt. It is said to be a tree 60 feet high; the plants are 
in young bud with inflorescence scarcely developed. The foliage 
closely resembles that of the type of C. cuspidata, but the inflores- 
cence appears to be more strongly velutinous than in the Martius 
specimen. 

Peru (probably). Amazonian Brazil. 

Calyptranthes densiflora Poepp. ex Berg, Linnaea 27: 30. 1855. 

A tree 4-7 meters high, sparsely covered in the inflorescence and 
on the lower leaf-surfaces with brownish-yellow fusiform sessile but 
rather loosely appressed dibrachiate hairs up to 0.8 mm. long on 



FLORA OF PERU 601 

the midvein and about 0.5 mm. long on the inflorescence; leaves 
elliptic or ovate, 3-5.5 cm. wide, 7-13 cm. long, 2-2.5 times as long 
as wide, acuminate, the base acute to rounded, the margins decur- 
rent on the stout petiole 1-1.5 mm. thick, 5-6 mm. long; midvein 
impressed or sharply sulcate above, prominent beneath; lateral veins 
15-20 pairs, slender and close with numerous intermediate veins, 
obscure above, more distinct beneath; marginal vein about equaling 
the laterals, 1.5-2 (-3.5) mm. from margin; leaves nearly concolorous, 
lustrous and smooth and obscurely impressed-puncticulate above, 
slightly more yellowish-green or -brown beneath, the surface rather 
sparingly dotted with small glands, often with pale appressed hairs 
and some brownish-yellow dibrachiate hairs; inflorescence axillary, 
the axis flattened, 2-4 mm. long and at least 2-3 mm. wide; flower- 
ing branches 2, spicate, (3-) 8-12 cm. long, opposite, arising from 
the lowest nodes of the axis, the peduncle about half as long as the 
entire branch, angled but scarcely compressed, 2-2.5 mm. thick just 
below the first node; flowers sessile in clusters of 10-20 at each of 
3-5 nodes and at the tips, the clusters sub-opposite or usually some 
or all of them alternate, the axis often zigzag; bracts about half as 
many as the flowers and simulating an involucre about the clusters, 
1-2.5 mm. wide, 2-3 mm. long, obovate, cucullate, glabrous on the 
inner surface; buds 2-2.5 mm. long, obovoid, gland-dotted, not 
apiculate, strigose with lustrous golden or somewhat arachnoid hairs; 
calyptra 1.5-2 mm. across, explanate; hypanthium hollow nearly to 
the base of the bud, the walls thin and collapsing after anthesis; style 
5-6 mm. long; stamens 30-40, up to 5 mm. long, the anthers 0.4-0.5 
mm. long; fruit (according to Poeppig) smaller than a pea, globose, 
crowned with the tubular hypanthium. F.M. Neg. 31509. 

San Martin: Zepelacio near Moyobamba, King 3307, 3747. In 
sylvis ad Mission Tocache, Poeppig 2019 (type, herb. Wien). 

In Poeppig's original specimen, which I have seen through the 
kindness of Dr. Rechinger, the spikes are apparently somewhat un- 
developed and perhaps abnormal; they are about 1.5 cm. long, each 
with 25-40 flowers closely packed in a sub-cylindric cluster. 

Calyptranthes krugioides McVaugh, Fieldiana Bot. 29: 182. 
1956. 

Tree to 20 meters high, the branchlets and inflorescence thickly 
rufous-hirsute with coarse flexuous hairs about 0.5 mm. long; mid- 
vein above densely and veins beneath sparingly hairy in young foli- 
age; leaves elliptic or ovate, 2.5-6.5 cm. wide, 10-16 cm. long, 2.5-3 
(-4) times as long as wide, about equally and often abruptly narrowed 



602 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

to the slenderly or caudately acuminate tip, and to the rounded or 
subcuneate base, the margins decurrent on the stout petiole 1.5-2 
mm. thick and 5-11 mm. long; principal veins impressed above, the 
mid vein sulcate but raised in a narrow central line in this groove; 
veins prominent beneath, the principal laterals 10-12 pairs, the mar- 
ginal vein about equaling the laterals, 2-4 mm. from the margin; 
leaves drying brown, dull, the upper surface smooth and without 
evident glands, the lower lighter in color, minutely dark-dotted; in- 
florescences axillary, paired, spikelike, 5-6 cm. long, the axis with 
3-4 flowering nodes, about 1 mm. in diameter but appearing more 
than 2 mm. thick because of the hairs; flowers distant, 8-13 in each 
spike, at the upper nodes solitary and sessile, at the lower nodes in 
groups of 3 on a common branch 3 mm. long; bracteoles ovate, acute, 
about 2.5 mm. long; buds broadly obovoid to nearly globose, 5-6 mm. 
long, obscurely apiculate, completely concealed by the hairs; calyp- 
tra about 3.5 mm. high; hypanthium cup-shaped, sparingly strigose 
within; style 8 mm. long, glabrous or with a few hairs at base; sta- 
mens 125-150, nearly as long as the style, the anthers 0.5 mm. long; 
petals 3, elliptic, rounded at the apex, 3 mm. wide and 4 mm. long 
in the mature bud; fruit not seen. The name is given in reference to 
the superficial resemblance between this species and Krugia ferru- 
ginea, of northern South America and the West Indies. Univ. of 
Mich. Neg. 465. 

Loreto: Iquitos, edge of lake, October 11, 1929, alt. 120 meters, 
Williams 3675 (F, type). Amazonian Brazil (basin of Rio Jurud, 
Krukoff5041). 

Calyptranthes longifolia Berg, in Mart. Fl. Bras. 14, pt. 1: 
46. 1857. C. pleophlebia Diels, Verh. Bot. Ver. Brandenb. 48: 188. 
1907. 

Shrub to 3 meters high, thickly appressed-hairy in the inflores- 
cence, on the young branchlets and buds and petioles, with coarse 
rufous mostly dibrachiate hairs up to 1.5 (-2) mm. long; leaves at 
least when young with a few long hairs beneath; leaves elliptic or 
lanceolate, 4-6.5 (-8) cm. wide, 10-25 cm. long, 2.5-4.5 times as 
long as wide, gradually or abruptly acuminate at tip, rounded or 
often subcuneate at base, the margins decurrent on the stout short 
petiole 1.5-2.5 mm. thick, 4-6 (-8) mm. long; midvein impressed 
above, prominent beneath; lateral veins 15-20 pairs, like the mar- 
ginal vein usually somewhat impressed above and rather prominent 
beneath; marginal vein about equaling the laterals, somewhat arched 



r. FLORA OF PERU 603 

between them, 2-5 (-10) mm. from the margin; leaves rather dull 
when dry, at least the lower surface turning pale brown; upper sur- 
face smooth or inconspicuously impressed-puncticulate; lower surface 
sparingly glandular; inflorescence a pair of few-flowered stout cymes 
2-5 cm. long, the axis 1.5 mm. thick, from the lowest nodes of an 
axillary branch, this branch aborting or (usually, in the specimens 
seen) elongating and becoming leafy at tip, the cymes appearing 
lateral and supra-axillary; cymes about 15- or fewer-flowered, with 
1-5 (usually 2-3) nodes, each with an oblong or bluntly triangular 
bract, 3-3.5 mm. wide and 4-15 mm. long, at base; flowers sessile, 
mostly in crowded laterally compressed bracteate clusters of 3 to 5 
at the tips of the branches; bracts ovate, rounded on the back, acute, 
2-3.5 mm. long, persistent until the fruit is grown; buds about 2.5 
mm. long, obovoid, very shortly apiculate, completely hidden by the 
abundant hairs; calyptra explanate at anthesis, about 2 mm. wide; 
style undeveloped in specimens examined; stamens about 75, 3-4 
mm. long; fruit purple (Killip & Smith), probably globose, about 
1.5 cm. in diameter, 1-seeded, the seed kidney-shaped, 1 cm. long. 
As already pointed out by Amshoff (Med. Bot. Mus. Rijksuniv. 
Utrecht 86: 150. 1942), the flowers of a closely related species, 
Calyptranthes fasciculata Berg, are often unisexual. A similar con- 
dition apparently prevails in the present species, and in C. speciosa 
var. gigantifolia. F.M. Negs. 23383, 23388. 

San Martin: Chazuta, Rio Huallaga, Klug 4103. Loreto: Yuri- 
maguas, Poeppig 2162, type; Williams 4666. Stromgebiet des Uca- 
yali von 10 S. bis zur Mundung, Tessmann 3432 (F.M. Neg. 23394). 
Cerro de Escaler, 1300 meters, Ule 6751, type of C. pleophlebia. 
Junin: Colonia Perene, Killip & Smith 24933. Bolivia. 

Calyptranthes macrophylla Berg, in Mart. Fl. Bras. 14, pt. 1 : 
45. 1857. 

A shrub or tree to 5-6 meters high, the inflorescence covered with 
dark rusty fusiform, closely appressed dibrachiate hairs 0.3 mm. 
long; lower leaf-surface covered with very numerous, closely ap- 
pressed lustrous pale hairs 0.1-0.2 mm. long, mixed with a few 
darker dibrachiate hairs, these on the midrib up to 0.5 mm. long; 
leaves ovate or ovate-lanceolate, 5.5-9 cm. wide, 18-24 cm. long, 
2.7-4.2 times as long as wide, narrowed from the middle or below 
to the long-acute or gradually long-acuminate tip, rounded at base, 
the margins decurrent on the stout pubescent petiole 2.5 mm. thick, 
10-12 mm. long; midvein impressed or sulcate above, raised its own 
thickness beneath; lateral veins very slender, 30-35 pairs, very 



604 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

slightly elevated both sides; marginal vein nearly straight, about 
equaling the laterals, 1.5-2 mm. from the inrolled margin; blades 
lustrous, greenish-brown or darker and minutely impressed-punctic- 
ulate above, pale tan and somewhat lustrous beneath from the cov- 
ering of small hairs which also nearly cover the numerous small dark 
glands; inflorescence axillary, the axis flattened and short, 3-10 mm. 
long, 2-3 mm. wide; flowering branches 1.5-6.5 cm. long, only the 
two lateral developed, or also a third representing the principal axis 
of the flowering branch; each branch 5- to many-flowered, the flowers 
sessile or on pedicels 2.5 mm. long; bracts lance-linear, about 5 mm. 
long, probably deciduous at anthesis; buds obconic, scarcely apicu- 
late, 3-4 mm. long; disk in fruit sunken, glabrous, about 3 mm. wide; 
fruit globose, 7-11 mm. in diameter, the hypanthium prolonged into 
a cylindrical neck 1-1.5 mm. long, 2-2.5 mm. wide; calyptra some- 
times persistent on the fruit, explanate, 2.5 mm. across. Flowering 
specimens of this species have apparently not been recently collected, 
but the species itself is well known from the upper Amazon region of 
Brazil and from nearby Venezuela. The Krukoff specimen cited 
below is included with some doubt as to its identity; the leaves are 
sparsely pubescent, and slightly more prominently acuminate than 
in the other specimens seen; the flowers are more numerous, and the 
pubescence of the inflorescence is of relatively loose and pale hairs; 
possibly it represents a distinct species but this cannot be determined 
with certainty from the material, which is in bud only, and with 
over-mature and weathered foliage. F.M. Neg. 31510. (See also a 
note under C. speciosa.} 

Rio Acre: Near mouth of Rio Macauhan, Krukoff 5498. Ama- 
zonian Brazil to Venezuela. 

Calyptranthes maxima McVaugh, Fieldiana Bot. 29: 182. 
1956. 

A small tree, the branchlets and inflorescence puberulent with 
minute appressed brownish dibrachiate hairs about 0.1 mm. long, 
the inflorescence (and especially the hypanthium) pale scurfy as well; 
leaves elliptic, thin in texture, 7.5-10.5 cm. wide, 25-39 cm. long, 
3.5-4 times as long as wide, about equally narrowed to both ends, 
acuminate at tip, appearing cordate-auriculate at base when dry; 
petioles dark and stout, about 3 mm. thick, 4 mm. long, the thick- 
ened portion extending 5-6 mm. beneath the blade and there merg- 
ing into the unmodified midvein, the whole petiole depressed below 
the plane of the base of the leaf and lying in a short steep-sided fur- 
row; midvein centrally sulcate, but elevated on the upper surface, 



FLORA OF PERU 605 

prominent beneath; lateral veins very slender, 25-35 pairs in addi- 
tion to some intermediates, convex on both sides, more conspicuous 
beneath; marginal vein about equaling the laterals, nearly straight, 
2-4 mm. from margin; blades pale green and lustrous above, paler 
and with small dark glandular dots beneath; inflorescence from the 
terminal node of a stout terminal bracteate shoot 1.5 cm. long; bracts 
of the shoot scarious, coriaceous, lance-attenuate, 6 mm. wide, 15-25 
mm. long; bracteoles scarious, linear, blunt-tipped, 2.5 mm. wide 
and 7 mm. long, deciduous at anthesis; inflorescence branches soli- 
tary, 6-10 cm. long, 1 to 2 times compound, 6- to 11-flowered, on 
stout compressed peduncles 3.5 mm. wide below the first node; lower 
divisions of the primary branch 8 mm. long, 3-flowered; all flowers 
except the terminal one on flat pedicel-like branchlets 2-3 mm. long, 
2 mm. wide; buds 7-8 mm. long, obovoid and nearly flat- topped, the 
calyptra nearly flat, scarcely apiculate, 5-6 mm. across, glabrous 
inside; hypanthium glabrous within, prolonged 2.5 mm. beyond the 
summit of the ovary; stamens about 200, 5 mm. long, densely fring- 
ing the orifice of the hypanthium in a zone 1.5 mm. wide; anthers 
0.6-0.8 mm. long; style stout, about 8 mm. long, 0.4 mm. in diam- 
eter, enlarged (perhaps deformed) distally; ovary bilocular, the 
ovules in each locule 2, ascending. Known only from the Trapecio 
amazonico, in Colombian territory very near to the Peruvian bound- 
ary, Schultes & Black 8526, type. 

Peru (probably). Amazonian Colombia. 

Calyptranthes multiflora Berg, in Mart. Fl. Bras. 14, pt. 1: 
42. 1857. ?C. poeppigiana Berg, I.e. 45. 

A tree, pubescent in the inflorescence with numerous erect rufous 
hairs up to 0.2 mm. long, and usually with a few appressed, slightly 
longer and sometimes dibrachiate hairs near the base of the pan- 
icles; petioles and mid veins of young leaves near the base sparingly 
pubescent; leaves elliptic or narrowly ovate, 2-4 (-6) cm. wide, 
5-10 cm. long, (1.7-) 2.4-3 times as long as wide, about equally 
narrowed to the short-acuminate tip and the cuneate or subcuneate 
base, the margins decurrent on the short stout petiole 2-2.5 (-3.5) 
mm. long; midvein prominent beneath, not impressed above but 
plane or forming a low flat or convex ridge 0.5-1 mm. wide; lateral 
veins about 10 pairs, but often seeming to be 20 or more closely 
parallel veins on a side because of the many smaller intermediates; 
marginal vein about equaling the laterals, somewhat arched between 
them, usually less than 1 mm. from margin (up to 4 mm.); leaves 
dull in drying, nearly concolorous or the lower surface paler, both 



606 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

sides with small inconspicuous glandular dots, the veins obscure 
above, evident beneath at least in mature leaves; inflorescence a 
pair of panicles, up to 8.5 cm. long and 2-3 cm. wide, from the basal 
nodes of a short axillary axis (or the axis elongating 3-4 mm. and 
producing a second and shorter pair of panicles from the next node) ; 
panicles 3 times compound, usually with about 5 irregular nodes, 
the lower branches up to 7- or 9-flowered; pedicels of the ultimate 
branchlets none, or up to 2 mm. long; flowers sessile above the brac- 
teoles, but the base of the hypanthium often narrowed into a pedicel- 
like base; bracts and bracteoles linear, glabrous, deciduous before 
anthesis, the former 2-4 mm. long, the latter 1-1.5 mm.; buds 2-2.5 
mm. long, glabrous, roughened by the prominent glands, apiculate, 
obovoid and usually attenuate at base; calyptra about one- third the 
length of the bud; petals 2 or more, spatulate, erose, 1 mm. long and 
nearly as wide; style 5 mm. long; stamens about 100, up to 6 mm. 
long; fruit not seen. F.M. Neg. 23384. 

Loreto: Timbuchi, Williams 1087. Manfinfa, Williams 1089. 
Amazonian Brazil, Venezuela and Colombia. 

Calyptranthes poeppigiana Berg, the type of which came from 
Ega, Brazil (Poeppig, s.n., in herb. Wien), is indistinguishable from 
C. multiflora except that the inflorescences are reduced to paired or 
glomerate spikelike branches 1.5-3 cm. long. Dr. Rechinger kindly 
made the type of C. poeppigiana available for my study, but I have 
not seen any other specimens resembling it, and I suppose that it 
represents an aberrant form of the relatively widespread C. multi- 
flora. Univ. of Mich. Neg. 1018. 

Calyptranthes paniculata R. & P. Fl. Peruv. Prodr. 74. t. 13. 
1794. C. fragrans in herb. Ruiz, ex Berg, Linnaea 27: 20. 1855. 

Tree to 30 meters high, completely glabrous or the vegetative 
buds strigose with reddish fusiform dibrachiate hairs up to 0.5 mm. 
long, and a few similar hairs persistent at the base and about the 
nodes of the inflorescence; leaves elliptic to lance-ovate or ovate, 
2.5-4 (-6.5) cm. wide, 8-12.5 cm. long, (2-) 2.5-3.2 times as long as 
wide, bluntly short-acuminate, at base cuneate or sub-cuneate, or 
occasionally somewhat rounded, the margins decurrent on the petiole 
1.5 mm. thick, 7 mm. long; midvein broadly sulcate above, promi- 
nent beneath; lateral veins very slender, about 12 (-15) pairs, ob- 
scure above, slightly elevated beneath; marginal vein about equaling 
the laterals and arched between them, 2-3 (-6) mm. from margin, 
with an outer, very fine, submarginal vein beyond it; leaves in dry- 



FLORA OF PERU 607 

ing pale green or brown above, somewhat lustrous and impressed- 
puncticulate, the lower surface greenish-brown, sparingly and mi- 
nutely gland-dotted; inflorescence axillary, the axis tangentially 
flattened, 2.5-3 mm. long and wide, the 2 flowering branches from 
the lowest nodes, 8-10 cm. long, 3 to 4 times compound, many- 
flowered, divaricately and irregularly branched, the peduncle terete 
or nearly so, 1-1.5 mm. thick; branches alternate or opposite, the 
nodes often enlarged, the ultimate branchlets divaricate or somewhat 
reflexed, 2-10 mm. long, distally enlarged and bearing 1-3 sessile 
terminal flowers; bracts deciduous at anthesis, ciliate, 1-1.5 mm. 
long; buds 3-3.5 mm. long, obovoid or constricted about the mid- 
dle, narrowed to the broad sessile base; calyptra about half as long 
as the persistent hypanthium, broadly infundibuliform, 1.3-1.5 mm. 
wide, with a conic apiculum 0.5 mm. long; hypanthium after dehis- 
cence campanulate, 1.5-2 mm. across the orifice; fruit and flowers 
unknown. F.M. Neg. 23387. 

Huanuco: Pozuzo, Ruiz (type, in herb. Madrid?, not seen). 
Rio Acre: near mouth of Rio Macauhan, Krukoff 5258, 5487. Local- 
ity uncertain: Ruiz (isotype of C. fragrans, herb. Geneva ex herb. 
Berol. ex herb. Lambert); Ruiz 24/45 (herb. Chicago ex herb. 
Madrid). 

Calyptranthes plica ta McVaugh, Fieldiana Bot. 29: 182. 1956. 

A shrub 4.5 meters high, nearly glabrous except that the inflores- 
cence is appressed-hirsutulous with rufous partly dibrachiate hairs 
up to 0.6 mm. long, the petioles and the midveins on the lower sur- 
face of the leaf puberulent; leaves large, sessile, obovate, 10-12 cm. 
wide, 30-38 cm. long, 2.5-3 times as long as wide, short-acuminate, 
narrowed to a width of 4-5 cm. near base and the margins there 
broadly decurrent and plicate on the enlarged petiolar base which is 
5 mm. thick on the lower surface; mid vein impressed or deeply sul- 
cate above, prominent beneath; lateral veins 20-25 pairs, impressed- 
above, prominent beneath; marginal vein about equaling the lat- 
erals and but slightly arched between them, 2-4 mm. from the mar- 
gin; upper surface of blade dull and obscurely impressed-punctate; 
lower surface dull and probably paler, without apparent glands; in- 
florescence a divaricately branched, 3 times compound, bracteate 
panicle up to 7 cm. long, the panicle-branches very slender, up to 
1.5 mm. in diameter, the nodes with conspicuous persistent divari- 
cate boat-shaped attenuate bracts 2-3 mm. long (the lowest up to 
1.5 mm. wide, 5 mm. long); flowers 50-100, mostly sessile in pairs 
along the spikelike secondary branches of the panicle, or in 3's at 



608 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

the tips of the branches; immature buds obovoid, 3.5 mm. long, the 
calyptra pointed and somewhat apiculate, the hypanthium turbinate, 
glabrous within, prolonged at least 1 mm. beyond the summit of the 
ovary; stamens about 60; style not seen in buds examined; petals 
none. A distinctive species, but unfortunately known only from 
the type specimen, which includes a single inflorescence with half- 
grown buds, and another, detached inflorescence with very much 
younger, undeveloped buds. 

Peru (probably). Amazonian Brazil (Amazonas, Mun. Sao Paulo 
de Olivenca, Krukoff 8432, type). 

Calyptranthes pulchella DC. Prodr. 3: 257. 1828. 

A tree to 8 meters high, glabrous except the vegetative buds and 
youngest branchlets strigose (usually very sparingly) with reddish 
hairs up to 0.5 mm. long; branchlets 2-winged, the wings up to 0.5 
mm. high, arising just above the axil at one node and terminating 
between the leaf -bases at the node above, in an auricle-like prolonga- 
tion 1-1.7 mm. long; leaves obovate or broadly elliptic, 1.5-3.3 cm. 
wide, 2.5-6 cm. long, 1.5-2 times as long as wide, the apex rounded, 
acute but blunt-tipped, or broadly and shortly acuminate, the base 
acute or cuneate, the margins decurrent on the stout channeled peti- 
ole 1 mm. thick, 2-3 mm. long; midvein sulcate and mostly impressed 
above, prominent beneath; lateral veins very slender and scarcely 
apparent, about 10-12 pairs with several intermediates; marginal 
vein like the laterals, 0.5-1 mm. from the margin; leaves dull in dry- 
ing, pale green and indistinctly impressed-puncticulate above, pale 
brown and dotted with small dark glands beneath; inflorescence 
axillary, the axis either abortive and 1 mm. long, or elongate and 
leafy; flowering branches 2, 3-5.5 cm. long, opposite, arising from 
the lowest nodes of the axis, filiform or very slender, terete at base, 
gland-dotted, the peduncle distally enlarged and compressed, 2-2.5 
cm. long, up to a little more than 1 mm. wide below the first node, 
often more slender; flowers few, mostly 3-11, sessile on the axis and 
at the tip or the lowest on lateral branches 3-6 mm. long; bracts and 
bracteoles not seen; buds 4-7 mm. long, 2-3 mm. thick, oblong or 
constricted above the middle, prominently gland-dotted and obconic 
above the broadly sessile base, obtusely pointed and rounded or ob- 
scurely apiculate, the calyptra urceolate, 2-3 mm. long; style gla- 
brous, 6-8 mm. long (often with a subglobose small gall near the 
middle, the gall caused by a member of the Cynipidae); stamens 
about 125, up to 5-6 mm. long, the anthers 0.3 mm. long; fruit 
globose, 8 mm. in diameter, thickly glandular-verruculose. The 



FLORA OF PERU 609 

Peruvian plant resembles the type of C. pulchella 5 parviflora Berg, in 
Mart. Fl. Bras. 14, pt. 1 : 516. 1859, namely Spruce's no. 2729 from near 
Panure" on the Rio Vaupe"s. The type of C. pulchella var. pulchella, 
a collection made by Martius in the State of Minas Geraes, has the 
inflorescence and hypanthium sparsely red-strigose, but a compari- 
son made at Munich in 1954 between this specimen and an isotype 
of d parviflora indicates that these are conspecific. The Peruvian 
specimens have larger buds than the plants collected by Martius and 
Spruce; for notes on the varieties of this species see Mem. N. Y. 
Bot. Card. 10: 76-77. 1958. F.M. Neg. 19897 (var. pulchella}. 
Univ. of Mich. Neg. 1031 (var. parviflora). 

Loreto: Mishuyacu, Klug 1373. Timbuchi, Williams 959. 
Southern Brazil; Amazonian lowlands, Brazil to Colombia. 

Calyptranthes rufotomentosa McVaugh, Fieldiana Bot. 29: 
183. 1956. 

A shrub, the inflorescence densely tomentose with dark reddish- 
brown tangled hairs up to 1 mm. long; leaves glabrous, oblanceolate, 
4-5 cm. wide, 15-21 cm. long, about 4 times as long as wide, nar- 
rowly acuminate with blunt-pointed acumen, the margins attenuate 
toward the base from above the middle, then abruptly narrowed and 
passing into the ventral surface of the nearly terete, stout, dark gla- 
brous petiole 3 mm. thick, 7 mm. long; midvein elevated above in a 
flattened ridge or merely convex, elevated beneath its own thick- 
ness; lateral veins 10-15 pairs, rather distant and without conspic- 
ous intermediates, convex but not prominent above, more evident 
beneath; marginal vein about equaling the laterals, nearly straight, 
prominent beneath, 3-4 mm. from margin; blades dark and lustrous 
above, and obscurely impressed-punctate, the lower surface dull and 
paler; inflorescence (not seen attached) about 6 cm. long, 3 times 
compound, about 50-flowered, the flowers clustered toward the tips 
of the branches, sessile; bracts scarious, glabrous or glabrescent dor- 
sally, the larger ones linguaeform, 3 mm. wide, 8 mm. long, subper- 
sistent; buds probably 3-4 mm. long before opening; hypanthium in 
open flowers urceolate, 4 mm. long, rounded at base, 3-3.5 mm. 
across the mouth, glabrous within, produced 1.5 mm. beyond the 
summit of the ovary; calyptra conic, apiculate, 1 mm. high, 1.5-2 
mm. across; stamens about 75, 2 mm. long, the anthers 0.3 mm. long; 
style not seen; petals 3(?), narrow and slender-pointed, 0.6 mm. wide, 
1.7 mm. long. 

Peru (probably). Amazonian Brazil (Amazonas, Sao Paulo de 
Olivenca, Ducke 2240, type). 



610 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Calyptranthes ruiziana Berg, Linnaea 27: 22. 1855. 

Shrub or small tree to 5 meters high, the branchlets not or very 
narrowly winged, the branchlets, inflorescence and vegetative buds 
thinly covered with yellowish- or reddish-brown fusiform appressed 
dibrachiate hairs mostly about 0.3 mm. long (up to 0.6 mm. long 
or more, on leaf -bases) ; lower leaf-surfaces with few dark hairs and 
usually with rather numerous persistent pale appressed hairs; leaves 
elliptic, 2.5-4 cm. wide, 5.5-10 cm. long, 2-3 times as long as wide, 
acuminate, the base acute, the margins cuneately decurrent on the 
stout petiole 1-1.3 mm. thick, 6-7 mm. long; midvein sulcate above, 
prominent beneath; lateral veins 12-15 pairs or apparently more 
because of the numerous parallel intermediate veins, obscure above, 
somewhat elevated but not very prominent beneath; marginal vein 
about equaling the laterals and little arched between them, 0.7-2 mm. 
from the margin; blades smooth and somewhat lustrous, green or 
drying brown above, impressed-puncticulate but this often obscure 
in mature leaves; lower surface yellow-green or whitish, obscurely or 
apparently not at all dotted; inflorescence axillary, the axis tangen- 
tially flattened, glabrous, 2 mm. wide, very short or up to 10 mm. 
long; flowering branches 2, from the lowest nodes, 3 times compound, 
many-flowered, (4-) 6-10 cm. long, the panicle 4-5 cm. wide at base, 
the branches opposite or alternate, the flowers sessile, aggregated 
toward the tips of the branches and mostly in 3- to 10-flowered 
clusters, these short-pedunculate except the terminal; bracts decid- 
uous before anthesis; buds 2-2.5 mm. long, obovate, shortly apicu- 
late or nearly rounded at tip, the base obconic, strigose; hypanthium 
after dehiscence campanulate, 1.5-1.7 mm. long, the orifice 1.3-1.7 
mm. wide in flower and fruit; calyptra concave, 1.3-1.7 mm. wide; 
style 4-4.5 mm. long; stamens about 60, up to 5 mm. long, the an- 
thers about 0.3 mm. long; fruit globose or oblate, 4-6 mm. in diam- 
eter, thickly gland-dotted, the hypanthium prolonged into a short 
neck more than 1 mm. wide and nearly 1 mm. long. 

Loreto: Along Rio Itaya, Williams 175. Along Rio Mazan near 
Iquitos, Williams 8148. Without locality: Ruiz (type, in herb. 
Berlin, not seen); Ruiz 5105 (probable isotype, herb. US ex herb. 
Berlin ex herb. Lambert) . 

Tentatively referred to this species, but with slightly larger, 
nearly eglandular fruit, and narrower leaves, is Killip & Smith's 
no. 29199, from San Lorenzo, Loreto, between mouths of Rio Pas- 
taza and Rio Huallaga. A plant which will be found here in the key, 
but perhaps represents another species, is Killip & Smith's no. 28293 



FLORA OF PERU 611 

(herb. NY), from between Yurimaguas and Balsapuerto, on the 
lower Rio Huallaga. The leaves are up to 11 cm. wide and more 
than 20 cm. long, with markedly impressed veins (including the 
marginal and the 15 or more pairs of laterals). The lower panicle 
branches are 5 cm. long. The calyptra is conic, apiculate, 1 mm. 
long; fruit 6-7 mm. in diameter, with style 4.5 mm. long. 

Calyptranthes sessilis McVaugh, Fieldiana Bot. 29: 183. 1956. 

A small shrub, glabrous except a few minute flat brownish hairs 
at the base of the hypanthium; leaves ovate, cordate, sessile (the 
broad compressed petiolar base up to 3 mm. long and wide), 4-7 cm. 
wide, 9-12 cm. long, 1.8-2.5 times as long as wide; blades acuminate, 
the acumen rounded at very tip; midvein prominent beneath, im- 
pressed above; lateral veins about 15 pairs, impressed above; mar- 
ginal vein impressed above, about equaling the laterals and arched 
between them, 3-6 mm. from the margin, and with a smaller vein 
parallel to it and between it and the margin; leaves dull in drying, 
the upper surface obscurely glandular at maturity, the lower more 
plainly gland-dotted; inflorescence axillary or falsely terminal, con- 
sisting of paired narrow panicles 6 cm. long, these opposite and aris- 
ing from the two sides of a primordium in the axil of each of the 
terminal leaves; panicles sometimes bracteate at base, the bracts 
lanceolate, keeled, 3 mm. wide, 6 mm. long; peduncle 2-3 cm. long, 
somewhat compressed and 1.7 mm. wide below the first node; nodes 
about 5, somewhat irregular, the lower ones bearing paired pedicel- 
late clusters of 3 flowers each, or all nodes with the pedicels 1-flow- 
ered and opposite or subopposite; bracteoles and bracts (except the 
basal) not seen; flowers sessile, or on stout pedicels up to 2 mm. long, 
0.5 mm. thick; buds fusiform, 6-7 mm. long and 2 mm. thick, taper- 
ing about equally to the slender hypanthium and the cylindrical apex, 
or somewhat abruptly contracted into the beak; operculum 4-4.5 
mm. long; petals none; style 8.5 mm. long; stamens 60-75, about 
8 mm. long; anthers broadly elliptic, about 1 mm. long; fruit globose, 
7 mm. thick; seed 6 mm. long, the radicle about half as long, com- 
pressed to one side of the cotyledons. Calyptranthes spruceana Berg, 
in Mart. Fl. Bras. 14, pt. 1: 45. 1857, based on Spruce 1551 from the 
lower Rio Negro, is very similar. In that species, however, the buds 
are said to be globose and 5 mm. in diameter, the leaves are obtusely 
rounded at the tips, the veins are not impressed on the upper sur- 
face, and the fruits are more clustered near the tips of the inflores- 
cence. Univ. of Mich. Neg. 446. 



612 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Loreto: Yurimaguas, Williams 4583 (F, type); Killip & Smith 
27603. 

Calyptranthes Simula ta McVaugh, Fieldiana Bot. 29: 184. 
1956. 

Shrub or tree to 9 meters high, the inflorescence [and probably 
the vegetative buds and young leaves and branchlets] with a thin 
covering of flat fusiform yellowish T brown appressed hairs mostly 
about 0.2-0.3 mm. long, a few similar hairs persisting on the lower 
leaf-surface; leaves ovate or lanceolate, 3.5-6 cm. wide and 9-15 cm. 
long, or on vigorous branches 7-8 cm. wide, 18-25 cm. long, all 2.5- 
3 (-3.6) times as long as wide, somewhat narrowed toward the apex 
from the middle or below and prominently triangular-acuminate, 
gradually rounded to the subcuneate base, the margins decurrent on 
the stout dark petiole 1.5-2.5 mm. thick, 7-12 mm. long; midvein 
broadly sulcate above, prominent beneath; lateral veins 20-25 or 
more pairs in addition to several intermediate ones, all slender, 
slightly raised on both surfaces, prominulous beneath; marginal vein 
similar to the laterals and arched between them, 2-3 (-5) mm. from 
margin, a very fine submarginal vein visible beyond it; blades dry- 
ing green, the upper surface smooth and impressed-puncticulate, the 
lower yellow-green, dull, finely dark-dotted; inflorescence axillary, 
the short axis much flattened tangentially, up to 3 mm. wide and 
about 4 mm. long, the paired flowering branches from the lowest 
node, 6-11 cm. long, the peduncle 1.5-2 (-4) cm. long, somewhat 
angled or compressed, 1.5-2 mm. wide below the first node; panicle 
narrow, or broader and loosely many-flowered, 3 to 4 times com- 
pound, the lower branches up to 3-5 cm. long; flowers numerous, 
sessile, mostly in 3's on short peduncles along the branches and near 
the tips; branches opposite or mostly alternate; bracts deciduous 
before anthesis; buds 2-2.5 mm. long, obovoid, turbinate and more 
or less rufous-strigose near base, and often arachnoid-whitened as 
well, the apex rounded or shortly apiculate; hypanthium after de- 
hiscence broadly campanulate, 1.3-1.5 mm. long, the orifice about 
1.5 mm. wide; calyptra explanate, 1.3-2 mm. wide; style 5-6 mm. 
long or a little less; stamens 40-50, 4-6 mm. long, the anthers 0.3 mm. 
long. This species is known only from specimens in which the flow- 
ering branches are borne in the terminal axils of the twigs, on old 
wood or what appears to have been a vigorous shoot of the preced- 
ing season. The normal size and shape of the leaves on average 
twigs cannot be determined with certainty, nor can the winged or 
non-winged condition of the young branchlets be ascertained. It is 



FLORA OF PERU 613 

possible that this plant is conspecific with Calyptranthes tessmannii, 
which it resembles in many respects. From the specimens at hand, 
however, the two are abundantly separable by many characters. 

Loreto:'lquitos, Aug. 2-8, 1929, Killip & Smith 26916, 27352. 
Iquitos, Tessmann 5372 (G); Peru-Colombia boundary, forest near 
Rio Putumayo, Sept. 26-Oct. 10, 1930, G. King 1614 (MICH, type). 

A collection from Tarapoto, San Martin, Williams 6539, is prob- 
ably the same species; it bears immature globose or oblate fruit about 
1 cm. in diameter; the pubescence is exactly that of C. simulata and 
the leaves are very like those of that species except that the lateral 
veins are hardly more than 15 pairs; the fertile axis is continuous 
and leafy and the flowering branch is lateral from its base. 

Calyptranthes speciosa Sagot, Ann. Sci. Nat. ser. 6, 20: 187. 
1885, var. gigantifolia (McVaugh) McVaugh, Mem. N. Y. Bot. 
Card. 10: 79. 1958. C. gigantifolia McVaugh, Fieldiana Bot. 29: 
181. 1956. 

Tree up to 20 meters high, the inflorescence densely appressed 
pubescent with coarse rufous mostly dibrachiate hairs up to 0.5 mm. 
long, the leaves glabrous or pubescent beneath especially when young 
with a few pale flaccid hairs or with a few dibrachiate hairs inter- 
mixed; leaves elliptic or oblong, 6-15 cm. wide, 14-38 cm. long, 2.5-4 
times as long as wide, rather abruptly narrowed at both ends, the 
tip narrowly acuminate, the base rounded and the margins decur- 
rent on the stout petiole 8-12 mm. long, 1.5-3.5 mm. thick; mid vein 
and usually the marginal vein impressed above, all veins prominent 
beneath; lateral veins 20-30 pairs; marginal vein continuous, about 
equaling the laterals and somewhat arched between them, 2-4 mm. 
from the nearly plane margin, usually with a smaller outer vein be- 
yond it; leaves often browning in drying, the upper surface smooth, 
sparingly and often obscurely impressed-puncticulate, the lower sur- 
face gland-dotted; inflorescence axillary and usually 4 times com- 
pound, often appearing supra-axillary when the central axis elongates 
and becomes leafy, or terminal when no more than one axillary bud 
develops at a terminal node; basal and longest branches of the pan- 
icle 3 times compound, 6-10 cm. long with up to 100 flowers, divari- 
cately branched, the branches compressed and up to 2 mm. wide 
below the nodes; buds 3-5 mm. long, sessile, obovate, rounded or 
obscurely apiculate at tips, clustered near the tips of the branchlets; 
calyptra about half as high as the persistent hypanthium; bracts and 
bracteoles mostly deciduous before anthesis, lanceolate to ovate, 
1-2.5 mm. long or the lowest bracts persistent, foliaceous, ovate, 



614 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

1.5 cm. long, 6-8 mm. wide; style apparently defective in all flowers 
seen, 1-1.5 mm. long; stamens 150-175, about 5 mm. long, cream- 
color or golden-yellow (Klug), the anthers about 0.5 mm. long; fruit 
globose, not seen mature, but by analogy with related species prob- 
ably 8-10 mm. in diameter. Colombian specimens collected by 
Mutis (nos. 1940, 2234, 2951, 3961, and 5754) appear to belong to 
this variety also. Also closely related is Calyptranthes macrophylla 
Berg, based on Poeppig 2739, from Ega, Brazil. This latter species, 
however, differs markedly in leaf-characters: its leaves are more 
coriaceous, acute rather than acuminate, less prominently veined; 
the marginal vein is close to the margin and not impressed above; 
the lower surface is densely pubescent with minute appressed pale 
hairs and with numerous dibrachiate hairs intermixed. In this last 
respect C. macrophylla agrees precisely with C. speciosa var. speciosa 
of the Guianas. C. gigantifolia is scarcely distinguishable from typ- 
ical C. speciosa and surely does not merit recognition as a distinct 
species. Its leaves are glabrous or nearly so beneath (densely and 
finely pubescent, as noted above, in C. speciosa) ; the midvein is im- 
pressed on the upper surface (in C. speciosa sometimes impressed but 
more often plane or convex) ; the marginal vein lies 2-4 mm. from the 
margin (as against usually less than 1 mm. in C. speciosa) ; the flowers 
in C. gigantifolia appear to be somewhat larger than those of C. speci- 
osa and with more numerous stamens (150-175 as against 80), but 
seem otherwise indistinguishable. The inflorescence in the two 
plants, with its somewhat unusual supernumerary branches, appears 
to be identical. Univ. of Mich. Neg. 486. 

San Martin: Juanjui, Klug 4277 (US, type). Loreto: Florida, 
Klug 2332, 2347. Bolivia, Amazonian Colombia. 

Calyptranthes tessmannii Burret, in herb., ex McVaugh, 
Fieldiana Bot. 29: 184. 1956. 

A shrub or tree, the branchlets and vegetative buds, inflorescence 
and petioles loosely appressed pubescent with yellow-brown fusi- 
form dibrachiate hairs up to 0.5 mm. long, a few hairs persisting 
on the lower surface of the leaves, especially on the mid veins; branch- 
lets narrowly wing-angled, the wings arising above the axil at one 
node and terminating between the leaf-bases at the node above, the 
terminal auricle wider than the wing, up to 0.5 mm. wide, 1.7-2.5 
mm. long; leaves elliptic, 3-4.5 cm. wide, 8-11.5 cm. long, about 2.5 
times as long as wide, nearly equally rounded to both ends, the tip 
abruptly and caudately acuminate, the acumen 3 mm. wide at base, 
1.3-2.5 cm. long; base of blade acute or slightly rounded, the mar- 



FLORA OF PERU 615 

gins cuneately decurrent on the stout petiole 1-1.5 mm. thick, 8-10 
mm. long; mid vein impressed above, prominent beneath; lateral 
veins about 15 pairs, slightly elevated on both surfaces, prominu- 
lous beneath; marginal vein about equaling the laterals and arched 
between them, prominulous beneath, 1-2.5 mm. from margin, with 
a faint submarginal vein sometimes discernible beyond it; blades 
dark or dull green above, smooth and impressed-punctate, the lower 
surface paler, yellowish-green, dotted with small glands; fertile 
branches leafy, axillary, the primary axis 5 mm. long or less, tan- 
gentially much flattened, 3 mm. wide below the first node, from 
which arise the 2 opposite narrow panicles 5-10 cm. long and up to 
2.5 cm. wide at base; peduncle 2-3.5 cm. long, somewhat com- 
pressed, 1.5-2.5 mm. wide below the first node; lower branches 
about 1.5 cm. long, 10-flowered, the branches from the upper nodes 
shorter, often alternate, the flowers sessile, fewer, in short-peduncu- 
late clusters; bracts narrowly elliptic or oblanceolate, 3 mm. long, 
partly persistent until anthesis; buds 3-3.5 mm. long, fusiform, ap- 
pressed brown-hairy, the narrow apiculate tip 1 mm. long (up to 
2 mm. on the calyptra after dehiscence) ; calyptra about 2 mm. wide, 
the hypanthium after dehiscence broadly infundibuliform or cam- 
panulate, less than 2 mm. long; style 6 mm. long; stamens about 50, 
as long as the style, attached about at the line of dehiscence and 
falling almost with the calyptra; anthers about 0.3 mm. long. 
F.M. Neg. 23395. 

Loreto: Stromgebiet des Maranon von Iquitos aufwarts bis zur 
Santiago-Miindung am Pongo de Manseriche, ca. 77 30' west, G. 
Tessmann 4832 in 1924 (G, type). 

Calyptranthes tridymantha Diels, Verh. Bot. Ver. Brandenb. 
48: 188. 1907. 

Shrub 3-9 meters high, loosely tomentose on the young branch- 
lets and vegetative buds and inflorescence, with golden-yellow di- 
brachiate hairs up to more than 1 mm. long, the basal stalk of the 
hair erect and often as long as the spreading or ascending branches; 
mature leaves often with a few hairs persisting beneath; branchlets 
narrowly 2-winged, the wing up to 0.5 mm. wide, arising above the 
axil at one node and terminating between the leaf-bases at the node 
above, in an auricle-like prolongation 2 mm. long; leaves elliptic, 
ovate, or occasionally obovate, 2-3.5 cm. wide, 3-5.5 cm. long, 1.8- 
2.4 times as long as wide, the apex rounded to obscurely and bluntly 
acuminate, the base rounded, or somewhat narrowed from below 
the middle and abruptly contracted to the very base, the margins 



616 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

shortly decurrent on the petiole 1-1.5 mm. thick, 2-2.5 mm. long; 
midvein impressed above, prominent beneath; lateral veins 10-12 
pairs, slightly elevated and visible on both sides, but very slender 
and with numerous intermediate veins nearly the same size; mar- 
ginal vein distinctly double, the inner about equaling the laterals 
and slightly arched between them, 1-1.5 mm. from margin; leaves 
dull green and obscurely or not at all impressed-puncticulate above, 
pale brown beneath, the glands not or scarcely apparent; inflores- 
cence axillary, the axis abortive, the flowering branches 2, 3-5 cm. 
long, each few- (about 9-) flowered, opposite from the lowest nodes, 
tomentose and very minutely gland-dotted, the peduncle 2-2.5 cm. 
long, slightly compressed distally and about 1 mm. wide near sum- 
mit; lateral secondary branches usually a single pair, 6-20 mm. long; 
flowers sessile, in terminal triads; bracts deciduous before anthesis, 
subfoliaceous, less than 1 cm. long; bracteoles not seen; buds prob- 
ably about 5 mm. long, shortly apiculate; calyptriform calyx [i.e., 
the calyptra?] 2-2.5 mm. across (Diels); style longer than the sta- 
mens; stamens 6 mm. long, white (Diels) ; petals none. A distinctive 
species by virtue of the golden-yellow tomentum of erect, stalked 
and branched hairs. F.M. Neg. 23396. 

Rio Acre: Near mouth of Rio Macauhan, Krukoff 5549. Ama- 
zonian Brazil. 

3. MYRCIA DC. 

Reference: Berg, Linnaea 27: 35-80, 82-129. 1855-1856; and in 
Mart. Fl. Bras. 14, pt. 1: 59-143, 150-210. 1857. 

Calyx-lobes 5 (rarely 4), distinct and imbricated in bud and in 
flower, and usually persistent on the fruit; hypanthium variously or 
not at all prolonged beyond the summit of the ovary. Petals usu- 
ally 5, often half as long as the stamens or longer, and conspicuous. 
Inflorescence-axis usually prolonged, with several pairs of lateral 
branches. 

A large genus of tropical America; Myrcia DC. and Aulomyrcia 
Berg, according to Berg, together comprised more than 400 species. 
The supposed distinction between these two groups lies in the hypan- 
thium, which in Myrcia is described as "scarcely produced beyond 
the ovary, and constricted beneath the calyx," and in Aulomyrcia 
is described as "strongly produced beyond the ovary, but not con- 
stricted beneath the calyx." Although numerous species can be 
assigned to one genus or the other on the basis of this character, 
it seems quite impossible to apply the character generally because 



FLORA OF PERU 617 

various other species occupy positions intermediate between the ex- 
tremes, and the placement of a given species in Myrcia rather than 
Aulomyrcia, or the reverse, can hardly be made objectively. I have 
therefore followed Bentham, in Benth. & Hook. f. Gen. PI. 1: 716. 
1865; Kiaerskou, Enum. Myrt. Bras. 63. 1893; and Urban, Bot. 
Jahrb. 19: 577-582. 1895, in relegating Aulomyrcia to synonymy. 

Leaves sessile, cordate. 

Leaves ovate, 13-17 cm. wide, 1.7-2 times as long as wide, the 
veins impressed above; inflorescence decompound, stout, 20 
cm. long or more; buds 7 mm. long or more; northeastern 
Peru (Loreto) M. obumbrans 

Leaves lanceolate, 3.5-5 cm. wide, 2.5^4 times as long as wide, 
the lateral veins not impressed above; inflorescence few- 
flowered, 4-7 cm. long; buds 4-6 mm. long; northern Bolivia 
(La Paz) M. connata 

Leaves definitely petiolate, cuneate or rounded at base or occasion- 
ally subcordate. 

Inflorescence at anthesis (and usually in fruit) conspicuously 
bracteate, the bracts ovate, pointed, 6-12 mm. long; calyx- 
lobes 2.5-6 mm. long, lanceolate or ovate, much longer than 
wide; branchlets long-hirsute. 

Hairs of branchlets reddish-brown, up to 3.5 mm. long; calyx- 
lobes 4, the larger ones 6 mm. long M. huallagae 

Hairs of branchlets yellowish-brown, up to 2 mm. long; calyx- 
lobes 5, rarely 4, the larger ones 2.5-4 mm. long. 

M . bracteata 

Inflorescence with small inconspicuous bracts which are deciduous 
before the flowers open or occasionally at least in part per- 
sistent; calyx-lobes 3 mm. long or usually less, rounded to 
sub-truncate or triangular, mostly as wide as, or wider than, 
long; branchlets various. 

Leaves 2.5-7 cm. long, narrowly elliptic or oblanceolate, mostly 
5-6 times as long as wide, blunt- tipped M. salicifolia 

Leaves broader, usually larger and not more than 3 times as 
long as wide, if narrow and elongate the tips acuminate or 
narrowly acute. 

Summit of the ovary, and interior of the prolonged and cup- 
like hypanthium, glabrous; fruit usually globose, 5-6 mm. 
in diameter. 



618 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Outer surface of the hypanthium glabrous, the branches of 
the inflorescence glabrous or sparingly pubescent. 

Leaves small, less than 3 cm. long, elliptic to obovate, 
rounded to blunt-pointed at tip, mostly cuneate at 
base; inflorescence 3-4 cm. long, often exceeding the 
leaves; flowers mostly 11 or fewer. . .M. myrtillifolia 

Leaves mostly 4-10 cm. long, elliptic or ovate, rarely 
obovate; inflorescence larger, if as little as 5 cm. 
long then equaling or shorter than the leaves; 
flowers usually more numerous. 

Pubescen