LIBRARY OF THE
UNIVERSITY OF ILLINOIS
TROPICAL AMERICAN MYRTACEAE
NOTES ON GENERIC CONCEPTS
AND DESCRIPTIONS OF
PREVIOUSLY UNRECOGNIZED SPECIES
VOLUME 29, NUMBER 3
CHICAGO NATURAL HISTORY MUSEUM
NOVEMBER 30, 1956
TROPICAL AMERICAN MYRTACEAE
NOTES ON GENERIC CONCEPTS
AND DESCRIPTIONS OF
PREVIOUSLY UNRECOGNIZED SPECIES
Professor of Botany, University of Michigan
VOLUME 29, NUMBER 3
CHICAGO NATURAL HISTORY MUSEUM
NOVEMBER 30, 1956
PRINTED IN THE UNITED STATES OF AMERICA
BY CHICAGO NATURAL HISTORY MUSEUM PRESS
Tropical American Myrtaceae
The following notes have been prepared as a general introduction
to a formal treatment of the Myrtaceae of Peru which is now in the
course of preparation. A large part of the present paper is devoted
to the characterization and description of approximately 80 species
and sub-specific taxa, mostly from Peru, which appear to be new to
science. The keys which are set forth below are intended primarily
for the student who wishes to place the newly described species
properly among their congeners or to follow the lines of reasoning
by which I became convinced that the specimens represented
undescribed taxa. Both keys and descriptions have been abridged
from those which were originally prepared for the Flora of Peru, so
that extra-Peruvian species have not been included in the keys
unless there seems reason to believe that ultimately they may be
found in that country.
The American representatives of the Myrtaceae have long been
considered a "difficult" group, and one in need of much systematic
study. This is partly because of the undoubtedly very large number
of species in the American tropics, and partly because of certain
features inherent in the plants themselves (e.g., flowers and fruits
of the same species are rarely obtainable at the same time). Dis-
tinctions between species and even between genera are sometimes
nice ones, for flowers as well as vegetative structures are relatively
uniform throughout the whole family. Identification of flowering
material is difficult for the casual student because distinctions
between genera have been made primarily on characters of the
mature embryo. Embryo characters are not always easy to observe
in dried material, even in specimens with mature fruit, and they are
impossible to make out from flowering specimens.
One of the principal impediments in the way of serious work on
the American Myrtaceae during the last century, paradoxically
enough, seems to have been the existence of two important and
indeed monumental treatises by Otto Karl Berg (1815-1866),
146 FIELDIANA: BOTANY, VOLUME 29
a Professor of Pharmacy at the University of Berlin. 1 Berg laid the
foundations for all later taxonomic work on this family in the New
World, but his monographs are difficult to use except after prolonged
study and collation. Neither one was provided with a workable key
to species, and the generic keys can be used for specimens in the
fruiting condition only. Berg's taxonomic philosophy led him to
believe that most species were sharply restricted in range to single
phytogeographic regions, which in turn increased inordinately the
number of species he described as new.
Probably the greatest obstacle confronting the would-be student
has been the vast number of species involved. According to Berg
himself, 2 the known American species in 1859 comprised a total of
1726, of which he personally had proposed 1008. The largest genera
were Eugenia (537 species), Aulomyrcia (240), Myrcia (184) and
Psidium (101). The published enumeration listed 696 species in
Brazil, with the greatest concentration in the southeastern part of
that country (390 species in Minas Gerais and Goiaz together, 265 in
Rio de Janeiro, and 175 in Parana and Sao Paulo). Only 161 species
were listed from the whole Amazonian region, and only 54 from
Peru and Bolivia together. Because of the large numbers of species
in certain genera and in certain areas, and because of the way in
which Berg organized his monographs, it is for any practical purpose
impossible to identify an unknown specimen unless one already has
some knowledge of the Myrtaceae, or unless the specimen is in good
fruiting condition and comes from an area whence few species are
Berg's monographs in spite of the difficulties attendant upon
their use were major contributions to systematic botany. His
major theoretical work was perhaps the clarification of the taxonomy
of the tribe Myrteae, which is discussed below. His generic concepts
were probably somewhat narrower than those of most modern
workers, but he was a keen and accurate observer. He recognized
as new a great many species which were indubitably distinct, and
a great many others which now appear to have been based upon
rather trivial characters. When one considers that many of the
1 Revisio Myrtacearum Americae, Linnaea 27: 1-472; index, op. cit. 786-795.
1855-1856 (also distributed as a separate, without index). Mantissa I. ad re-
visionem Myrtacearum Americae, op. cit. 29: 207-264. 1858. Mantissa II. ad
revisionem Myrtacearum Americae, op. cit. 30: 647-713. 1861. Mantissa III.
ad revisionem Myrtacearum Americae, op. cit. 31: 247-262. 71862. Myrtaceae,
in Mart. Fl. Bras. 14, pt. 1: 1-656, pis. 1-82. 1857-1859.
2 Conspectus distributions Myrtacearum Americae hue usque cognitarum,
Mart. Fl. Bras. 14, pt. 1: 619-622. 1859.
McVAUGH: TROPICAL AMERICAN MYRTACEAE 147
species recognized by Berg were known to him only through single
and often imperfect even sterile specimens, and that only rarely
was he able to see both flowers and fruit in the same species, it is
at once apparent that the quality of his work was extraordinarily
When the present work on the Myrtaceae of Peru was begun, it
was soon apparent that the number of species native within the
political boundaries of the country would approximate 150. This
suggested that a large number of the species were new to science, as
Berg had listed no more than 54 from Peru and Bolivia together, and
subsequent authors had added hardly more than a score of species
to the flora of Peru. Another alternative seemed possible, namely,
that many of the Peruvian species would prove to be identical with
others which had been described previously, by Berg or by others,
from extra- Peruvian material. It therefore seemed necessary to
undertake at least a cursory survey of all tropical American repre-
sentatives of the family, in order to establish and delimit the princi-
pal patterns of distribution.
In the course of this survey I have examined most of the material
which is available at the United States National Herbarium and at
Chicago Natural History Museum; the latter is very rich in South
American Myrtaceae and has many isotypes and type fragments
from the classical collections. I have also had the privilege of
studying selected specimens from the New York Botanical Garden;
the Harvard University herbarium; the Naturhistorisches Museum,
Vienna; and the herbarium of the Universidad Nacional Mayor de
San Marcos, Lima. With the aid of a grant from the Horace H.
Rackham School of Graduate Studies, University of Michigan, I
was enabled to spend the months of July and August, 1954, in the
study of these plants as they are represented in European herbaria.
Most of Berg's types, or duplicates of these, are preserved at Munich
(in the Brazilian herbarium of Martius), at Geneva (where are also
the types of the DeCandolle Prodromus), at Paris, or at Brussels.
Many isotypes are to be found at the British Museum (Natural
History), or at Kew. I was able to visit each of the above herbaria
at least for a short time, and I am especially grateful to the authorities
at Geneva, at Paris, and at the two British herbaria, for permission
to borrow certain critical material for further study. Although I
was unable to visit Vienna, Dr. K. H. Rechinger very kindly located
for me, and forwarded to me for study, most of the unique specimens
which Berg had described from among the collections there. A few
of Berg's types which were at Berlin, and apparently not represented
148 FIELDIANA: BOTANY, VOLUME 29
by duplicates elsewhere, were presumably destroyed in World
The following generalizations relative to range patterns in tropi-
cal American Myrtaceae are presented as tentative, after study of
the materials discussed above; it is thought that future work will not
markedly affect the principal conclusions, but concepts of individual
species may be radically changed after study of the groups to which
they belong, and names applied to Peruvian species may on this
account have to be changed in some instances.
Distribution of South American Myrtaceae
1. The Chilean Myrtaceae are all endemic, and none of the
species ranges as far north as Peru. This is not unexpected, in view
of the peculiar isolation which Chile enjoys, and it is mentioned here
chiefly because Berg wrongly attributed a number of Chilean species
to Peru, and Peruvian species to Chile. This error came about
because of lack of sufficient knowledge of the itineraries of certain
collectors, particularly Dombey and Poeppig, who visited both
countries. The Myrtaceae of Chile are probably better known than
those of any other area of comparable extent in South America,
thanks chiefly to the efforts of Eberhard Kausel of Santiago, who
has published numerous papers on the subject. 1
2. A majority of the species of southern Brazil (i.e., from south
of an arbitrary line drawn at about 15 S. Lat., and including all of
Brazil south of Minas Gerais, southern Goiaz and southern Mato
Grosso) do not extend northward into the Amazon basin and are
not likely to be found in Peru.
3. Species of the Bolivian lowlands, even the lowlands of
northern Bolivia, have little in common with most Peruvian species
but show strong affinities with species ranging from southern Brazil
to Uruguay and northern Argentina. The line between Bolivian
and Peruvian lowlands appears to be crossed by relatively few
4. Species from the northeastern states of Brazil (e.g., Bahia,
Piauhy, and Ceard) are often distinct from those of southern Brazil
1 Cf. Contribution al estudio de las Mirtaceas Chilenas, Rev. Argent. Agron.
9: 39-68. 1942; op. cit. 221-243. 1942; op. cit. 11: 320-327. "1944" (1945); Notas
mirto!6gicas, Lilloa 13: 125-149. "1947" (1948). The first two papers comprise
a revision of the Chilean genera and species; the third is in the form of a supple-
ment incorporating some new information. The final paper presents miscel-
laneous new observations and is concluded with a new key to the Chilean genera.
McVAUGH: TROPICAL AMERICAN MYRTACEAE 149
and usually seem distinct from those of the Amazon region but may
show affinities with both. This area is much in need of further
5. Some species of the Guiana lowlands, and most species of the
Amazon lowlands, seem potentially wide-ranging and should be
considered in any treatment of the Peruvian flora. At least a few,
and probably more, species certainly range widely from the West
Indies through the Guianas and most of lowland Brazil, Peru
6. Most of the myrtaceous species found by early collectors
on the upper Amazon and its tributaries, as far down as Ega (Teffe"),
have since been re-collected in Peru. A large number of distinctive
species, however, have been collected along the Rio Negro and other
rivers which drain into the Amazon from the north but have not
been collected along the upper reaches of the Amazon itself. It
appears that many species which range northward into Venezuela
in the Rio Negro drainage, although they may reach the Amazon
near the mouth of the Rio Negro, do not extend far up the Amazon
and are not to be expected in Peru.
7. Plants of the inter-Andean valleys, and from high elevations
up to near the limits of vegetation, seem in Peru to be mostly
endemic, although the Peruvian species are often very similar to
those of the more northern parts of the Andes.
In view of the above conclusions I have, in describing the follow-
ing novelties from Peru and elsewhere, considered in the most careful
detail all the species previously described from the entire Amazon
basin ; I have paid close attention to species described from northern
South America, and more than casual attention to species described
from eastern and southern Brazil. Further taxonomic studies of
the very numerous published species from these latter areas, how-
ever, may bring about changes in concepts and in nomenclature
which will affect some Peruvian species also. It is further possible
that a few Peruvian species will prove to be identical with some
of those described by Linnaeus, Jacquin, Lamarck, and other early
One of Berg's principal contributions to the taxonomy of the
Myrtaceae was the documentation of the fact that the tribe Myrteae,
including practically all the American members of the family, can be
150 FIELDIANA: BOTANY, VOLUME 29
divided into three coherent groups (subtribes) based on characters
of the embryo. This had been discerned much earlier by DeCandolle,
but the specimens available to that author were much too few in
number to enable him to make the general conclusions which were
set forth by Berg. Berg recognized three subtribes, which he called
Myrcioideae, Eugenioideae, and Pimentoideae, and which included,
respectively, 11, 12, and 17 genera. 1
More recent workers have not always recognized all the genera
admitted by Berg. Bentham (Benth. & Hook, f . Gen. PL 1 : 712-720.
1865) reduced the numbers of genera in the subtribes to 3, 3, and 7,
respectively. Niedenzu (Nattirl. Pflanzenfam. Ill, pt. 7: 64-86.
1893) reversed the trend and admitted 5, 4, and 13 genera, re-
spectively. Conservative modern opinion would agree upon the
recognition of at least the following generic groups (Chilean genera
not included) :
1. MYRCIINAE. Cotyledons foliaceous, contortuplicate; radicle
(a) Calyptranthes Sw. Buds closed, calyx calyptrate.
(b) Marlierea Camb. Calyx closed or barely open in bud, split-
ting irregularly in anthesis.
(c) Myrcia DC. Calyx with 5 (rarely 4) distinct lobes.
Other genera proposed by Berg depend upon single characters
of the anthers (Cerqueiria, Gomidesia) or of the calyx (Aulomyrcia,
Calycampe, Calyptromyrcia, Eugeniopsis, Rubachia). None of these
has been much studied or widely accepted except Aulomyrcia, which
has been taken up in recent years by Miss Amshoff in her studies
of the Myrtaceae of northern South America. The genus Myrc-
eugenia Berg, with a disjunct range in Chile and Uruguay, is
apparently a distinct group. The others mentioned above, however,
are but doubtfully distinct from Myrcia. Most of the species are
south-Brazilian, and proper elucidation of the genera must wait
upon taxonomic studies in this region.
1 According to the International Code (Article 29), these subtribes are properly
known as Myrciinae Berg, Eugeniinae Berg, and Pimentinae Berg. The pertinent
rule states: "When the name of a taxon . . . has been published with an improper
termination . . . the ending must be changed to accord with the rule, without
change of authority." The names of these three subtribes were first published
with the proper termination by Niedenzu (Natiirl. Pflanzenfam. Ill, pt. 7: 62.
1893) but that author substituted Myrtinae for Pimentinae, contrary to the
provisions of Article 29. The names Myrciinae and Eugeniinae have usually been
attributed to Niedenzu, e.g., in Dalla Torre & Harms (Gen. Siphon. 348. 1903),
but are properly attributed to Berg.
McVAUGH: TROPICAL AMERICAN MYRTACEAE 151
2. EUGENIINAE. Cotyledons fleshy, distinct or conferruminate
or the embryo undivided ; radicle very short.
(a) Eugenia L. Calyx-lobes usually 4, distinct; hypanthium not
prolonged above the summit of the ovary; ovules numerous.
(b) Calyeorectes Berg. Buds closed, irregularly splitting; hy-
panthium prolonged; ovules numerous.
(c) Myrciaria Berg. Calyx-lobes 4; hypanthium prolonged, cir-
cumscissile at base; ovules 2 in each locule.
Certain additional genera, not recognized by Berg, have been
accepted by Amshoff (e.g., in Fl. Suriname 3: 56-158. 1951). These
(d) Plinia L. Buds closed or nearly so, the calyx splitting
irregularly; flowers glomerate, involucrate; hypanthium pro-
longed; ovules 2 in each locule.
(e) Catinga Aubl. Calyx-lobes 4, thickened distally and coher-
ent in the bud; hypanthium not prolonged; ovules numerous.
Of the remaining genera accepted by Berg, none has been widely
accepted. Two of these (Phyllocalyx, Stenocalyx) depend upon
a character of the inflorescence which is easily recognized but of
doubtful importance. Myrcianthes, according to Bentham (op. cit.
715), was wrongly placed in the Eugeniinae and belongs rather with
Myrtus; and Mitranthes, according to Bentham (I.e. 717) belongs
with Calyptranthes in the Myrciinae. Schizocalyx, Siphoneugena and
Hexachlamys are based on calyx characters which scarcely serve
to distinguish the first from Calyeorectes and the last two from
Eugenia. Acca certainly belongs in the Pimentinae, and is treated
below. In recent years it has been shown that Aulacocarpus prob-
ably belongs to the Melastomaceae.
3. PIMENTINAE. Embryo spiral or curved; radicle elongate;
cotyledons very short.
(a) Campomanesia R. & P. Ovary usually 6- to many-locular;
calyx-lobes 5, membranaceous; embryo spirally involute, the
testa membranaceous; ovules 2- or rarely 4-seriate.
(b) Psidium L. Ovary 2- to many-locular; calyx closed in the
bud or with 5 short lobes; embryo circinate; ovules many-
152 FIELDIANA: BOTANY, VOLUME 29
(c) Calycolpus Berg. Ovary usually 4- to 5-locular; calyx-lobes
5, often foliaceous, spreading or reflexed in bud; embryo
(d) Pimenta Lindl. Ovary bilocular; calyx-lobes 4 or 5; embryo
cyclic or sub-spiral; ovules 1-6, pendulous; inflorescence
(e) Ugni Turcz. Calyx-lobes 4 or 5; embryo arcuate; peduncles
1-flowered; anthers sagittate, the connective dilated.
Cf) Myrteola Berg. Calyx-lobes usually 4; embryo arcuate;
peduncles 1-flowered; anthers subrotund, the filaments
(g) Blepharocalyx Berg. Calyx-lobes 4, ciliate, deciduous;
embryo sub-spiral or spiral, the testa membranaceous;
peduncles dichotomous, mostly 3- or more-flowered.
The rest of the genera accepted by Berg have been little studied.
The monotypic Paivaea differs from Campomanesia chiefly in the
patelliform-dilated base of the young calyx. Also monotypic are
Calyptropsidium and Psidiopsis, which differ from Psidium each
by a single character of the calyx and which are difficult to defend
as independent genera. Pseudocaryophyllus was scarcely known
to Berg except from descriptions and illustrations and was founded
at least in part upon a misunderstanding of the morphology of one
species. Amomis has been placed by many authors, including
Bentham and Niedenzu, in the synonymy of Pimenta. Acrandra
appears to be distinct by virtue of its apiculate anthers. Britoa,
Abbevillea and Lacerdaea were placed by Bentham in the synonymy
of Campomanesia, from which they differ by calyx characters of
The genus Myrtus is not included in the above list. Berg as-
signed 34 American species to Myrtus, in addition to 5 species about
which he had some reservations. At least half the species included
by Berg in Myrtus have been shown to belong to other genera, and
the rest have not been critically studied. This matter will be dis-
cussed below, with reference to the Peruvian genera in particular.
The Myrtaceous Inflorescence
Students of the Myrtaceae have to a considerable degree passed
over the inflorescence in their search for characters which may be
used in the delimitation of taxa. The subject of inflorescence
McVAUGH: TROPICAL AMERICAN MYRTACEAE 153
morphology was indeed discussed at some length by Bentham in an
early paper 1 and before him by Grisebach, but it has been little
stressed by more recent workers. It is therefore thought to be
worth while to discuss the myrtaceous inflorescence here, and to
follow this discussion by a summary of the features which seem to
be of taxonomic significance.
Most of the species of the Myrciinae can be recognized as such,
even in the absence of both flowers and fruit, from the inflorescence
alone; it is even possible to make many identifications to genera
on this basis. Most of the American genera of the Eugeniinae and
Pimentinae can also be recognized by a combination of inflorescence
characters with calyx characters which are apparent either in
flowering or in fruiting material. Various markedly different in-
florescences occur among the American Myrtaceae, but the differ-
ences are, at least to some extent, superficial. All known types are
derivable from a single basic pattern. Branching patterns of
vegetative stems are fundamentally similar to those of flowering and
fruiting stems, and an understanding of each is essential to an
understanding of the morphology of the inflorescence.
In the tribe Myrteae the leaves and branches are normally
decussate; that is, in pairs at the nodes, with those at any one node
standing at right angles to those at the nodes above and below it,
and the leaves at alternate nodes standing directly above or below
one another. This arrangement prevails not only in simple (un-
branched) axes, but also in branching systems. Branching at the
lowest node of an axillary branch, for example, will occur in the
plane of the tangent at right angles to the plane of branching at the
node from which the axillary branch arises. The sequence is readily
observed in leafy branch-systems, in bracteate inflorescences devoid
of ordinary foliar structures, and in transitions from vegetative to
reproductive branches. The inflorescence is thus interpreted as an
axillary branch of which the primary branchlets are normally
decussate, exactly in the manner of sterile branches. Deviations
from the usual pattern, e.g., the alternate branching sometimes
seen in Myrcia near the tips of the panicles, an occasional unilateral
panicle in Calyptranthes, or occasional solitary flowers in normally
racemosely flowered species of Eugenia, are assumed to represent
reduced or derived conditions.
It is sometimes difficult to recognize the proper limits of a single
inflorescence. In some genera, e.g., Myrcia, almost any new branch
1 Notes on Myrtaceae, Journ. Linn. Soc. Bot. 10: 101-166. 1869.
FIELDIANA: BOTANY, VOLUME 29
FIG. 1. Decussate leaves and branches in the Myrtaceae. Large circles
represent the principal branch; small circles, subsidiary branching axes (i.e.,
a leafy branch, the axis of a raceme, or a pedicel); separate lenticular figures,
petioles of ordinary leaves; angular lines, bracts. Bracts and bracteoles are in
arising from the axil of a foliage leaf may represent a potential
inflorescence. At the one extreme the whole branch may be modified
in the direction of flower production, all its leaves reduced to minute
deciduous bracts and its secondary branchlets irregularly disposed;
such a branch is readily defined as a single inflorescence. The same
new axillary branch, on the other hand, may bear normal leaves
at most of the nodes and miniature inflorescences from certain axils
McVAUGH: TROPICAL AMERICAN MYRTACEAE
only (e.g., the lower ones). The partly leafy branch is the homo-
logue of the completely fertile branch; what, then, represents the
inflorescence of this species? In practice this can usually be deter-
mined by correlation with leaves of a certain size, i.e., those occur-
ring on permanent mature twigs; leaves are often reduced in size
when occurring on temporary (inflorescence) branches.
FIG. 2. Schematic representation of flowering in Eugenia. Leafless raceme
from the lower node; partially fertile branch from the upper node. Bracts and
bracteoles in solid black. See description in text.
In other genera, e.g., in Eugenia, the flowers may occur in
a short leafless raceme (the whole subtended by a foliage leaf), or
in the same species they may be solitary at the lowest (leafless)
156 FIELDIANA: BOTANY, VOLUME 29
nodes of a new branch which bears normally developed foliage
leaves at the upper nodes. In the first case it may be assumed that
the fertile branch failed to develop beyond the lower nodes; in the
other case that the axis continued to grow and produce leaves. The
short leafless raceme is the homologue of the leafy branch with
flowers at the lowest (and leafless) nodes, but it is difficult to re-
gard the latter as an "inflorescence."
Finally, it should be stated that the primitive myrtaceous in-
florescence is assumed to be one in which the primary axis, and each
of the subordinate axes, is terminated by a flower.
The inflorescence (or to put it perhaps more precisely, the
potentially floriferous branch which develops from the axil of a leaf)
may be modified in aspect and morphology in one or more of the
1. Reduction of leaves to small bracts which may be colored or
scarious, persistent or deciduous.
2. Abortion of the terminal bud of the primary axis or those
of the secondary axes.
3. Reduction in length of the primary axis, with or without
a corresponding reduction in the number of nodes.
4. Termination of the primary axis by the formation of a flower
at the first node (inflorescence a dichasium) or at one of the suc-
5. Reduction of secondary axes to 1-flowered bracteolate pe-
duncles (inflorescence a raceme).
6. Dehiscence of secondary axes from one or both sides of
a node, with the production of irregularly branched panicles.
7. The regular reduction, from the base to the apex of the
inflorescence, of internode length and number of nodes in all sub-
In the following paragraphs are described the principal modi-
fications which result from combinations of the above:
1. The raceme. This type is characteristic of the subtribe
Eugeniinae and occurs occasionally in genera of the Pimentinae.
The raceme is an axillary and usually leafless inflorescence with
a single primary axis and several or many nodes; the axis is theoreti-
cally indeterminate but usually aborts at the tip and lacks a terminal
flower; the internodes may be elongated or much abbreviated; the
McVAUGH: TROPICAL AMERICAN MYRTACEAE
branches from the axis are in decussate pairs, all about the same
length and reduced to bracteolate "pedicels" terminated each by
a solitary flower. In exceptional cases the racemes are compound,
e.g., in Eugenia florida, in which the individual flowers of the
raceme are wholly or partly replaced by small racemosely flowered
branches, or in E. stipitata, in which each peduncle may bear a simple
dichasium instead of a single flower.
FIG. 3. Modified racemose inflorescences in Eugenia. Left, E. stipitata
subsp. sororia: the fertile branch is a short raceme in which some of the peduncles
are 3-flowered. Right, E. punicifolia, in which the flowers are reduced to a single
pair in each raceme. Bracts and bracteoles in solid black.
When the axis of the raceme is much reduced in length and the
nodes are approximate, the number of nodes may be correspondingly
reduced and the flowers may appear to be umbellate or glomerate.
The most extreme reduction occurs in species like Eugenia puni-
cifolia, in which the flowers are a single pair in each axil, one arising
from each side of the abortive axillary bud.
In some species of Eugenia, as mentioned above, a new leafy
branch may have the lowest two or three nodes fertile and bracteate
rather than leafy, but the upper nodes sterile and normally leafy;
in such species the flowers may thus appear to be solitary at leafless
nodes. In the same species, from other leafy nodes, there may arise
abbreviated racemes in which the leaves are all reduced to small
bracts and the axis is abortive.
A similar situation prevails in Campomanesia lineatifolia (Pimen-
tinae), a species in which the flowers usually occur at the two lowest
158 FIELDIANA: BOTANY, VOLUME 29
nodes of an otherwise leafy branch but in which the branch may be
reduced to a short leafless axis comprising only the two fertile nodes.
In any inflorescence of the racemose type each pedicel is sub-
tended by a bract. If the number of flowers in a raceme is reduced
to one, as may rarely happen, it is still ordinarily possible to observe
the pedicellar bract, or at least the bract-scar, and usually possible
in addition to observe the abortive terminal bud of the inflorescence.
In species which normally bear solitary flowers in the axils of
ordinary leaves (see discussion below), the pedicel bears no bract
at base and the flower itself represents the termination of the axis.
2. The myrcioid panicle. With few exceptions this type is
found in the principal genera of the Myrciinae, with some modi-
fications as noted below. The axillary panicle is compound, or
decompound, with the primary axis and each of the subsidiary
branches terminating in a flower. The principal branches are op-
posite and decussate, the lower ones elongate and usually themselves
compound and many-flowered, and the ultimate branchlets of the
panicle usually with the flowers aggregated in threes (i.e., in simple
dichasia) near the tips. The transition between elongate basal
branches and simple dichasia at the tips is accomplished by
gradual reduction, in successive branches, of the number of nodes
and the length of the internodes. Branching toward the tips of the
panicle may be irregular because of abortion of one or both buds
at a given node, or because of the dehiscence (at the point of origin)
of one or both branchlets after the initiation of growth; either of
these processes may result in, or be accompanied by, distortion or
zigzag growth of the axis which remains.
Because of the irregular development of the myrcioid panicle
it is usually not practicable to state the usual number of flowers
in the panicle of a given species, except within broad limits. A
large panicle may contain 300 flowers or even more; a small panicle
in the same species may contain no more than 50. In some species
the number of flowers may be reduced to 3, 2, or even one. On
the whole it is a fair generalization that a "many-flowered" species
will probably prove upon examination to be a member of the
Myrciinae, whereas among the Eugeniinae an inflorescence with as
many as 50 flowers is a rarity.
Usually the myrcioid panicle can be distinguished at a glance
from a compound racemosely branched panicle (which may be
"many-flowered," as in Eugenia florida) by the long lower branches.
In a racemosely branched panicle the pedicels and the internodes
FIG. 4. Diagram of the myrcioid panicle. Position of flowers and branches
indicated to the extent necessary to make clear the pattern of branching which
is repeated in each part of the panicle. Branches in one plane are shown; those
at right angles to this plane are omitted, but their positions are indicated by
160 FIELDIANA: BOTANY, VOLUME 29
are essentially of uniform length, so that the flowers appear uni-
formly distributed in the inflorescence. In the myrcioid panicle,
especially after anthesis, the observer receives the general impression
of numerous elongated slender branches which bear short-stalked
or sessile flowers or flower clusters at irregular intervals, and par-
ticularly near their tips.
Among the myrcioid genera in our flora it is possible to recognize
two tendencies in the branching of the inflorescence. In the genus
Myrcia, and in the segregate genera Aulomyrcia and Gomidesia, the
principal axis of the inflorescence is as long as, or a little longer than,
the primary lateral branches; the panicle, as a result of this, is about
as long as wide. In Marlierea and Calyptranthes the principal axis
may abort at the node where the lowest lateral branches emerge, so
that the inflorescence as a whole appears to consist of paired spikes
or panicles arising from the same axil. This condition is found
regularly in some species, occasionally in others, and rarely or
never in a third group. It is thus by no means consistent in the
genera in which it occurs, but as far as I am aware it never occurs in
3. The dichasium. This type characterizes the Pimentinae and
is found in certain genera of the Eugeniinae (e.g., Anamomis Griseb.).
The primary axillary axis is immediately determinate; it terminates
in a flower at the first node. The flowers may be thus normally
solitary and subtended by normal foliage leaves. Invariably the
flower is subtended by a pair of opposite bracteoles which indicate
the potentiality of lateral branching at these points, and in various
species solitary flowers are often found on the same plant with
3- or 7-flowered dichasia.
Branching of the primary axis may take place at the node just
beneath the terminal flower (i.e., from the axils of the subtending
bracteoles), with the production of a simple (3-flowered) dichasium.
By further symmetrical development of the lateral branches, the
inflorescence may become a 7-, 15- or rarely 31-flowered dichasium.
In the forks of a compressed 15- or 31-flowered dichasium the ter-
minal flowers may fail to develop to maturity.
In most species with predominantly solitary flowers or few-
flowered dichasia, a majority of the mature leaves bear flowers in
their axils, and there appears to be no marked restriction of fertility
to the lower nodes of a shoot. When solitary flowers and dichasia
occur on the same plant, the most vigorous nodes and leaves appear
McVAUGH: TROPICAL AMERICAN MYRTACEAE 161
FIG. 5. Details of branching at the tips of the myrcioid panicle. Left,
a cluster of three nearly sessile buds forming a "false dichasium," in which the
lateral flowers arise from the second node below the terminal flower; the buds
at the first node are abortive. Right, another species, in which the lateral flowers
of the dichasium are borne on slender pedicel-like branches from the node im-
mediately below the terminal flower.
FIG. 6. A dichasium in Eugenia. In some species of Eugenia and Psidium,
solitary flowers and 3-flowered dichasia may be found on the same plant.
to be correlated with inflorescences bearing the larger numbers of
flowers. In certain species of Psidium, and perhaps in some species
of Eugenia also, the dichasia may arise from leafy axils, or in the
same species from leafless nodes at the bases of leafy shoots; this
condition is apparently less frequent among species in which the
flowers are borne in dichasia than among the racemose-flowered
162 FIELDIANA: BOTANY, VOLUME 29
The Myrtaceous Leaf
The keys and descriptions which follow include some terms
which are used repeatedly in certain restricted senses; the following
notes describe the ways in which these terms apply specifically to
the Peruvian Myrtaceae:
The leaves of most American Myrtaceae have a straight midvein
which extends the entire length of the blade; the leaves are pinnately
veined, and the midvein is very markedly thicker than any of the
lateral veins. The midvein almost without exception is conspicuous
on the lower (dorsal) surface of the leaf, and considerably raised
above the surface as a convex line; in some species it is elevated to
such an extent that its diameter at right angles to the leaf is as great
as the diameter in the plane of the leaf. The midvein on the upper
(ventral) surface of the leaf may be conspicuous and somewhat
elevated or at the other extreme nearly invisible. The vein proper
may be convex, flat, somewhat concave, or with a narrow groove
or channel running the entire length; in a few cases the midline of
the vein may be elevated in a narrow ridge. In addition to the above
modifications in the surface of the vein proper, the whole vein may
be impressed; that is, depressed in a trough or furrow below the
general surface of the leaf. In general, therefore, to say that a vein
is impressed refers to the position of the whole vein in relation to the
rest of the leaf, not to any modification of the vein itself.
The lateral veins arise at intervals from the midvein and extend
toward the margin, forming usually an acute angle with the distal
portion of the midvein. The lateral veins are said to occur in pairs,
but this is something of a misnomer, because the veins on opposing
sides of the midvein are more often alternate than opposite. The
word "pairs" is used to avoid a longer and more clumsy expression;
i.e., it is preferable to say "12 pairs" than "12 on each side of the
midvein." The lateral veins are often nearly unbranched, that is,
they lack major branches and appear to extend without inter-
ruption from the midvein to the margin or near it; in some species
the lateral veins fork in the distal half or the distal third. The
secondary veins which connect the laterals are often very small and
appear to the eye as a small network between larger veins.
As a lateral vein approaches the leaf margin it invariably curves
toward the apex of the leaf and joins the next succeeding vein. In
this way are produced various kinds of marginal veins which are
approximately parallel to the leaf margin. In some species the
McVAUGH: TROPICAL AMERICAN MYRTACEAE 163
lateral veins diminish gradually in thickness from base to apex and
recurve markedly toward the midvein before joining the succeeding
laterals. In this case no marginal vein, as such, can be distinguished,
and in extreme cases the lateral veins become so much attenuated
toward the tips that one can with difficulty make out the points of
union with the succeeding veins. In other species the lateral veins
are straight and of uniform diameter, but their distal ends are
connected by a series of arches or loops which together comprise
a marginal vein; this arched marginal vein may be thinner than the
laterals, but in many species its diameter is as great as that of the
laterals. In the species which are thought to present the most
highly evolved vein pattern, the marginal vein is a characteristic
and conspicuous feature of the leaf, arching almost not at all between
the lateral veins and thus nearly straight except that it follows the
curve of the leaf margin. When the marginal vein is of this kind,
the laterals may extend directly from midvein to marginal vein and
enter the latter almost at right angles, with little or no apparent
tendency to turn toward the apex of the leaf.
The number of pairs of lateral veins is used in various instances
to distinguish species. The number is constant within rather broad
limits only, and the actual number present in a given leaf is often
doubtful. A precise count can often be made in species which have
strong rib-like veins and few or no intermediate veins, but such
species are in the minority. Far more abundant are species in which
the major pairs of veins are but little stronger than the numerous
slender intermediates which may curve or branch or join another
vein. In doubtful cases the difficulty can often be resolved by
counting the number of arches in the marginal vein; as a general
rule these arches extend in unbroken curves from one major lateral
vein to the next, and small intermediate veins cause little or no
break in the smooth curve of the arch. At best the numbers of veins
as stated are mere approximations.
The Genera of Myrtaceae Which Occur in Peru
The following key is intended as an indication of the taxonomic
arrangement which is suggested by consideration of the morpho-
logical features of these plants. The key will not "work" for species
which are in one way or another anomalous, e.g., for species of
Myrcia with 4 calyx-lobes. It will suffice, however, to indicate gen-
eric affinities for a great majority of all species in the Peruvian flora.
164 FIELDIANA: BOTANY, VOLUME 29
1. Inflorescence myrcioid (i.e., compound, many-flowered, with the lower
branches opposite and elongate; the upper branches progressively shorter
and closer together; the uppermost often alternate because of abortion of one
of a pair; the flowers tending to be aggregated toward the tips of the branch-
lets and often solitary or in simple dichasia at the tips); calyx-lobes, if
developed, usually 5 (Myrciinae).
2. Calyx-lobes 5 (rarely 4), distinct even in the bud; principal axis of the panicle
well developed, about as long as the primary lateral branches Myrcia.
2. Calyx closed in bud, or only the tips of the lobes free, in anthesis circum-
scissile or splitting irregularly; principal axis of the panicle often abortive
at the lowest node and the panicle bifurcate.
3. Calyx closed in bud, circumscissile, the calyptra often remaining attached
at one side Calyptranthes.
3. Calyx closed in bud, or the tips of the (usually 5) calyx-lobes free, the flower
opening by irregular longitudinal slits between the lobes Marlierea.
1. Inflorescence bifurcate (usually with sessile flowers in the forks), or racemose
(the flowers or branches all in decussate pairs on an unbranched axis which
aborts at tip), or the flowers solitary.
4. Inflorescence racemose (species with glomerate or subumbellate flowers
should be sought here; inspection will usually show that these flowers are in
approximate, decussate pairs on a much shortened axis); calyx-lobes 4 and
distinct, or rarely the buds closed or merely the tips of the lobes free (includes
most of Eugeniinae).
5. Hypanthium prolonged into a tube above the summit of the ovary; flowers
mostly 4, sessile or nearly so, in conspicuously bracteate clusters; ovules
2 in each locule.
6. Calyx-lobes distinct, the tube circumscissile at base Myrciaria.
6. Calyx splitting irregularly and longitudinally in anthesis, the bud completely
closed or the tips of the calyx-lobes free Plinia.
5. Hypanthium not prolonged above the ovary; calyx-lobes distinct; flowers
few or many, rarely conspicuously bracteate (the bracteoles subtending in-
dividual flowers often persistent and evident) ; ovules usually numerous.
4. Inflorescence dichotomously branched, or the flowers solitary.
7. Inflorescence dichotomously branched.
8. Flowers 4-merous; calyx-lobes distinct and imbricate.
9. Flowers often on old leafless branches, in regularly thrice-dichotomous cymes
with a sessile flower in each fork; bracts persistent, in cuplike pairs; calyx-
lobes persistent; flowers red, the stamens and style much longer (6-15 times)
than the hypanthium Myrrhinium.
9. Inflorescence axillary, on new leafy branches; dichasia usually 3- or 7-
flowered, the forking branches variable in number and length; bracts usually
deciduous; flowers white or yellowish, the stamens and style proportionately
10. Calyx-lobes deciduous Blepharocalyx.
10. Calyx-lobes persistent Eugenia.
8. Flowers 5-merous; calyx splitting irregularly in anthesis, the buds closed or
the lobes short and widely separated Psidium.
7. Flowers solitary.
11. Calyx-lobes 4.
12. Small-leaved shrubs (leaves less than 1 cm. long) of the high Andes; bracteoles
foliaceous, elongate and persistent; anthers subrotund, the filaments filiform.
McVAUGH: TROPICAL AMERICAN MYRTACEAE 165
12. Shrubs or trees with larger leaves, of middle and low elevations; bracteoles
mostly deciduous, never persistent and foliaceous Eugenia.
11. Calyx-lobes 5, or the calyx opening irregularly.
13. Shrubs with small coriaceous leaves mostly less than 2 cm. long; high-
montane Andean species; calyx-lobes 5, distinct; peduncles all 1-flowered;
bracteoles foliaceous, elongate and persistent; anthers sagittate, the con-
nective dilated Ugni.
13. Shrubs of middle and low elevations, with larger leaves; flowers solitary or
in dichasia, the bracteoles never foliaceous and persistent; filaments filiform,
the anthers not sagittate.
14. Calyx-lobes distinct, membranaceous, spreading in flower and in fruit;
leaves with 3-4 pairs of the lowest lateral veins closely grouped and the upper
increasingly distant Campomanesia.
14. Calyx closed in the bud, or with short appressed lobes, the tube splitting in
anthesis along irregular longitudinal lines; veins nearly uniformly spaced.
One Peruvian genus, Acca, has purposely been omitted from the
above key because its inclusion would complicate the key and make
it more difficult of comprehension. One species of Acca has
the flowers solitary or in simple dichasia; the other species has the
flowers solitary or in congested racemes. The calyx-lobes are 4 in
number in each species, and so are anomalous in the Pimentinae.
The genus is indeed of uncertain systematic position but is easily
recognized by the villous or tomentose foliage and the large red
Notes on Individual Peruvian Genera
The following conclusions have been reached after study of the
criteria used by earlier workers, as set forth briefly above, and after
the evaluation of these criteria in the light of new data, particularly
from the inflorescence. Study of the vast myrtaceous flora of
southeastern Brazil may demonstrate the wisdom of taking a some-
what less provincial point of view with respect to the submergence
of such genera as Abbevillea, Calyptromyrcia, Gomidesia, Rubachia,
Siphoneugena, and others. The notes below are based primarily on
materials from the Amazon basin and the northern Andes and should
be considered with this in mind.
1. The myrcioid genera. It seems clear that the Myrciinae are
represented in Peru by three genera only. I cannot agree with Miss
Amshoff that Aulomyrcia is a valid genus, even though it is possible
to recognize certain species as members of the "genus." The
characters of Aulomyrcia, which are readily observed in a series of
small-flowered species typified by A. multiflora (Lam.) Berg, are
two only: the hypanthium is prolonged above the summit of the
ovary but is not constricted beneath the calyx; the disc and the
166 FIELDIANA: BOTANY, VOLUME 29
inner surface of the hypanthium are usually glabrous. Among
the Peruvian species, in my opinion, the distinction between Myrcia
and Aulomyrcia breaks down completely; depending upon the species
selected, one can demonstrate almost any desired stage of the tran-
sition between the glabrous and prolonged hypanthium and calyx
of Aulomyrcia on the one hand and the hairy ovary and short calyx
of Myrcia proper.
The status of Rubachia is not entirely clear and must be clarified
by study of the several Brazilian species. The one Peruvian species
referred by Berg to this genus is known in immature flowering
condition only, and its relationships appear to be with other Peru-
vian species of Myrcia. The genus Gomidesia, as far as I am aware,
does not occur within or near the borders of Peru, although it has
been reported from Bolivia; like Rubachia and Blepharocalyx, it is
primarily a genus of southeastern Brazil.
The distinction between Myrcia and Marlierea is a somewhat
nebulous one, and it is probable that Marlierea comprises a phylo-
genetically diverse assemblage of species which have been arbitrarily
assigned to the genus because of the character of the irregularly
splitting calyx. In spite of this doubt as to the monophyletic origin
of Marlierea, its species are often recognizable as such by the per-
sistent bracts and the abortion of the primary axis of the inflo-
rescence. On the basis of the somewhat weak association of these
characters of calyx, bracts and inflorescence, it seems best for the
present to recognize the genus as distinct.
Calyptranthes, like Marlierea, shows some evidence of internal
diversity, but the calyx-character is such a distinctive one, and is
correlated in so many species with the presence of dibranchiate
hairs, that the genus is one of the most readily recognized among the
2. The eugenioid genera. The genera of the Eugeniinae are
somewhat difficult to delimit, partly because of the paucity of
material available for study and partly because of the many and
morphologically diverse species. Whereas in the myrcioid groups
the morphology of the inflorescence, the structure of the embryo,
and the number of ovules in each locule remain essentially constant
from genus to genus and even from species to species, in the eugen-
ioid taxa quite the reverse is true. The number of species is large,
and most of them have been assigned at one time or another to the
genus Eugenia; any attempt at orderly classification of the whole
group necessitates a decision as to which taxa shall be removed
McVAUGH: TROPICAL AMERICAN MYRTACEAE 167
from the inclusive Eugenia, and as to the ultimate circumscription
of Eugenia itself.
The satisfactory partition of Eugenia on a worldwide basis has
not yet been accomplished. It was thought at one time that the
New World species could be separated from those of the Old World
on the basis of floral characters. Merrill and Perry, 1 in their studies
of Asiatic Myrtaceae, pointed out that such a separation was not
practicable, but indicated that in their opinion the American species
(Eugenia sens, str.) could be effectively distinguished from the
Asiatic (the genus Syzygium Gaertn.) as follows:
Eugenia: Embryo apparently undivided, pseudomonocotyledon-
ous; seed-coat smooth and free from the pericarp; inflorescence
centripetal, with the pedicels 1-flowered; calyx-limb less prolonged.
Syzygium: Cotyledons distinct; seed-coat roughish, loosely or
closely adhering to the pericarp; inflorescence usually centrifugal,
with the panicles branching by threes or with secondary cymes;
calyx-limb more prolonged.
An opposing point of view has been taken more recently by
Henderson 2 in his study of Eugenia (sens, lat.) in Malaya. In this
author's opinion little dependence can be placed on the seed-
characters to which Merrill and Perry attached primary importance.
Henderson maintains Eugenia as a single worldwide genus to include
not only Syzygium (which contains the great majority of the Old
World species) but also Acmena DC. and Cleistocalyx Blume, both
of which were recognized by Merrill and Perry as independent
genera. Henderson also suggests tentatively that "if the Old and
New World species are to be separated, better characters might be
found in the structure of the inflorescences and flowers."
Ingle and Dadswell, 3 in a paper on the wood anatomy of the
Myrtaceae, support on anatomical grounds the view that at least two
genera are probably represented in the inclusive Eugenia. A taxon
which these authors call "Eugenia B" (that is, including Acmena,
Cleistocalyx, Syzygium, and some Asiatic and Pacific species still
referred to Eugenia) is shown to differ fundamentally from "Eugenia
A" (that is, the New World species, plus the few Old World species
referred to the genus Jossinia). Ingle and Dadswell examined
Arnold Arb. 19: 99-100, 205-208. 1938; Mem. Amer. Acad. Arts,
Sci. 18, pt. 3 [Mem. Gray Herb. 4]: 135-140. 1939.
2 Gardens' Bull. Singapore 12, pt. 1: 1-17. 1949.
3 The anatomy of the timbers of the south-west Pacific area. III. Myrtaceae.
Austral. Journ. Bot. 1: 353-401. pis. 1-10. 1953.
168 FIELDIANA: BOTANY, VOLUME 29
wood-samples of seven species of Eugenia thought to be tropical
American in origin, and numerous samples of various species from
the southwest Pacific. Although these authors do not support
Merrill and Perry in the recognition of the small genera Acmena,
Cleistocalyx and Jossinia, all are in general agreement that there
is a notable hiatus between two great groups of species, one primarily
of the Old World and the other of the New, that have been in the
past indiscriminately assigned to Eugenia.
The separation of "Eugenia A" and "Eugenia B" has received
additional, if somewhat inconclusive, support from studies in the
comparative morphology of pollen, by Kathleen M. Pike. 1 Miss
Pike found the pollen of three species of "Eugenia A" (none of which
was American) to be quite distinct from that of "Eugenia B." She
was unable to distinguish the pollen of Cleistocalyx from that of
Syzygium, but could distinguish these genera from Acmena on the
basis of pollen morphology.
An increasing body of evidence thus points to the conclusion that
the New World eugenioid species are generically distinct from most
of the Old World species formerly referred to Eugenia. It is probable
that the name Syzygium Gaertn. will be conserved for the principal
group of these Old World species, and it may be that Syzygium will
attain general recognition as an independent genus. The propriety
of such a course, as pointed out by Henderson, cannot be determined
with much certainty unless someone is willing to undertake the
thankless and difficult task of a worldwide revision of Eugenia.
In the meantime it seems clear that the American species can
profitably be studied apart from their Old World relatives, since as
far as I am aware no one has suggested any close relationships at
the specific level.
Within the large group of American eugenioid species are several
distinctive sub-groups which some authors have recognized as
(a) Eugenia sens. str. This is the group which includes the type
of the genus, E. uniflora L. In the approximately 70 species recog-
nized by Berg (as members of his genera Phyllocalyx and Steno-
calyx), the hypanthium is not prolonged beyond the ovary, the
ovules are numerous, and the embryo is undivided; the calyx-lobes
are often elongated and foliaceous or subfoliaceous, and the flowers
occur partly in short racemes and partly in bracteate axils at the
1 Pollen morphology of Myrtaceae from the south-west Pacific area, Austral.
Journ. Bot. 4: 13-53. pi. 1. 1956.
McVAUGH: TROPICAL AMERICAN MYRTACEAE 169
bases of new shoots. Berg based the genera Phyllocalyx and Steno-
calyx on the characters of calyx-lobes and flower arrangement. It
does not seem desirable to recognize these segregate genera (aside
from the fact that one of them, Stenocalyx, was illegitimate when
published for it included the type species of Eugenia), because the
inflorescence character on which they are primarily based is one
which occurs occasionally in racemose-flowered species of Eugenia.
It is possible to demonstrate various intermediate conditions be-
tween the one extreme, in which all flowers are in the axils of bracts
at the bases of new shoots, and the other, in which the flowers are
normally in short racemes (as in Eugenia egensis DC.). Phyllocalyx
and Stenocalyx together, however, form a somewhat doubtfully
recognizable taxon, already designated by Niedenzu (Natiirl.
Pflanzenfam. Ill, pt. 7: 81. 1893) as Eugenia, subgenus Macrocalyx.
The range of the group is from the West Indies southward through-
out the lowlands of eastern South America. As yet no species has
been found in Peru.
(b) Typical Eugenia in the sense of most authors, not of Lin-
naeus. A large group of species with multiovulate ovary, the
hypanthium little or not at all prolonged beyond the ovary, and
the flowers in short or long racemes or apparently glomerate because
of extreme reduction of the raceme axis. These include most of the
Biflorae, Corymbiflorae, Glomeratae, Racemosae, Racemulosae and
Umbellatae of Berg, a total according to him of about 360 species
in America. The embryo, as far as known, is undivided, as in typical
Eugenia; most species are known from flowering material only,
however, and it is not possible to make a general statement about
embryo structure in this group.
(c) The genus Anamomis Griseb. A group of 40 to 50 species,
ranging from the West Indies southward, chiefly along the Andes,
to Bolivia and Argentina. The ovary is multiovulate, the hy-
panthium is little or not at all prolonged, the flowers are solitary or
in simple or compound dichasia, and the cotyledons as far as known
are 2 and distinct. The species of this taxon were included by Berg
in his group Dichotomae, of Eugenia. On the basis of geographical
restriction, the structure of the inflorescence, and the morphology
of the embryo, Anamomis seems to be more deserving of independent
generic status than Plinia, Myrciaria or Catinga. A revision is
needed, however, of the entire group of taxa including Anamomis,
the south-Brazilian Myrcianthes Berg, and the Chilean genera
Reichea Kausel and Myrceugenella Kausel. Myrceugenella, for
170 FIELDIANA: BOTANY, VOLUME 29
example, differs from Anamomis chiefly in having the radicle about
as long as (instead of about one-third as long as) the cotyledons.
Reichea and Myrcianthes have pentamerous rather than tetramerous
flowers, but otherwise seem to differ little from Anamomis. At
present, moreover, most of the species of Anamomis are unknown
in the fruiting condition (mature fruits have been seen for 5 of the
19 known species which occur in Peru), and in view of the importance
which has been attached to the myrtaceous embryo as an indicator
of relationship, it seems unwise to transfer numerous species to this
genus, on the basis of the inflorescence alone, before the taxonomy
of the group has been carefully studied.
The nomenclature of the group is somewhat involved. If study
indicates that but a single genus should be recognized, the oldest
name for this is apparently Lama A. Gray, published in 1854 and
thus antedating any of the generic names published by Berg. If
Myrcianthes and Anamomis prove to be congeneric, the former name
has priority. If Anamomis proves to be distinct from all the others,
the name is valid for the Andean-West Indian group of species
which is considered here.
There appears to be little more than superficial similarity be-
tween the species of Anamomis and those of the Old World genus
Syzygium Gaertn. The inflorescence in Syzygium is often terminal;
if lateral it may be strictly racemose (with the terminal flower
developed, not abortive as in most American Eugenias), or more
often it may simulate the myrcioid panicle, with the principal
branches opposite, elongate, and terminating in single flowers or
small dichasia. The consistent occurrence of a regularly dicho-
tomous dichasium is apparently unique in Anamomis and its rela-
tives (cf. similar inflorescences in Pimentinae). 1
1 Since the above was set in type I have received, through the kindness of
Dr. Carl Skottsberg, a copy of a very recent paper in which the whole matter
of generic segregations in the American Myrtaceae is discussed by Dr. Eberhard
Kausel (Beitrag zur Systematik der Myrtaceen, Ark. Bot. 3: 491-516. 1956).
Kausel proposes to erect a new family, Leptospermaceae, for the capsular-fruited
genera usually assigned to Myrtaceae, and to restrict the use of the name Myr-
taceae to those fleshy-fruited genera which comprise the tribe Myrteae DC.
The Myrtaceae in this narrower sense are divided by Kausel, on the basis of
characters of the seed and embryo, into the following taxa, which are presumably
intended as subfamilies, although apparently not formally designated as such:
Eugenioideae, Plinioideae, Cryptorhizoideae, Myrtoideae, and Myrcioideae.
Five new genera are proposed, of which one is assigned to Eugenioideae, three to
Plinioideae, and one to Myrtoideae. In the Plinioideae Kausel recognizes Ana-
momis Griseb. and four other genera which have the inflorescence an axillary
dichasium, viz., Pseudanamomis Kausel, Reichea Kausel, Myrcianthes Berg,
and Amyrsia Raf. with type species Myrtus foliosa H.B.K. The superficially
similar Pseudomyrcianthes Kausel is assigned to the Eugenioideae and Myrceu-
genella Kausel to the Myrcioideae.
McVAUGH: TROPICAL AMERICAN MYRTACEAE 171
(d) The genus Myrciaria Berg. A group probably of about
40 species with few (usually 2) ovules in each locule of the ovary,
the hypanthium markedly prolonged beyond the ovary and circum-
scissile at the base, the flowers subsessile in small (racemose) clusters
and the cotyledons distinct or nearly so. According to Berg the
genus included 65 species; Bentham pointed out (Benth. & Hook,
f. Gen. PI. 1: 720. 1865) that several of the original species probably
belonged to some myrcioid genus. After the removal of these
anomalous species the genus Myrciaria appears to comprise a co-
herent and natural group, which ranges from Panama southward,
chiefly in the lowlands east of the Andes. Siphoneugena Berg is
a very small south-Brazilian genus of uncertain status which differs
from Myrciaria chiefly in having the ovary multiovulate.
(e) The genus Plinia L., in the sense of Urban (Repert. Sp.
Nov. 15: 412-413. 1919). A small assemblage of species in which
the locules of the ovary are 2-ovulate, the hypanthium is prolonged,
the calyx is partially or completely closed in bud and splits longi-
tudinally at anthesis, the flowers are usually in sessile bracteate
clusters, and the cotyledons are distinct. About half a dozen closely
related species are known from northern continental South America;
Urban described in his later years a considerable number of species
of Plinia from the West Indies, but most of these are small-leaved
plants with little apparent similarity to the South American species.
On the basis of the distinctive calyx and the small number of ovules,
the South American species are readily recognized as members of
this group, and I have assigned several Peruvian species to it.
(f) A heterogeneous group of perhaps 15-20 species, in which
the ovary is multiovulate, the hypanthium is mostly not or scarcely
prolonged beyond the ovary, the buds are closed or the calyx-lobes
coherent in the bud, the flowers are in short axillary racemes, and
the embryo is undivided. Except for the united or coherent calyx-
lobes, any of these species could pass for a member of the genus
Eugenia, and many of them have been attributed at times to that
genus. The taxon characterized above includes Calycorectes Berg
(with buds closed) and probably also Schizocalyx Berg, and in Peru
and Amazonian Brazil the genus Catinga Aubl. (in the sense of
Amshoff, in Fl. Suriname 3: 105. 1951). The status of Catinga is not
clear to me, and I hesitate to recognize it as an independent genus.
It differs from Eugenia in one character only, namely, that the four
calyx-lobes are coherent in the bud for part or all of their length and
172 FIELDIANA: BOTANY, VOLUME 29
are usually thickened and cucullate at the tips. 1 The species
assigned to Catinga presumably differ from Calycorectes because in
the former the lobes are loosely coherent along narrow lines near
the edges but separate cleanly at anthesis into symmetrical thin-
edged lobes, instead of rupturing irregularly as in Calycorectes.
To me this is not a convincing character upon which to base a genus,
especially since the supposed species of Catinga are not otherwise
very different from the many species of Eugenia which have similar
foliage and similar inflorescences. The genus Calycorectes itself is
not well understood; it is indeed restricted by Amshoff (op. cit. 104)
to a single species, although more than 15 species have been de-
In summation, the Peruvian eugenioid species belong for the
most part to Eugenia, which is understood to include Phyllocalyx
Berg and Stenocalyx Berg, and, for the present at least, Anamomis
Griseb. and Catinga Aubl. Myrciaria Berg is apparently a distinct
genus, and Plinia L. (in the sense of Urban) is recognized.
3. The pimentoid genera. The genera of Pimentinae which
have membranaceous seed-coats, namely Blepharocalyx and Cam-
pomanesia, are represented in Peru by a single species each; evalu-
ation of their generic limits should be undertaken only after study
of the species of eastern and southeastern Brazil, where these genera
are represented by additional species. The remaining genera, those
with hard or bony seeds, fall into two principal groups, namely,
Psidium and Myrtus and their respective segregates. The separation
between these two has been traditionally upon the basis of calyx
morphology. Myrtus has been a form-genus including especially all
small-leaved species (like the type species, M. communis L., of
Europe) with 1-flowered peduncles, 2- to 3-locular ovary and 5
(rarely 4) distinct calyx-lobes. Psidium was originally a form-genus
including mostly larger-leaved species (like the type species, P.
guajava L., the guava) with 1- or 3-flowered peduncles, the ovary
usually more than 2-locular, and the calyx closed or nearly closed
in bud and splitting irregularly in anthesis.
In tropical America Psidium is a relatively "good" genus, with
numerous and distinctive species, but unfortunately the character
of the closed and irregularly breaking calyx applies to relatively few
of these. Many species described by DeCandolle, Berg, and later
authors have an open, lobed calyx. The lobes are usually small, and
1 For additional notes on the status of Catinga, see below (p. 203), in the
discussion of Eugenia acrensis.
McVAUGH: TROPICAL AMERICAN MYRTACEAE 173
the hypanthium splits longitudinally in the sinuses as the flowering
period passes and the fruit expands; this constitutes, in fact, one of
the principal ways in which a species of Psidium may be recognized.
Calyptropsidium Berg and Psidiopsis Berg differ in minor features
of the calyx and are not convincingly distinct from Psidium.
Seemingly intermediate between Psidium and Myrlus is Caly-
colpus, which according to Riley 1 contains 12 species. This is
a genus which has almost no characters. According to Riley it may
be distinguished from Campomanesia, Myrtus and Psidium by its
calyx-lobes, "which are patent in the flower-bud, whereas in Cam-
pomanesia and Myrtus they are appressed to the corolla when in
bud, and in Psidium they form a closed calyx which splits into
segments as the flower expands." It is true that a species of Caly-
colpus can usually be recognized as such by the rather large flowers
which are solitary in the axils, by the calyx-lobes which are disposed
as noted above, and by the coriaceous leaves, which usually blacken
in drying. Superficially, however, a small Calycolpus looks not
unlike a large specimen of Myrtus communis, and if any revision of
Myrtus is undertaken the status of Calycolpus should be considered
at the same time. As far as I am aware, no species of this genus
ranges as far southwest as the borders of Peru; the Rio Negro forms
the approximate limit of the group in this direction.
The genus Myrtus itself is in need of study. Bentham (Benth.
& Hook. f. Gen. PL 1: 714. 1865) stated that the genus as he knew it
contained probably about 50 species, mostly American but occurring
on all major land areas except possibly tropical Asia. Actually
Myrtus in the sense of Bentham was something of a catch-all. It
included the small Andean groups now generally referred to the
genera Myrteola Berg and Ugni Turcz. ; Anamomis and Myrcianthes
of the Eugeniinae; Myrceugenia of the Myrciinae; the Bergian
genera Blepharocalyx and Pseudocaryophyllus of the Pimentinae;
additional Chilean genera now regarded by Kausel as completely
distinct; and various additional American plants, practically all of
which have since been referred to genera other than Myrtus.
At least one recent author has retained a few American species
in Myrtus, 2 but an opposing point of view has been taken by Burret, 3
who has transferred most of the described species (from both Old
1 Kew Bull. 1926: 145-154. 1926.
2 Legrand, D. Las Mirtaceas del Uruguay, An. Mus. Hist. Nat. Montevideo,
ser. 2, 4, pt. 11: 21-24. 1936.
3 Myrtaceen-Studien, Notizbl. Bot. Gart. Berlin 15: 479-500. 1941.
174 FIELDIANA: BOTANY, VOLUME 29
and New Worlds) to other genera, retaining in Myrtus no more than
the original M. communis, one African species and about 14 species
in Florida and the Greater Antilles.
Pending further revision of the whole group as it exists in
America, I have chosen to recognize Ugni and Myrteola as local,
independent genera. These appear well founded when judged by
ordinary criteria of morphological distinctness, homogeneity, and
well-defined geographical range. It is by no means clear that Myrtus
is represented in the American flora by any species whatsoever, and
it seems unwise to attempt to force the several Myrtus-\ike American
groups into a generic concept based so largely upon a single European
Finally, a word should be said about Acca, which Berg placed
among the Eugeniinae. Bentham stated his opinion that the genus
belonged rather with Psidium, and the seed-characters of the two
species of Acca confirm this disposition. The genus is well marked,
with style and androecium unlike anything known in the Myrteae
unless it be Feijoa Berg, which Burret (Repert. Sp. Nov. 50: 49.
1941) has relegated to the synonymy of Acca.
Systematic Treatment of Genera and Species
1. MARLIEREA Camb. 1
1. 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.
2. Inflorescence, including the hypanthium, strongly velutinous or at least with
numerous loosely ascending reddish or reddish-yellow silky hairs up to
1 mm. long.
3. Inflorescence strongly velutinous; flowers large, the buds 7 mm. long; calyx-
tips free; leaves acute or acuminate, 13-18 cm. long. . .M. velutina McVaugh.
3. Inflorescence loosely silky-hairy; flowers small, the buds 2.5 mm. long,
closed, apiculate; leaves caudate-acuminate, 4.5-7 cm. long.
M. caudata McVaugh.
2. Inflorescence glabrous or essentially so; at least the tips of the calyx-lobes
evident in the bud.
4. Leaves 9-14 cm. long, 2-3.5 times as long as wide; lateral veins 12-15 pairs,
the transverse veins obscurely reticulate; leaves finely and obscurely dark-
dotted; calyx-lobes in bud minute, ciliate M. scytophylla Diels.
4. 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 millimeter; calyx-lobes in bud distinct, the
inner ones broadly scarious-margined, 2.5 mm. wide. .M. areolala McVaugh.
1 One species, Marlierea insignis, is described below but is not included in the
key; it is known from Amazonian Colombia (Rio Apaporis) and is probably not
a member of the Peruvian flora.
McVAUGH: TROPICAL AMERICAN MYRTACEAE 175
1. Midvein sharply and narrowly impressed on the upper surface, or in one
species concave or broadly sulcate.
5. 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; mid vein concave or sulcate; hypanthium tomentose within.
M. spruceana Berg.
5. Inflorescence pubescent, often sparsely so, with short, pale or reddish hairs
0.5 mm. long or less; buds glabrous or essentially so, 1.5-3.5 mm. long;
mid vein sharply and narrowly impressed; hypanthium glabrous within.
6. Leaves cordate-auriculate, nearly sessile, the petioles 3-4 mm. long; in-
florescence finely hispidulous with minute stiff erect hairs; buds closed, with
a prominent narrow apiculum; staminal ring short-hairy.
M. subulata McVaugh.
6. Leaves acute to cuneate or somewhat rounded at base, on petioles 4-10 mm.
long; inflorescence pubescent with appressed or ascending hairs; buds closed
or the calyx-lobes distinct; staminal ring glabrous.
7. Petioles transversely rimose, the reddish-brown or whitish papery layers
separating but persistent; calyx in bud with 4 very small separate deltoid
tips; lower branches of the panicle straight and elongated, spikelike, with
numerous sessile flowers and short squarrose bracts.
M. umbraticola (HBK.) Berg.
7. Petioles smooth and with unbroken surface, usually dark in color; buds
closed, or with 5 distinct calyx-lobes; inflorescence various.
8. Inflorescence thinly pubescent with pale hairs; bracts and bracteoles decidu-
ous before anthesis; lateral veins of the leaves not impressed above; buds
3-3.5 mm. long, closed at apex M. imperfecta McVaugh.
8. Inflorescence pubescent with lustrous rufous hairs; bracts and bracteoles
persistent, squarrose; lateral veins impressed above; buds 1.5-2 mm. long,
with 5 distinct calyx-lobes M. squarrosa McVaugh.
Marlierea areolata McVaugh, sp. nov.
Subglabra, multiflora, foliis 7.5 cm. longis; venis utroque latere 6-8, venulis
reticulatis; laminis utrinque crebro pellucido-punctatis; nervo medio supra piano
vel convexo; calycis lobis liberis 4, hypanthio demum explanato, ad germinis
A shrub or tree with elliptic, bluntly acuminate leaves and relatively few
lateral veins, the foliage distinctive also because of the prominently reticulate
veinlets and the numerous large translucent glands; calyx-lobes in unequal pairs,
the inner ones broadly scarious-margined, about 1.7 mm. long and 2.5 mm. wide;
style 4 mm. long; stamens 75-100.
The large and distinct calyx-lobes, and the deciduous bracts and
bracteoles, suggest that this species may have some affinity to the
genus Myrcia (Aulomyrcia) . It is here referred, however, to Mar-
lierea, because of the flattening of the hypanthium after anthesis,
and the accompanying distortion and splitting of the calyx. As an
additional minor character may be mentioned the terminal or
falsely terminal inflorescence in this species; this character recurs
throughout the genera Calyptranthes and Marlierea, whereas in
Myrcia the panicles are more often from the lower axils.
176 FIELDIANA: BOTANY, VOLUME 29
Peru, Loreto: Stromgebiet des Ucayali von 10 S. bis zur Miin-
dung, G. Tessmann 3264, anno 1923 (G, type). This specimen was
determined in the herbarium by Burret as a new species of Myrcia,
but as far as I can learn this has never been published. F.M. Neg.
Marlierea caudata McVaugh, sp. nov.
Arbor, ramulis paniculisque ferrugineo-sericeis; foliis parvis, usque ad 7 cm.
longis, longe acuminatis; paniculis paucifloris, alabastra 2.5 mm. longa, clausa,
A tree up to 7 meters high, the pubescence of reddish or reddish-yellow silky
hairs; leaves 4.5-7 cm. long, 2-3 times as long as wide (including the narrow
acuminate tip 1-2 cm. long); principal branches of the inflorescence 3-5 cm. long,
each with 20 flowers or fewer; calyx splitting irregularly into 4 oblong lobes; style
4.5 mm. long; stamens about 100; fruit subglobose, 8-11 mm. in diameter.
Peru, Loreto: Mishuyacu, near Iquitos, elev. 100 meters, forest,
G. King 235, Oct.-Nov., 1929 (F 624286, type; NY; US); forest
between [lower] Rio Nanay and Rio Napo, June 6, 1929, L. Williams
Marlierea imperfecta McVaugh, sp. nov.
Arbor, pubescens, foliis 12-25 cm. longis acuminatis, petiolis 5-8 mm. longis
canaliculatis; nervo medio supra impresso, venis lateralibus non impressis; panic-
ulis multifloris, bracteis bracteolisque deciduis; alabastris clausis, 3-3.5 mm.
A small tree, thinly pubescent with pale or reddish-based hairs; leaves elliptic,
2.5 times as long as wide, with 10-15 pairs of lateral veins; inflorescence 3 times
compound, the principal branches 8-14 cm. long; calyx splitting irregularly into
4 lobes; style 4-4.5 mm. long; stamens about 100.
Peru, Loreto: Mishuyacu, near Iquitos, elev. 100 meters, forest,
Jan., 1930, G. Klug 787 (US 1455778, type; NY; F).
Marlierea insignis McVaugh, sp. nov.
Arbor mediocris, insigniter velutina; ramulis, petiolis paniculisque, pilis
rigidis erectis usque ad 1 mm. longis, aureo-fulvis, dense obtectis; foliis ellipticis
15-25 cm. longis acuminatis; venis utroque latere 12-15, planiusculis vel impressis;
calyce in alabastro ut videtur clauso, ad florendi tempus irregulariter rumpente;
paniculae ramis 4-6 cm. longis crassis, paucifloris; fructu globoso, diametro
Tree up to 10 meters high, densely velutinous with golden-brown hairs, the
hairs stiff, erect, up to 1 mm. long, completely covering the branchlets, vegetative
buds, inflorescence (including the fruit), and mid vein at least beneath; lower
leaf-surface loosely hirsute with similar hairs; leaves broadly elliptic, 7-13 cm.
wide, 15-25 cm. long, rounded to apex and then abruptly short-acuminate, the
McVAUGH: TROPICAL AMERICAN MYRTACEAE 177
acumen acute with concave sides, 5-15 mm. long; base of blade rounded, the
margins at very base decurrent on the petiole, which is up to 4 mm. thick (including
hairs) and 10 mm. long; midvein flat or concave above, velutinous or glabrate,
prominent beneath; lateral veins 12-15 pairs, plane or impressed above, prominent
beneath; marginal vein about equaling the laterals, strong and little arched
between them, 3-6 mm. from margin; leaves dull and pale green above, yellow
green beneath and there obscurely and minutely dark-dotted; inflorescence
branches 3-6 cm. long, 3 mm. thick (including the hairs), solitary in the axils
or the fertile branch aborting at the first node and producing a pair of nearly
equal lateral branches; individual branches spike-like, with up to 7 nearly sessile
flowers; flowers not seen; fruit globose or nearly so, about 1.5 cm. long, 1.3-2
cm. in diameter; calyx apparently closed in bud, with a conical top, glabrous
within this, and breaking irregularly in anthesis; inner disc at base of style 2 mm.
in diameter, the hypanthium produced 1-1.5 mm. above the summit of the ovary
and hairy within, the numerous (probably 150-200) stamens produced near the
summit of the bud in a ring 1.5 mm. wide; anthers 0.3 mm. long; style 5 mm. long;
ovules 2 in each of 2 locules.
Colombia, Amazonas-Vaupe"s: Rio Apaporis, Soratama, elev. ca.
250 meters, June 21, 1951, R. E. Schultes & I. Cabrera 12766 (US
2220050, type; MICH) ; Aug. 24, 1951, Schultes & Cabrera 13722 (US).
Marlierea squarrosa McVaugh, sp. nov.
Frutex, rufo-pubescens; foliis 9-13 cm. longis acuminatis, venis supra im-
pressis, petiolis 5-7 mm. longis levibus; floribus subspicatis, bracteis divaricatis
persistentibus; alabastris 1.5-2 mm. longis glabris; calycis lobis 5 valde inaequal-
A shrub 2 meters high, with elliptic leaves 2.5 times as long as wide, and
many-flowered panicles; flowers small, the calyx splitting irregularly from the
base of the lobes to the summit of the ovary; style 3.5 mm. long; stamens about 50.
This species, like Marlierea 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 (when dry) and free calyx-lobes
of some species of Myrcia. It might conceivably be a hybrid in-
volving Marlierea umbraticola, which also has impressed veins, but
as yet M . umbraticola has not been collected within the borders of
Peru. The following is the only known collection of M. squarrosa.
Peru, Loreto: Mishuyacu, near Iquitos, elev. 100 meters, forest,
G. King 169, Oct.-Nov., 1929 (US 1455168, type; NY; F).
Marlierea subulata McVaugh, sp. nov.
Arbor, hispidula; foliis subsessilibus, 13-18 cm. longis, ovatis lanceolatisve ;
nervo medio supra impresso; paniculis angustis paucifloris; alabastris subglabris,
3 mm. longis, clausis, apiculatis; disco staminali pubescente.
178 FIELDIANA: BOTANY, VOLUME 29
A tree 5 meters high, with cordate-auriculate nearly sessile leaves about 3
times as long as wide; bracts subulate, up to 3 mm. long, more or less persistent
through anthesis; calyx splitting irregularly into 4 lobes; style 6 mm. long; stamens
75-100; fruit up to 1 cm. long and 1.5 cm. in diameter.
Peru, Loreto: Mishuyacu, near Iquitos, elev. 100 meters, forest,
May, 1930, G. King 1341 (F 627613, type; NY; US).
Marlierea velutina McVaugh, sp. nov.
Fulvo-brunneo-velutina, foliis 13-18 cm. longis acuminatis, nervo medio
supra piano vel basi sulcato; paniculis multifloris; floribus grandis, alabastris ut
videtur 7 mm. longis; hypanthio circiter 8-sulcato; calyce irregulariter rumpente.
A shrub or tree, densely beset with coarse yellowish-brown hairs up to 1 mm.
long; leaves elliptic to lanceolate, about 3 times as long as wide; inflorescence
6-10 cm. long, 2 or 3 times compound; calyx-lobes 4, at first united except the free
tips 1 mm. long; style 7 mm. long; stamens probably about 200.
A distinctive species suggesting some of the members of the
genus Gomidesia because of the color and character of the pu-
Brazil, Guapore": Falls of Madeira, H. H. Rusby 2683, Oct., 1886
(F, type; US). Univ. of Mich. Neg. 449.
2. CALYPTRANTHES Sw.
1. 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 brownish dibranchiate hairs 0.1 mm.
long; leaves 25-39 cm. long, narrowly elliptic, appearing cordate-auriculate
at base, nearly sessile, the stout petiole 4 mm. long. . .C. maxima McVaugh.
1. Flowers smaller, the buds if 5 mm. long or more fusiform and apiculate, or
noticeably hirsute or velutinous; inflorescence glabrous to hairy; leaves
usually smaller and slender-petiolate, if sessile or essentially so the buds not
2. Leaves sessile, with veins impressed on the upper surface; blades cordate
at base, or the margins much produced and puckered.
3. Leaves ovate, cordate, 9-12 cm. long; inflorescence glabrous, the buds
fusiform, 6-7 mm. long C. sessilis McVaugh.
3. Leaves obovate, with the margins near base produced into puckered folds,
the blades 30-38 cm. long; inflorescence appressed-hirsutulous, the buds
obovoid, 3.5 mm. long C. plicata McVaugh.
2. Leaves petiolate, cuneate to acute or somewhat rounded at base, the veins
usually not impressed above.
4. Inflorescence of paired spikes, the individual flowers sessile along the axis,
or the lowest in sessile or very short-peduncled groups of three.
5. Buds glabrous; leaves 2.5-6 cm. long, often obovate with rounded or some-
times short-acuminate tips; flowers 3-5 (-11) in each spike. . C. pulchella DC.
5. Buds strigose or hirsute; leaves 7-16 cm. long, elliptic or ovate, usually
prominently and often narrowly acuminate; flowers more numerous.
McVAUGH: TROPICAL AMERICAN MYRTACEAE 179
6. Branchlets and inflorescence, including the buds, thickly rufous-hirsute; buds
broadly obovoid to nearly globose, 5-6 mm. long, concealed by the hairs;
flowers 8-13 in each spike C. krugioides McVaugh.
6. Branchlets and inflorescence with appressed yellow or brown dibranchiate
hairs; buds obovoid or broadly fusiform, 2-2.5 mm. long, rather sparingly
7. 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 im-
pressed above, slender and closely parallel C. densiflora Berg.
7. Spikes 3-5.5 cm. long, the flowers 25 or fewer in each, in small sessile clusters
of 1-3 each; leaves with narrow acumen 1.5-2 cm. long; lateral veins im-
pressed above, the principal ones prominent beneath and contrasting with
the less conspicuous intermediates C. brevispicata McVaugh.
4. Inflorescence of paired panicles or compound dichasia, the basal branches
elongate and again branched, or occasionally 1-flowered only.
8. Branches of the inflorescence variously pubescent (sometimes thinly so)
with appressed or erect hairs; hypanthium variously strigose to tomentose
except in C. multiflora.
9. 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 Berg.
9. Hypanthium strigose or variously velutinous or tomentose, if nearly glabrous
the midvein sulcate or narrowly impressed above.
10. Inflorescence a compact umbelliform cyme 2-5 cm. long with 15 flowers or
fewer, the ovate boat-shaped bracts sub-foliaceous, persistent; inflorescence,
including the flowers, thickly hirsute with coarse sessile dibranchiate hairs
up to 1.5-2 mm. long and attached near one end C. longifolia Berg.
10. Inflorescence paniculate or by reduction racemose, often 3 to 4 times com-
pound and many-flowered (if short and few-flowered not hirsute as above);
bracts all deciduous before anthesis, or a few (usually the basal ones) per-
sisting; hairs of the inflorescence various, mostly sessile and less than 1 mm.
11. Hairs of the inflorescence golden-yellow, dibranchiate, 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; branchlets 2-winged; flowers mostly 10 or fewer on each branch.
C. tridymantha Diels.
11. Hairs of the inflorescence sessile and, if abundant, usually red or rusty, all
somewhat appressed and mostly less than 0.5 mm. long, or with very short
hairs intermixed, or the inflorescence velutinous or tomentose.
12. Leaves large, mostly more than 15 cm. long (often 20-30 cm.), with 20-35
pairs of lateral veins; inflorescence with abundant rufous pubescence; buds
obovate or obconic, scarcely apiculate, 3-5 mm. long.
13. Blades tapering from the middle or below, to a slender apex; lateral and
marginal veins scarcely apparent on the lower leaf surface, which is covered
with very numerous closely appressed pale hairs up to 0.2 mm. long; inflo-
rescence 5 cm. long or less, few-flowered C. macrophylla Berg.
13. Blades abruptly and narrowly acuminate; lateral and marginal veins forming
a conspicuous pattern on the lower surface, which is glabrous or sparingly
appressed-pubescent; inflorescence 6-10 cm. long, many-flowered.
C. gigantifolia McVaugh.
12. 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 pu-
180 FIELDIANA: BOTANY, VOLUME 29
14. Flowers small, the buds 2-2.5 mm. long, obovoid, the apex rounded or shortly
apiculate; panicles mostly 3 times compound, many-flowered, the branches
sparingly covered with appressed pale or sometimes reddish hairs; hypan-
thium strigose, sometimes very sparingly so; paired panicles from an abortive
flattened axis 1 cm. long or less.
15. 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 pairs of lateral veins; style 4-4.5 mm. long.
C. ruiziana Berg.
15. Lower leaf surfaces glabrous except for a few dark hairs; leaves ovate or
lanceolate, usually widest somewhat below the middle, 9-15 (-25) cm. long,
with 20-25 pairs of veins; style 5-6 mm. long C. simulata McVaugh.
14. Flowers larger, the buds 3-6 mm. long, variously shaped; panicles compound,
or by reduction racemoid, the branches and the hypanthium usually con-
spicuously appressed-hairy, velutinous, or tomentose, with ferruginous or
dark reddish hairs; panicles paired or with a central axis.
16. Inflorescence a pair of spikes with all flowers sessile, or the lower branches
1-2 cm. long and 1-flowered, or with 3 sessile flowers at the tip; buds 5-6 mm.
long, abundantly hirsute, the hypanthium hairy within.
C. krugioides McVaugh.
16. Inflorescence paniculate, usually many-flowered and 3 times compound;
buds 3-4 mm. long, appressed-hairy to tomentose, the hypanthium glabrous
17. Mid vein impressed above; hypanthium appressed-hairy, the hairs of the
inflorescence rusty-brown; buds fusiform, 3-3.5 mm. long; panicle narrow,
the lowest branches about 1.5 cm. long C. tessmannii McVaugh.
17. Midvein convex above (and then sometimes sulcate) or raised in a narrow
ridge; hypanthium loosely velutinous or tomentose; buds obovoid or ellipsoid;
lower branches of the panicle relatively long.
18. Leaves 15-21 cm. long, about 4 times as long as wide, the straight marginal
vein and the 10-15 short divaricate lateral veins prominent beneath; petiole
very stout, 3 mm. thick and 7 mm. long; buds broadly ellipsoid, heavily
tomentose with dark red hairs, the hypanthium urceolate in anthesis.
C. rufotomentosa McVaugh.
18. Leaves 10-16 cm. long, about 2.5 times as long as wide, the marginal vein
not well defined, or consisting of a series of loops or arches between the 8-12
pairs of arcuate laterals; petiole 1 mm. thick, 8-10 mm. long; buds obovoid,
sparingly or rather densely velutinous with rusty brown or reddish hairs, the
hypanthium probably turbinate in anthesis C. cuspidata DC.
8. Branches of the inflorescence, and the hypanthium, completely glabrous or
with a very few scattered hairs about the base and the nodes of the panicle.
19. Panicles with 20 flowers or fewer, sometimes reduced and raceme-like or
spike-like; peduncle and rachis filiform or very slender, often terete and
nearly straight, usually less than 1 mm. thick; branchlets narrowly 2-winged.
20. Flowers mostly sessile, 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 DC.
20. 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 Berg.
19. Panicle many-flowered, 3 to 4 times compound, the peduncle usually some-
what angular near summit and 1-1.5 mm. thick, the rachis often irregularly
enlarged and zigzag; wings, if produced on branchlets, not persisting through
a growing season.
McVAUGH: TROPICAL AMERICAN MYRTACEAE 181
21. Lower leaf surface obscurely gland-dotted, the dots about 15 per square
millimeter; leaves relatively narrow, mostly 2.5 times as long as broad, or
longer; panicle branches irregularly alternate C. paniculata R. & P.
21. Lower leaf surface prominently dark-dotted, the dots more than 50 per
square millimeter; leaves broader, mostly 2.3 times as long as broad, or less;
panicle branches, both large and small, often verticillate or fasciculate.
C. crebra McVaugh.
Calyptranthes brevispicata McVaugh, sp. nov.
Arbor pubescens, foliis longe acuminatis, venis supra impressis, subtus
prominulis, venulis interjectis tenuioribus; spicis 3-5.5 cm. longis binis, floribus
usque ad 25 sessilibus, fasciculatim 1-3-nis e nodis oriundis.
A small tree with elliptic-lanceolate leaves 11-15 cm. long, sparingly pu-
bescent with yellowish brown hairs; a species superficially suggesting C. densiflora
Berg, but the spikes shorter and with far fewer flowers than in that species; buds
fusiform, about 2.5 mm. long; style 6 mm. long; stamens about 50.
Peru, Loreto : Florida, Rio Putumayo, at mouth of Rio Zubineta,
elev. 200 meters, forest, March-April, 1931, G. King 2040 (NY; US
Calyptranthes crebra McVaugh, sp. nov.
Arbor subglabra, floribus paniculatis, paniculae ramis ramulisque verticillatis
vel fasciculatis; foliis ellipticis vel ovatis, 1.8-2.3-plo longioribus quam latioribus,
subtus creberrime atro-punctatis.
A tree 4-10 meters high, nearly glabrous, with leaves 8.5-10.5 cm. long;
a species superficially like C. paniculata R. & P., but readily distinguished from
that species by the key characters. Buds obovate, apiculate, about 2 mm. long;
style 5 mm. long; stamens 60-75; fruit globose, about 5 mm. in diameter.
Peru, Loreto: Near Iquitos, in forest, elev. 100 meters, Oct.-Nov.,
1929, G. King 77 (F; NY; US), 78 (F 624362, type; NY; US), 607
(F; NY; US).
Calyptranthes gigantifolia McVaugh, sp. nov.
Arbor grandis, foliis 14-38 cm. longis acuminatis, subtus glabris vel parce
pilis appressis pubescentibus, venis prominentibus; paniculis pilis appressis rufis
vestitis, 6-10 cm. longis multifloris; alabastris 3-5 mm. longis.
A tree up to 20 meters high, resembling C. macrophylla Berg, but that species
having the leaves more coriaceous, acute rather than acuminate, and less promi-
nently veined; the marginal vein in C. macrophylla is close to the margin and not
impressed above; the lower surface in C. macrophylla is densely pubescent with
minute appressed pale hairs and with numerous intermixed dibranchiate hairs;
in this last respect C. macrophylla agrees precisely with C. speciosa Sagot, of the
Guianas, and it may be identical with that species, as already suggested by
182 FIELDIANA: BOTANY, VOLUME 29
Colombian specimens collected by Mutis (nos. 1940, 2234, 2951,
3961, and 5754) appear to be conspecific with the Peruvian speci-
mens cited below.
Peru, San Martin: Juanjui, G. King 4277 (A; F; US 1458692,
type). Loreto: Florida, Klug 2332 (F; G; GH; NY; US), 2347
(F; G; GH; NY; US).
Calyptranthes krugioides McVaugh, sp. nov.
Arbor grandis, ramulis inflorescentibusque rufo-hirsutis, foliis 10-16 cm.
longis, floribus 8-13, ultimis solitariis, infimis ternis, alabastris 5-6 mm. longis
obovoideis vel subglobosis.
A tree up to 20 meters high, named for its superficial resemblance to Krugia
ferruginea, a native of the West Indies and northern South America. The few-
flowered paired spikes, 5-6 cm. long, are conspicuously rufous-hirsute, and the
buds completely obscured by the hairs; style 8 mm. long; stamens 125-150;
petals 3, 4 mm. long in the bud.
Amazonian Brazil: Basin of Rio Jurua, B. A. Krukoff 5041
(NY; US). Peru, Loreto: Iquitos, edge of lake, elev. 120 meters,
Oct. 11, 1929, L. Williams 3675 (F 618382, type). Univ. of Mich.
Calyptranthes maxima McVaugh, sp. nov.
Arbor parva, puberulenta; ramulis paniculisque sparsiuscule pilis adpressis
minutis, 0.1 mm. longis dibranchiatis, obsitis; foliis ellipticis maximis, 25-39 cm.
longis, acuminatis; petiolo incrassato, 3 mm. diametro, 4 mm. longo; nervo medio
supra elevato sulcatoque; paniculis paucifloris, floribus puberulentis furfurace-
isque, maximis; alabastris 7-8 mm. longis; calyptra explanata, vix apiculata;
staminibus circiter 200.
A tree with leaves thin but extremely large for the genus, and the flowers
large and coarse; leaves narrowly elliptic, acuminate, 3.5-4 times as long as wide,
with 25-35 pairs of slender lateral veins; petiole 4 mm. long, its thickened portion
extending also 5-6 mm. beneath the cordate-auriculate base of the blade; flowers
up to 10 on each main branch of the panicle, arising from a stout flattened axis
up to 3.5 mm. wide; stamens relatively short for the flower, 5 mm. long, fringing
the summit of the hypanthium.
Colombia, Amazonas: Trapecio amazonico, Loretoyacu River,
elev. about 100 meters, Oct., 1946, R. E. Schultes & G. Black 8526
(US 1996504, type).
Calyptranthes plicata McVaugh, sp. nov.
Frutex, foliis maximis obovatis, 30-38 cm. longis, sessilibus, marginibus ad
basin decurrentibus plicatisque; paniculis tenuibus usque ad 7 cm. longis, pilis
appressis runs, partim dibranchiatis obtectis; paniculae bracteis divaricatis per-
sistentibus, 2-3 (-5) mm. longis; alabastris 3.5 mm. longis, obovoideis, acutis,
sessilibus; staminibus circiter 60.
McVAUGH: TROPICAL AMERICAN MYRTACEAE 183
A shrub about 4.5 meters high, with stem 5 cm. in diameter, nearly glabrous
except the appressed-pubescent divaricately branched panicle which bears 50-100
flowers; leaves 10-12 cm. wide, 2.5-3 times as long as wide, with 20-25 pairs of
conspicuous lateral veins; a distinctive species because of the large sessile leaves
with broadly decurrent and puckered basal margins.
The inflorescence, with its persistently bracteate nodes, divari-
cate branches and relatively few sessile flowers, suggests that of
Calyptranthes longifolia Berg but is more slender and elongate;
vegetatively, the two species are dissimilar. Known only from the
type, which bears one inflorescence in young bud, and a second,
detached inflorescence with very much younger, undeveloped buds.
Brazil, Amazonas: Municipality Sao Paulo Olivenca, near Pal-
mares, on terra firma, low land, high forest, Sept. 11-Oct. 26, 1936,
B. A. Krukoff8432 (NY, type).
Calyptranthes rufotomentosa McVaugh, sp. nov.
Frutex, panicula rufo-tomentosa et foliis elongatis glabris; foliis oblanceolatis,
15-21 cm. longis, nervo medio supra elevato, petiolo incrassato; panicula circiter
50-flora, 6-7 cm. longa; hypanthio urceolato, ad florendi tempus 4 mm. longo.
A shrub, glabrous except for the heavy dark red tomentum which completely
covers the inflorescence; leaves narrow, and narrowly acuminate, 4-5 cm. wide,
4 times as long as wide, the straight marginal vein and the rather widely spaced
10-15 pairs of lateral veins conspicuous on the lower surface; flowers clustered
toward the tips of the panicle branches; stamens about 75; style apparently
undeveloped in the flowers examined; calyptra conic, apiculate, 1 mm. long.
Brazil, Amazonas: Sao Paulo de Olivenca, caatinga de arvores
baixas em areia branca, quasi pura, Oct. 15, 1942, A. Ducke 2240
Calyptranthes sessilis McVaugh, sp. nov.
Frutex, glaber, foliis sessilibus ovatis 9-12 cm. longis acuminatis; venis supra
impressis; floribus in paniculis angustis perpaucis; alabastris fusiformibus 7 mm.
Buds 2 mm. thick, the operculum 4-4.5 mm. long; style 8.5 mm. long; stamens
60-75; fruit globose, 7 mm. in diameter.
This species is similar to C. spruceana Berg, but in that species
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 surface, and the fruits are more clustered toward the tips of
Peru, Loreto: Yurimaguas, L. Williams 4583 (F 623440, type),
Killip & Smith 27603 (NY; US). Univ. of Mich. Neg. 446.
184 FIELDIANA: BOTANY, VOLUME 29
Calyptranthes simulata McVaugh, sp. nov.
Frutex vel arbor, pubescens; foliis ovatis lanceolatisve 9-15 (-25) cm. longis
subglabris; venis utroque latere 20-25; paniculis binis e ramo compresso abortive
1 cm. longo oriundis; paniculae ramis infimis usque ad 3-5 cm. longis; alabastris
2-2.5 mm. longis obovatis, apice rotundatis vel brevi-apiculatis; hypanthio
A shrub or tree to 9 meters high, belonging to the group of species with
middle-sized and nearly glabrous leaves and many small flowers in much-branched
panicles. The calyptra is explanate, 1.3-2 mm. wide; style 5-6 mm. long; stamens
Known only from specimens in which the flowering panicles
are borne in the terminal axils of the twigs, on old wood. The normal
size and shape of the leaves on sterile twigs cannot be determined
with certainty, nor can the presence or absence of wings on the
branchlets be verified. It is possible that Calyptranthes simulata
and C. tessmannii are conspecific, but on the basis of the flowering
specimens at hand the two species are readily separable by the use
of the key characters.
Peru, Loreto: Iquitos, Aug. 2-8, 1929, Kittip & Smith 27352
(NY; US), 26916 (F; NY; US); Iquitos, G. Tessmann 5372 (G; NY).
Peru-Colombia boundary: forest near Rio Putumayo, Sept. 26-Oct.
10, 1930, G. King 1614 (F; GH; MICH, type; NY; US).
A collection from Tarapoto, San Martin, Peru, Wittiams 6539
(F), may be the same species. It bears immature globose or oblate
fruits about 1 cm. in diameter; the pubescence is exactly that of
C. simulata and the leaves are similar in shape to those of that
species. The specimen suggests C. tessmannii, however, in that the
lateral veins hardly exceed 15 in number, and the panicle arises
laterally from the base of a leafy shoot.
Calyptranthes tessmannii Burret, in herb., ex McVaugh,
Arbor vel frutex, pubescens; foliis ellipticis, 8-11.5 cm. longis, subglabris,
venis utroque latere ca. 15; paniculis duabus solitariis, oppositis, e nodo infimo
rami lateralis foliosi oriundis; paniculae ramis infimis 1.5 cm. longis; alabastris
3-3.5 mm. longis fusiformibus, apice conspicue apiculatis; hypanthio strigoso.
A species which appears distinctive because of the narrow panicles arising
from the lowest node of a leafy shoot; the relatively narrow and fusiform buds
with long-apiculate tips are also characteristic. The style is about 6 mm. long;
stamens about 50. Flowering specimens are readily distinguished from Calypt-
ranthes multiflora, C. ruiziana and C. simulata by the key characters.
Peru, Loreto : Stromgebiet des Maranon von Iquitos . . . am
Pongo de Manseriche, G. Tessmann 4832 in 1924 (G, type). F.M.
McVAUGH: TROPICAL AMERICAN MYRTACEAE 185
3. MYRCIA DC.
1. Leaves sessile, cordate,
la. 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 (Berg) McVaugh.
la. 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 McVaugh.
1. Leaves definitely petiolate, cuneate or rounded at base or occasionally
2. 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.
3. Hairs of the branchlets reddish brown, up to 3.5 mm. long; calyx-lobes 4,
the larger ones 6 mm. long M. huallagae McVaugh.
3. Hairs of the branchlets yellowish brown, up to 2 mm. long; calyx-lobes 5,
rarely 4, the larger ones 2.5-4 mm. long M. bracteata (Rich.) DC.
2. Inflorescence with small inconspicuous bracts which are deciduous usually
before the flowers open; calyx-lobes 3 mm. long or usually much less, rounded
to subtruncate or triangular, mostly as wide as, or wider than, long;
4. Summit of the ovary and interior of the prolonged and cuplike hypanthium
glabrous; fruit usually globose, 5-6 mm. in diameter. [This includes 7 species
which were assigned by Berg to Aulomyrcia, and one additional species,
Myrcia bipennis (Berg) McVaugh, which is treated below.]
4. Summit of the ovary and usually the interior of the hypanthium hairy;
hypanthium variously or not at all prolonged beyond the ovary, the center
of the flower thus flat or variously depressed; fruit various.
5. Outer surface of the hypanthium glabrous, the entire plant glabrous or nearly
so; fruiting hypanthium prolonged into a neck 2 mm. long at the apex of
the fruit M. subglabra McVaugh.
5. Outer surface of the hypanthium hairy, usually strongly so; fruiting hy-
panthium, as far as known, not prolonged into a neck.
6. Inflorescence slender, the axis often terete or, if compressed, mostly less than
1.5 mm. wide (measured just below the lowest branches); small-flowered
species, the disc 1-2 (rarely to 2.5) mm. wide; fruit ellipsoid or oblong, 5-9
mm. long. [This includes M. poeppiyiana Berg, M. paivae Berg, M. splendens
(Sw.) DC., M. sylvatica (Mey.) DC., and slender forms of M. fallax (Rich.)
6. Inflorescence relatively stout, the axis and branches compressed or strongly
flattened; axis at least 1.5 mm. wide and often 2.5-4 mm. wide (measured
just below the lowest branches); flowers small or large; fruit various.
7. Calyx in bud and in fruit truncate, the lobes one-third to one-half as long
as wide, together forming a nearly straight-margined band or crown en-
circling the summit of the hypanthium; hypanthium prolonged up to 2 mm.
beyond the summit of the ovary. [Includes 2 species, both referred to
Aulomyrcia by Berg: A. chilensis Berg, A. neesiana (DC.) Berg.]
7. Calyx-margin not truncate, the lobes together not forming a recognizable
unit but individually spreading, with evident sinuses between them; indivi-
dual lobes broadly rounded or triangular, as long as wide, or a little wider
than long; hypanthium various, usually not prolonged.
186 FIELDIANA: BOTANY, VOLUME 29
8. Inner surface of the calyx-lobes glabrous or with a few scattered hairs, often
with large glandular dots; small-flowered species, the disc 1.3-2 (rarely 2.5)
mm. wide; style (as far as known) 5 mm. long or less.
9. Upper leaf surface with 5-8 large convex glandular dots per square milli-
meter; principal veins impressed M. perlusa DC.
9. Upper leaf surface without apparent glands, or with minute impressed glands;
veins not impressed, or the laterals or midvein somewhat so.
10. Upper leaf surface thickly but minutely impressed-puncticulate; lateral veins
(15-) 20-25 pairs; inflorescence often with the principal branches from near
the base M. fascicularis Berg.
10. Leaves not impressed-puncticulate; lateral veins 10-15 pairs; inflorescence
usually not branching from near the base.
11. Lateral veins impressed, 10-12 pairs, much stronger than the intermediate
ones; midvein hairy, flat or somewhat convex above; inflorescence hirsute
with yellowish hairs up to 1 mm. long; hypanthium elongate in bud, with
about 10 narrow longitudinal ridges M. ambivalens McVaugh.
11. Lateral veins not impressed, often slightly elevated in drying, about 12-15
pairs, these often little stronger than the intermediate ones; midvein gla-
brous or pubescent, sulcate or slightly impressed; inflorescence sparingly
short-pubescent (rarely hirsutulous) ; hypanthium narrowly conic in bud,
not ridged. [Includes M. fallax (Rich.) DC., and vigorous forms of M.
splendens (Sw.) DC.]
8. Inner surface of the calyx-lobes appressed-pubescent; small- or large-flowered
species; style various.
12. Lower leaf surface finely bullate; pubescence of inflorescence soft, ochraceous
or rusty M. deflexa (Poir.) DC.
12. Lower leaf surface smooth or verruculose, not bullate; pubescence various.
13. Branches of the inflorescence, and the hypanthium, abundantly tomentose
or hirsutulous with reddish hairs.
14. Inflorescence rufous-tomentose; leaves 10-15 cm. long, broadly and bluntly
acuminate, or obtusely pointed, lustrous and nearly glabrous beneath, with
prominent large glands M. atrorufa McVaugh.
14. Inflorescence pale rufous-hirsutulous with hairs of different lengths inter-
mixed; leaves 12-26 cm. long, abruptly acuminate, dull and pale beneath,
eglandular but with numerous appressed hairs. .M. egensis (Berg) McVaugh.
13. Branches of the inflorescence, and the hypanthium, with white or yellow,
appressed or spreading, mostly straight hairs; hairs sometimes confined to
the hypanthium and then occasionally reddish or tawny.
15. Lateral veins, and often the marginal vein also, impressed on the upper
leaf surface [see also M. madida, a large-leaved species in which the lateral
veins may dry as fine lines slightly below the surface level].
16. Leaves hirsute beneath, at least on the veins, with soft yellow hairs up to
1 mm. long; inflorescence similarly hirsute with spreading or ascending hairs;
lateral veins about 10 (rarely -15) pairs; disc 3-3.5 mm. wide.
M. mollis (HBK.) DC.
16. Leaves sparingly appressed-pubescent beneath, or with a few long hairs on
the veins; inflorescence sparingly hispidulous to appressed-pubescent with
short hairs; lateral veins mostly more than 15 pairs.
17. Leaves rounded, obtuse, or emarginate at apex, middle-sized (mostly less
than 15 cm. long), relatively broad (mostly not more than twice as long as
17a. Calyx-lobes about 1 mm. long; leaves rounded or emarginate at apex, 1.3-1.5
times as long as wide; inflorescence and branchlets sparingly hairy.
M . fasciata McVaugh.
McVAUGH: TROPICAL AMERICAN MYRTACEAE 187
17a. Calyx-lobes 2-2.5 mm. long; leaves rounded to obtuse or obscurely apiculate,
about twice as long as wide; inflorescence and branchlets velutinous.
M. dispar McVaugh.
17. Leaves markedly acuminate or acute, mostly larger (16-27 cm. long), rela-
tively narrow (mostly 2-4 times as long as wide).
18. Leaves bullate above, between the 15-20 pairs of ladder-like veins; petiole
1.5-2 mm. long; lateral veins prominent beneath near the mid vein only;
small-flowered species, the disc 1.5-2 mm. wide M. fenestrata DC.
18. Leaves flat or nearly so in drying; petioles 3-8 mm. long; lateral veins promi-
nent their whole length; large-flowered species, the disc 3-4 mm. wide.
19. Leaves appressed-pubescent beneath; midvein sulcate above; flowers aggre-
gated on very short angular branchlets; hypanthium strongly angled in
drying, whitened by thick felted hairs M. pentagona McVaugh.
19. Leaves nearly glabrous and markedly verruculose beneath; midvein forming
a narrow ridge above; inflorescence loosely branched, the flowers mostly in
threes on slender branchlets; hypanthium conic, strigose.
M. crassimarginata McVaugh.
15. Lateral and marginal veins not impressed on the upper surface, usually
slightly raised in drying.
20. Leaves hirsute beneath, at least on the veins, with soft yellow hairs up to
1 mm. long; inflorescence similarly hirsute with spreading or ascending hairs;
disc (2.5-) 3-3.5 mm. wide; lateral veins about 10 (rarely -15).
M. mollis (HBK.) DC.
20. Leaves appressed-pubescent or nearly glabrous beneath; inflorescence ap-
pressed-pubescent or short-hirsutulous with hairs less than 0.5 mm. long;
disc and veins various.
21. Flowers aggregated near the tips of the panicle branches, those of the terminal
clusters sessile or on very short pedicels up to 1 mm. long and nearly as thick;
disc 3 mm. wide or less; lateral veins 15-20 pairs.
22. Hypanthium whitened with very short pale felted hairs; disc 2.5-3 mm.
wide; leaves 18-23 cm. long; buds 4-4.5 mm. long. . . .M. madida McVaugh.
22. Hypanthium covered thinly by appressed short silky hairs; disc 2-2.5 mm.
wide; leaves 7-14 cm. long; buds 3 mm. long M. concava McVaugh.
21. Lateral flowers of the terminal clusters on slender pedicels 2-6 mm. long;
disc 3-4 mm. wide; leaves 16 cm. long or less, with 10-15 pairs of lateral veins.
23. Branches of the inflorescence short-hirsutulous; marginal vein about as
strong as the laterals and strongly arched between them.
M. albobrunnea McVaugh.
23. Branches of the inflorescence appressed-strigose or nearly glabrous; lateral
veins diminishing distally and usually not forming a distinct marginal vein.
M. dichasialis McVaugh.
Myrcia albobrunnea McVaugh, sp. nov.
Arbor, ochraceo-hirsutula, foliis acuminatis 9-16 cm. longis; venis supra vix
manifestis; nervo marginali prominente; paniculis multifloris, rhachi valida 1.5-
2.5 mm. lata, floribus ultimis tenuiter pedicellatis; calycis lobis intus extusque
pubescentibus; germine hirsute, disco 4 mm. lato.
A tree 3-8 meters high, the ovate leaves with 10-15 pairs of lateral veins;
flowers large, the style 7.5-8 mm. long, the stamens more than 200. Flowers
(according to Klug) white and brown. The center of the flower is much depressed
(i.e., the hypanthium is prolonged about 1 mm. beyond the summit of the ovary),
as in the supposed genus Aulomyrcia.
188 FIELDIANA: BOTANY, VOLUME 29
This species strongly resembles M. dichasialis, which is a native
also of the vicinity of Iquitos, and it is with some hesitancy that
I describe them both as new. Myrcia albobrunnea differs from
M. dichasialis in having the branchlets markedly hirsutulous in-
stead of glabrous or sparingly appressed-strigose. The flowers in
M. albobrunnea are slightly larger, the leaves are less markedly
reticulate on the upper surface, the midvein is hairy above, and the
marginal vein is well defined although strongly arched between the
laterals; the lateral flowers of the terminal triads are somewhat
more conspicuously pedicellate in M. dichasialis.
Peru, Loreto: Iquitos, elev. 100 meters, woods, Killip & Smith
27006, Aug. 3-11, 1929 (F; US) ; Mishuyacu, elev. 100 meters, forest,
Feb.-Mar., 1930, G. King 1030 (F; US 1456120, type).
Myrcia ambivalens McVaugh, sp. nov.
Frutex, ramulis paniculisque pilis flavidis, flexuosis, usque ad 1 mm. longis
obsitis; foliis 5-12 cm. longis, acuminatis; paniculis multifloris validis; nervo
medio supra piano vel vix elevato; venis utroque latere 10-12, supra impressis;
calycis lobis intus glabris; disco piloso, ut videtur 2 mm. lato; hypanthio circiter
Leaves ovate, elliptic or obovate, nearly concolorous and eglandular, slightly
lustrous above. The type is in bud; the flowers are small: buds 4 mm. long, the
calyx-lobes 1-1.5 mm. long, the stamens about 200.
A distinctive species because of the indument, the comparatively
strong and few veins of the leaves, and the elongate and sulcate
Peru, Loreto: Yurimaguas, in forest, Nov. 7, 1929, L. Williams
4706 (F), 4737 (F 624825, type). Univ. of Mich. Neg. 473.
Myrcia atrorufa McVaugh, sp. nov.
Frutex, rufo-pubescens vel -tomentosus; foliis obtuse acuminatis vel obtusis,
subtus lucidis, conspicue glandulosis; paniculis validis, rhachi usque ad 2.5 mm.
lata; calycis lobis intus pubescentibus; disco 2.5-3 mm. lato, piloso, concavo.
A shrub 2 meters high with ovate leaves 10-15 cm. long, the foliage distinctive
because of the usually bluntly pointed tips of the blades, the coarse reticulum
formed by the small veins on the upper surface, and the prominent large glands
beneath. The crisped rufous hairs of the branchlets and inflorescence are unlike
those of other Peruvian species. Style probably 4 mm. long; stamens 125-150;
fruit subglobose, about 1 cm. in diameter.
Peru, Huanuco: Between Huanuco and Pampayacu, R. Kanehira
17, Jan. 30, 1927 (A; F; GH). Puno: Prov. Carabaya, trail from
Santo Domingo to Chabuca mine, elev. 1,900 meters, moist open
McVAUGH: TROPICAL AMERICAN MYRTACEAE 189
places with dense vegetation, much fog and rain, R. D. Metcalf
30671, May 30-June 1, 1942 (US 1876047, type).
Myrcia bipennis (Berg) McVaugh, comb. nov. Myrciaria
bipennis Berg, Linnaea 31: 259. ?1862.
A most distinctive species, but quite out of place in the genus
to which Berg assigned it. The seeds in the recent collections made
in Amazonian Brazil by Holt & Blake (no. 556) and in Venezuela by
Williams (no. 14763) are definitely myrcioid. No flowering material
was available to Berg, and apparently none has been collected since
his time; the immature buds on the type specimens (Spruce, no.
3770) suggest the genus Marlierea, and the morphology of the
inflorescence suggests Marlierea more than Myrcia. The hypan-
thium on the mature fruit, however, is straight-sided within and
with a definite rim (as in the Aulomyrcia group of Myrcia), and the
calyx-lobes are well formed, not irregularly split at base as in
Myrcia concava McVaugh, sp. nov.
Arbor, paniculis multifloris, strigosis, 7-11 cm. longis, rhachi valida, usque
ad 2 mm. lata; foliis acuminatis, 7-14 cm. longis; venis utrinque prominulis, non
impressis; floribus subsessilibus, germine piloso, disco 2-2.5 mm. lato, hypanthio
extus sericeo-strigoso, calycis lobis intus pubescentibus.
A tree 6-8 meters high with elliptic or ovate leaves about 2.5 times as long
as wide; buds 3 mm. long; hypanthium-base not attenuate, but abruptly con-
tracted into the pedicel; style 4.5 mm. long; stamens about 150.
A species which is separated by no very obvious characters from
those of the Myrcia fallax-splendens complex, but which differs from
these in its short, stout inflorescence, its conspicuous calyx-lobes,
which are pubescent within and concave and spreading as the petals
fall, and its sparingly strigose and broad-based hypanthium.
Peru, Loreto: Mishuyacu, near Iquitos, forest, elev. 100 meters,
G. King 454 (F; NY; US), 800 (F; NY; US 1455791, type).
Myrcia connata McVaugh, sp. nov.
Frutex 5-6 m. altus, strigosus; foliis sessilibus lanceolatis, 9-16 cm. longis,
cordatis; nervo medio supra impresso; venis utroque latere 12-15, inconspicuis;
paniculis paucifloris validis; calycis lobis intus pilosis; disco 3 mm. lato, piloso,
A shrub 5-6 meters high, the branchlets, leaf-buds and inflorescence (es-
pecially the calyx and hypanthium) strigose with straight yellowish hairs up to
0.5 mm. long; at least the young leaves sparingly strigose beneath; leaves sessile,
lanceolate, 3.5-5 cm. wide, 9-16 cm. long, about 2.5-4 times as long as wide, acute
190 FIELDIANA: BOTANY, VOLUME 29
or obscurely acuminate, blunt at very tip, cordate at base, the auricles of opposite
pairs overlapping; midvein impressed above, prominent beneath; lateral veins
12-15 pairs, slender, inconspicuous on both sides, more prominent beneath;
marginal vein about equaling the laterals and arched between them, 3-5 mm.
from the margin; small veins markedly reticulate in prevailingly right-angled
patterns; blades somewhat lustrous above, paler and dull beneath, the glandular
dots scarcely apparent; inflorescence a panicle 4-7 cm. long, about 12-flowered,
once- or twice-compound, the peduncle 2.5-4.5 cm. long, 2 mm. wide below the
first node; lower branches of the panicle up to 1.2 cm. long; flowers sessile; buds
4-6 mm. long, the hypanthium about 2 mm. long, heavily appressed-hairy;
calyx-lobes 5, broadly rounded, appressed-hairy on both sides, 1.3-1.5 mm. long,
2.5 mm. wide; disc about 3 mm. wide, sunken, the hypanthium prolonged 1-1.5
mm. beyond the summit of the ovary; style probably about 10 mm. long; stamens
150-200, 7-8 mm. long, the anthers 0.6 mm. long; petals hairy outside, about
7-8 mm. long and 5 mm. wide; fruit fleshy, about 1 cm. in diameter, a little longer
Bolivia, La Paz: Prov. S. Yungas, basin of Rio Bopi, San Barto-
lom< (near Calisaya), elev. 750-900 meters, July 1-22, 1939, B. A.
Krukoff 10382 (NY, type; US; Y).
Myrcia crassimarginata McVaugh, sp. nov.
Arbor, ramulis paniculisque pilis brevibus ochraceis puberulentis, ramulis
novellis velutinis; foliis acuminatis, subtus minute siliceo-verruculosis, venis
impressis; nervo marginali prominente, nervo medio supra elevato planiusculo;
paniculis multifloris validis, rhachi usque ad 3 mm. lata; floribus ultimis tenuiter
pedicellatis; germine piloso; disco 3-4 mm. lato; calycis lobis intus strigosis.
A tree 5 meters high, the leaves elliptic, 16-20 cm. long, 2-2.5 times as long
as wide; the foliage is characteristic because of the prominent and strongly im-
pressed veins, and the ridge formed by the midvein on the upper leaf-surface.
Flowers large, the buds 5 mm. long; style 6-8 mm. long; stamens about 300.
A collection made by Tessmann (no. 4319), at the mouth of
Rio Santiago, on high land, is apparently the same species, but in
the specimen seen (at NY) the leaves are appressed-pubescent and
smooth beneath (not verruculose) .
Peru, Loreto: Fortaleza, near Yurimaguas, forest, elev. about
140 meters, Dec., 1932, G. King 2821 (A, type; F; G; GH; NY; US).
Myrcia dichasialis McVaugh, sp. nov.
Arbor, foliis ovatis brevi-acuminatis 8-15 cm. longis, venis supra non impressis,
longiori a margine distantia arcuatim unitis; paniculis multifloris validis, rhachi
2-2.5 mm. lata, ramulis appresse-strigosis vel subglabris, floribus ultimis tenuiter
pedicellatis; germine hirsute; disco 3-4 mm. lato; calycis lobis intus pubescentibus.
Perhaps a shrub, or a tree up to 10 meters high, with ovate leaves 2-3 times
as long as wide, the marginal vein usually not distinct from the laterals which
diminish distally as they arch inward and fuse with the next succeeding ones.
Flowers large; style about 7 mm. long; stamens about 200.
McVAUGH: TROPICAL AMERICAN MYRTACEAE 191
The distinctions between this species and M. albobrunnea are
mentioned above, in the discussion following the description of
Peru, Loreto: Iquitos, Mexia 6508 (F; G; US). Gamitanacocha,
Rio Mazan, J. M. Schunke 134 (A; F 997268, type; US), 188 (A;
F; US). Stromgebiet des Ucayali von 10 S. bis zur Miindung,
G. Tessmann 3411 (G). Along Rio Itaya, L. Williams 91 (F).
Myrcia dispar McVaugh, sp. nov.
Arbor velutina, paniculis ramulisque dense pilis aureo-flavidis usque ad 0.8
mm. longis obtectis; foliis obtusis vel breve apiculatis, 10-15 cm. longis, subtus
minute siliceo-verruculosis strigosisque; venis impressis; nervo marginali supra
non impresso; paniculis circiter ut videtur 20-floris, validis, rhachi usque ad 2.5
mm. lata; floribus sessilibus; germine piloso; disco 3 mm. lato; calycis lobis intus
strigosis, 2-2.5 mm. longis.
A tree said by Krukoff to be 45 feet high; leaves coriaceous, elliptic to ovate
or obovate, very inconspicuously if at all acuminate, about twice as long as wide;
panicles stout and few-flowered, notably shorter than the leaves, even in fruit
retaining considerable amounts of the indument which covers the branchlets;
fruit globose, 1-1.5 cm. in diameter, crowned by the erect calyx; flowers not seen
but apparently large for the genus.
Differs from other known species in the short, stout inflorescence,
relatively large and few flowers, coarsely veined blunt leaves, and
copious distinctive indument. It is associated in the key with
Myrcia fasciata, but it bears little resemblance to that species
except in the key characters. Its actual relationships are not
apparent to me.
Brazil, Acre: Basin of Rio Jurua, upper Rio Jurupary, Lat.
8-9 S., Long, about 70 W., on terra firma, July 15, 1933, B. A.
Krukoff 5232 (NY, type).
Myrcia egensis (Berg) McVaugh, comb. nov. Aulomyrcia
egensis Berg, in Mart. Fl. Bras. 14, pt. 1: 99. 1857. A. macrophylla
Berg, in Mart. I.e.
The type of A. egensis, Poeppig's no. 2551, from Ega, Brazil,
bears immature fruit. Except for somewhat shorter petioles it
seems not to differ significantly from a flowering specimen collected
by Klug (no. 3569) in Peru. I have not made a direct comparison
between the type of A. macrophylla and more recently collected
materials, but from notes made upon the type in Munich in 1954,
and from a photograph of it (F.M. Neg. 19818), I believe it to be
conspecific with the other two collections cited above.
192 FIELDIANA: BOTANY, VOLUME 29
Myrcia fascia ta McVaugh, sp. nov.
Arbor strigosa, foliis rotundo-ovatis, coriaceis, venis impressis; paniculae
ramulis compressis, pedunculo superne usque ad 3-4 mm. lato; germine piloso;
disco circiter 2.5 mm. lato; hypanthio pilis plurimis atro-fuscis obsito; calycis
lobis intus pubescentibus.
A tree to 12 meters high, with lustrous coriaceous broad leaves which are
rounded or emarginate at apex; flowers about 25, in small groups near the tips
of the much-flattened panicle branches; mature flowers not seen, but the buds
2 mm. long; fruit subglobose, about 1 cm. in diameter.
Known only from the following collections, one of which bears
very immature buds, and the other nearly mature green fruit.
Ecuador, Azuay: Forested slopes between Cruz Pamba and Loma
de Canela, in region of Rio Sadracay, tributary of Rio Mehuir,
north of Molleturo, elev. 2,315-2,500 meters, June 12, 1943, J. A.
Steyermark 52969 (F 1391179, type), 52961 (F).
Myrcia huallagae McVaugh, sp. nov. M. lanceolate, 7 grandi-
folia Berg, in Mart. Fl. Bras. 14, pt. 1: 155. 1857.
Frutex debilis, hirsutus, pilis rufis erectis usque ad 3.5 mm. longis obsitus;
foliis elliptico-lanceolatis, basi subauriculatis; floribus paucis, bracteis bracteo-
lisque persistentibus, ovatis, usque ad 12 mm. longis, suffultis; calycis lobis 4,
exterioribus majoribus; disco concavo, 3.5 mm. lato; M. bracteatae maxime affinis.
A shrub or woody vine up to 3-4 meters high, with leaves 8-12 cm. long, often
3 times as long as wide; the raceme-like inflorescences are up to 4 cm. long, much
shorter than the leaves; style 6 mm. long; fruit ellipsoid, 13 mm. long.
Closely akin to Myrcia bracteata and scarcely to be distinguished
from that species except by the characters given in the key and by
the somewhat larger size of most vegetative and reproductive parts.
It is here recognized as a species because it is separated from M.
bracteata by a series of tangible though perhaps minor features, and
because it occupies a restricted geographical range.
Peru, Loreto: Yurimaguas, Kittip & Smith 27976 (US), 28188
(US), L. Williams 3837 (F), 3888 (F), 4715 (F). In sylvis ad Hua-
llagam, Feb., 1831, Poeppig 2267 (W, type, and type of M. lanceo-
lata 7 grandifolia) .
Myrcia tnadida McVaugh, sp. nov.
Arbor, minute puberula, foliis acuminatis, 9-23 cm. longis; venis utrinque
prominulis, non impressis; paniculis multifloris, validis, rhachi compressa, usque
ad 3.5 mm. lata; floribus subsessilibus, germine piloso, disco 2.5-3 mm. lato,
hypanthio extus basin versus pilis brevissimis incano-sympilematis obtecto,
calycis lobis intus pubescentibus.
A tree to 12 meters high, nearly glabrous but with very small appressed pale
hairs which are numerous and densely aggregated on the leaf-buds and the base
McVAUGH: TROPICAL AMERICAN MYRTACEAE 193
of the hypanthium, whitening the surface and in drying somewhat felted together
as if from moistening (madida, drenched, sodden). Leaves elliptic to ovate or
obovate, about 3 times as long as wide; style 6 mm. long; stamens about 125;
fruit black, ellipsoid, 10-13 mm. long.
Known only from the following collections:
Peru, Loreto: Rio Napo near Mazan, elev. 110 meters, over-
hanging river, Mexia 6448, Jan. 27, 1932 (F 718476, type; G; US).
Manfinfa, upper Rio Nanay, L. Williams 1088 (F).
Myrcia obumbrans (Berg) McVaugh, comb. nov. Rubachia
obumbrans Berg, in Mart. Fl. Bras. 14, pt. 1: 28. 1857. Marlierea
obumbrans (Berg) Niedz. in Natiirl. Pflanzenfam. Ill, pt. 7: 76. 1893.
Berg assigned this species to Rubachia because of his observation
that after an thesis the calyx split longitudinally below the sinuses.
I have not been able to confirm this, but the plant bears marked
resemblances to other Peruvian species of Myrcia and seems to have
little in common, either morphologically or geographically, with the
6 south-Brazilian species of Rubachia. The genus Rubachia is in-
deed of doubtful validity; Bentham (Benth. & Hook. f. Gen. PI. 1:
717. 1865) relegated it to the synonymy of Marlierea; Niedenzu, as
noted above, made the formal transfer of Rubachia obumbrans to
Marlierea. On the basis of the material which I have seen (Killip
& Smith 28180; Williams 4581), I judge that the calyx in this species
is that of Myrcia, not of Marlierea. These specimens, however, are
in bud only, and additional collections in flower or young fruit are
needed to confirm these observations.
Myrcia pentagona McVaugh, sp. nov.
Arbor, ramulis, paniculis et praesertim hypanthio, pilis mollibus sympilematis,
albicatis; foliis acuminatis, 20-30 cm. longis, venis impressis; paniculis multifloris,
validis, rhachi usque ad 4 mm. lata; floribus sessilibus, e ramulis subtetragonis;
germine piloso, 3-4 mm. lato; hypanthio 5-gono; calycis lobis intus pubescentibus.
A tree 5 meters high, with elliptic leaves 3-4 times as long as wide and 18-25
pairs of strong lateral veins and a strongly impressed marginal vein. The peculiar
indument and the strongly angled hypanthium are characteristic. Flowers large:
style 6-7 mm. long; stamens about 150.
Peru, Loreto: Florida, Rio Putumayo, mouth of Rio Zubineta,
elev. 180 meters, forest, May-July, 1931, G. King 2152 (F 668840,
type; G; GH; NY; US).
Myrcia splendens (Sw.) DC., var. chrysocoma McVaugh,
194 FIELDIANA: BOTANY, VOLUME 29
Arbor, M. splendenti maxime affinis, sed distincta: Indumento densiore,
longiore, pilis appressis vel ascendentibus, lucidis, sericeis, fulvis vel ochraceis,
usque ad 1 mm. longis, calycis lobos superantibus.
This is the silky-pubescent extreme of the forms of Myrcia
splendens; a somewhat less pubescent form is M. sericea Berg, and
another is the silky-strigose M. saxicola Berg. The present variety
is known only from the type collection:
Peru, San Martin: Pongo de Cainarachi, Rio Cainarachi (a tribu-
tary of the Rio Huallaga), elev. 230 meters, in forest, Sept.-Oct.,
1932, G. King 2622 (A; F; G; GH; NY; US 1457061, type).
Myrcia subglabra McVaugh, sp. nov.
Arbor, subglabra (gemmis petalisque exceptis), foliis 8-20 cm. longis, apice
graduatim acuminatis, basi marginibus decurrentibus; nervo medio supra im-
presso, venis 15-18 supra planiusculis vel elevatis; laminis utrinque lucidis, grosse
pellucido-punctatis; paniculis multifloris validis, rhachi 2 mm. lata; germine
pallide piloso; hypanthio glabro, supra germen valde (2 mm.) producto, urceolato;
calycis lobis utrinque glabris.
A tree 6 meters high, with strongly flattened branchlets and leaves elliptic-
lanceolate or narrowly ovate, 2.5-4 times as long as wide. The plant is readily
identified by the prominently glandular and reticulately veined leaves and by the
narrowly prolonged neck of the hypanthium, which exceeds in length those of all
other species of Myrcia known to me. Flowers relatively large: disc 2.5-3 mm.
wide; style 6-7 mm. long; stamens about 200; fruit ellipsoid-oblong, about 1 cm.
Thus far known only from the Mapiri region of northern Bolivia,
where it has been collected several times in the subtropical forested
areas, at elevations of 570-850 meters.
Bolivia, La Paz: San Carlos, Nov. 18, 1926, 0. Buchtien 956
(F 928708, type; GH; NY; US), 942 (F; GH; NY; US), 943 (F;
MICH; US), 944 (US), 957 (US); Charopampa, Buchtien 74 (F; G),
1909 (G); San Antonio, Buchtien 1909 (US); Copacabana, Krukoff
4. EUGENIA L. 1
1. Leaves ternate, or opposite at some nodes, 2 cm. long or less; flowers solitary,
axillary; northern Peru (Cajamarca) to Colombia E. triquetra Berg.
1. Leaves opposite, or occasionally sub-opposite.
2. (See second "2," p. 200.) Principal axis of the inflorescence racemosely
branched, the branches or flowers in decussate pairs or groups; terminal
flower of the axis usually wanting; axis elongate to extremely shortened, so
1 Three species, extra-Peruvian in range, are described below but are not
included in the key. These are Eugenia percrenata (Mato Grosso, Brazil),
E. valvata (central Ecuador) and E. variareolata (eastern Colombia).
McVAUGH: TROPICAL AMERICAN MYRTACEAE 195
that the flowers may appear to be racemose, or in axillary fascicles, umbels
or glomerules; flowers, if occasionally solitary, arising from the base (the
lowest radial node) of an abortive axillary bud or from the basal bracteate
nodes of new branchlets which are leafy above; bracteoles at base of flower
usually broad and persistent, often connate.
Racemes irregularly compound, the primary branches cymosely 3 (or
rarely 7)- flowered, or 1-flowered; hypanthium on a slender stipe (pseudo-
stalk) often longer than itself, the linear deciduous bracteoles thus well below
the flower, and the central flower of the cymose clusters apparently pedi-
cellate E. stipitata McVaugh.
Racemes simple, the primary branches (pedicels) 1-flowered; rarely the
racemes racemosely compound, or with 2-3 flowers arising together from
a node of the axis, or with 1-2 additional, subordinate racemes arising from
the lowest (tangential) nodes of the primary one; hypanthium not slender-
stipitate, the bracteoles close to and often surrounding the base of the flower.
(See second "4," p. 196.) Axis of the raceme relatively slender and elongate,
3-5 cm. long or more; if shorter, then the internodes much longer than the
diameter of the axis, and some or all of them 5 mm. long or more.
Axis of the raceme 1.5 cm. long or less, the ascending slender pedicels often
as long as, or longer than, the axis; inflorescence appressed-strigose or -silky.
[Includes E. biflora (L.) DC., E. inundata DC., and E. macrocalyx (Rusby)
Axis of the raceme 2 cm. long or more or, if shorter, the pedicels 4-7 mm.
long, widely spreading and much shorter than the axis; pubescence and
foliage various, the leaves neither mucronate nor small and obovate-cuneate.
Hypanthium glabrous without, the plants nearly glabrous, usually finely
pubescent in the inflorescence; small-flowered species with slender, loosely
Lateral veins of the leaves 12-15 pairs, wide-spreading and nearly straight,
the marginal vein nearly straight between the laterals and amply differenti-
ated from them, 2-4 mm. from the margin; inflorescence closely appressed-
pubescent, the calyx-lobes and bracteoles fimbriate-ciliate.
E. calva McVaugh.
Lateral veins 10 pairs or fewer, curved and ascending, at least the basal ones
diminishing distally and not forming a definite marginal nerve; inflorescence
glabrous, or finely hispidulous, or with minute appressed dibranchiate hairs;
calyx-lobes and bracteoles sparsely short-ciliate E. florida DC.
Hypanthium, at least the base, thickly strigose or pubescent without; plants
variously pubescent, the inflorescence usually markedly so; flowers and
Leaves rounded or obtuse at tips, broadly elliptic-ovate, 7 cm. long or less;
plants densely ferruginous-tomentose; flowers up to 4 pairs in a stout raceme
2 cm. long, or some flowers solitary near the base of new leafy shoots.
E. racemiflora Berg.
Leaves usually markedly acuminate, relatively narrow and more than 7 cm.
long; plants various, not ferruginous-tomentose (in E. atroracemosa rufous-
velutinous); flowers all racemose.
Bracts, bracteoles and calyx-lobes thickly beset with dark raised rounded
glands, but inconspicuously if at all pubescent.
Pubescence of appressed reddish-brown, partly dibranchiate hairs, intermixed
with shorter erect hairs; connate bracteoles forming an involucre-like cupule
2.5 mm. across; larger calyx-lobes 2.5 mm. long; style 5-6 mm. long; stamens
about 100; coastal Ecuador (and Peru?) E. pustulescens McVaugh.
196 FIELDIANA: BOTANY, VOLUME 29
10. Pubescence of appressed grayish-white hairs; cupule 4 mm. across; calyx-
lobes 4 mm. long; style 8 mm. long; stamens 250-300; Amazonian Peru.
E. polyadena Berg.
9. Bracts, bracteoles and calyx-lobes not at all glandular or inconspicuously
so, but variously and often conspicuously pubescent.
11. Midvein elevated above in a narrow median line; inflorescence slender, the
axis mostly not more than 1 mm. thick, closely appressed-pubescent with
reddish brown mostly dibranchiate hairs E. dibranchiata McVaugh.
11. Midvein impressed; inflorescence various.
12. Axis of the raceme slender, 0.5-1 mm. thick (measured just below the lowest
node); leaves mostly less than 10 cm. long; flowers small, the disc 2 mm.
wide or less.
13. Leaves tomentose beneath, sub-vernicose and rough above, with many
glandular dots E. curvipilosa McVaugh.
13. Leaves glabrous or essentially so at maturity, the upper surface smooth,
rather dull, eglandular E. limbosa Berg.
12. Axis of the raceme stouter, 1-2 mm. thick (measured just below the lowest
node); leaves mostly more than 10 cm. long; flowers larger, the disc 2.5 mm.
wide or more.
14. Lateral and marginal veins impressed above, prominent beneath.
15. Lateral veins 8-12 pairs; leaves caudate-acuminate; disc 6 mm. wide; veins
of the lower leaf surface appressed-pubescent with tawny hairs.
E. longicuspis McVaugh.
15. Lateral veins about 15-20 pairs; leaves moderately acuminate; disc 3-3.5
(-6) mm. wide; veins of the lower leaf surface velutinous with erect rufous
hairs E. atroracemosa McVaugh.
14. Lateral veins (about 15-20 pairs) and marginal vein elevated slightly but
inconspicuous on both surfaces, the blades flat; disc 2.5-3 mm. wide; lower
leaf surface, including veins, with very short, closely appressed colorless
hairs E. riparia DC.
4. (See first "4," p. 195.) Axis of the raceme much abbreviated, 2 cm. long or
usually much less; if more than 1 cm. long the nodes approximate and the
internodes 3 mm. long or less, and hardly longer than the thickness of the
stout angled rachis; flowers sometimes solitary at the lowest nodes of new
leafy branchlets, and in axillary clusters or short racemes on the same plant.
16. Leaves markedly bullate, large and broad (5-10 cm. wide), elliptic to obovate;
racemes mostly 10-18 mm. long, the axis quadrangular, with up to 8-13
pairs of slender-pedicellate flowers E. tetrasticha Berg.
16. Leaves flat, or the principal veins impressed above; blades variously shaped.
17. Outer corky layers of the petiole irregularly loosening and flaking off, the
whole much roughened and appearing 3-5 mm. thick; leaves large, 20-35
cm. long, 3-5 times as long as wide; flowers clustered on old wood, the axis
4 mm. long or less.
18. Leaves mostly oblanceolate, cordate-auriculate and subsessile (petiole 3-5
mm. long and about as thick); lateral veins about 15; midvein impressed
above; calyx-lobes 4-5 mm. long E. multirimosa McVaugh.
18. Leaves elliptic-oblong, rounded at base, the petiole 10-18 mm. long and 3-4
mm. thick; lateral veins 20-30; midvein convex above; calyx-lobes 6-8
mm. long E. tumulescens McVaugh.
17. Petiole smooth or wrinkled, not thick and exfoliating, rarely more than
2 mm. thick; leaves and flowers various.
19. Large-leaved, large-flowered, coarse species (leaves 23-35 cm. long or more;
petioles 2-4 mm. thick; buds 1-2 cm. long; anthers 1-1.8 mm. long).
McVAUGH: TROPICAL AMERICAN MYRTACEAE 197
Lateral veins 25-30 pairs; marginal vein about as strong as the laterals;
leaves about 4 times as long as wide; buds probably 2 cm. long, the sub-
orbicular calyx-lobes up to 15 mm. long; bracteoles elliptic, probably de-
ciduous E. scalariformis McVaugh.
Lateral veins 9-12 pairs, prominent and ascending but diminishing distally
and scarcely forming a marginal vein; leaves 2.3-3 times as long as wide;
buds about 1 cm. long, the triangular or oblong calyx-lobes 4-6 mm. long;
bracteoles obovate, enveloping the bud, their narrow connate bases forming
a collar about the pedicel E. myrobalana DC.
Leaves usually smaller, or with a definite marginal vein; flowers various, the
buds mostly less than 1 cm. long; bracteoles usually small and persistent.
Calyx-lobes foliaceous, elongate, erect, 7-9 mm. long at anthesis, separated
by broad rounded sinuses; leaves 9-18 cm. long, with about 10 pairs of veins;
anthers 0.3 mm. long E. macrocalyx (Rusby) McVaugh.
Calyx-lobes not or scarcely foliaceous, mostly broad, concave and imbricate;
if more than 6 mm. long the margins variously connate or overlapping and
the anthers 1-1.5 mm. long; leaves various.
Leaves with a glabrous cartilaginous margin (visible from above), this
formed by a heavy convex vein about twice as thick as the lateral veins;
blades elliptic-oblong, 6-8.5 cm. wide, 12-14 cm. long; plants velutinous
with coarse reddish-brown hairs; flowers in sessile glomerules.
E. percincta McVaugh.
Leaves without a heavy vein at the margin; plants not coarsely rufous-
velutinous; flowers various.
Calyx-lobes large (the larger ones 5-8 mm. long or more), often as broad as
long, much imbricated but distinct from the first, glabrous within; flowers
probably always in sessile clusters on old wood.
Inflorescence minutely appressed-pubescent with glistening brown hairs;
leaves papillose-roughened above; style 9-10 mm. long.
E. tenuimarginata McVaugh.
Plants glabrous except the ciliate margins of the perianth lobes and bracte-
oles; leaves smooth above; style 15 mm. long or more.
Leaves 8-14 cm. wide and about 1.5 times as long; disc 4-5 mm. wide; anthers
2-2.3 mm. long E. chartacea McVaugh.
Leaves 4-7.5 cm. wide and 3-3.5 times as long; disc 6-7 mm. wide; anthers
1-1.3 mm. long E. illepida McVaugh.
Calyx-lobes smaller (the larger ones 6 mm. long or less) or if longer (up to
9-10 mm. long), then oblong, usually hooded at the tips and loosely connate
in the bud; inner surface of calyx-lobes glabrous or pubescent; flowers
Bracteoles early deciduous (just before, or usually much before anthesis) or
wanting (unknown in E. quebradensis), subulate or linear or sometimes
lanceolate to ovate, narrowed at base, not connate.
Inflorescence, especially the hypanthium, heavily pubescent or tomentose
with gray, white, or light yellowish hairs.
Branchlets and inflorescence tomentose with crisped, matted hairs; flowers
sessile in small clusters; leaves lanceolate, 0.8-1.5 cm. wide, 5-6 times as
long; Lambayeque E. quebradensis McVaugh.
Branchlets and inflorescence strigose or velutinous, the hairs mostly straight
and ascending; flowers pedicellate; leaves ovate to elliptic or oblong.
Leaves 2-6 cm. long; flowers mostly 1-2 pairs, often from the lowest axils
of new leafy branches.
198 FIELDIANA: BOTANY, VOLUME 29
30. Bracteoles ovate or lanceolate, blunt; pedicels, hypanthium and calyx
velutinous; Junln E. barbata McVaugh.
30. Bracteoles subulate; hypanthium closely strigose, the pedicel sparsely so, the
calyx glabrate; Bolivia E. mandonii McVaugh.
29. Leaves 8-24 cm. long; flowers up to 6 pairs in axillary racemes; bracteoles
subulate; Amazon basin E. prosoneura Berg.
27. Hairs of the inflorescence red or copper color or dark purplish brown.
31. Hairs coarse, dark purplish brown; leaves narrowly elliptic, 14-24 cm. long.
E. gomesiana Berg.
31. Hairs slender, reddish or pale copper color; leaves broader, mostly less than
15 cm. long.
32. Bracts 1 mm. long, broadly rounded at tips and convex on the backs, in-
conspicuous; calyx-lobes 2.5 mm. long, pubescent except at tips, triangular,
the long points longer than the corolla in bud E. aerosa McVaugh.
32. Bracts elongate, 1.5-7 mm. long, delicate, loosely imbricated in 4 ranks
at the bases of racemes and new shoots; calyx-lobes short and broadly
rounded, 1.5 mm. long, glabrous both sides, much shorter than the corolla
in bud E. patrisii Vahl.
26. Bracteoles persistent, usually until after the fruit falls, lanceolate or broader,
broad-based, often connate and involucre-like.
33. Calyx-lobes oblong or elliptic, longer than the petals in bud, hooded and
thickened at tips, connate in bud below the middle but separating and
reflexed at anthesis, glabrous within; anthers 1-1.5 mm. long.
34. Buds 12-15 mm. long; calyx-lobes 8-10 mm. long in flower; stamens 300
or more E. schunkei McVaugh.
34. Buds 9 mm. long or less; calyx-lobes 3-6 mm. long in flower (up to 9 mm.
long in fruit) ; stamens 75-175.
35. Lateral veins about 20 pairs; buds 6 mm. long, closed at the tip; pedicels
8-20 mm. long E. hexovulata McVaugh.
35. Lateral veins 6-15 pairs; calyx-lobes distinct in the bud at least distally;
pedicels 2-8 mm. long (or a little more in fruit).
36. Buds 4.5-5 mm. long; calyx-lobes 3-3.5 mm. long; style 6-7 mm. long.
E. cuspidifolia DC.
36. Buds 6-9 mm. long; calyx-lobes 3.5-6 mm. long (to 9 mm. in fruit); style
(unknown in E. acrensis) 10 mm. long.
37. Calyx permanently velutinous; lateral veins 12-15 pairs, the basal not
strongly ascending; leaves with numerous small convex glands above.
E. acrensis McVaugh.
37. Calyx glabrate; lateral veins 6-10 pairs, the basal ones often strongly ascend-
ing and not forming a definite marginal vein; leaves eglandular and very
smooth above E. feijoi Berg.
33. Calyx-lobes various, not hooded or thickened at the tips or connate below
the middle; anthers usually less than 1 mm. long.
38. Inflorescence, including the hypanthium, appressed-pubescent, velutinous
or tomentose with reddish or reddish-brown hairs.
39. Bracteoles ovate, 1.5 mm. long, not connate.
42. Calyx-lobes 3-4 mm. long, concave, dehiscent after anthesis; buds 4 mm.
long; inflorescence and branchlets closely and finely tomentose with very
small dark red-brown hairs; leaves 16-24 cm. long, the arcuate-ascending
lateral veins scarcely forming a marginal vein E. curvivenia McVaugh.
42. Calyx-lobes 4-6 mm. long, flattened, or cucullate and reflexed or spreading
after anthesis; hairs appressed-ascending or spreading, lustrous, coppery or
pale red, up to 0.5 mm. long; marginal veins about as strong as the laterals.
McVAUGH: TROPICAL AMERICAN MYRTACEAE 199
Leaves 4.5-8 cm. long, with 7-10 pairs of lateral veins; calyx-lobes flat and
spreading after anthesis, the inner pair truncate, obovate.
E. crucicalyx McVaugh.
Leaves 15-21 cm. long, with 12-15 pairs of lateral veins; calyx-lobes reflexed
and cucullate after anthesis E. acrensis McVaugh.
Bracteoles broadly ovate to rotund, fused by the basal margins and forming
an involucre-like cupule beneath the flower.
Veins impressed above, elevated and conspicuous beneath; leaves pale and
smooth or appressed-pubescent beneath; inflorescence softly rufous-tomen-
tose; margins of the bracteoles and calyx-lobes delicate and fracturing even in
bud E. macrophylla Berg.
Veins slightly convex on both surfaces, not conspicuous on either one; lower
leaf surface glistening, irregularly cellular-honeycombed or obscured by tiny
hairs; inflorescence puberulent with short crisped hairs; bracteoles and calyx
relatively tough and unbroken even in age.
Calyx-lobes 6 mm. long; leaves 12-19 cm. long, with 12-15 pairs of lateral
veins, appearing loosely cellular-honeycombed beneath when viewed with a
lens E. heterochroma Diels.
Calyx-lobes 1.5-2 mm. long; leaves 6-13 cm. long, with 6-10 pairs of lateral
veins, the lower surface obscured by tiny glistening hairs.
E. versicolor McVaugh.
Inflorescence, particularly the hypanthium, glabrous, or pubescent with pale
white or yellowish hairs (hairs of young shoots sometimes reddish).
Leaves coarsely impressed-punctate and dark above, the lateral veins not
at all or scarcely apparent; inflorescence usually with some elongate nodes,
appressed-pubescent with pale hairs; blades 10 cm. long or less. [Including
E. biflora (L.) DC. and E. inundata DC.]
Leaves not impressed-punctate, the lateral veins usually apparent above,
the blades often more than 10 cm. long; nodes of raceme approximate.
Midvein elevated above in a sharply defined pubescent ridge; bracts 1-3.5
mm. long, 4-ranked at base of pedicels E. subterminalis DC.
Midvein impressed to convex above; bracts not 4-ranked, usually incon-
Flowers usually 2, one from each side of the axillary bud; leaves mostly 3-6
cm. long, obtuse E. punicifolia (HBK.) DC.
Flowers fasciculate or in approximate pairs in short racemes, usually 4 or
more; leaves mostly larger and acuminate.
Inflorescence, including hypanthium, densely hispidulous with erect pale
hairs about 0.1 mm. long; raceme up to 8 mm. long, with 2-5 pairs of flowers
on pedicels 2-4 mm. long; leaves elliptic, 4-7 cm. long, subcaudate-acuminate.
E. micranthoides McVaugh.
Inflorescence, at least the hypanthium, glabrous or sparingly strigose;
leaves and inflorescence various.
Branchlets, petioles and inflorescence somewhat strigose with white appressed
hairs; flowers middle-sized, the buds 5-10 mm. long, the disc 3-4 mm. wide;
leaves 9-18 cm. long, 2.5-3 times as long as wide.
Buds 7-10 mm. long; marginal vein evident, 2-3 mm. from margin; petiole
heavily strigose with hairs up to 0.5-0.8 mm. long, spirally longitudinally
furrowed and transversely wrinkled; hypanthium with a few long hairs.
E. spruceana Berg.
Buds 5-6 mm. long; marginal vein not distinct from the laterals; petiole
nearly glabrous, merely irregularly wrinkled in drying; hypanthium with
numerous very fine short hairs E. atrosquamata McVaugh.
200 FIELDIANA: BOTANY, VOLUME 29
48. Branchlets, petioles and inflorescence glabrous, or pubescent with very small
dibranchiate hairs, minute erect hairs or sparse stiff and ascending reddish
or pale hairs; flowers smaller, the buds 5 mm. long or less, the disc 1-2.5
50. Leaves elliptic-ovate, 5-6 cm. long or less; midvein flat above; plants sparingly
strigose with reddish or yellowish hairs up to 0.5 mm. long, the hypanthium
glabrous; flowers tiny, the calyx-lobes 1 mm. long, the disc 1 mm. wide;
stamens 36 or fewer E. malpighioides (HBK.) DC.
50. Leaves usually more than 5 cm. long, rarely ovate; pubescence not as above;
flowers larger; stamens 60-75 or more.
51. Flowers sessile or nearly so, in clusters on the stems, the pedicels 2 mm. long
or less (up to 4 mm. in fruit); styles (as far as known) 10 mm. long or more;
hypanthium often obconic E. nigra DC.
51. Flowers on definite slender pedicels mostly 4-10 mm. long; style mostly less
than 10 mm. long; hypanthium mostly campanulate.
52. Midvein on upper surface of leaf flat or convex (then sometimes with a shallow
median furrow near base), never narrowly impressed. [Includes E. egensis
DC., E. flavescens DC., E. dittocrepis Berg, and E. ochrophloea Diels.]
52. Midvein impressed on the upper surface, the actual vein narrow and often
obscured in the bottom of a deep fold or furrow.
53. Leaves mostly elliptic-lanceolate, long-pointed, very smooth and glabrous
above; lateral veins inconspicuous, sometimes reddish; plant nearly glabrous,
the branchlets and pedicels sometimes minutely and sparsely hispidulous.
E. schomburgkii Benth.
53. Leaves elliptic to obovate, shortly and often bluntly acuminate, the veins
usually somewhat elevated and forming an evident reticulum on the upper
surface; pubescence various.
54. Inflorescence glabrous or minutely pubescent with pale erect hairs; leaves
elliptic, blunt-pointed or obscurely acuminate E. tapacumensis Berg.
54. Inflorescence more or less appressed-puberulent with minute and partly
55. Leaves oblanceolate or obovate, 14-18 cm. long, coarsely veiny-reticulate.
E. discrete, McVaugh.
55. Leaves elliptic, 7-11 cm. long, finely veiny-reticulate.
E. quadrijuga McVaugh.
2. (See first "2," p. 194.) Flowers solitary or in 3- to many-flowered dichasia,
the terminal flower or flowers usually present and sessile; flowers, if solitary,
arising directly from the axil of a foliage leaf, not from the lower bracteate
nodes of new branches which are leafy above or from the bracteate nodes of
abortive axillary buds; bracts and bracteoles mostly linear or lanceolate,
scarious, deciduous at anthesis or often much before this.
56. Primary branches of the inflorescence racemosely arranged, i.e., in decussate
pairs (see first "3," p. 195) E. stipitata McVaugh.
56. Cymes (dichasia) arising directly from the leaf axils, sometimes irregularly
branched but the branches not in decussate pairs; bracteoles usually closely
subtending the flowers.
57. Flowers 5-merous; leaves coriaceous, yellow-green and vernicose, subsessile,
orbicular to broadly ovate, 1.5-7.5 cm. long and wide; Pacific slope, Lima.
E. quinqueloba McVaugh.
57. Flowers 4-merous; leaves various; species mostly of the inter-Andean valleys.
58. Flowers solitary (see second "58," p. 201, for plants with flowers partly
solitary and partly in threes or more numerous).
McVAUGH: TROPICAL AMERICAN MYRTACEAE 201
59. Leaves 6 mm. long or less, rigidly coriaceous, vernicose, prominently im-
pressed-punctate on both sides; flowers numerous toward the tips of the
branches, on peduncles 2-3 mm. long E. minimifolia McVaugh.
59. Leaves mostly 1-2.5 cm. long, the texture various; blades not impressed-
punctate beneath, sometimes obscurely so above; flowers relatively few, not
crowded in the terminal axils, the pedicels (or peduncles) usually much
60. Hypanthium with strong longitudinal angles; leaves often suborbicular,
grayish green and closely appressed-pubescent beneath. . .E. oreophila Diels.
60. Hypanthium not angled; leaves rarely suborbicular, neither grayish green
nor closely appressed-pubescent.
61. Hypanthium narrowly obconic, attenuate at base; pedicels 12-20 mm. long;
calyx-lobes glabrous within; leaves about 2.5 cm. long, mostly acutely
narrowed to both ends E. myrtomimeta Diels.
61. Hypanthium narrowly campanulate or broader, blunt or rounded at base
and usually broader than the markedly compressed pedicel; calyx-lobes
appressed-pubescent within (sometimes sparingly so in E. myrsinoides with
pedicels 2-6 mm. long); leaves various.
62. Leaf margins pale, cartilaginous-thickened and revolute, often irregularly
roughened and apparently denticulate because of the prominent glands;
flowers always solitary, the pedicels filiform, little compressed, scarcely 1 mm.
wide at summit, 10-20 mm. long or more E. cartilaginea McVaugh.
62. Leaf margins scarcely thickened or roughened, the glands scarcely or not
at all apparent in mature leaves; peduncle 1- or 3-flowered, usually markedly
compressed, enlarged distally and 1 mm. wide or more.
63. Branchlets coarsely appressed-pubescent but the inflorescence glabrous;
peduncle 1- or 3-flowered, if 1-flowered 5-15 mm. long, compressed, up to
1.5 mm. wide; stamens about 250; Cuzco E. indifferens McVaugh.
63. Branchlets from nearly glabrous to pubescent, if coarsely pubescent the
inflorescence not markedly less so; hypanthium densely strigose; peduncle
1- or 3-flowered; stamens 50-100.
64. Leaves mostly less than 1.5 cm. long, obovate, and cuneate at base; flowers
all solitary, on pedicels mostly 2-6 mm. long; hypanthium mostly glabrous;
northern Peru and Ecuador E. myrsinoides (HBK.) Diels.
64. Leaves 1.2-4 cm. long, broadly elliptic to ovate, rounded at base; peduncles
65. Branchlets glabrous or sparingly strigose; Ecuador and Colombia. [Includes
E. hallii Berg and E. foliosa (HBK.) DC.]
65. Branchlets densely strigose with straight appressed white or brownish hairs;
southern Peru (Arequipa) E. ferreyrae McVaugh.
58. Inflorescence a 3- to many-flowered dichasium, the central flowers usually
present and sessile (see first "58," p. 200, for plants with solitary flowers
at least in part).
66. Veins of the lower leaf-surface raised in a conspicuous reticulum, the veinlets
enlarged, with appearance of having softened and fused; terminal flowers
sub-pedicellate E. fimbriata (HBK.) DC.
66. Veins, if apparent, slender and inconspicuously reticulate; pubescence and
inflorescence various; terminal flower usually closely sessile.
67. Inflorescence usually reddish purple, glabrous or essentially so, stout, the
peduncle 2-3.5 mm. wide near summit; flowers usually 3-7, large, the style
10-11 mm. long E. rhopaloides (HBK.) DC.
67. Inflorescence green or brownish in drying, glabrous or variously pubescent;
peduncle 2 mm. wide at summit or usually less; flowers smaller, the style
8.5 mm. long or less.
202 FIELDIANA: BOTANY, VOLUME 29
68. Inflorescence 7-flowered (or the smaller ones on the same plant 3-flowered)
or repeatedly branched and many-flowered; plants usually markedly strigose
or otherwise pubescent, especially the inflorescence and hypanthium.
69. Inflorescence loosely pubescent or tomentose with soft curved or spreading
hairs; dichasia usually with more than 7 flowers.
70. Pubescence of whitish spreading hairs 0.5-1 mm. long; leaves mostly obovate,
5 cm. long or less; northern Peru (Huanuco, Cajamarca).
E. lindleyana (HBK.) DC.
70. Pubescence of rufous curved matted hairs; leaves elliptic-ovate, 6.5-11 cm.
long; Bolivia E. pearcei McVaugh.
69. Inflorescence thinly appressed-puberulent or strigose with shorter hairs;
leaves and number of flowers variable.
71. Leaves less than 2 cm. long, obovate, cuneate; flowers small (style 3.5-4.5
mm. long), in terminal clusters of 2- to 3-forked cymes; Bolivia.
E. osteomeloides (Rusby) McVaugh.
71. Leaves, if less than 2 cm. long, usually elliptic or suborbicular, rarely cuneate;
style 5-7 mm. long; dichasia axillary, 3- to 7-flowered; central Peru to
72. Branchlets thinly appressed-puberulent; leaves elliptic-oblong, 2.5-5 cm.
long; disc 3.5-4 mm. wide, the hairy staminal ring prominent; calyx-lobes
2.5-3 mm. long; stamens more than 200 E. limbata (HBK.) DC.
72. Branchlets coarsely pubescent with spreading-ascending hairs; leaves ellip-
tic to suborbicular, 0.5-2 cm. long; disc 2.5 mm. wide, the staminal ring
inconspicuous, nearly glabrous; stamens 50-60; central Peru (Ayacucho;
Apurimac) E. bifurcata McVaugh.
68. Inflorescence 3-flowered (or if occasionally 7-flowered, glabrous) or the
flowers solitary in some or all of the axils; hypanthium glabrous to variously
73. Hypanthium with strong longitudinal angles; leaves often suborbicular,
grayish green and closely appressed-pubescent beneath; Cuzco.
E. oreophila Diels.
73. Hypanthium not angled; leaves various, rarely suborbicular, neither grayish
green nor closely appressed-pubescent.
74. Branchlets nearly or quite glabrous, sometimes thinly strigose; northern
Peru (Libertad) to Colombia. [Includes E. compressa (HBK.) DC., E.
discolor (HBK.) DC., E. foliosa (HBK.) DC., and E. hallii Berg.]
74. Branchlets with numerous appressed or spreading stiff hairs up to about
0.5 mm. long; southern Peru (Arequipa; Cuzco).
75. Inflorescence glabrous; stamens about 250; Cuzco. .E. indifferens McVaugh.
75. Inflorescence rather densely strigose; stamens about 50; coastal hills, Are-
quipa E. ferreyrae McVaugh.
Eugenia acrensis McVaugh, sp. nov.
Frutex, gemmis racemisque pilis lucidis cupreis, adpressis obtectis; foliis 15-21
cm. longis acuminatis, venis utroque latere 12-15; racemis abbreviatis; bracteolis
persistentibus, ovatis nee connatis, 1.5 mm. longis; calycis lobis intus glabris, in
alabastro corollam cooperientibus, ad florendi tempus reflexis cucullatisque.
A shrub 3.5 meters high, nearly glabrous except for the coppery-velutinous
inflorescence and vegetative buds; leaves elliptic, 15-21 cm. long, 2.5-3 times as
long as wide, with somewhat impressed veins, the marginal vein about as strong
McVAUGH: TROPICAL AMERICAN MYRTACEAE 203
as the laterals and arched between them; calyx at anthesis splitting in the sinuses
down to the level of the staminal ring; stamens about 200; anthers 1.2 mm. long.
This species, with E. cuspidifolia DC. and E. feijoi Berg, belongs
to a somewhat ill-defined group characterized by elongate anthers
and hooded and partially or completely connate calyx-lobes which
are reflexed and cucullate after anthesis. According to Amshoff
(Rec. Trav. bot. ne'er!. 42: 21. 1949, and in Pulle, Flora of Suriname 3,
pt. 2: 58, 105. 1951), these species comprise the genus Catinga Aubl.
Plants of this affinity are rather abundant in the upper Amazon
Basin, but the available collections are few, and the taxonomy of the
group is in need of study and clarification. The degree of coherence
of the calyx-lobes in the bud varies from almost none in the present
species to almost complete union in the newly described E. hex-
ovulata, and in specimens identified by Amshoff as Catinga moschata
Brazil, Acre: Basin of Rio Purus, near mouth of Rio Macauhan
(tributary of Rio Yaco), Lat. 9 20' S., Long. 69 W., on terra firma,
Aug. 22, 1933, B. A. Krukoff 5619 (NY, type).
Eugenia aerosa McVaugh, sp. nov.
Arbor, ramulis gemmis paniculisque, pilis pallide cupreis, adpressis obtectis;
foliis 13-15 cm. longis acuminatis, venis utroque latere 8-10; racemis abbreviatis;
bracteolis deciduis; calycis lobis deltoideis, 2.5 mm. longis, ut videtur valvatis, ad
florendi tempus reflexis.
A tree up to 15 meters high with elliptic-oblong leaves 2.3-3 times as long
as wide, resembling E. patrisii but lacking the 4-ranked imbricated bracts of that
species; buds 6 mm. long, pyriform, the long-pointed calyx-lobes exceeding the
globe of the petals until anthesis; style 10 mm. long; stamens about 100; fruit
ellipsoid, 2.3 cm. long.
The single fruiting specimens, probably belonging to this species,
are from Krukoff's no. 6870, from Livramento, Amazonas, Brazil.
Peru, Loreto: Mishuyacu, near Iquitos, forest, elev. 100 meters,
May-June, 1930, G. King 1535 (F 627573, type; NY; US). Iquitos,
Aug., 1925, G. Tessmann 5355 (fragment, F; G; NY).
Eugenia atroracemosa McVaugh, sp. nov.
Arbor, rufo- vel atro-velutina, foliis 11-18 cm. longis acuminatis, venis
utroque latere circiter 20; racemis 5-7.5 cm. longis validis, floribus 12-20; disco
3-3.5 mm. lato.
A tree 4 meters high with oblong, ovate or oblanceolate leaves mostly 2-3
times as long as wide, readily recognized by the stout reddish brown or almost
black velutinous racemes; flowers moderately large, the buds 7 mm. long; calyx-
lobes 2.5-3 mm. long; style 6-6.5 mm. long; stamens 125-150.
204 FIELDIANA: BOTANY, VOLUME 29
Peru, San Martin: Zepelacio, near Moyobamba, elev. 1,100-1,200
meters, Oct.-Nov., 1933, G. Klug 3368 (A; F 736442, type; GH; NY;
US). Brazil, Amazonas: Sao Paulo de Olivenca, Krukoff 8738 (NY).
Eugenia atrosquamata McVaugh, sp. nov.
Arbor vel frutex, appresse pubescens, ramulis petiolisque et racemis, pilis
albidis usque ad 0.2 mm. longis obsitis; foliis 9-15 cm. longis obtuse acuminatis;
nervo medio supra planiusculo vel concavo; venis utroque latere 6-10, supra
planiusculis vel convexis; racemis abbreviatis; alabastris 5-6 mm. longis; calycis
lobis inaequalibus, exterioribus 3.5 mm. longis, 4 mm. latis; bracteolis persistenti-
bus glabris, ovatis, 1.5 mm. longis et latis.
A shrub or tree with elliptic leaves about 2.5 times as long as wide, the rela-
tively few lateral veins ascending but usually not forming a definite marginal vein.
Flowers 4-6 pairs, middle-sized, with dark contrasting bracts and bracteoles;
disc 3 mm. wide; style 7 mm. long; stamens about 100.
This plant is contrasted in the key with E. spruceana Berg, which
it somewhat resembles, but its actual relationships are obscure
Peru, Loreto: Iquitos, May, 1925, G. Tessmann 5130 (G, type;
NY). Colombia, Amazonas: Trapecio amazonico, Loretoyacu
River, elev. 100 meters, March, 1946, R. E. Schultes 7133 (US).
Univ. of Mich. Neg. 439.
Eugenia barbata McVaugh, sp. nov.
Ut videtur frutex, ramulis hispidulis, floribus cinereo-flavido-velutinis; foliis
3-5 cm. longis, acutis vel obtusis, venis inconspicuis; racemis abbreviatis, floribus
plerumque 2, interdum 4; bracteolis deciduis, obtusis; alabastris 9-10 mm. longis
Probably a shrub, 4-8 meters high with coriaceous broadly elliptic leaves and
markedly hairy, rather large flowers in small nearly sessile clusters or solitary
at the lower nodes; calyx-lobes suborbicular, 4-5 mm. long; style 9-10 mm. long;
stamens about 250.
This plant superficially resembles E. pungens Berg, a species
which ranges from the lowlands of Bolivia to Uruguay; in E. pungens,
however, the leaves are normally aristate, and the pubescence is
Peru, Junin: Valle del Rio San Bernardo, elev. 2,200-2,300
meters, en monte bajo, abierto, compuesto de arbustos, Apr. 2,
1913, A. Weberbauer 6558 (F; GH; US 1497251 pro parte, type;
USM). Pariahuanca, elev. 1,800 meters, bosque compuesto de
arboles y arbustos, Apr. 8, 1913, Weberbauer 6594 (F; GH; US;
McVAUGH: TROPICAL AMERICAN MYRTACEAE 205
Eugenia bifurcata McVaugh, sp. nov.
Frutex parvifolius, appresse pubescens vel strigosus; foliis subrotundatis,
0.5-2 cm. longis; floribus subcorymbosis, dichasiis 3-7-floris; stylo 5-7 mm.
longo; disco 2.5 mm. lato, fere glabro; staminibus 50-60.
A shrub to 3 meters high, with elliptic to suborbicular, ovate or obovate
leaves up to 1.5 times as long as wide, the blades coriaceous and obscurely veined.
The rather slender, several-flowered, corymbosely aggregated dichasia 2-3 cm.
long are distinctive. Flowers small, the buds 3-4 mm. long.
This species is superficially similar in flower and inflorescence
characters to E. osteomeloides (Rusby) McVaugh, a Bolivian species,
but differs in a number of individually trivial respects, the most
notable of which are contrasted above in the key to species.
Peru, Apurimac: Valley of the Rio Pampas, Lat. 13 20' S. to
13 30' S., elev. 2,600 meters, A. Weberbauer 5842 (F; GH; US
1497208, type) ; Pincos, in rain-green shrubland, elev. 2,700 meters,
Feb. 19, 1939, Stork & Horton 10678 (F). Ayacucho: Hills from
River Pampas to Ocros, elev. 9,000-10,000 feet, R. Pearce (BM).
Eugenia calva McVaugh, sp. nov.
Arbor, appresse pubescens, glabrescens, floribus glabris; foliis 9-15 cm. longis
acuminatis, venis utroque latere 12-15, nervo marginali aperto; racemis 4.5-5.5
cm. longis; alabastris 2-3 mm. longis glabris, calycis lobis fimbriatis, intus pu-
A tree 6 meters high with lance-ovate or elliptic leaves 2.5-3 times as long
as wide; because of the glabrous flowers this species is not likely to be confused
with any other except E. florida DC., from which it differs in the characters set
forth in the key to species. Flowers small, the larger calyx-lobes 1.5 mm. long;
stamens about 100.
Peru, Loreto: Florida, Rio Putumayo, mouth of Rio Zubineta,
elev. 180 meters, forest, May-July, 1931, G. King 2146 (F 668811,
type; G; GH; NY; US).
Eugenia cartilaginea McVaugh, sp. nov.
Frutex vel arbor parvifolius, appresse hispidulus vel gemmis strigosis; foliis
1-2.5 cm. longis, coriaceis, obtusis vel acutis; marginibus pallidis, incrassatis,
paulum revolutis, glandulosis; floribus solitariis, axillaribus, vel infimis ex axillis
bractearum caducarum oriundis; hypanthio obconico; pedicellis filiformibus
10-20 mm. longis; calycis lobis intus pubescentibus.
A shrub or small tree 4-6 meters high with lustrous ovate or elliptic leaves
1.5-2 times as long as wide, the margins often irregularly roughened by the
prominent glands; bracteoles deciduous; flowers rather small, the disc 2.5-3 mm.
wide; style 5-6 mm. long; stamens about 75.
This species evidently belongs with the complex which includes
some species with 3-flowered dichasia, e.g., E. discolor (HBK.) DC.
206 FIELDIANA: BOTANY, VOLUME 29
It is similar on the one hand to E. orthostemon Berg, a species which
ranges from central Ecuador to Colombia. In E. orthostemon,
however, pubescence is of more general occurrence on the flowers
and pedicels, and the leaves are larger (up to 3.5 cm. wide and 6-7
cm. long) ; the leaves also lack the cartilaginous margins, prominent
glands and mucronate tips which characterize those of E. carti-
laginea. The latter is similar in many respects, on the other hand,
to E. myrsinoides (HBK.) Diels, from which it may be distinguished
by the characters given in the key.
Peru, Junin: Valley of the Rio Mantaro near Huachicna, elev.
2,300 meters, Weberbauer 6548 (F; GH; NY; US; USM). Huan-
cavelica: Pampas-Salcabamba trail, elev. 2,500 meters, Stork &
Horton 10443 (F 1180795, type).
Eugenia chartacea McVaugh, sp. nov.
Arbor, glabra (disco sparse piloso; calycis lobis petalisque minute ciliatis);
foliis late ellipticis vel ovatis, 11-21 cm. longis obtuse acuminatis; venis 7-8, supra
convexis sed sulcatis; nervo marginal! aperto; petiolo 2 mm. crasso canaliculate;
racemis abbreviatis; alabastris 15-18 mm. longis; calycis lobis rigidis, chartaceis,
imbricatis, 10-12 "mm. longis; antheris 2-2.3 mm. longis.
A tree 15 meters high with very broad leaves about 1.5 times as long as wide,
relatively few veins, and small clusters of large flowers produced on old wood.
Disc quadrangular, 4-5 mm. wide; style probably 15 mm. long or more; stamens
Brazil, Amazonas: Basin of Rio Jurua, on high terra firma, near
mouth of Rio Embira (tributary of Rio Tarauaca), Lat. 7 30' S.,
Long. 70 15' W., June 21, 1933, B. A. Krukoff 4951 (NY, type; US).
Eugenia crucicalyx McVaugh, sp. nov.
Frutex, ramulis gemmisque et racemis pilis appressis, pallide rufulis, obtectis;
foliis 4.5-8 cm. longis brevi-acuminatis; venis utroque latere 7-10, utrinque
elevatis; racemis abbreviatis; hypanthio paulum angulato, rufo-velutino; calycis
lobis 5-6 mm. longis, quam hypanthio 2-3-plo longioribus, intus glabris, ad
florendi tempus patentibus; bracteolis 1.5 mm. longis persistentibus, ovatis,
distinctis; stylo 8 mm. longo.
A shrub with concolorous, lustrous elliptic leaves about twice as long as wide
and a well-defined marginal vein; buds 7-8 mm. long; calyx-lobes in unequal
pairs, the inner ones obovate and truncate, the outer ovate-triangular, bluntly
pointed; disc 2-2.5 mm. wide; stamens 200-250.
Peru, San Martin: Tarapoto, elev. 360-900 meters, forest, Dec. 4,
1929, L. Williams 6216 (F 627015, type). Univ. of Mich. Neg. 466.
Eugenia curvipilosa McVaugh, sp. nov.
Frutex, pallide brunneo-pilosus; foliis 4-7 cm. longis breve lateque acuminatis,
supra sub-vernicosis scabriusculisque, subtus tomentosis; racemis 1-2 cm. longis
McVAUGH: TROPICAL AMERICAN MYRTACEAE 207
tenuibus; floribus 6-12, divaricatis, pedicellis 4-6 mm. longis; hypanthio strigoso,
disco 1.5-2 mm. lato.
A shrub 3 meters high, with subcoriaceous elliptic-ovate leaves 1.5-2 times
as long as wide, the midvein and the about 10 lateral veins somewhat impressed
above; bracteoles persistent, connate, glabrescent, not strongly glandular; flowers
small, the buds 4 mm. long, style about 4(?) mm. long, stamens 50-60.
Superficially suggesting E. limbosa Berg, but readily differenti-
ated from that species by the leaf and pubescence characters used
in the key.
Peru, Cajamarca: Valley of the Rio Llaucan, near Pion, elev.
1,700-1,800 meters, June, 1915, A. Weberbauer 7137 (F 628083,
type; GH). Univ. of Mich. Neg. 447.
Eugenia curvivenia McVaugh, sp. nov.
Frutex, minute fusco-rufo-tomentosus; foliis 16-24 cm. longis acuminatis
mox glabrescentibus; venis utroque latere circiter 10, impressis, arcuatim adscen-
dentibus, apicem versus sensim extenuatis; nervo marginali nullo; racemis ab-
breviatis; bracteolis 0.8-1.5 mm. longis persistentibus, non connatis; floribus
minusculis, alabastris 4 mm. longis, calycis lobis 3-4 mm. longis distinctis, deciduis;
disco 2 mm. lato; stylo 5 mm. longo.
A shrub 6 meters high with large, broadly elliptic-ovate coriaceous veiny
leaves 2-2.5 times as long as wide, stout dark petioles, and (in proportion to the
leaves) very small flowers in small clusters in the axils; stamens 50-60.
Brazil, Amazonas: Basin of Rio Jurua, near mouth of Rio
Embira, tributary of Rio Tarauaca, 7 30' S. Lat., 70 15' W. Long.,
June 27, 1933, B. A. Krukoff 5045 (NY, type; US).
Eugenia dibranchiata McVaugh, sp. nov.
Arbor, innovationibus petiolisque et racemis, pilis rufis appressis, partim
dibranchiatis, obsitis; foliis glabris vel appresse strigosis, 8.5-14 cm. longis anguste
acuminatis; nervo medio supra elevato; venis utroque latere 10-15; nervo margin-
ali interrupto, exiguo; racemis 3-6 cm. longis tenuibus; bracteolis obtusis, con-
natis, strigosis vel ciliatis, nee glandulosis; disco 2.5-3 mm. lato; stylo 7 mm.
A tall forest tree with inconspicuously veined elliptic leaves 2-3 times as long
as wide, and slender axillary racemes with 4-7 pairs of flowers; hypanthium
pubescent without; calyx-lobes about 2 mm. long, pubescent on both sides;
stamens 60-75, up to 10 mm. long.
This species is apparently most closely related to E. florida DC.;
the leaves suggest in color and texture those of E. florida, but the
marginal vein is more uniformly developed in E. dibranchiata, and
the midvein is elevated, whereas in E. florida it is flat or somewhat
impressed. Specimens suggesting intermediates between these two
species have been collected near Palmares, Brazil, not far from the
208 FIELDIANA: BOTANY, VOLUME 29
Peruvian border (Krukoff no. 8322), and also at Zepelacio, near
Moyobamba, Peru (Klug no. 3304). These plants have rather
well-developed marginal veins, and the pubescence and floral charac-
ters are nearly as in E. dibranchiata (except that the hypanthium is
glabrous in Klug's collection), but the midveins are impressed as in
E. florida and the young leaves are pubescent along the midvein
as in that species. It appears also that E. dibranchiata is related to
the species complex which includes E. riparia DC., but differs in the
thin and concolorous leaves, poorly developed marginal vein, ele-
vated midrib, and the shorter and more compact racemes with smaller
flowers, relatively longer styles and a far greater proportion of
Peru, Loreto: Caballo-Cocha, Aug. 13, 1929, L. Williams 2449
(F 615017, type); Williams 2415 (F), 2416 (F). La Victoria,
Williams 2934 (F). Leticia, Williams 3145 (F).
Eugenia discreta McVaugh, sp. nov.
Arbor, puberulenta, pilis appressis minimis, magna pro parte dibranchiatis,
obsita; foliis oblanceolatis obovatisve, 14-18 cm. longis acuminatis; venis utroque
latere 10-15, supra elevatis; nervo medio supra impresso; venulis reticulatis;
racemis abbreviatis; floribus 4-6 minusculis pedicellatis, alabastris 5 mm. longis;
bracteolis ovatis obtusis, 2.5 mm. longis persistentibus connatis; disco 2 mm.
lato; staminibus circiter 75.
A tree to 11 meters high with reticulate- veiny leaves 2.5-3 times as long as
wide. The leaves tend to become liver-colored beneath in drying but to remain
olive-green above. The calyx-lobes are unequal, 1-2.5 mm. long, the style 7-9
mm. long; fruit globose, said to be yellow.
Peru, Loreto: Gamitanacocha, Rio Mazan, on river bank, Jan.
15, 1935, J. M. Schunke 40 (A; F; US 1458946, type). Brazil,
Amazonas: Near mouth of Rio Embira, Krukoff 4884 (US).
The Peruvian species related to the above are poorly represented
in herbaria and are consequently difficult to interpret taxonomically.
The group may be characterized as having small, slender-pedicellate,
glabrous flowers in axillary "glomerules" or "fascicles" (actually
very short racemes), nearly glabrous foliage, the midvein impressed
above, and the marginal vein relatively far from the margin with
a distinct sub-marginal vein beyond it, the fruit globose and 1 cm.
in diameter or less. The species in question include E. schom-
burgkii Benth., E. tapacumensis Berg, E. maculata Berg, and two
newly proposed species, E. discreta and E. quadrijuga. A species of
Amazonian Brazil, E. agathopoda Diels (Verh. Bot. Ver. Brandenb.
48: 192. 1907), is similar but has larger flowers, longer racemes and
McVAUGH: TROPICAL AMERICAN MYRTACEAE 209
Because of the small amount of material available for study, no
really workable key to the above species can be constructed. The
flowers of all the species are so much alike that they provide no good
diagnostic characters, and the fruits are almost unknown. In the
present treatment Eugenia schomburgkii includes chiefly glabrous or
nearly glabrous plants with lanceolate and long-pointed leaves which
are very smooth on the upper surface; E. maculata is distinguished
from broad-leaved extremes of E. schomburgkii, and from the other
species involved, by its relatively well-developed raceme axis which
may be as much as 5 mm. long. Eugenia tapacumensis is a plant
with subcoriaceous, elliptic, bluntly pointed leaves and rather fine,
inconspicuous veins; E. discreta, known only from the type, has
mostly oblanceolate and rather large, reticulate-veiny leaves, and
sparse, appressed dibranchiate hairs in the inflorescence. The
remaining species, E. quadrijuga, is described from four collections
which may represent one species or conceivably as many as four.
The collections agree very well in most features, but differ in leaf
shape from long-elliptic (as in Ule's no. 9661, which suggests E.
schomburgkii) to broadly elliptic (as in Klug's no. 3153, which
suggests E. tapacumensis). The pubescence differs slightly from
one of the four collections to another, but all have in common the
minutely bristly ascending hairs of the pedicels.
Eugenia ferreyrae McVaugh, sp. nov.
Frutex, ramulis dichasiisque, et innovationibus, insigniter strigosis; foliis
1-2 cm. longis coriaceis, obtusis vel rotundatis vel emarginatis; nervo medio
impresso, venis et nervo marginali inconspicuis; floribus solitariis, vel plerumque
pedunculis 3-floris, 7-10 mm. longis, compressis, apicem versus 1 mm. latis;
hypanthio ut videtur campanulato, strigoso; calycis lobis intus strigosis vel
demum glabrescentibus; staminibus ut videtur circiter 50; embryone cotylis
A shrub with stiff smallish, broadly elliptic, ovate or obovate leaves 1.2-1.9
times as long as wide, sometimes drying bluish- or grayish-green above and dark
reddish-brown beneath, the upper surface sparingly impressed-puncticulate;
calyx-lobes broad and coriaceous, 2-3 mm. long; disc 3 mm. wide; style 7-9 mm.
long; fruit probably globose, about 7-10 mm. in diameter, the seeds 1 or 2, about
7 mm. long.
This species is of unusually great interest, as coming from an
arid area of southern cis- Andean Peru, where no other native species
of Myrtaceae is known to occur. Unfortunately it is known but
from two collections, and one fruit only has been available for study.
The plant resembles in general habit and leaf morphology some of
the Chilean Myrtaceae, e.g., Reichea coquimbensis (Barn.) Kausel,
210 FIELDIANA: BOTANY, VOLUME 29
but that species has pentamerous flowers and a very short erect
radicle, whereas in Eugenia ferreyrae the radicle is accumbent,
tapering-cylindrical, about 0.5 mm. thick at base, and 2.5 mm. long.
The embryo of E. ferreyrae, with its separate cotyledons and prom-
inent radicle, is anomalous in the genus Eugenia, sens, str., but
apparently agrees well with those of other species of the genus
Anamomis Griseb. Until a more general survey of fruit and seed
characters in Anamomis can be made, however, I prefer to refer the
present species and its close relatives to Eugenia.
Peru, Arequipa: Prov. Caraveli, Lomas de Chaparra, elev. 500-
560 meters, Oct. 19, 1946, Ramdn Ferreyra 1483 (USM, type);
southeast of the port of Chala, highway to Chaparra, elev. 600-750
meters, Oct. 10, 1955, Ferreyra 11450 (MICH). Univ. of Mich.
Eugenia hexovulata McVaugh, sp. nov.
Arbor, appresse pubescens, ramulis petiolis umbellisque, et folii pagina inferiore,
pilis densis rufis dibranchiatis obtectis; foliis 12-24 cm. longis acuminatis; nervo
medio supra elevato; venis utroque latere circiter 20, utrinque paulum elevatis;
nervo marginali aperto; racemis abbreviatis, umbelliformibus, usque ad 15-floris;
alabastris 6 mm. longis clausis, obtusis vel obscure apiculatis, hypanthio 8-
angulato; calyce ad florendi tempus usque ad verticem germinis in lobos 4 sub-
aequales longitudinaliter direpto; lobis ovatis, acutis, 5 mm. longis, intus glabris;
bracteolis persistentibus, appressis.
A tree 15 meters high, with lance-oblong leaves about 3 times as long as wide;
disc 2 mm. wide, the staminal ring extending nearly to the depressed center;
style 6-7 mm. long; stamens 100-125, the anthers linear, 1-1.2 mm. long; ovary
bilocular, the ovules 3 in each locule, collateral.
A plant of uncertain relationships; the small number of ovules
is unusual but not unknown in the Eugeniinae. An affinity to
E. feijoi Berg, E. cuspidifolia DC. and their relatives (the supposed
genus Catinga Aubl.) is suggested by the characters of inflorescence,
bracteoles and pubescence, the angled hypanthium, the long narrow
anthers and the broad staminal ring. The venation in E. hex-
ovulata, however, is anomalous in Catinga, and the closed buds of the
present species appear to be nearly unique.
Peru, Loreto : Pumayacu, between Balsapuerto and Moyobamba,
elev. 600-1,200 meters, forest, Aug.-Sept., 1933, G. King 3194
(F 715574, type; NY). Univ. of Mich. Neg. 462.
Eugenia illepida McVaugh, sp. nov.
Arbor, glabra (disco piloso; bracteolis, petalis et calycis lobis ciliatis); foliis
4-7.5 cm. latis, 12-26 cm. longis, leviter acuminatis; petiolo 1-2 mm. crasso; venis
McVAUGH: TROPICAL AMERICAN MYRTACEAE 211
utroque latere 10-15, utrinque paulum elevatis; nervo marginali aperto; racemis
abbreviates; alabastris 12 mm. longis, supra hypanthium globosis; calycis lobis
rigidis, glandulosis, late imbricatis, inaequalibus; lobis interioribus 8 mm. longis,
14 mm. latis; bracteolis persistentibus nee connatis; disco 6-7 mm. lato; antheris
1-1.3 mm. longis.
A tree to 20 meters high, with relatively thin oblanceolate or elliptic leaves
3-3.5 times as long as wide. Flowers in axillary clusters, on pedicels up to 15 mm.
long; flowers large, the style 16-17 mm. long, the stamens probably about 300.
Brazil, Acre: Near mouth of Rio Macauhan (tributary of Rio
Yaco), Lat. 9 20' S., Long. 69 W., on terra firma, Aug. 13, 1933,
B. A. Krukoff 5482 (NY; US), Aug. 27, 1933, Krukoff 5675 (NY;
US 1662164, type). Krukoff 's no. 5482 is represented by unbroken
specimens, with the inflorescence in place; in these specimens, the
anthers are invariably abnormal because of the attacks of gall-
forming insects, and no. 5675, with apparently normal flowers, is
designated as type.
Eugenia indifferens McVaugh, sp. nov.
Arbor vel frutex, ramulis foliisque novellis pubescentibus vel strigosis, pe-
dunculo hypanthioque glabris; foliis rigide coriaceis 1.3-3 cm. longis obtusis,
maturitate eglandulosis; venis inconspicuis; pedunculis 1-3-floris compressis,
5-15 mm. longis, apicem versus usque ad 1.5 mm. latis; hypanthio late turbinato;
calycis lobis intus sericeis; staminibus circiter 250.
A tree or shrub with ovate, obscurely veined leaves as broad as long, or up
to twice as long as broad; most of the flowers solitary; buds 3-5 mm. long; disc 3.5
mm. wide, glabrous, the style 6.5-7.5 mm. long.
Peru, Cuzco: Urubamba Valley, Hacienda Fanccac, elev. 2,760
meters, Sept. 10, 1928, F. L, Herrera 2099 (F; US 1422430, type).
Univ. of Mich. Neg. 442.
Eugenia longicuspis McVaugh, sp. nov.
Arbor, ramulis racemisque pilis appresse-adscendentibus obtectis; foliis 18-21
cm. longis, cuspidato-acuminatis; nervo medio supra impresso; venis utroque
latere 8-12, supra impressis, subtus strigosis; nervo marginali aperto, supra
impresso; racemis 6-9 cm. longis validis; disco 6 mm. lato.
A tree 9 meters high with obovate-oblong leaves about 3 times as long as
wide, readily recognized by the caudate tips 2-3 cm. long, and the prominent
pattern formed on the lower surface by the lateral and marginal veins. Flowers
3-5 pairs, large for this group of species; bracteoles connate, persistent, together
5 mm. wide; style not seen; stamens more than 200.
The only collection surely referable to this species is the type,
which bears flowers from which the corolla, androecium and style
have fallen. A specimen which seems intermediate between this
and E. atroracemosa McVaugh is Williams' no. 4747, from Yurima-
212 FIELDIANA: BOTANY, VOLUME 29
guas, Peru, which bears immature buds and one raceme with half-
grown fruits. This has the stout, short-pedicelled racemes, short-
acuminate leaves and non-connate bracteoles of E. atroracemosa, but
the disc (6-7 mm. wide and glabrous at the center), stamens (about
200) and style (9 mm. long) of E. longicuspis. The ovules are about
7 in each locule, and collateral. This may represent an undescribed
species, but the material is imperfect and is referred with doubt
to E. longicuspis. A second collection from near Iquitos, Williams'
no. 8018, is perhaps also referable to this species, but the leaves tend
to be elliptic and are not markedly caudate-acuminate, the upper
surface is impressed-puncticulate, the pedicels are 5 mm. long and
the disc is 4.5 mm. wide; the fruit, nearly at maturity, is subglobose,
about 1.5 cm. in diameter.
Peru, Loreto: Mishuyacu, near Iquitos, elev. 100 meters, forest,
Feb.-Mar., 1930, G. King 855 (F; NY; US 1455842, type).
Eugenia macrocalyx (Rusby) McVaugh, comb. nov. Caly-
corectes macrocalyx Rusby, Mem. N. Y. Bot. Gard. 7: 313. 1927.
A distinctive species, but quite out of place in Calycorectes,
which was described as having the calyx closed in bud, then splitting
longitudinally. The present species belongs rather to the genus
Phyllocalyx Berg, which I believe is better regarded as a part of the
inclusive genus Eugenia.
Eugenia mandonii McVaugh, sp. nov.
Arbor vel frutex, ramulis floribusque et praesertim hypanthio strigosis; foliis
2-4 cm. longis obtusis acutisve vel subacuminatis; venis utroque latere 5-8
inconspicuis; pedunculis solitariis, 1-floris, oppositis, ex infimis nodis 2 ramulorum
hornotinorum oriundis, vel rhachi raro abortiva, abbreviata, efoliata, 1-4-flora;
bracteolis subulatis 2 mm. longis strigosis, deciduis; calycis lobis rotundatis
2.5 mm. longis, intus strigosis.
A tree or shrub with small coriaceous elliptic-ovate leaves 2-3 times as long
as wide, strigose and vernicose above, with impressed midvein; pedicels 8-16 mm.
long, subtended by small foliaceous bracts or much smaller scarious bracts 1.5-2
mm. long; buds 6 mm. long, obovate; disc 4-4.5 mm. wide; style 6.5-9 mm. long;
stamens about 200.
Bolivia: Prov. Larecaja, viciniis Ananea, in nemoribus, June,
1856, G. Mandon 634 (G, type; NY). Univ. of Mich. Neg. 438.
Eugenia micranthoides McVaugh, sp. nov.
Arbor vel frutex, minute pubescens; ramulis racemisque dense pilis erectis
0.1 mm. longis obsitis; foliis ellipticis 4-7 cm. longis subcaudatis supra eglandu-
losis; nervo medio supra sulcato; venis utroque latere circiter 10, utrinque obscuris;
McVAUGH: TROPICAL AMERICAN MYRTACEAE 213
racemis brevis, 2-8 mm. longis tenuibus, 4-10-floris; pedicellis 2-4 mm. longis;
bracteolis persistentibus; calycis lobis 1 mm. longis.
A shrub or small tree with tiny racemes, small flowers and small narrow leaves
2-3 times as long as wide, their narrowly acuminate tips 1-1.5 cm. long; disc
about 2 mm. wide; style 4.5-5 mm. long; stamens about 50.
This plant is strikingly similar in characters of pubescence and
in morphology of the inflorescence to E. micrantha (HBK.) DC., but
in that species the leaves are narrower, prominently veined and
glandular, the midvein is not impressed above, the flowers are
smaller and the bracts are persistent. The type locality of E.
micrantha is near Honda, in the valley of the Rio Magdalena,
Colombia. A note at the end of the original description, however
(HBK. Nov. Gen. & Sp. 6: 145 [folio ed. 115]. 1823), reads: "Bon-
plandius haec specimina in Peru via lecta esse memorat." Ap-
parently this reference to Peru is erroneous, for the species is well
known in Colombia but has not been found subsequently in Peru.
Peru, Lore to: Yurimaguas, Parana Pura, in forest, Oct.-Nov.,
1929, L. Williams 3825 (F), 4102 (F), 4620 (F 623009, type).
Eugenia minimifolia McVaugh, sp. nov.
Frutex multiramosus compactus, appresse strigosus; foliis 2.5-6 mm. longis
vernicosis rigide coriaceis, obtusis, subaveniis, utrinque impresso-punctatis; pe-
dunculis 1-floris solitariis, 2-3 mm. longis.
A much-branched sclerophyllous shrub 2 meters high, with the numerous
white flowers abundant toward the tips of the branches; leaves obovate or elliptic,
1.5-2 times as long as wide; buds 3.5 mm. long; calyx-lobes broad, 1 mm. long,
glabrous both sides; style 5.5-6 mm. long; stamens about 125.
Peru, Huancavelica : Prov. Tayacaja, hills to the left of the Rio
Mantaro, above the bridge of Chiquiac, low open mountain sides
covered with evergreen shrubs, elev. 2,600-2,700 meters, Mar. 15,
1913, A. Weberbauer 6500 (F; GH; US 1497222, type; USM).
Eugenia multirimosa McVaugh, sp. nov.
Frutex vel arbor, glomerulis (racemis abbreviatis) puberulis; ramis rufidis,
exophloeo longitudinaliter rumpente et secedente; foliis subsessilibus, basi cordato-
auriculatis, plerumque oblanceolatis, ut videtur 25-35 cm. longis, 3-5-plo longior-
ibus quam latioribus; nervo medio supra impresso; venis utroque latere circiter
15; petiolo rimoso 3-5 mm. longo, ut videtur 3-5 mm. crasso (exophloeo suberoso
soluto); racemis abbreviatis; bracteolis persistentibus 1.5 mm. longis; calycis
lobis suborbiculatis, majoribus 4-5 mm. longis; disco 3.5-4.5 mm. lato.
A shrub or small tree, well marked by the nearly sessile, cordate-auriculate
and oblanceolate leaves, and by the tendency for the outer cortical layers of the
petioles and pedicels to separate irregularly; flowers about 6, on stout pedicels
8-11 mm. long, not large for the size of the leaves, the hypanthium in anthesis
214 FIELDIANA: BOTANY, VOLUME 29
3 mm. long; stamens probably about 300; fruit said to be orange, ellipsoid, about
3 cm. long.
Peru, Loreto: Yurimaguas, along lower Rio Huallaga, elev. 135
meters, dense forest, Aug. 23-Sept. 7, 1929, Kittip & Smith 29019
(NY; US 1462448, type); Yurimaguas, Kittip & Smith 28020 (F;
NY); Timbuchi, L. Williams 1017 (F).
Eugenia osteomeloides (Rusby) McVaugh, comb. nov. Myr-
tus osteomeloides Rusby, Mem. Torr. Bot. Club 6: 36. 1896. M.
myrciopsis 0. Ktze. Rev. Gen. 3, pt. 2: 92. 1898. E. myrciopsis
(O. Ktze.) K. Schum. in Just, Bot. Jahresb. 26, pt. 1: 359. 1900.
This is a Eugenia of the affinity of Anamomis Griseb., not
a member of the Pimentinae. The cotyledons are large, distinct and
plano-convex; the radicle is cylindrical and about a third as long as
the seed. The type specimens of Myrtus myrciopsis, from Santa
Cruz, Bolivia, are somewhat more densely pubescent than other
plants from the same area. Possibly the number of hairs is related
to the fact that Kuntze's plants are in young bud; there do not
appear to be other significant differences between M. myrciopsis and
Eugenia pearcei McVaugh, sp. nov.
Frutex sempervirens, rufo-tomentosus; foliis coriaceis, 6.5-11 cm. longis
obtusis, subtus tomentosis; venis utroque latere circiter 10, arcuatim adscenden-
tibus, apicem versus sensim extenuatis, venulis nee incrassatis; dichasio plerumque
7-15-floro, floribus in dichotomis sessilibus; pedunculo 2.5-5 cm. longo, compresso,
apicem versus usque ad 2 mm. lato; alabastris 4-5 mm. longis; calycis lobis utrin-
que tomentosis; stylo 5 mm. longo vel longiore; disco 3-4 mm. lato.
An evergreen shrub 2-3 meters high, with lustrous thick elliptic-ovate leaves
about twice as long as wide, impressed-punctate above; flowers middle-sized, the
calyx-lobes 3 mm. wide, 1.5-2.5 mm. long; stamens 200-250; petals lightly tomen-
A very distinctive species, but unfortunately the only known
specimen has been severely damaged by insects.
Bolivia: Hills near Pata, 6,000 feet, Dec., 1864, R. Pearce (BM).
Univ. of Mich. Neg. 483.
Eugenia percincta McVaugh, sp. nov.
Frutex vel arbor, dense fulvo-velutinus; foliis elliptico-oblongis 12-14 cm.
longis, 6-8.5 cm. latis deltoideo-acuminatis, crassimarginatis; racemis perbrevibus,
rhachi subnulla et floribus 4 approximatis; calycis lobis 4-5 mm. longis latisve
concavis, imbricatis, utrinque velutinis; bracteolis deciduis; stylo 14-16 mm.
McVAUGH: TROPICAL AMERICAN MYRTACEAE 215
A tree or shrub, well marked by the heavy covering of tawny hairs, the
strong cartilaginous margins of the leaves, and the conspicuous glomerules of
rather large flowers; disc 4-4.5 mm. wide; stamens about 150.
Brazil, Guapore": Falls of Madeira, Oct., 1886, H. H. Rusby 2084
(NY; US 1416665, type). Univ. of Mich. Neg. 453.
Eugenia percrenata McVaugh, sp. nov.
Frutex, subglaber; foliis 6-9 cm. longis caudatis, marginibus conspicue
crenatis; nervo medio supra planiusculo vel elevato; racemis 4-10-floris ab-
breviatis; bracteolis persistentibus; calycis lobis suborbiculatis vel oblongis, 1.5-2
mm. longis; disco 1.5 mm. lato.
A shrub 5 meters high, glabrous except for a few strigose hairs on the bracts
and on vegetative buds; leaves elliptic, 2.5-3.5 cm. wide, and 3-3.5 times as long
as wide, the apex caudate-acuminate, the base cuneate and attenuate to a petiole
1-1.5 mm. thick and 3-4 mm. long; margins deeply crenate with 6-10 notches on
each side, the notches 1 mm. deep with a gland 0.5-0.8 mm. wide at the base of
each; midvein flat, or centrally and sharply keeled above, pale and elevated
beneath; lateral veins about 12 pairs, obscure above, somewhat raised beneath;
marginal veins about equaling the laterals and arched between them, 1-2 mm. from
margin; blades when dry dull and dark in color and obscurely gland-dotted above,
brown and conspicuously glandular beneath; inflorescence a short axillary raceme,
the axis 2-5 mm. long, with 2-5 approximate decussate pairs of flowers; bracts
1 mm. long, ovate, ciliate, rounded on the backs; pedicels 5-10 mm. long, some-
what compressed distally and there 1 mm. wide; bracteoles 0.7 mm. long, per-
sistent and nearly erect, ovate but trough-shaped and so appearing lance-ovate,
acute; hypanthium 2 mm. long, constricted at base above the bracteoles; calyx-
lobes suborbicular or oblong, rounded at apex, strongly reflexed after anthesis,
1.5-2 mm. long and wide; disc about 1.5 mm. wide, somewhat quadrangular;
style 3.5 mm. long; stamens about 50, up to 3 mm. long; anthers 0.8 mm. long;
petals ovate, 3 mm. wide, 3.5 mm. long, yellowish white (Krukoff ) ; ovary bilocular,
the ovules about 15 in each locule, radiating from a centrally affixed placenta.
Brazil, Mato Grosso: Near Tabajara, upper Machado River
region, in terra firma, Nov. 18, 1931, B. A. Krukoff 1368 (MICH,
type; NY; US).
This species resembles in habit and in morphology of leaf and
inflorescence both Eugenia egensis and E. flavescens, but it is readily
distinguished from these and from other known species by the
caudate-acuminate and markedly crenate leaves, and by the reflexed
Eugenia pustulescens McVaugh, sp. nov.
Arbor vel frutex, plusminusve dense strigosus, pilis ferrugineo-rufidis ap-
pressis partim dibranchiatis, etiam pilis erectis brevioribus immixtis, obsitus;
foliis 7-9 cm. longis brevi-acuminatis; racemis usque ad 6 cm. longis validis;
bracteis bracteolisque et calycis lobis vix pubescentibus, sed glandulis fuscis
convexis crebro obsitis; stylo 5-6 mm. longo; staminibus circiter 100.
216 FIELDIANA: BOTANY, VOLUME 29
A shrub or tree with elliptic leaves 2-2.7 times as long as wide, inconspicuously
veined. The very prominent glands in the inflorescence serve to distinguish this
species from all others except E. polyadena Berg, which as noted in the key to
species has differences in pubescence and considerably larger flowers, as well as
a different geographical range. Buds 5 mm. long; larger calyx-lobes 2-2.5 mm.
long; flowers up to about 18.
Ecuador: El Recreo, Prov. Manabi, y S. Lat., H. F. A. Eggers
15787 (GH; US 1361943, type), Eggers 14957, Apr. 29, 1897 (F).
Locality uncertain: Fl. H[?uayaquil.j no. 408, [anno] 1803 (Herb.
The last specimen cited above is of uncertain origin but may
have formed a part of the Flora Huayaquilensis, of which some
specimens were distributed, probably by Pavon. The collector may
have been Tafalla, but even this is uncertain. Another specimen
of this same species, from the Moricand herbarium and now at
Geneva (G), is labelled "Eugenia sp nova, Peru," and in another
hand, "Pavon." This is probably one of a considerable series which
Moricand received from Pavon in 1827, and which included plants
from various parts of America, collected in part by Pavon himself,
and in part by others. From the state of preservation of this
particular specimen, from its morphology and from its degree of
maturity, it appears that the plant originally formed a part of the
collection cited above, which is attributed tentatively to Tafalla.
The Flora Huayaquilensis is known to include species from various
parts of what is now Ecuador, including the higher Andes (e.g.,
Myrteola microphylla var. microphylla} , so that it is impossible to
make a definite statement that the above specimens of Eugenia
pustulescens came from any particular part of Ecuador. In view of
the known modern localities where the species grows, however, it
seems most probable that the early collection or collections came
from the coastal lowlands of Ecuador; this, if true, suggests that the
species may justifiably be excluded from the known flora of Peru.
Eugenia quadrijuga McVaugh, sp. nov.
Frutex vel arbor, plusminusve appresse pubescens; foliis ellipticis, 7-11 cm.
longis obtuse acuminatis; venis utroque latere circiter 10, utrinque elevatis;
nervo medio supra impresso; venulis tenuiter reticulatis; racemis abbreviatis;
floribus 2-8 pedicellatis, pedicellis minute hispidulis, pilis brevissimis adscenden-
tibus obsitis; alabastris 4.5-5 mm. longis; bracteolis longiusculis, 1.5 mm. longis
persistentibus nee connatis; disco 1.5-2.5 mm. lato; staminibus circiter 60-75.
A shrub or tree up to 15 meters high; for discussion of the variability and
relationships of this species, see above under E. discreta; calyx-lobes up to 2.5-3
mm. long; style 6-8 mm. long; fruit globose or pyriform, probably about 1 cm.
McVAUGH: TROPICAL AMERICAN MYRTACEAE 217
Peru, Loreto: Pumayacu, between Balsapuerto and Moyobamba,
G. King 3153 (A; F; G; GH; US). Brazil, Acre: near mouth of Rio
Macauhan (tributary of Rio Yaco), Lat. 9 20' S., Long. 69 W., on
terra firma, Aug. 21, 1933, B. A. Krukoff 5594 (NY; US 1664166,
type; Y); same locality, Krukoff 5415 (NY; US). Seringal S.
Francisco, E. Ule 9661 (G; US).
Eugenia quebradensis McVaugh, sp. nov.
Arbor vel frutex, tomentosus, vel gemmis ramulisque novellis sericeo-veluti-
nis, pilis sordidis usque ad 1 mm. longis obtectis; foliis lanceolatis, 5.5-7.5 cm.
longis, 5-6-plo longioribus quam latioribus; racemis abbreviates, floribus majus-
culis sessilibus glomeratisque; bracteolis ignotis, ut videtur deciduis; calycis lobis
rotundatis, 3-3.5 mm. longis; disco 6 mm. lato.
A tree or shrub, with the aspect of some species of Psidium, well marked by
the tomentum, the narrow rigidly coriaceous leaves and the large sessile flowers;
hypanthium 4-5 mm. long and wide, with 4 strong winglike angles; style probably
10-15 mm. long; stamens about 300.
Known only from the type, which is from a botanically little-
Peru, Lambayeque: Prov. Chiclayo, quebrada del Rio Sana,
monte seco, Dec., 1928, N. Esposto s.n. (USM, type). U. S. Nat.
Mus. Neg. 4466.
Eugenia quinqueloba McVaugh, sp. nov.
Arbor pilosa, ramulis dichasiisque pilis appressis vel adscendentibus usque
ad 0.8 mm. longis obtectis; pilis pellucidis, luminibus rufidis; foliis subsessilibus,
orbiculatis vel late ovatis, supra flavido-viridibus vernicosisque; dichasiis 3-7-
floris; alabastris 5-6 mm. longis; floribus 5-meris; calycis lobis inaequalibus,
rotundatis, majoribus 3, 3 mm. longis; disco 3-4 mm. lato.
A tree 4-6 meters high, unique in its sessile and often suborbicular rigid
leaves up to 7.5 cm. long and wide, its 5-merous flowers in small dichasia, and its
isolated position in the Department of Lima. Dichasium up to about 3 cm. long;
style 6 mm. long; stamens 75-100; petals white or yellowish (Ferreyra); fruit
probably ellipsoid, 1 cm. long or more.
Peru, Lima: Prov. Huarochiri, arriba de San Bartolome", monte
bajo, elev. 2,900-3,000 meters, Nov. 5, 1954, R. Ferreyra 10417
(MICH, type); same locality, Ferreyra 10424 (MICH).
A very distinct species of uncertain relationships. Were it not for
the 5-merous flowers, the characters of the embryo, placentation and
inflorescence would align E. quinqueloba perfectly with the group of
species that I take to represent the genus Anamomis Griseb. Surely
E. quinqueloba has little affinity to the genus Myrcianthes, a small
eugenioid group of eastern warm-temperate South America, in
218 FIELDIANA: BOTANY, VOLUME 29
which its 5-merous flowers would place it according to Berg's system.
Possibly it is somewhat more closely related to the monotypic
Chilean genus Reichea Kausel, which has, however, mostly solitary
flowers, a very short erect radicle, and apparently somewhat different
placentation. (In E. quinqueloba the ovules are about 20 in each of
the two locules, radially and externally directed from a short,
centrally affixed placenta; the testa of the [somewhat immature]
seed is free and membranaceous, the cotyledons distinct, fleshy,
plano-convex, the radicle accumbent and at least half as long as the
cotyledons; for the corresponding details in Reichea, see Kausel in
Lilloa 13: 129-130. 1947.) I have assigned the present species to
Eugenia in spite of the 5-merous flowers, pending revision and ex-
amination of the generic characters of the whole group of Andean
species which apparently comprise the major portion of the so-called
genus Anamomis, as well as the characters of the Chilean genera
Reichea and Myrceugenella. See also some remarks above, under
Eugenia scalariformis McVaugh, sp. nov.
Arbor vel frutex; foliis adultis subglabris, 33-35 cm. longis, 4-plo longioribus
quam latioribus; venis utroque latere 25-30; nervo marginali aperto, valido, vix
arcuato; petiolo 4 mm. crasso, 12-15 mm. longo; ramis annotinis ut videtur flo-
riferis, floribus pedicellatis, ?glomeratis; alabastris ut videtur 2 cm. longis; bracteo-
lis 10 mm. longis, appressis, deciduis; calycis lobis rotundatis, imbricatis con-
cavisque, interioribus 15 mm. longis; staminibus ut videtur circiter 500, antheris
1.5-1.8 mm. longis linearibus.
A shrub or tree with very long elliptic or oblanceolate acuminate leaves with
numerous scalariform lateral veins; the inflorescence is densely felted with short
flaccid pale brown mostly dibranchiate hairs; hypanthium 8-angled, 10 mm. long;
disc 10 mm. wide, glabrous; style 2.5 cm. long; petals 25-32 mm. long.
Superficially resembles Eugenia tumulescens, from which it may
be distinguished by the smooth petiole, the more numerous veins,
the pedicels, which are longer than the hypanthium, and the much
Peru, Loreto: Stromgebiet des Maranon, Santiago Miindung am
Pongo de Manseriche, G. Tessmann 4328 (G, type). F.M. Neg.
Eugenia schunkei McVaugh, sp. nov.
Arbor, subglabra (gemmis rufo-strigosis; calycis lobis bracteolisque minute
ciliatis; disco sparse piloso); foliis (ramulorum terminalium validorum) 14-23
cm. longis acuminatis; venis utroque latere circiter 10, apicem versus sensim
extenuatis, superioribus venam marginalem formantibus; racemis abbreviatis;
alabastris 12-15 mm. longis pyriformibus; bracteolis 1-2.5 mm. longis persistenti-
McVAUGH: TROPICAL AMERICAN MYRTACEAE 219
bus nee connatis; calycis lobis 8-10 mm. longis oblongis, apicem versus incras-
satis cucullatisque, intus glabris, ad florendi tempus reflexis; staminibus 300 vel
ultra, antheris 1.5 mm. longis linearibus in alabastro erectis.
A tree 5 meters high with oblong-lanceolate leaves often 3.5 times as long as
wide and the marginal vein evident in the distal one-third of the blade; racemes
with 5-6 approximate decussate pairs of flowers, often several together on short
spurlike excrescences; disc 4 mm. wide; style 12 mm. long.
A species which is evidently allied to E. feijoi Berg and others
which are sometimes referred to the genus Catinga, but with very
much larger flowers than any other known species of this group.
Peru, Loreto: Rio Mazan, Quebrada Luno, on river bank, elev.
110 meters, Feb., 1935, Jose M. Schunke 184 (A; US 1459093, type).
Univ. of Mich. Neg. 450.
Eugenia stipitata McVaugh, sp. nov.
Arbor, hispidula; foliorum venis 6-10, arcuatim adscendentibus, inter se
arcuatis sed venam marginalem vix formantibus; racemi ramis oppositis, 1-floris,
vel ramis dichotomis 3 (vel raro 7) -floris, flore in dichotomiis stipitato; pedicellis
1-floris 10-20 mm. longis, longitudinaliter acutangulatis striatisque; bracteolis
1-2 mm. longis linearibus deciduis; hypanthio turbinato; disco quadrangulato,
piloso; calycis lobis rotundatis 4, imbricatis, intus appresse pubescentibus, ad
florendi tempus reflexis; germine 4-loculari, ovulis in quoque loculo circiter 10,
ut videtur biseriatim angulo loculorum interno affixis.
A species of uncertain systematic position, apparently without
any close relatives. The branching of the inflorescence appears to
be unique among the American species of Myrtaceae. The structure
of the ovary suggests that of the Subtribe Pimentinae, but the seeds
(known in subsp. sororia only) are definitely eugenioid in structure
although relatively more numerous than is usual in the Eugeniinae.
The species occurs in two well-marked populations, either one of
which would probably be described as an independent species if it
were found geographically isolated. These populations, described
below as subspecies, are readily separated by the characters given
in the key, but they have so many qualitative characters in common
that they are surely to be considered as conspecific.
1. Folia ovata vel late elliptica, 5-6 cm. lata, 1.8-2.3-plo longiora quam latiora,
venis supra impressis, pagina inferiore pilis erectis acutisque et usque ad 0.5
mm. longis crebro obsita; pedicelli plerumque in medio fere, vel infra medium
bibracteolati; stylus 7-8.5 mm. longus, glaber; calycis lobi 4-5 mm. longi.
1. Folia elliptica, 2.5-4.5 cm. lata, 2.2-3.3-plo longiora quam latiora, venis
supra vix manifestis, nee impressis; pagina inferiore maturitate glabra, vel
venis solum hispidulis, vel pilis minutissimis 0.1 mm. longis crebro obsita;
pedicelli 3-5 mm. ultra hypanthium bibracteolati; stylus 5-6.5 mm. longus,
basi pilosus; calycis lobi 2.5-3 mm. longi subsp. sororia.
220 FIELDIANA: BOTANY, VOLUME 29
Eugenia stipitata McVaugh, subsp. stipitata. E. stipitata
McVaugh, as to type.
More markedly hispidulous than the subsp. sororia, with larger and broader
leaves, more conspicuous veins and larger flowers. Stamens 100-150. The fruit
Peru, Loreto: San Antonio, on Rio Itaya, Killip & Smith 29469
(NY; US), Williams 3397 (F). La Victoria, Williams 2787 (F).
Mishuyacu, near Iquitos, forest, elev. 100 meters, Jan., 1930, G.
King 788 (F 624179, type; NY; US). Brazil, Amazonas: Mun.
Humayta, near Livramento, B. A. Krukoff 6591 (NY; US). Univ.
of Mich. Neg. 464.
Eugenia stipitata McVaugh, subsp. sororia McVaugh, subsp.
Differs from subsp. stipitata as noted under that taxon and in the key. Sta-
mens about 75; fruit oblate, velutinous, about 1.5 cm. across; seeds 6-15, reniform,
3-7 mm. long, the embryo completely undivided or the cotyledons slightly sepa-
rated at the chalazal end; testa membranaceous.
Peru, San Martin: Juanjui, Alto Rio Huallaga, elev. 400 meters,
forest, Oct., 1934, G. King 3834 (F; GH, type; US). Tarapoto,
Williams 5486 (F), 5667 (F). Bolivia, ?Beni: Junction of rivers
Beni and Madre de Dios, H. H. Rusby 597 (F; MICH; NY; US).
Brazil, Amazonas: Near mouth of Rio Embira, B. A. Krukoff 4859
(US). Colombia, Meta: Villavicencio, Bro. Apollinaire Myrt. no. 2
(US). Univ. of Mich. Neg. 474.
Eugenia tenuimarginata McVaugh, sp. nov.
Arbor vel frutex, foliis ramulisque adultis glabris, racemis minute appresse-
pubescentibus; foliis 10-14 cm. longis obtuse acuminatis, supra scabrido-papil-
losis; venis utroque latere 6-8, supra paullum elevatis; racemis abbreviatis;
bracteolis connatis persistentibus; alabastris 8-9 mm. longis; calycis lobis rotun-
datis, late imbricatis, intus glabris, marginibus fragilibus, hyalinis, tenuioribus;
lobis interioribus majoribus 5-8 mm. longis latisque; disco 6 mm. lato; stylo 9-10
A tree or shrub, with elliptic, elliptic-ovate or -obovate leaves about twice
as long as wide, the veins rather prominent beneath, and the surfaces somewhat
papillose. The flowers are large, probably always in clusters on old wood; stamens
about 250; anthers 1-1.2 mm. long.
Peru, Loreto: Mouth of Rio Santiago, on high land, G. Tess-
mann 4213, anno "1924" (fragment, F; G, type; NY). Univ. of
Mich. Neg. 437.
McVAUGH: TROPICAL AMERICAN MYRTACEAE 221
Eugenia tumulescens McVaugh, sp. nov.
Frutex tumulescens, ramis crassiusculis, cortice atro-rufido squamuloso
secedente; racemis tomentulosis; foliis petiolatis, ellipticis oblongisve, 20-38
cm. longis, 2.5-5-plo longioribus quam latioribus; nervo medio utrinque elevato;
venis utroque latere 20-30; petiolo rimoso 10-18 mm. longo, ut videtur 3-4 mm.
crasso (exophloeo suberoso soluto); ramis annotinis floriferis, racemis abbreviatis;
bracteolis 4-5 mm. longis oblongis, ad florendi tempus erectis; disco 3.5-4 mm.
A shrub said to form mounds 70 cm. high, with elongate stiff veiny leaves
which are acuminate at tip and narrowed or rounded at base to the corky-thickened
petioles. Flowers 6-8, small for the size of the leaves; hypanthium 3 mm. long,
obtusely 8-ridged and expanded into the gamosepalous calyx-base 1 mm. long;
style 10-12 mm. long; stamens 250-300, the anthers 2.4-2.6 mm. long; fruit
long-ovoid, salmon-yellow (according to Froes), probably 3-5 cm. long.
Brazil, Amazonas: Rio Cauabury, between Rio la and Rio
Maturaca, E. G. Holt & E. R. Blake 438, Nov. 3-7, 1930 (US);
Porto Curucuhy, Rio Negro, terreno arenoso alto, beira do rio,
R. Froes 21106, Oct. 6, 1945 (MICH, type; NY).
Eugenia valvata McVaugh, sp. nov.
Frutex vel arbor usque ad 5 m. alta, glabra; foliis plerumque 5-6 cm. longis
ellipticis obtusis, venis inconspicuis; racemis 1-4 abbreviatis, floribus pedicellatis,
in glomerulis umbelliformibus conglobatis; bracteolis persistentibus 1-1.5 mm.
longis ciliatis; calycis lobis ciliatis subdeltoideis, ut videtur valvatis, inter se ad
florendi tempus longitudinaliter rumpentibus.
An intricately branched shrub or small tree 3-5 meters high, glabrous except
the bristly receptacular disc and the ciliate perianth-lobes and bracteoles; leaves
elliptic, coriaceous, (1.5-) 2-3.5 cm. wide, (3-) 5-6 cm. long, about twice as long
as wide, obtuse or obscurely acuminate at tip, rounded or gradually narrowed
at base to the petiole 3-5 mm. long; margins somewhat pale-cartilaginous and
revolute; midvein sulcate above, prominent beneath; lateral veins 6-10 pairs,
very slender, scarcely apparent in mature leaves; marginal vein 2-3 mm. from
margin, about as strong as the laterals and somewhat arched between them; upper
surface of blade smooth, lustrous, the lower surface paler; glandular dots numerous,
small, apparent in young leaves but hardly at all on mature foliage; inflorescence
an abbreviated axillary raceme, or usually a cluster of 2-4 racemes from each axil,
the axis of the raceme 3-6 mm. long, bearing 5-7 approximate, decussate pairs
of flowers on pedicels (5-) 11-15 mm. long and up to 0.8 mm. wide at the some-
what compressed apex; bracts thin, deltoid or ovate, reddish brown, 0.7-1.5 mm.
long; bracteoles lanceolate or ovate, persistent, appressed to the base of the
hypanthium, 1-1.5 mm. long; hypanthium subcylindric, 1-1.5 mm. in diameter,
1.5-2.5 mm. long, somewhat enlarged distally, then abruptly widened into the
base of the spreading-ascending rounded-deltoid calyx-lobes; lobes membrana-
ceous, markedly convex without in the bud, ciliate at tips or in the distal half only,
united by the proximal margins and separating at the time of anthesis by longi-
tudinal splits up to 1-1.5 mm. long; globe of petals in the opening bud 1.5-2
times as long as the calyx; disc 3 mm. wide; style 5-6.5 mm. long; stamens about
222 FIELDI ANA: BOTANY, VOLUME 29
75, about as long as the style, the anthers 0.6-0.8 mm. long; petals white or pinkish,
about 5 mm. long; ovary bilocular, the ovules 12-20 in each locule, attached
radially to the central septum.
A species which is distinctive because of the umbelliform clusters
of flowers and the splitting of the calyx; the calyx-lobes show
scarcely a trace of any imbricate condition even in the youngest
bud, and are well separated by irregular short breaks below the
sinuses by the time the flower opens. Fruit of this species is not
definitely known, but Hitchcock's collection includes a single de-
tached fruit, with persistent bracteoles and calyx-lobes, which is
apparently dark in color, globose and about 1 cm. in diameter.
Ecuador, Chimborazo: Canon of the Rio Chanchan, about 5 km.
north of Huigra, elev. 5,000-6,500 feet, moist forested valleys in the
afternoon fog-belt, May 19-28, 1945, W. H. Camp E-3280 (MICH) ;
canon of the Rio Chanchan, open deforested slope with small patches
of scrub in the draws, directly above Huigra, elev. 7,000 feet, May
29-31, 1945, W. H. Camp E-3512 (MICH, type); Huigra, elev.
1,200 meters, A. S. Hitchcock 20733 (US). Canar: Santa Rosa de
Cafiar, J. N. Rose 22655 (US); between Tambo and Suscal, north
rim of the valley of Rio de Canar, Camp E-2757 (MICH).
Eugenia variareolata McVaugh, sp. nov.
Arbor vel frutex, rufo-tomentosus; foliis maximis, 30-45 cm. longis obovatis
oblanceolatisve, supra lucidis, subtus minute ceroso-papillosis glaucisque; margin-
ibus cartilagineis; venulis utrinque reticulatis; racemis abbreviatis; bracteolis
2.5-3 mm. longis persistentibus; calycis lobis intus glabris, ovatis, 7-9 mm. longis,
ad florendi tempus reflexis, deciduis; disco 4.5 mm. lato.
A tree or shrub, the branchlets, inflorescence and petioles closely tomentose
with coarse flexuous lustrous tangled dark red-brown hairs up to 0.5 mm. long,
a few hairs persistent on the veins of the lower leaf-surface; leaves obovate or
oblanceolate, 10-15 cm. wide, 30-45 cm. long, about 3 times as long as wide,
rounded from above the middle to a broad short acumen, and narrowed from near
or above the middle to near the base, where abruptly contracted, subcordate, with
low rounded basal lobes; cartilaginous margin (about as thick as the marginal vein
on the upper surface) passing abruptly into the flat summit of the petiole, which
is 3-4 mm. thick, 10-15 mm. long; midvein convex and sometimes shallowly
sulcate above, 1.5-2 mm. wide at base, prominent beneath; lateral veins 12-15
pairs, ascending, convex but sometimes also impressed above, prominent beneath;
marginal vein (5-) 10-20 mm. from margin, evident but appearing as a series of
asymmetric loops formed by the laterals; minor veins prominulous outside the
marginal vein, including 2 successively smaller submarginal veins forming rather
symmetrical arches connected to the inner veins by small right-angled veinlets;
veinlets on both surfaces forming angular areoles of varying sizes; upper surface
of blade smooth, lustrous, green or drying brown, the lower surface reddish brown
or paler, glaucous, minutely waxy-papillose; both surfaces obscurely gland-dotted;
McVAUGH: TROPICAL AMERICAN MYRTACEAE 223
inflorescence an abbreviated axillary raceme (or 2-3 racemes from the same axil),
the axis up to 5 mm. long, bearing as many as 4 approximate, decussate pairs of
flowers on pedicels 1-2 mm. thick, 10-12 mm. long; bracts scarious, ovate or
subrotund, 1-2 mm. long; bracteoles broadly ovate to suborbicular, broad at base,
persistent, 2.5-3 mm. long; hypanthium bluntly 4-angled, 2-3 mm. long, hemi-
spheric to subglobose; calyx-lobes membranaceous, thin-margined, ovate, bluntly
pointed, 4-6 mm. wide near base, 7-9 mm. long, glabrous within, reflexed at
flowering time and finally dehiscent; disc quadrangular, 4.5 mm. wide, its center
1.5 mm. wide, deeply depressed (1 mm.), with red-hirsute margin; style glabrous,
more than 10 mm. long; ovary bilocular, the ovules about 35 in each locule,
attached radially to the central septum.
A most distinctive species, of which unfortunately neither buds,
complete flowers nor fruits are known.
Colombia, Meta: Villavicencio, elev. 450 meters, Jan., 1856,
J. J. Triana 14 (BM, type; COL; NY). Univ. of Mich. Neg. 485.
Eugenia versicolor McVaugh, sp. nov.
Arbor, ramulis racemisque puberulis, pilis pallide rufis, crispiusculis, brevibus,
obtectis; foliis 6-13 cm. longis acuminatis, venis utroque latere 6-10; racemis
abbreviatis; bracteolis persistentibus, connatis; calycis lobis late rotundatis, intus
appresse pubescentibus, 1.5-2 mm. longis; stylo 7-9 mm. longo; folii pagina
inferiore ferruginea vel cinerea, pilis crebris minutissimis nitidis obtecta.
A tree to 15 meters high with elliptic leaves 2-3 times as long as wide, rela-
tively few veins, and umbelliform clusters of flowers; the lustrous upper surface
of the leaves contrasts markedly with the rusty or ashy color of the lower surface;
buds 4.5-6 mm. long; disc 2-2.5 mm. wide; stamens about 200.
Compared in the key with E. heterochroma Diels, and perhaps
related to that species or to E. ferreiraeana Berg, the type of which
came from near the mouth of the Rio Negro, Brazil.
Brazil, Amazonas: Basin of Rio Solimoes, Mun. Sao Paulo de
Olivenca, basin of Creek Belem, high forest, terra firma, Oct. 26-
Dec. 11, 1936, B. A. Krukoff 8910 (MICH; US). Colombia,
Amazonas : Trapecio amazonico, Loretoyacu River, elev. 100 meters,
Nov., 1945, R. E. Schultes 6959 (US, type).
5. PLINIA L.
1. Pedicels 4-5 mm. long; flowers 2-4 pairs in short racemes; buds completely
closed, apiculate, 7-8 mm. long; plants appressed-hispidulous with no long
silky hairs P. clausa McVaugh.
1. Flowers sessile or subsessile, in sessile clusters subtended by sterile bracts;
buds, if closed, 12 mm. long; plants variously hirsutulous or silky-pilose
in the inflorescence.
2. Buds 12 mm. long, completely closed, whitened and felted with long ap-
pressed hairs; stamens about 500; leaves 4-6.5 cm. wide.
P. inflata McVaugh.
224 FIELDIANA: BOTANY, VOLUME 29
2. Buds 6-7 mm. long or less, silky-pilose or hirsutulous, the calyx-lobes free
at tips; stamens 125-150 (number unknown in P. pinnata); leaves various.
3. Leaves hirsutulous beneath, with hairs about 0.5 mm. long, the veins with
some longer hairs up to 2 mm. long; hypanthium 2-2.5 mm. across; style
4.5-6 mm. long P. pinnata L.
3. Leaves with minute hairs 0.2 mm. long on the lower surface, the young
branchlets and leaves having also some hairs up to 4 mm. long; hypanthium
3.5-4.5 mm. wide; style 9-11 mm. long P. duplipilosa McVaugh.
Plinia clausa McVaugh, sp. nov.
Arbor vel frutex, ramulis foliis novellis petiolisque ochraceo-hispidulis;
alabastris albidis, clausis, apiculatis, 7-8 mm. longis; racemis usque ad 3 mm.
longis, 4-8-floris, floribus pedicellatis; hypanthio supra germen circiter 4 mm.
producto; staminibus 200-250.
A tree or shrub with nearly glabrous, elliptic and narrowly acuminate leaves
7-10 cm. long, 2-2.3 times as long as wide; the flowers are in short racemes with
conspicuous membranaceous bracts and bracteoles 2.5-4 mm. long; stamens
arising from a broad zone occupying most of the distal half of the bud, their bases
intermixed with short silky hairs, the inner surface of the hypanthium glabrous
Peru, Loreto: Soledad (lower Rio Itaya, near Iquitos), July, 1925,
G. Tessmann 5287 (NY, type).
Plinia duplipilosa McVaugh, sp. nov.
Arbor, ramulis foliisque novellis pilosis, pilis albidis rectis tenuibusque, usque
ad 4 mm. longis, obsitis; glomerulis dense sericeo-pilosis; ramulis petiolisque, et
folii paginae inferioris venis minute pubescentibus, pilis erectis 0.2 mm. longis
immixtis; foliis 10-13 cm. longis acuminatis; glomerulis 4-floris, sessilibus, bracte-
atis; bracteis 4-seriatis, inferioribus sterilibus; bracteolis 5-6 mm. longis, pilosis
ciliatisque; alabastris 6.5 mm. longis, calyce 4-dentato, dentibus deltoideis 1.5
mm. longis; hypanthio supra germen 2.5-3 mm. producto; stylo 9-11 mm. longo;
A tree with elliptic-ovate leaves about 2.5 times as long as wide and 6-10
pairs of lateral veins; the rather large silky flowers are in clusters of 4 in leafless
axils on old wood, the clusters subtended by 4-ranked sterile bracts; buds con-
cealed by straight hairs 2.5 mm. long; calyx and hypanthium glabrous within,
the limb at maturity nearly quadrangular, 5-6 mm. on a side, the receptacular
cup 3.5-4.5 mm. wide.
The flowers and inflorescence in Klug's collection are very like
those of the type, but the leaves are longer and narrower, more
prominently veined beneath, and with up to 15 pairs of veins.
Apparently Cuatrecasas' no. 7092, from the lowlands of eastern
Colombia, is also conspecific; the flowers in this collection have more
elongate bracts and free calyx-tips, but are otherwise much like
McVAUGH: TROPICAL AMERICAN MYRTACEAE 225
Peru, Loreto: Yurimaguas, elev. 135 meters, dense forest, Aug.-
Sept., 1929, Killip & Smith 28007 (NY; US 1461669, type); Mishu-
yacu, near Iquitos, elev. 100 meters, forest, G. Klug 1155 (US).
Plinia inflata McVaugh, sp. nov.
Arbor hispidula, pilosa etiam, pilis eburneis 1-1.5 mm. longis, tenuissimis,
obsita; glomerulis ut videtur 4-floris, pilis longis concretis dealbatis; foliis 9-15
cm. longis, 4-6.5 cm. latis acuminatis; bracteis sterilibus 3-4-jugis, 4-seriatis;
alabastris clausis, apiculatis, 12 mm. longis, calyce demum in lobos 4 subaequales,
10-12 mm. longi, longitudinaliter direpto; hypanthio supra germen 2 mm. pro-
ducto; staminibus circiter 500; petalis minusculis 3 mm. longis.
A tree to 9 meters high, with elliptic-oblong leaves about 2.5 times as long
as wide, and about 15 pairs of very slender lateral veins; the plant appears to
differ from others in the same genus in its larger flowers, which are markedly
whitened by the matted hairs; the splits between the calyx-lobes extend some-
what deeper than the inner (proximal) margin of the broad hairy staminal ring,
which is 6-7 mm. wide and extends distally to a line 1.5-2 mm. from the apex
of the bud; style not seen.
Brazil, Amazonas: Basin of Rio Madeira, Mun. Humayta, near
Tres Casas, low terra firma, rare, Sept. 28, 1934, B. A. Krukoff 6365
(NY, type; US); same locality, Oct. 8, 1934, Krukoff 6525 (NY).
6. PSIDIUM L.
1. Calyx-lobes 7-9 mm. long and distinct, or the lobes prolonged into narrow
appendages 6-14 mm. long.
2. Calyx-lobes 4, ovate, 7-9 mm. long, tomentose without, appressed to the
bud and covering it, apparently valvate P. ulei Diels.
2. Calyx-lobes 5, separating irregularly as segments 4.5-5 mm. long and wide,
each tipped by an erect or spreading narrow foliaceous nearly glabrous
appendage 1.5-3 mm. wide, 6-14 mm. long P. caudatum McVaugh.
1. Calyx-lobes 3 mm. long or less, short, broad and rounded, or the buds com-
pletely closed before anthesis and dehiscing irregularly.
3. Leaves crenate, obtuse and cuspidate, narrow (1-2 cm. wide, 2-7 cm. long);
plants finely pubescent; flowers solitary; buds glabrous, about 12 mm. long.
P. maribense DC.
3. Leaves entire, or sometimes irregularly undulate, rarely less than 2.5 cm.
wide and if so acute or acuminate; pubescence and flowers various.
4. Plants completely glabrous, even to the young vegetative buds; branchlets
compressed, not angled; leaves 3-5 cm. wide, 7-14 cm. long, narrowed from
below or near the middle to the acute and mucronate tip; calyx open, slightly
flaring in bud, the broadly rounded lobes 2 mm. high. . .P. densicomum DC.
4. Plants with evident and usually abundant pubescence (sometimes on young
growing parts only); branchlets often angled; leaves and calyx various.
5. Lateral veins 12-20 pairs, usually impressed above, prominent beneath and
well differentiated from the smaller intermediate veins; young growth
heavily pubescent; branchlets 4-angled or sometimes terete; buds closed,
10-16 mm. long, pointed, not cuspidate; peduncles 1 (very rarely 3) -flowered.
226 FIELDIANA: BOTANY, VOLUME 29
6. Leaves lanceolate or narrowly ovate, 1.5-3 cm. wide, 3-4 times as long as
wide, gradually narrowed to the acute or acuminate tip; fruit with 8-15
irregular longitudinal ridges P. rutidocarpum G. Don.
6. Leaves elliptic or oblong, 3-6 cm. wide, 2-3 times as long as wide, more
abruptly narrowed to the obtusely pointed or rounded tip; fruit globose or
pyriform, smooth P. guajava L.
5. Lateral veins 6-15 pairs (mostly 10 pairs or fewer), if more numerous then
slender and scarcely differentiated from the intermediate veins, or the plants
very sparsely pubescent only; branchlets and flowers various.
7. Lower leaf-surface hirsutulous, the numerous hairs erect or nearly so, 0.5-1
8. Leaves mostly 6-10 cm. long; pubescence of branchlets reddish; buds 10-12
mm. long, almost completely closed; peduncle 1.5-3 cm. long, 3-flowered.
P. guineense Sw.
8. Leaves mostly 5-6.5 cm. long; pubescence tawny yellow; buds 3.5-4 mm.
long, with 5 broad low calyx-lobes 1 mm. long; peduncle 1 cm. long or less,
3- to 7-flowered P. fulvum McVaugh.
7. Lower leaf-surface glabrous or sparingly pubescent or strigose.
9. Leaves small for the genus, mostly 5.5 cm. long or less, often acute at both
ends; buds 5-7 mm. long, open, the 5 broad low calyx-lobes much shorter
than the corolla; branchlets terete or sometimes longitudinally channeled.
10. Dichasia 3-flowered, with very slender divaricate branches; calyx-lobes 1 mm.
long or less; marginal vein of leaf strongly arcuate, 1.5-4 mm. from the
margin at the points where the arches join the lateral veins; glands not
apparent even in young leaves P. pedicellatum McVaugh.
10. Flowers solitary; calyx-lobes more than 2.5 mm. long; marginal vein about
1 mm. from margin, not strongly arcuate; leaves gland-dotted on both sides
at least when young P. arayan (HBK.) Burret.
9. Leaves larger, 5-10 cm. long or even longer; buds 10-16 mm. long, closed
at apex; branchlets various.
11. Branchlets quadrangular, wing-angled; buds shortly apiculate; lateral veins
10 pairs or fewer; leaves usually with very numerous dark raised glands
beneath; Amazonian lowlands, widely distributed P. acutangulum DC.
11. Branchlets compressed, sometimes with low rounded ridges, not wing-angled;
buds with linear or subulate apiculum 2-5 mm. long; lateral veins 10-15
pairs; leaves with numerous small open glandular depressions of varying
sizes, on both surfaces; Pacific slopes, Tumbez P. rostratum McVaugh.
Psidium caudatum McVaugh, nom. nov. Psidiopsis moritzi-
ana Berg, Linnaea 27: 351. 1856, non Psidium moritzianum Berg,
I.e. 359. Calycolpus moritzianus (Berg) Burret, Repert. Sp. Nov. 50:
As far as I am aware, no one has been able to point out any
distinguishing feature of Psidiopsis except the prolonged tips of the
calyx-lobes. This is certainly noteworthy but scarcely indicative
of a profound evolutionary hiatus between the one species which is
so marked, and the other species of Psidium.
Psidium fulvum McVaugh, sp. nov.
Arbor vel frutex, fulvo-hirsutulus; foliis integris ovatis, 5-6.5 cm. longis
obtusis; venis utroque latere 8-10; pedunculis usque ad 1 cm. longis, 3-7-floris;
McVAUGH: TROPICAL AMERICAN MYRTACEAE 227
alabastris 3.5-4 mm. longis; calycis lobis 5, 1 mm. longis, 2-2.5 mm. latis; disco
5-angulato, 3-4 mm. lato; germine 3-loculari, sporophoris axillaribus peltatis
A shrub or tree, rather conspicuously and densely tawny-hirsute with hairs
up to 0.5 mm. long; leaves broadly ovate, 1.5-2 times as long as wide; flowers
small; style glabrous, not seen fully expanded, probably peltate; stamens about
Peru, Amazonas: Chachapoyas, A. Mathews ["derniere collect."]
(BM, type; G). Univ. of Mich. Neg. 484.
Psidium pedicellatum McVaugh, sp. nov.
Arbor, puberula; foliis integris immaturis 3-5.5 cm. longis, brevi-acuminatis;
venis utroque latere 7-10; pedunculis usque ad 1.5 cm. longis, ramis filiformibus
5-6 mm. longis divaricatis; dichasiis 3-floris; alabastris 5 mm. longis; calycis
margine arcuato-undulato 5-lobato, lobis 0.8-1 mm. longis; disco glabro, 2.5 mm.
lato; germine 2-loculari, sporophoris axillaribus peltatis 20-ovulatis.
A tree 12 meters high with scant puberulence, the leaves broadly elliptic-ovate
and 1.5 (-2) times as long as wide; glands not apparent even in young leaves;
dichasia usually 3-flowered, the flowers occasionally solitary; flowers small, the
style 4-5 mm. long, the stigma subpeltate; stamens about 150.
Ecuador, Santiago-Zamora : Along Quebrada Honda, vicinity of
Rancho Achupallas, elev. 2,500-2,700 meters, along river, Oct. 10,
1943, J. A. Steyermark 54571 (F 1391169, type). Univ. of Mich.
Psidium rostratum McVaugh, sp. nov.
Arbor vel frutex, pubescens vel strigosus; foliis integris 10-14 cm. longis
acutis vel apice obtusis; venis utroque latere 10-15; pedunculis 1-floris, axillaribus,
vel infimis ex nodis efoliatis, vel rhachi abortiva, oriundis; alabastris 12-16 mm.
longis clausis, longe apiculatis; calyce intus dense strigoso; disco 12-14 mm. lato.
A shrub or tree to 10 meters high, rather sparingly pubescent, with irregularly
ovate to oblong or even obovate leaves about twice as long as wide, and irregularly
glandular-pitted on both surfaces; flowers large, the style 10-13 mm. long, the
stigma peltate; stamens probably about 300.
Peru, Tumbez: Mountains east of Hacienda Chicama, in decidu-
ous bushwood, elev. 900-1,000 meters, Feb. 19-24, 1927, A. Weber-
bauer 7648 (F 571783, in flower; F 571784, type, in bud). Univ. of
Mich. Negs. 471 (type), 494.
7. MYRTEOLA Berg
1. Plants glabrous and subherbaceous; stamens 12 or fewer; northern Peru to
Venezuela M. oxycoccoides (Benth.) Berg.
1. At least the young branches densely pubescent, or if exceptionally the whole
plant glabrous, an erect shrub with 30 or more stamens.
228 FIELDIANA: BOTANY, VOLUME 29
2. Leaf-margins strongly revolute; leaves densely strigose or setose beneath;
stamens more than 30; northern Peru and Ecuador. [M. acerosa (Berg)
Burret; M. microphylla (Humb. & Bonpl.) Berg, var. microphylla.]
2. Leaf-margins not or scarcely revolute; leaves glabrous beneath or sparingly
3. Stamens 20 or fewer; leaves mostly broadest below the middle, with a tend-
ency to become bullate at least in age; plants with extensive rhizomes, the
flowering branches prostrate or erect, often less than 20 cm. high (up to
1 meter); bracteoles 1.5-3 mm. long; calyx-lobes 1.5-2 mm. long.
4. Leaves narrowly lanceolate, 1.5-2 mm. wide, 6-7 mm. long, bullate from the
first, the midvein not apparent beneath.
M. vaccinioides var. carabaya McVaugh.
4. Leaves ovate, 2.5-5 mm. wide, 4-8 mm. long, bullate in age or not at all, the
midvein apparent at least in young leaves.
M. vaccinioides (HBK.) Berg, var. vaccinioides.
3. Stamens 30-65; leaves broadest near the middle or but slightly below it, not
bullate or only exceptionally so, the midvein apparent beneath; erect shrubs
mostly 1-2 meters high; bracteoles 3.5-6 mm. long; calyx-lobes 2.5-3.5
mm. long. [M. microphylla var. glabrata Berg; M. weberbaueri Diels.]
Myrteola vaccinioides (HBK.) Berg, var. carabaya McVaugh,
Frutex, strigoso-hispidulus, rhizomatus; ramis floriferis 6-15 cm. longis erectis;
foliis anguste lanceolatis, subtus bullatis, nervo medio haud manifesto; bracteolis
2-2.5 mm. longis; staminibus circiter 15.
Further revisionary study in this genus, or examination of ad-
ditional material of the present plant, may well indicate that this
is an independent species. Unquestionably, however, it is akin to
M. vaccinioides, which it much resembles in habit, in stamen-number,
and in all vegetative characters except the leaf-shape.
Peru, Puno: Prov. Carabaya, June-July, 1847, H. A. Weddell
4667 (P, type). Bolivia, La Paz: Unduavi, en bosques, elev. 3,300
meters, Feb. 12, 1907, 0. Buchtien 647 (NY). Univ. of Mich.