JUL 2 4 1984

FIELDIANA
Botany

Published by Field Museum of Natural History

Volume 36, No. 10 September 9, 1975

Tropical American Plants, XVII

Louis O. WILLIAMS

CURATOR EMERITUS, DEPARTMENT OF BOTANY
FIELD MUSEUM OF NATURAL HISTORY

We acknowledge with thanks the long continued support of the
National Science Foundation which has made possible intensive
field work in Guatemala and in other countries of Central America,
with the resultant accumulation of basic research materials already
in use by many botanists. More importantly this support has made
possible the floristic studies which the source of "Flora of
Guatemala" and may well provide the basis for.a>^JJt$ptic flora of
central Central America. - \AB*^^

COMPOSITAE, TRIBE EtipATO^ifiAB^ '
IN GUATEMALA AND ADJACENT* REGIONS

The Eupatorioid Compositae of GwaxqgRaRrare one of the most
interesting groups of plants in the country and one of the most
visible in the highlands at the end of the wet season. Most are
species that occur in regions where a wet and a dry season is well
pronounced. However, there are species characteristic of the wet
montane regions and of the wet lowland Atlantic area. Eupatorioids
are predominantly montane plants and in the Central America
highlands they are abundant in both species and individuals, but
nowhere so abundant as in western highland Guatemala.

There will be 10 Eupatorioid genera included in the "Flora of
Guatemala" and only Eupatorium is large. Stevia, a very large and
complicated genus in Mexico, is in Guatemala in limited numbers
and there was no burst of endemic species as happened with
Eupatorium.

Library of Congress Catalog Card Number: 61-17730
US ISSN 0015-0746

Publication 1210 77

_. BIOLOGY LIBRARY

78 FIELDIANA: BOTANY, VOLUME 36

AGERATUM

The genus Ageratum has been studied critically twice in this
century. First was Dr. B. L. Robinson's (Proc. Am. Acad. 49: 438-
491. 1913) study of Ageratum (and the allied genera Alomia and
Oxylobus) and, second, the very recent monograph by Miles F.
Johnson (Ann. Mo. Bot. Card. 58: 6-88. 1971). This is a bit of a
record for tropical American plants. Dr. Johnson did not consider
those plants usually placed in Alomia, saying that they were
distinguished from Ageratum by lack of a pappus. More recently
King and Robinson (Phytologia 24: 108-111. 1972) maintained the
genus Alomia HBK. in a limited sense, but say that the genus in
the broad sense "contains six very distinct elements." In their
account four species survive as the genus Alomia. In a following
paper (King and Robinson, Phytologia 24: 112-117. 1972) several
species of Alomia were transferred to Ageratum. In a third paper
(King and Robinson, Phytologia 24: 118-119. 1972) the genus
Blakeanthus, based on a species of Alomia, was established.

In the "Flora of Guatemala" Ageratum will be used in
approximately the traditional sense, with the addition of those
Guatemalan plants sometimes referred to Alomia, including
Blakeanthus.

Field studies made in Honduras indicated that epappose
Alomia microcarpa (Benth.) Rob. and pappose Ageratum con-
yzoides L. growing in the same old corn field were not
distinguishable, except by presence or absence of pappus. It was
concluded that the two species are synonymous (perhaps differing
by a single gene) and this indicates the direction of generic
consolidation which we follow. Ageratum pinetorum and A.
littorale, as noted below, both have pappose and epappose
populations.

Ageratum cordatum (Blake) L. Wms. comb. nov. Alomia
cordate Blake, Proc. Biol. Soc. Wash. 60: 41. 1947. Blakeanthus
cordatus King & Robinson, Phytologia 24: 119. 1972.

I do not find King and Robinson's reasons for establishing a
genus based on this species very impressive. The species is
reasonably distinctive and fits into the genus Ageratum with no
difficulty.

Ageratum isocarphoides (DC.) L. Wms. comb. nov.

WILLIAMS: TROPICAL AMERICAN PLANTS, XVII

79

FIG. 1. Ageratum cordatum. A, branch from plant, x Vfc; B, capitulum, X 6, with
flowers at anthesis; C, flower at anthesis, x 22; D, flower with corolla removed, x
22; E, portion of corolla, dissected, x 22.

Coelestina isocarphoides DC. Prodr. 5: 107. 1836. Alomia isocar-
phoides Rob. Proc. Am. Acad. 49: 449. 1913.

This name appears in Hemsley, Biol. Cent. Am. Bot. 2: 81.
1881, as "Ageratum isocarphoides, DC." Hemsley apparently did not
intend this name as a transfer and a basionym is not given. The

80 FIELDIANA: BOTANY, VOLUME 36

name as used is a nomen nudum. The Index Kewensis attributes the
name to Hemsley without basionyms. This is covered in the
International Code of Botanical Nomenclature, Article 33, note 1.
Recommendation 72A of the rules states that "Authors should
avoid adoption of an illegitimate epithet previously published for
the same taxon." However, since there has been no confusion of the
plant intended, I prefer to use the name Ageratum isocarphoides.

Johnson in his monograph of Ageratum excluded this species,
referring it to Alomia as B. L. Robinson had done years before.
Johnson did include Ageratum echioides (Less.) Rob., a very closely
allied species which both De Candolle and Robinson pointed out,
and A. echinoides and A. isocarphoides certainly should be placed
in the same genus. Johnson has annotated two Guatemalan
specimens, Steyermark 50731 and 51500, as Ageratum echioides but
these seem to belong in Ageratum isocarphoides.

Ageratum littorale Gray, Proc. Am. Acad. 16: 78. 1880;
Johnson, Ann. Mo. Bot. Gard. 58: 70. 1971. A. littorale var.
hondurense Rob. Proc. Am. Acad. 49: 468. 1913.

This species occurs on Roatan Island, Honduras, from whence
came the type of the var. hondurense. A recent collection from
Roatan is Molina 20693 which is apparently the second known
collection from Central America. The pappus on this collection is a
minute corona. Robinson (1913, p. 440) commented on the calvous
forms of Ageratum littorale and states that it would be artificial to
transfer them to Alomia. We are placing all Central American
alomias into Ageratum.

Ageratum pinetorum (L. Wms.) King & Robinson, Phytologia
24: 115. 1972. Alomia pinetorum L. Wms. Fieldiana, Bot. 31: 25, fig.
4. 1964.

The type collection of this epappose species was a large one,
with 25 duplicates and these now widely distributed. In a re-
examination of the type, one plant on the herbarium sheet was seen
to bear a pappus of setiferous scales! The two plants on the type
sheet certainly differ in no other way. Another collection of this
species, Standley 27410, from approximately the type locality, has
quite epappose achenes (but with a corona). Standley's collection
was determined by Johnson (who excluded Alomia from Ageratum)
as Ageratum corymbosum f. longipetiolatum (Rob.) Johnson.
Eupatorioid Compositae are not always easy!

WILLIAMS: TROPICAL AMERICAN PLANTS, XVII 81

Ageratum platylepis (Rob.) King & Robinson, Phytologia 24:
113. 1972. Alomia platylepis Rob. Proc. Am. Acad. 49: 448. 1913.
Alomia quatemalensis Rob. I.e. syn. nov., not Ageratum guatema-
lense Johnson, 1971. Ageratum ben/amin-lincolnii King &
Robinson, I.e.

Dr. Robinson described these two species and distinguished
them by characters that have proven to be untrustworthy or
questionable even on the isotypes which I have studied. The ovate
or ovate-oblong ecostate phyllaries of Alomia platylepis Rob. are
oblong-lanceolate or linear-lanceolate and costate exactly as in the
isotype of Alomia guatemalensis Rob.

Ageratum robinsonianum (L. Wms.) L. Wms. comb. nov.
Alomia robinsonianum L. Wms. Fieldiana, Bot. 31: 27. 1964.

This Mexican species is transferred to Ageratum as there seems
to be no question that the condition of the pappus is not valid for
distinguishing Alomia from Ageratum.

BRICKELLIA

Brickellia pacayensis Coulter, Bot. Gaz. 16: 98. 1891;
Robinson, Contr. Gray Herb. 1: 95, fig. 74. 1917. Coleosanthus
pacayensis Coulter, Bot. Gaz. 20: 46. 1895.

Originally described from Pacaya Volcano in Guatemala the
species is now known from south Mexico to Honduras. Like many of
the species in the subsection Coleosanthus (Cassini) Rob., it is
somewhat difficult to circumscribe. There is apparent intergression
between B. pacayensis and the complex of Guatemalan material
that I call B. paniculata.

Brickellia paniculata (Miller) Rob. Proc. Am. Acad. 42: 48.
1906. Eupatorium paniculatum Miller, Card. Diet. ed. 8, No. 15.
1768. E. rigidum Benth. PL Hartw. 88. 1841, not Sw. (type from
valley of Guatemala, Hartweg 598). Brickellia hartwegii Gray, PL
Wright. 1: 85. 1852. Coleosanthus rigidus O. Kuntze, Rev. Gen. PL
1: 328. 1891. Brickellia argyrolepis Rob. Mem. Gray Herb. 1: 90, fig.
69. 1917 (type from Costa Rica, collection not specified). B.
guatemalensis Rob. I.e. 92, fig. 71 (type from Alta Verapaz,
Tuerckheim II. 2145). B. adenocarpa Rob. I.e. 94, fig. 73 (type from
Quezaltenango, Holway 466). B. adenocarpa var. gladulipes Rob.
I.e. (type from Quezaltenango, Holway 92).

82 FIELDIANA: BOTANY, VOLUME 36

Dr. B. L. Robinson in his "Monograph of the Genus Brickellia"
(Mem. Gray Herb. 1: 1-151. 1917) separated the species once
considered as Coleosanthus Cassini into section Bulbostylis subsec-
tion Coleosanthus. There were 26 species in the subsection with
many of the species perhaps too much alike. In Guatemala
botanists have distinguished five or six of these species. With a very
great increase in the study material available, it now seems to me
that this number may be reduced to two: B. paniculata, which is a
polymorphic aggregation, and B. pacayensis, mentioned above,
which may intergrade with B. paniculata. I have studied the
problem only briefly with the purpose of finding a stable name for
the Central American material. It seems likely that other species
from Mexico differ only slightly from my concept of B. paniculata.
the oldest name in the subsection. This small group of brickellias
should provide material for an interesting cytotaxonomic study.

EUPATORIUM

The genus Eupatorium is one of the largest genera in the
Guatemalan flora. At the end of the rainy season, from November
to January, it is found everywhere in the highlands. The abundant,
many-headed inflorescences make the genus visible and attractive.
We (Williams, Molina, and Williams) spent the two collecting
seasons, 1972-1973 and 1973-1974, making a special effort to get as
many collections of Compositae as possible during their principal
flowering season. The number of collections made and the variety
of species found in Guatemala by our colleagues and ourselves
indicate that Guatemala with the borderlands of Chiapas (Mexico)
and British Honduras (now Belize), is the richest area in the world
for Eupatorium. Mexico has a greater number of species, but in a
land area 18 times as large.

Eupatorium, so abundant in Mexico and Guatemala, decreases
rapidly in numbers, as well as in individuals, proceeding south-
eastward down the mountains to Panama. The genus increases in
numbers again in South America, but probably nowhere is it so
prominent a component of the flora as in Guatemala.

Eupatorium, like other large genera distributed from the
tropical, wet or dry, lowlands to the high montane regions, shows
much altitudinal stratification and distribution. The species found
in the lowlands are more likely to be widespread; those of middle
elevations may be distributed from Mexico to Costa Rica or
Panama; and those from the highest montane forest regions may

WILLIAMS: TROPICAL AMERICAN PLANTS, XVII 83

well be endemic. Quite a few species, like Eupatorium monticola
described below, are found in the high mountains of south Mexico,
skip across the Isthmus of Tehuantepec, and are found again in the
high mountains of Chiapas and Guatemala, but continue no further
southeastward along the Central American isthmus.

The "Flora of Guatemala" will be the first large floristic work
to appear, in fact, the only considerable study of any regional group
of Eupatorium since King and Robinson initiated their studies of
the generic limits within the tribe Eupatorieae. The decision to use
the broad generic concepts in working out the Eupatorieae for
Guatemala, instead of following the proposals of King and
Robinson, is mine. I am quite sure that the late Dr. Standley would
agree.

Eupatorium is a large genus with perhaps as many as 600
species, most of them located in the Americas, but with a few
species in Europe, Asia, and Africa. Guatemala has more than 85 of
these species.

King and Robinson have now proposed or resurrected more
than 50 Eupatorioid genera, of which not less than 18 have been
applied to species occuring in Guatemala. In preparing the account
of Eupatorium I have considered these generic concepts and have
found them both useful and detrimental in grouping species for
study. It seems obvious to me that these concepts are of uneven
quality. The genus Neomirandea, proposed by King and Robinson,
contains species that might better have been placed into the
segregate Ageratina. Ageratina, itself, might have been divided in
two on quite obvious characters in the phyllaries, and these
concepts would have been quite as good as some of those based on
more abstruse characters. King and Robinson have placed elements
which seem to me to be conspecific in different genera, suggesting
that they may have been based on studies that were too superficial
or of inadequate material.

Eupatorium in Guatemala is a large, though manageable, genus
and certainly a fascinating one. To break it down into a group of
sometimes nebulous segregates would hardly increase the ease with
which the "Flora" is used. A flora should be useable to people of
many disciplines, not only to systematic botanists, and the scientific
output should be intelligible to all of them.

Eupatorium aschenbornianum Schauer, Linnaea 19: 720.
1847. E. donnell-smithii Coulter, Bot. Gaz. 16: 95. 1891. E. donnell-

84 FIELDIANA: BOTANY, VOLUME 36

smithii var. parvifolium Donn.-Sm. Bot. Gaz. 16: 69. 1891.
Ageratina aschenborniana King & Robinson, Phytologia 19: 212.
1970.

The type of E. aschenbornianum originated from the Toluca
valley in Mexico and the species is widely distributed from central
and south Mexico south to Panama. The types of E. donnell-smithii
and the variety parvifolium both originated in Guatemala. The
species is closely related to E. pazcuarense, with which it is often
confused. It may be distinguished, however, by the smaller heads
(rarely longer than 5 mm.), with the phyllaries as long as the
pappus; the leaves are triplinerved from the base of the blade, but
this character does not seem consistent.

Ku pa tori urn araliaefolium Lessing, Linnaea 6: 403. 1831.
Neomirandea araliaefolia King & Robinson, Phytologia 19: 307.
1970.

Macbride brought a fragment from the type to Field Museum
from Berlin. The photograph of the type is numbered 16202. It is of
interest to note that Scheide and Deppe's 1242 is also the basis for
Eupatorium sordidum Lessing, an error in numeration since the
localities given for the two species are different. The species is
usually epiphytic, but sometimes a shrub in the forest. The flowers
are white and the species is most common at middle elevations, but
grows from near sea level to somewhat less that 2,000 m.; the range
is from Vera Cruz in Mexico to Costa Rica. The species described
below, E. oreophilum and E. orogenes, are allied species.

Eupatorium burgeri (King & Robinson) L. Wms. comb. nov.
Ageratina burgeri King & Robinson, Phytologia 24: 85. 1972.

A Costa Rican species apparently well distinguished from the
eupatoria that I know. The type was collected by Molina et al.
17782, in 1966.

Eupatorium capillipes Benth. in Oersted, Vid. Medd. Kjoben-
havn 1852: 79. 1853. Ageratina helenae King & Robinson,
Phytologia 24: 90. 1972. Ageratina molinae King & Robinson, I.e.
93.

A quite abundant species in Guatemala and occasional from El
Salvador to Costa Rica. It should be found in adjacent Mexico. I
assume that King and Robinson did not know of E. capillipes

WILLIAMS: TROPICAL AMERICAN PLANTS, XVII 85

Benth. as they have not transferred it to any other genus. Both
Ageratina helenae and A. molinae are described as shrubs. Isotypes
available to me show the plants to be herbs, perhaps even annuals,
and this is my recollection of them in the field.

Eupatorium carmonis Standl. & Steyerm. Field Mus. Bot. 22:
303. 1940.

The type of this species was collected by Standley (63734) not
far from Antigua and a second collection was made at the Iximche*
ruins near Tecpan, Chimaltenango, Molina et al. 16125. The species
is closely related to E. capillipes Benth. I am able to distinguish it
only by the leaves which are truncate to broadly cuneate at the
base and 3-5-plinerved from the base. These are not very impressive
characters.

Eupatorium collinum DC. Prodr. 5: 164. 1836. E. neaeanum
DC. I.e. 160, syn. nov. Krystenia collina Greene, Leaflets 1: 9. 1903.
Chromolaena collina King & Robinson, Phytologia 20: 208. 1970.

This species is widely distributed through Mexico and south
through Guatemala, El Salvador, and Honduras. There is a
variation in Nicaragua and Costa Rica, undescribed to my
knowledge, which has much smaller heads with fewer flowers arid
with the phyllaries only slightly shorter than the florets.

De Candolle described two Mexican eupatoriums in the
Prodromus, E. collinum and E. neaeanum, of which phototypes are
available (F33612 and F33608). These photographs with descriptions
indicate that these taxa are well within the range of variation of the
species. This variation includes relatively compact shrubs with quite
small leaves, as small as 3-5 cm. long, and puberulent on the under
surface. In contrast, there are more robust plants with large leaves
(to 15 cm. long in Guatemalan specimens) that are almost glabrous
below and with the stems only puberulent. De Candolle placed E.
neaeanum into a group of species with 25-30 flowers in a head; and
E. collinum into a group with 30-100 flowers in a head. I find no
other characters in the photographs or the descriptions and for the
"Flora of Guatemala" I place the variable and abundant material
under E. collinum DC., the name which has been widely used.

Eupatorium eras sir ameum Rob., Proc. Am. Acad. 35: 332.
1900. Ageratina crassiramea King & Robinson, Phytologia 19: 228.

86 FIELDIANA: BOTANY, VOLUME 36

1970. Pachythamnus crassirameus King & Robinson, Phytologia 23:
154. 1972.

This species occurs from southern Mexico to Guatemala, El
Salvador, and Nicaragua. The material which we have is inadequate
and the only Guatemalan specimen with leaves is without flowers.
The altitudinal range of these specimens which have been
determined as E. crassirameum is rather too great from 2,500 m.
or more (Guatemalan specimen with leaves), with most specimens
found from 800 to 1,000 m., and one from Nicaragua at about 200
m. The ecological range is also too great from epiphytic in the
mixed forest in Guatemala to a dry, hot lava flow in Nicaragua.

The specimens all have thick, fleshy, and brittle stems which
are leafless at flowering time. No other species of Central America
is said to have these characteristics. It is reasonable to expect that
more than one species is represented.

Eupatorium cupressorum Standl. & Steyerm. Field Mus.
Bot. 23: 183. 1944. Ageratina cupressorum King & Robinson,
Phytologia 24: 89. 1972, as A. cupressora.

This species previously was known only from the cypress
forests in the department of Totonicapan (Standley 84001). We
collected it from three additional departments during our 1972-1973
field trip: from the department of Chimaltenango (on the border of
the department of Solola) at 2,000 m.; from the department of Baja
Verapaz at 1,600-1,800 m.; and from the department of Quiche at
1,800-2,000 m. The collection numbers are Williams, Molina
^Williams 41571, 41577, 41600, and 42140, soon to be distributed.

The phyllaries of the species are essentially equal with two or
three very short basal ones. The flowers in the heads are about 20.
In the original description the phyllaries were said to be 2-3-seriate;
the flowers in the heads 40 or more.

Eupatorium glaberrimum DC. Prodr. 5: 144. 1836. E.
oerstedianum Benth. in Oersted, Vid. Medd. Kjoebenhavn 1852: 74.
1853. E. vernonioides Coulter, Bot. Gaz. 20: 45. 1895. Eu-
patoriastrum opadoclinium Blake, Journ. Wash. Acad. 28: 479.
1938. Chromolaena glaberrima King & Robinson, Phytologia 20:
208. 1970. C. oerstedianum King & Robinson, I.e. C. opadoclinia
King & Robinson, I.e. Eupatorium opadoclinium McVaugh, Contr.
U. Mich. Herb. 9: 388. 1972.

WILLIAMS: TROPICAL AMERICAN PLANTS, XVII 87

It is unfortunate that the well-known name Eupatorium
oerstedianum on perhaps thousands of specimens in the herbaria of
the world must be substituted by a name that is relatively little
known but which was published several years previously. There is a
photograph (F 16261) of E. glaberrimum, the specimen that was in
the Berlin Herbarium from the original Haenke collection. In Field
Museum there is a "Reliquiae Haenkeanae" specimen that might
have come from the same collection. It was received undetermined
and was determined as E. oerstedianum Benth. by Standley. This is
not surprising since Standley knew Eupatorium oerstedianum very
well and E. glaberrimum possibly not at all.

The species is distributed from well north of the Isthmus of
Tehuantepec in Mexico to Costa Rica. In this range there are plants
that are almost glabrous to ones in which the stems and leaves are
densely pubescent. Intermediates are to be found and the
geographic ranges of the extreme phases are not separated. The
heavily pubescent phase (Eupatoriastrum opadoclinium) is most
abundant in south Mexico but occurs also in Honduras and Costa
Rica. The pales subtending the flowers in the heads, a presumed
character of Eupatoriastrum, only occasionally are found in this
species.

Eupatorium hebebotryum (DC.) Hemsl. Biol. Cent. Am. Bot.
2: 95. 1881; Robinson in Standley, Contr. U. S. Nat. Herb. 23: 1446.
1926. Critonia hebebotrya DC. Prodr. 5: 141. 1836.

The species is based upon a specimen collected in Mexico by
Haenke (phototype F33602), perhaps from the state of Morelos,
according to Dr. Robinson. It is considered to be distributed from
central Mexico to Costa Rica and specimens available seem to bear
this out, but the species belongs in a complex group where critical
studies would be helpful.

Eupatorium imitans Rob. Contr. Gray Herb. 68: 20. 1923.
Fleishmannia imitans King & Robinson, Phytologia 19: 203. 1970.

This species is an occasional plant along the edges of streams or
rivers, often growing at the water edge. It generally occurs at low
elevations, but occasionally is found up to 1,800 m. Eupatorium
imitans is one of a half-dozen species usually found along streams
and often covered at flood stage.

The species usually has the phyllaries arranged in three or four

88 FIELDIANA: BOTANY, VOLUME 36

series and, consequently, belongs in a group of species referred to
Fleishmannia by King and Robinson. However, the phyllaries often
are essentially equal in length and greater in number than in the
other species referred to that generic concept. The flower heads are
larger than those of other species and have a large number of
florets. Therefore, this species would seem to be intermediate
between King and Robinson's concepts of Fleishmannia and
Ageratina.

Eupatorium jejunum Standl. & Steyerm. Field Mus. Bot. 23:
183. 1944.

A delicate little wet-season annual not uncommon at middle
elevations in Guatemala. Superficially it seems related to the
complex of species near E. microstemon Cass., and especially to E.
sinclairii Benth. However, the phyllaries are equal in length rather
than graduated and seriated as in those species.

Eupatorium leucocephalum Benth. PL Hartw. 86. 1841. E.
mendax Standl. & Steyerm. Field Mus. Bot. 23: 185. 1944.
Fleishmanniopsis leucocephala King & Robinson, Phytologia 21:
403. 1971. F. mendax King & Robinson, I.e.

I find no reason to believe that the immature type of E.
mendax is different from E. leucocephalum.

Eupatorium lucentifolium L. Wms. sp. nov.

Arbusculae vel frutices usque ad 9 m. altae caulibus leviter complanatis obscure
puberulentis. Folia penninervia Integra subcoriacea lanceolato-ovata vel oblongo-
ovata abrupte acuminata basi acuta vel rotundata lucentes glabra manifeste
reticulata; inflorescentia terminalis multicapitata; capitulis sessilibus vel breviter
pedicellatis vulgo 6 mm. longis 5-floribus; phyllaria 3-4-seriatim segmenta exteriora
anguste ovata et interiora oblongo-ovata vel lineari-oblonga obtusa; achaenia 5-
porcata glabra; corolla tubularis glabra lobis angusti ovatis.

Shrubs or trees to 9 m. tall but usually less, stems slightly flattened at the
younger somewhat immature nodes, obscurely puberulent with appressed hairs.
Leaves penninerved with 6-8 pairs of secondary nerves, entire, subcoriaceous,
somewhat bullate, lanceolate-ovate or oblong-ovate, abruptly acuminate, somewhat
acute or rounded at the base, shining, glabrous except minutely puberulent on the
midvein below, the blade 7-15 cm. long and 2.5-6 cm. broad, tertiary nerves
prominently reticulate, especially above, petiole slender, appressed puberulent, 1-2
cm. long; inflorescence a terminal multicapitate obscurely puberulent profuse
corymbose panicle to 25 cm. long and as broad; the heads sessile or the pedicels not
more than 1 mm. long; the heads mostly about 6 mm. long and usually with 5 florets;
phyllaries 3-4-seriate with the outer ones narrowly ovate and the inner ones oblong-
ovate to linear-oblong, obtuse, the innermost about 5 mm. long and about as long as

WILLIAMS: TROPICAL AMERICAN PLANTS, XVII 89

the pappus; achenes prismatic and 5-ridged, glabrous, about 2 mm. long; pappus
about 4 mm. long, of 30-40 bristles; corolla pink or white, tubular, glabrous, about 4
mm. long and 0.3 mm. in diameter, the lobes 0.2-0.3 mm. long and narrowly ovate;
anthers minute, about 0.6 mm. long, appendaged at the apex; style at anthesis
exserted 3-4 mm. beyond the corolla, slightly thickened at the apex.

Mexico: flowers pink or white, tree 30 ft. tall, heavily wooded
slopes at Lago Tsiskaw on the Guatemalan border, 30 km. east of
La Trinitaria, municipio of La Trinitaria, alt. 4,500 ft. (1,360 m.),
April 14, 1965, Breedlove 9765 (type, F; MICH).

Guatemala: shrub 2 m. tall, forested quebrada along Rfo
Carcha between Coban and San Pedro Carcha, alt. about 1,360 m.,
March 26-27, 1941, Standley 89769 (F).

Closely related to Eupatorium nubigenum from which it may
be distinguished by the relatively broader entire leaves with
prominent tertiary reticulations (not minutely papillate), and by
the appressed puberulent stems and inflorescences. The leaves are
not pellucid-dotted as in E. nubigenum

The specimen from Guatemala is immature, but almost surely
this species. Dr. Breedlove's collection is very close geographically
to the department of Huehuetenango.

Eupatorium luxii Rob. Proc. Am. Acad. 36: 480. 1901. E.
pansamalense Rob. I.e. 482, syn, nov. E. oresbioides Rob. Proc. Am.
Acad. 44: 618. 1909. E. prionophylhim var. asymetrum Rob. Contr.
Gray Herb. 61: 11. 1920. Neobartlettia luxii King & Robinson,
Phytologia 21: 296. 1971. N. pansamalensis King & Robinson, I.e.
Bartlettina luxii King & Robinson, Phytologia 22: 161. 1971. B.
pansamalensis King & Robinson, I.e. fi. oresbioides King &
Robinson, I.e.

The three names involved in the synonomy were all proposed
by Dr. Robinson from highland Guatemalan specimens and each of
them is somewhat different from the others. However, the large
number of collections now available seem to close the gaps between
them and, in addition, there is another form, unknown to Dr.
Robinson, in which the phyllaries are somewhat lanate. The type
collection of E. luxii has the phyllaries quite broadly lanceolate or
lanceolate-triangular and acute, puberulent, black-purple when
fresh, and retaining this coloration in most dry specimens. The
phyllaries mostly reach to the middle of the pappus or a bit more.
Typically Eupatorium oresbioides is similar to E. luxii, but with
most of the phyllaries a little narrower and a little longer in

90 FIELDIANA: BOTANY, VOLUME 36

relation to the pappus and the longer ones, at least, obtuse. A form
has the phyllaries sordid-sublanate. Typically E. pansamalense has
leaves somewhat larger than those of E. oresbioides, but has
phyllaries much the same.

Since Eupatorium luxii and E. pansamalense were published at
the same time, I retain E. luxii which commemorates an early
Guatemalan collector, and indicate E. pansamalense as a new
synonym.

Eupatorium macrum Standl. & Steyerm. Field Mus. Bot. 23:
184. 1944.

The original specimen came from Chiquimula in Guatemala
and has been found recently in the Peten. The species is
occasionally abundant in the central mountains and valleys of
Honduras. We have distributed one collection of this species
Williams & Molina 23251 to more than 25 herbaria. The species
is related to E. leucocephalum Benth., but may be distinguished by
the sessile heads, the slender rather than enlarged ends of the
styles, and the pubescent leaves.

Eupatorium mairetianum DC. Prodr. 5: 167. 1836. E.
rafaelense Coulter, Bot. Gaz. 16: 97. 1891. E. mairetianum var.
adenopodum Rob. Proc. Am. Acad. 51: 534. 1916. Ageratina
mauretiana King & Robinson, Phytologia 19: 224. 1970. A.
rafaelensis King & Robinson, I.e. 225.

This is a difficult and not uncommon species at middle and
higher elevations in Guatemala. Whether or not all of the
Guatemalan material is one species or two, we must leave to the
monographer. There are apparent differences in the phyllaries, in
the nervation of the leaves, in the relative length of the heads and
phyllaries, in the base of the leaves. I have been unable to find a
way to use these differences for they seem not to be consistent.

Eupatorium magistri L. Wms. sp. nov.

Arbusculae vel lianae glabrae, caulis teretibus. Folia ovata vel elliptico-ovata
Integra acuta vel breviter acuminata. basibus rotundatis vel obtusis, petioli graciles;
inflorescentiae terminates vel axillares, paniculatae paucicapitatae; capituli 10-
floribus 8 mm. longi; involucre multiphyllariata 3-5-seriata, phyllariis ovatis ad
angusti-lanceolata; achaenia glabra 5-porcata; corolla cylindrica, lobis lanceolatis.

Shrubs or perhaps lianas to be found in the wet forests, glabrous, the stems
terete with opposite branches; leaves ovate to elliptic-ovate, acute or short
acuminate, rounded or obtuse at the base, 5-plinerved with the lower pair of nerves

WILLIAMS TROPICAL AMERICAN PLANTS, XVII 91

less conspicuous, blade 3-7 cm. long and 1.5-2.5 cm. broad, entire or nearly so, the
petiole slender, 0.5-1.5 cm. long; inflorescences terminal or axillary on the lateral
branches, paniculate, each 2-6 cm. long and with relatively few sessile or short-
pedicellate heads; the heads about 8 mm. long and 3-4 mm. broad each with 10
florets; involucre of many phyllaries in 3-5-series, the outer ones much shorter, from
about 1 mm. long and broadly ovate, obscurely ciliolate, the inner phyllaries longer,
5-7 mm. long and to narrowly lanceolate and acute; mature achenes glabrous, 5-
ridged, about 2 mm. long; pappus about 4.5 mm. long, as long as the corolla and with
30-40 bristles minutely barbellate at the apex; corolla about 4.5 mm. long and 0.3-0.4
mm. in diameter, cylindric, the lobes lanceolate, about 0.5 mm. long; anthers slender,
about 1.5 mm. long, appendaged at the apex; style at anthesis somewhat thickened at
the apex and exserted about 2 mm. beyond the corolla.

Guatemala: Cubilguitz, Alta Verapaz, August, 1907, Tuerck-
heim II. 1912 (type, F).

The species is related to E. bartlettii Rob. and was so
determined by Standley. It is distinguished by the smaller 5-
plinerved leaves. The heads are somewhat larger and with 10
florets.

In naming this species it is a pleasure to recall Benjamin
Lincoln Robinson, master of the genus Eupatorium and outstanding
student of the Compositae.

Eupatorium molinae L. Wms. nom. nov. Neomirandea
ovandensis King & Robinson, Phytologia 19: 309. 1970, not
Eupatorium ovandense Grashoff & Beam an, 1969.

The type of the species is from Mt. Ovando, state of Chiapas,
Mexico (Matuda 3917). The species has been found in Guatemala in
the department of San Marcos (Williams et al. 26203, 26863). It is
an epiphytic plant with attractive lavendar or purple flowers and a
species of the montane cloud forest. My colleague of many years in
the field in Central America, Prof. Antonio Molina R., years ago
had indicated the second of our collections mentioned above as an
undescribed species. It is a pleasure to rename the plant for him.

This species, while described in the segregate genus Neo-
mirandea by King and Robinson, should probably have been
assigned to the segregate Ageratina if they had wished to be
consistent.

Eupatorium monticola L. Wms. sp. nov.

Frutices vel arbusculae usque ad 5 m. alta, ramulis teretes dense ferrugineo-
tomentosis; folia membranacea, ovata vel deltoideo-ovata acuta vel acuminate,
breviter cuneata ad petiolos, crenato-serrata, 5-7-plinervia, supra glabra vel sparse
pubescentia subtus breviter villosulosa vel glabrescentes; petioli ferrugineo-tomentosi;

92 FIELDIANA: BOTANY, VOLUME 36

inflorescentiae corymbosae multicapitatae; capitula campanulata pedicellata 20-30-
floribus, plus duplo longioribus quam phyllaria; phyllaria subaequilonga linearia
acuta; achaenia 5-angulata ad angulos scaberula; corollae subcylindricae, lobis
lanciformis.

Stout shrubs or small trees 0.5-5 m. tall, the young branches terete, covered with
reddish, purplish or ferrugineous tomentum of somewhat viscid hairs; leaves opposite,
membranaceous, ovate to deltoid-ovate, acute or acuminate, shortly cuneate to the
petiole, crenate-serrate, 5-7-plinerved, with some nerves arising well above the base of
the blade, glabrous to usually sparsely puberulent above, usually short villosulous
below especially along the nerves or glabrescent, blades 5-17 cm. long and 3-10 cm.
broad (from very high elevations 3 cm. long or even less), petioles ferruginous
tomentose becoming glabrous, 1-4 cm. long; inflorescence corymbose, rounded, 6-15
cm. broad, dense with many heads or in age somewhat lax, with reduced leaves or
bracteolate; heads narrowly campanulate, pedicellate, with 20-30 lilac or purple
(rarely white) flowers, 8-10 mm. long at anthesis; involucre less than half as long as
the heads; phyllaries subequal, linear, acute, mostly 4-5 mm. long; achenes about 3
mm. long, 5-ridged, scaberulous on the angles, black; pappus with about 25 bristles,
about 4 mm. long; corolla subcylindric, gradually enlarged upward, about 5 mm. long,
lobes lanceiform, about 0.5 mm. long; styles exserted about 2 mm. at anthesis.

Guatemala: flowers rose, tree 3-5 m. high, mixed cypress-oak-
pine forest area, Sierra Madre Mountains about 10-15 km. south of
Totonicapan, Dept. Totonicapan, alt. 2,800 m., December 20, 1972,
Williams, Molina & Williams 41503 (type F; EAP, MICH, others).

ADDITIONAL SPECIMENS. -

Mexico: Breedlove 7998; Conzatti 2260; Pringle 6011; Purpus
2644; Smith 385, 1191.

Guatemala: Kellerman 5182; Molina 21223; Molina, Burger &
Wallenta 16444; Standley 60936, 61047, 81650, 81754, 81765, 85380;
Steyermark 34708, 35679; Williams, Molina & Williams 21969,
22901, 22918, 22978, 25470, 25878, 25881, 25883, 41471; Williams et
al. 26991, 27035.

The species is a common and attractive shrub or small tree in
the western highlands of Guatemala with an altitudinal range of
2,600 to 3,700 m. Those at the highest elevations are often not more
than one-half meter tall and with the leaves sometimes not more
than 3 cm. long.

The species has most often been confused with Eupatorium
subpenninervium Sch.-Bip. (a synonym of E. pazcuarense HBK.)
and with E. vernale Vatke & Kurtz, both species in which the
phyllaries are about as long as the heads, instead of half as long.
The leaf venation is different from that of E. pazcuarense and
apparently from that of E. vernale.

FlG. 2. Eupatorium monticola. A, habit, X Vi; B, portion of inflorescence, X 2;
C, flower, X 9, with much enlarged apex of a pappus bristle; D, corolla with styles,
x 9; E, achene, X 9; F, corolla dissected, X 9; G, anther much enlarged. Drawn
from the type.

93

94 FIELDIANA: BOTANY, VOLUME 36

Eupatorium nubigenum Benth. PL Hartw. 85. 1841. E.
hospitale Rob. Proc. Am. Acad. 43: 32. 1907. E. microdon Rob. I.e.
54: 252. 1918. Critonia hospitalis King & Robinson, Phytologia 22:
49. 1971. Critoniadelphus microdon King & Robinson, I.e. 53. C.
nubigenus King & Robinson, I.e. 53.

I find no differences between the three species which are given
above other than characteristics that I consider to be of minor
importance. Eupatorium nubigenum (type from Guatemala) has
pellucid veins in the leaves; E. microdon (type from Guatemala)
has pellucid dots in the leaves; E. hospitale (type from Mexico) has
both pellucid dots and lines or veins. I must admit that these
conditions exist, but there are specimens that I cannot distinguish
otherwise in which pellucid dots or lines seem to be lacking.
Specimens with immature leaves badly dried often are opaque.
Eupatorium hospitale has the achenes hispidulous, while the other
two do not. However, both minutely hispidulous and glabrous
achenes can be found in the inflorescence of our specimen of
Matuda 5262 (opaque leaves) from Chiapas. Contemporary synan-
therologists have placed species which I consider synonymous into
different genera, as indicated by the synonomy above.

Eupatorium nubivagum L. Wms. sp. nov.

Herbae subalpinae usque ad 0.5 m., erectae ascendentes caulibus pubescentibus;
folia ovata vel lanceolato-ovata acuta serrata leviter pubescentia triplinervia;
inflorescentiae paniculae corymbosae; capitulis campanulatis plusminusve 40-floris,
involucra biseriata, phyllarus valde aequalibus lineari-oblongis vel lineari-oblanceo-
latis acutis erosis trinervis; achaenia nigra 5-porcata leviter strigillosa, pappus

barbellatus albus; corollae anguste campanulatae apicibus leviter pilosis, lobis

t riangulari-ovatis acutis.

Subalpine herbs to 0.5 m. tall, erect or ascending, the stems densely crisped
puberulent, leafy, terete; leaves ovate or ovate-lanceolate, acute, serrate, sparsely
crisped puberulent on both surfaces, triplinerved from the acute base, blade to 4 cm.
long and 2.5 cm. broad, reduced upward, the petioles to about 2 cm. long;
inflorescences terminal, corymbose panicles to about 7 cm. long and nearly as broad,
the heads about 5-20 in each corymb; heads campanulate, 7-9 mm. long and nearly as
broad, with about 40 florets or fewer; involucres biseriate lacking small basal
phyllaries, the phyllaries subequal, linear-oblong or linear-oblanceolate, acute, erose
above, ciliolate and the outer ones puberulent, prominently 3-nerved, about 5 mm.
long and 1-1.5 mm. broad, much shorter than the pappus and florets; achenes black,
5-ridged, sparingly strigillose, 2.5-3 mm. long; pappus white, barbellate, about 4 mm.
long; corollas tubular to about the middle, narrowly campanulate above, sparsely
pilose at the apex, 3.5-4 mm. long; lobes triangular-ovate, acute, about 0.4 mm. long;
style exserted about 1 mm. at anthesis, terete and slightly clavate.

FIG. 3. Eupatorium nubivagum L. Wms. A, habit, X V4; B, inflorescence, X 1; C,
capitulum, partially dissected, x 3; D, flower, x 12, with greatly enlarged detail; E,
corolla dissected, x 12; F, anther, much enlarged.

95

96 FIELDIANA: BOTANY, VOLUME 36

Guatemala: herb to 0.5 m., open wet meadows, Sierra
Cuchumatanes between Paquix and Llanos de San Miguel, road to
San Juan Ixcoy, Dept. Huehuetenango, alt. 3,300 m., November 17,
1967, Molina 21238 (type, F; EAP).

A species of the subalpine meadows closely related to
Eupatorium muelleri Sch.-Bip., which is a species of middle
elevations, 1,000 m. in south Mexico to about 2,000 m. in
Guatemala. The larger heads with phyllaries two-thirds the length
of the heads, narrowly campanulate corollas, and triplinerved leaves
all serve to distinguish this species from E. muelleri. This species
has smaller heads and phyllaries nearly as long as the heads,
corollas expanded upward but not campanulate, and the leaves
penninerved.

Eupatorium oreophilum L. Wms. sp. nov.

Lianae vel frutices epiphyticae aut terrestres, glabrae vel obscure pubescentes.
Folia lanceolata vel lanceolato-ovata, acuminata, petiolis graciles; inflorescentiae
paniculae multicapitatae, capitulis 2-4-floribus; phyllaria 3-4-seriata, extus ovata
acuta atrofusca, intus lineari-lanceolata acuta apice interdum lacerata; achaenia
prismatica 5-7- porcata glabra; corollae purpureae campanulatae, faucibus hirsutulis.

Epiphytic vines or terrestrial shrubs up to 3-4 m. long or tall, the stems and
inflorescences at first obscurely sordid puberulent, young stems somewhat ancipitous,
becoming terete; leaves lanceolate to lanceolate-ovate, acuminate, penninerved with
5-7 pairs of obscure lateral nerves, the mature blades 5-13 cm. long and 3-7 cm. broad,
somewhat fleshy, glabrous, petioles slender and mostly 2-5 cm. long; inflorescences
terminal, densely-headed compound panicles to about 25 cm. long and as broad;
heads up to 3 times as long as broad, mostly about 12 mm. long, flowers mostly 2-4 in
each head; phyllaries glabrous, 3-4-seriate, the outer series ovate, acute, about 1.5-2
mm. long, the inner series progressively longer and narrower, the innermost linear-
lanceolate, acute and the apex often lacerate and to about 5-6 mm. long; achenes
prismatic, 5-7-ridged, glabrous; pappus 30-40 bristles and these about 4 mm. long, not
barbellate; corolla 4-5 mm. long, campanulate, the tube 2.5-3 mm. long, the linear-
lanceolate acute lobes about 1.5 mm. long, the throat with a dense ring of hairs;
anthers at anthesis exserted, about 2 mm. long, the filaments free and very slender,
attached at the throat of the corolla; styles filiform, at anthesis long exserted.

Costa Rica: Cerro de la Muerte in subparamo, provincia de
Cartage, alt. 3,200 m., June 20, 1966, Anderson & Mori 113 (F);
epiphyte with rather short, woody, sappy stems, foliage coriaceous,
Palmira, Prov. Alajuela, alt. 2,300 m., June 21, 1938, Austin Smith
NY794 (F, NY); epiphyte on large pasture tree, 3 m. long, soft
wooded stem, foliage slightly fleshy, buds pale purple, Palmira,
Prov. Alajuela, alt. 2,300 m., July 16, 1938, Austin Smith H948 (F);
epiphytic vine in cloud forest below "El Jardin," Cerro de la

WILLIAMS: TROPICAL AMERICAN PLANTS, XVII 97

Muerte, Cordillera de Talamanca, province of Cartago, alt. 2,700 m.,
May 20, 1956, Williams 20198 (type, F; EAP).

This species belongs in a group of eupatoria that are quite
occasionally epiphytic. Some of them have been treated as
Neomirandea (Phytologia 19: 305-310. 1970). Eupatorium oreophi-
lum is distinguished from the several species related to E.
araliaefolium by the very narrow heads with 3-4-seriate involucres.
It has, so far as I know, the fewest flowers per head of any of the
allied species. The achenes usually have more than five ridges or
costae and this is somewhat unusual.

The number of heads in the enormous inflorescences must
often be more than a thousand.

The locality where I collected the type specimen nearly 20
years ago, then a virgin cloud forest, has been almost completely
devastated and is now a pasture for cattle.

Eupatorium orogenes L. Wms. sp. nov.

Frutices vel lianae epiphyticae. Folia carnosa penninervia elliptica vel elliptico-
oblonga acuta vel acuminata glabra, petiolis brevibus; inflorescentiae terminales
thyrsiformes multicapitatae; capitula anguste campanulata quinquefloribus; in-
volucra obscure 2-3-seriata; phyllaria lanceolata usque ad angusti-oblonga subglabra,
capitulls semiaequilongis; achaenia glabra 5-porcata; corollae subcylindricae sursum
expansae.

Epiphytic shrubs or vines, the young stems and inflorescences sordid hispidulous,
glabrescent with age. Leaves fleshy, penninerved and the 3-5 pairs of lateral nerves
inconspicuous on both sides, blade elliptic to elliptic-oblong, acute or short
acuminate, glabrous or nearly so, the blades 6-11 cm. long and 1.5-4.5 cm. broad,
acute and tapering to the base; petioles up to 1 cm. long, rather thick; infloresences
terminal, large and flat-topped, many-headed thyrses, to about 15 cm. high and as
broad, hirsutulous, pedicels to twice as long as the heads; heads narrowly
campanulate, with about 5 florets, mostly 7-8 mm. long; involucre with the phyllaries
in 2-3 series, about half as long as the head; phyllaries 7-8, lanceolate to narrowly
oblong, essentially glabrous, the outer ones about 2 mm. long and acute, the inner
ones about 4 mm. long, the acute apex somewhat fimbriolate; achenes glabrous, 5-
angled, about 2.5 mm. long; pappus as long as the corollas; corolla deep purple to
violet, subcylindric, expanded upward, glabrous inside and outside, about 5-6 mm.
long, the lobes narrowly triangular, acute, about 0.7-1 mm. long.

Costa Rica: epiphytic shrub, inflorescence pale violet, disturbed
primary forest high over Rio Grande de Orosi, 8 km. south of
Tapanti, Prov. Cartago, alt. 1,500 m., May 7, 1967, Lent 933 (type,
F); flowers beet-purple, epiphtic shrublet on remanant trees in
pasture, San Rafael, 2 km. east of Vara Blanca, Prov. Heredia, alt.
1,800 m., April 26, 1969, Lent 1634.

98 FIELDIANA: BOTANY, VOLUME 36

The attractive purple-flowered species of the Eupatorium
araliaefolium group, mostly epiphytic shrubs, are proving to be
widely scattered in Central America. The present species is related
to E. oreophilum L. Wms., described above, from which it is
distinguished by the short-petiolate leaves, the less prominently
seriate involucres, and the glabrous inner face of the corolla. It is
related also to E. araliaefolium Less., but less closely so again the
short petioles are obvious, the number of flowers in head are fewer,
and the phyllaries fewer and differently arranged. The species most
closely related would seem to be the Mexican-Guatemalan E.
molinae mentioned above, a species vegetatively very similar but
differing in floral detail. The heads of E. molinae have about twice
as many florets, the phyllaries and florets larger and somewhat
different. The two species occupy quite different ranges.

Eupatorium pazcuarense HBK. Nov. Gen. Sp. 4: 123. 1820. E.
subpenninervium Sch.-Bip. ex Klatt, Leopoldina 20: 89. 1884. E.
skutchii Rob. Contr. Gray Herb. 104: 27. 1934. Kyrstenia pazcua-
rensis Greene, Leafl. Bot. Ob. & Grit. 1: 9. 1903. Ageratina
pazcuarensis King & Robinson, Phytologia 19: 215. 1970. A.
skutchii King & Robinson, I.e. 217.

Eupatorium pazcuarense was first collected on the shores of
Lake Patzcuaro in Michoacan, Mexico. A phototype (F 37369) from
the specimen in Paris shows the plant that is occasional in Mexico
and very abundant in the highlands of Guatemala. The heads are
mostly 7-11 mm. long, the phyllaries only slightly shorter than the
pappus and in a single series or in two subequal series, very often
with one or two very short phyllaries, almost subtending bractlets,
at the base of the head; the leaves ovate or ovate-lanceolate, acute
or acuminate at the apex and obtuse or even subcordate at the base,
coarsely dentate and variously and sparsely pubescent, triplinerved
from above the base of the blade. The plant has white florets and is
said to be herbaceous. Much Mexican material identified with this
name appears to belong elsewhere.

Eupatorium skutchii Rob. is not distinguishable from E.
pazcuarense, but was considered to be a coarse shrub and has
triplinerved leaves originating at the base of the blade. We have
made numerous collections in the western highlands which we have
distributed under this name. It has been considered to be a herb as
often as a shrub. Whether the leaves are tri- or 5-plinerved from the
base of the leaf blade or from above the base seems to be variable
and all intergradations are to be found.

WILLIAMS: TROPICAL AMERICAN PLANTS, XVII 99

Eupatorium subpenninervium Sch.-Bip. ex Klatt. Described
from south Mexico and based on Liebmann's collection number 90,
the type in Paris (phototype F 37427) and an isotype in Field
Museum seem to indicate that this differs not at all from E.
pazcuarense. Considerable south Mexican and Guatemalan materi-
al has been referred to E. subpenninervium, but it differs in several
respects from E. pazcuarense. It is described above as Eupatorium
monticola.

Eupatorium perpetiolatum (King & Robinson) L. Wms.
comb. nov. Pseudokyrsteniopsis perpetiolata King & Robinson,
Phytologia 27: 241, fig. 1973.

King and Robinson have pointed out that this species was
distributed by us as Eupatorium pycnocephaloides Rob., which it
obviously is not, and have described it as a new genus and species as
indicated above. The species is quite a distinctive one due to the
thickened petiole bases which, however, are not unique in this
species of Eupatorium, and to the constricted throat of the corolla.
It fits into the genus Eupatorium without undue violence. The
plant is Guatemalan from the department of Quich6, Williams,
Molina & Williams 22457.

Eupatorium pinabetense Rob. Proc. Am. Acad. 36: 482. 1901.
Neobartlettia pinabetensis King & Robinson, Phytologia 21: 296.
1971. Bartlettina pinabetensis King & Robinson, Phytologia 22: 161.
1971.

This species is closely related to Eupatorium tuerckheimii Klatt
(1884) and is distinguished from it only by the following minor
characters: the branches of the inflorescence are pubescent and the
serrations of the leaf margins are more abundant and, while not so
prominently digitaliform as in E. tuerckheimii, they are thickened.
There are about 10 flowers in the heads of this species, while in E.
tuerckheimii they are much more numerous.

Eupatorium pinetorum Williams & Molina sp. nov.

Arbusculae vel frutices usque ad 4 m. altar, rainuli teretes porcati glabri. Folia
lanceolata acuminata obscure serrulata ad bases acuta vel leviter attenuate glabra
penninervia vel subplinervia, petioli graciles; inflorescentiae corymboso-paniculatae
multicapitatae; capitula breviter pedicellate aut sessile 5-floribus; phyllaria chartacea
2-3 nervata triseriata, externa oblongo-ovate obtusa, interne elliptica obtusa vel
leviter acuta; achaenia nigra hirtella 5-porcata; corollae tubulares, lobis lineari-
oblongis.

100 FIELDIANA: BOTANY, VOLUME 36

Weak trees or perhaps shrubs to 4 m. tall, the stems terete but prominently
ridged, glabrous or obscurely puberulent above and in the inflorescence, the nodes
short, up to 3 cm. long. Leaves lanceolate, acuminate, obscurely serrulate, acute and
somewhat attenuate to the base, glabrous except apparent domatia beneath in the
axils of the principal nerves, penninerved or subplinerved with mostly 3 pairs of
lateral nerves, the principal pair arising well above the base of the blade, blades
mostly 9-11 cm. long and 2.5-4 cm. broad, petioles slender and mostly 1.5 cm. long;
inflorescences corymbose panicles to 10 cm. long and as broad, multicapitate, the
pedicels and peduncles spreading puberulent; heads about 8 mm. long, short
pedicellate or sessile, with 5 florets; phyllaries about 10, chartaceous, with 2 (or 3)
indurated nerves, triseriate, the outermost oblong-ovate, obtuse, about 1-1.5 mm.
long, the innermost narrowly elliptic, obtuse or somewhat acute, 5-6 mm. long;
achenes black at maturity, hirtellous, with 5 lighter colored ridges, 2.5-3 mm. long;
pappus of about 30 barbellate bristles 4 mm. long; corolla (probably white) tubular
and slightly inflated at the base, about 4.5 mm. long and 0.5 mm. in diameter near
the base, the lobes linear-oblong, about 0.5 mm. long; styles slender, exserted about 1
mm. at anthesis.

Honduras: tree 4 m. tall, rocky hillside near El Espino (near
Honduran-Nicaraguan frontier), Dept. Choluteca, alt. 1,000 m.,
December 30, 1963, Williams, Molina & Williams 26318 (type, F;
EAP).

This highland pine forest species is related to the lowland
Eupatorium campechense Rob., which we know only from Mexico
(Campeche), the Peten in Guatemala, and British Honduras. This
species is more puberulent in the inflorescence, has the leaves
penninerved or if subplinerved with three prominent pairs of lateral
nerves originating well above the base (the upper pair of nerves
toward the apex). It is the only Eupatorium which we recall that
has domatia beneath in the axils of the nerves.

The leaves on the type specimen appear somewhat translucent,
perhaps due to having been preserved originally by a formaline
method.

Eupatorium prunellaefolium HBK. Nov. Gen. & Sp. 4: 123.
1820. E. salinum Standl. & Steyerm. Field Mus. Bot. 23: 188. 1944.
Ageratina prunellaefolia King & Robinson, Phytologia 19: 215.
1970.

It is with some hesitation that I have reduced E. salinum in the
manuscript for the "Flora of Guatemala." The type is from a
relatively low elevation, 650 m., while other specimens from Mexico
and Guatemala come from some 3,000 m. Such great differences in
elevation of habitats are not unusual in Eupatorium but should be
checked out. However, I find that the type of E. salinum fits very
well into the variations of E. prunellaefolium.

WILLIAMS: TROPICAL AMERICAN PLANTS, XVII 101

Eupatorium psoraleum Rob. Proc. Boston Soc. Nat. Hist. 31:
253. 1904. Neomirandea psoralea King & Robinson, Phytologia 19:
308. 1970.

Costa Rica: epiphytic [in] tree 15 m. up in host, flowers white,
primary forest above Rio Grande de Orosi, 8 km. south of Tapanti,
Prov. Cartago, alt. 1,550 m., August 27, 1967, Lent 1234; Finca
Hnos. Cruz, Coliblanco [=Cariblanco?], Prov. Cartago, alt. 2,600m.,
July 26, 1951, Reark 264 (EAP; F).

These specimens determined from the characters are the first
ones that we have distinguished. Both were originally determined as
E. araliaefolium Less. Dr. Robinson's description fits very well and
is incisive in one or two characters. He describes the branches as "6-
angularibus striatis squamulis minutis transversis tectis." The
transverse squamulae are in no other species that I know and the
resemblence is with E. daleoides as Dr. Robinson remarked.

The type of this species is presumed to be in the National
Herbarium in San Jose, Costa Rica but this has not been confirmed.

Eupatorium quercetorum L. Wms. sp. nov.

Arbusculae usque ad 2.5 m. vel ultrae, ramulis gracilibus puberulentis vel glabris.
Folia rlliptira vel elliptico-oblanceolata acuminata serrata penninervia petiolis
gracilibus; inflorescentiae terminates cymoso-paniculatae capitulis 20-30 ornatis;
capitula cylindrica 15-30-flora; phyllaria 5-6-seriata exteriores ovata interiores
linearia obtusa; achaenia glabra 5-porcata; corollae subcampanulatae lobis ovato-
triangularibus; styli filiformes exserti.

Shrubs to 2.5 m. or more, the branches slender, terete, pubescent becoming
glabrous. Leaves elliptic to elliptic-oblanceolate, acuminate, serrate, acute to the
petiolate base, penninerved with 4-5 pairs of nerves, glabrous above, puberulent along
the nerves below, the blade 4-9 cm. long and 1.5-2.5 cm. broad, the petiole slender,
puberulent, 4-10 mm. long; inflorescences terminal, cymose-paniculate, arachnoid-
puberulent, with 20-30 heads, to 5 cm. long and as broad; heads cylindric, 8-10 mm.
long and 2-3 mm. in diameter, with 15-20 flowers in each head; phyllaries 5-6-seriate,
puberulent, the outermost suborbicular-ovate, obtuse, about 2 mm. long, the inner
series becoming longer and narrower, the innermost linear, obtuse, about 7 mm. long,
reaching almost to the top of the pappus; achenes glabrous, 5-ridged; pappus about 4
mm. long, of about 30 nearly smooth bristles; corolla white, subcampanulate at the
throat, about 4.5 mm. long and 1.2 mm. broad at the throat, the lobes ovate-
triangular, acute; anthers appendaged at the apex, linear-oblong, about 1.5 mm. long;
style filiform, surpassing the corolla about 4 mm. at anthesis.

Mexico: flowers white, plant about 8 ft. tall, slope with
Quercus, 5 km. above Soyalo along the road to Bochil, municipio of
Soyalo, Chiapas, alt. 4,100 ft., November 2, 1965, Breedlove 14046
(type, F; MICH); flowers purple, shrub 6 ft. on slopes along Tana

102 FIELDIANA: BOTANY, VOLUME 36

Te River near Sahal K'esh, paraje of Mahben Chauk, municipio of
Tenejapa, alt. 2,900 ft., November 27, 1964, Breedlove 7675 (F,
MICH).

Closely related to Eupatorium glaberrimum DC., a common
species in the oak-pine forests from Mexico to Costa Rica. It is
easily distinguished by the petiolate leaves attenuate to an acute
base; the heads are smaller and with half as many flowers. The
specimens were distributed as Eupatorium oerstedianum Benth.,
which I consider to be a synonym of E. glaberrimum DC.

The species is to be expected in Guatemala since one of the
specimens cited was collected not far from the border. It will be
included in the "Flora of Guatemala."

Eupatorium scoparioides L. Wms. sp. nov.

Arbusculae parvae scopariaeformes usque ad 1 m. altae, ramuli graciles striati
dense puberulentes; folia triangulari-ovata acuta vel acuminata, base truncata vel
subcordata glabra vel subtus leviter pubescentes et glanduloso-punctata;
inflorescentiae spicatae vel spicato-paniculatae; capituli parvi plusminusve 5-flori;
involucri bi-triseriati; phyllaria extus glandulosa exteriores linear!- lanceo la ta vel
ovata acuta interiores lineari-oblonga truncata vel leviter acuta; achaenia strigillosa
negra perparva; corolla alba cylindrica vel apice leviter subcampanulata 1.5-2 mm.
longa, lobis minutis; styli breviter exserti.

Small profusely branched shrubs to about 1 m. tall, the multistriate branches
slender, terete, densely spreading puberulent, the hairs sometimes minutely dendroid
toward the apices; the petiolate leaves triangular-ovate, acute or acuminate, truncate
to subcordate at the base, trinerved from the base of the blade, glabrous above or
nearly so, puberulent on the nerves below and densely punctate-glandular over the
entire surface, blades 1.5-3 cm. long and 12.3 cm. broad at the base, the petioles
slender, puberulent, mostly 5-6 mm. long; inflorescences spreading puberulent,
terminal and axillary, spicate or spicate-paniculate, bracteate, to about 8 cm. long,
the heads in small cymules, pedicellate; heads 3-4 mm. long, with about 5 florets;
involucre bi- or triseriate, with only 10-12 prominently glandular-dotted phyllaries,
outer phyllaries linear-lanceolate to ovate, acute, mostly less than 1 mm. long, the
inner phyllaries linear-oblong, truncate or rarely acute and somewhat expanded at
the apices, mostly about 2 mm. long; achenes 5-ridged, strigillose, black at maturity,
about 1-1.5 mm. long; pappus of 25-30 bristles, about 1.5 mm. long; corolla white,
cylindric but subcampanulate at the throat, about 1.5-2 mm. long, the lobes minute,
linear-lanceolate, about 0.2 mm. long; styles exserted about 1 mm. at anthesis.

Guatemala: flores blancas, planta hasta 1 m., comun, sabanas y
pinares abiertos de Poptun, Depto. Pete'n, alt. 500 m., November 11,
1965, Molina 15595 (type, F; EAP).

This is one of the most distinctive of the species of Eupatorium
in Central America. There are no closely allied species known to
me. The small heads with few bi- or triseriate phyllaries, the form of

WILLIAMS: TROPICAL AMERICAN PLANTS, XVII 103

the inflorescence, the very small flowers, the glandular-punctate
leaves and phyllaries, the sometimes dendroid hairs on the stems,
are a combination of characters that set the species apart.

Eupatorium semialatum Benth. PI. Hartw. 76. 1840.

This species was described from near Zunil, Department of
Quezaltenango, Guatemala. It has been known under the name
Eupatorium ligustrinum DC., which is certainly a related species,
and a mass of material from south Mexico and Guatemala has
accumulated. An excellent phototype of Eupatorium ligustrinum
from the Delessert Herbarium in Geneva indicates that this species
based on Berlandier 2143 from Tamaulipas is quite different from
ours. Bentham's epithet will be used in the "Flora of Guatemala."

Eupatorium sodali L. Wms. nom. nov. Piqueria standleyi
Rob. Contr. Gray Herb. 104: 4. 1934, not Eupatorium standleyi Rob.
Koanophyllon standleyi King & Robinson, Phytologia 22: 151. 1971.
1971.

Since the specific name "standleyi" may not be used again in
Eupatorium, the name "sodali" is applied and is intended to refer to
Standley, my associate and companion in the tropics for many
years.

The present species illustrates the artificiality of the class-
ification among some Compositae and more especially of the tribe
Eupatorieae. This species originally was placed with quite unlike
plants in the genus Piqueria by one of the most competent students
of American Compositae. The deciding factor in placing it there
undoubtedly was the lack of a pappus, yet pappus-bearing
specimens are now known and the species fits quite well into
Eupatorium, although it does disrupt in a minor way the pappus
character ascribed to Eupatorium.

There were two species in Guatemala that had been called
Piqueria, P. trinervis Cav., the type species of the genus Piqueria,
and P. standleyi Rob. Piqueria standleyi superficially is similar to
certain Andean South American species but Dr. Robinson, who had
revised the genus (Proc. Am. Acad. 42: 4-16. 1906), when he
described P. standleyi said that he had failed to relate it to any of
the South American piquerias.

It is well known that epappose plants appear in species of
Compositae that are normally pappose. This often happens in the
tribe Eupatorieae, especially in Ageratum.

104 FIELDIANA: BOTANY, VOLUME 36

I had just finished with the revision of Eupatorium for the
"Flora of Guatemala" when I took up Piqueria. I suspected that
Piqueria standleyi might be an epappose Eupatorium and began a
search among those species that seemed related, hoping to find one
that would be the same as Piqueria standleyi but pappus-bearing. I
did not find a species that satisfied this requirement but did find
two collections from El Salvador that are identical to the type of
Piqueria standleyi, except that they bear a pappus. Standley &
Padilla 3416 is rather past prime condition but a few achenes
remain. This specimen was determined as P. standleyi by Standley
who perhaps did not notice the achenes. The other specimen,
Calderon 2465, is .at an thesis and there can be no question that it
represents P. standleyi. This specimen was determined as Eu-
patorium solidaginoides HBK., probably by Standley. During the
revision of Eupatorium for the "Flora" I questioned the determinat-
ion, but passed it by since the specimen was from El Salvador.

Since there seems to be no question that Piqueria standleyi is a
Eupatorium but usually without pappus, I provide the name above
and transfer the species to Eupatorium.

Eupatorium solidaginoides HBK. Nov. Gen. & Sp. 4: 126.
1920. Koanophyllon solidaginoides King & Robinson, Phytologia
22: 151. 1971.

The species is a difficult one for it must be placed into two
different sections of the key to Eupatorium, based upon the
interpretation of the arrangement of the phyllaries in the
capitulum. In some specimens the phyllaries seem to be essentially
equal with 2-3 short bract-like phyllaries at the base. In other
specimens the phyllaries seem to be triseriate. The species obviously
is related to E. pycnocephalioides and to E. microstemon, as well as
to such species as E. mimicum and its allies. The species, as I
understand it, is wide ranging, both in latitude and in altitude. It is
found from Mexico to northern South America and the altitudinal
range in Central America is from sea level to 1,800 m.

Eupatorium sorensenii (King & Robinson) L. Wms. comb,
nov. Koanophyllon sorensenii King & Robinson, Phytologia 23:
395. 1972.

The species is closely related to Eupatorium montigenum
Standl. & Steyerm. and E. hypomalacum Rob., both Guatemalan
species.

FlG. 4. Eupatorium sodali.(Piqueria standleyi). A, habit, X 2Vz; B, segment of an
inflorescence, x 2'-2; C, head of immature flowers, x 10; D, head of epappose flowers
at anthesis, x 10; E, epappose floret, X 17; F, epappose floret dissected, X 17; G,
pappose floret, x 17; H, Paul Carpenter Standley (1884-1963), for whom this plant
was originally named, and renamed, when about 50 years old.

106

106 FIELDIANA: BOTANY, VOLUME 36

Eupatorium tomentellum Schrad. Ind. Sem. Hort. Goett.
1833: 3, t. 3. 1833. Ageratina tomentella King & Robinson,
Phytologia 19: 227. 1970.

This species is known in a rather limited area in Guatemala
along the base of the Cuchumatanes Mountains from Aguacatan
(Huehuetenango) to Sacapulas (Quiche). It is often found in oak-
pine forest, but it is more common on very dry, hot slopes. Several
collections have been made in recent years. Eupatorium ovillum
Standl. & Steyerm. is a closely related species known only from the
highlands of western Guatemala.

Eupatorium tuerckheimii Klatt, Leopoldina 20: 95. 1884.
Neobartlettia tuerckheimii King & Robinson, Phytologia 21: 297.
1971. Bartlettina tuerckheimii King & Robinson, Phytologia 22: 162.
1971.

Eupatorium tuerckheimii is closely related to E. pinabetense
Rob., mentioned above. The isotype (Tuerckheim 77) shows curious
long silky pubescence on the receptacle. I have been unable to find
this pubescence on other specimens of this species, nor in E.
pinabetense.

Eupatorium tunii L. Wms. sp. nov.

Lianae glabrae usque ad 2.5 cm. diametrales. Folia lanceolato-ovata vel ovata
acuta aut breviter acuminata triplinervia, petiolis graciles; inflorescentiae axillares
aut terminales vulgo cymulae compactae multicapitatae; capituli sessiles 4-6-floribus;
involucrum 4-6-seriatum, phyllariis valde inaequalibus ovatis usque ad anguste
ellipticis; achaenia obscure hirtella, 5-porcata; corolla cylindrica, lobis lanceolatis
acutis.

Glabrous vines to 2.5 cm. in diameter, the branches opposite. Leaves lanceolate-
ovate to ovate, acute or very shortly acuminate, glabrous, triplinerved with the
nerves beginning 4-5 mm. above the base of the blade, the blade 4.5-9 cm. long and 2-
5 cm. broad, petioles slender, mostly about 1 cm. long; inflorescence axillary or
terminal, usually very compact cymules 2-3 cm. in diameter and each containing up
to about 50 heads; heads sessile or nearly so, somewhat more lax in fruit, bearing 4-6
florets, 7-11 mm. long and 2-2.5 mm. in diameter; involucre 4-6-seriate, the phyllaries
very unequal, noticeably puberulent at the apices, lowermost phyllaries ovate and
about 1 mm. long, the innermost lanceolate to narrowly elliptic, acute and 5-6 mm.
long; achenes obscurely hirtellous on the angles, becoming glabrous, brownish-black,
5-ridged, about 2.5-3 mm. long; pappus 5-6 mm. long, equalling the corollas, very
obscurely barbellate or smooth; corolla cylindric and slightly expanded upward,
about 6 mm. long, the lobes lanceolate, acute, about 0.5 mm. long.

Guatemala: flores amarillentas, bejuco en foresta baja, en
orillando aguada del aereopuerto, Parque Nacional, Tikal, Peten, 8
sep. 1970, Rolando Tun Ortiz 1306 (type F; EAP, MICH).

WILLIAMS: TROPICAL AMERICAN PLANTS, XVII 107

British Honduras: florets greenish, vine 20 ft.. 1 in. in diameter,
odor of chrysanthemum, Mullins River Road, alt. 50 ft., April 6,
1929, Schipp 137 (F).

Mexico: Fortuno, Coatzacoalcos River, Vera Cruz, alt. 30-50 m.,
March, 1937, Llewelyn Williams 8431 (F).

This species belongs in what now appears to be a small complex
of vines found in the lowlands of Guatemala, British Honduras and
apparently on the coast of Vera Cruz. Included is Eupatorium
bartlettii Rob., E. magistri L. Wms., described above and this
species. Eupatorium tunii is easily distinguished from the other two
species by the very dense inflorescences, instead of relatively lax
ones, and by the detail of the shape and venation of the leaves. All
three species are native of Guatemala and British Honduras.
Eupatorium bartlettii and E. tunii have capitula containing 4-6
flowers and have triplinerved leaves while E. magistri has capitula
containing about 10 flowers and 5-plinerved leaves.

The species is named for Orlando Tun Ortiz, who has made
about 2,000 collections in the department of Peten thereby
increasing our knowledge of that remote part of Guatemala.

Eupatorium vitalbae DC. Prodr. 5: 163. 1836. Heterocondylus
vitalbae King & Robinson, Phytologia 24: 392. 1972.

Originally described from the vicinity of Rio de Janeiro the
species now is known over much of the low tropics of South
America. It is in Panama and common in Costa Rica. It is known in
Nicaragua and in Honduras a few miles from the Guatemalan
frontier. Since it should be found in Guatemala, it will be included
in the flora. We have two collections by Haenke, one marked Peru,
the other Mexico. No other record is available to me from Mexico.

Eupatorium viscidipes Rob. Proc. Am. Acad. 36: 484. 1901.
Fleishmannia viscidipes King & Robinson, Phytologia 19: 206. 1970.

Two recent collections of this species, Molina et al. 30291 and
Williams et al. 43020 (both taken in January, 1974 and of which
there is abundant material for examination), show achenes with
normal pappus, with reduced pappus and without pappus all in the
same head. No heads were found in which more than one or two
achenes were calvous.

MACVAUGHIELLA

Macvaughiella standleyi (Steyerm.) King & Robinson, Sida

108 FIELDIANA: BOTANY, VOLUME 36

3: 282. 1968. Schaetzellia standleyi Steyerm. Field Mus. Bot. 23:
107. 1944.

There are two species in this genus. The Central American one
is dubiously distinct from M. mexicana (Sch.-Bip.) King &
Robinson, but we have not seen adequate material from Mexico to
judge.

This species is a common one in eastern Guatemala and in
Honduras at middle elevations, usually in open oak-pine forest
areas. Plants of M. standleyi occur on the volcanic cliffs just above
the sea in El Salvador. The plants from this locality are more
luxuriant than those from middle elevations in Guatemala and
Honduras, but I have found no other way to distinguish them.

MIKANIA

Mikania aromatica Oersted, Overs. Dansk. Vidensk. Selsk.
Forh. 10. 1863. Willughbaea globosa Coulter, Bot. Gaz. 20: 46. 1896.
Mikania globosa Coulter, 1. c. in synon.

A photograph of the original specimen in Copenhagen shows
that this is the rather common Mexican (Vera Cruz, Oaxaca,
Chiapas) and Central American species, extending from Guatemala
to Costa Rica. The heads in the inflorescences are contracted into
globose glomerules; the leaves are coriaceous and both the
secondary and tertiary nervation is prominulous. Kew Index is
incorrect in giving the species as Brazilian. The Costa Rican M.
tonduzii Rob. perhaps is to be referred to this species.

Mikania houstoniana (L.) Rob. var. guatemalensis (Standl.
& Steyerm.) L. Wms. comb. nov. Mikania guatemalensis Standl. &
Steyerm. Field Mus. Bot. 23: 105. 1944.

A minor variation from the coastal region of the Gulf of
Honduras, in Guatemala, British Honduras, and Honduras. It is
distinguished from the typical variety by having the inflorescence
glabrous, not puberulent; the achenes puberulent to nearly
glabrous, and gland-dotted or atomiferous, not glabrous and
glandless; the leaves mostly acute at the base or at most obtuse,
not truncate or cordate.

There is a specimen from Costa Rica very similar to those of
northern Central America, Williams et al. 28577, but it comes from
the cloud forest region at 3,200 m. while the others are known from
600 m. or less.

FlG. 5. Macvaughiella standleyi. A, branch of the plant, X V4; B, detail of a
portion of an inflorescence, X 3; C, flowers preanthesis and at anthesls, X 10, and
with enlarged detail; D, flower dissected, x 20; E, phyllaries, x 10; F, Rogers
McVaugh, American botanist (1909- ).

Mikania stipulifera L. Wins. sp. nov.

Lianae suffruticosae; folia late ovato-cordata longe acuminata sparse strigosa
petiolis gracilibus ornatis; stipulis foliaceis reniformibus; inflorescentiae pedunculatae
axilares paniculato-cymosae; capitulae cylindricae 4-floribus; phyllaria imbricata
anguste lanceolata acuta vel breviter acuminata glabrecentes; achaenia prismatica
glabra; pappus obscure barbellatus; corollae tubus cylindricus, limbus campanula tus
4-lobatus.

109

110 FIELDIANA BOTANY, VOLUME 36

Lianas of unknown length. Stems terete, sparsely puberulent or glabrous,
internodes 10-12 cm. long; leaves broadly ovate-cordate, long acuminate, the blade
about 4 cm. long and 3 cm. broad, sparsely strigillous, becoming glabrous below;
petioles slender, about 2 cm. long; stipules reniform. foliaceous. to about 0.7 cm. long
and 1.5 cm. broad, strigillous; inflorescence axillary, long pedunculate cymose
panicles to about 10 cm. long; heads narrowly cylindric, pedicellate, subtended often
by a linear bracteole half as long as the head, heads about 10 mm. long; phyllaries
imbricated, narrowly lanceolate, acute and short acuminate, glabrous or nearly so,
about 6-7 mm. long and 1.5 mm. broad; flowers white, 4 in each head; achenes about
3 mm. long, prismatic, atomiferous, glabrous; pappus bristles about 6 mm. long, very
obscurely barbellate; corolla tube cylindric, about 2 mm. long, the limb campanulate,
lobate, to 1.5 mm. long.

Honduras: liana, white flowers, cloud forest on Mt. San
Juancito, Dept. Morazan, alt. 6,000 ft., June 19, 1948, Classman
1684 (type F).

Easily distinguished from all other Central American mikanias
by the persistent foliaceous stipules, the inflorescences at the nodes,
and the linear-lanceolate, acute, phyllaries.

UNIVERSITY OF ILLINOIS URBANA