Contributions from the United States National Herbarium
Volume 56: 1—184

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Paul M. Peterson

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LIBRARIES

A Revision of Festuca (Poaceae: Loliinae)
in South American Paramos

Daniel Stancik
and
Paul M. Peterson

om

Smithsonian Institution
Scholarly Press

WASHINGTON, D.C.
2007

ABSTRACT

Stan¢ik, Daniel and Paul M. Peterson. A revision of Festuca (Poaceae: Loliinae) in South American paramos. Smithsonian Contri-
butions from the United States National Herbarium, Volume 56, 184 pages (including 97 figures). A taxonomic revision of Festuca
L. for the South American paramos of Brazil, Colombia, Ecuador, northern Peru, and Venezuela is given. Fifty-six species and six
subspecies of Festuca are recognized in the study area. Fifty-three species are native to the flora region and three are introduced and
cultivated. Keys for determining the species, a classification, synonymies, descriptions, leaf anatomical descriptions, observations,
distribution and habitat, specimens examined, illustrations, and leaf surface micrographs are provided. One species: F turimiqui-
rensis Stancik & P.M. Peterson, and one subspecies: F. tolucensis subsp. culata Stancik & P.M. Peterson, are newly described.
The following three names were lectotypified: Bromus caldasii Kunth [=Festuca caldasii (Kunth) Kunth], Festuca fratercula Rupr.
ex E. Fourn. (~Festuca amplissima Rupr. subsp. amplissima), and Festuca ovina subvar. jamesonii St.-Yves (=Festuca glumosa
Hack. ex E.B. Alexeev).

KEY worps: Anatomy, Brazil, Classification, Colombia, Ecuador, Festuca, Loliinae, Poaceae, Taxonomy, Venezuela.

RESUMEN

e

Stan¢ik, Daniel and Paul M. Peterson. A revision of Festuca (Poaceae: Loliinae) in South American paramos. Smithsonian Con-
tributions from the United States National Herbarium, xxx paginas (incluyendo 97 figuras). Se presenta una revision taxonomica
del género Festuca (Poaceae) para los paramos del Sudamérica cubriendo la regién de Brazil, Colombia, Ecuador, norte del Pert y
Venezuela. Se reconocen 56 especies y seis subespecies. Cincuenta y tres especies son nativas de la region, tres son introducidas y
cultivadas. En el trabajo se presenta una clave para la determinacion de las especies, una clasificacion del género en la region estu-
diada, sindnimos, descripciones, descripcion de la anatomia foliar, distribucion y ecologia de las especies, lista de los especimenes
estudiados, ilustraciones y microfotografias de la epidermis foliar. Se describe una nueva especie: F. turimiquirensis Stancik &
P.M. Peterson, y una subespecie F. tolucensis subsp. culata Stancik & P.M. Peterson. También se designan lectotipos para tres
nombres: Bromus caldasii Kunth [=Festuca caldasii (Kunth) Kunth], Festuca fratercula Rupr. ex E. Fourn. (=Festuca amplissima
Rupr. subsp. amplissima) y Festuca ovina subvar. jamesonii St.-Yves (=Festuca glumosa Hack. ex E.B. Alexeev).

PALABRAS CLAVES: Anatomia, Brasil, Clasificacion, Colombia, Ecuador, Festuca, Loliinae, Poaceae, Taxonomia, Venezuela.

Published by Smithsonian Institution Scholarly Press

P.O. Box 37012

MRC 957

Washington, D.C. 20013-7012
www.scholarlypress.si.edu

The periodical Contributions from the U.S. National Herbarium was first published in 1890 by the United States Department of
Agriculture. From July 1, 1902, forward it was published as a Bulletin of the United States National Museum. The series was discon-
tinued after volume 38 (1974) and revived with volume 39 (2000) as a venue for publishing taxonomic papers, checklists, floras, and
monographs produced by staff and associates at the U.S. National Herbarium. It is externally peer reviewed and published at irregular
intervals. Subscriptions and other correspondence should be addressed to Smithsonian Institution Scholarly Press. The present issue
is available in print for free while supplies last; PDF files are available at http://www.sil.si.edu/smithsoniancontributions.

Contributions from the United States National Herbarium (ISSN 0097-1618)

Cover design by Alice R. Tangerini; front, habit of Festuca fimbriata Nees; back, habit of Festuca fragilis (Luces) B. Bricefio.
Illustrated by Anna Skoumalova.

POSTMASTER: Send address changes to Smithsonian Institution Scholarly Press, P.O. Box 37012, MRC 957, Washington, D.C.
20013-7012.

Library of Congress Cataloging-in-Publication Data

Stancik, Daniel.
A revision of festuca (Poaceae: Loliinae) in South American paramos / Daniel Stanéik and Paul M. Peterson.
p. cm. — (Contributions from the United States National Herbarium ; 56)

Includes bibliographical references and index.

Summary in Spanish.
1. Fescue—South America—Classification. 2. Fescue—South America—lIdentification. I. Peterson, Paul M. II. Title.
QK495.G74S728 2007
584’.9—dc22 2007026985

The paper used in this publication meets the minimum requirements of the American National Standard for Permanence of
Paper for Printed Library Materials Z39.48—1992.

CONTENTS

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Ree geste nae ENe SUNN 5 ee eee Rh Fen Gk ee a a Fed t cites sdeasuscaasestoectadseabceavarsacevdedssacssse 6
reenter e M e FEM a NEE, PA Deis hecaiivasadeaa leva Tivele ovaguseledtvodavsedsuaocl¥eesets 6

pee rE ER rer SN te TARE See deta see sd ric cee ta de sak iste clas sadusucashacedavesdeudadscisaestadictessensdesacoentabooes yj

mm Ons oa yds TEIN dee ene, SN TE nec a es Aa cecrde cat eestve des bed insdacdnalitusseterenslidsasapacscessddcoevsedevec 8

Sea ERs err et BN NY Fe os Se doe SEA SREI NS Sv dod daub badd acudee seabed Shu ldase desde cencdouseasdee 8

ee me NETS TEIN Sense ee eeN a oi cas saaT sa ee Se Aideastsucdadsssusssdeidessesudesasstdasseascssaveseccecessavees 1]
KEY TO THE SPECIES OF FESTUCA IN BRAZIL, COLOMBIA, ECUADOR, AND VENEZUELA.....ccccccceeseeeeees 11
SURVEY OF THE SPECIES AND SUBGENERIC TAXA OF FESTUCA IN SOUTH AMERICAN PARAMOS .......22.2. 16
DESCRIPTIONS OF SPECIES OF FESTUCA FROM SOUTH AMERICAN PARAMOS .........cseccccccceseescscsueeesceeees 17
STEED TEED Cy. UNS TE Re, est oR By aE PR ob Sie 1 YL ORe By an a 146

SCANNING ELECTRON MICROGRAPHS OF LEAF EPIDERMAL SURFACES ...........0..0000..... 147

meer re ee cae ey aN ee The eee scab tM stand ei ta ona tadabdecseendhaisdeacesvectecvsechssecsesetadedvadevactie 173
empires Pam T NTT 5 9 oa eh hss needa, cuban es sade acne eaMa eas Seda sobecccesastasesdnccobsceuessuclesccsedsces 173
APHABETICAL INDEX OF ACCEPTED SPECIES AND SUBSPECIES>.......000...<.0000ccceccssssessccsneseasetacncaesons Wes
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A Revision of Festuca (Poaceae: Loliinae)
in South American Paramos

Daniel Stancik! and Paul M. Peterson?

INTRODUCTION

Composed of more than 400 species, Festuca
is one of the largest genera in the Poaceae (Clayton
& Renvoize 1986; Watson & Dallwitz 1992). The
genus has cosmopolitan distribution and it is an
important component of grass ecosystems of the
temperate zone as well as the alpine grasslands
of the tropical zone. Although the diversity of the
genus /estuca is centered in the Holarctic zone of
Eurasia and North America, with about 140 species
(Darbyshire et al. 2003), the South American Andes
represent a significant center of Festuca diversity.

HISTORICAL OVERVIEW.—In South America, taxo-
nomic studies of Festuca have been concentrated
in the southern part of the continent, 1.e. Argentina,
Bolivia, Chile, and Peru (Phillipi 1859-60, 1865;
Pilger 1898, 1906; Hitchcock 1927; Saint-Yves
1927; Parodi 1935, 1953; Macbride 1936; Infantes
1952; Foster 1966; Tiirpe 1969; Vareschi 1970;
Correa 1978; Matthei 1982; Tovar 1993; Brako
& Zarucchi 1993; Zuloaga et al. 1994; Renvoize
1998; Soreng et al. 2003). Taxonomic information
on Festuca in the paramo zone of South America
(Venezuela, Colombia, Ecuador, N Peru, SE Bra-
zil) has been quite sporadic, although botanical
collections have increased in the last 50 years.
The oldest specimens of Festuca from Venezuela
were collected during 1906-1911 by A. Jahn,
H. Pittier. The first list of Venezuelan grasses was
published in 1937 (Anonymous 1937) and only
two species of Festuca (F. livida and F: tolucensis)
were listed. In 1942, Luces distinguished between
the Mesoamerican and the Venezuelan popula-
tions of F- livida and described a new Venezuelan
species, F! fragilis (Luces) Bricefio. Two species
(F. fragilis and F- tolucensis) were mentioned in the

first Venezuelan paramo flora by Vareschi (1970).
Later, Alexeev (1986) in his work on Festuca for
Venezuela, Colombia, and Ecuador mentioned only
one species from Venezuela (F. tolucensis). The
recently published study of Bricefo and Murillo
(1994) treats Festuca in the state Mérida (Ven-
ezuela) and lists five native species (F. pinetorum
Swallen, F’ elviae Bricefio, F: tolucensis, F. fragilis,
F. coromotensis Bricefio). Stancik and Peterson
(2003) indicated that F’ pinetorum was misidentified
and therefore does not occur in Venezuela. In addi-
tion, Davidse et al. (1994) mentioned the species
F. amplissima Rupr.; the only known Venezuelan
specimen (Tillett & Honig 746-765, MO) was
collected in Serrania de Perija on the Colombian-
Venezuelan boundary. The earliest collections of
Festuca from Ecuador and Colombia were made by
A. von Humboldt and A.J.A. Bonpland (Expedition
to Ecuador in 1802-1803; see Humboldt et al. 1816
and Jorgensen et al. 1999) and J.C. Mutis and FJ.
Caldas (Real Expedicion Botanica 1783-1808; see
Pinto-Escobar 1976, 1985; and Blanco & del Valle
1991). Seven species are present in Humboldt’s
collection, five of which were later described by
C.S. Kunth as new (F: andicola, F: caldasii, F- dasy-
antha, F- procera, F: quadridentata). The Festuca
specimens collected by T. Haenke (Expedicion
Malaspina 1789-1794) are of even earlier date, but
none of them are from Ecuador or Colombia. The
uncertain origin of the type specimen of Festuca
presliana Hitche. (= Bromus depauperatus J. Pres)
is discussed in Stancik & Peterson (2003).

Hitchcock (1927), in his overview of grasses
of Ecuador, Peru, and Bolivia, mentioned 17 spe-
cies. The Ecuadorian species included F. dasyantha,

' Department of Botany, Charles University of Prague, Benaska 2, 128 01 Prague 2, Czech Republic. dan_stancik@yahoo.com

? Department of Botany MRC-166, National Museum of Natural History, Smithsonian Institution, Washington, DC 20013-7012,

USA. peterson@si.edu

6 Festuca in South American Paramos

F ulochaeta, F. eminens, F: procera, F: sublimes,
F. andicola, F. dichoclada, F. dolichophylla,
F. breviaristata. Festuca eminens 1s also cited for
Colombia. Hitchcock’s publication represents the
first overview of grasses for this region but does not
reflect the entire diversity, since more than 60 species
are known from the region today. Additionally, the
accepted “broad species” concept results in the inclu-
sion of species (such as F. ulochaeta, F: dichoclada,
F- dolichophylla), that do not appear in Ecuador.

The detailed work of Saint-Yves (1927) has
little practical use for Festuca taxonomists in South
America, as the work suffers from insufficient spec-
imens studied and appears to be focused on minute
details rather than estimating species variability and
pointing out differences. The taxa were recognized
in a confused system of varieties and forms. Later,
descriptions of new species were published only
occasionally (Swallen 1948) and as checklists of
local floras and herbaria collections (Pinto-Escobar
1966, 1985; Acosta-Solis 1969; Burger 1981; Cleef
1981). There treatments mechanically accepted
the conclusions of previous authors (principally
of Hitchcock) and confusion surrounding species
delimitations have continued (e.g., the citation of
F. dolichophylla for Colombia).

New insight into the taxonomy of Festuca in
northern South America was brought by Alexeev
(1984, 1986), particularly in terms of lectotypification
of previously described species. and description of
new taxa at sub-generic and specific levels. Alexeev
(1986) cited 19 native species from Ecuador and
Colombia. The work was based on the examination
and revision of a limited number of herbarium speci-
mens without the advantage of field work and it could
not reflect the entire diversity of the genus in South
America. Alexeev’s conclusions were reflected in the
checklists of local floras published later (JOrgensen
& Ulloa Ulloa 1994; Jorgensen & Leon Yanez 1999;
Luteyn 1999; Rangel 2000). Interest in the study of
the Brazilian paramo is recent (Safford 1999a, b).

TAXONOMY AND Diversity.—Fifty-three native
and three introduced species are recognized in the
study area. Among them: five species (F dinirica,
F. guramacalana, F: hatico, F. turimiquirensis and
F. venezuelana) and two subspecies (F. tolucensis
subsp. culata and F: tolucensis subsp. perijae) are
recognized as new for Venezuela; twelve species
(F. boyacensis, F: chita, F. chitagana, F. cocuyana,

F. hatico, F- monguensis, F- nereidaensis, F. pilar-
franceii, F- toca, F- sanctae-martae, F: sumapana,
and F’ woodii) and one subspecies (F. amplissima
subsp. magdalenaensis) as new for Colombia; six
species (F. carchiense, F: holubii, F- imbaburensis,
F. laegaardii, F: oroana, and F: soukupii) and three
subspecies (F’ chimborazensis subsp. micacochensis,
F. parciflora subsp. loxana, and F: vaginalis subsp.
cayambae) as new to Ecuador; and, one species
(F. renvoizii) as new to Peruvian jalca (i.e., humid
alpine grass ecosystem of the northern Peru).

The species found in South America are attrib-
uted to the six subgenera and seven sections of the
genus Festuca. About 80% of South American Fes-
tuca species belong to the large cosmopolitan subg.
Festuca. Remarkably, three subgenera (Festuca
subg. Helleria, subg. Mallopetalon, and subg. Erosi-
florae) and three sections (Festuca sect. Ruprechtia,
sect. Glabricarpae, and sect. Cataphyllophorae) are
endemic or largely endemic to South America.

The distribution of species diversity is not
homogeneous in South America. The Andes
between central Peru and northern Argentina has
the highest species concentration (almost 50% of
all South American taxa). In the northern Andes,
there are two areas with high species diversity,
central Ecuador (Pichincha with 14 species, and
Chimborazo with 13 species) and northern Colom-
bia (Boyaca with 14 species). High levels of local
endemicity were observed in regions with the high-
est species diversity, especially in Boyaca.

Similarity of species composition was also
analyzed to elucidate the relationships among
the South American regions. As expected, strong
relationships exist between the Festuca species of
Ecuador and Colombia, and the Festuca species
from northern South America (“paramo group”)
are clearly distinct from the those of central and
southern South America.

BioceocraPuy.—The known range of several
Festuca species is expanded contributing to a better
understanding of the biogeography of the genus in
South America.

Festuca amplissima was found in northern
Colombia (Sierra Nevada de Santa Marta) and on
the Colombian—Venezuelan boundary (Serrania de
Perija). This species represents the biogeographical

Festuca in South American Paramos 7

connection between Mesoamerican and South Amer-
ican flora. The species was originally known from
Costa Rica, Guatemala, Mexico, and Panama.

Festuca fragilis was found in Nevado del
Cocuy (Dept. Boyaca, Colombia) and was previ-
ously reported only from the state of Mérida in Ven-
ezuela, ca. 250 km from the Colombian locality.

Festuca ulochaeta was found in the Cordil-
lera Oriental of the Colombian Andes as well as
the northern Andes and Cordillera de la Costa of
Venezuela, and in Costa Rica. Formerly, this species
was known only from the region of Brazilian Shield
of southeastern Brazil and northeastern Argentina.
This information might be important in determining
the origins and phylogeny of other South American
species of Festuca sect. Subulatae.

The first collections of the genus Festuca in
Colombian Cordillera Occidental are reported:
F. asplundii (Farallones de Cali and Macizo de
Tamana), F. sodiroana (Paramo Frontino), and
F. andicola (Paramo Frontino).

Festuca caldasii, F- glumosa, and F: sodiro-
ana, which were formerly regarded as endemic to
Ecuador, were confirmed in Colombia, showing
the biogeographical relationship between Colom-
bian Cordillera Central and Ecuadorian Andes.
Similarily, Festuca asplundii, which was originally
considered endemic to Colombia and Ecuador, was
confirmed in the Andes of northern Peru.

Festuca dinirica and F- guaramacalana are
the first records of the genus reported from the
mountains of National Park Dinira and from the
massif of National Park Guaramacal in Venezuela,
respectively, and represent the most easterly Andean
records of Festuca.

Important extra-Andean findings of Festuca in
Venezuela were made in Cordillera de la Costa en
Venezuela. The localities Colonia Tovar and Ser-
rania de Turimiquire represent the most easterly,
mountain forests that include Festuca in northern
South America. These localities are located 250
km and 600 km east of the nearest Andean locality,
respectively.

The following species that were formerly
mentioned in the floras of Venezuela, Colombia,

Ecuador, or Peru were not confirmed: F. andicola
(excluded from Peru), F’ casapaltensis Ball (excluded
from Ecuador), F’ dasyantha Kunth (excluded from
Colombia), F: dichoclada Pilg. (excluded from
Ecuador), F: dolichophylla J. Presl (excluded from
Colombia and Ecuador), F’ ortophylla Pilg. (excluded
from Ecuador), F peruviana Infantes (excluded from
Ecuador), F: pinetorum (excluded from Venezuela),
F. quadridentata Kunth (excluded from Colom-
bia), F. rigescens (J. Presl) Kunth (excluded from
Ecuador), F’ vaginalis (Benth.) Laegaard (excluded
from Colombia), and F: ulochaeta Nees ex Steud.
(excluded from Ecuador).

MATERIAL AND METHODS

The work presented here summarizes the
results of taxonomic and biogeographic research
on the genus Festuca in paramos of South America
that was carried out between 1998 and 2004. Exten-
sive field and herbarium studies were performed to
assess the diversity of the genus in this geographic
area of South America. The area studied included
Brazil, Colombia, Ecuador, northern Peru, and Ven-
ezuela. An exhaustive bibliographic search for lit-
erature on the taxonomy of Festuca was performed,
with special focus on the genus in Mesoamerica
and South America. The protologues, local floras,
and monographs were consulted to document all
reports of Fectuca from this area. More than 1,500
specimens of Festuca from the study area and
1,000 specimens from other Mesoamerican and
South American countries were studied. Specimens
are located at AAU, B, BA, BAA, BOG, BOL,
CAUP,.C, CAR, COL, CUVC, CTES; — FAUC,
FMB, HUA, HERZ, JAUM, JBGP, K, LOJA, LP,
LPB, MA, MEDEL, MER, MERC, MERF, MY,
NY, QCA, QCNE, QAP, QPLS, P, PORT, PR,
PRC, PSO, R: RB, TOLT, TULY, U; USS; UPTC,
US, UTMC, VALLE, VEN, and W (abbreviations
according to Holmgren et al. 1998 onwards). The
type specimens of all recognized species and nearly
all types of their synonyms were examined.

The taxonomic analysis is based on exten-
sive morphological and anatomical comparisons.
Twenty-six morphological characters were selected
as useful for the descriptions and differentiation
among the species. The selected characters were:
height of plants, formation of shoot innovations,
number and position of the nodes, presence/absence
of auricles, length and diameter of the leaf blades,

8 Festuca in South American Paramos

shape and length of ligule, color and structure of
veins, vestiture of culms, length of panicle, length
of spikelets (measured total length of spikelet) and
number of florets, length, shape and vestiture of
glumes, lemma and palea, length of anthers, and
length of hilum. After preliminary observations
(Stancik 1999), five anatomical characters of the
leaf cross sections (number of vascular bundles,
number of ribs, distribution of the sclerenchyma,
relative density, and length of trichomes) were
also included in the study. Anatomical sections
were prepared from dry herbarium specimens after
hydratation in a soap solution and observed under
microscope at 100 magnification.

ACKNOWLEDGEMENTS

We would like to thank the Faculty of Science,
Charles University of Prague, for institutional sup-
port during realization of this research. We also
thank the grant agency of Czech Republic (projects
GACR 206/03/0197, GACR 206/06/P293), Minis-
try of Education of Czech Republic (project MSM
0021620828), and the Smithsonian Institution for
financial support. Our thanks belong also to Anna
Skoumalova for providing plant illustrations. We
wish to thank the Laboratory of Electron Micro-
scopy (LEM) of the Institute of Microbiology,
Academy of Sciences of the Czech Republic (Dr.
O. Benda, O. Kofronova, majority of photos on Fig.
72-97), LEM of Aarhus University (A. Sloth — Fig.
76E, 77A,B, 85C,D, 89C,D, 94A,B, 97C), LEM
of Royal Botanic Gardens Kew (Dr. H. Banks
— Fig.74A—D, 93E, F) for assistance during prepa-
ration of micro photos. This study could not have
been completed without the kind assistance of the
curators and researchers at the following herbaria:
AAU, B, BA, BAA, BOG, BOLV, CAUP, C, CAR,
COL, CONC, CUVC, CTES, FAUC, FMB, GB,
HUA, HERZ, JAUM, JBGP, K, L, LOJA, LP, LPB,
MA, MEDEL, MER, MERC, MERF, MO, MY, NY,
QCA, QCNE, QAP, QPLS, P, PORT, PR, PRC,
PSO, R, RB, SGO, TOLI, TULV, U, UIS, UPTC,
US, UTMC, VALLE, VEN, W. Finally, the Smith-
sonina Institution Scholarly Press is acknowledged
for preparing the final version for printing.

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org/science2/IndexHerbariorum.asp.

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1816. Nova Genera et Species plantarum.
(quarto ed.). Typographia d’Hautel, Paris.
Pp. 150-155.

Infantes, J.G.V. 1952. Estudio botanico, quimico y
fitogeografico del genero Festuca. Rev. Cien-
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Jorgensen, P.M and C. Ulloa Ulloa. 1994. Seed
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Jorgensen, M. and S. Leon-Yanez. 1999. Catalogue
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Festuca in South American Paramos 11

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en la Republica Argentina. Monogr. Syst. Bot.
Missouri Bot. Gard. 47: 1-178.

TAXONOMIC TREATMENT

wesmped 1... sp. Pl. 1: 73.1753. Tyee: & ovina L.,
Sp. Pl. 1: 73-74. 1753. (lectotype: designated
by Nash, Ill. Fl. U.S. Canad., ed. 2, 1:269. 1913;
also Jarvis et al., Watsonia 16: 300. 1987).

Perennial herbs, densely or loosely caespitose
(forming often large tussocks) or rhizomatous, com-
monly monoecious, exceptionally dioecious (not in
South America). Culms unbranched 0.1—2 m high,
below panicle glabrous or finely scabrous, with 1
or more (2—6) glabrous nodes. Innovations extra-
vaginal, intravaginal or mixed. Sheaths with free
or partially united margins, non-auriculate (except
in Festuca subg. Schedonorus); basal sheaths occa-
sionally thickened into a bulb; ligules membranous;
normally less than 1 mm, rarely to 10 mm long; leaf
blades flat but often conduplicate or setaceous and
then basal, without cross nerves, 0.2—-15 mm wide,
basal cataphyls rarely present. Inflorescences an
open or contracted panicle. Spikelets usually with 2
or several bisexual or unisexual florets and an apical
rudiment, chasmogamous, some species cleistoga-
mous or viviparous 1.e., vegetative proliferation of
the spikelets, laterally compressed, disarticulating
above the glumes and below the florets; glumes lan-
ceolate to ovate, pointed, carinate or non-carinate,

shorter than adjacent lemma; lower glume shorter,
1(rarely 3)-nerved; upper glume 3(rarely 5)-nerved;
lemmas commonly membranous (rarely coria-
ceous), rounded (exceptionally carinate), 5-nerved,
entire (rarely bidentate), acute, shortly mucronate
or awned; awn continuous with midnerve, terminal,
rarely subterminal, up to 15 mm long; paleas mem-
branous, two-carinate, two-dentate, as long as the
lemma or a little shorter, scabrous at keels, awnless;
lodicules two, small, about 1 mm long, two-dentate,
hyaline.; stamens 3, anthers 0.4—6 mm long; ovaries
glabrous, rarely hairy at apex, styles terminal with-
out an apical appendage; stigmas 2, white. Fruit a
caryopsis, free or with lemma adherent, with a linear
or sometimes oblong hilum.

Leaf blade anatomy.—Cross sections
with five or more vascular bundles, with or with-
out adaxial ribs over the vascular bundles; bul-
liform cells present or absent; sclerenchyma under
abaxial epidermis continuous or discontinuous,
adaxial sclerenchyma fascicles present or absent;
sclerenchyma sometimes extending to the vascular
bundles abaxially and also sometimes adaxially
forming girders; hairs on adaxial epidermis dense
or scattered, 20—200 um long, prickles on abaxial
epidermis present or absent.

KEY TO THE SPECIES OF FESTUCA IN BRAZIL, COLOMBIA, ECUADOR, AND VENEZUELA

la. Plants rhizomatous; culms solitary 30-100 cm tall or loosely caespitose herbs with culms 80—250

cm tall; plants often in mountain forests .........

lb. Plants densely caespitose, without rhizomes (only several species exceptionally forming short
rhizomes inside the tussocks); culms short 8—50(—60) cm tall or culms tall 60-170 cm tall; plants

Shea Wa nOGMalE SLAs ANS sted, sie hongscicuieleds eter pers eehle ahead Mdbiiwel, Mnowareedtig kus WMatiesdes. od 24
2a. Cataphylls (basal shoots coriaceous or with coriaceous-membranous scales) present................. 2
Bc aR SN eB aera See ang ach oc cl reaealdalatta ie tcl nad stele ah duc Vaiieawhondansbosexts 6
3a. Panicle branches densely hairy; ligules 1—1.5(—2) mm long...............cceeeeeseeeeeee 27. F. laegaardii

3b. Panicle branches glabrous or scabrous; ligules less than 1 mm long..............ccccecccceeseeeseeeeeeeeteeeees 4

12

Aa.

Ab.

Sa.

Sb.

6a.

6b.

7a.
Tilsys

8a.
8b.

9a.
Ob.

10a.
10b.

lla.
11b.

12a.
(2b:

iar

[3ib:

14a.

14b.

15a.
15b.

16a.
16b.

iva

17b.

18a.

Festuca in South American Paramos

Spikelets with 4-6 florets; lower glumes 2—3 mm long; upper glumes 34 mm long; lemmas 4-5 mm

lone tawned;paleasas long as adjacent lemima ..522222 28 Ai ieiese aeeee corners eee ee 30. F. toca
Spikelets with 24 florets; lower glumes more than 3 mm long; upper glumes more than 4 mm long;
lemmas more than 5 mm long, short-awned; paleas shorter than adjacent lemma...................0... 5

Panicles nutant and nodding, 9-12 cm wide; lower glumes 4—5 mm long; upper glumes 5.5—6.5 mm

long; lemmas 6—7 mm long; anthers I[—1-6 mm long’. :2ee ee eee 23. F. chitagana
Panicles erect, 0.5—1 cm wide; lower glumes 3—3.6 mm long; upper glumes 4.5—5 mm long; lemmas
6~-6.5 mm long xanthers' 3: Soman Lome A os ascesce ates scene eocosee ce eae eee 26. F. hatico
Auricles large; more than 1 mm) long; faleate .....0.......c.c.c00.ccsee.ceereeceree stot
Auricles absent or small, less than 1 mm long and inconspicuous, not falcate................:cseeeeeeee 8
Margins of auricles hairy; upper glumes 4.5—6.5 mm long................seeeeeeeeees 12. F. arundinacea
Margins of auricles without hairs; upper glumes 3.5—4 mm long@...............::::ceeee 13. F. pratensis
Apex Of TOdiCules Wainy ccsccicssasesteagtesssau-sgenndoeneccoscuerssosme ee toe ee eee ee 18. F. fimbriata
Apex of lodicules @labrouss.........c.<2c<c-.2c5cncecssndacaasensocenecectebsessavanseduacotecececasebeeasd eee eee rrr 9
Lower glumes 0-8—2.5 mitt 1OMG..5.2. 2... saSececccesceocaceathasonseeb acess oats sdscesesewee eee 10
Lower glumes longer than 2.5 mim OME.........:......ccccsesscccesoonccessseseeesecenescetvenntestee rn is
Lemma awns more than 4 mam JOMg 20.2.2... 2..c..cc0eccacesendceeeseatessesencaccesscnaseasonnaececeees seen 1]
Lemma awns less than 4 mim TOM 0.0.0.5. 2200. ecscccsoecsdesneeacssatscecesaceeseassneeeeneseea.ceasee rn 2
Ligules 3—3.5 mm long; lemma awns 9-12 mam long .....2.... 2.2.2 sseiccs gaceeose 4. F. flacca
Ligules 1-2 mm long; lemma awns 5—7 mim long ee..se ee ee 7. F. tovariensis
Spikelets 10-13 mm long; lemmas 7—8.5 mm long; anthers 3—3.5 mm long......... 10. F. reclinata
Spikelets 7-10 mm long; lemmas 4.5—6.5 mm long; anthers 0.8—1.5 mm long.................:0se 13
Leaf blades flat: panicles nutant: lemmas aywnless - sees ee eee ee 6. F. sodiroana
Leaf blades conduplicate or involute, sometimes flat; panicles erect; lemmas with awns 0.5—
LS) POM os cacs essa sed cn somes aaina ana o Reece dia Ha Ae chica page ct ames an oth eg Jt ee re ec rr 14
Culms solitary with long rhizomes; glumes and lemmas purple; panicles 0.5—1 cm wide...............
sevasaiedooses ade odo ceed e¥eti ad tcl dele nebau nt eeu abecbeacGits esr elaas ie Alea Aa Se ee 19. F. andicola
Culms loosely tufted; glumes and lemmas green; panicles 1—2 cm wide.................0 3. F. elviae
Ligules 2.5—13 mm long:.)....4)402.5 (ahi here ee 16
Ligules 0.5—1.5(—2) tin Lome 5.002 Sects seca evades cpa sonst eee ic ieee ee 19
Ligules 7-13 mm long; spikelets 11-14 mm long; lemmas awnless .................:ccscssccsseeseereeeeees 17
Ligules 2.5—-5 mm long; spikelets 14-17 mm long; lemma awns 1—8 mm long...................:006 18

Lower glumes (3.5—)4—5 mm long; upper glumes 5—6.5 mm long; margins of lemmas erose,
10 0. 0 [<1 « nn pm ne MEA OO oy eyo nce ccaccuaoeoncsactonaccactoconsostice 15. F. quadridentata
Lower glumes 5—7.5 mm long; upper glumes 7-8 mm long; margins of lemma entire ...................
stensdeesorsanasdanscenssasousoceutansich nite net sedisteel ay ReeUngn ee Aes sae estas aaa eae Ne Merete 14. F. dichoclada

18b.

19a.

19b.

20a.

20b.

21a.

21b.

22a.

22b.

23a.

23b.

24a.

24b.

Dae
25b.

26a.
26b.

Pia.

27b.

28a.

28b.

29a.

29b.

30a.

30b.

31a.
Silib:

Festuca in South American Paramos KS

Upper glumes 7—9 mm long; lemmas 10—11 mm long; anthers 3—3.5 mm long .............00ccccceeeee.
Re ene ae ad, cee Se ae ah es eases clan eupimtan eh aie hl, ie when oe i ib hase’ 16. F. venezuelana
Romemalnmess5— 5-5 (—4,) main lomeicnis. ci ecto ees etnset noes. eeeridadcte) hb eiees.. 20
Oe Fag DTN sh CHAT MN a h a ka snoy a ied Sao da gbtReeo vavanecdnsazesecen oundsesalGNeeencxs 21

Leaf blades 3—5 mm wide; upper glumes 4.5—6 mm long; lemmas awnless or with awns 0.5 mm
Rasa ES a naa eee enn ee cone in ores onoeptdn. oY ts psaplutln, ceceunues) tettdsh 2yadsieal 2. F. coromotensis
Leaf blades 5-11 mm wide; upper glumes 3.5-4.5 mm long; lemma awns 7—15 mm long,
AICO S mene sete taken tenh etait 92 linea MWe Mt nicely as sande th sme sce site Macabeal 8. F. ulochaeta

Upper glumes 5—5.5 mm long; lemmas 6.5—7 mm long; awns 2.5—3.5 mm long........ 20. F. rubra
Upper glumes 6—8 mm long; lemmas 8—10 mm long; lemma awns 0-1 mm long...................... Ze

Leaf blades conduplicate or involute, 0.6—0.7 mm in diameter; ligules 1—-1.5 mm long; lemmas
Epeaoibe) Iga AW | G)IN SS Aeemnene Mirae Ee mee ea. 2 oarcinail dnc deel -daceniiseesdedinuennesesededeniensdcsmegnecdseses 5. F. guaramaca
Leaf blades flat, 3-12 mm wide; ligules 0.5—1 mm long; lemmas 8—9 mm long......................4.. 23

Caespitose plants with rhizomes, 120—180 cm tall; lemma awnless; anthers 3 mm.........................
Spree Peer (yb Serr nna toned a: Benes eran, aclu hy chon cds) Ar wet lie nate Nee wld dada 1. F. amplissima
Culms solitary, rhizomatous, 70—90 cm tall; lemma short-awned, the awns 0.5—1 mm long; anthers
re ABITIBI OMS) teen eens eee tes aarni se Se 8 Be ole | boas 2. haan ctii nada aua lee dedecumieuty leeds 11. F. woodii

Cataphylls (basal shoots coriaceous or with coriaceous-membranous scales) present................ DS
EET ESOVES IIS 1 0 SETI o hacia Bone Ene heehee tes Soe EEE SRE oi ee ee et ee 30

emits with/a-dense coverine.of Lome hairs :2e...6....8 cc. cswetd cece teens enn ceavetecdenee 25. F. dasyantha
MASE ATOMS Ol) SCADTOUS 2 =. 55) acco ccs dues ee Pet USN Sunk eee he. eae ede eee Beh 26

Leaf blades 0.8—1.6 mm in diameter; abaxial schlerenchyma continuousS...............:::cccseeeeeeeeeees 2H,
Leaf blades 0.4—0.7 mm in diameter; abaxial schlerenchyma discontinuous ..............:c::ceeeeeeeees 22

ER eR ee Pere era vine e ing Meets ates chs) da lnetlsoa dese Cee OIRRY ANIM Seth lull shiners edeatbane es 28. F. pilar-franceii
Cataphylls 0.5—1.5 cm long; ligules more than 0.3 mm long; lemmas 6—8 mm long.................. 28

Lower glumes 5.5—6.5 mm long; upper glumes 7—8.5 mm long; anthers 0.8—1.1 mm long.............
EE ener ee Pate me, eee ate, As) yh cutee dnd So. seelae ht oon cee ees isaee eA, ete 35. F. chita
Lower glumes 3-4.5 mm long; upper glumes 4.5—6.5 mm long; anthers 2—3.5 mm long................
EY Oe Nias Mm eae Stes Toss ti teen) Si teen Nd AS ee hee alee ol aes copie Le PY 24. F. colombiana

Pe ee se 2 oasis ete AO ies Sime eedel geese. Tul. Meee apaee ct cee i bes cet hie ati a 3)
Plants growing in small tussocks, culms 15—60(—80) cm tall and 0.05—0.2 m in diameter near
[SPESE dd cccke Re eae Se eet a8 ee SB Het OMe nT ESE Sn SUR ROR NTP EO EY me el SRNR SNe 43
LISS 0 AIOE SIS 1b OE, eee es Mane Lae Ane a ee yi Sone aR eS EEE SON Me Toney ee 32

ecm est Ole SPO AUTEN Vea eee one yeh en sls ye ccs sweden Ueiegatien ath, Sevens tietvad sia adi echacel ciavs Deion nie 36

14

Festuca in South American Paramos

32a.
32b.

33a.

33b.

34a.

34b.

35a.

35D:

36a.

36b.

37a.
37b.

38a.

38b.

39a.

39D.

40a.
A4Ob.

Ala.

Alb.

42a.

42b.

43a.

43b.

44a.

Leaf blades 0.5—0.7 mm in diameter; ligules predominantly acuten./. 220002... .0et eee 33
Leaf blades 0.8—1.5 mur iin diameter: lisulls truncate . ccc.ccctecessccsoccsnesevesteceesesssseeeceseees sso eee 35

Some spikelets transformed into vegetative shoots (viviparous plants), sexual organs in such spikelets
ADS CINE: cecesavssiececsn hi diecaaalbens uch eceateee enue aen take ceon deme ecceuautmeu cases pcan anak anne ne: tae eae 52. F. subulifolia
Spikelets perfect, well developed, no spikelets vegetatively prolifering ................::cccesseeeeeeees 34

Spikelets 8-14 mm long; lower glumes 6—8.5 mm long; upper glumes 6.5—9.5 mm long; lemmas
610 in lone Re a ie RO, EE ea hae 54. F. tolucensis
Spikelets 6.5—9 mm long; lower glumes 3.5—5.5 mm long; upper glumes 4-6 mm long; lemmas
56/5 (28) tata NOM 5522. Sessa aeccs va dap oiguecaticen eso ue wel trast ween too eeeeitoucoat aun Sones 52. F. subulifolia

Spikelets perfect, not vegetatively prolifering, 8—10 mm long, with 34 florets; lemmas 6.5—8.5 mm
long. with awins’ O71 mimlOMme: 250.2 ee.e aetestees nc oeaees en neree tee ceeeeaten eee oe 37. F. densipaniculata
All spikelets with vegetative proliferation; lemmas transformed into vegetative shoots, sexual organs
AUD SSL vp tahes snc snesat ca vet ae eee baamacea etree nee Lenten ot bed et ee eteesen lec eetew ete sa te ean eee eek et eee 29. F. asplundii

Leaf blades flat or conduplicate, 2—5 mm in diameter; spikelets 12—15 mm long.....................:e0008
stalin bn eee a etc aR A ER SI AEs SN. 42. F. glyceriantha

Leaf blades involute, less than 2 mm in diameter; spikelets shorter than 12 mm long................ 37

Anthers 11:5: mam long 5. 2.2ccesvascacereedelondyed dear aeewaeaubecenotiete cc ceceeceeeeensee cee: Reet eee eer 38
Anthers:2=3 tram lomo... 2.52 Beas wickcec cet tens eceaceerenanpnadi deed eseneedneunsanone cane ee 39

Culms about 50 cm tall; leaf blades 0.6—0.8 mm in diameter; spikelets with 3-4 florets; lower glumes

4mm long: upper elumes?s mami Tome: =. e ees ceeaese ee eee eee 46. F. monguensis
Culms 80—100 cm tall; leaf blades 1.2—1.5 mm in diameter; spikelets with 5 florets; lower glumes
6=6.5 mm long; upper glumes. 7—S:mimilome <cscceeesce eee ee 47. F. nereidaensis

Leaf blades 1.1—1.8(—4) mm in diameter; lower glumes 2.4—2.6 mm long; upper glumes 3.6—
3:9 man LONG ae... Ae BO AE RE, CTE ete 48. F. oroana
Leaf blades 0.4—1 mm in diameter; lower glumes 3—7 mm long; upper glumes 4-8 mm long.... 40

Ligules longer than. 1-mim; panicles Opet s...42.0.0s.c.nscceesoesevqeceesseteieseset nee eae ee 41
Ligules shorter than 1 mm; panicles comtracted!:...22.5...200.0-205.-ca0b 2) ceeessescsee 42
Ligules (1.5—)2—3.5 mm long; lower glumes 3—4 mm long; upper glumes 4—5.5 mm long.............
veeceosonbacisusdpaii «nalts wOvauy alae ate ote at eee ORE oe tnaitee Nit Unt ti eet ie ee eenae ema 55. F. turimiquensis
Ligules (0.5—)1—2(—2.5) mm long; lower glumes (3.5—)4—6(—6.5) mm long; upper glumes (5.5-)
6—7.5G-8) mom LOMB. 2: gesscccesite sassy eee nee ee oe 36. F. cleefiana
Leaf blades 0.4—0.6 mm in diameter; ligules 0.5—1 mm long; spikelets with 3-4 florets; lemmatal
awns |.5—4 mm long.) 2328.4. ee ee eee 40. F. dinirica
Leaf blades 0.7—0.1 mm in diameter; ligules 0.1—0.3 mm long; spikelets with 5—7(—8) florets; lem-
matal awns O=1 mim... asec eee 38. F. cundinamarcae
Spikelets about 3 cm long, with 5—7 florets; lemmas 17—20 mm long ....................+ 17. F. fragilis
Spikelets less than 2 em long, lemmas'less than 15 mm lomersi22 52 oes seee eee 44
Lower glumes 1.2—2 mm long; upper glumes 2—2.7 mm long; lemmas 4.5—5.5 mm long...............

Serene ener er ee Ti shag saonyacdusacosbdasoaocecoonasconcisovaoenseons 21. F. soukupii

A4b.

45a.

A5b.

46a.
46b.

47a.

A7b.

48a.

A8b.

49a.

49b.

50a.

50b..

1a.

Sb!

52a.

520.

53a.

53D:

54a.

54b.

55a.

55b.

Festuca in South American Paramos 15

Fa ne sad ENR ene eens Re Aen aos he ae eet MN Rk oer unl 45
SFE OI GLUE 2111707 1 ac cee eee em ee ere eC ne i a er eT 46
“er EeL! DEVE SS. SOR TESTIS Gee ae ey eal 2S hs Roc ee a 8 CECE EAR 0 a rr 52
Bereta evs La AMIS | OO ee LEN ATIE AAR et net eae ee eee See he ied dy tab dc Geb dade iedusaceseassesessoccuezuecaabede 47
Mematiatayilless OG awiis SmOrten thai | TMI so .2 ooo. ccc coeicaceececadcactaseccdoeacecossccclscicevasacaecaccusveccccevescee 48

Culms 8—15(—30) cm tall; lower glumes 2.5—3.5 mm long; upper glumes 3.5—5 mm long; lemma

“TED, CODD Spa TO 1 A ceo Me ee 34. F. chimborazensis
Culms 15-55 cm tall; lower glumes 44.5 mm long; upper glumes 5.5—6.5 mm long; lemma with
FPO TNS 4 OA CUO 0 sy ge Nn eee ec ee 44. F. huamachucensis

Leaf blades less than 0.8 mm in diameter; lower glumes 2.54.5 mm long; upper glumes 4-6.5 mm
SES ECPOVETEUS SS) O07 ey gh tn Ve TO SS) 0 cp nee, nee eT en 49
Leaf blades (0.5—)0.8—3.5 mm in diameter; lower glumes 6.5—10 mm long; upper glumes 6.5—
ws) TEDEED LCV SST TTS 7s COIS Vet ene oe CE 50

Ligules 0.5—0.7 mm long; upper glumes 44.2 mm long; lemmas 5.5—6(—7) mm long...................

soc seas sossabaagioseagtiage acetone ar Ne ari io eee gor ne ene rn et eee 45. F. imbaburensis
Ligules 1.2—-1.5 mm long; upper glumes 5—5.5 mm long; lemmas 7—7.5 mm long..................0:008

Leaf blades involute, 0.5—0.8 mm in diameter; panicles about 0.5 cm wide; spikelets 8—9.5 mm

CLES SITOITIE IS, 7/5 (OMS 010 ha FY 0 ae le ne 56. F. vaginalis
Leaf blades conduplicate, up to 3.5 mm wide; panicles 0.8—2 cm wide; spikelets 10—13 mm; lemmas
Seem CRIEN Ue eet MO FC AA Bod uy ode Mend ac scch sz h Cleese) Seat avons) Mac cddvanes cast abecsS Mel Adee eee caseee othe. Dil

Culms 20—50 cm tall; leaf blades 0.8—2 mm in diameter; ligules 0.6—1 mm long; panicles 0.8—1.5 cm

A OWVCn PUIMICS 7.) —O UMAN (OIG, 2555-22502. 20.20. <sapcnrediageretaccdaaviasieooasesedtbeeloteee 37. F. cocuyana
Culms 60—80 cm tall; leaf blades 2—3.5 mm wide; ligules 1—1.6 mm long; panicles 1.5—2 cm wide;
OPO PELE SITE SS SIS GOV 10) 1278 ee am ee 51. F. sanctae-martae
ereules 0:3—0:5 min lone-calius and rachilla clabrous........:....0..0i00 i as ee Ree 53
Ligules 0.5—1.2 mm long; callus and rachilla scabrous or sparsely hairy.............c:cceseeeesteeeeeeees 54
Culms 15-20 cm tall; spikelets 9.5—11 mm long; lower glumes 4—5 mm long; upper glumes 5.5—6 mm
long; paleas as long as adjacent lemma; anthers 0.6—0.8 mm long... 53. F. sumapana
Culms 30-60 cm tall; spikelets 8—9.5 mm long; lower glumes 3—3.7 mm long; upper glumes 4-5 mm
long; paleas shorter than adjacent lemma; anthers 0.9—1.2 mm long................... 32. F. boyacensis

Leaf blades 0.4—0.7 mm in diameter with 5 vascular bundles; paleas as long as adjacent lemma...
a lecjepssrs isd et a fee DAS aOR Raitt Niece ee ee POSTER PEE COT Pe tr RMe <- Te AE 4 Mh, Pea 5

RS are ame RE Peet io ern UN Teed oto Woks duc totnvcne al ec ovser TeAcAOhase cou cvvRe ee eee EG! 49. F. parciflora
Leaf blades green; spikelets 9-10.5 mm long; lower glumes 3.5—4.5 mm long; upper glumes
ee ees eMUA UTI GHA Oe a rar Me cenaest tay Te Soran neeay tape or ieee aeraeivesteaseunsusitenes sasenaceye MeRVerte te 33. F. carchiense

16 Festuca in South American Paramos

56a. Culms 50-60 cm tall; ligules 0.6—0.8 mm long; lower glumes 4 mm long; upper glumes 5.5 mm

long; santhers) £3: mimalome 2a ee ee oe ie sca eea torecmaaee 43. F. holubii
56b. Culms 15—55 cm tall; ligules 0.8—1.2 mm long; lower glumes 4.5—6 mm long; upper glumes 6—6.5
mim-lone:anthers 0:8=00 aims Lome sees esos cess tecehees eee ee 41. F. glumosa

SURVEY OF THE SPECIES AND SUBGENERIC TAXA OF
FESTUCA IN SOUTH AMERICAN PARAMOS

1. Subg. Drymanthele V.1. Krecz. & Bobrov, Fl. URSS 2: 532. 1934. Type: F. drymeja Mert. & W.D.J.
Koch
1.1. Sect. Ruprechtia E.B. Alexeev, Novosti Sist. VysS. Rast. 17: 42. 1980. Tyre: F’ amplissima Rupr.
1. Festuca amplissima
2. Subg. Subulatae (Tzvelev) E.B. Alexeev, Bjull. Moskovsk. Ob8¢. Isp. Prir., Otd. Biol. 82(3): 96.
1977. Type: F) subulata Trin.
2.1. Sect. Subulatae Tzvelev, Bot. Zhurn. 56(9): 1253. 1971. Type: FE. subulata Trin.
2. Festuca coromotensis
3. Festuca elviae
4. Festuca flacca
5. Festuca guaramaca
6. Festuca sodiroana
7. Festuca tovariensis
8. Festuca ulochaeta
2.2. Sect. Glabricarpae E.B. Alexeev, Bot. Zhurn. 67(9): 1291. 1982. Type: F. breviglumis Swallen
9. Festuca caldasii
10. Festuca reclinata
11 Festuca woodii
3. Subg. Schedonorus (P. Beauv.) Peterm., Deutschl. Fl. 643. 1849. Type: F. arundinaceae Schreb.
3.1. Sect. Schedonorus (P. Beauv.) Koch, Syn. Fl. Germ. Helv. 810. 1837. Type: F. arundinaceae
Schreb.
12. Festuca arundinacea
13. Festuca pratensis
4. Subg. Erosiflorae E.B. Alexeev, Novosti Sist. Vyssh. Rast. 23: 11. 1986. Type: F) quadridentata
Kunth
14. Festuca dichoclada
15. Festuca quadridentata
16. Festuca venezuelana
5. Subg. Helleria E.B. Alexeev, Novosti Syst. Vyssh. Rast. 17: 42. 1980. Type: F: livida (Kunth) Willd.
ex Spreng.
17. Festuca fragilis
6. Subg. Mallopetalon (Doll) E.B. Alexeev, Bot. Zhurn. 69(3): 346. 1984. Type: F: fimbriata Nees.
18. Festuca fimbriata
7. Subg. Festuca
7.1. Sect. Aulaxyper Dumort, Observ. Gramin. Belg.: 104. 1824. Type: F rubra L.
19. Festuca andicola
20. Festuca rubra
21. Festuca soukupii
7.2. Sect. Cataphyllophorae E.B. Alexeev, Bot. Zhurn. 69(11): 1546. 1984. Tyre: F: procera Kunth
22. Festuca azucarica
23. Festuca chitagana
24. Festuca colombiana
25. Festuca dasyantha
26. Festuca hatico

Festuca in South American Paramos 17

27. Festuca laegaardii
28. Festuca pilar-franceii
29. Festuca procera
30. Festuca toca

7.3. Sect. Festuca
31. Festuca asplundii
32. Festuca boyacensis
33. Festuca carchiense
34. Festuca chimborazensis
35. Festuca chita
36. Festuca cleefiana
37. Festuca cocuyana
38. Festuca cundinamarcae
39. Festuca densipaniculata
40. Festuca dinirica
41. Festuca glumosa
42. Festuca glyceriantha
43. Festuca holubii
44. Festuca huamachucensis
45. Festuca imbaburensis
46. Festuca monguensis
47. Festuca nereidaensis
48. Festuca oroana
49. Festuca parciflora
50. Festuca renvoizei
51. Festuca sanctae-martae
52. Festuca subulifolia
53. Festuca sumapana
54. Festuca tolucensis
55. Festuca turimiquirensis
56. Festuca vaginalis

DESCRIPTIONS OF SPECIES OF FESTUCA FROM SOUTH AMERICAN PARAMOS

1. Festuca amplissima Rupr. subsp. amplissima,
Bull. Acad. Roy. Sci. Bruxelles 9(2): 236. 1842.
(Figs. 2, 72A—D). Tyre: Mexico. Veracruz, Pic
d’Orizaba, 10,000 ft, Jun—Oct 1840, H. Galeotti
5766 (lectotype: BR; isotype: W!).

Festuca fratercula Rupr. ex E. Fourn. Mexic. PI.
2:124. 1886. Type: Mexico. Veracruz, Cordil-
lera, Pico de Orizaba, 12,000 ft, Jun—Oct 1840,
H. Galeotti 5778 (lectotype: LE!, designated
here).

Uniola effusa E. Fourn., Mexic. Pl. 2: 122. 1886. Type:
Mexico. San Nicolas, E. Bourgeau 1032 (holo-
type: P; isotype: US-865711 fragm. ex P!).

Uniola muelleri E. Fourn., Mexic. Pl. 2: 122. 1886.
Type: Mexico. Veracruz, Orizaba, Mueller 2115
(holotype: LE; isotype: US-091892 fragm. ex
LE!),

Rhizomatous perennials, innovations extra-
vaginal. Culms 120-180 cm tall, erect, scabrous;
nodes 3—6. Leaf sheaths coriaceous, scabrous,
brown, striate, older basal sheaths disintegrating
into fibres; auricles absent; ligules 0.5—1 mm long,
coriaceous, apex truncate; blades 35—70 x 0.4—1.2
cm, flat to involute at tip, green, abaxially scabrous,
with prickles on adaxial and abaxial surfaces.
Panicles 30-40 cm long, 25—40 cm wide, lax,
pyramidal, branched; branches scabrous. Spikelets
12-15 mm long, lanceolate, florets 4 or 5; rachilla
densely hairy; glumes 4.5—-8 mm long, membranous,
narrowly lanceolate, green, scabrous; lower glumes
4.5—6 mm long, 1|-nerved; upper glumes 6-8 mm
long, 3-nerved; lemmas 8—9 mm long, 5-nerved,
membranous to coriaceous, lanceolate, green, sca-
brous and densely hairy, apex entire, awnless; callus

18 Festuca in South American Paramos

sparsely hairy; paleas as long as adjacent lemma,
membranous, papillose, keels scabrous; anthers
about 2.9 mm long; ovary apex glabrous. Caryopses
lanceolate; hilum 4/5 as long as the grain.

Leaf blade anatomy.—Cross sections with
many vascular bundles; adaxial ribs small; scleren-
chyma under both abaxial and adaxial epidermis,
discontinuous, extending to the vascular bundles
forming girders; adaxial epidermis with bulliform
cells and without hairs.

Observations —Festuca amplissima is the only
species represented in Festuca sect. Ruprechtia. It
differs from other species of F’ subg. Montanae
by lacking cataphylls and by having glabrous
ovaries.

Distribution and habitat—This species was
confirmed in Mexico, Guatemala, Costa Rica, and
Panama. Davidse et al. (1994) also mentioned this
species from Venezuela. In Central America and
Mexico this species is known from forests (Pinus
spp. and Quercus spp.) and forest clearings.

Additional specimens examined. COSTA
RICA. Cartago: Canton de Turrialba, Parque
Nacional Chirripo, Cuenca del Matina, estacion
Crestones, sendero a Ventisqueros, 09°29°52’°N,
83°29'20"W, 3400-3500 m, 11 May 1996, Gambo
et al. 819 (MO). San José: Trail to the Valle de
los Leones, lower part of Valle de los conejos,
along the upper river Talari paramo formation
with Chusquea sp. and areas of short burned for-
est, 83°31°W, 09°27'N, 3250-3450 m, 21-23
Aug 1971, Burger & Gomez &246 (NY); Canton
de Pérez Zeledén, Parque Nacional Chirripo,
cuenca Terraba - Sierpe Chirripo, base Crestones
paramo, 8°27°25"N, 83°30'38"W, 3460 m, 12 Jul
1996, Alfaro 1027 (MO); Along the trail from
Canaan to Chirripo via Los Angeles above N the
Rio Talari, elfin forest of dead oak trees, 09°30'N,
83°31°W, 3200-3400 m, 19-22 Jan 1970, Burger
etal. 7428 (F). GUATEMALA. Chimaltenango:
Volcan de Agua, 10000 ft, 22 Jul 1937, JRJ 895
(F); Volcan de Acatenango, above Las Calderas,
2700-2900 m, 1 Mar 1939, Standley 61885 (F);
above Las Calderas, 2700-2900 m, 1 Mar 1939,
Standley 61599 (F). Sacatepequez: Volcan de
Agua, above Santa Maria de Jesus, 2250-3000
m, 2 Nov 1939, Standley 65227 (F). MEXICO.
Distrito Federal: Ajusco, Rancho Vieja Cuilotepec
Tlalpan, 2730 m, 19 Sep 1985, E. Manriqu et al.
1147 (MO); Valle de Mexico, 2500 m, 1 Jan 1952,
Matuda 25909 (MO). Chiapas: Mun. San Cristdbal
las Casas, at Piedracitas, 7 km E of San Cristobal

along road to Zontehuitz, slopes with Quercus and
Pinus, 7800 ft, 20 Jul 1965, D. Breedlove 11151
(F); Mun. Tenejapa, Paraje Matsab, 9000 ft, 5 Dec
1966, Alush Shilom Ton 957 (F). Guerrero: Mun.
Heliodoro Castillo, ladera E del Cerro Teotepec, 68
km de Filo de Caballo por terraceria a El Paraiso,
Bosque de Abies, Quercus, Pinus, Alnus y Clethra,
2730 m, 23 Nov 1991, Gonzalez-Ledesma et al. 484
(F); Mun. de Chilpancingo, Parque Omiltemi a 35
km W of Chilpancingo bosque mesofilo, 2450 m,
21 Nov 1991, Gonzalez-Ledesma et al. 478 (MO).
Jalisco: 14-18 km SW of Tequila on Volcan de
Tequila Quercus, Arbutus forest, 2400-2600 m,
11 Jul 1974, D. Breedlove 39265 (MO). Micho-
acan: Mountain near Patzcuaro, 8500 ft, 23 Nov
1891, C.G. Pringle 3945 (NY); Mt. Tancitaro,
rocky ledge, 10300 ft, 25 Jul 1941, Leavenworth
et al. 1213 (F, MO, NY). Oaxaca: Distr. De Ixtlan,
Mun. De Ixtlan, Rancho Teja, 0.25 km N of the
upper casita, 2250 m, 4 Aug 1981, Martin 630
(US). Puebla: Mun. San Nicolas de los Ranchos
6 km al SE de Paso de Cortez, brecha a Xalitzintla,
20°18'N, 98°44’ W, 3400 m, 14 Sep 1988, Tenorio
15093 (US). Veracruz: Pico de Orizaba, 2500 m,
Liebmann 6109, 6110 (C); Sep 1841, Liebmann
12904 (C). PANAMA. Chiriqui: Volcan Baru, in
and near Potrero Muleto, abundant floor of crater
9400-10000 ft, 17 Nov 1978, B. Hammel 5619
(MO); Volcan Chiriqui, Boquete Distr., 10400 ft,
18 Jul 1938, Davidson 1043 (F, MO); W slope of
Volcan Chiriqui, 11 km directly WNW of Boquete,
ridge looking down on Potrero Muleto, elfin forest
dominated by Ericaceae, 3000 m, 20 Nov 1975,
G. Davidse & D’Arcy 10244 (NY).

1b. Festuca amplissima subsp. sierrae Stancik,
Preslia, Prague, 75: 341. 2003. (Fig. 1). Type:
Colombia. Magdalena, Sierra Nevada de Santa
Marta, Hoya del Rio Donachui, Cancurtia, SE
slopes, 2400-2650 m, 10 Oct 1959, J. Cuatre-
casas & Castaneda 24736 (holotype: COL!;
isotype: US!).

This subspecies differs from typical form by hav-
ing a larger, denser, and widely spreading panicles.

Observations.—The outstanding character
of larger panicles and the isolated geographical
position in the northern Andes (Sierra Nevada de
Santa Marta, Serranna de Perija), were the primary
reasons for recognizing this subspecies.

Distribution and habitat—In Colombia and
Venezuela Festuca amplissima subsp. sierrae

Festuca in South American Paramos 19

WV
NOH x \\y

\i NAT \ Wy

Ni VINNY,

{
\ Ny \ \ IN vu \)

\

Lm
—

Yo TZZ=

) yy
ANY i
N) Y) NS NY
N\ \N vA

100 x

= = ee
——

Figure 1. Festuca amplissima subsp. sierrae. A. Stylized growth form. B. Habit. C. Spikelet. D. Glumes. E. Lemma. F. Leaf blade
cross-section. A-E, Cuatrecasas & Castaneda 24736 (COL). F, Barclay & Juajibioy 7033 (COL).

occurs in matorral and shrub lands and the grass
transitional zone of the paramo at an altitude of
2200-3400 m.

Additional specimens examined. COLOM-
BIA. Magdalena: Sierra Nevada de Santa Marta,

alrededores de cabeceras de Rio Ancho, paramo
de Macotama, 3490 m, 16 Feb 1959, Barclay &
Juajibioy 7033 (COL, MO, US); Paramo de Maco-
tama, NW side of valley, grassy slope, 3490 m,
13-18 Dec 1959, Barclay & Juajibioy 7017 (MO);

20 Festuca in South American Paramos

Mun. Ciénaga, Cabeceras del Rio Sevilla, 3500 m,
Carbono 2474 (UTMC); transecto del Buritaca-filo
La Cumbre, 3020 m, 15 Aug 1977, Rangel et al. 945
(COL); about 30 mi inland from Dibulla, 3850 m,
Jul 1923, Seifriz 489 (US). VENEZUELA. Zulia:
Serrania de Perija, Serrania de los Motilones, Distr.
Perija, headwaters of Rio Negro, 3000 m, 27 Jun
1974, Tillet & Honig 746—765 (MO).

2. Festuca coromotensis B. Bricefio, Ernstia 4 (3-4):
76. 1994. (Figs. 2, 3, 72E—F, 73A & B). Type:
Venezuela. Mérida, Dept. Libertador, Parque
Nacional Sierra Nevada, Laguna La,Coromoto,
3300 m, 3 Jul 1987, B. Briceno & G. Adamo
2003 (holotype: MERF!; isotype: MERC!).

Loosely tufted, rhizomatous perennials with
extravaginal innovations. Culms 100-120 cm tall,
erect, scabrous; nodes 4 or 5 on lower 1/2. Leaf

sheaths membranous, glabrous, striate, brown; auri-
cles absent; ligules 0.5—0.7 mm long, membranous to
coriaceous, apex truncate, ciliate; blades 15—25 cm
long, 3—5 mm wide, flat, green, abaxially scabrous.
Panicles 20-25 cm long, 2—5 cm wide, flexuous,
pendant; branches scabrous. Spikelets 9-11 mm
long, florets 3 or 4; rachilla ca. 1.2 mm long, pilose;
glumes 2.5—6 mm long, narrowly lanceolate, coria-
ceous, purplish, pilose, apex acute; lower glumes
2.5—3.5 mm long, |-nerved; upper glumes 4.5—-6 mm
long, 3-nerved; lemmas 7.5—8.5 mm long, 5-nerved,
lanceolate, membranous to coriaceous, margins short
pilose, apex entire, awned or awnless, pilose, the awn
0—0.5 mm long; paleas as long as the lemma, pilose;
lodicules ca. 0.8 mm long, oblong; anthers 2—2.5 mm
long; ovary apex glabrous. Caryopses lanceolate;
hilum 4/5 of the total length.

Leaf blade anatomy.—Cross-sections with
numerous vascular bundles; adaxial ribs small,

VENEZUELA

COLOMBIA

e ECUADOR

-80

Figure 2. Distribution of Festuca amplissima (@), F. coromotensis (&), and F. flacca (m).

Festuca in South American Paramos 24

H +

Figure 3. Festuca coromotensis. A. Stylized growth form. B. Habit. C. Ligule. D. Spikelet. E. Glumes. F. Lemma. G. Lemma with
palea and rachilla. H. Leaf blade cross-section. A—H, Stancik 4173 (PRC).

inconspicuous; sclerenchyma below abaxial and Observations —The habit of F: coromotensis is
adaxial epidermis, discontinuous, forming girders; _ very similar to F’ sodiroana. However, the spikelets
bulliform cells absent; abaxial epidermis with scat- of F. sodiroana have shorter glumes (lower glumes
tered prickles; adaxial epidermis with sparse hairs, — 1.3—1.8 mm, upper glumes 2.2—3.5 mm), shorter lem-
the hairs ca. 0.1 mm long. mas (5—6.5 mm), and shorter anthers (0.8—1.2 mm).

D2 Festuca in South American Paramos

Distribution and habitat.—This species is
endemic to Venezuela, known only from the Andean
states of Mérida and Tachira. Festuca coromotensis
occurs in the Andean forest and matorral communi-
ties between 3000-3500 m.

Additional specimens examined. VENEZU-
ELA. Mérida: Entre Los Arbolitos y Los Nevados,
Paramo El Toro, 3100 m, 12 Oct 1988, B. Briceno
et al. 2328 (Herbarium Bricefio); Mun. Santo
Domingo, Laguna Negra, patches of Polylepis,
08°47’13.4’N, 70°48 '26.1” W, 3500 m, 6 Nov 2000,
D. Stancik 4173 (CAR, COL, PRC); Mun. Santo
Domingo, Laguna Mucubaji, schrubby patches,
08°47°19.5"N, 70°48°36.1°W, 3470 m, 6 Nov
2000, D. Stancik 4174 (AAU, CAR, COL, PRC);
Mun. Tabay, Laguna Coromoto, mountain forest,
08°35'41.7'N, 71°01 26.9" W, 3100-3200 m, 7 Nov
2000, D. Stancik 4180 (AAU, CAR, COL, US,
VEN, W); Mun. Tabay, Laguna Coromoto, moun-
tain forest, 3000—3100 m, 7 Nov 2000, D. Stancik
4177B (PRC). Tachira: Paramo entre Enconejada
para Pogonero, 3000 m, Sep 1955, Baldillo 3496
(VEN); Mun. La Grita, Par. La Negra, 8°15°11.2°N,
71°52’41.5°W, 3200 m, 10 Nov 2000, D. Stancik
4281 (AAU, CAR, COL, VEN); 8°15'26.4°N,
71°52°16.5°W, 3100 m, D. Stancik 4282 (CAR,
COL, PRC).

3. Festuca elviae B. Bricefio, Ernstia 4 (3-4): 77,
f. 2-4. 1994. (Figs. 4, 6, 73C—F). Tyre: Vene-
zuela. Mérida, Libertador: Paramo La Culata,
camino hacia Laguna Tapada, 3300 m, 19 Oct
1984, B. Briceno & G. Adamo 1114 (holotype:
MERF!).

Loosely tufted, rhizomatous, perennials with
extravaginal innovations. Culms 40—80(—-100) cm
tall, erect, glabrous, nodes 2 or 3 in basal half. Leaf
sheaths membranous, brown, glabrous; auricles
absent; ligules 0.7-1.5 mm long, membranous,
apex truncate, ciliate or dentate; blades 15—20 cm
long, 1-6 mm wide, flat, sometimes involute, green,
abaxially scabrous. Panicles 10-17 x 1—2 cm, con-
tracted, erect; branches scabrous. Spikelets 7-9 mm
long, florets 3-5; rachillas 1-1.4 mm long, glabrous;
glumes 0.8—3 mm long, narrowly lanceolate, mem-
branous, green, glabrous, apex acute; lower glumes
0.8-1.5 mm long, 1-nerved; upper glumes 2-3 mm
long, 3-nerved; lemmas 4.5—5.5(—-6) mm long,
5-nerved, lanceolate, membranaceous-coriaceous,
green, apex glabrous or scabrous, entire, sometimes
shortly awned, the awn 0.5—1(—2) mm long; paleas

almost as long as the lemma, apex scabrous; anthers
0.9-1.2 mm long; ovary apex glabrous. Caryopses
oblong; hilum nearly the entire length.

Leaf blade anatomy.—Cross sections with
numerous vascular bundles and ribs on abaxial
surface; sclerenchyma under both abaxial and
adaxial epidermis discontinuous and extending to
the vascular bundles; bulliform cells absent; abaxial
epidermis with scattered prickles.

Observations.—Spikelets with short glumes
and short-awned lemmas suggest affinity with the
Colombian and Ecuadorian species, F’ andicola
and F. sodiroana. However, Festuca andicola and
F. sodiroana have slightly longer lower glumes
(1.3-1.8 mm), nodding culms, and narrower
panicles (0.4—1 cm).

Distribution and habitat.—This species is
endemic to Venezuela, known only from the state of
Mérida. It occurs in the humid or swampy margins
of the Andean forest, along streams, in clearings,
and in pastures, between 3000-3700 m.

Additional specimens examined. VENEZU-
ELA. Mérida: Dept. Rangel, SE de la entrada a la
Lag. Mucubaji, 3550 m, 28 Aug 1980, B. Briceno
& Adamo 191, 196 (MERF, MO); 3550 m, 2 Oct
1996, B. Briceno et al. 3390 (Herbarium Breceno);
Dept. Rangel, SE del Hotel Los Frailes, 3100 m,
22 May 1981, B. Briceno & Adamo 274 (MERF,
MO); Dept. Rangel, camino hacia Laguna Las
Canoas, 3100 m, 10 Jul 1981, B. Briceno & Adamo
306 (MERF); Paramo Los Granates, 3420 m, 30
Aug 1985, B. Briceno & Adamo 1320 (MERF);
Paramo Misinta, 3100 m, 3 Dec 1986, B. Briceno
& Adamo 1705 (MERF); Mun. Santo Domingo,
Laguna Mucubaji, swamps around the lagoon,
08°47°33"N, 70°48'56"W, 3600 m, 6 Nov 2000,
D. Stancik 4171 (AAU, CAR, COL, PRC); Laguna
Mucubaji, shrubby patches, 3470 m, 8°47°20°N,
70°48'36"W, D. Stancik 4175 (AAU, CAR, COL,
PRC). Mun. Mucuchies, Paramo de Piedras Blan-
cas, 8°48'31"N, 70°55’08”"W, 3700 m, 4 Nov 2000,
D. Stancik 4215 (AAU, CAR, COL, PRC); Mun.
La Culata, Paramo La Culata, 8°45’°N, 71°33 '°W,
3300 m, 12 Nov 2000, D. Stanctk 4250 (CAR, COL,
PRC); 3100 m, D. Stanctk 4257 (CAR, COL, PRC);
8°45'N, 71°03’ W, 3100 m, 12 Nov 2000, D. Stanctk
4256 (CAR, COL, PRC, US); Dept. Libertador, NE
con respecto al Valle de la Culata., Dept. Liberta-
dor, paramo La Culata, Laguna Tapada, 3300 m,
7 Jun 1984, B. Briceno & Adamo 948 (Herbarium
Bricefio), 28 Sep 1984, B. Briceno & Adamo 1086
(MERF); Pass on the Mérida-Barinas Hwy, paramo

Festuca in South American Paramos 23

Figure 4. Festuca elviae. A. Habit. B. Inflorescence (detail). C. Ligule. D. Spikelet. E. Glumes. F. Lemma. G. Lemma with palea
and rachilla. H. Leaf blade cross-section. A-H, Briceno & Adamo 948 (PRC).

24 Festuca in South American Paramos

above Laguna Grande et Universidad de los Andes,
3500 m, 19 Nov 1971, G. Davidse et al. 3223 (MO);
Valle de los Calderones y Paramo de los Pozones,
3400 m, 19 Oct 1995, B. Briceno et al. 3345 (her-
barium Bricefio); Laguna Coromoto, mountain
forest, 8°35°41°N, 71°01°27’W, 3000-3100 m,
7 Nov 2000, D. Stancik 4178 (AAU, CAR, COL,
PRC, US, W); Entre La Mucuy e Laguna Coromoto,
3000 m, 27 Mar 1994, B. Briceno et al. 2601 (her-
barium Briceno).

4. Festuca flacca Hack. ex E.B. Alexeev, Bot.
Zhurn. (Moscow & Leningrad) 69: 1543.
1984. (Figs. 2, 5, 74A—D). Tyre: Ecuador.
Pichincha, in silv. super. pr. Tablahungi, 1887,
Sodiro 36/11 (holotype: W!; isotypes: QPLS!,
W!, US!).

Festuca subulata var. fraseriana St.- Yves, Candollea
3: 451. 1928. Type: Amerique du Sud, Equateur,
sin. loc., 1860, Fraser s.n. (holotype: G!).

Loosely tufted perennials with extravaginal
innovations. Culms 70—120(—150) cm tall, erect,
glabrous; nodes 3—7 nodes in distal half. Leaf
sheaths membranous, brown, striate, scabrous,
occassionally hairy, margins free; auricles absent;
ligules 3—3.5 mm long, membranous, apex acute;
blades 25—30 x 0.3—1.1 cm, flat, green, abaxially
scabrous. Panicles 20-35 x 5—15 cm, lax, few-
flowered, linear-oblong to ovate; branches scabrous.
Spikelets 7-8 mm long, florets 2—4 with a rudiment;
rachilla pilose; glumes 1.2—3.5 mm long, green, gla-
brous, narrowly lanceolate, apex acuminate; lower
glumes 1.2—2.4 mm long, 1-nerved; upper glumes
2.5—3.5 mm long, 3-nerved, sometimes scabrous
on dorsally; lemmas 6.5—7 mm long, 5-nerved,
membranous to coriaceous, lanceolate, green, occa-
sionally purplish-green, papillose, apex scabrous,
awned, the awn 9-12 mm long, scabrous, fine and
straight; paleas almost as long as the lemma, gla-
brous and inconspicuously scabrous on margins and
keels, apex hairy; lodicules lanceolate, acuminate;
anthers 1.1—1.4 mm long; ovary apex sparsely hairy.
Caryopses lanceolate; hilum 4/5 the length.

Leaf blade anatomy.—Cross sections with
numerous vascular bundles and without ribs above;
sclerenchyma under both abaxial and adaxial epi-
dermis, discontinuous, extending to the vascular
bundles forming girders; bulliform cells present;
abaxial epidermis (or both sides) with scattered
prickles.

Observations.—Alexeev (1984) incorrectly
cited the page of the original publication of Sodiro
(1889), where this taxon is mentioned for the first
time then later described by Alexeev as F. flacca.
Festuca flacca is morphologically similar to other
long-awned species that have short glumes, such as:
F. ulochaeta from Brazil, Colombia, and Venezuela;
F- cochabambana from Bolivia; and, F. tovariensis
from Bolivia and Peru. However, Festuca ulochaeta
has flexuous lemma awns, F tovariensis has trun-
cate ligules, and F’ cochabambana has pubescent
leaf sheaths.

Distribution and habitat.—Festuca flacca 1s
endemic to Ecuador, known from the central and
northern Andean departments (Carchi, Cotopaxi,
Imbabura and Pichincha). It occurs in mountain
Andean forests between 2900-3500 m.

Additional specimens examined. ECUADOR.
Carchi: Hacienda San Rafael, faldas occidentales
de la Cordillera Oriental, 2900-3000 m, Acosta-
Solis 21026 (US); wooded hills about 5 mi S of
Tulcan, 2500 m, A.S. Hitchcock 21042 (US). Coto-
paxi: Parque Nacional Cotopaxi, pine plantation,
00°37’S, 78°27’ W, 3400 m, S. Laegaard 54540A
(AAU, QCA), S. Laegaard 54540B (AAU, QCA).
Cotopaxi, 3550 m, E. Asplund 6359 (NY, S, US).
Imbabura: Lake Cuicocha - Islote Chica, 3150 m,
E. Asplund 7156 (NY, S, US); Laguna Mojanda,
10 km SSW of Otavalo, 00°10°N, 78°18'°W,
2900-3150 m, Sparre 13562 (S). Pichincha: Road
Pifo—Papallacta, km 15, Polylepis incana forest,
00°16’S, 78°17°W, 3530 m, 16 Apr 1992, S. Lae-
gaard 102316 (AAU, QCA, QCNE); Pifo, 3000 m,
Mille s.n. (QPLS); Volcan Pasachoa, above house of
Fundacion Natura, 00°27’S, 78°30’ W, 2900-3300
m, 16 Sep 1985, S. Laegaard 55260 (AAU, QCA,
QCNE); 3200 m, 27 Apr 1985, S. Laegaard 54164
(AAU, QCA, QCNE); 19 Sep 1985, S. Laegaard
55281 (AAU, QCA, QCNE); Mt. Pichincha — St.
Gertriadis, Sodiro s.n. (QPLS); Mt. Pichincha —
Tablahuras, Sodiro s.n. (QPLS); Mt. Pichincha,
Sodiro s.n. (US); Slope of Pichincha above Lloa,
3200 m, E. Asplund 7535 (AAU); E. Asplund 7559
(K, P, S,US); 2900-3100 m, Acosta-Solis 20054
(US); Concepcion, near Hda. Monyas, 10200 ft,
Bell 17 (S); Mun. Amaguafia, Volcan Pasochoa,
bosque andino, 3400 m, 14 Sep 2000, D. Stancik
3668 (PRC, QCA, US); Mun. Otavalo, road from
Laguna Mojanda to Cochasqui, shrubby margin of
the road, 00°04’55.1’°N, 78°17'49.8°W, 3450 m, 19
Oct 2000, D. Stancik 4104 (PRC, QCA).

25

Festuca in South American Paramos

SS

SS

.
———— SQN SSS
SSAay A
Ss

F. Lemma. G. Lemma with palea

Figure 5. Festuca flacca. A. Stylized growth form. B. Habit. C. Ligule. D. Spikelet. E. Glumes.

and rachilla. H. Leaf blade cross

Stancik 3668 (PRC).

>

-section. A—H

26 Festuca in South American Paramos

-80 -75

COLOMBIA

-80 -75

200 400 Kilometers

-70 -65 -60

Figure 6. Distribution of Festuca sodiroana (@), F- elviae (A), and F: guaramacalana ().

5. Festuca guaramacalana Stancik, Novon
14(3): 343. 2004. (Figs. 6, 7, 74E—-F, 75A &
B). Type: Venezuela. Trujillo, Mun. Bocono,
Parque Nacional Guaramacal, 09°14°15’N,
70°11°14"W, Andean mountain forest, margin
of the brook with Neurolepis sp., Cortaderia
sp., 2880 m, 29 Nov 2000, D. Stancik 4286
(holotype: PRC!; isotypes: CAR!, COL!).

Rhizomatous perennial, forming small tus-
socks with extravaginal innovations. Culms 100—
130 cm tall, erect, glabrous; nodes 3 or 4 nodes in
distal half. Leaf sheaths membranous-coriaceous,
purplish-brown, striate, fibrous at base, margins
free; auricles absent; ligules 1-1.5 mm long, mem-
branous, sometimes coriaceous, apex acute; blades
30-40 x 0.6—0.7 cm, conduplicate to involute,
green, sometimes olive-green, abaxially glabrous.
Panicles 15—20 x 2—5 cm, flexuous, pendant, elon-

gate, branched; branches finely scabrous. Spikelets
13-15 mm long, narrowly lanceolate, florets 5—7;
rachilla pilose; glumes 3.7—7 mm long, narrowly
lanceolate, purplish, membranous to coriaceous,
sparsely scabrous, apex acute; lower glumes 3.7—
4.7 mm long, 1|-nerved; upper glumes 5—7 mm long,
3-nerved; lemmas 9.5—10 mm long, membranous
to coriaceous, lanceolate, 5-nerved, purplish, papil-
lose, apex entire, mucronate or shortly awned, the
awn 0.5—1 mm long; callus sparsely pilose; paleas
finely pilose, almost as long as the lemma; lodicules
ovate, two-dentate; anthers 2.5—2.8 mm long; ovary
apex glabrous. Caryopses lanceolate; hilum 4/5 as
long as the grain.

Leaf blade anatomy.—Cross-sections with
about 11 vascular bundles and 5 ribs above; scle-
renchyma under both abaxial and adaxial epidermis
discontinuous and extending to some vascular
bundles forming girders; bulliform cells absent;

Festuca in South American Paramos 27

H
|
y)
tN!
i
IIl\\
\f i 7
y,
CIN Ht
H
\ \ |
| \
f WAN
Aa |
\)} |
H
WWW I
\ A
{ ) \ |
i} i ! t
| iN \ i
|
4 h yA
\ ,
\ Nii} I Y Qi
I\ Hl if it
\ |
\\\y Mi Vix
Y H iy
f . \) it]
Vr |
Ky i}
if) 5
“Alli \
X\ i
\
NU j
VAL TA
\
| | NV
I\ { |
| i A
\\ ]
\

any
il
|
\)
\
\\

Figure 7. Festuca guaramacalana. A. Growing form. B. Habit. C. Ligule. D. Spikelet. E. Glumes. F. Lemma. G. Lemma with
palea and rachilla. H. Leaf blade cross-section. A—H, Stancik 4286 (PRC).

28 Festuca in South American Paramos

adaxial epidermis with scattered microhairs, ca.
0.09 mm long.

Observations.—Festuca guaramacalana
belongs to Festuca sect. Subulatae and among the
South American members of this section, F) guara-
macalana has the largest spikelets and longest
glumes (Stancik & Peterson 2002). The lemmatal
awns of F! guaramacalana are short (0.5—1 mm)
and this is in contrast with most other members of
sect. Subulatae, although F: sodiroana typically has
acute lemmas. Morphologically, F’ coromotensis
appears to be most similar to F’ guaramacalana
but the former has shorter ligules (0.5—0.7 mm
long), ligules with truncate apices, shorter spikelets
(9-11 mm long) with 3 to 4 florets, and shorter
lemmas (7.5—8.5 mm long).

Distribution and habitat.—Festuca guara-
macalana is endemic to Venezuela and is known
only from Parque Nacional Guaramacal where it
occurs near small brooks with Neurolepis sp. and
Cortaderia sp. between 2600-2880 m. This species
is a narrow endemic and is vulnerable to extinction
due to loss of habitat.

Additional specimens examined. VENEZU-
ELA. Trujillo: Mun. Bocono. Parque Nacio-
nal Guaramacal. Paramo El Pumar. 2870 m,
2004, M. Ramirez & N. Cuello 3523, 3524, 3528
(PORT).

6. Festuca sodiroana Hack. ex E.B. Alexeev, Bot.
Zhurn. (Moscow & Leningrad) 69: 1545.
1984. (Figs. 6, 8, 75C-—F). Type: Ecuador.
Pichincha, silv. super. et pasq., 1884, Sodiro
36/6 (holotype: W!; isotypes: MO-923640!,
PRC ex QPLS!, QPLS!, US!).

Rhizomatous perennials with extravaginal
innovations. Culms (40—)50—120(—150) cm tall,
erect, glabrous; nodes 2 or 3(-4) in distal half. Leaf
sheaths membranous, brown, striate, pilose at base,
margins free; auricles absent; ligules 0.2—1.1 mm
long membranous to coriaceous, apex truncate,
ciliate; blades 15—25 x 0.4—0.7 cm, flat, green, abax-
ially glabrous. Panicles 15—20 x 1—2 cm, lax, con-
tracted and something nutant, branched; branches
scabrous. Spikelets 7.5—9(—10) mm long, florets 4
or 5(—6); rachilla pilose; glumes 1.3—2.9(-3.5) mm
long, lanceolate, green, membranous to coriaceous,
glabrous, apex acute; lower glumes 1.3—1.8 mm long,
l-nerved; upper glumes 2.2—2.9(—3.5) mm long,
3-nerved; lemmas 5—6(—6.5) mm long, 5-nerved,
membranous to coriaceous, lanceolate, green,

awnless, apex acute, entire, muticous, scabrous;
paleas almost as long as the lemma, glabrous or
inconspicuously scabrous, apex hairy; lodicules
lanceolate, acuminate; anthers 0.8—1.2 mm long;
ovary apex glabrous. Caryopses lanceolate; hilum
4/5 of total length.

Leaf blade anatomy.—Cross-sections with
numerous vascular bundles, without ribs above;
sclerenchyma under both abaxial and adaxial epi-
dermis, discontinuous, extending to the vascular
bundles; bulliform cells present; abaxial epidermis
with scattered prickles, adaxial epidermis without
hairs or with scattered hairs.

Observations.—Specimens that belong to this
species were originally determined by Hackel and
Sodiro as either F. sodiroana or F. pichinchae, but
these names were not validly published. Alexeev
recognized that these specimens belonged to the
same species and validated the name F. sodiroana.
Festuca sodiroana differs from others in sect.
Subulatae by having muticous lemmas.

Distribution and habitat.—Known from
southern Ecuador to northern Colombia where it
is found in all three cordilleras. It occurs in forest
clearings, margins of brooks in Andean mountain
forests, roads banks, and trails at an altitude of
2600-3800 m. Festuca sodiroana is known to
occur in vegetation communities: Neurolepido
aristatae—Oreopanacion nitidii (Cleef et al. 1983)
and Chusquea scandentis—Weinmannion roilottii
(Cleef et al. 1983).

Additional specimens examined. COLOM-
BIA. Antioquia: Mun. Urrao, Paramo Frontino,
Llano Grande, 3520 m, 10 Sep 1986, Roldan et al.
360 (HUA); Campanas, La Laguna, 3500-3800 m,
3 Mar 1989, MacDougal et al. 4504 (HUA, MO).
Cauca: Macizo Colombiano, Paramo de Las Papas,
entre Boqueron y La Hoya, 2910 m, 11 Sep 1958,
Idrobo et al. 3026 (COL); Parque Nacional Puracé,
termales de San Juan, 3100-3300 m, 6 Apr 1985,
Wood 4792 (COL, K); Paramo de Moras, between
Mosoco y Pitayo, 3000-3500 m, Feb 1906, Pittier
1511 (US). Mt. Puracé, 3100-3300 m, 16 Jun 1922,
Killip 6709 (US). Cundinamarca: Paramo de
Tablazo, 3200 m, 8 Apr 1984, Wood 4347 (COL,
FMB, K). Narino: El Encano, road from Pasto to
Virgen, vereda Caltapamba, 3100 m, 23 Mar 1999,
D. Stancik 2992 (COL, PRC, PSO, US); D. Stancik
2991 (COL, PRC, PSO); El Encano, road to village
Colon, km 4, 2650 m, 13 Mar 1999, D. Stancik
2866 (COL, PRC, PSO); km 6, 2900 m, 13 Mar
1999, D. Stancik 2858 (COL, PRC, PSO); Mun.

29

Festuca in South American Paramos

Figure 8. Festuca sodiroana. A. Stylized growth form. B. Habit. C. Ligule. D. Spikelet. E. Glumes. F. Lemma. G. Lemma with

palea and rachilla. H. Leaf blade cross-section. A-H, Stancik 2992 (PRC).

30 Festuca in South American Paramos

Cumbal, vereda Las Huertas, 3600 m, 9 Mar 1999,
D. Stanctk 2754 (COL, PRC, PSO); Cumbal, lake-
shore, 4000 m, 24 Mar 1941, Sneider 430 (NY);
Mun. Pasto, paramo Puerto Frio, between villages
Las Almas and Alisales, 2900 m,14 Mar 1999, D.
Stancik 2870 (COL, PRC, PSO); Mun. Tuquerres,
Volcan Azufral, road from vereda San Roque Alto,
3500 m, 9 Mar 1999, D. Stancik 2772 (COL, PRC,
PSO); 2650 m, 9 Mar 1999, D. Stancik 2776 (COL,
PRC, PSO); Vereda El Carmelo and La Florida,
Volcan Dona Juana, 2900 m, 18 Mar 1999, D.
Stancik 2902 (COL, PRC, PSO); Paramo de Bord-
oncillo, Mun. Santiago, vereda San Antonio de
Bellavista, 1°11°N, 77°06’ W, 3200-3400 m, 18
Mar 1964, Alberto et al. 144 (COL); Mun. Guachu-
cal, paramo de Infernillo, 3200 m, 5 Mar 1999, D.
Stancik 2632 (COL, PRC). Norte de Santander:
between Pamplona and Mutiscua, 2700 m, 1 Apr
1984, Wood 4326 (COL, FMB, K). Putumayo:
Paramo de Tabano, 15 May 1935, Archer 3402
(US). Santander: Valle California, 2800 m, 14 Sep
1985, Wood 5066 (FMB, K). Tolima: Mun. Ibague,
Nevado del Tolima, 04°37.4’N, 75°19.8°W, 3400
m, 8 Jun 2000, D. Stancik 3595 (COL, PRC, PSO);
3400 m, 16 Dec 1984, Wood 4648 (COL, FMB, K);
W slope of Paramo Rico, 3200 m, 15-19 Jan 1927,
Killip & Smith 17861 (NY). sin. loc.: Cerro
Nevada, Bos. La Pefia, 2900 m, Lindig 1116 (K, P);
Lindig 1117 (P); Mutis 5544 (US). ECUADOR.
Azuay: Road Cuenca — Saraguro, km 6, S of
Cumbe, 03°04’S, 79°00'°W, 3150 m, S. Laegaard
105132 (AAU); Parque Nacional Cajas—Laguna
Llaviuco, 3150 m, 22 Apr 1990, P.M. Peterson
8872, CR. Annable & M.E. Poston (K, MO, QCA,
QCNE, US); Ganadel, 3250 m, E. Asplund 17846
(S); between Cuenca and Hiugra, 2700-3000 m,
A.S. Hitchcock 21682 (US); road Sigsig—Guada-
quiza, 02°09’S, 78°43’ W, 3300 m, 29 May 1992, S.
Laegaard et al. 103032 (AAU, QCA, QCNE);
Mun. Cuenca, Parque Nacional Cajas, Laguna
Llaviucu, 02°50°30"S, 79°08’°45’ W, 3100-3150 m,
3 Aug 1999; Palice | (@R@); Palices (PR).
Palice 4 (PRC). Bolivar: Road Guaranda—Pueblo
Viejo, km 21.6, 01°35’S, 79°09’ W, 2500 m, 6 Mar
1988, S. Laegaard & S.A. Renvoize 70580 (AAU);
km 20.1, 01°35’S, 79°05’ W, 2700 m, S. Laegaard
& S.A. Renvoize 70577 (AAU, K, QCA, QCNE).
Cafar: Road Cafiar—Biblian, Fuganillas, 02°36’S,
7T8°54°W, S. Laegaard 52754 (AAU, QCA, QCNE);
Tipococha—Hacienda Shical, 3000 m, Acosta-Solis
16963 (US); Acosta-Solis 16974 (US). Carchi: El
Angel, Hacienda la Esperanza, 00°39’N, 77°54 'W,

3300 m, S. Laegaard 53118 (AAU, QCA); El
Voladero, 00°38'°N, 67°53'°W, 3400-3800 m,
Davalos 22 (US); wooded hills about 5 mi S of
Tulcan, 2500 m, A.S. Hitchcock 21094 (US); Road
Las Juntas—El Angel, 00°46’N, 77°46’ W, 3180 m,
11 Mar 1992, S. Laegaard 101711 (AAU, QCA,
QCNE); La Rinconada, 3200 m, EF. Asplund 7195
(F, S); 3000 m, A.S. Hitchcock 20798 (NY, US).
Cotopaxi: Road Pilalo—Zumbagua, 15 km above
Pilalo, 00°59’S, 78°58°W, 3350 m, Holm-Nielsen
24604 (AAU); km 9, 0°59°S, 78°57’ W, 3150 m, 7
Apr 1992, S. Laegaard 102227 (AAU, QCA,
QCNE); Road Pilalo—Latacunga, 00°57’S, 78°58’ W,
3400 m, Holm-Nielsen 1482 (AAU, MO, S); Road
Quevedo-—Latacunga, 3600 m, Harling et al. 8904
(GB); Rio Chalupas, 00°50’S, 78°21°W, 3700 m,
S. Laegaard 101781 (AAU, QCA, QCNE); Mun.
Lasso, Volcan Cotopaxi, margin of the forest at the
road to National Park entrance, 00°39’6’S,
78°30°55"W, 3530 m, 28 Sep 2000, D. Stancik
3885, 3886 (PRC, QCA). Chimborazo: Road
Chunchi—Zhub, km 22, 02°22’S, 78°57’ W, 2780 m,
27 May 1992, S. Laegaard et al. 103010 (AAU,
QCA); Urbina towards Mt. Chimborazo, 3750 m,
E. Asplund 7878 (S); Pallatanga—comunidad Jesus
del Gran Poder, 01°58’S, 78°56’ W, 2800-3200 m,
Clark et al. 1372 (QCNE). Imbabura: Road
Otavalo—Selva Alegre, km 25.2, 00°16’S, 78°24’ W,
3300 m, S. Laegaard et al. 70813 (AAU, K, QCA);
Paramo de Angochagua, 2900-3600 m, Acosta-
Solis 18839 (US); Cayambe—Laguna San Marcos,
11200 ft, Cazalet & Pennington 5417 (K, NY, US);
Mun. Urcuqui, road to Cerro Yanaurcu, 00°26'28'°N,
78°15'24'"W, 4100 m, 15 Oct 2000, D. Stancik 4095
(PRC, QCA); Mun. Otavalo, road from Laguna
Mojanda to Cochasqui, 00°04’55’N, 78°17'50"W,
3450 m, 19 Oct 2000, D. Stancik 4105 (PRC,
QCA); D. Stanctk 4113 (PRC, QCA). Loja: Road
to Fierra Urcu, 03°33’S, 79’15’W, 3100 m, S. Lae-
gaard et al. 18855 (AAU, LOJA, QCA, QCNE);
Road Saraguro—Yacuambi, 03°34’S, 79°06’ W, 3050
m, S. Laegaard 20688 (AAU, LOJA); between San
Lucas and Ona, 2200-3100 m, A.S. Hitchcock
21512 (US); Udo de Sabanilla, 04°27°S, 79°09'W,
2700 m, 1 Sep 1998, S. Laegaard et al. 19095
(LOJA); Mun. Saraguro, road to Fierra Urcu,
03°42°40"S, 79°18°12"W, 3400-3450 m, 24 Aug
2000, D. Stancik 3771, 3781 (PRC, QCA); 3000—
3100 m, D. Stanctk 3792 (PRC, QCA). Morona-
Santiago: road Gualaceo-Limon, km 12 of pass,
03°01°S, 78°37°W, 2590 m, S. Laegaard et al.
103079 (AAU); Mun. Alao, way from Alao to

Festuca in South American Paramos EY

Parque Nacional Sangay, 3500-3700 m, 23 Jul
1999, D. Stancitk 3292 (PRC, QCA, W); Parque
Nacional Sangay, patches of the forest above the
Quebrada Tablamitza, 3600 m, 22 Jul 1999, D.
Stancik 3352 (PRC, QCA). Napo: km 45 on road
Salcedo—Napo, 6 km NE, 00°56’S, 78°23 'W,
3600 m, S. Laegaard 53372 (AAU, QCA, QCNE).
Pichincha: Near Quito, Jun 1922, Harteman 39
(US); Paramo Guamani, 00°15’S, 78°12’W, 3700
m, S. Laegaard 105069 (AAU); 3530 m, S. Lae-
gaard 102318 (AAU, QCA, QCNE); Volcan Pich-
incha, 3250 m, Asplund 6155 (AAU, MO); above
Lloa, 3200 m, E. Asplund 7537 (F, S); 3250 m, E.
Asplund 6155 (F, MO, S, US); Mt. Pichincha,
Sodiro s.n. (NY, MO); 3400 m, Sodiro s.n. (US);
Milles et al. 281 (US); Sodiro s.n. (US); 3600 m,
Sodiro 282 (US); 3300 m, E. Asplund 16163 (S);
At base of Mt. Pichincha—La Carolina, Sodiro s.n.
(S); road Chillogallo—Chiriboga, km 3 E of San
Juan, 3100 m, Sparre 16950 (S); Sparre 16920 (S);
San Juan towards Quito, 3400 m, E. Asplund 16094
(S); Tambillo, 2700 m, E. Asplund 6215 (AAU, F,
MO, S, US); road Pifo—Pintag, 00°19’S, 78°17°W,
3100 m, S. Laegaard 102293 (AAU, QCA, QCNE);
road Santo Domingo—Quito, between Saloya and
Chiriboga, 2800 m, Harling et al. 10415 (GB);
Volcan Corazon, 11000 ft, Prescott 913 (NY); road
km 13.5 ESE of Machachi, and 10 km NE towards
Sangolqui, 3350 m, 25 May 1990, PM. Peterson
9313 & E.J. Judziewicz (K, MO, QCA, QCNE,
US); road Quito—Nono, 00°06’S, 78°31°W, 3300 m,
17 Jun 1984, S. Laegaard 52285 (AAU, QCA,
QCNE); Nono, 2600 m, E. Asplund 7462 (S); Vol-
can Pasochoa, 00°21’S, 78°29’ W, 3100-3400 m, 23
Feb 1992, S. Laegaard 101406 (AAU, QCA);
00°27°S, 78°30°W, 3200 m, S. Laegaard 55282
(AAU, QCA, QCNE); Sodiro s.n. (QPLS); High-
way Aloag—St. Domingo, 2900 m, Sparre 15076
(S); Mun. Pifo, Paramo Guamani, Polylepis forest,
00°19°S, 78°15’ W, 3700 m, 19 Jun 1999, D. Stancik
3010 (PRC, QCA); Mun. Amaguajna, volcan Pasa-
choa, 3400 m, 14 Sep 2000, D. Stancik 3667 (PRC,
QCA); Mun. San Antonio, Res. Geobotanica Pulu-
lahua, 00°02’20’N, 78°29°33’W, bottom of the
crater, 2700 m, 13 Sep 2000, D. Stancik 3656
(PRC, QCA). Tungurahua: sin loc., Spruce 5938
(K, W). Mun. Bafios, Volcan Tungurahua, on way
to refugium, E side of the volcan, 3300 m, 29 Jul
1999, Palice 9 (PRC); Mun. Pillaro, Las Llangana-
tis, 01° 09°43”S, 78°15’9" W, 3600 m, 28 Sep 2000,
D. Stancik 3985 (PRC, QCA). Zamora—Chinchipe:
road Loja—Zamora, ca. 2-6 km E of pass, 03°59’S,

79°09°W, 2750 m, S. Laegaard 18748 (AAU,
LOJA); 2600 m, S. Laegaard 18727 (AAU, LOJA);
Road Loja—Zamora, | km E of pass, 04°00’S,
79°09'W, 2700 m, S. Laegaard 18483 (AAU,
LOJA, QCA, QCNE).

7. Festuca tovariensis Stancik & P.M. Peterson, Sida
20(1): 24. 2002. (Figs. 9, 11). Type: Peru. Huan-
cavelica, Prov. Tayacaja, Chuspi—Hda. Tocas,
entre Colchabamba y Paucarbamba, monte bajo,
2800 m, 22 Apr 1954, O. Tovar Serpa 2057
(holotype: US-2181286!; isotype: USM!).

Loosely tufted perennials with extravaginal
innovations. Culms 70—90 cm tall, erect, glabrous;
nodes 2-4 nodes in basal half. Leaf sheaths mem-
branous, brown, margins free; auricles absent;
ligules 1-2 mm long, membranous, margins ciliate,
apex truncate; blades 8-15 cm long, 1.5-4.5 mm
wide, flat, green, abaxially scabrous with ribs on
abaxial surface. Panicles 15—20 x 7—10 cm, open,
pendant; branches scabrous. Spikelets 7.5—9.5 mm
long, florets 3; rachillas 1.1—1.4 mm long, densely
pilose; glumes 1.5—4.5(—5) mm long, narrowly
lanceolate, coriaceous, purplish, glabrous, apex
acute sometimes scabrous; lower glumes 1.5—1.8
mm long, 1-nerved; upper glumes 3.5—4.5(—5)
mm long, 1—3-nerved; lemmas 5.5—6.5 mm long,
lanceolate, coriaceous to membranous, inconspicu-
ously 5-nerved, purplish-green, scabrous, apex
entire, awned, the awn 5—7 mm long, terminal, sca-
brous, straight; paleas as long as the lemma, keels
scabrous, apex hairy; lodicules ca. 0.8 mm long,
lanceolate, acuminate; anthers 1.5—1.6 mm long;
ovary apex sparsely hairy. Caryopses lanceolate;
hilum 2/5—1/2 of total length.

Observations.— Festuca tovariensis 1s mor-
phologically similar to other long-awned species
that have short glumes, such as: F. ulochaeta from
Brazil, Colombia, and Venezuela; F. cochabambana
from Bolivia; and F! flacca from Ecuador.

Distribution and habitat.—This species is
known only from the Andean forest zone of north-
ern and central Peru between 2500-3850 m.

Additional specimens examined. PERU.
Ancash: Prov. Yungay, Parque Nacional Huascaran,
Llanganuco sector, Maria Josefa trail between
Chinancocha and Pucayacu, 09°05’S, 77°39’ W,
3700-3850 m, 5 Jul 1985, Smith 10561 (MO).
Ayacucho: Prov. Huanta/La Mar, Tambo, Wolken-
Nebel-Buschwald, 37 km to Ayna, 3250 m, 23 Mar
1977, Ellenberg 7024 (USM). Cajamarca. Prov.

32 Festuca in South American Paramos

Figure 9. Festuca tovariensis. A. Stylized growth form. B. Habit. C. Ligule. D. Spikelet. E. Glumes. F. Lemma. G. Lemma with
palea and rachilla. H. Caryopsis. A-H, Vega et al. 1544 (F).

Festuca in South American Paramos 33

Cajamarca, Yumagual, entre San Juan y El Gavilan-
Gavilan, ladera con arbustos, 2500 m, 6 Oct 1975,
Vega et al. 1544 (F); Prov. Contumaza, alrededores
de Guzmango, ladera, 2600 m, 25 Jul 1992, Saga-
steguii 14785 (F); Carretera a Yumagual, 2500 m,
26 Jun 1966, Vega 249 (USM).

8. Festuca ulochaeta Nees ex Steud., Syn. Pl. Glu-
mac. 1:305. 1854. (Figs. 10, 11, 76A—D). Tye:
Brazil. Sellow s.n. (isotypes: B!, K!)

Festuca leptothrix Trin. ex Doll, Fl. Bras. 2(3):
115. 1878. Type: Brazil. Sao Paulo, G.H. von
Langsdorffs.n. (holotype: LE-TRIN-2818.01!;
isotypes: K!, US-91399 fragm!).

Vulpia ulochaeta Nees ex Doll, Fl. Bras. 2(3): 115.
1878, nom. inval. Type: Brazil. Sellow s.n.
(holotype: B!, BAA-3489 fragm ex B).

Locsely tufted perennials with extravaginal
innovations. Culms 60-120 cm tall, erect, gla-
brous; nodes 24 nodes. Leaf sheaths membranous,
brown, striate, glabrous; auricles absent; ligules
0.5—1(-2) mm long, membranous, apex truncate;
blades 15-30 x 0.5—1.1 cm, flats, green, abaxially
scabrous. Panicles 15—25(—35) x 15—25 cm, flexu-
ous, pendant, ovate; branches scabrous. Spikelets
9-12 mm long, florets 3—5; rachilla shortly hairy;
glumes 2.5—4.5(—6) mm long, membranous to
coriaceous, narrowly lanceolate, green, apex acute,
upper third scabrous; lower glumes 2.5—3.5(-4) mm
long, 1-nerved; upper glumes 3.5—4.5(—6) mm long,
3-nerved; lemmas 6—8(—9) mm long, 5-nerved,
membranous to coriaceous, lanceolate, green, papil-
lose, apex scabrous, awned, the awn 7-15 mm long,
scabrous, flexuous; paleas almost as long as the
lemma, glabrous, margins and upper third scabrous;
lodicules lanceolate; anthers 1.1—1.5(—2) mm long,
ovary apex sparsely hairy. Caryopses lanceolate;
hilum 3/4 of total length.

Leaf blade anatomy.—Cross-sections with
numerous vascular bundles, without ribs above;
sclerenchyma under both abaxial and adaxial epi-
dermis, discontinuous and extending to thevascular
bundles; bulliform cells absent; abaxial epidermis
with scattered prickles.

Observations.—Festuca ulochaeta has spike-
lets with short glumes and long-awned lemmas,
characteristics shared with F’ flacca, F. cuzcoensis,
and F? cochabambana. Festuca ulochaeta differs
from these other species by having lemmas with
extremely long (7—15 mm) and distinctly flexuous
(versus straight) awns.

Distribution and habitat.—Festuca ulochaeta
is known from the humid forests of SE Brazil and
NE Argentina between 700-2200 m. In Colombia
and Venezuela this species occurs in clearings, mar-
gins of the streams, and roadsides in Andean forests
(Cordillera Oriental and Aragua, Mérida) between
2600-3100 m. Festuca ulochaeta is reported from
Costa Rica for the first time here.

Additional specimens examined. ARGEN-
TINA. Misiones: Dpto. Gral. Manuel Belgrano,
Santo Andresito, 26°12’S, 53°40’ W, 720 m, 15 Feb
1996, O. Morrone & A.M. Cialdella 854 (CTES,
MO). Salta: Dpto. Santa Victoria, camino de Toldos
a Lipeo, a 15 km de Toldos, 1650 m, 11 Nov 1974,
A.M. Ttirpe 2932 (W); Arroyo Latas, 20 Feb 1924,
L. Parodi 5673 (US). BRAZIL. Minas Gerais:
Xanxere, Pinheiral, 9 km E of Xanxere, 600-800 m,
26 Feb 1957, Smith & Klein 11836 (B, R, US); Serra
de Caparao, 2100—2220 m, 4 May 1925, A. Chase
9659 (W); 2100 m, 4 May 1925, A. Chase 9672
(US); 2100-2200 m, A. Chase 9673 (US); Monte
Verde, Sep 1997, Wagner 5010 (ICN); Vila Mon-
teverde, Pico da Pedra Salada, Burman 880 (SP).
Parana: ca. 85 km of Guarapuava, 800-1050 m,
6 Mar 1967, Linderman et al. 4658 (K, W, US);
Ypiranga, 9 Feb 1904, Dusen 3624 (R); Dusen 3515
(R); Piraquara—Estrada a Monte Algre, Apr 1950,
G. Hatschbach 1912 (BAA); Tres Barras, 27 Jan
1916, Dusen 17561 (BAA); Iraty, in silvula subu-
liginosa, 26 Feb 1909, Dusen 7808 (US); Curitiba,
woods along stream, Estacao Experimental, 13 Feb
1946, J.R. Swallen 8540 (US); Curitiba, colonia
Muricy-—S. José dos Pinhais, na beira da Estrada, |
Mar 1965, Scito & Kuniyoshi 1315 (K); Curitiba,
Orla do brejo, 15 Jan 1975, Maguire 3487 (K); Orla
Guarapuava, fazenda Capao Redondo, 20—23 Mar
1946, J.R. Swallen 8859 (US); Mun. Palmas, 1100
m, 16 Feb 1958, G. Hatschbach 4715 (US); Curitiba,
Parque Barigui, area degradada, proxima a orla da
foresta con Araucaria, 25°22°S, 49°13°W, 2 Apr
1997, Kozera & Izernhagen 476 (NY); Piraguara, ad
marginam, | Jul 1909, Dusen 7783 (K, NY); Mun.
Rio Branco do Sul, Dombrowski 2432 (ICN, K);
Sao Mateus do Sul, Rio Potinga, 760 m, do interior
da mata, 9 Feb 1966, G. Hatschbach 13820 (K).
Rio de Janeiro: Serra de Itatiaia, 1960 m, 16 Jan
1925, A. Chase 8276 (RB, US); Parque Nacional de
Itatiaia, entre Agulhas Negras e Abrigo Massena,
campo alto 11—12 Feb 1985, Burman 905 (SP). Rio
Grande do Sul: Highway BR-116 to Lajes, 1000
m, 10 Mar 1976, G. Davidse et al. 11074 (K, SP,
VEN); Entre J. Kroeff et Roncinka, 18 Mar 1964,

34 Festuca in South American Paramos

SS

A

1mm

Figure 10. Festuca ulochaeta. A. Stylized growth form. B. Habit. C. Inflorescence. D. Ligule. E. Spikelet. F. Glumes. G. Lemma.
H. Lemma with palea and rachilla. I. Leaf blade cross-section. A—I, Stancik 4179 (COL).

Festuca in South American Paramos 35

Brescia & Marches 4212 (K, P); Farroupilha, in
araucarieto, 700 m, 15 Feb 1957, Camargo 59992
(B); Vila Oliva, Caxias do Sul, 24 Feb 1954, Rambo
54999 (B, BAA); Rambo 54989 (B, US); Mun.
Cruz Alta, campos de Cruz Alta, 500 m, Feb 1906,
Jurgens 6258 (W, US); Serra de Caparas, Espirito
Santo, 27 Nov 1929, A. Chase 10101 (W, US); Sao
Leopoldo—Boni Jesus, Dutra 331 (ICN, R); Dutra
402 (R, US); Taimbe, Sao Francisco de Paula, 18
Dec 1950, Rambo 49315 (BAA); Sao Francisco de
Paula—Tainhas, Dec 1953, Barreto 1871 (BAA);
Mun. de Caxias do Sul, Ana Rech, 780 m, 18 Mar
2000, Scur 656 (US); Taimbesinho, in araucarieto
aperto, 20 Feb 1953, Rambo 54011 (US); Esmer-
alda, estaccao ecologica De Aracuri, 2 Dec 1979,
Winge et al. 1362 (ICN); Winge 1272 (ICN); km
60 W of Passo Fundo along highway BR 285 to
Vacaria, intersection with Rio Igeiro, 11 Mar 1976,
G. Davidse & D’Arcy 11166 (SP); Sao Francisco da
Paula to Eletra, Araucaria woodland 31 Jan 1965,

Clayton 4491 (K, SP); Cambara do Sul, 29°00’S,
50°00°W, 27 Jan 1948, Rambo 36446 (K); Cam-
bara do Sul, Itaimbezinho, Bela Vista, 1 Dec 1981,
Wagner 944 (ICN); Itaimbezinho, Bela Vista, Valls
2386 (ICN); Wagner et al. 298 (ICN); 1 Dec 1981,
Wagner 946 (ICN); Wagner 947 (ICN); Valls 56
(ICN); Lagoa dos Patos, Saco de Tapes, Dec 1980,
Goergem & Wagner 50197 (ICN); Mun. de Caxias
do Sol-Ana Rech 780 m, 18 Mar 2000, Scur 656
(MO). Santa Catarina: Campo Ere, 29 Feb 1964,
Castellanios 24729 (COL, K); Cacador-Curitibafios,
33 km SE of Cagador on the road to Lebon Regis (47
km), 700-900 m, 16 Mar 1957, Smith & Klein 12188
(R, US); Serra de Boa Vista, Sao José, 3 Mar 1961,
Reitz & Klein 10832 (B, US); 10 Nov 1960, Reitz
& Klein 10399 (B, US); Palmares, Campos Novos,
900 m, 11 Apr 1963, Reitz & Klein 14611 (B, US);
Mun. Cacador, km W of Cacador, ruderal, 900-1000
m, 6 Feb 1957, Smith & Klein 10887 (R); Bog. 8
km N of Cagador, 950-1100 m, 7 Feb 1957, Smith

50 -40 -30

10

-80 -70 60

600 Kilometers
+ + -30

Figure 11. Distribution of Festuca ulochaeta (@), F- tovariensis (&), and F- caldasii (m).

36 Festuca in South American Paramos

& Klein 10956 (R, US); Mun. Joacaba, campos of
palmas, 55 km W of Cacador, 1000-1200 m, 18 Feb
1957, Smith & Klein 11384 (R, US); Mun. Chapeco,
Fazenda Campo Sao Vicente, 24 km W of Campo
Ere, 900-1000 m, 20 Feb 1957, Smith & Klein 11611
(R, US); Mun. Lajes, E of Capao Alto, 900—1000 m,
12 Feb 1957, Smith & Klein 11335 (R, US); Mun.
Campo Alegre, lower fazenda of ernesto Scheide,
900 m, 10 Mar 1957, Smith & Klein 12024 (R,
US); Morro Juco Prudente, steep slopes of wooded
arroyo, | Jan 1946, /.R. Swallen 8033 (US); Coxilha
Rica, open woods, 6 Jan 1946, J.R. Swallen 8168
(US); Picadas, km 181 da ERF, Papanduva, 1000
m, 26 Feb 1962, Reitz & Klein 12509 (US); Sao
Joaquim, Urupema, matinha, 1200 m, 24 Dec 1962,
Reitz & Klein 14585 (US); Rio Cagador, 21 Jan
1946, J.R. Swallen 8231 (MO, NY, US); km 21, NE
of Santa Cecilia, along Highway BR-116 to Curitiba,
open grassland with rocks outcrops, marshy area and
patches of trees, 1250 m, 3 Oct 1976, G. Davidse
et al. 11082 (NY, SP); km 18, NE of Santa Cecilia,
along Highway BR-116 to Curitiba, grassy roadside,
1200 m, 3 Oct 1976, G. Davidse et al. 11087 (NY,
SP); Campo Dos Padres, Bom Retiro, 1900 m, 20
Dec 1953, Reitz 2611 (NY). Sao Paulo: 1816-1821,
Saint-Hilaire 312 (P); Saint-Hilaire 304 (P); Sao
Paulo prope Apiahy, Puiggari s.n. (W); Serra de
Bocaina, Apr 1951, Segadas 2827 (R); Campos do
Jordao, subida para o pico do Itapeva, Kuhlmann
2242 (SP). COLOMBIA. Cundinamarca: Paramo
del Tablazo, W of Subachoque, 3100 m, Wood 3842
(COL, FMB, K); between La Calera—Choachi, ca. 1
km above Mundo Nuevo, 2600 m, 1983, Wood 5104
(COL, K). sin. loc., Lindig 1862 (P). COSTA RICA.
Limon: Canton de Talamanca, Sabanas de Durika,
| km aguas abajo de la confluencia de los Rio Uk y
Rio Kuk, 83°19°30°W, 09°25 '20°N, 2250 m, 20 Oct
1989, Herrera 3730 (K, MO). VENEZUELA. Ara-
gua: Colonia Tovar, trayecto El Lagunazo - Colonia
Tovar, 2100 m, Feb 1953, Aristeguieta 763 (VEN).
Mérida: Mun. Tabay, Laguna Coromoto, moun-
tain forest, 3000-3100 m, 7 Nov 2000, D. Stancik
4177 (AAU, CAR, COL, PRC, W); 2800-3000 m,
D. Stancik 4179 (CAR, COL, PRC, US); Laguna La
Coromoto, 3200-3300 m, 3 Jul 1987, B. Briceno &
Adamo 2011 (Herbarium Bricefio).

9. Festuca caldasii (Kunth) Kunth, Revis. Gramin.
1: 132. 1835. (Figs. 11, 12, 76K & F,77A & B).
Bromus caldasii Kunth Nov. Gen. Sp. (quarto
ed.) 1: 151. 1816. Schedonorus caldasii (Kunth)

Roem. & Schult., Syst. Veg. 2: 709. 1819.

Festuca quadridentata var. caldasii (Kunth)

St.-Yves, Candollea 3: 266. 1927. Festuca

quitensis Willd. ex Kunth, Enum. Pl. 1: 407.

1833, nom. inval. Type: Ecuador. crescit locis

altis regni Quitensis, prope Chillo, Humboldt &

Bonpland s.n. (lectotype: P!, designated here;

isolectotypes: B!, US-865519 fragm. ex P!).
Bromus procerus Kunth, Nov. Gen. Sp. (quarto ed.)

1: 150. 1816. Schedonorus procerus (Kunth)

Roem. & Schult., Syst. Veg. 2:708. 1819.

Bromus procerus Humb. ex Spreng., Syst. Veg.

1:357. 1825. nom. illegit.

Festuca quadridentata subsp. eminens (Kunth) St.-
Yves, Candollea 3: 266. 1927. Festuca eminens
Kunth, Révis. Gramin. 1: 132. 1829. Type:
Ecuador. Pichincha, 2410 m, Feb, Humboldt
& Bonpland 2296 (holotype: P!; isotypes:
B!, BAA-1206 fragm. ex B, P!, US-2875405
fragm. ex P!).

Rhizomatous perennials with extravaginal
innovations. Culms 70-150 cm tall, erect, sca-
brous; nodes 3 or 4 in distal half. Leaf sheaths
coriaceous, scabrous, chestnut-brown to brown,
striate, fibrous near base, margins free; ligules
2.5—3 mm long, membranous, abaxially hairy,
apex acute, lacerate; blades 20-35 x 0.3-0.9 cm,
flat, green, scabrous, with prickles on both abaxial
and adaxial epidermis. Panicles 13-17 x 5—8 cm,
ovate; branches erect or sometimes spreading,
scabrous. Spikelets 15—17 mm long, oval, florets
5—7(—8); rachilla pilose; glumes 3.5—6.5 mm long,
membranous, lanceolate, green with transparent
membranous margins; lower glumes 3.5—4(—-5) mm
long, 1-nerved; upper glumes 4.5—6.5 mm long,
3-nerved; lemmas 10—14 mm long, lanceolate,
membranous to coriaceous, 5-nerved, green, sca-
brous or shortly densely hairy, apex entire, awned,
the awn 1—3 mm long; callus glabrous; paleas 4/5
as long as the lemma, membranous, scabrous; lodi-
cules 0.8—1 mm long, oblong-lanceolate; anthers
(2.8-)3.5—4.5 mm long; ovary apex glabrous.
Caryopses not seen.

Leaf blade anatomy.—Cross-sections with
many vascular bundles, with just small ribs; scle-
renchyma under both abaxial and adaxial epider-
mis, discontinuous; bulliform cells present, few;
epidermis without hairs.

Observations.—Festuca caldasii 1s morpho-
logically similar to F’ woodii, a species that occurs
in northern Colombia. However, /. woodii has

37

Festuca in South American Paramos

S YY

E. Glumes. F. Lemma. G. Lemma with

Figure 12. Festuca caldasii. A. Stylized growth form. B. Habit. C. Ligule. D. Spikelet.

palea and rachilla. H. Leaf blade cross-section. A-H, Laegaard 20405 (F).

38 Festuca in South American Paramos

smaller spikelets (12-14 versus 15-17 mm long),
shorter lemmas (8.5—9 versus 10—14 mm long), and
shorter lemma awns (1 versus 1—3 mm long). In
Costa Rica and Panama there are two other species
in sect. Glabricarpae: F- breviglumis Swallen and
F. chiriquensis Swallen. However, both of these
species have shorter (0.5—1 mm long) and acute
ligules.

Distribution and habitat.—Festuca caldasii
occurs sporadically from southern Colombia
(Cauca, Narifio) to southern Ecuador (Caniar, Chim-
borazo, Loja, Pichincha). This species is known
from the margins of Andean forests and matorral
vegetation between 1900-3000 m.

Additional specimens examined. COLOM-
BIA. Cauca: Tierra Dentro, below Pitaio, 2400
m, Feb 1906, Pittier 437 (US); Rio Paez Valley,
between Huila and Bitonco, 1900-2300 m, Feb
1906, Pittier 1320 (US); near Jambalo, 2200 m, Feb
1906, Pittier 1444 (US). Narino: Mun. Buesaco,
a 2 km de la poblacion, 2000 m, Ramirez 1404
(COL); Mun. Pasto, Morasurco hill, 2550 m, 24
Feb 1986, Wood 5309 (COL, FMB, K). ECUA-
DOR. Canar: El Tambo—Carshao road, km
1.5—3, 02°28'S, 78°59 W, 3100 m, 10 Jun 1999,
S. Laegaard & Sklenar 20302 (LOJA); Chimbo-
razo: Canyon of Rio Chanohan, about 5 km N
of Hiugra, 5000-6500 ft, CAMP (F, K, NY, US);
road Sipambe—Hiugra, km 10, 02°15’S, 78°57’°W,
2050 m, S. Laegaard 20405 (AAU, PRC); Huigra,
1200 m, A.S. Hitchcock 20746 (US); km 6 NE of
Pallatanga and 5.4km W on road to Chillanes,
1950 m, PM. Peterson & E.J. Judziewicz 9259
(QCA, US). Cafion of the Rio Chanchan, about 5 km
N of Huigra, moist forested valleys in the afternoon
fog belt, 5000-6500 ft, 19-28 May 1945, Camp
3327 (K). Loja: Road Catacocha-La Toma, km
28, 03°58’S, 79°31 °W, 2200-2250 m, S. Laegaard
102535 (AAU, QCA); Zumba road, km 6 above
Jimbura, 04°40’, 79°26’W, 2400-2450 m, S. Lae-
gaard 105256 (AAU); road Amalusa—Jimbura,
km 7-9, 04°36’S, 79°28’W, 1900 m, S. Laegaard
105235 (AAU, QCA). Pichincha: Cotocollao,
Sodiro s.n. (QPLS, US); Sodiro s.n. (W).

10. Festuca reclinata Swallen, Contr. U.S. Natl.
Herb. 29(6): 254. 1949. (Figs. 13, 14, 77C—F).
Type: Colombia. Santander: Paramo de Almorz-
adero, Cordillera Oriental, 3500-3700 m, 20
Jun 1940, J. Cuatrecasas & H.G. Barriga 9970
(holotype: US-1798714!; isotype: COL!).

Rhizomatous perennials forming small tussocks
with extravaginal innovations. Culms 30-40 cm tall,
decumbens to erect, glabrous; nodes 2 or 3 in distal
half; leaf sheaths membranous, greenish-white,
scabrous, upper sheaths closed for 1/2 the length;
auricles absent; ligules 1-2.5 mm long, membra-
nous, apex acute, ephemeral; blades 5—15 x 0.3-0.5
cm, flat, green, abaxially scabrous. Panicles 9-10 cm
x 2-3 cm, flexuous, ovate, branched; branches gla-
brous. Spikelets 10—13 mm long, obovate, florets 4;
rachilla papillose; glumes 1.3—3.5 mm long, mem-
branous, lanceolate, green, glabrous, upper margins
hairy; lower glumes 1.3—2 mm long, 1-nerved; upper
glumes 3—3.5 mm long, 3-nerved; lemmas 7—8.5 mm
long, 5-nerved, lanceolate, membranous, green, sca-
brous, apex two-dentate, awned between the teeth,
the awn 1—2 mm long; callus glabrous; paleas 4/5
as long as the lemma, membranous, keels scabrous;
lodicules lanceolate; anthers 3—3.5 mm long; ovary
apex glabrous. Caryopses not seen.

Leaf blade anatomy.—Cross-sections with
many vascular bundles, with small ribs; scleren-
chyma under both abaxial and adaxial epidermis,
discontinuous, small, extending to the vascular
bundles forming girders; bulliform cells absent;
epidermis sparsely hairy.

Observations.—Festuca reclinata superficially
resembles Aphanelytrum procumbens Hack. The
spikelets and panicles of F. reclinata are similar
to Aphanelytrum, but the glumes in the former are
clearly nerved (versus unnerved), which is the differ-
ence between the two genera. Alexeev (1986) placed
F. reclinata in sect. Glabricarpae and we consider
the Alexeev’s decision to be only provisional.

Distribution and habitat.—Festuca reclinata 1s
known only from the type locality in the Colombian
Cordillera Oriental, Dept. Santander were it was
found in a paramo.

11. Festuca woodii Stancik, Darwiniana 41(1-4):
107. 2003. (Figs. 14, 15, 78A—D). Type:
Colombia. Boyaca, Sierra Nevada del Cocuy,
Hda. La Esperanza, 3700 m, in crevices of
limestone pavement on a steep, open dip
slope. Vigorously tufted perennial more than
1 m, inflorescence purple brown, 29 Oct 1985,
J.RI. Wood 5254 (holotype: COL!, isotypes:
FMB], K!).

Rhizomatous perennials with extravaginal
innovations. Culm 70—90 cm tall, erect, scabrous;

Festuca in South American Paramos 39

Figure 13. Festuca reclinata. A. Growing form. B. Ligule. C. Spikelet. D. Glumes. E. Lemma. F. Lemma with palea and rachilla.
G. Leaf blade cross-section. A-G, Cuatrecasas & Barriga 9970 (COL).

40 Festuca in South American Paramos

-75

COLOMBIA

-85

N

400 Kilometers

Figure 14. Distribution of Festuca pratensis (e), F. woodii ( &), and F. reclinata (m).

nodes 2 or 3; leaf sheaths membranous, glabrous,
grayish, fibrous and ephemeral at base, margins
free; auricles absent; ligules ca. 0.5 mm long, mem-
branous, apex truncate; blades 50-60 x 0.25—0.5
cm, flat, conduplicate near apex, green, scabrous,
with prickles on both abaxial and adaxial surface.
Panicles ca. 20 cm long and 15 cm wide, flexuous,
ovate, branched; branches scabrous. Spikelets 12—
14 mm long, ovate, florets 4 or 5; rachilla densely
hairy; glumes 4.5—7.5 mm long, membranous, nar-
rowly lanceolate, green, sparsely scabrous; lower
glumes 4.5—5.5 mm long, 1-nerved; upper glumes
6—7.5 mm long, 3-nerved; lemmas 8.5—9 mm long,
5-nerved, membranous to coriaceous, lanceolate,
green, scabrous and short-hairy, apex two-dentate,
awned, the awn 0.5—1 mm long; callus glabrous;
paleas as long as the lemma, membranous, papil-
lose; anthers 3.5—4 mm long; ovary apex sparsely
short-hairy. Caryopses not seen.

Leaf blade anatomy.—Cross-sections with
many vascular bundles and small ribs above; scle-
renchyma under both abaxial and adaxial epidermis,
discontinuous, extending to the vascular bundles
forming girders; bulliform cells present; epidermis
without hairs.

Observations.—Festuca woodii is morphologi-
cally similar to F’ caldasii known from southern
Colombia and Ecuador. However, F: woodii has
shorter ligules (0.5 versus 2.5—3 mm long), truncate
(versus acute) ligules, smaller spikelets (12-14
versus 15—17 mm long), and longer glumes.

Distribution and habitat.—Festuca woodii
is endemic to the Colombian Cordillera Oriental
(Boyaca, Cundinamarca) occuring in matorral and
grass paramo vegetation types on rocky slopes and
calcareous outcrops between 2700-3700 m.

Additional specimens examined. COLOMBIA.
Cundinamarca: Bogota, 2730 m, 16 Sep 1915,

Festuca in South American Paramos 4]

a “ <<

oS, y,

_——<$<———S—=
SY <
== = = = —
— WS y >
y
"A

>
—

Figure 15. Festuca woodii. A. Stylized growth form. B. Habit. C. Spikelet. D. Glumes. E. Lemma. F. Leaf blade cross-section.
A-F, Wood 5254 (COL).

42 Festuca in South American Paramos

Apollinaire & Arthur 18 (US). Boyaca: Sierra
Nevada del Cocuy, Hda. La Esperanza, 3700 m,
29 Dec 1985, Wood 5354 (FMB).

12. Festuca arundinacea Schreb., Spic. Fl. Lips.
57. 1771. (Figs. 16, 17, 78E & F, 79A & B).
Bromus arundinaceus (Schreb.) Roth, Tent.
Fl. Germ. 2: 141. 1789. Schedonorus arundi-
naceus (Schreb.) Dumort., Observ. Gramin.
Belg. 106. 1824, nom. conserv. Festuca elatior
var. arundinacea (Schreb.) Wimm., Fl. Schles.
3: 59. 1857. Festuca elatior subsp. arundi-
nacea (Schreb.) Celak., Prodr. Fl. Bohmen 1:
51. 1867. Festuca elatior subsp. arundinacea
(Schreb.) Hack., Monogr. Festuc. Eur. 152.
1882. Lolium arundinaceum (Schreb.) Dar-
bysh., Novon 3(3): 241. 1993. Tyre: Scheuzer,
Agrostographia, tab. 5, fig. 18. 1719. (lecto-
type: designated by Reveal, Terrell, Wiersema
& Scholz, Taxon 40: 136. 1991).

Festuca elatior L., Sp. Pl. 1: 75. 1753, nom. rej. Poa
elatior (L.) Moench, Enum. PI. Hess. 37. 1777.
Avena secunda Salisb., Prodr. Stirp. 22. 1796.
Bromus elatior (L.) Koeler, Descr. Gram. 214.
1802. Schedonorus elatior (L.) P. Beauv., Ess.
Agrost. 99, 156, 162, 177. 1812. Festuca pra-
tensis vat. elatior (L.) Gaudin, Fl. Helv. 1: 293.
1828. Bucetum elatius (L.) Parnell, Grasses
Scotl. 107, pl. 46. 1842. Tragus elatior (L.)
Panz. ex B.D. Jacks., Ind. Kew. 2: 1098. 1895.
Gnomonia elatior (L.) Lunell, Amer. Midl.
Nat. 4: 224. 1915. (lectotype: LINN-92.17,
designated by Terrell, Brittonia 19: 131. 1967;
again by Linder, Bothalia 16: 59. 1986).

Poa phoenix Scop., Fl. Carniol., ed. 2, 1: 74. 1771.
Schedonorus phoenix (Scop.) J. Holub, Preslia
70: 113: 1998:

Bromus littoreus Retz., Fl. Scand. Prodr. 19. 1779.
Schedonorus \ittoreus (Retz.) Tzvelev, Nov.
Sist. Vyssh. Rast. 31: 258. 1998.

Festuca littoralis Wahlenb., nom. illeg. hom., Nova
Acta Regiae Soc. Sci. Upsal. 8: 211. 1821.

Festuca pseudosclerophylla Krivot., Bot. Mater.
Gerb. Bot. Inst. Komarova Akad. Nauk SSSR
17: 73. 1955. Leucopoa pseudosclerophylla
(Krivot.) Bor in K. H. Rech., Fl. Iranica 70:
13. 1970:

Poa hybrida var. vallesiaca Bronm., Repert. Spec.
Nov. Regni Veg. 16: 301. 1919.

Festuca elatior subsp. arundinacea var. genuina
subvar. orientalis Hack., Monogr. Festuc.
Europ. 154. 1882. Festuca orientalis (Hack.)

Krecz. & Bobrov, Fl. URSS 2: 531. 1934.
Festuca arundinacea subsp. orientalis (Hack.)
Tzvelev, Fl. URSS 18: 17. 1970.

Festuca regeliana Pavl., Byull. Moskovsk. Obshch.
Isp. Prir., Otd. Biol. 41(1): 80. 1938.

Loosely tufted to shortly rhizomatous peren-
nials with extravaginal innovations. Culms (50-)
100-150 cm tall, erect, scabrous; nodes 3. Leaf
sheaths coriaceous, striate, glabrous or scabrous;
auricles present, falcate, margins ciliate; ligules
ca. 0.5 mm long, membranous to coriaceous, apex
truncate; blades 25—30 cm long, 5—11 mm wide, flat,
green, abaxially scabrous. Panicles 10-17 x 4-6 cm,
narrow, with erect branches; branches scabrous.
Spikelets 9-12(—15) mm long, florets 6—8(—10);
rachilla scabrous; glumes 3.3—6.5(—7) mm long,
lanceolate, membranous to coriaceous, green, gla-
brous, apex acute; lower glumes 3.3—5(—5.5) mm
long, 1-nerved; upper glumes 4.5—6.5 mm long,
3-nerved, sometimes scabrous on back; lemmas
6—7.5(—8) mm long, 5-nerved, lanceolate, membra-
nous to coriaceous, green, apex scabrous on midrib,
mucronate or short-awned, the awn 0.5—1.5 mm
long; paleas almost as long as the lemma, scabrous
on margins and keels; lodicules lanceolate, acumi-
nate; anthers 2.7—3.5 mm long; ovary apex glabrous
or sparsely hairy.

Leaf blade anatomy.—Cross-sections with
numerous vascular bundles, with small ribs above;
sclerenchyma discontinuous, extending to the
vascular bundles under both abaxial and adaxial
epidermis; bulliform cells present.

Observations.—This species is often treated
by many authors as a separate genus, Schedonorus
arundinaceus (Soreng et al. 2003).

Distribution and habitat.—Festuca arundi-
nacea 1s introduced from Europe and cultivated in
pastures, fields and rarely escaping along roadsides
between 2300-3600 m.

Additional specimens examined. COLOMBIA.
Cundinamarca: Mun. de Suba, Hda. Las Mer-
cedes, 16 May 1964, E. Forero et al. 45 (COL); 8
Oct 1964, E. Forero et al. 82 (COL); 15 Oct 1964,
E. Forero et al. 92 (COL); between Guasca and La
Calera, Vereda Santa Helena, 2950 m, 9 Dec 1984,
Wood 4639 (COL, K). Narifio: Mun. Ipiales, “Las
Lajas”, 2740 m, 8 Aug 1939, Garcia-Barriga 7838A
(COL). ECUADOR. Cajiar: S of El Tambo, km
1.5—3 on road to Carshao, 02°28’S, 78°59’ W, 3100
m, S. Laegaard & Sklenar 20301 (AAU); Mun. El
Tambo, road from El Tambo to Ingapirca, km 1,

Festuca in South American Paramos 43

Figure 16. Festuca arundinacea. A. Stylized growth form. B. Habit. C. Inflorescence. D. Ligule. E. Spikelet. F. Glumes. G. Lemma.
H. Lemma with palea and rachilla. I. Leaf blade cross-section. A-I, Stancik 4102 (PRC).

44 Festuca in South American Paramos

burned matorral and paramo, 3400-3500 m, 29 Aug
2000, D. Stanctk 3798 (PRC, QCA). Chimborazo:
km 5 N of Tixan, meadows, 02°06’S, 78°46’°W,
3280 m, S. Laegaard 101818 (AAU, QCAQCNE);
S. Laegaard et al. 103003 (AAU, QCA, QCNE); km
22 on road Alausi—Riobamba, 02°08’S, 78°46’ W,
3250 m, S. Laegaard 18667 (AAU,LOJA, QCA,
QCNE); pasture near Riobamba, Sodiro s.n. (QPLS).
Cotopaxi: Salcedo, 2650 m, Acosta-Solis 10206 (F,
US). Loja: La Argelia (Escuela de Agronomia),
04°02’S, 79°12°W, 2100 m, S. Laegaard 18704
(LOJA). Morona-Santiago: Hda. Huargualla—Hda.
San Eduardo, 01°57’S, 78°32.2’W, 3600 m, 19 Jul
1999, D. Stancik 3309 (PRC, QCA); D. Stancik
3310 (PRC, QCA). Pichincha: Quito, 00°10’S,
78°30'W, 2850 m, 30 Mar 1998, S. Laegaard 18623
(LOJA); Quito—Santa Catalina, 00°22°S, 78°21 °W,
Feb 1981, Vivar & Marin 1305 (LOJA); Quito—El
Batan, 2850 m, Acosta-Solis 19821 (US); Acosta-
Solis 19832 (US); Acosta-Solis 19835 (US); W
side of Mt. Pichincha, 3070 m, Sodiro s.n. (QPLS);
Pichincha, Aug 1888, Sodiro s.n. (QPLS); 3000 m,
1904, Sodiro s.n. (US); Road Lloa—Guagua Pichin-
cha, km 6, 00°13’S, 78°35’ W, 3600 m, S. Laegaard
102716 (AAU). Mun. Otavalo, Ruchanda-—road to
Quito, 3100 m, 14 Jul 1999, D. Stancik 3221 (PRC,
QCA); Mun. Otavalo, road from Laguna Mojanda
to Cochasqui, 00°04°55’N, 78°17°50°W, 3450 m,
19 Oct 2000, D. Stancik 4102 (PRC, QCA). Tungu-
rahua: Tungurahua, Aug 1901, Sodiro s.n. (QPLS,
US); Slope of Mt. Tungurahua above Bajos, 2300
m, FE. Asplund 8425 (S); Pillaro, 2850 m, E. Asplund
8150 (NY, QCA, S). Zamora-Chinchipe: Old
road Loja—Zamora ca. 1 km E of pass, 03°59'S,
79°09'W, 2750 m, 26 Apr 1998, S. Laegaard 18744
(AAU, LOJA, QCA, QCNE). VENEZUELA.
Barinas: 74 km NW of Barinas on Hwy 1 and
71 km NE of Mérida, slopes with Festuca, Espe-
letia, Loutegia, Stevia, and Sporobolus, 2740 m,
24 Nov 1991, PM. Peterson 11181 (US). Mérida.
Entre Pedregal y Apartaderos, praderas de fuentes
del Chama, asociado a Juncus, Rumex etc., 3240 m,
23 Aug 1981, Ponce & Trujuillo 245 (MY); Dept.
Rangel, Paramo de Mucubaji, Mesa del Caballo,
El Pedregal, 3350 m, 12 Jun 1981, B. Bricerio &
Adamo 286 (Herbarium Bricefio, PRC).

13. Festuca pratensis Huds., Fl. Angl. 37. 1762.
(Fig. 14). Festuca fluitans var. pratensis (Huds.)
Huds., Fl. Angl. (ed. 2) 47. 1778. Schedonorus
pratensis (Huds.) Beauv., Ess. Agrostogr. 99,
163, 177. 1812. Bromus pratensis (Huds.)

Spreng., Syst. Veg. 1: 359. 1825, nom. illeg.
Bucetum pratense (Huds.) Parnell, Grass.
Scotland 105, t. 46. 1842. Festuca elatior var.
pratensis (Huds.) A. Gray, Man. Bot. (ed. 5) 634.
1867. Festuca elatior subsp. pratensis (Huds.)
Hack., Bot. Centralbl. 8: 407. 1881. Festuca
elatior subsp. pratensis (Huds.) Hack., Monogr.
Festuc. Eur. 150. 1882, isonym. Tragus pratensis
(Huds.) Panz. ex B.D. Jacks., Index Kew. 2:1099.
1895. Lolium pratense (Huds.) Darbysh., Novon
3(3): 242. 1993. Type: Great Britain (BM-SL),
Buddle s.n., (lectotype: BM-SL!, Herb. Sloane.
125.16, designated by Reveal, Terrell, Wiersema
& Scholz, Taxon 40:135. 1991).

Festuca poaeoides Michx., Fl. Bor.-Amer. 1: 67.
1803, nom. illeg. Festuca poaeoides vat.
americana Pers., Syn. Pl. 1: 94. 1805. Festuca
americana (Pers.) F. G. Dietr., Nachtr. vollst.
Lex. Gartn. 3: 332.1817. Schedonorus ameri-
canus (Pers.) Roem. & Schult., Syst. Veg. 2:
706. 1817.

Schedonorus radicans Dumort., Obs. Gramin.
belg.: 106. 1824. Festuca radicans (Dumort.)
Steud., Syn. Pl. Glum. 1: 309. 1854.

Festuca glabra Spreng., Syst. Veg. 1: 353. 1824,
nom. illeg. hom.

Rhizomatous perennials with extravaginal
innovations. Culms 60—120 cm tall, erect, scabrous;
nodes 3 in the basal half. Leaf sheaths (lowermost)
coriaceous, brown, striate, scabrous, margins free,
sometimes fibrous; auricles falcate, without hairs
on margins; ligules 0.3—0.5 mm long membra-
nous, sometimes coriaceous, apex truncate; blades
20-25 x 0.4—0.8(—1) cm, flat, green, abaxially sca-
brous. Panicles 15—20 x 3—5 cm, contracted, narrow;
branches scabrous. Spikelets 9-11 mm long, florets
5 or 6; rachilla scabrous; glumes 2.7—-4 mm long,
lanceolate, membranous, sometimes coriaceous,
green, glabrous, apex acuminate; lower glumes
2.7—3 mm long, 1-nerved; upper glumes 3.74 mm
long, 3-nerved; lemmas 6—6.5 mm long, 5-nerved,
membranous to coriaceous, lanceolate, green, apex
scabrous, mucronate; paleas almost as long as the
lemma, scabrous on margins and keels; lodicules
lanceolate, acuminate; anthers 2.7—3.5 mm long;
ovary apex glabrous or sparsely hairy.

Leaf blade anatomy.—Cross-sections with
numerous vascular bundles, without ribs above;
sclerenchyma under both abaxial and adaxial epi-
dermis, discontinuous, extending to the vascular
bundles forming girders; bulliform cells present.

Festuca in South American Paramos 45

N

300 600 Kilometers

Figure 17. Distribution of Festuca dichoclada (¢), F: arundinacea ( &), and F: venezuelana (m).

Observations.—Modern authors generally
treat this taxon as a separate genus, Schedonorus
pratensis (Soreng et al. 2003).

Distribution and habitat.—This species is
introduced from the Europe and sporadically cul-
tivated at an altitude around 3000 m.

Additional specimens examined. ECUADOR.
Chimborazo: Riobamba, Sodiro s.n. (US). Pich-
incha: Quito—Santa Catalina, 00°22’S, 78°31°W,
Vivar & Marin 1306 (LOJA); Canton Tabacundo—
Picalqui, 2750 m, Acosta-Solis 16268 (US).

14. Festuca dichoclada Pilg., Bot. Jahrb. Syst. 37:
514. 1906. (Fig. 17). Festuca quadridentata
var. dichoclada (Pilg.) St.-Yves, Candollea
3: 265. 1927. Type: Peru. Ancachs (Ancash),
in declivibus montium Cordillera blanca
supra Caraz in faucibus umbrosis fruticibus
altis obtectis, 3300-3600 m, 9 Jun 1903,
A. Weberbauer 3230 (holotype: B!; isotypes:

BAA-1196 fragm. ex B, US-2875396 fragm.
exe,

Large rhizomatous, loosely caespitose perenni-
als with intravaginal innovations. Culms 150-200
cm tall, erect, scabrous; nodes 3 or 4 in distal half.
Leaf sheaths coriaceous, scabrous; auricles absent;
ligules 8-12 mm long, membranous, apex acute;
blades 40—100 =< 0.5—1.4 cm, flat occasionally par-
tially conduplicate, green, abaxially scabrous, with
prickles on abaxial and adaxial epidermis. Panicles
20-40 x 20—25 cm, ovate; branches scabrous. Spike-
lets 11-14 mm long, oblong, florets 3—5; rachilla
hairy; glumes 5—9 mm long, lanceolate, greenish-
white, smooth to papillose, apex acute; lower
glumes 5—7.5 mm long, l-nerved; upper glumes
6.5—9 mm long, 3-nerved; lemmas 8.5—10 mm long,
lanceolate, 5-nerved, membranous to coriaceous,
scabrous, apex entire, awnless; callus glabrous;
paleas as long as the lemma, smooth to papillose;

46 Festuca in South American Paramos

lodicules ovate, obtuse; anthers 3.5—4.2(-4.7) mm
long; ovary apex glabrous. Caryopses oblong; hilum
nearly as long as the caryopsis.

Leaf blade anatomy.—Cross-sections with
many vascular bundles and ribs above; scleren-
chyma under both abaxial and adaxial epidermis,
discontinuous; bulliform cells present; epidermis
without hairs.

Distribution and habitat—Known only from
northern and central Peru. It occurs along margins
of Andean forests, clearings, rocky slopes, and
paramos between 3000—4000 m.

Additional specimens examined. PERU.
Ancash: Prov. Bolognesi, Jerusalem, cerro al E de
Aquia, arcilloso—pedregoso en monte pluvifolio,
3200-3250 m, 18 May 1950, R. Ferreyra 7526
(US); Casca, abajo de Chiquian, 3100-3200 m,
9 May 1950, R. Ferreyra 7281 (US); Prov. Yun-
gay, Huascaran Parque Nacional, between Lake
Llanganuco and Portachuelo, 77°36’W, 09°03’S,
4200-—4329m, 16 Aug 1984, Smith 8233 (US).
Prov. Huaraz, Huascaran Parque Nacional, Que-
brada Shallap, 77°24’ W, 09°30’S, 3700-4000 m,
22 May 1985, Smith et al. 10769 (F); Prov.
Huaylas, Huascaran Parque Nacional, environs of
Auquispuquio, 77°57’ W, 08°49°S, 3900-4000 m,
4 Sep 1986, Smith et al. 12110 (F); Prov. Yungay,
Huascaran Parque Nacional, Llanganuco sec-
tor, Maria Josefa trail between Chinancocha and
Pucayacu, 77°39’ W, 09°05’S, 3700-3850 m, 5 Jul
1985, Smith 10537 (F); Prov. Carhuaz, Huascaran
Parque Nacional, Quebrada Ulta, on road to Ulta
Pass, 77°32°W, 09°07’S, 4000-4400 m, 29 Jul
1985, Smith 11389 (F). Cajamarca: Prov. Con-
tumaza, Guzmango, cerro Campanillas, 3050 m,
1 Jun 1959, SagasteguiiSagasteguii 2991 (US);
Prov. Cajamarca, SAIS José Carlos Mariategui,
km 20—40 on Sunchubamba—San Juan road, jalca
with small patches of ceja de selva in rocky area,
3300-3500 m, 5 Jun 1984, Smith 7534 (US); Prov.
Cajamarca, Distr. San Juan, carretera San Juan—
Cajamarca, arriba de San Juan, ladera arcillosa-
pedregosa, 2350 m, 6 Dec 1993, Vega 730 (F);
Prov. Cajamarca, Cumbe Mayo, 21 km al W de
Cajamarca, ladera con arbustos disersos, 3100 m, 4
Nov 1977, Vega 1966 (F). Huancavelica: Conaica,
Carhuay—pampa arriba de Conaica, 3700-3750 m,
18 Mar 1951, O. Tovar 143 (US). Junin: Prov.
Huancayo, Hda. Acopalca, 4000 m, 20 Jul 1945,
Infantes 435 (US). La Libertad: Prov. De Otuzca,
cerca a Usquil, falda de cerro, junto con Chusquea,
Loasa, Cajophora, 3000-3100 m, 9 Jun 1950,

R. Ferreyra 7642 (K, US); R. Ferreyra 7664 (K,
US); Otuzco, entre piedras, 2600 m, 6 Apr 1990,
Leiva & Leiva 96 (F); Otuzco, cerro de los Enamo-
rados (al norte de Salpo), ladera, 3440 m, 16 Jun
1993, Leiva 797 (MO); Near Usquil, Utusco, hill-
side, 3200 m, 9 Jun 1950, Anderson 1271 (US).

15. Festuca quadridentata Kunth, Nov. Gen.
Sp. (quarto ed.) 1: 154. 1816. (Figs. 18, 21,
79C-F, 80A). Festuca flexuosa Willd. ex
Kunth, Enum. Pl. 1: 407. 1833, nom. inval.
pro syn. Type: Ecuador. Chimborazo, 2860
m, Humboldt & Bonpland s.n. (holotype: P!;
isotypes: B!, P!).

Large tussocked perennials with intravagi-
nal innovations. Culms 170—200 cm tall, erect,
scabrous; nodes 3 or 4 in distal half. Leaf sheaths
coriaceous, striate, scabrous; auricles absent; lig-
ules (8—)9—13 mm long, membranous, apex acute;
blades 40-150 x 0.7—1.4 cm, flat to conduplicate,
green, scabrous with prickles on abaxial and adax-
ial epidermis. Panicles 20-40 x 15—20 cm, ovate;
branches scabrous. Spikelets 11—13(—15) mm long,
oblong, florets (3—)4 or 5(—6); rachilla with short,
scattered hairs; glumes (3.5—)4—6.5 mm long,
lanceolate, greenish-white, smooth sometimes
papillose, apex acute; lower glumes (3.5—)4—5 mm
long, 1-nerved; upper glumes 5—6.5 mm long,
3-nerved; lemmas 8.5—9.5 mm long, lanceolate,
5-nerved, membranous to coriaceous, scabrous
or papillose, apex erose and dentate, awnless or
sometimes mucronate; callus glabrous; paleas as
long as the lemma or longer, smooth or papillose;
lodicules ca. 0.7 mm long, ovate, apex obtuse;
anthers 3.5—4.2(-4.7) mm long; ovary apex gla-
brous. Caryopses oblong; hilum nearly as long as
the caryopsis.

Leaf blade anatomy.—Cross-sections with
many vascular bundles, corresponding to the num-
ber of ribs above; sclerenchyma under both abaxial
and adaxial epidermis, discontinuous; bulliform
cells present; epidermis without hairs.

Observations.—Festuca quadridentata 1s
morphologically similar to the Peruvian species
F: dichoclada, however the former differs by having
shorter lower glumes (5—7.5 mm long in F- dicho-
clada), shorter upper glumes (3.5—5 mm versus
6.5—9 mm), and dentate lemmas (versus entire).

Distribution and habitat.—This species is
endemic to central Ecuador (Cafiar, Chimborazo,
Morona-Santiago, Pichincha) where is known from

47

Festuca in South American Paramos

Wi

.
EA

FZ)

i
yy Yj

jl

F. Lemma. G. Lemma

E. Glumes.
if 20308 (PRC); H, Stancik 3317 (PRC).

Figure 18. Festuca quadridentata. A. Stylized growth form. B. Habit. C. Ligule. D. Spikelet.
with palea and rachilla. H. Leaf blade cross-section. A-G, Laegaard & Sklena

48 Festuca in South American Paramos

Andean mountain forests, between (1500—)3000-
3400 m.

Additional specimens examined. ECUADOR.
Canar: S of El Tambo, ca. 1.5—3 km road to Car-
shao, 02°28’S, 78°59’ W, 3100 m, 10 Jun 1999, S.
Laegaard & Sklenar 20308 (AAU, LOJA, PRC);
10 Jun 1999, S. Laegaard & Sklenar 20287 (AAU,
LOJA, PRC). Chimborazo: near Alao, along Rio
Alao, 01°52’S, 78°30°W, 3200-3400 m, 22 Sep
1985, S. Laegaard 55290 (AAU, QCNE); km
10 N of Sipambe, 02°08’S, 78°52°W, 3400 m,
11 Nov 1985, S. Laegaard 55567 (AAU, QCA);
road Alao—Chambo, km 10, 01°50'S, 78°34 'W,
3000 m, 11 Oct 1985, S. Laegaard 55411 (AAU,
QCA); in prov. Riobamba, Sodiro s.n. (QPLS);
Alao, Hab. Llactapamba, 3200 m, Acosta-Solis
7581 (F, US); eastern Cordillera of Riobamba, 3200
m, Rimbach 54 (F, US). Morona-Santiago: Parque
Nacional Sangay, Hda. Huargualla—San Eduardo,
02°0025'S, 78°27 W, 37000m, DD. Stancil 33:17
(AAU, QCA, PRC, US). Pichincha: Gualea, Sodiro
s.n. (QPLS); 1500 m, Mille s.n. (QPLS). Sin. loc.
Sodiro 20/5 (P).

16. Festuca venezuelana Stancik, Darwiniana
41(1-4): 111, f. 15b-1. 2003. (Figs. 17, 19,
80B-—F). Tyre: Venezuela. Tachira, Mun. La
Grita, Paramo La Negra, cross of the roads to La
Grita and Pogonero, 08°13 '22°N, 71°52’51°W,
shrubby margin of the road with Asteraceae,
Melastomataceae, Cordia sp., 2800 m, 11
Nov 2000, D. Stancik 4262 (holotype: PRC!,
isotypes: AAU!, CAR!, COL!, W!).

Tussocked perennials with extravaginal inno-
vations. Culms 130-180 cm tall, erect, scabrous;
nodes 3. Leaf sheaths coriaceous, fibrous, scabrous,
brown; auricles absent; ligules 3—5 mm long, mem-
branous, apex acute; blades 25-40 x 0.5—1.1 cm,
flat, green, abaxially scabrous. Panicles 20-25 x
10—15 cm, lax, nutant; branches scabrous. Spikelets
14-16 mm long, oblong-lanceolate, florets 5—7;
rachillas 1—1.2 mm long, scabrous; glumes 3—9
mm long, narrowly lanceolate, coriaceous, margins
membranous, apex acute; lower glumes 3-4.5 mm,
l-nerved; upper glumes 7—9 mm long, 3-nerved,
apex scabrous; lemmas 10—11 mm long, 5-nerved,
lanceolate, coriaceous, brownish-green, scabrous,
apex two-dentate and awned, the awn 2-8 mm long,
some plants awnless; paleas as long as the lemma,
scabrous on margins and keels; lodicules lanceolate,
acuminate; anthers 3—3.5 mm long; ovary apex

glabrous. Caryopses oblong-lanceolate; hilum 3/4
of caryopsis length.

Leaf blade anatomy.—Cross-sections with
many vascular bundles, with shallow ribs; scleren-
chyma under both abaxial and adaxial epidermis,
discontinuous; bulliform cells present; epidermis
with prickles on abaxial and adaxial surfaces,
macrohairs absent.

Observations.—Festuca venezuelana 1s mor-
phologically similar to the Bolivian F: steinbachii.
However, F: steinbachii has shorter ligules (ca.
1 mm long), shorter spikelets (11-13 mm long),
and shorter awns (0.5—1.3 mm long).

Distribution and habitat.—Festuca venezu-
elana ranges from Colombian Cordillera Oriental
(N de Santander) to the western part of the Venezu-
elan Andes (Tachira). It occurs in Andean forest
clearings between 2800-3400 m.

Additional specimens examined. COLOMBIA.
Norte de Santander: between Mutiscua and Pam-
plona, edge of wood, 3400 m, 32 Feb 1927, Killip
19719 (K, US). VENEZUELA. Tachira: Grita,
Porqueras-Aldea Agua Diaz, 22 Sep 1942, Tamayo
2326 (US, VEN).

17. Festuca fragilis (Luces) B. Bricefio, Ernstia
4(3-4): 7 8-79. 1994. (Figs. 20, 21, 81A—F).
Helleria fragilis Luces, J. Wash. Acad. Sci.
32(6): 157, f. 1. 1942, nom. inval. Hellerochloa
fragilis (Luces) Rauschert, Taxon 31(3): 561.
1982. Type: Venezuela. Mérida, Paramo de
Tucani, 4500 m, 17 Dec 1910, A. Jahn 62
(holotype: VEN!; isotype: US!).

Small tussocked perennials with intravaginal
innovations. Culms 20—25 cm tall, erect, finely sca-
brous; nodes 1, basal. Leaf sheaths membranous to
coriaceous, stramineous, glabrous, inconspicuously
striate; auricles absent; ligules 4-5 mm long, mem-
branous, apex acute, ciliate; blades 10—15 cm long,
0.6—0.7 mm wide, conduplicate to involute, abaxi-
ally scrabrous, olive-green, only 2 or 3 per culm,
apex acute. Panicles 5—8 x 1-2 cm, contracted,
dense, lanceolate, branches scabrous. Spikelets ca.
3 cm long, obovate, florets 5—7; rachilla densely
hairy; glumes 7.5-13 mm long, membranous,
lanceolate, margins dentate, apex acute; lower
glumes 7.5—9.5 mm long, 1-nerved; upper glumes
10-13 mm long, 3-nerved; lemmas 17—20 mm long,
5-nerved, lanceolate, membranous, upper 1/2 and
margins scabrous, apex shortly two-dentate and
awned, the awn 5—7 mm long; callus glabrous;

49

Festuca in South American Paramos

F. Glumes. G. Lemma.

Figure 19. Festuca venezuelana. A. Stylized growth form. B. Habit. C. Inflorescence. D. Ligule. E. Spikelet.

H. Lemma with palea and rachilla. I. Leaf blade cross

section. A-I, Stancik 4263 (PRC).

Festuca in South American Paramos

50

Add:

tg
PEA 77
WF pe

ALO
LEG,
SGT
Kel
L/

Ii
y A ig
H i ( J
| xX
iil O

Figure 20. Festuca fragilis. A. Habit. B. Ligule. C. Spikelet. D. Glumes. E. Lemma. F. Lemma with palea and rachilla. G. Leaf

blade cross-section. A—G, from Stancik 4192 (PRC).

Festuca in South American Paramos 51

-80 -75

COLOMBIA

ECUADOR

-80 -75

N

200 400 Kilometers

Figure 21. Distribution of Festuca quadridentata (@), F- fragilis (A), and F. rubra (m).

paleas 3/4 as long as the lemma, membranous,
lanceolate, papillose, upper third and along keels
scabrous; lodicules lanceolate; anthers 1.5—2.2 mm
long; ovary apex glabrous. Caryopses lanceolate;
hilum 2/3 of total length.

Leaf blade anatomy.—Cross-sections with 7
vascular bundles and 5 ribs above; sclerenchyma
under abaxial and adaxial epidermis, continuous,
girders absent; epidermis with hairs on adaxial
surface, scattered, the hairs 0.03 mm long.

Observations.—The Mexican species,
F livida (F: subg. Helleria) is morphologically simi-
lar to F: fragilis but differs from it by having shorter
culms (10—15 cm tall), shorter lemmas (8-14 mm
long), and shorter lemmatal awns (1—4 mm long).

Distribution and habitat.—Festuca fragilis
is known only from the state of Mérida, Venezu-
ela and the Sierra Nevada del Cocuy, Colombia,

from dry rocky slopes of super-paramo between
3600-4800 m.

Additional specimens examined. COLOMBIA.
Arauca: Sierra Nevada del Cocuy, Laguna La
Plaza, 4300 m, 31 Dec 1985, Wood 5259 (COL).
VENEZUELA. Merida: Mun. Mucuchies, Paramo
de Piedras Blancas, mountain ridge towards Alto
Mucunano, 08°49'54’"N, 70°57'33"W, 4350 m,
4 Nov 2000, D. Stancik 4182 (CAR, COL, PRC,
US, W); Laguna Negra, 08°49°16'N, 70°57°02’W,
4270 m, D. Stancik 4192 (AAU, CAR, COL, PRC);
Mun. Merida, Sierra Nevada—Pico Espejo, 8°42°N,
71°12’W, 4 Nov 2000, D. Stancik 4248 (AAU,
CAR, COL, PRC, W); Barclay & Juajibioy 10180
(US); Pico de Las Piedras Blancas, 4740 m, 25
May 1952, Vareschi 1232 (VEN); Camino a Pico
Bolivar, 4100-4300 m, 10 Oct 1953, Little 15717
(MER); paramo Media Luna, pendiente N del Pico

52 Festuca in South American Paramos

El Toro, 08°32'27'N, 71°04°19"W, super-paramo
con Lachemilla equisetifolia y Coespeletia morit-
ziana, 4350 m, 21 Dec 1994, Berg 470 (K); Sierra
Nevada de Santo Domingo, between Apartaderos
and Timotes, E facing rocky slope, 21—26 Nov
1959, Barclay et al. 9677 (US); Laguna Coromoto,
3600 m, Oct 1956, Aristeguieta 2596 (VEN); 3400
m, Oct 1956, Aristeguieta 2603 (VEN); Paramos
mas altos cerca de El Gavilan, 4200 m, 25 Jan
1929, Pittier 13276 (VEN, US); Paramo de Pie-
dras Blancas, 4300 m, 25 Sep 1952, Vareschi et
al. 1233 (VEN); 4200 m, 28 Feb 1976, Baruch 46
(VEN); 4400 m, 8 Dec 1979, Barreto 656 (MERC);
4000-4600 m, 31 Oct 1981, B. Briceno et al. 358
(MERC); 4150-4250 m, 13 Nov 1976, Ricardi &
Carrez 5930 (MER); Paramo de Mucuchies, Pico
Aguila, Pico de Aguila, 3800-3900 m, 22 Jan 1986,
Bono 5662 (VEN); 4300 m, 17 Oct 1984, Bono
4328 (VEN); 4700 m, 12 Feb 1976, Teran 13215
(MERF); 4118m, 6 Dec 1980, Badillo et al. 7562
(MY); Pico Espejo, 4765m, 17 Dec 1969, J. Steyer-
mark & Koyama 102389 (VEN); 4680-4780 m, 31
Dec 1961, Teran & Melchior 833 (MER, HERZ);
Teran & Melchior 848 (MER); 4400 m, 16 Dec
1952, Bernardi 268 (MER); Loma Redonda—Alto
de La Cruz, 4040-4300 m, 11 Nov 1994, B. Briceno
3074 (Herbarium Bricefio); Mun. Rangel, Paramo
El Banco, entrada por la Toma baja, 4420 m, 21
Oct 1997, B. Briceno & Molinillo 346] (Herbarium
Bricefio); Paramo Mucubaji, camino a Mucunuque,
3600 m, 25 Oct 1980, B. Briceno & Adamo 244
(Herbarium Bricenio, US); 4200-4300 m, 4 Nov
1992, Meier 3025 (VEN); Distr. Campo Elias,
Sierra La Culata, Pico El Campanario, 4325m, 20
Oct 1972, Teran 7732 (MERF); 3600-4100 m, 26
Oct 1972, Teran 7909 (MERF);

18. Festuca fimbriata Nees, Fl. Bras. Enum. Pl. 2(1):
472.1829. (Figs. 22, 23, 82A—E). Type: Brasilia
meridionales, Montevideo, F: Sellow s.n. (holo-
type: B!; isotypes: K!, LE-TRIN-2806.01!, LE-
TRIN-2806.02!, US-557541 ex B!, US-1441522
ex B!, US-1126679 fragm. ex W!, W!).

Festuca ampliflora Doll, Fl. Bras. 2(3): 116, pl.
34. 1878. Type: Brazil. Minas Gerais, Caldas,
Regnell III 1409 (lectotype: S! designated by
Alexeev, Bot. Zhurn. (Moscow & Leningrad)
69: 348. 1984; isolectotypes: P!, US-2875375
fragm. ex M!).

Rhizomatous perennials with extravaginal
innovations. Culms (60—)90—150(—250) cm tall,

erect, glabrous; nodes 3 or 4 in distal half; cataphylls
sometimes present, about 4-6 cm long. Leaf sheaths
membranous, brown, striate, glabrous; auricles
absent; ligules 0.1—0.5(—1) mm long, coriaceous,
apex truncate, ciliate; blades 20-50 x 0.4—1.5 cm,
flat, green, abaxially scabrous with prickles on
abaxial epidermis. Panicles 20-40 x 10—15 cm,
ovate; branches scabrous. Spikelets 11—15(—20) mm
long, oblong or ovate, florets (S—)6—7(—9); rachillas
1—1.3 mm long, glabrous; glumes (3.5—)4-8(9) mm
long, lanceolate, scabrous, apex acute; lower glumes
(3.5—)4—6(—7) mm long, 1-nerved; upper glumes
5.5—6(—7.5) mm long, 3-nerved; lemmas 6-8
(—9) mm long, membranous to coriaceous, lanceo-
late, 5-nerved, scabrous, papillose, apex entire, awn-
less; callus glabrous; paleas as long as the lemma,
scabrous; lodicules ca. 1mm long, lanceolate, with
scattered hairs at apex; anthers 2.5—3.5(4) mm
long; ovary apex pilose. Caryopses lanceolate; hilum
nearly as long as the grain.

Leaf blade anatomy.—Cross-sections with
numerous vascular bundles with small ribs above;
sclerenchyma under both abaxial and adaxial epi-
dermis, discontinuous and extending to the vascular
bundles; bulliform cells present; epidermis with
scattered prickles.

Observations.—Festuca fimbriata 1s charac-
terised by having pilose lodicules. This unusual
feature probably represents a plesiomorphy since
elsewhere in the genus it is only known to occur in
the Australian F’ subg. Austrofestuca.

Distribution and habitat.—Festuca fimbriata
is restricted to the area between northern Uruguay,
northeastern Argentina, and southeastern Brazil,
in forest openings, wet depressions, and swamps
between (70—)700—1650(—2400)m.

Additional specimens examined. ARGEN-
TINA. Corrientes: Dept. Santo Tomé, Ea. Timbo,
costa del Rio Uruguay, 26 km SE de Colina
Garabi, 16 Sep 1980, O. Ahumada & A. Schinini
4085 (CTES); Arroyo Chimiray y Ruta 40, 8 Oct
1976, Quarin 3409 (CTES); Dept. Santo Tomé,
Garruchos, 22 Oct 1954, Burkart 19654 (BAA);
Estancia “Garruchos”, swamps and wet medows,
T.M. Pedersen 819 (C, MO); Ruta 37, 5 km E de
Gdor. Virasoro, 14 Nov 1974, A. Schinini & Car-
nevali 10523 (US). Misiones: Dept. Capital, Ruta
provincial No. 1, 12 km S de Posadas, 16 Nov 1974,
A. Schinini & Carnevali 10687 (CTES). BRAZIL.
Minas Gerais: Jan 1880, Regnell s.n. (W); Caldas,
28 Jan 1846, Widgren s.n. (K, US); Monte Verde,
Sep 1997, Longhi & Witten 5011 (ICN). Parana:

a3

Festuca in South American Paramos

Figure 22. Festuca fimbriata. A. Stylized growth form. B. Habit. C. Inflorescence. D. Ligule. E. Spikelet. F. Glumes. G. Lemma.

H. Lemma with palea and rachilla. I. Lodicule. J. Leaf blade cross-section. A—J

, .M. Pedersen 11475 (MO).

54 Festuca in South American Paramos

4

0 500 1000 Kilometers
el +

Figure 23. Distribution of Festuca fimbriata (@), and F: andicola (A).

Piraquara, Pinhais,, 25 Nov 1968, G. Hatschbach
20379 (C, K); Mun. Balsa Nova, Bicudo, 6 Dec
1962, G. Hatschbach 9551 (US); 26 Feb 1909,
Dusen 8972 (K, US); Mun. Curitiba, Atuba, mar-
gin of swamp, 23 Jan 1965, Clayton 4301 (K);
Castro, Rio Cunhaporanga, varzea, 26 Nov 1968,
G. Hatschbach 20381 (K); Mun. Jaguariaiva, Rio
Jaguariaiva, campo, 13 Nov 1974, G. Hatschbach
35575 (K). Rio de Janeiro: Mun. Itatiaia, Pratelei-
ras, 2300 m, Mar 1937, Brade & Kuhlman 15631
(B, COL, PRC, RB); Serra de Itatiaia, 1900 m, Mar
1894, Ule 243 (R); Rio de Janeiro, Alto de Itatiaia,
2200-2400 m, 17 Jan 1925, A. Chase 8302 (US).
Rio Grande do Sol: Aparados da Rocinha, Bom
Jesus, 18 Jan 1950, Rambo 45452 (US); Vacaria, 9
Jan 1946, J.R. Swallen 8189 (MO, US); Mun. Rio
Pardo, Fasenda Soledad (Jurgens), 70 m, 1905,
Jurgensen G85 (US); Bom Jesus, entre Ausentes e
Poste Fiscal da Serra da Rocinha, 4 Dec 1971, Valls
et al. 1893 (US); Facenda Carauna—Bom Jesus,

Dutra 543 (R); Mun. Julio de Castilhos, between
Abacatu & Venado, 13 Nov 1976, 7M. Pedersen
11475 (C, CTES, K, MO, P); Mun. Vacaria, Jul
1954, Barreto 577 (BAA); S. Luiz, 24 Nov 1952,
Rambo 53471 (B, US); Faz Englert Sao Francisco
de Paula, 1 Jan 1952, Rambo 54704 (B, US); Serra
de Rocinha, Bom Jesus, 1200 m, 18 Jan 1950,
Rambo 45466 (B); 3 Feb 1953, Rambo 53816 (B,
US); Cambara do Sul, Itaimbezinho, 15 Dec 1973,
Hitchcock 28 (ICN); Taimbezinho, Nov 1977, Boe-
chat 41059 (ICN); Cambara do Sul, Itaimbezinho,
7 Jan 1977, Sampaio & Arzivenco 341, 362 & 404
(ICN); Bom Jesus—Faz. Arechavaleta, Cavauna,
Jan 1903, Dutra 538 (ICN); Bom Jesus—estr. Para
Vacaria, 15 km apos ponte de saida de B. Jesus, 30
Nov 1975, Longhi & Sampaio 368 (ICN). Santa
Catarina: Mun. Mafra, Campo Novo, 7 km from
Mafra, 9 Dec 1971, Smith & Klein 15761 (ICN, K,
R, US); Campo Novo, Mafra, 750 m, 12 Dec 1962,
Klein 3872 (K, US); Mun. Porto Uniao, 45 km S,

Festuca in South American Paramos 55

900-1100 m, 5 Feb 1967, Smith & Klein 10855
(US); Campo de Areao, Santa Cecilia, 1100 m, 19
Dec 1962, Reitz & Klein 14172 (US); Mun. Caga-
dor, 8 km N of Cacador, 950-1100 m, 7 Feb 1957,
Smith & Klein 10958 (US); Mun. Sao Joaquim,
source of Rio Capivaras, Serra do Oratorio, 10 km
E of Bom Jardim da Serra (Cambajuva), 1200 m,
16 Jan 1957, Smith & Reitz 10127 (US); Mun. Bom
Retiro, Fazenda Campo dos padres, 1650 m, 25 Jan
1957, Smith & Reitz 10428 (US); Mun. Marombas,
Curitibafios, Banhado, 900 m, 9 Jan 1962, Reitz &
Klein 11812 (US); Base of Morro Juco Prudente,
1 Jan 1946, J.R. Swallen 8089 (MO, US); Mun.
Bom Jardim de Serra, 10 km S of Bom Jardim at
Rio Capivaras, 15 Dec 1971, Smith & Klein 15805
(K, MO, R, US); Mun. Curitibafios, 28 Nov 1971,
Smith & Klein 15474 (K, R, US); Mun. Lajes, 8 km
S of Painel, 19 Dec 1971, Smith & Klein 15916 (K,
R, US); 27 km S of Lajes, 20 Dec 1971, Smith &
Klein 15936 (K, R, US); Mun. Chapeco, campo,
bog and pinheiral, Fezenda Campo Sao Vicente, 24
km W of Campo Ere, 900—1000 m, 26 Dec 1956,
Smith et al. 9389 (R, US); Mun. Ere, 6 km W of
Campo Ere, 26°22°S, 53°06’ W, 900—1000 m, 6 Dec
1964, Smith & Klein 13674 (K, P, R, US); Campo
de Chatorio, Ule 616 (W); Walde am Capirone,
Serra Geval, Mar 1891, Ule 1959 (P); Sao Biento,
22 Jun 1885, Schwaike s.n. (P); Mun. Rio Pardo,
70 m, Nov 1908, Jurgens 175 (W); Campos Dos
Padres, 22 Jan 1957, Rambo 60082 (B); Curitiba-
fos, Banhadoi, 900 m, 10 Jan 1962, Reitz & Klein
11948 (B, US). SAo Paulo: Campos do Jordao, in
paludosis, 1600 m, Feb 1946, Leite 3490 (US);
Mun. Sao Roque, 50 km W from, Sao Paulo, 850 m,
12 Feb 1987, Tsugaru & Otsuka B-2265 (MO).
PARAGDAY. Caazapa: Tavai, bosque cercano
al hospital, 28 Oct 1988, Degen 861 (FCQ, MO);
galery forest 1 km N of hospital, 26°10’S, 55°27’ W,
28 Oct 1988, Zardinii 7701 (FCQ, MO). Itapua:
Cordill. San Rafael, Estero Yukeri, Bernardi 18618
(MO). URUGUAY. Caticeiras Rivera, 11 Dec
1907, Berro 5994 (BAA). Tacuarembo. Campos
de Tacuarembo, terrenos arenosos, Arechavalet
s.n. (W); Montevideo, 1826, D’Orbygny 39 (P);
Cerro Largo, Cerro de Las Cuentas, 29 Dec 1938,
Rosengurtt B-2764 (US).

19. Festuca andicola Kunth, Nov. Gen. Sp. (quarto
ed.) 1: 153.1816. (Figs. 23, 24, 83C & D).
Festuca racemosa Willd. ex Spreng., Syst.
Veg. 1: 352. 1825, nom. inval. Tyre: Ecuador.
Carchi, Paramo de Puntas, 3220 m, Humboldt

& Bonpland s.n. (holotype: P!; isotypes: B!,
P!, US-2875373 fragm. ex P!).

Perennials with well developed rhizomes,
innovations extravaginal. Culms 50-70 cm tall,
erect, solitary, glabrous; nodes 2 or (3) in basal half.
Leaf sheaths membranous, brown, striate; auricles
absent; ligules 0.5-0.9 mm long, membranous to
coriaceous, apex truncate, ciliate; blades 10-17 x
0.1—0.4 cm, flat or conduplicate, green, abaxially
glabrous. Panicles 12—17 x 0.5—1 cm, contracted,
erect, elongate; branches mostly glabrous. Spikelets
7-9 mm long, florets 3 or 4(—5); rachilla glabrous or
with scattered hairs; glumes 1.4—-2.5 mm long, mem-
branous to coriaceous, oblong-lanceolate, purplish,
apex obtuse ciliate; lower glumes 1.4—1.8 mm long,
1-nerved; upper glumes 2—2.5 mm long, 3-nerved,
apex short-ciliate; lemmas 5—5.5 mm long, 5-nerved,
membranous to coriaceous, lanceolate, purplish,
upper third scabrous, apex acute or short—-awned,
the awn or mucro 0.5—0.7 mm long; callus glabrous;
paleas as long as the lemma, membranous, glabrous,
scabrous on keels, apex hairy; lodicules lanceolate,
acuminate; anthers 0.8—1.1 mm long; ovary apex
glabrous. Caryopses not seen.

Leaf blade anatomy.—Cross-sections with
7—9(-11) vascular bundles and 5—7(—9) ribs above;
sclerenchyma under abaxial epidermis discon-
tinuous, some fascicles extending to the vascular
bundles forming girders; adaxial epidermis with
scattered hairs, the hairs 0.05—0.09 mm long.

Observations.—Saint-Yves (1927, 180) men-
tions Festuca dissitiflora Steud., a nom. nud., as
belonging to F’ andicola. Examination of the voucher
of F. dissitiflora housed at US (W. Lechler 1829,
US-2875397 fragm. ex GOET!) confirms that this
species has nothing to do with F- andicola and that
it is asynonym of F! rigescens (J. Presl) Kunth. His-
torically, the name F’ andicola had been applied to
a heterogeneous group, and this group was divided
into two species by Stan¢cik (2003) where a new type
and name, F) soukupii, were designated. Festuca
andicola and F: soukupii are morphologically similar
to the Venezuelan species, F. e/viae, and the Peruvian
F: tenuiculmis Tovar. Festuca tenuiculmis differs
from the rest of the group by having ramified, opened
panicles. Festuca andicola differs from the others
by having solitary culms with rhizomes, whereas
F) soukupii is caespitose forming small bunches.

Distribution and habitat.— Festuca andicola
ranges from southern Ecuador to northern Colom-
bia. Tovar (1972) mentions F: andicola as being

56 Festuca in South American Paramos

Figure 24. Festuca andicola. A. Habit. B. Ligule. C. Spikelet. D. Glumes. E. Lemma. F. Lemma with palea and rachilla. G. Leaf
blade cross-section. A-G, Stancik 3551 (PRC).

Festuca in South American Paramos 57

present in Peru but no Peruvian specimens were
found during preparation of this revision. It occurs
in swamps and margins of streams in the grass
paramo zone and Andean mountain forests between
2900—3800(—4000) m. This species is associated, in
addition to many others, with communities of the
order Marchantio-Epilobietalia (Cleef 1981).
Additional specimens examined. COLOMBIA.
Antioquia: Mun. Urrao, Paramo Frontino, La
Cueva, 3450 m, 11 Sep 1986, Roldan et al. 396
(COL, HUA, MO); Llano Grande, 3400 m, 10 Sep
1986, Roldan et al. 307 (HUA). Boyaca: Mun.
Aquitania, Paramo Los Curies, 3500 m, 8 Feb
1999, D. Stancik 2190 (COL, FMB, PRC); Mun.
Mongui, Paramo de Laguna La Colorada, 3600 m,
21 Jan 1999, D. Stancik 2024 (COL, PRC); Mun.
Santa Rosa, Paramo Alto Lamadero, 3200 m, 30
Feb 1988, D. Stancik 1455 (COL, PRC); Paramo
de Pisba, carratera Socha—La Punta, 3570 m, 11 Jun
1972, Cleef 4376 (COL, U, US); Mun. Arcabuco,
way from Laguna Iguaque to Laguna Ojo de Agua,
3650 m, 15 Oct 1998, D. Stancik 948 (COL, FMB,
PRC); Mun. Toca, Paramo Cortadero, 5°30'N,
73°15’ W, 3350 m, 14 Nov 1998, D. Stancik 1365
(COL, FMB, PRC). Caldas: Mun. Manizales,
Parque Nacional Los Nevados, 4020 m, 18 Sep
1999, D. Stancik 3412 (COL, PRC); way from
Rio Nereidas to Casa del Cisne, km 5—7, 4000 m,
18 Sep 1999, D. Stancik 3399 (COL, PRC); Los
Nevados, Villamaria, 3 Feb 1985, Sahn & Hernan
1060 (FMB). Cauca: Macizo Colombiano, Valle
de las Papas, alrededores de Valencia, 2910 m,
11 Sep 1959, Idrobo et al. 3678 (COL). Cun-
dinamarca: Mun. San Juan de Sumapaz, Parque
Nacional Sumapaz, Alto de Toquilla, 3800 m, 15
Nov 1999, D. Stancik 3554 (AAU, COL, PRC); 15
Nov 1999, D. Stancik 3551 (COL, PRC); road from
San Juan to Usme, km 5—7, 3650 m, 15 Nov 1999,
D. Stancik 3547 (COL, PRC); Parque Nacional
Chingaza, mina Palacio, 3700 m, 11 Oct 1992,
Figueredo 119 (COL, HPUJ); 3700 m, 5 Aug 1992,
Figueredo 126 (HPUJ); Muncipio Fomeque, Valle
de los Frailejones, 3150 m, 12 Jun 1989, Bernal
& Jimenez 1127 (HPUJ); Monserrate, Aug 1859,
Lindig 14 (P); Alto San Juan, 4000 m, R. Fosberg
20749 (US). Meta: Paramo de Sumapaz, Hoya El
Nevado, Laguna La Guitarra, 3405m, 24 Jan 1973,
Cleef 8285 (COL). Narifio: Mun. Tuquerres, Vol-
can Azufral, road from Vereda San Roque to Laguna
Verde, km 3, 2650 m, 9 Mar 1999, D. Stancik 2775
(COL, PRC, PSO); El Encano, vereda Catapamba,
2900 m, 23 Mar 1999, D. Stancik 2983 (AAU,

COL, PRC, PSO, W); Cumbal, 3300 m, 16 Apr
1986, Benavides 6497 (COL, PSO). Norte de
Santander: Mun. Chilos, vereda El Hatico on road
Pamplona—Bucaramanga, 3500-3700 m, 1 Dec
2000, D. Stancik (COL, PRC). Risaralda: Laguna
del Otun, quebrada Juntas, 3750 m, 29 Aug 1985,
Sanchez & Hernandez 950 (FMB). Santander:
Paramo de Berlin, cerca del Picacho, 3290 m, 29
Sep 1966, Robinson & Beltran 3142 (US). ECUA-
DOR. Carchi: road Las Juntas—El Angel, km 14,
3400 m, 11 Mar 1992, S. Laegaard 101731 (AAU,
QCA, QCNE). Chimborazo: Mun. Riobamba,
Volcan Chimborazo, sector Cruce de Los Arenales,
01°28°14"S, 78°54’06"W, 4300 m, 20 Sep 2000,
D. Stancik 3712 (PRC, QCA); D. Stanctk 3705
(AAU, PRC, QCA). Cotopaxi: Volcan Cotopaxi,
road to Limpiapungo, 00°37’S, 78°27’ W, 3850
m, S. Laegaard 5866 (AAU, QCA). Imbabura:
Cordillera Oriental, Paramo de Angochagua, 2900—
3600 m, Acosta-Solis 18836 (US); Mun. Urcuqui,
road to Cerro Yanaurcu, 00°28'13’N, 78°18'45’W,
4100 m, 15 Oct 2000, D. Stancik 4092 (PRC, QCA).
Pichincha: Lloa, 3000 m, E. Asplund 7524 (AAU,
F, S); Paramo de Mojanda—Laguna Negra, 00°08’S,
78°16’ W, 3800 m, 14 May 1985, S. Laegaard 54327
(AAU, QCA, QCNE); Road Pifo—Papallacta, 3 km
W of Paso de la Virgen, 00°18°S, 78°14’ W, 3700-
3900 m, 7 Aug 1985, S. Laegaard 54892 (AAU,
QCA); Cerro Ungui, 2860 m, Hartman 7a (US);
Mun. Amiguafia, Volcan Pasachoa, 00°29°51°S,
78°29'25’W, 3700 m, 14 Sep 2000, D. Stancik 3685
(PRC, QCA); Mun. Amaguana, Volcan Pasachoa,
00°30'23”S, 78°29'28°W, 3350 m, 14 Sep 2000, D.
Stancitk 3683 (AAU, PRC, QCA). Tungurahua:
Cotalo, 2900 m, Acosta-Solis 9879 (US); Slope of
Mt. Tungurahua above Banos, 2800 m, E. Asplund
8427 (NY, S).

20. Festuca rubra L., Sp. Pl. 1: 74. 1753. (Figs.
21,25, 83A & B). Tyre: In paludosis prati regi
Upsalia, Anonymous (lectotype: GB!, designa-
ted by Jarvis et al., Watsonia 16:302. 1987.)

Loosely tufted and rhizomatous perennials
with extravaginal (rarely intravaginal) innovations.
Culms 20—50 cm tall, erect, glabrous; 1 node in
basal half. Leaf sheaths membranous to coriaceous,
brown, striate, glabrous or pilose, margins united
at base; auricles absent, ligules 0.3—0.5 mm long,
membranous to coriaceous; blades 7—15 cm long,
0.7—1 mm wide, conduplicate to involute, green,
sometimes glaucous, abaxially glabrous, apex

58 Festuca in South American Paramos

\

YZ

S /,
Co J
Ch) *

a a
Se aD

Figure 25. Festuca rubra. A. Habit. B. Ligule. C. Spikelet. D. Glumes. E. Lemma. F. Lemma with palea and rachilla. G. Leaf

blade cross-section. A—G, Stancik 3457 (PRC).

Festuca in South American Paramos 59

obtuse. Panicles ca. 10 cm long, 3—5 cm wide,
erect or spreading; branches scabrous. Spikelets
10-12 mm long, florets 5 or 6; rachilla glabrous
or with scattered hairs; glumes 4—5.5 mm long,
membranous to coriaceous, lanceolate, purple,
apex acute; lower glumes ca. 4 mm long, |-nerved;
upper glumes 5—5.5 mm long, 3-nerved; lemmas
6.5—7 mm long, 5-nerved, membranous to coria-
ceous, lanceolate, green to purplish, apex scabrous,
awned, the awn 2.5—3.5 mm long; callus glabrous;
paleas almost as long as the lemma, scabrous on
keels, apex hairy; lodicules oblong-lanceolate;
anthers 3.5—3.7 mm long; ovary apex glabrous.
Caryopses not observed.

Leaf blade anatomy.—Cross-sections with 7
vascular bundles and 5 ribs above; sclerenchyma
discontinuous under both abaxial and adaxial epi-
dermis, not forming girders; adaxial epidermis with
scattered hairs, the hairs 0.02 mm long.

Distribution and habitat.—This species is
introduced from Europe and rarely cultivated.

Additional specimens examined. COLOMBIA.
Caldas: Mun. Neira. Vereda Chupaderos, Sector
Campamentos, pasture, 3650 m, D. Stancik 3457
(COL, PRC). ECUADOR. Chimborazo: Paramo
de Urbina, 25 km N of Riobamba, ca. 2 km W of
Panamerican Hwy., 3500 m, 28 Jan 2000, S. Lae-
gaard et al. 20994 (AAU, LOJA, QCA, PRC).
Pichincha: Picalqui al S de Tabacundo, 2750 m,
Acosta-Solis 16358 (F, US).

21. Festuca soukupii Stancik, Folia Geobot.
Phytotax. 39(1): 103, f. 2, 1-5. 2004. (Figs.
26, 27, 83E & F). Type: Ecuador. Imbabura,
Mun. Cayambe, Volcan Cayambe, 00°31.6’N,
78°55.6°W, swamps below the refuge with
Loricaria sp., Festuca glumosa, Distichia
muscoides Nees & Meyen, etc., 4450 m, 20
Dec 2000, D. Stancik 4162 (holotype: PRC!;
isotypes: AAU!, QCA!).

Small tussocked perennials with intra- or
extravaginal innovations. Culms 10—50 cm tall,
erect, glabrous; nodes 1(—2) in basal half. Leaf
sheaths membranous, brown, striate, densely
hairy and appearing shaggy, margins free; auricles
absent; ligules 0.3—0.5 mm long, membranous
to coriaceous, apex obtuse, short-ciliate; blades
5-25 cm long, 0.5—0.8 mm wide, conduplicate to
involute, green, abaxially glabrous, apex obtuse.
Panicles 2.5—13(—17) cm long, 0.3—0.5 cm wide,
erect, contracted, elongate; branches hairy. Spike-

lets 6—8(—9) mm long, florets 3 or 4(—5); rachilla
glabrous or with short scattered hairs; glumes
1.2—2.5(—2.7) mm long, membranous to coriaceous,
ovate to lanceolate, purplish, apex obtuse, ciliate;
lower glumes 1.2—2 mm long, 1-nerved; upper
glumes 2—2.5(—2.7) mm long, 3-nerved, apex short-
ciliate; lemmas 4.5—5.5 mm long, membranous to
coriaceous, lanceolate, 5-nerved, purplish, papil-
lose, apex awned, sometimes scabrous, the awn
(0.5—)0.7—1.7 mm long; callus glabrous; paleas as
long as the lemma, glabrous, scabrous on keels,
apex hairy; lodicules oblong-lanceolate; anthers
0.8—1.1 mm long; ovary apex glabrous. Caryopses
lanceolate; hilum 3/5 of total length.

Leaf blade anatomy.—Cross-sections with 5—7
vascular bundles and 3—5 ribs above; sclerenchyma
under abaxial epidermis discontinuous, under adax-
ial epidermis absent [larger plants with 7—9(-11)
vascular bundles and adaxial schlerenchyma, some-
times forming 24 girders]; adaxial epidermis with
scattered hairs, the hairs 0.03—0.05 mm long.

Observations.—Festuca soukupii is mor-
phologically similar to F andicola. However,
F. andicola has well developed rhizomes (versus
tussock forming) and glabrous sheaths (versus
densely hairy).

Distribution and habitat.—Festuca soukupii
ranges from the Cordillera Central of northern
Colombia, through Ecuador to northern Peru. It is
known from humid grass paramo and super-paramo
vegetations between 3700-4500 m. In addition, this
species is found in large tufts along trails and roads
up to the forest zone between 2600-3200 m.

Additional specimens examined. COLOMBIA.
Caldas: Mun. Manizales, Parque Nacional Los
Nevados, Hda. La Esperanza, 3500-3700 m, 18
Sep 1999, D. Stancik 3416 (COL, PRC); Mun.
Neira, vereda Chupaderos, sector “Campamentos”’,
3650 m, 26 Sep 1999, D. Stancik 3455 (COL,
PRC); Paramo de Quindio, 3700-4200 m, 20 Aug
1922, Pennell 9951 (US). Cauca: Mun. Popayan,
Parque Nacional Puracé, Pilimbala, 02°22.1°N,
76°24.06'W, 3250 m, 6 Jul 2000, D. Stancik 3610
(COL, PRC); Macizo Colombiano, Valle de las
Papas, alrededores de Valencia, 2910 m, 11 Sep
1958, Idrobo et al. 3682 (COL, US). Narifio: Mun.
Cumbal, Volcan Nevado de Cumbal, NE slopes,
3700 m, 9 Mar 1999, D. Stancik 2744 (AAU, COL,
PSO, PRC); Mun. El Encano, between vereda
Catapamba and road Pasto—El Encano, 2800 m, 23
Mar 1999, D. Stancik 2981 (COL, PSO, PRC);
Plain N of Guachucal, 3000 m, 1 Mar 1986, Wood

Festuca in South American Paramos

F. Lemma. G. Lemma with palea and rachilla. H. Leaf
ik 4162 (PRC).

ca soukupii. A, B. Habit. C. Ligule. D. Spikelet. E. Glumes.
s-section. A, C—G, Stancik 3887 (PRC); B, Stanctk 3079 (PRC); H, Stané

Figure 26. Festu

blade cros

Festuca in South American Paramos 61

COLOMBIA

-80 -75

N

0 200 400 600 Kilometers
ee +

-70 -65 -60

Figure 27. Distribution of Festuca soukupii (¢), F-: chitagana ( A), and F. hatico (m).

5331 (COL, K). Risaralda: Mun. Pereira, Parque
Nacional Los Nevados, below Nevado del Cisne,
way from Casa del Cisne to Laguna de Otun, km 5,
4100 m, 19 Sep 1999, D. Stancik 3380 (PRC, W));
below Nevado Santa label, crossroads to Villa
Maria, Hda. Potosi and Casa del Cisne, 4000—
4100 m, 20 Sep 1999, D. Stancik 3373 (COL, PRC,
US). Tolima: N side of Nevado del Ruiz, 3800 m,
19 Feb 1984, Wood 4247 (COL, FMB, K); Mun.
Ibague, Nevado del Tolima, La Cueva, 04° 39’N,
76°19.5'°W, 3900 m, 8 Jun 2000, D. Stancik 3596
(COL, PRC). ECUADOR. Azuay: Parque Nacio-
nal Cajas, slopes near Laguna Luspa, 3700 m, 21
Apr 1990, P.M. Peterson 8858, C.R. Annable &
M.E. Poston (QCA, QCNE, US); along Rio Miguir,
3480 m, 21 Apr 1990, PM. Peterson 8870, C.R.
Annable & M.E. Poston (QCA, QCNE, US); around
lake Llaviaucu, 02°56’S, 79°10’ W, 3400 m, Oxford
Expedition 30 (K); El Pan, mountain zone, 2800 m,

Harlington 1197 (S); around Lagoon Taglacocha—
Tres Cruces, 02°47’S, 79°13°W, 3900-4000 m,
1 Sep 2000, D. Stancik 3858 (PRC, QCA); road
Cuenca—Saraguro, km 42, 03°10°S, 79°02°W,
3450 m, 19 Feb 1985, S. Laegaard 53580 (AAU,
QCA); road Cuenca—Limon, towards antennas,
03°10’S, 79°02’W, 3350 m, 21 Nov 1999, S. Lae-
gaard et al. 20824 (LOJA, AAU); Mun. Nabon,
antennas on the road Loja—Cuenca, 03°12’S,
79°02’ W, 3300-3450 m, 30 Aug 2000, D. Stancik
3802, 3808 (QCA, PRC). Bolivar: pass on road
Guaranda—Riobamba, 01°35’S, 78°50’ W, 4050—
4150 m, 10 Jul 1990, S. Laegaard 71737 (AAU,
QCNE); highway to Guaranda km 66.5 and to
Fecundo km 2.2, 4300 m, 03 May 1990, PM.
Peterson 8996 & C.R. Annable (MO, QCA, QCNE,
US); Simiatung, Hacienda Talahua, 3400 m, Pen-
land & Summers 583 (F). Caniar: road Canar—
Chunchi, 02°24’S, 78°59’ W, 3200 m, 05 Mar 1985,

62 Festuca in South American Paramos

S. Laegaard 53837 (AAU, QCA, QCNE); El
Tambo, road El Tambo—Ingapirca, 3400-3500 m,
D. Stanctk 3800 (PRC, QCA). Carchi: km 21.2
NE of El Angel and 27.8 km SW of Tulcan, above
Rio Bolo, 3320 m, 14 May 1990, PM. Peterson
9139, E.J. Judziewicz & R.M. King (MO, QCA,
US); km 11.4 NE of El Angel on road to Tulcan,
3240 m, PM. Peterson 9118, E.J. Judziewicz &
R.M. King (US); km 14 along road Las Juntas—El
Angel, 3400 m, 11 Mar 1997, S. Laegaard 101718
(AAU, QCA, QCNE). Cotopaxi: road Zumbahua—
Pujili, km 39, 00°53’S, 78°47°W, 3750-3800 m,
S. Laegaard 102087 (AAU); Paramo de Laguna
Salayampe, E of Latacunga, 00°56’S, 78°25’W,
4000-4100 m, S. Laegaard 54148 (AAU); Volcan
Cotopaxi, N slope, 00°39’S, 78°26’°W, 4200 m,
Sklenar & Kosteckova 1905 (AAU); Parque Nacio-
nal Cotopaxi, Laguna Limpiopungo, 00°40’S,
78°30’ W, 3800 m, 3 Apr 1982, Fegan & Falconi 9
(QCA); Falda NNW, 4180 m, 03 Jul 1986, Ehren-
burg 60 (QCA); Valley NW of Limpiapungo,
00°38°S, 78°28 W, 3900-4000 m, 25 Feb 1992,
S. Laegaard 101432 (AAU, QCA); N slope on the
left side of the road to refuge, 00°39’S, 78°26’ W,
4200 m, 19 Jul 1995, Sklenar & Kosteckova 85-8,
86-9, 87-6 (NY); Quebrada ca. 3 km de carratera,
00°40’S, 78°30'W, 3400 m, 08 May 1982, Bravo
56, 58 (QCA); railway station Cotopaxi, 3400 m,
E. Asplund 6478 (S, US); Cotopaxi, 2500 m,
E. Asplund 6400 (S); road Salcedo—Napo, km 27—
39, 4000-4100 m, Sparre 15708 (S); Mun. Chaupi,
NE slope of Volcan Illiniza, 37°59"S, 78°42°42°W,
4000-4050 m, 12 Oct 2000, D. Stancik 4023, 4028,
4031 (PRC, QCA); Mun. Lasso, volcan Cotopaxi,
00°39’6"S, 78; 78°30'55"W, 3530 m, 28 Sep 2000,
D. Stanctk 3108, 3128, 3879, 3887 (AAU, PRC,
QCA). Chimborazo: Volcan Chimborazo, E side,
01°28'S, 78°46’ W, 4260 m, Sklendr & Sklenarova
2262 (AAU); 4300 m, Sklenar & Sklenarova 2168
(AAU); km 42.7 SW of Ambato on Highway to
Guaranda, 4020 m, PM. Peterson 8973 & C.R.
Annable (MO, QCA, QCNE, US); km 10 E of Lago
Colta on road to Pallatango, 3725m, P.M. Peterson
9209, E.J. Judziewicz, R.M. King & P.M. Jorgensen
(K, MO, QCA, QCNE, US); between Urbina and
Mt. Chimborazo, 3600-4500 m, A.S. Hitchcock
21981 (US); 3750 m, E. Asplund 7892 (K, S);
3900 m, E. Asplund 8447 (P, S); Paramo de Urbina
km 25 N of Riobamba, ca. 2 km W of Panamerican
Hwy., 01°29°S, 78°42°W, 3500 m, S. Laegaard et
al. 20993 (AAU); S slope of Mt. Chimborazo,
3800 m, Fagerlind & Wibon 934-bis (S); km 9 NE

of San Juan de Velasco on road to Lago Colta,
3600 m, 21 May 1990, PM. Peterson 9235, E.J.
Judziewicz, R.M. King & P.M. Jorgensen (QCA,
QCNE, US); Daldal Valley km 10 E of Licto,
01°48’S, 78°32’ W, 3700 m, Ramsay et al. 104A (K);
00°15’S, 78°28°W, 3450 m, 19 Aug 1987, Ramsay
& Smith 245, 247 (K, QCA, QCNE); along Rio
Alao, 01°52’S, 78°30°W, 3200-3400 m, S. Lae-
gaard 55301 (AAU); near pass between Volcan
Chimborazo and Carihuairazo, 01°27’S, 78°48’ W,
4400 m, S. Laegaard & Sanchez 20027 (AAU);
Carihuasco above Mashahuasca, 01°29’S, 78°49’ W,
4500 m, S. Laegaard & Sklenar 20363 (AAU); road
Guamote—Macas km 15, 02°00’S, 78°40°W, 3750
m, S. Laegaard & Sklenar 20335 (AAU, LOJA);
Mun. Riobamba, volcan Chimborazo, sector Cruce
de los Arenales, 01°27°51”S, 78°53 58" W, 4150 m,
20 Sep 2000, D. Stancik 3713 (PRC, QCA). Imba-
bura: Hacienda Mojanda on road to Otavalo,
2900-3000 m, Sparre 13516 (AAU, S); Laguna
Grande, 00°08'N, 78°16’ W, 3725-3750 m, S. Lae-
gaard 52374A (AAU, QCA); Laguna Cuicocha,
3100 m, F. Asplund 20217 (S); Mun. Urcuqui, road
to Cerro Yanaurcu, 00°26'28"N, 78°15'24’W,
4100 m, 15 Oct 2000, D. Stancitk 4090B, 4093
(PRC, QCA); Mun. Cayambe, volcan Cayambe,
00°00'26.5’N, 78°01°21" W, 4300 m, 20 Oct 2000,
D. Stancik 4158, 4162 (PRC, QCA); Mun. Urcuqui,
road to Cerro Yanaurcu, 00°37°46'N, 78°41 °45’°W,
4100 m, 15 Oct 2000, D. Stancik 4094 (PRC,
QCA). Loja: km 10 along road to Fierra Urcu,
03°41’S, 78°18’W, 3400 m, S. Laegaard et al.
19066 (AAU, LOJA, QCA); Cerro de Arcos W of
road Manu—Zaruma, 03°34’S, 79°28’°W, 3500—
3600 m, S. Laegaard & Aguirre 20609 (AAU);
Paramo de Carboncillo, km 11 S of Ofia, 2800 m,
S. Laegaard 19739 (AAU, QCA); Mun. Saraguro,
road to Fierra Urcu, 03°41°S, 79°16’ W, 3000—
3100 m, 24 Aug 2000, D. Stancik 3787, 3780 (PRC,
QCA, US). Morona-Santigo: Hda. Huargualla—
Hda. San Eduardo, way to Parque Nacional Sangay,
02°00.25'S, 78; 27° W, 3700 mz; 20uigigiS 2
D. Stancik 3320 (PRC, QCA); Parque Nacional
Sangay, Plaza Culebrillas, 01°58°S, 78°25’W,
3500-3600 m, 22 Jul 1999, D. Stancik 3358, 3359
(PRC, QCA). Napo: Valle Vicioso E of Volcan
Cotopaxi, 3600 m, Holm-Nielsen & Balslev 23755
(AAU); El Tambo SE of Volcan Cotopaxi, 00°42’S,
78°18'W, 3650 m, S. Laegaard 55535 (AAU).
Pichincha: road Pifo—Pintag, 2.5 hours horseride
above Inga Moserat, 00°19’S, 78°17°W, 3950 m,
S. Laegaard 102268, 102237 (AAU, QCA); Paramo

Festuca in South American Paramos 63

de Guamani, 3960 m, 23 Nov 1991, Leon 1230
(QCA); 4200-4250 m, S. Laegaard 101384 (AAU);
4300 m, S. Laegaard & S.A. Renvoize 70519
(AAU); 4100 m, S. Laegaard 103107 (AAU,
QCA); road Quito—Santo Domingo near San Juan,
00°17°S, 78°37’ W, 3450-3500 m, S. Laegaard
52605, 52607 (AAU, QCA); Andinum Quitensium
Pichincha, Jameson 296 (P, W); from Pichincha,
4000ft, Jun 1859, Jameson 14 (W); Quito—Panecillo,
2900 m, E. Asplund 6020 (S, US); Mun. Otavalo,
shrubby margin and pajonal on the road from
Otavalo to Laguna Mojanda, 00°09°54’°N,
78°17°20°W, 3450 m, 19 Oct 2000, D. Stancitk 4111
(PRC, QCA); Nevado Cayambe, W side of the
volcano, 00°01°N, 78°01’ W, 4200 m, Sklenar &
Kosteckova 66-10 (US); road to Refuge, 4300 m,
S. Laegaard & S.A. Renvoize 70519 (AAU); Par-
amo de Trujillo between peaks of Illinizas and
Cotopaxi, 3450 m, Barclay & Juajibioy 7961 (MO,
US); Tunel de agua Quito—Papallacta, 00°22’S,
78°08 W, 16 Mar 1994, Aguirre & Merino 4219
(AAU, LOJA); E slope of Illiniza Sur, 4300 m,
Sklenar & Kosteckova 16-1 (US); 4200 m, Sklenar
& Kosteckova 15-5 (NY); Volcan Illiniza, N slope,
4000-4100 m, Sparre 15635 (S); road Lloa—Guagua
Pichincha, km 10, 00°13’S, 78°35’W, 4170 m,
Laegaard et al. 102737 (AAU); Rucu Pichincha,
4400 m, E. Asplund 17313 (S); NE slope, 4300 m,
Sklenay & Kosteckova 1-24 (NY); 4600 m,
E. Asplund 8606 (S); slope above Lloa, 3200 m,
E. Asplund 7558 (S); WNW slope of Antisana,
4600 m, Halloy B-54 (AAU); between La Libertad
and San Juan, 3200 m, E. Asplund 16259 (S); Canal
on W side of Volcan Atacazo, 00°20’S, 78°38’W,
3750-3800 m, S. Laegaard 55673 (AAU, QCA);
slopes of Ruminahui, 3900 m, Sparre 15871 (S);
Mun. Pifo, paramo de Guamani, 00°19’S, 78°15’ W,
3700 m, 19 Jun 1999, D. Stancik 3004 (AAU, PRC,
QCA); Mun. Amaguanfia, Volcan Pasachoa,
00°30°23”"S, 78°29'28"W, 3350 m, 14 Sep 2000,
D. Stancik 3686 (PRC, QCA). Tungurahua: km
10 S of Mocha, 3600 m, Harling et al. 10533 (GB);
Cotalo, 2900 m, Acosta-Solis 9884 (US); Acosta-
Solis 9877 (US); Mt. Carihuayrazo, 4400 m,
E. Asplund 8470 (S); Mun. Pillaro, Las Llanganatis,
around Aucacocha lagoon, 01°8°55’S,
78°20'00.4’"W, 3800 m, 28 Sep 2000, D. Stancik
3903 (PRC, QCA). Zamora—Chinchipe: Road
Loja—Zamora, ca. 2-6 km E of pass, 03°59’S,
79°09'°W, 2600 m, S. Laegaard 18736 (AAU,
LOJA, QCA, QCNE). PERU. Cajamarca: 52 km
N of Cajamarca on Hwy 3N towards Bambamarca,

small lagoon with Ranunculus and open grasslands
with Deyeuxia, 3780 m, 16 Mar 2000, PM. Peter-
son 14907 & N. Refulio Rodriguez (MO, US,
USM); Prov. Chota, Miracosta, entre Miracosta y
Pampa del Lirio, 3380 m, 11 Dec 2000, Vega et al.
10320 (F, MO); Pampa Larga, al N de la explota-
cion minera Yanacocha, 3900 m, 14 May 1994,
Vega 7143 (F); Jalca de Kumullca, ruta a Celendin,
3650 m, 7 Feb 1975, Vega et al. 1638 (F); Cerro
Sexcemayo, al W de Cajamarca, jalca graminosa,
3500 m, 4 Feb 1991, Vega 5419 (F); Prov. Celendin,
Cajamarca—Celendin road, especially near large
rock outcrops, 3000-3450 m, 28 May 1984, Smith
et al. 7314, 7321a, 7332 (MO); Celendin, Jalca de
Kumulca, mountain pass on the road Cajamarca—
Celendin, 07°02°31°S, 78°15°33”W, humid jalca
vegetation with tussock grasses, 3700 m, 26 Aug
2004, Sklenar & Cruz 8706 (PRC).

22. Festuca azucarica E.B. Alexeev, Bot. Zhurn.
(Moscow & Leningrad) 69: 1546. 1984. (Figs.
28, 29, 84A—D). Type: Colombia. Valle de
Cauca, Cordillera Central, cerro Pan de Azucar,
3700 m, 26 Feb 1969, J. Cuatrecasas, Espinal
& Ramos 27562 (holotype: US-278!; isotypes:
COL!, U-5508!).

Dense tussock forming perennials with intra-
vaginal innovations. Culms 70—80 cm tall, erect, gla-
brous; nodes 1 or 2 in distal half; cataphylls 0.5—1.5
cm long, short, coriaceous. Leaf sheaths coriaceous,
brown, glabrous; auricles absent; ligules 0.3—1(—1.5)
mm long, coriaceous, apex truncate, short-ciliate;
blades 30-50 cm long, 0.9-1.4 mm wide, condu-
plicate, rigid, olive-green, abaxially glabrous, apex
mucronate. Panicles 10—25 x 1—3(—6) cm, lanceolate
to ovate, mostly contracted; branches glabrous or
finely scabrous. Spikelets 9-10 mm long, lanceo-
late, florets 4—6; rachilla densely hairy; glumes
(2.5—)3—5(—6) mm long, coriaceous, narrowly lan-
ceolate, dark purple, sparsely scabrous; lower glumes
(2.5—)3—3.5(-4) mm long, 1-nerved; upper glumes
4-5(—6) mm long, 3-nerved; lemmas (5.5—)6—7.5 mm
long, 5-nerved, membranous to coriaceous, lanceo-
late, dark purple, scabrous and densely hairy, apex
entire, awnless or short-awned, the awn 0.3—0.5 mm
long; callus sparsely hairy; paleas as long as the
lemma, papillose, upper third hairy; anthers 2.5—3.5
mm long; ovary apex glabrous. Caryopses lanceolate;
hilum 1/2—3/5 as long as the grain.

Leaf blade anatomy.—Cross-sections with
9—15 vascular bundles and 7—14 ribs; sclerenchyma

64

Festuca in South American Paramos

2cm

1 cm

WW | Wi
Figure 28. Festuca azucarica. A. Stylized growth form. B. Habit. C. Cataphylls. D. Ligule. E. Spikelet. F. Glumes. G. Lemma.
H. Lemma with palea and rachilla. I. Leaf blade cross-section. A-I, Stancik 3377 (PRC).

Festuca in South American Paramos 65

COLOMBIA

N

400 600 Kilometers |

Figure 29. Distribution of Festuca colombiana (¢@), F: dasyantha (A), F: azucarica (@), and F-laegaardii (@).

under abaxial epidermis continuous or discontinu-
ous, united with all the vascular bundles, adaxial
sclerenchyma present, extending to all vascular
bundles; bulliform cells absent; epidermis densely
hairy adaxially, the hairs 0.4—1.3 mm long.

Observations.—Festuca azucarica is mor-
phologically similar to F’ pilar-franceii, a species
endemic to the Colombian Cordillera Oriental.
However, F: azucarica differs by having short cato-
phylls 0.5—1.5 cm long (versus 2—5 cm in F. pilar-
franceii) and longer lemmas (5.5—)6—7.5 mm long
(versus 4.5—6 mm). Festuca dasyantha, F. colom-
biana, F: laegaardii, and F: toca are all species in
F. sect. Cataphyllophorae that are characterized by
the presence of cataphylls.

Distribution and habitat.—Festuca azucarica
is endemic to the Colombian Cordillera Central
(Caldas, Quindio, Risaralda, Tolima, and Valle de
Cauca). It is one of the dominant species of the

matorral zone and grass paramo and occurs between
3500-4000 m. This species is found in many differ-
ent plant communities that contain Calamagrostis
recta (Kunth) Trin. ex Steud., Festuca aff. sub-
limis & Diplostephium rupestre (Kunth) Wedd.
(Cuatrecasas1934), Espeletietum hartwegianae-
Calamagrostiosum effusum (Cuatrecasas 1934),
Festuco dolichophyllae-Calamagrostietum effusae
(Salamanca-V. 1991) and Espeletia hartwegiana
subsp. centroandina Cuatrec. and Calamagrostis
recta (Cleef et al. 1983).

Additional specimens examined. COLOM-
BIA. Caldas: Mun. Manizales, Parque Nacional
Los Nevados, way from Casa del Cisne to Rio
Nereidas, km 5, 3800—4000 m, 18 Sep 1999,
D. Stancik 3404, 3405 (COL, PRC); Nevado El
Ruiz Libano, 4000 m, 21 Jul 1958, Barclay 6431
(COL, US); 4500 m, Aug 1984, Yepes-Agredo 760
(COL); 3400-3500 m, 17 Dec 1917, Pennell 3003

66 Festuca in South American Paramos

(US); Nevado de Santa Isabel, Quebrada de Leon,
3800 m, 24 Nov 1946, J. Cuatrecasas 23137 (U,
US, VALLE); Cabeceras del Rio Otun, Laguna
Taburetes, 3580 m, 24 Nov 1946, J. Cuatrecasas
23178 (US, VALLE); Nevado del Ruiz, 8 Oct
1983, Wood 4033 (AAU, K); Parque Nacional Los
Nevados, 5 May 1940, J. Cuatrecasas 9292 (COL).
Quindio: Mun. Salento, vereda Cocora, below
Nevado del Quindio, 22 May 1989, J. Luteyn et al.
13051 (COL); Pijao, Paramo de Chili, 3600 m, 17
Sep 1998, Correa et al. 129 (COL); Mun. Salento,
Paramo de Romerales, 3680 m, 30 Oct 1994, Velez
etal. 4514 (HUQ). Risaralda: Mun. Pereira, Parque
Nacional Los Nevados, below Nevado Santa Isabel,
crossroads to Villa Maria, Casa del Cisne and Hda.
Potosi, 4000—4100 m, 20 Sep 1999, D. Stanctk
339), 3376, 3377 (COL, PRE): Mun. Pereira,
Parque Nacional Los Nevados, 3510 m, 27 Jul
1958, Jaramillo-Mejia & Cleef 5738 (COL); Exped.
Botan. Novae-Granatae, sin. loc., Mutis 5545, 5555
(MA, US). Tolima: Mun. Santa Isabel, Quebrada
Africa, 3900 m, 19 Feb 1980, Jaramillo-Mejia et
al. 6181 (COL); 3800 m, 10 Feb 1980, Diaz-Pie-
drahita & Rangel 2037 (COL); Los Valles, finca La
Cascada, cabeceras del Rio Anaime, 4000 m, 10 Feb
1980, Echeverry 1973 (COL, TOLI); Mun. Ibague,
Nevado del Tolima, around La Cueva, 04°39’N,
75°19.5°W, grass paramo, 3900 m, 7—8 Jun 2000,
D. Stanctk 3599 (COL, PRC).

23. Festuca chitagana Stancik, Darwiniana
41(1-4): 130, f. 12g—k. 2003. (Figs. 27, 30).
Type: Colombia. Santander, Mun. Cerrito,
paramo del Almorzadero, km 15 on the road
from Cerrito to Chitaga, 3700 m, 25 Feb 1999,
D. Stancik & Medina 2577 (holotype: PRC!;
isotypes: COL!, FMB!).

Rhizomatous perennials forming small tus-
socks with intra- and extravaginal innovations.
Culms 60-80 cm tall, erect, glabrous; nodes 2-4
in distal half; cataphylls short, coriaceous, gray-
brown; Leaf sheaths membranous to coriaceous,
brownish-gray, glabrous, striate; auricles absent;
ligules 0.5—1 mm long, membranous to coriaceous,
apex truncate, ciliate; blades 15—20 cm long,
0.7-0.9 mm wide, involute, abaxially glabrous,
olive-green, apex obtuse. Panicles 10-12 cm ~x
9—12 cm, triangular, flexuous, nutant, branches
glabrous. Spikelets 6-9 mm long, lanceolate, flo-
rets 2 or 3; rachilla sparsely short-hairy; glumes
4—6.5 mm long, membranous, glabrous, apex acute;

lower glumes 4—5 mm long, narrowly lanceolate,
1-nerved; upper glumes 5.5—6.5 mm long, oblong-
lanceolate, 3-nerved; lemmas 6—7 mm long,
5-nerved, lanceolate, membranous to coriaceous,
hairy, apex acute or short-awned, the awn 0.5—1 mm
long; callus glabrous; paleas 4/5 as long as the
lemma, lanceolate, membranous, densely hairy;
lodicules 0.8—1 mm long, lanceolate; anthers
1—1.6 mm long; ovary apex glabrous. Caryopses
lanceolate; hilum 4/5 as long as the grain.

Leaf blade anatomy.—Cross-sections with 7—9
vascular bundles and 5—7 ribs above; sclerenchyma
under abaxial epidermis discontinuous; adaxial
sclerenchyma and girders absent; adaxial epidermis
sparsely hairy, the hairs ca. 0.07 mm long.

Observations.—Festuca chitagana 1s morpho-
logically similar to the Mesoamerican species F' tala-
mancensis Davidse that is known only from Costa
Rica. However, F) talamancensis has shorter lower
glumes (3.5—3.8 mm versus 4—5 mm in F: chitagana),
shorter upper glumes (5—5.3 mm versus 5.5—6.6), and
an oblong caryopsis (versus lanceolate) with a hilum
only 3/5 as long as the grain (versus 4/5).

Distribution and habitat.— This species is
endemic to Colombia and is known only from
the northern Cordillera Occidental (Paramo de
Frontino) and Oriental (Paramo del Almorzadero,
Sierra Nevada del Cocuy). It occurs in the grass
paramo zone with shrubs mostly on stony substrates
between 3400-4000 m.

Additional specimens examined. COLOMBIA.
Antioquia: Mun. Urrao, Paramo de Frontino,
Churumblum—La Mosca, 3400 m, 14 Nov 1984,
X. Londono et al. 550 (COL, MEDEL). Boyaca:
Mun. El Cocuy—Giiican, Parque Nacional El Cocuy,
Las Cabafias Kanwara, way to Pico Ritacuba along
river Playita, 4070 m, D. Stancik & Carvajal 1866
(PRC).

24. Festuca colombiana E.B. Alexeev, Bot.
Zhurn. (Moscow & Leningrad) 69: 1546, f.
2, 7-8. 1984. (Figs. 29, 31, 84K—F, 85A-B).
Type: Colombia.:Cundinamarca, Paramo de
Sumapaz, Chisaca, Laguna Negra, orilla sur,
pedregal humedo con Sphagnum sp., 3800 m,
11 Dec 1971, A.M. Cleef 181 (holotype: US-
2785658!; isotypes: COL!, P!, U!, VEN!).

Short rhizomatous perennials forming small
tussocks with intra- and extravaginal innovations.
Culms 40—80(—120) cm tall, erect, finely scabrous;
nodes | or 2(—3) in distal half; cataphylls small

Festuca in South American Paramos 67

Figure 30. Festuca chitagana. A. Stylized growth form. B. Habit. C. Cataphylls. D. Ligule. E. Spikelet. F. Glumes. G. Lemma.
H. Lemma with palea and rachilla. I. Leaf blade cross-section. A-I, Stancik & Carvajal 1866 (PRC).

68

Festuca in South American Paramos

———— —=
S< a
We, SSG SSE,

——=
—_——

<=

~S

— =

SS

=

————

a eS

Figure 31. Festuca colombiana. A. Stylized growth form. B. Habit. C. Cataphylls. D. Ligule. E. Spikelet. F. Glumes. G. Lemma
H. Lemma with palea and rachilla. I. Leaf blade cross-section. A-I, Stanctk 1889 (AAU).

Festuca in South American Paramos 69

(about 1—2 cm long), coriaceous. Leaf sheaths
membranous to coriaceous, glabrous, stramineous
to grayish, margins free; auricles absent; ligules
0.1—0.3(—0.5) mm long, coriaceous, apex truncate,
short-ciliate; blades (15—)20—30(-40) cm long,
0.4-0.6(—0.8) mm wide, conduplicate to involute,
rigid, abaxially glabrous, green, apex acute. Pan-
icles 10—20(—25) x 1-2 cm, contracted; branches
densely short-villose. Spikelets 9-12 mm long,
oblong-lanceolate, florets 4-6(—7); rachilla densely
pilose; glumes 3—5.5(—6.5) mm long, lanceolate,
coriaceous, purplish, upper 1/4 scabrous, apex acute;
lower glumes 3—-4(-4.5) mm long, 1-nerved; upper
glumes 4.5—5.5(—6.5) mm long, 3-nerved; lemmas
(5.5—)6—7(—7.5) mm long, 5-nerved, lanceolate,
membranous to coriaceous, purplish, papillose,
upper 1/3 sparsely scabrous, apex entire, mucronate
or awned, the awn up to 2 mm long; paleas as long as
the lemma, papillose, upper 1/3 hairy; lodicules ca.
1.1 mm long, lanceolate, acuminate; anthers 2—3.5
mm long; ovary apex glabrous. Caryopses oblong-
lanceolate; hilum 2/3 as long as the grain.

Leaf blade anatomy.—Cross-sections with 7—
9(—13) vascular bundles and 5—11 ribs; sclerenchyma
under abaxial epidermis discontinuous, rarely con-
tinuous, extending to all the vascular bundles, adaxial
sclerenchyma extending to every second vascular
bundle; bulliform cells absent; adaxial epidermis
densely hairy, the hairs 0.02—0.09 mm long.

Observations.—This species has a relatively
large area of distribution and plants from differ-
ent localities are morphologically variable. It is
morphologically similar to F! azucarica and F-
pilar-franceii, as mentioned previously.

Distribution and habitat.—Festuca colom-
biana is endemic to Colombia where it is known
only from the Cordillera Oriental (Boyaca, Cun-
dinamarca, Meta, Santander). It can be found
growing in grass paramos between 3200-4000 m.
This species is known from the following vegeta-
tion communities: Espeletio argentae-Arcytophyl-
letum nitidi (Rangel & Ariza 2000), Diplostephio
Phylicoidis-Espeletietum grandi-florae (Rangel &
Ariza 2000), Calamagrostietum planifoli-effusae
(Vargas & Zuluaga 1985), Espeletio argentae-
Calamagrostiosum effusum (Cuatrecasas 1934), and
Bartsio santolinifoliae-Calamagrostietum effusae
(Rangel & Ariza 2000).

Additional specimens examined. COLOM-
BIA. Boyaca: Mun. Arcabuco, Santuario Iguaque,
Laguna Iguaque, 3450-3500 m, 15 Oct 1998,
D. Stancik 928 (COL, FMB, PRC); Mun. Saboya,

Paramo Saboya, 3000 m, 24 Oct 1998, D. Stancik
1065,1069 (CO, FMB, PRC); Mun. Aquitania,
Paramo Sarna, 3400 m, 20 Jan 1999, D. Stancik
1969 (COL, FMB, PRC); D. Stancitk 1978 (COL,
PRC); 3100 m, 14 Jan 1999, D. Stanctk 1923 (COL,
FMB, PRC); D. Stancik 2003 (COL, PRC); Mun.
Aquitania, Paramo Los Curies, 3500 m, 8 Feb 1999,
D. Stanctk 2168, 2192 (COL, FMB, PRC); 3500 m,
8 Feb 1999, D. Stanctk 2174, 2218 (COL, FMB,
PRC); Mun. Samaca, Paramo Rabonal, 200 m, |
Nov 1998, 5D: Stancik 1323,,1324, 1325 (COL,
FMB, PRC); D. Stancik 1297 (COL, PRC); vereda
Ruchcal, 1 Nov 1998, D. Stanctk 1287, 1288, 1290,
1302, 1319, 1321 (COL, FMB, PRC); Mun. Aquita-
nia, road Aquitania—Sisvaca, km 6, 2900 m, 10 Feb
1999, D. Stanctk 2256 (COL, FMB, PRC); 3100 m,
13 Jan 1999, D. Stancik 1924 (COL, FMB, PRC,
W); 3350 m, 14 Jan 1999, D. Stancitk 1921 (COL,
FMB, PRC); Mun. El Cocuy, Parque Nacional
Nevado del Cocuy, Las Cabafias Kanwara, 3950
m, 30 Dec 1998, D. Stanctk 1875 (COL, FMB,
PRC); paramo on the road to Pico Ritcuba, km 2-3
from Cabanas, 4000 m, 30 Dec 1999, D. Stancik
& Carvajal 1811, 1848, 1891 (COL, FMB, PRC);
D. Stancik & Carvajal 1874 (COL, PRC); El
Cocuy—Giican, Parque Nacional El Cocuy, sector
Lagunillas, road from Casa Herrera to lagoons, km
3-4, 3950 m, 30 Dec 1998, D. Stancik 1851, 1854
(COL, FMB, PRC); Hac. La Esperanza—Valle de
los Frailejones, 3750 m, 30 Dec 1998, D. Stancik &
Carvajal 1783, 1787 (COL, FMB, PRC); Hac. La
Esperanza, rocky slope in the front of house, fields
and rest of the Polylepis forest, 3650 m, 30 Dec
1999, D. Stancik 1842 (COL, FMB, PRC); Mun.
Mongui, Laguna La Colorada, 3550 m, 21 Jan 1999,
D. Stancik 2025 (COL, FMB, PRC); Mun. Toca,
Paramo Cortadero, 05°30’N, 73°15’ W, 3300 m,
14 Nov 1998, D. Stancik 1352, 1355, 1357, 1391,
1402 (COL, FMB, PRC); Mun. Paipa, cuchilla El
Paramo, 3200 m, 3 Dec 1998, D. Stancik 1512,
1516, 1530, 1531, 1546 (COL, FMB, PRC); NW of
Duitama, Paramo de La Rusia, 3400 m, 2 Jul 1984,
Wood 4479 (COL, K). Cundinamarca: El Chico,
Bogota, Dec 1946, Black 46-605 (NY); Mt. Guada-
lupe above Bogota, bushy slope, 2800-3300 m, 9
Dec 1917, Pennell 1923 (NY); Paramo de Guasca,
entre Guasca y Gacheta, 3320 m, | Jun 1959, Bar-
clay & Juajibioy 6517 (MQ); Paramo entre Cogua y
San Cayetano, cercania de la Laguna Seca, 3700 m,
12 Nov 1972, Cleef 6258 (COL, US); Chapinero
near Bogota, 3000-3100 m, 23 Sep 1917, Pennell
2018 (MO, US); Paramo de Monserrate, 3300 m,

70 Festuca in South American Paramos

12 Sep 1987, Sanchez 283 (COL); Parque Nacional
Sumapaz, Laguna Chisaca, 3700-3800 m, 10 Feb
1961, Pinto-Escobar & Hernandez 523 (COL);
3706 m, Pedraza et al. 491 (COL, PRC); Cuchilla
La Rabona, 4020 m, 11 Feb 1972, Cleef 1587 (COL,
U); Mun. Usme, Laguna Negra, 3750 m, 20 Feb
1986, Rangel & Aguirre 3720 (COL); Paramo de
Chisaca, 3910 m, 11 Nov 1958, Barclay & Juajibioy
6109 (COL, MO, US); 3650-3700 m, 29 Oct 1959,
J. Cuatrecasas & Jaramillo-Mejia 25742 (COL);
Laguna Negra, 3800 m, 11 Dec 1971, Cleef 181
(COL, P, U, US); 3500 m, 16 Jul 1998, D. Stancik
303 (COL, PRC); Mun. Guatavita, vereda Carbo-
nel Alto, 3200 m, 30 Oct 1999, D. Stanctk 3500
(COL, PRC); Guadalupe, 3000 m, Jul 1913, Apol-
linaire & Arthur 121 (US); Mun. Bogota, Alto de
la Viga, 3550 m, 1 Nov 1999, D. Stanctk 3538,
3540, 3542 (COL, PRC); Mun. Tausa, cuchilla
Los Cuervos—Laguna Verde, 3650 m, D. Stanctk
3496 (COL, PRC); Mun. Choachi, Paramo de
Cruz Verde, laguna Verjon, 3450 m, 19 Oct 1985,
Castellanos et al. 3 (BOG); 3720 m, Aug 1935,
Garcia-Barriga 1016, 1019 (AAU, COL); Choachi,
2900 m, Lindig 1013 (K, P); 2900-3200 m, Aug
1859, Lindig 1053 (US); 3200 m, 18 Dec 1915,
Apollinaire & Arthur 9 (US); Paramo de Palacio,
2 km al lado de Mina, 3750 m, 29 Nov 1972, Cleef
& Urribe 6694 (COL, U, US); Paramo de Guasca,
E] Santuario, 3200 m, 26 Jan 1972, J. Cuatrecasas
3550 (MA); Mun. San Juan de Sumapaz, road to
Usme, km 5-7, 2950 m, 1 Nov 1998, D. Stanctk
1289 (COL, PRC); 15 Nov 1999, D. Stancik 3544,
3545, 3546, 3560 (COL, PRC); Mun. Santa Rosa de
Viterbo, Paramo Alto Lamadero, Laguna Sagrado
Corazon, 3200 m, 30 Nov 1998, D. Stancitk 1436
(COL, FMB); Mun. Siachoque, paramo Siachoque,
5°28’51"N, 73°12'44’W, 23 Jan 1999, D. Stancik
2073 (COL, PRC); Las Tronaderas, 3770 m, 24 Jan
1999, D. Stancik 2063 (COL, PRC); vereda Car-
nichogue Arriba; 3400 m, 24 Jan 1999, D. Stanctk
2053 (COL, PRC); 23 Jan 1999, D. Stancik 2052
(PRC, COL); 1998, D. Stancik 2408 (COL, PRC);
Vereda San Antonio, 3600 m, 16 Oct 1999, D.
Stancik 3491, 3492, 3493, 3494 (COL, PRC);
cuchilla El] Muchacho, 3600 m, 16 Oct 1999, D.
Stancik 3480 (COL, FMB, PRC); Mun. Zipaquira,
paramo Guerrero, 3500 m, 16 Oct 1999, D. Stancik
348] (COL, FMB, PRC); Paramo de Chisaca, way
to Chisaca, near Rio Tunjuelito, 3750 m, 20 Sep
1966, T.R. Soderstrom 1229 (COL, K, TULV);
Paramo El Tablazo, frequent in recently burned
paramo, 3200 m, 2 Apr 1983, Wood 3612 (K).

Meta: Macizo de Sumapaz, Hoya de la Quebrada
Clarincito, Los Frailes, 3720 m, 2 Jul 1981, Diaz-
Piedrahita 2367 (COL). Santander: Mun. Cer-
rito, Paramo del Almorzadero, vereda Mortyno—La
Cascada, 3400 m, 25 Feb 1999, D. Stancik 2527,
2550, 2562, 2563, 2564, 2565 (COL, FMB, PRC);
25 Feb 1999, D. Stancitk 2560, 2561 (COL, PRC);
Mun. Concepcion, vereda Juradito, Paramo de Gal-
lina, 3200 m, 24 Feb 1999, D. Stancik 2487, 2507,
2508, 2511, 2512 (COL, FMB, PRC); Vicinity of
Vetas, rocky hillside, 3100—3250 m, 20 Jan 1927,
Killip 17329 (US); Exped. Botan. Novae-Granatae,
sin. loc., Mutis 5570 (MA).

25. Festuca dasyantha Kunth, Nov. Gen. Sp.
(quarto ed.) 1: 154-155. 1816. (Figs. 29, 32,
85C & D). Tyre: Ecuador. Cotopaxi, in Montis
Cotopaxi, 4090 m, Humboldt & Bonpland s.n.
(holotype: P!; isotypes: B!, BAA-1187 fragm.
ex B!, B-W-2071!, US-2875395 fragm!, W!).

Tufted perennials with short rhizomes and
intravaginal innovations. Culms 40-100 cm tall,
erect, glabrous; nodes 2 or 3(—4) in basal half;
cataphylls 1-4 cm long, coriaceous, dark brown,
striate. Leaf sheaths membranous to coriaceous,
brown, striate, glabrous or scabrous; ligules 0.30.5
mm long, membranous to coriaceous, margins
ciliate, apex truncate; blades 15—35 cm long, 0.7—
1.4(—2.2) mm wide, mostly conduplicate, green,
abaxially glabrous. Panicles 15—30 x 1—3 cm, nar-
row, erect, branches glabrous or sparsely scabrous.
Spikelets 10-11 mm long, florets 4 or 5; rachilla
densely pilose; glumes 3—5 mm long, lanceolate,
membranous to coriaceous, green, distal 1/3
often densely hirsute, apex acute; lower glumes
3—3.5 mm long, 1-nerved; upper glumes 4.5—5 mm
long, prominently 3-nerved; lemmas 5.5—6 mm
long, lanceolate, 5-nerved, membranous, green,
densely hairy, apex entire, acute or short-awned, the
awn, up to 1 mm long; paleas as long as the lemma,
glabrous, upper 1/3 hirsute; lodicules lanceolate,
acuminate; anthers 2.3—3.3 mm long; ovary apex
glabrous. Caryopses not seen.

Leaf blade anatomy.—Cross-sections with
9-17 vascular bundles; sclerenchyma discontinuous
under both abaxial and adaxial epidermis, extending
to every other or nearly all vascular bundles forming
girders; bulliform cells not observed; adaxial epi-
dermis densely hairy, the hairs 0.7—1.3 mm long.

Observations.—Festuca dasyantha differs
from all other species in F: sect. Cataphyllophorae,

Fl

Festuca in South American Paramos

ESAS eS SSS
_——SSS

\s)

Figure 32. Festuca dasyantha. A. Stylized growth form. B. Habit. C. Cataphylls. D. Ligule. E. Spikelet. F. Glumes. G. Lemma.
H. Lemma with palea and rachilla. I. Leaf blade cross-section. A-H, Laegaard 101260 (AAU); I, Mille 664 (QPLS).

q2 Festuca in South American Paramos

such as F: procera, F: colombiana, and F- laegaardii,
by having glumes and lemmas bearing long hairs.

Distribution and habitat.—Festuca dasyantha
is endemic to central Ecuador (Carchi, Imbabura,
Pichincha). It occurs in grass paramos between
2800-3000 m.

Additional specimens examined. ECUADOR.
Carchi: Road el Angel—Mira, km 3, 00°37’'N,
77°58'W, 2950 m, S. Laegaard 101260 (AAU,
QCA). Imbabura: Entre Yuquin y Cebadal,
Sigsipamva, 2800 m, Acosta-Solis 18729 (US).
Pichincha: Pifo, pascuis andinis et subandinis,
3000 m, Mille 664 (QPLS); 2700 m, Mille 659
(QPLS); between Pifo and Los Coralles, 2800 m,
E. Asplund 6978 (K, NY, S, US); Quito—Panecillo,
Sodiro s.n. (MO, NY, QPLS, US).

Note.—Specimen Mutis 5554 (US ex MA)
comes from Ecuadorian collection of F.J. Caldas and
dating to the period of Mutis’s stay in Colombia.

26. Festuca hatico Stancik, Darwiniana 41(1-4): 119,
f. 121-p. 2003. (Figs. 27, 33, 86A & B). Type:
Colombia. Norte de Santander, Mun. Chilos,
Vereda Hatico on road Bucaramanga—Pamplona,
3500-3700 m, 1 Dec 2000, D. Stancik 4290
(holotype: PRC; isotype: COL).

Tussocked perennials with rhizomes and intra-
and extravaginal innovations. Culms 60—70 cm tall,
erect, glabrous; nodes 2 or 3 in distal half; cataphylls
coriaceous, gray-brown, striate. Leaf sheaths coria-
ceous, brownish-stramineous, glabrous, striate;
auricles absent; ligules 0.3—0.5 mm long, mem-
branous to coriaceous, bilobed; blades 30-40 cm
long, 0.7—0.9 mm wide, conduplicate to involute,
abaxially glabrous, green, apex obtuse. Panicles
10-20 x 0.5—1 cm, contracted, narrow; branches
finely scabrous. Spikelets ca. 9 mm long, lanceo-
late, florets 3 or 4; rachilla short, papillose; glumes
3.2-5 mm long, coriaceous, glabrous, apex acute;
lower glumes 3—3.6 mm long, narrowly lanceolate,
l-nerved; upper glumes 4.5—5 mm long, lanceo-
late, 3-nerved; lemmas 6—6.5 mm long, 5-nerved,
membranous to coriaceous, oblong-lanceolate,
papillose, apex scabrous and two-dentate, awned,
the awn 0.3—-0.7 mm long; callus glabrous; paleas
4/5 as long as the lemma, lanceolate, membranous,
papillose, upper 1/2 hairy; lodicules ca. 1.7 mm
long, triangular; anthers ca. 3.5 mm long; ovary
apex glabrous. Caryopses not seen.

Leaf blade anatomy.—Cross-sections
usually with 9 vascular bundles and 7 ribs above;

sclerenchyma under abaxial and adaxial epidermis
discontinuous, extending to some vascular bundles
forming girders; adaxial epidermis hairy, the hairs,
0.4—0.9 mm long, often numerous.

Observations.—F estuca hatico is morphologi-
cally similar to Festuca colombiana. However, the
latter species has wider panicles (1—2 cm versus
0.5—1 cm), spikelets with 4—7 florets (versus 3 or
4), pilose rachillas (versus papillose), and paleas as
long as the lemmas (versus 4/5 as long).

Distribution and habitat.—Festuca hatico is
endemic to the northeastern Andes and is known
from Colombian Cordillera Oriental (Norte de
Santander, Santander) and Venezuela (Lara). It can
be found growing in shrubby patches of the paramo
zone between 3100-3900 m. —

Additional specimens examined. COLOMBIA.
Norte de Santander: Mun. Chilos, Vereda Hatico
on road Bucaramanga—Pamplona, 3500-3700 m,
1 Dec 2000, D. Stancik 4290 (COL, PRC).
Santander: East slope of Paramo de las Coloradas,
above La Baja, 3900 m, 27 Jan 1927, Killip 18479
(US). VENEZUELA. Lara: Mun. Humocaro Alto,
Parque Nacional Dinira, 9°35’39"N, 70°07 12’W,
3170 m, D. Stancik 4289 (CAR, COL, PRC).

27. Festuca laegaardii Stancik, Folia Geobot. Phy-
totax. 39(1): 107, f. 4, 6-10. 2004. (Figs. 29,
34, 86C—F). Tyre: Ecuador. Tungurahua/Napo,
Mun. Pillaro, Las Llanganatis, 01°09°37.5’S,
78°14'50.8"W, Valle de los Frailejones, mar-
gins of the swamp dominated by Carex sp. and
dryer shrub formation, 3500 m, 28 Sep 2000,
D. Stancik 3983 (holotype: PRC!; isotypes:
AAU!, QCA!).

Rhizomatous and loosely caespitose perennials
with extravaginal innovations. Culms (15—)30—100
cm tall, erect, glabrous; nodes 2 or 3(—5) in dis-
tal half; cataphylls present, short, membranous,
brown, striate. Leaf sheaths membranous, purplish-
brown, striate, hairy; ligules 1—-1.5(—2) mm long,
membranous to coriaceous, apex truncate, ciliate;
blades 20—25 x 0.5—1.1 cm, conduplicate, abaxially
glabrous, rarely hairy. Panicles (5—)10—15(—18) x
1-2 (-11) cm, compressed, rarely flexuous,
branched; branches densely hairy. Spikelets (7.5—)
8—10(—12) mm long, florets (3—)4 or 5(—6); rachilla
sparsely pilose; glumes 2.8—5.5(—6) mm long,
coriaceous, narrowly lanceolate, purple, upper 1/3
densely hairy, apex acute; lower glumes 2.8—3.7
(-4) mm long, 1-nerved; upper glumes 4.5—5.5

Festuca in South American Paramos a)

~ as Ge eee
\ UN

x
Ae! Se

— KA bk

Figure 33. Festuca hatico. A. Stylized growth form. B. Habit. C. Cataphylls. D. Ligule. E. Spikelet. F. Glumes. G. Lemma.
H. Lemma with palea and rachilla. I. Leaf blade cross-section. A-I, Stanctk 4290 (PRC).

74 Festuca in South American Paramos

Figure 34. Festuca laegaardii. A. Stylized growth form. B. Habit. C. Cataphylls. D. Ligule. E. Spikelet. F. Glumes. G. Lemma.
H. Lemma with palea and rachilla. I. Leaf blade cross-section. A-I, Stanctk 3983 (PRC).

Festuca in South American Paramos 1

(6) mm long, 3-nerved; lemmas 5.5—6.5 mm long,
5-nerved, lanceolate, membranous to coriaceous,
green-purple, apex acute, entire, upper 1/3 hairy;
paleas almost as long as the lemma, glabrous or
inconspicuously scabrous, apex hairy; lodicules
obovate; anthers 2.4—2.8(—3) mm long; ovary apex
glabrous. Caryopses not observed.

Leaf blade anatomy.—Cross-sections with
7-11 vascular bundles and 3-7 ribs above; scleren-
chyma under both abaxial and adaxial epidermis
discontinuous, extending to some vascular bundles
forming girders; bulliform cells not observed;
adaxial epidermis usually with scattered hairs, the
hairs 0.45—0.7 mm long.

Observations.—Festuca laegaardii is mor-
phologically similar to Festuca dasyantha. How-
ever, the latter species has densely hairy adaxial
epidermis(versus scattered hairs in F’ /aegaardii),
lemmas densely hairy throughout (versus only
short hairy on the upper third), shorter ligules (0.3—
0.5 mm long), and glabrous panicle branches.

Distribution and habitat.— Festuca laegaardii
is endemic to Ecuador and is known only from
Azuay, Cafiar, Carchi, and Napo. This species
occurs in humid and swampy patches in grass
paramos between 2900-3400 m.

Additional specimens examined. ECUADOR.
Azuay: Road Gualaceo—San Juan Bosco, ca. 3 km
from pass, 03°00’S, 78°39°W, 3250-3350 m, S.
Laegaard 53965 (AAU, QCA); Road Cuenca — Sara-
guro, km 68, 03°20’S, 78°10’°W, 3350 m, S. Lae-
gaard 53597 (AAU, QCA). Cafiar: Road Biblian—

Canar, 3400 m, Harling et al. 8635 (GB). Carchi:

Alor—Hda. San Rafael, 2900-3000 m, Acosta-Solis
21031 (US). Morona-Santiago: Parque Nacional
Sangay, Plaza Culebrillas, 01°58’S, 78°25 ’W, forest
grassy patches, 3500-3700 m, D. Stancik 3365 (PRC,
QCA). Napo: Laguna San Marcos NE of Volcan
Cayambe, 00°07’N, 77°58’ W, 3370 m, B. Allgaard
et al. 34039 (AAU, PRC); Papallacta—NE de la
laguna principal, 3400 m, Jaramillo et al. 11855
(QCA); Jaramillo et al. 11856 (MO, QCA).

28. Festuca pilar-franceii Stancik, Darwiniana
41(1-4): 121, f. 14a—-e. 2003. (Figs. 35, 37,
87A & B). Tyre: Colombia, Cundinamarca,
Mun. Bogota D.C., sector Santa Rosa—Laguna
Chisaca, 3500-3650 m, 7 Aug 1998, D. Stancik
3585 (holotype: PRC!; isotype: COL!).

Densely tussocked perennials with intravaginal
innovations. Culms 70—100(—120) cm tall, erect,

glabrous; nodes 1 or 2 in distal half; cataphylls
2-5 cm long, membranous to coriaceous. Leaf
sheaths membranous to coriaceous, brown-stramin-
eous, glabrous; auricles absent; ligules 0.1-0.3 mm
long, coriaceous, apex truncate; blades 30-60 cm
long, 0.7—1.1 mm in diameter, conduplicate to
involute, abaxially glabrous, green, apex obtuse.
Panicles (12—)15—20(—25) x 1-7 cm, oblong-
lanceolate, branched, flexuous, erect; branches
glabrous or finely scabrous. Spikelets (8—)9—-11 mm
long, lanceolate, florets 4—6(—7); rachilla densely
pilose; glumes 2—5 mm long, narrow lanceolate,
coriaceous, glabrous, purple or purplish-green, apex
acute; lower glumes 2—3(—3.5) mm long, 1-nerved;
upper glumes 3—5 mm long, 3-nerved; lemmas
4.5—-6 mm, lanceolate, 5-nerved, membranous to
coriaceous, purplish-green, scabrous or hairy, apex
mucronate or short-awned, the awn < | mm long;
paleas as long as the lemma, papillose, upper 1/3
hairy; lodicules lanceolate, acuminate; anthers
2.5—3.5 mm long; ovary apex sparsely hairy. Cary-
opses not seen.

Leaf blade anatomy.—Cross-sections with 11—
15 vascular bundles and 9-13 ribs; sclerenchyma
under abaxial epidermis continuous, extending to
all the vascular bundles forming girders; bulliform
cells absent; adaxial epidermis densely hairy, the
hairs 0.8—1.2 mm long.

Observations.—Festuca pilar-franceii is mor-
phologically similar to F. azucarica, a species from
the Cordillera Central. In the Cordillera Oriental,
F- cleefiana is another species that is morphologically
similar to F) pilar-franceii. However, F: cleefiana
lacks cataphylls, has longer ligules [(0.5—)1—2(—2.5)
mm], and has longer lemmas (6—8 mm).

Distribution and habitat.—This species is
endemic to the Colombian Cordillera Oriental
(Macizo de Bogota, Paramo de Sumapaz). It occurs
in the swampy patches and margins of lagoons in
grass paramos between 3600-4000 m. Festuca
pilar-franceii is associated with two communities:
Baccharis revoluta & Cortaderia cf. nitida (Sturm
& Rangel 1985) and Chusquea tesellata, Espeletia
grandiflora, and Calamagrostis effusa (Sturm &
Rangel 1985).

Additional specimens examined. COLOM-
BIA. Cundinamarca: Cabrera, Paramo de Cruz
Verde, 3720 m, VIII 1935, Garcia-Barriga 1018
(COL); Sumapaz, a 2 km de la division de car-
retera a San Juan, 3960 m, 3 Oct 1978, Rangel
1650 (COL); km 51-60 via San Juan, 10 Feb 1986,
Torres-Romero & Lozano 2982 (COL); Laguna

76

Festuca in South American Paramos

ida A
1 eho elles

\ x | )

V7

———
per
—__

SS
SX
=

—_

=

x mS NS ~
wee a <a t

A _
LL DV
GLP Ak

Figure 35. Festuca pilar-franceii. A. Stylized growth form. B. Habit. C. Cataphylls. D. Ligule. E. Spikelet. F. Glumes. G. Lemma
H. Lemma with palea and rachilla. I. Leaf blade cross-section. A-I, Stanctk 315 (PRC).

Festuca in South American Paramos ol.

Chisaca, 3625m, 25 Aug 1972, Cleef 5259 (COL,
U, US); Paramo Chisaca, 3750-3960 m, Oct 1966,
T.R. Soderstrom 1307 (US, TULV); near headwa-
ter of Rio San Juan, 18 km E of Cabrera, 4°5'N,
74°12’W, 4000 m, 11 Aug 1943, R. Fosberg 20745
(US); Macizo de Bogota, Laguna Negra, 3720 m,
11 Sep 1961, J. Cuatrecasas & Jaramillo-Mejia
25865 (COL, US); 3600-3700 m, III 1973, Cleef
3605 (COL, P, U); 16 Jul 1998, D. Stanctk 210, 217,
Pie 422), 226,231, 241, 248, 250) 256, 2357,
295,264,205, 267, 208, 270,277, 276, 302, 311,
mie wis 4, 315, 316, 317,318, 319, 320, 32]
(COL, PRC); 9 Aug 1999, D. Stancik 3559, 3562,
3563, 3564, 3565, 3569, 3568, 3579, 3580, 3583,
3584, 3586, 3587, 3589, 3590 (COL, PRC); Mun.
Bogota, Laguna Chisaca, 9 Aug 1999, D. Stancik
wae 07. 370, 3972, 3973, 3574, 3575, 3576,
3979.55 76, 3901, 3582, 3565, 3388 (COL, PRC);
6 Jan 1997, D. Stancik 491 (COL, PRC); 15 Nov
1999, D- Stanctk 3559 (COL, PRC, US); 7 Aug
1998, D. Stancik 228 (COL, PRC); D. Stancik 3494
(COL, PRC); D. Stancik 3561 (COL, PRC); Parque
Nacional, Sumapaz, vereda Santa Rosa, ladera de la
quebrada Bijuacales, localidad 20, 3500 m, 07 Aug
1998, Pedraza et al. 228, 242, 278 (COL, PRC);
Paramo de Sumapaz, between St. Rita and San Juan,
3600 m, 15 Jul 1984, Wood 4539 (K, US). Meta:
Paramo Sumapaz, Hoya de Quebrada Sitidles, 0.5
km al SW de Laguna La Primavera, 3550 m, 26 Jan
1972, Cleef 1048 (COL, U); Cerro Nevado de Suma-
paz, 3615m, 29 Jan 1972, Cleef 1272 (COL); lado
SW, 3590 m, 12 Jan 1973, Cleef 7704 (COL).

29. Festuca procera Kunth, Nov. Gen. Sp. (quarto
ed.) 1: 154. 1816. (Figs. 36, 37, 87C—F).
Diplachne procera (Kunth) Spreng., Syst.
Veg. 1: 351. 1825. Festuca orgyalis Willd. ex
Spreng., Syst. Veg. 1: 351. 1825, nom. inval.
Festuca scabra Willd. ex Steud., Nomencl.
Bot. (ed. 2) 1: 632. 1840, nom. inval. Festuca
procera Nees & Meyen, Nov. Actorum Acad.
Caes. Leop.-Carol. German. Nat. Cur. 19(1):
166-167. 1843, nom. illeg. hom. Festuca
orgyalis Willd. ex E. Fourn., Biol. Cent.-
Amer., Bot. 3: 582. 1885, nom. illeg. superfl.
Type: Ecuador. Pichincha, Crescit locis alsis,
subfrigidis regni Quitensos prope Chillo, San
Antonio de Lulumbamba et Lloa, inter alt.
1280-1470 hexap., floret Aprili, Majo, Hum-
boldt & Bonpland s.n. (lectotype: B!, designa-
ted by Alexeev, Novosti Sist. Vyss. Rast. 23:
12. 1986; isolectotype: P!).

Festuca peruviana E.B. Alexeev, Bjull. Moskovsk.
Ob8S¢. Isp. Prir., Otd. Biol. 89(4): 114. 1984,
hom. illeg. Tyre: Peru. Tabina, Jul 1854, Lech-
ler 2114 (holotype: LE!; isotype: S).

Festuca sublimis Pilg., Bot. Jahrb. Syst. 25(5):
718. 1898. Type: Ecuador. Azuay, Pucara de
Chisalo, 1 Feb 1874, A. Sttibel 297 (holotype:
B!; isotypes: BAA-1341 fragm. ex B!, US-
81580 ex B!).

Tussocked perennials with intra- and extra-
vaginal innovations. Culms 80-170 cm tall, erect,
glabrous; nodes 1; cataphylls 1-4 cm long, mem-
branous to coriaceous, grayish-brown, striate. Leaf
sheaths membranous to coriaceous, brownish-gray,
glabrous or with short hairs; ligules 0.3—-0.7 mm
long, coriaceous, margins ciliate, apex truncate;
blades 30—50 cm long, 0.8—1.6 mm wide, condu-
plicate, abaxially glabrous or sometimes scabrous,
green, apex obtuse. Panicles 15—25 x 3—15(—20) cm,
oblong to ovate, lax, densely flowered; branches
glabrous or rarely sparingly scabrous. Spikelets
11-15 mm long, oblong-lanceolate, florets (4—)6
or 7; rachilla densely hairy; glumes (2—)3—5.5 mm
long, membranous, glabrous below, upper 1/3
hairy, purplish-green, apex acute; lower glumes
(2—)3-3.5 mm long, lanceolate, 1-nerved; upper
glumes (4—)4.5—5.5 mm long, oblong, 3-nerved;
lemmas (6—)6.5—8 mm long, lanceolate, 5-nerved,
membranous to coriaceous, purplish-green, papil-
lose, upper 1/3 scabrous, apex mucronate or short-
awned, the awn 0.3—0.8 mm long; callus glabrous
sometimes sparsely hairy; paleas as long as the
lemma, lanceolate, membranous, hairy along upper
1/4 and keels, apex with longer hairs; lodicules ca.
1 mm long, obovate; anthers 2.8—3.5 mm long;
ovary apex glabrous or with sparse hairs. Caryopses
not observed.

Leaf blade anatomy.—Cross-sections with
(11—)15—21 vascular bundles and (5—)7—11 ribs
above; sclerenchyma under abaxial epidermis con-
tinuous or discontinuous extending to all vascular
bundle; adaxial epidermis discontinuous, extending
to every second vascular bundle; adaxial epidermis
with dense covering of curly to straight hairs, the
hairs 0.15—0.2 mm long.

Observations.—Festuca procera differs from
other species of F: sect. Cataphyllophorae by form-
ing large tussocks with tall culms (80—170 versus
60-80 cm tall in other members), long, densely
flowered panicles, and long spikelets [(9—)11—13
(—15) versus 6—11(—12) mm in other members]. This

78 Festuca in South American Paramos

INS

IN

(mam
SSS S

Ss =

———

—

— = =
SS

—
—
SS
——

SS

Ni
ss

Figure 36. Festuca procera. A. Stylized growth form. B. Habit. C. Cataphylls. D. Ligule. E. Spikelet. F. Glumes. G. Lemma. H.
Lemma with palea and rachilla. I. Leaf blade cross-section. A-I, Stanctk 3313 (PRC).

Festuca in South American Paramos 79

COLOMBIA

| | 300 600 Kilometers
ES

-70

Figure 37. Distribution of Festuca procera (@), F- pilar-franceii (A), and F:boyacensis (m).

species is morphologically quite variable. Speci-
mens of F: procera from southern Ecuador (region
Azuay and Canar) present striking differences in
abaxial surface texture of the leaf blades. Some
specimens have sparingly scabrous leaf blades,
inflorescence branches that are scabrous, and nar-
rower panicles. In addition, all of these unusual
specimens lack cataphylls, although this may be an
artifact of the collection. Taxonomic status of this
group of specimens needs further study.

Distribution and habitat.—This species ranges
from Colombia (Cordillera Central and Oriental),
to Ecuador, and Peru. It occurs along margins
of the Andean mountain forests and in matorral
and transitional zones of grass paramo between
3400-3900 m.

Additional specimens examined. COLOMBIA.
Boyaca: Cafion de Chicamocha, Mun. Sativasur,
17 Oct 1992, Etter 662 (COL); Mun. El Cocuy—

Giiican, Parque Nacional El Cocuy, Hda. La Espe-
ranza, Valle de los Frailejones, 3800 m, 30 Dec
1998, D. Stancik 1788 (COL, FMB, PRC). Cauca:
Mun. San Sebastian, Macizo Colombiano, Valle de
las Papas, 2910 m, 11 Sep 1958, Jdrobo et al. 3737
(COL, US); Mun. Puracé, Parque Nacional Puracé,
quebrada San Nicolas, 3100 m, 6 Apr 1985, Wood
4797 (COL, K). Norte de Santander: Quebrada
del Rio Chitagu, 3400-3500 m, 20 Jul 1940,
J. Cuatrecasas & Garcia-Barriga 10036 (COL);
3400-3500 m, 20 Jul 1940, J. Cuatrecasas & Gar-
cia-Barriga 10037 (COL); Pamplona—Cerro al NE,
2770 m, 3400-3500 m, 26 Jul 1940, J. Cuatrecasas
& Garcia-Barriga 10207 (COL). ECUADOR.
Azuay: ca. 7 km S of road to Nabon, 03°07'S,
79°19'W, 3250-3350 m, S. Laegaard 101856
(AAU). Bolivar: km 14.5 E of Guaranda on road
to Riobamba, 2950 m, P.M. Peterson 9297 & EJ.
Judziewicz (K, MO, QCA, QCNE). Caniar: km 11.6

80 Festuca in South American Paramos

NW of Tambo on Panamerican Highway and 2.2
km E of road to microwave station, 3220 m, PM.
Peterson 8846, C.R. Annable & M.E. Poston (MO,
QCA, US); S of El Tambo, ca. 1.5—3 km along road
to Carshao, 02°28°S, 78°59’ W, 3100 m, S. Laegaard
& Sklenar 20296 (AAU, LOJA, PRC); El Tambo,
3400-3500 m, D. Stancitk 3793 (AAU, PRC, QCA);
D. Stancik 3794 (PRC, QCA); Road Pimo-Santa
Rosa, 3000 m, Acosta-Solis 21439A, B (US); Par-
amo road ca. 10 km WNW of Canar, 02°30’S,
79°00’ W, 3550 m, S. Laegaard 71031 (AAU, PRC,
QCA). Cotopaxi: at Laguna Quilotoa, 00°51’S,
78°53'W, 3500-3600 m, S. Laegaard 101350
(AAU, QCA); 3480-3500 m, S. Laegaard 101355
(AAU, QCA); 3700-3800 m, S. Laegaard 101340
(AAU, QCA); 00°52’S, 78°53°W, 3500-3750 m,
Ceron 21862 (QAP); road Angamarca—Corazon km
11 “O109"S; 787 W. 2880 m5 Apr 199255.
Laegaard 102164 (AAU, QCA); Parque Nacional
Cotopaxi, 3200 m, zuriete 129 (AAU, QCA); 3500
m, Escalona & Gallegos 287 (US); Parque Nacio-
nal Cotopaxi, 2.4 km W of entrance, 2870 m, PM.
Peterson 8734, C.R. Annable & M.E. Poston (K,
MO, QCA, QCNE, US); ca. 3 km de la entrada,
00°40°S, 78°30°W, 3200 m, 3 Jan 1982, Argiiello
2, 33, 37, 440 (QCA); Sosa et al. 43 (QCA); 13
Nov 1982, Balslev & T.: de Vries 3471 (QCA); km
2 N of Lasso on Panamerican Highway, 3270 m,
14 Apr 1990, P.M. Peterson 8747, C.R. Annable &
MLE. Poston (K, MO, QCA, QCNE, US); Panam
Highway, ca. 6 km N of Lasso, quebrada ENE
Pastocalle, 3400 m, Sparre 15827, 15836 (S);
Latacunga, paroquilla Sigchos, 2800 m, 30 Jun
1990, Ceron 10605 (QAP); Latacunga, paroquilla
Belisario, Quevedo—Cerro Putzalagua, 00°57’S,
78°33 W, 2800-3500 m, Ceron 25977 (QAP).
Chimborazo: road from San Fernando toward Mt.
Chimborazo and El Arenal, 3930 m, Barclay &
Juajiboy 8178 (COL, MO, US); Arenales de
Palmira, ca. 7 km S of Guamote, 01°59’S, 78°43 ’W,
3150 m, S. Laegaard et al. 18672 (AAU, LOJA,
QCA,QCNE); ca. 7 km S of Palmira, 02°00’S,
78°43 W, 3170 m, 20 Mar 1992, S. Laegaard
101814 (AAU, QCA, QCNE); 02°02’S, 78°45’W,
3200-3300 m, S. Laegaard 71695 (AAU, QCA,
QCNE); W of Arenales de Palmira, 02°01’S,
78°45 W, 3330 m, S. Laegaard 105102 (AAU); km
8.9 N of Palmira, 3260 m, PM. Peterson 8815 (K,
MO, QCA, QCNE, US); km 10 on road Alao—
Chamlo, 01°50’S, 78°34’ W, 3000 m, S. Laegaard
55410 (AAU, QCA); km 4.6 of Pungala on road to
Alao, 2970 m, PM. Peterson 9179, E.J. Judziewicz,

R.M. King & P.M. Jorgensen (K, MO, QCA,
QCNE, US); km 9 S of Guamote, then 1.6 km SW
on track to Alausi, 3300 m, PM. Peterson 9316 &
E.J. Judziewicz (K, MO, QCA, QCNE, US); Chim-
borazo, Chuquipoyo, 3900 m, Andre 3919 (K, US);
ca. km 10 S of Palmira, 3250 m, 11 Nov 1985, S.
Laegaard 55566 (AAU, QCA); ca. 5 km S of
Palmira, 02°06’S, 78°45’W, 3300 m, S. Laegaard
20379 (AAU, LOJA); Mt. Chimborazo, Sodiro s.n.
(QPLS); Puela, 01°30’°N, 78°30’ W, 2315m, Ceron
15102 (QAP). Imbabura: Laguna Cuicocha,
00°18°N, 78°22°W, 3000-3100 m, S. Laegaard &
S.A. Renvoize 70873, 70882 (AAU, K, QCA,
QCNE); 2900-3100 m, 24 May 1991, Penafiel et
al. 69 (MO, QCNE, PRC); 3100 m, E. Asplund
20176 (S); road Yahuarcocha—Mariano Acosta km
9, 00°22’N, 78°03 W, 2900 m, 7 Feb 1992, S. Lae-
gaard 101131 (AAU, QCA); Entre Muyurco
y Tallachupa, Nevado Cotacachi, 3000 m, Acosta-
Solis 19006 (US); 3200-3300 m, Penafiel et al.
1187 (MO); Mun. Otavalo, road from Laguna
Mojanda to Cochasqui, 00°04’55’N, 78°17°50’W,
3450 m, 19 Oct 2000, D. Stancik 4106 (PRC,
QCA). Loja: km 1 E of Guachamana on road
Macara—Catacocha, 2670 m, 9 Jun 1990, P.M.
Peterson 9512 & E.J. Judziewicz (K, MO, QCA,
QCNE, US). Morona—Santiago: Hda. Huargualla—
Hda. San Eduardo, 01°57’S, 78°32’ W, 3600 m, 19
Jul 1999, D. Stancik 3312, 3313, 3314,ERE,
QCA); way to Parque Nacional Sangay, 02°15’S,
78°27 W, 3700 m, 19 Jul 1999, D. Stancik 3317
(PRC, QCA). Pichincha: road to Yanaurco on N
side of Volcan Pichincha, 00°07’S, 78°34’ W, 3600
m, 17 Jun 1981, S. Laegaard 52301 (AAU, QCA);
Volcan Pichincha, 00°09°S, 78°32°W, 3200-3400
m, 5 Feb 1988, S. Laegaard 70002 (AAU, QCA,
OCNE); Sodiro s.n. (QPLS, US); km 41 on road
Quito—Latacunga, 3350 m, 13 Apr 1990, PM.
Peterson 8717, C.R. Annable & M.E. Poston
(QCNE, US); km 41, 3350 m, Buendia 2 (QCA);
Environments of Quito, Jameson 191 (K); Pane-
cillo, Sep 1887, Sodiro s.n. (QPLS); vicinity of
Quito, 3100 m, E. Asplund 6151 (AAU, MO, S,
US); Prop. Quito, Sodiro s.n. (QPLS, US); 3250
m, E. Asplund 6154 (AAU, S); 3500 m, 13 Jan
1926, E. Asplund 6139 (S, US); Road Pifo—Pintag,
above Inga Monserrat 00°19’S, 78°17W, 3400—
3500 m, S. Laegaard 102285 (AAU, QCA, QCNE);
Sep 1890, Mt. Pichincha, Sodiro s.n. (MO, NY,
QPLS); along road to the antennas, 00°09’S,
78°32’ W, 3200-3400 m, S. Laegaard 70016 (AAU,
QCA, QCNE); Mille 283, 285 (US); SE of Ungui,

Festuca in South American Paramos 81

3000 m, Firmin 116 (US); above Quito, 3300 m,
E. Asplund 16161 (S); 5 Feb 1988, S. Laegaard
70014 (AAU, QCA, QCNE); between Quito and
Conocoto, 3000 m, E. Asplund 10375 (S); between
Quito—Lloa, 3200 m, E. Asplund 7392 (S), 3600
m, Sparre 16006 (S); Volcan Cayambe, entrance
to Parque Nacional, 00°03’S, 78°04°W, 3550 m,
S. Laegaard & S.A Renvoize 70485 (AAU, K,
QCA, QCNE); 3870 m, Escalona & Gallegos E384
(MO); Antisana, road to Hacienda Pinatura,
3000-3100 m, 3 Mar 1988, S. Laegaard & S.A
Renvoize 70542 (AAU, K, QCA, QCNE); 00°26’S,
78°20’ W, 3400 m, S. Laegaard 102857 (AAU,
QCA, QCNE). Tungurahua: road Ambato-—
Guaranda, km 25, 01°18’S, 78°48°W, 3300 m, 5
May 1999, S. Laegaard & Sanchez 20033 (AAU,
LOJA, QCA); 01°24’S, 78°51°W, 4000-4100 m,
S. Laegaard 54802, 54809 (AAU, QCA); Pillaro,
along irrigation ditch, 2850 m, 10 Aug 1939, E.
Asplund &135 (NY, QCA, S, US). Bafios, 1800 m,
Zeeocp 1923. A.S. Hitchcock 21926 (NY, US);
Llangahua, 3300 m, Acosta-Solis 16678 (US);
Pucara de Chisalo, Stibel 297 (K). PERU. Ancash:
Prov. Pallasca, 5km S of road towards Cabafia
steep slope with Puya, Oxalis, Festuca, Tillandsia,
3210 m, 27 Mar 1997, PM. Peterson 13922 & N.
Refulio Rodriguez (K, MO, US, USM). Arequipa:
Volcan de Misti, open slopes and canyon sides,
3100-3300 m, 4 Nov 1925, Pennell 13231 (F).
Cajamarca: Prov. Cajamarca, a 9 km de la Caja-
marca, sobre carretera Cajamarca—Pacasmayo,
2800 m, 6 Nov 1984, Vega & Ruiz 3584 (MO);
surrounding of Cajamarca, 2750 m, 21 May 1980,
Becker et al. 1234 (LPB); Prov. Chota, near Las
Palmas, ca. 24 km NE de Chota, fragmented moun-
tain forest, 78°37’ W, 06°29’S, 2789m, 17 Apr
1993, M. Dillon et al. 6359A (F, MO); Distr. Banos
del Inca, a 3 km arriba de Banos del Inca, caretera
Cajamarca—Celendin, 16 Mar 1984, Vega 3239 (F);
km 13 de la carretera Cajamarca—Cumbe Mayo,
frequent, 3000 m, 4 Sep 1984, Vega 3324 (F);
Quebrada de la Esperanza, carratera a Cumbe
Mayo, quebrada que converga al Valle de Caja-
marca, arribe Cerro Sta. Apolonia, 2800 m, 4 Mar
1984, Vega et al. 3283 (F); Distr. San Juan, car-
retera Cajamarca—San Juan, 2350 m, 6 Dec 1993,
Vega 723 (F); Distr. De Chetilla, ruta a Llullapuquio
bosque perennifolio, 2650-2750 m, 21 May 1986,
Vega et al. 4117 (F); Quebrada de la Esperanza,
carratera a Cumbe Mayo, quebrada que converga
al Valle de Cajamarca, arribe Cerro Sta. Apolonia,
2800 m, 4 Mar 1984, Vega et al. 3283 (F); a 1 km

arriba de San Juan, siguiendo la carretera a Caja-
marca, 2300 m, 6 Feb 1985, Vega et al. 3855 (F);
Sexcemayo, al W de la ciudad de Cajamarca, 3500
m, 20 Jun 1991, Vega et al. 5745 (F); a la altura del
Paso El Gavilan, 3200 m, 18 Apr 1976, Vega et al.
1393 (F); Entre La Encanada y Kumullca, 3300 m,
17 Jun 1975, Vega et al. 1598 (F); Prov. Celendin,
Guanambra, a 11 km de la carretera Celendin—Caja-
marca, 2830 m, 3 Dec 1988, Vega 466] (F); al E
de la ciudad de Celendin, 2600 m, 5 Apr 1980, Vega
3830 (F); Celendin—Cajamarca road, 78°12’W,
06°58’S, 2800-3400 m, 25 Feb 1984, Smith 6222
(MO); above Celendin, 2750 m, 23 Mar 1988, S.A.
Renvoize 4879 (K); Bajando el paso de Gelig, sobre
la carretera Celendin—Balsas, 3000 m, 15 Jun 1981,
Vega 2576 (F); Prov. Chota, 5 km N of Chota on
road towards Conchan, 2560 m, 18 Mar 2000, P
M. Peterson 14968 & N. Refulio Rodriguez (MO,
US, USM); 20 km NW of Bambamarca on Hwy 3
N towards Chota, 2900 m, 18 Mar 2000, PM.
Peterson 14953, 14955, 14957 & N. Refulio Rodri-
guez (K, MO, US, USM); Chota—Tacabamba road,
6—8 km from Chota, 78°38’ W, 06°32’S, 2650 m,
20 Feb 1983, Smith et al. 3627 (K, MO); Distr.
Llacanora, sobre la carretera a Cajabamba, 2760
m, 6 Jul 1980, Vega 2250 (F); Prov. Cutervo, 10
km NW of Chiguirip and 19 km SE of Cutervo,
2630 m, 19 Mar 2000, PM. Peterson 14989, 14992
& N. Refulio Rodriguez (K, MO, US, USM); Prov.
Hualgayoc, cerro La Llama de las Ventanillas, al
SW de Bambamarca, 3890 m, 5 Mar 1991, Vega et
al. 5696 (F); Prov. San Marcos, Yanupacha, al E
de Ichocan, 3100 m, 26 Mar 1994, Vega et al. 6933
(F). Cuzco: Prov. Paucartambo, altura de Teleban,
3700-3800 m, 16 Jul 1990, Cano 3780 (F); Huayna
Pichu, rocks, 68°07°W, 16°17’S, 3000 m, 5 Aug
1937, Tutin 1285 (BM). Junin: Huancayo, 18.5
km NW of Huancayo up Rio Shullcas, just above
Acopalca, 3890 m, 12 Apr 1997, PM. Peterson
14224 & O. Tovar (K). La Libertad: Prov. Bolivar,
arriba de Longotea, ruta Bolivar, 2800 m, 28 Aug
1989, Vega 5013 (F). Lambayeque: Prov. Ferre-
nafe, Incahuasi, 3400 m, 22 Jun 1986, Quiroz 1929
(F, MO). Piura. Prov. Huancabamba, San Antonio,
km 25 de la carretera Huancabamba-—Salala,
2850 m, 5 Jan 1990, Vega et al. 5185 (F); 30 km N
of Huancabamba and 3 km N of Salala, 2980-—
3200 m, 2 Apr 2000, PM. Peterson 15178 &
N. Refulio Rodriguez (K, US, USM).

30. Festuca toca Stancik, Darwiniana 41(1—4): 123,
f. 10h—m. 2003. (Figs. 38, 39, 88A & B). Tyre:

82

Figure 38. Festuca toca. A. Habit. B. Inflorescence. C. Cataphylls. D. Ligule. E. Spikelet. F. Glumes. G. Lemma. H. Lemma with
palea and rachilla. I. Leaf blade cross-section. A-I, Stancik 1418 (PRC).

Festuca in South American Paramos

Aid

100 x

PER TIRS:

2mm

Festuca in South American Paramos 83

Colombia. Boyaca, Mun. Toca, road from Pesca
to Toca, km 4—-5, Paramo Cortadero, 5°30'N,
73°15’ W, 2700 m, 14 Nov 1998, D. Stancik 1404
(holotype: PRC!; isotypes: COL!, FMB!).

Rhizomatous perennials with extravaginal
innovations. Culms 50—90 cm tall, erect, solitary,
glabrous or finely scabrous; nodes 2-4; cataphylls
short, coriaceous, gray-brown. Leaf sheaths mem-
branous, brown, striate; auricles absent; ligules 0.1—
0.5 mm long, membranous, apex truncate; blades
10-35 cm long, 0.5—1.1 mm wide, conduplicate or
conduplicate to involute, green, abaxially glabrous.
Panicles 6—15(—20) x ca. 0.5 cm, erect, contracted,
elongate; branches scabrous. Spikelets 6-8 mm
long, florets 4 or 5(—6); rachilla glabrous or with
scattered hairs; glumes 2—4 mm long, lanceolate,
membranous to coriaceous, purplish, apex acute,
scabrous; lower glumes 2—3 mm long, 1-nerved;
upper glumes 3-4 mm long, 3-nerved; lemmas

4—5 mm long, lanceolate, 5-nerved, membranous
to coriaceous, purplish, upper 1/3 scabrous, apex
mucronate; callus glabrous; paleas as long as the
lemma, membranous, papillose, upper 1/3 hairy,
lodicules lanceolate, acuminate; anthers 1.1—1.6
mm long; ovary apex glabrous. Caryopses lanceo-
late; hilum 3/5 of total lenght.

Leaf blade anatomy.—Cross-sections with
7—15 vascular bundles and 5—13 ribs above; scle-
renchyma under abaxial epidermis discontinuous,
sometimes abaxial continuous, extending to some
(rarely to all) vascular bundles forming girders;
adaxial epidermis with scattered hairs, the hairs
0.2—0.9 mm long.

Observations.—Festuca toca differs from the
remaining species of F: sect. Cataphyllophorae by
having solitary culms. Festuca toca is morphologi-
cally similar to F' andicola (F: sect. Aulaxyper), but
differs by having longer glumes (lower glumes
2—3 versus 1.4—1.8 mm; upper glumes 34 versus

-80 -75 -70 -65 -60
9 &
‘ a f
Se 2b e eB
-| a
10 10
* be VENEZUELA
A
PANAMA “
5 —45
@ @
COLOMBIA
©
0 0
3 EC iia BRAZIL
5 4-5
aE le Sree : 7
0 300 . _ 600 Kilometers
-10 -10

-80 -75

-70 -65 -60

Figure 39. Distribution of Festuca asplundii (@), F. toca (A), and F. cocuyana (i).

84 Festuca in South American Paramos

2—2.5 mm) and longer anthers (1.1—1.6 versus
0.8—1 mm).

Distribution and habitat.—This species is
endemic to Colombia where it is known only from
the Cordillera Oriental (Boyaca, Norte de Santander).
It occurs in swampy patches in grass paramo zone
between 3100-3500 m. Festuca toca is present in
different vegetation communities such as: Lysipomia
sphagnophilla subsp. minor (Cleef 1981).

Additional specimens examined. COLOMBIA.
Boyaca: Mun. Duitama, paramo Pan de Azucar,
3450 m, 30 Nov 1998, D. Stancik 1418 (COL,
PRC); Paramo de Rusia, 3400 m, 2 Jan 1984, Wood
4481 (COL, FMB, K); 3570 m, 13 Dec. 1972, Cleef
7139 (COL); summit of road from Duitama to
Abendanos, 3490 m, 12 Feb 1972, D. Stanctk 2371
(COL, PRC); Hda Los Tres Corrales, 5°55°3.2°N,
73°4°13.5°W, 3490 m, Hernandez 1022A (COL);
Mun. Toca, road from Pesca to Toca, km 4-5,
paramo Cortadero, 5°30°N, 73°15’°W, 2700 m,
14 Nov 1998, D. Stancik 1404 (COL, FMB, PRC);
3350 m, 14 Nov 1998, D. Stancik 1365 (COL,
FMB, PRC); Mun. Susacon, road km 20, margin
of Rio Susacon, 3150 m, 24 Feb 1999, D. Stancik
2513 (COL, PRC); Paramo Guantiva (Sanjuanero),
3300 m, 10 Aug 1977, Ruiz & Romero 3 (COL);
Mun. Paipa, Cuchilla El Paramo, 3200 m, 4 Dec
1998, D. Stancik 1533 (COL, PRC); Mun. Arca-
buco, Santuario Iguaque, way from Laguna Iguque
to Laguna Ojo de Agua, 3650 m, 15 Oct 1998,
D. Stancik 948 (COL, FMB, PRC). Norte de
Santander: Paramo de Santurban, 3200 m, 4 Jan
1984, Wood 4507 (COL, FMB, K).

31. Festuca asplundii E.B. Alexeev, Bjull. Mos-
kovsk. ObS¢. Isp. Prir., Otd. Biol. 89(4): 116.
1984. (Figs. 39, 40, 88C—F). Tyre: Ecuador.
Pichincha, Iter Regnellianum quartum, Flora
Aequatoriensis, prov. Pichincha, vicinity of
Quito, Rucu Pichincha, 4650 m, 31 Aug 1939,
E. Asplund 8586 (holotype: S!).

Tussocked perennials with intravaginal innova-
tions. Culms up to 80 cm tall, erect, scabrous; nodes
1 near base. Leaf sheaths wide, coriaceous, stra-
mineous, scabrous; ligules 0.5—1.5 mm long, coria-
ceous, apex truncate or emarginate, short-ciliate;
blades ca. 40 cm long, 1—1.4 mm wide, conduplicate
to involute, rigid, somewhat abaxially scabrous,
green, apex obtuse. Panicles 12—27 x 3-4(-10) cm,
contracted, rarely with open branches; branches

scabrous. Spikelets sterile, forming 1(—2) shoots
(i.e., vegetative proliferation), florets 14, up to 6
when shoots lacking; glumes 4.5—6(—7) mm long,
coriaceous, dark purple, scabrous dorsally, distal
margins (upper 1/2) membranous, apex acute; lower
glumes 4.5—5(—6) mm long, lanceolate, 1-nerved;
upper glumes 5—6(—7) mm long, oblong-lanceo-
late, 3-nerved; lower lemmas sterile, 4.5—6(—7)
mm long, 5-nerved (sometimes inconspicuously),
lanceolate, scabrous or papillose dorsally, margins
membranous, apex entire or slightly two-dentate
and short-awned, the awns up to 0.8 mm long, dark
purple; lodicules and sexual organs absent; upper
lemmas converted into prolifering shoots 1—2.5 cm
long, 1—3-leafed are prolonged into leafy bracts.

Leaf blade anatomy.—Cross-sections with
11-17 vascular bundles and 9-15 ribs; scleren-
chyma under abaxial epidermis continuous extend-
ing to all the vascular bundles; adaxial epidermis
discontinuous extending to every other vascular
bundle; bulliform cells absent; abaxial epidermis
with prickles and densely hairy adaxially, the hairs
0.04—0.09 mm long.

Observations.—This species does not seem
to be morphologically similar to any other species
of Festuca from South America. St.-Yves (1927)
considered F. asplundii to be a viviparous form
of F. procera only lacking cataphylls. Individual
specimens of F. asplundii are highly variable in
spikelet size, panicle shape, and ligule charac-
teristics. We include numerous specimens from
Peru in F- asplundii, although Alexeev considered
this species to be endemic only to Colombia and
Ecuador. He attributes Peruvian specimens to the
unclearly described species, F! ancachsana and
invalidly described, F’ peruviana (Alexeev 1986).
In Festuca, vivipary occurs rarely in developing
spikelets, although, it can sometimes be found in
a few individuals of F: subulifolia, F. procera, and
F. tolucensis.

Distribution and habitat.—Festuca asplundii 1s
known from northern Peru, Ecuador, and Colombia
(Cordillera Central and Cordillera Occidental). It is
found in swampy and humid patches of grass par-
amo and the lower zones of superparamo between
3300-4300 m. It is a dominant and co-dominant
species in communities of Espeletia hartwegiana
subsp. centroandina y Calamagrostis recta (Cleef et
al. 1983), Festuca dolichophylla, Senecio latiflorus
& Pentacalia vernicosa (Sturm and Rangel 1985),
Festuco dolichophyllae-Calamagrostietum effusae

85

Festuca in South American Paramos

Figure 40. Festuca asplundii. A. Stylized growth form. B. Habit. C. Inflorescence. D. Ligule. E. Spikelet. F. Glumes. G. Lemmas.

H. Leaf blade cross-section. A—-H, Stancik 3675 (PRC).

86 Festuca in South American Paramos

(Salamanca et al. 1991), Senecionetum rufescentis-
Agrostiosum (Cuatrecasas 1934), Calamagrostis
effusa and Loricaria cf. complanata (Rangel et al.
1995), Espeletia pycnophylla & Arcytophyllum
capitatum (Rangel and Luteyn 1995), Gynoxyo-
Polylepidetum sericae (Salamanca et al. 1991),
Loricaria thuyoides & Pentacalia vernicosa (Rangel
et al. 1995), Loricaria thuyoides & Arcytophyllum
capitatum (Rangel et al.1995), Calamagrostis effusa
& Festuca aff. dolichophylla (Salamanca 1991),
and Calamagrostis recta, Festuca aff. sublimis &
Diplostephium rupestre (Cuatrecasas 1934).
Additional specimens examined. COLOMBIA.
Caldas: Mun. Manizales, Parque Nacional Los
Nevados, entrance, 4020 m, 18 Sep 1999,
D. Stancik 3410, 3411 (COL, PRC); road from
Casa del Cisne to Rio Nereidas, km 5, 3800—4000
m, 18 Sep 1999, D. Stancik 3401, 3402 (COL,
PRC); Carretera Manizales—Nevado del Ruiz, km
72,4100 m, 1 Mar 1977, Forrero et al. 3644 (COL,
MO); Carretera entre Manizales y el Nevado del
Ruiz, 4000 m, 5 May 1959, Pinto-Escobar 415
(COL, PSO); Paramo de Herves, division de
Aguato, 3300 m, Feb 1852, Triana 810 (COL);
3500 m, Feb 1852, Triana s.n. (US); alrededores
del Nevado del Ruiz, 4140-4300 m, 9 Oct 1978,
Rangel et al. 1800 (COL); Nevado del Ruiz, 8 Oct
1983, Wood 4030 (AAU, K); Paramo El Ruiz,
3700-4200 m, 26 Dec 1936, Chardon 5010 (COL);
3400-3700 m, 17 Dec 1917, Pennell 3064 (K, MO,
US); Nevado del Ruiz, arenales de Ruiz, 3 km a de
la Olleta, 4250 m, 14 Oct 1972, Cleef et al. 5983
(COL, U, US); 3900-4200 m, 5 May 1940, J. Cua-
trecasas 9311 (COL, US); Carretera El Ruiz—Otun,
Hda. Buenos Aires, 3940 m, 20 May 1979, Sala-
manca AC172 (COL); 3500 m, Dec 1974, Liana 71
(MEDEL); Paramo de Quindio, 4300-4500 m, 20
Aug 1922, Pennell 9890 (K, US). Cauca: Mun.
Popayan, Parque Nacional Puracé, Pilimbala,
02°20.07'N, 76°23.65 W, 4200-4300 m, 6 Jul 2000,
D. Stancik 3611 (COL, PRC); Puracé, Laguna San
Rafael, 3300 m, 6 Apr 1985, Wood 4802 (COL,
FMB, K). Narifio: Mun. Cumbal, Nevado del
Cumbal, N-NE slope, 4100 m, 9 Mar 1999, D.
Stanctk 2724 (COL, PRC); Mun. Tuquerres, Volcan
Azufral, Laguna Verde, 3750 m, 9 Mar 1999, D.
Stancik 2759 (COL, PRC); Mun. Pasto, Volcan
Galeras, 3920 m, 6 Aug 1977, Pinto-Escobar 1845
(COL); 3900 m, 29 Oct 1983, Wood 4062 (COL);
Volcan Galeras, 3900 m, Wood 4063 (FMB, K).
Risaralda: Mun. Santuario, SE del Cerro Venta-

nas, Macizo del Tamara, 3820 m, 11 Feb 1983,
Torres-Romero et al. 1875 (COL); Mun. Pereira,
Parque Nacional Los Nevados, Laguna del Otun,
3800 m, 16 Feb 1980, Jaramillo-Mejia et al. 5634
(COL); Nevado de Sabanta Isabel, 4300 m, 17 Feb
1980, Jaramillo-Mejia et al. 5683 (COL). Mun.
Santa Rosa, Volcan Santa Rosa, 4280 m, 20 Feb
1980, Jaramillo-Mejia et al. 5770 (COL); Paramo
entre Termales y Libano, 4000 m, 21 Dec 1958,
Barclay & Juajibioy 6456 (COL, MO, US). Tolima:
Mun. Ibague, 04°38.5’N, 75°19.1°W, 4250 m, 8 Jun
2000, D. Stancik 3606 (COL, PRC); Mun. Santa
Isabel, vertiente oriental abajo del paso de El Otun,
4200 m, 7 Feb 1980, Jaramillo-Mejia 6237 (COL);
Quebrada Africa, 4200-4300 m, 5 Feb 1980, Diaz-
Piedrahita & Jaramillo-Mejia 1821 (COL, UV).
Valle de Cauca: Mun. Cali, Farallones de Cali,
03°19.78'N, 76°41.7°W, 3950 m, 9 Jul 2000 D.
Stancik 3612, 3613, 3614 (COL, PRC); 3650-3700
m, D. Stancik 3637 (COL, PRC). ECUADOR.
Azuay: Parque Nacional Cajas, Totorococha—
Mazan Valley, 02°53’S, 79°10’ W, 4000 m, 12 Nov.
1987, Ramsay & Smith 502 (K, QCA, QCNE);
Cuenca—Molleturo road, 3600-4200 m, 26 Jul 1982,
Clements et al. 2170 (AAU, QCA, QCNE); Paramo
de Soldados ca. 14 km above Soldados, 02°08 54'S,
79°08°17°W, 3900 m, 9 Jan 2000, S. Laegaard et
al. 20931 (AAU, LOJA); between Huagrarancha
and Loma de Galapagos, 3140—3505m, J. Steyer-
mark 53481 (US); Mun. Cuenca, Parque Nacional
Cajas, N side of Lagoon Taglococha, 02°47’S,
79°15'W, 4100-4200 m, 1 Sep 2000, D. Stancik
3849, 3871 (PRC, QCA). Carchi: Volcan Los
Chiles, along road ca. 1 km E of pass, 00°49'N,
77°57’ W, 3950-3980 m, S. Laegaard 101695
(AAU, QCNE); D. Stancik 3241, 3279 (PRC,
QCA); 00°48'N, 77°56’ W, 3850-4000 m, S. Lae-
gaard 54972 (AAU, QCNE); SW slope of Volcan
Chiles, 3740 m, B. Ollgaard & Balslev 8545
(AAU); S slope, Ramsay 896 (K, QCA); Paramo
del Angel, 3700 m, S. Largaard 101284 (AAU,
QCA); sector El Volador, 00°38’°N, 67°53’W,
3400-3800 m, Davalos 19 (US). Chimborazo: E
slope of Mt. Chimborazo, 4450 m, E. Asplund 7927
(S); 4250 m, FE. Asplund 7806 (S); El Altar, N side
of the Volcan, 4200-4300 m, Sklenar & Kosteckova
1106 (QCNE); Paramo de los Altares, Collanes
valley, 4200 m, Ramsay & Smith 415B (K, QCA,
QCNE); N side of Volcan, 4200-4400 m, Sklenar
& Kosteckova 1028 (AAU); 4300 m, Sklenar &
Kosteckova 97-10 (QCA); N side, 4400 m, Sklendr

Festuca in South American Paramos 87

& Kosteckova 91-8 (US); from campsite above Rio
Alao 8.5 km E of Guardiana Alao, 3350-3550 m,
P.M. Peterson 9204, E.J. Judziewicz, R.M. King &
P.M. Jorgensen (K, MO, QCA, QCNE, US); W of
pass Alao-Huamboya, 01°48’S, 78°25’ W, 3750—
3800 m, S. Laegaard 55425 (AAU, QCA). Cerro
Yanaurcu, 02°14’S, 78°30’ W, 4000-4200 m,
Sklenar & Kosteckova 1478 (AAU); Paramo de Las
Tres Cruces, N and E of Alao, 3500 m, Barclay &
Juajibioy 8804 (MO, US); between Mt. Chimbo-
razo and Urbina, 3600-4500 m, A.S. Hitchcock
21978 (US). Cotopaxi: Cerro Verde, road San
Miguel—Puerto Nuevo, km 29, 3950-4050 m, B.
OAllgaard & Balslev 9924 (MO). Imbabura: Pro-
antag, 3500-3800 m, Acosta-Solis 19199 (US);
Mun. Urcuqui, road to Cerro Yanaurcu, 00°28.5’N,
78°20’ W, 4150 m, 15 Oct 2000, D. Stancik 4097
(PRC, QCA). Mun. Cayambe, Volcan Cayambe,
00°00’°31.6°N, 78°00'55.6’ W, 4450 m, 20 Oct 2000,
D. Stancitk 4150, 4151 (PRC, QCA). Loja: Jion-
bura—Yumba road, ca. 3 km W of pass, 04°44’S,
79°25 'W, 3450 m, S. Laegaard et al. 18606 (AAU,
LOJA, QCA, QCNE); along road to Fierra Urcu,
ca. 10 km from main road Loja—Saraguro, 03°33’S,
79°15’ W, 3400 m, S. Laegaard et al. 18859 (AAU,
LOJA, QCA, QCNE); Cordillera de Amaluza,
Laguna de Jimbira, 04°42’S, 79°25’W, 3400 m,
Cabrera 438 (LOJA); Laguna Chuquiraga E of
Amaluza, 04°37’S, 79°22’ W, 3300 m, S. Laegaard
etal. 19280 (AAU, LOJA, QCNE); Mun. Saraguro,
road to Fierra Urcu, 03°42'40"S, 79°18'12’W,
3400-3450 m, 24 Aug 2000, D. Stancik 3763, 3767
(PRC, QCA). Napo: Parque Nacional Las Llanga-
natis, base of Cerro Hermoso on W side, near lake
78°18°W, 01°14’S, 3850 m, 13 Nov 1999, D.A.
Neill et al. 12042 (MO); W of Cerro Hermoso, near
saddle between headwaters of Rio Verde and Rio
Topo, 78°19.5°W, 01°11’S, 3950 m, D.A. Neill et
al. 11992 (MO); camino desde el Paramo de Sogu-
illas hasta Aucacocha, 78°19’ W, 01°08’S, 3940 m,
16 Nov 1999, Narvazez et al. 548 (MO); Paramo
de Soguillas, near Las Torres de Llanganatis,
3850-4000 m, 16 May 1982, B. Allgaard & Holm-
Nielsen 38728 (MO). Pichincha: W side of Sin-
cholagua, 13—14,500 ft, Feb 1880, Whymper 1326
(K). W side of the mountain ridge ca. 2 km W of
Cerro Saraurcu, 4200 m, Sklenar & Kosteckova
1817 (AAU); Nevado Cayambe, W side, 4200 m,
Sklenar & Kosteckova 1142 (QCA); SW slope,
4400 m, Sklenar & Kosteckova 1890 (AAU); SW
slope, 4250 m, Molau & Eriksen 3244 (QCA); NE

side of Cayambe Mountain, ca. 4900 m, Cazalet &
Pennington 5763 (K, US); 3 km S of Refugio on
the SW slope, 4220 m, PM. Peterson 9076, E.J.
Judziewicz & R.M. King (K, MO, QCA, QCNE,
US); along road to refuge, 00°04’S, 77°57’W,
4300 m, S. Laegaard & S.A. Renvoize 70511
(AAU, K, QCA, QCNE); W side, 4200 m, Sklenar
& Kosteckova 64-19 (US); N slope, 3750-3800 m,
B. Allgaard et al. 34202 (K, MO); base of Volcan
Sincholagua, 00°34’S, 78°22’W, 4200 m, Holm-
Nielsen 6612 (AAU, S); Cerro Sincholagua,
3900-4000 m, Balslev et al. 3901 (AAU, MO,
QCA); W side, 13,000—-14,500 ft, Whymper s.n.
(K); Paramo de Guamani, E. Asplund 17174 (S);
Laguna Linda, 3 km W of Paso de Guamani,
3900 m, Sparre 17719 (S); 01°19°S, 78°12’°W,
4000 m, S. Laegaard 18449 (AAU, LOJA, QCA,
QCNE); 00°56’S, 78°23’ W, 3600 m, S. Laegaard
53343 (AAU, QCA, QCNE); 4050 m, B. Allgaard
& Balslev 10122 (MO); Laguna de Papallacta,
3750 m, B. Ollgaard & Balslev 8170 (MO);
antenna, 4000-4100 m, B. Ollgaard & Balslev
10074 (M); Aguirre et al. 4206 (LOJA); 4050 m,
S. Laegaard 51318 (AAU, QCA); S of Paso de la
Virgen, 00°21°S, 78°13’ W, 4050-4250 m, S. Lae-
gaard 55680 (AAU, QCA); 4200-4250 m, S.
Laegaard 101383 (AAU, QCA); W slopes, 3800—
4100 m, Balslev 1615 (AAU); 4060 m, Fierrp 24
(QCA); 4050 m, Naranyo 4 (QCA); 3960 m, Len
1139 (QCA); Volcan Cotacachi, 00°35’S, 78°20'W,
4150 m, Ramsay & Smith 785 (K, QCNE); 4200 m,
Sklenar & Kosteckova 112-25 (QCA); vicinity of
Quito, Rucu Pichincha, 465 m, FE. Asplund 8576
(MO, NY, QCA, S); Volcan Antisana—Laguna
Micacocha, B. Ollgaard & Balslev 8868 (MO);
3850-3950 m, S. Laegaard 101595 (AAU, QCA,
QCNE); valley NW of N peak of Volcan, 4000-—
4300 m, Grubb et al. 639 (K); Paramo de Mojanda,
between Laguna Grande and L. Negra, 00°08'N,
78°16’ W, 3700-3800 m, S. Laegaard 54581 (AAU,
QCA, QCNE); Mun. Pifo, Paramo de Guamani,
grass paramo, 00°19°S, 78°12°W, 4300 m, 19 Jun
1999, D. Stanctk 3044, 3050 (PRC, QCA); Mun.
Amaguana, summit of Volcan Pasachoa, 00°28’S,
78°29 'W, 4200 m, 14 Sep 2000, D. Stancik 3675
(PRC, QCA, US); 4150 m, D. Stancik 3703 (PRC,
QCA). Sucumbios: Paramo Mirador, SW of
Playon de San Francisco, 3400-3600 m, P.M.
Peterson 9154, E.J. Judziewicz & R.M. King (US).
Tungurahua: Cordillera de Llanganatis, El Tambo
near Lake Yanacocha, 3650 m, EF. Asplund 10000

88 Festuca in South American Paramos

(S); below summit of Pan de Azucar, 4100 m,
B. Ollgaard et al. 38541 (AAU, MO, NY, QCA);
B. Ollgaard et al. 38540 (AAU); N slope towards
Rio Golpe just N of Chosca Aucacocha, 3600 m,
B. @llgaard et al. 38670 (AAU, MO, NY, QCA);
Las Tolas, ca. 5 km SE of Laguna Pisayambo,
01°07’S, 78°21°W, 3900-4000 m, S. Laegaard et
al. 20733 (AAU, LOJA); Paramo de Jaramillo,
01°08°10"S, 78°08°22"W, 4000-4250 m, S. Lae-
gaard 53289 (AAU, QCNE); Faldas de Cazahuala,
3200 m, Acosta-Solis 5097 (US); Mun. Pillaro, Las
Llanganatis, Rio Milin, 01°07°37°S, 78°21°W,
3750 m, 28 Sep 2000, D. Stanctk 3904 (PRC,
QCA). Zamora-Chinchipe: Sendero Amaluza—
Palanda, cerca de la Laguna Arrebatadas, 3350 m,
Larsen et al. 21 (AAU, NY, QCA). PERU. Ama-
zonas: Prov. Chachapoyas, upper slope of Cerro
Campanario, 3600-3900 m, 3 Aug 1962, J. Wur-
dack 1572 (US). Ancash: Yungay Prov., Huascaran
Parque Nacional Llanganica Sector, Quebrada
Ancachs at portachuelo, 09°03 ’S, 77°35’ W, 4770—
4870 m, 31 Dec 1984, Smith et al. 8879 (QCA, U);
Yungay Prov., Huascaran Parque Nacional, 4700 m,
Smith 11318 (MO, QCNE); Prov. Huari, Catac-
Chavin road, 40 km from Catac, Huascaran Parque
Nacional, 9°42’S, 77°13 'W, 4360 m, 18 Aug 1984,
Smith 8284 (MO, US); Prov. Recuy, Huascaran
Parque Nacional, camino over Cahuish Tunnel,
puna, 09°41°S,77°15°W, 4550-4500 m, 7 Oct
1985, Smith et al. 11127 (MO); Prov. Yungay,
Huascaran Parque Nacional, Morococha at largest
lake 08°55"S, 77-35 We 4550 im: Io) Jan 1985:
Smith et al. 9225 (MO). Cajamarca. Prov. Caja-
marca. 52 km N of Cajamarca on Hwy 3 N towards
Bambamarca, and ca. 1 km W on a small road,
Carretera Lagunas Yanacanchilla, 3800 m, 16 Mar
2000, P.M. Peterson 14892 & N. Refulio Rodriguez
(MO, US, USM); Prov. Hualgayoc, desvio de la
carretera Coymolache—Chugur, Cerro Tantahuatay,
3700 m, 29 Apr 1994, Vega 7087 (F); Cajamarca—
Bambamarca road, 55 km N of Cajamarca,
06°55°S, 78°35’ W, 3750 m, 6 Feb 1984, Smith et
al. 7428 (MO). La Libertad: Prov. Bolivar, entre
Longotea y Bolivar, en jalca, 3600 m, 2 Jun 1960,
Lopez & Sagasteguii 3340 (US); Prov. Pataz, Paso
de Alaska, Carretera a Tayabamba, 3900 m, 24 Jun
1974, Lopez & Sagasteguii 8180 (MO). San Mar-
tin. Prov. Mariscal Caceres, grassland in NW sec-
tor Rio Abiseo Parque Nacional, 07°40’S, 77°30’ W,
3650 m, B. Leon & Young 1662 (MO).

32. Festuca boyacensis Stancik, Darwiniana
41(1-4): 128, f. 7f#k. 2003. (Figs. 37, 41, 89A
& B). Type: Colombia. Boyaca, Mun. Aquita-
nia, paramo Los Curies, 3500 m, 8 Feb 1999,
D. Stancik & Medina 2166 (holotype: PRC!;
isotypes: COL!, FMB!).

Tussocked perennials with intravaginal inno-
vations. Culms 30-60 cm tall, erect, glabrous;
nodes | near base with 2 short leaves. Leaf sheaths
membranous, stramineous, occasionally brown-
ish, glabrous; auricles absent; ligules 0.3—0.6 mm
long, membranous, apex truncate, ciliate; blades
15—20 cm long, 0.4—0.7 mm wide, conduplicate to
involute, glaucous, abaxially finely scabrous, apex
acute. Panicles 6-15 x 0.5—1 cm, contracted, nar-
row; branches scabrous. Spikelet 8—9.5 mm long,
lanceolate, florets 3 or 4; rachilla glabrous; glumes
3—5.2 mm long, membranous, glabrous, margins
finely ciliate, purplish, apex acute; lower glumes
3—3.7 mm long, lanceolate, 1-nerved; upper glumes
4—5 mm long, ovate, 3-nerved; lemmas 6—7.5 mm
long, lanceolate, 5-nerved, membranous, glabrous,
purplish, awned, the awn 1—2 mm long; callus gla-
brous; paleas 5/6 as long as the lemma, glabrous,
keels ciliate; lodicules lanceolate, acuminate; anthers
0.9-1.2 mm long; ovary apex glabrous. Caryopses
lanceolate; hilum 1/2 as long as the grain.

Leaf blade anatomy.—Cross-sections with 5—7
vascular bundles and 3—5 ribs above; sclerenchyma
only abaxially present, discontinuous or almost con-
tinuous; bulliform cells absent; adaxial epidermis
with sporadic hairs, the hairs 0.01—0.1 mm.

Observations.—Festuca boyacensis is mor-
phologically similar to the Ecuadorian species,
FE. chimborazensis. However, F: chimborazensis
has smaller culms [8—15(—28) versus 30-60 cm],
smaller panicles (2—8 versus 6—15 cm), and glumes
with obtuse (acute in F’ boyacensis) apices.

Distribution and habitat.—Festuca boyacen-
sis is endemic to Colombia, known only from the
Cordillera Oriental (Boyaca). It occurs in swampy
patches and margins of lakes and springs in grass
paramos between 3000-3500 m.

Additional specimens examined. COLOMBIA.
Boyaca: Mun. Santa Rosa de Viterbo, paramo Alto
Lamadero, 3200 m, 30 Nov 1999, D. Stanctk 1493
(COL, MB, PRC).

33. Festuca carchiense Stancik, Folia Geobot. Phy-
totax. 39(1): 98, f. 1, 1-5. 2004. (Figs. 42, 44,

Festuca in South American Paramos

tk 1493 (PRC).

is. A. Stylized growth form. B. Habit. C. Ligule. D. Spikelet. E. Glumes. F, Lemma. G. Lemma with
s-section. A—-H, Stané.

Figure 41. Festuca boyacens
palea and rachilla. H. Leaf blade cros

90 Festuca in South American Paramos

=

SK =
Spe Sas
SSSA
OTS FR
Vt SJ
E =
Ly

Figure 42. Festuca carchiense. A. Stylized growth form. B. Habit. C. Ligule. D. Spikelet. E. Glumes. F. Lemma. G. Lemma with
palea and rachilla. H. Leaf blade cross-section. A-H, Laegaard 101162 (AAU).

Festuca in South American Paramos 9]

89C & D). Tyre: Ecuador. Carchi, km 11 along
road Las Juntas (Tulcan)-El Angel, 77°50’°W,
00°43’N, 3330 m, 11 Mar 1992, S. Laegaard
101716 (holotype: AAU!; isotypes: PRC},
QCA!, QCNE!).

Densely tussocked perennials with intravagi-
nal innovations. Culms 50-60 cm tall, erect, gla-
brous; nodes 1, basal. Leaf sheaths membranous,
brownish-gray, glabrous, inconspicuously striate;
ligules 0.5—0.8 mm long, apex short-ciliate; blades
25-40 cm long, 0.4—0.7 mm wide, conduplicate
to involute, finely abaxially scabrous, green, apex
obtuse. Panicles 10—20 x ca. 0.7 cm, contracted,
slender; branches scabrous. Spikelets 9-10.5 mm
long, lanceolate, florets 3 or 4; rachilla with scattered
hairs; glumes (3.5—)4—6.5 mm long, membranous,
purplish-green, upper 1/5 scabrous; lower glumes
(3.5—)4 4.5 mm long, oblong-lanceolate, 1-nerved,
apex acute; upper glumes 5—6.5 mm long, oblong,
3-nerved, apex acute; lemmas (6—)7—7.5 mm long,
lanceolate, 5-nerved, membranous, purplish-green,
scabrous distally, awned, the awn 0.8—1.5 mm long;
callus with scattered hairs; paleas as long as the
lemma, membranous, upper 1/3 and keels scabrous
to hairy; lodicules ca. 0.7 mm long, lanceolate;
anthers 1.1—-1.3 mm long; ovary apex glabrous.
Caryopses not observed.

Leaf blade anatomy.—Cross-sections with 5
vascular bundles, 3(—5) ribs above; sclerenchyma
discontinuous and small under abaxial epidermis,
absent under adaxial epidermis; adaxial epidermis
with scattered hairs, the hairs ca. 0.09 mm long.

Observations.—F estuca carchiense is morpho-
logically similar to F glumosa and F: imaburensis.
(See details in F’ glumosa section).

Distribution and habitat.—This species is
known only from northern Ecuador (Carchi,
Imbabura) and it occurs in grass paramo between
3200-3800 m.

Additional specimens examined. ECUADOR.
Carchi: Paramo del Angel, lakes at Los Voladeros,
00°41 °N, 77°53'W, 3700 m, S. Laegaard 101294
(AAU, QCA); 3200-3800 m, S. Laegaard 55769
(AAU). Imbabura: road Yahuarcocha—Mariano
Acosta, km 20, Paramo de Mariano Acosta,
00°20°S, 78°00’ W, 3600-3650 m, 8 Feb 1992,
S. Laegaard 101162 (AAU, QCA, PRC).

34. Festuca chimborazensis E.B. Alexeev, Bot.
Zhurn. (Moscow & Leningrad) 69: 1549.

1984. (Figs. 43, 44, 90A & B). Type: Ecuador.
Chimborazo, Iter Regnellianum quartum, Flora
Aequatoriensis, prov. Chimborazo, southern
slopes of Mt. Chimborazo, hard dry ground
along a rivulet, 3900 m, 21 Aug 1939, E.
Asplund 8446 (holotype: S!; isotypes: NY!,
QCA!).

Densely tufted perennials with intravagi-
nal innovations. Culms 8—15(30) cm tall, erect,
glabrous; nodes 1, basal with a single short leaf.
Leaf sheaths coriaceous, grayish, finely striate,
glabrous; ligules 0.3—0.4 mm long, membranous,
apex truncate, short-ciliate; blades 4-15 cm long,
0.4-0.6 mm wide, conduplicate to involute,
curved to mostly straight, glaucous, apex obtuse.
Panicles 2-10 cm long, 5—7 mm wide, contracted,
narrow; branches scabrous on all surfaces. Spike-
lets 8-10(—-11.5) mm long, lanceolate, florets
3 or 4(—5); rachilla with scattered short hairs;
glumes 2.5—5 mm long, 2/5—1/2 the length of the
spikelet, membranous to coriaceous, finely ciliate
on margins, whitish-green, apex obtuse; lower
glumes 2.5—3.5 mm long, oblong, 1-nerved; upper
glumes 3.5—5 mm long, ovate, 3-nerved; lemmas
6—6.5(—7) mm long, lanceolate, 5-nerved, membra-
nous, glabrous, whitish-green, sometimes purplish,
awned, apex acuminate, the awn 1—2 mm long;
callus glabrous; paleas 5/6 as long as the lemma,
glabrous, ciliate on keels; lodicules ca. 0.8 mm
long, lanceolate, acuminate; anthers 0.7—0.8 mm
long; ovary apex glabrous. Caryopses lanceolate;
hilum 1/2—3/5 as long as the grain.

Leaf blade anatomy.—Cross-sections with
5(—7) vascular bundles and 3(—5) ribs above; scle-
renchyma present abaxially, continuous or discon-
tinuous; bulliform cells absent; adaxial epidermis
with few hairs, the hairs ca. 0.09 mm long.

Observations.—Morphologically it appears
that F) boyacensis from the Colombian Cordillera
Oriental is most similar to F: chimborazensis. Other
species that appear similar morphologically are
F: glumosa, F. imbaburensis, F: carchiense, and
F. sumapana. All of these species are more robust,
i.e., with larger culms and have straight and often
scabrous leaf blades.

Distribution and habitat.—This species is
endemic to Ecuador (Bolivar, Chimborazo, Coto-
paxi, Pichincha, Tungurahua). It is known from
humid or swampy patches in the superparamo
growing between 3500-4500 m.

92 Festuca in South American Paramos

y
NS SS

}

Py
AT AY
ely
AWA
4

; Hob j
eee fy ei 7 {
4 Z ir)

[1mm Fed) Atk

tS

Figure 43. Festuca chimborazensis subsp. chimborazensis. A. Habit. B. Ligule. C. Inflorescence (detail). D. Spikelet. E. Glumes.
F. Lemma. G. Lemma with palea and rachilla. H. Leaf blade cross-section. A-H, Stancik 3883 (PRC).

Festuca in South American Paramos 93

-80 -75

COLOMBIA

VENEZUELA

ECUADOR

-80 -75

N

| 200 400 Kilometers

-70 oA

Figure 44. Distribution of Festuca chimborazensis (@), F: carchiense ( &), F- cleefiana (m), and F. dinirica (@).

KEY TO THE SUBSPECIES OF FESTUCA CHIMBORAZENSIS

1. Leaf blades 4-9 cm long, curved; leaf cross-sections with only 5 vascular bundles; culms 8-15 cm
tall; panicles 2-8 cm long, ca. 5 mm wide; upper glumes 4.5—5 mm IONnG..............ccceeseeeseeeeeeeeeeees

PROCTER HEHEHE HEHE SESE ES EEE EEE HESS EE EE SHEE EE EEEEESEEHEEEHEEEEEEEEEEES

.... 34a. F. chimborazensis subsp. chimborazensis

1. Leaf blades 11-15 cm long, mostly straight; leaf cross-sections with (5—)7 vascular bundles; culms
22-40 cm tall; panicles 8-10 cm long, ca. 7 mm wide; upper glumes 3.54 mm long................5:

POPS EEEPO EE ET HEHEHE HEHEHE HEE EEE EEE EEEEEE THEE EE EEEEEHEEHEHEHEEEHEEEHEES

34a. Festuca chimborazensis subsp. chimbora-
zensis (Fig. 43).

Culms 8-15 cm tall. Leaf blades 4—9 cm long,
curved; leaf cross-sections with only 5 vascular
bundles. Panicles 2-8 cm long, ca. 5 mm wide.
Spikelets with upper glumes 4.5—5 mm long.

sot 34b. F. chimborazensis subsp. micacochensis

Additional specimens examined. ECUADOR.
Bolivar: Road Ambato—Guaranda, 12 km W of
intersection between old and new road, 01°26’S,
78°56’ W, 4150 m, | Aug 1985, S. Laegaard 54820
(AAU, QCNE); 55.4 km SW of Ambato, 4050 m,
3 May 1990, PM. Peterson 8979 & C.R. Annable
(MO, QCA, QCNE, US); 66.5 km SW of Ambato,

94 Festuca in South American Paramos

4300 m, PM. Peterson & C.R. Annable &995 (MO,
QCA, US); km 4 on the road Los Arenales—Salinas,
01°24’S, 78°55’°W, S. Laegaard 55346 (AAU);
66.5 km SW of Ambato on Hwy to Guaranda and
2.2 km on road to Fecundo Vela, sandy volcanic
soil in Festuca-Calamagrostis grassland, 4300 m,
5 Mar 1990, PM. Peterson & C.R. Annable
8996A (MO). Chimborazo: Southern slope of
Mt. Chimborazo, 3900 m, FE. Asplund 8411, 8446
(NY, QCA, S); 3800 m, Fagerlind et al. 934bis
(S); along Whymper road ca. 15 km S of Cruce de
Los Arenales, 01°32°S, 78°52°W, 3950 m, 18 Sep
1998, S. Laegaard 19155 (AAU, QCNE); Urbina
towards Mt. Chimborazo, 3700 m, E. A’plund 7892
(K, NY); sector Cruce de los Arenales, swampy,
01°28°14"S, 78°54'06"W, 4300 m, 20 Sep 2000,
D. Stancik 3711, 3719 (PRC, QCA); 01°27°0.8°S,
78°53 °57.5'W, 4150 m, 20 Sep 2000, D. Stancik
3710, 3720 (AAU, PRC, QCA). Cotopaxi: Volcan
Cotopaxi, N side of the mountain, superparamo
vegetation, sandy slope, 00°39’S, 78°42’ W, 4250—
4450 m, Sklenar & Kosteckova 858 (AAU); Parque
Nacional Cotopaxi, NW of Limpio Pungo, 00°37'S,
78°27'°W, 3850-4000 m, S. Laegaard 52125
(AAU); Hacienda Pauzacha S of Volcan Cotopaxi,
00°44'S, 78°49°W, 3650 m, S. Laegaard 55736
(AAU); along road El Chaupi—Pastocalle, 00°41’S,
78°39'W, 3500 m, 2 Jul 1985, S. Laegaard 54594
(AAU, QCA, QCNE); Angamarca road 5 km from
junction to road Latacunga—La Mana, 01°00’S,
78°55'°W, 4250-4280 m, S. Laegaard 102156
(AAU); 21.5 km E of Pilalo, 3820 m, 15 Apr 1990,
P.M. Peterson 8768, C.R. Annable & M.E. Poston
(MO, QCA, US); Tililac, 4000 m, Acosta-Solis
16765 (US); Mun. Chaupi, NE slope of Volcan
Illiniza Norte, 00°38’S, 78°42°W, 4000-4050 m,
12 Oct 2000, D. Stanctk 4024, 4031B, 4033B
(PRC, QCA); swampy patches, 00°38’S, 78°42’ W,
4000-4050 m, 12 Oct 2000, D. Stanctk 4034 (AAU,
PRC, QCA, US); Mun. Lasso, Parque Nacional
Cotopaxi, NNE slope of Volcan Cotopaxi, valley
of Quebrada Taniloma, 00°36.8’S, 78°24.3’°W,
3750 m, 21 Jun 1999, D. Stancik 3080 (AAU, PRC,
QCA, W); E slope of Volcan Cotopaxi, 00°38’S,
78°25'W, 4300 m, 28 Sep 2000, D. Stancik 3880
(AAU, PRC, QCA); plain below SE slope of Volcan
Cotopaxi, 0°37°S, 78°24’ W, 3800-3900 m, 28 Sep
2000, D. Stancik 3883 (PRC, QCA). Pichincha:
Iter Regnellianum quartum, Flora Aequatoriensis,
prov. Pichincha, Sincholagua, 4200 m, 7 Jul 1939,
E. Asplund 8641 (S-paratype); mountain ridge ca.
2 km to the W from Cerro Saraurcu, 4200-4350 m,

Sklenar & Kosteckova 1322 (AAU); Volcan
Iliniza, NE slope below the refugio, 00°39’S,
78°42'°W, 4430 m, Holm-Nielsen 24871 (AAU);
Entre Pifo y boqueron de Cerro Corrales, Paramo
de Guamani, 4050 m, Barclay & Juajibioy 8905
(COL, MO); Volcan Antisana, between campsite
IMAP and Laguna Micacocha, 00°33’S, 78°12’ W,
3850-3950 m, S. Laegaard 101583 (AAU, QCA,
QCNE); N side, ca. 12 km along road from Haci-
enda Antisana, 00°27’S, 78°10°W, S. Laegaard
102892 (AAU); 00°30’S, 78°00°W, 4100 m, Grubb
et al. 620 (NY, US); NE slope of Rucu Pichincha,
00°10’S, 78°34’ W, 4300 m, Sklenar & Kosteckova
1-24 (NY). Tungurahua: Comunidad Rumipata,
pasture in paramo, 00°22’S, 78°55’W, 4000 m,
Brandbyge 42597 (AAU, MO).

34b. Festuca chimborazensis subsp. micacochen-
sis Stancik, Folia Geobot. Phytotax. 39(1): 105,
f. 3, 6-10. 2004. Type: Ecuador. Pichincha/
Napo, Volcan Antisana, between Campamento
IMAP and Laguna Micacocha, 00°33’S,
78°12'W, naked soil on trail, 3850-3950 m,
7 Mar 1992, S. Laegaard 101612 (holotype:
AAU!; isotypes: PRC!, QCA!, QCNE!).

Culms 22-40 cm tall. Leaf blades 11-15 cm
long, mostly straight; leaf cross-sections with (5—)7
vascular bundles. Panicles 8-10 cm long, ca. 7 mm
wide. Spikelets with upper glumes 3.5—4 mm long.

Distribution and habitat.—Festuca chim-
borazensis subsp. micacochensis has so far only
been recorded from three sites: western slopes of
Antisana, northern slopes of Chimborazo, and at
Guagua Pichincha. At all three sites this subspe-
cies 1s sympatric with F! chimborazensis subsp.
chimborazensis.

Additional specimens examined. ECUADOR.
Bolivar: 55.4 km SW of Ambato on Highway to
Guaranda, 4050 m, 3 May 1990, PM. Peterson
8982 & C_R. Annable (K, MO, QCA, QCNE, US);
km 42.7, 4020 m, 3 May 1990, P.M. Peterson 8973
& C.R. Annable (MO, QCA, QCNE, US); along
main road ca. 1 km W of Cruce de Los Arenales,
01°25’S, 78°54’ W, 4050-4100 m, S. Laegaard
101517 (AAU). Pichincha: Pifo—Pintag road, in
valley 2.5 hours horseride above Inga Monserrat,
00°19’S, 78°17’ W, 3625—3725m, 11 Apr 1992,
S. Laegaard 102253 (AAU, QCA, QCNE); road
Lloa—Guagua Pichincha, km 11, 00°12’S, 78°35’ W,
4300 m, S. Laegaard et al. 102744 (AAU); S. Lae-
gaard et al. 102739 (AAU, QCA).

Festuca in South American Paramos 95

35. Festuca chita Stancik, Darwiniana 41(1-4):
129, f. 13a—f. 2003. (Figs. 45, 49). Type:
Colombia. Boyaca, Cafion del Chicamocha,
Mun. Chita, Vereda Los Colorados, 3300 m, 14
May 1991, Etter 66/ (holotype: COL!).

Tussocked perennials with intravaginal inno-
vations. Culms 40—60 cm tall, erect, glabrous;
nodes 1, basal; cataphylls small, coriaceous. Leaf
sheaths membranous, brown to purplish-brown,
glabrous; auricles absent; ligules 0.30.5 mm long,
membranous, apex truncate, short-ciliate; blades
10—15 cm long, 0.4—0.6 mm wide, involute, green,
abaxially glabrous, apex obtuse. Panicles ca. 8
cm long, ca. 3 cm wide, contracted; branches gla-
brous. Spikelets 8—9 mm long, lanceolate, florets
3 or 4; rachilla densely hairy; glumes 5.5—8(—8.5)
mm long, coriaceous, dark purple; lower glumes
5.5—6.5 mm long, narrowly lanceolate, 1-nerved;
upper glumes 7—8(—8.5) mm long, lanceolate,
3-nerved, sparsely scabrous; lemmas 6.5—7.5 mm
long, lanceolate, 5-nerved, coriaceous, dark purple,
scabrous and short-hairy, entire, awned, the awn
0.5—1.5 mm long; callus glabrous or sparsely hairy;
paleas 2/3—4/5 as long as the lemma, glabrous,
apex hairy; anthers 0.8—1.1 mm long; ovary apex
glabrous. Caryopses lanceolate; hilum 1/2 as long
as the grain.

Leaf blade anatomy.—Cross-sections with
7-11 vascular bundles and 5 ribs, sclerenchyma
under abaxial epidermis discontinuous, extending
to all the vascular bundles, adaxial sclerenchyma
present, extending to vascular bundles only excep-
tionally; bulliform cells absent; adaxial epidermis
hairy, the hairs ca. 0.5 mm long.

Geographical distribution.—This species is
endemic to the Colombian Cordillera Oriental, at
Cafion del Chicamocha and is only known from the
type locality at an altitude of 3300 m. The Canon
del Chicamocha is a dry valley without typical
paramo vegetation.

Additional specimens examined. COLOMBIA.
Boyaca: Cafion del Chicamocha, Mun. Chita,
Vereda Los Colorados, 3300 m, 27 Jan 1927, Killip
18479 (COL).

36. Festuca cleefiana E.B. Alexeev, Bot. Zhurn.
(Moscow & Leningrad) 69: 1548. 1984. (Figs.
44, 46, 90C—F). Tyre: Colombia. Boyaca,
Paramo de la Rusia, NW-N de Duitama Aislada,
3580 m, 7 Dec 1972, A.M. Cleef 6826 (holo-
type: US-2785718!; isotypes: COL!, U!).

Densesly tussocked perennials with intra-
vaginal innovations. Culms 70—130(—150) cm tall,
erect, glabrous sometimes finely scabrous; nodes
1 or 2(—3). Leaf sheaths, membranous to coria-
ceous, grayish-brown, glabrous; auricles absent;
ligules (0.5—)1—2(-2.5) mm long, membranous to
coriaceous, apex mostly obtuse, short-ciliate; blades
30—50(—60) cm long, 0.6—0.9(—1.1) mm wide, con-
duplicate, abaxially glabrous, green, apex obtuse.
Panicles 10—20(—25) x 3—10(-15) cm, flexuous,
branched, erect, oblong; branches finely scabrous.
Spikelets 8-11 mm long, oblong, florets 4-6(—7);
rachilla long, densely pilose; glumes (3.5—)4—7.5
(—8) mm long, narrowly lanceolate, coriaceous,
purple or purplish-green, upper 1/4 scabrous, apex
acute; lower glumes (3.5—)4—6(—6.5) mm long,
1-nerved; upper glumes (5.5—)6—7.5(—8) mm long,
3-nerved; lemmas 6—7.5(—8) mm long, lanceolate,
membranous to coriaceous, 5-nerved, purplish-
green, scabrous to densely hairy, apex short two-
dentate, awned, the awn 0.5—2.5 mm long; paleas
as long as the lemma, papillose, upper 1/3 hairy;
lodicules oblong-lanceolate, acuminate; anthers
(2—)2.5—3 mm long; ovary apex glabrous. Cary-
opses oblong-lanceolate; hilum 3/4 as long as the
grain, linear.

Leaf blade anatomy.—Cross-sections with
7—-9(—11) vascular bundles and 5-9 ribs; scle-
renchyma under abaxial epidermis continuous
or discontinuous, extending to all the vascular
bundles; adaxial epidermis present, extending to
all or every other vascular bundle; bulliform cells
absent; adaxial epidermis densely hairy, the hairs
0.02—0.07 mm long.

Observations.—Festuca cleefiana 1s an easily
recognizable species, forming relatively large tufts
with flexuous, ramified panicles and spikelets with
typically long glumes.

Distribution and habitat.— Festuca cleefi-
ana is endemic to Colombian Cordillera Oriental
(Cundinamarca, Boyaca, Santander). It is a typical
species of the grass paramo occurring between
3300-4000 m. This species is known from the
communities of Festuca dolichophylla & Pas-
palum bonplandianum (Rangel & Sturm 1995),
Sphagno magellanici-Chusquetum tesellatae
(Rangel & Sturm 1995), Hypericum juniperinum
(Cleef 1981).

Additional specimens examined. COLOM-
BIA. Cundinamarca: Mun. Bogota, valley
between Usme and Nazareth, 4°13’N, 74°11 °W,
3480 m, 16 Jul 1998, D. Stancik 272 (PRC); Mun.

96 Festuca in South American Paramos

= = eo
—S— — ae

==

CE Zi
LGM e
Cee
————_
SSS

Figure 45. Festuca chita. A. Stylized growth form. B. Habit. C. Spikelet. D. Glumes. E. Lemma. F. Leaf blade cross-section. A-F,
Etter 661 (COL).

Festuca in South American Paramos 97

Figure 46. Festuca cleefiana. A. Stylized growth form. B. Habit. C. Ligule. D. Spikelet. E. Glumes. F. Lemma. G. Lemma with
palea and rachilla. H. Leaf blade cross-section. A-H, Stancik 1421 (PRC).

Villapinzon, Paramo La Calera, 5°12’N, 73°33 ’W,
3500 m, 22 Jul 1978, D. Stancitk 409, 466, 467
(COL, PRC). Boyaca: Mun. Aquitania, Paramo
El] Guarne, 3500 m, 8 Feb 1999, D. Stancitk 2160,
2188, 2189 (COL, FMB, COL); Mun. Aquitania,
road to Sisvaca, km 6, 2900 m, 10 Feb 1999,

D. Stancik 1920 (COL, FMB, PRC); D. Stancik
2257 (COL, FMB, PRC); km 2, 3350 m, 14 Jan
1999, D. Stancik 1950 (PRC); 3240 m, 14 Jan
1999, D. Stancik 1964 (COL, FMB, PRC); 3450
m, 14 Jan 1999, D. Stancik 1935 (COL, FMB,
PRC); Mun. Aquitania, Paramo Sarna, 3400 m,

98 Festuca in South American Paramos

20 Jan 1999, D. Stancik 2000 (COL, PRC); Mun.
Arcabuco, Santuario Iguaque, Laguna Iguaque,
3600 m, 14 Oct 1998, D. Stanctk 924, 926, 945
(COL, FMB, PRC); Lagun Ojo de Agua, 3650 m,
14 Oct 1998, D. Stancitk 927 (COL, FMB, PRC);
Mun. Arcabuco, SW slope of Cerro Volcan,
3000 m, 20 Nov 1998, D. Stancik 1407 (COL,
FMB, PRC); Mun. Belén, Paramo de Gina, 3250
m, 6 Feb 1999, D. Stancik 2130, 2136 (COL, PRC);
3300 m, 6 Feb 1999, D. Stancik 2137, 2138 (PRC);
Mun. Duitama, Paramo de La Rusia, 5°54’27°N,
73°4'44’W, D. Stancik 1004 (COL); 3750 m, 12
Feb 1999.2). Stanctk, 2367, 2372,2373 (COL,
PRC); 3580 m, 3540 m, Hernandez 1311 (COL);
Paramo Pan de Azucar, 3400 m, 30 Nov 1998, D.
Stancik 1421, 1422 (COL, FMB, PRC); 3350 m,
30 Nov 1998, D. Stanctk 1423 (COL, FMB, PRC);
3400 m, 30 Nov 1998, D. Stancik 1482 (COL,
FMB, PRC, US); Mun. El Cocuy, Parque Nacio-
nal El Cocuy, Las Cabanas Kanwara, 4070 m, 30
Dec 1998, D. Stanctk 1827, 1865 (COL, FMB,
PRC); 4350 m, 30 Dec 1998, D. Stancik 1828,
1830 (COL, FMB, PRC); pass between Cocuy
and Chita, 4000 m, 3 Feb 1985, Wood 5141 (COL,
K); Mun. Chinavita, Cerro Mamapacha, 2900 m,
11 Dec 1998, D. Stancik 1602, 1635, 1640 (COL,
FMB, PRC); 2850 m, 11 Dec 1998, D. Stancik
1629 (COL, PRC); 11 Dec 1998, D. Stancik 1576,
1577,, 1628 (COLG EMBs BR); 11 Wee 199s:
D. Stancik 1580 (COL, PRC); Mun. Mongui,
Laguna La Colorada, 3550 m, 21 Jan 1999, D.
Stancitk 2020 (COL, FMB, PRC); 3650 m, 21 Jan
1999, D. Stancik 2045 (COL, FMB, PRC); Mun.
Paipa, Cuchilla El Paramo, 3200 m, 4 Dec 1998,
D. Stanctk 1503; 1532, 1535, 1956 (ER©): Mun.
Santa Rosa de Viterbo, Alto Lamadero, 3200 m,
30 Nov 1998, D. Stanctk 1438, 1457, 1463, 1465
(COL, PRC); Mun. Siachoque, Paramo Siachoque,
05°30’N, 73°8’W, 3650 m, 24 Jan 1999, D. Stancik
2051, 2050, 2054 (COL, FMB, PRC); Mun.
Socota, Paramo Pisva, lagoon Chorro Negro, 3500
m,.11 Feb 1999, D. Stanctk 2346, 2350 (COL,
FMB, PRC); Alto de Cardon, 3540 m, 11 Feb
1999, D. Stancik 2314 (COL, FMB, PRC); Alto de
Calarca, 3500 m, 11 Feb 1999, D. Stancik 2313,
2332 (COL, FMB, PRC); Mun. Sotaquira, Paramo
Chontales, 3050 m, 15 Dec 1998, D. Stancik
1688, 1689, 1690 (COL, FMB, PRC); 14 Nov
1998, D. Stanctk 1339 (COL, FMB, PRC); Mun.
Pesca Paramo Cortadero, vereda Puerta Chiquita,
3500-3700 m, 15 Oct 1981, Bejarano 44 (COL);
Paramo de Belén, 3500 m, 15 Nov 1999, Rangel

et al. 3515 (COL); Mun. Toca, Paramo Cortadero,
5°30’N, 73°15’ W, 3450 m, 14 Nov 1998, D. Stancik
1339 (COL, FMB, PRC); 3350 m, 14 Nov 1998,
D. Stanctk 1366 (COL, FMB, PRC); 3600 m, 14
Nov 1998, D. Stancitk 1373 (COL, FMB, PRC);
Mun. Labranzagrande, Paramo Franco, 3300 m,
9 Feb 1999, D. Stancik 2224, 2230, 2232, 2236,
2240 (COL, FMB, PRC); Mun. Labranzagrande,
Valle de Rio Cushiana, 5 km from Toquilla,
2950 m, 9 Feb 1999, D. Stancik 2249 (COL, FMB,
PRC). Santander: Mun. Concepcion, vereda
Juradito, Paramo de Gallina, 3500 m, 24 Feb 1999,
D. Stancik 2495, 2502, 2503, 2505, (COL, FMB,
PRC); 24 Feb 1999, D. Stancik 2489 (PRC).

37. Festuca cocuyana Stancik, Darwiniana 41
(1-4): 133, f. 12a—e. 2003. (Figs. 39, 47,
91A—-D). Type: Colombia. Boyaca, Mun.
Cocuy & Guican, Parque Nacional El Cocuy,
Alto de las Cuevas. Grass paramo with Cala-
magrostis effusa, Espeletia sp., Acaulimlva sp..,
etc., 3850 m, 30 Dec 1999, D. Stancik 1886
(holotype: PRC!; isotypes: COL!, FMB!).

Tussocked perennials with intravaginal innova-
tions. Culms 20—50 cm tall, erect, glabrous; nodes 1,
basal. Leaf sheaths membranous to coriaceous, stra-
mineous, glabrous, finely striate; auricles absent;
ligules 0.6—1 mm long, membranous to coriaceous,
apex truncate; blades 15—20 cm long, 0.8—1.5
(—2) mm wide, conduplicate or flat, abaxially gla-
brous. Panicles 8—12 cm long, 0.8—1.5 mm wide,
contracted, narrow, elongate. Spikelets 10-13 mm
long, lanceolate, florets 2 or 3; rachilla pubescent;
glumes 7.5—10.5 mm long, almost as long as the
spikelet, lanceolate, membranous, scabrous along
midnerve, apex acute; lower glumes 7.5—8 mm
long, 1-nerved; upper glumes 7.5—10.5 mm long,
3-nerved; lemmas 8—10 mm long, lanceolate,
5-nerved, membranous, awnless or short-awned, the
awn ca. 0.5 mm long; callus glabrous or sparsely
hairy; paleas 3/4 as long as the lemma, papillose,
distally scabrous; lodicules 1—1.4 mm long, lanceo-
late, two-dentate; anthers 0.8—1.1 mm long; ovary
apex glabrous. Caryopses lanceolate; hilum 3/4 as
long as the grain, linear.

Leaf blade anatomy.—Cross-sections with
(8—)10—11 vascular bundles and 5-7 ribs above;
sclerenchyma under both abaxial and adaxial
epidermis, discontinuous; sclerenchyma girders
absent; bulliform cells absent; adaxial epidermis
with scattered hairs, the hairs ca. 0.02 mm long.

99

Festuca in South American Paramos

E. Glumes. F. Lemma. G. Lemma with

Figure 47. Festuca cocuyana. A. Stylized growth form. B. Habit. C. Ligule. D. Spikelet.

palea and rachilla. H. Leaf blade cross-section. A-H, Stancik 1886 (PRC).

100

Observations.—Festuca cocuyana is morpho-
logically similar to F. sanctae-martae, an endemic
species of Sierra Nevada de Santa Marta. Festuca
cocuyana differs from F: sanctae-martae by having
shorter culms (20-50 verses 60—80 cm), shorter leaf
blades (15—20 verses 25—35 cm), and shorter lower
glumes (7.5—8 verses 8.5—9.5 mm).

Distribution and habitat.— It is endemic to
the Sierra Nevada del Cocuy in the Colombian
Cordillera Oriental. It occurs between 3800-4300
m, in grass paramo dominated by Calamagrostis
effusa and Espeletia spp., and on rocky slopes in
the superparamo.

Additional specimens examined. COLOMBIA.
Arauca: Sierra Nevada del Cocuy, cabeceras de la
Quebrada El Playon, Patio Bolos 2.5 km, al NW
de la Laguna La Plaza, hoya S. José, 4300 m, 10
Mar 1973, Cleef 9078 (COL); S of the Lagoon La
Plaza, 4300 m, 31 Dec 1985, Wood 5260 (COL,
FMB, K).

38. Festuca cundinamarcae E.B. Alexeev, Bot.
Zhurn. (Moscow & Leningrad) 69: 1548.
1984. (Figs. 48, 49, 91E & F,92A & B). Tyre:
Colombia. Cundinamarca, Macizo de Bogota,
Cerro Diego Largo, paramo, 3540 m, 14 Jan
1940, J. Cuatrecasas 7966 (holotype: S!; iso-
types: COL!, U!, US!).

Tussocked perennials with intravaginal
innovations. Culms 60—100(—120) cm tall, erect,
scabrous; nodes | on distal half. Leaf sheaths coria-
ceous, stramineous, glabrous; ligules 0.1—0.3 mm
long, coriaceous, apex truncate, short-ciliate;
blades 30-50 cm long, 0.7—1(—1.2) mm wide,
conduplicate, rigid, abaxially glabrous, dark
green, apex obtuse or mucronate. Panicles 10—20 x
0.5—1.5 cm, contracted, narrow; branches scabrous.
Spikelets 9-13 mm long, oblong-lanceolate, florets
5—7(—8); rachilla sparsely short-pilose; glumes 3—
6 mm long, lanceolate, membranous to coriaceous,
green, scabrous distally, apex acute; lower glumes
34.5 mm long, 1-nerved; upper glumes (4.5—)5—6
mm long, 3-nerved; lemmas (5.5—)6—7(—7.5) mm
long, lanceolate, 5-nerved, membranous to coria-
ceous, glabrous, upper 1/3 scabrous, apex entire,
usually awned, the awn up to 1 mm long; paleas
as long as the lemma, papillose, upper 1/3 hairy;
lodicules ca. 1.1 mm long, lanceolate, acuminate;
anthers 2-3 mm long; ovary apex glabrous. Cary-
opses oblong-lanceolate; hilum 2/3 as long as the
grain, linear.

Festuca in South American Paramos

Leaf blade anatomy.—Cross-sections with
9-15 vascular bundles and 7—13 ribs; sclerenchyma
under abaxial epidermis discontinuous, extending
to all or every other the vascular bundles, adaxial
sclerenchyma present, often extending to the
vascular bundles forming girders; bulliform cells
absent; adaxial epidermis densely hairy, the hairs
0.09—0.2 mm long.

Distribution and habitat.—Festuca. cun-
dinamarcae is endemic to Colombia, where it is
known only from the Cordillera Oriental (Cun-
dinamarca, Boyaca, Meta). It is a common species
of grass paramos growing between 2700-3700 m.
This species occurs in communities such as: Pen-
tacalio nitidae-tessellatae (Sanchez & Rangel
1990), Calamagrostietum planifoli-effusae (Var-
gas & Zuloaga 1985), Espeletietum argenteae-
Calamagrostiosum effusum (Cuatrecasas 1934),
Bartsio santolinifoliae-Calamagrostietum effusae
(Rangel & Ariza 2000).

Additional specimens examined. COLOMBIA.
Boyaca: Boyaca, 2700 m, 15 Apr 1964, Saravia
3912 (COL, PRC); Mun. Pesca, Puerta Chiquita,
Paramo de Cortadera, 3500—3700 m, 15 Oct 1981,
Bejarano 22 (UPTC); Mun. Aquitania, road Suse—
Sisvaca, km 1.5—2, 3050 m, 13 Jan 1999, D. Stancik
1939, 1951, 1952 (COL, FMB, PRC); 14 Jan 1999,
D. Stancik 1944 (PRC); Paramo Sarna, 3350 m,
20 Jan 1999, D. Stancik 1987, 1988 (COL, FMB,
PRC); 3400 m, 20 Jan 1999, D. Stancik 2005 (COL,
FMB, PRC); 3200 m, 20 Jan 1999, D. Stanctk 1985,
1986 (COL, FMB, PRC); Mun. Aquitania, Paramo
Suse, 3350 m, 14 Jan 1999, D. Stanctk 1929 (COL,
PRC); 14 Jan 1999, D. Stancitk 1945 (PRC); 14
Jan 1999, D. Stancik 1922 (COL, FMB, PRC);
Mun. Belén, Paramo Gina, 3250 m, 6 Feb 1999,
D- Stancik 2109, 2115, 2116, 2HI7, 2120 CO
PRC); 6 Feb 1999, D. Stanctk 2174 2/s6( GO
FMB, PRC); Mun. Mongui, Laguna La Colorada,
3300 m, 21 Jan 1999, D. Stancik 2019 (COL, FMB,
PRC); Mun. Toca, Paramo Cortadero, 05°30'N,
73°15’ W, 2700 m, 14 Nov 1998, D. Stancik 1402
(COL, FMB, PRC); 3200 m, 14 Nov 1998, D.
Stanctk 1347, 1360, 1363, 1396 (COL, FMB, PRC);
4 Dec 1998, D. Stancik 1513 (COL, FMB, PRC);
Mun. Samaca, Paramo Rabonal, 2000 m, 1 Nov
1998, D. Stancik 1286 (COL, FMB, PRC); Mun.
Toquilla, Valle of Rio Cushiana, 2950 m, 9 Feb
1999, D. Stancik 2250 (COL, FMB, PRC); Mun.
Tutasa, Paramo de Carnicerias, 3300 m, 26 Feb
1999, D. Stanctk 2597, 2599, 2612 (PRC); Mun.
Paipa, cuchilla El Paramo, 3250 m, 3 Dec 1998,

Festuca in South American Paramos 101

Figure 48. Festuca cundinamarcae. A. Stylized growth form. B. Habit. C. Ligule. D. Spikelet. E. Glumes. F. Lemma. G. Lemma
with palea and rachilla. H. Leaf blade cross-section. A-H, Stancik 3539 (PRC).

102

-75

Festuca in South American Paramos

COLOMBIA

200 400 Kilometers

-65

Figure 49. Distribution of Festuca glumosa (@), F. cundinamarcae ( A), F- chita (m), and F:turimiquirensis (@).

D. Stancik 1523 (COL, FMB, PRC); 4 Dec 1998,
D. Stancik 1511 (COL, PRC); Mun. Susacon, km
20, margin of river Susacon, 3150 m, 23 Feb 1999,
D. Stancik 2472 (PRC); 23 Feb 1999, D. Stancitk
2471, 2473 (COL, FMB, PRC). Cundinamarca:
Cordillera de Bogota, 2900 m, Nov 1856, Triana
s.n. (US); Bogota, Paramo de Monserrate, vereda
El Verjon, 3000-3200 m, 29 Jul 1988, Garzon
617 (COL); 3160 m, 15 Feb 1980, Zuluoga 107
(COL); 3000-3300 m, 16 Apr 1944, Killip et al.
38050 (COL); 3125m, 24 May 1968, Valder-
rano 34 (UPTC); 3125m, 26 Apr 1968, Sturm
33 (UPTC); Mun. Bogota, Alto de la Viga, 1 Feb
2000, D. Stancik 3539 (COL, PRC); Paramo de
Cruz Verde, 3400-3500 m, 15 Sep 1940, J. Cua-
trecasas 10446 (COL, US); 04°34’N, 74°01°W,
3500-3700 m, 17 Jun 1995, S. Laegaard & May-
orga 17536 (AAU, COL); Sumapaz—Santa Rosa,
3600 m, 15 Jul 1984, Wood 4539 (COL, FMB);

Hoya de Quebrada Sitiales, 1.5 km NW de Laguna
La Guitarra, 26 Jan 1972, Cleef 1090 (COL, U);
Paramo Chingaza, sector Rio La Playa, Valle de
Fraillejon, 3250 m, 14 Nov 1992, Guerrero 152
(COL); near La Calera, 3500 m, 1983, Wood 3523
(AAU, COL, K); Paramo de Palacio, La Sibéria,
3400 m, 30 Feb 1961, Rodriguez 4130 (COL);
Paramo between Bogota and Choachi, 3320 m,
7 Jan 1974, G. Davidse et al. 5545 (COL, US);
Paramo entre Cogua y San Cayetano, cercania de
la Laguna Seca, 3700 m, 12 Feb 1972, Cleef 6285
(COL); Paramo de Guasca, 3200-3300 m, 2 Jun
1940, J. Cuatrecasas 9475 (COL, US). Meta:
Paramo Sumapaz, Hoya Sitiales, laguna La Prima-
vera, 3580 m, 25 Jan 1972, Cleef 982A (COL).

39. Festuca densipaniculata E.B. Alexeev, Bot.
Zhurn. (Moscow & Leningrad) 69: 1551.
1984. (Figs. 50, 52, 92C—F). Type: Ecuador.

Festuca in South American Paramos 103

WV

SZ
SSS

lie

Z SIE we
SL YAW

—S

Z
“a2

MN
Ap

Figure 50. Festuca densipaniculata. A. Stylized growth form. B. Habit. C. Ligule. D. Spikelet. E. Glumes. F. Lemma. G. Lemma
with palea and rachilla. H. Leaf blade cross-section. A-H, Asplund 8397 (S).

104

Chimborazo, southern slope of Mt. Chimbo-
razo, sandy ground, 4600 m, 18 Mar 1939,
E. Asplund 8397 (holotype: S!).

Tussocked perennials with intravaginal innova-
tions. Culms 20—50 cm tall, erect, finely scabrous;
nodes 1, basal. Leaf sheaths coriaceous, stramineous,
glabrous or scabrous; ligules 0.8—1.5(—1.7) mm long,
membranous, apex truncate, short-ciliate; blades
17-25 cm, long, 0.8—1.2 mm wide, conduplicate,
abaxially scabrous, glaucous, apex pointed. Panicles
5—10 x 1.5—2 cm, contracted, dense, lanceolate to
oblong; branches finely scabrous throughout. Spike-
lets 8—9(—10) mm long, elliptic to oblong, florets 3
or 4; rachilla densely hairy; glumes 4—6 mm long,
membranous to coriaceous, green and somewhat
glaucous, apex acute; lower glumes 4—5 mm long,
lanceolate to oblong-lanceolate, 3-nerved, glabrous
or upper 1/3 scabrous; upper glumes (4.5—)5—6 mm
long, oblong to oblong-lanceolate, 5-nerved, gla-
brous or upper 1/2 scabrous; lemmas (6.5—)7—8.5
mm long, lanceolate, S—7- nerved, membranous,
green or sometimes purplish, glaucous, upper 1/2 or
entire surface scabrous, awned, the awn 0.7—1 mm
long; callus glabrous; paleas as long as the lemma,
lanceolate, membranous, upper 1/3 and along keels
scabrous to hairy; lodicules 1.4—2 mm long, oblong;
anthers (2.3—)2.7—2.8 mm long; ovary apex gla-
brous. Caryopses lanceolate to oblong-lanceolate;
hilum 1/2 as long as the grain, linear.

Leaf blade anatomy.—Cross-sections with
(7—)11—15 vascular bundles and 5-7 ribs above;
sclerenchyma under abaxial epidermis continuous,
adaxial sclerenchyma present, extending to every
other vascular bundle forming girders; abaxial epi-
dermis with a dense covering of prickles, adaxial
epidermis with frequent hairs, the hairs 0.2-0.4 mm
long.

Observations.—The type specimen of F. den-
sipaniculata was originally annotated by Alexeev
as “Festuca densiflora” but he published it as
F. densipaniculata because of the earlier hom-
onym, F: densiflora Tovar. There are only two
known collections of this species. Leaf anatomi-
cal characteristics suggest that F’ subulifolia and
F. asplundii may be closely related. Otherwise, the
status of this species is not readily apparent and
molecular studies will be needed for elucidation
of its affinities.

Distribution and habitat.—Festuca densipan-
iculata is known only from grass paramos of central
Ecuador between 4000-4600 m.

Festuca in South American Paramos

Additional specimens examined. ECUADOR.
Chimborazo: S slope of Mt. Chimborazo, sandy
ground, 4600 m, EF. Asplund &396 (S-paratype).
Napo: Antisana, 22 Jul 1997, Sklenar & Kosteckova
2738 (PRC).

40. Festuca dinirica Stancik, Novon 14(3):
341-343, f. 1f. 2004. (Figs. 44, 51). Type:
Venezuela. Lara, Mun. Humocaro Alto, Parque
Nacional Dinira, 09°35’39"N, 70°07’ 12’W,
grass paramo with shrubby patches with
Rhynchospora sp., Chusquea sp., Blechnum
sp., Calamagrostis effusa, Espeletia sp., and
Diplostephium sp., 3170 m, 30 Nov 2000,
D. Stancik 4287 (holotype: PRC!; isotypes:
CAR!, COL!).

Tussocked perennials with intra- and extra-
vaginal innovations. Culms 50—60 cm tall, erect,
glabrous; nodes 1, in distal half. Leaf sheaths coria-
ceous, brown, glabrous, fibrous near base; auricles
absent; ligules 0.5—1 mm long, membranous,
apex truncate; blades 20—25 cm long, 0.4—0.6 mm
in diameter, conduplicate to involute, abaxially
glabrous, green. Panicles 10-12 x 0.5—1.5 cm,
contracted, erect; branches glabrous. Spikelets
8—10 mm long, lanceolate sometime oblong-
lanceolate, florets 3 or 4; rachillas 1—-1.2 mm long,
sparsely hairy; glumes 3.4—6 mm long, lanceolate,
membranous to coriaceous, violet, apex acute;
lower glumes 3.4—4 mm long, 1-nerved; upper
glumes 5.5—6 mm long, 3-nerved; lemmas 6-7 mm
long, lanceolate, 5-nerved, coriaceous, violet,
scabrous, awned, the awn 1.5—4 mm long; callus
glabrous; paleas as long as the lemma, lanceolate,
membranous, scabrous; anthers 2—2.5 mm long;
ovary apex glabrouse. Caryopses lanceolate; hilum
4/5 as long as the grain, linear.

Leaf blade anatomy.—Cross-sections usu-
ally with 5 vascular bundles and 3—5 ribs above;
sclerenchyma under abaxial and adaxial epidermis
discontinuous; adaxial epidermis hairy, the hairs
ca. 0.09 mm long.

Observations.—The spikelet structure of
F. dinirica resembles F. tolucensis, another fre-
quent species of the Venezuelan paramos. However
F. dinirica clearly differs from F: tolucensis by its
smaller culms (40-60 versus 60-80 cm), mixed
innovations (versus intravaginal), brown, striate,
and fibrous leaf sheaths (versus stramineous and
non-fibrous), shorter ligules (0.5—1 mm versus
1.8—3.5 mm), and anatomically with discontinuous

Festuca in South American Paramos 105

Figure 51. Festuca dinirica. A. Habit. B. Ligule. C. Spikelet. D. Glumes. E. Lemma. F. Lemma with palea and rachilla. G. Leaf
blade cross-section. A—G, Stancik 4288 (PRC).

106

-75

Festuca in South American Paramos

-70

COLOMBIA

aoa

-80 -75

400 600 Kilometers :

Figure 52. Distribution of Festuca glyceriantha (@), F- holubii (A), F: densipaniculata (m), and F- monguensis (@).

abaxial sclerenchyma and all vascular bundles
free (versus continuous abaxial sclerenchyma with
some girders persent). The characteristics of mixed
innovations with brown, striate, fibrous sheaths in
F: dinirica suggest a close relationship with F’ sect.
Aulaxyper. In South America, this section is not very
common and is still poorly studied. The Colombian
and Ecuadorian species, F: andicola, is geographi-
cally the nearest species within F: sect. Aulaxyper.
Distribution and habitat.—Festuca dinirica is
endemic to Venezuela (Lara) and occurs at an alti-
tude of about 3200 m where it forms small tussocks
in shrubby-grassy paramos and is associated with
various species of Blechnum, Espeletia, Hypericum,
Calamagrostis, Chusquea, and Rhynchospora.
Additional specimens examined. VENEZU-
ELA. Lara: Mun. Humocaro Alto, Parque Nacio-
nal Dinira, 09°35’39”"N, 70°07'12”W, 3170 m, 30
Nov 2000, D. Stancik 4288 (CAR, COL, PRC).

41. Festuca glumosa Hack. ex E.B. Alexeev, Bot.
Zhurn. (Moscow & Leningrad) 69(11): 1549.
1984. (Figs. 49, 53, 93A—D). Tyre: Ecuador.
In pasq. andinis, 1886, A. Sodiro 36/4 (holo-
type: W!)

Festuca ovina subvar. jamesonii St.-Yves, Can-
dollea 3: 166. 1927. Type: Ecuador Andes
de Quito, 15000 ft, Jameson 230 (lectotype:
G-DC!, designated here).

Densely tussocked perennials with intra-
vaginal innovations. Culms 15—55 cm tall, erect,
finely scabrous; nodes 1, basal with 2 leaves. Leaf
sheaths membranous to coriaceous, firm, stramin-
eous glabrous, inconspicuously striate; ligules
0.8—1.2 mm long, apex two-dentate, short-ciliate;
blades 10—25 cm long, 0.8—1.4 mm wide, invo-
lute, abaxially scabrous, glaucous, apex obtuse.
Panicles 8-10 x 0.6—1 cm, linear to lanceolate,

Festuca in South American Paramos 107

Figure 53. Festuca glumosa. A. Stylized growth form. B. Habit. C. Ligule. D. Spikelet. E. Glumes. F. Lemma. G. Lemma with
palea and rachilla. H. Leaf blade cross-section. A-H, Stancik 3392 (PRC).

108

contracted, dense, branches scabrous. Spike-
lets 8.5-10 mm long, lanceolate, florets 3 or 4;
rachilla with scattered hairs; glumes 4.5—6.5 mm
long, lanceolate, membranous to coriaceous,
keeled, purplish-green, upper 1/3 scabrous along
keel, apex acute; lower glumes 4.5—6 mm long,
1-nerved; upper glumes 6—6.5 mm long, 3-nerved;
lemmas 7—8 mm long, lanceolate, 5-nerved,
membranous, keeled, purplish-green, scabrous
distally, awned, the awn 0.5—1.5 mm long; cal-
lus with scattered hairs; paleas 4/5 as long as
the lemma, lanceolate, membranous, upper 1/3
and keels scabrous to hairy; lodicules lanceolate;
anthers 0.8—0.9 mm long; ovary apex glabrous.
Caryopses lanceolate; hilum 1/2—3/5 as long as
the grain, linear.

Characters F. glumosa
Leaf width (mm) 0.8—1.4
Leaf length (cm) 10-25
Ligule length (mm) 0.8-1.2
Panicle length (cm) 8-10
Spikelet length (mm) S50
Number of florets 3 or 4
Lower glume length (mm) 4.5—-6
Upper glume length (mm) 6—6.5
Lemma length (mm) 7-8
Awn length (mm) 0.51.5
Anther length (mm) 0.8—0.9

Distribution and habitat.—Festuca glumosa
ranges from central Ecuador to the northern Cor-
dillera Central in Colombia and occurs in (dry)
grasslands and rocky slopes of superparamos
between 4000-4600 m. Festuca glumosa is known
from the following communities: Diplostephio
eriophori-Loricarietum colombianae (Salamanca et
al. 1991), Cerastio floccosi-Pentacalietum gelidae
(Salamanca et al. 1991), Lupino alopecuroidis-
Agrostietum araucanae (Salamanca et al. 1991),
Baccharido caespitosae-Agrostietum araucanae
(Salamanca et al. 1991), and Aciachne acicularis-
Calamagrostis intermedia (Verweij 1995).

Additional specimens examined. COLOMBIA.
Caldas: Mun. Manizales, Parque Nacional Los Neva-
dos, between Crater La Oletta and el Ruiz, way to
Rio Nereidas, 4200-4500 m, 18 Sep 1999, D. Stancik
3393 (COL, PRC, W)); D. Stancik 3392 (COL, PRC);
D. Stancik 3413 (COL, PRC); 19 Sep 1999, Casa del
Cisne, 4180 m, 19 Sep 1999, D. Stancik 3383 (COL,
PRC); way from Rio Nereidas, km 5—7, 4000 m, 18

Festuca in South American Paramos

Leaf blade anatomy.—Cross-sections with
7 (rarely 5) vascular bundles and 5 ribs above;
sclerenchyma usually discontinuous under abaxial
epidermis and small or absent under adaxial epi-
dermis; adaxial epidermis with scattered hairs, the
hairs ca. 0.09 mm long.

Observations.—A specimen of F: glumosa
labeled as Sodiro 36/4 was found at QPLS. How-
ever, this specimen is not the type specimen since
the collection date is Sep 1887 and the locality
is “crescit in pasquis on Pichincha”. Apparently,
Sodiro used a single number to indicate the same
species. Morphologically similar species are
F. carchiense and F. imbaburensis. The species
may be distinguished by the characters shown in
the following tabular comparison.

F. carchiense F. imbaburensis

0.4—0.7 0.50.6
25-40 9-20
0.5—0.8 0.5—0.7
10-20 6-12
9—-10.5 8-10
3 or 4 2 or 3
44.5 3-3.5
5-6.5 44.2
TA 5.5-6
0.8-1.5 0.5—0.7
1.1-1.3 1-1.5

Sep 1999, D. Stancik 3394 (COL, PRC); Mun. Vil-
lamaria, Crater Olleta, 04°54’N, 75°21 W, 4600 m,
27 Jan 1988, L.G. Clark & X. Londono 376 (COL,
TULYV, UPTC, US); Refugio del Ruiz, carratera hacia
“E] Siléncio”’, 4310 m, 7 Oct 1978, Rangel et al. 1732
(COL); Cordillera Central, camino entre Buenos
Aires y la Laguna Verde, 4020-4350 m, 9 Oct 1978,
Rangel et al. 1791 (COL); Paramo entre Termales
y Nevado Libano, 4620 m, 8 Dec 1958, Barclay &
Juajibioy 6313 (COL); Risaralda: Mun. Pereira,
below Nevado El Cisne, way from Casa del Cisne to
Laguna Otun, km 5, 4100 m, 19 Sep 1999, D. Stancik
3378 (COL, PRC); Volcan de Santa Rosa, vertiente
oriental, 4500 m, 20 Feb 1980, Jaramillo-Mejia et
al. 5747 (COL); Entre Laguna El Otun y Paramo de
Santa Rosa, 4375 m, 4 Feb 1980, Jaramillo-Mejia
et al. 6149 (COL); below Nevado de Santa Isabel,
4000 m, 20 Sep 1999, D. Stancik 3373B (COL,
PRC); 4000-4100 m, 20 Sep 1999, D. Stancik 3374
(COL, PRC), 4300 m, 13 Jul 1980, Salamanca et
al. 40 (US); junto a la Laguna del Beso, 4300 m,

Festuca in South American Paramos

25 Nov 1946, J. Cuatrecasas 23201 (US); Parque
Nacional Los Nevados, vertiente W del Nevado de
Quindio, 4450 m, Salamanca et al. SS143 (U). Tol-
ima: Mun. Ibague, Nevado del Tolima, 04°38.5'N,
75°19.1°W, 4250 m, 8 Jun 2000, D. Stancik 3601,
3602, 3607 (COL, PRC); Mun. Santa Isabel, paso
del Otun a la Quebrada Africa, 3900 m, 6 Feb 1980,
Jaramillo-Mejia et al. 6210 (COL); 4200-4300 m,
Diaz—Piedrahita & Jaramillo-Mejia 1821 (COL).
ECUADOR. Chimborazo: El Altar, N side, 4300
m, Sklenar & Kosteckova 1931 (AAU); Paramo de
los Altares, 01°40’S, 78°24’ W, 4200 m, 3 Sep 1987,
Ramsay & Smith 410 (K, QCNE); Volcan Chimbo-
razo, lower refuge, 01°28°S, 78°50’ W, 4800-4840 m,
11 May 1992, S. Laegaard 102797 (AAU, QCA,
QCNE); S. Laegaard 102803 (AAU); 05 Mar 1988,
S. Laegaard & S.A. Renvoize 70555 (AAU, K, QCA,
QCNE); E slope, 4550 m, EF. Asplund 7926 (S);
Cerro Yuibug—Pailacajas, E side, 4100 m, Sklenar &
Sklenaova 2990 (AAU); W side of Cerro Yanaurcu,
4300 m, Sklenar & Kosteckova 1853 (AAU). Imba-
bura: Nevado Cotacachi, SW side, 4400 m, Sk/lenar
& Kosteckova 4300-4400 m, S. Laegaard 54515B
(AAU, QCA); Mun. Cayambe, Volcan Cayambe,
swamps below the Refugium, 00°00’N, 78°01°W,
4450 m, 20 Oct 2000, D. Stancik 4149 (PRC, QCA);
rocky superparamo, 00°00°31”S, 78°00°34.4’W,
4600 m, 20 Oct 2000, D. Stancik 4129, 4133, 4134,
4166 (AAU, PRC, QCA); D. Stancik 4122 (PRC,
QCA); Mun. Urcuqui, Cerro Yanaurcu, rocks and
pajonal, 00°29’N, 78°21” W, 4300 m, 15 Oct 2000,
D. Stancik 4039 (AAU, PRC, QCA). Pichincha:
Volcan Cayambe, 14000—14500 ft, Whymper 1330
(K); NE side, 15000 ft, Cazalet & Pennington 5749B
(US); Cazalet & Pennington 5749 (K, NY); road
to refuge, 4300-4400 m, 1 Mar 1988, S. Laegaard
70491 (AAU, K, QCA); SW side, 4600 m, Sklenar
& Kosteckova 55-2 (US); Paso de la Virgen, 4400 m,
S. Laegaard 53869 (AAU, QCA, QCNE); Paramo de
Guamani, 4200-4250 m, S. Laegaard 55718 (AAU,
QCA); 4200-4250 m, S. Laegaard 55696 (AAU);
00°20’S, 78°12’ W, 4050 m, 2 Mar 1999, S. Laegaard
etal. 19630 (AAU, QCA, QCNE); NE slope of Rucu
Pichincha, 00°10’S, 78°34’ W, 4500 m, Sklenar &
Kosteckova 1903 (AAU); NE slope, 4300-4500
m, Sklenar & Kosteckova 1933 (AAU); W slope,
4400-4500 m, 6 Mar 1988, Molau & Eriksen
3298 (AAU, GB, QCA, QCNE); 4500 m, Sklenar
& Kosteckova 9-1 (QCNE); 4350 m, E. Asplund
17334 (S); Guagua Pichincha, 00°12’S, 78°35’W,
4500-4600 m, S. Laegaard et al. 102760 (AAU);
Lloa—Guagua Pichincha road, km 10, 00°13’S,

109

78°35'W, 4170 m, S. Laegaard et al. 102725
(AAU); Pichincha, 4700 m, E. Asplund 8594 (NY,
QCA); 4100-4500 m, A.S. Hitchcock 21058 (NY,
US); Sodiro s.n. (QPLS); Volcan Antisana, N side,
00°27°S, 78°10°W, 4700 m, S. Laegaard 102889
(AAU); Nevado Cayambe, SW slope, 4400 m,
Sklenar & Kosteckova 1893 (AAU); SW side,
4500-4700 m, Sklenar & Kosteckova 735 (AAU);
4220 m, 11 May 1990, PM. Peterson 9086, E.J.
Judziewicz & R.M. King (K, MO, QCA, QCNE, US);
4400 m, 01 Jul 1995, Sklenar & Kosteckova 49-19
(QCNE); 4600 m, 2 Jul 1995, Sklenar & Kosteckova
56-2 (QCNE); E slope of Illiniza Sur, 4300 m,
Sklenar & Kosteckova 1863 (AAU); Cerro Saraurcu,
W side, 4100-4200 m, Sklenar & Kosteckova 1836
(AAU); Mt. Corazon, E slope, 4500 m, E. Asplund
17504 (S); Mun. Pifo, Paramo Guamani, grassy
paramo, 00°19’S, 78°12°W, 4300 m, 19 Jun 1999,
D. Stancik 3031, 3034 (AAU, PRC, QCA); 4000 m,
D. Stancik 3048 (PRC, QCA). Tungurahua: Cordil-
lera de Llanganatis, Paramo de Jaramillo, 01°10’S,
78°22’ W, 4000-4250 m, S. Laegaard 53293 (AAU);
S. Laegaard 53307 (AAU, QCA).

42. Festuca glyceriantha Pilg., Bot. Jahrb. Syst.
37: 516. 1906. (Figs. 52, 54, 93E & F). Tyre:
Peru. Ancash: Yungay, Yanganuco, 4100 m,
15 Jun 1903, A. Weberbauer 3275 (holotype:
B!; isotypes: BAA-1228 fragm. ex B!, US-
2875417!).

Tussocked perennials with intravaginal
innovations. Culms (30—)80—120 cm tall, erect,
glabrous; nodes 1 or 2, basal. Leaf sheaths coria-
ceous, dark brown, glabrous, striate, old sheaths
fibrous; ligules 0.5—1 mm long, apex acute, ciliate;
blades 20-60 cm long, 2—5 mm wide, conduplicate
or flat, abaxially glabrous, green or pale-green.
Panicles 15—17 x 1.5—2 cm, contracted, lanceo-
late, discontinuous; branches scabrous. Spikelets
12-15 mm long, lanceolate, florets 4(—5); rachil-
las 2—2.5 mm long, glabrous, rarely scabrous;
glumes 3.5—6.5 mm long, coriaceous, glabrous,
violet, margins membranous; lower glumes 3.5—5
mm long, lanceolate, 1-nerved, apex acute; upper
glumes (4.5—)5—6.5 mm long, lanceolate, 3-nerved,
apex acute; lemmas 6—8.5 mm long, lanceolate,
5-nerved, coriaceous, violet, upper 1/3 with brown
stripes, margin membranous, awned, the awn 0.5—
2.5 mm long; paleas almost as long as the lemma,
membranous, slightly scabrous; anthers 3—3.5 mm
long; ovary apex glabrous. Caryopses not seen.

110 Festuca in South American Paramos

= SS = =

Ve
SSS
SSS
S

——

Zz
<< =
<a : =

Figure 54. Festuca glyceriantha. A. Stylized growth form. B. Habit. C. Ligule. D. Spikelet. E. Glumes. F. Lemma with palea and
rachilla. G. Lemma. H. Leaf blade cross-section. A-H, Vega 2012 (F).

Festuca in South American Paramos iT |

Leaf blade anatomy.—Cross-sections with
(7—)11—15 vascular bundles and corresponding
ribs above; sclerenchyma under abaxial epidermis
discontinuous, extending to the vascular bundles,
adaxial sclerenchyma present, extending often to
the adaxial epidermis hairy; the hairs short.

Observations.—Festuca glyceriantha is very
distinct and easily recognized since it has dark
brown, fibrous sheaths (often resembling bulbs) and
brown stripes on the upper 1/2 of the lemma.

Distribution and habitat.—Festuca glycerian-
tha is known from northern and central Peru where
it occurs in alpine grass vegetation (jalca) between
3300—4100(-4700) m.

Additional specimens examined. PERU.
Ancash: Prov. Huaylas, Huascaran Parque
Nacional, Quebrada Los Cedros, Alpamayo trail,
4020-4700 m, 3 Nov 1985, Smith et al. 9939 (MO);
08°50’S. 77°44’ W, 4020-4700 m, 3 Nov 1985,
Smith 9942 (F, MO); Smith et al. 9944 (MO); Paron
Vall., E of lake, 08°59’S, 77°38’ W, 4200 m, 27 Sep
1985, Smith 11493 (MO); Prov. Huari, Huascaran
Parque Nacional, quebrada Rurichinchay, 09°23’S,
77°16’ W, 3840-3870 m, 13 Jun 1986, Smith et al.
12630 (US); Cordillera Blanca, 28 km SE of San
Luis on road towards Huar1, 4230 m, 23 Mar 1997,
P.M. Peterson 13875 & N. Refulio Rodriguez (US,
USM); Prov. de Bolognesi, Caillarpacsa, arriba de
Chuiguian, monte fluvial, 3900 m, 4 May 1952,
Cerrate 1540 (US). Cajamarca: Prov. Celendin,
lugar Challuayaco, sobre la carretera a Celendin, E
del Paso de Cumullca, 3550 m, 5 Dec 1984, Vega
et al. 3485 (F); Desvio a Huanico, sobre la ruta
Cajamarca—Celendin, 3500 m, 4 Oct 1987, Vega
4371 (F); Sorochuco, arriba del Punre, 06°57’S,
78°16’ W, 3340 m, 8 Nov 2001, Vega 10889 (F,
MO); El Sendamal, entre la Encanada y Celendin,
3520 m, 7 Mar 1975, Vega et al. 1695 (F); Cerca
al Centro Turistico Cumbe Mayo, 3460 m, 21 Apr
2001, Vega 10509 (F, MO); Cajamarca—Celendin
road, 07°02’S, 78°14’ W, 3450 m, 15 Jul 1983,
Smith et al. 4278 (MO); Prov. Cajamarca, cerro
Campanario, entre Cajamarca y Cooperativa
Atahualpa, 3500 m, 8 Mar 1993, Vega 66/6 (F);
Prov. Cajamarca, Cerro Maqui—Maqui, al N de
Cajamarca, ruta Shanta alta, 4120 m, 29 Jan 1994,
Vega et al. 6694 (F); Cerro Campanario, en la
SAIS Atahualpa, al NW de Cajamarca, 3500 m, 4
Nov 1994, Vega et al. 6974, 6975 (F); arboretum
de CICAFOR, 3400 m, 16 May 1981, Vega et al.
2527 (F); Cerro Sexcemayo, al W de Cajamarca,
3430 m, 4 Feb 1991, Vega 5427 (F); 3500 m, 20 Jun

1991, Vega et al. 5744 (F); Jalca de Kumullca, entre
La Encanada y Celendin, 3600 m, 27 May 1977,
Vega et al. 2012 (F); Coymolache ruta Cajamarca—
Hualgalloc, 3850 m, 7 Jan 1977, Vega et al. 2061
(F); 16 km W of Central Plaza of Cajamarca up road
towards Cumbemayo, 3440 m, 31 Mar 1997, PM.
Peterson 14011 & N. Refulio Rodriguez (MO, US,
USM); Prov. Hualgayoc, Perlamayo, Tres Lagunas,
subiendo hacia Tambillo, 3200-2850 m, 22 Jul
1986, Vega 4168 (F); arriba de la ciudad de Hual-
gayoc a 31 km de Bambamarca, 3500 m, 26 Mar
1985, Vega et al. 3797 (F); Paso de Coymolache,
km 81 de la carretera Cajamarca-Bambamarca, al
W de Hualgayoc, 3900 m, 13 Jun 1980, Vega 2263
(F); al W de Hualgayoc, 3 km sobre la carretera a
Hualgayoc—Cajamarca, 3600 m, 15 Jun 2001, Vega
10764 (F, MO); Prov. San Miguel, 64 km N of
Cajamarca and 3 km SW of El Cobro, on route 3N
towards Bambamarca, 3530 m, 16 Mar 2000, PM.
Peterson14925 & N. Refulio Rodriguez (MO, US,
USM); Prov. San Pablo, 32 km NW of Cajamarca
on road towards San Pablo via Porcon, 3470 m,
14 Mar 2000, PM. Petersonl4855 & N. Refulio
Rodriguez (MO, US, USM); 2 km SW of Hualgayoc
on road towards El Cobro and Cajamarca, 3530 m,
17 Mar 2000, PM. Peterson 14936 & N. Refulio
Rodriguez (MO, US, USM); Prov. De Hualgayoc,
cerca a Coymolache entre Hualgayoc y Cajamarca,
jalca, 3800 m, 16 Aug 1952, R. Ferreyra 8566
(US). Junin: NW part of Junin, Goyllarisquisca,
4000 m, 7 Nov 1923, A.S. Hitchcock 22328 (US).
La Libertad: Prov. Santiago de Chuco, 23 km SW
of Huamachuco on road towards Alto de Tamboras
and Pampas, 3540 m, 29 Mar 1997, P.M. Peterson
13961 & N. Refulio Rodriguez (MO, US, USM);
Santiago—Shoreyo road, 35 km from Santiago,
08°03’S, 78°18’ W, 4120 m, 26 Aug 1982, Smith
2337 (MO); Los Quinuales (al N de Quiruvilca),
ladera, 3775 m, 24 Mar 1994, Leiva & Leiva
1094 (MO); Carrion, Trujillo-—Huamachuco road,
07°56'S, 78°10’°W, 3700 m, 13 Feb 1983, Smith
et al. 3312 (MO). Prov. Huamachuco, Munmalca,
Hda. Cochabamba, 3200 m, 26 Jun 1958, Lopez &
Sagasteguii 2827 (US). Piura: Prov. Huancabamba,
Chulucanitas bajo, 3150 m, 5 Mar 1980, Vega
5271).

43. Festuca holubii Stancik, Folia Geobot. Phy-
totax. 39(1): 102, f. 1, 6-10. 2004. (Figs. 52,
55). Type: Ecuador. Loja, Cerro de Arcos W
of road Manu—Zaruma, 03°34’S, 79°28 '°W,
3250-3600 m, moist paramo, 14 Sep 1999, S.

(a2 Festuca in South American Paramos

Figure 55. Festuca holubii. A. Stylized growth form. B. Habit. C. Ligule. D. Spikelet. E. Glumes. F. Lemma. G. Lemma with palea
and rachilla. H. Leaf blade cross-section. A-H, Laegaard &Aguirre 20611 (PRC).

Festuca in South American Paramos

Laegaard & Aguirre 20611 (holotype: AAU!;
isotypes: LOJA!, PRC!, QCA!, QCNE!).

Tussocked perennials with intravaginal inno-
vations. Culms, 50-60 cm tall, erect, finely sca-
brous; nodes 1, basal. Leaf sheaths membranous,
brownish-gray, glabrous, striate; ligules 0.6—0.8
mm long, two-lobed, apex short-ciliate; blades
15—25 cm long, 0.8 mm wide, conduplicate to invo-
lute, finely abaxially scabrous, green, apex obtuse.
Panicles 8—14 cm long, ca. 0.7 cm wide, contracted,
narrow, sparsely branched; branches scabrous on
ribs. Spikelets 9-10 mm long, lanceolate, florets
3 or 4; rachilla glabrous or sparsely hairy; glumes
4—5.5 mm long, lanceolate to oblong-lanceolate,
membranous to coriaceous, keeled, purplish, gla-
brous, apex obtuse and scabrous; lower glumes ca.
4 mm long, 1-nerved; upper glumes ca. 5.5 mm
long, 3-nerved; lemmas ca. 7 mm long, oblong-
lanceolate, 5-nerved, membranous to coriaceous,
purplish-green, distally keeled, scabrous, awned,
the awn 1—2 mm long; callus glabrous; paleas 5/6
as long as the lemma, lanceolate, membranous,
scabrous only on keels; lodicules 0.9—1 mm long,
oblong, bilobed; anthers ca. 1.3 mm long; ovary
apex glabrous. Caryopses lanceolate; hilum 3/5—3/4
as long as the grain, linear.

Leaf blade anatomy.—Cross-sections with 7
vascular bundles and 5 ribs above; sclerenchyma
discontinuous, only under abaxial epidermis;
adaxial epidermis with numerous hairs, the hairs
ca. 0.09 mm long.

Observations.—Festuca holubii is morpho-
logically similar to the Peruvian F! divergens.
However, the former differs from the latter by hav-
ing taller culms (50—60 versus 20-35 cm), longer
panicles (8—14 versus 6—7 cm), shorter spikelets are
(9-10 versus 12—14 mm), and shorter lemmas (ca.
7 versus 10-12 mm).

Distribution and habitat.—Festuca holubii is
known only from the type collection in southern
Ecuador and it occurs in grass paramos between
3200-3600 m.

44. Festuca huamachucensis Infantes, Revista
Ci. (Lima) 54: 107. 1952. (Figs. 56, 57). Tyee:
Peru. La Libertad, Huamachua Prov., 7 Aug
1951, J. Infantes Vera 3538 (holotype: herb.
Infantes; isotype: US-2044946!).

Tussocked perennials with intravaginal innova-
tions. Culms 15—55 cm tall, erect, glabrous; nodes

F13

1 or 2, basal. Leaf sheaths membranous, brown,
glabrous, striate; ligules 0.5—1 mm long, apex
truncate, ciliate; blades 5—20 cm long, 0.6—0.8 mm
wide, conduplicate to involute, abaxially glabrous,
green, sometimes slightly curved. Panicles 15—17 x
0.5—1 cm, narrow, contracted, with few branches
and these only near base; branches sparsely
scabrous. Spikelets 8.5—10 mm long, lanceolate,
florets 3 or 4, probably cleistogamous; rachillas
0.5—0.7 mm long, glabrous or sparsely scabrous;
glumes 4—6.5 mm long, lanceolate, membranous,
glabrous, apex acute; lower glumes 44.5 mm
long, 1-nerved; upper glumes 5.5—6.5 mm long,
3-nerved; lemmas, 7—7.5 mm long, lanceolate,
5-nerved, membranous, glabrous, awned, the awn
4—6 mm long; paleas almost as long as the lemma,
membranous; anthers 0.8—0.9 mm long; ovary apex
glabrous. Caryopses lanceolate; hilum 2/5 as long
as the grain, linear.

Leaf blade anatomy.—Cross-sections with
5(—7) vascular bundles, 3—5 ribs above; scleren-
chyma under abaxial epidermis discontinuous,
forming small fascicles, adaxial sclerenchyma
absent or exceptionally forming small fascicles;
adaxial epidermis with short scattered hairs.

Distribution and habitat.—Festuca huamachu-
censis is endemic to northern Peru where it has been
found growing on wet alpine grasslands (jalca).

Additional specimens examined. PERU.
Ancash: Prov. Recuay, Huascaran Parque Nacio-
nal, quebrada Queshque, lateral valley toward Rio
Pachacoto, 09°50S’, 77°18’ W, 4500-4600 m, 18
Mar 1986, Smith et al. 1185] (MO). Cajamarca:
Prov. Cajamarca, 52 km N of Cajamarca, on Hwy
3N towards Bambamarca, and ca. 1 km W ona
small road “Carretera Lagunas Yanacanchilla”,
3800-3900 m, 16 Mar 2000, PM. Peterson14901
& N. Refulio Rodriguez (MO, US); Laguna Maqui—
Maqui, pajonal de jalca, 3910 m, 4 Oct 1994,
Vega et al. 6955 (F); Cerro Piedras Gachas, entre
Yanacocha y Llaucan, 3900 m, 23 Apr 1994, Vega
et al. 6994 (F); cima de cerro con Piedras grandes,
3900 m, 24 Apr 1994, Vega et al. 7041 (F); cima
del Cerro Maqui—Maqui, ladera arenosa, 4150 m,
20 Mar 1994, Vega et al. 6914 (F); Pampa Larga,
al N de la explotacion minera Yanacocha, ladera de
jalca graminosa, 3900 m, 14 May 1994, Vega 7124
(F); Jalea de Cumullca, lado occidental, 3600 m,
31 May 1984, Vega et al. 3494 (F); ruta a Guagal,
km 10, 3600 m, 27 May 1977, Vega et al. 2018
(F); Ladera de jalca, 3900 m, 14 May 1994, Vega
et al. 7128 (F); Prov. Celendin, desvio Huanico,

114

Festuca in South American Paramos

Figure 56. Festuca huamachucensis. A. Habit. B. Ligule. C. Spikelet. D. Glumes. E. Lemma. F. Lemma with palea and rachilla.

G. Leaf blade cross-section. A-G, Vega 6994 (F).

sobre la ruta Cajamarca—Celendin, 3500 m, 4 Oct
1987, Vega 4378 (F); 07°02°31"S, 78°15°33’W,
3700 m, 26 Aug 2004, Sklenar & Zapata Cruz 8705
(PRC); Prov. Hualgayoc. Desvio de la carretera
Coymolache—Chugur, 3750 m, 30 Apr 1994, Vega

7111 (F). La Libertad: Prov. Santiago de Chuco,
42 km S of Huamachuco on road towards Pampas,
4000 m, 29 Mar 1997, PM. Peterson 13951 &
N. Refulio Rodriguez (K, MO, US, USM); Prov.
Carrion, al pie del Nevado de Huaylillas, jalca,

Festuca in South American Paramos 115

07°52°S, 78°01 °W, 3840 m, 21 May 2001, Zapata
16506, 16508 (F).

45. Festuca imbaburensis Stancik, Folia Geobot.
Phytotax. 39(1): 100, f. 3, 1-5. 2004. (Figs.
57, 58, 94A & B). Tyre: Ecuador. Imbabura,
Mun. Urcuqui, road to Cerro Yanaurcu,
00°28'29’N, 78°20'04’W, grass paramo with
swampy patches, 4150 m, 15 Oct 2000, D.
Stancik 4098 (holotype: PRC!; isotypes:
AAU!, QCA!).

Tussocked perennials with intravaginal inno-
vations. Culms (15—)20—50 cm tall, erect, finely
scabrous; nodes 1, basal. Leaf sheaths membra-
nous to coriaceous, stramineous, glabrous; ligules
0.5—0.7 mm long, membranous, apex truncate,
short-ciliate; blades 9-20 cm long, 0.5—0.6 mm
wide, conduplicate to involute, somewhat rigid,
green, glaucous, abaxially glabrous, apex obtuse.

Panicles 6—12 x ca. 0.5 cm, contracted, with few
scabrous branches, each with a few spikelets.
Spikelets 8—10 mm long, lanceolate, florets 2 or 3;
rachilla glabrous; glumes 34.2 mm long, keeled
on back, membranous, dark purple, glabrous or
upper 1/4 sparsely papillose, apex obtuse; lower
glumes 3—3.5 mm long, oblong, 1-nerved; upper
glumes 44.2 mm long, ovate, 3-nerved; lem-
mas 5.5—6(—7) mm long, lanceolate, 5-nerved,
membranous to coriaceous, dark purple, upper
1/4 scabrous, awned, the awn 0.5—0.7 mm long;
callus glabrous; paleas shorter than the lemma,
lanceolate, membranous, upper 1/4 and along keels
densely scabrous to pilose; lodicules ovate, mark-
edly bilobed; anthers 1—1.5 mm long; ovary apex
glabrous. Caryopses not observed.

Leaf blade anatomy.—Cross-sections with
5—7 vascular bundles and 3-5 ribs above; scle-
renchyma under abaxial epidermis discontinuous,
in small fascicles, adaxial sclerenchyma absent;

COLOMBIA

N

0 300 600 Kilometers

Figure 57. Distribution of Festuca huamachucensis (), F: imbaburensis ( &), F: nereidaensis (m), and F. sanctae-martae (@).

116 Festuca in South American Paramos

Figure 58. Festuca imbaburensis. A. Habit. B. Ligule. C. Spikelet. D. Glumes. E. Lemma. F. Lemma with palea and rachilla. G.
Leaf blade cross-section. A—G, Stancik 3899 (PRC).

Festuca in South American Paramos

adaxial epidermis with scattered hairs, the hairs
0.09 mm long.

Observations.—Festuca imbaburensis is
morphologically similar to F’ glumosa but differs
by having glabrous leaf blades (versus scabrous),
narrower blades (0.5—0.6 versus 0.8—1.4 mm),
shorter ligules (0.5—0.7 versus 0.8—1.2 mm), gla-
brous rachillas (versus pilose), and shorter lemmas
(5.5—6(—7) versus 7-8 mm).

Distribution and habitat.— Festuca imbabu-
rensis ranges from central Ecuador to southern
Colombia. It occurs in swampy patches associated
with grass paramos and superparamos between
3400-4200 m.

Additional specimens examined. COLOMBIA.
Narino: Mun. Cumbal, Volcan Nevado de Cumbal,
N-NE slopes, 3700 m, 9 Mar 1999, D. Stancik 2749
(COL, PRC, US); Mun. Tuquerres, Volcan Azufral,
road from vereda San Roque alto to Laguna Verde,
km 4, 3700 m, 9 Mar 1999, D. Stancik 2783 (COL,
PRC); road from vereda San Roque alto to Laguna
Verde, km 5, 3800 m, 9 Mar 1999, D. Stancik
2786 (AAU, COL, PRC). ECUADOR. Azuay:
Parque Nacional Cajas, trail from Molleturo to
Paraguillas, 02°49’S, 79°15’W, 4200-4300 m,
S. Laegaard 102666 (AAU); Mun. Cuenca, Parque
Nacional Cajas, Cerro Amarrillo, 2°45’S, 79°15’ W,
4300-4400 m, 13 Jul 1997, D. Stanctk 2056 (PRC,
QCA); Cajas Parque Nacional, E flanks and summit
area of Cerro Amarillo, 4451m, lower superparamo
with Loricaria sp., Azorella sp., Werneria sp.,
79°15'°W, 02°45’S, Sklenar 2596 (PRC, QCA).
Bolivar: road Los Arenales—Salinas, km 4, 01°24’S,
78°55 W, 4300 m, S. Laegaard 55368 (AAU). Car-
chi: Paramo del Angel, sector Voladero, 00°38’N,
67°53’ W, 3400-3800 m, Davalos 18 (US); Volcan
Los Chiles, 00°49°N, 77°57’ W, 3950-3980 m,
S. Laegaard 101692 (AAU, QCA); Mun. Tulcan,
volcan Chiles, SW slope, 00°48°N, 77°57’'W,
4055m, 14 Jul 1999, D. Stancik 3228 (AAU, PRC,
QCA, US, W); D. Stancik 3235, 3267 (AAU, PRC,
QCA). Chimborazo: E] Altar, N side of Volcan,
4200-4400 m, Sklenar & Kosteckova 956 (AAU);
4200 m, Sklenar & Kosteckova 1929 (AAU);
Sklenar & Kosteckova 1031 (US); Collanes val-
ley, Paramo de Los Altares, 4000 m, Ramsay &
Smith 343 (QCA, QCNE); Alao—Paramo de Laguna
Negra, 4100 m, Barclay & Juajibioy 8763 (MO,
US); E side of Volcan Chimborazo, 4300 m, Sklenar
& Kosteckova 2168 (AAU); km 8.5 E of Guardiana
Alao above Rio Alao, 3350-3550 m, PM. Peterson

bry

9197, E.J. Judziewicz, R.M. King & P.M. Jorgensen
(MO, US); E of Licto, 4200 m, 30 Oct 1987,
Ramsay 1035 (K, QCA, QCNE). Cotopaxi: road
San Miguel (Salcedo)—Puerto Nuevo, 29 km from
San Miguel, 3950-4050 m, B. Ollgaard & Balslev
9921 (AAU, F, MO, NY, S); Paramo de Cotopaxi,
T: de Vries s.n. (AAU); Parque Nacional Cotopaxi,
4220 m, 21 Jun 1999, D. Stancik 3098 (AAU, PRC,
QCA); Mun. Lasso, Parque Nacional Cotopaxi,
NE side of volcano, 00° 39.3’S, 78°25’ W, 4220 m,
26 Jun 1999, D. Stancik 3101 (AAU, PRC, QCA).
Imbabura: Paramo de Mariano Acosta, km 25 on
the road Yahuarcocha—Mariano Acosta, 00°20’S,
78°00°W, 3650-3750 m, S. Laegaard 101194
(AAU); Mun. Urcuqui, road to Cerro Yanaurcu,
S slope, 00°28°56"N, 78°20'28.5"W, 4300 m, 15
Oct 2000, D. Stancik 4081 (AAU, PRC, QCA);
D. Stancik 4090 (AAU, PRC, QCA, US); D. Stancik
4091 (AAU, PRC, QCA); D. Stancik 4092B, 4098
(PRC, QCA). Pichincha: road Pifo—Pintag, in
valley 2.5 hours horseride above Inga Monserat,
00°19°S, 78°17°W, 3950 m, S. Laegaard 102262
(AAU); along road El Chaup—Pastocalle, 00°41’S,
78°39 W, 3200-3700 m, S. Laegaard 54548 (MO);
road Quito—Paramo Guamani, 00°17’S, 78°09’ W,
B. Allgaard & Balslev 10073 (AAU, F, NY, MO,
S); antenna N of pass, 00°17°S, 78°09’ W, 4260xm,
B. Ollgaard & Balslev 10100 (AAU, NY); 00°17'S,
78°11 W, 4250-4400 m, S. Laegaard et al. 53859
(AAU, QCNE); 4050 m, S. Laegaard 101373
(AAU, QCA); peak ca. 6 km S of Paso de la Vir-
gen, 00°21’°S, 78°13’ W, 4200-4250 m, S. Laegaard
55698 (AAU); 4100 m, S. Laegaard 103105 (QCA);
3900 m, Ramsay et al. 153 (QCA, QCNE); N side
of Volcan Antisana, ca. km 12 along road to Haci-
enda Antisana, 00°27’S, 78°10’ W, 4400-4500 m,
S. Laegaard 102871 (AAU); Mun. Pifo, Paramo
de Guamani, 00° 20’S, 78°12’ W, 4000 m, 19 Jun
1999, D. Stancitk 3054B (PRC, QCA); D. Stancik
3054A (AAU, PRC, QCA); Superparamo vegeta-
tion on the SW side of Cotacachi, 0°21°N, 78°21 °W,
4300-4400 m, 12 Aug 2004, Sklenar 8204 (PRC).
Tungurahua: along trail from Mesa Tablon to
Limpiopungu, S of Laguna Pisayambo, 01°07’S,
78°21'°W, 4000-4050 m, S. Laegaard 19452
(AAU); Llanganatis, by Lake Aucacocha, 3700 m,
Edwards s.n. (P); Mun. Pillaro, Las Llanganatis,
01°07°55.5” S, 78°20°40"W, 3900 m, 28 Sep 2000,
D. Stancik 3898 (PRC, QCA); D. Stancik 3899
(AAU, PRC, QCA); 01°09°37.5’S, 78°14°51°W,
3500 m, D. Stancik 3990 (PRC, QCA).

Festuca in South American Paramos

118

ee ee ee ee ee ee ee ee ee ee ee. Pe _

is. A. Stylized growth form. B. Habit. C. Ligule. D. Spikelet. E. Glumes. F. Lemma. G. Lemma with
s-section. A-H, Stancik & Galvis2026 (PRC).

monguens
palea and rachilla. H. Leaf blade cros

Figure 59. Festuca

Festuca in South American Paramos

46. Festuca monguensis Stancik, Darwiniana
41(1-4): 139, f. 8a—e. 2003. (Figs. 52, 59,
94C—F). Tyre: Colombia. Boyaca, Mun. Mon-
gui, paramo de la Laguna La Colorada, 3550
m, 21 Jan 1999, D. Stancik & Galvis 2026
(holotype: PRC!; isotype: COL!, FMB!).

Tussocked perennials with intravaginal innova-
tions. Culms ca. 50 cm tall, erect, scabrous; nodes
2 on upper 1/2. Leaf sheaths coriaceous, stramine-
ous, finely striate, glabrous; auricles absent; ligules
0.7—-1(—1.5) mm long, coriaceous, apex truncate,
short-ciliate; blades 20-25 cm long, 0.6—0.8 mm
wide, conduplicate to involute, abaxially scabrous,
green, apex obtuse. Panicles 5—15 x 0.5—0.8 cm,
contracted, lanceolate, with few branches; branches
scabrous. Spikelets 9-11 mm long, lanceolate,
florets 3 or 4; rachilla glabrous or with short,
scattered hairs; glumes 4—5 mm long, lanceolate,
coriaceous, purplish-green, scabrous near apex,
margins membranous, apex acute; lower glumes
ca. 4 mm, |-nerved; upper glumes ca. 5 mm long,
3-nerved; lemmas 6—7 mm long, lanceolate,
5-nerved, membranous, purplish-green, papillose,
scabrous near apex, awned, the awn 0.5—2.5 mm
long; callus glabrous; paleas as long as the lemma,
lanceolate, membranous, papillose, hairy along
keel and near apex; lodicules 1.3 mm long, oblong,
two-dentate; anthers 1.2—1.3 mm long; ovary apex
glabrous. Caryopses not seen.

Leaf blade anatomy.—Cross-sections usually
with 7 vascular bundles, 5(—7) ribs above; scle-
renchyma under abaxial epidermis discontinuous
sometime extending to the vascular bundles, scle-
renchyma absent adaxially; adaxial epidermis with
scattered hairs, the hairs ca. 0.09 mm long.

Observations.—Festuca monguensis differs
from other species of F: sect. Festuca by having
short panicles on distinctive long culms.

Distribution and habitat.— Festuca monguen-
sis is endemic to Colombia and is known only
from type locality in Cordillera Oriental (Boyaca).
It is known from grass paramo dominated with
Calamagrostis effusa (Kunth) Steud. between
3400-3600 m.

Additional specimens examined. COLOMBIA.
Boyaca: Mun. Mongui, paramo de la Laguna La
Colorada, 3550 m, 21 Jan 1999, D. Stancik & Galvis
2025 (COL, PRC); 3450 m, 21 Jan 1999, D. Stancik
& Galvis 2028 (COL, FMB, PRC).

119

47. Festuca nereidaensis Stancik, Darwiniana
41(1-4): 139, f. 8fk. 2003. (Figs. 57, 60,
95A-D). Type: Colombia. Caldas, Mun.
Manizales, Parque Nacional Los Nevados, road
from Casa del Cisne to Rio Nereidas, km 5,
3800-4000 m, 18 Dec 1999, D. Stancik 3401B
(holotype: PRC!; isotype: COL!).

Tussocked perennials with intravaginal innova-
tions. Culms 80—100 cm tall, erect, finely scabrous
just below panicle; nodes 2 or 3 on upper 1/2. Leaf
sheaths membranous to coriaceous, stramineous,
glabrous; auricles absent; ligules 0.7—-1 mm long,
membranous, apex truncate, ciliate; blades 25-35 cm
long, 1.2—1.5 mm wide, conduplicate, abaxially gla-
brous, green, apex acute. Panicles 15—20 x 1—1.5 cm,
contracted, lanceolate, branches finely scabrous.
Spikelets 11-12 mm long, lanceolate, florets 5;
rachilla short, sparsely short-hairy; glumes 6-8 mm
long, lanceolate, coriaceous, margins membranous,
dark, glabrous, apex acute; lower glumes 6-6.5 mm
long, 1-nerved; upper glumes 7—8 mm long, 3-nerved;
lemmas 7—7.5 mm long, lanceolate, 5-nerved, coria-
ceous, finely papillose, awned, apex pointed, the awn
0.5—0.7 mm long; callus glabrous; paleas almost as
long as the lemma, lanceolate, membranous, papil-
lose, upper 1/2 scabrous, apex hairy; lodicules 1.3 mm
long, lanceolate, two-dentate; anthers 1.3—1.6 mm
long; ovary apex glabrous. Caryopses not seen.

Leaf blade anatomy.—Cross-sections with
10-12 vascular bundles and 9 ribs above; scleren-
chyma under abaxial epidermis almost continu-
ous, sometimes extending to the vascular bundles,
adaxial sclerenchyma present but not extending to
the vascular bundles; adaxial epidermis hairy, the
hairs ca. 0.45 mm long.

Distribution and habitat.—Festuca nereidae-
nsis is endemic to Colombia and is known only from
the type locality of Nevado del Ruiz in Cordillera
Central. This species occurs in grass paramo zones
between 3800-4000 m.

48. Festuca oroana Stancik, Folia Geobot. Phy-
totax. 39(1): 104, f. 2, 6-10. 2004. (Figs. 61,
64, 95E & F). Type: Ecuador. El Oro, along
mule-track Tambillo-Manu, mountain forest,
grazed and/or burned, dense tussock, 03°30’S,
79°32’ W, 3200-3320 m, 21 Feb 1988, 21 Feb
1988, S. Laegaard 70309 (holotype: AAU!;
isotypes: MO!, PRC!, QCA!, QCNE!).

Festuca in South American Paramos

120

Figure 60. Festuca nereidaensis. A. Stylized growth form. B. Habit. C. Ligule. D. Spikelet. E. Glumes. F. Lemma. G. Lemma with

palea and rachilla. H. Leaf blade cross-section. A-H, Stancik 3401B (PRC).

Festuca in South American Paramos 121

<= :

—S

Lig,

Z,

EGA

AA Fee,

SS
— 2

SS
v >

Figure 61. Festuca oroana. A. Stylized growth form. B. Habit. C. Ligule. D. Spikelet. E. Glumes. F. Lemma. G. Lemma with palea
and rachilla. H. Leaf blade cross-section. A-H, Laegaard 70309 (PRC).

122

Tussocked perennials with intravaginal innova-
tions. Culms ca. 100 cm tall, erect, scabrous; nodes
2(-3) on lower 1/2. Leaf sheaths membranous to
coriaceous, lightly brownish-gray, finely scabrous,
margins free; auricles absent; ligules 0.5—1(—1.5)
mm long, membranous, apex truncate, short-cili-
ate; blades 35-50 cm long, 1.1—1.8(-4) mm wide,
conduplicate to flat, abaxially glabrous, green.
Panicles ca. 20 cm long, ca. 25 cm wide, pyramidal,
spreading, branches and pedicels densely minutely
scabrous. Spikelets 9-10 mm long, lanceolate
to oblong-lanceolate, florets 4 or 5(—6); rachilla
densely hairy, the hairs short; glumes 2.4—3.9
mm long, membranous to coriaceous, green, apex
acute; lower glumes 2.4—2.6 mm long, lanceolate,
l-nerved, glabrous; upper glumes 3.6—3.9 mm
long, oblong to oblong-lanceolate, 3-nerved; lem-
mas 5.5—6(—6.5) mm long, lanceolate, 5-nerved,
membranous, greenish, papillose or upper 1/3 finely
scabrous, awned, the awn 0.5—1 mm long; callus
hairy; paleas as long as the lemma, lanceolate,
membranous, margins short-hairy, apex glabrous;
lodicules oblong-lanceolate; anthers 2.7—2.8 mm
long; ovary apex glabrous. Caryopses not seen.

Leaf blade anatomy.—Cross-sections with
about 17 vascular bundles and 9 ribs above; scle-
renchyma under abaxial epidermis discontinuous,
adaxial sclerenchyma extending to the vascular
bundles forming girders; adaxial epidermis densely
hairy, the hairs, 0.2—0.4 mm long.

Observations.—In South America, there are
no species morphologically similar to F. oroana.
Perhaps F: fiebrigii is superficially similar but differs
by having cylindrical panicles (versus pyramidal in
F- oroana), longer glumes and lemmas (lower glume
ca. 4 versus 2.4-2.6 mm; upper glumes 5—5.5 versus
3.6—3.9 mm; lemmas 6—7 versus 5.5—6 mm).

Distribution and habitat.—Festuca oroana is
known only from type locality in Southern Ecuador.
It occurs in Andean mountain forests at an altitude
of about 3200 m.

49. Festuca parciflora Swallen, Contr. U.S.
Natl. Herb. 29(6): 255. 1949. (Figs. 62, 64).

Festuca in South American Paramos

Type: Ecuador. Azuay, paramo, in vicinity of
Toreador, between Molleturo and Quinoas,
3810-3930 m, 15 Jun 1943, J. A. Steyermark
53092 (holotype: US-1911635!).

Tussocked, loosely caespitose or short-
rhizomatous perennials with intravaginal innova-
tions. Culms 28-60 cm tall, erect, scabrous or
glabrous; nodes 1, basal. Leaf sheaths membranous
to coriaceous, brownish-gray, glabrous, inconspicu-
ously striate; ligules 0.8—1 mm long, membranous
to coriaceous, apex truncate, short-ciliate; blades
13—23 cm long, 0.4—0.6 mm wide, conduplicate to
involute, slightly abaxially scabrous, glaucous, apex
obtuse. Panicles 3-15 cm long, ca. 0.5 cm wide,
contracted, narrow with few erect branches; branches
scabrous on ribs. Spikelets 8.5—9 mm long, lanceo-
late, florets 2 or 3(—4); rachilla with scattered hairs;
glumes 3—5 mm long, keeled 1n upper part, membra-
nous, purplish-white, upper 1/4 sparsely tuberculate;
lower glumes 3—3.5 mm long, lanceolate, 1-nerved;
upper glumes 4—5 mm long, ovate, 3-nerved; lemmas
6—7 mm long, lanceolate, 5-nerved, membranous,
keeled, purplish-green, upper 1/3 scabrous, awned,
the awn 0.5—1.5 mm long; callus with scattered hairs;
paleas as long as the lemma, lanceolate, membranous,
upper part and along keels densely scabrous to pilose;
lodicules lanceolate; anthers 0.8—1.1(—1.4) mm long;
ovary apex glabrous. Caryopses lanceolate; hilum 4/5
as long as the grain, linear.

Leaf blade anatomy.—Cross-sections with 5
vascular bundles and 3 ribs above; sclerenchyma
discontinuous under abaxial epidermis, adaxial
sclerenchyma absent; adaxial epidermis with scat-
tered hairs, the hairs 0.09 mm long.

Observations.—This species is morpho-
logically similar to F’ glumosa, F- imbaburensis, F-
carchiense, and F: sumapana but differs by having
slender, scabrous leaf blades, smaller spikelets, and
smaller floral parts.

Distribution and habitat.—Festuca parciflora
is endemic to southern Ecuador (Azuay, Loja) and
is known from the grass paramo zone between
3600-4200 m.

KEY TO THE SUBSPECIES OF FESTUCA PARCIFLORA

la. Culms 40-50 cm tall, scabrous; plants tussocked; panicles 7—11(—15) cm tall ...........ceeeceeeeeeeeneeees

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1b. Culms ca. 25 cm tall, glabrous; plants short-rhizomatous to loosely caespitose; panicles 3—7 cm

5d cane eee teen 49b. F. parciflora subsp. loxana

123

Festuca in South American Paramos

d

a. F. Lemma with palea an

. A. Habit. B. Ligule. C. Spikelet. D. Glumes. E. Lemm

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124

49a. Festuca parciflora subsp. parciflora (Fig.
62).

Tussocked perennials. Culms 40—50 cm tall,
scabrous; panicles 7—11(—15) cm tall.

Additional specimens examined. ECUA-
DOR. Azuay: Parque Nacional Cajas, N side of
Laguna Luspa, 02°48’S, 79°15’W, 3850 m, | Sep
2000, D. Stanéik 3845 (AAU, PRC); D. Stancik
3846 (AAU, PRC, US, W); Mun. Cuenca, Parque
Nacional Cajas, between El Refugio and Lagoon
Taglacocha—Tres Cruces, 02°47°S, 79°13 W, 3900—
4000 m, | Sep 2000, D. Stanctk 3859 (AAU, PRC);
N side of Lagoon Taglacocha, 4100—4200 m, | Sep
2000, D. Stancik 3873 (AAU, PRC, US).

49b. Festuca parciflora subsp. loxana Stancik,
Folia Geobot. Phytotax. 39(1): 107. 2004.
Type: Ecuador. Loja, Cerro de Arcos W of road
Manu—Zaruma, 03°34'S, 79°28’ W, 3250-3600
m, 14 Sep 1999, S. Laegaard & Aguirre 20608
(holotype: AAU!; isotypes: LOJA!, QCA!).

Rhizomatous or loosely cespitose perennials.
Culms ca. 25 cm tall, glabrous. Panicles 3—7 cm
tall.

Additional specimens examined. ECUADOR.
Loja: Surroundings of Laguna Chuquiraga E of
Amaluza, 04°37°S, 79°22’ W, 3300 m, S. Laegaard
et al. 19285 (AAU).

50. Festuca renvoizei Stancik, Novon 17(1):
100-104. 2007. (Figs. 63, 64). Typr: Peru. Caja-
marca, Las Lagunas, 50 km from Cajamarca on
road to Bambamarca, heavily grazed and bur-
ned grass paramo with crystalline rock outcrops
and numerous lagoons of various size, growing
in very wet area beside lagoon, 4000 m, 29
Mar 1988, S.A. Renvoize & S. Laegaard 5031
(holotype: AAU!; isotypes: CPUN!, K!).

Tussocked perennials with intravaginal innova-
tions. Culms 40-50 cm tall, erect, scabrous; nodes
1, basal. Leaf sheaths membranous, brownish-gray,
abaxially glabrous, margins free; auricles absent;
ligules 1.2-1.5 mm long, membranous, short-
ciliate; blades 20-30 cm long, 0.5—0.7 mm wide,
conduplicate to involute, mostly glabrous, green,
apex obtuse. Panicles 11-17 cm long, 0.5—0.7
cm wide, contracted, slender; branches scabrous
throughout. Spikelets 9-10.5 mm long, lanceolate,
florets 2 or 3; rachilla glabrous; glumes 3.5—5.5 mm

Festuca in South American Paramos

long, membranous, dark purple, mostly glabrous,
apex acute; lower glumes 3.5—4.5 mm long, lan-
ceolate, 1-nerved; upper glumes 5—5.5 mm long,
oblong-lanceolate, 3-nerved; lemmas 7—7.5 mm
long, lanceolate, 5-nerved, membranous, dark
purple, scabrous distally, awned, the awn 0.5—1
mm long; callus glabrous; paleas 4/5 as long as the
lemma, membranous, upper 1/3 and along keels
hairy; lodicules ca. 0.8 mm long, lanceolate; anthers
1—-1.2 mm long; ovary apex glabrous. Caryopses
not observed.

Leaf blade anatomy.—Cross-sections with
5—8 vascular bundles, 5 ribs above; sclerenchyma
under abaxial epidermis discontinuous and small,
adaxially absent; adaxial epidermis with scattered
hairs, the hairs ca. 0.1 mm long.

Distribution and habitat.—Festuca renvoizei
is known only from Department Cajamarca in
northern Peru from grass paramos in very wet areas
near lagoons at 4000 m.

Additional specimens examined. Peru. Caja-
marca: Parte baja del Cerro Shillas Negras y
Laguna Totora, 3920 m, 23 Apr 1994, Vega et al.
7011 (F, CPUN).

51. Festuca sanctae-martae Stancik, Preslia
75(4): 343-345, f. 2. 2004. (Figs. 57, 65,
96A-—D). Tyre: Colombia. Magdalena, Sierra
Nevada de Santa Marta, SE slope, Hoya del
Rio Donachui, Laguna de Calocribe (E of
Meollaca), paramo, 3600—3700 m, 30 Sep
1959, J. Cuatrecasas & Castaneda 24532
(holotype: COL!; isotype: US!).

Tussocked perennials with intravaginal inno-
vations. Culms 60-80 cm tall, erect, glabrous or
finely scabrous under panicle; nodes 1, basal and
1 or 2 shorter leaves. Leaf sheaths membranous to
coriaceous, stramineous, glabrous, finely striate;
auricles absent; ligules 1—1.6 mm long, membra-
nous to coriaceous, apex truncate; blades 25-35 cm
long, (0.8-)2—3.5 mm wide, conduplicate or flat,
abaxially glabrous. Panicles 10-16 = 1.5—2 cm,
contracted, elongate. Spikelets 10-13 mm long,
lanceolate, florets 2 or 3; rachilla pubescent; glumes
8.5-10.5 mm long, almost as long as the spikelet,
lanceolate, membranous, white to purplish-white,
scabrous along midnerve, apex acute; lower
glumes 8.5—9.5 mm long, 1-nerved; upper glumes
8.5-10.5 m long, 3-nerved; lemmas 9-10 mm
long, lanceolate, 5-nerved, membranous, awnless
or short-awned, the awn 0.5—1 mm long; callus

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wil

Figure 63. Festuca renvoizei. A. Stylized growth form. B. Habit. C. Ligule. D. Spikelet. E. Glumes. F. Lemma. G. Lemma with
palea and rachilla. H. Leaf blade cross-section. A-H, Renvoize & Laegaard 5031 (AAU)

126

-75

Festuca in South American Paramos

COLOMBIA

-80

-75

N

0 100 200 Kilometers
—
Ga ee er rT!

-70

Figure 64. Distribution of Festuca vaginalis (¢), F: parciflora ( &), F: oroana (m), and F- renvoizei (@).

glabrous or sparsely hairy; paleas 3/4 as long as
the lemma, papillose, upper 1/3 scabrous; lodicules
1—1.4 mm long, lanceolate; anthers 0.8—1.1 mm
long; ovary apex glabrous. Caryopses lanceolate;
hilum 3/4 as long as the grain, linear.

Leaf blade anatomy.—Cross-sections with
(8—)10—11 vascular bundles, with 5—7 ribs above;
sclerenchyma under both abaxial and adaxial epi-
dermis, discontinuous, sclerenchyma girders absent;
bulliform cells not observed; adaxial epidermis with
scattered hairs, the hairs ca. 0.02 mm long.

Observations.—Festuca sanctae-martae
is morphologically similar to F. glumosa and
F. cocuyana. Festuca sanctae-martae and F: cocuy-
ana differ from F: glumosa by having conduplicate
(versus involute) leaf blades with 8—11 (versus 5—7)
vascular bundles, lower glumes 7.5—9.5 mm long
(versus 4.5-6 mm), upper glumes 7.5—-10.5 mm
long (versus 6—6.5 mm), and lemmas 8—10 mm long

(versus 7-8 mm). Festuca cocuyana differs from
F. sanctae-martae by having shorter culms (20-50
versus 60—80 cm), shorter leaf blades (15—20 versus
25-35 cm), and shorter lower glumes (7.5—8 versus
8.5—9.5 mm).

Distribution and habitat.—Festuca sanctae-
martae is endemic to the Sierra Nevada de Santa
Marta in northern Colombia. This species is
known from the grass paramos dominated with
Calamagrostis effusa and shrubby matorral zones
between 3600-4300 m.

Additional specimens examined. COLOMBIA.
Magdalena: Sierra Nevada de Santa Marta, valley
descending SW from Picos Reina and Ojeda, around
Lagoons Naboba, Mamito, and Mamo, 4200-4300 m,
J. Cuatrecasas & Castaneda 24563 (COL, US).

52. Festuca subulifolia Benth., Pl. Hartw. 262.
1846. (Figs. 66, 67, 96E—-F). Festuca tolucensis

127

Festuca in South American Paramos

-- 0 == =u

Figure 65. Festuca sanctae-martae. A. Stylized growth form. B. Habit. C. Ligule. D. Spikelet. E. Glumes. F. Lemma. G. Lemma

with palea and rachilla. H. Leaf blade cross-section. A-H, Cuatrecasas & Castaneda 24532 (COL).

128 Festuca in South American Paramos

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Figure 66. Festuca subulifolia. A. Stylized growth form. B. Habit. C. Inflorescence with proliferating spikelets. D. Ligule. E.
Spikelet. F. Glumes. G. Lemma. H. Lemma with palea and rachilla. I. Leaf blade cross-section. A-B, D-I, Stancik 3013 (PRC);
C, Stanctk 3336 (PRC).

Festuca in South American Paramos 129

-85 -80 -75 -70 -65 -60

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Figure 67. Distribution of Festuca tolucensis (@), F- subulifolia (&), and F’ sumapana (@).

var. subulifolia (Benth.) St.-Yves, Candollea
2: 302, 316. 1925. Type: Ecuador. Pichincha,
Hacienda de Antisana, Hartweg 1455 (lecto-
type: K!, designated by Alexeev, Novosti Sist.
Vyss. Rast. 23: 18. 1986; isolectotypes: B!, NY
ex Kil Pl. W?!).

Festuca breviaristata Pilg., Bot. Jahrb. Syst. 25(5):
717-718. 1898. Type: Ecuador. Pichincha,
Mount Puntas, 4400 m, Oct-Nov 1971, A. Stu-
bel 207 (holotype: B!; isotypes: US-2875385
fragm. ex B!, S!).

Festuca cajamarcae Pilg., Bot. Jahrb. Syst. 37:
513. 1906. Festuca distichovaginata var.
cajamarcae (Pilg.) St.-Yves, Candollea 3 :222.
1927. Type: Peru. Cajamarca, Pas Coymolache,
4000-4100 m, May 1904, A. Weberbauer 3974
(holotype: B!; isotypes: BAA-1171 fragm.!,
US-2875389!).

Festuca ferreyrae Tovar, Publ. Mus. Hist. Nat.
Javier Prado, Ser. B, Bot. 32: 8. 1984. Type:

Peru. Amazonas, Prov. Chachapoyas, Lei-
mebamba, en pajonal, 17 Apr 1964, R. Fer-
reyra 15512 (holotype: US-2565588!).

Dense tussocks (25—)50—80 cm tall; innova-
tions intravaginal; culms erect, finely scabrid or
glabrous, with one basal node and 2 leaves. Sheaths
coriaceous, firm, stramineous (or greyish), glabrous
(or finely scabrid); ligule coriaceous-membranous,
two-lobed, shortly ciliate, (0.5—)1—2.5 mm long; leaf
blades linear, conduplicate, pungent, 10-60 cm long,
0.4—0.8 mm in diameter, rough, green to glaucous.
Panicle usually contracted, dense, lanceolate to
oblong, some times spreading, ovate, (6—)12—35
long, 1—2(—10) cm wide; branches finely scabrid.
Spikelets 6.5-9 mm long, ovate, with 3-5 florets;
rachilla covered (densely) by hairs; glumes coria-
ceous-membranous, acute, green or purplish-green,
upper third (or all) scabrous, nerves prominent;
lower glume lanceolate, (3—)3.5—4.5(—5.5) mm long,

130

1-nerved; upper glume oblong, 4—5.5(—6) mm long,
3-nerved; lemmas 5—6.5(—8) mm long, 5-nerved,
coriaceous-membranous (on margins), oblong-
lanceolate, purplish-green, densely scabrid to rough
on all surfaces, shortly two-dentate, with awn
0.5—2.5 mm long; callus glabrous; palea membra-
nous, lanceolate, two-carinate, at least upper third
and along keels scabrid to hairy, two-dentate, as
long as the lemma (or longer); lodicules obovate,
two-dentate, about 1.1 mm long; stamens 3, anthers
(2.1—)2.4—2.8(3.2) mm; ovary apex glabrous. Cary-
opsis oblong-lanceolate; hilum linear, 4/5 of total.
Leaf blade cross section typically with 7 vascular
bundles and 5 ribs above; sclerenchyma under
abaxial epidermis continuos with all or almost all
vascular bundles, adaxial sclerenchyma present at
3 vascular bundles and forming girders. (about 30%
of studied specimens with 9-13 vascular bundles
forming 7—11 ribs and with 3-6 girders); abaxial
epidermis densely covered by prickles, hairs more or
less dense on adaxial epidermis, 0.1—0.2 mm long.

Leaf blade anatomy.—Cross-sections typ1-
cally with 7 vascular bundles and 5 ribs above;
sclerenchyma under abaxial epidermis continuous
extending to (almost) all vascular bundles; adaxial
sclerenchyma present and extending to 3 vascular
bundles forming girders, ca. 30% of studied speci-
mens with 9—13 vascular bundles forming 7—11 ribs
and with 3-6 girders); abaxial epidermis densely
covered by prickles, densely hairy on adaxial epi-
dermis, the hairs 0.1—0.2 mm long.

Observations.—Festuca subulifolia is mor-
phologically similar to F. tolucensis, however
F. subulifolia differs by having large girders in leaf
blade cross sections and shorter truncate ligules.
Festuca subulifolia is a highly variable species that
is often a dominant or co-dominant in the grass
paramo zone. The variation in form and size of the
panicles and floral parts cannot be taxonomically
interpreted easily and it is probably a result of the
diversity of natural conditions and of paramo man-
agement, 1.e., burning and grazing. Characteristics,
such as two-dentate lemmas with a relatively long
awn and finely scabrous leaf blades, are fairly stable
with not much variation. After comparing the types
of Peruvian material of F’ cajamarcae and F. fer-
reyrae, with that of F| subulifolia, it was concluded
that these types represent slightly different forms of
the same species. Therefore the former two names
are here treated as synonyms.

Distribution and habitat.—Festuca subulifolia
is known from Peru, Ecuador, and Colombia. In

Festuca in South American Paramos

Colombia Festuca subulifolia is known only in
southern part of Cordillera Central (Narifio, Cauca).
It is often a dominant species of grass paramos
between 2900-3700 m. This species is known
from various plant communities such as: Agrostis
cf. haenkeana (Cuatrecasas 1934), Calamagrostio
effusae-Calamagrostietum macrophyllae (Duque &
Rangel 1989), Aciachne acicularis-Calamagrostis
intermedia (Verweij 1995), Ugno myricoidis-
Espeletietum hartwegianae (Rangel & Ariza 2000),
Espeletio hartwegianae-Calamagrostietum effusae
(Salamanca 1991), and Calamagrostis effusae-
Espeletia hartwegiana subsp. centroandina (Rangel
& Franco 1985).

Additional specimens examined. COLOMBIA.
Cauca: Mun. Popayan, Parque Nacional Purace,
Volcan Puracé, 02°20.07'N, 76°23.65'°W, 4200—
4300 m, 6 Jul 2000, D. Stancik 3609 (COL, PRC);
Pilimbala, 02°22.1’N, 76°24.06’W, 3350 m, 6 Jul
2000, D. Stancik 3608 (COL, PRC), 3450 m, 2 Apr
1939, Kjell von Sneidern 2144 (K, US); Macizo
Colombiano, Valle de las Papas, alrededores de
Valencia, 3500-3700 m, 15 Sep 1981, Idrobo et al.
3834 (COL, P, US); Hda. Los Andes, 3150 m, 17
Sep 1958, Barclay 5865 (COL); alrededores de la
Laguna de Cusiyaco, 3360 m, 7 Oct 1958, Barclay
& Juajibioy 5942 (COL, MO); 3360 m, 7 Oct 1958,
Barclay & Juajibioy 5945 (COL, MO, US); 12 Oct
1958, Barclay & Juajibioy 6052 (COL, MO);
Paramo del Puracé, E slope, around Laguna San
Rafael, 3320 m, 11 Oct 1961, J. Cuatrecasas &
Willard 26292 (COL, US); 3300 m, 27 Dec 1988,
Ortiz 1272 (COL); Laguna San Rafael, 3300 m, 6
Apr 1985, Wood 4806, 4807 (COL, K); Paramo El
Hinchadera, 3700 m, 24 Jul 1943, J. Cuatrecasas
14694 (US, VALLE); Cabeceras del Rio Paéz,
Laguna del Paéz, 3450 m, 4 Dec 1944, J. Cuatreca-
sas 19059 (US, MO, VALLE); Paramo de Moras,
9 Mar 1999, D. Stancik 2685 (COL, PRC); 3000—
3500 m, Feb 1906, Pittier 1409 (US); Mt. Pan de
Azucar, 3500-3700 m, 16 Jun 1922, F Pennell
7057 (US). Narino: El Encano, km 6 in direction
to village Colon, 2900 m, 13 Mar 1999, D. Stancik
2860 (COL, PRC, PSO); 13 Mar 1999, D. Stanctk
2856 (COL, PRC); Mun. Guachucal, vereda Que-
tanbu, Paramo Infiernillo, 3200 m, 9 Mar 1999,
D. Stancik 2613, 2618 (COL, PRC, PSO); 3100 m,
9 Mar 1999, D. Stanctk 2617, 2619 (COL, PRC,
PSO); D. Stancik 2631 (COL, PRC); Mun. Pasto,
paramo Puerto frio, between villages Las Almas
and Alisales, 2900 m, 14 Mar 1999, D. Stancik
2876 (COL, PRC, PSO); Mun. Pasto, Volcan de

Festuca in South American Paramos

Galeras, 3400 m, 16 Mar 1999, D. Stanctk 2911
meet PRC. PSO); 3200 m: 16 Mar 1999, D.
Stanctk 2909 (COL, PRC, PSO); 9 Mar 1999, D.
Stanctk 2659 (COL, PRC, PSO); 3600 m, 16 Mar
1999, D. Stanctk 2918 (COL, PRC, PSO); 2900 m,
16 Mar 1999, D. Stanctk 2910 (COL, PRC, PSO);
16 Mar 1999, D. Stanctk 2908 (COL, PRC, PSO);
Mun. Pasto, Morazurco, vereda San Francisco,
3100 m, 21 Mar 1999, D. Stancik 2962 (COL, PRC,
PSO); 21 Mar 1999, D. Stancik 2963 (COL, PRC,
PSO); Village Piedrancha, Paramo Infernillo, 3200
m, 9 Mar 1999, D. Stancik 2659 (COL, PRC, PSO);
9 Mar 1999, D. Stancik 2656, 2657 (COL, PRC,
Eee); 2 Stancik 2658 (COL, PRC); Mun.
Tuquerres, Volcan Azufral, vereda San Roque Alto,
3600 m, 9 Mar 1999, D. Stancik 2764 (COL, PRC,
PSO); D. Stancik 2766 (COL, PRC); 9 Mar 1999,
D. Stancik 2802 (COL, PRC, PSO); 9 Mar 1999,
D. Stancik 2789 (COL, PRC); E slope, 01°07°N,
77°40’ W, 3100-3300 m, 15 Jun 1995, J. Luteyn et
al. 12834 (COL, MO); 12 May 1989, J. Luteyn et
al. 12842 (COL); Paramo Bordoncillo, 3250 m, 5
Mar 1963, Espinal 1035 (COL, PSO, US); Chiles,
cerca de poblacion Chiles, 3200 m, 30 Oct 1955,
Fernandez-Pérez 2931 (COL); Paramo de Uillin-
sayaco, between Pasto y Sibundoy, 15 Jul 1957,
Barclay 4529 (COL, MO); Volcan Galeras, 2900
m, 30 Oct 1980, Benavides 2660 (PSO); 3500 m,
21 Mar 1986, Benavides 6554 (PSO); 3400 m, 25
Aug 1964, Mora-Osejo 3195 (PSO); 3600 m, 5 Feb
1965, Mora-Osejo 3503 (PSO); Cumbal, 4000 m,
19 Mar 1941, Kjell von Sneider 362 (US); 19 Mar
1941, Kjell von Sneider 435 (US); 3700 m, 24 Mar
1941, Kjell von Sneider 415 (US); Mun. Pasto,
Santuario Volcano Galeras, 2900-3600 m, 16 Mar
1999, D. Stanctk 2914, 2917 (COL, PRC); Mun.
Cumbal, Nevado del Cumbal, N-NE slope, from
vereda Las Vuertas, 3700 m, 5 Sep 1999, D. Stancik
2748 (COL, PRC). ECUADOR. Azuay: Paramo
SW of Cuenca, 02°55’S, 79°08’ W, 3500 m, 1 Sep
1984, S. Laegaard 52837 (AAU, K, QCA, QCNE);
Road Gualaceo—Sucua, 03°00’S, 78°40’ W, 3450 m,
23 Oct 1984, S. Laegaard 53213 (AAU, QCA);
along new road Cuenca—Saraguro, km 14 N of
sideroad to Nabon, 03°12’S, 79°02’ W, 3420 m,
S. Laegaard 102376 (AAU); 03°07’S, 79°19°W,
3250-3350 m, 22 Mar 1992, S. Laegaard 101856
(QCA, QCNE); Parque Nacional Cajas, road to
Molleturo, km 38.4, 02°46’S, 79°14’W, 4110-
4350 m, PM, Jorgensen et al. 2099 (MO, PRC);
P.M. Jorgensen 2074 (MO, PRC); Laguna Torrea-
dora, 02°46'50’S, 79°13'40°W, 4100-4150 m,

Li.

2 Aug 1999, Palice 6, 7, 8 (PRC); N side of Lagoon
Taglococha, 02°47’S, 79°15’W, 4100-4200 m,
1 Sep 2000, D. Stancik 3847, 3848, 3850, 3872
(PRC, QCA); slopes near Laguna Luspa and
Laguna Canutillos, 3700 m, 21 Apr 1990, PM.
Peterson 8865, C.R. Annable & M.E. Poston (K,
MO). Totorococha—Mazan Valley, 02°53’S,
79°10°W, 3900 m, Ramsay & Smith 510 (K, QCA,
QCNE), vicinity of Toreador, 3810-3930 m,
J. Steyermark 53146 (F, NY, US); J. Steyermark
53182 (US); Paramo de Tinajillas, ca. 10 km S of
Cumbe, 3300 m, Harling 11255 (GB); km 13.7 S of
Cumbe, 3470 m, PM. Peterson 8876, C.R. Annable
& M.E. Poston (US); Paramo de Patococha, entre
Gualaceo y Limon, 3450-3500 m, Barclay & Jua-
Jibioy 8641 (MO, US); 03°00’S, 78°38’ W, 3400 m,
S. Laegaard et al. 103067 (AAU, QCA, QCNE);
km 45.4 W of Indanza on road to Cuenca, 3300 m,
P.M. Peterson 8932, C.R. Annable & M.E. Poston
(K, MO, QCA, QCNE, US); between el Refugio y
Lagoon Taglacocha, 02°47’S, 79°13’W, 3900-
4000 m, | Sep 2000, D. Stancik 3862, 3863, 3864,
3865, (PRC, QCA); Lagoon Luspa, S side, 02°48’S,
79°15°W, 3850 m, 1 Sep 2000, D. Stancik 3837
(PRC, QCA); Mun. Nabon, road Loja—Cuenca,
Cerro Ungahipucera, 03°12’S, 79°02°W, 3300-
3450 m, 30 Aug 2000, D. Stancik 3804, 3805, 3811,
3815 (PRC, QCA); Bolivar: pass on road Gua-
randa—Riobamba, NE of road, 01°35’S, 78°50’°W,
4050-4150 m, 10 Jun 1990, S. Laegaard 71739,
71741 (AAU, QCNE); km 27, 01°38’S, 78°51 °W,
3900-4100 m, S. Laegaard 55001 (AAU, QCA);
highway Ambato—Guaranda km 55.4, 4050 m, P
M. Peterson 8983 & C.R. Annable (MO, QCA,
QCNE, US); km 60.4, 4090 m, PM. Peterson 8985
& C.R. Annable (K, MO, QCA, QCNE, US).
Caniar: at the antennas of Culebrillas, ca. 17 km
from Panamerican Hwy., 02°26'S, 78°57°W,
4000 m, 4 Feb 2000, S. Laegaard 21016 (AAU,
LOJA); road Canar—Chunchi, 02°24’S, 78°59'W,
3200 m, S. Laegaard 53834 (AAU, QCA, QCNE);
nudo de Cordillera Oriental and Occidental,
3240 m, Barclay & Juajibioy 8325 (COL, US); km
11.4 NW of Tambo, S facing slope, 3220 m, P.M.
Peterson 8846, C. R. Annable & M.E. Poston
(QCNE). Carchi: Paramo del Angel, sector Los
Voladores, 00°41'N, 77°53’°W, 3700-3750 m,
S. Laegaard 101264A (AAU, QCA); NE of El
Angel, 3700-3900 m, S. Laegaard 53125 (AAU,
QCA, QCNE); 3600 m, Harlington 4067 (S);
3400-3800 m, Davalos 21, 28 (US); 3200-3800 m,
S. Laegaard 55771 (AAU, QCA); 00°41°N,

132

78°54’ W, Holm-Nielsen 5427 (AAU, COL, F, MO,
S); 00°SO0'N, 77°55’W, 3800 m, Grignon 84189
(AAU); 3750-3850 m, Holm-Nielsen 5383 (F);
3800 m, Acosta-Solis 10538 (US); km 13 on road
Las Juntas—El Angel, 00°43 'N, 77°50°W, 3380 m,
S. Laegaard 101726B (AAU, QCA, QCNE);
3300 m, E. Asplund 7055 (S); 3900 m, E. Asplund
17022 (S); km 4 W of Tufifio, Monte Redondo
area, 3475m, J. Luteyn & Luteyn 5713 (AAU, MO,
NY, US); NE of road El Angel towards Tulcan,
3240 m, PM. Peterson 9122, E.J. Judziewicz &
R.M. King (K, MO, QCA, QCNE, US); 13 May
1990, PM. Peterson 9127, E.J. Judziewicz & R.M.
King (K, MO, QCA, QCNE, US); 3320 m, PM.
Peterson 9140, E.J. Judziewicz & R.M. King (K,
MO, QCA, US); Nudo de Boliche—Voladero,
3900 m, Penland & Summers &89 (F); Road
Tulcan—El Carmelo, 3300 m, Harling & Andersson
12538 (GB); Paramo del Angel, 30 Dec 1931,
Benoist 4636 (P); Mun. Tulcan, volcan Chiles,
around Laguna Verde, 00°48 °N, 77°56’ W, 4000 m,
14 Jul 1999, D. Stancitk 3240 (PRC, QCA). Chim-
borazo: Parque Nacional Sangay—comunidad
Alao—Llactapamba, 01°58’S, 78°28°W, 3500 m,
Ceron et al. 11813 (MO, QCNE); Altar, A peak—
Valley of Collanes, 13000 ft, Whymper 1638 (K);
Collanes valley, 01°40°S, 78°24’ W, 4200 m, Ram-
say & Smith 422 (K); Panamerica Highway
Riobamba—Ambato, 01°30’S, 78°42’°W, 3500 m,
S. Laegaard 55401 (AAU, QCA); W of pass
Alao—Huamboya, 01°48’S, 78°25°W, 3750-3800
m, S. Laegaard 55417 (AAU, QCA); E of Guar-
diana Alao, km 8.5 via Huamboya, 3350-3550 m,
P.M. Peterson 9189, E.J. Judziewicz, R.M. King &
P.M. Jorgensen (MO, US); Paramo de Chacheaco,
E de Chunchi, 3800 m, Barclay & Juajibioy 8264
(MO, US); Daldal Valley E of Licto, km 10,
01°48°S, 78°32°W, 3850 m, Ramsay et al. 1057
(QCA); Mt. Chimborazo, E slope, 4250 m, E. As-
plund 8402 (NY, QCA, S), 3650 m, E. Asplund
7880 (F, S); road to Pallatanga, km 10 E of Lago
Colta, 3725 m, PM. Peterson 9215, E.J. Judziewicz,
R.M. King & P.M. Jorgensen (QCA, QCNE, US);
along Whymper road, ca. 10 km S of Cruce de los
Arenales, 01°30’S, 78°52’ W, 4100 m, S. Laegaard
19148 (AAU, LOJA, QCNE); Chimborazo-—
Reserva faunistica, 01°30’S, 78°49°W, 4150 m,
Ceron et al. 19819 (QAP); Pungala—comunidad
Alao, 01°52’S, 78°30°W, Ceron 28957 (QAP);
along Rio Alao, 01°52’S, 78°30’ W, 3200-3400 m,
S. Laegaard 55304 (AAU, QCA); Paramo de
Urbina, 3600 m, Acosta-Solis 21195, 21208, 21216

Festuca in South American Paramos

(US); 3600-4500 m, A.S. Hitchcock 21944, 21951
(US); 3700 m, E. Asplund 7803 (S); 3600 m, A.S.
Hitchcock 22030 (NY, US); road El Pocuy—El
Arenal, 3800 m, Acosta-Solis 166989 (US); Chim-
borazo, 4100-4400 m, Rauh & Hirsch E321, E331
(US); W slope of Chimborazo, 01°30’S, 77°50’ W,
4000 m, Ramsay & Smith 1016 (K); E side,
3480 m, Barclay & Juajibioy 8819 (COL, MO,
US); lado A, 3500 m, Barclay & Juajibioy 8162
(COL, MO, US); Bosque Andino de Cubillin,
3300-3400 m, Acosta-Solis 7554 (US); paramo
along the road to Guaranda, 4150 m, EF. Asplund
8191 (S); road Pallatanga—Cajabanba, km 47 from
Pallatanga, 3750-3800 m, B. Ollgaard & Balslev
8926 (AAU, MO, NY); B. Ollgaard & Balslev
8962 (F); road to lago Colta, km 9 NE of San Juan,
3600 m, 21 May 1990, PM. Peterson 9245, E.J.
Judziewicz, R.M. King & P.M. Jorgensen (K, MO,
QCA, QCNE, US); Entre Culebrillas y Yanayacu,
3300-3450 m, Acosta-Solis 7614, 7623 (F, S, US);
Mun. Guaranda, W side of the volcan Chimborazo,
01°28’S, 78°52’ W, 4100 m, 5 Jul 1999, D. Stancik
3162, 3164, 3169, 3189, 3190, 3193 (PRC, QCA);
NNW side of volcan, above vereda Rio Colorado,
4150 m, 2 Jul 1999, D. Stanctk 3176 (PRC, QCA);
Mun. Riobamba, Volcan Chimborazo, sector Cruce
de los Arenales, 01°28'14.6"S, 78°54 '06'W,
4300 m, 20 Sep 2000, D. Stancik 3705B, 3707,
3708, 3709, 3714, 3715, 3716, 3717 (PRC, QCA.
Cotopaxi: Parque Nacional Cotopaxi, falda NNW,
4000 m, Ehrenburg I, 27, 36 (QCA); E of Loma
Ingapirca, 00°40°S, 78°30°W, 4000 m, 6 Nov
1982, Balslev et al. 3435, 3438 (QCA); Balslev et
al. 3437 (MO, QCA, US); falda N de Cotopaxi,
4000 m, Balslev et al. 3675, 3733, 3734, 3738
(QCA); Cerro Sinfana, 00°40’S, 78°28’ W, 15 Dec
1990, Ceron 12609 (QAP); railway station Coto-
paxi, 3500 m, E. Asplund 16807 (S); 3400 m,
E. Asplund 6471 (F, MO, S, US); 3500 m, E. As-
plund 18286 (S); 3350 m, 13 Apr 1990, P.M. Peter-
son 8721 (K, MO, QCA, QCNE, US); 3550 m,
E. Asplund 6369 (F, S); quebrada de Agualongo,
00°40’S, 78°30’ W, 3000 m, 10 Jan 1994, Ceron
18088 (QAP); Laguna Limpiopungo, 3800 m, 14
Apr 1990, PM. Peterson 8744, C.R. Annable &
M.E. Poston (K, MO, QCA, QCNE, US); along
road of pine-plantation, 00°37’S, 78°27 W, 3435
m, 10 May 1984, S. Laegaard 52107 (AAU, QCA,
QCNE); 00°38’S, 78°33’ W, 3400 m, S. Laegaard
69261 (AAU); 00°40’S, 78°30’ W, 3900 m, Munoz
207 (QCA); quebrada de Agualongo, 3400 m, 2
Feb 1992, Montesdeoca 280, 282, 284, 295, 339,

Festuca in South American Paramos

348, 585 (AAU); 00°40’S, 78°30°W, 4550 m,
Argtiello 348 (QCA); 4300 m, Argitiello 354,
357 (QCA); T: de Vries s.n. (AAU); Hacienda Pau-
zacha S of Volcan Cotopaxi, 00°44’S, 78°29°W,
3650 m, 2 Dec 1985, S. Laegaard 55734 (AAU,
QCA); Cotopaxi, 3550 m, EF. Asplund 6370 (AAU,
F, MO, S, US); 3600 m, E. Asplund 6349 (AAU, F,
MO, S, US); 3600 m, E. Asplund 6348 (AAU, F,
S); road Zumbahua—Pujili, km 33, 00°53’S,
78°48 W, 3850-3900 m, 4 Apr 1992, S. Laegaard
102103 (AAU, QCA); 00°53’S, 78°48°W, 3850—
3900 m, S. Laegaard 102100 (AAU); Angamarca
road, km 5 from junction to road Latacunga—La
Mana, 01°00’S, 78°55’W, 4000-4100 m, S. Lae-
gaard 102134B (AAU); between Limpio Punja y
Ruminahui, 00°38’S, 78°28°W, 3900-4200 m, S.
Laegaard 55748 (AAU); E of Pilalo, km 19,
3600 m, 15 Apr 1990, PM. Peterson 8759, CR.
Annable & M.E. Poston (K, MO, QCA, QCNE,
US); km 21.5, 3820 m, PM. Peterson 8770, C.R.
Annable & M.E. Poston (MO, QCA, QCNE, US);
SW of El Chaupi, km 6.6 on road to base of Illini-
zas, 3800 m, 1 May 1990, PM. Peterson 8951 &
C.R. Annable (K, MO, QCA, QCNE, US); Canton
Latacunga—Volcan Rutzalagua, 00°57’S, 78°33 W,
2900-3500 m, 14 Feb 1994, Ceron et al. 25697
(QAP); Paramo de Apagua entre Zumbagua y
Pilalo, 4200 m, Barclay & Juajibioy 8085 (MO,
US); Salcedo—Napo road, 00°55’S, 78°28’°W,
3800-3850 m, J. Luteyn 13452 (MO, NY, QCA,
QCNE, US); Pilalo—Latacunga road, 00°57’S,
78°58 W, 3400 m, Holm-Nielsen 1485 (AAU, NY,
US); road Quevedo—Latacunga, Zumbagua,
3700 m, Harling et al. 8924 (GB); Mulatos paramo,
lagunas, 13000 ft, Prescott 656 (NY); Panamerican
Hwy., km 6 A Lasso, quebrada vegetation ENE
Pastocalle, 3400 m, Sparre 15828 (S); Mun.
Chaupi, NE slope of volcan IIliniza Norte, 00°38'S,
78°43 ’'W, 4000-4050 m, 12 Oct 2000, D. Stancik
4026, 4027, 4030, 4032 (PRC, QCA); Mun. Lasso,
Parque Nacional Cotopaxi, NE side of volcano,
00°37.8'S, 78°24.26'W, 3850 m, 21 Jun 1999,
D. Stanctk 3119, 3146, 3147, 3149, 3150, 3151,
3152, 3153, 3155, 3884, 4026 (PRC, QCA); around
Museum, 3500 m, D. Stancik 3158 (PRC, QCA);
Parque Nacional Cotopaxi entrance, 00°39’S,
78°31°W, 3530 m, 28 Sep 2000, D. Stancik 3888,
3889 (PRC, QCA); SE slope of Cotopaxi, 00°37’S,
78°24.5’W, 3800-3900 m, 28 Sep 2000, D. Stancik
3882 (PRC, QCA). Imbabura: Paramo Mariano
Acosta, road Yahuarcocha—Mariano Acosta, km 20,
00°20’S, 78°00’ W, 3650-3750 m, 9 Feb 1992,

133

S. Laegaard 101181, 101164B (AAU, QCA);
Laguna Grande de Mojanda, 00°08’S, 78°17’ W,
3850 m, S. Laegaard 55635 (AAU); Volcan
Cayambe, S side, 78°10’ W, 00°15’N, 4300 m,
Sklenar & Kosteckova 1881 (AAU); vicinity of
Mojanda, 15 km N of Malchinguin, S of Otavalo,
4000-4500 m, 11 Oct 1974, Gentry 12685 (MO);
Volcan Cotacachi, 00°22’N, 78°20’°W, 4200-—
4300 m, S. Laegaard 54506 (AAU); Laguna Cui-
cocha, 3100-3300 m, Penafiel et al. 527, 667, 720
(MO, QCNE); 3100-3400 m, Peviafiel et al. 402
(MO, QAP, QCNE); 3300-3350 m, Penafiel 1067
(QAP); Proantag, estrivaciones occidental de la
Cordillera Oriental, 3500-3800 m, Acosta-Solis
19196 (US); Hacienda Yura Cruz, 10 km N of
Ibarra, 00°22’°N, 78°05’W, 3700-3800 m, Holm-
Nielsen 6462 (AAU, NY, MO, S); quebrada Rumi-
pamba, 3700-4000 m, Ceron 29480 (QAP);
Cayambe—Laguna San Marcos, 11200 ft, Cazalet
& Pennington 5434 (K, NY, US); Cerro Imbabura,
faldas occidentales, 2800-4000 m, Acosta-Solis
17670 (US); Mun. Urcuqui, road to Cerro
Yanaurcu, 00°28°13°N, 78°18°45"°W, 4100 m, 15
Oct 2000, D. Stancik 4089, 4101 (PRC, QCA);
Mun. Cayambe, volcan Cayambe, 00°00’4°N,
78°01°40.4"W, 4200 m, 20 Oct 2000, D. Stancik
4163, 4164 (PRC, QCA); Mun. Otavalo, road from
Otavalo to Laguna Mojanda, 00°07°57°N,
78°16'27'W, 3800 m, 19 Oct 2000, D. Stancik 4115
(PRC, QCA); Mun. Urcuqui, road to Cerro
Yanaurcu, 00°28’N, 78°17.5°W, 3900 m, 15 Oct
2000, D. Stancik 4096, 4164 (PRC, QCA). Loja:
Road to Fierra Urcu, ca. 10 km from main road
Loa—Saraguro, 03°33’S, 79°15’W, S. Laegaard et
al. 18868 (AAU, LOJA, QCA, QCNE); 03°43’S,
79°19’ W, 3650 m, S. Laegaard & Sklenar 20282A
(AAU, LOJA); 03°’S, 79°19’°W, 3520 m, Sklenar
& S. Laegaard 7092 (AAU); Cerro de Arcos, W of
road Manu—Zaruma, 03°34’S, 79°28°W, 3500-
3600 m, S. Laegaard et al. 20613 (AAU, LOJA);
Mun. Saraguro, road to Fierra Urcu, 03°42°40°S,
79°18'12”"W, 3400-3450 m, 24 Aug 2000,
D. Stanctk 3769: 3770, 3772,°3773; 3774 (RG,
QCA). Morona — Santiago: Hda. Huargualla—Hda.
San Eduardo, way to Parque Nacional Sangay,
O27025'S,) 18 27 he Oo U0 im 30 Sol 1998.
D. Stancik 3326, 3327 (PRC, QCA); Parque Nacio-
nal Sangay, Plazabamba, 02°0.64’S, 78°26’ W,
3600 m, 20 Jul 1999, D. Stancik 3337 (PRC, QCA);
confluence of Quebrada Plazabamba Chico with Q.
Plazanbamba, 02°0.48’S, 78°25.4’W, 3000 m, 21
Jul 1999, D. Stancik 3344 (PRC, QCA); ridge

134

above Q. Plazabamba Chico, 3600 m, 20 Jul 1999,
D. Stanctk 3351 (PRC, QCA); Plaza Culebrillas,
01°58’S, 78°25’W, swampy plate with grassy
vegetation and shrubby patches, 3500-3600 m, 22
Jul 1999, D. Stancik 3355 (PRC, QCA); Plaza
Culebrillas, 01°58’S, 78°25’ W, 3500-3600 m, 22
Jul 1999, D. Stancik 3353, 3354, 3360, 3363, 3367
(PRC, QCA). Napo: Road Salcedo—Napo, ca. 6 km
NE of km 45, 3900 m, S. Laegaard 53373 (AAU,
QCA, QCNE); Paramo de Miranda, 00°34’N,
77°39’ W, 3700-3900 m, 23 May 1985, S. Laegaard
54409 (AAU, QCA, QCNE); 3900-4100 m,
S. Laegaard 54416 (AAU, QCA, QCNE); Cerro
Antisana, SW slope, 4200 m, Black 32 (AAU);
below Laguna Micacocha, paramo de Loma Gorda,
3850 m, Holm-Nielsen 20789 (AAU); W side of
Volcan Puntos, 00°12’S, 78°10’ W, 4150-4200 m,
S. Laegaard 54731 (AAU); Los Llanganati, entre
Ainchilibi y Rio Portrero al E de Romo Paramo,
2500-3620 m, Barclay & Juajibioy 9160 (COL,
US); 3600-3700 m, Barclay & Juajibioy 9142
(COL, MO, US). Pichincha: Volcan Pichincha, via
occidental, 2800-3830 m, Barford & Blicher-
Mathiesen 41546 (AAU, QCA, QCNE); 3500-
5000 m, Gentry 12378 (MO, QCA); Guagua
Pichincha, 4000 m, Harlington 4552 (S); 3500—
4500 m, Ceron 28250 (QAP); N slope, ca. 5 km
WSW of Cotocollao, 00°08’S, 78°33°W, Sparre
13709 (S); Sodiro s.n. (QPLS); Pichincha—Cerro
Ventanillas, 00°09’S, 78°32’°W, 3850 m, S. Lae-
gaard 51060 (AAU, QCA, QCNE); Lloa—Guagua
Pichincha, km 10, 4170 m, S. Laegaard et al.
102733 (AAU, QCA, QCNE); Sodiro s.n. (QPLS);
Jameson 70 (K); Hacienda Montecielo on S slope,
3400-3800 m, Sparre 17408 (S); Jameson s.n. (K);
3400 m, FE. Asplund 6134 (S); 8500 ft, Spruce 5509
(K, P, US, W); Sodiro s.n. (US); 3700 m, Balslev
23600 (AAU); N slope, ca. 5 km WSW Cotocollao,
3600-3750 m, Sparre 13709 (AAU); Volcan Pasa-
choa, 00°27’S, 78°30’ W, 3700-3900 m, S. Laegaard
55279 (AAU, COL, F, GB, K, LOJA, MO, NY,
QCA, QCNE, VEN, W); 3700-3900 m, S. Lae-
gaard 55279 (COL, MO); 00°27’S, 78°30’ W, 3900
m, S. Laegaard 55276 (AAU, QCA); 00°27’S,
78°30°W, 3600 m, S. Laegaard 55286 (AAU);
3000-3500 m, Ceron & Alarcon 12260 (MO, PRC,
QAP, QCNE); 3500-4300 m, Ceron & Alarcon
4805 (QCNE); 00°27’S, 78°28°W, 2850-3900 m,
Paredes 41 (QCA); 3800 m, Penland & Summers
969 (F); Paramo Papallacta, 3900 m, Maf 57
(AAU); Guamani, 01°19’S, 78°12’ W, 3600-3800 m,
E. Asplund &718 (QCA, S); 3200 m, Mille s.n.

Festuca in South American Paramos

(QPLS); Mille 284 (US); E. Asplund 9634 (S); E.
Asplund 8724 (MO, NY, QCA, S, US); 3900 m, E.
Asplund 17176 (S); 4000 m, Vargas et al. 2083
(AAU, MO, QCNE); Ramsay 426 (K); 4200 m,
Quintana 2 (QCA); 00°15’S, 78°12’ W, 3800 m, 7
Oct 1987, Ramsay & Smith 742 (K, QCA, QCNE);
3800 m, Harling et al. 10477 (GB); Andibus Qui-
tensis, Jameson s.n. (K, S); vicinity of Quito,
3200 m, EF. Asplund 6140 (S); Quito—Andes,
Jameson 91 (K, W); 3500 m, E. Asplund 6138 (F,
S); Volcan Antisana, falda W, 4200 m, Ehrenburg
191 (QCA); 4150 m, Grubb et al. 656 (NY, K); W
side, 00°28’S, 78°12’ W, 4200 m, Balslev et al. 4070
(QCA); Hacianda de Antisana, Hartweg 1455 (K,
NY, US); between campamento IMAP and Laguna
Micacocha, 00°33’S, 78°12°W, 3850-3950 m,
S. Laegaard 101577 (AAU); between Mt. Antisana
and Mt. Sincholagua, E. Asplund 8647 (NY, QCA,
S); E. Asplund 8645 (S); E. Asplund 8644 (QCA,
S); road from Otavalo to Laguna Mojanda, 3810 m,
J.F. Smith 1997 (AAU, F, MO, NY, QCNE, US);
J.F. Smith 1999 (NY, QCA, QCNE); D. Stancik
4103 (PRC); 00°08’S, 78°16’°W, 3725-3750 m,
S. Laegaard 52373 (AAU, QCA); Laguna Negra,
00°08’S, 78°16’ W, 3800 m, S. Laegaard 54351
(AAU, QCA, QCNE); Il liniza Sur, E slope, 00°40’S,
48°42’ W, 4200 m, Sklenar & Kosteckova 14-9 (US);
Illiniza Norte, 4000-4100 m, Sparre 15781 (S);
Mount Atacazo, 4700 m, Harteman 67 (US);
00°02'°N, 78°37° W, 4250 m, S. Laegaard 53871
(AAU, QCA, QCNE); 00°20’S, 78°38°W, 3750-
3800 m, S. Laegaard 55672 (AAU, QCA), SW
slope, km 19 from San Juan, 00°21°S, 78°39’ W,
2900 m, Holm-Nielsen 25163 (AAU); 00°20’S,
78°36'W, 3600 m, Espinosa 81 (QCA); road
Chillogalo—San Juan, 00°18°S, 78°39°W, 3100-
3260 m, 8 Sep 1945, Jaramillo & Zak 623 (AAU,
MO, PRC, QCA); Sincholagua, E. Asplund 8639
(QCA, S); Pululahua—San Bartolo al E de Moras-
pungo, 00°01’S, 78°29’ W, 3050 m, 13 Dec 1990,
Ceron & Montesdoca 12010 (QAP); Mont Ungui,
3400 m, Firmin 137 (F, US, W); El Corazon,
11000 ft, Prescott 834, 839, 848 (NY); 4100 m,
E. Asplund 17522 (S); E side of Corazon, 13-—
14,000 ft, Feb 1880, Whymper 1322 (K); Pogonales
de Pichincha, 13 May 1930, Benoist 2515 (P); Mun.
Pifo, Paramo de Guamani, 00°19’S, 78°15’ W, 3700
m, 19 Jun 1999, D. Stanctk 3013 (PRC, QCA, W);
Mun. Amaguafia, E slope of volcan Pasachoa,
00°27'56"S, 78°28'40"W, 4150 m, 14 Sep 2000,
D. Stanctk 3671, 3677, 3687, 3688, 3690, 3691,
3692, 3704 (PRC, QCA). Sucumbios: SW de

Festuca in South American Paramos

Playon de San Francisco, Paramo Mirador, 3400—
3600 m, 15 May 1990, PM. Peterson 9155, E.J.
Judziewicz & R.M. King (MO, QCA, QCNE, US).
Tungurahua: Cordillera de Llanganatis, Lake
Yanacocha, 3600 m, E. Asplund 9967 (QCA); El
Tambo near Lake Yanacocha, 3650 m, E. Asplund
9964 (S); Parque Nacional Llanganatis, via
Salcedo—Tena; 3600 m, 17 Apr 1998, Vargas et al.
2583 (MO, QCNE); Lake Aucacocha, 3700 m, Aug
1969, Edwards 70 (K); Las Tolas—5 km SE of
Laguna Pisayambo, 01°07’S, 78°21’ W, 3700 m, 12
Jan 1999, S. Laegaard & Grignon 19395 (AAU,
LOJA, QCA); road Ambato—Guaranda, 01°25’S,
78°51 °W, 4070 m, Brandbyge 42133 (AAU, QCA);
Comunidad Rumipata, 00°22’S, 78°55’ W, 4000 m,
Brandbyge 42586 (MO, AAU); Cerro Sangay, 4000
m, Stiibel 257 (S); Mun. Pillaro, Las Llanganatis,
around Aucacocha lagoon, 01°9’S, 78°20°W, 3800
m, 28 Sep 2000, D. Stancik 3901, 3902 (PRC,
QCA); Las llanganatis—Pisayambo, 01°07’S,
78°20’ W, 3600 m, 28 Sep 2000, D. Stancitk 3900
(PRC, QCA). PERU. Amazonas: Prov. Cha-
chapoyas, encima de Leimebamba, pajonal,
3400-3500 m, 17 Mar 1964, R. Ferreyra 15518
(US); upper slope and summit of Cerro Cam-
panario, 3600-3900 m, 3 Aug 1962, J. Wurdack
1571 (US). Ancash: Prov. Huaylas, Huascaran
Parque Nacional, quebrada Alpamayo at foot of
snowfree peak above Lago Jancarirish, 08°53’S,
77°41 W, 4350-4500 m, 3 Sep 1985, Smith et al.
9783 (MO); Prov. Recuay, Huascaran Parque
Nacional, Rio Pachacoto drainage between mineral
springs and Pumashimi, 09°53’S, 77°17’ W, 4200-—
4300 m, 16 Mar 1986, Smith et al. 11799 (MO);
Prov. Yungay, Huascaran Parque Nacional, Llan-
ganuco sector, quebrada Demanda, trail to Chacra-
raju base camp and Brogue glacier, 09°02’S,
77°36 W, 4100-4400 m, 4 Dec 1985, Smith et al.
10253 (MO); Huascaran Parque Nacional, Llanga-
nuco sector, quebrada Ancosh, 09°03’S, 77°35’ W,
4500 m, 4 Feb 1985, Smith et al. 10229 (MO).
Cajamarca: Prov. Cajamarca, dist. Cajamarca,
entre Cajamarca y Cumbe Mayo, km 14, Al arbo-
retum Cumbe Mayo de Cicafor, 3400 m, 18 Apr
1981, Vega et al. 2464 (F); Cajamarca-Bambamarca
road, in puna, 07°00’S, 78°33’ W, 3800 m, 17 Feb
1983, Smith 3482 (MO); Bajando el Paso El
Gavilan hacia San Juan, el E de la carretera Caja-
marca—Pacasmayo, 3000 m, 18 May 1986, Vega
4043 (F); a la altura del Paso El GavilanGavilan,
3200 m, 18 Apr 1976, Vega 1385 (F); Jalca de
Kumullca, entre La Encanada y Celendin, 3600 m,

135

27 May 1977, Vega et al. 2011 (F); Sorochuco,
Carretera Michiquillay—El Punre, 07°00’S,
78°18°W, 3570 m, 9 Aug 2001, Vega et al. 10904
(F); 9 Sep 2001, Vega et al. 10906 (F); humid jalca
vegetation with tussock grasses, 07°02’S, 78°15’ W,
26 Aug 2004, Sklenar & Zapata Cruz 8630 (PRC);
Cumbe Mayo, 21 km al W de Cajamarca, ladera
con arbustos dispersos, 3100 m, 4 Nov 1977, Vega
1969 (F); Coymolache ruta Cajamarca—Hualgalloc,
3850 m, 7 Jan 1977, Vega et al. 2052 (F); Pampa
Larga, al N de la explotacion minera Yanacocha,
ladera graminosa de jalca, 3900 m, 14 May 1994,
Vega 7136 (F); Prov. Hualgayoc, desvio de la car-
retera Coymolache—Chugur, jalca graminosa, 3700
m, 28 Apr 1994, Vega et al. 7062 (F); Prov. Hual-
gayoc, Coymolache, 06°46’S, 78°38’ W, 4000 m,
15 Jun 2001, Vega et al. 10795 (F, MO); Prov.
Hualgayoc, El Tingo, desvio a Chugur, 3350 m, 22
Jul 1986, Vega 4145 (F). Cuzco. Paso de Tres
Cruces, Cerro de Cusilluyoc, 3800-3900 m, 5 Mar
1925, Pennell 13845 (NY); Sacsahuaman, rocky
stream-bank, 3500-3600 m, 24 Apr 1925, Pennell
13552 (NY); Prov. Espinar, Hda Cuyo, 4200-4500
m, 24 Mar 1956, Vargas 11176 (US). La Libertad.
Prov. Bolivar, ascenso ao Nevado de Cajamarquilla,
07°08’S, 77°42’W, 3000 m, Vega 1118] (F, MO);
entre el desvio a Uchumarca y Santa Luisa, 07°04’S,
77°49'W, 3700 m, Vega 11190 (F, MO); Prov.
Otuzco, Trujillo-Huamachuco road, 10-15 km
before Shorey, 07°59’S, 78°22’ W, 3300 m, 13 Feb
1983, Smith et al. 3278 (MQ); Prov. Pataz, grass-
land in Manachaqui valley, ca. 10 km NE of Pataz,
07°40’S, 77°30’ W, 3600 m, 22 Mar 1988, B. Leon
& Young 1679 (MO); Manachaqui valley, ca. 10 km
NE of Pataz, 77°30’'W, 07°40’S, 3600 m, 28 Feb
1988, B. Leon & Young 1101 (MO); Prov. Santiago
de Chuco, al oeste del cementerio de Quiruvilca,
3870 m, 21 May 2001, Leiva & Leiva 2525 (MO);
Paso de Alaska. Carretera a Tayabamba, 3900 m,
24 Jun 1974, Lopez & Sagasteguii 8180 (MO, US).
San Martin. Prov. Huallaga, distr. Saposoa, entre
la Artesa y Rangra Pata, 3600-3800 m, 23 Aug
2001, Quipuscoa & Vilchez 2611 (F, MQ); Prov.
Mariscal Caceres, Pampa de Cuy, NW sector Rio
Abiseo Parque Nacional, 07°40’S, 77°30’ W, 3550
m, 3 Jun 1988, B. Ledn & Young 1364 (MO); Cho-
chos valley, NW corner of Rio Abiseo Parque
Nacional, 3425 m, 31 May 1986, Young & Watson
3484 (MO).

Note. Ocassionally there are proliferating plants
with clearly transformed spikelets and sexual organs.
Panicles are compact with short deformed branches;

136

spikelets are unisexual, transformed into vegetative
shoots, only exceptionally with some spikelets car-
rying rudiments of sexual organs; and some florets
are without anthers and ovaries. These plants can be
distinguished from F. asplundii by having narrower
leaf blades (0.4—0.8 versus 1—1.4 mm wide) and
viviparous panicles that are abnormally developed
(versus normally developed in F’ asplundii).

Viviparous specimens examined. COLOMBIA.
Narino: Mun. Tuquerres, Volcan Azufral, road from
vereda San Roque Alto to Laguna Verde, km 6,
3850 m, 9 Mar 1999, D. Stancik 2790 (COL, PRC,
PSO). ECUADOR. Morona-Santiago: Parque
Nacional Sangay, Plazabamba, 02°00.64’S, 78;
26'W, 3600 m, 19 Jul 1999, D. Stancik 3335, 3336
(PRC, QCA). Pastaza: E of pass Alao—-Huamboya,
01°48°S, 78°25’ W, 3700-3900 m, S. Laegaard
55452 (AAU, PRC, QCA). Tungurahua: Llanga-
natis, Las Tolas, 5 km SE of Laguna Pisayambo,
01°07°S, 78°21°W, 3700 m, 12 Jan 1999, S. Lae-
gaard 19395 (AAU, LOJA, QCA); S of Laguna
Verde at Cerro Hermoso, 01°14°S, 78°18 'W,
3850 m, 11 Nov 1980, Holm-Nielsen & Jaramillo
28415 (AAU, QCA).

53. Festuca sumapana Stancik, Darwiniana
41(1-4): 144, f. 7l-p. 2003. (Figs. 67, 68).
Type: Colombia. Meta, Paramo de Sumapaz,
Cerro Nevado del Sumapaz, 4130 m, 16 Jan
1973, Cleef 7930 (holotype: COL!; isotypes:
U!, US!).

Tussocked perennials with intravaginal innova-
tions. Culms 15—20 cm tall, erect, glabrous; nodes 1,
basal and 1 leaf. Leaf sheaths membranous, grayish-
stramineous, glabrous; auricles absent; ligules
0.3-0.5 mm long, membranous, apex two-lobate;
blades 8-12 cm long, 0.5—0.7 mm wide, involute,
abaxially scabrous, green, apex obtuse. Panicles
5—7 x 0.40.6 cm, lanceolate, contracted; branches
scabrous. Spikelets 9.5—11 mm long, lanceolate,
florets 3 or 4; rachilla glabrous; glumes 4-6 mm
long, lanceolate, keeled, membranous, purplish-
green, scabrous dorsally, apex acute; lower glumes
4—S mm long, 1-nerved; upper glumes 5.5—6 mm
long, 3-nerved; lemmas 6.5—7 mm long, lanceolate,
5-nerved, keeled, membranous, purplish-green,
papillose, awned, the awn 0.5—1.5 mm long; callus
glabrous; paleas as long as the lemma, lanceolate,
membranous, margins scabrous; lodicules lanceo-
late; anthers 0.6—0.8 mm long; ovary apex glabrous.
Caryopses not seen.

Festuca in South American Paramos

Leaf blade anatomy.—Cross-sections usually
with 5 vascular bundles, 3 ribs above; sclerenchyma
under abaxial epidermis discontinuous, adaxially
absent; adaxial epidermis with scattered hairs, the
hairs ca. 0.07 mm long.

Observations.—Festuca sumapana is morpho-
logically similar to F' glumosa but differs in having
shorter ligules (0.3—0.5 versus 0.8—1.2 mm), nar-
rower leaf blades (0.5—0.7 versus 0.8—1.4 mm), and
shorter lower glumes (4—5 versus 4.5—6 mm).

Distribution and habitat.—Festuca sumapana
is endemic to Colombia where it is known only
from the type locality, Paramo de Sumapaz (Cun-
dinamarca). It occurs at an altitude about 4000 m
in the superparamo zone.

54. Festuca tolucensis Kunth, Nov. Gen. Sp. (quarto
ed.) 1: 153. 1816. (Figs. 67, 69, 97A—C). Type:
Cresit in montosis, scopulosis, apricis regni
Mexicani, inter Islahuaca et Toluca, 1380
hexap., Sep, Humboldt & Bonpland s.n. (holo-
type: B!; isotypes: B!, BM, P!).

Festuca aequipaleata E. Fourn., Biol. Cent.-Amer.,
Bot. 3(20): 581. 1885. Festuca aequipaleata
E. Fourn., Mexic. Pl. 2: 125. 1886, isonym.
Type: Mexico. Orizaba, 14,000 ft, Liebmann
6108 (lectotype: C!, designated by Alexeev,
Novosti Sist. VysS. Rast. 21: 47. 1984; isolec-
totypes: C!, K!).

Festuca liebmannii E. Fourn., Mexic. Pl. 2: 124.
1886. TyPE: Mexico. Liebmann 517 (holotype:
P; isotype: US!).

Festuca multiculmis Steud., Syn. Pl. Glumac. 1:
310. 1854. Type: Mexico. Mt. Toluca, Heller
306 (holotype: P!). |

Tussocked perennials with intravaginal innova-
tions. Culms 60-80 cm tall, erect, finely scabrous;
nodes 1, basal. Leaf sheaths membranous-coriaceous,
stramineous to grayish, glabrous; auricles absent;
ligules 1.8—3.5 mm long, membranous, apex acute;
blades 20-30 cm long, 0.5—0.7 mm wide, involute,
green to olive-green, abaxially scabrous, apex acute.
Panicles 20—25 x 0.5—1.5 cm, lanceolate, contracted,
branches finely scabrous. Spikelets 7—12(—15) mm
long, lanceolate sometimes oblong-lanceolate, florets
3 or 4(—6); rachilla densely hairy; glumes 6—9.5 mm
long, lanceolate, coriaceous, green, markedly sca-
brous, apex acute; lower glumes (2—)6—8.5 mm
long, 1-nerved; upper glumes (4—)6.5—9.5 mm long,
3-nerved; lemmas 6—9.5(—10.5) mm long, lanceolate,
5-nerved, coriaceous, green, scabrous on all surfaces,

hy

Festuca in South American Paramos

qk

Figure 68. Festuca sumapana. A. Stylized growth form. B. Habit. C. Ligule. D. Spikelet. E. Glumes. F. Lemma. G. Lemma with

palea and rachilla. H. Leaf blade cross-section. A-H, Cleef 7930 (COL).

138 Festuca in South American Paramos

HM

Ni
Mn NIA
INNUN

|
hie

\

‘
\\

—~

= :
a [
iN

=

—_-

———e
>~

ee ara

SEES

—

—S

——=——=

——

=
——
——
—— —
es

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Figure 69. Festuca tolucensis subsp. tolucensis. A. Stylized growth form. B. Habit. C. Ligule. D. Spikelet. E. Glumes. F. Lemma.
G. Lemma with palea and rachilla. H. Leaf blade cross-section. A-H, Stancitk 4279 (PRC).

Festuca in South American Paramos

awned, the awn 1—1.5 mm long; callus glabrous;
paleas almost as long as the lemma, lanceolate, mem-
branous, scabrous, short-hairy; anthers 2.5—3 mm
long; ovary apex glabrous. Caryopses lanceolate;
hilum 5/6 as long as the grain, linear.

Leaf blade anatomy.—Cross-sections typically
with 5—9 vascular bundles and 3—5 ribs above;
sclerenchyma under abaxial epidermis continu-
ous, extending to somevascular bundles, adaxial
sclerenchyma present in 3—S ribs, isolated; abaxial
epidermis with a dense covering of prickles; adaxial
epidermis hairy, the hairs 0.03—0.05 mm long.

Observations.—Festuca tolucensis is morpho-
logically similar to F subulifolia and both species

139

form tussocks of fine-leaved plants. Besides differe-
ing in leaf cross-section, F: tolucensis has spikelets
7-12(-15) mm long (versus 6.5—9 in F. subulifolia),
lower glumes 6.5—9.5 (versus 3.5—5.5) mm long,
and upper glumes 6.5—9.5 (versus 4-6) mm long.

Distribution and habitat.—Festuca tolucensis
has a wide distribution and ranges from Mexico
to Venezuela. In Colombia, Festuca tolucensis
is only known from the Serrania de Perija. We
recognize three subspecies as differentiated in
the tabular comparison and key below. It is a
dominant species found in dry rocky slopes or
swampy margins of lagoons in grass paramos
between 3000—4000 m.

F. Distribution Glume length (mm) Lemma Spikelets Vascular bundles
tolucensis Lower Upper’ Length No. florets Length No. Girders
subsp. (mm) (mm)

tolucensis MEX-VEN (4-)5-6 (5-)6-8 6.5-7.5 4-5 7-10 7-9 1—3(—5)
perijae ROL, VEN 5.5-6.5. 6.5-7.5.. 6-7 2 or 3 10 5 0

culata VEN 2-2.5 4-5 6-7 4or 5 10-11 5 |

la.

1b.

2a.

2b.

KEY TO THE SUBSPECIES OF FESTUCA TOLUCENSIS

Lower glumes 2—2.5 mm long; upper glumes 4-5 mm long; spikelets 10—11 mm long..................

armen 9 Sonn etn Ser t SE) see aca (eerie hs 2... cbt efebecatcd ae eeeitend 54c. F. tolucensis subsp. culata

Lower glumes (4—)5—6.5 mm long; upper glumes (5—)6—8 mm long; spikelets 7-10 mm long.... 2

Small tussocks (less than 40 cm tall), leaves with 5 vascular bundles, spikelets with 2 or 3 florets;
pom) IN TA ee cee Bae Ee eee en ee Ree ee 54b. F. tolucensis subsp. perijae
Tussocks more than 40 cm tall, leaves with 7—9 vascular bundles, spikelets with 4-5 florets; 7-10 mm
eee a Nee esc Ee WA eho can nse he Raletueaivs 54a. F. tolucensis subsp. tolucensis

54a. Festuca tolucensis subsp. tolucensis (Fig. 69).

Additional specimens examined. COSTA
RICA. San José: Iraza, 1300 m, 14 Jun 1932, Storek
2889 (MO); Canton de Pérez Zeledén, Parque Nacio-
nal Chirripo, Cordillera de Talamanca, Rio Talari,
cerca del cruce al Cerro Ventisqueros, 3600—3450
m, 27 Jan 1996, Morales 5194 (MO); Canton de
Pérez Zeledon, Parque Nacional Chirripo, cuenca
Terraba—Sierpe Chirripo, base Crestones, 09°27.5'N,
83°30.5'°W, 3460 m, 12 Jul 1996, Alfaro 1029
(MO); Canton de Pérez Zeledon, Parque Nacional
Chirripo, cuenca Terraba—Sierpe, Sandero a Valle
Los Leones, 09°26'N, 83°30.5’°W, 3100-3400 m,
18 Jul 1998, Alfaro 1771 (MO). MEXICO. D.F.:
Ajusco, Sep 1928, Lyonnet 260 (NY); Lomas, Oct

1932, Lyonnet 2973 (MO). Volcan Toluca, 4150 m,
18 Oct 1953, Sohns & Matuda 991 (NY); W side
in volcanic ash, competing with Sedum minimum,
3940 m, 9 Nov 1955, J. Clausen 34 (NY); crater,
above timberline, dwarf alpine plants, 14000 ft, 17
Aug 1972, Dziekanowski & Bolingbroke 1911 (MO,
NY); 1846, Heller 62, 306 (P, W); 4150 m, 18 Oct
1953, Sohn & Matuda 99] (P). Iztaccihuatl, side of
Mt. at La Joya, 3990 m, 7 Jun 1960, J. Beaman 3482
(NY); Monte de Rio Frio, km 49, road from Mexico
City to Puebla, 4000 m, 31 Jul 1929, Mexia 2684
(NY); Mt. Popocatepetl, 3400 m, 8 Jun 1910, A.S.
Hitchcock 496 (C, MO, NY, W); Tlaxcala vertiente
N del Volcano La Malinche, zacatonal de Festuca y
Calamagrostis, 3920 m, 16 Sep 1986, Gonzalez et
al. 206 (F, MO); Mun. de Amecameca, bosque de

140

pino, 3900 m, 30 Nov 1980, Juan & Alva 101 (MO).
Chiapas: Summit of Volcano Tacana, Mun. Union
Juaréz, scattered dwarfed Pinus sp., 2200 m, 30 Jul
1972, D. Breedlove 26723 (NY); Volcan Tacana,
near summit sandy, gravelly soil above timberline,
4090 m, 8 Dec 1959, J. Beaman 3216 (NY); Mt.
Tacana, 2000-4038 m, VIII 1938, Matuda 2337,
2360 (NY). Jalisco: Nevado de Colima, side of
mountain below Puerto de Colima, sandy soil in
sparsely vegetated meadow, 4000 m, 26 Aug 1958,
J. Beaman 2363 (NY); S of Ciudad Guzman (Zapot-
lan), near summit, in rockslide, gravel, around rocks,
4000 m, 7 Feb 1956, Gregory & Eiten 301 (MO,
NY, P); Mt. Nevada, 14300 ft, 23—24 Sep 1910, A.S.
Hitchcock 7165 (NY). Michoacan: Tancitaro region,
Mt. Tancitaro, 10000-11000 ft, 25 Jul 1941, Leav-
enworth & Hoogstraal 1218 (MO, NY). Puebla:
Arroyo Paso Buey, NW side of volcano Pico de
Orizaba and 7 km NW of the summit, 0.5 km SE of
Miguel Hidalgo, 97°18 °30"W, 19°04°30'N, 3300 m,
9 Jul 1986, M. Nee & Soule 33022 (MO, NY); Faldas
del Pico Orizaba, 3300 m, 17 Oct 1971, Hernandez
et al. 1336 (F, MO). Veracruz: Mun. Perote, km
1—2 above Escobillo on the NW slopes of Cofre de
Perote 3300 m, 21 Jan 1984, Taylor et al. 170 (F,
MO, NY); Pico Orizaba, NE side of Mt., 3980 m, 16
Aug 1958, J. Beaman 2284 (NY); Mt. Orizaba, sandy
plains, 14000 ft, 7 Aug 1891, Seaton 228 (NY); 18 km
de Perote camino a la cima del Cofre, 3790 m, 10 Feb
1984, Gonzalez et al. 137 (MO); Pico de Orizaba,
14000 ft, Sep 1941, Liebmann 2886, 3030, 6117 (C);
Wawra 947 (W); Ross 1276 (W). VENEZUELA.
Mérida: Sierra Nevada—Laguna La Coromoto,
3300-3400 m, 18 Feb 1966, Schulz & Rodriguez 317
(US); 13000 ft, Feb 1846, Funck & Schlim 1132 (P);
4000-4400 m, 24 Sep 1952, Humbert 26353 (COL,
MER, P, US); 4000-4400 m, Humbert 26806 (MER);
Humbert 26506 (P, US); pastizal, 3700 m, 25 Sep
1970, Castelano & Monasterio 20 (VEN); Laguna
La Coromoto, Plantaginetum, 3200 m, 24 Feb 1955,
Vareschii 3853 (VEN); 3400 m, Oct 1956, Aristegui-
eta 2602 (VEN, US); 3300 m, 3 Jul 1987, Briceno &
Adamo 1988 (MERF); Valle de Mucubaji, abajo del
Pico Mucufiuque, 08°46’35’N, 70°48 °57°W, 3730 m,
3 Sep 1998, Berg 98-29-16 (K); Laguna Mucubaji,
swamps around the lagoon, 8°47.5’N, 70°49’'W,
3600 m, 6 Nov 2000, D. Stancik 4172 (CAR, COL,
PRC); 3570 m, 23 Aug 1980, Bricenio & Adamo 184
(Herbarium Bricefio, MERF, MO, PRC, US); camino
hacia Mucunuque, 4000-4300 m, 4 Nov 1992, Meier
et al. 3026 (VEN); Laguna de Mucubaji, 3600 m,
Dec 1952, Aristeguieta 964 (US); Paramo de Mucub-

Festuca in South American Paramos

aji, 3500 m, 23 Aug 1980, Briceno & Adamo 187
(MO); S side of Hwy 7, 5 km E of Laguna Victoria,
3000 m, 22 Aug 1972, Hanselmann & Loleveless 296
(MO); Laguna de Mucubaji, 8 Nov 1952, Aristegui-
eta s.n. (VEN); 3900 m, 9 Sep 1958, Vareschi 7021
(VEN); 8 Nov 1952, Aristeguieta 947 (VEN). Mun.
Tabay, Laguna La Coromoto, 8°35.5’N, 71°01’W,
3300 m, 7 Nov 2000, D. Stancik 4181 (CAR, COL,
PRC); Laguna La Coromoto, 3300-3400 m, 15
Dec 1966, Schulz 317 (MER); Sierra St. Domingo,
Pico de Mucunuqui, al S de Laguna Grande, 4400
m, 28 Nov 1959, Barclay & Juajibioy 9895 (COL);
Paramo de Mucuchies, Pico de Aguila, 4418 m, 21
Nov 1959, Barclay & Juajibioy 9656 (COL, MO,
US); Sierra del Norte, 6 Oct 1952, Humbert 26824
(COL, MER, P, US); Mun. Mucuchies, Paramo de
Piedras Blancas, Lagua Negra, 8°49°N, 70°57’ W,
4270 m, 4 Nov 2000, D. Stancik 4191 (CAR, COL,
PRC, W); 4200 m, D. Stancitk 4207 (CAR, COL,
PRC, W); D. Stancik 4176 (AAU, CAR, COL,
PRC); 8°48.5°N, 70°55.5°W, 3800 m, D. Stancik
4225 (CAR, COL, PRC); Laguna Tapada, 8°49°N,
70°56'43"W, 4100 m, D. Stancik 4242 (AAU, CAR,
COL, PRC); 4100 m, D. Stancik 4243 (CAR, COL,
PRC, W); Laguna Negra, 9 Sep 1952, Aristeguieta
1003 (VEN); Laguna Los Patos, Laguna El Infer-
nito, 3700 m, Aug 1956, Aristeguieta 2464 (VEN).
Paramo de Piedras Blancas, 4400 m, 8 Dec 1979,
Barreto 645 (MERC); 4000-4600 m, 9 May 1985,
Briceno et al. 1146 (MERC); 4000-4600 m, 14
Nov 1981, Briceno et al. 440 (Herbarium Bricefo,
MERC); sector Las Cruces, 3700-3900 m, 8 Sep
2000, Briceno 3711 (Herbarium Briceno); Mun. La
Culata, Paramo La Culata, Briceno & Adamo 1067
(MERF); 3300 m, 7 Jun 1984, Briceno & Adamo
949 (MERF); Mun. Mucuchies, Paramo de Misinta,
NE de Mucuchies, 4000 m, 26 Oct 1984, Briceno &
Adamo 1139 (MERF); Paramo Las Monsalves, selva
de Polylepis, 3900 m, 10 Nov 1952, Vareschi 2232,
2134, 2244 (VEN); 3920 m, Vareschi 2248 (VEN);
Pico Espejo—Timoncito, paramo, 4300-4700 m, 16
Dec 1952, Bernardi 275 (NY). Tachira: Mun. La
Grita, Paramo La Negra, 08°15’N, 71°53°W, 3200 m,
11 Nov 2000, D. Stancik 4280 (CAR, PRC); D.
Stancik 4279 (CAR, COL, PRC); Sierra Nevada,
camino a Pico Bolivar, cerca de Refugio Moya,
4100-4300 m, 10 Oct 1953, Littele 15712 (MER);
Pico Bolivar y Espejo, 15-18 Dec 1959, Barclay
& Juajibioy 10214 (US); Sierra Nevada, Loma
Redonda, Paramo Media Luna, 4100 m, 3 Feb 1995,
Berg 649 (MERC); Berg & Steinmetz 1&7 (Her-
barium Briceno, MERC); Loma Redonda, 3940 m,

Festuca in South American Paramos 14]

7 Oct 1994, Briceno 2854 (Herbarium Bricenio); Alto
de La Cruz, 4040-4300 m, 11 Nov 1994, Briceno et
al. 3077 (Herbarium Bricefio); Mun. Rangel, Laguna
La Ciega, 4200 m, 29 Sep 2000, Bricentio & Molinillo
3766 (Herbarium Bricefio); Mun. Rangel, Paramo El
Banco, 4420 m, 21 Oct 1997, Bricefio & Molinillo
3447 (Herbarium Bricefio); Paramo de Misinta,
NE de Mucuchies, 4000 m, 5 Jun 1987, Briceno &
Adamo 1131 (Herbarium Bricefio); Paramo EI Toro,
Los Arbolitos y Los Nevados, 3100 m, 12 Oct 1988,
Briceno et al. 2330 (Herbarium Bricefio); cresta Pico
el Toro, 4200 m, 25 Nov 1994, Briceno 3119 (Her-
barium Bricefio, PRC); via El Leon, 3460-3580 m, 4
Nov 1994, Briceno 311] (Herbarium Bricefio, PRC);
Valle de los Calderones, Paramo de los Pozones, 3350
m, 19 Oct 1995, Bricerio 3357 (Herbarium Bricefio);
Laguna Verde—Laguna Suero, 3990-4190 m, 29 Mar
1994, Briceno et al. 2667 (Herbarium Bricefio);
Laguna El Espejo, 3800-3950 m, 9 Dec 1994,
Briceno 3178 (Herbarium Bricefio); Laguna de Los
Anteojos, 3900 m, 2 Dec 1994, Bricenio et al. 3158
(Herbarium Bricefio); Laguna La Fria, 3460-3580
m, 4 Nov 1994, Briceno et al. 3054 (Herbarium
Bricefio, PRC). Trujillo: Paramo de la Cristalina,
2900 m, Oct 1910, Jahn 18 (US, VEN); 20 Oct 1910,
Jahn 43 (VEN); Tuname—Guirigoy, 3200 m, Aug
1958, Aristeguieta & Medina 3531 (VEN); Guirigoy,
hacia La Parida, 3400 m, Aug 1958, Aristeguieta &
Medina 3558 (VEN); Dept. Escuque, Cabimbi, Teta
de Niquitao—Laguna Negra, 3800-3900 m, 19 Aug
1988, Briceno s.n. (Herbarium Bricefio).

54b. Festuca tolucensis subsp. perijae Stancik,
Darwiniana 41(1-4): 146, f. lla—f. 2003.
Type: Colombia. Magdalena, Sierra de Perija,
E of Manaure, Cerro Avion, 3550-3450 m, 8
Nov 1959, J. Cuatrecasas & Castaneda 25136
(holotype: COL!; isotype: US!).

Panicles slender and compact. Spikelets
9-10 mm long; lower glumes 5.5—6.5 mm; upper
glumes 6.5—7.5 mm long; lemmas 6—7 mm long.

Leaf blade anatomy.—Abaxial epidermis con-
tinuous, not extending to the vascular bundles.

Distribution and habitat.—Fesctuca tolucensis
subsp. perijae is known only from the Serrania
de Perija on the Colombian—Venezuelan border,
and is found in paramos at an altitude between
3000-3500 m.

Additional specimens examined. COLOMBIA.
Magdalena: Sierra de Perija, plain between Cerro
Venado and Cerro Avion, 3270-3350 m, J. Cuatreca-

sas & Castaneda 25133 (COL, US). VENEZUELA.
Zulia: Sierra de Perija—Serrania de Valledupar, cam-
pamento “Monte Viruela’, 10°25°13”N, 72°52’42’W,
3100 m, S.S. Tillett 747-1122 (COL).

54c. Festuca tolucensis subsp. culata Stancik &
P.M. Peterson, subsp. nov. Type: Venezuela.
Mérida, Mun. La Culata, Paramo La Culata,
3300 m, 8°45°42°N, 71°03’ W, 3300 m, 12
Nov 2000, D. Stanctk 4259 (holotype: PRC!;
isotypes: CAR!, COL!).

Haec subspecies a Festuca tolucensis subsp.
tolucensis et F: tolucensis subsp. perijae spiculis
10-11 (non 7-10) mm, glumis minoribus (inferi-
oribus 2—2.5 non 4—6.5 mm, superioribus 4—5 non
6—7.5 mm) distinguitur.

Spikelets 10—11 mm long; lower glumes 2-2.5
mm long; upper glumes 4—5 mm long.

Leaf blade anatomy.—Abaxial epidermis
continuous, extending just to a single vascular
bundle.

Distribution and habitat.—Festuca tolucensis
subsp. culata is known only from Mérida where
the subspecies occurs in rocky paramos of La
Culata and El Escorial, at an altutude of about
3000-3350 m.

Additional specimens examined. VENEZU-
ELA. Merida: Paramo El Escorial, 3100-3420 m,
10 Jul 1987, Briceno & Adamo 2052 (Herbarium
Bricefio, PRC). Mun. La Culata, Paramo La Culata,
3300 m, 8°45°42’N, 71°03’°W, 3300 m, 12 Nov
2000, D. Stancik 4258 (CAR, COL, PRC).

55. Festuca turimiquirensis Stancik & P.M. Peter-
son, sp. nov. (Figs. 49, 70). Type: Venezuela.
Anzoategui, Distr. Libertad, ridges and tops
of Montanas Negras, along the Sucre and
Anzoategui border, 20 km NE of Bergantin,
NE of Buenos Aires, Serrania de Turimiquire,
10°04’30°N, 64°11 °W, 2000-2350 m, bamboo-
ericaceous scrub and elfin cloud forest, 28 Nov
1981, G. Davidse et al. 19610 (holotype: MO!,
isotypes: NY!, VEN!).

Haec species a F. tolucensis glumis minoribus
(inferibus 3—4 non 5—8.5 mm, superioribus 4—5.5
non 5—9.5 mm), aristis longioribus et paniculae
majoribus (6—7 cm lat., non 0.5—1.5 cm), et scler-
enchymate abaxiali non continui differt.

Tussocked perennials with intra- and extra-
vaginal innovations. Culms 70-80 cm tall, erect,

———
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——
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————

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Festuca in South American Paramos 143

glabrous; nodes 3 or 4 near base. Leaf sheaths
membranous, brown, glabrous, striate; ligules
1.5—4 mm long, apex acute; blades 40-50 cm
long, 0.6—0.7 mm wide, conduplicate, abaxially
glabrous, green. Panicles 15—17 x 6—7 cm, open,
ovate; branches glabrous. Spikelets 10—11 mm long,
ovate, florets 4; rachillas 1.2—1.4 mm long, hairy;
glumes 3—5.5 mm long, membranous, upper 1/3
scabrous, apex acute; lower glumes 3-4 mm long,
lanceolate, 1-nerved; upper glumes 4—5.5 mm long,
lanceolate, 3-nerved; lemmas 7—8.5 mm long, lan-
ceolate, 5-nerved, membranous, upper 1/2 scabrous,
awned, the awn 0.7—2.5 mm long; paleas as long
as the lemma, membranous, scabrous; anthers ca.
3 mm long; ovary apex sparsely hairy. Caryopses
lanceolate; hilum 5/6 as long as the grain, linear.

Leaf blade anatomy.—Cross-sections usually
with 7 vascular bundles, 5 ribs above; sclerenchyma
under abaxial epidermis discontinuous, forming large
fascicles, adaxial sclerenchyma present and extending
to every second vascular bundle; adaxial epidermis
with scattered hairs, the hairs ca. 0.1 mm long.

Observations. This Venezuelan species is geo-
graphically isolated and known only from the type
collection. Festuca turimiquirensis differs from
F. tolucensis by having smaller spikelets (lower
glumes 3-4 versus 5—8.5 mm long, upper glumes
4—5.5 versus 5—9.5 mm long), opened panicles (6—7
versus 0.5—1.5 cm wide), and the abaxial scleren-
chyma is discontinuous (versus continuous).

Distribution and habitat.—Festuca turimiqui-
rensis 1s restricted to the Serrania de Turimiquire
(Anzoategui) of Venezuela where it occurs in
grasslands on rocky slopes in open areas between
2000-2350 m.

56. Festuca vaginalis (Benth.) Laegaard, Novon
8(1): 30. 1988. (Figs. 64, 71, 97D). Poa
vaginalis Benth., Pl. Hartw. 261. 1846. Type:
Ecuador. Pichincha, Hacienda de Antisana,
K.T. Hartweg 1450 (holotype: K!; isotype:
US-88715!).

Tussocked perennials with intravaginal innova-
tions. Culms 20—60 cm tall, erect, glabrous; nodes
1, basal and 1 or 2 short leaves. Leaf sheaths coria-
ceous, stramineous, occasionally grayish, glabrous;
ligules 0.8—1.2 mm long, coriaceous, oblong or
triangular, apex short-ciliate; blades (4-)7—-17 cm
long, 0.4—0.8 mm wide, conduplicate, glaucous.
Panicles 10-15 x 0.5 cm, contracted, narrow,
whitish or purplish-white, occasionally pale green;
Spikelets 8—9.5 mm long, lanceolate, florets 2 or 3;
rachilla short-pubescent; glumes (5.5—)6.5—9 mm
long, almost as long as the spikelet, membranous,
white or with purple stripes, midnerve scabrous
distally, apex obtuse; lower glumes (5.5—)6.5—8
(—9) mm long, 1-nerved; upper glumes 6.5—9 mm
long, 3-nerved; lemmas 7.5—8.5 mm long, oblong,
5-nerved, membranous, apex short two-dentate,
awned, the awn 0.5—1 mm long; callus glabrous;
paleas 3/4 as long as the lemma, scabrous on keels;
lodicules lanceolate; anthers 0.6—0.9 mm long;
ovary apex glabrous. Caryopses lanceolate, hilum
2/3 as long as the grain, linear.

Leaf blade anatomy.—Cross-sections with
(7—)9-11 vascular bundles and 5—9 ribs above;
sclerenchyma discontinuous under both abaxial and
adaxial epidermis, girders absent; bulliform cells
absent; adaxial epidermis with scattered hairs, the
hairs 0.075—0.09 mm long.

Observations.—Festuca vaginalis is morpho-
logically distinct from the other species treated
in this revision and we do not know which spe-
cies might be its closest sister. Festuca vaginalis
has been mentioned as occurring in Ecuador
and Colombia (Luteyn 1999, Rangel 2000);
although its occurrence in Colombia has not been
confirmed.

Distribution and habitat.— Festuca vaginalis
is endemic to Ecuador. It occurs on volcanic rocks
in superparamos and dry grass paramos between
3500-5000 m. It also has spread secondarily onto
disturbed areas of dry grass paramos and along road
embankments, so called “arenal.”

KEY TO THE SUBSPECIES OF FESTUCA VAGINALIS

la. Culms 30-60 cm long, robust; leaf blades ca. 0.8 mm wide; panicles whitish or purplish-white....

COOP T HSE ETHH ESET ESTE ESSE ESTHESEEEHETEHHHEEH OHHH HH SHEESH EHEHEHEHETEOTOS

tee Ten oerr 56a. F. vaginalis subsp. vaginalis

1b. Culms 20-30 cm tall, not robust; leaf blades 0.4—0.5(—0.6) mm wide; panicles pale-green............

POPS OH SOSH SSE SSHTHEHEHHESHESOS HEHEHE TH THESE HHEHEHEHEEEHEHESEHESEHESESESESE

sea iia te alae a 56b. F. vaginalis subsp. cayambae

Festuca in South American Paramos

144

Figure 71. Festuca vaginalis subsp. vaginalis A. & B. Habit. C. Ligule. D. Spikelet. E. Glumes. F. Lemma. G. Lemma with palea

and rachilla. H. Leaf blade cross-section. A-H, Stanctk 3134 (COL).

Festuca in South American Paramos

145

56A. Festuca vaginalis subsp. vaginalis (Fig. 71).

Culms 30-60 cm long, robust. Leaf blades ca.
0.8 mm wide. Panicles whitish or purplish-white.

Leaf blade anatomy.—Cross-sections with
9-11 vascular bundles.

Additional specimens examined. ECUA-
DOR. Bolivar: Volcan Chimborazo, W side,
1°28°S, 78°48’°W, 4800 m, 14 Sep 1995, Sklenar
& Kosteckova 143-2 (QCNE); road Salinas—Los
Arenales, km 6, 01°22’S, 79°00’W, 4000 m, S.
Laegaard 55327 (AAU); valley 4 km S of Cruces
de los Arenales, 4270-4300 m, 3 Mar 1992, S. Lae-
gaard 101559 (AAU, QCA); 01°25’S, 78°54’ W,
4160-4200 m, S. Laegaard 101495 (AAU); km
66.5 SW of Ambato on Hwy to Guaranda and 1.2
on road Fecundo Vela, km 1.2 km, 4290 m, 03
May 1990, PM. Peterson 8987 & C.R. Annable
(K, MO, US, QCNE). Cotopaxi: Angamarca road,
km 5 from junction to Latacunga—La Mana road,
01°00°S, 78°55°W, 4250-4280 m, S. Laegaard
102149 (AAU); Volcan Cotopaxi, road to Refu-
gium, 00°40’S, 78°26’W, 4350 m, S. Laegaard
103127 (AAU); SW slope, 4100 m, FE. Asplund
7497 (F, K, NY, S, US); El Tambo SE of Volcan
Cotopaxi, 00°42’S, 78°18’ W, 3650 m, S. Laegaard
55536 (AAU); Hacienda Pauzacha, 00°44’S,
78°29" W, 3650 m, S. Laegaard 55735 (AAU);
road Zumbahua—Pujili, km 39, 00°53’S, 78°47’ W,
3750-3800 m, S. Laegaard 102096 (AAU); Parque
Nacional Cotopaxi, Laguna Limpiopungo, 3800 m,
P.M. Peterson 8739, C.R. Annable & M.E. Poston
(MO, QCA, QCNE, US); plate below NE side of
volcano, 00°37’S, 78°24’ W, 3800 m, 26 Jun 1999,
D. Stancik 3137 (PRC, QCA); NE side of volcano
Cotopaxi, 00°37.5’S, 78°24.2°W, 3820 m, 26 Jun
1999, D. Stancik 3134 (PRC, QCA); valley NW
of Limpiopungo to foot of Volcan Rumifiahui,
00°38°S, 78°28°W, 4300 m, 25 Feb 1992, S. Lae-
gaard 101460 (AAU, QCNE). Laguna Quilotoa,
00°51°S, 78°53’W, 3480-3820 m, S. Laegaard
101335A (AAU); Panamerican Hwy., km 2 S of
Cruce de Cordillera, 00°39’S, 78°35°W, 3500 m,
13 Aug 1984, S. Laegaard 52656 (AAU, QCNE);
S. Laegaard 52659 (AAU); S of Cruce de Are-
males, km 5, 01°32'S, 78°52°W, 3950 m, 18 Sep
1998, S. Laegaard 19159 (AAU, LOJA); Rancho
Santa Maria, 00°43’S, 78°29’W, 3750-3950 m,
S. Laegaard 52202 (AAU); Mun. Chaupi, NE slope
of volcan Illiniza Norte, 00°38'°11”S, 78°43'1"W,
4450 m, 12 Oct 2000, D. Stancik 4013 (PRC, QCA);
SE slope of volcan, 00°38’'45"S, 78°25’32”"W,

4300 m, 28 Sep 2000, D. Stancik 3881 (PRC, QCA,
US). Chimborazo: Volcan Chimborazo, lower
Refugio, 01°28’S, 78°50’W, 4840 m, S. Laegaard
102796 (AAU,QCNE); 5000 m, 11 May 1992, Rauh
& Hirsch E315 (US); road from Refugium, km
4.7, 01°27°S, 78°52’ W, 4330-4400 m, S. Laegaard
102795 (AAU, QCNE); Chimborazo, Sodiro s.n.
(US); lower Refugium, 01°27’S, 78°50’ W, 4800 m,
5 Mar 1988, S. Laegaard et al. 70553 (QCA);
Sodiro s.n. (QPLS); 01°27’S, 78°50’W, 4800 m,
S. Laegaard et al. 70554 (AAU, QCNE); paroguilla
San Juan, 01°30’S, 78°49’ W, 4150 m, 4 Aug 1992,
Ceron et al. 19822 (QAP); Cruce de los Arenales,
km 5, 01°27’S, 78°52°W, 4330-4400 m, S. Lae-
gaard 102795 (AAU, QCNE); km 4.7 from lower
refugio of Volcano Chimborazo, arenales, 01°27’S,
78°50°W, 4500 m, 5 Mar 1988, S. Laegaard & S.A.
Renvoize 70558 (K); Panamerican Hwy. Riobamba—
Ambato, km 22, 01°30’S, 78°42’ W, 3500 m, 11 Oct
1985, S. Laegaard 55400 (AAU, QCA); S side of
Volcano, 4300 m, 1929, Rimbach 97a (US); Urbina
— paramo on E flank of Mt. Chimborazo, 3600 m,
A.S. Hitchcock 22019 (K, NY, US); S slope, 3800 m,
Fagerlind & Wibom 934bis (S); Fagerlind & Wibom
932 (S); Dec 1890, Sodiro s.n. (QPLS); 4600 m,
E. Asplund 8399 (S); Nudo deIgualata—Sanancajas,
3600 m, Acosta-Solis 21187 (US); km 18.2 ESE of
Lago Colta on road to Pallatanga, 3700 m, 21 May
1990, PM. Peterson 9228, E.J. Judziewicz, R.M.
King & PM. Jorgensen (MO, QCA, QCNE, US);
Prov. Riobamba, praed Toledo, Sep 1891, Sodiro
s.n. (P); Mun. Guaranda, W side of volcan Chim-
borazo, road to Refugium, 01°28°S, 78°52’ W, 4100
m, 5 Jul 1999, D. Stancik 3163 (PRC, QCA); Mun.
Riobamba, volcan Chimborazo, around upper Refu-
gio, 5000 m, 20 Sep 2000, D. Stancik 3706 (PRC,
QCA); sector Cruce de los Arenales, 01°28°15’S,
78°54'6" W, 4300 m, 20 Sep 2000, D. Stanctk 3718
(PRC, QCA). Imbabura: Paramo on the N side
of Nevado Cayambe, 0°03°07°N, 77°59'27°W, 17
Aug 2004, Sklenar 8067, 8115 (PRC). Morona—
Santiago: Hda. Hargualla—Hda. San Eduardo, way
to Parque Nacional Sangay, 02°0.25’S, 78°27'°W,
3700 m, 19 Jul 1999) D: Stanctk 3323 (PRC,
QCA); Parque Nacional Sangay, a confluence of
Quebrada Plazabamba Chico with Q. Plazabamba,
02°0.48’'S, 78°25.4°W, 3600 m, 21 Jul 1999,
D. Stancik 3341 (PRC, QCA). Pichincha: Cerro
Antisana, W. Huagrahialina Campamento, 4200 m,
Black 278, 279 (AAU); 4600 m, E. Asplund 17338
(NY, S); 00°32’S, 78°12’W, type locality, 4030 m,
S. Laegaard 102848 (AAU); between Antisana and

146

Sincholagua, FE. Asplund 3648 (US); Rucu Pichin-
cha, NE slope, 00°10’S, 78°34’ W, 4300-4500 m,
Sklenar & Kosteckova 198 (AAU); Lloa—Guagua
Pichinha road, km 11, 00°12’S, 78°35’W, 4310 m,
9 May 1992, S. Laegaard 102743 (AAU, QCNE);
around Refugium, 00°12’S, 78°35°W, 4550 m, S.
Laegaard 102758 (AAU); 4600 m, FE. Asplund 7397
(K, NY, S, US); Pichincha, Cotocollao, Sodiro s.n.
(QPLS); 3200 m, EF. Asplund 6620 (S, US); Paramo
de Guamani, 00°18’S, 78°14’ W, 3750 m, S. Lae-
gaard 51386 (AAU, QCNE); 00°20’S, 78°14’W,
3700 m, S. Laegaard 19598 (AAU, LOJA, PRC,
QCNE); 3600-3800 m, E. Asplund 8721 (S);
4050 m, Barclay & Juajibioy 8905 (COL, US);
Andium Quitensium, Jameson 230 (P); Mun. Pifo,
Paramo de Guamani, 00°20’S, 78°12°W, 4000 m,
19 Jun 1999, D. Stancik 3014 (PRC, QCA); 3700 m,
D. Stancik 3020 (PRC, QCA, US); D. Stancitk 3001
(PRC, QCA). Tungurahua: Volcan Chimborazo,
SW of San Fernando, 3500 m, 23 Jul 1959, Barclay
& Juajibioy 166 (COL, US); above Mocha, 3300
m, E. Asplund 7872 (S).

56b. Festuca vaginalis subsp. cayambae Stancik,
Folia Geobot. Phytotax. 39(1):107. 2004.
Type: Ecuador. Pichincha, along road to
Volcan Cayambe, near entrance to National
Park, 78°04’ W, 00°03’S, 3550 m, 1 Mar 1988,
S. Laegaard & S.A. Renvoize 70488 (holotype:
AAU!; isotypes: QCA!, QCNE!, K!).

Culms 20-30 cm tall, not robust. Leaf blades
0.4—0.5(—0.6) mm wide. Panicles pale-green.

Leaf blade anatomy.—Cross-sections with 7
vascular bundles.

Festuca in South American Paramos

Additional specimens examined. ECUADOR.
Bolivar: km 60.4 SW of Ambato on Hwy to Guar-
anda, 4090 m, PM. Peterson & C.R. Annable 8984
(QCNE, US); 01°25’S, 78°52"W, 3950 mi, 15 Fes
1999, S. Laegaard 19589 (AAU, PRC); km 66.5,
3760 m, 3 May 1990, P.M. Peterson & C.R. Annable
9000 (K, MO, QCA, US); km 33, 01°18’S, 78°48’ W,
3700 m, 1 Mar 1992, S. Laegaard 101505 (AAU,
QCNE); Cruce de los Arenales, 01°25’S, 78°54’ W,
4160-4200 m, S. Laegaard 101493 (AAU, PRC).
Canar: Panamerican Hwy. ca. 5 km S of Cafiar,
02°33'S,78°56'W, 3500 m, S. Laegaard 105123
(AAU); 3350 m, S. Laegaard 101828 (AAU). Chim-
borazo: Mun. Guaranda, pasture around Vereda
Rio Colorado, 4200 m, 2 Jul 1999, D. Stancik 3182
(PRC, QCA, W). Cotopaxi: Mun. Chaupi, NE slope
of volcan IIliniza Norte, 00°37°59"S, 78°42°42”W,
4000-4050 m, 12 Oct 2000, D. Stancitk 4033, 4035
(PRC, QCA). Pichincha: Hacienda Antisana and
Mt. Sincholagua, E. Asplund 8648 (S).

EXCLUDED NAME

Festuca sublimis forma vivipara St.-Yves, Can-
dollea 3: 255. 1927. Type: Perms Pucarde
Ladrillos, entre Pacasmayo et Mayobamba,
3400 m, Sttibel Fi. Peru 30 (syntype: B?);
Ecuador. Ilizna, Tisiche, Stiuibel Fl. Ecuador
301 (syntype: B?).

Comments.—The status of this taxon is in
question since we were unable to locate any of
the syntypes. Alexeev (1986) cited this taxon as a
synonym of F. asplundii but did not indicate that
he saw any of the original material.

SCANNING ELECTRON MICROGRAPHS
OF LEAF EPIDERMAL SURFACES

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ae .
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Mint Di

2

P93. oa

|
:
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cies

Figure 72. Leaf blade surfaces. A-D. Festuca amplissima subsp. amplissima. A. Abaxial epidermis with ribs, densely covered with
prickles. B. Abaxial epidermis, detail view of long cell with sinuous walls, rounded silica bodies, and prickles. C. Adaxial epidermis
with long cells and regularly occurring stomata. D. Adaxial epidermis with long and short (silica bodies) cells, stomata (surrounded
by wax), and prickles on the margin. E & F. F. coromotensis. E. Abaxial epidermis with regular ribs. F. Abaxial epidermis with detail
of oval (sinuous) silica bodies and prickles. A & B, Barclay 7033 (COL); C & D, Liebmann 6110 (C); E, F, Stancik 4180 (PRC).

Festuca in South American Paramos

A See

am

wiht ets Wing
cosalintedidl gece Ea

mpeg gor”

ie ill iy pas
|

4 Rawr tls
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i
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Figure 73. Leaf blade surfaces. A & B. Festuca coromotensis. A. Adaxial epidermis with ribs, sparsely covered with short macro-
hairs. B. Adaxial epidermis, detail view of silica bodies, stomata and macro-hairs. C—F. F. elviae. C. Abaxial epidermis with long
cells and regularly occurring silica bodies. D. Abaxial epidermis, detail view of silica bodies with short cork cell. E. Adaxial epi-
dermis with regular ribs and short macro-hairs. F. Adaxial epidermis, detail view of stomata and cell walls covered by fine crystals

of wax. A & B, Stancik 4180 (PRC); C-E, Stancik 4178 (PRC); F, Stancik 4171 (PRC).

149
B. Adaxial

detail view of stoma and prickle. D. Adaxial epidermis, detail
F. Abaxial epidermis with detail of circular

Abaxial epidermis with regular ribs

E & F, Stancik 4286 (PRC).

. Adaxial epidermis,

Festuca in South American Paramos

>

Festuca flacca. A. Abaxial epidermis with small ribs densely covered with prickles.

Leaf blade surfaces. A-D.
epidermis covered with prickles and stomata. C

view of wax crystals. E &

silica bodies. A—D,

F. F. guaramacalana. E

Cik 4180 (PRC);

Stan

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74

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Figure

Festuca in South American Paramos

"
%
1
if

Fo An

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Figure 75. Leaf blade surfaces. A & B. Festuca guaramacalana. A. Adaxial epidermis with ribs, covered with macro-hairs. B. Adaxial
epidermis, detail view of macro-hairs. C-F. F. sodiroana. C. Abaxial epidermis with small ribs and occasionally occurring prickles.
D. Abaxial epidermis, detail view of long and short (silica bodies) cells. E. Adaxial epidermis with regular ribs and sparse short
macro-hairs. F. Adaxial epidermis, detail view of stoma. A & B, Stancik 4286 (PRC); C—F, Stancik 2632 (PRC).

Festuca in South American Paramos

LES TF WRU AY CR 2004

SEWN I7 BU AV CR 2005

Figure 76. Leaf blade surfaces. A-D. Festuca ulochaeta. A. Abaxial epidermis with small ribs. B. Abaxial epidermis, detail view
of prickle and silica bodies with cork cells. C. Adaxial epidermis with small ribs and regurally occurring stomata and short macro-
hairs. D. Adaxial epidermis, detail view of stomata and macro-hairs. E & F. F. caldasii. E. Abaxial epidermis densely covered with

prickles. F. Abaxial epidermis with detail of oval (sinuous) silica bodies and prickles. A—D, Stancik 4179 (PRC); E & F, Laegaard
20405 (AAU).

152 Festuca in South American Paramos

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Figure 77. Leaf blade surfaces. A & B. Festuca caldasii. A. Adaxial epidermis with small ribs and regurally distributed stomata
and macro-hairs. B. Adaxial epidermis, detail view of stomata covered by wax. C-F. F. reclinata. C. Abaxial epidermis with long
cells and regularly occurring stomata and slender prickles. D. Abaxial epidermis, detail view of prickles and stomata. E. Adaxial
epidermis with small ribs; regurally occurring prickles and stomata. F. Adaxial epidermis with detail of prickles and wax covering
cell walls. A & B, Laegaard 102535 (AAU); C-F, Cuatrecasas & Barriga 9970 (COL).

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epidermis with pronounced ribs densely covered with prickles. C. Adaxial epidermis with detail view of stomata laying between

two ribs. D. Adaxial epidermis with detail view of prickles and waxes covering cell walls. E & F. F: arundinacea. E. Abaxial
epidermis with regular ribs and rows of prickles. F. Abaxial epidermis with detail view of stomata and silica bodies. A-D, Wood

Figure 78. Leaf blade surfaces. A-D. Festuca woodii. A. Abaxial epidermis with ribs, densely covered with prickles. B. Adaxial
5254 (COL); E & F, Stancik 322] (PRC).

Festuca in South American Paramos

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Figure 79. Leaf blade surfaces. A & B. Festuca arundinacea. A. Adaxial epidermis with rows of stomata between small ribs.
B. Adaxial epidermis, detail view of long cells interrupted by short cells (silica bodies and cork cells). C—F. F. quadridentata.
C. Abaxial epidermis covered by prickles. D. Abaxial epidermis with detail view of prickles and short cells. E. Adaxial epidermis

with pronounced ribs densely covered with prickles and wax. F. Adaxial epidermis with detail of wax and prickles. A & B, Stancik
3221 (PRC); C-F, Laegaard & Sklenar 20308 (PRC).

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C. Abaxial epidermis with detail view of silica bodies and prickles. D. Adaxial

stomata and silica bodies. E. Adaxial epidermis. F. Adaxial epidermis with detail view of
Laegaard & Sklenar 20308 (PRC); B-F, Stancik 4263(PRC).

Festuca in South American Paramos

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80. Leaf blade surfaces. A. Festuca quadridentata. Abaxial epidermis with detail view of wax. B—F. F. venezuelana. B. Ab-

, stomata and silica bodies. A

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Festuca in South American Paramos

Figure 81. Leaf blade surfaces. A-F. Festuca fragilis. A. Abaxial epidermis densely covered with prickles and silica bodies. B,
C. Abaxial epidermis with detail view of silica bodies and prickles. D. Adaxial epidermis with ribs and macro-hairs. Stomata on
the bottom of ribs. E. Adaxial epidermis with detail view of macro-hairs and stomata. F. Adaxial epidermis with detail view of wax
covering cell walls. A-C, Wood 5259 (COL); D-F, Stanctk 4248 (PRC).

Adaxial epidermis, detail view of
Brade & Kuhlman 15631 (PRC);

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Figure 82. Leaf blade surfaces. A-E. Festuca fimbriata. A. Abaxial epidermis with ribs. B. Abaxial epidermis, detail view of silica
bodies, prickles and stomata. C. Adaxial epidermis with pronounced ribs covered densely with prickles. D. Adaxial epidermis,

stoma and wax on the cell walls. F. F rubra. Abaxial epidermis with long and short cells. AE,

detail view of ribs with prickles and silica bodies on the top of ribs, stomata between the ribs. E.
F, Stanctk 3457 (PRC).

Festuca in South American Paramos

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Figure 83. Leaf blade surfaces. A & B. Festuca rubra. A. Adaxial epidermis with ribs covered with macro-hairs. B. Adaxial epi-
dermis, detail view of macro-hairs and stomata on the base of the ribs. C & D. F andicola. C. Abaxial epidermis of the involute
blade with macro-hairs on the margin. D. Adaxial epidermis, detail view of macro-hair and wax on the cell walls. E & F. F soukupii.
E. Abaxial epidermis of the involute blade with silica bodies and macro-hairs on the margin. F. Adaxial epidermis with macro-hairs
on the ribs. A & B, Stancik 3457 (PRC); C & D, Laegaard 52807 (AAU); E &F, Stancik 3455 (PRC).

Festuca in South American Paramos

Figure 84. Leaf blade surfaces. A-D. Festuca azucarica. A. Abaxial epidermis with long and short cells. B. Abaxial epidermis,
detail view of rounded silica bodies. C. Adaxial epidermis with ribs covered with long macro-hairs. D. Adaxial epidermis, detail
view of stomata at the bottom of rib (middle part of this image). E & F. F. colombiana. E. Abaxial epidermis with long and short
cells. F. Adaxial epidermis, macro-hairs on the lateral walls of the ribs. A-D, Stancitk 3405 (PRC); E & F, Stancik 2561(PRC).

Festuca in South American Paramos

Figure 85. Leaf blade surfaces. A & B. Festuca colombiana. A. Adaxial epidermis, detail view of stomata and macro-hairs. B. Adaxial
epidermis, detail view of wax. C & D F. dasyantha. C. Abaxial epidermis with small ribs, long and short cells. D. Adaxial epidermis
with ribs covered with long macro-hairs. E & F. F. hatico. E. Abaxial epidermis with small ribs. F. Abaxial epidermis, detail view
of silica bodies. A & B, Stancik 1289 (PRC); C & D, Laegaard 101260 (AAU); E & F, Stancik 4289 (PRC).

Festuca in South American Paramos

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Figure 86. Leaf blade surfaces. A & B. Festuca hatico. A. Adaxial epidermis with ribs, densely covered with macro-hairs. B. Adaxial
epidermis, detail view of macro-hairs and wax deposits. C—F. F: laegaardii. C. Abaxial epidermis with small ribs. D. Abaxial epider-
mis, detail view of long and short cells (primarily silica bodies). E. Adaxial epidermis with ribs covered with macro-hairs, stomata
on the base of the ribs. F. Adaxial epidermis, detail view of stomata. A & B, Stancik 4289 (PRC); C—F, Stancik 3365 (PRC).

Festuca in South American Paramos

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Figure 87. Leaf blade surfaces. A & B. Festuca pilar-franceii. A. Abaxial epidermis, detail view of silica bodies. B. Adaxial epider-
mis, detail view of ribs covered with long macro-hairs. C—F. F. procera. C. Abaxial epidermis with small ribs. D. Abaxial epidermis
with long and short (silica bodies) cells. E. Adaxial epidermis with ribs covered by long macro-hairs. F. Adaxial epidermis, detail
view of macro-hair and wax deposits. A & B, Stancitk 241 (PRC); C—F, Stancik 4106 (PRC).

Festuca in South American Paramos

Figure 88. Leaf blade surfaces. A & B. Festuca toca. A. Abaxial epidermis with small rib and silica bodies. B. Adaxial epidermis,
detail view of rib covered with macro-hairs, stomata are on the base of the rib. C-F. F. asplundii. C. Abaxial epidermis sparsely
covered with prickles. D. Abaxial epidermis, detail view of prickles and silica bodies. E. Adaxial epidermis with ribs covered with
macro-hairs. F. Adaxial epidermis, detail view of macro-hairs and stomata on the base of the rib. A & B, Stancik 1418 (PRC); C-F,
Stancik 3611 (PRC).

Festuca in South American Paramos

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Figure 89. Leaf blade surfaces. A & B. Festuca boyacensis. A. Abaxial epidermis, detail view of silica bodies. B. Adaxial epider-
mis, detail view of stomata and macro-hairs and cell walls covered with wax. C & D. F. carchiense. C. Abaxial epidermis, view of
long and short cells and sparse prickles. D. Adaxial epidermis with ribs covered with short macro-hairs. E & F. F. chimborazensis

subsp. chimborazensis. E. Abaxial epidermis with small ribs. F. Abaxial epidermis, detail view of silica bodies. A & B, Stancik
2166 (PRC); C & D, Laegaard 101716 (AAU); E & F, Stancik 4033B (PRC).

Festuca in South American Paramos

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Figure 90. Leaf blade surfaces. A & B. Festuca chimborazensis. A. Adaxial epidermis with ribs with stomata between long cells,
sparsely covered with short macro-hairs. B. Adaxial epidermis, detail view of prickles (margin of the leaves), macro-hairs and
stomata. C—F. Festuca cleefiana. C. Abaxial epidermis. D. Abaxial epidermis, detail view of silica bodies. E. Adaxial epidermis

with ribs covered with macro-hairs. F. Adaxial epidermis, detail view of stomata on the base of the ribs. A & B, Stancik 4033B
(PRC); C-F, Stancik 2020 (PRC).

Festuca in South American Paramos

Pa

Figure 91. Leaf blade surfaces. A-D. Festuca cocuyana. A. Abaxial epidermis with long and short cells. B. Abaxial epidermis, detail
view of silica bodies. C. Adaxial epidermis with several ribs covered with macro-hairs, sometimes with silica-bodies or stomata
between long cells. D. Adaxial epidermis with detail view of macro-hairs, stomata and wax. E & F. F. cundinamarcae. E. Abaxial
epidermis. F. Abaxial epidermis with detail view of silica bodies. A-D, Cleef 9078 (COL); E & F, Stancik 2114 (PRC).

Festuca in South American Paramos

Figure 92. Leaf blade surfaces. A & B. Festuca cundinamarceae. A. Adaxial epidermis with ribs densely covered with macro-hairs.
B. Adaxial epidermis, detail view of stomata at the base of the ribs. C-F. F. densipaniculata. C. Abaxial epidermis densely covered
with prickles. D. Abaxial epidermis with detail view of prickles and wax. E. Adaxial epidermis with regular ribs covered with
macro-hairs. Stomata on the base of ribs (right margin of the image). F. Adaxial epidermis with detail of macro-hairs and stomata.
A & B, Stancik 2114 (PRC); C-F, Asplund 8397 (S).

Festuca in South American Paramos

Figure 93. Leaf blade surfaces. A-D. Festuca glumosa. A. Abaxial epidermis of the involute leaf with rows of prickles. B. Abaxial
epidermis, detail view of silica bodies and prickles. C. Adaxial epidermis with ribs covered with short macro-hairs. D. Adaxial
epidermis with detail view of macro-hairs and stomata. Cell walls covered with wax. E & F. F. glyceriantha. E. Adaxial epidermis

with regular ribs covered with macro-hairs. F. Adaxial epidermis with detail of wax. A-D, Stancik 4180 (PRC); E & F, Peterson
14936 (US).

Festuca in South American Paramos

LES S7 WES PS CF 2005

Figure 94. Leaf blade surfaces. A & B. Festuca imbaburensis. A. Adaxial epidermis with ribs covered with macro-hairs. Cell walls
covered with wax deposits. B. Adaxial epidermis, detail view of wax, stomata and macro-hair. C-F. F monguensis. C. Abaxial
epidermis of the involute leaf, with small ribs and prickles on the margin. D. Adaxial epidermis with detail view of prickles and
silica bodies. E. Adaxial epidermis with regular ribs covered with macro-hairs. F. Adaxial epidermis with detail of macro-hairs and
stomata (at the bottom of ribs). A & B, Stancik 4098 (AAU); C-F, Stancik 2025 (PRC).

Festuca in South American Paramos

ips

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Piso ‘.

Figure 95. Leaf blade surfaces. A-D. Festuca nereidaensis. A. Abaxial epidermis with long and short cells. B. Abaxial epidermis,
detail view of silica bodies. C. Adaxial epidermis with ribs covered densely with macro-hairs. D. Adaxial epidermis with detail
view of macro-hairs. Stomata on the bottom of the ribs (not seen). E & F. F. oroana. E. Adaxial epidermis with regular ribs covered
densely with macro-hairs. F. Adaxial epidermis with detail view of macro-hairs and dense coverage of wax. A-D, Stancik 3401B
(PRC); E & F, Laegaard 70309 (AAU).

Festuca in South American Paramos

Figure 96. Leaf blade surfaces. A-D. Festuca sanctae-martae. A. Abaxial epidermis with small ribs and prominent long and short
cells. B. Abaxial epidermis, detail view of silica bodies. C. Adaxial epidermis with small ribs and short macro-hairs. D. Adaxial
epidermis with detail view of macro-hairs and stomata between long cells. E & F. F. subulifolia. E. Abaxial epidermis with detail
view of prickles and silica bodies. F. Adaxial epidermis with small ribs covered with macro-hairs and stomata at the bottom between
ribs. A-D, Cuatrecasas & Castafieda 24532 (COL); E & F, Stancik 2914 (PRC).

172 Festuca in South American Paramos

Figure 97. Leaf blade surfaces. A—C. Festuca tolucensis subsp. tolucensis. A. Abaxial epidermis densely covered with prickles
and silica bodies. B. Adaxial epidermis with ribs covered with macro-hairs, stomata at the bottom between the ribs. C. Adaxial
epidermis, detail view of stomata and macro-hairs. D. F. vaginalis. Adaxial epidermis with ribs covered with short macro-hairs and
deposits of wax on the cell walls. A-C, Stancik 4279 (PRC); D, Laegaard 101559 (AAU).

Festuca in South American Paramos Ls

Fig.
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Appendices
INDEX TO FIGURES
co. LTS ENG OL SVTOON ONT AVIS ST (1 en ee Pe RR 3)
2. Distribution map of Festuca amplissima, F: coromotensis, and F. flACCQ .....c..cccccccecccceseeeeseceesees 20
rae SAE OMEOIELCSTICO)COLOMOLCIISIS: vin nceee.ens05sre<cet cab went ces oceu thnks \orlsua lesa sos esestwtsatovesavect | dencedhess a) |
Maa sA Tet MODE IMC STIECO CN VIO ab. fe acca ao a5, Ons ac se deta ie eaue os Sataalcebade vtwgisceedic ne ccesaustongualedecdeldleebduass 23
MaDe UIOMOM CSIC A ACCOM cn. coccancksseanckosaneshnn-ceveeehowie eh ate tededs sevtunlsas dase bd bicavidebGueabbiuneabaleas DS
6. Distribution map of Festuca sodiroana, F. elviae, and F. guaramacdlan .......cccccccccccccceeesceeeeeee 26
RUS MALOMEON MEST CA SUGTAIMOACGIGNA .....0......csckccessacseseecdchleidsecsiectacsesseseosvonsevascsuasecusededsedeneares 27,
PMNS SMO Le MCSTUCO SOGIOONG -ciasn exca.csansnnestusnessnatesduccsavtesieeecare <thasdpedvcbin le lSsbaessacetsabdbesscodeeees 29
PMNS Obi OMUOLIIESTUCE TOVATICHISIS »...cce.s0c-casaueseacdeoos:cesesscceisdebdicatdorebueesadiesssceesavondctdasaicbavbeceddsteas 32
MBN RES UALTO ME OMA CSTIICH WI OCI OCLC. eiecere ic. a.0pncscesunedebssadeschcacs-ntaQdestvoudvesevluchéssbevetiavidethdbessacdobees 34
11. Distribution map of Festuca ulochaeta, F: tovariensis, and F) CAIAASTI........0..ccccccccccccccceesteceees 39
Mem NEA std OMO MTC STU CO COLAASIU 52.0 c2cee.anoh csaeiantecteacdecceeaeetviagicasausenicendvantessecveacesiwslestadecessacedoevees 37
PMB SALOMON CSE CHT CCIM oinn.ccsccanniiedaenn ad eteveads neni sats oisyuescudeesahsceasdedesbodedavsevestie¥eoteuneieedeoleen 39
14. Distribution map of Festuca pratensis, F’ woodii, and F. reclinata............cccccccccsseessseesesssesseeees AQ
Mem Mats AIO TROD CSIIICH WOOL sec ccseks orien anenuie sane cdessgOoes dedad sinseicioadeauuaacUnssedsddsuvnas eda athds Séedenpeboacandass 4]
Mem sitrANIO MTOM CSIUCA ATUNGINACCOE.. vvcos.casesancdeos auncucevandsu bavassieresatcndusabeahesennsGoleadsssabeethabueceebess 43
17. Distribution map of Festuca dichoclada, F: arundinacea, and F: venezuelana..............c000000000 45
PSUS iTeAN OPO CSTICH GUAGTIGENIGLG 5.2.0. <..0c005.2nccctiediecasen gece sped len caveonddndés ohuceduasndissdesiactsosdencdeceee 47
ees rA FIOM OMMCSTUCH VENCZUCIONG :.....1.0:0ncn-ncadestsunaesaaedscstedbdeesvessachadiedaestasoensevenatidedsuceteciad Mees 49
PM sore ONM CSCO [PASTS races cn ezcccaco: vennevaedeceseuctnde signe csduastessses nddpbacsts dese chelsuasseechene dacdlhes 50
2 Disinbution map of Festuca quadridentata, F fragilis, and F. rubra ...........00c0csesseessoocdessencseees Sil
i WITUSISEIIMCIL CLE) ASL LCTOD 1/00) 101 (0 yn sire REP oe: Ee ee eee eee eT ee ee 58)
Poe Disiabuvion map OF Festuca fimbriata, and F. ANGICOIlA...2:..c....00s.cisenesesnsscveonntuvavanccntsateanacetceens 54
Pe IS PALI OMNON II CSICO GNGICON Os scctasnct ats: Jos ace these Sonttceiadesbadeedn te {svancs vesudhadevarcadabaveterasbacdenbes ons 56
PE Sem AI OI OMe Il CSPI CO GOTO rec ect ae cde oeistice etatns done Ssdnedeie ast sab toisaive svc sdvaseus ndawovadeneneeeed sabtele vebdvons 58
Pia TS Deol OMVONG!d CSTICOSOUKU DU er te ain. tea diets dna nceebesarceuts dandeuusiuad, sestbibiese. Sougbdnteds ve bead udinedesedeeees 60
27 Distribution map of Festuca soukupil, chitagana, and F. hatico::....2.1..1c0ccisaeaseoensesecesoneee® 61
PMNS PRALIOMEON ESCA AZUCORIC Cia setae. 2h, deteetsc Ue ugesbsetacagesnduetie oan scdabdaik gebnadevedseeedesteabesdoenbeew ed 64
29. Distribution map of Festuca colombiana, F: dasyantha, F- azucarica, and F: laegaardi.......... 65
SOM SthabOM Ole eSTICO CHIGOANO 4.09%. Scenics doe hcvdcanscuaduesten¥-cot isoesuos ds 0 heysase tosh oe tadsatet es tasien soot tees 67
SPSS TANIOMOMMCSINCA COLOMDIGNGE sax.c.02.00sscneanneiesesdbos oaiee owds scents ones oceoassebatinedapscueee-Malscdosapeeed 68
SP MUSt KA OMROMMCSTUCA MASVGNENG si J ace. 9... cacate ab dusesabeaeiss <oth aesgeenua due talsuaee Svaedeaebaei tes, toate tance eek TA
Seeds treatlomOlelCSMICONOTIEO.. 8 ecco5: autor Saticnn sel cauesedaited tues «Ue iiontar bela stigeisb aepeptoactedsoeescauthotanereneas 1)
SA MUStratOmM Of MEStUCA 1ACCOOUGID cathe cee cake ection, acasa tees senses Sekuest taoce chasse aquarabanedzwdyedsctasinog o-updeoes 74
SP MUStrA nM OMUOL CSCI PIU] B= fNOMCCLE am... eoc sa asea bem sandeesechsesce aes. c0sbi sede sstarabhuedse nasete- tasted meee! 76
Soolilustration Of MeStUCa DKOCCHG: df sracansces incu ioe- Sic tate tide teepestie basses seedosanaals esas ones peene eaReenae tenes 78
37. Distribution map of Festuca procera, F: pilar-franceii, and F. DOYACeNSIS ..........0cccccereeeeereees 79
SS MMMSHAatOM Of MESHHCO TOCA s.. .1. 2:21. owe Denies chit Bye. demands. teueiaten shes tele Resets A oes ie Baas 82
39) Distributionanap of Kesticaasplundit, H toca, and Fx COCUVGNG.N <x. cwansss-eansntenadcebs-teede-4<cteeae<s 83
AO. ustration Of HestucaGs plum ii is, dss: osQrwepeaccraeien Seat catous vesaiisg ae tea see ae RE Maggi. soe 85
a Wistrationvol Festuca DOV ACCNSIS a sscsrs. 2s Lace mxeanion cipeaasascteeceeteie te socks MEER eet oee tes As 89
AD Mlustration: of Festited: CAnrChiCns €: dike sya ssitat ace aia pekabo, Cans meee at eth bye Echoes 90
a3, dilustration- ol 1 estica ChimbOraZzensis x. 0% Bax. avaninte, a ckouseoen ten teas eed eens phoee eee osheee ale 04s o2
44. Distribution map of Festuca chimborazensis, F- carchiense, F: cleefiana, and F: dinirica........ 93
45 ulllustration Ol F CstuCa- Chita. seancmn. Meee a seem rds. Mere eGen, tay. Cenc deN ini, shad teks tesgse)anctde eu: 96
AG. Mtustrationiot FestucasClecpiands. WN yess teate aeuew oe daeet tat, deeetaain he Sede meeenes eked eate tes. beaks cae: oF
AF aUStratlOnvOl TT ESTUCGCOCUY GN Gicei crs, epee, Cheenaci aaah seven hey eames een tad. 20, Osa 99

174

Fig.
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Fig.
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Fig.

Festuca in South American Paramos

48. Illustration. of Festuca cundindmaréde sii.2i0.8. ct de ete a 101
49, Distribution map of Festuca glumosa, F: cundinamarcae, F- chita, and F. turimiquirensis.... 102
50. Illustration of Féstiica densipanieulat]a os, .édeccesecscdinescdin cavisiees ideas ac tee ee eds 103
51. Ilhustration of FP@stuCa GiniriGa seccissccccsveis cscs. Qi dactea ee ites a 105
52. Distribution map of Festuca glyceriantha, F: holubii, F. densipaniculata, and F- monguensis ....

bésedncctassocusdavesddeulsaton ss SQbaehagadaaadeedwus sNtuandseeeeskonawwadedte pad even seeeee eaae eee ata Lane STEM Be Rams aE ahs (aaa a 106
53. Ilustration Of Festuca SIUMOSG «0.003: essscsecis.catenetaasgedhe ts cutaae tenadeteahestateten seeder 107
54. Illustration of Festuca @lyCerigntha sssccscecvescsccasectaneeticasccetecenasectetaaiceen athe 110
55. Uhistration of Festuca NOMI «.ccccccccsseseaes.s.sicaseatabeas sins veis edecaneaaa ates ta 112
56. Illustration of Festuca hUAMACHENSIS sz :.sccccssisicecssevaaseteessaasteccnaucocecats caesacetotedaasl ety eee 114
57. Distribution map of Festuca huamachucensis, F: imbaburensis, F- nereidaensis, and F- sanctae-

IMT IGG 5.4.4 sxsssszacaccdonsascasadhten saatesanicadees ianeasbaghdadaasa sTEd eG Ode aRReE eRe 115
58. Illustration of Festuca imbabDurensis «. ..cc.ci.cccesecccccsessascceatsscseneds Gesteeanddeecsk ds pase ee 116
59. Illustration of Festuc@ MONGUCNBIS ....5220..0060.cecsesesedeaca betaes azian das sedge aaeh dead 118
60. Ilhustration of Festuca Nereidaensts.......s100cccscastectecctaatetacessseaiiesd.vaceeissiasteecetiedee ee 120
61. Hhustration. of PestuCa OFOGNG ..c.sssscccchscssiec.ssasacenscevdGecsneseasia ihenadacctagitucedene 121
62. Illustration of Festuca parciflOr[g.....cscccciss..ccssssaedestesissaseectsatatiatiiaaeshecietanst ee 123
63. Illustration Of Festuca rENVOIZEL «.22..:.ccceccscc.cccvseceeesteusvedéesaaestetensstazehsanedeii ances tee ei 125
64. Distribution map of Festuca vaginalis, F. parciflora, F: oroana, and F. renvoiZii..............00+ 126
65. Illustration of Festuca SANCtAC-MGP1GE ......i..cccesccccesceseecccetsatescacsasanes candssuesiaiiasan ee 127
66. Illustration of Festuca suUbUlifOlidci.cccsscccccccccccccssesguetctietecsiassssaecsecdtéiasiseactele cheese 128
67. Distribution map of Festuca tolucensis, F. subulifolia,and F. SUMAPAn .........eseccceeseeeeesseeeeess 125
68. Illustration Of Festuca SUMADGNG ..icsissssicccccessccsiesssecsasdscistbasiieasacderiasigivalodial eee i337
69. Illustration of Festuca toluCensts ..scccicccscdss.ces.sessscsncassams naetetaseditasadaean nies 138
70. Illustration of Festuca turiMmiqQuirensts ....00...iscseccsvesdesesectessessttdisseraateastiees ee 142
71. Tlhustration of Festuca vaginalis .scsc.cccissasscccsssanstdecsssessestessecandeacecdoncccaessnagdestesehe ee 144
72. Leaf blade surfaces of Festuca amplissima and F. COPOMOEENSIS ........ccsccccccccceeseesssceeeecesensees 147
73. Leaf blade surfaces of Festuca coromotensis anid F. €lvidé.....:..cis.ssiccsssssecceasnssessseee 148
74. Leaf blade surfaces of Festuca flacca and F. ZUArAMACAIANA.......11cccccccsssesecssssssscssssnstecessnsnees 149
75. Leaf blade surfaces of Festuca guaramacalana and F. SOdiIvOANG ........cccccccccessssssseceeseesnsnsnsees 150
76. Leaf blade surfaces of Festuca ulochaeta and F. Caldasit ....:..:ic.cicssssecsssescesssastes ee IS]
77. Leaf blade surfaces of Festuca caldasii and F. reclindtd «...2.0:..::.0..:00:0::).4.0 ee 152
78. Leaf blade surfaces of Festuca woodii and F. Arundinacea .........ccccccccssesscccsessstccccesssteseenncczes 153
79. Leaf blade surfaces of Festuca arundinacea and F. quadridentata........ecsccccccscccssssseceesseeeeeees 154
80. Leaf blade surfaces of Festuca quadridentata and F. veneZuelan ........ccccccsssecccseesseceeeennneceees 155
81. Leaf blade surfaces of Festuca fragilis..s.ccccssccscacssceew.e2t sch tetictesihctanaee ate. ee 156
82. Leaf blade surfaces of Festuca fimbriata and F. rubra... ....cicictectasstis cst 137
83. Leaf blade surfaces of Festuca rubra, F- andicola, and F; SOUKUpIi...........0.ccccccccesseeeesseeeeeeee 158
84. Leaf blade surfaces of Festuca azucarica and F. COLOMDIONG...........ccccctessesscscsssneccccssuccnceneeate 159
85. Leaf blade surfaces of Festuca colombiana, F- dasyantha, and F. hati ..........cccccscccccsesseeeees 160
86. Leaf blade surfaces of Festuca hatico and F laegaardii 2.2.0.1). uitectet:tinssec 161
87. Leaf blade surfaces of Festuca pilar-franceli and F. PrOCerd ........cscccsessecccesscceesnnccessnneeeeeseeees 162
88. Leaf blade surfaces of Festuca toca and Fo GSpluniit c2cccccczesteesctstecaetsteestaseee te 163
89. Leaf blade surfaces of Festuca boyacensis, F: carchiense, and F- chimborazensis..............-+- 164
90. Leaf blade surfaces of Festuca chimborazensis and Festuca Cleefidnd ..........ccccsccesseceeseeeeseees 165
91. Leaf blade surfaces of Festuca cocuyana and F: cundinamarcde .i220.2.0051:21.2-2 166
92. Leaf blade surfaces of Festuca cundinamarceae and F: densipaniculat ..........s:cccccccesseeeereees 167
93. Leaf blade surfaces of Festuca glumosa and F elyCeriantha.......0.:..scc..cs.c1esteaescesesieeee ee 168
94. Leaf blade surfaces of Festuca imbaburensis and F. MONQUENSIS ........1:cccccesscesseceseeeseeeneeeenees 169
95. Leaf blade surfaces of Festuca nereidaensis and F. Ofroana 2... ieee 170
96. Leaf blade surfaces of Festuca sanctae-martae and F. SUDULIfOLA ........ccccccescccsseeseeeseeneeeseeees 171
97. Leaf blade surfaces of Festuca tolucensis and F. vaginalis... 22 172

Festuca in South American Paramos ‘735

ALPHABETICAL LIST OF ACCEPTED SPECIES AND SUBSPECIES

Number in parentheses corresponds to the number of the species in the taxonomic treatment.

Festuca amplissima subsp. amplissima (1a)

Festuca amplissima subsp. sierrae (1b)

Festuca andicola (19)

Festuca arundinacea (12)

Festuca asplundii (31)

Festuca azucarica (22)

Festuca boyacensis (32)

Festuca caldasii (9)

Festuca carchiense (33)

Festuca chimborazensis subsp. chimborazensis
(34a)

Festuca chimborazensis subsp. micacochensis
(34b)

Festuca chita (35)

Festuca chitagana (23)

Festuca cleefiana (36)

Festuca cocuyana (37)

Festuca colombiana (24)

Festuca coromotensis (2)

Festuca cundinamarcae (38)

Festuca dasyantha (25)

Festuca densipaniculata (39)

Festuca dichoclada (14)

Festuca dinirica (40)

Festuca elviae (3)

Festuca fimbriata (18)

Festuca flacca (4)

Festuca fragilis (17)

Festuca glumosa (41)

Festuca glyceriantha (42)

Festuca guaramacalana (5)

Festuca hatico (26)

Festuca holubii (43)

Festuca huamachucensis (44)

Festuca imbaburensis (45)

Festuca laegaardii (27)

Festuca monguensis (46)

Festuca nereidaensis (47)

Festuca oroana (48)

Festuca parciflora subsp. loxana (49b)
Festuca parciflora subsp. parciflora (49a)
Festuca pilar-franceii (28)

Festuca procera (29)

Festuca pratensis (13)

Festuca quadridentata (15)

Festuca reclinata (10)

Festuca renvoizei (50)

Festuca rubra (20)

Festuca sanctae-martae (51)

Festuca sodiroana (6)

Festuca soukupii (21)

Festuca subulifolia (52)

Festuca sumapana (53)

Festuca toca (30)

Festuca tolucensis subsp. culata (54c)
Festuca tolucensis subsp. perijae (54b)
Festuca tolucensis subsp. tolucensis (54a)
Festuca tovariensis (7)

Festuca turimiquirensis (55)

Festuca ulochaeta (8)

Festuca vaginalis subsp. cayambae (56b)
Festuca vaginalis subsp. vaginalis (56a)
Festuca venezuelana (16)

Festuca woodii (11)

176

Festuca in South American Paramos

ALPHABETICAL INDEX TO COLLECTORS

Number in parentheses corresponds to number of the species in the taxonomic treatment as indicated in
the Alphabetical List of Accepted Species and Subspecies preceeding this section.

Acosta-Solis 21026, 20054 (4); 16963, 16974, 18839
(6); 110206, 19821, 19832, 19835 (12); 6268 (13);
7581 (15); 9879, 18836 (19); 16358 (20); 9884,
9877 (21); 18729 (25); 21031 (27); 16678, 19006,
21439A, B (29); 5097, 19199 (31); 16765 (34a);
7554, 7614, 7623, 10538, 17670, 19196, 21195,
21208, 21216, 166989 (52); 21187 (56a)

Aguirre & Merino 4219 (21)

Aguirre et al. 4206 (31)

Ahumada, O. & A. Schinini 4085 (18)

Alberto et al. 144 (6)

Alfaro 1027 (1a)

Alfaro 1029, 1771 (54a)

Alush Shilom Ton 957 (1a)

Anderson 1271 (14)

Andre 3919 (29)

Anonymous (20)

Apollinaire & Arthur 18 (11); 9, 121 (24)

Archer 3402 (6)

Arechavalet s.n. (18)

Argtiello 348, 354, 357 (52)

Aristeguieta & Medina 3531, 3558 (54a)

Aristeguieta 2596 (17)

Aristeguieta 2603 (17)

Aristeguieta 763 (8)

Aristeguieta 947, 964, 1003, 2464, 2602, s.n. (54a)

Asplund, E. 6359, 7156, 7535, 7559 (4); 6155, 6215,
7195, 7462, 7537, 7878, 16094, 16163, 17846
(6); 8150, 8425 (12); 7524, 8427 (19); 6020,
6400, 6478, 7558, 7892, 8447, 8470, 8606, 16259,
17313, 20217 (21); 6139, 6151, 6154, 7392, 8135,
10375, 16161, 20176 (29); 7806, 7927, 8576,
8586, 10000, 17174 (31); 8446 (34); 7892, 8411,
8446, 8641 (34a); 8397, 8396 (39); 7926, 8594,
17334, 17504 (41); 6134, 6138, 6140, 6369, 6370,
6349, 6348, 6471, 7055, 7803, 7880, 8191, 8402,
8639, 8645, 8644, 8647, 8718, 8724, 9634, 9964,
16807, 9967, 17022. 17176, 17522, e286; G2).
3648, 6620, 7397, 7497, 7872, 8399, 8721, 17338
(56a); 8648 (56b)

Badillo et al. 7562 (17)

Baldillo 3496 (2)

Balslev & T. de Vries 3471 (29)

Balslev 1615 (31); 23600 (52);

Balslev et al. 3901 (31); 3435, 3438, 3437, 3675, 3733,
3734, 3738, 4070 (52)

Barclay & Juajibioy 7017, 7033 (1b); 10180 (17); 7961
(21); 6109, 6517 (24); 8178 (29); 6456, 8804

(31); 8905 (34a); 6313 (41); 8763 (45); 5942,
5945, 6052, 8085, 8162, 8264, 8325, 8641, 8819,
9142, 9160 (52); 9656, 9895, 10214 (54a); 8166,
8905 (56a)

Barclay 6431 (22); 4529, 5865 (52)

Barclay et al. 9677 (17)

Barford & Blicher-Mathiesen 41546 (52)

Barreto 1871 (8); 656 (17); 577 (18); 645 (54a)

Baruch 46 (17)

Beaman, J. 2284, 2363, 3216, 3482 (54a)

Becker et al. 1234 (29)

Bejarano 44 (36); 22 (38)

Bell 17 (4)

Benavides 6497 (19); 2660, 6554 (52)

Benoist 2515, 4636 (52)

Berg & Steinmetz 187 (54a)

Berg 470 (17); 98-29-16 (54a)

Bernal & Jimenez 1127 (19)

Bernardi 268 (17); 18618 (18); 275 (54a)

Berro 5994 (18)

Black 46-605 (24); 32 (52); 278, 279 (56a)

Boechat 41059 (18)

Bono 5662 (17)

Bourgeau, E. 1032 (1a)

Brade & Kuhlman 15631 (18)

Brandbyge 42597 (34a); 42133, 42586 (52)

Bravo 56, 58 (21)

Breedlove, D. 11151, 39265 (1a); 26723 (54a)

Brescia & Marches 4212 (8)

Bricefio 2854, 3111, 3119, 3178, 3357, 3711 (54a)

Briceno & Adamo 184, 187, 949, 1067, 1131, 1139, 1988
(54a); 2052 (54c)

Briceno & Molinillo 3447 (54a)

Briceno et al. 440, 1146, 2330, 2667, 3054, 3077 (54a)

Bricefio, B. & G. Adamo 2003 (2); 191, 196, 274, 306,
948, 1086, 1114, 1320, 1705 (3); 2011 (8); 286
(12); 244 (17)

Briceno, B. & Molinillo 3461 (17)

Briceno, B. 3074 (17)

Bricefio, B. et al. 2328 (2); 2601, 3345, 3390 (3); 358 (17)

Buddle s.n (13)

Buendia 2 (29)

Burger & Gomez 8246 (1a)

Burgeret al. 7428 (1a)

Burkart 19654 (18)

Burman 880, 905 (8)

Cabrera 438 (31)
Camargo 59992 (8)

Festuca in South American Paramos £77.

Camp 3327 (9)

Cano 3780 (29)

Carbono 2474 (1b)

Castelano & Monasterio 20 (54a)

Castellafios 24729 (8)

Castellanos et al. 3 (24)

Cazalet & Pennington 5417 (6); 5763 (31); 5749, 5749B
(41); 5434 (52)

Ceron & Alarcon 4805, 12260 (52)

Ceron & Montesdoca 12010 (52)

Ceron 10605, 15102, 21862, 25977 (29); 12609, 18088,
28250, 28957, 29480 (52)

Ceron et al. 11813, 19819, 25697 (52); 19822 (56a)

Cerrate 1540 (42)

Chardon 5010 (31)

Chase, A. 8276, 9659, 9672, 9673, 10101 (8); 8302
(18)

Clark et al. 1372 (6)

Clausen, J. 34 (54a)

Clayton 4491 (8); 4301 (18)

Cleef & Urribe 6694 (24)

Cleef 4376, 8285 (19); 181, 1587, 6258 (24); 1048, 1272,
3605, 5259, 7704 (28); 7139 (G0); 9078 (37);
982A, 1090, 6285 (38); 7930 (53)

Cleef et al. 5983 (31)

Cleef, A.M. 181 (24); 6826 (36)

Clements et al. 2170 (31)

Correa et al. 129 (22)

Cuatrecasas, J. & Castaneda 24736 (1b); 24532, 24563
(51); 25136, 25133 (54b)

Cuatrecasas, J. & Garcia-Barriga 10037, 10207 (29)

Cuatrecasas, J. & H.G. Barriga 9970 (10)

Cuatrecasas, J. & Jaramillo-Mejia 25742 (24); 25865
(28)

Cuatrecasas, J. & Willard 26292 (52)

@uateecasas, J) 9292. 23137, 23178 (22); 9311 G1);
7966, 9475, 10446 (38); 23201 (41); 14694,
19059 (52)

Cuatrecasas, J., Espinal & Ramos 27562 (22); 3550 (24)

D’Orbygny 39 (18)

Davalos 22 (6); 19 (31); 18 (45); 21, 28 (52)

Davidse, G. & D’Arcy 10244 (1a); 11166 (8)

Davidse, G. et al. 3223 (3); 11074, 11082, 11087 (8);
5545 (38); 19610 (55)

Davidson 1043 (1a)

Degen 861 (18)

Diaz-Piedrahita & Jaramillo-Mejia 1821 (31);
1821(41)

Diaz-Piedrahita & Rangel 2037 (22)

Diaz-Piedrahita 2367 (24)

Dillon, M. et al. 6359A (29)

Dombrowski 2432 (8)

Dusen 3624, 3515, 7783, 7808, 17561 (8); 8972 (18)

Dutra 331, 402 (8); 538, 543 (18)

Dziekanowski & Bolingbroke 1911 (54a)

Echeverry 1973 (22)

Edwards s.n. (45); 70 (52)

Ehrenburg 60 (21); 1, 27, 36, 191 (52)
Ellenberg 7024 (7)

Escalona & Gallegos 287, E384 (29)
Espinal 81, 1035 (52)

Etter 662 (29); 661 (35)

Fagerlind & Wibom 934-bis (21); 934bis, 932 (56a)

Fagerlind et al. 934bis (34a)

Fegan & Falconi 9 (21)

Fernandez-Pérez 2931 (52)

Ferreyra, R. 7526, 7281, 7642, 7664 (14); 8566 (42);
SSI Sayles (SY)

Fierrp 24 (31)

Figueredo 119, 126 (19)

Firmin 116 (29); 137 (52)

Forero, E. et al. 45, 82, 92 (12)

Forrero et al. 3644 (31)

Fosberg, R. 20749 (19); 20745 (28)

Fraser s.n. (4)

Funck & Schlim 1132 (54a)

Galeotti, H. 5766, 5778 (1a)

Gamboet al. 819 (1a)

Garcia-Barriga 7838A (12); 1016, 1019 (24); 1018 (28)
Garzon 617 (38)

Gentry 12378, 12685 (52)

Goergem & Wagner 50197 (8)
Gonzalez et al. 137, 206 (54a)
Gonzalez-Ledesma et al. 484, 478 (1a)
Gregory & Eiten 301 (54a)

Grignon 84189 (52)

Grubb et al. 639 (31); 656 (52)
Guerrero 152 (38)

Halloy B-54 (21)

Hammel, B. 5619 (1a)

Hanselmann & Loleveless 296 (54a)

Harling & Andersson 12538 (52)

Harling 11255 (52)

Harling et al. 8904, 10415 (6); 10533 (21); 8635 (27);
8924, 10477 (52)

Harlington 1197 (21); 4067, 4552 (52)

Harteman 39 (6); 67 (52)

Hartman 7a (19)

Hartweg 1455 (52)

Hatschbach, G. 1912, 4715, 13820 (8); 9551, 20379,
20381, 355'7351(L8)

Heller 306 (54); 62, 306 (54a)

Hernandez 1022A (30); 1311 (36)

Hernandez et al. 1336 (54a)

Herrera 3730 (8)

Hitchcock 28 (18); 496 (54a)

Hitchcock, A.S. 21042 (4); 20798, 21094, 21512, 21682
(6); 20746 (9); 21981 (21); 21926 (29); 21978

178 Festuca in South American Paramos

(31); 21058 (41); 21944, 21951, 22030 (52);
22328 (42); 7165 (54a); 22019 (56a)

Holm-Nielsen & Balslev 23755 (21)

Holm-Nielsen & Jaramillo 28415 (52)

Holm-Nielsen 1482, 24604 (6); 6612 (31); 24871 (34a);
1485, 5383, 5427, 6462, 20789, 25163 (52)

Humbert 26353, 26506, 26806, 26824 (54a)

Humboldt & Bonpland 2296, s.n. (9); s.n. (15); s.n. (19);
Shi: (25)$ san (29)

Idrobo et al. 3026 (6); 3678 (19); 3682 (21); 3737 (29);
3834 (52)

Infantes 435 (14)

Izuriete 129 (29)

J.F. Smith 1997; 1999 (52)

Jahn 18, 43 (54a)

Jahn, A. 62 (17)

Jameson 14, 296 (21); 191 (29); 230 (41); 70, 91, s.n
(52); 230 (56a)

Jaramillo - Mejia & Cleef 5738 (22)

Jaramillo & Zak 623 (52)

Jaramillo et al. 11855, 11856 (27)

Jaramillo-Mejia 6237 (31)

Jaramillo-Mejia et al. 5634, 5683, 5770 (31); 5747,
6149, 6210 (41)

Jorgensen, P.M. 2074 (52)

Jorgensen, P.M. et al. 2099 (52)

JRJ 895 (1a)

Juan & Alva 101 (54a)

Jurgens 6258 (8); 175 (18)

Jurgensen G85 (18)

K.T. Hartweg 1450 (56)

Killip & Smith 17861 (6)

Killip 6709 (6); 19719 (16); 17329 (24); 18479 (26);
18479 (35)

Killip et al. 38050 (38)

Kjell von Sneidern 362, 435, 415, 2144 (52)

Klein 3872 (18)

Kozera & Izernhagen 476 (8)

Kuhlmann 2242 (8)

L. Parodi 5673 (8)

L.G. Clark & X. Londofio 376 (41)

Laegaard & Aguirre 20611 (43)

Laegaard & Sklenar 20308, 20287 (15)

Laegaard, S. & Aguirre 20609 (21); 20608 (49b)

Laegaard, S. & Grignon 19395 (52)

Laegaard, S. & Mayorga 17536(38)

Laegaard, S. & S.A. Renvoize 70580, 70577 (6); 70519
(21); 70485, 70542, 70873, 70882 (29); 70511
(31); 70555 (41); 70558 (56a); 70488 (56b)

Laegaard, S. & Sanchez 20027 (21); 20033 (29)

Laegaard, S. & Sklenar 20302 (9); 20301 (12); 20335,
20363 (21); 20296 (29); 20282A (52)

Laegaard, S. 54540A, 54540B, 102316, 54164, 55260,
55281 (4); 18748, 18727, 18483, 20688, 52285,
52754, 53118, 53372, 55282, 101406, 101711,
101781, 102227; 102293, 102318, POS0Ge9
105132 (6); 20405, 102535, 105256, 105235 (9);
18623, 18667, 18704, 18744, 101818, 102716
(12); 55290, 55411, 55567 (15); 5866, 54327,
54892, 101731 (19); 18736, 19739, 52374A,
52605, 52607, 53580, 53837, 54148, 55301,
55535, 55673, 71737, 101384, 101432, 101718,
102087, 102268, 102237, 103107 (21); 101260
(25); 53965, 53597 (27); 20379, 52301, 54802,
54809, 5556670002, 70014, 70016, 71031, 71695,
55410, 101131, 101350, 101355, 101340, 101814,
101856, 102164, 102285, 102857, 105102 (29);
18449, 51318, 53289, 53343, 54581, 54972,
55425, 55680, 101284, 101383, 101595, 101695
(31); 55769, 101162, 101294, 101716 (33); 19155,
52125, 54594, 54820, 55346, 55736, 101583,
102156, 102892 (34a); 101517, 101612, 102253
(34b); 53293, 53307, 53869, 54518B, 55696,
55718, 70491, 102797, 102803, 102889 (41);
19452, 54548, 55368, 55698, 101194, 101373,
101692, 102262, 102666, 102871, 103105 (45);
70309 (48); 19148, 19395, 21016, 51060, 52107,
52373, 52837, 53125, 53213; 35373; saeae"
53871, 54351, 54409, 54416, 54506, 54731,
55001, 55276, 55279, 55286, 55304, 55401,
55417, 55452, 55635, 55672, 55734, 55748,
55771, 69261, 71739, 71741, 101181, 101164B,
101264A, 101577, 101726B, 101856, 102103,
102100, 102134B, 102376 (52); 19159, 19598,
51386, 52202, 52656, 52659, 55327, 55400,
55536, 55735, 101335A, 101460; "01495;
101559, 102096, 102149, 102743, 102758,
102795, 102796, 102848, 103127 (56a); 19589,
101493, 101505, 101828, 105123 (56b)

Laegaard, S. et al. 18855, 19095, 70813, 103010, 103079,
103032 (6); 103003 (12); 20994 (20); 19066,
20824, 20993, 102737 (21); 18672 (29); 18606,
18859, 19280, 20733, 20931 (31); 102744, 102739
(34b); 19630, 102760, 102725 (41); 53859 (45);
19285 (49b); 18868, 20613, 102733, 103067 (52);
70553, 70554 (56a)

Langsdorff s.n. (8)

Larsen et al. 21 (31)

Leavenworth & Hoogstraal 1218 (54a)

Leavenworth et al. 1213 (1a)

Lechler 2114 (29)

Leite 3490 (18)

Leiva & Leiva 96 (14); 1094 (42); 2525 (52)

Leiva 797 (14)

Len 1139 G1)

Leon, B. & Young 1662 (31); 1101, 1364, 1679 (52)

Liebmann 6109, 6110, 12904 (1a); 517, 6108 (54); 2886,
3030, 6117 (54a)

Festuca in South American Paramos 179

Linderman et al. 4658 (8)

Lindig 1116, 1117 (6); 1862 (8); 14 (19); 1013, 1053
(24)

Littele 15712 (54a)

Little 15717 (17)

Llana 71 (31)

Longhi & Sampaio 368 (18)

Longhi & Witten 5011 (18)

Lopez & Sagasteguii 3340, 8180 (31); 2827 (42); 8180
(52)

Luteyn, J. & Luteyn 5713 (52)

Luteyn, J. 13452 (52)

Luteyn, J. et al. 13051 (22); 12834, 12842 (52)

Lyonnet 260, 2973 (54a)

MacDougal et al. 4504 (6)

Maf 57 (52)

Maguire 3487 (8)

Manrique, E. et al 1147 (1a)

Martin 630 (1a)

Matuda 25909 (1a); 2337, 2360 (54a)

Meier 3025 (17)

Meier et al. 3026 (54a)

Mejia et al. 6181 (22)

Mexia 2684 (54a)

Mille s.n. (4); s.n. (15); 664, 659 (25); 283, 285 (29);
s.n. (52)

Milles et al. 281 (6)

Molau & Eriksen 3244 (31); 3298 (41)

Montesdeoca 280, 282, 284, 295, 339, 348, 585 (52)

Morales 5194 (54a)

Mora-Osejo 3195, 3503 (52)

Morrone, O. & A.M. Cialdella 854 (8)

Mueller 2115 (1a)

Munoz 207 (52)

Mutis 5544 (6); 5570 (24)

Naranyo 4 (31)

Narvazez et al. 548 (31)

Nee, M. & Soule 33022 (54a)
Neill, D.A. et al. 11992, 12042 (31)

Ollgaard, B. & Balslev 10074, 8170, 8545, 8868, 9924,
10122 (31); 9921, 10073, 10100 (45); 8926 (52)

Ollgaard, B. & Holm- Nielsen 38728 (31)

Ollgaard, B. et al. 34039 (27); 34202, 38540, 38541,
38670 (31)

Ortiz 1272 (52)

Oxford Expedition 30 (21)

Palice 1, 3, 4, 9 (6); 6, 7, 8 (52)

Pedersen, T.M. 819, 11475 (18)

Pedraza et al. 491 (24); 228, 242, 278 (28)
Pefnafiel 1067 (52)

Pefiafiel et al. 69 (29); 402, 527, 667, 720 (52)
Penland & Summers 583 (21); 889, 969 (52)

Pennell 9951 (21); 3003 (22); 1923, 2018 (24); 13231
(29); 3064, 9890 (31); 13552, 13845 (52)

Pennell, F. 7057 (52)

Peterson, P.M. 11181 (12); 8815 (29); 8721 (52)

Peterson, P.M. 14224 & O. Tovar (29)

Peterson, P.M. 14907 & N. Refulio Rodriguez (21); 14892
& N. Refulio Rodriguez (31); 13875, 13961,
14011, 14936, 14855, 14925 & N. Refulio Rodri-
guez (42); 13951, 14901 & N. Refulio Rodriguez
(43); 13922, 14953, 14955, 14957, 14968, 14989,
14992, 15178 & N. Refulio Rodriguez (29)

Peterson, P.M. 8872, C.R. Annable & M.E. Poston (6);
8858, 8870, C.R. Annable & M.E. Poston (21);
8717, 8734, 8747, 8846, C.R. Annable & M.E.
Poston (29); 8768, C.R. Annable & M.E. Poston
(34a); 8744, 8759, 8770, 8846, 8865, 8876, 8932,
C.R. Annable & M.E. Poston (52); 8739, C.R.
Annable & M.E. Poston (56a)

Peterson, P.M. 8973, 8996 & C.R. Annable (21); 8979,
8995, 8996A & C.R. Annable (34a); 8982, 8973
& C.R. Annable (34b); 8951, 8983, 8985 & C.R.
Annable (52); 8987 & C.R. Annable (56a); 8984,
9000 & C.R. Annable (56b).

Peterson, P.M. 9118, 9139, E.J. Judziewicz & R.M. King
(21); 9076, 9154, E.J. Judziewicz & R.M. King
(31); 9086, E.J. Judziewicz & R.M. King (41);
9122, 9127, 9140, 9155, E.J. Judziewicz & R.M.
King (52)

Peterson, P.M. 9209, 9235, E.J. Judziewicz, R.M. King &
P.M. Jorgensen (21); 9179, E.J. Judziewicz, R.M.
King & P.M. Jorgensen (29); 9204, E.J. Judzie-
wicz, R.M. King & P.M. Jorgensen (31); 9197,
E.J. Judziewicz, R.M. King & P.M. Jorgensen (45);
9189, 9215, 9245, E.J. Judziewicz, R.M. King &
P.M. Jorgensen (52); 9228, E.J. Judziewicz, R.M.
King & P.M. Jorgensen (56a)

Peterson, P.M. 9313 & E.J. Judziewicz (6); 9259 & E.J.
Judziewicz (9); 9297, 9316, 9512 & E.J. Judzie-
wicz (29)

Pinto-Escobar & Hernandez 523 (24)

Pinto-Escobar 415, 1845 (31)

Pittier 1511 (6); 437, 1320, 1444 (9); 13276 (17); 1409
(52)

Ponce & Trujuillo 245 (12)

Prescott 913 (6); 656, 834, 839, 848 (52)

Pringle, C.G. 3945 (la)

Puiggari s.n. (8)

Quarin 3409 (18)

Quintana 2 (52)

Quipuscoa & Vilchez 2611 (52)
Quiroz 1929 (29)

Rambo 36446, 49315, 54011, 54999, 54989 (8); 45452,
45466, 53471, 53816, 54704, 60082 (18)
Ramirez 1404 (9)

180 Festuca in South American Paramos

Ramirez, M. & N. Cuello 3523, 3524, 3528 (5)

Ramsay & Smith 245, 247 (21); 415B, 502, 785 (31); 410
(41); 343 (45); 422, 510, 742, 1016 (52)

Ramsay 896 (31); 1035 (45); 426 (52)

Ramsay et al. 104A (21); 153 (45); 1057 (52)

Rangel & Aguirre 3720 (24)

Rangel 1650 (28)

Rangel) et aly 945:°(1b);) 1800 Gl); 35151G6);, 1732,
1791 (41)

Rauh & Hirsch E321, E331 (52); E315 (56a);

Regnell III 1409, s.n. (18)

Reitz & Klein 10832, 10399, 12509, 14611, 14585 (8);
11812, 11948, 14172 (18)

Reitz 2611 (8)

Renvoize, S.A. & S. Laegaard 5031 (50)

Renvoize, S.A. 4879 (29)

Ricardi & Carrez 5930 (17)

Rimbach 54 (15); 97a (56a)

Robinson & Beltran 3142 (19)

Rodriguez 4130 (38)

Roldan et al. 360 (6); 396, 307 (19)

Rosengurtt B-2764 (18)

Ross 1276 (54a)

Ruiz & Romero 3 (30)

Sagasteguil 14785 (7); 2991 (14)

Sahn & Hernan 1060 (19)

Saint-Hilaire 312, 304 (8)

Salamanca AC172 (31)

Salamanca et al. 40, SS143 (41)

Sampaio & Arzivenco 341, 362 & 404 (18)

Sanchez & Hernandez 950 (19)

Sanchez 283 (24)

Saravia 3912 (38)

Schinini, A. & Carnevali 10523, 10687 (18)

Schulz & Rodriguez 317 (54a)

Schulz 317 (54a)

Schwaike s.n. (18)

Scito & Kuntyoshi 1315 (8)

Scur 656 (8)

Seaton 228 (54a)

Seifriz 489 (1b)

Sellow s.n. (8)

Sellow, F. s.n. (18)

Sklenaf & Cruz 8706 (21)

Sklenar & Kosteckova 1-24, 15-5, 16-1, 66-10, 85-8,
86-9, 87-6, 1905 (21); 64-19, 91-8, 97-10, 112-25,
1028, 1106, 1142, 1478, 1817, 1890 (31); 1-24,
858, 1322 (34a); 2738 (39); 49-19, 55-2, 56-2, 9-1,
735, 1836, 1853, 1863; 1893, 1903n 19S e933
(41); 956, 1031, 1929, 2168 (45); 14-9, 1881 (52);
143-2, 198 (S6a)

Sklenar & S. Laegaard 7092 (52)

Sklenar & Sklenafova 2168, 2262 (21); 2990 (41)

Sklenar & Zapata Cruz 8705 (43); 8630 (52)

Sklenar 2596, 8204 (45); 8067, 8115 (56a)

Smith & Klein 10887, 10956, 11384, 11335, 11611,
11836, 12024, 12188 (8); 10855, 10958, 13674,
15474, 15761, 15805, 15916, 15936 (18)

Smith & Reitz 10127, 10428 (18)

Smith 10561 (7); 7534, 8233, 10537, 10769, 11389,
12110 (14); 6222 (29); 8284, 11318 (31); 9942,
11493 (42); 3482 (52)

Smith et al. 9389 (18); 7314, 7321a, 7332 (21); 3627
(29); 7428, 8879, 9225, 11127 (31); 3312, 4278,
9939, 9944, 12630 (42); 11851 (43); 3278, 9783,
10229, 10253, 11799 (52)

Sneider 430 (6)

Soderstrom, T.R. 1229 (24); 1307 (28)

Sodiro 36/11, s.n. (4); 36/6, 282, s.n. (6); s.n. (9); s.n.
(12); sn. (13); 20/5; s.n. (15); Sit (25) 5 siaee:
s.n. (41); s.n (52); s.n. (56a)

Sodiro, A. 36/4 (41)

Sohn & Matuda 991 (54a)

Sosa et al. 43 (29)

Sparre 13562 (4); 15076, 16950, 16920 (6); 15635,
15708, 15871 (21); 15827, 15836, 16006 (29);
17719 (31); 13709, 15781, 15828, 17408 (52)

Spruce 5938 (6); 5509 (52)

Stancik, D. & Carvajal 1866 (23); 1783, 1787, 1811,
1848, 1874, 1891 (24)

Stancik, D. & Galvis 2025, 2026, 2028 (46)

Stancik, D. & Medina 2577 (23); 2166 (32)

Stancik, D. 4173, 4174, 4177B, 4180, 4281 (2); 4171,
4175, 4178, 4215, 4250, 4256, 4257 (3); 3668,
4104 (4); 4286 (5); 2632, 2754, 2772, 2776, 2858,
2866, 2870, 2902, 2992, 2991, 3010/3292) 3352.
3595, 3667, 3656,3771, 3781, 3792, 3885, 3886,
3985,4095, 4105, 4113 (6); 4177, 4179 (8); 3221,
3309, 3310, 3798, 4102 (12); 3317 (15); 4262 (16);
4182, 4192, 4248 (17); 948, 1455, 1365, 2024,
2190, 2775, 2983, 3412, 3399, 3554, 355 eGe5,
3683, 3705, 3712, 4092, s.n. (19); 3457 (20); 2744,
2981, 3004, 3108, 3128, 3320, 3358, 3359, 3373,
3380, 3416, 3455, 3596, 3610, 3686, 3713, 3787,
3780, 3800, 3802, 3808, 3858, 3879, 3887, 3903,
4023, 4028, 4031, 4090B, 4093, 4094, 4111, 4158,
4162 (21); 3375, 3376, 3377, 3404, 3405, 3599
(22); 303, 928, 1065, 1069, 12387; 123851289;
1290, 1297, 1302, 1319, 1321, 13235 132451325,
1352, 1355, 1357, 1391, 1402; 1436; 15 aioe:
1530, 1531, 1546, 1842, 1851, 1854, 1921, 1923,
1924, 1969, 1978, 2003, 2025, 2052, 2053, 2063,
2073, 2168, 2174, 2192, 2218, 2256, 2408, 2487,
2507, 2508, 2511,-2512, 2527,255052560;2 360"
2562, 2563, 2564, 2565, 3480, 3481, 3491, 3492,
3493, 3494, 3496, 3500, 3538, 3540, 3542, 3544,
3545, 3546, 3560 (24);4290, 4289 (26); 3365,
3983 (27); 210, 217, 2118, 224, 225, 226; 228725
241, 248, 250, 256, 257, 258, 264, 265, 267, 268,
270, 277, 278, 302, 311, 312, 313, 314; Sie
317, 318, 319, 320, 321, 491, 3494, 355973565

Festuca in South American Paramos 181

3562, 3563, 3564, 3565, 3569, 3568, 3579, 3580,
3583, 3584, 3585, 3586, 3587, 3589, 3590, 3566,
SION 35 109572555755 35745 BOD, 39 16, 3501.
3518, 358i; 3582,. 3585, 3588 (28): 1788, 3312,
3313, 3314, 3317, 3793, 3794, 4106 (29); 2724,
2759, 3044, 3050, 3241, 3279, 3401, 3402, 3410,
3411, 3606, 3611, 3612, 3613, 3614, 3637, 3675,
3703, 3763, 3767, 3849, 3871, 3904, 4097, 4150,
4151 (31); 1493 (32); 948, 1365, 1404, 1418, 1533,
2371 (30); 3080, 3710, 3711, 3719, 3720, 3880,
3883, 4024, 4031B, 4033B, 4034 (34a); 272, 409,
466, 467, 924, 926, 945, 927, 1339, 1366, 1373,
1407, 1421, 1422, 1423, 1438, 1457, 1463, 1465,
ie 1503, 1532, 1555, 1556; 1576, 1577, 1580,
1628, 1602, 1635, 1640, 1629, 1688, 1689, 1690,
1827, 1865, 1828, 1830, 1920, 1935, 1950, 1964,
2000, 2020, 2045, 2051, 2050, 2054, 2130, 2136,
2137, 2138, 2160, 2188, 2189, 2224, 2230, 2232,
2236, 2240, 2495, 2502, 2503, 2505, 2249, 2257,
PANS 2332, 2314..2346; 2350, 2367, 2372,.2373,
2489 (36); 1886 (37); 1286, 1347, 1360, 1363,
196. 1402-15115 1513, 1523-1922, 1929; 1939,
1951, 1952, 1944, 1945, 1985, 1986, 1987, 1988,
2005, 2019; 2109; 2115, 2116, 2117, 2120, 2114,
PNGn2 250, 247 | 24732472, 2597, 2599, 2612,
3539 (38); 4287, 4288 (40); 3031, 3034, 3048,
3373B, 3374, 3378, 3383, 3392, 3393, 3394, 3413,
3601, 3602, 3607, 4039, 4122, 4129, 4133, 4134,
4166, 4149 (41); 2056, 2749, 2783, 2786, 3054A,
30548, 3098, 3101, 3228, 3235, 3267, 3898, 3899,

3990, 4081, 4090, 4091, 4092B, 4098 (45); 3401B
(47); 3846, 3859, 3873 (49a); 2613, 2618, 2617,
2619, 2631, 2656, 2657, 2658, 2659, 2685, 2748,
2764, 2766, 2789, 2790, 2802, 2856, 2876, 2860,
2962, 2963, 2908, 2909, 2910, 2911, 2918, 2914,
2917, 3013, 3119, 3146, 3147, 3149, 3150, 3151,
S152. 3153> 3155,.3158, 3162, 3164, 3169, 3189,
BOs 3 193-3176, 3240; 3326, 3327, 3335, 3336,
3337, 3344, 3351, 3353, 3354, 3355, 3360, 3363,
3367, 3608, 3609, 3671, 3677, 3687, 3688, 3690,
3691, 3692, 3704, 3705B, 3707, 3708, 3709, 3714,
Sls 310,504 7.9709, 37 10, 37725 3773, 3714,
3804, 3805, 3811, 3815, 3837, 3847, 3848, 3850,
3862, 3863, 3864, 3865, 3872, 3882, 3884, 3888,
3889, 3900, 3901, 3902, 4026, 4027, 4030, 4032,
4089, 4096, 4101, 4103, 4115, 4163, 4164 (52);
4172, 4176, 4181, 4191, 4207, 4225, 4242, 4243,
4279, 4280 (54a); 4258, 4259 (54c); 3001, 3014,
3020, 3134, 3137, 3323, 3341, 3706, 3718, 3881,
4013 (56a); 3182, 4033, 4035 (56b)

Standley 61885, 61899, 65227 (1a)

Steyermark, J. & Koyama 102389 (17)

Steyermark, J. 53481 (31); 53146, 53182 (52)

Steyermark, J. A. 53092 (49a)

Storek 2889 (54a)

Stiibel 297 (29); 257 (52)

Stiibel, A. 297 (29); 207 (52)

Sturm 33(38)

Swallen, J.R. 8033, 8168, 8231, 8540, 8859 (8); 8089,
8189 (18)

Tamayo 2326 (16)

Taylor et al. 170 (54a)

Tenorio 15093 (1)

Teran & Melchior 833, 848 (17)
Teran 77132. 7909; 13215:(17)
Tillet & Honig 746-765 (1b)
Tillett, S.S. 747-1122 (54b)
Torres-Romero & Lozano 2982 (28)
Torres-Romero et al. 1875 (31)
Tovar Serpa, O. 2057 (7)

Tovar, O. 143 (14)

Triana 810, s.n. (31); s.n. (38)
Tsugaru & Otsuka B-2265 (18)
Turpe, A.M. 2932 (8)

Tutin 1285 (29)

Ule 243, 616, 1959 (18)

Valderrano 34 (38)

Valls 56, 2386 (8)

Valls et al. 1893 (18)

Vareschi 1232 (17); 2134, 2232, 2244, 2248, 3853,
7021 (54a)

Vareschi et al. 1233 (17)

Vargas 11176 (52)

Vargas et al. 2083, 2583 (52)

Vega & Ruiz 3584 (29)

Vega 249 (7); 1966 (14); 5419, 7143 (21); 723, 2250,
2576, 3239, 3830, 4661, 5013 (29); 7087 (31);
2263, 4168, 4371, 5271, 5427, 6616, 10509,
10764, 10889 (42); 4378, 7111, 7124 (43); 1385,
1969, 4043, 4145, 7136, 11181, 11190 (52)

Vega et al. 1544 (7); 1638, 10320 (21); 411, 1393, 1598,
32833 385); 1.85, 5096, 5745, 6953 @9)- 1695,
2012, 2061, 2527, 3797, 5744, 6694, 6974, 6975
(42); 2018, 3494, 6914, 6955, 6994, 7041, 7128
(43); 7011 (50); 2011, 2464, 7062, 10795, 10904,
10906 (52)

Vélez et al. 4514 (22)

Vivar & Marin 1305 (12); 1306 (13)

Vries, T. de s.n. (45); s.n. (52)

Wagner 944, 946, 947, 5010 (8)

Wagner et al. 298 (8)

Wawra 947 (54a)

Weberbauer, A. 3230 (14); 3275 (42); 3974 (52)

Whymper 1326, s.n. (31); 1330 (41); 1322, 1638 (52)

Widgren s.n. (18)

Winge et al. 1272, 1362 (8)

Wood 4326, 4347, 4792, 5066, 4648 (6); 3842, 5104 (8);
5309 (9); 4639 (12); 5259 (17); 4247, 5331 (21);

182 Festuca in South American Paramos

4033 (22); 3612 (24); 4539 (28); 4797 (29); 4481, | Yepes-Agredo 760 (22)
4507 (30); 4030, 4062, 4063, 4802 (31); 5141 (36); Young & Watson 3484 (52)
5260 (37); 3523, 4539 (38); 4806, 4807 (52)
Wood, J.R.I. 5254, 5354 (11); Zapata 16506, 16508 (43)
Wurdack, J. 1572 (31); 1571 (52) Zardinii 7701 (18)
Zuluoga 107 (38)
X. London et al. 550 (23)

Festuca in South American Paramos 183

INDEX TO SCIENTIFIC NAMES

Accepted names are in regular type; main entry for each is in bold face; synonyms are in italics; figures

are in parentheses.

Avena secunda Salisb. 42

Bromus arundinaceus (Schreb.) Roth 42
caldasii Kunth 36

depauperatus J. Pres| 5

elatior (L.) Koeler 42

littoreus Retz. 42

pratensis (Huds.) Spreng. 44
procerus Kunth 36

Bucetum elatius (L.) Parnell 42
pratense (Huds.) Parnell 44
Diplachne procera (Kunth) Spreng. 77

Festuca L. 11

aequipaleata E. Fourn. 136

americana (Pers.) F. G. Dietr. 44

ampliflora Doll 52

amplissima Rupr. 5, 6, 13, 16, 18, (19, 20, 147)

amplissima Rupr. subsp. amplissima 17

amplissima subsp. magdalenaensis 6

amplissima subsp. sierrae Stancik 18

andicola Kunth 5, 6, 7, 12, 16, 22, (54), 55, (56, 158)

arundinacea Schreb. 12, 16, 42, (43, 45, 153, 154)

arundinacea subsp. orientalis (Hack.) Tzvelev 42

asplundii E.B. Alexeev 7, 14, 17, (83), 84, (85), 104,
136, (163)

Festuca sect. Aulaxyper Dumort 16, 83, 106

azucarica E.B. Alexeev 13, 16, 63, (64, 65, 159)

boyacensis Stancik 6, 15, 17, (79), 88, (89), 91, (164)

breviaristata Pilg. 6, 129

breviglumis Swallen 16, 38

cajamarcae Pilg. 129, 130

caldasii (Kunth) Kunth 5, 7, 12, 16, (35), 36, (37), 38,
40, (151, 152)

carchiense Stancik 6, 15, 17, 88, (90), 91, (93), 108,
122, (164)

casapaltiensis Ball 7

Festuca sect. Cataphyllophorae E.B. Alexeev 6, 16,
65, 70, 77, 83

chimborazensis E.B. Alexeev 15, 17, 88, 91, (92, 93,
164, 165)

chimborazensis subsp. chimborazensis (92), 93, 94

chimborazensis subsp. micacochensis 6, 93, 94

chita Stancik 6, 13, 17, 95, (96, 102)

chitagana Stancik 6, 12, 16, (61), 66, (67)

cleefiana E.B. Alexeev 14, 17, 75, (93), 95, (97, 165)

cocuyana Stancik 6, 15, 17, (83), 98, (99), 100, 126,
(166)

colombiana E.B. Alexeev 13, 16, (65), 66, (68), 69,
72, (159, 160)

coromotensis B. Bricefio 5, 13, 16, 20, 21, (20, 21),
22, 28, (147, 148)

cundinamarcae E.B. Alexeev 14, 17, 100, (101, 102,
166, 167)

dasyantha Kunth 5, 7, 13, 16, (65), 70, (71), 72, 75,
(160)

densipaniculata E.B. Alexeev 14, 17, 102, (103), 104,
(106, 167)

dichoclada Pilg. 6, 7, 12, 16, 45, (45), 46

dinirica Stancik 6, 7, 14, 17, (93), 104, (105), 106

dissitiflora Steud. 55

distichovaginata var. cajamarcae (Pilg.) St.-Yves
129

dolichophylla J.Presl. 6, 7

Festuca subg. Drymanthele V.I. Krecz. & Bobrov 16

drymeja Mert. & W.D.J. Koch 16

elatior L. 42

elatior subsp. arundinacea (Schreb.) Celak. 42

elatior subsp. arundinacea (Schreb.) Hack. 42

elatior subsp. arundinacea var. genuine subvar. ori-
entalis Hack 42

elatior subsp. pratensis (Huds.) Hack. 44

elatior var. arundinacea (Schreb.) Wimm 42

elatior var. pratensis (Huds.) A. Gray 44

elviae B. Bricefio 5, 12, 16, 22, (23, 26), 55, (148)

eminens Kunth 6, 36

Festuca subg. Erosiflorae E.B. Alexeev 16

ferreyrae Tovar 129, 130

fimbriata Nees 12, 16, 52, (53, 54, 157)

flacca Hack. ex E.B. Alexeev 12, 16, (20), 24, (25),
33, (149)

flexuosa Willd. ex Kunth 46

fluitans var. pratensis (Huds.) Huds. 44

fragilis (Luces) B. Bricefio 5, 7, 14, 16, 48, (50, 51),
51, (136)

fratercula Rupr. ex E. Fourn. 17

glabra Spreng. 44

Festuca sect.Glabricarpae E.B. Alexeev 6, 16, 38

glumosa Hack. ex E.B. Alexeev 7, 16, 17, 91, (102),
106, (107), 108, 117, 122, 126, 136, (168)

glyceriantha Pilg. 14, 17, (106), 109, (110), 111,
(168)

guaramacalana Stancik 6, 7, 13, 16, 26, (26, 27), 28,
(149, 150)

hatico Stancik 6, 12, 16, (61), 72, (73, 160, 161)

Festuca subg. Helleria E.B. Alexeev 6, 16, 51

holubii Stancik 6, 16, 17, (106), 111, (112), 113

huamachucensis Infantes 15, 17, 113, (114, 115)

imbaburensis Stancik 6, 15, 17, 91, 108, 115, (115,
116), 117, 122, (169)

laegaardii Stancik 6, 11, 17, 65, (65), 72, (74), 75,
(161)

184

leptothrix Trin. ex Doll 33
liebmannii E. Fourn. 136
littoralis Wahlenb. 42
livida (Kunth) Willd. ex Spreng. 5, 16, 51
Festuca subg. Mallopetalon (Doll) E.B. Alexeev 6,
16
“monguensis Stancik 6, 14, 17, (106, 118), 119, (169)
multiculmis Steud. 136
nereidaensis Stancik 6, 14, 17, (115), 119, (120, 170)
orgyalis Willd. ex E. Fourn. 77
orgyalis Willd. Ex Spreng. 77
orientalis (Hack.) Krecz. & Bobrov 42
oroana Stancik 6, 14, 17, 119, (121), 122, (126, 170)
orthophylla Pilg. 7
ovina L. 11
ovina subvar. jamesonii St.-Yves 106
parciflora subsp. loxana 6, 122, 124
parcifiora subsp. parciflora 122, (123), 124
parciflora Swallen 15, 17, 122, (126)
peruviana E.B. Alexeev 7, 77,
pilar-franceii Stancik 6, 13, 17, 65, 69, 75, (76, 79,
162)
pinetorum Swallen 5, 7
poaeoides Michx. 44
poaeoides var. americana Pers. 44
pratensis Huds. 12, 16, (40), 44
pratensis var. elatior (L.) Gaudin 42
presliana Hitche. 5
procera: Kunth 3.06; pbs Gn oa) Zeta) Ore ane os
84, (162)
procera Nees & Meyen 77
pseudosclerophylla Krivot. 42
quadridentata Kunth 5, 7, 12, 16, 46, (47, 51, 154,
155)
quadridentata subsp. eminens (Kunth) St.-Yves 36
quadridentata var. caldasii (Kunth) St.-Yves 36
quadridentata var. dichoclada (Pilg.) St.-Ives 45
quitensis Willd. ex Kunth 36
racemosa Willd. ex Spreng. 55
reclinata Swallen 12, 16, 38, (39, 40, 152)
regeliana Pavl. 42
renvoizei Stancik 6, 15, 17, 124, (125, 126)
rigescens (J. Presl) Kunth. 7, 55
rubra LE 13,16; (SD) 57. (O8s lsietas)
Festuca sect. Ruprechtia E.B. Alexeev 6, 16, 18
sanctae-martae Stancik 6, 15, 17, 100, (115), 124,
WAS (ZT. Ni)
scabra Willd. ex Steud. 77
Festuca sect. Schedonorus (P. Beauv.) Koch 16
Festuca sect. Schedonorus (P. Beauv.) Peterm. 11, 16
sodiroana Hack. ex E.B. Alexeev 7, 12, 16, 21, 22,
(26), 28, (29, 150)
soukupii Stancik 6, 14, 16, 55, 59, (60, 61, 158)
steinbachii 48

Festuca in South American Paramos

sublimis forma vivipara St.-Yves 146

sublimis Pilg. 77

subulata Trin. 16

subulata var. fraseriana St.-Yves 24

Festuca subg. Subulatae (Tzvelev) E.B. Alexeev 16

Festuca sect. Subulatae Tzvelev 16, 28

subulifolia Benth. 14, 17, 84, 104, 126, (128, 129),
1305 1395.7)

sumapana Stancik 6, 15, 17, 91, 122, (129), 136,
(137)

tenuiculmis Tovar 55

toca Stancik 6, 12, 17, 65, 81, (82, 83), 83, 84, (163)

tolucensis Kunth 5, 14, 17, 84, 104, (129), 130, 136,
139, 141, 143

tolucensis subsp. culata Stancik & P.M. Peterson 6,
139, 141

tolucensis subsp. perijae Stancik 6, 139, 141

tolucensis subsp. tolucensis (138), 139, (172)

tolucensis var. subulifolia (Benth.) St.-Yves 126

tovariensis Stancik & P.M. Peterson 12, 16, 24, 31,
@2535))

turimiquirensis Stancik & P.M. Peterson 6, 14, 17,
(102), 141, (142), 143

ulochaeta Nees ex Steud. 6, 7, 13, 16, 24, 31, 33, (34,
355) 151)

vaginalis (Benth.) Laegaard 7, 15, 17, (126), 143,
(172)

vaginalis subsp. cayambae Stancik 6, 143, 146

vaginalis subsp. vaginalis 143, (144), 145

venezuelana Stancik 6, 13, 16, (45), 48, (49, 155)

woodii Stancik 6, 13, 16, 36, 38, 40, (40, 41, 153)

Gnomonia elatior (L.) Lunell 42

fragilis Luces 48
fragilis (Luces) Rauschert 48

Leucopoa pseudosclerophylla (Krivot.) Bor 42
Lolium arundinaceum (Schreb.) Darbysh. 42
pratense (Huds.) Darbysh. 44
Poa elatior (L.) Moench 42
hybrida var. vallesiaca Bronm. 42
phoenix Scop. 42
Schedonorus americanus (Pers.) Roem. & Schult. 44
arundinaceus (Schreb.) Dumort. 42
caldasii (Kunth) Roem. & Schult 36
elatior (L.) P. Beauv. 42
littoreus (Retz.) Tzvelev 42
phoenix (Scop.) J. Holub 42
pratensis (Huds.) Beauv. 44, 45
procerus (Kunth) Roem. & Schult. 36
radicans Dumort. 44
Tragus elatior (L.) Panz. ex B.D. Jacks. 42
pratensis (Huds.) Panz. ex B.D. Jacks. 44
Uniola effusa E. Fourn.17
muelleri E. Fourn. 17
Vulpia ulochaeta Nees ex Doll 33