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SEGUIR LEYENDO
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OCTUBRE 29 DE 2018
CUCBA | UNIVERSIDAD DE GUADALAJARA

                                     9
                                                         ISSN: 2007-5049
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Quercus centenaria en el municipio de San Sebastián del Oeste, Jalisco.
                                    Fotografía de L.M. González-Villarreal,
                                    4-5 de octubre de 2011.

Nota del editor                                                    Editor’s note
Con la intención de llegar a un público más extenso                With the intention to make it possible for more readers
que hacen uso de las tecnologías actuales, se decidió              to have easy access to our publications we have
publicar la revista ibugana exclusivamente en                      decided to publish our bulletin ibugana exclusively
formato digital. En México, el Instituto Nacional de               in digital format. This does not imply that it is a new
Derechos de Autor, establece que se reinicie la serie con          journal and therefore libraries should not designate a
un ISSN distinto y a partir del “número uno” para la               new title for ibugana. However, the mexican
versión electrónica. Esto no significa que se trate de             Instituto Nacional de Derechos de Autor requires
otra revista, por ello no será necesario alterar los               distinct ISSN number beginning with “number one” for
registros de la versión impresa que de ella se tengan en           the first electronic volume. Please note this difference
las bibliotecas.                                                   in future citations.

Esta versión electrónica puede consultarse de manera               The electronic version is available to anyone in:
libre en la dirección: http://ibugana.cucba.udg.mx y               http://ibugana.cucba.udg.mx. The page is designed
está diseñada para imprimirse en papel tamaño carta                to print on letter size paper (8.5 × 11 inches).
(21.59 × 27.94 cm).
                                                                   We welcome articles regarding any aspects of botany
Serán bienvenidos todos los trabajos en las diferentes             for review and possible publication. Information for
áreas de la botánica para su revisión y posible publi-             contributors is available at the address cited above.
cación; la “información para los autores” se encuentra             Proposals should be sent to:
en la dirección antes citada. Las propuestas deben                 ibugana@cucba.udg.mx
dirigirse a: ibugana@cucba.udg.mx

ibugana, No. 9, octubre de 2018, es una publicación editada por la Universidad de Guadalajara a través del Departamento
de Botánica y Zoología, del Centro Universitario de Ciencias Biológicas y Agropecuarias. Las Agujas, Zapopan, Jalisco, kilómetro
15.5 carretera Guadalajara-Nogales, C.P. 45101, tel.: (33) 3777-1192 ext. 32973, http://ibugana.cucba.udg.mx,
editores.ibugana@gmail.com, editor responsable: Luz María González-Villarreal. Reserva de Derechos al Uso Exclusivo [En
trámite], ISSN: 2007-5049, otorgados por el Instituto Nacional del Derecho de Autor.

Las opiniones expresadas por los autores no necesariamente reflejan la postura del editor de la publicación.

Queda estrictamente prohibida la reproducción total o parcial de los contenidos e imágenes de la publicación sin previa
autorización de la Universidad de Guadalajara.
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Contenido

Universidad de Guadalajara

Rectoría General

Miguel Ángel Navarro Navarro      3 Three novelties in Clethra (Clethraceae: sect. Cuellaria)
Rector                            from the cloud forests of southern Mexico and western
                                  Guatemala
Carmen Enedina Rodríguez          Luz María González-Villarreal       Citar
Armenta
Vicerrector Ejecutivo             21 Nuevos registros de ocho especies de Tigridieae
                                  (Iridaceae) en México
José Alfredo Peña Ramos
                                  Guadalupe Munguía-Lino, Arturo Castro-Castro y Aarón Rodríguez   Citar
Secretario General

Centro Universitario              47 Dos nuevas especies de encinos (Quercus:
de Ciencias Biológicas y          Fagaceae), adicionales para la Flora de Jalisco y Áreas
Agropecuarias                     Colindantes, en el Occidente de México
                                  Luz María González-Villarreal       Citar
Carlos Beas Zárate
Rector                            73 The Field in the Frame: Rediscovering the
                                  Photographs of Thomas Baillie MacDougall
Ramón Rodríguez Macías
                                  Catherine Phillips   Citar
Secretario Académico

Adrián Gómez Medrano
Secretario Administrativo

Departamento de Botánica y
Zoología

Jesús Jacqueline Reynoso Dueñas
Jefe de Departamento

Instituto de Botánica

Ofelia Vargas Ponce
Director

Luz María González-Villarreal
Editor Jefe

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Consejo editorial

Paul E. Berry                       Jerzy Rzedowski                                      Es una publicación electrónica de

University of Michigan              Instituto de Ecología, A.C.                             la Universidad de Guadalajara,
Ann Arbor, Michigan, E.U.A.         Pátzcuaro, Michoacán, México.                        que tiene el propósito de difundir

                                                                                           el conocimiento de la botánica,
Thomas F. Daniel                    Tom L. Wendt                                             entendida en sentido amplio,
California Academy of Sciences      University of Texas                                      así como los resultados de los
San Francisco, California, E.U.A.   Austin, Texas, E.U.A.
                                                                                       trabajos de investigación científica

                                                                                         desarrollados en sus propias y en
M. Socorro González Elizondo        José Luis Villaseñor
                                                                                                         otras instituciones.
Instituto Politécnico Nacional      Instituto de Biología, UNAM
Durango, Durango, México.           C.U., México, D.F.
                                                                                            Se publican trabajos originales

                                                                                              e inéditos en español, inglés,
María de Lourdes Rico Arce
                                                                                        portugués y francés; cada artículo
Royal Botanic Gardens, Kew
Richmond Surrey, Inglaterra.                                                           contiene un resumen en español y

                                                                                        en inglés, además del propio de la

                                                                                        lengua en que esté escrito. No hay

                                                                                          límites en el número de páginas

                                                                                         ni en la cantidad de fotografías a

                                                                                                                       color.

                                                                                         Diseño editorial

                                                                                          Orgánica Editores
                                                                                                 Saulo Cortés |
                                                                                         José Manuel Sánchez
                                                                                   Enrique Díaz de León 514-2b,
                                                                                              Guadalajara, Jal.
                                                                                             T (33) 3825-8528 |
                                                                                                 (33) 3825-8545
                                                                                    www.organicaeditores.mx

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Three novelties in Clethra (Clethraceae: sect. Cuellaria)
from the cloud forests of southern Mexico and western
Guatemala
Luz María González-Villarreal1,2                                                                                         Citar

1
 Wisconsin State Herbarium,                           Abstract                              Resumen
Department of Botany, University of                   Three new endemic species of Cle­     Se describen e ilustran tres espe-
Wisconsin-Madison, Birge Hall, 430
                                                      thra (Clethraceae) are described      cies nuevas endémicas de Clethra
Lincoln Drive, Madison, WI 53706 USA.
                                                      and illustrated. Clethra motozin­     (Clethraceae), Clethra motozintla­
2
 Herbario Luz María Villarreal de Puga,               tlana and C. tacanensis occur in      na y C. tacanensis del sur de Chia-
Instituto de Botánica, Departamento                   southernmost Chiapas, Mexico,         pas, México, y C. zunila de Guate-
de Botánica y Zoología, Centro                        and C. zunila in western Guatema-     mala occidental. Las tres especies
Universitario de Ciencias Biológicas                  la. All three are highland endemic    prosperan en el bosque de neblina
y Agropecuarias, Universidad de
                                                      species that inhabit montane cloud    a elevaciones de 2000−3100 m, en
Guadalajara. Apartado postal 1–139,
                                                      forests at 2000−3100 m elevation      volcanes de interés desde el pun-
Zapopan 45101, Jalisco, México.
                                                      on floristically interesting volca-   to de vista florístico y en cadenas
@   luz.gvillarreal@academicos.udg.mx                 noes and mountain chains. These       montañosas de esos países. Estas
                                                      species have been confused with       plantas fueron confundidas o re-
                                                      or considered related to C. pache­    lacionadas con C. pachecoana con
                                                      coana, with which they are com-       la que se hace una comparación y
                                                      pared and for which an updated        además se da una descripción ac-
                                                      description is given. Additionally,   tualizada. Asimismo se presenta
                                                      a key to distinguish these taxa and   una clave para identificar las espe-
                                                      a map showing their geographical      cies y un mapa mostrando su dis-
                                                      distributions are provided.           tribución geográfica.

                                                      Keywords: endemism, foliar tri-       Palabras clave: endemismos, tri-
                                                      chomes, Mesoamerica, new spe-         comas foliares, Mesoamérica, nue-
                                                      cies, taxonomy.                       vas especies, taxonomía.

Introduction

T   he genus Clethra L. (Clethraceae) is distribut-
    ed in temperate and tropical regions of both
hemispheres. It consists of shrubs and trees and
                                                                        dowski (2001). Mexico, with about 30 species,
                                                                        is the richest country, and more than half of its
                                                                        species are endemic. The diversity of the genus
is well represented in mountainous areas. The                           is mostly concentrated in the state of Chiapas,
number of taxa included in it varies depending                          where 18 species occur, of which six are local
on the author. According to Sleumer (1967) it                           endemics. It is followed by the states of Oaxaca
contains 64 species, whereas Fior et al. (2003)                         and Veracruz (González-Villarreal 2006). Stand-
mentioned 85, but the most accurate estimate is                         ley & Williams (1966) reported eight species for
approximately 120, as cited by Calderón de Rze-                         Guatemala, which has the highest diversity of all

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LUZ MARÍA GONZÁLEZ-VILLARREAL

Central American nations. However, the work           ly with reddish-brown or golden trichomes. The
of Sleumer (1967) and studies currently being         vestiture is comprised of two strata. The inferi-
carried out by the author show that there are         or stratum is made up of minute, more or less
actually 12 species, including one of those here      appressed, stellate trichomes, and the superior
proposed.                                             stratum of longer, stellate trichomes and fascic-
                                                      ulate trichomes that are either sessile, stipitate
   In his revision of the genus Clethra Sleumer       or pedestaled, and often with acicular or filiform
(1967) emphasized the importance of a certain         trichomes intermixed or these mainly concen-
characters, particularly seed morphology. On          trated on the main veins. The series includes 15
the basis of these features he proposed an infra-     species distributed throughout Mexico, Central
generic classification in which he classified the     America, South America, Trinidad and Jamaica
64 species into two sections: Clethra (25 spp.),      (González-Villarreal 2007).
distributed in East Asia and North America, and
section Cuellaria (39 spp.), represented mostly in
the Western Hemisphere. Section Cuellaria was         Materials and Methods
further divided by Sleumer into two subsections:
Cuellaria and Pseudocuellaria, the latter including   Herbarium material from the following collec-
only C. arborea Aiton from the island of Madeira.     tions was studied: A, ARIZ, BM, CAS, CHAPA,
However, molecular data and seed morphology           DS, ENCB, F, GUADA, IBUG, INIF, K, L, LL,
show that C. arborea is not well-supported as be-     MEXU, MICH, MO, NY, RSA, TEX, U, US and
longing to section Cuellaria and should be trans-     WIS; acronyms are according to Thiers (2015).
ferred to section Clethra (Fior et al. 2003).         The majority of the specimens were sent on loan
                                                      to the Wisconsin State Herbarium (WIS), where
   Sleumer (1967) used variation in leaf in-          this work was carried out.
dumentum to divide subsection Cuellaria into
four series: Glabrae, Tomentellae, Tomento­               The new species were recognized only from
sae, and Ferrugineae. In adopting his proposal,       herbarium material. However, field explorations
González-Villarreal (1996, 2007) has studied tri-     were conducted in March of 2009 in the state of
chome morphology in detail and learned that it        Chiapas, Mexico to gather additional informa-
indeed serves to distinguish between groups of        tion and photographs of Clethra motozintlana. All
species, allowing the assignment of the species       measurements of floral and vegetative charac-
proposed here to series Tomentellae and Tomen­        ters included in the following descriptions were
tosae.                                                obtained from herbarium specimens. The flower
                                                      dissections were prepared by boiling the materi-
   The series Tomentellae is primarily charac-        al in water for a few minutes. The examination
terized by the abaxial leaf surface, which is pale,   of pubescence and flowers was aided by a stereo
yellowish or ochraceous, and although smooth          microscope, Olympus SZ61 with 45× magnifica-
to the touch and glabrous in appearance, in real-     tion. A dichotomous key were developed to help
ity is totally covered by minute, appressed, stel-    identify the three species here proposed. Geo-
late trichomes. Also, the midvein and secondary       graphical distributions of the taxa were deter-
veins are covered with scattered, acicular tri-       mined on the basis of the examined specimens,
chomes that are mostly antrorsely orientated,         and a map was made from the elevational mod-
but sometimes are glabrous or with a few fascic-      el obtained from Global Administrative Areas
ulate trichomes. This series contains 16 species,     (http://www. gadm.org/).
and although most diverse in Mexico, it also oc-
curs in Central America, the Antilles and Cuba,       Taxonomic treatment
where it is represented only by Clethra cubensis
A.Rich. (González-Villarreal 2007). In contrast,      During a revision of the genus Clethra in Mexico
the series Tomentosae is mostly characterized         and Central America, the following new species
by the abaxial leaf surface being soft or harsh to    were encountered, and the description of C. pa­
the touch and tomentulose, tomentose or woo-          checoana Standl. & Steyerm. was amplified.

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THREE NOVELTIES IN CLETHRA (CLETHRACEAE: SECT. CUELLARIA) FROM THE CLOUD FORESTS OF SOUTHERN MEXICO AND…

Clethra motozintlana L.M.González, sp. nov.                  tiny, multiradiate trichomes, midvein somewhat
(Figures 1−2, map 1).                                        prominent; secondary veins (8−) 10−15 on each
                                                             side, slender, reddish-brown contrasting with the
   Type: Mexico, CHIAPAS, Mpio. Motozintla                   pale surface, usually arch-ascending, branching
de Mendoza, N and W slope of Cerro Mozotal                   and anastomosing to form interconnected arches
below the microwave tower along the road from                near the margins, bearing appressed, stellate tri-
Huixtla to El Porvenir and Siltepec, elev. 3000 m,           chomes and sometimes a few acicular trichomes,
30 Dec 1972 (fl, fr), D.E. Breedlove & R.F. Thorne           veinlets forming a fine, not clearly distinct net-
31190 (holotype: MICH; isotypes: DS, INIF, LL,               work. Inflorescence a short panicle of (2−) 5−10
MEXU226306, MO2610502, RSA).                                 racemes, (2.5−) 5−15 cm long, loosely-flowered,
                                                             up to ca. 20 flowers per cluster; rachis slender, red-
    Clethra motozintlana is morphologically sim-             dish-brown, densely covered by multiradiate and
ilar to C. pachecoana, but it differs in having              fasciculate trichomes with 2−5 erect rays; bracts
broadly elliptic or obovate to obovate-oblong                ovate, somewhat conduplicate, 1.5−2.5 (−3) mm
leaf blades (vs. narrow elliptic) with the base              long, as long as the flowering pedicels or short-
not folded; the adaxial surface evidently punc-              er, the basal bracts early deciduous, 5.5−7.5 mm
tate; and the abaxial surface with scattered, red-           long; pedicels slender, (0.8−) 2−3 mm long, re-
dish, multiradiate trichomes in addition to the              curved in flower; flower buds subglobose; sepals
appressed, stellate trichomes. The inflorescenc-             ovate to ovate-oblong, (1.5−) 2.5−3 (−3.5) × (1−)
es are loosely-flowered with ovate bracts (vs.               1.5−2 mm, apex of the internal lobes acute and the
densely-flowered with lanceolate bracts) as long             external obtuse, margins ciliate, reddish-brown
as the pedicels or shorter (vs. longer than the              pubescent, canescent-puberulous inside; petals
pedicels), and the petals are glabrous inside (vs.           entirely free, oblong or spathulate, (2.5−) 3.5−4.5
sparingly pilose).                                           (−5) × (1−) 2−3.5 mm, margins deeply fimbriate,
                                                             long-ciliate, glabrous inside; filaments filiform,
    Tree small to medium-sized, 8−15 (−20) m tall,           2−3 mm long, slightly enlarged at the base, flat-
trunk often slender, 20−30 cm in diameter; bark              tened laterally; anthers sagittate, 0.7−1.3 mm long;
smooth to slightly fissured; branchlets slender,             ovary 3-carpellate, 1−2 mm in diameter, densely
fragile, ca. 4 mm in diameter, light brown, tomen-           sericeous; style short, (1−) 1.5−2 mm long. Fruit
tulose, beginning to shed with age. Leaves subco-            a small, depressed-globose, trigonous capsule,
riaceous, bicolored; petiole slender, (0.3−) 1−1.5           when mature 2.5−4 × 3−6 (−7) mm, on a pedicel
(−2) cm long, reddish-brown, puberulous but                  3−4 mm long. Seeds numerous, flattened, winged
glabrescent with age; leaf blades very variable in           all around, 2.0 × 1.5 mm, including the wing.
shape even on the same branchlet, mostly broadly
elliptic, or obovate to obovate-oblong, (1.5−) 4−7              Distribution and Habitat: Clethra motozintlana
(−13) × (0.8−) 2−5 (−7.5) cm, apex acute or ob-              is endemic to Chiapas, Mexico and known from
tuse to rounded, sometimes acuminate or apicu-               Cerro Malé, Cerro Boquerón, Cerro Mozotal, and
late, base mostly cuneate or narrowed to abruptly            the Tacaná Volcano within the Sierra Madre de
rounded, margins serrate in young leaves, entire             Chiapas; it probably also occurs in Guatemala. It
in mature ones, undulate, rarely with 4−6 low                grows at elevations from (1350-) 2000 to 3000 m,
serrations usually confined to the distal third of           often along roads or on steep slopes in evergreen
the blade; adaxial surface dark green, when young            cloud forests with the characteristic elements A­
covered with tiny, amber-colored, multiradiate               bies guatemalensis, Alnus acuminata subsp. arguta,
trichomes, glabrescent to glabrous with age but              Arbutus glandulosa, Clethra glaberrima, C. oleoides,
clearly punctate, the midvein furrowed, densely              Comarostaphylis arbutoides, Cornus disciflora, Chi­
covered with multiradiate and acicular trichomes             ranthodendron pentadactylon, Drimys granadensis,
or glabrescent, secondary veins impressed, the               Fuchsia spp., Gaultheria montana, Ostrya virgi­
veinlets not evident; abaxial surface pale-brown             niana var. guatemalensis, Symplocos limoncillo, and
or somewhat ochraceous, glabrous in appear-                  species of Pinus and Quercus.
ance but actually covered with dense, minute, ap-
pressed, stellate trichomes and scattered, reddish,

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LUZ MARÍA GONZÁLEZ-VILLARREAL

Figure 1. Clethra motozintlana. A. Young branchlet. B. Leaves, showing both surfaces. C. Paniculate inflorescense. D. Matu-
re branchlet. E. Fruits. Drawn by Manuel Peinado.

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THREE NOVELTIES IN CLETHRA (CLETHRACEAE: SECT. CUELLARIA) FROM THE CLOUD FORESTS OF SOUTHERN MEXICO AND…

                                                                 B

                                                          A
                                                                 D

                                                          C

                                                                 F
                                                          E

Figure 2. Clethra motozintlana. A. Habit. B. Bark. C-D. Comparison of both leaf surfaces. E. Infructescence. F. View taken
between Motozintla de Mendoza and Niquivil, Chiapas, Mexico where this species grows.

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LUZ MARÍA GONZÁLEZ-VILLARREAL

    Clethra motozintlana is a slender tree with dark              Phenology: Clethra motozintlana was collect-
green leaves that are glossy on the adaxial surface            ed with flowers in November and December, and
and yellowish or gray-buff beneath. It is sympatric            with fruits in December and March, although both
with two other Clethra species, one of which is C.             phases likely occur year-round.
glaberrima Lundell (series Glabrae). Both of them
have been collected on the road to El Porvenir and                Etymology: The species is named after the
Siltepec and along the ridge above Siltepec on the             Municipality of Motozintla de Mendoza, Chi-
road to Huixtla at 2100 m. The other sympatric                 apas, Mexico, where the plant grows on mountain
species is C. oleoides L.O.Williams (series Tomen­             peaks.
tellae), which has also been reported at 3000 m
on cerro Mozotal. This is the second highest peak                 A member of series Tomentellae, Clethra moto­
(3012 m elev.) in Chiapas, the highest being Ta-               zintlana has been confused with C. pachecoana, as
caná Volcano. Chiapas is well known for its high               can be seen in herbarium material as well as spec-
plant diversity, but agricultural land use is affect-          imens cited by Sleumer (1967) and González-
ing cloud forest habitat, and many species, such               Villa­rreal (1996). Clethra motozintlana and C. pa­
as C. motozintlana, may face conservation threats.             checoana share fragile, reddish-brown branchlets

Map 1. Geographical distribution of Clethra motozintlana, C. pachecoana, C. tacanensis, and C. zunila. Source: Elevational
model obtained from Global Administrative Areas, gadm.org.

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THREE NOVELTIES IN CLETHRA (CLETHRACEAE: SECT. CUELLARIA) FROM THE CLOUD FORESTS OF SOUTHERN MEXICO AND…

bearing relatively small, bicolored leaves with              5153 (both WIS); El Rosario-Tolimán road be-
appressed, stellate trichomes on the abaxial sur-            tween Huixtla and Motozintla de Mendoza, 15°
face and short inflorescences that are commonly              18’ N, 92° 23’ W, 6 Feb 1990, M. Heath & A. Long
no longer than 15 cm. However, leaves of C. mo­              2127 (GUADA); Cerro Mozotal, 29 km al SW de
tozintlana are subcoriaceous, very variable leaf             Motozintla camino a El Porvenir, 15° 26’ 13” N,
shape, ranging from broadly to narrowly elliptic             92° 28’ 00” W, elev. 2850−3000 m, 26 Feb 1988,
or obovate to obovate-oblong with the base most-             E. Martínez et al. 22405 (MEXU); road from Mo-
ly cuneate and the adaxial surface clearly punc-             tozintla to Niquivil, 7 km from Motozintla de Men-
tate. In contrast, C. pachecoana has coriaceous              doza, Barrio Allende, elev. 1350 m, 7 Feb 1990, P.J.
leaves that are consistently narrowly elliptic, the          Stafford et al. 256 (BM, MEXU); ibid., P.J. Stafford
base of which is truncate to cuneate but conspic-            et al. 265 (BM, MO); from El Rosario to Cerro Bo-
uously and persistently folded for 1−2 mm. The               querón, 2 km S of Ojo de Agua, elev. 1750 m, 8
inflorescences in C. motozintlana are loosely-flow-          Feb 1990, P.J. Stafford et al. 324 (BM); from ejido
ered with ovate bracts that are as long as or some-          Boquerón to Cerro Boquerón, 15° 15’ N, 92° 17’
times shorter than the flowering pedicels; also,             W, elev. 2400 m, 9 Feb 1990, P.J. Stafford et al. 384
the petals are glabrous inside. In contrast, in C.           (BM). Mpio. Unión de Juárez: Volcán Tacaná,
pachecoana the inflorescences are densely-flow-              subiendo por Talquián, elev. 2000 m, 8 Jul 1991,
ered with much more delicate, recurved pedicels;             L.M. González-Villarreal et al. 4151 (IBUG, WIS).
the lanceolate bracts are longer than the flower-
ing pedicels; and the petals are pilose inside. As           Clethra pachecoana Standl. & Steyerm., Field
noted by Sleumer (1967), the length of the bracts            Mus. Nat. Hist., Bot. Ser. 22(4): 259−260. 1940.
relative to that of the pedicels at a given stage is         (Figure 3, map 1).
more or less constant in each species. Geographi-
cally, C. pachecoana is distributed throughout the               Type: Guatemala, Depto. Sacatepéquez, slopes
mountains from central Guatemala to El Salvador              of Volcán de Agua, above Santa María de Jesús,
at the same elevations as C. motozintlana in the             damp forest, elev. 2250−3000 m, 11 Feb 1939
Chiapas-Guatemala border region.                             (fl, fr imature), P.C. Standley 65123 (holotype:
                                                             F984781).
    Additional specimens examined: Mexico,
CHIAPAS, Mpio. Motozintla de Mendoza: N                          Tree small to medium-sized, (3−) 5−15 (−30) m
and W slopes of Cerro Mozotal below microwave                tall, trunk often slender; branchlets slender, fragile,
tower along road from Huixtla to El Porvenir and             4−5 mm in diameter, reddish-brown, puberulent,
Siltepec, elev. 3000 m, 18 Nov 1971, D.E. Breedlove          beginning to shed with age. Leaves coriaceous,
& A.R. Smith 22772 (DS, ENCB); SW side of Cerro              bicolorous; petiole slender, 0.6−1.5 cm long, often
Mozotal, 11 km NW of the jct. of the road to Mo-             dark on dry specimens, puberulent; leaf blades
tozintla to El Porvenir and Siltepec, elev. 2100 m,          mostly narrowly elliptic, sometimes obovate-ob-
29 Dec 1972, D.E. Breedlove & R.F. Thorne 31209              long, (2.5−) 5−10 × (1−) 1.5−3 (−3.5) cm, apex
(CHAPA, DS, ENCB, LL, MICH, MO, NY); NW                      acute to acuminate, rarely obtuse, base truncate
slope of Cerro Malé, 3−4 km W of El Porvenir along           to cuneate, conspicuously and persistently folded
road from Huixtla to Siltepec, elev. 2800 m, 19              for 1−2 mm, margins entire, sometimes slightly
Sep 1976, D.E. Breedlove 40391 (DS, MO); ibid., 26           revolute or serrulate with 6−12 teeth especially
Nov 1981, D.E. Breedlove & B. Bartholomew 55974              on young leaves; adaxial surface dull green, when
(CAS, MO); NE of Cerro Boquerón, from El Ro-                 young covered with tiny, amber-colored, multi-
sario to Niquivil, elev. 2255 m, 29 Nov 1986, D.E.           radiate trichomes, glabrescent to glabrous with
Breedlove & J. Sigg 66148 (CAS); Cerro Mozotal,              age, all the veins furrowed, veinlets sometimes
NW of Motozintla de Mendoza, elev. 2750 m, 7                 evident; abaxial surface pale-brown, glabrous
Jul 1991, L.M. González-Villarreal et al. 4104, 4105,        in appearance but actually covered with dense,
4106 (all IBUG, WIS); Toninchihuan, 2 km antes               minute, appressed, stellate trichomes, some with
de llegar a Niquivil, carretera Motozintla-Niquiv-           an amber-colored center; secondary veins 10−15
il, 15° 15’ 58” N, 92° 13’ 31.3” W, elev. 2620 m,            on each side, slender but somewhat prominent,
14 Mar 2009, L.M. González-Villarreal et al. 5152,           reddish-brown contrasting with the pale surface,

         | OCTUBRE 29 DE 2018 | VOL. 9 | pp. 3–19                                                                  9
LUZ MARÍA GONZÁLEZ-VILLARREAL

                                                                                       B

                                                                             A         C

                                    D      E                                                                                     F

Figure 3. Clethra pachecoana. A. Collection from vicinity of the Pacaya Volcano, Guatemala, A. Molina & A.R. Molina 27651
(MICH). B. Detail, showing the folded leaf bases. C. Leaves, showing both surfaces. D. Branchlet with a paniculate inflores-
cence. E. Detail, showing the flowers. F. Infructescences.

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THREE NOVELTIES IN CLETHRA (CLETHRACEAE: SECT. CUELLARIA) FROM THE CLOUD FORESTS OF SOUTHERN MEXICO AND…

usually arch-ascending or straight, branching and                Common Names: Guatemala: escobo (Stand­
anastomosing to form interconnected arches near              ley 61823), sapotilla (Steyermark 32779, 32793),
the margins, bearing few acicular trichomes, vein-           xiorsh (Huehuetenango) in Flora of Guatemala
lets forming a fine, not clearly distinct network.           (Standley & Williams 1966), zapotillo (Molina &
Inflorescence a short panicle of (4−) 6−8 racemes,           Molina 27651); nance macho in Listado de Árbo­
5−12 cm long, densely-flowered; rachis slender,              les nativos e introducidos de El Salvador (2009).
reddish-brown, puberulous; bracts lanceolate,
(1−) 3−4 mm long, longer than the flowering pedi-               Clethra pachecoana, described as a handsome,
cels, pubescent, early deciduous; pedicels slender,          slender tree up to 30 m tall with a very dense
1−3 mm long, recurved in flower, straight in fruit;          crown, has relatively small, narrowly elliptic
flower buds obovoid; sepals ovate-oblong, 1.8−2.5            leaves that are dark green above and paler beneath
× 1−1.5 mm, apex of the internal lobes acute and             with buff-brown nerves and folded bases. Vigor-
the external obtuse, margins ciliate often whitish,          ous sterile shoots tend to have leaves with serrate
reddish-brown pubescent, canescent-puberulous                margins and an often glabrate surface beneath, as
inside; petals entirely free, obovate to spathulate,         seen in the collections Standley 57781, 61823 and
3.5−4 × 2−3 mm, margins whitish, fimbriate-cili-             Steyermark 32779.
ate, sparingly pilose in the lower part inside; fila-
ments filiform, 2−2.5 mm long, slightly enlarged                 Dry material often exhibits dark petioles. Cle­
at the base, flattened laterally; anthers obcordate,         thra pachecoana has been confused with C. motoz­
ca. 0.6 mm long, with a small appendage at the               intlana and C. tacanensis, but is clearly distinguish-
base; ovary 3-carpellate, 1−1.5 mm in diameter,              able based on the differences and comments given
densely sericeous; style short, 1.8−2 mm long.               at the end of these species and in the key.
Fruit a small, depressed-globose, trigonous cap-
sule, when mature 2−3 × 3−5 mm, on a pedicel                     Specimens examined: El Salvador. Depto.
2−4 mm long. Seeds numerous, flattened, winged               Santa Ana: El Común, N slopes of Santa Ana Vol-
all around, 1.2−1.5 mm in diameter.                          cano, elev. 6500 ft [1981 m], 17 Feb 1958, P.H.
                                                             Allen & R. Armour 6795 (F); ibid., facing Lake
   Distribution and Habitat: Clethra pachecoa­               Coatepeque, elev. 6000−7000 ft [1829−2134 m],
na is distributed throughout a volcanic range at             25 Sep 1958, P.H. Allen 6956 (F, NY, TEX, US);
(1600–) 2000 to 3100 m in elevation, mostly in               ibid., P.H. Allen 6959 (NY); Volcán de Santa Ana,
Guatemala but extending to the southeast into                elev. 1590−2340 m, 19 Feb 1946, M.C. Carlson 709
El Salvador at its known southern limit. It reach-           (F). Guatemala. Depto. Baja Verapaz: Unión
es the highest elevations on the Acatenango, de              Barrios, km 150 de la carretera a Cobán, 4 Nov
Agua, Atitlán, and Pacaya Volcanoes in Guate-                1973, E. Contreras 11572 (MEXU). Depto. Chi-
mala and is also very common on the Santa Ana                maltenango: Sie­rra de Santa Elena, Jun 1937, W.
Volcano in El Salvador, the highest volcano in               Seifriz 17 (US); Chichavac finca, elev. 2400−2700
the country. It inhabits cloud forest, humid oak-            m, Nov-Dec 1930, A.F. Skutch 85 (MICH, US);
pine forest, and open grassy slopes, where it is             ibid., 23 Nov 1933, A.F. Skutch 701 (F, MICH,
often associated with Abies guatemalensis, Chi­              US); region of Las Calderas, elev. 1800−2100 m,
rathodendron pentadactylon, Cupressus lusitanica,            22 Nov 1938, P.C. Standley 57781 (F); ibid., P.C.
Parathesis spp., Pinus spp., and Synardisia venosa.          Standley 57807 (F, L-fragment, U, US); slopes of
Clethra pachecoana is one of the trees that fre-             Volcán de Acate­nango, above Las Calderas, elev.
quently comes up after forests are cut. In fact,             2400−2700 m, 3 Jan 1939, P.C. Standley 61823
the genus Clethra has a reputation for being pio-            (F). Chimaltenango/Sacatepéquez: Volcán de
neer plants in secondary forest.                             Acatenango, elev. 2400 m, 27 Nov 1993, G. Castil­
                                                             lo et al. 2035 (F); ibid., 13 Feb 1993, Luarca & Léliz
   Phenology: Clethra pachecoana was collected               2665 (MEXU); ibid., elev. 2500−3100 m, 15 Sep
with flowers from November to February, and                  2004, A.M. Ortiz 12 (MEXU). Chimaltenango/
with fruits in April.                                        Quiché/Sololá: Cerro Chichoy near Chichoy, ca.
                                                             14° 48’ N, 91° W, elev. 2800 m, 26−27 Jan 1949,
                                                             L.O. Williams & A. Molina R. 15360 (F). Depto.

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LUZ MARÍA GONZÁLEZ-VILLARREAL

Escuintla: Vicinity of Volcán Pacaya, elev. 2000       dish-brown tomentum on both sides; petiole
m, 28 Sep 1972, A. Molina & A.R. Molina 27651          somewhat thick, (0.6−) 1−1.5 cm long, red-
(ENCB, F, MICH, U). Depto. Guatemala: F.N.             dish-brown, with hirsute indumentum, glabres-
“La Aurora”, 1940, I. Aguilar 450 (F); Volcán Pa-      cent with age; leaf blades broadly elliptic to obo-
caya, above Las Calderas, elev. 1800−2400 m, 30        vate or oblong, (3.5−) 5−9 × (1.5−) 2−4.5 cm,
Nov 1938, P.C. Standley 58333 (F, NY); slopes of       apex rounded to obtuse or acute, bases consis-
Volcán Pacaya, between San Francisco Sales and         tently folded up to 5 mm, margins entire; adaxi-
the base of the active cone, elev. 1800−2300 m, 20     al surface dull, covered mostly with multiradiate
Dec 1940, P.C. Standley 80689 (F). Depto. Jalapa:      trichomes mixed with stellate trichomes of 6−9
montaña de Xalapán, elev. 2600 m, 12 May 1994,         rays, sessile-fasciculate trichomes of 6−8 rays and
Marroquín 2200 (MEXU); between Miramundo               stipitate-fasciculate trichomes of 2−4 rays, the
and summit of Montaña Miramundo between Ja-            rays long and erect or twisted, surface glabrescent
lapa and Mataquescuintla, 6 mi S of Miramundo,         but minutely punctate with age, secondary veins
elev. 2000−2500 m, 5 Dec 1939, J.A. Steyermark         reddish-tomentose, bearing acicular trichomes;
32779 (F); Montaña Miramundo, between Mira-            abaxial surface dull, harsh to the touch, pale to
mundo and Buena Vista, between Jalapa and Lago         reddish-brown tomentose, covered with min-
Ayarza, elev. 2000−2200 m, 6 Dec 1939, J.A. St­        ute, appressed, stellate trichomes as an inferior
eyermark 32793 (F). Depto. Sololá: southern            stratum, superior stratum with dispersed, stipi-
slopes of Volcán Atitlán, elev. 1700−3800 [?] m,       tate-fasciculate trichomes of 2−4 rays, mixed with
11 Jun 1942, J.A. Steyermark 47393 (F).                sessile-fasciculate trichomes of 6−8 rays, the rays
                                                       long and erect, twisted or somewhat collapsed,
Clethra tacanensis Lundell ex L.M.González, sp.        sometimes overlapping but not tangled; sec-
nov. (Figure 4, map 1).                                ondary veins 10−15 on each side, slender, red-
                                                       dish-brown contrasting with the paler surface,
   Type: Mexico, CHIAPAS, Mpio. Unión de               arch-ascending or straight, branching and anas-
Juárez, Volcán Tacaná, Chiquihuite, elev. 2800 m,      tomosing to form interconnected arches near
27 Mar 1939 (fr), E. Matuda 2820 (holotype: LL;        the margins, bearing long acicular trichomes,
isotypes: A, F, K, LL, MICH (2), MO, NY).              veinlets very fine lines forming a network. Inflo-
                                                       rescence a short panicle of 6−8 racemes, 5−10 cm
    Clethra tacanensis is morphologically similar      long, densely-flowered; rachis somewhat thick,
to C. salvadorensis Britton but it differs in having   reddish-brown, densely hirsute-tomentose; bracts
coriaceous leaf blades with the adaxial surface        subulate, somewhat conduplicate, 1.5−2.5 (−4) ×
mostly covered by multiradiate trichomes (vs.          1 mm, as long as the flowering pedicels or slightly
acicular trichomes) evidently punctate with age;       longer, densely reddish-brown tomentose, early
and the abaxial surface reddish-brown, harsh-to-       deciduous; pedicels short, 1.5−2 (−2.5) mm long,
mentose (vs. pale or yellowish, soft tomentose)        straight, tomentose; sepals ovate to ovate-oblong,
with dispersed, sessile-fasciculate trichomes of       2−3 × 1−1.5 mm, apex of the internal lobes obtuse
6−8 rays and stipitate-fasciculate trichomes of        to rounded, the external with long-ciliate mar-
2−4 rays (vs. mostly acicular trichomes) as the        gins, glabrous inside; petals entirely free, obo-
superior stratum. The inflorescences are short         vate to spathulate, (2−) 3−3.5 × 1.5 mm, margins
panicles 5−10 cm long (vs. elongate racemes            fimbriate-ciliate, glabrous or sparingly pilose in
10−15 (−20) cm long). The bracts 1.5−2.5 (−4)          the lower part inside; filaments filiform, 1.5−2
mm long, as long as the flowering pedicels or          mm long, slightly enlarged at the base, flattened
slightly longer [vs. (3.5−) 4−5 (−8) mm long, lon-     laterally; anthers sagittate, ca. 0.8 mm long, with
ger than the flowering pedicels].                      a small appendage at the base; ovary 3-carpellate,
                                                       1−2 mm in diameter, densely sericeous; style
   Tree small to medium-sized, 6−8 m tall;             slender, short, 1.8−2.2 mm long. Fruit a small,
branchlets somewhat thick, 5−10 mm in diam-            depressed-globose, trigonous capsule, when ma-
eter, the young ones hirsute with reddish-brown        ture, 3−4 × 4−6 mm, on a pedicel 2−2.5 mm long.
trichomes, beginning to shed with age. Leaves          Seeds numerous, flattened, winged all around,
coriaceous, bicolored, when young with red-            2−2.5 × 1−1.8 mm, including the wing.

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THREE NOVELTIES IN CLETHRA (CLETHRACEAE: SECT. CUELLARIA) FROM THE CLOUD FORESTS OF SOUTHERN MEXICO AND…

Figure 4. Clethra tacanensis. A. Branchlet with an infructescence. B. Detail, showing the folded leaf bases. C. Infructescen-
ce. D. Fruit. Drawn by Manuel Peinado.

          | OCTUBRE 29 DE 2018 | VOL. 9 | pp. 3–19                                                                          13
LUZ MARÍA GONZÁLEZ-VILLARREAL

   Distribution and Habitat: Clethra tacanensis          There is another Matuda collection (2753)
is endemic to the Tacaná Volcano at the south-        from the Departamento de Soconusco, Mu-
ernmost end of the Mexico-Guatemala border,           nicipio Unión de Juárez, Volcán Tacaná, elev.
where it grows in cloud forest at elevations be-      1400 m, 17−30 Mar 1939 at A, B, BEROL, CAS, F,
tween 2500−2800 (−3000) m. Breedlove’s (1981)         L-fragm., LL, MICH, MO, NY, UC, and US. Orig-
statement that “there are many local endemics in      inally identified as Clethra pachecoana, this gath-
this region, especially in the higher-elevation as-   ering bears Lundell’s annotation “C. tacanensis
sociations about Volcán Tacaná” is supported by       sp. nov.” However, Sleumer (1967: 162) noticed
the description of this new species.                  that it represents another species and cited it as
                                                      C. lanata Mart. & Gal. In fact, Matuda 2753 is C.
   Phenology: Clethra tacanensis was collected        salvadorensis, with narrower leaves growing in
with fruits in February and March.                    the same region but at lower elevation.

   Etymology: The specific epithet refers to the         Clethra lanata is distributed from Michoacán
Tacaná Volcano and recognizes its rich diversity      to Oaxaca, whereas C. salvadorensis is a Central
and high endemism. The area was declared a Bio-       American species that reaches Mexico only in the
sphere Reserve in 2003.                               southern part of Chiapas.

   Clethra tacanensis has been misidentified as          Clethra tacanensis is morphologically similar
C. pachecoana, a species of series Tomentellae,       to C. salvadorensis (also of series Tomentosae) in
likely because they share relatively similarly        having tomentose leaves with a consistently fold-
shaped leaves that are somewhat small in size         ed bases. However, leaves of C. salvadorensis are
(up to 10 cm long) with strongly folded bases,        subcoriaceous with the adaxial surface mainly
as well as short inflorescences of densely-flow-      with acicular trichomes and abaxial surface pale
ered panicles with 6−8 racemes, mostly less than      or yellowish, soft-tomentose, with mostly acic-
10 cm long. In fact, the Steyermark’s collec-         ular trichomes, mixed with fascicular trichomes
tions numbered 36205 and 36215 were cited by          of 2 rays as the superior stratum. In contrast, C.
Sleumer (1967: 121) as C. pachecoana. However,        tacanensis has coriaceous leaves with the adaxial
C. tacanensis is placed here under series Tomen­      surface mostly with multiradiate trichomes and
tosae, being readily distinguished by having red-     abaxial surface reddish-brown, harsh-tomen-
dish-brown, tomentose leaves with the abaxial         tose, with sessile-fasciculate trichomes of 6−8
surface harsh to the touch. The vestiture is com-     rays mixed with stipitate-fasciculate trichomes
prised of two strata.                                 of 2−4 rays as superior stratum. The inflores-
                                                      cences of C. salvadorensis are mostly elongate ra-
   This new species was recognized in 1939 for        cemes, 10−15 (−20) cm long with bracts (3.5−)
the first time by Cyrus L. Lundell, who named         4−5 (−8) mm long, longer than the flowering
it Clethra tacanensis. However, a formal de-          pedicels; flowers densely pilose inside. In con-
scription was not forthcoming, this name being        trast, C. tacanensis has short panicles, 5−10 cm
known only on herbarium sheets such as Matuda         long with bracts 1.5−2.5 (−4) mm long, as long
2820, the type. The collections Matuda 2820 at        of the flowering pedicels or longer; flowers gla-
A, F, MICH, and NY and Matuda S-231 at A and          brous or sparingly pilose inside. Clethra salva­
MICH also were labeled by Sleumer in 1965 as          dorensis is a widespread species occurring from
“Clethra tacanensis Lundell, not published” with      southern Chiapas, Mexico through Guatemala
the additional note: “Clethra pachecoana x lanata.    to Costa Rica. It grows at elevations from 1000
Cl. tacanensis Lundell not published.” It is pos-     to 1800 m, on forest margins, open pine and/
sible that Matuda’s collection numbered S-231         or Quercus woods, coffee plantations and semi-­
may be part of the type, bearing as it does oth-      evergreen forest.
erwise identical data and fruits at the same stage
of maturity.                                             In the Flora of Guatemala (Standley & Wil-
                                                      liams 1966) and Flora Mesoamericana (Vickery
                                                      2004), Clethra lanata and C. salvadorensis were

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THREE NOVELTIES IN CLETHRA (CLETHRACEAE: SECT. CUELLARIA) FROM THE CLOUD FORESTS OF SOUTHERN MEXICO AND…

treated as synonyms of C. mexicana, a species                coriaceous, concolorous; petiole slender, (0.7−)
res­tricted to central Mexico that is characterized          1−1.5 cm long, hirsute, glabrescent with age;
by coriaceous, noticeably ferrugineous-tomen-                leaf blades elliptic-oblong or obovate, 4−7.5 ×
tose leaves; racemose, red-brown inflorescen­                2−4 cm, apex obtuse to rounded, base cuneate,
ces with a stout rachis; and large fruits (up to ca.         scarcely folded, margins serrate; adaxial surface
10 mm in diameter). However, Sleumer (1967)                  shiny green, covered mostly with sessile-fascicu-
accepted C. lanata, placing C. salvadorensis in its          late trichomes of 2−4 rays, mixed with dispersed
long list of synonyms, and it was also recognized            acicular trichomes, these erect, ca. 0.8 mm long,
by Hamilton (2001) for Flora of Nicaragua. Ac-               somewhat thick, and some stellate trichomes of
cording to González-Villarreal (2007), C. lanata             4−7 rays; abaxial surface reddish-brown tomen-
and C. mexicana are completely different spe-                tose, harsh to the touch, covered with minute,
cies, and C. salvadorensis is the only synonym               appressed, stellate trichomes as an inferior stra-
of C. lanata. Nevertheless, as the result of recent          tum, superior stratum with uniformly scattered,
studies, the author has concluded that there are             acicular trichomes, these long, erect, amber-col-
three different taxa.                                        ored, mixed with some stipitate-fasciculate tri-
                                                             chomes of 2−4 (−5) rays, the rays erect, all veins
   Additional specimens examined: Guatemala.                 raised; secondary veins 10−15 on each side,
DEPTO. SAN MARCOS, Mpio. Cacahoatan, be-                     reddish-brown tomentose, contrasting with the
tween La Vega ridge along Río Vega, and north-east           paler surface, usually arch-ascending, branching
slopes of Volcán Tacaná, to 3 mi from Guatemala-             before the margin, veinlets forming a fine dis-
Me­xico boundary, in vicinity of San Rafael, elev.           tinct network. Inflorescence a short panicle of 5
2500−3000 m, 20 Feb 1940, J.A. Steyermark 36205              racemes, 3−6 cm long, densely-flowered; rachis
(F); ibid., J.A. Steyermark 36215 (F, MICH). Me­             somewhat thick, densely yellowish-brown to-
xico. CHIAPAS, Mpio. Unión de Juárez, Volcán                 mentose, covered with acicular trichomes and
Tacaná, Chiquihuite, elev. 2800 m, 27 Mar 1939,              fasciculate trichomes of 2−3 rays; bracts deltoid,
E. Matuda S-231 (A, ARIZ, MICH).                             1−1.2 mm long, shorter than the flowering ped-
                                                             icels, densely tomentose, early deciduous; ped-
Clethra zunila L.M.González, sp. nov. (Figure 5,             icels short, 1.5−2.5 mm long, straight, densely
map 1).                                                      tomentose; sepals ovate-oblong, 2−3 mm long,
                                                             apex of the internal lobes obtuse and the exter-
   Type: Guatemala, Depto. Quetzaltenango,                   nal with ciliate margins, densely pubescent on
Volcán Zunil, elev. 2500−3800 m, 22 Jan 1940                 both surfaces; petals entirely free, spathulate,
(fl), J.A. Steyermark 34651 (holotype: F1043249;             2.5−3 × 1.5−1.8 mm, margin whitish, long-cili-
isotype: MICH).                                              ate, fimbriate, sparingly long-haired on the low-
                                                             er part inside; filaments filiform, 1−2 mm long,
   Clethra zunila is morphologically similar to C.           slightly enlarged at the base, flattened laterally;
lanata, but it differs by having the adaxial leaf            anthers sagittate, less than 1 mm long, with a
surface mostly with sessile-fasciculate trichomes            small appendage at the base; ovary 3-carpellate,
of 2−4 rays mixed with dispersed acicular and                1.5−2 mm wide, densely long-sericeous; style
some stellate trichomes of 4−7 rays (vs. most-               slender, short, 1.3−1.5 mm long. Fruit and seeds
ly acicular trichomes). The inflorescences are               unknown.
short panicles 3−6 cm long (vs. elongated pani-
cles 10−20 (−30) cm long). The bracts are del-                  Distribution and Habitat: Clethra zunila is
toid, 1−1.2 mm long, shorter than the flowering              endemic to the Zunil Volcano [Pico Zunil] in
pedicels (vs. lanceolate or subulate, 2−6 (−8) mm            western Guatemala. The type was collected at
long, longer than the flowering pedicels).                   elevations of 2500−3800 [? possibly 3500] m,
                                                             where it was found in dry thickets above a sulfur
   Shrub or small tree, 6 m tall; branchlets slen-           terrace. Pico Zunil has been documented to sup-
der, 5−7 mm in diameter, reddish-brown, hir-                 port relatively high numbers of endemic species
sute tomentose, trichomes mostly acicular, up to             (Quedensley & Véliz 2009).
1.5 mm long, beginning to shed with age. Leaves

         | OCTUBRE 29 DE 2018 | VOL. 9 | pp. 3–19                                                             15
LUZ MARÍA GONZÁLEZ-VILLARREAL

Figure 5. Clethra zunila. A-B. Branchlets with paniculate inflorescences. C. Leaf, showing the abaxial surface. D. Flower.
Drawn by Manuel Peinado.

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THREE NOVELTIES IN CLETHRA (CLETHRACEAE: SECT. CUELLARIA) FROM THE CLOUD FORESTS OF SOUTHERN MEXICO AND…

   Phenology: Clethra zunila was collected in                the transition zones from pine forest to tropical
flower in January.                                           deciduous forest, pine-oak forest, gallery forest,
                                                             and cloud forest (González-Villarreal 1996).
    Etymology: The species is named after the Zu-
nil Volcano, where it is endemic.                            Key to distinguish some species of Clethra
                                                             from the montane cloud forests of
   Steyermark described this plant in the field              southern Mexico and western Guatemala
as “a shrub 10 ft tall. Leaves subcoriaceous, firm,
rich shining green above with buff-brown mid-                1. Abaxial leaf surface pale-brown to
rib, gray-brown beneath with prominent pale                     yellowish or somewhat ochraceous,
orange-brown midrib and lateral nerves.” Origi-                 smooth to the touch, glabrous in
nally, the type collection was labeled as “Clethra              appearance but covered only with
tacanensis Lundell, sp. nov.” and later by Sleumer              dense, minute, appressed, stellate
(the specimen at MICH) as “Clethra, possibly hy-                trichomes as the inferior stratum,
brid pachecoana x lanata.”                                      sometimes with few acicular tricho-
                                                                mes on the main veins........series Tomentellae
   Steyermark 34999 from Zunil, collected at
2500 m and cited by Sleumer (1967: 162−163)                  2. Leaves very variable in shape,
as Clethra lanata, probably is part of the same C.              broadly to narrowly elliptic, obova-
zunila population, but it was not seen.                         te, or sometimes obovate-oblong,
                                                                the base mostly cuneate, narrowed
   Clethra zunila is morphologically similar to C.              to abruptly rounded, not folded;
lanata both in the series Tomentosae. Although                  adaxial leaf surface punctate and
the pubescence on the abaxial leaf surface of                   abaxial surface with dispersed,
the first is harsh to the touch and soft in the se­             reddish, tiny, multiradiate trichomes
cond, they share acicular trichomes mixed with                  in addition to the appressed, stellate
stipitate-fasciculate trichomes of 2−4 (−5) rays)               trichomes; inflorescences loosely-
as the superior stratum. However, leaves of C.                  flowered, bracts ovate, somewhat
lanata are subcoriaceous with petiole 1.5−2.5                   conduplicate, 1.5−2.5 (−3) mm long,
(−4.5) cm long. The adaxial leaf surface bearing                as long as the flowering pedicels or
mostly acicular trichomes mixed with stellate                   shorter; sepals (1.5−) 2.5−3 (−3.5) ×
trichomes. In contrast, C. zunila has coriaceous                (1−) 1.5−2 mm; petals glabrous insi-
leaves with petiole 1−1.5 cm long. The adaxial                  de.............................................. C. motozintlana
leaf surface mostly with sessile-fasciculate tri-
chomes of 2−4 rays mixed with dispersed acicu-               2. Leaves mostly narrowly elliptic, so-
lar and some stellate trichomes of 4−7 rays. The                metimes obovate-oblong, the base
inflorescenses of C. lanata are elongate panicles               truncate to cuneate, persistently fol-
of 5−10 racemes, 10−20 (-30) cm long with lan-                  ded; adaxial leaf surface not puncta-
ceolate or subulate bracts, 2−6 (-8) mm long,                   te and abaxial surface with only
longer than the flowering pedicels; flowers with                appressed, stellate trichomes; inflo-
the filaments (1.5−) 2−3 mm long and the style                  rescences densely-flowered, bracts
(1.5−) 2−3 (−4) mm long. In contrast, C. zunila                 lanceolate, (1−) 3−4 mm long, lon-
has short panicles of 5 racemes, less than 10 cm                ger than the flowering pedicels; se-
long with deltoid bracts, 1−1.2 mm long, short-                 pals 1.8−2.5 × 1−1.5 mm; petals spa-
er than the flowering pedicels; flowers with the                ringly pilose inside....................C. pachecoana
1−2 mm long and style 1.3−1.5 mm long. Geo-
graphically, C. lanata occurs mostly in the Sierra           1. Abaxial leaf surface reddish-brown
Madre del Sur, from Michoacán to Oaxaca over                    tomentose, harsh to the touch, cove-
a wide range of elevation, from 350 to 2300 m,                  red with dispersed, fasciculate and
in a variety of habitats. At the lower elevation                acicular trichomes, as the superior
in the Curatella-Bursonima savannah and also in                 stratum in addition to the appres-

         | OCTUBRE 29 DE 2018 | VOL. 9 | pp. 3–19                                                                             17
LUZ MARÍA GONZÁLEZ-VILLARREAL

     sed, stellate trichomes as the infe-                                  shorter than the flowering pedicels;
     rior stratum, with dense acicular tri-                                sepals densely pubescent inside;
     chomes on the main veins.................                             style 1.3−1.5 mm long........................C. zunila
     ...............................................series Tomentosae

3. Trichomes of the abaxial leaf surfa-                                 Acknowledgments
   ce stipitate-fasciculate of 2−4 rays,
   mixed with sessile-fasciculate of                                    This paper is the result of a M.S. degree and
   6−8 rays, as the superior stratum;                                   more recently of the revision of the genus Cle­
   adaxial leaf surface dull, covered                                   thra in Central America, both carried out at
   mostly with multiradiate trichomes                                   the University of Wisconsin-Madison, U.S.A.
   mixed with stellate trichomes of 6−9                                 in conjunction with the University of Guadala-
   rays, sessile-fasciculate trichomes of                               jara, Mexico. I wish to express my gratitude to
   6−8 rays and stipitate-fasciculate tri-                              these institutions for their support in providing
   chomes 2−4 rays, with age puncta-                                    facilities. I am especially thankful to my former
   te; leaves with margins entire and                                   major professor Dr. Hugh H. Iltis† for his sup-
   base strongly folded; bracts subu-                                   port and guidance throughout so many years; to
   late, 1.5−2.5 (−4) mm long, as long                                  Kenneth Cameron, Director of the Wisconsin
   as the flowering pedicels or slightly                                State Herbarium (WIS), for letting me work in
   longer; sepals glabrous inside; style                                the herbarium during academic stays; Theodore
   1.8−2.2 mm long.......................... C. tacanensis              S. Cochrane for his careful review and correc-
                                                                        tions made to the English manuscript; and Mark
3. Trichomes of the abaxial leaf surfa-                                 A. Wetter for assisting with loans. I am grateful
   ce acicular, mixed with some stipita-                                to the curators of herbaria cited for use of their
   te-fasciculate of 2−4 (−5) rays, as the                              collections. Thanks also to Sergio Zamudio and
   superior stratum; adaxial leaf surface                               Victor Steinman for useful comments and sug-
   shiny, covered mostly with acicular                                  gestions. To Manuel Peinado for the line draw-
   trichomes mixed with stellate tri-                                   ings and Miguel A. García-Martínez for the map.
   chomes of 4−7 rays and sessile-fas-                                  Finally, I thank two anonymous reviewers for
   ciculate trichomes of 2−4 rays, with                                 providing reviews of the manuscript.v
   age not punctate; leaves with mar-
   gins serrulate and base slightly fol-
   ded; bracts deltoid, 1−1.2 mm long,

Literature cited                                Calderón de Rzedowski, G. 2001.              cales) with notes on biogeog-
                                                   Clethraceae. Pp. 522−524. In:             raphy: evidence from plastid
Anónimo. 2009. Árboles nativos                     Rzedowski, G.C. & J. Rzedows-             and nuclear DNA sequenc-
   e introducidos de El Salvador.                  ki (eds.). Flora fanerogámi­              es. Int. J. Plant Sci. 164(6):
   Parte 1: Angiospermae – Fa-                     ca del Valle de México, 2a. ed.           997−1006. Available from:
   milias A a L. Englera 29: 97.                   Instituto de Ecología, A.C. y                                   Conocimiento y Uso de la Bio-           González-Villarreal, L.M. 1996.
   17/04/2018.                                     diversidad, Pátzcuaro (Micho-             Clethra (Clethraceae) Section
Breedlove, D.E. 1981. Introduction                 acán), México. 1406 pp.                   Cuellaria in Mexico: Taxo­
   to the Flora of Chiapas. Pp.                 Fior, S., P.O. Karis & A.A. Ander-           nomy, ecology and biogeo­
   1−35. Flora of Chiapas. Part 1.                 berg. 2003. Phylogeny, taxon-             graphy. M.S. thesis. University
   California Academy of Scienc-                   omy, and systematic position              of Wisconsin-Madison, Madi-
   es, San Francisco, CA, USA.                     of Clethra (Clethraceae, Eri-             son, WI, USA. 382 pp.

18                                                                                                | OCTUBRE 29 DE 2018 | VOL. 9 | pp. 3–19
THREE NOVELTIES IN CLETHRA (CLETHRACEAE: SECT. CUELLARIA) FROM THE CLOUD FORESTS OF SOUTHERN MEXICO AND…

González-Villarreal, L.M. 2006.                        Pool & O.M. Montiel (eds.).         Standley, P.C. & L.O. Williams. 1966.
  Novelties in Clethra (Clethra-                       Monogr. Syst. Bot. Missouri Bot.       Clethraceae. Flora of Guate-
  ceae) from Mexico. ibugana                           Gard. 85(1): 615−616.                  mala. Fieldiana: Botany 24(8):
  13(1): 11−25. Available from:                     Quedensley, T.S. & M.E. Véliz Pérez.      74−81.
                                     (Lamiaceae), a new species             A Global Directory of Public
González-Villarreal, L.M. 2007.                        from a cloud forest in western         Herbaria and Associated Staff.
  Foliar trichome variations in                        Guatemala. J. Bot. Res. Inst.          New York Botanical Garden’s
  Clethra Subsect. Cuellaria (Cle-                     Texas 4(1): 27−31. Available           Virtual Herbarium. Available
  thraceae) from Mexico. ibu­                          from: . 29/09/2015.
  from:                                   ia clethracearum. Bot. Jahrb.          Flora Mesoamericana on-line.
Hamilton, C.W. 2001. Clethraceae.                      Syst. 87(1): 36−116; 87(2):            . 12/11/15.

         | OCTUBRE 29 DE 2018 | VOL. 9 | pp. 3–19                                                                             19
20   | OCTUBRE 29 DE 2018 | VOL. 9
Nuevos registros de ocho especies de Tigridieae
(Iridaceae) en México
Guadalupe Munguía-Lino1, 2, , Arturo Castro-Castro4 y Aarón Rodríguez2, 3
                                  @

                                                                                                                       Citar
1
  Cátedras CONACYT – Universidad de              Resumen                                Abstract
Guadalajara.                                     Se presentan localidades nuevas        We report new locations of Sessi-
2
  Herbario Luz María Villarreal de Puga,
                                                 de Sessilanthera latifolia, Tigridia   lanthera latifolia, Tigridia alpestris
Instituto de Botánica.
3
  Laboratorio Nacional de Identificación         alpestris subsp. obtusa, T. catari-    subsp. obtusa, T. catarinensis, T.
y Caracterización Vegetal, Consejo               nensis, T. gracielae, T. immaculata,   gracielae, T. immaculata, T. po-
Nacional de Ciencia y Tecnología,                T. potosina, T. pugana y T. suarezii   tosina, T. pugana, and T. suarezii
Departamento de Botánica y Zoología,             en México. Se comentan los rasgos      in Mexico. The distinctive cha-
Centro Universitario de Ciencias Bioló-          distintivos de cada taxón y se am-     racteristics of each species are
gicas y Agropecuarias, Universidad de
                                                 plía el conocimiento de su distri-     highlighted and the knowledge
Guadalajara. Apartado postal 1−139,
Zapopan 45101, Jalisco, México.                  bución.                                about its geographical distribution
                                                                                        is updated.
4
 Cátedras CONACYT – Centro Inter-                Palabras clave: distribución, en-
disciplinario de Investigación para el           demismo, México, microedemis-          Key words: distribution, ende-
Desarrollo Integral Regional. Unidad
                                                 mo, riesgo, Sessilanthera, Tigridia.   mism, Mexico, microendemism,
Durango, Instituto Politécnico Nacional,
Sigma 119, Fracc. 20 de noviembre II,                                                   threatened, Sessilanthera, Tigridia.
Durango 34220, Durango, México.

@   gmlinno@gmail.com

Introducción

L  a tribu Tigridieae B.M.Kittel (Iridaceae) es
   un grupo monofilético (Rodríguez & Sytsma
2006; Goldblatt et al. 2008). En México está re-
                                                                    Durante el proceso de revisión de material de
                                                                    herbario y de recolecciones en campo para reali-
                                                                    zar un análisis biogeográfico de la tribu en Norte-
presentada por 12 géneros y 60 especies (34.8 %                     américa, se encontraron ejemplares procedentes
de su diversidad total), de las cuales 46 son endé-                 de localidades no conocidas.
micas, por lo que el país es considerado como un
centro de diversificación del grupo (Rodríguez                          El objetivo del presente trabajo es dar a co-
& Systma 2006; Munguía-Lino et al. 2015). Tigri-                    nocer localidades nuevas de Sessilanthera latifo-
dia es el género más ampliamente distribuido, ya                    lia (Weath.) Molseed & Cruden, Tigridia alpestris
que está presente en 25 entidades federativas del                   Molseed subsp. obtusa Molseed, T. catarinensis
territorio. Cipura campanulata Ravenna, Nema-                       Cruden, T. gracielae Aarón Rodr. & Ortiz-Cat., T.
stylis tenuis (Herb.) S.Watson y Tigridia pavonia                   immaculata (Herb.) Ravenna, T. potosina López-
(L.f.) DC. tienen las áreas de distribución más                     Ferr. & Espejo, T. pugana Aarón Rodr. & Ortiz-
extensas. Por el contrario, 30 taxa se encuentran                   Cat. y T. suarezii Aarón Rodr. & Ortiz-Cat. en los
restringidos a uno o dos estados, de los que 14 se                  estados de Guerrero, Jalisco, México, Morelos,
conocen de uno o dos municipios y tienen me-                        Oaxaca, Puebla y San Luis Potosí.
nos de 10 registros (cuadro 1).
                                                                       Adicionalmente, se proporciona una descrip-
   El conocimiento de las especies de Tigridieae                    ción morfológica actualizada de cada especie. La
y su distribución en México son incompletos.                        ampliación del conocimiento relativo a la distri-

          | OCTUBRE 29 DE 2018 | VOL. 9 | pp. 21–46                                                                         21
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