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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.
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 | OCTUBRE 29 DE 2018 | VOL. 9 1
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 2 | OCTUBRE 29 DE 2018 | VOL. 9
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 | OCTUBRE 29 DE 2018 | VOL. 9 | pp. 3–19 3
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. 4 | OCTUBRE 29 DE 2018 | VOL. 9 | pp. 3–19
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, | OCTUBRE 29 DE 2018 | VOL. 9 | pp. 3–19 5
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. 6 | OCTUBRE 29 DE 2018 | VOL. 9 | pp. 3–19
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. | OCTUBRE 29 DE 2018 | VOL. 9 | pp. 3–19 7
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. 8 | OCTUBRE 29 DE 2018 | VOL. 9 | pp. 3–19
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. 10 | OCTUBRE 29 DE 2018 | VOL. 9 | pp. 3–19
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. | OCTUBRE 29 DE 2018 | VOL. 9 | pp. 3–19 11
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. 12 | OCTUBRE 29 DE 2018 | VOL. 9 | pp. 3–19
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 14 | OCTUBRE 29 DE 2018 | VOL. 9 | pp. 3–19
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. 16 | OCTUBRE 29 DE 2018 | VOL. 9 | pp. 3–19
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
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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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