Mexican botany Act Print ISSN 0187-7151

Act. Bot. Mex No.91 April Patzcuaro. 2010

Articles

Taxonomic revision of Agave potatorum Zucc complex. (Agavaceae): new taxa and neotipificación

Taxonomic review of the Agave potatorum Zucc. (Agavaceae) complex: new taxa and neotypification

Joshua Abisai Garcia-Mendoza

National Autonomous University of Mexico, Botanical Garden, Institute of Biology, Apdo. Postal 70-614, 04510 Mexico, DF Mexico. [email protected]

Received in May 2009 Accepted in February 2010

SUMMARY

Agave taxonomic potatorum complex belonging to the group is reviewed Hiemiflorae are described and illustrated two new taxa to science. Agave nussaviorum and A. nussaviorum subsp. deltoidea the state of Oaxaca. The new species is compared with Agave seemanniana Jacobi and A. potatorum Zucc. Information on distribution, uses, phenology and a key to the separation of the taxa provided.

Keywords: Agavaceae, Agave nussaviorum, potatorum Agave, Mexico, Oaxaca.

ABSTRACT

The Agave potatorum complex, of group Hiemiflorae, is revised. Two new taxa from Oaxaca, Agave nussaviorum and A. nussaviorum subsp. deltoidea, are Described and illustrated. The new species is Agave seemanniana Mexican botany Act - Taxonomic review of Agave potatorum Zucc complex. (Agavaceae): new taxa and neotipificación

Compared With Jacobi and A. potatorum Zucc. ITS Information Concerning distribution, ethnobotany, and phenology, and an identification key to the taxa of the complex are provided.

Key words: Agavaceae, Agave nussaviorum, potatorum Agave, Mexico, Oaxaca.

During the preparation of taxonomic tried Agave (Agavaceae) for the Flora of the Tehuacan Valley-Cuicatlán, it was considered necessary to address the delimitation of Agave potatorum complex. The species belongs to Hiemiflorae group (Gentry, 1982) characterized by having leaves ovate to broadly lanceolate, blue-green, with serrated margin to racemosa paniculate inflorescence, flowers grouped in umbels, sometimes congested, densely bracteolate, tépalos longer than the inserted tube and half filaments or above the tube. Plants usually without suckers and preferably reproduction by seed. The Hiemiflorae group contains 13 species distributed in southern Mexico (Puebla and Veracruz) to Nicaragua.

Agave potatorum was described by Zuccarini in 1832, from Mexico shipped by Karwinski and cultivated in the Botanical Garden of Munich plant. Subsequently, Zuccarini (1833) provided information on the growth habit, leaf measures, shape, size of the teeth and inflorescence branching (according to Karwinski). The area of origin of the species is not known; Zuccarini mentions "Crescit in Mexican empire '", although it is highly likely to have been collected near Tehuacan, Puebla, since it was published along with two other agaves the same valley: Agave macroacantha Zuce, and A. karwinskii Zucc. Then more they related to the same species names, some with pictures, but all based on cultivated plants in various European gardens were published. Gentry (1982) met under Agave potatorum a highly variable number of plants growing in the states of Oaxaca and Puebla; therefore, finding that correlated morphological characters of consistently applied a broad concept for the species. The aforementioned author studied wild populations of Agave potatorum broad sense, that collected in 1952, 1963, 1965, 1967 and 1976 seasons in which he herborizó 13 copies with their duplicates. These collections, coupled with another 18 more, deposited in herbaria DES, ILL, MEXU, MICH, MO and US, were not enough to taxonomically define the observed entities, which prevented him segregate taxa at specific level or subspecific.

After work Gentry numerous field trips to observe plants in their habitats were carried herborizar a greater number of copies and ex situ specimens grown in the Botanical Garden of the Institute of Biology of the National Autonomous University of Mexico, to assess its behavior and rule out possible morphological plasticity. Finally, it was concluded that these are three different species. In this article a new taxonomic delimitation of Agave potatorum proposed, are described and illustrated a new species and a new subspecies for science and is considered necessary to relocate some specimens of A. potatorum cited by Gentry (1982), now considered within A. nussaviorum or A. seemanniana.

The type of Agave potatorum was never formally elected. Gentry (1982) mentions that in 1969 found two specimens in the herbarium of Munich (M), which are only part of the inflorescence. The oldest, "1867, Hortus Monacensis" represents a small form with short side branches, similar to that observed by said author. The other copy "03/07/79, HbM" is a longer inflorescence, side stalk 8 cm long. Gentry believes that "Since These specimens date from 33 and 46 years after the publication of Zuccarini's species, types as Their suitability is doubtful. However, neotypes They can be Regarded as". Gentry did not choose the neotipo formally, so we preferred to propose a new nomenclatural type to support adequately the name.

Zuce potatorum Agave Plant 15: 2 (Beiblatt 2): 96. 1832. Agave Karw artichoke. exSalm-Dyck, Hort. Dyck Agave .: 307 1834. elegans hort. ex Salm-Dyck, Bonplandia 7: 89. 1859. Agave latifolia hort. ex Salm-Dyck, Bonplandia 7: 89. 1859. Agave pulchra hort. ex Salm-Dyck, Bonplandia 7: 89. 1859. Agave saundersii Hook.f., Bot. Mag. 91. t. 5493. 1865. Agave verschaffeltii Lem. ex Jacobi, Hamburger Garten-Blumenzeitung 21: 122. 1865. Agave A. polymorpha var scolymus. Terrace, Prim. Contr. Monogr. Agave, p. 38. 1885. Agave potatorum var. Verschaffeltii (Lem. Ex Jacobi) A.Berger, Die Agaven. p.186. 1915. Fig. 1 . Mexican botany Act - Taxonomic review of Agave potatorum Zucc complex. (Agavaceae): new taxa and neotipificación

Perennials, solitary, stemless, rarely stoloniferous 1-3 tillers; compact rosettes, 30-50 cm high, 40-60 cm diameter. Leaves 30-60 (400) per plant, 15-35 cm long, (5-) 8-10 cm wide, ovate, oblong or lanceolate, succulent, erect, flexible to rigid, flat or somewhat concave towards the apex, thickened toward the base, glaucous green-glaucous; sinuate to crenate margin, nipples 4-7 (-10) mm high; teeth of the middle, 4-6 (-12) mm long, 4-6 (-8) mm wide, retrorse, antrorsos or straight, brown to gray, separated by 1-3 cm; thorn 3-4 cm long, 0.4-0.7 cm wide, straight or winding, acute, broadly grooved to flat in the beam, decurrente by less than 1 cm, brown to gray. Inflorescence 3-5 (-6) m high, fertile portion in the middle or upper third, paniculata, open, with (7-) 11-25 primary branches 10-30 cm long, separated by 15-30 cm, to sometimes secondary branches 0.5-4 cm long; green to reddish-brown stalk, bract peduncle 12-20 cm long, 3.5-4 cm wide at the base, triangular, chartaceous, with Mexican botany Act - Taxonomic review of Agave potatorum Zucc complex. (Agavaceae): new taxa and neotipificación

some little teeth on the margin, spine 0.5-2.5 cm long, 0.7-1 cm wide, brown -rojiza; bracts of primary branches 5-8 cm long, 2-5 cm wide, ovate, amplexicaules. Flowers (15-) 20-40 per umbel, (5-) 5.5-7 cm long, campanulate to something urceoladas, succulent, yellow-green with purple or red dyes, green ovary; pedicels 1-1.5 cm long; ovary 2.5-3 cm long, 4-6 mm wide, cylindrical to slightly triquetrous, neck 2-3 (-5) mm long, tube (1) 1.3-1.6 cm long (0.8-) 1.3-1.8 cm diameter the distal portion, sulcate; tépalos 1.2-1.5 (-2) cm long, 3-6 mm wide, triangular or oblong, erect, thick, external 1-2 mm longer than domestic, involute margin, apex cucullate hard, the keel- shaped internal; filaments 3-4 (-4.5) cm long, flattened adaxially, inserted in the middle or third quarter of the tube top, beige, sometimes with purple tints, anthers 1.5-2.5 cm long, 2-3 mm wide, straight, versatile, yellowish; Style 5.5-6 cm long, stigma clavate, lobed. Capsules (3.5-) 4-6 cm long, 1.5-2 cm wide, oblong, stipitate, rostrate. Seeds 5-6 mm long, 4-5 mm wide, lunulares, with a barely perceptible wing, flat, black.

Kind. Mexico, Puebla: 4-10 miles SW of Tehuacan along road to Zapotitlán, 5600-6000 feet (1700-1800 m), arid thorn forest over limestone hills, August 1963, HS Gentry, A. & J. Arguelles Barclay 20427 (neotipo , here designated: MEXU !; isoneotipos: ARIZ !, US)!.

Specimens examined. . Mexico, OAXACA City Center: Cerro El Barrio (El Rosario) San Antonio, C. Conzatti et al. 4099 (MEXU, US); Monte Alban, A. Ramirez Laguna sn (MEXU); San Felipe, near City of Oaxaca, JN Rose and W. Hough 4592 (US) Coixtlahuaca District. 7 km SW of Calapa bridge Tepelmeme municipality of Villa de Morelos, A. Garcia-Mendoza and E. Solano 7256, 7257 (MEXU, MO); 3 km S of El Rodeo, A. Garcia-Mendoza et al. 7525 (MEXU); Otate Canyon, SW of Hijadero Aria, P. Tenorio et al. 17103 (MEXU); Maguey hill to W San Antonio Abad, road to La Mexicana, municipality of Santiago Ihuitlán Pens, P. . Tenorio 17539 (MEXU) Cuicatlán District: 2 km in a straight line to the northeast of San Juan Tonaltepec, municipality of Santiago Nacaltepec, G and G Juarez Cruz CA 669 (MEXU); Chico River Canyon, Cuicatlán E, F. Miranda 4619 (MEXU) Etla District. Long Earth, limits with San Juan del Estado, CA Cruz et al. 1815 (MEXU); 1 km to the north of the turnoff to the silks, A. Garcia- Mendoza 5704 (MEXU); same locality, cultivated in the Botanical Garden, A. Garcia-Mendoza 6105 (MEXU); 4.5 km W of El Parian way Nochixtlán, municipality of San Jeronimo Sosola, A. Garcia-Mendoza and E. Solano 7307 (MEXU); E 0.5 km silks, DH Lorence et al. . 4633 (MEXU) Huajuapan District: 5 km ahead of Santiago Chazumba Tehuacan highway (km 58), A. . Salinas 7644 (MEXU) Ixtlán District: NE Natividad Mendez Capulalpam Township, river S. Figueroa and B. Y. Guzman R. 766 (MEXU); 2 km S of The Student A. And R. Garcia-Mendoza Torres 1959 (MEXU); 6-8 miles NE of Cd Oaxaca along road to Ixtlan, Gentry HS 12057, 12079 (ARIZ, MEXU, US).; Yavesía, I. . Trejo 28 (MEXU) Miahuatlán District: 12 km S of Cuixtla road to Santa María Coatlán, A. Garcia-Mendoza and L. de la Rosa 6359 (MEXU) Mixe District. 6 km N of Tamazulapan Mixes, Totontepec way, A. Garcia-Mendoza et al. . 1279 (MEXU) Tlacolula District: 2 km N of San Dionisio Ocotepec, A. And R. Garcia-Mendoza Torres 1971 (MEXU); 10 km N of Mitla road to Ayutla Mixes, A. And R. Garcia-Mendoza Torres 1973 (MEXU); 5 km SE of Matatlán, Mitla-Totontepec road, cultivated in the Botanical Garden, A. Garcia- Mendoza 6554 (MEXU); 4 km SW of Santa Rosa de Buenavista, municipality of San Sebastián Abasolo, A. Garcia- Mendoza et al. 6815 (MEXU); Nine Tips hill, about 8 km to the north of San Baltasar Guelavila, A. Garcia-Mendoza and H. Espinosa 7426 (MEXU); Yagul natural monument, A. Garcia-Mendoza et al. 7595 (MEXU); 8 miles E of Mitla, Gentry HS 22344 (MEXU); Santiago Matatlán the S, SC and MA Solano Vara 196 (ARIZ); 10 km to the east of Santa Maria Albarradas, R. Torres and C. Martinez 12538 (MEXU); near Cueva Blanca, J. Schoenwetter JSOX-78 (US) District Sola de Vega. 12.1 km N of Sola de Vega on the road to Oaxaca, B. Bartholomew et al. . 3271 (MEXU) Zaachila District: 3 km S of San Cristobal Cuatro Venados, Minas river path, A. Garcia-Mendoza et al. 8192 (MEXU). Zimatlán District: place La Cantera, A. Miranda and O. Hernandez 738 (MEXU); Canada Frijolón, 8 km NE of The Brotherhood, A. Miranda and O. Hernandez 754 (MEXU). PUEBLA. A 7 km from San Andrés Cacaloapan municipality Tepanco Lopez, I. Calzada 23189 (MEXU); Azumbilla 8.5 km going towards La Esperanza, M. Cházaro and BL Mostul sn (MEXU); Puebla, A. Chimal 37 (MEXU); Tehuacan Endlich 1909, 1929a (MO); Tlayúa quarry, ca 2 km east to Tepeji, R. Felger 8520 (DES); 6 km NW of Azumbilla road Tehuacan municipality Chapulco, A. Garcia-Mendoza and F. Palma 5629 (MEXU); 5 km S of Tehuacan, Zapotitlan road, cultivated in the Botanical Garden, A. Garcia-Mendoza 6121 (MEXU); Virgin, 8 km to the north of Zapotitlan, cultivated in the Botanical Garden, A. Garcia-Mendoza 6560 (MEXU); 13 km S of Puerta Cañada, municipality of Cañada Morelos, A. Garcia-Mendoza et al. 6664 (MEXU); 5 km S of Puerto Las Palmas, Izúcar municipality of Matamoros, A. Garcia- Mendoza et al. 6860 (MEXU); Loma Colorada, municipality Palmar de Bravo, A. Garcia-Mendoza et al. 7090 (MEXU, MO); 10 km S of Molcaxac, A. Garcia-Mendoza et al. 7505 (MEXU); 22-24 km NE of Tehuacan, along highway to Veracruz, Gentry HS 12288 (ARIZ, MEXU, US); Tehuacan, D. Sn Griffiths (MO); Agua Chiquita spot in the nearby river Axamilpa municipality Tepeji Rodriguez, E. Guízar and G Herrera 4090 (MEXU); Cuixosto spot in the nearby river Axamilpa municipality Tepeji Rodriguez, E. A. Miranda Guízar and 4755 (MEXU); The Escobillera hill, 1 km from Sabino Farol, municipality of Caltepec, E. A. Miranda Guízar and 4901 (MEXU); Izúcar de Matamoros, F. Miranda 2157 (MEXU); 10 miles S of Izúcar de Matamoros, along highway Between Matamoros and Acatlan, EC Ogdeny CL 51201 Gilly (ARIZ, MEXU, US); Tehuacan elevations saw La Mesa, A. Ramirez Laguna sn (MEXU); near Tehuacan, JN Rose and R. There are 5950 (US); JN Rose et al. 10024 (US); JN Rose yJ.S. Rose 11261, 11269, 11421.11426 (US); Soap hill, the W of La Mesa, municipality of Caltepec, P. Romero C. Tenorio and 4734 (MEXU, Mexican botany Act - Taxonomic review of Agave potatorum Zucc complex. (Agavaceae): new taxa and neotipificación

MO); Dead hill, SW of San Simon, municipality of Caltepec, P. Romero C. Tenorio and 5052 (MEXU, MO); 10 km NE of Azumbilla road Esperanza, municipality of Chapulco, P. Tenorio 15200 (MEXU); Tehuacan W. Trelease sn 12 Aug 1903 15 Aug 1903, February 5, 1905, February 10, 1905 (MO); W of Tehuacan, W. Trelease sn (MO).

Phenology. It blooms from late August to November and fruits from November to March.

Habitat and distribution. Agave potatorum develops flat spots or slope, on sandy soils derived from limestone, in an altitudinal range of from the 1300-2400 m, in arid and semiarid climates. Part of the tropical deciduous forest, the desert scrub with columnar and desert scrub with Quercus cacti. Its distribution covers the eastern end of the basin of the Balsas River Valley of Tehuacan-Cuicatlán, foothills of the hills and mountains Mixtec surrounding the Central Valleys of Oaxaca.

Etymology: The specific epithet is Latin potator drinker -oris (wine).

Common name and uses: papalometl, papalomé, papalomé maguey Tobala ha tsääjts NAK (Mixe) (Vásquez, 2006). It is mainly used for the production of mezcal liquor prized for its soft organoleptic qualities. The flowers are used in food like the flower stem development when started. In traditional medicine, the leaves are used to reduce swelling.

Agave potatorum is characterized by compact rosettes, with many flat sheets with small nipples and teeth; paniculata inflorescence with primary branches 10-30 cm long, separated by 15-30 cm, present in the middle or upper third of the inflorescence; bracts shorter than the primary branches; campanulate flowers with a slightly urceolado tube, stamens inserted at the middle of the tube or the third upper room, pipe connection: tépalos 1: 1 or even the tube becomes up to 2 times longer than the tepals. At the end of the Central Valley of Oaxaca in Tlacolula Mixe districts and east, and Sola de Vega Miahuatlán south potatorum Agave has a greater variation in vegetative and reproductive organs; This is probably due to flowering synchronously and is sympatric with Agave seemanniana, species with which probably exchanged genetic material. These specimens with intermediate morphological characteristics of hybrid origin likely listed below.

Miahuatlán District: 2 km SW of San Jerónimo Coatlán, A. Garcia-Mendoza et al. 4663 (MEXU); 7 to 8 miles SE of Miahuatlán over new road along Sierra Madre del Sur, Gentry HS 12106 (ARIZ, MEXU, US); 11.6 km SW of San Pablo Coatlán, road Miahuatlán - Piedra Larga, R. Torres and A. 10775 Campos (MEXU, MO) Mixe District. 4 km E of Tlahuitoltepec, 3 km N of Tamazulapan Mixes, A. And R. Garcia-Mendoza Torres 2000 (MEXU) District Sola de Vega. Yocúa, 4 km N of Sola de Vega, A. . Garcia-Mendoza and Espinosa H. P. 7458, 7459 (MEXU) Tlacolula District: 10 km S of Cuajimoloyas, A. Garcia-Mendoza 5672 (MEXU); 8 km to the north of Diaz Ordaz, A. Garcia- Mendoza et al. 5782 (MEXU); same locality, cultivated in the Botanical Garden, A. Garcia-Mendoza 6317, 6559 (MEXU); 9 km S of Matatlán, at the base of the hill Nine Points, A. H. Garcia-Mendoza and Espinosa 7429 (MEXU); 8 miles E of Mitla along road to Ayutla, Gentry HS 22344 (MEXU).

Agave nussaviorum Garcia-Mend. Fig. 2 . Mexican botany Act - Taxonomic review of Agave potatorum Zucc complex. (Agavaceae): new taxa and neotipificación

Ex affinitate A. potatorum Zucc. et A. seemanniana Jacobi, ab A. potatorum differt racemosa inflorescentia firn, subsessilibus umbellis proximissimis inter se, paucifloris, floribus 1/3 1/6 apicalis partis ad occupantibus; bracteis quam ramis longioribus; rosulis apertis prophylaxis, minus foliis Numerosis; mamillis marginalized Instructo magnis, dentibus in mamillarum apicibus decurrentibus, floribus brevitubulatis, tubi longitudine quoad rationes longitudinis tepalorum 1: 1.5-2.5; ab A seemanniana differt partibus vegetativis reproductivisque magnitudine Minori foliis Numerosis Magis, et racemosa lanceolatis oblongis inflorescentia vel.

Perennials, solitary, stemless; open to compact rosettes, 30-50 cm high, 40-60 cm diameter. 25-40 leaves per plant, (12-) 20-40 cm long, (5-) 7-ll cm wide, lanceolate, oblong to ovate, succulent, erect to recurved, flexible, Mexican botany Act - Taxonomic review of Agave potatorum Zucc complex. (Agavaceae): new taxa and neotipificación

somewhat concave towards the middle and apex, yellow-green, blue-green towards the base; crenate margin, prominent nipples towards the middle of the sheet, 5-10 mm high, variously curved; teeth (5-) 8-12 mm long, (5- ) 8-10 mm wide, with wide base that covers the top of the bottle, retrorse, antrorsos or straight, brown to gray, 2- 3.5 cm apart, in interstitial occasions with little teeth, towards the base of the blade teeth are small and close together; spine 2-3 (-4) cm long, 0.3-0.5 (-0.8) cm wide, winding to straight, grooved in the beam, decurrente by 1-2 cm, brown to gray. Inflorescence (1.5-) 3-5 (-6) m high, fertile portion 1/3 to 1/6 of the distal part, racemosa, firn, flowers 20-40 (-60) umbels, branches less than 3 cm long, spaced apart by 3-5 cm; purple green stalk; bract peduncle 12-17 cm long, 3.5-4.5 cm wide at the base, triangular, chartaceous, with some little teeth on the margin, spine 2-3.5 cm long, 0.3-0.5 (l) cm wide, reddish-brown; bracts of 2.5-3.5 umbels (-5) cm long, 2-3.5 cm wide, ovate, chartaceous, amplexicaules, longer than the branches, thorn less than 5 mm. Flowers 6-12 per umbel, 4.5-5.5 (-7) cm long, slightly bell-shaped, succulent, yellow with purple tints, green ovary; pedicels 2-5 (-10) mm long; ovary 2.3-3 - (- 3.5) cm long, 4-6 mm wide, cylindrical neck 2-3 (-5) mm long, tube 0.7-1 (4.3) cm long, 0.7-1 cm diameter portion distal slightly sulcate; tépalos 1.5-2.5 cm long, 2-5 mm wide, triangular or oblong, thick, slightly longer external to internal, involute margin, apex cucullate hard, the keel-shaped internal; filaments (2.5-) 3.5-4.5 (-5.5) cm long, inserted 2-3 mm below the base of the tepals, yellow with purple tints, anthers 2-2.5 cm long, 2 mm wide, versatile, yellowish; style 4-6 cm long, stigma clavate, lobed. Capsules 4-6 cm long, 2-2.5 cm wide, oblong to ovate, stipitate, rostrate. Seeds 5-6 mm long, 4-5 mm wide, lunulares, with a very narrow wing, flat, black.

Kind. Mexico, Oaxaca: Nochixtlán municipality of Santo Domingo Yanhuitlán 1 km NW of Yanhuitlán, 2238 m, eroded Quercus forest area, September 5, 2008, District A. Garcia-Mendoza, S. F. Martínez Franco and 9204 (holotype: MEXU; isotypes: ENCB, IEB, MO, OAX, US).

Paratypes. . Mexico, OAXACA District Coixtlahuaca: 195.5 km carr. Oaxaca - Tehuacan, near Monte Verde, municipality of Santa Maria Nativitas, A. Garcia-Mendoza et al. 9206 (ENCB, MEXU) Huajuapan District: palmetum, H. Bravo Hollis sn (MEXU); 10 km NW of Huajuapan, road Acatlan A. Garcia-Mendoza et al. 3628 (MEXU); El Mirador, 9 km NW of Huajuapan, A. Garcia-Mendoza and J. Reyes 4933 (MEXU); Glen Lencho, 6 km SW of San Marcos Arteaga, Huajuapan highway - Tonala, A. Garcia-Mendoza and F. Cruz 5804 palm (MEXU); same locality, cultivated in the Botanical Garden, A. Garcia-Mendoza 6118 (MEXU); 6 km SE of Huajuapan highway Tamazulapan, A. Garcia-Mendoza et al. 6328 (MEXU); 1 km to the NO on Guadalupe Cuautepec deviation, A. Garcia-Mendoza et al. 7096 (MEXU, MO); 1 km on the deviation Cuautepec Guadalupe, A. Garcia- Mendoza and E. Solano 7336 (MEXU, MO); 6 Miles NW of Huajuapan along highway, Gentry HS 22401 (ARIZ, MEXU, US); ca. 2 miles N of Huajuapan along highway to Puebla, HS Gentry and M. Gentry 23656 (ARIZ, DES, US); Palo Flor site, 4 km NW of Santo Domingo Yodohino, municipality Santo Domingo Yodohino, J. Lopez Moreno 279 (MEXU); Huajuapan, km 80, H. Quero sn (MEXU); Rancho La Cruz, 31 km W of Huajuapan, R. Torres et al. . 14162 (MEXU) District Juxtlahuaca: 4.5 km from San Sebastián Tecomaxtlahuaca, path to Seco river and stream Bell, JI 21808 Calzada (MEXU); Yucha hills Cuanshii, municipality of San Juan Mixtepec, J. . Reyes 1227 (MEXU) Nochixtlán District: Glen Yucuduza, municipality of Asuncion Nochixtlán, D. Aquino 201 (MEXU); 12 km E of Nochixtlán to 1 km N of river Salinas, A. Garcia-Mendoza and Rosa L. of 6175 (MEXU); about 2 miles NW of Yanhuitlán along highway, Gentry HS 12111 (ARIZ, MEXU, US); 1 mile NW of Yanhuitlán along highway, Gentry HS 22399 (ARIZ, MEXU, US); Pan-Am below. Hwy. At Puerto Herrera (420 km) just NW of Yanhuitlán, litigation HH et al. 1147 (ENCB); 21 km von Llano Verde, std. to Jaltepec Morelos, B. Leuenberger and C. 2773 Schiers (MEXU); 1 mile north of La Joya, along highway from Cd Oaxacato Acatlán, EC 51193 Ogden and CL Gilly (MEXU, US).; Methoxy, F. . Palma Cruz 13 (MEXU) District Teposcolula: 5.3 km from the junction of the river del Oro on the aqueduct of the CFE, road Tamazulapan - Huajuapan, JI 23786 Calzada (MEXU); 7 km SW of San Andrés Lagunas way Teposcolula A. Garcia-Mendoza 893 (MEXU); Anama around 3 km S of San Vicente Nuñú, municipality of San Vicente Nuñú, A. Garcia-Mendoza et al. 5250; 6 km W of Teposcolula way to San Pedro Yucunama, A. Garcia-Mendoza 8837 (MEXU); About 15 mi NW of Tejupan, Gentry HS 12124 (MEXU) Tlaxiaco District. 3 km NW of Tlaxiaco and 3 km from the turnoff to Santiago Nundiche, municipality of Santiago Nundiche, A. Castaneda et al. 95 (MEXU).

Phenology. It blooms from late August to December and fruits from November to March.

Habitat and distribution. Nussaviorum Agave takes place in the mountains of the Mixteca Alta, in flat spots or hillsides, on soils derived from limestone or sandstone, in an altitudinal range of 1700-2500 m. It is part of Quercus-Pinus forest with Juniperus scrub and sclerophyllous Rhus, Brahea, Arctostaphylos, Nolina, Comarostaphylis, etc. Distribution to the tops of the mountains is restricted Coixtlahuaca districts, Huajuapan, Juxtlahuaca, Nochixtlán, Teposcolula and Tlaxiaco.

Etymology. The specific epithet refers to the Oaxacan Mixteca Alta region and its inhabitants (ñuu Savi = land of villagers clouds or clouds), geographic range of the species. Mexican botany Act - Taxonomic review of Agave potatorum Zucc complex. (Agavaceae): new taxa and neotipificación

Common names and uses. Papalometl, papalomé, "yavi ticushi" (Mixtec). It is mainly used as a food and medicinal plant. The Mixtecs made the​​ "yahui ndodo" bittersweet food greatly appreciated in the past; Also, the young quiote oven baked and eaten as a snack; flowers (cacayas or tibilos) are consumed in stews and mixed with cooked corn dough for tortillas. The juice of the leaves are attributed medicinal properties to fight diseases of the circulatory system such as blood clots and diabetes. The tatemadas sheets (semicocidas fire) are used to relieve inflammation caused by blows or falls in animals and man. Lopez (2008) mentions having up to seven different uses, among others, the inflorescence is used as fodder, dry quiote in construction and fuel.

Agave nussaviorum differs from A. potatorum by racemosa type of inflorescence branches, very close together and very short umbels few flowers, the fertile part covers 1/3 to 1/6 of the distal part of the inflorescence, with bracts longer the branches. The rosettes have fewer leaves, they are more open and greenish-yellow color, the leaves usually have large nipples and deriving teeth on the tip of the nipple, sometimes interstitial little teeth are observed. Flowers are subsésiles with short tubes in a ratio of 1: 1.5-2.5 with respect to tépalos; inserted filaments 2-3 mm below the base of the tépalos. The habitat corresponds to the temperate climate is cool and humid, with Quercus-Pinus forest, Pinus forest and sclerophyllous scrub. In some eroded and poor soil sites, the rosettes are globular and small, with inflorescences not greater than 2 m high.

Agave seemanniana differs because this species has all his greatest vegetative and reproductive organs, have fewer leaves, so spatulate or broadly lanceolate, glaucous color, recurved, outside the winding sheet, big teeth, mostly apart and inflorescence a larger branches with primary, secondary and even tertiary. Agave nussaviorum coexists with A. convallis, A. atrovirens, A. and A. salmiana angustifolia (in a few places) and north with A. potatorum.

You agave nussaviorum likely have a hybrid origin between A. potatorum (morphologically similar species) and some species of the subgenus Littaea as A. convallis or A. angustiarum; however, no evidence to support that process. Currently, A. nussaviorum occupies a geographic area adjacent to A. potatorum grows in a different habitat, where they successfully reproduce sexually. In this process it would be possible to also consider the human management, since they are widely used species, although for different purposes. Some morphological evidence suggesting hybridization are presented in the following specimens collected in areas disturbed by human activities.

Ixtlan District: hillside above the Rio Grande, Yahuiche E, A. F. Garcia-Mendoza and Martinez 7055 (MEXU) Nochixtlán District. Tinú Arroyo, 10 km SE of Nochixtlán, A. . Garcia-Mendoza and E. Solano 7300 (MEXU) District Teposcolula: The Organito 1 km N of Rancho Peñasco, San Pedro Nopala, A. Garcia-Mendoza and Rosa L. of 6158 (MEXU).

The above information corresponds to the typical subspecies A. . nussaviorum ssp nussaviorum since then is proposed as new and is described:

Agave nussaviorum Garcia-Mend. subsp. deltoidea Garcia-Mend., subsp. nov. Fig. 3 . Mexican botany Act - Taxonomic review of Agave potatorum Zucc complex. (Agavaceae): new taxa and neotipificación

A typica subspecies differt deltoideis foliis longioribus latioribusque, continuous marginalized stratum denticulis vel quam 5mm margine brevioribus adpressis, et spina longa 6-10 cm, longe decurrenti.

Perennials, solitary, stemless; open rosettes, 50-70 cm high, 60-80 cm diameter. 20-40 leaves per plant, 20-40 cm long, (8-) 14-20 cm wide at the heel end deltoid widely lanceolate, succulent, rigid, flat or concave, yellow- green, sometimes with cross-zoned bands glaucous; length, straight and horny or sparsely toothed margin; teeth (1-) 4-5 mm long, on a broad basis (0.6-) 1-1.3 cm wide, retrorse, adpresos outside or rare free time and recurved, gray, separated by 1-2 cm; spine (4-) 6-10 cm long, 0.5-0.8 cm wide, decurrent for 10-14 cm or continuous with the margin, ribbed on the back, grayish. Inflorescence 4.5-6.5 m high, stem 3-5 m, green, fertile Mexican botany Act - Taxonomic review of Agave potatorum Zucc complex. (Agavaceae): new taxa and neotipificación

portion in the last upper third, racemosa, firn, flowers clustered in umbels subsésiles 30-60 1.5-2.5 cm long, separated by 3-5 cm ; bract peduncle 12-18 cm long, 2-5 cm wide at the base, triangular, with some little teeth on the margin, chartaceous, spine (2-) 3-4 cm long (0.3-) 0.6-l cm wide , reddish-brown bracts branches 4 cm long, 3 cm wide, ovate, longer than the branches. Flowers 4-10 per umbel, 4.5-6 (-7) cm long, somewhat bell- shaped, greenish-yellow, green ovary; subsessile with pedicel up to 5 mm long; ovary 2.5-2.8 cm long, 4-8 mm wide, cylindrical neck 3-5 mm long, 1-1.5 cm diameter tube, 1-1.3 cm wide at the distal portion; tépalos (1.5-) 2.5-2.8 cm long, 3-8 mm wide, triangular or oblong, external with involute margin, apex cucullate hard, the keel- shaped internal; filaments (3.5-) 4-5 cm long, inserted 2-4 mm below the base of the tepals, yellow, anthers 1.5- 2.5 cm long, 2-3 mm wide, versatile, yellowish; style 5-5.5 cm long, lobed stigma. Capsules 3.5-4 cm long, 1.5-2 cm wide, oblong, rostrate. Seeds 5-6 mm long, 3-4 mm wide, lunulares, with a very narrow wing, flat, black.

Kind. Mexico, Oaxaca: Teposcolula, municipality of San Pedro Nopala, closed the Peña del Tigre, 10 km N of Nopala, 2600 m, rosetophilous scrub, April 30, 1995, District A. F. Garcia-Mendoza and Martinez 6051 (holotype: MEXU; isotype: OAX).

Paratypes. . Mexico, OAXACA Teposcolula District: Pericón hill, 5 km NW of Nopala, municipality of San Pedro Nopala, A. Garcia-Mendoza et al. 3959 (MEXU); Cerro La Peña del Tigre, 10 km N of Nopala, A. F. Garcia-Mendoza and Martinez 6057 (MEXU); same locality, cultivated in the Botanical Garden, A. Garcia-Mendoza 7237 (MEXU); Malintzin hill, 8 km to the north of Nopala, A. Garcia-Mendoza et al. 6689 (MEXU).

Phenology. Flowers from September to October and fruits from December to April.

. Habitat and distribution Agave nussaviorum . var deltoid sites developed in stony soils derived from igneous rocks, at altitudes of 2400-2600 m; rosetophilous is part of scrubland dominated by convallis Agave, Agave nussaviorum, Dasylirion serratifolium, Furcraea longaeva, Hechtia sp., Nolina sp. and mixtecana Yucca. is apparently endemic to the type locality and nearby areas.

Etymology. The specific epithet refers to the shape of the leaves.

The subspecies deltoid differs from the typical subspecies by longer and wider leaves deltoid, with continuous margin horny little teeth or less than 5 mm long, appressed apart, spine 6-10 cm long, long decurrent. It develops east range of the typical subspecies, in a different habitat, at higher altitudes and soils derived from volcanic rock, where it is scarce and restricted to igneous substrates.

Agave seemanniana Jacobi, Abh. Schles. Ges. Vaterl. Cult., ABTH. Naturwiss. 1868: 154. 1869 Agave scolymus var. seemanniana (Jacobi) A. Terrace, Prim Contr.. Monogr. Agave, p. 38. 1885. Agave caroli-schmidtii A. Berger, Die Agaven. p. 199. 1915. Agave guatemalensis A. Berger, Die Agaven. p. 201. 1915. Agave seemanniana var. perscabra Trel., Trans. Acad. St. Louis Sci. 23: 137. 1915. Agave tortispina . Trel, Trans. Acad. St. Louis Sci. 23: 135. 1915. Agave pygmaea Gentry, Agaves Cont N. Amer.. p. 494. 1982. Agave seemanniana subsp. pygmaea (Gentry) B. Ullrich, Brit. Cactus Succ. J. 10: 1992. 27. Figs. 4D , 5D ; Gentry, 1982: 496, fig. 17.23. Mexican botany Act - Taxonomic review of Agave potatorum Zucc complex. (Agavaceae): new taxa and neotipificación Mexican botany Act - Taxonomic review of Agave potatorum Zucc complex. (Agavaceae): new taxa and neotipificación

Perennials, solitary, stemless; open rosettes, (0.8-) 1-1.5 m high, (1.3-) 1.5-2 m diameter. Sheets (16-) 30-40 per plant, 40-55 cm long, (10-) 14-22 cm wide, spatulate, broadly lanceolate or ovate, succulent, recurved to erect, narrow at base, glaucous, green-glaucous or yellow-green, zoned; crenate margin, nipples (0.7-) 1-1.5 (-2) cm tall; teeth (0.6-) 1-1.5 cm long, 1-1.4 cm wide, with wide base that covers the top of the bottle, retrorse at right, brown to gray, separated by (3-) 4-5.5 cm; spine (3) 4.5-5 cm long, 0.5-1 cm wide, winding, decurrente 2-6 cm, broadly channeled into the beam. Inflorescence paniculate, 4-6 (-7) m tall, open, fertile portion in the upper half, silhouette oblong, flowers in 15-30 lateral umbels, primary branches 15-30 (-50) cm long, separated by 15 -30 cm, secondary branches (1-) 3-7 (-10) cm, occasionally present tertiary branches 1-1.5 cm; bract peduncle 15-22 cm long, 5.5-7 cm wide at the base, triangular, sometimes with some little teeth on the margin, chartaceous, spine (2-) 5-10 mm long, bracts of primary branches 8-10 cm long, 4-5 cm wide, deltoid. Flowers (40) 60-80 Mexican botany Act - Taxonomic review of Agave potatorum Zucc complex. (Agavaceae): new taxa and neotipificación

umbel, (5-) 6-7 cm long, slightly bell-shaped, yellow, green ovary; pedicels 1-1.5 cm long; ovary 2.5-3.5 cm long, 5-7 mm wide, triquetrous a cylindrical neck 3-5 mm long, 0.7-1 tube (4.2) cm diameter 0.7-1 (4.2) cm wide at the distal portion; tépalos 1.5-2 cm long, 3-5 mm wide, slightly longer the keel-shaped internal and external, involutes, sometimes tinged with purple, linear, erect, thick, apex cucullate; filaments 3-4.5 cm long, inserted 1-2 (-3) mm below the base of the tepals, yellow, anthers 1.5-2 cm long, 2-3 mm wide, versatile, yellow; Style 4.5-6 cm long, lobed stigma. Capsules 4.5-5 (-6.5) cm long, 2-2.5 cm wide, oblong, shortly stipitate, short face. Seeds 5-7 mm long, 4-6 mm wide, lunulares without wing, flat, black.

Kind. Honduras, 8-12 km SE of Tegucigalpa along road to Zamorano, open rocky slope with pine, approx 1965, Gentry HS 20684 (neotipo appointed by Gentry (1982): US isoneotipos: DES !, MEXU!).

Specimens examined. . Mexico, OAXACA Cuicatlán District: Cerro del Cosahuico, Chilar San Jose, CA Cruz Espinoza 491 (MEXU); Santiago Dominguillo, CA Cruz Espinoza 2037 (MEXU); Santiago Quiotepec, CA Cruz Espinoza 2144 (MEXU); 4 km S of Tomellín way to San Pedro Jaltepetongo, A. Garcia-Mendoza et al. 7356 (MEXU); 6 km SE of Tomellín way to San Pedro Jaltepetongo, A. Garcia-Mendoza et al. 7389 (MEXU); 16 km SE of Dominguillo, by road Tehuacán - Oaxaca, F. Gonzalez-Medrano et al. 1747 (MEXU); Tomellín Canyon, JN Rose Rose and JS 11527 (US); Tomellín Canyon, W. Trelease sn (MO). Miahuatlán District: 6 km to the north of San Jerónimo Coatlán, A. Garcia-Mendoza et al. 4594 (MEXU); 0.5 km W of San Lorenzo Mixtepec, E. Hunn 500 (MEXU). Pochutla District: Zacatal Cerro Lobo, municipality of San Miguel del Puerto, S. Salas et al. 5100 (MEXU, SERO); same locality, G Salazar 7029 (MEXU). Sola de Vega District: San Sebastian de Las Grutas, F. Palma C. 52 (MEXU). Tehuantepec District: 2 km to the north of La Concepción, 18 km S of San Miguel Tenango, A. E. Garcia-Mendoza and Martinez 3592, 3594 (MEXU); 5 km S of San Miguel Tenango, A. Garcia- Mendoza and E. Martínez 3599 (MEXU); 15 km S of Buenos Aires route to San Miguel Tenango, cultivated in the Botanical Garden, A Garcia-Mendoza 6327, 6555 (MEXU); E slope of Cerro Guiengola, L. C. Torres et al. 344 (MEXU); El Ocotal, Cerro Guiengola, L. C. Torres et al. 944 (MEXU); Cerro El Gavilan, 10 km to the east of Santiago Lachiguiri, J. 3577 S. Kings (MEXU). Tlaxiaco District: 3 km S of Chalcatongo, Yosondúa way, A. Garcia-Mendoza et al. 6837 . (MEXU) Villa Alta District: Cima del Cerro del Aguila, 4 km S of Santo Domingo Xagacia, A. Garcia-Mendoza et al. 5770 (MEXU); Santo Domingo Xagacia, F. Palma 74 (MEXU). Yautepec District: Portillo de Nejapa, A. Garcia-Mendoza 5691 (MEXU); Portillo de Nejapa, plant cultivated in the Botanical Garden, A. Garcia-Mendoza 6104 (MEXU); Hose, 6 km E of The Shrimp, A. H. Garcia-Mendoza and Espinosa 7421 (MEXU); 690 km highway to Tehuantepec along 10 miles SE of shrimp, Gentry HS 21886 (ARIZ, MEXU); 8 miles SE of shrimp along highway to Tehuantepec, and M. Gentry Gentry HS 23655 (ARIZ); Between Tequisistlán and Totolapan, on road 195, 23 km NW of Rio Hondo, B. Leuenbergery C. Schiers 2538 (MEXU); Climb the tower microwave San Cristobal, 12 km SE of The Shrimp, R. 12588 Torres (MEXU).

Phenology. Flowers from September to December and fruits from December to April.

. Habitat and distribution seemanniana Agave takes place on mountain slopes, on sandy soils derived mainly of limestone, between 700-1700 meters; part of the deciduous forest and ecotone towards the subtropical forest of Quercus-Pinus. In Oaxaca is distributed in the basin of the Tomellín and Grande rivers in the district of Cuicatlán and in the mountains of the Sierra Madre del Sur, mainly in the districts of Tlaxiaco, Miahuatlán, Pochutla, Yautepec and Tehuantepec.

Common names and uses. Bilia, flat, Dob-be blah dua, Tobala, papalometl. It is used for the production of mezcal. In the districts of Miahuatlán and Tlaxiaco also it used to prepare pulque (A. Garcia-Mendoza et al. 4594, 6837 and E. Hunn 500).

Agave seemanniana rosettes is characterized by over one meter high, open, with few leaves recurved, spatulate to broadly lanceolate, margin with prominent nipples and large teeth and deriving bases. The inflorescence has primary branches, secondary and tertiary occasionally. The flowers are characterized by short tubes inserted filaments 1-2 (-3) mm below the base of Tépalos stamens are relatively short, and in excess to the tube by 1.5- 2.5 cm. As potatorum Agave, blooms in the fall, with viable seeds during winter and spring. Gentry (1982) mentions that lives from Chiapas to Nicaragua; but nonetheless, he collected plants in Oaxaca, copies assigned to Agave potatorum (HSG 21886, 23655, see also in his pictured below 17.21). seemanniana Agave coexists with A. potatorum, A. angustifolia and A. convallis.

Taxa treated in this study as well as other species of Agave with coexisting, can be identified with the following key: Mexican botany Act - Taxonomic review of Agave potatorum Zucc complex. (Agavaceae): new taxa and neotipificación

In Figures 4 and 5 and comparative photographs shown in Figure 6 the distribution of taxa treated. Mexican botany Act - Taxonomic review of Agave potatorum Zucc complex. (Agavaceae): new taxa and neotipificación

Acknowledgements

curators of herbal ARIZ, DES, ENCB, MEXU, MO, OAX, SERO and US We are grateful for allowing consult their collections. To Drs. Fernando Chiang, Eloy Galvan Raquel Solano and critically reviewing the manuscript. Latin diagnoses were made by​​ the first reviewer. Special thanks to Sonia and Francisco Franco Martínez Martínez Aguirre for their constant and invaluable cooperation in the fieldwork. The Figures 1 and 2 are input from Elvia Esparza and three Albino Moon. Photos of Agave seemanniana courtesy of Dr. Gerardo Salazar. Mexican botany Act - Taxonomic review of Agave potatorum Zucc complex. (Agavaceae): new taxa and neotipificación

LITERATURE CITED

Gentry, HS 1982. Agaves of Continental North America. University of Arizona Press. Tucson, Arizona. 670 pp. [ Links ]

Lopez, MJ 2008. ethnobotanical study in the municipality of Santo Domingo Yodohino, District of Huajuapan de Leon, Oaxaca. Bachelor thesis. Autonomous University Chapingo. Chapingo, State of Mexico. 220 pp. [ Links ]

Vasquez, G. 2006. Integrated use of pulque maguey ( Agave salmiana ) in Santa Maria Tlahuitoltepec Mixe Oaxaca. Bachelor thesis. Autonomous University Chapingo. Chapingo, State of Mexico. 103 pp. [ Links ]

Zuccarini, JG 1833. Über einige aus den Pflanzen Gattungen Agave und Fourcroya. Phys. Nova Acta Med. Acad. You fall. Leop.-Carol. Nat. Cur. 16 (2): 659-679, tab. 48-51. [ Links ]

Institute of Ecology, AC Bajio Regional Center

Av. Lazaro Cardenas 3052, Patzcuaro, Michoacan, MX, 61600, (52-434) 342-2698

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