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Taxonomic and nomenclaturas status of the Mexican species in the Tillandsia viridiflora complex (Bromeliaceae).

During the course of two botanical field trips through the state of Oaxaca in the years 2005 and 2006, the late Jurgen Lautner, Renate Ehlers and collaborators collected several plants of a species of Tillandsia L. in the Sierra Mixe, which were provisionally identified as T. viridiflora (Beer) Baker. Additionally in 2006, Thorsten Kromer and Elisabeth Otto collected one specimen of the same Tillandsia species on the slopes of the Santa Marta volcano, located in the southeastern region of Los Tuxtlas, in the state of Veracruz. In order to confirm the identity of these plants, we studied cultivated and herbarium material, and also the related taxonomic literature. After a careful study of literature relating to these plants (Baker, 1889; Smith & Downs, 1977; Weber, 1984; Utley & Burt-Utley, 1994; Beaman & Judd, 1996), an analysis of the original descriptions and plates of the names related to Tillandsia viridiflora and its putative synonyms T. macropetala Wawra, T. orizabenzis Baker, T. longiflora Sesse & Moc. and T. billbergiae (Lem.) Baker (Schlechtendal, 1845; Beer, 1856; Lemaire, 1869; Morren, 1873, 1880; Wawra, 1887; Baker, 1888; Sesse & Mocino, 1887, 1894; Mez, 1935), and study of all relevant specimens and types deposited in the main Mexican (CHAP, ENCB, FCME, IBUG, IEB, MEXU, OAX, UAMIZ, XAL) and foreign (GH, K, LL, MA, MICH, MO, OXF, P, SEL, TEX, UC, US, WU) herbaria, we come to the conclusion that the plants collected at both localities mentioned above, and also some herbarium material formerly identified as T. viridiflora, T. grandis and/ or T. heterophylla E. Morren must be assigned to T. macropetala, a name previously considered by some authors as a taxonomic synonym of T. grandis (Mez, 1935) or T. viridiflora (Smith & Downs, 1977; Espejo & Lopez-Ferrari, 1994; Espejo et al., 2004). We believe that each of these three species constitutes a phenetically distinctive entity, following a morphological-taxonomic species concept. However, besides their differences in habit and the appearance of leaves and peduncle bracts, especially obvious in living plants (Fig. 1, 2 C-D), T. macropetala seems most closely related to T. viridiflora, as both species are more similar in size and epiphytic, whereas T. grandis is much bigger and lithophytic (Fig. 2A; Table 1).

With the reestablishment of Tillandsia macropetala as a valid species, the number of taxa of the T. viridiflora complex present in Mexico reaches three. Additionally, because of its close morphological similarity, this species complex probably includes the Caribbean T. paniculata (L.) L. and T. baliophylla Harms. Following the classification of Smith & Downs (1977), all these belong to Tillandsia subgenus Pseudoalcantarea Mez. However, Beaman & Judd (1996) suggested that this subgenus is paraphyletic, and transferred T. grandis and T. paniculata to subgenus Tillandsia. Furthermore, Barfuss et al. (2005) in a phylogenetic analysis of subfamily Tillandsioideae suggested that T. baliophylla and T. viridiflora also do not form a clade. Therefore, although we have clarified species limits in this complex, it is clear that much more phylogenetic work is necessary before we can be confident as to their phylogenetic relationships. Below we include an identification key for the three Mexican taxa, their correct names, complete synonymy, descriptions, and a distribution map.

[FIGURA 1 OMITIR]

[FIGURA 2 OMITIR]

Key to the Mexican species of the Tillandsia viridiflora complex

1 Inflorescence simple; leaves narrowly triangular, 25-50 cm long; petals ca. 9 cm long; plants 0.65-1 m high.... T. viridiflora

1 Inflorescence once branched, very rarely simple in T. macropetala but then the leaves strap-shaped; plants 1.3-4 m high.

2 Plants 2-4 m high, usually lithophytic; inflorescence branches (7)15-30, with a stipe 15-36 cm long, strongly curved; leaves 60-120 cm long; peduncle bracts not imbricate; petals 10.3-10.5 cm long.... T. grandis

2 Plants 1.3-1.75 m high, usually epiphytic; inflorescence branches 3-7, with a stipe 5-11 cm long, straight to slightly curved; leaves 17-45 cm long; peduncle bracts imbricate; petals 10.7-12.3 cm long.... T. macropetala

Tillandsia grandis Schltdl., Linnaea 18: 424-426. 1845 "1844". TYPE: Veracruz, in praeruptis, hacienda de la Laguna, Aug. fruct., VIII.1829, C. Schiede s.n. (Lectotype [W. Weber, 1984] B photo!; Isolectotypes: B photo!, BM, HAL 45629 photo!, LE, OXF). Fig. 2 A.

Rosulate stemless lithophytic or very rarely epiphytic herbs, flowering 2-4 m tall, with solitary tank rosettes to 1.7 m in diameter. Leaves numerous; sheaths brown to light brown on the adaxial surface, dark brown on the abaxial surface, widely elliptic to widely ovate, 10-25 cm long, 10-16 cm wide, densely brown punctulate-lepidote on both surfaces; blades green, paler beneath, strap-shaped, 60-120 cm long, 8-15 cm wide at the base, punctulate-lepidote on both surfaces, acuminate, erect to ascending. Inflorescence terminal, erect, compound, once branched, with 15 to 30 spikes, peduncle terete, to 1 m long, 2.5-3 cm in diameter, partially covered by the bract sheaths; peduncle bracts not imbricate, green, foliaceous, similar to the basal leaves; sheaths 5-6 cm long, blades 5-7 cm long; spikes compressed, oblong, 14-25 cm long, 5-7 cm wide, long stipitate, stipes strongly curved, 15-36 cm long, 8-10 mm in diameter, bracteate, slightly compressed; primary bracts vaginiform, ovate-triangular, 5.5-7 cm long, 4.5-5 cm wide, acuminate; floral bracts dark green, widely elliptic, 3.5-4 cm long, 1.6-2 cm wide, imbricate but the rachis visible after anthesis, nervate, ecarinate, glabrous outside, brown punctulate-lepidote inside, margin erose, apex obtuse to rounded; flowers distichous, erect, 15-20 per spike, actinomorphic, corolla helicoiform, subsessile, pedicels strong, to 1 mm long; sepals free, green, ovate to triangular, 3.4-4 cm long, 1.4-1.6 cm wide, ecarinate, glabrous outside, brown-punctulate lepidote inside, margin erose, apex rounded to acute; petals light green with brown-purple dots, oblong, strap shaped, twisted, 10.3-10.5 cm long, 9-10 mm wide, rounded at the apex; stamens subequal, filaments free, light green to white, filiform, 10-11 cm long, anthers green to green-whithish, linear, curved, 9-12 mm long; ovary green, ovoid, 9-11 mm long, ca. 6 mm in diameter, style green, filiform, 10-11 cm long, stigma green. Capsule green, fusiform, rostrate, 4-6.5 cm long, 6-7 mm in diameter; seeds dark brown, fusiform, ca. 5 mm long, with a white coma ca. 4 cm long.

Specimens examined: Mexico, Guanajuato, municipio de Atarjea, arroyo del rio Blanco, 1300 m, 7.III.1991, E. Ventura & E. Lopez 9153 (IEB, MEXU, XAL). Hidalgo, municipio de Zimapan, 10 km al NW de Zimapan, 1000 m, 30.IV.1965, L. Gonzalez Q. 2376 (ENCB, MICH); municipio de Zimapan, aproximadamente 800 m al sur de las Adjuntas, 16.I.1991, V. M. Huerta 1176 (IEB); municipio de Zimapan, barranca de Toliman, 1520 m, 4.V.1954, E. Matuda 32204 (MEXU, US); municipio de Zimapan, between Zimapan and barranca de Toliman, on road to mine Loma del Toro, 1200 m, 25.IV.1947, H. E. Moore jr. 2602 (GH, UC, US). Oaxaca, municipio de Huautla de Jimenez, cerca de cascada de Chochotla, Huautla, 1700 m, 7.IV.1974, E. Matuda 38623 (LL, MEXU); municipio de Huautla de Jimenez, aproximadamente I. 5 km de Aguacatitla, por la terraceria a Santa Cruz de Juarez, 1589 m, 12.V.2001, X. Munn & T. Kasey 1182 (MEXU); municipio de San Juan Bautista Cuicatlan, a 33 km al SE de Cuicatlan, camino a Oaxaca, 1524 m, 8.I.2001, E. Martinez & V. Torres 33479 (MEXU); municipio de San Juan Bautista Cuicatlan, Santa Maria Almoloyas - Santa Catarina Tlaxila, 1600 m, 10.XII.1991, A. Salinas et al. 6671 (MEXU); municipio de Santiago Nacaltepec, a 49 km al NE de San Francisco Telixtlahuaca, camino a Santiago Dominguillo, 6.II.1966, C. Delgadillo 217 (MEXU, MO, XAL); municipio de Santiago Nacaltepec, El Mirador, headwaters of rio Quiotepec, between Santiago Nacaltepec and Santiago Dominguillo, 6.II.1966, W. R. Ernst 2487 (MEXU, US); municipio de Tepelmeme Villa de Morelos, cerro paraje Ladron (subiendo por Pasto Chino y barranca Copalillo), 1750 m, 14.IX.1994, A. Salinas & E. L. Cruz 7836 (MEXU). Puebla, municipio de Caltepec, mogote Leon, lado NE del cerro Grande, 2000 m, 10.II.1984, P. Tenorio 5434 (MEXU, OAX, TEX); municipio de San Jose Miahuatlan, Agua los Granados, ca. 20 km al W de San Jose Axusco, 1100 m, 24.IX.1990, A. Salinas et al. 5793 (MEXU); municipio de Tehuacan, near Tehuacan, 840 m, 30.VIII.1905, J. N. Rose et al. 10124 (US). Queretaro, municipio de Landa de Matamoros, Tangojo, a la orilla del rio Moctezuma, 400 m, 15.IV.1988, S. Zamudio 6318 (IEB, MEXU, MICH, MO, UAMIZ); municipio de San Joaquin, canada La Culebra, 1 km al NE de la Tinaja, 1600 m, 27.VIII.1978, S. Zamudio 3234 (IEB). Veracruz, municipio de Coatepec, cerro de Achichuca, entre Tuzamapan y Jalcomulco, 900 m, 21.III.1979, G. Castillo & R. Ortega 489 (ENCB, XAL); municipio de Fortin, Fortin de las Flores (cultivated at posada Loma), 850 m, 20.VII.1967, L. I. Nevling & A. Gomez-Pompa 197 (MEXU); municipio de Fortin, between Fortin de las Flores and Orizaba, above the Moctezuma brewery hydroelectric plant, 11.VII.1982, J. Utley & K. Burt-Utley 7052 (MEXU); municipio de Nogales, cerros calizos al sur y suroeste de Orizaba, 1200 m, R. Hernandez & R. Dirzo 2087 (ENCB, MEXU); municipio de Nogales,1-2 km despues de Nogales, sobre la carretera de cuota Orizaba - Puebla, 1350 m, 23.III.1997, A. R. Lopez-Ferrari et al. 2446 (IEB, MEXU, UAMIZ, XAL); municipio de Nogales, a 2 km al E de Nogales, autopista a Cordoba, 900 m, 28.IV.1988, E. Martinez & C. Ramos 22647 (CHAP, IEB, MEXU, MO); municipio de Orizaba, near Orizaba, 1938, M. B. Foster VI (GH); municipio de Orizaba, cerro de San Miguel, Orizaba, 1200 m, 9.III.1971, R. Hernandez & R. Cedillo 1136 (GH, MEXU); municipio de Orizaba, limite N de la ciudad de Orizaba, cerro de calizas junto a Tugrablok S. A., 1300-1500 m, 25.III.1986, D. Lorence 5000 (MEXU); municipio de Puente Nacional, barranca de Panoaya, 790 m, 25.IV.2004, A. R. Lopez-Ferrari et al. 3118 (IEB, UAMIZ); municipio de Puente Nacional, barranca situada 2 km al SE de Palmillas, 550 m, 22.V.1985, M. E. Medina & S. A. Contreras 104 (ENCB, MEXU, XAL); municipio de Totutla, barranca de Tlalpalan, 1100 m, I.1982, M. Chazaro & T. Mejia 2067 (MEXU, XAL).

Tillandsia macropetala Wawra, Wiener III. Gart.-Zeitung 12: 241-244, Fig. 50. 1887. TYPE: E. Morren in Liege hortus s.n. (not located). Lectotype, here designated: Figure 50 in Wiener Illustrierte Garten-Zeitung 12. 1887 (Fig. 3). Epitype, here designated: municipio de Soteapan, faldas del volcan Santa Marta, 18[grados]19' N, 94[grados]50' W, 1300 m, 19.VIII.2006, T. Kromer & E. Otto 2866 (MEXU, SEL, UAMIZ, XAL). Figs. 1A-B, 2 C.

Rosulate stemless epiphytic or very rarely lithophytic herbs, flowering 1.3-1.75 m tall, with solitary tank rosettes to 1.2 m in diameter. Leaves numerous; sheaths brown to light brown on both surfaces, widely elliptic, 13-15 cm long, 8-8.5 cm wide, densely brown punctulate-lepidote on both surfaces; blades green, paler beneath, strap shaped, 17-45 cm long, 3.5-7 cm wide at the base, punctulate-lepidote on both surfaces, acuminate, erect to ascending. Inflorescence terminal, erect, compound, once branched, with 3 to 7 spikes, very rarely simple, peduncle terete, 55-83 cm long, 5-8 mm in diameter; peduncle bracts imbricate, green, foliaceous only the distal vaginiform ones; sheaths 3.5-4.2 cm long, blades 7-14 cm long; spikes compressed, oblong, 9-23.5 cm long, 3-6 cm wide, stipitate, stipes straight to slightly curved, 5-11 cm long, 3.5-5 cm in diameter; primary bracts vaginiform, ovate-triangular, 5.5-7 cm long, 4.5-5.3 cm wide, acuminate; floral bracts green, glaucous, widely elliptic, 3.3-4.3 cm long, 1.4-2.4 cm wide, imbricate but the rachis visible after anthesis, nervate, ecarinate, glabrous outside, brown punctulate-lepidote inside, margin erose, apex rounded to acute; flowers distichous, erect, (5)15-20 per spike, actinomorphic, corolla helicoiform, subsessile; sepals free, green, oblong, 3-3.4 cm long, 1.3-1.5 cm wide, nervate, ecarinate, glabrous outside, brown-punctulate lepidote inside, margin erose, apex rounded to acute; petals light green, strap shaped, twisted, 10.7-12.3 cm long, 14-17 mm wide, rounded at the apex; stamens subequal, filaments free, light green to white, filiform, 10.4-11.7 cm long, anthers green to green-whitish, linear, curved, 7.5-8 mm long; ovary green, ovoid, 9-12 mm long, ca. 6 mm in diameter, style green, filiform, 10.7-11.4 cm long, stigma green. Capsule green, fusiform, rostrate, 5.3-6 cm long, ca. 1 cm in diameter; seeds dark brown, fusiform, ca. 4.5 mm long, with a white coma ca. 3 cm long.

[FIGURA 3 OMITIR]

Specimens examined: Mexico, Chiapas, municipio de Cintalapa, southeast of Cerro Baul on the border with state of Oaxaca, 16 km northwest of Rizo de Oro, along a logging road to colonia Figueroa, 1600 m, 21.IV.1972, D. E. Breedlove 24770 (ENCB, MEXU); municipio de Cintalapa, cerro Baul, 1600 m, 2.V.1999, M. A. Perez Farrera 1954 (UAMIZ); municipio de Tenejapa, near paraje Yashanal, 2460 m, 28.I.1981, D. E. Breedlove 49664 (MEXU): municipio de Villaflores, on the southeast side of cerro Tres Picos and the ridges near summit, 2100-2500 m, 28.III.1973, D. E. Breedlove 34445 (ENCB, MEXU, MICH, MO, TEX); municipio de Villaflores, zona nucleo Tres Picos, 1350 m, 21.III.2004, A. Reyes et al. 6495 (UAMIZ). Oaxaca, municipio de Guevea de Humboldt, recorrido hacia el cerro de la Pena Blanca, al SW de la Cumbre, la cual esta a 13.5 km al N de Guevea de Humboldt, 1300 m, 30.III.1991, A. Campos & R. Torres 3614 (CHAP, IEB, MEXU); municipio de Guevea de Humboldt, sierra Mixe, between Santa Maria Guienagati and Santiago Lachiguiri, 1500-1550 m, 20.II.2005, J. Lautner et al. EM051104 (MEXU, SEL, WU); ibid., 1500-1550 m, 9.II.2006, J. Lautner et al. EM060602 (UAMIZ); municipio de Guevea de Humboldt, camino a Pena Blanca, entrando por la Cumbre, la cual esta a 11 km al N de Guevea de Humboldt, 1230 m, 30.III.1994, R. Torres & L. Cortes 14360 (GH, IEB, MEXU); municipio de Ixtlan de Juarez, 39 km al S de Valle Nacional, sobre la carretera a Oaxaca, 1800 m, 26.VI.1975, J. Rzedowski 33348 (ENCB); municipio de Ixtlan de Juarez, 39 km al S de Valle Nacional, sobre la carretera a Oaxaca, 1800 m, 27.XII.1975, J. Rzedowski 33826 (ENCB); municipio de Ixtlan de Juarez, 10 km al S de Esperanza, carretera Valle Nacional - Oaxaca, 2260 m, 7.VIII.2002, J. Santana & L. Pacheco 907A (UAMIZ); municipio de San Andres Yaa, a 7 km de la desviacion a San Andres Yaa, al S de Villa Alta, camino a Oaxaca, 2030 m, 16.III.1982, R. Cedillo & R. Torres 1201 (ENCB, MEXU, MO); municipio de San Felipe Usila, 5.6 km en linea recta al S (172[grados]) de Santa Cruz Tepetotula, 1970 m, 25.II.1994, C. Gallardo et al. 919 (MEXU); municipio de San Felipe Usila, vereda Monte Pan hacia el campamento del tramo de la carretera, 1650 m, 12.XI.2007, G. Ibarra et al. 3965 (MEXU); municipio de San Felipe Usila, 5.6 km en linea recta al S (172[grados]) de Santa Cruz Tepetotutla, 2000 m, 22.XII.1993, A. Rincon et al. 242 (MEXU); municipio de San Juan Juquila Vijanos, 700 m del entronque Talea - Juquila, sobre la terraceria Maravillas - Talea, 1930-1975 m, 7.III.1977, X. Munn et al. 275 (MEXU); municipio de San Miguel Chimalapa, arroyo Rana, Cerro Azul, 16.III.1948, T. MacDougall s.n. (GH, US); municipio de San Pedro Yolox, 11.5 km despues de la Esperanza, rumbo a Ixtlan, carretera Oaxaca - Tuxtepec, 2068 m, 20.XI.2003, A. Espejo et al. 6700 (UAMIZ); municipio de Santiago Comaltepec, 5 km al W de la Esperanza, carretera Tuxtepec - Oaxaca, 2000 m, 22.XII.1993, R. Torres & E. Martinez 11331 (MEXU); municipio de Santiago Comaltepec, 5 km al W de la Esperanza, carretera Tuxtepec - Oaxaca, 1760 m, 26.I.1988, R. Torres & E. Martinez 11344 (MEXU); municipio de Santiago Comaltepec, 5.7 miles S of Vista Hermosa or 24 miles S of Valle Nacional along hwy 175, 6.VIII.1981, J. Utley & K. Burt-Utley 6728 (MEXU); municipio de Santiago Comaltepec, Vista Hermosa, 20 miles SW of Valle Nacional, NE Oaxaca (Chinantecas Mountains), 1220 m, O. C. van Hyning 5959 (US); municipio de Totontepec Villa de Morelos, rumbo Villa Alta, 16 km al W de Totontepec, 1860 m, 14.IX.1986, E. Ramirez & P. Ramirez 552 (IBUG, IEB, MEXU). Puebla, municipio de Ahuacatlan, desviacion a Amixtlan 3.6 km al S de Ahuacatlan, 1150 m, 14.IV.1985, P. Tenorio et al. 8774 (MEXU); municipio de Ahuacatlan, Agua Dulce, 4 km al SE de Ahuacatlan, brecha a Zapotitlan, 1180 m, 17.V.1987, G. Toriz et al. 414 (MEXU); municipio de Teziutlan, Agua de Obispo, 1450 m, 9.V.1970, F. Ventura 1061 (ENCB, IEB, MEXU); municipio de Xicotepec, 3 km al W de Xicotepec de Juarez (Villa de Juarez), 1100 m, 26.III.1968, L. Gutierrez 168 (ENCB); municipio de Xicotepec, cerca Villa Juarez, al NW, 8.IV.1944, F. Miranda 3205 (MEXU); municipio de Zacapoaxtla, Apulco, cerca de Zacapoaxtla, 1300 m, 10.III.1973, W. Boege 2744 (MEXU); municipio de Zacapoaxtla, cascada de la Gloria, cerca de Apulco, 850 m, 1400 m, 12.IV.1974, J. Rzedowski 31850 (ENCB). Veracruz, municipio de Alto Lucero de Gutierrez Barrios, cerro de la Cima, entre la Sombra y Tierra Blanca, 1700 m, 10.IV.1981, G. Castillo & F. Vazquez 1598 (XAL); municipio de Atzalan, Alseseca, 6.XII.1969, F. Ventura 187 (ENCB); municipio de Chiconquiaco, los Baules, al E de parada Paredes, 1650 m, 9.III.1988, C. Gutierrez 3066 (XAL); municipio de Coatepec, Cinco Palos, 3 km al NW, 1700 m, 21.XI.1993, V. E. Luna et al. 721 (XAL); municipio de Coatepec, Loma Alta, 4 km al E de Cinco Palos, 1900 m, 13.III.1994, V. E. Luna et al. 1423 (XAL); municipio de Coatepec, near Coatepec, 11.IV.1957, M. B. Foster & O. C. van Hyning 3002 (US); municipio de Huatusco, 50 m al este de Dos Puentes, carretera Huatusco - Totutla, 1300 m, 29.VIII.1979, S. Avendano & G. Castillo 486 (XAL); municipio de Jilotepec, Jilotepec, on moist cliffs, 500 ft, 30.III.1959, O. C. van Hyning 598 (US); municipio de Juchique de Ferrer, cerro de Villa Rica, cerca de la hacienda de la Flor, 1280 m, G. Castillo et al. 1779 (XAL); municipio de Orizaba, Orizaba, M. Botteri 65 (GH); municipio de Orizaba, Orizaba, 2550-3000 ft, V.1961, F. Kubisch 14 (US); municipio de Perote, barranca de Patula (cerca de Perote) [barranca de las Minas, Perote], 1400 m, 5.VII.1976, E. Matuda 38720 (CHAP, ENCB, MEXU, MO, XAL); municipio de Perote, barranca de planta mina, Perote, 1400 m, 5.VII.1976, E. Matuda et al. 38734 (MEXU); municipio de Soteapan, cerro Platanillo, ejido de Santa Marta, sierra de Santa Marta, 1500 m, 21.XII.1978, I. Calzada 5094 (XAL); municipio de Soteapan, volcan de Santa Marta, 1720 m, 20.XII.1978, R. V. Ortega 1121 (XAL); municipio de Tatahuicapan de Juarez, faldas del volcan Santa Marta, 1300 m, 19.VIII.2006, T. Kromer & E. Otto 2866 (MEXU, SEL, UAMIZ, XAL); municipio de Tatahuica pan de Juarez, faldas del volcan Santa Marta, 1450 m, 28.VIII.2008, T. Kromer et al. 3713 (CITRO, MEXU); municipio de Tlalnelhuayocan, entre Los Capulines y Rancho Viejo, 1500 m, 24.XI.1990, P. Zamora 2796 (XAL); municipio de Xalapa, Sedeno - Rancho Servin, 6.VI.1993, G. Carmona 50 (XAL); municipio de Yecuatla, Los Capulines, near Paz de Enriquez, ca. 8 km (by air) N of Chiconquiaco, 1400 m, 13.I.1984, K. Taylor et al. 139 (XAL); municipio de Yecuatla, 6 km al NW de Santa Rita, 22.II.1975, M. Vazquez V-1878 (ENCB, MEXU); municipio de Yecuatla, el Haya, 1300 m, 11.II.1971, F. Ventura 3095 (ENCB, MICH).

Smith & Downs (1977) cited Vriesea macropetala hort. (Rev. Hort. 69: 336. 1897) as synonym of Tillandsia viridiflora. However, this name is only mentioned in a publication of the Societe National dHorticulture de France without any description or mention of a basionym. Probably, Smith & Downs (1977) supposed that the name could be a transfer from Tillandsia macropetala Wawra, because of the same epithet, although we think that there is no clear evidence and it might be a nomen nudum.

Tillandsia viridiflora (Beer) Baker, J. Bot. 26: 81. 1888. Platystachys viridiflora Beer, Fam. Bromel. 81. 1856 "1857". TYPE: Mexico, "diese Pflanze wurde in der Gartnerei des Grafen Attems zu Graz aus Samen gezogen, welchen Herr C. Heller aus Mexico sendete. Sie bluhte das erste Mal im November 1854 und ist jetzt in meinem Besitze" (not located). Figs. 1 C-D, 2 D.

Tillandsia orizabensis Baker, J. Bot. 26: 105. 1888. TYPE: Veracruz, Orizaba, region d'Orizaba au Borrego, E. Bourgeau 3055 (Lectotype (Smith & Downs, 1977): K!; Isolectotypes: P!, P!); without precise locality, L. Hahn s.n. (Syntype: K!).

Tillandsia longiflora Sesse & Moc., Fl. mexic. 88-89. 1894 "1887"; Fl. mexic. ed. 2: 81. 1894. TYPE: Hidalgo, habitat supra arbores montium Huehuetlae. Floret Octobri. Totonaci Tacongxnexanat adpellant, M. Sesse & J. Mogino s n. [No. Cuatrecasas 5459; 196-PB] (Holotype: MA 600161!, MA 606715!) (Blanco Fernandez de Caleya et al., 2010; Stafleu & Cowan, 1985).

Rosulate stemless epiphytic herbs, flowering 0.65-1 m tall, with solitary tank rosettes to 40 cm in diameter. Leaves numerous; sheaths brown to light brown, elliptic to oblong, 10-14 cm long, 5.5-6.5 cm wide, densely but inconspicuosly punctulate-lepidote on both surfaces; blades green, slightly glaucous, paler beneath, narrowly triangular, 25-50 cm long, 3-4 cm wide at the base, glabrous on the adaxial surface, sparsely punctulate-lepidote on the abaxial surface, acuminate, erect to ascending. Inflorescence terminal, erect, simple, peduncle terete, 45-50 cm long, 5-7 mm in diameter, totally covered by the bract sheaths; basal peduncle bracts green to reddish, imbricate, vaginiform, only the basal ones foliaceous; sheaths 3.5-4.5 cm long, blades 1-30 cm long; spike compressed, oblong to lanceolate, 17-30 cm long, 6-7 cm wide; floral bracts dark green, glaucous, widely elliptic, 3.2-4 cm long, 2-2.4 cm wide, imbricate but the rachis visible after anthesis, smooth to nervate, ecarinate, glabrous outside, sparsely brown punctulate-lepidote inside, apex rounded; flowers distichous, ascendent, 13-22 per spike, actinomorphic, corolla helicoiform, subsessile; sepals free, green, oblanceolate, 3-3.4 cm long, ca. 1 cm wide, nervate, ecarinate, glabrous outside, sparsely brown-punctulate lepidote inside, margin hyaline and erose, apex acute; petals light green, narrowly oblong, strap shaped, twisted, ca. 9 cm long, ca. 1 cm wide, rounded to acute at the apex; stamens subequal, longer than petals, filaments free, light green to white, filiform, 10-10.5 cm long, anthers green to green-whithish, linear, curved, 9-11 mm long; ovary green, ovoid, ca. 14 mm long, ca. 5 mm in diameter, style green, filiform, 10-11 cm long, stigma green. Capsule green, fusiform, rostrate, 5.7-6 cm long, ca. 9 mm in diameter; seeds dark brown, fusiform, ca. 4 mm long, with a white coma 2.2-2.8 cm long.

Specimens examined: Mexico, Chiapas, municipio de Berriozabal, 13 km north of Berriozabal, near pozo Turipache and finca el Suspiro, 900 m, 12.I.1973, D. E. Breedlove & A. R. Smith 31519 (ENCB); municipio de Berriozabal, Berriozabal - las Vistas, 15.X.1950, F. Miranda 6708 (MEXU); municipio de La Trinitaria, 15 km east northeast of Dos Lagos above Santa Elena, 1000 m, 29.XII.1981, D. E. Breedlove 56607 (MEXU); municipio de La Trinitaria, 15 km east northeast of Dos Lagos above Santa Elena, 1170 m, 19.I.1982, D. E. Breedlove & F. Almeda 57551 (MEXU); municipio de La Triniatria, 13 km al E de Tziscao, 1000 m, 9.VIII.1985, T. Chehaibar et al. 202 (UAMIZ); municipio de Ocosingo, en la comunidad lacandona de Naha, 27 km al sureste de Palenque, por la carretera fronteriza al crucero Chancala, despues 55.6 km por el camino de terraceria hacia Monte Libano, 950 m, 20.XII.1993, A. Duran & S. Levy 55 (MEXU); municipio de Ocosingo, en laguna Ocotalito, a 12 km al N de Monte Libano, camino a Chancala, 980 m, 2.II.1986, E. Martinez 17008 (MEXU); municipio de Ocosingo, laguna Ocotalito, 12 km al N de Monte Libano, camino a Naja, 950 m, 12.IV.1986, E. Martinez & A. Garcia-Mendoza 17980 (MEXU, MO); municipio de Ocosingo, a 10 km al NE de Monte Libano, camino a Chancala, 980 m, 17.VI.1986, E. Martinez & M. A. Soto 18835 (MEXU); municipio de Ocosingo, en laguna Ocotalito a 12 km al N de Monte Libano, camino a Naja, 950 m, 23.V.1987, E. Martinez M-21248 (MEXU); municipio de Ocosingo, alrededor del ex poblado del Chamizal, 830 m, 3.XII.1976, P. Valdivia 2422 (XAL); municipio de Oxchuc, la Cascada, el rio Corralito, 1250 m, 10.IV.1983, A. Mendez Ton (Shilom Ton) 5820 (IEB, MEXU); municipio de Yajalon, arroyo de Grava, 800 m, 16.III.1983, A. Mendez Ton (Shilom Ton) 5684 (MEXU); without precise locality, VI.1981, R. W. Read & P. E. Desautles 84-103 (US). Hidalgo, municipio de Chapulhuacan, 3 km al N de Chapulhuacan, 850 m, 15.III.1960, J. Rzedowski 12322 (ENCB); municipio de Huehuetla, Huehuetla, X, M. Sesse & J. Mocino s.n. (MA); municipio de Lolotla, a 1.55 km al S de Chalma, 1230 m, 27.X.2001, A. Ponce 230 (FCME). Oaxaca, municipio de San Lucas Zoquiapam, Cerro del Fraile, near Huautla de Jimenez, 1830 m, 2.VIII.1938, R. E. Schultes & B. P. Reko 410 (GH); municipio de San Pedro Teutila, el Faro, paraje torre 130 de la L. T. Temascal II - Oaxaca Potencia, 1263 m, 20.I. 2005, G. Juarez & C. A. Cruz 1111 (MEXU); municipio de San Pedro Teutila, el Faro, paraje torre 125 de la L. T. Temascal II Oaxaca Potencia, 670 m, 19.I.2005 C. A. Cruz & G. Juarez 2481 (MEXU); municipio de Santiago Lachiguiri, recorrido de crucero Guadalupe a las cuevas, 12.5 km al NE de Santiago Lachiguiri, 1200 m, 10.V.1991, A. Campos & R. Torres 3679 (MEXU). Puebla, municipio de Xicotepec, Villa Juarez, 1200 m, 3.VII.1969, C. Beutelspacher s.n. (MEXU); municipio de Xicotepec, Loma Linda, 6 km al SE de Xicotepec, carretera a la Union, 1130 m, 11.VI.1985, P. Tenorio & C. Romero de T. 8963 (ENCB, MEXU); municipio de Xicotepec, 5 km al NE de Xicotepec, carretera a La Ceiba, 1200 m, 24.II.1987, P. Tenorio et al. 12603 (MEXU); municipio de Xicotepec, 5 km al NE de Xicotepec, carretera a La Ceiba, 1200 m, 24.II.1987, G. Toriz et al. 288 (MEXU); municipio de Zongozotla, Zongozotla, 0.5 km camino a Huitzila, 840 m, 4.IX.1985, J. L. Martinez & F. Vazquez 735 (XAL). Queretaro, municipio de Jalpan de Serra, 1-1.5 km al E de El Saucito, 1070 m, 7.XII.1989, E. Carranza 2249 (IEB); municipio de Jalpan de Serra, 2-3 km al N de La Parada (Valle Verde), 1150-1200 m, 31.X.1990, B. Servin 639 (IEB); municipio de Landa de Matamoros, km 6 de la brecha de Agua Zarca a Neblinas, 850-1100 m, 24.VIII.1988, S. Zamudio 6716 (IEB); municipio de Pinal de Amoles, ca. 4 km al SE de Santa Agueda, 1190 m, 18.IV.1989, E. Carranza 1640 (IEB). San Luis Potosi, municipio de Rayon, Las Canoas, 17.X.1891, C. G. Pringle 5100 (GH); municipio de Tamazunchale, cerro de San Francisco, Tamazunchale, 1020 m, P. Maury 6258 (GH). Tamaulipas, municipio de Gomez Farias, reserva de la biosfera Rancho del Cielo, in the vicinity of the station, 1170 m, 2.V.1995, P. Hietz 107 (XAL). Veracruz, municipio de Alto Lucero de Gutierrez Barrios, Rancho Nuevo, al W de Plan de las Hayas, 1300 m, 6.IV.1981, G. Castillo & F. Vazquez 1291 (XAL); municipio de Alto Lucero de Gutierrez Barrios, la Piedra Cuata, entre Plan de las Hayas y Rancho Nuevo, 1400 m, 7.IV.1981, G. Castillo & F. Vazquez 1352 (XAL); municipio de Atoyac, montagnes d'Atoyac, XI.1903, A. Duges 20 (GH); municipio de Atoyac, mountains of Atoyac, X.1899, A. Duges s.n. (US); municipio de Atzalan, Comunidad Cruz Gorda, congregacion San Salvador, 990 m, 17.VI.2008, T. Kromer et al. 3320 (CITRO, MEXU, XAL, XALU); municipio de Chiconquiaco, Santa Rita Chiconquiaco, 1500 m, 28.II.1975, V. Sosa 88 (XAL); municipio de Cordova, Vallee de Cordoba, 18.XII.1865, E. Bourgeau s.n. (P); municipio de Huatusco, puente antiguo, ca. 5.6 km antes de llegar a Huatusco, viniendo de Totutla, 1290 m, 20.VII. 2007, A. Espejo et al. 7054 (UAMIZ); municipio de Jilotepec, El Esquilon, cerro del Cuajilote, 26.XII.1978, G. Castillo 411 (XAL); municipio de Jilotepec, 2.5 km NE of Jilotepec, along road to Naolinco, 1350 m, 10.VII.1982, G. Diggs & M. Nee 2776 (GH, MO, XAL); municipio de Mecayapan, Santa Marta, al N de Ocotal, 1450 m, 17.VIII.1980, F. Ramirez 1186 (XAL); municipio de Mecayapan crater del volcan Santa Marta, por Ocotal Chico, 900 m, 12.III.1981, F. Ramirez & F. Vazquez 1535 (XAL); municipio de Mecayapan, faldas del volcan Santa Marta, 1090 m, 19.VIII.2006, T. Kromer & E. Otto 2865 (MEXU); municipio de Misantla, Misantla, VI.1866, L. Hahn s.n. (P); municipio de Naolinco, cascada de Naolinco, 1530 m, 3.VII.2003, A. Mendoza et al. 704 (UAMIZ); municipio de San Andres Tuxtla, dentro del crater del volcan San Martin Tuxtla, lado S, 1450 m, 31.V.1984, I. Calzada 10666 (XAL); municipio de San Andres Tuxtla, cerro Coyolar al N de Catemaco, 29.V.1985, I. Calzada 11805 (XAL); municipio de San Andres Tuxtla, cima del cerro Mastagaga, al N del ejido Ruiz Cortines, sierra de los Tuxtlas, 30.V.1985, I. Calzada 11825 (XAL); municipio de San Andres Tuxtla, falda del volcan San Martin, 1300 m, 14.IV.2005, T. Kromer et al. 1956 (MEXU, XAL); municipio de San Andres Tuxtla, ejido Ruiz Cortines, falda del volcan San Martin Tuxtla, 1000 m, 1.VII.2005, T. Kromer & A. Acebey 2299 (MEXU); municipio de Soteapan, crater del volcan Santa Marta, SE del rio Xochiapan, 670 m, 15.V.1985, I. Calzada 11434 (XAL); municipio de Soteapan, Santa Marta, ladera W del volcan Santa Marta, F. Ramirez 969 (XAL); municipio de Tatahuicapan de Juarez, faldas del volcan Santa Marta, 1350 m, 28.VIII.2008, T. Kromer et al. 3714 (MEXU); municipio de Tepetzintla, en la Sierra de Otontepec, al NE de Tepetzintla, 1200 m, 27.VIII.1981, G. Castillo & A. Benavides 2249 (XAL); municipio de Tepetzintla, sierra de San Juan Otontepec, 900 m, 13.XII.1981, G. Castillo & A. Benavides 2384 (XAL); municipio de Tepetzintla, sierra de Otontepec, 1250 m, 14.XII.1981, G. Castillo & A. Benavides 2542 (XAL); municipio de Tepetzintla, Sierra de Otontepec, al NE de Tepetzintla, 1170 m, 25.VIII.1981, F. Vazquez & L. Tapia 208 (XAL); municipio de Tepetzintla, sierra de la Pena Blanca, sierra de Tantima, 21.IX.1989, P. Zamora et al. 1226 (XAL); municipio de Totutla, el Mirador, 21 km E of Huatusco, at km 45 along hwy to Puente Nacional, 23.VIII.1977, T. Croat 43973 (MEXU, MO); municipio de Totutla, Mirador, 1841-43 m, F. Liebmann 7 (UC); municipio de Totutla, Mirador, VIII.1930, C. A. Purpus 15763 (UC); municipio de Xalapa, pres de Jalapa, 4.IX.1866, L. Hahn 594 (P); municipio de Xico, gorge at puente Acabaloya, ca. 1 km SE of Xico Viejo and 5 km NW of Xico, along trail between the two, 1600 m, 31.III.1983, M. Nee & K. Taylor 26268 (XAL); municipio de Zongolica, reserva ecologica La Quinta, Amatitla, 1250 m, 6.III.2008, T. Kromer et al. 3131 (CITRO, MEXU, XAL).

It is also important to mention that sometimes plants of Tillandsia heterophylla (Fig. 2 B) are confused in herbarium material with T. viridiflora and/or T. macropetala because of the branched inflorescences and overall morphology of the floral bracts. However, in living plants, the flowers of T. heterophylla have infundibuliform and slightly zygomorphic white corollas with included stamens, while the flowers of T. viridiflora and T. macropetala have helicoiform actinomorphic green corollas, spirally twisted petals, and the stamens are exposed. In addition the plants of T. heterophylla are conspicuously glaucous and pruinose.

Finally, we suggest that the name Vriesea billbergiae Lem. (Tillandsia billbergiae (Lem.) Baker) should probably be excluded from the synonymy of T. viridiflora because the original description of the former mentioned the presence of a "gland" at the base of petals: "... [segmentis] internis basi ima glandula maxima rotundato-biloba tenuissima instructis This character is not present in any species of Tillandsia and we think that this name eventually will be assigned to some species of Vriesea Lindl., as this genus in its current circumscription is separated from Tillandsia solely on floral characters (Till, 2000). However, as the description could be wrong, it will be necessary to study the type specimen (Type: Ghiesbreght in Verschaffelt Hortus, Ghent, no date, Mexico) in order to confidently exclude the name.

All three species of the Tillandsia viridiflora complex show a wide distribution in the central and southern states of Mexico (Fig. 4), where they principally occur in or near montane rain or moist pine-oak forests along the slopes of the Sierra Madre Oriental, Sierra de Juarez, and Sierra de Chiapas. However, they can also be found in humid to moderately dry lowland forests. T. grandis occurs at 400 m to 2000 m elevation and grows primarily lithophytically on very steep, inaccessible limestone cliffs, but it is very occasionally epiphytic. T. macropetala can be found at elevations of (150)1100 m to 2500 m, whereas T. viridiflora occurs from 670 m to 1830 m elevation, both species usually occur as epiphytes and only rarely are found terrestrially. Since the distributions of T. grandis and T. viridiflora range from Mexico to Nicaragua (Beaman & Judd, 1996), it is very likely that a future review of herbarium material in combination with the characters used in our identification key also will demonstrate the existence of T. macropetala in Central America, as the latter species is widespread in the southern states of Mexico, and thus probably not endemic to the country.

Their flowering periods are only slightly different, with Tillandsia grandis producing flowers between March and July, whereas T. macropetala and T. viridiflora start blooming in January and last until August (Espejo et al., 2005). In addition to their similar flower morphology and phenology, all three species show inconspicuous greenish inflorescences and a nocturnal anthesis, in contrast to most Tillandsia species that have brightly colored bracts with contrasting flowers which are pollinated by hummingbirds (Gardner, 1986; Benzing, 2000; Kessler & Kromer, 2000). Hietz & Hietz-Seifert (1994) suggest sphingid moths as the most likely visitors, because of their long spreading stamina and pistil exerted from an open actinomorphic flower. However, personal observations by the senior author, combined with the identification of hexose-rich nectar in T. viridiflora by Kromer et al. (2008), suggest bat-pollination, although the plants in the complex do not show the typical chiropterophilous floral syndrome with bell-shaped, large zygomorphic flowers, characteristic of the bat-pollinated Werauhia species (Kromer et al., 2007). Furthermore, since the flowers of T. macropetala and T. viridiflora are still open during the next morning, they might also be visited by hummingbirds and show a bimodal pollination system.

[FIGURA 4 OMITIR]

ACKNOWLEDGMENTS

We thank Amparo Acebey, Manfred Kretz, Ulrich Lautner, Gunther Noller, Elisabeth Otto, and Wolfgang Schindhelm for fieldwork assistance, Walter Till, Bruce K. Holst, and two anonymous reviewers for valuable comments on the manuscript, Lilia Ruiz-Ruiz for preparing the figures 1-3, Nancy Martinez-Correa for preparing the distribution map and the curators of CHAP, ENCB, FCME, GH, IBUG, IEB, K, LL, MA, MEXU, MICH, MO, OAX, OXF, P, SEL, TEX, UAMIZ, UC, US, WU and XAL for providing the facilities to consult specimens and data. Sergio Zamudio sent us images of T. viridiflora deposited at IEB. To Elizabeth Bascom Scholarship from the Missouri Botanical Garden granted to A. R. Lopez-Ferrari to study the Mexican Bromeliaceae collections deposited at the main U.S. herbaria. The study of T. Kromer was supported by a postdoctoral grant from the Universidad Nacional Autonoma de Mexico and a research grant from the Bromeliad Society International.

Recibido en marzo de 2011.

Aceptado en noviembre de 2011.

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THORSTEN KROMER (1), ADOLFO ESPEJO-SERNA (2,5), ANA ROSA LOPEZ-FERRARI (2), RENATE EHLERS (3) & JURGEN LAUTNER (4) ([dagger])

(1) Universidad Veracruzana, Centro de Investigaciones Tropicales, interior de la ex-hacienda Lucas Martin, privada de Araucarias s.n., colonia 21 de Marzo, 91119 Xalapa, Veracruz, Mexico.

(2) Universidad Autonoma Metropolitana-Iztapalapa, Division de Ciencias Biologicas y de la Salud, Departamento de Biologia, Herbario Metropolitano, Apdo. postal 55-535, 09340 Mexico, D.F., Mexico.

(3) Herrenbergerstr 14, 70563 Stuttgart, Germany.

(4) Albrecht von Haller Institute of Plant Sciences, Untere Karspule 2, 37073 Gottingen, Germany. ([dagger]) Died 2 October 2009.

(5) Autor para la correspondencia: aes@xanum.uam.mx
Table 1. Comparison of selected morphological characters to
distinguish the Mexican species of the Tillandsia viridiflora
complex.

                  Tillandsia grandis   Tillandsia macropetala

Plants                2-4 m high          1.3-1.75 m high

Leaf blades       strap shaped, 60-    strap shaped, 17-45 cm
                   120 cm x 8-15 cm          x 3.5-7 cm

Inflorescence       once branched,       once branched, 3-7
                     15-30 spikes      spikes, rarely simple

Peduncle             to 1 m long           55-83 cm long

Peduncle bracts     not imbricate,     imbricate, foliaceous,
                      foliaceous        only the distal ones
                                             vaginiform

Petals              10.3-10.5 cm x         10.7-12.3 cm x
                       9-10 mm                14-17 mm

                  Tillandsia viridiflora

Plants                0.65-1 m high

Leaf blades        narrowly triangular,
                    25-50 cm x 3-4 cm

Inflorescence             simple

Peduncle              45-50 cm long

Peduncle bracts   imbricate, vaginiform,
                   only the basal ones
                        foliaceous

Petals                 9 cm x 10 mm
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