Dioon edule Lindley (1843: 59)

Hernández-Tapia, Jessica E., Jiménez-Ramírez, Jaime & Vovides, Andrew P., 2020, Taxonomic revision of the genus Dioon (Zamiaceae), Phytotaxa 442 (4), pp. 267-290 : 277-279

Dioon edule Lindley (1843: 59) . Holotype:— Cultivated from material collected in Mexico, Lindley s.n. (CGE).

= Zamia maeleni Miquel (1844: 97) .

= Platyzamia rigida Zuccarini (1845: 25) . Holotype:— MEXICO. In Imperio Mexicano, without date, Karwinski von Karwin s.n. (M).

= Dioon imbricatum Miquel (1847: 36) . Lectotype (designed by Stevenson & Sabato 1986):—Tab. IV, d-e in Verh. Eerste Klasse Kon. Ned. Instituut Wet. Ser. 3, 4: 1851.

= Dioon edule var. imbricatum (Miquel) Miquel (1868: 427) . Homonyme: Dioon edule var. imbricatum Abel (1886: 167) . Basonym: Dioon edule fo. imbricatum (Miquel) Miquel (1861:22) .

= Dioon strobilaceum Lemaire (1863: 4) . Homonyme: D. strobilaceum Lemaire ex A. DC. (1868: 537) . Lectotype (Designed by Stevenson & Sabato 1986): Description.

= Macrozamia littoralis Liebmann ex Hemsley (1883: 191) . Type:—Icon Palmarum lamina 51–53 s.n.

= Macrozamia pectinata Liebmann ex Hemsley (1883: 191) . Type:—Icon Palmarum lamina 54–56 s.n.

= Dioon edule var. lanuginosum Wittmack (1897: 153) . Lectotype (designed by Stevenson & Sabato 1986). Basonym: Dioon edule fo. l anuginosum (Wittmack) J. Schuster (1932: 127).

Plants 1–3 m tall, sometimes up to 5 m tall. Stems erect to semipostrate branching in old specimens, 22–30 cm diameter. Cataphylls triangular-acuminate to cuspidate, lanate, 5–10 cm long, 2 –4 cm wide. Leaves 5–40 per crown, 55–150 cm long, glabrescent, trichomes white-brown. Leaflets 60–140 pairs, linear-lanceolate, the apex acuminate, flat-convex, not imbricate, sometimes slightly imbricate, covering less than one third of the width, subopposite to opposite, margin entire, with 9–12 veins, the apical leaflets 0.8–3.9 cm long, intermediate ones 4–14 cm long and the basal ones 0.6–7 cm long, all leaflets 0.4–1.5 cm wide, 0.1–0.4 cm distance between articulations, without pinnacanths, sometimes with them, tomentose on the underside or absent; rachis straight to rarely twisted at the apex, terete, tomentose to glabrate; petioles terete, 10–32 cm long, 0.5–2.1 cm wide, lanate at the base and tomentose, sometimes glabrate. Male strobili cylindrical, 35 cm long, 4–8.5 cm wide, slightly stalked, apparently sessile; microsporophylls cuneate, tomentose to slightly lanate, sometimes glabrate, recurved and mucronate in the distal region, 1.8–3 cm long, 2–3 cm wide, sori with 3–4 sporangia, rarely 5. Female strobili ovoid to cylindrical 14–35 cm long, 10–20 cm wide, peduncle short; megasporophylls deltoid-acuminate to trulate, flat to recurved, 9.2 cm long, 2–4 cm wide, lanate on the outer surface, glabrate on the inner surface, the apex sometimes glabrate and the base lanate, sometimes lanate on both surfaces, except for the regions where the seeds are developing. Seeds ovoid, sometimes spherical, 1–3 cm long, 1.5–2.5 cm wide, sarcotesta bright yellow, medium yellow to orange-yellow.

Distribution and habitat: —This species has a wide distribution range and can be found along the Gulf of Mexico and in the Sierra Madre Oriental, in the states of Hidalgo, Querétaro, San Luis Potosí, Veracruz and Tamaulipas. It can be found in different types of vegetation such as oak forest, tropical deciduous forest, xeric shrublands and coastal dunes. It thrives in semiarid to humid warm-temperate climates, in an altitudinal range of 0–1550 m. The specimens of CDMX correspond to cultivated plants.

Uses and common names:—It has several uses. In some regions, seeds are grinding into a flour-like product. Leaves are used as ornament in churches during religious congregations. The testa of seeds is also used for making toys called rezumbadores. The plant also has ornamental uses. Some common names of this specie are chamal, chamalito, quiotamal, sotol, palma de Dolores, palma de macetas, palma de Santa Teresita, palama de serrucho, palma de la virgen, palmita, tullidora ( Bonta et al. 2019, Bonta & Osborne 2007).

Taxonomic notes:—It can be distinguished from the other species owing to the non-imbricate leaflets with entire margins and megasporophylls trulate to deltoid-acuminate. Dioon edule morphologies, proposed by Whitelock (2004), were evaluated and were not valid since no morphological pattern was found for each locality, finding several morphologies in a single locality. Therefore, the species was managed as a polymorph.

The name Dioon edule var. edule and Dioon edule fo. edule is in fact an invalid name because it lacks publication and type specimen.

Specimens examined:— MEXICO. Hidalgo: Jacala, 832 m, 21/March/2003, Contreras-Medina R. et al. 54 (MEXU, FCME); Jacala, 900 m, without date, Contreras-Medina R. et al. 53 (FCME); Tlanchinol, 582 m, no date, Rendón R. Ma. s.n. (FCME). CDMX: Polanco, Carbajal M. 45 (MEXU); Jardín Botánico CU, Cuspinera s.n. (FCME). Querétaro: Cadereyta de Montes, 1455 m, 12/June/2001, Walters T. et al. TW 2001–06 (XAL); Jalpan de Serra, 1100 m, 16/February/1985, Fernández R. 2690 (ENCB); Jalpan de Serra, 1250 m, 17/August/1985, Servín B. 428 (MEXU); Jalpan de Serra, 900 m, 29/December/1977, Vovides A. P. 336 (XAL); Jalpan de Serra, 650 m, 15/May/1993, Zirahuén-Ortega R. 565 (XAL, MEXU); Landa de Matamoros, 1403 m, 8/September/2006, Ceja J. et al. 1793, 1794 (UAMIZ); Landa de Matamoros, 1455 m, 12/January/2001, Walters T. et al. TW 2001–06 (MEXU). San Luis Potosí: Ciudad del Maíz, 25/September/1960, Medellín-Leal F. 611 (ENCB); Ciudad del Maiz, 914 m, 18/July/1955, R. M. & A. F. Tryon 5157 (MEXU); Ciudad del Maiz, 850 m, 7/November/1960, Rees J. 1612 (XAL); Ciudad del Maiz, 650 m, 13/January/2001, Walters T. 2001–09–A, 2001–09–B (XAL); Tamasopo, 700 m, 30/June/1962, Medellín-Leal F. 1307 (ENCB); Tamasopo, 1300 m, 30/June/1962, Medellín-Leal F. 1328 (ENCB); Tamasopo, 770 m, 30/July/1962, Medellín-Leal F. 1307 (MEXU); Tamasopo, 384 m, 19/January/2008, Rubio-Méndez G. 2 (XAL); Tamasopo, 447 m, 19/January/2008, Rubio-Méndez G. 3 (XAL); Tamasopo, 447 m, 20/April/2008, Rubio-Méndez G. 4 (XAL); Tamasopo, 750 m, 11/September/1967, Rzedowski J. 24570, 24434 (ENCB); Tamasopo, 700m, 28/05/1959, Rzedowski J. 10627 (ENCB); Tamasopo, 700 m, 15/August/2003, Vovides A. P. 1464 (XAL); Rayón, 1078 m, 19/ September/2007, Rubio-Méndez G. 1 (XAL); Ciudad Valles, 300 m, Bracamontes Ma. R. 267 (FCME); Ciudad Valles, 944 m, 28/April/1974, Fyrell P. & Magdill R. 2345 (ENCB); Ciudad Valles, 300 m, 5/August/1971, Gibson A. C. & Gibson L. C. 2659 (ENCB); Ciudad Valles, 370 m, 20/July/1983, Stevenson D. W. 564 P, 564 J, 564 O, 564 K, 564 N (MEXU). Tamaulipas: Ciudad Victoria, 1143 m, 1/June/2004, Corona S. 3 (MEXU); Ciudad Victoria, 610 m, 15/ February/1949, Clausen R. T. & Edwards J. L. 7377 (ENCB); Ciudad Victoria, 400 m, 15/November/1985, Jiménez J. 356 (ENCB); Ciudad Victoria, 1200 m, 1/February/1991, Pérez E. & Zamudio S. 2087 (XAL, MEXU); Ciudad Victoria, 1470 m, 3/July/2002, Zamudio S. 11876 (XAL, MEXU); Jaumave, 1700 m, 1/August/1972, McGrill L. et al. 9720 (ENCB); Ocampo, 623 m, 14/January/2001, Walters T. 2001–12–A (XAL); San Carlos, 450 m, 21/July/1930, Barlett H. 10538 (ENCB); San Carlos, 700 m, 23/March/1986, Martínez M. 979 (ENCB); Santa Cruz de Bezana, 17 m, 22/February/1972, Díaz-Moreno J. M. s.n. (ENCB); Villa Mainero, 1350 m, 26/May/1974, Duncan T. 2398 (ENCB). Veracruz: Actopan, 300 m, 17/November/1984, Acosta R. & Acosta N. 72 (XAL); Actopan, 200 m, 29/ September/2003, Nicolalde-Morejón F. et al. 1413 (XAL); Actopan, 20 m, 22/May/1988, Vovides A. P. 1146 (XAL); Actopan, 20 m, 5/April/2001, Vovides A. P. 1394 (XAL); Alto Lucero de Gutiérrez Barrios, 130 m, 4/December/1974, Rees J. 1638 (MEXU, XAL); Alto Lucero de Gutiérrez Barrios, 100 m, September/1976, Rees J. 1674 (MEXU, XAL); Alto Lucero de Gutiérrez Barrios, 130 m, 4/November/1974, Rees J. 1639 (MEXU, XAL); Alto Lucero de Gutiérrez Barrios, 1194 m, 8/June/1994, Sevilla D. DJS697 (XAL); Alto Lucero de Gutiérrez Barrios, 586 m, 25/ July/2008, Vázquez M. MVT8678 (MEXU, XAL); Coatepec, 680 m, 19/March/1983, Nee M. & Taylor K. 26032–a (XAL); Coatepec, 560 m, 7/December/1974, Rees J. 1645 (XAL); Coatepec, 600 m, 30/October/1980, Vovides A. P. 626 (XAL); Coatepec, 844 m, 17/September/1990, Zamora P. 2616 (XAL); Colipa, 300 m, 2/December/1974, Rees J. 1644 (XAL); Emiliano Zapata, 1437 m, 18/June/1994, Castillo-Campos G. et al. 12408 (XAL); Emiliano Zapata, 845 m, 14/July/2016, Castañeda-Zarate M. Colorada-Durán W. B. MCZ-1267 (MEXU); Emiliano Zapata, 500 m, 19/November/1974, Rees J. 1621, 1622 (XAL); Fortín de las Flores, 950 m, 10/April/1969, Nevling & Gómez-Pompa 625 (MEXU); Jalcomulco, 560 m, 30/June/2005, Carrillo-Reyes & Ortiz-Catedral L. 4620 (XAL); Jalcomulco, 350 m, 26/November/1991, Castillo G. & Zamora P. 7524 (XAL); Jalcomulco, 340 m, 12/July/1991, Castillo G. & Zamora P. 8262 (XAL); Jalcomulco, 1580 m, 21/January/1976, Rees J. 1669 (XAL); Vega de Alatorre, 550 m, 23/July/1981, Castillo G. & Benavides A. 2114 (XAL); Xalapa, 1270 m, 29/August/2007, Luna V. E. 1578, 1579, 1580, 1581, 1582, (XAL); Xalapa, 1308 m, 8/April/1994, Zamora P. 3813 (XAL); Xalapa, 900 m, 2/October/1975, Zola M. G. 830 (XAL, MEXU).

Dioon holmgrenii De Luca, Sabato & Vázquez Torres (1981: 552) . Type:— MEXICO. Oaxaca: Juquila, 720 m, 29/ May/1979, Vázquez-Torres V-2304 (holotype: XALU!).

Plants 6–7 m tall, unbranched, stems 30–40 cm diameter. Cataphylls triangular-acute, lanate, 5–9 cm long, 2–4 cm wide. Leaves 8–20 per crown, 100–150 cm long, glabrescent, trichomes whitish-brown. Leaflets 74–140 pairs, linear-lanceolate, acuminate, convex, not-imbricate, subopposite, margin spinose with 3–4 prickles, rarely 1–6 prickles, veins 15, the apical leaflets 1.5–5.1 cm long, the intermediate ones 7.2–12.5 cm long and the basal ones 1.8–3.3 cm long, all leaflets 0.3–0.9 cm wide, distance between articulations 0.1–0.5 cm, without pinnacanths; rachis straight to twisted at the apex, semiterete, tomentose to glabrate; petioles semiterete, 13–30 cm long, lanate at the base. Male strobili cylindrical, 40 cm long, 7.5 cm wide, peduncles short, appearing sessile; microsporophylls cuneate, tomentose, mucronate distally, 1.2 cm long, 2 cm wide. Female strobili ovoid to cylindrical, 30–50 cm long, 20–30 cm wide, peduncles short, appearing sessile; megasporophylls trulate, flat, 5–12 cm long, 2.8–4.5 cm wide, lanate on the outer surface, the inner surface glabrate. Seeds subglobose to ellipsoid, 3.6 cm long, 3 cm wide, sarcotesta yellow.

Distribution and habitat.—It is distributed in tropical deciduous forest, oak forest, pine-oak forest and in the ecotone between these types of vegetation in the state of Oaxaca. The altitudinal range is 450–1500 m and can be found in coastal dunes, but not at sea level. It thrives in warm-humid to temperate climates. The specimens of Veracruz are cultivated plants.

Uses and common names.—It is used only as an ornamental plant. In the regions where it is found is called mirasol, palma sol and plumilla (Bonta & Osborn 2007).

Taxonomic notes.—It is distinguished mainly for its non-imbricate, convex leaflets with 3–4 marginal prickles, the apex of the cataphylls is acute, the rachis is twisted at the apex and the megasporophyll is trulate. It is akin to Dioon planifolium , but this species presents cataphylls with an acuminate apex, flat leaves, leaflet margin entire or with 1 marginal prickle and the megasporophylls are deltoid-acuminate.

Specimens examined:— MEXICO. Oaxaca: San Bartolomé Loxicha , 800 m, without date, Nicolalde-Morejón F. et al. 1468 ( XAL) ; San Bartolomé Loxicha , 730 m, 23/February/2000, Salas-Morales S. 2777 ( OAX, SERO, MEXU) ; San Bartolomé Loxicha , 580 m, 23/February/2000, Salas-Morales S. 2779 ( SERO) ; San Bartolomé Loxicha , 978 m, 21/May/2004, Walters T. et al. TW 2004–21 A ( XAL, MEXU) ; San Gabriel Mixtepec , 500 m, 25/May/2005, Nicolalde-Morejón et al. 1475 ( XAL) ; San Gabriel Mixtepec , 600 m, 26/October/1973, Rees J. 1602 ( XAL) ; San Gabriel Mixtepec , 600 m, 22/October/1973, Rees J. 1604 ( XAL) ; San Pedro Jucahtengo , 1025 m, 9/February/2000, Salas-Morales S. 2686 ( SERO, MEXU) ; San Pedro Mixtepec , 488 m, 16/October/1993, Brigata T. Walters (7–1) ( XAL) ; San Pedro Mixtepec , 565 m, 20/May/2004, Walters T. et al. TW 2004–20 A ( SERO, XAL, MEXU) ; San Pedro Mixtepec , 980 m, 21/May/2004, Walters T. et al. TW 2004–21 B ( SERO) ; Santa Cruz Itundujia , 1360 m, 23/January/2008, Salas-Morales S. 6366 ( SERO) ; Santa Cruz Itundujia , 1461 m, 23/June/2007, Velasco-Gutiérrez 1986 ( SERO, MEXU) ; Zapotitlán Palmas , 2022 m, 9/December/1983, Torres R. 4286 ( MEXU) . Veracruz: Xalapa , 1270 m, 29/August/2007, Luna V. E. 1583, 1584 ( XAL) .