Ceratozamia robusta Miq., Tijdschr. Wis-Natuurk. Wetensch. Eerste Kl. Kon. Ned. Inst. Wetensch. 1: 42. 1847

Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco & Stevenson, Dennis Wm., 2022, Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus, PhytoKeys 208, pp. 1-102 : 1

26. Ceratozamia robusta Miq., Tijdschr. Wis-Natuurk. Wetensch. Eerste Kl. Kon. Ned. Inst. Wetensch. 1: 42. 1847

Fig. 27C

Ceratozamia mexicana Brongn. var. robusta (Miq.) Dyer, Biol. Cent.-Amer., Bot. 3: 193. 1884. Type. Based on: Ceratozamia robusta Miq.

Type.

Mexico. Chiapas: NW of Tuxtla Gutiérrez on road through San Fernando 23.5 km from route 190, 1,000 m, 9 Jul 1983, D.W. Stevenson 549A (neotype, designated by Stevenson and Sabato 1986, pg. 581: NY! [00001119]) .

Description.

Stem 30-100 cm long, 20-40 cm in diameter, epigeous, erect and decumbent. Cataphylls 5.0-9.0 × 3.5-7.5 cm wide at the base, persistent, long triangular, reddish brown, densely brown tomentose at emergence, partially tomentose at maturity, apex acuminate. Leaves 10-50, 138-266 cm long, ascending, dark brown at emergence, green and glabrous at maturity. Petiole 56-85 cm long, terete, linear, brown in young leaves, green in mature leaves; with 27-55 robust prickles, 0.48-0.70 cm long. Rachis 60-190 cm long, terete, linear, brown in young leaves, green in mature leaves, with prickles. Leaflets 20-45 pairs, opposite to subopposite, insertion in one plane, lanceolate, in general longitudinally planar, not basally falcate, papyraceous, flat, green with adaxial side glabrous and abaxial side glaucous, distal end with entire margins, acuminate and asymmetrical at the apex, attenuate at base, with conspicuous and light-green veins; median leaflets 28-42 × 2.5-4.0 cm, 3.0-4.5 cm between leaflets; articulations 0.2-0.8 cm wide, green. Pollen strobili 50-70 cm long, 7.0-8.5 cm in diameter, solitary, cylindrical, erect, light green to yellowish green with blackish trichomes at emergence, yellowish cream with reddish brown trichomes at maturity; peduncle 10-15 cm long, 2.5-3.2 cm in diameter, tomentose, reddish brown to brown; microsporophylls 2.33-3.0 × 1.14-1.90 cm, obconic with a non-recurved distal face and deeply lobate fertile portion, infertile portion 0.45-0.65 cm long and rounded with curved horns 0.26-0.40 cm long, 0.55-0.75 cm and a right angle between the horns. Ovulate strobili 26-40 cm long, 11-14.5 cm in diameter, solitary, cylindrical, erect, dark green at emergence, dark green with scarce blackish trichomes at maturity, acuminate apex; peduncle 5.0-11.0 cm long, 2.5-4.0 cm in diameter, erect, tomentose, reddish brown; megasporophylls 153-280, 8-14 orthostichies with 17-20 sporophylls per orthostichy, 1.7-2.5 × 2.3-3.5 cm, with a prominent distal face, horns straight and 0.38-0.50 cm long, 1.35-1.60 cm between horns with a right angle between the horns. Seeds 2.5-3.5 cm long, 1.9-2.5 cm in diameter, ovate, sarcotesta whitish yellow to yellow when immature, light brown at maturity.

Distribution and habitat.

Ceratozamia robusta is widely distributed in Chiapas State, Mexico and northern Guatemala (Fig. 22B ). It occurs in evergreen tropical forest between 400 to 1,300 m on karstic rocks with humus soil.

Etymology.

The epithet is derived from the robust appearance of the species.

Common names.

Mexico. Chiapas: Amendu (M.A. Pérez-Farrera 30), piña del tlacoache (A.P. Vovides & J.I. Calzada 481).

Uses.

The seeds are used as food, and the leaves to decorate altars.

Conservation status.

( IUCN 2021). Ceratozamia robusta is listed as “Endangered” under criteria A2acd. This species has a wide distribution, but some populations previously considered to be part of this entity have been assigned to described species durint the last ten years. Therefore, a reassessment of its conservation status is required.

Discussion.

Ceratozamia robusta is similar to C. subroseophylla and distinguished by its dark green ovulate strobilus with sparse blackish trichomes on megasporophylls at maturity, prominent distal face, straight horns with a right angle between them, differentiating it easily from C. subroseophylla . The pollen strobilus of C. robusta is the largest in the genus (up to 70 cm long).

Specimens examined.

Guatemala. Alta Verapaz: 200 m, 13 Jul 2002 , S. Hernández 757 (MO); 1,800 -2,000 m, 27 Feb 1939, C.L. Wilson 262 (F); 300-500 m, 17 Apr 1942, J.A. Steyermark 45734 ( US); 879 m, 15 Mar 2008 , M. Véliz et al. 19998 (BIGU). Huehuetenango: 2,000 m, 10 Sep 1942 , J.A. Steyermark 52046 (NY, US); 1,150 m, 26 Jul 1943 , J.A. Steyermark 49506 (F; MO; NY); 400 m, 27 Jul 1942, J.A. Steyermark 49682 (NY); 900- 1,300 m, 3 Sep 1942, J.A. Steyermark 51818 (F; NY; US); 900- 1,000 m, 6 Mar 2009 , M.J.M. Christenhusz et al. 5600 (MO); 1,161 m, 21 Dec 2010, L. Vélasquez et al. 1566 (BIGU). Mexico. Chiapas: Mun. Berriozabal , 1,129 m, 6 Mar 2014 , F. Nicolalde-Morejón et al. 1970-1975 (CIB); 1,129 m, 6 Mar 2014, L. Martínez-Domínguez et al. 41-47, 49, 50 (CIB), 48 (CIB, MEXU). Mun. Copainalá, 1,100 m, M. Gutiérrez & T. Acero 240 (CHIP). Mun. Ocosingo , 900 m, 16 Mar 1981 , A.P. Vovides & J.I. Calzada 481 (XAL); 14 Apr 1967 , D.E. Breedlove 15687 (ENCB); 700 m, 18 Apr 1985, E. Martínez S. 12067 (MEXU); 860 m, 24 Feb 1984, J. García F. 720 ( CH, CIB, XAL). Mun. Ocozocoautla de Espinosa , 818 m, 11 Nov 1997 , R. Álvarez G. 6 (CHIP). Mun. Osumacinta , 692, 18 Jul 2008 , R. Gallegos Ramos 211 (CHIP). Mun. San Fernando , 680 m, 29 Apr 1995 , A.P. Vovides & M.A. Pérez-Farrera 1266ª (XAL); 22 Sep 1993 , Brigada T. Walters s/n (XAL); 9 Jul 1983, D.W. Stevenson et al. 549 B - F (NY); 1,000 m, 9 Jul 1983, D.W. Stevenson et al. 550 A-C (NY), 24 Apr 1999, M.A. Isidro V. 388 (CHIP); 29 Apr 1995, M.A. Pérez-Farrera 293 (MEXU); 12 Jun 2009, N. Martínez-Meléndez 2692 (MEXU); 1,200 m, 5 Dec 2000, R.A. Palestina & I. Acosta 2707 (XAL). Mun. Tuxtla Gutierrez , 1,235 m, 19 Mar 1994 , M.A. Pérez-Farrera 30 ( CH, MEXU, XAL); 1,200 m, 1 Nov 1995 , M.A. Pérez-Farrera 820 (CHIP).

Figures

Figure 22. Distribution of Ceratozamia species. A C. osbornei B C. robusta C C. sabatoi D C. sancheziae.

Figure 27. Plants of Ceratozamia in habitat. A C. norstogii B C. oliversacksii C C. robusta D C. sabatoi E C. santillanii F C. tenuis G C. totonacorum H C. vovidesii I C. zaragozae.