Ceratozamia fuscoviridis W.Bull., Retail List. 154: 4. 1879

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

10. Ceratozamia fuscoviridis W.Bull., Retail List. 154: 4. 1879

Figs 3A , 9C , 15J

Ceratozamia mexicana Brongn. f. fuscoviridis (W.Bull.) J.Schust., Pflanzenr. (Engler) Heft 99, 4 fam 1: 132. 1932, as C. mexicana var. longifolia f. fuscoviridis . Type: Based on Ceratozamia fuscoviridis W.Bull.

Type.

Cultivated at Glasnevin, Ireland "Hort. Bot. Glasnevin", 21 Mar 1878 (accessioned 1881), D. Moore s.n. (neotype, designated by Calonje and Sennikov 2017, pg. 161: K! [K000501714, K000501712, K000501713]). Mexico. Hidalgo: Mun. Molango, km 3 carretera Molango-Xochicoatlán, 1,860 m, 31 Mar 2015, L. Marti ́nez-Domínguez et al. 493 ♀ (epitype, designated by Martínez-Domínguez et al. 2018a: 105: CIB! [acc. # 17465UV]; isoepitype: MEXU! [acc. # 1520508, 1520282, 1520519]) .

Description.

Stem 20-90 cm long, 25-40 cm in diameter, epigeous, erect and decumbent. Cataphylls 2-4.5 × 2-3.5 cm wide at the base, persistent, narrowly triangular, reddish brown, densely brownish tomentose at emergence, tomentose at maturity, apex acuminate. Leaves 10-70, 92-215 cm long, ascending, light green and dark brown at emergence, with whitish gray trichomes, glabrous at maturity. Petiole 40-95 cm long, terete, linear, dark green in mature leaves; with 14-60 (66) thin prickles, 0.09-0.33 cm long. Rachis 65-150 cm long, terete, linear, green in mature leaves, with prickles. Leaflets 28-67 pairs, opposite to subopposite, insertion in one plane, lanceolate, abaxially curved longitudinally, basally falcate, papyraceous, flat, green, with adaxial and abaxial sides glabrous, distal end with entire margins, acuminate and symmetrical at the apex, attenuate at base, with conspicuous and green-light brown veins; median leaflets 16.6-42 × 1.3-2.1 cm, 0.6-2 cm between leaflets; articulations 0.6-1.3 cm wide, green and brown. Pollen strobili 22-30 cm long, 5-8 cm in diameter, solitary, cylindrical, erect, brownish yellow at emergence, greenish brown with reddish brown trichomes at maturity; peduncle 5-14.5 cm long, 1.6-2.3 cm in diameter, tomentose, reddish brown to brown; microsporophylls 1.6-2.4 × 1.2-1.6 cm, obconic with a non-recurved distal face and a deeply lobate fertile portion, infertile portion 0.45-0.56 cm long and orbicular with straight horns 0.16-0.29 cm long, 0.53-0.80 cm and a right angle between the horns. Ovulate strobili 24-35 cm long, 8.5-15 cm in diameter, solitary, cylindrical, erect, green with brown trichomes at emergence, brown-green with dark brown trichomes at maturity, acuminate apex; peduncle 4-15.5 cm long, 1.8-2.5 cm in diameter, erect or pendulous, tomentose, brown to reddish brown; megasporophylls 49-195, 7-15 orthostichies with 7-15 sporophylls per orthostichy, 2.3-3.5 × 3.0-4.0 cm, with a truncate distal face, horns straight and 0.70-0.99 cm long, 1.7-2.1 cm between horns and an obtuse angle between the horns. Seeds 2-2.6 cm long, 1.5-2 cm in diameter, ovate, sarcotesta whitish yellow to yellow when immature, light brown at maturity.

Distribution and habitat.

Ceratozamia fuscoviridis is endemic to south to central Carso Huasteco in Mexico from Hidalgo to northwest of Veracruz States (Fig. 17B ) in cloud forests between 1,300-1,900 m elevation.

Etymology.

The epithet refers to the dark-brown color of the leaf at emergence.

Common names.

Mexico. Veracruz: teocintle, teocintli and tepecintli ( Bonta et al. 2019).

Uses.

This species has decorative uses associated with religious ceremonies and national holidays. The leaves are commonly used to make arches in the entrances of some Roman Catholic churches or decorate the walls during national holidays.

Conservation status.

( IUCN 2021). Ceratozamia fuscoviridis is listed as "Critically Endangered" under criteria B1ab(i,iii,iv,v). This species requires a reassessment of its conservation status because new populations have recently been recorded, including its current circumscription that includes populations from Veracruz State. Based on the number of populations, modelled potential geographical distribution from ecological niche and its occurrence in a Natural Protected Areas (Los Mármoles National Park), C. fuscoviridis could be listed as "Endangered (EN)" ( Martínez-Domínguez et al. 2021).

Discussion.

Ceratozamia fuscoviridis is polymorphic within populations because individual plants may have either a light green or dark-brown leaf color at emergence. Individuals with dark-brown leaf at emergence also have a brownish abaxial side. This species is most geographically proximate to C. chamberlainii , but it can be distinguished by lanceolate and papyraceous leaflets, a petiole armed with long and thin prickles; and ovulate strobili that are brownish green with dark trichomes at maturity.

Osborne et al. (2006) intended to validate Ceratozamia fuscoviridis D.Moore, which was invalidly published in 1878. However, in 2017, Calonje & Sennikov published the correction to this name using a brief description by William Bull in horticultural catalogues. This work is the valid publication for the specie and the name " C. fuscoviridis D.Moore" is an isonym with no nomenclatural status.

Specimens examined.

Mexico. Hidalgo: Mun. Eloxochitlán, 18 Mar 1995, I. Luna-Vega 54716 (FCME). Mun. Metztitlán, 30 Dec 1992, J.L. López-García 449 (ENCB, IBUG, MEXU). Mun. Molango de Escamilla , 1,380 m, 29 May 1999, A.P. Vovides 1298 (XAL); 1,400 m, 29 May 1999, A.P. Vovides 1301 (XAL); 1,500 m, 24 Jul 2008, A. Vite-Reyes et al. 6 (XAL); 1,860 m, 31 Mar 2015, F. Nicolalde-Morejón et al. 2209 - 2211 (CIB); 1,860 m, 31 Mar 2015, L. Martínez-Domínguez et al. 485 -492, 494- 514 (CIB); T.W. Walters 2001-03-A (XAL). Mun. Tenango de Doria , 1,700 m, 12 Mar 1993, I. Luna-Vega 914 (FCME). Mun. Tlanchinol, 1,450 m, 25 Aug 1992, I. Luna-Vega s/n (FCME); 1,420 m, 7 Oct 1992, I. Luna-Vega 625 (FCME, XAL); 25 Aug 1992, I. Luna-Vega 789 (XAL); 1,312 m, 31 Mar 2015, F. Nicolalde-Morejón et al. 2204-2208 (CIB); 1,312 m, 31 Mar 2015, L. Martínez-Domínguez et al. 455-484 (CIB). Mun. Zacualtipan de Angeles , 23 Jan, 1983, J. Rees 389 (CHAPA, FCME, MEXU) ; 1,360 m, 4 Dec 1974, J. Rees 1611 (CHIP, XAL); 4 Dec 1974, J. Rees 6339 (IEB). Veracruz: Mun. Huayacocotla, 1,913 m, 23 Feb 2005, D. Saavedra Millán 64 (FCME); 1,850 m, 13 Mar 1980, J. Palma G. 63 (XAL); 1,844 m, 1 Apr 2015, F. Nicolalde-Morejón et al. 2212 - 2214 (CIB); 1,700 m, 24 Mar 1981, L. Ballesteros & F. Ballesteros 460 (XAL); 1,550 m, 23 Apr 1981, L.G. Juárez G. 47 (XAL); 1,844 m, 1 Apr 2015, L. Martínez-Domínguez et al. 515 - 544 (CIB); 1,900 m, 11 Feb 1972, R. Hernández M. 1507 (MEXU, XAL); 26 Feb 1975, V. Sosa 59 (XAL).

Figures

Figure 3. Comparison of qualitative characters of microsporophylls A Ceratozamia fuscoviridis B C. brevifrons C C. chamberlainii D C. kuesteriana.

Figure 9. Color of leaves at emergence A Ceratozamia sancheziae B C. kuesteriana C C. fuscoviridis D C. miqueliana.

Figure 15. Plants of Ceratozamia in habitat A C. alvarezii B C. aurantiaca C C. becerrae D C. brevifrons E C. chamberlainii F C. chimalapensis G C. decumbens H C. delucana I C. euryphyllidia J C. fuscoviridis K C. hondurensis L C. huastecorum.

Figure 17. Distribution of Ceratozamia species A C. euryphyllidia B C. fuscoviridis C C. hildae D C. hondurensis.