Ceratozamia subroseophylla Mart.-Domínguez & Nic.-Mor., Phytotaxa 268(1): 35. 2016

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

30. Ceratozamia subroseophylla Mart.-Domínguez & Nic.-Mor., Phytotaxa 268(1): 35. 2016

Figs 12D , 25 , 26

Ceratozamia dominguezii Pérez-Farr. & Gut.Ortega, Taxonomy 1: 353. 2021. Type: Mexico. Veracruz: Mun. Uxpanapa, 130 m, 29 May 2021, M.A. Pérez-Farrera 4013 (holotype: HEM [n.v]; isotypes: MEXU [n.v], XAL [n.v]).

Type.

Mexico. Veracruz: Mun. Santiago Tuxtla, hill in front Sinapán, 425 m, 15 Jul 2014, L. Marti ́nez-Domínguez 158 ♀ (holotype: CIB! [acc. # 16893UV]; isotypes: MEXU! [MEXU01446538-MEXU01446542], NY!) .

Description.

Stem 30-250 (500) cm long, 18-45 cm in diameter, epigeous, erect and decumbent. Cataphylls 5-7 × 2-2.5 cm wide at the base, persistent, triangular, reddish brown, densely brown tomentose at emergence, glabrous at maturity, apex acuminate. Leaves 10-83, 72-370 cm long, ascending, yellowish brown at emergence, with whitish gray trichomes, glabrous at maturity. Petiole 50-150 cm long, terete, linear, greenish brown or dark brown at emergence, green in mature leaves; with 25-55 robust prickles, 0.40-0.85 cm long. Rachis 80-255 cm long, terete, linear, greenish brown or dark brown at emergence, green in mature leaves, with prickles. Leaflets 23-48 pairs, opposite to subopposite, insertion in one plane, lanceolate, abaxially curved longitudinally along distal half, not basally falcate, papyraceous, flat, dark green with adaxial side glabrous and abaxial side glaucous, acuminate and symmetrical at the apex, attenuate at base, with conspicuous and light-green veins; median leaflets 18-45 × 2.5-4 cm, 1.7-6.5 cm between leaflets; articulations 0.7-1.9 cm wide, brown in young leaves and green in mature leaves. Pollen strobili 15-30 cm long, 3.1-5.4 cm in diameter, solitary, cylindrical, erect, green with reddish trichomes at emergence, greenish yellow at maturity with dark brown trichomes; peduncle 9.5-19 cm long, 1.5-2.7 cm in diameter, tomentose, reddish brown; microsporophylls 1.47-2.8 × 1.01-1.4 cm, obconic with a non-recurved distal face and lobate fertile portion, infertile portion 0.49-0.65 cm long and rounded with straight horns 0.20-0.30 cm long, 0.48-0.53 cm and an acute angle between the horns. Ovulate strobili 15.5-40 cm long, 7-11.6 cm in diameter, solitary, cylindrical, erect, yellow with reddish to purple trichomes at emergence, green and megasporophylls with base pale pink and dark brown to reddish brown trichomes at maturity, mucronate apex; peduncle 9.8-17.5 cm long, 1.8-2.3 cm in diameter, erect, tomentose, light brown; megasporophylls 110-210, 9-12 orthostichies with 11-20 sporophylls orthostichy, 1.8-2.5 × 2.5-4.47 cm, with a prominent distal face, horns straight 0.34-1.0 cm long and 0.70-1.40 cm between horns with a right angle between the horns. Seeds 2.9-3.8 cm long, 0.80-1.75 cm in diameter ovate, sarcotesta whitish pink when immature, light brown at maturity.

Distribution and habitat.

Ceratozamia subroseophylla occurs from southeastern Veracruz State to Tabasco State, Mexico including the montane zone of the region Santiago Tuxtla and Uxpanapa (Fig. 24B ). It occurs on soils of volcanic origin and karstic rocks in evergreen tropical rain forest between 111 and 1,050 m.

Etymology.

The specific epithet efers to the rosaceous (i.e., pale pink) color at the base of megasporophylls.

Common names.

Mexico. Veracruz: Hymniom pekmuk (Popoluca ethnic group) (Leonti 542).

Uses.

The stem is boiled to make medicinal tea to treat kidney stones (Leonti 542).

Preliminary conservation status.

Ceratozamia subroseophylla has several populations throughout its distribution range, but anthropogenic land-use changes affect the populations of Veracruz and Tabasco. According to the IUCN Red List criteria, we recommend “Endangered” (EN) for this species.

Discussion.

Ceratozamia subroseophylla is distinguished from C. leptoceras and C. oliversacksii by its green-brownish petiole and rachis with abundant and robust prickles, and its lanceolate and papyraceous leaflets (Fig. 25 ). In addition, C. subroseophylla has affinity with C. robusta , but it is easily identified by the ovulate strobilus which has mucronate apex, rosaceous base of megasporophylls rosaceous with dark brown to reddish brown trichomes at maturity (Fig. 26 ).

Populations from Uxpanapa (Veracruz) were recently described as Ceratozamia dominguezii , however, these fall within the range of variation of C. subroseophylla as circumscribed here. Our circumscription is based on comparative morphology, both vegetative and reproductive structures, and phenology (Figs 25 , 26 ).

Specimens examined.

Mexico. Tabasco: Mun. Macuspana, 185 m, 13 Jul 2021, F. Nicolalde-Morejón et al. 3674-3680 (CIB); 185 m, 13 Jul 2021, L. Martínez-Domínguez et al. 2295-2300 (CIB), 190 m, 13 Jul 2021, L. Martínez-Domínguez et al. 2301 (CIB, MEXU). Veracruz: Mun. Catemaco, 1953, H. Bravo 26 (MEXU); 18 May 1995, M.A. Garci ́a B. et al. 813 (XAL). Mun. Hidalgotitlán, 150 m, 16 Jan 1975, Brigada Vazquez 1760 (MO, XAL); 140 m, 16 Dec 1974, J. Rees 1655 (XAL); 16 Apr 1975, M. Vázquez-Torres 1760 (MEXU). Mun. Hueyapan de Ocampo , 400 m, 14 May 2000, Leonti 542 (MEXU). Mun. Jesús Carranza, 100 m, 19 Feb 2009, D. Jimeno-Sevilla 1045 (MEXU, MO, XAL); 131 m, 27 Sep 2020, L. Martínez-Domínguez et al. 2040 (CIB); M. Vázquez-Torres 2430 (CHAPA, ENCB, MEXU, NY, SLPM, XAL). Mun. Mecayapan, 950 m, 26 Jan 1992, M. Va ́zquez-Torres et al. 4122 (CIB). Mun. Santiago Tuxtla, 500 m, 5 Jul 1983, D.W. Stevenson et al. 539 A-L (NY), 540 F-H (NY); 420 m, 28 Mar 2014, F. Nicolalde-Morejo ́n & L. Marti ́nez-Domínguez 1985-1987 (CIB); 250-500 m, 30 Sep 1983, J.F. Ortega O. et al. 358 (XAL); 420 m, 28 Mar 2014, L. Marti ́nez-Domínguez & F. Nicolalde-Morejo ́ n 78, 79 (NY), 80, 81 (XAL), 82-87 (CIB); 420 m, 7 Jun 2014, L. Marti ́nez-Domínguez & F. Nicolalde-Morejo ́n 129 (CIB); 425 m, 15 Jul 2014, L. Marti ́nez-Domínguez 136-157, 159 (CIB); 500 m, 12 May 1965, M. Sousa 2420 (MEXU); 23 Aug 1962, R.F. Andrle 64 (US); 497 m, 17 Jan 2001, T.W. Walters TW-2001-17 (MEXU, XAL). Mun. Soteapan, 21 Jun 1963, G.N. Ross 57 ( US); 1,050 m, 17 Mar 1968, M. Sousa 3645 (MEXU); 500 m, 18 May 1986, M. Va ́zquez-Torres et al. 3579 (CIB). Mun. Tatahuicapan de Jua ́rez, 849 m, C.I. Carvajal-Herna ́ndez & M. Jua ́rez F. 796 (CIB). Mun. Uxpanapa, 131 m, 27 Sep 2020, L. Marti ́nez-Domínguez et al. 2040 (CIB, MEXU); 111 m, 27 Sep 2020, L. Marti ́nez-Domínguez et al. 2041 (CIB), 2042 (CIB, MEXU), 2043-2048 (CIB); 111 m, 11 Jun 2021, L. Marti ́nez-Domínguez et al. 2273 (CIB) .

Figures

Figure 12. Seeds of Ceratozamia A C. sancheziae B C. alvarezii C C. brevifrons. D C. subroseophylla.

Figure 24. Distribution of Ceratozamia species. A C. santillanii B C. subroseophylla C C. tenuis D C. totonacorum.

Figure 25. Vegetative characters of Ceratozamia subroseophylla A adult plant (from Uxpanapa, Veracruz) B young leaf (from Santiago Tuxtla, Veracruz) C new leaves of adult plant (from Santiago Tuxtla, Veracruz) D prickles on rachis (from Uxpanapa, Veracruz).

Figure 26. Reproductive characters of Ceratozamia subroseophylla A immature pollen strobilus (from Santiago Tuxtla, Veracruz) B pollen strobilus at maturity (from Santiago Tuxtla, Veracruz) C immature ovulate strobilus (from Uxpanapa, Veracruz) D immature ovulate strobilus (from Santiago Tuxtla, Veracruz) E ovulate strobilus at maturity (Uxpanapa, Veracruz) F seeds.