A New Species of Ceratozamia (Zamiaceae) from Queretaro and Hidalgo, Mexico Andrew P. Vovides Institute) de Ecologia, A.C., Apartado Postal 63, 91000 Xalapa, Veracruz, Mexico Mario Vazquez Torres Instituto de Investigaciones Biologicas, Apartado Postal 294, 91000 Xalapa, Veracruz, Mexico Hart Schutzman Department ol Environmental Horticulture, University of Florida, Fifield Hall, Gainesville, Florida 32611-0670, U.S.A. Carlos G. Iglesias Instituto de Ecologia, A.C., Apartado Postal 63, 91000 Xalapa, Veracruz, Mexico Abstract. A new species, Ceratozamia sabatoi, pauca, usque 6 pinnate, glabra; petiolus subteres vel cy- lindricus, 25-45 cm longus, parte infima dilatatus, validis is described and illustrated. Its closest affinities are spinis armatus; rachis semiteres in dimidio inferiore paucis with C. kuesteriana from Tamaulipas, but it differs spinis armata, supra fere inermis vel inermis, in cuspidem from that species in both cone and leaf morphology. 5-11 mm longern excurrens; foliola subopposita vel al- The chromosome number of Ceratozamia sabatoi terna, 15-35 juga, remota linearis, 12-25 longa, 1.2- is 2n = 16, and the karyotype is similar to other 2.4 cm lata, basi cuneata, apicem symetrice, cuspidata, margine integerrima, 9-14 (x = 12)nervis; strobilus mas- species of the genus. culinus lineari-cylindricus 12-20 cm longus, 2-2.4 cm latus; pedunculus tomentosus 6-14 cm longus, 0.8-1 cm During botanical explorations and ecological im¬ latus; strobilus femeninus cylindricus 8 10 cm longus, 4- pact studies in the area that will be affected by the 6 cm latus; pedunculus tomentosus 4-8 cm longus, 4- 5.5 cm latus; semina 1.5 longa, 1.3 cm lata; 2n = 16. Zimapan hydro-electric dam project, a small-trunked Ceratozamia was found in pine-oak forest by Ri¬ Small palmlike plants; trunk partly subterranean, cardo Zirahuen, a biologist collaborating with the globose, readily producing offshoots, becoming cy¬ aforementioned project in the state of Queretaro. lindrical with age up to 25 cm long, 17.5 cm diam. This taxon also appears in an adjacent population protected with persistent leaf bases, dark brown in in mixed oak forest on the Hidalgo side of the border color. Leaves 2-6, pinnate, spirally arranged form¬ at a similar elevation. ing an open crown, up to 80 cm long, 52 cm wide; Following further expeditions to both localities to petiole and rachis ascending to horizontal, armed procure live, vegetative, and fertile material to es¬ with short stout prickles, 0.08-0.4 cm long (x = tablish at the Jardin Botanico Clavijero (Botanic 0.15, n = 40), petiole tomentose at base. Leaflets Garden of the Instituto de Ecologia) for observation, 12-136 (x = 69, n = 33 leaves) lanceolate, nar¬ it was concluded that this species is new to science. rowly obovate to subulate, glabrous, margin entire, It is compared to Ceratozamia kuesteriana Regel, subrevolute, apex pungent, base attenuate, dark to with which it is closely related, but differs in habit, light green on adaxial surface, lighter green on ab- leaf, female cone characteristics, and the light green axial, 9-29 cm long (x = 17, n = 47), 0.7-2.4 cm color ol emergent leaves. wide (x = 1.2, n = 47); articulation zone 0.2 0.5 cm wide (x = 0.35, n = 43), venation ± visible on Ceratozamia sabatoi Vovides, Vazquez Torres, adaxial surface, more prominent on abaxial, number Schutzman & Iglesias, sp. nov. TYPE: Mexico. of veins 9-14 (x = 12, n = 20), distance between Queretaro: 15 Apr. 1991, A. P. Vovides 1205 veins 0.75-1.4 mm. Microstrobili green when im¬ female (holotype, XAL). Figure 1. mature becoming light to dark brown at dehiscence, 6.5-23 cm long (x = 15.8), 1.9-3 cm diam. (x = Truncus globosus ad cylindricum, hypogaeus vel semi- hypogaeus, humilis ad 25 cm altus; cataphylla lanata, 2.3) ; peduncle tomentose, 1.5-11 cm long (x = triangularia 4.55 cm longa, basi 2.5-3.5 cm lata; folia 6.3) , 0.5-1 cm diam. (x = 0.7, all measurements Novon 3: 502-506. 1993. Volume 3, Number 4 Vovides et al. 503 1993 Ceratozamia sabatoi Figure 1. a-k, Ceratozamia sabatoi Vovides, Vazquez Torres, Schutzman & Iglesias. —a. Habit of plant. —b. Leaf cataphyll. —c. Base of petiole. —d. Detail of leaflets and rachis. —e. Male cone cataphyll. —f. Male cone at dehiscence. —g. Microsporophyll. —h. Female cone cataphyll. —i. Mature female cone. —j. Megasporophyll. — k. Seeds. 504 Novon lb. Median leaflet width less than 2.8 cm, persistent II II II II ii ii .. leaf bases dark brown, not appressed to trunk || || |l II II n ii || 2a. Petiole and rachis coppery in adult leaves; leaflets falcate to subfalcate, linear-lanceo¬ late; veins 6-9; megastrobili reddish-brown to dark green, longer than 13cm . Figure 2. Diploid idiogram of Ceratozamia sabatoi (2n .C. kuesteriana = 16), bar = 2 /itm. 2b. Petiole and rachis light to dark green in adult leaves; leaflets not falcate to subfal¬ cate, not linear-lanceolate; veins 9 14; me¬ gastrobili blue-green to blue-brown, less than n = 6); microsporophylls numerous, spirally ar¬ 13cm long .C. sabatoi ranged forming apparently vertical rows, cuneiform, bicornate at distal end, fertile portion covering %- Chromosomal Studies % of abaxial surface excluding horns and stalk, 0.9 The chromosome number and karyotype were 1.4 cm long (x = 1.1), 0.4-0.7 cm wide (x = 0.6, n = 6); microsporangia numerous in sori of 2 to 3, determined from three established specimens held dehiscence by longitudinal slit, 0.9-1.3 mm diam. at the Jardin Botanico Fco. J. Clavijero under the (x = 1.1, n = 12). Megastrobili cylindrical to barrel¬ accession numbers: 91-028, 91-040, 91-041 and shaped slightly tapering toward apex, light blue- vouchers deposited at (XAL). The root tip mitosis green when immature, turning green to light blue- technique was used described by Vovides (1983) brown at maturity, 6.0-12 cm long (x = 9.5), 3.4- and chromosome classification based on centromere 5.6 cm diam. (x = 4.8); peduncle tomentose, 2.0- position that of Levan et al. (1964) modified by 10 cm long (x = 5.4), 0.6-1.3 cm diam. (x = 0.8); Schlarbaum and Tsuchiya (1984). The diploid idi¬ megasporophylls numerous, spirally arranged form¬ ogram (Fig. 2) was constructed by taking the av¬ ing apparently vertical rows, cuneiform-peltate, dis¬ erage arm lengths of the best three metaphase cells tal ends hexagonal, thickened, bicornate, with red¬ examined (Fig. 3). Arm lengths, total chromosome dish tomentum near horns, 1.7-2.6 cm long (x = length, chromosome index (short arm divided by 2.1), 1.1-2.8 cm wide (x = 1.7, all measurements long arm), and symmetry index (length of longest pair divided by length of shortest pair) were com¬ n = 7). Seeds ovate variably angulate, sarcotesta creamy-white when immature becoming blue-green puted using the average arm lengths from the three to light brown when mature, sclerotesta light beige, metaphase cells (Table 1). The karyotype shows 11 smooth, 8-10 visible ridges radiating from micro- median region (m) chromosomes, 1 submedian (sm) pyle, 1.3-1.9 cm long (x = 1.5), 1.1-1.4 cm diam. chromosome, 1 subterminal region (st) chromosome, (x = 1.3, n = 6). Chromosome number 2n = 16. and 2 terminal point (T) chromosomes. Satellite number and position vary with cells observed and We assign the specific epithet in honor of the a maximum of 5 satellites were recorded, but not late Sergio Sabato, distinguished professor at the considered in the calculations. University of Naples, Italy, for his outstanding and prolific fieldwork and experimental biology on neo¬ Habitat tropical Zamiaceae. The vegetation at the Queretaro locality is mainly Paratypes. MEXICO. Queretaro: 15 Apr. 1991, A. pine-oak forest dominated by Pirius teocote Schle- P. Vovides, K. Norstog & P. Fawcett 1200, 1201, 1202, chtendal & Chamberlain, Quercus crassifolia Hum¬ 1204, all 9, 1190, 1191, 1192, 1194 all 6 (XAL). Hi¬ dalgo: 12 Feb. 1991, A. P. Vovides, M. V azquez T. <£ boldt & Bonpland, Q. macrophylla Martius & Gal- C. lglesias 1156, 1157, 1158, 1159, 1160 all 9, 1161, eotti, Q. germana Chamberlain & Schlechtendal, 1162, 1163, 1164, 1165, 1166, 1168, 1169, 1170, and Q. xalapensis Humboldt & Bonpland on reddish 1171, 1172, 1173, 1175, 1176, 1177, 1178, 1179, clay (laterite) soils. The vegetation of the Hidalgo 1180, 1181, 1182, 1183, 1184, 1185, 1186 all <5 (XAL). locality, a mixed oak forest, appears much richer The following key permits the separation of Cer¬ due to higher rainfall. The dominant tree species atozamia sabatoi from C. kuesteriana and C. rni- are; Arbutus xalapensis Humboldt, Bonpland & crostrobila. Kunth, Buddleja cordata Humboldt, Bonpland & Kunth, Carya ovata (Miller) K. Koch, Litsea glau- cescens Humboldt, Bonpland & Kunth, Meliosma Diagnostic Key alba (Schlechtendal) Walpers, Persea sp., Primus serotina Ehrenberg, Quercus germana Chamber- la. Median leaflet width greater than 2.8 cm, per¬ sistent leaf bases light brown, tightly appressed lain & Schlechtendal, Q. sartorii Liebmann, Q. to trunk .C. microstrobila polymorpha Schlechtendal & Chamberlain, and Q. Volume 3, Number 4 Vovides et al. 505 1993 Ceratozamia sabatoi r • •(cid:9632)‘v ' TABLE 1. Karyotypic data at metaphase in root-tip mitoses of Ceratozamia sabatoi (mean of three rneta- phase cells), m = median, msm = median-submedian, sm = submedian, st = subterminal, T = terminal. Arm length Chro¬ (Mm) mo¬ Centro¬ some Long Short Index r mere pair (L) (S) Total (S/L) value position i 6.67 6.62 13.29 0.99 1.01 III 6.92 6.10 13.02 0.88 1.13 m 2 6.78 6.15 12.93 0.91 1.10 m 6.67 6.15 12.82 0.92 1.08 m 3 6.61 5.96 12.57 0.90 1.11 m 5.67 5.30 10.97 0.93 1.07 m 4 5.53 5.41 10.94 0.98 1.02 m 5.56 5.15 10.71 0.93 1.08 m 5 5.43 4.78 10.21 0.88 1.14 m 5.54 4.18 9.72 0.75 1.33 msm Figure 3. Chromosomes of Ceratozamia sabatoi at mi¬ 6 5.14 4.24 9.38 0.82 1.21 in totic metaphase, bar = 2 4.92 4.15 9.07 0.84 1.19 m 7 5.14 1.96 7.10 0.38 2.62 sm 4.99 1.66 6.65 0.33 3.01 st xalapensis Humboldt & Bonpland. The soil is a 8 6.64 0.00 6.64 0.00 0.00 T dark humus-rich clay on limestone. 6.71 0.00 6.71 0.00 0.00 T Symmetry index = 0.51. Average chromosome in¬ Discussion dex = 0.72. Total chromosome length = 162.73. Precise information on localities has been pur¬ posely omitted in order to discourage indiscriminate commercial collecting, which could lead to the ex¬ probably due to differential contraction at meta¬ tinction of this endangered species. phase. The number and position of satellites varies Even though some vegetative characteristics, es¬ between the species (Vovides, 1983). Two satellites pecially the leaves, show on occasion morphological have been reported by Vovides (1985) in C. kues¬ similarities between Ceratozamia kuesteriana and teriana and C. sabatoi has five. C. sabatoi, the consistent differences in cones, emer¬ Acknowledgments. We thank Ricardo Zirahuen, gent leaves, seeds, and habitat lead us to consider Christina Jocabeth P., and colleagues of the Com- the two taxa as separate species. Geographically ision Federal de Electricidad (CFE) of the Zimapan speaking, several species are found in the areas Hydroelectric Project for permission, assistance, and between C. kuesteriana and C. sabatoi, including use of helicopter in the field during the early ex¬ C. zaragozae Medellin, C. hidae Landry & Wilson, ploration of the habitat. We thank CONACyT for C. microstrobila Vovides & Rees, and C. mexicana partial funding of this investigation through project var. robusta (Miquel) Dyer, species whose popula¬ No. P020CCOR-904133. We also thank Edmundo tions are well delimited. Ceratozamia sabatoi occurs Saavedra for the excellent botanical illustration of amongst low shrubs in oak forests and mixed oak- this species. pine forests on volcanic soils in relatively dry hab¬ itats; C. kuesteriana, however, is found in much Literature Cited wetter cloud-forests on karst topography. The chromosome count and karyotype are con¬ Levan, A., K. Fredga & A. A. Sandberg. 1964. No¬ menclature for centromeric position on chromo¬ sistent with that reported for the genus (2n =16) somes. Hereditas 52: 201-220. by Marchant (1968), Vovides (1983; 1985) and Marchant, C. J. 1968. Chromosome patterns and nu¬ Moretti (1990). The karyotype is nearly typical for clear phenomena in the cycad families Stangeriaceae the genus Ceratozamia (12m + 2sm + 2T), which and Zamiaceae. Chroinosoma (Berl.) 24: 100-134. appears to be stable. The msm and st chromosomes Moretti, A. 1990. Karyotypic data on north and central American Zamiaceae (Cycadales) and their phylo¬ fall into these categories only by 0.03 and 0.01 gm genetic implications. Amer. J. Bot. 77: 1016-1029. respectively and, considering the considerable length Schlarbaum, S. E. & T. Tsuchiya. 1984. The chro¬ of cycad chromosomes, these discrepancies are mosomes of Cunninghamia konishi, C. lanceolata. 506 Novon and Taiwania cryptomerioides (Taxodiaceae). PI. -. 1985. Systematic studies on Mexican Zami¬ Syst. Evol. 145: 169-181. aceae II. Additional notes on Ceratozamia kuester- Vovides, A. P. 1983. Systematic studies on the Mexican iana from Tamaulipas, Mexico. Brittonia 37: 226- Zamiaceae I. Chromosome numbers and karyotypes. 231. Amer. J. Bot. 70: 1002-1006. Volume 3, Number 1, pp. 1-92 of NOVON was published on 19 March 1993. Volume 3, Number 2, pp. 93-219 of NOVON was published on 1 July 1993. Volume 3, Number 3, pp. 221-310 of NOVON was published on 30 September 1993. Volume 3, Number 4, pp. 311-506 of NOVON was published on 29 December 1993.