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Taxonomic Revision of Zamia in Mega-Mexico

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Taxonomic Revision of Zamia in Mega-Mexico Taxonomic revision of Zamia in Mega-Mexico 1 1 FERNANDO NICOLALDE-MOREJÓN ,ANDREW P. V OVIDES , 2 AND DENNIS W. STEVENSON 1 Departamento de Biología Evolutiva, Instituto de Ecología, A. C. km 2.5 Antigua Carretera a Coatepec No. 351, Xalapa 91070 Veracruz, Mexico 2 The New York Botanical Garden, Bronx, NY 10458-5120, USA; e-mail: [email protected] Abstract. The genus Zamia is revised for Mega-Mexico, with 22 species recognized and described. The study presents a taxonomic clarification for the genus in Mesoa- merica, a contribution that provides the foundation for a future monograph for Zamia in the Neotropics. The largest proportion of species richness and endemism for the genus is concentrated in southeastern Mexico, among the states of Chiapas, Oaxaca, Tabasco, and Veracruz, an area that is considered highly diverse in floristic terms. Distribution maps and a key to species are also provided, as well as complete des- criptions of the specimens examined, including information on nomenclatural types, habitats, synonymies, and etymologies. A lectotype is designated for Zamia loddigesii, and neotypes for Z. galeottii, Z. leiboldii var. angustifolia, and Z. variegata. Zamia spartea is illustrated for the first time, and chromosome numbers for Z. herrerae are reported and illustrated. Finally, scanning electron micrographs of leaflet trichome character states are presented, along with a discussion of their systematic implications within the group. Key Words: Endemism, Mexico, gymnosperms, cycads, floristic richness, Zamia. Resumen. El género Zamia es revisado para Mega-México, con 22 especies descritas y una species dubia. El estudio está orientado al esclarecimiento taxonómico del género en Mega-México, una contribución que siente los fundamentos para una futura monografía para Zamia en el Neotrópico. La mayor riqueza y endemismo para el género se concentra en el sureste de México, entre los estados de Chiapas, Oaxaca, Tabasco y Veracruz, área de alta biodiversidad florística. Mapas de distribución y una clave para las especies son presentadas, como también descripciones completas, tipos, hábitat, sinónimos, etimología y especimenes examinadas. Se designa un lectotipo para Zamia loddigesii, y neotipos para Z. galeottii, Z. leiboldii var. angustifolia y Z. variegata. Zamia spartea es ilustrada por primera vez y números cromosómicos para Z. herrerae son presentados. Finalmente, se presentan fotografías de tricomas al mi- croscopio electrónico y una discusión de sus implicaciones en la sistemática del grupo. According to Stevenson (1992), Zamiaceae Central America that is floristically similar to comprises eight genera distributed in tropical the southern part of Mexico (Mega-Mexico) and subtropical Africa, Australia, Greater while Microcycas A. DC. (1 sp.) and Chigua Antilles, North, Central and South America. D.W. Stev. (2 spp.) are endemic to Cuba and Five genera with 94 species are known from Colombia, respectively. The type genus for the Neotropics (Hill et al., 2007). The genera the family, Zamia L. (59 spp.), is distributed Ceratozamia Brongn. (21 spp.) and Dioon throughout tropical and sub-tropical America Lindl. (13 spp.) are both endemic to Mexico as well as the Caribbean, with exception to and a neighboring biogeographic region of the Lesser Antilles. Brittonia, 61(4), 2009, pp. 301–335 ISSUED: 1 December 2009 © 2009, by The New York Botanical Garden Press, Bronx, NY 10458-5126 U.S.A. 302 BRITTONIA [VOL 61 Zamia is the widest distributed genus of the Mexico’ hereafter, given that the slight dif- order Cycadales in the Neotropics. Its northern ferences in the boundaries of ‘Mega-Mexico 1’ range starts in Georgia and Florida (U.S.A.), with respect to ‘Mega-Mexico 2’ do not affect reaching Bolivia and the Mato Grosso of the biogeographic aspects of our description Brazil in South America (Balduzzi et al., of the species. This study is intended to 1982;Sabato,1990; Norstog & Nicholls, provide a taxonomic clarification of the genus 1997; Stevenson, 2001a, b). A remarkable in Mesoamerica and to provide the basis for a morphological and cytological variation has future monograph of Zamia. been documented (Vovides, 1983; Moretti & Sabato, 1984; Moretti, 1990a, b; Stevenson et – al., 1995 1996a; Vovides & Olivares, 1996; Materials and methods Norstog & Nicholls, 1997), and also high levels of genetic variation (González-Astorga The present taxonomic revision is based on et al., 2006). As a consequence of this more than 450 specimens from the following complexity, the taxonomy of Zamia is contro- herbaria: B, BM, CIB, CHIP, CICY, ECOSUR, versial; although the genus comprises 75 ENCB, F, FCME, FLAS, FTG, HEM, IBUG, species, their circumscription and limits have IEB, K, LE, MEXU, MO, NY, SERO, U, yet to be determined (Hill et al., 2007). UADY, UAMIZ, US, W, WIS, XAL, XALU, The most recent exhaustive taxonomic and ZEA. Unfortunately, we were not able to treatment for Zamia was published by Schus- obtain vouchers on loan from NAP; therefore, ter (1932) and later work by Sabato (1990) material from this herbarium is not cited. and Stevenson (1987; 1991a, b; 1993; 2001a, Chromosome counts for Zamia herrerae b; 2004) underlined nomenclatural and taxo- Calderón & Standley were performed on five nomic anomalies in Zamia, principally owing individuals held at the Jardín Botánico Fco. to insufficient fieldwork and a scarcity of Javier Clavijero, Instituto de Ecología, A.C. good-quality botanical collections. The taxo- (JBC). These plants has been previously nomic history for the genus in Mexico began collected at the Acacoyagua and Tonalá with the publication of Zamia fufuracea L. f. regions, located in the state of Chiapas and in 1789 from the central-south coastal region represent the species range in southern Mex- of Veracruz, which also represents the first ico. Plants from its full biogeographic range, cycad species described from the American which would include El Salvador and Guate- continent. During the 19th century, six spe- mala, were not available for this study. A cies were subsequently described for Mexico; modified root-tip squash method was used for Z. fischeri Miq., Z. katzeriana (Regel) Rettig, examining somatic metaphase cells described Z. lawsoniana Dyer, Z. loddigesii Miq., Z. by Vovides (1983) with a 12 to 15 hour ice- spartea A. DC. and Z. verschaffeltii Miq., water (0°C) follow-up soak after the 0.2% whereas the remaining known species were colchicine pretreatment at ambient tempera- documented and characterized during the ture (Schutzman et al., 1988). Counts were 20th century, with a marked tendency for made from the best 10–15 metaphase cells taxonomic activity during the last thirty years and karyotype noted according to the classi- (Vovides et al., 1983; Schutzman et al., 1988; fication of Schlarbaum and Tsuchiya (1984). Stevenson et al., 1995–1996a, b; Schutzman Photomicrographs were produced using a & Vovides, 1998; Vovides, 1999). Zeiss Fomi III photomicroscope fitted with The present taxonomic revision includes planapochromatic objectives and Kodak Plus- endemic species of the cycad genus Zamia that X pan ASA 125 film. occur in ‘Mega-Mexico 2’, a term coined by Scanning electron micrographs (SEM) Rzedowski (1991) that associates the Central were taken on young leaflet material from American territories of Guatemala, Belize, living plants cultivated in the JBC. Samples Honduras and northern Nicaragua to the were placed on sample stubs with double Mexican states of Nayarit on the Pacificto sided adhesive tape and then introduced into southern Tamaulipas on the Gulf of Mexico using a dessicator for 24 hours. All samples were biotic (mainly floristic) criteria. Rzedowski’s sputter coated with gold-palladium at 1.5 kv concept will be referred to as simply ‘Mega- at 5 mA for 8 minutes with a Jeol Fine Coat 2009] NICOLALDE ET AL.: ZAMIA IN MEGA-MEXICO 303 JFC 1100 sputter coater. Observations were notable, with branches reaching up to 80 cm made with a Jeol JSM-5600LV SEM. long in adult plants. In all cases, the types have been examined TRICHOMES.—Trichomes of cycads leaves by one or more of the authors. are bi-celled, consisting of a small basal cell and a longer free apical portion (Stevenson, Results 1981). All the trichomes analyzed here show the same bifurcate pattern, with one arm HABITAT proportionally longer than the other (Fig. 1A). Of the 22 species included in this revision, Each trichome presents a rounded basal cell 18 occur in specific habitats, accounting for and a more extensive cylindrical bifurcate free the restricted distribution of the majority of portion, which in most cases observed had the taxa. The species with the widest distri- evidence of collapse and twisting (Fig. 1B–F). bution are associated with two or more The pubescence was significantly denser in vegetation types, namely (a) Zamia paucijuga emerging than in older adult leaves with the Wieland, found in pine-oak, oak and tropical latter becoming completely glabrous. An dry forests; (b) Z. polymorpha D.W. Stev., A. exception to this condition is found in Zamia Moretti & Vázq. Torres, located in evergreen furfuracea, which maintains a great part of its tropical rainforest, sub-deciduous tropical original indument on the abaxial surface of forest and their secondary succession stages; each leaflet. (c) Z. herrerae, which occurs in evergreen According to Stevenson (1981), four types tropical rainforest, sub-deciduous tropical of trichomes occur in Zamia; of these, the forest and tropical dry forest and their transparent ramified and the colored ramified secondary succession stages and finally, to a are the types found in the present work. In lesser extent; and (d) Z. loddigesii, present in contrast to what was found in the aforemen- evergreen tropical rainforest, but more com- tioned study by Stevenson, no trend or monly in sub-deciduous tropical forest and its correlation was found between aerial stems secondary succession stages. (i.e., Z. soconuscensis or Z. inermis) and bifurcate trichomes with equal length branches. Trichomes with unequal sized MORPHOLOGY prevail among taxa with both aerial and HABIT.—All adult cycad stems are pachycau- subterranean stems.
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