Taxonomic revision of Zamia in Mega-Mexico€¦ · Tabasco, and Veracruz, ... spartea is illustrated for the ﬁrst time, and chromosome numbers for Z. herrerae are reported and illustrated

Taxonomic revision of Zamia in Mega-Mexico

FERNANDO NICOLALDE-MOREJÓN1, ANDREW P. VOVIDES

1,AND DENNIS W. STEVENSON2

1Departamento de Biología Evolutiva, Instituto de Ecología, A. C. km 2.5 Antigua Carretera aCoatepec No. 351, Xalapa 91070 Veracruz, Mexico

2The 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 recognizedand described. The study presents a taxonomic clarification for the genus in Mesoa-merica, a contribution that provides the foundation for a future monograph for Zamiain the Neotropics. The largest proportion of species richness and endemism for thegenus 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. Zamiaspartea is illustrated for the first time, and chromosome numbers for Z. herrerae arereported and illustrated. Finally, scanning electron micrographs of leaflet trichomecharacter states are presented, along with a discussion of their systematic implicationswithin the group.

Key Words: Endemism, Mexico, gymnosperms, cycads, floristic richness, Zamia.

Resumen. El género Zamia es revisado para Mega-México, con 22 especies descritasy una species dubia. El estudio está orientado al esclarecimiento taxonómico delgénero en Mega-México, una contribución que siente los fundamentos para una futuramonografía para Zamia en el Neotrópico. La mayor riqueza y endemismo para elgénero 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 unaclave para las especies son presentadas, como también descripciones completas, tipos,hábitat, sinónimos, etimología y especimenes examinadas. Se designa un lectotipopara Zamia loddigesii, y neotipos para Z. galeottii, Z. leiboldii var. angustifolia y Z.variegata. Zamia spartea es ilustrada por primera vez y números cromosómicos paraZ. herrerae son presentados. Finalmente, se presentan fotografías de tricomas al mi-croscopio electrónico y una discusión de sus implicaciones en la sistemática delgrupo.

According to Stevenson (1992), Zamiaceaecomprises eight genera distributed in tropicaland subtropical Africa, Australia, GreaterAntilles, North, Central and South America.Five genera with 94 species are known fromthe Neotropics (Hill et al., 2007). The generaCeratozamia Brongn. (21 spp.) and DioonLindl. (13 spp.) are both endemic to Mexicoand a neighboring biogeographic region of

Central America that is floristically similar tothe southern part of Mexico (Mega-Mexico)while Microcycas A. DC. (1 sp.) and ChiguaD.W. Stev. (2 spp.) are endemic to Cuba andColombia, respectively. The type genus forthe family, Zamia L. (59 spp.), is distributedthroughout tropical and sub-tropical Americaas well as the Caribbean, with exception tothe 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.

Zamia is the widest distributed genus of theorder Cycadales in the Neotropics. Its northernrange starts in Georgia and Florida (U.S.A.),reaching Bolivia and the Mato Grosso ofBrazil in South America (Balduzzi et al.,1982; Sabato, 1990; Norstog & Nicholls,1997; Stevenson, 2001a, b). A remarkablemorphological and cytological variation hasbeen documented (Vovides, 1983; Moretti &Sabato, 1984; Moretti, 1990a, b; Stevenson etal., 1995–1996a; Vovides & Olivares, 1996;Norstog & Nicholls, 1997), and also highlevels of genetic variation (González-Astorgaet al., 2006). As a consequence of thiscomplexity, the taxonomy of Zamia is contro-versial; although the genus comprises 75species, their circumscription and limits haveyet to be determined (Hill et al., 2007).The most recent exhaustive taxonomic

treatment for Zamia was published by Schus-ter (1932) and later work by Sabato (1990)and Stevenson (1987; 1991a, b; 1993; 2001a,b; 2004) underlined nomenclatural and taxo-nomic anomalies in Zamia, principally owingto insufficient fieldwork and a scarcity ofgood-quality botanical collections. The taxo-nomic history for the genus in Mexico beganwith the publication of Zamia fufuracea L. f.in 1789 from the central-south coastal regionof Veracruz, which also represents the firstcycad species described from the Americancontinent. During the 19th century, six spe-cies were subsequently described for Mexico;Z. fischeri Miq., Z. katzeriana (Regel) Rettig,Z. lawsoniana Dyer, Z. loddigesii Miq., Z.spartea A. DC. and Z. verschaffeltii Miq.,whereas the remaining known species weredocumented and characterized during the20th century, with a marked tendency fortaxonomic activity during the last thirty years(Vovides et al., 1983; Schutzman et al., 1988;Stevenson et al., 1995–1996a, b; Schutzman& Vovides, 1998; Vovides, 1999).The present taxonomic revision includes

endemic species of the cycad genus Zamia thatoccur in ‘Mega-Mexico 2’, a term coined byRzedowski (1991) that associates the CentralAmerican territories of Guatemala, Belize,Honduras and northern Nicaragua to theMexican states of Nayarit on the Pacific tosouthern Tamaulipas on the Gulf ofMexico usingbiotic (mainly floristic) criteria. Rzedowski’sconcept will be referred to as simply ‘Mega-

Mexico’ hereafter, given that the slight dif-ferences in the boundaries of ‘Mega-Mexico 1’with respect to ‘Mega-Mexico 2’ do not affectthe biogeographic aspects of our descriptionof the species. This study is intended toprovide a taxonomic clarification of the genusin Mesoamerica and to provide the basis for afuture monograph of Zamia.

Materials and methods

The present taxonomic revision is based onmore than 450 specimens from the followingherbaria: B, BM, CIB, CHIP, CICY, ECOSUR,ENCB, F, FCME, FLAS, FTG, HEM, IBUG,IEB, K, LE, MEXU, MO, NY, SERO, U,UADY, UAMIZ, US, W, WIS, XAL, XALU,and ZEA. Unfortunately, we were not able toobtain vouchers on loan from NAP; therefore,material from this herbarium is not cited.Chromosome counts for Zamia herrerae

Calderón & Standley were performed on fiveindividuals held at the Jardín Botánico Fco.Javier Clavijero, Instituto de Ecología, A.C.(JBC). These plants has been previouslycollected at the Acacoyagua and Tonaláregions, located in the state of Chiapas andrepresent the species range in southern Mex-ico. Plants from its full biogeographic range,which would include El Salvador and Guate-mala, were not available for this study. Amodified root-tip squash method was used forexamining somatic metaphase cells describedby Vovides (1983) with a 12 to 15 hour ice-water (0°C) follow-up soak after the 0.2%colchicine pretreatment at ambient tempera-ture (Schutzman et al., 1988). Counts weremade from the best 10–15 metaphase cellsand karyotype noted according to the classi-fication of Schlarbaum and Tsuchiya (1984).Photomicrographs were produced using aZeiss Fomi III photomicroscope fitted withplanapochromatic objectives and Kodak Plus-X pan ASA 125 film.Scanning electron micrographs (SEM)

were taken on young leaflet material fromliving plants cultivated in the JBC. Sampleswere placed on sample stubs with doublesided adhesive tape and then introduced intoa dessicator for 24 hours. All samples weresputter coated with gold-palladium at 1.5 kvat 5 mA for 8 minutes with a Jeol Fine Coat

302 BRITTONIA [VOL 61

JFC 1100 sputter coater. Observations weremade with a Jeol JSM-5600LV SEM.In all cases, the types have been examined

by one or more of the authors.

Results

HABITAT

Of the 22 species included in this revision,18 occur in specific habitats, accounting forthe restricted distribution of the majority ofthe taxa. The species with the widest distri-bution are associated with two or morevegetation types, namely (a) Zamia paucijugaWieland, found in pine-oak, oak and tropicaldry forests; (b) Z. polymorpha D.W. Stev., A.Moretti & Vázq. Torres, located in evergreentropical rainforest, sub-deciduous tropicalforest and their secondary succession stages;(c) Z. herrerae, which occurs in evergreentropical rainforest, sub-deciduous tropicalforest and tropical dry forest and theirsecondary succession stages and finally, to alesser extent; and (d) Z. loddigesii, present inevergreen tropical rainforest, but more com-monly in sub-deciduous tropical forest and itssecondary succession stages.

MORPHOLOGY

HABIT.—All adult cycad stems are pachycau-lous and may be columnar and arborescent orsubterranean and tuber-like. The genera Dioon,Microcycas, Ceratozamia and LepidozamiaRegel are usually columnar arborescent in habit,while the subterranean forms are characteristicof Bowenia Hook. ex Hook. f., Chigua, andStangeria T. Moore. Cycas L., EncephalartosLehm.,MacrozamiaMiq. and Zamia have bothstem morphologies, either subterranean tuber-like or columnar arborescent (Stevenson, 1980).In this context, the species of Zamia inMesoamerica represent both growth forms, withthe subterranean tuber-like habitat predominant.Only four species, Z. inermis, Z. onanreyesii, Z.soconuscensis and Z. tuerckheimii, have arbo-rescent stems reaching up to 100 cm or more inheight. Some species branch dichotomouslywith age (namely, Z. fischeriMiq., Z. furfuracea,Z. inermis Vovides, Rees & Vázq. Torres, Z.soconuscensis Schutzman, Vovides & Dehgan,Z. loddigesii and Z. herrerae), with the coastaldune species Z. furfuracea being the most

notable, with branches reaching up to 80 cmlong in adult plants.TRICHOMES.—Trichomes of cycads leaves

are bi-celled, consisting of a small basal celland a longer free apical portion (Stevenson,1981). All the trichomes analyzed here showthe same bifurcate pattern, with one armproportionally longer than the other (Fig. 1A).Each trichome presents a rounded basal celland a more extensive cylindrical bifurcate freeportion, which in most cases observed hadevidence of collapse and twisting (Fig. 1B–F).The pubescence was significantly denser inemerging than in older adult leaves with thelatter becoming completely glabrous. Anexception to this condition is found in Zamiafurfuracea, which maintains a great part of itsoriginal indument on the abaxial surface ofeach leaflet.According to Stevenson (1981), four types

of trichomes occur in Zamia; of these, thetransparent ramified and the colored ramifiedare the types found in the present work. Incontrast to what was found in the aforemen-tioned study by Stevenson, no trend orcorrelation was found between aerial stems(i.e., Z. soconuscensis or Z. inermis) andbifurcate trichomes with equal lengthbranches. Trichomes with unequal sizedprevail among taxa with both aerial andsubterranean stems. In this context, weconsider that a more extensive and detailedsampling of the genus Zamia across theNeotropics would be necessary to corroborateany correlation between trichome morpholo-gy and stem habit.REPRODUCTIVE STRUCTURES.—Although the

characters employed for the identification ofspecies of Zamia have been obtained fromleaf morphology (Miquel, 1861, 1869; Regel,1857, 1876; de Candolle, 1868; Schuster,1932; Eckenwalder, 1980; Vovides et al.,1983; Newell, 1986; Schutzman & Vovides,1998; Schutzman et al., 1988; Stevenson,1993, 2001a, b, 2004; Nicolalde-Morejón etal., 2008), the evaluation of reproductivecharacters, especially those corresponding toovulate strobili, is essential to discriminateamong closely related taxa. Outstandingattributes that should be considered in thiscase are (i) the form and shape of the coneapex (Stevenson, 1987); (ii) the peduncleposition with respect to the vertical axis of

303NICOLALDE ET AL.: ZAMIA IN MEGA-MEXICO2009]

the cone, when mature (Schutzman et al.,1988); and (iii) the overall color of the cone(see descriptions and Figs. 2, 3). In contrast tothe ovulate strobili, pollen strobili showscarce variation at the species level, and theirutility to discriminate among species thatmight possess high degrees of genealogicalaffinity is relatively low. For the descriptionof the pollen reproductive axes, the terminol-

ogy introduced by Mundry and Stützel (2003)has been followed.CHROMOSOME NUMBERS.—Zamia shows the

highest chromosome numbers and karyotypevariation throughout the order Cycadales,with 2n counts of 16, 17, 18, 22, 23, 24, 25,26, 27, and 28 (Marchant, 1968; Norstog,1980, 1981; Vovides, 1983; Moretti &Sabato, 1984; Moretti, 1990a, b; Vovides &

FIG. 1. Trichomes. A–B. Zamia furfuracea. C–D. Z. katzeriana. E–F. Z. polymorpha.


FIG. 2. Illustrations of ovulate strobili. A. Zamia inermis. B. Z. vazquezii. C. Z. paucijuga. D. Z. lacandona. E. Z.spartea.


FIG. 3. Illustrations of ovulate strobili. A. Z. fischeri. B. Z. furfuracea. C. Z. cremnophila. D. Z. katzeriana. E. Z.polymorpha.


Olivares, 1996). In contrast, all Ceratozamiaspecies studied so far have stable diploidchromosome numbers (2n=16) and karyo-types, as do all the Dioon species analyzed todate with stable diploid chromosome numb-ers (2n=18) and karyotypes (Marchant, 1968;Vovides, 1983, 1985; Moretti, 1990a, b).Chromosome counts are presented and

illustrated (Fig. 4) here for the first time forZamia herrerae. This species has, but to alesser extent, cytotype polymorphisms similarto those found for Z. paucijuga (Moretti &Sabato, 1984) and Z. polymorpha (Vovides &Olivares, 1996; Stevenson et al., 1995–1996b). Zamia herrerae has 2n=23, 24 fromtwo populations along its Mexican range inChiapas (Fig. 4). Both the m (median regionof the chromosomes) and T (telocentric)chromosomes vary in number (6–11 T, 4–6 m) and are large (6–11 μm for T and 8–12μm for m) and their arms can be longer thanhalf the spindle axis, which can cause mitoticinstability during telophase (Schubert, 2007).Karyotype differences are probably due tocentric fissions occurring on some of thelarger m chromosomes, giving rise to telo-centrics with part of the centromere stillpresent. In this context, there are two generalhypotheses to explain karyotypic evolution in

Zamia: first, Norstog’s hypothesis (Norstog,1980, 1981) relating karyotype simplificationand symmetry with progressive fusion oftelocentric chromosomes, which predicts lowdiploid number in taxa with a high number ofmetacentric chromosomes, and secondly thatof more recent research on Zamia, whichpostulates centric fission rather than fusionproducing a progressively higher diploid num-ber and asymmetric karyotypes with a highnumber of telocentric chromosomes (Moretti& Sabato, 1984; Vovides & Olivares, 1996;Caputo et al., 2004). For a more in-depthdiscussion of mechanisms of chromosomeevolution in seed plants, see Jones (1998).The highly asymmetric karyotypes and

somatic chromosome numbers in both Z.paucijuga and Z. polymorpha, which alsoappear to be the pattern for Z. herrerae, seemto be correlated with the highest morpholog-ical variation and widest geographic distribu-tion of the genus in Mesoamerica. Zamiaherrerae has a range of about 1000 km,spanning El Salvador, Guatemala, and Chia-pas (Mexico); therefore, we suggest investi-gating the distribution of chromosomecharacter states in this species throughout itsrange. This karyotype asymmetry contrastswith their congeners of a more restricted

FIG. 4. Chromosomes of Zamia herrerae. A. 2n=23. B. 2n=24.


distribution, which often have less morpho-logical variation and a tendency towardsconstant chromosome number and karyotype,e.g., Z. cremnophila Vovides, Schutzman &Dehgan, Z. fischeri, Z. inermis, Z. katzeriana,Z. purpurea and Z. soconuscensis (all 2n=16),as well as Z. furfuracea, Z. spartea and Z.vazquezii (all 2n=18). Vovides and Olivares(1996) and Jones (1998) comment thatatypical chromosome number increase attrib-uted to fission is probably a result of stressfulinfluences.

DISTRIBUTION AND ENDEMISM

Seventy-five percent of the species in thisrevision are endemic to the type locality andnearby areas. They are limited to two or threeclose populations with low population densi-ties. These attributes are consistent withRabinowitz’s (1981) evaluation criteria forspecies rarity, which mainly considers infor-mation related to geographic range, habitatspecificity and local population size. Incongruence with these criteria, the endemicZamia species of Mesoamerica are consideredrare, threatened or endangered and are listedunder the IUCN Red List (IUCN, 2005; Hillet al., 2007).With 21 endemic cycad species in three

genera, Mexico has the highest cycad diver-sity and number of endemics of the region.Six species of Zamia are known in Guate-mala, two of which are endemic (Z. monticolaChamberlain and Z. tuerckheimii J. DonnellSmith); in Honduras, three species are knownof which three recently described species areendemic (Z. oreillyi C. Nelson, Z. sandovaliiC. Nelson, and Z. onanreyesii C. Nelson & G.Sandoval). Two further species are known forBelize, of which one is endemic (Z. prasinaW. Bull) and finally, El Salvador is repre-sented by one broadly distributed species, Z.herrerae, with a range that runs along thePacific seaboard through Guatemala and theSierra Madre de Chiapas in Mexico. Otherspecies of the genus in Mexico with broaddistributions are (a) Z. paucijuga, distributedalong the Pacific seaboard of Mexico rangingfrom Nayarit (northwest Mexico) to Oaxaca(southwest); (b) Z. polymorpha, distributedwidely throughout the Yucatán penninsula inMexico, Belize, and the Petén region of

Guatemala; (c) Z. loddigesii, ranging alongthe Gulf of Mexico seaboard from Tamauli-pas (northeast Mexico) to Tabasco (south-east); and (d) Z. variegata Warsz., distributedbetween Guatemala and southern Mexico.The southern and southeastern regions of

Mexico, comprising the states of Veracruz,Oaxaca, Chiapas, and Tabasco, are the mostdiverse area of Mexico for the genus Zamia,with seven endemic species (Z. cremnophila,Z. lacandona, Z. loddigesii, Z. katzeriana, Z.purpurea, Z. polymorpha, and Z. spartea).The Gulf of Mexico region has three micro-endemics Z. furfuracea, Z. inermis and Z.vazquezii, whereas two micro-endemic spe-cies, Z. fischeri and Z. soconuscensis, areknown respectively from the Sierra MadreOriental and the Sierra Madre de Chiapas.Finally, in spite of the status of Z. pauci-

juga as a Mexican endemic, its distribution isextremely wide within Mega-Mexico, cover-ing a range of about 1000 km between thestates of Oaxaca and Nayarit. The northernlimit of Z. paucijuga in the latter staterepresents the northernmost distribution forthe genus along the Pacific seaboard of theNeotropics.

Taxonomic treatment

Zamia L., Sp. Pl. ed. 2: 1659. 1763. nom.cons. Type: Zamia pumila L.Palma-Felix Adanson, Fam. Pl. 2: 21, 587.1763.

Aulacophyllum Regel, Gartenflora 25: 140.1876.

Stems hypogeous and epigeous, erect todecumbent, sometimes dichotomouslybranched in mature plants. Cataphylls charta-ceous to membranaceous, stipulate, persistentor deciduous, base triangular, apex longacuminate to aristate, tomentose, generallyreddish-brown to yellowish. Ptyxis erect toinflexed. Leaves stipulate, ascending todescending to spreading, reddish-brown orgreen when emerging; petiole sometimesblackish in young leaves, terete or subterete,without prickles or heavily to lightly armedwith straight or bifurcate prickles; rachissubterete generally with few prickles alongthe proximal third or without prickles, withup to 60 pairs of leaflets. Leaflets articulate,


sessile, papyraceous to coriaceous, linear,linear-lanceolate, lanceolate, ovate, obovateobpyriform to elliptic, opposite to suboppo-site, falcate or non-falcate, imbricate tonon-imbricate, generally acute at apex andsymmetric, attenuate at base, margins entireto dentate along upper third, subrevolute,articulations green, yellowish or dark brownin young leaflets. Pollen strobili usually 1–2(−4), with sterile tip, erect, cylindrical toconical, light brown to purple, tomentulose,apex acute to apiculate, generally with dense-ly tomentose peduncles; pollen sporangio-phores cuneiform, distal face truncate

hexagonal, 0.3–0.55 cm long, fertile abaxialsurface 2 lobed with 2–14 bisporangiatesynangia per lobe, sporangia dehiscent bylongitudinal slit. Ovulate strobili usuallysolitary, erect to decumbent, cylindrical toellipsoid, purple to yellowish, generallytomentulose, apex acute to apiculate; pedun-cle densely tomentose; ovulate sporangio-phores cuneiform-peltate to scutiform, distalend truncate-hexagonal when not scutiform.Seeds ovoid, sarcotesta white to pink whenimmature, red at maturity, sclerotesta smoothbut sometimes with several furrows runninglongitudinally from micropylar end.

Key to the species of Zamia in Mega-Mexico

1. Leaflets chartaceous to papyraceous.2. Leaflet margin dentate.

3. Leaflets elliptic, adaxial surface with yellow to cream variegation . . . . . . . . . . . . . . . . . . . . . Z. variegata3. Leaflets long-lanceolate, without variegation.

4. Leaflets imbricate, peduncle of pollen strobili decumbent, up to 16 cm long. . . . . . . . . . . . . . Z. oreillyi4. Leaflets not imbricate, peduncle of pollen strobili erect, up to 8 cm long . . . . . . . . . . . . . . .Z. herrerae

2. Leaflet margin serrulate to entire.5. Leaflet margin entire, chartaceous.

6. Leaflets oblong-lanceolate, glossy, 4–6 cm wide; ovulate strobili iridescent blue-green at matu-rity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Z. tuerckheimii

6. Leaflets linear-lanceolate, not glossy, 0.6–1.5 cm wide, ovulate strobili dark-brown at matu-rity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Z. soconuscensis

5. Leaflet margin serrulate, papyraceous.7. Leaflets sub-falcate basally, long-acuminate, strongly apically curved. . . . . . . . . . . . . Z. monticola7. Leaflets straight, acuminate, not curved at apex.

8. Leaflets elliptic to lanceolate, ovulate strobili cylindrical to ovoid, dark-green and glabrous whenmature; from San Luís Potosí and Querétaro, Mexico . . . . . . . . . . . . . . . . . . . . . . . Z. fischeri

8. Leaflets ovate to obpyriform, ovulate strobili ovoid-cylindrical, gray to brown tomentulose whenmature; endemic to Veracruz, Mexico . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Z. vazquezii

1. Leaflets coriaceous.9. Leaflet margin entire to serrulate.10. Leaflet margin entire, petiole unarmed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Z. inermis10. Leaflet margin serrulate, petiole prickly.

11. Leaflets long acuminate apically; stems arborescent . . . . . . . . . . . . . . . Z. onanreyesii11. Leaflets rounded to acute apically; stems subterranean.

12. Leaflets obovate to oblanceolate, keeled adaxially, apex rounded . . . . Z. furfuracea12. Leaflets linear to oblanceolate, flat adaxially, apex acute.

13. Leaflets linear, 0.4–0.6 cm wide . . . . . . . . . . . . . . . . . . . . . . . . . .Z. spartea13. Leaflets lanceolate, ≥0.7 cm wide.

14. Leaflets falcate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Z. sandovallii14. Leaflets not falcate.

15. Leaflets lanceolate-oblanceolate, ovulate strobili dark-brown tomentu-lose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Z. polymorpha

15. Leaflets linear-lanceolate, ovulate strobili brown to yellowish.16. Ovulate strobili ellipsoid to cylindrical, apex acute to apiculate,

yellowish-brown; from the Pacific seaboard of Mexico . . .Z. paucijuga16. Ovulate strobili ellipsoid to conical, apex acute, yellowish; from the Gulf

of Mexico seaboard . . . . . . . . . . . . . . . . . . . . . . . . . . . Z. loddigesii9. Leaflet margin distinctly dentate.

17. Leaflets channeled adaxially between veins, appearing plicate . . . . Z. purpurea17. Leaflets smooth, not channeled adaxially between veins, not appearing plicate.

18. Leaflets linear-lanceolate, imbricate, petiole strongly armed with straight orbifurcate prickles up to 6 mm long . . . . . . . . . . . . . . . . . .Z. cremnophila


18. Leaflets lanceolate to oblanceolate, not imbricate, petiolearmed with smallstraight prickles, generally between 2–4 mm.19. Leaflets with brilliantly shining cuticle on adaxial surface;ovulate strobili

decumbent when mature . . . . . . . . . . . . . . . . . . . . . . . . Z. katzeriana19. Leaflets without brilliantly shining cuticle on adaxial surface; ovulate

strobili erect when mature.20. Distal leaflets sub-falcate, petiole with bulbous base, blackish in young

leaves. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Z. lacandona20. All leaflets straight, petiole with no bulbous base, greenish in young

leaves.21. Leaflets oblong to lanceolate, bright grass-green, without conspic-

uously denticulate along margins; ovulate strobili green glabrouswhen mature, apex acuminate . . . . . . . . . . . . . . . . Z. prasina

21. Leaflets long-lanceolate, conspicuous dentate, up to 4 mm long;ovulate strobili brown tomentulose when mature, apex long-apiculate. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Z. standleyi

Zamia cremnophila Vovides, Schutzman &Dehgan., Bot. Gaz. 149(3): 351. 1988. Type:Mexico. Tabasco: 18 Aug 1981, M. A.Magaña& S. Zamudio 343 (holotype: MEXU;isotypes: CSAT, FCME, MO, UAMIZ).

Stem hypogeous, generally unbranched, 5–34 cm long, 4–11.3 cm in diam. Cataphyllschartaceous, persistent, base triangular, apexaristate, 5.4 cm long, 2.9 cm at base, reddish-brown tomentose. Ptyxis inflexed. Leaves 2–3(4), 57.5–179×40.2–76.1 cm, descending, red-dish-brown when emerging; petiole 10.2–95.3 cm long, blackish in young leaves,subterete, heavily armed with straight or bifur-cate prickles up to 6 mm long; rachis subterete,up to 84 cm long, with few prickles along theproximal third. Leaflets 5–28 pairs, sessile,coriaceous, lanceolate, opposite to subopposite,imbricate, apex acute, base attenuate, marginsdentate along distal third, subrevolute; articula-tions dark brown in young leaflets, 0.8–1.2 cm;the median leaflets 22.7–38×3.1–4.4 cm wide.Pollen strobili usually 1–2, erect, cylindrical, upto 8 cm long and 1.4 cm in diam, light brown,apex acute; peduncle densely light browntomentose, 4.5 cm long, 1.2 cm in diam; pollensporangiophores cuneiform, distal face truncate-hexagonal, 0.35 cm long, fertile abaxial surfacewith 3 bisporangiate synangia per lobe. Ovulatestrobili usually solitary, erect, ellipsoid, 13.2 cmlong, 4.9 cm in diam, brown to reddish,tomentulose, apex acuminate; peduncle denselybrown tomentose, 2.9 cm long, 1.6 cm in diam;distal face hexagonal-truncate with a horizontallongitudinal depression, 1–1.2 cm high, 1.1–1.4 cm wide. Seeds ovoid, sarcotesta white

when immature, red at maturity, 2–2.8 cm long,1.4–2 cm in diam, sclerotesta smooth.Chromosome number.—2n=16 (Schutzman

et al., 1988).Diversity and genetic structure.—The av-

erage of alleles per locus is A=1.98, thepercentage of polymorphic loci is P=94.3, theexpected heterozygosity is HE=0.347 andthe genetic differentiation between the twopopulations currently under study is Fst=0.093 (González-Astorga et al. unpubl. data).Distribution and habitat.—Endemic to

Tabasco, Mexico (Fig. 5), on karstic rocks andcliffs of the Sierra El Madrigal, between 50–150 m. The vegetation type where this speciesgrows is evergreen tropical rain forest or bosquetropical perennifolio of Rzedowski (1978).Etymology.—The specific epithet is derived

from the Greek word for cliff-friend(κρɛμνóσ = cremnos = cliff, and �ιλοσ =filos = friend/lover; Schutzman et al., 1988),because of its unusual habitat.Distinguishing features.—The species is

characterized by its exclusive habit on therocky walls of limestone cliffs, in addition todescendent leaves. Petioles are densely armedwith prickles that are sometimes branched,and also lanceolate, imbricate leaflets that arevisibly dentate along the distal third.

Additional specimens examined.— MEXICO.TABASCO: Teapa, F. Nicolalde-Morejón et al. 1497(XAL), M. A. Pérez-Farrera 293 (HEM), M. A. Pérez-Farrera 900 (HEM, MEXU).

Zamia fischeriMiq. in Lem., Hort. Vanhoutt. 1:20. 1845. Type: ex Horto Petropolitano in H.Houtte. vecta,Miquel s.n. (neotype, designatedby Stevenson & Sabato, 1986: U). (Fig. 7)


Stem subterranean, dichotomously branchingin older plants, up to 30 cm long, 4–8 cm indiam. Cataphylls membranaceous, persistent,base triangular, apex aristate, 4.5 cm long,1.2 cm at base, yellowish tomentose. Ptyxisinflexed. Leaves 1–5(–8), 15–45×8–20 cm,ascending to spreading, dark-brown whenemerging; petiole 9–14 cm long, blackish inyoung leaves, unarmed, terete; rachis subterete,up to 31 cm long, unarmed. Leaflets 20–35pairs, sessile, papyraceous, elliptic to oblanceo-late, alternate to subopposite, apex acute sym-metric, base cuneate, margins serrulate alongdistal third, subrevolute; articulations light-brown in young leaflets, 0.3–0.4 cm wide,median leaflets 5–9×1.5–4.5 cm. Pollen stro-bili usually 1–3, erect, conical, 4–6 cm long,1.5–2.2 cm in diam, gray tomentose, apexacute; peduncle yellowish tomentose, 2.8 cmlong, 0.9–1.1 cm in diam; pollen sporangio-phores cuneiform, distal face hexagonal-truncate, 0.3 cm long, fertile abaxial surfacewith 3–4 bisporangiate synangia per lobe.Ovulate strobili usually 1–2, erect, cylindrical

to ovoid, up to 9 cm long, 4.5 cm in diam, brownto reddish tomentulose when young, dark-greenand glabrous whenmature, apex acute; pedunclebrown tomentose, 3 cm long, 1.1 cm in diam;megasporangiophores peltate, distal face hexag-onal-truncate when immature, scutiform whenmature 0.8–1.2 cm high, 1.3–1.9 cm wide. Seedovoid, 1.6 cm long, 1.3 cm in diam, sarcotestapink when young and orange at maturity.Chromosome number.—2n=16 (Marchant,

1968; Moretti et al., 1991; Stevenson et al.,1995–1996a).Distribution and habitat.—Zamia fischeri

is endemic to Mexico (Tamaulipas, San LuisPotosí, Querétaro and Hidalgo; Fig. 5), be-tween 140–900 m elevation. The vegetationtype of its habitat is pine-oak forest, tropicaldeciduous forest and mountain tropical forest(sensu Rzedowski, 1978).Etymology.—The specific epithet is in

honor of Friedrich Fischer, a German cycadhorticulturist of the 19th century.Distinguishing features.—Zamia fischeri is

a small fern-like plant (up to 100 cm tall),

FIG. 5. Distribution of Zamia cremnophila ( ), Z. furfuracea ( ), Z. inermis (●), Z. katzeriana ( ), Z. lacandona(■), Z. monticola ( ), Z.onanreyesii ( ), Z. prasina ( ), Z. sandovallii (▲), Z. soconuscensis ( ), Z. vazquezii ( ).


with papyraceous but serrulate leaflets andunarmed petiole and rachis. Ovulate strobiligreenish upon maturity.

Additional specimens examined.—MEXICO.QUERÉTARO: Jalopan, López 438 (XAL), Servín 1471(XAL), Vovides 330 (XAL). SAN LUIS POTOSÍ: O. M.Clark 6839 (MO), J. Rees 1686 (XAL), Stevenson et al.566 (MEXU, NY), Vovides 753 (XAL); El Naranjo, F.Nicolalde-Morejón & J. González Astorga 1614 (XAL),1615 (XAL), 1616 (XAL), 1617 (XAL), 1618 (MEXU,XAL), 1619 (XAL); Ciudad Valles, F. Nicolalde-Morejón& J. González-Astorga 1620 (XAL), 1621 (XAL), 1622(XAL), 1623 (MEXU, XAL), 1624 (MEXU, XAL), 1625(MEXU, XAL), 1626 (MEXU, XAL). HIDALGO: Pisa-flores, O. Alcántara-Ayala & R. Mayorga-Saucedo 3325(FCME).

Zamia furfuracea L. f. in Aiton, HortusKew. 3: 477. 1789. Type: Palma america-na crassis rigidisque foliis, pl. 210, inHerm. Paradisus Batavus. 1698 (lectotype,designated by Stevenson & Sabato, 1986:pl. 210).

Zamia muricata var. obtusifolia Miquel, Tijdschr. nat.Gesch. Physiol. 10(1): 71–72. 1843. Zamia latifo-lia Loddiges ex Miquel, Tijdschr. wis-en natuurk.Wet. 2(4): 298. 1849. Type: Tab. VII, fig. a inLinnaea 19(4): 1847. (neotype, designated byStevenson & Sabato, 1986: Tab. VII, fig. a).

Zamia furfuracea var. trewii A. DC., Prodr. 16(2):541. 1868. Type: Palmifolia fructu clavato poly-pireno[polyspermo]. C. J. Trew, PI. Select. Tab.26. 1752 (holotype: Tab. 26, G; typotype, desig-nated by Stevenson & Sabato, 1986: BM).

Stem hypogeous, becoming epigeous withage, often dichotomously branched, up to60 cm long, 20 cm in diam. Cataphyllscoriaceous, persistent, base triangular, apexlong aristate, up to 10×3–4.5 cm at base,yellowish tomentose. Ptyxis inflexed. Leaves3 to many, 45–190×10–30 cm, diffuse,brown-yellowish when emerging; petiole17–50 cm long, brown-yellowish whenyoung, subterete, armed with small pricklesup to 3 mm long; rachis terete, up to 120 cmlong, with few prickles along the proximalthird. Leaflets 8–18 pairs, sessile, coriaceous,obovate to oblanceolate, opposite to subop-posite, imbricate, keeled, apex rounded tosub-acute, base attenuate, margins slightlyserrulate along the 1/2 distal portion, sub-revolute; articulations yellow in young andjuvenile leaflets, 0.4–0.7 cm wide; medianleaflets 14–20×4–7.5 cm. Pollen strobili

usually 2–4, erect, cylindrical, up to 17 cmlong, 1.8 cm in diam, yellowish to brown,apex acute; peduncle densely light-browntomentose, 12 cm long, 1.1 cm in diam;pollen sporangiophores cuneiform, distal facehexagonal-truncate, 0.45–0.55 cm long, fer-tile abaxial surface with 8–9 bisporangiatesynangia per lobe. Ovulate strobili usually 1per crown, erect, cylindrical, up to 25 cmlong, up to 10.5 cm in diam, yellowish-greentomentulose when immature, light brownupon maturity, apex apiculate; peduncle yel-lowish tomentose, up to 8 cm long, up to1.3 cm in diam; megasporangiophores pel-tate, distal end hexagonal-truncate, 0.9 cmhigh, 1.3 cm wide. Seeds ovoid, sarcotestayellowish-green when immature turning redat maturity, 1.6 cm long, 1.2 cm in diam,sclerotesta smooth.Chromosome number.—2n=18 (Moretti,

1990a, b)Diversity and genetic structure.—The av-


Mexico in central and south coastal Veracruz,along a coastal stretch of approximately150 kmin stable dunes and basalt cliffs (Fig. 5).Etymology.—The specific epithet alludes to

the persistent brown-yellowish trichomes ofthe leaves throughout the developmentalstages of the plant.Distinguishing features.—Leaves strongly

keeled with highly imbricate and coriaceousobovate to oblanceolate leaflets with brown-yellowish indumentum persisting with age.

Additional specimens examined.—MEXICO.VERACRUZ: Ibarra-Mariquez 316 (MEXU, MO),Ibarra-Mariquez 1952, 1959 (MEXU); Alvarado, Rees1650, 1651, 1652 (XAL), M. Vázquez-Torres et al. 4871(CIB); Catemaco, Calzada 1475 (MEXU), Calzada 2451(XAL), Cedillo 2610 (MEXU, XAL), González-Quintero1520 (ENCB), Ibarra 316 (MEXU), Lot 1277-14 (F,XAL), Menendez 115 (MEXU, MO, XAL), F. Nicolalde-Morejón et al. 1484 (XAL), 1485 (XAL), 1486 (XAL), J.Rees 1651 (IBUG, MEXU, XAL), J. Rees 1652 (XAL),Schatz & Nee 207 (XAL), Vovides 567 (MEXU, XAL);Mecayapan, Calzada et al. 11325 (XAL), Castillo-Campos 12732 (XAL); Lerdo De Tejada, Vovides 828(XAL), 829 (XAL), 830 (XAL), 831 (XAL), 832 (XAL),


833 (XAL), 839 (XAL); San Andrés Tuxtla, Castillo-Campos et al. 13881 (XAL), Chazaro B. 512 (XAL),Hammel & Merello 15499 (MO), Hernández-M 1216A(F, MEXU), Lorence 4978 (MEXU), Sousa 3099 (F,MEXU, MO), Vovides & Iglesias 1148 (XAL).

Zamia herrerae Calderón & Standl., Proc.Wash. Acad. Sci. 14(4): 93. 1924. Type: ElSalvador. Vicinity of Sonsonate, 17 Jul1923, S. Calderón 1682 (holotype: US).

Stem hypogeous, bifurcate in adult plants,6–26×4.5–13.5 cm in diam. Cataphylls char-taceous, persistent, base triangular, apexaristate, 4.6×1.2 cm wide at base, yellowishtomentose. Ptyxis inflexed. Leaves 2–4, erect,green to light-brown when emerging, 61–96×24.5–29.5 cm; petiole 19–34 cm long, brown-ish in young leaves, terete, armed withprickles up to 3 mm long; rachis subterete,up to 63 cm long, with few prickles along theproximal third. Leaflets 15–32 pairs, sessile,papyraceous, lanceolate, alternate to subop-posite, apex acute, base symmetric attenuate;margins dentate along distal 2/3, subrevolute;articulations dark brown when young, 0.3–0.5 cm wide; median leaflets 22–38×3.1–4.4 cm. Pollen strobili usually 2–3, erect,cylindrical to conical, 4.3–7.5 cm long, 1.3–2.1 cm in diam, light brown tomentulose,apex mucronate; peduncle densely light-brown tomentose, 6.2–7.8 cm long, 1.1–1.4 cm in diam; pollen sporangiophorescuneiform, distal face hexagonal, 0.35 cmlong, fertile abaxial surface with 3–4 bispor-angiate synangia per lobe. Ovulate strobiliusually solitary, erect, cylindrical to ovoid,7.1–11.6 cm long, 4.1–4.9 cm in diam,brown, tomentulose, apex acute; peduncledensely brown tomentose, 3.5–4.2 cm long,1.1–1.3 cm in diam; megasporangiophorespeltate, distal face hexagonal-truncate, 1.3–1.6 cm high, 2.2–2.8 cm wide. Seeds ovoid,sarcotesta pink when immature, red at matu-rity, 1.6–1.9 cm long, 1.2–1.5 cm in diam,sclerotesta smooth.Chromosome number.—2n=23, 24 (Fig. 4).Distribution and habitat.—The species

range is El Salvador, Guatemala, and southernMexico in Chiapas, between 100–600 melevations (Fig. 6). It generally grows intropical deciduous forest (sensu Rzedowski,1978) on deep clay soils. Also, this cycad may

be found in secondary growth forests andpastures.Etymology.—The specific epithet honors

Hector Herrera, a scientist from El Salvador.Distinguishing features.—Zamia herrerae is

distinguished by its long-lanceolate papyra-ceous leaflets with dentate margins along thedistal 2/3.

Additional specimens examined.—MEXICO.CHIAPAS: Escuintla, Matuda 16368 (MEXU), 16871(MEXU), 17332 (MEXU), 18332 (MEXU), M. A.Pérez-Farrera 143 (HEM), Schutzman 526 (XAL), 527(XAL), 528 (XAL); Tonalá, Farrera 2489 (CHIP), F.Nicolalde-Morejón & J. González-Astorga 1579 (XAL),1580 (XAL), 1581 (XAL), M. A. Pérez-Farrera 744(CIB).

GUATEMALA. P. C. Standley 67306 (F).

NICARAGUA. MANAGUA: Cultivated, A. Grijalva3658 (MO).

Zamia inermis Vovides, J.D. Rees & Vázq.Torres, Flora de Veracruz 26: 22. 1983. Type:Mexico. Veracruz: 6 Jun 1981, Vovides 666(holotype: XAL; isotype: F). (Fig. 8)

Stem epigeal, erect, dichotomously branch-ing in mature plants, 15–43 cm long, 8.6–26.4 cm in diam. Cataphylls chartaceous,persistent, base triangular, apex aristate, 2.1–2.6×4.1–5.6 cm wide at base, yellowishtomentose. Ptyxis inflexed to erect. Leaves 10–35, erect, light to yellowish-green when emerg-ing, 30–95×43.5–60 cm; petiole 18–41 cm long,greenish in young leaves, subterete, withoutprickles; rachis subterete, 15–19 cm long,unarmed. Leaflets 27–32 pairs, sessile, coria-ceous, linear-lanceolate, opposite to suboppo-site, apex acute, base attenuate; margins entire,subrevolute; articulations 0.4–0.6 cm wide;median leaflets 20–30.5×0.9–1.2 cm. Pollenstrobili cylindrical, usually 1–2 per crown, erect,up to 9.1 cm long, up to 2.8 cm in diam, beige-yellowish, apex acute; peduncle densely light-yellowish tomentose, up to 4.5 cm long, 1.1 cmin diam; pollen sporangiophores cuneiform,distal face hexagonal-truncate, 0.35 cm long,fertile abaxial surface with 5–6 bisporangiatesynangia per lobe. Ovulate strobili usually 1–2per crown, erect, cylindrical, 13–23 cm long, 8–9.8 cm in diam, light-brown to beige tomentu-lose, apex apiculate; peduncle brown tomentose,6–8 cm long, 1.2–1.4 cm in diam; megaspor-angiophores peltate, distal face hexagonal-trun-


cate, 0.7–0.9 cm high, 1.1–1.3 cm wide. Seedsovoid, sarcotesta pink when immature, red atmaturity, 1.7–2.5 cm long, 1.4–2.1 cm in diam,sclerotesta smooth.Chromosome number.—2n=16 (Vovides,

1983).Distribution and habitat.—Endemic to

Mexico in a small mountain range in centralVeracruz (Fig. 5) at 150–300 elevation onbasaltic soils. The vegetation type of the habitatis tropical deciduous forest (sensu Rzedowski,1978).Etymology.—The specific epithet alludes to

the absence of prickles on the petiole andrachis and the entire leaflet margins.Distinguishing features.—Zamia inermis

differs from its congeners in Mexico by atotal absence of prickles along the petiole andrachis, as well as having totally entire leafletsand light-yellowish to clear-beige tomentu-lum on the pollen and ovulate strobili.

Additional specimens examined.—MEXICO.VERACRUZ: Actopan, Acosta & Acosta 234 (XAL), F.

Nicolalde-Morejón & Vovides 1415 (XAL), 1416 (XAL),1417 (XAL), Schutzman 570 (XAL), 571 (XAL), 572(XAL), 575 (XAL), 576 (XAL), 577 (XAL), J. Rees etal. 681 (XAL).

Zamia katzeriana (Regel) Rettig, Garten-flora 45: 148. 1896. Ceratozamia katzeri-ana Regel, Acta Horti Petrop. 4(4): 298.1876. Type: ex Horto Katzer., Regel s.n.(lectotype, designated by Stevenson &Sabato, 1986: LE).

Zamia splendens Schutzman, Phytologia 55(5): 299.1984. Type: Cultivated in Fairchild Tropical Garden,Miami, accession no. FTG 76–1046, 11 Apr 1984, J.Watson s.n. (holotype: NY; isotypes: FLAS, FTG,MEXU).

Stem hypogeous, unbranched, up to 25 cmlong, up to 7 cm in diam. Cataphylls charta-ceous, semidecidious, base triangular, apexaristate, 5.3×1.4 cm at base, yellowishtomentose. Ptyxis inflexed. Leaves 1–2 (3),49–220×35–58 cm, ascending to descending,bright reddish-pink with lustrous cuticle when

FIG. 6. Distribution of Zamia fischeri (●), Z. herrerae ( ), Z. loddigesii ( ), Z. oreillyi ( ), Z. paucijuga ( ), Z.polymorpha (■), Z. purpurea (▲), Z. spartea ( ), Z. standleyi ( ), Z. tuerckheimii ( ), Z. variegata ( ).


emerging, dark green when mature; petiole20–130 cm long, terete, armed with fewsimple prickles; rachis subterete, up to84 cm long, unarmed. Leaflets 3–7 pairssessile, coriaceous, oblong-lanceolate, oppo-site to subopposite, adaxial surface withbrilliantly shining cuticle throughout life ofleaflet, apex acute, base attenuate; marginsdentate along distal third, subrevolute; artic-ulations brown in young leaflets turning greenwith age 0.6–1.6 cm wide; median leaflets18–35×3.5–12 cm. Pollen strobili 1–5, con-ical, light-brown tomentulose, erect uponemergence becoming prostrate to decumbentupon maturity, up to 3.9 cm long, 1.1 cm indiam, apex acute; peduncle light-brown to-mentose, up to 5.8 cm long, 1.2 cm in diam;pollen sporangiophores cuneiform, distal facehexagonal scutiform, 0.35 cm long, fertile

abaxial surface with 5–6 bisporangiate syn-angia per lobe. Ovulate strobili usuallysolitary, decumbent to erect, elliptic, 8–12 cm long, 4.5–6 cm in diam, brown toyellowish tomentose, apex aristate; pedunclebrown tomentose, 2.1–4.3 cm long, 1.1–1.3 cmin diam; megasporangiophores peltate, distalface hexagonal-truncate to scutiform, 1–1.2 cmhigh, 1.1–1.3 cm wide. Seeds ovoid, sarcotestapink when immature, red at maturity, 1.1–1.4 cm long, 1.6–1.8 cm in diam, sclerotestasmooth.Chromosome number.—2n=16 (Schutzman,

1984; Moretti, 1990a).Diversity and genetic structure.—The

average of alleles per locus is A=1.95, thepercentage of polymorphic loci is P=84.4,the expected heterozygosity is HE=0.280and the genetic differentiation between

FIG. 7. Zamia fischeri A. Habit. B. Leaflets. (From Stevenson 566, NY).


FIG. 8. Zamia inermis. A–B. Habit. C. Cataphyll. D. Pollen strobilus. E. Microsporophyll, abaxial and adaxialview. F–H. Ovulate strobilus. I. Ovulate sporangiophores. J. Seed. (From the holotype.)


the two populations currently under study isFst=0.194 (González-Astorga et al. unpubl.data).Distribution and habitat.—Endemic to

Mexico and known from the states ofChiapas, Tabasco and Veracruz (Fig. 5) at200–700 m in evergreen tropical forest (sensuRzedowski, 1978).Etymology.—Specific epithet in honor of

Katzer, inspector of the gardens in Paullowsk(Stevenson & Sabato, 1986).Distinguishing features.—Zamia katzeriana

is easily distinguished from its congeners byhaving leaves with highly lustrous or shinycuticles. Emerging leaves are a bright reddish-pink. Both pollen and ovulate strobili areborne on long peduncles that become descen-dent to prostrate.

Additional specimens examined.—MEXICO.VERACRUZ: Las Choapas, Martínez & Martínez-M. 825(HEM), F. Nicolalde-Morejón et al.1436 (XAL), 1437(XAL). TABASCO: Teapa, Hdez-Najarro 622 (CHIP), M.A. Magaña 1905 (MEXU), M. A. Pérez-Farrera s.n.(XAL), M. A. Pérez-Farrera 899 (HEM, MEXU),Walters s.n. (FTG accession 12-2, XAL). CHIAPAS: SanFernando, Vovides et al. 1266 (XAL), Palacios 383(CHIP), F. Nicolalde-Morejón & Pérez-Farrera 1420(XAL), M. A. Pérez-Farrera s.n. (XAL), Walters s.n.(FTG accession 23-2, XAL); Ocozocoautla, Gómez-Pompa 705 (FCME, MEXU), F. Nicolalde-Morejón etal. 1453 (XAL), 1454 (XAL), 1455 (XAL), 1456 (XAL),1457 (XAL), 1458 (XAL), 1459 (XAL), 1460 (XAL), M.A. Pérez-Farrera 29 (CHIP, CIB, MEXU); Tila, Vovideset al. 1340 (XAL), 1343 (XAL), 1341 (XAL).

Zamia lacandona Schutzman & Vovides,Novon 8(4): 441. 1998. Type: Mexico.Chiapas: Selva Lacandona, July 1984,Schutzman 517 (holotype: FLAS; isotype:XAL).

Stem hypogeous, unbranching, 6–14 cmlong, 5–9 cm in diam. Cataphylls coriaceous,persistent, base triangular, apex long-aristate,8.3×3.6 cm at base, reddish-brown tomen-tose. Ptyxis inflexed, reddish-brown. Leavesusually solitary, up to 3 under cultivation, 46–171×32–75 cm, ascending, reddish-brownwhen emerging; petiole 14–95.3 cm longwith bulbous base, blackish in young leaves,subterete, proximal section strongly chan-neled, armed with prickles up to 5 mm long;rachis subterete, 31–76 cm long, with fewprickles along proximal third. Leaflets 4–17

pairs, sessile, coriaceous, lanceolate, oppositeto subopposite, subfalcate, apex acute, baseattenuate, margins dentate along distal third,subrevolute; articulations dark-brown whenyoung, 0.4–1.1 cm wide; median leaflets15.6–37×2.9–6 cm. Pollen strobili usually2–3, erect, conical, 5.4–6.6 cm long, 1.5–1.7 cm in diam, light-brown, apex acute;peduncle light-brown tomentose, 5.8–6.9 cmlong, 1.2 cm in diam; pollen sporangiophorescuneiform, distal face hexagonal truncate,0.35 cm long, fertile abaxial surface with 4–5 bisporangiate synangia per lobe. Ovulatestrobili usually solitary, erect, ellipsoid,13.2 cm long, 4.9 cm in diam, dark-brown,tomentulose, apex acute to slightly apiculate;peduncle densely brown tomentose, 6.5 cmlong, 1.2 cm in diam; megasporangiophorespeltate, distal face hexagonal-truncate, 1–1.2 cm high, 1–1.5 cm wide. Seeds irregularlyovoid, sarcotesta pink when immature, red atmaturity, 2–2.4 cm long, 1.3–1.9 cm in diam,sclerotesta smooth.Chromosome number.—2n=16, 17, 18

(Schutzman & Vovides, 1998).Diversity and genetic structure.—The av-


Chiapas (Fig. 5) in the vicinity of theLacandon forest at 80–200 m elevation, inevergreen tropical forest (sensu Rzedowski,1978). It is also found in secondary succes-sion stages of the tropical forest.Etymology.—The epithet is derived from

the name of the 1.8 million hectare SelvaLacandona (Lacandona Jungle) in southeast-ern Chiapas, which itself bears the name ofthe Lacandona Maya Indians who inhabit theforest.Distinguishing features.—This species is

distinguished by having a solitary large leaf(up to three leaves may be maintained onplants under cultivation) with a stout erectpetiole and a strongly bulbous base. The leafis reddish-brown at emergence and petioledark purplish-brown turning dark-brown withage. Ovulate cone generally solitary, withacute apex.


Additional specimens examined.—MEXICO.CHIAPAS: Palenque, Schutzman 510 (XAL), 511 (XAL),512 (XAL), 513 (XAL), 514 (XAL), 515 (XAL), 516(XAL), 517 (XAL), 518 (XAL), 519 (XAL), 520 (XAL),F. Nicolalde-Morejón & N. Martínez 1418 (XAL), M. A.Pérez-Farrera 890 (HEM, MEXU), Walters s.n. (FTGaccession 14-2, XAL), M. Vázquez-Torres et al. 3925(CIB); San Jerónimo Tulija, Chavelas et al. ES=315(ENCB, MEXU), Schutzman 521 (XAL), 522 (XAL),524 (XAL), 523 (XAL), 525 (XAL).

Zamia loddigesii Miq., Tijdschr. Natuurl.Gesch. Physiol. 10: 72. 1843. Type: culti-vated by Van Houtte 3374 (lectotype, heredesignated: U).

Zamia galeotti De Vriese, in Hoven & De Vries.Tijdschr. Natuurl. Gesch. Physiol. 12: 24. 1845.Type: Mexico. Veracruz: 5 July 1983. D. W.Stevenson 538 (neotype, here designated: NY,isoneotype: XAL).

Zamia leiboldii Miq., Linnaea 19: 425. 1847. Zamialoddigesii var. leiboldii (Miquel) A. DC., Prodr. 16(2): 541. 1868. Type: E. Mexico in HortumLoehrianum Lipsiae attulit Liebold, 1845, Miquels.n. (holotype: U).

Zamia loddigesii var. angustifolia Regel, Bull. Soc.Nat. Moscou 30(1): 190. 1857. Type: ex hortoPetropolitano, 1856, Regel s.n. (holotype: LE)

Zamia loddigesii var. obtusifolia Regel, Bull. Soc.Nat. Moscou 30(1): 190. 1857. Type: t. 186, figs27–28 in Gartenflora 6: 1857. (lectotype, designat-ed by Stevenson & Sabato, 1986: LE).

Zamia mexicanaMiquel, Prodr. syst. Cycad. 13. 1861.Type: Eriozamia mexicana H. Belg., 1847, Miquels.n. (holotype: U).

Zamia leiboldii var. angustifolia Regel, Trudy Imp.S.-Petersburgsk. Bot. Sada 4(4): 307. 1876. Type:Mexico. Oaxaca: 15 Jul 1983. D. W. Stevenson 559(neotype, here designated: NY; isoneotype, XAL).

Zamia leiboldii var. latifolia Regel, Trudy Imp. S.Petersburgsk. Bot. Sada 4(4): 307. 1876. Type: exHorto Petropolitano, 1875, Regel s.n. (holotype:LE).

Zamia lawsoniana Dyer in Hemsley, Biol. Centr.-Amer., Bot. 3(16): 195. 1884. Type: Mexico.Oaxaca: Fielding 209 (holotype: OX; isotype: K)

Zamia cycadifolia Dyer in Hemsley, Biol. Cent.-Amer., Bot. 3(16): 195. 1884, non Jacquin 1809.Zamia loddigesii var. cycadifolia Schuster, Pflan-zenreich 99: 148. 1932. Type: Mexico. Bourgeau s.n. (holotype: K; isotype: C). nomen illegit.

Zamia sylvatica Chamberlain, Bot. Gaz. 81: 223.1926. Type: Mexico, Oaxaca, Tuxtepec, Sep 1910,C. J. Chamberlain s.n. (lectotype, designated byStevenson & Sabato, 1986: NY; isolectotype, F-3sheets).

Zamia loddigesii var. angustifolia (Regel) J. Schust.,in Engl., Pflanzenr. 4(1): 148. 1932. Type: México.Veracruz: savanne bei Mundo nuevo, Karwinski

1028b (lectotype, designated by Stevenson &Sabato, 1986: LE; isolectotype, LE).

Zamia loddigesii var. longifolia J. Schust., in Engl.,Pflanzenr. 4(1): 147. 1932. Type: Mexico. Vera-cruz: Colipa, Karwinski 1029 (lectotype, designat-ed by Stevenson & Sabato, 1986: LE; isolectotype:LE).

Stem hypogeous, branching dichotomouslywith age, 10–45 cm long, 8–15 cm in diam.Cataphylls chartaceous, persistent, base trian-gular, apex aristate, 8.4×3.7 cm at base,yellowish tomentose. Ptyxis inflexed. Leaves2–3 (4) ascending to spreading, 45–96×30–41 cm, light-green when emerging, green todark-green when mature; petiole 15–25 cmlong, green in young leaves, subterete, armedwith prickles up to 4 mm long; rachissubterete, up to 57 cm long, with few prickleson the proximal third. Leaflets 12–23 pairs,sessile, coriaceous, linear-lanceolate, oppositeto subopposite, apex acute, base attenuate,margins serrulate along distal third, subrevo-lute; articulations 0.4–0.7 cm wide; medianleaflets 16–26×1.8–3.1 cm. Pollen strobili 1–2 per crown, up to 6 (7) when multiplecrowned, erect, cylindrical, 8–14 cm long,1.8–3.5 cm in diam, light brown tomentulose,apex acute; peduncle light-brown tomentose,6 cm long, 1.2 cm in diam; pollen sporangio-phores cuneiform, distal face hexagonal-truncate, 0.3 cm long, fertile abaxial surfacewith 6–8 bisporangiate synangia per lobe.Ovulate strobili usually 1–2 per crown, erect,ellipsoid to conical, up to 16 cm long, up to6 cm in diam, beige-tomentulose, apex acute;peduncle brown-tomentose, up to 6 cm long,1.6 cm in diam; megasporangiophores peltate,distal face hexagonal-truncate, 0.7–1 cm high,1.9–2.6 cm wide. Seeds ovoid, sarcotesta pinkwhen immature, red at maturity, 1.4–1.8 cmlong, 0.8–1 cm in diam, sclerotesta smooth.Chromosome number.—2n=18 (Norstog,

1980; Moretti, 1990a, b).Diversity and genetic structure.—The av-

erage of alleles per locus is A=1.8, thepercentage of polymorphic loci is P=66.6,the expected heterozygosity is HE=0.266and the genetic differentiation between thetwo populations currently under study isFst=0.179 (González-Astorga et al. unpubl.data).Distribution and habitat.—Endemic to

Mexico and distributed widely within the


states of Tamaulipas, Hidalgo, Veracruz,Tabasco and parts of Oaxaca with a singleknown locality in Chiapas (Fig. 6). Thevegetation types are evergreen tropical forest,tropical deciduous and sub-deciduous forests(sensu Rzedowski, 1978), as well as a varietyof secondary succession and disturbed hab-itats such as pastures and cornfields, as wellas road-side vegetation.Etymology.—The specific epithet honors

Conrad Loddiges (1738–1826), a Germanhorticulturist who lived in London andcultivated American cycads.Nomenclatural notes.—The description

of Zamia loddigesii was lectotypified byStevenson & Sabato (1986). However, it isno longer permissible to designate a descrip-tion to serve as a type. Moreover, since thena specimen, Van Houtte 3374 at U, wasfound that was sent to Miquel by VanHoutte. This specimen predates the publica-tion of Z. loddigesii Miq. and matches thedescription. Thus, we are designating it thelectotype.Distinguishing features.—Zamia loddigesii

in contrast to Z. paucijuga, has been widelycollected along the Gulf of Mexico seaboardand on the Yucatan peninsula since the 19thcentury, largely by British and Russianbotanists, and recently by Mexican and NorthAmerican researchers. The high morphologi-cal variation presented by this species hasresulted in the publication of 10 affine names(Hill et al., 2007) that at times, has been abasis for the separation of natural popula-tions, i.e., that of the Yucatan peninsula(formerly Z. loddigesii sensu Vovides &Olivares, 1996) that is now a separate entityZ. polymorpha (Stevenson et al., 1995–1996b), a decision based on both vegetativeand reproductive characters that differ from Z.loddigesii mainly on pollen and ovulatestrobilus shape and indument color.Zamia loddigesii is similar to Z. polymor-

pha in leaf morphology; however, there aredifferences in the reproductive structures.Pollen cones of Z. loddigesii are beige incolor with a blunt apex whereas those of Z.polymorpha are dark reddish-brown or ma-roon with an acute apex. vulate cones of Z.loddigesii are cylindrical and beige in colorand those of Z. polymorpha are ovoid anddark maroon in color.

Additional specimens examined.—MEXICO.CHIAPAS: Ocozocoautla, M. A. Pérez-Farrera 81 (CHIP,HEM). HIDALGO: Atlapexco, San Juan 15 (XAL), 16(XAL), 17 (XAL), 18 (XAL), 19 (XAL), 20 (XAL), 21(XAL). PUEBLA: Sarukán et al. 4632 (FCME, MEXU).Oaxaca, Tuxtepec, J. Chamberlain s.n. (MO). TAMAULI-

PAS: R. L. Dressler 1858 (MO), Mayfiel et al. 791(MEXU); Aldama, F. Nicolalde-Morejón & J. González-Astorga 1585 (XAL), 1586 (XAL), 1587 (XAL), 1588(XAL). VERACRUZ: Chavelas et al. ES-4231 (MEXU),Dorantes et al. 964 (MEXU, MO), Dorantes et al. 1112(MEXU), Lot 733 (MEXU), Medrano et al. 2725(MEXU), Nevling & Gómez-Pompa 140 (MEXU),Santos 353 (XAL, XALU); Acayucan, Vovides et al.1376 (XAL), 1377 (XAL); Actopan, J. I. Calzada et al.6369 (MEXU, XAL), Lot 1027 (XAL), A. Vovides 754(XAL), 755 (XAL), 817 (XAL), 818 (XAL), 819 (XAL),820 (XAL), 821 (XAL), 822 (XAL), 823 (XAL), 824(XAL); Alto Lucero, J. Rees 1627 (XAL), 1629 (XAL),1630 (XAL), 1631 (XAL), 1632 (XAL), 1637 (XAL),Vovides 846 (XAL); Atoyac, Acevedo & Castillo-Campos 240 (XAL); Cotaxtla, González 82 (MEXU);Chicontepec, J. Rees 1615 (MEXU, XAL); Choapas,Vovides et al. 1373, 1374, 1735 (XAL); Coatepec, J.Rees & Vovides 1670 (XAL); Coatzacoalcos, Castillo &Acosta 16220 (XAL); Colipa, J. Rees 1634, 1635 (XAL);Cosautlan, Vovides 35 (XAL); Emiliano Zapata, J. Rees1763 (XAL), Stevenson et al. 538 (MEXU, NY), M.Vázquez-Torres 8071 (CIB); Huejultla, Stresser 291(MEXU); Hueyapan de Ocampo, Gómez-Pompa 4424(XAL), Vovides et al. 1378 (XAL), 1379 (XAL), 1380(XAL); Jalcomulco, Castillo & Zamora C. 7542 (XAL),Castillo & Gómez-Pompa 2588 (XAL), Castillo-Campos2727 (XAL); Mecayapan, Castillo et al. 13681 (XAL),13792 (XAL), 13843 (XAL), 13861 (XAL), 13865 (XAL),13866 (XAL), A. Calatayud & J. Martínez-Gándara 124(CIB); Moloacan, J. Rees 1656 (XAL); Puente Nacional,Castillo & Medina 4261 (XAL); Papantla, R. Cuevas et al.4652 (ZEA); Soteapan, Leonati 42 (MEXU); Soteapan, A.Calatayud & J. Benítez R 285 (CIB), M. A. Santos R. 352(CIB); Tampico Alto, Ortega & Ortega O. 2437 (XAL);Tezonapa, Robles 370 (XAL); Totutla, J. Rees 1661 (XAL);Yecuatla, J. Rees 1633 (XAL).

Zamia monticola Chamb., Bot. Gaz. 81:219. 1926. Type: cultivated from a singleseed collected opposite the crater of Nao-linco, near Xalapa, Veracruz, Mexico, Oct1925, C. J. Chamberlain s.n. (holotype:MO; isotype: NY).

Stem epigeal, up to 30 cm tall, 18–20 cm indiam. Cataphylls base triangular, apex linear-lanceolate, 3–6×1–2 cm wide at base. Leaves5–20, 100–200 cm long, erect to slightlycurved; petiole 50–75 cm long, terete, armedwith stout prickles in lower half; rachis terete,up to 100 cm long, with few prickles alongthe lower half. Leaflets 30–40 pairs, sessile,chartaceous to papyraceous, linear-lanceolate,


opposite to subopposite, subfalcate near thebase, apex long-acuminate and often stronglycurved, base attenuate, margins serrulate onlynear the base, subrevolute; articulations0.4–0.7 cm wide; the median leaflets 25–30×4–6 cm. Pollen strobili usually 2–6,erect, cylindrical to oblong, 12–20 cm longand 2–4 cm in diam, cream to light brown,apex acute; peduncle light brown tomentose,10–20 cm long; pollen sporangiophores cu-neiform, distal face hexagonal, 0.4 cm long,fertile abaxial surface 2-lobed with 10–16bisporangiate synangia per lobe. Ovulatestrobili unknown.Chromosome number.—UnknownEtymology.—The specific epithet alludes to

its mountainous habitat type, originallythought to be near Naolinco near the city ofXalapa, Veracruz in Mexico. It is now knownthat this species is not known from Mexicoand is endemic to Guatemala.Distribution and habitat.—Endemic to

Guatemala on rocky outcrops in primary andsecondary evergreen tropical rainforest (Fig. 6).Distinguishing features.—Characterized by

its consistently chartaceous to papyraceouslong acuminate leaflets that are stronglycurved near the apex with light serrulations.

Additional specimens examined.— GUATEMALA.ALTA VERAPAZ: H. Förther 2621/592 (NY).

Zamia onanreyesii C. Nelson & G. San-doval. Ceiba 49(1): 135. 2008. Type:Honduras. Cortés, 7 Jan 2008, O. Reyes406 (holotype: TEFH).

Zamia bussellii Schutzman, R. S. Adams, J. L.Haynes & Whitelock. The Cycad Newsletter 31(2/3), 22. 2008. Type: Honduras. Cortés, June2003, Whittington 2003/01 (holotype: FLAS).

Stem up to 2 m tall, up to 16 cm in diam.Ptyxis inflexed. Leaves 3–15(−44), 60–180×15–50 cm, erect to slightly curved, tomentu-lose when emerging; petiole 15–40 cm long,terete, sparsely to moderately armed withprickles; rachis terete, up to 40–120 cm long,with few prickles along proximal third. Leaf-lets up to 30 pairs, sessile, subcoriaceous,oblong-lanceolate, opposite to subopposite,long acuminate apically, base attenuate, mar-gins serrulate along distal third, subrevolute;articulations yellowish; median leaflets to

36×4 cm. Pollen strobili usually 1–3, erect,cylindrical, up to 27.5 cm long, up to 4 cm indiam, light brown to tan, apex acute; pedun-cle brown to tan, tomentose, up to 8.5 cmlong, up to 1.6 cm in diam; pollen sporan-giophore cuneiform, distal face hexagonaltruncate, fertile abaxial surface with up to 9bisporangiate sori per lobe. Ovulate strobiliusually solitary, erect, cylindrical, up to43 cm long, 12 cm in diam, brown togreenish, tomentulose, apex conical; peduncledensely brown tomentose, up to 5 cm long,up to 2.5 cm in diam, distal face hexagonal-truncate, 2.9 cm high, 4.7 cm wide. Seedsovoid, sarcotesta red at maturity, up to 3 cmlong, up to 2 cm in diam, sclerotesta smooth.Chromosome number.—Unknown.Distribution and habitat.—The species

range is Honduras (Fig. 6), between 0–1300 melevations in evergreen tropical forest.Etymology.—The specific epithet is in

honor of Onán Reyes, a Honduran biologist.Distinguishing features.—Stems arbores-

cent up to 2 m tall, leaflets coriaceous, longacuminate apically, with margin serrulate.Nomenclatural note.—Both Z. onanreyesii

and Z. busselllii were published in 2008. Thedates of issue are however different. The dateof issue for Z. onanreyesii is 6 September2008 and the date of issue for Z. bussellii is16 October 2008. Thus, under Article 29.1 ofthe Interantional Code of Botanical Nomen-clature (McNeill et al., 2007), Z. onanreyesiihas priority and is used here.

Additional specimen examined.—HONDURAS.Departamento Cortés: J. Haynes et al. 044A, 044B(TEFH).

Zamia oreillyi C. Nelson, Ceiba 46(1–2): 56.2005. Type: Honduras. Atlántida: 8 Apr2006, G. Sandoval et al. 1157 (holotype:TEFH). (Fig. 9)

Stem hypogeous, non-branching, up to25 cm long, up to 7.5 cm in diam. Leavesusually 1(2), up to 78.2 cm long, 31–35 cmwide, ascending to descending; petiole up to47 cm long, subterete, armed with smallprickles; rachis subterete, up to 35–40 cmlong, with few prickles along the proximalthird. Leaflets 29–31 pairs, sessile, papyra-ceous to sub-coriaceous, linear-lanceolate,opposite to subopposite, imbricate, apex


acuminate, base attenuate, margins dentate torarely entire along distal third, up to 0.3 cm,subrevolute; articulations brown in youngleaflets, 0.3–0.5 cm wide; the median leafletsup to 16 cm long, 1 cm wide. Pollen strobiliusually solitary, decumbent, cylindrical, up to2.5 cm long and 1 cm in diam, light brown,

apex acuminate; peduncle light brown tomen-tose, up to 15.9 cm long, 0.2 cm in diam;pollen sporangiophores cuneiform, distal facetruncate-hexagonal, 0.4 cm long, fertile ab-axial surface, with 2–3 bisporangiate synan-gia per lobe. Ovulate strobili unknown.Chromosome number.—Unknown

FIG. 9. Zamia oreillyi. A. Habit. B. Cataphyll. C–E. Leaflet variability. C. Broad and imbricate with dentatemargins. D. Narrow and imbricate with dentate margins. E. Narrow and slighly imbricate with entire margins. F.Ovulate strobilus. G. Pollen strobilus. H. Microsporophyll, abaxial view. (A-E, G-H from Balick 1711, NY).


Etymology.—The specific epithet is inhonor of Carlos Manuel O’Reilly, a Hon-duran biologist.Distribution and habitat.—Endemic to

Honduras, between 0–200 m in evergreentropical rainforest (Fig. 6).Distinguishing features.—Characterized by

strongly imbricate linear-lanceolate tooblong leaflets with dentate margins alongdistal third; pollen strobili usually solitary,decumbent, with fertile section up to 2.9 cmlong and decumbent peduncle up to 15.9 cmlong.

Additional specimen examined.—HONDURAS.ATLÁNTIDA. Balick 1711 (NY, TEFH).

Zamia paucijuga Wieland, American FossilCycads 2: 212. 1916. Type: Fig. 86 inAmerican Fossil Cycads 2: 212.1916.(lectotype, designated by Stevenson &Sabato, 1986: Fig. 86).

Stem hypogeous, branching dichotomouslywith age, 15–27 cm long, 8–13 cm in diam.Cataphylls coriaceous, persistent, base trian-gular, apex aristate, 4.5×3.4 cm at base,brown tomentose. Ptyxis inflexed. Leaves 2–3, ascending to descending 41–95×29–36 cmwide, brownish when emerging; petiole 10.2–32 cm long, green in young leaves, subterete,armed with prickles up to 4 mm long; rachissubterete, up to 56 cm long, with few pricklesalong the proximal third. Leaflets 5–28 pairssessile, coriaceous, lanceolate, opposite tosubopposite, apex acute, base attenuate, mar-gins serrulate to slightly dentate along distalthird, subrevolute; articulations brownish inyoung leaflets, 0.4–0.6 cm wide; the medianleaflets 14–19×2.3–3.4 cm. Pollen strobiliusually 1–2, erect, cylindrical, 6.3–11 cmlong, 2.1–2.6 cm in diam, light browntomentulose, apex acute; peduncle light-brown tomentose, 6.3 cm long, 1.2 cm indiam; pollen sporangiophores cuneiform, dis-tal face hexagonal-truncate, 0.3 cm long,fertile abaxial surface with 6–8 bisporangiatesynangia per lobe. Ovulate strobili usuallysolitary, erect, ellipsoid to cylindrical, 8.1 cmlong, 5.2 cm in diam, brown-yellowish,tomentulose, apex apiculate; peduncle dense-ly brown tomentose, 3.9 cm long, 1.3 cm indiam; megasporangiophores peltate, distal

face hexagonal-truncate, 0.6–0.8 cm high,1.1–1.5 cm wide. Seeds ovoid, sarcotestapink when immature, orange at maturity, 2–2.8 cm long, 1.5–1.7 cm in diam, sclerotestasmooth.Chromosome number.—2n=23, 34, 25, 26,

27, 28 (Moretti & Sabato, 1984).Distribution and habitat.—Endemic to

Mexico. Known from the states of Nayarit,Jalisco, Colima, Michoacan, Guerrero, andOaxaca (Fig. 6). Found in varied habitatsfrom dry open woodlands to understoryevergreen tropical forest.Etymology.—The specific epithet alludes to

few leaflet pairs per leaf.Distinguishing features.—Zamia paucijuga

is highly variable morphologically and inchromosome number and karyotype (Moretti,1990a, b). This may represent more than oneentity under the concept of Z. paucijugasensu stricto Wieland (1916). This speciesshares a number of characteristics with Z.loddigesii, being small plants with under-ground contractile stems and having coriaceousleaflets with marginal teeth. Nevertheless, itdiffers from Z. loddigesii in having highlycoriaceous leaflets with longer marginal teeth,and seeds with an orange sarcotesta at maturityas contrasted to the red sarcotesta of Z.paucijuga. Zamia paucijuga occurs along thePacific seaboard of Mexico whereas Z. loddi-gesii is known generally from the Gulf ofMexico seaboard.

Additional specimens examined.—MEXICO.COLIMA: McVaugh 15768 (FCME, MEXU). GUERRERO:

Acapulco, N. Noriega-Acosta 463, 546 (FCME), W.Thomas & J. L. Contreras 3744 (FCME); Chilpancingo,Kruse 902 (FCME, MEXU), R. M. Fonseca 1210(FCME), F. Nicolalde-Morejón et al. 1566, 1567, 1568,1569 (XAL), José Azueta, Vovides et al. 1426 (XAL);Petatlán, Vovides et al. 1427, 1428, 1429, 1430, 1431,1432, 1433, 1434 (XAL); Unión de Isidro Montes DeOca, Vovides et al. 1416, 1417, 1418, 1420, 1421 (XAL);La Unión, G. Lozano-Valdez 331 (FCME), J. Jiménez331 (FCME). JALISCO: Gómez-Pompa 4876 (MEXU); ElArenal, Castillo et al. 9822 (XAL); Cabo Corrientes,Castillo et al. 10147, 11733, 10280, 10466 (XAL), J.Ceja et al 1437 (UAMIZ), J. Ceja et al. 1470 (UAMIZ),F. Nicolalde-Morejón et al. 1524, 1525 (XAL); Cihuat-lán, J. Borocio R. s.n. (ZEA); Cuautitlán, Cochrane et al.10886 (IBUG, WIS, ZEA), R. Cuevas et al 7025 (ZEA),L. Guzmán & J. Santana M. 745, 947 (ZEA), F.Nicolalde-Morejón et al. 1528, 1529 (XAL), Pérez dela Rosa 1039 (FCME, IBUG, MEXU), 1040 (FCME,IBUG, XAL), 1041 (CIB), 1518 (IBUG), Ramírez 425(IBUG), M. Rosales & L. Cruz 75 (ZEA), J. Santana M.


et al. 5296 (ZEA); La Huerta, Cuevas et al. 4861 (IBUG,ZEA); San Sebastián, F. Nicolalde-Morejón et al. 1422(XAL), 1423 (XAL), 1424 (XAL), 1425 (XAL), 1426(XAL), 1427 (XAL), Pérez de la Rosa 1084 (IBUG,FCME, MEXU) 1097, 1098 (IBUG); Tuito, A. Flores etal. 614 (UAMIZ), F. Nicolalde-Morejón et al. 1429(XAL), Pérez de la Rosa 1438, 1439 (IBUG); Vallarta,Pérez de la Rosa 1413, 1415 (IBUG); Villa Purificación,Pérez de la Rosa 1885. (IBUG). NAYARIT: Gentry &Gilly 10496 (MEXU), McVaugh 19211 (FCME, MEXU),Vovides et al. 1487 (XAL), 1488 (XAL), 1489 (XAL),1490 (XAL), 1491 (XAL), 1493 (XAL); Compostela, F.Nicolalde-Morejón et al. 1521 (XAL), 1522 (XAL),1523 (XAL); Tepic, R. Dressler 1026 (MO), H. S. Gentry& C. I. Gilly 10498 (FCME). Oaxaca: Miranda 4205(MEXU), F. Nicolalde-Morejón et al. 1465 (XAL), 1466(XAL), 1467 (XAL), 1468 (XAL), 1469 (XAL), 1470(XAL), 1471 (XAL), 1472 (XAL), 1473 (XAL); Pochutla,Schutzman 543 (XAL), 544 (XAL), 545 (XAL), 546 (XAL),547 (XAL), 548 (XAL), 550 (XAL), 551 (XAL), 552 (XAL),553 (XAL), 554 (XAL), 555 (XAL), 556 (XAL), 557 (XAL),558 (XAL), 560 (XAL), 561 (XAL), 562 (XAL), 563 (XAL),565 (XAL), A. Nava-Zafra & J. Pascual 35 (SERO, FCME);Puerto Escondido, J. Rees 1603 (MO, XAL), Walters sn(FTG accession 7–14, XAL); San Gabriel Mixtepec, F.Nicolalde-Morejón et al. 1474 (XAL); San Pedro Pochulta, J.Lomelí et al. 2967 (MEXU).

Zamia polymorpha D.W. Stev., A. Moretti& Vázq. Torres, Delpinoa n.s. 37–38: 4.1995–1996 (issued 1998). Type: Belize.Cayo: 22 Jan 1989, D. W. Stevenson et al.1119 (holotype: NY; isotypes: BRH, FTG,MO, NY, U). (Fig. 10)

Stem hypogeous, branching dichotomouslywith age, up to 32 cm long, up to 14 cm indiam. Cataphylls chartaceous, persistent, basetriangular, apex aristate, 3–6.4×1.3–2.6 cm atbase, brown tomentose. Ptyxis inflexed.Leaves 2–3(4), ascending, 30–105×29–45 cm, brown when emerging, green whenmature; petiole 10.2–95.3 cm long, greenish inyoung leaves, subterete, armed with pricklesup to 4 mm long; rachis subterete, up to 67 cmlong, with few prickles along the proximalthird. Leaflets 3–12 pairs, sessile, coriaceous,lanceolate to oblanceolate, opposite to sub-opposite, apex acute, base attenuate, marginsserrulate along upper third, subrevolute; artic-ulations brown when young, green whenmature, 0.4–0.8 cm wide; the median leaflets17–35×2–3.5 cm. Pollen strobili usually 1–2,erect, conical, 6.5–7.3 cm long, 1.1–1.4 cm indiam, light to dark-brown tomentulose, apexacute; peduncle light brown tomentose, 6.8 cmlong, 1.2 cm in diam; pollen sporangiophorescuneiform, distal face hexagonal-truncate,

0.4 cm long, fertile abaxial surface with 4–5bisporangiate synangia per lobe. Ovulatestrobili usually solitary, erect, cylindrical toovoid, 8.7–16.3 cm long, 5–8.3 cm in diam,dark-brown tomentulose, apex acute; pedunclebrown tomentose, 4–7.5 cm long, 1.1 cm indiam; megasporangiophores peltate, distal facehexagonal-truncate, 0.8 cm high, 1.8–2.1 cmwide. Seeds ovoid, sarcotesta pink whenimmature, red at maturity, 1.4–2.1 cm long,0.5–0.9 cm in diam, sclerotesta smooth.Chromosome number.—2n=17, 22, 23, 24,

25, 26, 27, 28 (Stevenson et al., 1995–1996b,b; Vovides & Olivares, 1996).Distribution and habitat.—This species is

known from Belize, Guatemala and Mexicofrom 0–200 m elevation. In Mexico, its rangeincludes the states of Quintana Roo, Yucatán,Campeche, Tabasco and Chiapas (Fig. 6).Etymology.—The specific epithet alludes to

the extreme variation in leaf and leaflet mor-phology presented by this species (Stevensonet al., 1995–1996b).Distinguishing features.—Zamia polymor-

pha shares many morphological attributeswith Z. loddigesii of the Gulf of Mexicodrainage. However, there are clear differencesin the pollen and ovulate reproductive struc-tures. In Z. loddigesii, the pollen strobili arebeige in color with an acute apex and theovulate strobili are beige and cylindricalwhereas in Z. polymorpha the pollen strobiliare maroon with an acuminate apex and theovulate strobili are dark-maroon and ovoid.

Additional specimens examined.—MEXICO.CAMPECHE: Hernández et al. ES-184 (MEXU), Schutz-man 502 (XAL), 503 (XAL), 504 (XAL), 505 (XAL), 506(XAL), 507 (XAL), 508 (XAL), 509 (XAL); BenitoJuárez, Vovides et al. 1312 (XAL), 1313 (XAL), 1314(XAL), 1315 (XAL), 1316 (XAL), 1317 (XAL); Calakmul,Madrid et al. 736 (MEXU), Martínez 30420-A (MEXU);Ciudad del Carmen, Flores et al. 9586 (XAL); Champoton,Chan 3719 (CICY, XAL), Vovides 853 (XAL), 854 (XAL),855 (XAL), 1326 (XAL), 1328 (XAL), 1329 (XAL),1330 (XAL), 1331 (XAL), 1332 (XAL), 1333 (XAL), 1334(XAL), 1335 (XAL), 1336 (XAL), 1337 (XAL), 1338(XAL), 1339 (XAL), 1527 (XAL); Hopelchen, Ortega &Ucán 1562 (UADY, XAL), Ucán et al. 7293 (UADY,XAL), 7307 (UADY, XAL), 7398 (UADY, XAL).CHIAPAS: Palenque, Aguilar & Aguilar 1355 (MEXU), F.Nicolalde-Morejón & N. Martínez 1419 (XAL), Schutzman508 (XAL), Walters s.n. (FTG accession 13-2, XAL);Ocosingo, Walters s.n. (FTG accession 17-2, XAL).QUINTANA ROO: Cabrera et al. 2574 (MEXU), Davidse etal. 20075 (MEXU, MO), Téllez 1415 (MEXU), Trejo 225(CICY, MEXU); Adolfo Huerta, Álvarez et al. 9495


(MEXU); Chetumal, Vovides 852 (XAL), Flores & Burgos9635 (XAL), 9643 (XAL), Othon P. Blanco, Vovides et al.1318 (XAL), 1319 (XAL), 1320 (XAL), 1321 (XAL), 1322(XAL), 1323 (XAL), 1324 (XAL), 1325 (XAL). TABASCO:

Balancan, Matuda 3117 (MEXU), Méndez 214 (XAL),Novelo 169 (MEXU, XAL), Puig 788 (MEXU). Macu-spana, Vovides et al. 1344, 1345 (XAL). Yucatán, G. F.Gaumer 2430 (MO), Lundell & Gentle 827 (MEXU),May743 (CICY, MEXU); Tekon, Enríquez 94 (MEXU);

Tzucacab, Vovides et al. 1303 (XAL), 1306 (XAL), 1307(XAL), 1308 (XAL), Flores & Burgos 9642 (XAL);Valladolid, Vovides 856, 857 (XAL), Vovides et al. 867(XAL), 868 (XAL), 869 (XAL) 871 (XAL), 872 (XAL),873 (XAL), 874 (XAL), 875 (XAL), 881 (XAL), 877(XAL), 870 (MEXU, XAL), 876 (MEXU, XAL), 877(MEXU, XAL), 878 (MEXU, XAL), 880 (MEXU, XAL);Yaxcaba, Vovides et al. 1309 (XAL), 1310 (XAL), Vovides1311 (XAL).

FIG. 10. Zamia polymorpha. A. Habit (Stevenson et al. 1118, NY). B. Leaflets (from the holotype). C. Ovulatestrobilus. D. Pollen strobilus. E. Cataphyll.


GUATEMALA. PETEN: W. E. Harmon & J. A.Fuentes 5735 (MO).

BELIZE. BELIZE DISTRICT: Estrada 234 (CICY), D.L. Spellman 1548 (MO), C. Whitefoord 2603 (MO).CAYO DISTRICT: D. W. Stevenson et al. 1121 (FTG, MO,NY, U), 1122 (FTG, MO, NY, U), M. J. Balick et al.1803 (MO, NY), M. J. Balick et al. 2058 (NY), T. B.Croat 23732 (MO), D. R. Hotel & L. Thomas 1130(NY), J. S. Huston s.n. (MO), R. W. Long 3238 (MO), J.A. Ratter 5195 (MO), D. L. Spellman 1974 (MO), D. L.Spellman & W. W. Newey 1962 (K), J. R. Wiley 333(MO). Orange Walk, G. Davidse & A. E. Brant 32768(MO). Stann Creek, R. L. Walter 1099 (MO). Sin datos(K).

Zamia prasina W. Bull, Retail List: 20.1881. Type: cultivated from Br. Honduras,W. Bull s.n. (holotype: K).

Stem hypogeous to epigeous, rarelybranching. Cataphylls chartaceous, persistent,base triangular, apex aristate, 3–4.5×1.2–2.7 cm at base, brown tomentose. Ptyxisinflexed. Leaves 2–4(6), 57–100×25–35 cm,ascending to spreading, brown when emerg-ing; petiole 12–30 cm long, brown-greenishyoung leaves, subterete, sparsely to denselyarmed with prickles up to 4 mm long; rachissubterete, up to 70 cm long, with few pricklesalong the proximal third. Leaflets 12–18pairs, sessile, coriaceous, oblong to oblanceo-late, opposite to subopposite, apex acute toacuminate, base cuneate, margins serrate todenticulate in the upper two third, subrevo-lute; articulations brown-yellowish in youngleaflets, 0.3–0.6 cm wide; the median leaflets15–20×4–6 cm. Pollen strobili usually 1–2,erect, cylindrical to ovoid, up to 6–10 cmlong and 2–4 cm in diam, light brown, apexacute; peduncle densely light brown tomen-tose, 2–4 cm long, 1.5 cm in diam. Ovulatestrobili usually solitary, erect, cylindrical, 10–15 cm long, 5–7 cm in diam, green, glabrouswhen mature, apex acute; peduncle brown-greenish tomentose, 3.5 cm long, 1.5 cm indiam; megasporangiophores distal face hex-agonal-truncate, 1–1.4 cm high, 1.1–1.8 cmwide. Seeds ovoid, sarcotesta light red whenimmature, red at maturity, 1.5–2 cm long,0.5–0.8 cm in diam, sclerotesta smooth.Chromosome number.—Not known.Distribution and habitat.—Endemic to

Belize (Fig. 6) on rocky outcrops between100–200 m elevation in evergreen tropicalrainforest.

Etymology.—The specific epithet alludes tothe bright grass-green leaflets.Distinguishing features.—Plants with sub-

terranean or epigeal stems, distinctly serrulateleaflet margins and bright-green leaflets;ovulate strobili ovate, green and glabrouswhen mature.

Additional specimens examined.—BELIZE.TOLEDO: G. Davidse & A. E. Brant 32179 (MO),32232 (MO).

Zamia purpurea Vovides, J.D. Rees & Vázq.Torres, Flora de Veracruz 26: 28. 1983.Type: Mexico. Veracruz: 30 Jan 1982,Vovides 734 (holotype: XAL). (Fig. 11)

Stem hypogeous, dichotomously branchingwith age, up to 30 cm long, 4–6 cm in diam.Cataphylls membranaceous, deciduous, basetriangular, apex acuminate, 5.×1.8 cm atbase, reddish-brown tomentose. Ptyxis in-flexed. Leaves 1–6, ascending to spreading,34–90×32–38 cm, reddish-brown whenemerging, turning green to dark-green atmaturity; petiole 16–29 cm long, blackish inyoung leaves, subterete, armed with simpleprickles up to 4 mm long; rachis subterete,17–45 cm long, with few prickles along theproximal third. Leaflets 3–4(6) pairs, sessile,coriaceous, elliptic to lanceolate, opposite tosubopposite, veins highly prominent on ad-axial surface, though relatively inconspicuousveins have been reported on some individu-als, apex acute, base attenuate, marginsdentate along upper third, subrevolute; artic-ulations brown in young leaflets, 0.4–0.8 cmwide; the median leaflets 6–27×2–8.1 cm.Pollen strobili usually 1–2, erect, conical, 2–4.3 cm long, 0.5–1.1 cm in diam, light-browntomentulose, apex acute; peduncle light-brown tomentose, 2–3.5 cm long, 0.7–0.9 cm in diam; pollen sporangiophorescuneiform peltate, distal hexagonal face andtruncate to scutiform, 0.25 cm long, fertileabaxial surface with 2 bisporangiate synangiaper lobe. Ovulate strobili usually solitary,erect, conical, 6–9 cm long, 3–4.5 cm indiam, purplish-brown tomentulose when im-mature turning dark-purple glabrescent whenmature, apex acute; peduncle densely dark-brown tomentose, 2.9 cm long, 1.6 cm indiam, distal face hexagonal-scutiform, 0.9–


FIG. 11. Zamia purpurea. A. Habit. B. Pollen strobilus. C. Microsporophyll, abaxial view. D. Ovulate strobilus. E.Ovulate sporangiophores. F. Seed. (From Rees1 654, XAL.)


1.2 cm high, 1.6–2.1 cm wide. Seeds ovoid,sarcotesta pink when immature, red at matu-rity, 0.8–1.1 cm long, 0.6–0.8 cm in diam,sclerotesta smooth.Chromosome number.—2n=16 (Vovides,

1983)Diversity and genetic structure.—The

average of alleles per locus is A=2.10, thepercentage of polymorphic loci is P=100,the expected heterozygosity is HE=0.481 andthe genetic differentiation between the twopopulations currently under study is Fst=0.037 (González-Astorga et al. unpubl. data).Distribution and habitat.—Zamia pur-

purea is endemic to Mexico from the RíoUxpanapa drainage system within the statesof Veracruz and Oaxaca (Fig. 5) between 50–200 m in evergreen tropical forest (sensuRzedowski, 1978) classification.Etymology.—The specific epithet alludes to

the dark purple color of the ovulate strobili.Distinguishing features.—Characterized by

the prominent leaflet veins giving the coria-ceous leaflets a channeled appearance incontrast to the rest of its congeners inMesoamerica, and the dark purple color ofthe ripe ovulate strobili.

Additional specimens examined.—MEXICO.OAXACA: Santa María Chimalapa, S. H. Salas M. 982(SERO), Sánchez et al. 40 (B, MEXU); Santa MaríaLachixio, Cerón et al. 266 (XAL); San Juan Guichicovi, F.Nicolalde-Morejón & J. Torres 1503 (XAL), 1404 (XAL),M. Vázquez-Torres et al. 1470 (MO, XAL), Walters s.n.(FTG accession 10–1, XAL). VERACRUZ: M. Vázquez-Torres 4038 (CIB); Hidalgotitlán, Calzada 8374 (XAL),M. Vázquez-Torres et al. 224 (MO); Jesús Carranza, A.González-Christen s.n. (CIB), F. Nicolalde-Morejón & J.Torres 1502 (XAL), M. Vázquez-Torres et al. 2373 (CIB),M. Vázquez-Torres et al. V-2532 (CHAPA, CIB, XAL).

Zamia sandovalii C. Nelson, Ceiba 46(1–2):55.. 2005. Type: Honduras. Atlántida: Jan2006, G. Sandoval et al. 1154 (holotype:TEFH)

Stem hypogeous, non-branching, 15.7 cmlong, 10.1 cm in diam. Cataphylls persistent,base triangular. Leaves 1–3, ascending todescending up to 210×0.55 cm; petiole 68 cmlong, subterete, armed with small prickles;rachis subterete, up to 140 cm long, unarmed.Leaflets 68 pairs sessile, sub-coriaceous, lance-olate, opposite to subopposite, falcate, apexacuminate, base attenuate, margins serrulate

along distal third, subrevolute; the median leaf-lets 16–31×2–3 cm. Pollen strobili usually 2,erect, cylindrical, 11.2 cm long, 2.6 cm in diam,brown-reddish tomentulose, apex apiculate; pe-duncle light-brown tomentose, 16.7 cm long,1.2 cm in diam; pollen sporangiophores cunei-form, distal face hexagonal truncate, 0.8 cmlong, fertile abaxial surface with 2–3 bisporan-giate synangia per lobe. Ovulate strobili usuallysolitary, erect, ellipsoid to cylindrical, 12 cmlong, 4.5 cm in diam, brown-yellowish, tomen-tulose, apex long-acuminate; peduncle denselybrown tomentose, 1.5 cm long; megasporangio-phores peltate, distal face hexagonal-truncate,2–2.5 cm high, 1.8–2 cm wide. Seeds ovoid,sarcotesta white when immature, 1.2–1.8 cmlong, 0.5–0.7 cm in diam, sclerotesta smooth.Chromosome number.—UnknownDistribution and habitat.—Endemic to

Honduras (Fig. 6), between 200–350 m inevergreen tropical forest.Etymology.—The specific epithet honors

Germán Sandoval, biologist of the UniversidadNacional Autonoma de Honduras (Nelson,2005).Distinguishing features.—This species is

characterized by glabrous, sub-coriaceous,lanceolate, falcate leaflets; cylindric, ovulatestrobili with a strongly acuminate apex; andseeds with a white sclerotesta.

Additional specimens examined.—HONDURAS.ATLÁNTIDA: J. Haynes et al. 37 (TEFH), G. Sandovalet al. 1155 (TEFH), 1156 (TEFH).

Zamia soconuscensis Schutzman, Vovides &Dehgan, Bot. Gaz. 149(3): 347. 1988. Type:Mexico. Chiapas: Feb 1939, Matuda 2659(holotype: F; isotypes: CR, MEXU,MICH).

Stem epigeal, erect to decumbent in adultplants, branching dichotomously with age,30–65 cm long, 10–31.5 cm in diam. Cata-phylls chartaceous, persistent, base triangular,apex aristate, 7.1×2.6 cm at base, reddish-brown tomentose. Ptyxis inflexed to erect.Leaves 3–15 or more per apex, 120–190×45–62 cm, ascending, distal portion descendingto spreading, brown when emerging turninggreen at maturity; petiole 38–72 cm long,green-yellowish in young leaves, terete,armed with prickles up to 5 mm long; rachissubterete, up to 84 cm long, with few prickles


along the proximal third. Leaflets 41–52pairs, sessile, coriaceous, linear-lanceolate,alternate to subopposite, subfalcate, apexacute, base attenuate; margins entire, subre-volute; articulations brown in young leaflets,0.4–0.8 cm wide; the median leaflets 12–35×0.6–1.5 cm. Pollen strobili usually 1–3 perapex, erect, cylindrical to conical, 9–15 cmlong, 1.2–2.4 cm in diam, dark-brown tomen-tulose, apex apiculate; peduncle light-browntomentose, up 7.2 cm long, 1.2 cm in diam;pollen sporangiophores cuneiform, distal facehexagonal-truncate, 0.45 cm long, fertileabaxial surface with 5 bisporangiate synangiaper lobe. Ovulate strobili usually solitary,erect, cylindrical, 12–15 cm long, 6.1–7.3 cmin diam, dark-brown to reddish, tomentulose,apex aristate; peduncle brown puberulent,2.1 cm long, 1.2 cm in diam; megasporan-giophores peltate, distal face hexagonal-truncate, 0.7–0.9 cm high, 1.6–1.8 cm wide.Seeds ovoid to angular, sarcotesta white whenimmature turning salmon-pink when mature,sclerotesta light-beige, smooth with 6–8 lightfurrows running longitudinally and some-times dichotomizing, 2.1–2.6 cm long, 1.4–1.9 cm diam.Chromosome number.—2n=16 (Schutzman

et al., 1988).Distribution and habitat.—Endemic to

Chiapas, Mexico, between 900–1400 m inthe Soconusco mountain range of southernChiapas. It inhabits the understory herbaceouslayer of the transition zone between evergreentropical forest and cloud forest (Fig. 5).Etymology.—The specific epithet alludes to

the Sierra del Soconusco mountain range alsoknown as the Sierra Madre de Chiapas, beingthe region where this species is native(Schutzman et al., 1988).Distinguishing features.—Zamia soconus-

censis is the only species of the genus inMexico that approaches an arborescent habitwith leaves that can reach almost two meterslong that gracefully arch toward the terminalportion. The linear-lanceolate leaflets havetotally entire margins; the only other Mexicancongener with entire margins is Z. inermis.Ovulate strobili are short-pedunculate and sol-itary with a dark-brown velvety tomentulum.

Additional specimens examined.—MEXICO.CHIAPAS: Matuda 2087 (MEXU), 2535 (MEXU), 2590

(MEXU), 2656 (MEXU); Acacoyagua, García 149(CHIP, MEXU), M. A. Pérez-Farrera 141 (CIB, HEM,MEXU).

Zamia spartea A. DC., Prodr. 16 (2): 539.1868. Zamia loddigesii var. spartea (A.DC.) Schuster, Pflanzenr. 99: 148. 1932.Type: Mexico. Oaxaca: prope Acayucam,Verapa, Chimalapi, 1832, Alaman s.n.(holotype: G-DC). (Fig. 12).

Stem hypogeous, branching dichotomouslywith age, 5–40 cm long, 5–8 cm in diam.Cataphylls chartaceous, persistent, base trian-gular, apex aristate, 6×1.4 cm at base,yellowish tomentose. Ptyxis inflexed. Leaves2–5(8) per crown, 35–60×38–52 cm, ascend-ing to gracile, reddish-brown when emerging,turning green at maturity; petiole 12–21 cmlong, green-yellowish in young leaves, sub-terete, heavily armed with straight to some-times bifurcate prickles up to 4 mm long;rachis subterete, up to 42 cm long, with fewprickles along the proximal third. Leaflets15–27 pairs sessile, coriaceous, linear, alter-nate to subopposite, apex acute, base attenu-ate; margins serrulate along extreme distalportion, subrevolute; articulations light-orange when young, turning yellowish withage, 0.3–0.4 cm wide; the median leaflets 20–35×0.3–0.6 cm. Pollen strobili usually 2–3,erect, cylindrical, 6.5–8.5 cm long, 1.4–1.9 cm in diam, yellowish-beige tomentulose,apex acute; peduncle densely light-browntomentose, 6–8 cm long, 0.9–1.1 cm in diam;pollen sporangiophores cuneiform, distal facehexagonal-truncate, 0.3 cm long, fertile ab-axial surface with 10–14 bisporangiate syn-angia per lobe. Ovulate strobili usuallysolitary, erect, cylindrical to oval-cylindrical,9–12 cm long, 4.2–4.8 cm in diam, browntomentulose, apex acute; peduncle denselybrown tomentose, 3.4–3.8 cm long, 0.8–1.1 cm in diam; megasporangiophores peltate,distal face hexagonal-truncate, 0.8–1.1 cmhigh, 1–1.3 cm wide. Seeds ovoid, sarcotestapink when immature, red at maturity, 1.2–1.6 cm long, 0.7–0.9 cm in diam, sclerotestasmooth.Chromosome number.—2n=18 (Vovides,

1983; Moretti, 1990a).Distribution and habitat.—Endemic to

Oaxaca, Mexico. Known in southern areas of


FIG. 12. Zamia spartea. A. Habit. B. Leaflet. C. Pollen strobilus. D–E. Microsporophyll, abaxial and adaxialview. F–G. Petiole and cataphyll. H. Ovulate strobilus and peduncle. I. Ovulate sporangiophore. J. Ovulatesporangiophores, distal end truncate hexagonal. K. Seed with sarcotesta. L–N. Seeds.


the Isthmus of Tehuantepec (Fig. 5), between200–400 m elevation, associated with tropicaldeciduous forests (sensu Rzedowski, 1978).Etymology.—The epithet is from the broom

genus Spartium (Fabaceae), in reference tothe narrow and tapered leaflets of the cycad.Distinguishing features.—Small gracile

plants reaching up to 80 cm tall with verynarrow-linear leaflets with an almost entiremargin with a few very small serrulationsonly on the extreme distal portion.

Additional specimens examined.—MEXICO.OAXACA: Meave del Castillo & García 2388 (MEXU);Matías Romero, F. Nicolalde-Morejón & J. Torres 1505(XAL), 1506 (XAL), Schutzman 529 (XAL), 530 (XAL),531 (XAL), 532 (XAL), 533 (XAL), 534 (XAL), 535(XAL), 536 (XAL), 537 (XAL), 538 (XAL), 539 (XAL),540 (XAL), 541 (XAL), 542 (XAL), Vovides & Perales600 (XAL), Walters s.n. (FTG accession 9–3, XAL); SanJuan Guichicovi, N. Antonio-Barrera 83b (CIB), SantaMaría Chimalapa,A. Espejo et al 6485 (UAMIZ),R. GarcíaS. 341 (XAL, SERO), Torres 653 (XAL); San MiguelChimalapa,M. Vázquez-Torres 4039 (CIB).

Zamia standleyi Schutzman, Syst. Bot. 14(2): 214. 1989. Type: Honduras. Atlantida:Lanatilla Valley near Tela, Aug 1984, B.Schutzman 449 (holotype: FLAS; isotypes:ENA, FTG).

Stem hypogeous and tuberous, 6–14 cmlong, 5–9 cm in diam. Cataphylls chartaceous,persistent, base triangular, apex long-aristate,to 12 cm×1.5 cm at base, reddish-browntomentose. Ptyxis inflexed. Leaves 1–5, 20–100× to 55 cm, slightly to slightly recurved,tomentulose when emerging; petiole 35–60 cmlong, terete, sparsely to heavily armed withprickles; rachis terete, up to 70 cm long, withfew prickles along proximal third. Leaflets 10–15 pairs, sessile, subcoriaceous-coriaceous,long-lanceolate, opposite to subopposite, re-curved, apex acute, base attenuate, marginsdentate along distal third, subrevolute; articu-lations dark brown when young, 0.3–0.5 cmwide; median leaflets 20–45×1–4 cm. Pollenstrobili usually 1–3, decumbent, cylindrical,6–10 cm long, 1–2 cm in diam, light-browntomentulose, apex acute; peduncle light-browntomentose, 2–4 cm long, 1.1 cm in diam;pollen sporangiophores cuneiform, distal facehexagonal truncate, fertile abaxial surface with4 bisporangiate sori per lobe; Ovulate strobiliusually solitary, erect, cylindrical to slightly

ovoid, 8–12 cm long, 3–8 cm in diam, brown,tomentulose, apex long-apiculate; peduncledensely brown tomentose, 2.5–4 cm long,1.3 cm in diam, distal face hexagonal-truncate,0.7–1.2 cm high, 1.5–2.1 cm wide. Seedsovoid, sarcotesta pink when immature, red atmaturity, up to 3 cm long, up to 2 cm in diam,sclerotesta smooth.Chromosome number.—2n=16 (Schutzman,

1989).Distribution and habitat.—The species

range is Honduras and Guatemala (Fig. 6),between 0–200 m in evergreen tropical forest.Etymology.—The specific epithet honors

Paul C. Standley a prominent botanist ofthe flora of Mexico and Central America(Schutzman, 1989).Distinguishing features.—Its falcate leaflets

and conspicuous marginal teeth up to 4 mmlong, and long-apiculate ovulate strobili aswell as its cylindrical characterize this species.

Additional specimens examined.—GUATEMALA.PROVINCIA IZABAL: H. Förther 10234/252 (MSB, W).

HONDURAS. DEPARTAMENTO ATLÁNTIDA: Tela, V.Severen 1450 (NA), Standley 53721 (F, US); PuertoSierra, Wilson 537 (NY). DEPARTAMENTO SANTA BAR-

BARA: San Pedro Sula, Thieme 144 (US). DepartamentoYoro, Coyoles, T. G. Yuneker et al. 8186 (F, G, GH, MO,NY). DEPARTAMENTO CORTES: Montaña Santa Ana,Molina R. 3628 (F, GH).

Zamia tuerckheimii Donn. Sm., Bot. Gaz.(Crawfordsville) 35(1): 8. 1903. Type:Guatemala. Dept. Alta Verapaz: Cubilquitz,Jul 1900, von Tuerckheim 7786 (holotype:US; isotype: K).

Stem epigeal, rarely branched with age, upto 100 cm long, 10–12 cm in diam. Cata-phylls chartaceous, persistent, base triangular,apex aristate, 8.3×2.7 cm at base, lightbrown-greenish tomentose. Ptyxis inflexed toerect. Leaves 8–15, 100–200×20–45 cm,ascending, distal portion descending tospreading, green when emerging; petiole 30–50 cm long, green in young leaves, terete,armed with minute prickles to unarmed;rachis subterete, up to 150 cm long, unarmed.Leaflets 8–15 pairs, sessile, papyraceous,oblong-lanceolate, glossy, iridescent blue-green, alternate to subopposite, apex abruptlyacuminate, base attenuate; margins entire, sub-revolute; articulations green, 0.4–0.6 cm wide;


the median leaflets 14–18×4–6 cm x. Pollenstrobili usually solitary, erect, cylindrical, 12–15 cm long, 2–4 cm in diam, gray-brown atmaturity, apex acute; peduncle light-browntomentose, up 6.5 cm long, 1.4 cm in diam.Ovulate strobili usually solitary, erect, cylindri-cal, 12–18 cm long, 4–8 cm in diam, brownishwhen young, iridescent blue-green at maturity,apex aristate; peduncle light brown puberulent,3.4 cm long, 1.5 cm in diam; megasporangio-phores peltate, distal face hexagonal-truncate,1.4–2 cm high, 3–3.6 cmwide. Seeds elongate-ovoid, sarcotesta light red when immature, redat maturity, smooth, 2.1–2.6 cm long, 1.3–1.6 cm diam.Chromosome number.—2n=16 (Moretti,

1990b)Distribution and habitat.—Endemic to

Guatemala, between 250–1000 m in the AltaVerapaz department mountain range. It inhab-its the understory herbaceous layer of tropicalforest (Fig. 6).Etymology.—The specific epithet honors

Hans von Tüerkheim, who collected thisspecies for the first time in Guatemala.Distinguishing features.—The principal

distinguishing features of this species arearborescent stems up to one meter tall,papyraceous leaflets with entire margins andovulate cylindrical strobilus with abruptlyacuminate apex and aristate apex, the conebecoming blue-green iridescent when mature.

Additional specimens examined.—GUATEMALA.ALTA VERAPAZ: T. B. Croat 41647 (MO), H. Förther 11034(MSB, W), J. A. Steyermark 44484 (MO); Rubeltem, H.Förther 10918 (MSB, W).

Zamia variegata Warsz., Allg. Gartenzeitung32: 253. 1845. Type: Mexico. Chiapas:Lacandona, on border with Guatemala, 12 Jan1987 D. W. Stevenson 685 (neotype, heredesignated: NY; isoneotypes: U, XAL).

Zamia muricta var. picta Miquel, Tijdschr. Wis-ennatuurk. Wet.1(4): 198–199. 1848, non Von Houtte1846.Zamia pictaDyer inHemsley, Biol. Cent.-Amer.,Bot. 3(16): 194. 1884. Type: ex Horto Amsterdam(Z. picta H. Belg.),Miquel s.n. (holotype, U)

Stem hypogeous, 9–16 cm long, 4–10.5 cmin diam. Cataphylls chartaceous, base trian-gular, apex aristate, 4.6–7.1×2.1–2.4 at base,yellowish tomentose. Ptyxis inflexed. Leaves1–2(3), 40–291×24–44 cm, ascending; peti-

ole 35–177 cm long, dark-green with charac-teristic yellow variegation; petiole subterete,heavily armed with prickles up to 5 mm long;rachis subterete, up to 105 cm long, with fewprickles along the proximal third. Leaflets 3–10 pairs, sessile, papyraceous, elliptic, oppo-site to subopposite, dark-green with yellow orcream variegation, apex acute, base attenuate;margins dentate along distal third, subrevo-lute; articulations brown in young leafletsturning green at maturity, 0.4–0.8 cm wide;the median leaflets 12–22×3.1–8.8 cm. Pol-len strobili up to 6, erect, long-cylindrical,7–11 cm long, 1.9–2.5 cm in diam, yellow-ish-beige, apex acute; peduncle densely light-brown tomentose, 3–6 cm long, 0.7–0.9 cmin diam. Ovulate strobili 1–2 ovoid tocylindrical, 12 cm long, 4.5 cm in diam,gray-greenish tomentulose; megasporangio-phores peltate, distal face hexagonal-truncate,0.8–1.3 cm high, 1.7–2.1 cm wide. Seedsovoid, red at maturity, 1.1–1.5 cm long, 0.7–1 cm in diam, sclerotesta smooth.Chromosome number.—2n=21, 22 (Moretti

et al., 1991).Diversity and genetic structure.—The

average of alleles per locus is A=2.02, thepercentage of polymorphic loci is P=97.3,the expected heterozygosity is HE=0.355 andthe genetic differentiation between the twopopulations currently under study is Fst = 0.085(González-Astorga et al. unpubl. data).Distribution and habitat.—Zamia varie-

gata was described from plants collected inGuatemala by Warszewicz (Stevenson &Sabato, 1986). In Mexico, this plant isknown only from Chiapas, in lowland areasnear the Montes Azules Biosphere Reserve(Fig. 5). Its habitat is evergreen tropicalrainforest (sensu Rzedowski, 1978).Etymology.—The specific epithet alludes to

the variegated nature of the leaflets, anattribute unique to this species of Zamia.Distinguishing features.—Its yellow-cream

variegated papyraceous leaflets easily distin-guish Zamia variegata. The variegations are inthe form of irregular yellow blotches that aremost apparent on the adaxial side of the lamina.

Additional specimens examined.—MEXICO.Chiapas: Ocosingo, Castillo-Campos et al. 3848(XAL), 3855 (XAL), 3885 (XAL), M. Vázquez-Torreset al. 3924 (CIB); Margaritas, F. Nicolalde-Morejón et al.


1443 (XAL), 1444 (XAL), 1445 (XAL), 1446 (XAL),1447 (XAL), 1448 (XAL), 1449 (XAL), 1450 (XAL), 1451(XAL), 1452 (XAL); Lacandona, Stevenson 692 (NY,XAL).

GUATEMALA. Alta Verapaz: J. A. Steyermark45048 (F, NY). Izabal: J. J. Castillo & D. R. Hodel 2138(MO), M Véliz 6893 (BIGUA, MEXU).

Zamia vazquezii D.W. Stev., Sabato, A.Moretti & De Luca, Delpinoa n.s. 37–38:14. 1995–1996 (issued 1998). Type: Mexico.Veracruz: 22 Jan 1989, M. Vázquez-Torres etal. 3990 (holotype: NY; isotypes: FTG, MO,NY, U, CIB). (Fig. 13)

FIG. 13. Zamia vazquezii. A. Habit. B. Leaflet. C. Ovulate strobilus. D. Pollen strobilus. E. Cataphyll. (From theholotype.)


Stem hypogeous, dichotomously branchingwith age, 35 cm long, 12 cm in diam.Cataphylls chartaceous, persistent, base trian-gular, apex aristate, 5.9×3.8 cm at base,brown tomentose. Ptyxis inflexed. Leaves 4–6 to many, 35–100×16–29 cm, ascending,brownish when emerging, turning green atmaturity; petiole 21–45 cm long, terete,unarmed or rarely with prickles, rachis terete,up to 65 cm long. Leaflets 14–26 pairs,sessile, papyraceous, ovate to obpyriform,opposite to subopposite, apex acuminate,base cuneate; margins with numerous serra-tions along distal third, subrevolute, articu-lations brown in young leaflets, 0.5–0.7 cmwide; the median leaflets 7.1–14.6×2.8–4.1 cm. Pollen strobili usually 1–2 per apex,erect, ovoid to ovoid-cylindrical, up to10.6 cm long and 2.6 cm in diam, light-brown tomentulose, apex acute; peduncledensely light-brown tomentose, 4.5 cm long,1.2 cm in diam; pollen sporangiophore distalface hexagonal-truncate. Ovulate strobili usu-ally solitary, erect, cylindrical to ovoid-cylin-drical, up to 15 cm long, 7.3 cm in diam, grayto brown tomentulose, apex apiculate; pedun-cle densely brown tomentose, 5.3 cm long,1.4 cm in diam; megasporangiophores pel-tate, distal face hexagonal-truncate to scuti-form, 1.2 cm high, 2.9 cm wide. Seeds ovoid,sarcotesta pink when immature, orange-red tored at maturity, 1.6 cm long, 1.2 cm in diam,sclerotesta smooth.Chromosome number.—2n=18 (Stevenson

et al., 1995–1996a).Distribution and habitat.—Endemic to

Veracruz, Mexico (Fig. 5), found between50–350 m in evergreen tropical forest sensuRzedowski (1978) classification, on predom-inantly deep clayey soils.Etymology.—The specific epithet honors

Mario Vázquez-Torres, a Mexican biologistand cycad specialist.Distinguishing features.—Zamia vazquezii

has more leaves (up to six), longer with eachup to 100 cm long, and wider leaflets than Z.fischeri. The two species are disjunct indistribution and have different chromosomenumbers, Z. vazquezii 2n = 18 and Z. fischeri2n = 16 (Stevenson et al., 1995–1996a).

Additional specimens examined.—MEXICO.VERACRUZ: Papantla, M. Vázquez-Torres 4568 (CIB),Allen 1970–2141 (XAL); Tiguatlán, J. Rees 1617 (XAL).

Species dubium

Zamia verschaffeltii Miq., Verh. Kon. Ned.Akad. Wetensch., Afd. Natuurk. 2(4): 31.1870. Type: Mexico. “Zamia fusca latifoliaVersch.", Miquel s.n. (holotype, U).Zamia verschaffeltii forma latifolia Schuster,

Pflanzenr. 99: 138. 1932. Type: description (lecto-type, designated by Stevenson & Sabato, 1986).

Socorro is the localitymentioned by Schuster(1932) for Miquel’s (1870) source of Zamiaverschaffeltii, although no locality informationis given on the type specimen, which is the onlyspecimen at U where Miquel’s herbariumresides (Stafleu, 1966). The original descriptionby Miquel gives the source of material as “Eximperio Mexicano introduxit A. Verschaffelt,qui in Catalogis Z. fuscam latifoliam dixit” andmakes no mention of a specific locality.However, following the information given bySchuster (1932), we have conducted exhaustivesearches over the years at two possible locali-ties with the name of El Socorro, one inTabasco and the other at Ruta del Socorro inVeracruz, It is important to note that the wholearea and surrounding regions have been con-verted into vast sugar cane plantations so thathistorically Zamia could have been present butwould now be extirpated. We can confidentlysay that no species of Zamia were found in ornear the two localities of the name Socorro, andno other locality of this name was locatedwithin the distribution range of this speciescomplex in the study. Also, no individuals orpopulations of Zamia studied here conform tothe original description of Z. verschaffeltii;because we were unable to find another recordof Z. verschaffeltii since its publication in 1870,we believe that this species is probably extinct(Nicolalde-Morejón et al., 2008). It is interest-ing that Schuster actually described a formlatifolia Schuster does not cite any specimensof this form, but only that it is cultivated in“Garten Verschaffelt.” We have been unable tolocate any specimens of this form that wereseen by Schuster, and if one did exist at B it wasdestroyed. Therefore, all source materialremains enigmatic.

Acknowledgments

The first author thanks the Red Latin-oamericana de Botánica for the award of a


Ph. D. fellowship (RLB-06-D2; SystematicsProgram, Instituto de Ecología, A.C., Xalapa,Mexico) and the Mellon Foundation for astipend for a six months residence at the NewYork Botanical Garden during 2002. Thisresearch was supported partially by CONA-CyT-SEMARNAT grant No. 2002-CO1–0183 to AV and NSF Grants BSR-8607049and EF-0629817 to DWS. The authors thankFrancisco Vergara Silva, Jorge GonzálezAstorga, and Victoria Sosa for their com-ments on a previous version of this manu-script, and Pablo Carrillo-Reyes, EduardoRuiz for their comments on the diagnostickey to species of Zamia, Carlos Iglesias forassistance and guidance in the field, andEdmundo Saavedra for the illustrations ofZamia spartea. We thank the Curators andstaffs of the herbaria mentioned for makingtheir collections available for study, as well asthe staff of the Jardín Botánico Fco. J.Clavjiero of the Instituto de Ecología, A.C.for access to the living specimens of theMexican National Cycad Collection. Finally,the authors would like to thank Roy Osbornefor his salient discussions and diligence inreviewing this manuscript throughout thepreparation of this work.

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Taxonomic revision of Zamia in Mega-Mexico€¦ · Tabasco, and Veracruz, ... spartea is illustrated for the ﬁrst time, and chromosome numbers for Z. herrerae are reported and illustrated