Revision of Zamia in Mega Mexico

Taxonomic revision of Zamia in Mega-Mexico
FERNANDO NICOLALDE-MOREJN1, ANDREW P. VOVIDES1,

2
AND DENNIS W. STEVENSON
1
Departamento de Biologa Evolutiva, Instituto de Ecologa, A. C. km 2.5 Antigua Carretera a

Coatepec No. 351, Xalapa 91070 Veracruz, Mexico
2
The New York Botanical Garden, Bronx, NY 10458-5120, USA; e-mail: dws@nybg.org
Abstract. The genus Zamia is revised for Mega-Mexico, with 22 species recognized
and described. The study presents a taxonomic clarication for the genus in Mesoamerica, a contribution that provides the foundation for a future monograph for Zamia
in the Neotropics. The largest proportion of species richness and endemism for the
genus is concentrated in southeastern Mexico, among the states of Chiapas, Oaxaca,
Tabasco, and Veracruz, an area that is considered highly diverse in oristic terms.
Distribution maps and a key to species are also provided, as well as complete descriptions of the specimens examined, including information on nomenclatural types,
habitats, synonymies, and etymologies. A lectotype is designated for Zamia loddigesii,
and neotypes for Z. galeottii, Z. leiboldii var. angustifolia, and Z. variegata. Zamia
spartea is illustrated for the rst time, and chromosome numbers for Z. herrerae are
reported and illustrated. Finally, scanning electron micrographs of leaet trichome
character states are presented, along with a discussion of their systematic implications
within the group.
Key Words: Endemism, Mexico, gymnosperms, cycads, oristic richness, Zamia.
Resumen. El gnero Zamia es revisado para Mega-Mxico, con 22 especies descritas
y una species dubia. El estudio est orientado al esclarecimiento taxonmico del
gnero en Mega-Mxico, una contribucin que siente los fundamentos para una futura
monografa para Zamia en el Neotrpico. La mayor riqueza y endemismo para el
gnero se concentra en el sureste de Mxico, entre los estados de Chiapas, Oaxaca,
Tabasco y Veracruz, rea de alta biodiversidad orstica. Mapas de distribucin y una
clave para las especies son presentadas, como tambin descripciones completas, tipos,
hbitat, sinnimos, etimologa y especimenes examinadas. Se designa un lectotipo
para Zamia loddigesii, y neotipos para Z. galeottii, Z. leiboldii var. angustifolia y Z.
variegata. Zamia spartea es ilustrada por primera vez y nmeros cromosmicos para
Z. herrerae son presentados. Finalmente, se presentan fotografas de tricomas al microscopio electrnico y una discusin de sus implicaciones en la sistemtica del
grupo.
According to Stevenson (1992), Zamiaceae

comprises eight genera distributed in tropical
and subtropical Africa, Australia, Greater
Antilles, North, Central and South America.
Five genera with 94 species are known from
the Neotropics (Hill et al., 2007). The genera
Ceratozamia Brongn. (21 spp.) and Dioon
Lindl. (13 spp.) are both endemic to Mexico
and a neighboring biogeographic region of
Central America that is oristically similar to

the southern part of Mexico (Mega-Mexico)
while Microcycas A. DC. (1 sp.) and Chigua
D.W. Stev. (2 spp.) are endemic to Cuba and
Colombia, respectively. The type genus for
the family, Zamia L. (59 spp.), is distributed
throughout tropical and sub-tropical America
as well as the Caribbean, with exception to
the Lesser Antilles.
Brittonia, 61(4), 2009, pp. 301335

2009, by The New York Botanical Garden Press, Bronx, NY 10458-5126 U.S.A.
ISSUED: 1 December 2009
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Zamia is the widest distributed genus of the

order Cycadales in the Neotropics. Its northern
range starts in Georgia and Florida (U.S.A.),
reaching Bolivia and the Mato Grosso of
Brazil in South America (Balduzzi et al.,
1982; Sabato, 1990; Norstog & Nicholls,
1997; Stevenson, 2001a, b). A remarkable
morphological and cytological variation has
been documented (Vovides, 1983; Moretti &
Sabato, 1984; Moretti, 1990a, b; Stevenson et
al., 19951996a; Vovides & Olivares, 1996;
Norstog & Nicholls, 1997), and also high
levels of genetic variation (Gonzlez-Astorga
et al., 2006). As a consequence of this
complexity, the taxonomy of Zamia is controversial; although the genus comprises 75
species, their circumscription and limits have
yet to be determined (Hill et al., 2007).
The most recent exhaustive taxonomic
treatment for Zamia was published by Schuster (1932) and later work by Sabato (1990)
and Stevenson (1987; 1991a, b; 1993; 2001a,
b; 2004) underlined nomenclatural and taxonomic anomalies in Zamia, principally owing
to insufcient eldwork and a scarcity of
good-quality botanical collections. The taxonomic history for the genus in Mexico began
with the publication of Zamia fufuracea L. f.
in 1789 from the central-south coastal region
of Veracruz, which also represents the rst
cycad species described from the American
continent. During the 19th century, six species were subsequently described for Mexico;
Z. scheri Miq., Z. katzeriana (Regel) Rettig,
Z. lawsoniana Dyer, Z. loddigesii Miq., Z.
spartea A. DC. and Z. verschaffeltii Miq.,
whereas the remaining known species were
documented and characterized during the
20th century, with a marked tendency for
taxonomic activity during the last thirty years
(Vovides et al., 1983; Schutzman et al., 1988;
Stevenson et al., 19951996a, b; Schutzman
& Vovides, 1998; Vovides, 1999).
The present taxonomic revision includes
endemic species of the cycad genus Zamia that
occur in Mega-Mexico 2, a term coined by
Rzedowski (1991) that associates the Central
American territories of Guatemala, Belize,
Honduras and northern Nicaragua to the
Mexican states of Nayarit on the Pacic to
southern Tamaulipas on the Gulf of Mexico using
biotic (mainly oristic) criteria. Rzedowskis
concept will be referred to as simply Mega-
Mexico hereafter, given that the slight differences in the boundaries of Mega-Mexico 1
with respect to Mega-Mexico 2 do not affect
the biogeographic aspects of our description
of the species. This study is intended to
provide a taxonomic clarication of the genus
in Mesoamerica and to provide the basis for a
future monograph of Zamia.
Materials and methods

The present taxonomic revision is based on
more than 450 specimens from the following
herbaria: 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 to
obtain vouchers on loan from NAP; therefore,
material from this herbarium is not cited.
Chromosome counts for Zamia herrerae
Caldern & Standley were performed on ve
individuals held at the Jardn Botnico Fco.
Javier Clavijero, Instituto de Ecologa, A.C.
(JBC). These plants has been previously
collected at the Acacoyagua and Tonal
regions, located in the state of Chiapas and
represent the species range in southern Mexico. Plants from its full biogeographic range,
which would include El Salvador and Guatemala, were not available for this study. A
modied root-tip squash method was used for
examining somatic metaphase cells described
by Vovides (1983) with a 12 to 15 hour icewater (0C) follow-up soak after the 0.2%
colchicine pretreatment at ambient temperature (Schutzman et al., 1988). Counts were
made from the best 1015 metaphase cells
and karyotype noted according to the classication of Schlarbaum and Tsuchiya (1984).
Photomicrographs were produced using a
Zeiss Fomi III photomicroscope tted with
planapochromatic objectives and Kodak PlusX pan ASA 125 lm.
Scanning electron micrographs (SEM)
were taken on young leaet material from
living plants cultivated in the JBC. Samples
were placed on sample stubs with double
sided adhesive tape and then introduced into
a dessicator for 24 hours. All samples were
sputter coated with gold-palladium at 1.5 kv
at 5 mA for 8 minutes with a Jeol Fine Coat
2009]
NICOLALDE ET AL.: ZAMIA IN MEGA-MEXICO
JFC 1100 sputter coater. Observations were

made 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 specic habitats, accounting for
the restricted distribution of the majority of
the taxa. The species with the widest distribution are associated with two or more
vegetation types, namely (a) Zamia paucijuga
Wieland, found in pine-oak, oak and tropical
dry forests; (b) Z. polymorpha D.W. Stev., A.
Moretti & Vzq. Torres, located in evergreen
tropical rainforest, sub-deciduous tropical
forest and their secondary succession stages;
(c) Z. herrerae, which occurs in evergreen
tropical rainforest, sub-deciduous tropical
forest and tropical dry forest and their
secondary succession stages and nally, to a
lesser extent; and (d) Z. loddigesii, present in
evergreen tropical rainforest, but more commonly in sub-deciduous tropical forest and its
secondary succession stages.
MORPHOLOGY
HABIT.All adult cycad stems are pachycaulous and may be columnar and arborescent or
subterranean and tuber-like. The genera Dioon,
Microcycas, Ceratozamia and Lepidozamia
Regel are usually columnar arborescent in habit,
while the subterranean forms are characteristic
of Bowenia Hook. ex Hook. f., Chigua, and
Stangeria T. Moore. Cycas L., Encephalartos
Lehm., Macrozamia Miq. and Zamia have both
stem morphologies, either subterranean tuberlike or columnar arborescent (Stevenson, 1980).
In this context, the species of Zamia in
Mesoamerica represent both growth forms, with
the subterranean tuber-like habitat predominant.
Only four species, Z. inermis, Z. onanreyesii, Z.
soconuscensis and Z. tuerckheimii, have arborescent stems reaching up to 100 cm or more in
height. Some species branch dichotomously
with age (namely, Z. scheri Miq., Z. furfuracea,
Z. inermis Vovides, Rees & Vzq. Torres, Z.
soconuscensis Schutzman, Vovides & Dehgan,
Z. loddigesii and Z. herrerae), with the coastal
dune species Z. furfuracea being the most
303
notable, with branches reaching up to 80 cm

long in adult plants.
TRICHOMES.Trichomes of cycads leaves
are bi-celled, consisting of a small basal cell
and a longer free apical portion (Stevenson,
1981). All the trichomes analyzed here show
the same bifurcate pattern, with one arm
proportionally longer than the other (Fig. 1A).
Each trichome presents a rounded basal cell
and a more extensive cylindrical bifurcate free
portion, which in most cases observed had
evidence of collapse and twisting (Fig. 1BF).
The pubescence was signicantly denser in
emerging than in older adult leaves with the
latter becoming completely glabrous. An
exception to this condition is found in Zamia
furfuracea, which maintains a great part of its
original indument on the abaxial surface of
each leaet.
According to Stevenson (1981), four types
of trichomes occur in Zamia; of these, the
transparent ramied and the colored ramied
are the types found in the present work. In
contrast to what was found in the aforementioned study by Stevenson, no trend or
correlation was found between aerial stems
(i.e., Z. soconuscensis or Z. inermis) and
bifurcate trichomes with equal length
branches. Trichomes with unequal sized
prevail among taxa with both aerial and
subterranean stems. In this context, we
consider that a more extensive and detailed
sampling of the genus Zamia across the
Neotropics would be necessary to corroborate
any correlation between trichome morphology and stem habit.
REPRODUCTIVE STRUCTURES.Although the
characters employed for the identication of
species of Zamia have been obtained from
leaf 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-Morejn et
al., 2008), the evaluation of reproductive
characters, especially those corresponding to
ovulate strobili, is essential to discriminate
among closely related taxa. Outstanding
attributes that should be considered in this
case are (i) the form and shape of the cone
apex (Stevenson, 1987); (ii) the peduncle
position with respect to the vertical axis of
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FIG. 1. Trichomes. AB. Zamia furfuracea. CD. Z. katzeriana. EF. Z. polymorpha.
the cone, when mature (Schutzman et al.,

1988); and (iii) the overall color of the cone
(see descriptions and Figs. 2, 3). In contrast to
the ovulate strobili, pollen strobili show
scarce variation at the species level, and their
utility to discriminate among species that
might possess high degrees of genealogical
afnity is relatively low. For the description
of the pollen reproductive axes, the terminol-
ogy introduced by Mundry and Sttzel (2003)

has been followed.
CHROMOSOME NUMBERS.Zamia shows the
highest chromosome numbers and karyotype
variation 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 &
2009]
305
FIG. 2. Illustrations of ovulate strobili. A. Zamia inermis. B. Z. vazquezii. C. Z. paucijuga. D. Z. lacandona. E. Z.

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

polymorpha.
2009]
Olivares, 1996). In contrast, all Ceratozamia

species studied so far have stable diploid
chromosome numbers (2n=16) and karyotypes, as do all the Dioon species analyzed to
date with stable diploid chromosome numbers (2n=18) and karyotypes (Marchant, 1968;
Vovides, 1983, 1985; Moretti, 1990a, b).
Chromosome counts are presented and
illustrated (Fig. 4) here for the rst time for
Zamia herrerae. This species has, but to a
lesser extent, cytotype polymorphisms similar
to 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 from
two populations along its Mexican range in
Chiapas (Fig. 4). Both the m (median region
of the chromosomes) and T (telocentric)
chromosomes vary in number (611 T, 4
6 m) and are large (611 m for T and 812
m for m) and their arms can be longer than
half the spindle axis, which can cause mitotic
instability during telophase (Schubert, 2007).
Karyotype differences are probably due to
centric ssions occurring on some of the
larger m chromosomes, giving rise to telocentrics with part of the centromere still
present. In this context, there are two general
hypotheses to explain karyotypic evolution in
307
Zamia: rst, Norstogs hypothesis (Norstog,

1980, 1981) relating karyotype simplication
and symmetry with progressive fusion of
telocentric chromosomes, which predicts low
diploid number in taxa with a high number of
metacentric chromosomes, and secondly that
of more recent research on Zamia, which
postulates centric ssion rather than fusion
producing a progressively higher diploid number and asymmetric karyotypes with a high
number of telocentric chromosomes (Moretti
& Sabato, 1984; Vovides & Olivares, 1996;
Caputo et al., 2004). For a more in-depth
discussion of mechanisms of chromosome
evolution in seed plants, see Jones (1998).
The highly asymmetric karyotypes and
somatic chromosome numbers in both Z.
paucijuga and Z. polymorpha, which also
appear to be the pattern for Z. herrerae, seem
to be correlated with the highest morphological variation and widest geographic distribution of the genus in Mesoamerica. Zamia
herrerae has a range of about 1000 km,
spanning El Salvador, Guatemala, and Chiapas (Mexico); therefore, we suggest investigating the distribution of chromosome
character states in this species throughout its
range. This karyotype asymmetry contrasts
with their congeners of a more restricted
FIG. 4. Chromosomes of Zamia herrerae. A. 2n=23. B. 2n=24.
308
distribution, which often have less morphological variation and a tendency towards
constant chromosome number and karyotype,
e.g., Z. cremnophila Vovides, Schutzman &
Dehgan, Z. scheri, 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 that
atypical chromosome number increase attributed to ssion is probably a result of stressful
inuences.
DISTRIBUTION
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ENDEMISM
Seventy-ve percent of the species in this

revision are endemic to the type locality and
nearby areas. They are limited to two or three
close populations with low population densities. These attributes are consistent with
Rabinowitzs (1981) evaluation criteria for
species rarity, which mainly considers information related to geographic range, habitat
specicity and local population size. In
congruence with these criteria, the endemic
Zamia species of Mesoamerica are considered
rare, threatened or endangered and are listed
under the IUCN Red List (IUCN, 2005; Hill
et al., 2007).
With 21 endemic cycad species in three
genera, Mexico has the highest cycad diversity and number of endemics of the region.
Six species of Zamia are known in Guatemala, two of which are endemic (Z. monticola
Chamberlain and Z. tuerckheimii J. Donnell
Smith); in Honduras, three species are known
of which three recently described species are
endemic (Z. oreillyi C. Nelson, Z. sandovalii
C. Nelson, and Z. onanreyesii C. Nelson & G.
Sandoval). Two further species are known for
Belize, of which one is endemic (Z. prasina
W. Bull) and nally, El Salvador is represented by one broadly distributed species, Z.
herrerae, with a range that runs along the
Pacic seaboard through Guatemala and the
Sierra Madre de Chiapas in Mexico. Other
species of the genus in Mexico with broad
distributions are (a) Z. paucijuga, distributed
along the Pacic seaboard of Mexico ranging
from Nayarit (northwest Mexico) to Oaxaca
(southwest); (b) Z. polymorpha, distributed
widely throughout the Yucatn penninsula in
Mexico, Belize, and the Petn region of
Guatemala; (c) Z. loddigesii, ranging along

the Gulf of Mexico seaboard from Tamaulipas (northeast Mexico) to Tabasco (southeast); and (d) Z. variegata Warsz., distributed
between Guatemala and southern Mexico.
The southern and southeastern regions of
Mexico, comprising the states of Veracruz,
Oaxaca, Chiapas, and Tabasco, are the most
diverse 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 microendemics Z. furfuracea, Z. inermis and Z.
vazquezii, whereas two micro-endemic species, Z. scheri and Z. soconuscensis, are
known respectively from the Sierra Madre
Oriental and the Sierra Madre de Chiapas.
Finally, in spite of the status of Z. paucijuga as a Mexican endemic, its distribution is
extremely wide within Mega-Mexico, covering a range of about 1000 km between the
states of Oaxaca and Nayarit. The northern
limit of Z. paucijuga in the latter state
represents the northernmost distribution for
the genus along the Pacic seaboard of the
Neotropics.
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, Gartenora 25: 140.
1876.
Stems hypogeous and epigeous, erect to
decumbent, sometimes dichotomously
branched in mature plants. Cataphylls chartaceous to membranaceous, stipulate, persistent
or deciduous, base triangular, apex long
acuminate to aristate, tomentose, generally
reddish-brown to yellowish. Ptyxis erect to
inexed. Leaves stipulate, ascending to
descending to spreading, reddish-brown or
green when emerging; petiole sometimes
blackish in young leaves, terete or subterete,
without prickles or heavily to lightly armed
with straight or bifurcate prickles; rachis
subterete generally with few prickles along
the proximal third or without prickles, with
up to 60 pairs of leaets. Leaets articulate,
2009]
sessile, papyraceous to coriaceous, linear,

linear-lanceolate, lanceolate, ovate, obovate
obpyriform to elliptic, opposite to subopposite, falcate or non-falcate, imbricate to
non-imbricate, generally acute at apex and
symmetric, attenuate at base, margins entire
to dentate along upper third, subrevolute,
articulations green, yellowish or dark brown
in young leaets. Pollen strobili usually 12
(4), with sterile tip, erect, cylindrical to
conical, light brown to purple, tomentulose,
apex acute to apiculate, generally with densely tomentose peduncles; pollen sporangiophores cuneiform, distal face truncate
309
hexagonal, 0.30.55 cm long, fertile abaxial

surface 2 lobed with 214 bisporangiate
synangia per lobe, sporangia dehiscent by
longitudinal slit. Ovulate strobili usually
solitary, erect to decumbent, cylindrical to
ellipsoid, purple to yellowish, generally
tomentulose, apex acute to apiculate; peduncle densely tomentose; ovulate sporangiophores cuneiform-peltate to scutiform, distal
end truncate-hexagonal when not scutiform.
Seeds ovoid, sarcotesta white to pink when
immature, red at maturity, sclerotesta smooth
but sometimes with several furrows running
longitudinally from micropylar end.
Key to the species of Zamia in Mega-Mexico

1. Leaets chartaceous to papyraceous.
2. Leaet margin dentate.
3. Leaets elliptic, adaxial surface with yellow to cream variegation . . . . . . . . . . . . . . . . . . . . . Z. variegata
3. Leaets long-lanceolate, without variegation.
4. Leaets imbricate, peduncle of pollen strobili decumbent, up to 16 cm long. . . . . . . . . . . . . . Z. oreillyi
4. Leaets not imbricate, peduncle of pollen strobili erect, up to 8 cm long . . . . . . . . . . . . . . . Z. herrerae
2. Leaet margin serrulate to entire.
5. Leaet margin entire, chartaceous.
6. Leaets oblong-lanceolate, glossy, 46 cm wide; ovulate strobili iridescent blue-green at maturity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Z. tuerckheimii
6. Leaets linear-lanceolate, not glossy, 0.61.5 cm wide, ovulate strobili dark-brown at maturity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Z. soconuscensis
5. Leaet margin serrulate, papyraceous.
7. Leaets sub-falcate basally, long-acuminate, strongly apically curved. . . . . . . . . . . . . Z. monticola
7. Leaets straight, acuminate, not curved at apex.
8. Leaets elliptic to lanceolate, ovulate strobili cylindrical to ovoid, dark-green and glabrous when
mature; from San Lus Potos and Quertaro, Mexico . . . . . . . . . . . . . . . . . . . . . . . Z. scheri
8. Leaets ovate to obpyriform, ovulate strobili ovoid-cylindrical, gray to brown tomentulose when
mature; endemic to Veracruz, Mexico . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Z. vazquezii
1. Leaets coriaceous.
9. Leaet margin entire to serrulate.
10. Leaet margin entire, petiole unarmed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Z. inermis
10. Leaet margin serrulate, petiole prickly.
11. Leaets long acuminate apically; stems arborescent . . . . . . . . . . . . . . . Z. onanreyesii
11. Leaets rounded to acute apically; stems subterranean.
12. Leaets obovate to oblanceolate, keeled adaxially, apex rounded . . . . Z. furfuracea
12. Leaets linear to oblanceolate, at adaxially, apex acute.
13. Leaets linear, 0.40.6 cm wide . . . . . . . . . . . . . . . . . . . . . . . . . .Z. spartea
13. Leaets lanceolate, 0.7 cm wide.
14. Leaets falcate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Z. sandovallii
14. Leaets not falcate.
15. Leaets lanceolate-oblanceolate, ovulate strobili dark-brown tomentulose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Z. polymorpha
15. Leaets linear-lanceolate, ovulate strobili brown to yellowish.
16. Ovulate strobili ellipsoid to cylindrical, apex acute to apiculate,
yellowish-brown; from the Pacic seaboard of Mexico . . . Z. paucijuga
16. Ovulate strobili ellipsoid to conical, apex acute, yellowish; from the Gulf
of Mexico seaboard . . . . . . . . . . . . . . . . . . . . . . . . . . . Z. loddigesii
9. Leaet margin distinctly dentate.
17. Leaets channeled adaxially between veins, appearing plicate . . . . Z. purpurea
17. Leaets smooth, not channeled adaxially between veins, not appearing plicate.
18. Leaets linear-lanceolate, imbricate, petiole strongly armed with straight or
bifurcate prickles up to 6 mm long . . . . . . . . . . . . . . . . . .Z. cremnophila
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18. Leaets lanceolate to oblanceolate, not imbricate, petiolearmed with small

straight prickles, generally between 24 mm.
19. Leaets with brilliantly shining cuticle on adaxial surface;ovulate strobili
decumbent when mature . . . . . . . . . . . . . . . . . . . . . . . . Z. katzeriana
19. Leaets without brilliantly shining cuticle on adaxial surface; ovulate
strobili erect when mature.
20. Distal leaets sub-falcate, petiole with bulbous base, blackish in young
leaves. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Z. lacandona
20. All leaets straight, petiole with no bulbous base, greenish in young
leaves.
21. Leaets oblong to lanceolate, bright grass-green, without conspicuously denticulate along margins; ovulate strobili green glabrous
when mature, apex acuminate . . . . . . . . . . . . . . . . Z. prasina
21. Leaets long-lanceolate, conspicuous dentate, up to 4 mm long;
ovulate strobili brown tomentulose when mature, apex longapiculate. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Z. standleyi
Zamia cremnophila Vovides, Schutzman &

Dehgan., Bot. Gaz. 149(3): 351. 1988. Type:
Mexico. Tabasco: 18 Aug 1981, M. A.
Magaa & S. Zamudio 343 (holotype: MEXU;
isotypes: CSAT, FCME, MO, UAMIZ).
Stem hypogeous, generally unbranched, 5

34 cm long, 411.3 cm in diam. Cataphylls
chartaceous, persistent, base triangular, apex
aristate, 5.4 cm long, 2.9 cm at base, reddishbrown tomentose. Ptyxis inexed. Leaves 23
(4), 57.517940.276.1 cm, descending, reddish-brown when emerging; petiole 10.2
95.3 cm long, blackish in young leaves,
subterete, heavily armed with straight or bifurcate prickles up to 6 mm long; rachis subterete,
up to 84 cm long, with few prickles along the
proximal third. Leaets 528 pairs, sessile,
coriaceous, lanceolate, opposite to subopposite,
imbricate, apex acute, base attenuate, margins
dentate along distal third, subrevolute; articulations dark brown in young leaets, 0.81.2 cm;
the median leaets 22.7383.14.4 cm wide.
Pollen strobili usually 12, erect, cylindrical, up
to 8 cm long and 1.4 cm in diam, light brown,
apex acute; peduncle densely light brown
tomentose, 4.5 cm long, 1.2 cm in diam; pollen
sporangiophores cuneiform, distal face truncatehexagonal, 0.35 cm long, fertile abaxial surface
with 3 bisporangiate synangia per lobe. Ovulate
strobili usually solitary, erect, ellipsoid, 13.2 cm
long, 4.9 cm in diam, brown to reddish,
tomentulose, apex acuminate; peduncle densely
brown tomentose, 2.9 cm long, 1.6 cm in diam;
distal face hexagonal-truncate with a horizontal
longitudinal depression, 11.2 cm high, 1.1
1.4 cm wide. Seeds ovoid, sarcotesta white
when immature, red at maturity, 22.8 cm long,

1.42 cm in diam, sclerotesta smooth.
Chromosome number.2n=16 (Schutzman
et al., 1988).
Diversity and genetic structure.The average of alleles per locus is A=1.98, the
percentage of polymorphic loci is P=94.3, the
expected heterozygosity is HE =0.347 and
the genetic differentiation between the two
populations currently under study is Fst=
0.093 (Gonzlez-Astorga et al. unpubl. data).
Distribution and habitat.Endemic to
Tabasco, Mexico (Fig. 5), on karstic rocks and
cliffs of the Sierra El Madrigal, between 50
150 m. The vegetation type where this species
grows is evergreen tropical rain forest or bosque
tropical perennifolio of Rzedowski (1978).
Etymology.The specic epithet is derived
from the Greek word for cliff-friend
( = cremnos = cliff, and =
los = friend/lover; Schutzman et al., 1988),
because of its unusual habitat.
Distinguishing features.The species is
characterized by its exclusive habit on the
rocky walls of limestone cliffs, in addition to
descendent leaves. Petioles are densely armed
with prickles that are sometimes branched,
and also lanceolate, imbricate leaets that are
visibly dentate along the distal third.
Additional specimens examined. MEXICO.
TABASCO: Teapa, F. Nicolalde-Morejn et al. 1497
(XAL), M. A. Prez-Farrera 293 (HEM), M. A. PrezFarrera 900 (HEM, MEXU).
Zamia scheri Miq. in Lem., Hort. Vanhoutt. 1:

20. 1845. Type: ex Horto Petropolitano in H.
Houtte. vecta, Miquel s.n. (neotype, designated
by Stevenson & Sabato, 1986: U). (Fig. 7)
2009]
311
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 ( ).
Stem subterranean, dichotomously branching

in older plants, up to 30 cm long, 48 cm in
diam. Cataphylls membranaceous, persistent,
base triangular, apex aristate, 4.5 cm long,
1.2 cm at base, yellowish tomentose. Ptyxis
inexed. Leaves 15(8), 1545820 cm,
ascending to spreading, dark-brown when
emerging; petiole 914 cm long, blackish in
young leaves, unarmed, terete; rachis subterete,
up to 31 cm long, unarmed. Leaets 2035
pairs, sessile, papyraceous, elliptic to oblanceolate, alternate to subopposite, apex acute symmetric, base cuneate, margins serrulate along
distal third, subrevolute; articulations lightbrown in young leaets, 0.30.4 cm wide,
median leaets 591.54.5 cm. Pollen strobili usually 13, erect, conical, 46 cm long,
1.52.2 cm in diam, gray tomentose, apex
acute; peduncle yellowish tomentose, 2.8 cm
long, 0.91.1 cm in diam; pollen sporangiophores cuneiform, distal face hexagonaltruncate, 0.3 cm long, fertile abaxial surface
with 34 bisporangiate synangia per lobe.
Ovulate strobili usually 12, erect, cylindrical
to ovoid, up to 9 cm long, 4.5 cm in diam, brown

to reddish tomentulose when young, dark-green
and glabrous when mature, apex acute; peduncle
brown tomentose, 3 cm long, 1.1 cm in diam;
megasporangiophores peltate, distal face hexagonal-truncate when immature, scutiform when
mature 0.81.2 cm high, 1.31.9 cm wide. Seed
ovoid, 1.6 cm long, 1.3 cm in diam, sarcotesta
pink when young and orange at maturity.
Chromosome number.2n=16 (Marchant,
1968; Moretti et al., 1991; Stevenson et al.,
19951996a).
Distribution and habitat.Zamia scheri
is endemic to Mexico (Tamaulipas, San Luis
Potos, Quertaro and Hidalgo; Fig. 5), between 140900 m elevation. The vegetation
type of its habitat is pine-oak forest, tropical
deciduous forest and mountain tropical forest
(sensu Rzedowski, 1978).
Etymology.The specic epithet is in
honor of Friedrich Fischer, a German cycad
horticulturist of the 19th century.
Distinguishing features.Zamia scheri is
a small fern-like plant (up to 100 cm tall),
312
BRITTONIA
with papyraceous but serrulate leaets and

unarmed petiole and rachis. Ovulate strobili
greenish upon maturity.
Additional specimens examined.MEXICO.
QUERTARO: Jalopan, Lpez 438 (XAL), Servn 1471
(XAL), Vovides 330 (XAL). SAN LUIS POTOS: O. M.
Clark 6839 (MO), J. Rees 1686 (XAL), Stevenson et al.
566 (MEXU, NY), Vovides 753 (XAL); El Naranjo, F.
Nicolalde-Morejn & J. Gonzlez Astorga 1614 (XAL),
1615 (XAL), 1616 (XAL), 1617 (XAL), 1618 (MEXU,
XAL), 1619 (XAL); Ciudad Valles, F. Nicolalde-Morejn
& J. Gonzlez-Astorga 1620 (XAL), 1621 (XAL), 1622
(XAL), 1623 (MEXU, XAL), 1624 (MEXU, XAL), 1625
(MEXU, XAL), 1626 (MEXU, XAL). HIDALGO: Pisaores, O. Alcntara-Ayala & R. Mayorga-Saucedo 3325
(FCME).
Zamia furfuracea L. f. in Aiton, Hortus

Kew. 3: 477. 1789. Type: Palma americana crassis rigidisque foliis, pl. 210, in
Herm. Paradisus Batavus. 1698 (lectotype,
designated by Stevenson & Sabato, 1986:
pl. 210).
Zamia muricata var. obtusifolia Miquel, Tijdschr. nat.
Gesch. Physiol. 10(1): 7172. 1843. Zamia latifolia Loddiges ex Miquel, Tijdschr. wis-en natuurk.
Wet. 2(4): 298. 1849. Type: Tab. VII, g. a in
Linnaea 19(4): 1847. (neotype, designated by
Stevenson & Sabato, 1986: Tab. VII, g. a).
Zamia furfuracea var. trewii A. DC., Prodr. 16(2):
541. 1868. Type: Palmifolia fructu clavato polypireno[polyspermo]. C. J. Trew, PI. Select. Tab.
26. 1752 (holotype: Tab. 26, G; typotype, designated by Stevenson & Sabato, 1986: BM).
Stem hypogeous, becoming epigeous with

age, often dichotomously branched, up to
60 cm long, 20 cm in diam. Cataphylls
coriaceous, persistent, base triangular, apex
long aristate, up to 1034.5 cm at base,
yellowish tomentose. Ptyxis inexed. Leaves
3 to many, 45190 1030 cm, diffuse,
brown-yellowish when emerging; petiole
1750 cm long, brown-yellowish when
young, subterete, armed with small prickles
up to 3 mm long; rachis terete, up to 120 cm
long, with few prickles along the proximal
third. Leaets 818 pairs, sessile, coriaceous,
obovate to oblanceolate, opposite to subopposite, imbricate, keeled, apex rounded to
sub-acute, base attenuate, margins slightly
serrulate along the 1/2 distal portion, subrevolute; articulations yellow in young and
juvenile leaets, 0.40.7 cm wide; median
leaets 1420 47.5 cm. Pollen strobili
[VOL 61
usually 24, erect, cylindrical, up to 17 cm

long, 1.8 cm in diam, yellowish to brown,
apex acute; peduncle densely light-brown
tomentose, 12 cm long, 1.1 cm in diam;
pollen sporangiophores cuneiform, distal face
hexagonal-truncate, 0.450.55 cm long, fertile abaxial surface with 89 bisporangiate
synangia per lobe. Ovulate strobili usually 1
per crown, erect, cylindrical, up to 25 cm
long, up to 10.5 cm in diam, yellowish-green
tomentulose when immature, light brown
upon maturity, apex apiculate; peduncle yellowish tomentose, up to 8 cm long, up to
1.3 cm in diam; megasporangiophores peltate, distal end hexagonal-truncate, 0.9 cm
high, 1.3 cm wide. Seeds ovoid, sarcotesta
yellowish-green when immature turning red
at maturity, 1.6 cm long, 1.2 cm in diam,
sclerotesta smooth.
Chromosome number.2n=18 (Moretti,
1990a, b)
Mexico in central and south coastal Veracruz,
along a coastal stretch of approximately150 km
in stable dunes and basalt cliffs (Fig. 5).
Etymology.The specic epithet alludes to
the persistent brown-yellowish trichomes of
the leaves throughout the developmental
stages of the plant.
Distinguishing features.Leaves strongly
keeled with highly imbricate and coriaceous
obovate to oblanceolate leaets with brownyellowish indumentum persisting with age.
V ERACRUZ: Ibarra-Mariquez 316 (MEXU, MO),
Ibarra-Mariquez 1952, 1959 (MEXU); Alvarado, Rees
1650, 1651, 1652 (XAL), M. Vzquez-Torres et al. 4871
(CIB); Catemaco, Calzada 1475 (MEXU), Calzada 2451
(XAL), Cedillo 2610 (MEXU, XAL), Gonzlez-Quintero
1520 (ENCB), Ibarra 316 (MEXU), Lot 1277-14 (F,
XAL), Menendez 115 (MEXU, MO, XAL), F. NicolaldeMorejn 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), CastilloCampos 12732 (XAL); Lerdo De Tejada, Vovides 828
(XAL), 829 (XAL), 830 (XAL), 831 (XAL), 832 (XAL),
2009]
833 (XAL), 839 (XAL); San Andrs Tuxtla, CastilloCampos et al. 13881 (XAL), Chazaro B. 512 (XAL),
Hammel & Merello 15499 (MO), Hernndez-M 1216A
(F, MEXU), Lorence 4978 (MEXU), Sousa 3099 (F,
MEXU, MO), Vovides & Iglesias 1148 (XAL).
Zamia herrerae Caldern & Standl., Proc.

Wash. Acad. Sci. 14(4): 93. 1924. Type: El
Salvador. Vicinity of Sonsonate, 17 Jul
1923, S. Caldern 1682 (holotype: US).
313
be found in secondary growth forests and

pastures.
Etymology.The specic epithet honors
Hector Herrera, a scientist from El Salvador.
Distinguishing features.Zamia herrerae is
distinguished by its long-lanceolate papyraceous leaets with dentate margins along the
distal 2/3.
CHIAPAS: Escuintla, Matuda 16368 (MEXU), 16871
(MEXU), 17332 (MEXU), 18332 (MEXU), M. A.
Prez-Farrera 143 (HEM), Schutzman 526 (XAL), 527
(XAL), 528 (XAL); Tonal, Farrera 2489 (CHIP), F.
Nicolalde-Morejn & J. Gonzlez-Astorga 1579 (XAL),
1580 (XAL), 1581 (XAL), M. A. Prez-Farrera 744
(CIB).
GUATEMALA. P. C. Standley 67306 (F).
NICARAGUA. MANAGUA: Cultivated, A. Grijalva
3658 (MO).
Stem hypogeous, bifurcate in adult plants,

6264.513.5 cm in diam. Cataphylls chartaceous, persistent, base triangular, apex
aristate, 4.61.2 cm wide at base, yellowish
tomentose. Ptyxis inexed. Leaves 24, erect,
green to light-brown when emerging, 6196
24.529.5 cm; petiole 1934 cm long, brownish in young leaves, terete, armed with
prickles up to 3 mm long; rachis subterete,
up to 63 cm long, with few prickles along the Zamia inermis Vovides, J.D. Rees & Vzq.
Torres, Flora de Veracruz 26: 22. 1983. Type:
proximal third. Leaets 1532 pairs, sessile,
Mexico. Veracruz: 6 Jun 1981, Vovides 666
papyraceous, lanceolate, alternate to subop(holotype: XAL; isotype: F).
(Fig. 8)
posite, apex acute, base symmetric attenuate;
margins dentate along distal 2/3, subrevolute;
Stem epigeal, erect, dichotomously brancharticulations dark brown when young, 0.3
0.5 cm wide; median leaets 22383.1 ing in mature plants, 1543 cm long, 8.6
4.4 cm. Pollen strobili usually 23, erect, 26.4 cm in diam. Cataphylls chartaceous,
cylindrical to conical, 4.37.5 cm long, 1.3 persistent, base triangular, apex aristate, 2.1
2.1 cm in diam, light brown tomentulose, 2.6 4.15.6 cm wide at base, yellowish
apex mucronate; peduncle densely light- tomentose. Ptyxis inexed to erect. Leaves 10
brown tomentose, 6.27.8 cm long, 1.1 35, erect, light to yellowish-green when emerg1.4 cm in diam; pollen sporangiophores ing, 309543.560 cm; petiole 1841 cm long,
cuneiform, distal face hexagonal, 0.35 cm greenish in young leaves, subterete, without
long, fertile abaxial surface with 34 bispor- prickles; rachis subterete, 1519 cm long,
angiate synangia per lobe. Ovulate strobili unarmed. Leaets 2732 pairs, sessile, coriausually solitary, erect, cylindrical to ovoid, ceous, linear-lanceolate, opposite to suboppo7.111.6 cm long, 4.14.9 cm in diam, site, apex acute, base attenuate; margins entire,
brown, tomentulose, apex acute; peduncle subrevolute; articulations 0.40.6 cm wide;
densely brown tomentose, 3.54.2 cm long, median leaets 2030.50.91.2 cm. Pollen
1.11.3 cm in diam; megasporangiophores strobili cylindrical, usually 12 per crown, erect,
peltate, distal face hexagonal-truncate, 1.3 up to 9.1 cm long, up to 2.8 cm in diam, beige1.6 cm high, 2.22.8 cm wide. Seeds ovoid, yellowish, apex acute; peduncle densely lightsarcotesta pink when immature, red at matu- yellowish tomentose, up to 4.5 cm long, 1.1 cm
rity, 1.61.9 cm long, 1.21.5 cm in diam, in diam; pollen sporangiophores cuneiform,
distal face hexagonal-truncate, 0.35 cm long,
sclerotesta smooth.
Chromosome number.2n=23, 24 (Fig. 4). fertile abaxial surface with 56 bisporangiate
Distribution and habitat.The species synangia per lobe. Ovulate strobili usually 12
range is El Salvador, Guatemala, and southern per crown, erect, cylindrical, 1323 cm long, 8
Mexico in Chiapas, between 100600 m 9.8 cm in diam, light-brown to beige tomentuelevations (Fig. 6). It generally grows in lose, apex apiculate; peduncle brown tomentose,
tropical deciduous forest (sensu Rzedowski, 68 cm long, 1.21.4 cm in diam; megaspor1978) on deep clay soils. Also, this cycad may angiophores peltate, distal face hexagonal-trun-
314
BRITTONIA
[VOL 61
FIG. 6. Distribution of Zamia scheri (), Z. herrerae ( ), Z. loddigesii ( ), Z. oreillyi ( ), Z. paucijuga ( ), Z.

polymorpha (), Z. purpurea (), Z. spartea ( ), Z. standleyi ( ), Z. tuerckheimii ( ), Z. variegata ( ).
cate, 0.70.9 cm high, 1.11.3 cm wide. Seeds

ovoid, sarcotesta pink when immature, red at
maturity, 1.72.5 cm long, 1.42.1 cm in diam,
sclerotesta smooth.
Chromosome number.2n=16 (Vovides,
1983).
Mexico in a small mountain range in central
Veracruz (Fig. 5) at 150300 elevation on
basaltic soils. The vegetation type of the habitat
is tropical deciduous forest (sensu Rzedowski,
1978).
the absence of prickles on the petiole and
rachis and the entire leaet margins.
Distinguishing features.Zamia inermis
differs from its congeners in Mexico by a
total absence of prickles along the petiole and
rachis, as well as having totally entire leaets
and light-yellowish to clear-beige tomentulum on the pollen and ovulate strobili.
VERACRUZ: Actopan, Acosta & Acosta 234 (XAL), F.
Nicolalde-Morejn & Vovides 1415 (XAL), 1416 (XAL),

1417 (XAL), Schutzman 570 (XAL), 571 (XAL), 572
(XAL), 575 (XAL), 576 (XAL), 577 (XAL), J. Rees et
al. 681 (XAL).
Zamia katzeriana (Regel) Rettig, Gartenora 45: 148. 1896. Ceratozamia katzeriana 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 761046, 11 Apr 1984, J.
Watson s.n. (holotype: NY; isotypes: FLAS, FTG,
MEXU).
Stem hypogeous, unbranched, up to 25 cm

long, up to 7 cm in diam. Cataphylls chartaceous, semidecidious, base triangular, apex
aristate, 5.3 1.4 cm at base, yellowish
tomentose. Ptyxis inexed. Leaves 12 (3),
492203558 cm, ascending to descending,
bright reddish-pink with lustrous cuticle when
2009]
315
FIG. 7. Zamia scheri A. Habit. B. Leaets. (From Stevenson 566, NY).
emerging, dark green when mature; petiole

20130 cm long, terete, armed with few
simple prickles; rachis subterete, up to
84 cm long, unarmed. Leaets 37 pairs
sessile, coriaceous, oblong-lanceolate, opposite to subopposite, adaxial surface with
brilliantly shining cuticle throughout life of
leaet, apex acute, base attenuate; margins
dentate along distal third, subrevolute; articulations brown in young leaets turning green
with age 0.61.6 cm wide; median leaets
18353.512 cm. Pollen strobili 15, conical, light-brown tomentulose, erect upon
emergence becoming prostrate to decumbent
upon maturity, up to 3.9 cm long, 1.1 cm in
diam, apex acute; peduncle light-brown tomentose, up to 5.8 cm long, 1.2 cm in diam;
hexagonal scutiform, 0.35 cm long, fertile
abaxial surface with 56 bisporangiate synangia per lobe. Ovulate strobili usually
solitary, decumbent to erect, elliptic, 8
12 cm long, 4.56 cm in diam, brown to
yellowish tomentose, apex aristate; peduncle
brown tomentose, 2.14.3 cm long, 1.11.3 cm
in diam; megasporangiophores peltate, distal
face hexagonal-truncate to scutiform, 11.2 cm
high, 1.11.3 cm wide. Seeds ovoid, sarcotesta
pink when immature, red at maturity, 1.1
1.4 cm long, 1.61.8 cm in diam, sclerotesta
smooth.
Chromosome number.2n=16 (Schutzman,
1984; Moretti, 1990a).
Diversity and genetic structure.The
average of alleles per locus is A=1.95, the
percentage of polymorphic loci is P=84.4,
the expected heterozygosity is HE =0.280
and the genetic differentiation between
316
BRITTONIA
[VOL 61
FIG. 8. Zamia inermis. AB. Habit. C. Cataphyll. D. Pollen strobilus. E. Microsporophyll, abaxial and adaxial
view. FH. Ovulate strobilus. I. Ovulate sporangiophores. J. Seed. (From the holotype.)
2009]
the two populations currently under study is

Fst=0.194 (Gonzlez-Astorga et al. unpubl.
data).
Mexico and known from the states of
Chiapas, Tabasco and Veracruz (Fig. 5) at
200700 m in evergreen tropical forest (sensu
Rzedowski, 1978).
Etymology.Specic epithet in honor of
Katzer, inspector of the gardens in Paullowsk
(Stevenson & Sabato, 1986).
Distinguishing features.Zamia katzeriana
is easily distinguished from its congeners by
having leaves with highly lustrous or shiny
cuticles. Emerging leaves are a bright reddishpink. Both pollen and ovulate strobili are
borne on long peduncles that become descendent to prostrate.
VERACRUZ: Las Choapas, Martnez & Martnez-M. 825
(HEM), F. Nicolalde-Morejn et al.1436 (XAL), 1437
(XAL). TABASCO: Teapa, Hdez-Najarro 622 (CHIP), M.
A. Magaa 1905 (MEXU), M. A. Prez-Farrera s.n.
(XAL), M. A. Prez-Farrera 899 (HEM, MEXU),
Walters s.n. (FTG accession 12-2, XAL). CHIAPAS: San
Fernando, Vovides et al. 1266 (XAL), Palacios 383
(CHIP), F. Nicolalde-Morejn & Prez-Farrera 1420
(XAL), M. A. Prez-Farrera s.n. (XAL), Walters s.n.
(FTG accession 23-2, XAL); Ocozocoautla, GmezPompa 705 (FCME, MEXU), F. Nicolalde-Morejn et
al. 1453 (XAL), 1454 (XAL), 1455 (XAL), 1456 (XAL),
1457 (XAL), 1458 (XAL), 1459 (XAL), 1460 (XAL), M.
A. Prez-Farrera 29 (CHIP, CIB, MEXU); Tila, Vovides
et 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, 614 cm
long, 59 cm in diam. Cataphylls coriaceous,
persistent, base triangular, apex long-aristate,
8.33.6 cm at base, reddish-brown tomentose. Ptyxis inexed, reddish-brown. Leaves
usually solitary, up to 3 under cultivation, 46
1713275 cm, ascending, reddish-brown
when emerging; petiole 1495.3 cm long
with bulbous base, blackish in young leaves,
subterete, proximal section strongly channeled, armed with prickles up to 5 mm long;
rachis subterete, 3176 cm long, with few
prickles along proximal third. Leaets 417
317
pairs, sessile, coriaceous, lanceolate, opposite

to subopposite, subfalcate, apex acute, base
attenuate, margins dentate along distal third,
subrevolute; articulations dark-brown when
young, 0.41.1 cm wide; median leaets
15.6372.96 cm. Pollen strobili usually
23, erect, conical, 5.46.6 cm long, 1.5
1.7 cm in diam, light-brown, apex acute;
peduncle light-brown tomentose, 5.86.9 cm
long, 1.2 cm in diam; pollen sporangiophores
cuneiform, distal face hexagonal truncate,
0.35 cm long, fertile abaxial surface with 4
5 bisporangiate synangia per lobe. Ovulate
strobili 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 cm
long, 1.2 cm in diam; megasporangiophores
peltate, distal face hexagonal-truncate, 1
1.2 cm high, 11.5 cm wide. Seeds irregularly
maturity, 22.4 cm long, 1.31.9 cm in diam,
sclerotesta smooth.
Chromosome number.2n = 16, 17, 18
(Schutzman & Vovides, 1998).
Chiapas (Fig. 5) in the vicinity of the
Lacandon forest at 80200 m elevation, in
evergreen tropical forest (sensu Rzedowski,
1978). It is also found in secondary succession stages of the tropical forest.
Etymology.The epithet is derived from
the name of the 1.8 million hectare Selva
Lacandona (Lacandona Jungle) in southeastern Chiapas, which itself bears the name of
the Lacandona Maya Indians who inhabit the
forest.
Distinguishing features.This species is
distinguished by having a solitary large leaf
(up to three leaves may be maintained on
plants under cultivation) with a stout erect
petiole and a strongly bulbous base. The leaf
is reddish-brown at emergence and petiole
dark purplish-brown turning dark-brown with
age. Ovulate cone generally solitary, with
acute apex.
318
BRITTONIA

CHIAPAS: Palenque, Schutzman 510 (XAL), 511 (XAL),
512 (XAL), 513 (XAL), 514 (XAL), 515 (XAL), 516
F. Nicolalde-Morejn & N. Martnez 1418 (XAL), M. A.
Prez-Farrera 890 (HEM, MEXU), Walters s.n. (FTG
accession 14-2, XAL), M. Vzquez-Torres et al. 3925
(CIB); San Jernimo 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: cultivated by Van Houtte 3374 (lectotype, here
designated: 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. Zamia
loddigesii var. leiboldii (Miquel) A. DC., Prodr. 16
(2): 541. 1868. Type: E. Mexico in Hortum
Loehrianum Lipsiae attulit Liebold, 1845, Miquel
s.n. (holotype: U).
Zamia loddigesii var. angustifolia Regel, Bull. Soc.
Nat. Moscou 30(1): 190. 1857. Type: ex horto
Petropolitano, 1856, Regel s.n. (holotype: LE)
Zamia loddigesii var. obtusifolia Regel, Bull. Soc.
Nat. Moscou 30(1): 190. 1857. Type: t. 186, gs
2728 in Gartenora 6: 1857. (lectotype, designated by Stevenson & Sabato, 1986: LE).
Zamia mexicana Miquel, Prodr. syst. Cycad. 13. 1861.
Type: Eriozamia mexicana H. Belg., 1847, Miquel
s.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: ex
Horto 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, Panzenreich 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 by
Stevenson & Sabato, 1986: NY; isolectotype, F-3
sheets).
Zamia loddigesii var. angustifolia (Regel) J. Schust.,
in Engl., Panzenr. 4(1): 148. 1932. Type: Mxico.
Veracruz: savanne bei Mundo nuevo, Karwinski
[VOL 61
1028b (lectotype, designated by Stevenson &

Sabato, 1986: LE; isolectotype, LE).
Zamia loddigesii var. longifolia J. Schust., in Engl.,
Panzenr. 4(1): 147. 1932. Type: Mexico. Veracruz: Colipa, Karwinski 1029 (lectotype, designated by Stevenson & Sabato, 1986: LE; isolectotype:
LE).
Stem hypogeous, branching dichotomously

with age, 1045 cm long, 815 cm in diam.
Cataphylls chartaceous, persistent, base triangular, apex aristate, 8.43.7 cm at base,
23 (4) ascending to spreading, 459630
41 cm, light-green when emerging, green to
dark-green when mature; petiole 1525 cm
long, green in young leaves, subterete, armed
with prickles up to 4 mm long; rachis
subterete, up to 57 cm long, with few prickles
on the proximal third. Leaets 1223 pairs,
sessile, coriaceous, linear-lanceolate, opposite
to subopposite, apex acute, base attenuate,
margins serrulate along distal third, subrevolute; articulations 0.40.7 cm wide; median
leaets 16261.83.1 cm. Pollen strobili 1
2 per crown, up to 6 (7) when multiple
crowned, erect, cylindrical, 814 cm long,
1.83.5 cm in diam, light brown tomentulose,
apex acute; peduncle light-brown tomentose,
6 cm long, 1.2 cm in diam; pollen sporangiophores cuneiform, distal face hexagonaltruncate, 0.3 cm long, fertile abaxial surface
with 68 bisporangiate synangia per lobe.
Ovulate strobili usually 12 per crown, erect,
ellipsoid to conical, up to 16 cm long, up to
6 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, 0.71 cm high,
1.92.6 cm wide. Seeds ovoid, sarcotesta pink
when immature, red at maturity, 1.41.8 cm
long, 0.81 cm in diam, sclerotesta smooth.
Chromosome number.2n=18 (Norstog,
1980; Moretti, 1990a, b).
the expected heterozygosity is HE =0.266
and the genetic differentiation between the
two populations currently under study is
Fst=0.179 (Gonzlez-Astorga et al. unpubl.
data).
Mexico and distributed widely within the
2009]
states of Tamaulipas, Hidalgo, Veracruz,

Tabasco and parts of Oaxaca with a single
known locality in Chiapas (Fig. 6). The
vegetation types are evergreen tropical forest,
tropical deciduous and sub-deciduous forests
(sensu Rzedowski, 1978), as well as a variety
of secondary succession and disturbed habitats such as pastures and cornelds, as well
as road-side vegetation.
Conrad Loddiges (17381826), a German
horticulturist who lived in London and
cultivated American cycads.
Nomenclatural notes.The description
of Zamia loddigesii was lectotypied by
Stevenson & Sabato (1986). However, it is
no longer permissible to designate a description to serve as a type. Moreover, since then
a specimen, Van Houtte 3374 at U, was
found that was sent to Miquel by Van
Houtte. This specimen predates the publication of Z. loddigesii Miq. and matches the
description. Thus, we are designating it the
lectotype.
Distinguishing features.Zamia loddigesii
in contrast to Z. paucijuga, has been widely
collected along the Gulf of Mexico seaboard
and on the Yucatan peninsula since the 19th
century, largely by British and Russian
botanists, and recently by Mexican and North
American researchers. The high morphological variation presented by this species has
resulted in the publication of 10 afne names
(Hill et al., 2007) that at times, has been a
basis for the separation of natural populations, i.e., that of the Yucatan peninsula
(formerly Z. loddigesii sensu Vovides &
Olivares, 1996) that is now a separate entity
Z. polymorpha (Stevenson et al., 1995
1996b), a decision based on both vegetative
and reproductive characters that differ from Z.
loddigesii mainly on pollen and ovulate
strobilus shape and indument color.
Zamia loddigesii is similar to Z. polymorpha in leaf morphology; however, there are
differences in the reproductive structures.
Pollen cones of Z. loddigesii are beige in
color with a blunt apex whereas those of Z.
polymorpha are dark reddish-brown or maroon with an acute apex. vulate cones of Z.
loddigesii are cylindrical and beige in color
and those of Z. polymorpha are ovoid and
dark maroon in color.
319

CHIAPAS: Ocozocoautla, M. A. Prez-Farrera 81 (CHIP,
HEM). HIDALGO: Atlapexco, San Juan 15 (XAL), 16
(XAL), 17 (XAL), 18 (XAL), 19 (XAL), 20 (XAL), 21
(XAL). PUEBLA: Sarukn et al. 4632 (FCME, MEXU).
Oaxaca, Tuxtepec, J. Chamberlain s.n. (MO). TAMAULIPAS: R. L. Dressler 1858 (MO), Mayel et al. 791
(MEXU); Aldama, F. Nicolalde-Morejn & J. GonzlezAstorga 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 & Gmez-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
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 & CastilloCampos 240 (XAL); Cotaxtla, Gonzlez 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. Rees
1763 (XAL), Stevenson et al. 538 (MEXU, NY), M.
Vzquez-Torres 8071 (CIB); Huejultla, Stresser 291
(MEXU); Hueyapan de Ocampo, Gmez-Pompa 4424
(XAL), Vovides et al. 1378 (XAL), 1379 (XAL), 1380
(XAL); Jalcomulco, Castillo & Zamora C. 7542 (XAL),
Castillo & Gmez-Pompa 2588 (XAL), Castillo-Campos
2727 (XAL); Mecayapan, Castillo et al. 13681 (XAL),
13792 (XAL), 13843 (XAL), 13861 (XAL), 13865 (XAL),
13866 (XAL), A. Calatayud & J. Martnez-Gndara 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. Bentez 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 single
seed collected opposite the crater of Naolinco, near Xalapa, Veracruz, Mexico, Oct
1925, C. J. Chamberlain s.n. (holotype:
MO; isotype: NY).
Stem epigeal, up to 30 cm tall, 1820 cm in
diam. Cataphylls base triangular, apex linearlanceolate, 3612 cm wide at base. Leaves
520, 100200 cm long, erect to slightly
curved; petiole 5075 cm long, terete, armed
with stout prickles in lower half; rachis terete,
up to 100 cm long, with few prickles along
the lower half. Leaets 3040 pairs, sessile,
chartaceous to papyraceous, linear-lanceolate,
320
BRITTONIA
opposite to subopposite, subfalcate near the

base, apex long-acuminate and often strongly
curved, base attenuate, margins serrulate only
near the base, subrevolute; articulations
0.40.7 cm wide; the median leaets 25
30 46 cm. Pollen strobili usually 26,
erect, cylindrical to oblong, 1220 cm long
and 24 cm in diam, cream to light brown,
apex acute; peduncle light brown tomentose,
1020 cm long; pollen sporangiophores cuneiform, distal face hexagonal, 0.4 cm long,
fertile abaxial surface 2-lobed with 1016
bisporangiate synangia per lobe. Ovulate
strobili unknown.
Chromosome number.Unknown
its mountainous habitat type, originally
thought to be near Naolinco near the city of
Xalapa, Veracruz in Mexico. It is now known
that this species is not known from Mexico
and is endemic to Guatemala.
Guatemala on rocky outcrops in primary and
secondary evergreen tropical rainforest (Fig. 6).
Distinguishing features.Characterized by
its consistently chartaceous to papyraceous
long acuminate leaets that are strongly
curved near the apex with light serrulations.
Additional specimens examined. GUATEMALA.
ALTA VERAPAZ: H. Frther 2621/592 (NY).
Zamia onanreyesii C. Nelson & G. Sandoval. Ceiba 49(1): 135. 2008. Type:
Honduras. Corts, 7 Jan 2008, O. Reyes
406 (holotype: TEFH).
Zamia bussellii Schutzman, R. S. Adams, J. L.
Haynes & Whitelock. The Cycad Newsletter 31
(2/3), 22. 2008. Type: Honduras. Corts, June
2003, Whittington 2003/01 (holotype: FLAS).
Stem up to 2 m tall, up to 16 cm in diam.

Ptyxis inexed. Leaves 315(44), 60180
1550 cm, erect to slightly curved, tomentulose when emerging; petiole 1540 cm long,
terete, sparsely to moderately armed with
prickles; rachis terete, up to 40120 cm long,
with few prickles along proximal third. Leaflets up to 30 pairs, sessile, subcoriaceous,
oblong-lanceolate, opposite to subopposite,
long acuminate apically, base attenuate, margins serrulate along distal third, subrevolute;
articulations yellowish; median leaets to
[VOL 61
364 cm. Pollen strobili usually 13, erect,

cylindrical, up to 27.5 cm long, up to 4 cm in
diam, light brown to tan, apex acute; peduncle brown to tan, tomentose, up to 8.5 cm
long, up to 1.6 cm in diam; pollen sporangiophore cuneiform, distal face hexagonal
truncate, fertile abaxial surface with up to 9
bisporangiate sori per lobe. Ovulate strobili
usually solitary, erect, cylindrical, up to
43 cm long, 12 cm in diam, brown to
greenish, tomentulose, apex conical; peduncle
densely brown tomentose, up to 5 cm long,
up to 2.5 cm in diam, distal face hexagonaltruncate, 2.9 cm high, 4.7 cm wide. Seeds
ovoid, sarcotesta red at maturity, up to 3 cm
long, up to 2 cm in diam, sclerotesta smooth.
Chromosome number.Unknown.
Distribution and habitat.The species
range is Honduras (Fig. 6), between 01300 m
elevations in evergreen tropical forest.
honor of Onn Reyes, a Honduran biologist.
Distinguishing features.Stems arborescent up to 2 m tall, leaets coriaceous, long
acuminate apically, with margin serrulate.
Nomenclatural note.Both Z. onanreyesii
and Z. busselllii were published in 2008. The
dates of issue are however different. The date
of issue for Z. onanreyesii is 6 September
2008 and the date of issue for Z. bussellii is
16 October 2008. Thus, under Article 29.1 of
the Interantional Code of Botanical Nomenclature (McNeill et al., 2007), Z. onanreyesii
has priority and is used here.
Additional specimen examined.HONDURAS.
Departamento Corts: J. Haynes et al. 044A, 044B
(TEFH).
Zamia oreillyi C. Nelson, Ceiba 46(12): 56.

2005. Type: Honduras. Atlntida: 8 Apr
2006, G. Sandoval et al. 1157 (holotype:
TEFH).
(Fig. 9)
Stem hypogeous, non-branching, up to
25 cm long, up to 7.5 cm in diam. Leaves
usually 1(2), up to 78.2 cm long, 3135 cm
wide, ascending to descending; petiole up to
47 cm long, subterete, armed with small
prickles; rachis subterete, up to 3540 cm
third. Leaets 2931 pairs, sessile, papyraceous to sub-coriaceous, linear-lanceolate,
opposite to subopposite, imbricate, apex
2009]
321
FIG. 9. Zamia oreillyi. A. Habit. B. Cataphyll. CE. Leaet variability. C. Broad and imbricate with dentate
margins. 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).
acuminate, base attenuate, margins dentate to

rarely entire along distal third, up to 0.3 cm,
subrevolute; articulations brown in young
leaets, 0.30.5 cm wide; the median leaets
up to 16 cm long, 1 cm wide. Pollen strobili
usually solitary, decumbent, cylindrical, up to
2.5 cm long and 1 cm in diam, light brown,
apex acuminate; peduncle light brown tomentose, up to 15.9 cm long, 0.2 cm in diam;
truncate-hexagonal, 0.4 cm long, fertile abaxial surface, with 23 bisporangiate synangia per lobe. Ovulate strobili unknown.
322
BRITTONIA

honor of Carlos Manuel OReilly, a Honduran biologist.
Honduras, between 0200 m in evergreen
tropical rainforest (Fig. 6).
strongly imbricate linear-lanceolate to
oblong leaets with dentate margins along
distal third; pollen strobili usually solitary,
decumbent, with fertile section up to 2.9 cm
long and decumbent peduncle up to 15.9 cm
long.
Additional specimen examined.HONDURAS.
ATLNTIDA. Balick 1711 (NY, TEFH).
Zamia paucijuga Wieland, American Fossil

Cycads 2: 212. 1916. Type: Fig. 86 in
American Fossil Cycads 2: 212.1916.
(lectotype, designated by Stevenson &
Sabato, 1986: Fig. 86).
Cataphylls coriaceous, persistent, base triangular, apex aristate, 4.53.4 cm at base,
brown tomentose. Ptyxis inexed. Leaves 2
3, ascending to descending 41952936 cm
wide, brownish when emerging; petiole 10.2
32 cm long, green in young leaves, subterete,
armed with prickles up to 4 mm long; rachis
along the proximal third. Leaets 528 pairs
sessile, coriaceous, lanceolate, opposite to
subopposite, apex acute, base attenuate, margins serrulate to slightly dentate along distal
third, subrevolute; articulations brownish in
young leaets, 0.40.6 cm wide; the median
leaets 14192.33.4 cm. Pollen strobili
usually 12, erect, cylindrical, 6.311 cm
long, 2.12.6 cm in diam, light brown
tomentulose, apex acute; peduncle lightbrown tomentose, 6.3 cm long, 1.2 cm in
diam; pollen sporangiophores cuneiform, distal face hexagonal-truncate, 0.3 cm long,
fertile abaxial surface with 68 bisporangiate
synangia per lobe. Ovulate strobili usually
solitary, erect, ellipsoid to cylindrical, 8.1 cm
long, 5.2 cm in diam, brown-yellowish,
tomentulose, apex apiculate; peduncle densely brown tomentose, 3.9 cm long, 1.3 cm in
diam; megasporangiophores peltate, distal
[VOL 61
face hexagonal-truncate, 0.60.8 cm high,

1.11.5 cm wide. Seeds ovoid, sarcotesta
pink when immature, orange at maturity, 2
smooth.
Chromosome number.2n=23, 34, 25, 26,
27, 28 (Moretti & Sabato, 1984).
Mexico. Known from the states of Nayarit,
Jalisco, Colima, Michoacan, Guerrero, and
Oaxaca (Fig. 6). Found in varied habitats
from dry open woodlands to understory
evergreen tropical forest.
few leaet pairs per leaf.
Distinguishing features.Zamia paucijuga
is highly variable morphologically and in
chromosome number and karyotype (Moretti,
1990a, b). This may represent more than one
entity under the concept of Z. paucijuga
sensu stricto Wieland (1916). This species
shares a number of characteristics with Z.
loddigesii, being small plants with underground contractile stems and having coriaceous
leaets with marginal teeth. Nevertheless, it
differs from Z. loddigesii in having highly
coriaceous leaets with longer marginal teeth,
and seeds with an orange sarcotesta at maturity
as contrasted to the red sarcotesta of Z.
paucijuga. Zamia paucijuga occurs along the
Pacic seaboard of Mexico whereas Z. loddigesii is known generally from the Gulf of
Mexico seaboard.
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-Morejn et al. 1566, 1567, 1568,
1569 (XAL), Jos Azueta, Vovides et al. 1426 (XAL);
Petatln, Vovides et al. 1427, 1428, 1429, 1430, 1431,
1432, 1433, 1434 (XAL); Unin de Isidro Montes De
Oca, Vovides et al. 1416, 1417, 1418, 1420, 1421 (XAL);
La Unin, G. Lozano-Valdez 331 (FCME), J. Jimnez
331 (FCME). JALISCO: Gmez-Pompa 4876 (MEXU); El
Arenal, 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-Morejn et al. 1524, 1525 (XAL); Cihuatln, J. Borocio R. s.n. (ZEA); Cuautitln, Cochrane et al.
10886 (IBUG, WIS, ZEA), R. Cuevas et al 7025 (ZEA),
L. Guzmn & J. Santana M. 745, 947 (ZEA), F.
Nicolalde-Morejn et al. 1528, 1529 (XAL), Prez de
la Rosa 1039 (FCME, IBUG, MEXU), 1040 (FCME,
IBUG, XAL), 1041 (CIB), 1518 (IBUG), Ramrez 425
(IBUG), M. Rosales & L. Cruz 75 (ZEA), J. Santana M.
2009]
et al. 5296 (ZEA); La Huerta, Cuevas et al. 4861 (IBUG,

ZEA); San Sebastin, F. Nicolalde-Morejn et al. 1422
(XAL), 1423 (XAL), 1424 (XAL), 1425 (XAL), 1426
(XAL), 1427 (XAL), Prez de la Rosa 1084 (IBUG,
FCME, MEXU) 1097, 1098 (IBUG); Tuito, A. Flores et
al. 614 (UAMIZ), F. Nicolalde-Morejn et al. 1429
(XAL), Prez de la Rosa 1438, 1439 (IBUG); Vallarta,
Prez de la Rosa 1413, 1415 (IBUG); Villa Puricacin,
Prez 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-Morejn 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-Morejn 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),
565 (XAL), A. Nava-Zafra & J. Pascual 35 (SERO, FCME);
Puerto Escondido, J. Rees 1603 (MO, XAL), Walters sn
(FTG accession 714, XAL); San Gabriel Mixtepec, F.
Nicolalde-Morejn et al. 1474 (XAL); San Pedro Pochulta, J.
Lomel et al. 2967 (MEXU).
Zamia polymorpha D.W. Stev., A. Moretti

& Vzq. Torres, Delpinoa n.s. 3738: 4.
19951996 (issued 1998). Type: Belize.
Cayo: 22 Jan 1989, D. W. Stevenson et al.
1119 (holotype: NY; isotypes: BRH, FTG,
MO, NY, U).
(Fig. 10)
with age, up to 32 cm long, up to 14 cm in
diam. Cataphylls chartaceous, persistent, base
triangular, apex aristate, 36.41.32.6 cm at
base, brown tomentose. Ptyxis inexed.
Leaves 23(4), ascending, 30105 29
45 cm, brown when emerging, green when
mature; petiole 10.295.3 cm long, greenish in
young leaves, subterete, armed with prickles
up to 4 mm long; rachis subterete, up to 67 cm
third. Leaets 312 pairs, sessile, coriaceous,
lanceolate to oblanceolate, opposite to subopposite, apex acute, base attenuate, margins
serrulate along upper third, subrevolute; articulations brown when young, green when
mature, 0.40.8 cm wide; the median leaets
173523.5 cm. Pollen strobili usually 12,
erect, conical, 6.57.3 cm long, 1.11.4 cm in
diam, light to dark-brown tomentulose, apex
acute; peduncle light brown tomentose, 6.8 cm
long, 1.2 cm in diam; pollen sporangiophores
cuneiform, distal face hexagonal-truncate,
323
0.4 cm long, fertile abaxial surface with 45

bisporangiate synangia per lobe. Ovulate
strobili usually solitary, erect, cylindrical to
ovoid, 8.716.3 cm long, 58.3 cm in diam,
dark-brown tomentulose, apex acute; peduncle
brown tomentose, 47.5 cm long, 1.1 cm in
diam; megasporangiophores peltate, distal face
hexagonal-truncate, 0.8 cm high, 1.82.1 cm
wide. Seeds ovoid, sarcotesta pink when
immature, red at maturity, 1.42.1 cm long,
0.50.9 cm in diam, sclerotesta smooth.
Chromosome number.2n=17, 22, 23, 24,
25, 26, 27, 28 (Stevenson et al., 19951996b,
b; Vovides & Olivares, 1996).
Distribution and habitat.This species is
known from Belize, Guatemala and Mexico
from 0200 m elevation. In Mexico, its range
includes the states of Quintana Roo, Yucatn,
Campeche, Tabasco and Chiapas (Fig. 6).
the extreme variation in leaf and leaet morphology presented by this species (Stevenson
et al., 19951996b).
Distinguishing features.Zamia polymorpha shares many morphological attributes
with Z. loddigesii of the Gulf of Mexico
drainage. However, there are clear differences
in the pollen and ovulate reproductive structures. In Z. loddigesii, the pollen strobili are
beige in color with an acute apex and the
ovulate strobili are beige and cylindrical
whereas in Z. polymorpha the pollen strobili
are maroon with an acuminate apex and the
ovulate strobili are dark-maroon and ovoid.
CAMPECHE: Hernndez et al. ES-184 (MEXU), Schutzman 502 (XAL), 503 (XAL), 504 (XAL), 505 (XAL), 506
(XAL), 507 (XAL), 508 (XAL), 509 (XAL); Benito
Jurez, Vovides et al. 1312 (XAL), 1313 (XAL), 1314
(XAL), 1315 (XAL), 1316 (XAL), 1317 (XAL); Calakmul,
Madrid et al. 736 (MEXU), Martnez 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 &
Ucn 1562 (UADY, XAL), Ucn et al. 7293 (UADY,
XAL), 7307 (UADY, XAL), 7398 (UADY, XAL).
CHIAPAS: Palenque, Aguilar & Aguilar 1355 (MEXU), F.
Nicolalde-Morejn & N. Martnez 1419 (XAL), Schutzman
508 (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 et
al. 20075 (MEXU, MO), Tllez 1415 (MEXU), Trejo 225
(CICY, MEXU); Adolfo Huerta, lvarez et al. 9495
324
BRITTONIA
[VOL 61
FIG. 10. Zamia polymorpha. A. Habit (Stevenson et al. 1118, NY). B. Leaets (from the holotype). C. Ovulate
strobilus. D. Pollen strobilus. E. Cataphyll.
(MEXU); Chetumal, Vovides 852 (XAL), Flores & Burgos
9635 (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), Mndez 214 (XAL),
Novelo 169 (MEXU, XAL), Puig 788 (MEXU). Macuspana, Vovides et al. 1344, 1345 (XAL). Yucatn, G. F.
Gaumer 2430 (MO), Lundell & Gentle 827 (MEXU), May
743 (CICY, MEXU); Tekon, Enrquez 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), Vovides
1311 (XAL).
2009]
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).
325

the bright grass-green leaets.
Distinguishing features.Plants with subterranean or epigeal stems, distinctly serrulate
leaet margins and bright-green leaets;
ovulate strobili ovate, green and glabrous
when mature.
Additional specimens examined.BELIZE.
TOLEDO: G. Davidse & A. E. Brant 32179 (MO),
32232 (MO).
Zamia prasina W. Bull, Retail List: 20.

1881. Type: cultivated from Br. Honduras,
W. Bull s.n. (holotype: K).
Zamia purpurea Vovides, J.D. Rees & Vzq.

Torres, Flora de Veracruz 26: 28. 1983.
Type: Mexico. Veracruz: 30 Jan 1982,
Vovides 734 (holotype: XAL).
(Fig. 11)
Stem hypogeous to epigeous, rarely

branching. Cataphylls chartaceous, persistent,
base triangular, apex aristate, 34.51.2
2.7 cm at base, brown tomentose. Ptyxis
inexed. Leaves 24(6), 571002535 cm,
ascending to spreading, brown when emerging; petiole 1230 cm long, brown-greenish
young leaves, subterete, sparsely to densely
along the proximal third. Leaets 1218
pairs, sessile, coriaceous, oblong to oblanceolate, opposite to subopposite, apex acute to
acuminate, base cuneate, margins serrate to
denticulate in the upper two third, subrevolute; articulations brown-yellowish in young
leaets, 0.30.6 cm wide; the median leaets
152046 cm. Pollen strobili usually 12,
erect, cylindrical to ovoid, up to 610 cm
long and 24 cm in diam, light brown, apex
acute; peduncle densely light brown tomentose, 24 cm long, 1.5 cm in diam. Ovulate
strobili usually solitary, erect, cylindrical, 10
15 cm long, 57 cm in diam, green, glabrous
when mature, apex acute; peduncle browngreenish tomentose, 3.5 cm long, 1.5 cm in
diam; megasporangiophores distal face hexagonal-truncate, 11.4 cm high, 1.11.8 cm
wide. Seeds ovoid, sarcotesta light red when
immature, red at maturity, 1.52 cm long,
0.50.8 cm in diam, sclerotesta smooth.
Chromosome number.Not known.
Belize (Fig. 6) on rocky outcrops between
100200 m elevation in evergreen tropical
rainforest.
Stem hypogeous, dichotomously branching

with age, up to 30 cm long, 46 cm in diam.
Cataphylls membranaceous, deciduous, base
triangular, apex acuminate, 5.1.8 cm at
base, reddish-brown tomentose. Ptyxis inexed. Leaves 16, ascending to spreading,
3490 3238 cm, reddish-brown when
emerging, turning green to dark-green at
maturity; petiole 1629 cm long, blackish in
young leaves, subterete, armed with simple
prickles up to 4 mm long; rachis subterete,
1745 cm long, with few prickles along the
proximal third. Leaets 34(6) pairs, sessile,
coriaceous, elliptic to lanceolate, opposite to
subopposite, veins highly prominent on adaxial surface, though relatively inconspicuous
veins have been reported on some individuals, apex acute, base attenuate, margins
dentate along upper third, subrevolute; articulations brown in young leaets, 0.40.8 cm
wide; the median leaets 62728.1 cm.
Pollen strobili usually 12, erect, conical, 2
4.3 cm long, 0.51.1 cm in diam, light-brown
tomentulose, apex acute; peduncle lightbrown tomentose, 23.5 cm long, 0.7
0.9 cm in diam; pollen sporangiophores
cuneiform peltate, distal hexagonal face and
truncate to scutiform, 0.25 cm long, fertile
abaxial surface with 2 bisporangiate synangia
per lobe. Ovulate strobili usually solitary,
erect, conical, 69 cm long, 34.5 cm in
diam, purplish-brown tomentulose when immature turning dark-purple glabrescent when
mature, apex acute; peduncle densely darkbrown tomentose, 2.9 cm long, 1.6 cm in
diam, distal face hexagonal-scutiform, 0.9
326
BRITTONIA
[VOL 61
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.)
2009]
1.2 cm high, 1.62.1 cm wide. Seeds ovoid,

sarcotesta pink when immature, red at maturity, 0.81.1 cm long, 0.60.8 cm in diam,
sclerotesta smooth.
1983)
percentage of polymorphic loci is P=100,
the expected heterozygosity is HE =0.481 and
Distribution and habitat.Zamia purpurea is endemic to Mexico from the Ro
Uxpanapa drainage system within the states
of Veracruz and Oaxaca (Fig. 5) between 50
200 m in evergreen tropical forest (sensu
Rzedowski, 1978) classication.
the dark purple color of the ovulate strobili.
the prominent leaet veins giving the coriaceous leaets a channeled appearance in
contrast to the rest of its congeners in
Mesoamerica, and the dark purple color of
the ripe ovulate strobili.
OAXACA: Santa Mara Chimalapa, S. H. Salas M. 982
(SERO), Snchez et al. 40 (B, MEXU); Santa Mara
Lachixio, Cern et al. 266 (XAL); San Juan Guichicovi, F.
Nicolalde-Morejn & J. Torres 1503 (XAL), 1404 (XAL),
M. Vzquez-Torres et al. 1470 (MO, XAL), Walters s.n.
(FTG accession 101, XAL). VERACRUZ: M. VzquezTorres 4038 (CIB); Hidalgotitln, Calzada 8374 (XAL),
M. Vzquez-Torres et al. 224 (MO); Jess Carranza, A.
Gonzlez-Christen s.n. (CIB), F. Nicolalde-Morejn & J.
Torres 1502 (XAL), M. Vzquez-Torres et al. 2373 (CIB),
M. Vzquez-Torres et al. V-2532 (CHAPA, CIB, XAL).
Zamia sandovalii C. Nelson, Ceiba 46(12):

55.. 2005. Type: Honduras. Atlntida: Jan
2006, G. Sandoval et al. 1154 (holotype:
TEFH)
Stem hypogeous, non-branching, 15.7 cm
long, 10.1 cm in diam. Cataphylls persistent,
base triangular. Leaves 13, ascending to
descending up to 2100.55 cm; petiole 68 cm
long, subterete, armed with small prickles;
rachis subterete, up to 140 cm long, unarmed.
Leaets 68 pairs sessile, sub-coriaceous, lanceolate, opposite to subopposite, falcate, apex
acuminate, base attenuate, margins serrulate
327
along distal third, subrevolute; the median leaflets 163123 cm. Pollen strobili usually 2,
erect, cylindrical, 11.2 cm long, 2.6 cm in diam,
brown-reddish tomentulose, apex apiculate; peduncle light-brown tomentose, 16.7 cm long,
1.2 cm in diam; pollen sporangiophores cuneiform, distal face hexagonal truncate, 0.8 cm
long, fertile abaxial surface with 23 bisporangiate synangia per lobe. Ovulate strobili usually
solitary, erect, ellipsoid to cylindrical, 12 cm
long, 4.5 cm in diam, brown-yellowish, tomentulose, apex long-acuminate; peduncle densely
brown tomentose, 1.5 cm long; megasporangiophores peltate, distal face hexagonal-truncate,
22.5 cm high, 1.82 cm wide. Seeds ovoid,
sarcotesta white when immature, 1.21.8 cm
long, 0.50.7 cm in diam, sclerotesta smooth.
Honduras (Fig. 6), between 200350 m in
evergreen tropical forest.
Germn Sandoval, biologist of the Universidad
Nacional Autonoma de Honduras (Nelson,
2005).
Distinguishing features.This species is
characterized by glabrous, sub-coriaceous,
lanceolate, falcate leaets; cylindric, ovulate
strobili with a strongly acuminate apex; and
seeds with a white sclerotesta.
Additional specimens examined.HONDURAS.
ATLNTIDA: J. Haynes et al. 37 (TEFH), G. Sandoval
et 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 adult
plants, branching dichotomously with age,
3065 cm long, 1031.5 cm in diam. Cataphylls chartaceous, persistent, base triangular,
apex aristate, 7.12.6 cm at base, reddishbrown tomentose. Ptyxis inexed to erect.
Leaves 315 or more per apex, 12019045
62 cm, ascending, distal portion descending
to spreading, brown when emerging turning
green at maturity; petiole 3872 cm long,
green-yellowish in young leaves, terete,
328
BRITTONIA
along the proximal third. Leaets 4152

pairs, sessile, coriaceous, linear-lanceolate,
alternate to subopposite, subfalcate, apex
acute, base attenuate; margins entire, subrevolute; articulations brown in young leaets,
0.40.8 cm wide; the median leaets 1235
0.61.5 cm. Pollen strobili usually 13 per
apex, erect, cylindrical to conical, 915 cm
long, 1.22.4 cm in diam, dark-brown tomentulose, apex apiculate; peduncle light-brown
tomentose, up 7.2 cm long, 1.2 cm in diam;
hexagonal-truncate, 0.45 cm long, fertile
abaxial surface with 5 bisporangiate synangia
per lobe. Ovulate strobili usually solitary,
erect, cylindrical, 1215 cm long, 6.17.3 cm
in diam, dark-brown to reddish, tomentulose,
apex aristate; peduncle brown puberulent,
2.1 cm long, 1.2 cm in diam; megasporangiophores peltate, distal face hexagonaltruncate, 0.70.9 cm high, 1.61.8 cm wide.
Seeds ovoid to angular, sarcotesta white when
immature turning salmon-pink when mature,
sclerotesta light-beige, smooth with 68 light
furrows running longitudinally and sometimes dichotomizing, 2.12.6 cm long, 1.4
1.9 cm diam.
Chromosome number.2n=16 (Schutzman
et al., 1988).
Chiapas, Mexico, between 9001400 m in
the Soconusco mountain range of southern
Chiapas. It inhabits the understory herbaceous
layer of the transition zone between evergreen
tropical forest and cloud forest (Fig. 5).
the Sierra del Soconusco mountain range also
known as the Sierra Madre de Chiapas, being
the region where this species is native
(Schutzman et al., 1988).
Distinguishing features.Zamia soconuscensis is the only species of the genus in
Mexico that approaches an arborescent habit
with leaves that can reach almost two meters
long that gracefully arch toward the terminal
portion. The linear-lanceolate leaets have
totally entire margins; the only other Mexican
congener with entire margins is Z. inermis.
Ovulate strobili are short-pedunculate and solitary with a dark-brown velvety tomentulum.
CHIAPAS: Matuda 2087 (MEXU), 2535 (MEXU), 2590
[VOL 61
(MEXU), 2656 (MEXU); Acacoyagua, Garca 149

(CHIP, MEXU), M. A. Prez-Farrera 141 (CIB, HEM,
MEXU).
Zamia spartea A. DC., Prodr. 16 (2): 539.

1868. Zamia loddigesii var. spartea (A.
DC.) Schuster, Panzenr. 99: 148. 1932.
Type: Mexico. Oaxaca: prope Acayucam,
Verapa, Chimalapi, 1832, Alaman s.n.
(holotype: G-DC).
(Fig. 12).
Cataphylls chartaceous, persistent, base triangular, apex aristate, 6 1.4 cm at base,
25(8) per crown, 35603852 cm, ascending to gracile, reddish-brown when emerging,
turning green at maturity; petiole 1221 cm
long, green-yellowish in young leaves, subterete, heavily armed with straight to sometimes bifurcate prickles up to 4 mm long;
rachis subterete, up to 42 cm long, with few
prickles along the proximal third. Leaets
1527 pairs sessile, coriaceous, linear, alternate to subopposite, apex acute, base attenuate; margins serrulate along extreme distal
portion, subrevolute; articulations lightorange when young, turning yellowish with
age, 0.30.4 cm wide; the median leaets 20
350.30.6 cm. Pollen strobili usually 23,
erect, cylindrical, 6.58.5 cm long, 1.4
1.9 cm in diam, yellowish-beige tomentulose,
apex acute; peduncle densely light-brown
tomentose, 68 cm long, 0.91.1 cm in diam;
hexagonal-truncate, 0.3 cm long, fertile abaxial surface with 1014 bisporangiate synangia per lobe. Ovulate strobili usually
solitary, erect, cylindrical to oval-cylindrical,
912 cm long, 4.24.8 cm in diam, brown
tomentulose, apex acute; peduncle densely
brown tomentose, 3.43.8 cm long, 0.8
1.1 cm in diam; megasporangiophores peltate,
distal face hexagonal-truncate, 0.81.1 cm
high, 11.3 cm wide. Seeds ovoid, sarcotesta
pink when immature, red at maturity, 1.2
smooth.
1983; Moretti, 1990a).
Oaxaca, Mexico. Known in southern areas of
2009]
329
FIG. 12. Zamia spartea. A. Habit. B. Leaet. C. Pollen strobilus. DE. Microsporophyll, abaxial and adaxial
view. FG. Petiole and cataphyll. H. Ovulate strobilus and peduncle. I. Ovulate sporangiophore. J. Ovulate
sporangiophores, distal end truncate hexagonal. K. Seed with sarcotesta. LN. Seeds.
330
BRITTONIA
the Isthmus of Tehuantepec (Fig. 5), between

200400 m elevation, associated with tropical
deciduous forests (sensu Rzedowski, 1978).
Etymology.The epithet is from the broom
genus Spartium (Fabaceae), in reference to
the narrow and tapered leaets of the cycad.
Distinguishing features.Small gracile
plants reaching up to 80 cm tall with very
narrow-linear leaets with an almost entire
margin with a few very small serrulations
only on the extreme distal portion.
OAXACA: Meave del Castillo & Garca 2388 (MEXU);
Matas Romero, F. Nicolalde-Morejn & J. Torres 1505
(XAL), 1506 (XAL), Schutzman 529 (XAL), 530 (XAL),
531 (XAL), 532 (XAL), 533 (XAL), 534 (XAL), 535
540 (XAL), 541 (XAL), 542 (XAL), Vovides & Perales
600 (XAL), Walters s.n. (FTG accession 93, XAL); San
Juan Guichicovi, N. Antonio-Barrera 83b (CIB), Santa
Mara Chimalapa, A. Espejo et al 6485 (UAMIZ), R. Garca
S. 341 (XAL, SERO), Torres 653 (XAL); San Miguel
Chimalapa, M. Vzquez-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, 614 cm
long, 59 cm in diam. Cataphylls chartaceous,
persistent, base triangular, apex long-aristate,
to 12 cm1.5 cm at base, reddish-brown
tomentose. Ptyxis inexed. Leaves 15, 20
100 to 55 cm, slightly to slightly recurved,
tomentulose when emerging; petiole 3560 cm
long, terete, sparsely to heavily armed with
prickles; rachis terete, up to 70 cm long, with
few prickles along proximal third. Leaets 10
15 pairs, sessile, subcoriaceous-coriaceous,
long-lanceolate, opposite to subopposite, recurved, apex acute, base attenuate, margins
dentate along distal third, subrevolute; articulations dark brown when young, 0.30.5 cm
wide; median leaets 204514 cm. Pollen
strobili usually 13, decumbent, cylindrical,
610 cm long, 12 cm in diam, light-brown
tomentulose, apex acute; peduncle light-brown
tomentose, 24 cm long, 1.1 cm in diam;
hexagonal truncate, fertile abaxial surface with
4 bisporangiate sori per lobe; Ovulate strobili
usually solitary, erect, cylindrical to slightly
[VOL 61
ovoid, 812 cm long, 38 cm in diam, brown,

tomentulose, apex long-apiculate; peduncle
densely brown tomentose, 2.54 cm long,
1.3 cm in diam, distal face hexagonal-truncate,
0.71.2 cm high, 1.52.1 cm wide. Seeds
maturity, 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 0200 m in evergreen tropical forest.
Paul C. Standley a prominent botanist of
the ora of Mexico and Central America
(Schutzman, 1989).
Distinguishing features.Its falcate leaets
and conspicuous marginal teeth up to 4 mm
long, and long-apiculate ovulate strobili as
well as its cylindrical characterize this species.
Additional specimens examined.GUATEMALA.
PROVINCIA IZABAL: H. Frther 10234/252 (MSB, W).
HONDURAS. DEPARTAMENTO ATLNTIDA: Tela, V.
Severen 1450 (NA), Standley 53721 (F, US); Puerto
Sierra, Wilson 537 (NY). DEPARTAMENTO SANTA BARBARA: San Pedro Sula, Thieme 144 (US). Departamento
Yoro, Coyoles, T. G. Yuneker et al. 8186 (F, G, GH, MO,
NY). DEPARTAMENTO CORTES: Montaa 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, up
to 100 cm long, 1012 cm in diam. Cataphylls chartaceous, persistent, base triangular,
apex aristate, 8.3 2.7 cm at base, light
brown-greenish tomentose. Ptyxis inexed to
erect. Leaves 815, 100200 2045 cm,
ascending, distal portion descending to
spreading, 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.
Leaets 815 pairs, sessile, papyraceous,
oblong-lanceolate, glossy, iridescent bluegreen, alternate to subopposite, apex abruptly
acuminate, base attenuate; margins entire, subrevolute; articulations green, 0.40.6 cm wide;
2009]
the median leaets 141846 cm x. Pollen

strobili usually solitary, erect, cylindrical, 12
15 cm long, 24 cm in diam, gray-brown at
maturity, apex acute; peduncle light-brown
tomentose, up 6.5 cm long, 1.4 cm in diam.
Ovulate strobili usually solitary, erect, cylindrical, 1218 cm long, 48 cm in diam, brownish
when young, iridescent blue-green at maturity,
apex aristate; peduncle light brown puberulent,
3.4 cm long, 1.5 cm in diam; megasporangiophores peltate, distal face hexagonal-truncate,
1.42 cm high, 33.6 cm wide. Seeds elongateovoid, sarcotesta light red when immature, red
at maturity, smooth, 2.12.6 cm long, 1.3
1.6 cm diam.
Chromosome number.2n =16 (Moretti,
1990b)
Guatemala, between 2501000 m in the Alta
Verapaz department mountain range. It inhabits the understory herbaceous layer of tropical
forest (Fig. 6).
Hans von Terkheim, who collected this
species for the rst time in Guatemala.
Distinguishing features.The principal
distinguishing features of this species are
arborescent stems up to one meter tall,
papyraceous leaets with entire margins and
ovulate cylindrical strobilus with abruptly
acuminate apex and aristate apex, the cone
becoming blue-green iridescent when mature.
Additional specimens examined.GUATEMALA.
ALTA VERAPAZ: T. B. Croat 41647 (MO), H. Frther 11034
(MSB, W), J. A. Steyermark 44484 (MO); Rubeltem, H.
Frther 10918 (MSB, W).
Zamia variegata Warsz., Allg. Gartenzeitung

32: 253. 1845. Type: Mexico. Chiapas:
Lacandona, on border with Guatemala, 12 Jan
1987 D. W. Stevenson 685 (neotype, here
designated: NY; isoneotypes: U, XAL).
Zamia muricta var. picta Miquel, Tijdschr. Wis-en
natuurk. Wet.1(4): 198199. 1848, non Von Houtte
1846. Zamia picta Dyer in Hemsley, Biol. Cent.-Amer.,
Bot. 3(16): 194. 1884. Type: ex Horto Amsterdam
(Z. picta H. Belg.), Miquel s.n. (holotype, U)
Stem hypogeous, 916 cm long, 410.5 cm

in diam. Cataphylls chartaceous, base triangular, apex aristate, 4.67.12.12.4 at base,
12(3), 402912444 cm, ascending; peti-
331
ole 35177 cm long, dark-green with characteristic yellow variegation; petiole subterete,
heavily armed with prickles up to 5 mm long;
rachis subterete, up to 105 cm long, with few
prickles along the proximal third. Leaets 3
10 pairs, sessile, papyraceous, elliptic, opposite to subopposite, dark-green with yellow or
cream variegation, apex acute, base attenuate;
margins dentate along distal third, subrevolute; articulations brown in young leaets
turning green at maturity, 0.40.8 cm wide;
the median leaets 12223.18.8 cm. Pollen strobili up to 6, erect, long-cylindrical,
711 cm long, 1.92.5 cm in diam, yellowish-beige, apex acute; peduncle densely lightbrown tomentose, 36 cm long, 0.70.9 cm
in diam. Ovulate strobili 12 ovoid to
cylindrical, 12 cm long, 4.5 cm in diam,
gray-greenish tomentulose; megasporangiophores peltate, distal face hexagonal-truncate,
0.81.3 cm high, 1.72.1 cm wide. Seeds
ovoid, red at maturity, 1.11.5 cm long, 0.7
1 cm in diam, sclerotesta smooth.
Chromosome number.2n=21, 22 (Moretti
et al., 1991).
the expected heterozygosity is HE =0.355 and
populations currently under study is Fst = 0.085
(Gonzlez-Astorga et al. unpubl. data).
Distribution and habitat.Zamia variegata was described from plants collected in
Guatemala by Warszewicz (Stevenson &
Sabato, 1986). In Mexico, this plant is
known only from Chiapas, in lowland areas
near the Montes Azules Biosphere Reserve
(Fig. 5). Its habitat is evergreen tropical
rainforest (sensu Rzedowski, 1978).
the variegated nature of the leaets, an
attribute unique to this species of Zamia.
Distinguishing features.Its yellow-cream
variegated papyraceous leaets easily distinguish Zamia variegata. The variegations are in
the form of irregular yellow blotches that are
most apparent on the adaxial side of the lamina.
Chiapas: Ocosingo, Castillo-Campos et al. 3848
(XAL), 3855 (XAL), 3885 (XAL), M. Vzquez-Torres
et al. 3924 (CIB); Margaritas, F. Nicolalde-Morejn et al.
332
BRITTONIA
[VOL 61
FIG. 13. Zamia vazquezii. A. Habit. B. Leaet. C. Ovulate strobilus. D. Pollen strobilus. E. Cataphyll. (From the
holotype.)
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. Steyermark
45048 (F, NY). Izabal: J. J. Castillo & D. R. Hodel 2138
(MO), M Vliz 6893 (BIGUA, MEXU).
Zamia vazquezii D.W. Stev., Sabato, A.

Moretti & De Luca, Delpinoa n.s. 3738:
14. 19951996 (issued 1998). Type: Mexico.
Veracruz: 22 Jan 1989, M. Vzquez-Torres et
al. 3990 (holotype: NY; isotypes: FTG, MO,
NY, U, CIB).
(Fig. 13)
2009]
Stem hypogeous, dichotomously branching

Cataphylls chartaceous, persistent, base triangular, apex aristate, 5.93.8 cm at base,
brown tomentose. Ptyxis inexed. Leaves 4
6 to many, 351001629 cm, ascending,
brownish when emerging, turning green at
maturity; petiole 2145 cm long, terete,
unarmed or rarely with prickles, rachis terete,
up to 65 cm long. Leaets 1426 pairs,
sessile, papyraceous, ovate to obpyriform,
opposite to subopposite, apex acuminate,
base cuneate; margins with numerous serrations along distal third, subrevolute, articulations brown in young leaets, 0.50.7 cm
wide; the median leaets 7.114.6 2.8
4.1 cm. Pollen strobili usually 12 per apex,
erect, ovoid to ovoid-cylindrical, up to
10.6 cm long and 2.6 cm in diam, lightbrown tomentulose, apex acute; peduncle
densely light-brown tomentose, 4.5 cm long,
1.2 cm in diam; pollen sporangiophore distal
face hexagonal-truncate. Ovulate strobili usually solitary, erect, cylindrical to ovoid-cylindrical, up to 15 cm long, 7.3 cm in diam, gray
to brown tomentulose, apex apiculate; peduncle densely brown tomentose, 5.3 cm long,
1.4 cm in diam; megasporangiophores peltate, distal face hexagonal-truncate to scutiform, 1.2 cm high, 2.9 cm wide. Seeds ovoid,
sarcotesta pink when immature, orange-red to
red at maturity, 1.6 cm long, 1.2 cm in diam,
sclerotesta smooth.
Chromosome number.2n=18 (Stevenson
et al., 19951996a).
Veracruz, Mexico (Fig. 5), found between
50350 m in evergreen tropical forest sensu
Rzedowski (1978) classication, on predominantly deep clayey soils.
Mario Vzquez-Torres, a Mexican biologist
and cycad specialist.
Distinguishing features.Zamia vazquezii
has more leaves (up to six), longer with each
up to 100 cm long, and wider leaets than Z.
scheri. The two species are disjunct in
distribution and have different chromosome
numbers, Z. vazquezii 2n = 18 and Z. scheri
2n = 16 (Stevenson et al., 19951996a).
333
Species dubium
Zamia verschaffeltii Miq., Verh. Kon. Ned.
Akad. Wetensch., Afd. Natuurk. 2(4): 31.
1870. Type: Mexico. Zamia fusca latifolia
Versch.", Miquel s.n. (holotype, U).
Zamia verschaffeltii forma latifolia Schuster,
Panzenr. 99: 138. 1932. Type: description (lectotype, designated by Stevenson & Sabato, 1986).
Socorro is the locality mentioned by Schuster
(1932) for Miquels (1870) source of Zamia
verschaffeltii, although no locality information
is given on the type specimen, which is the only
specimen at U where Miquels herbarium
resides (Staeu, 1966). The original description
by Miquel gives the source of material as Ex
imperio Mexicano introduxit A. Verschaffelt,
qui in Catalogis Z. fuscam latifoliam dixit and
makes no mention of a specic locality.
However, following the information given by
Schuster (1932), we have conducted exhaustive
searches over the years at two possible localities with the name of El Socorro, one in
Tabasco and the other at Ruta del Socorro in
Veracruz, It is important to note that the whole
area and surrounding regions have been converted into vast sugar cane plantations so that
historically Zamia could have been present but
would now be extirpated. We can condently
say that no species of Zamia were found in or
near the two localities of the name Socorro, and
no other locality of this name was located
within the distribution range of this species
complex in the study. Also, no individuals or
populations of Zamia studied here conform to
the original description of Z. verschaffeltii;
because we were unable to nd another record
of Z. verschaffeltii since its publication in 1870,
we believe that this species is probably extinct
(Nicolalde-Morejn et al., 2008). It is interesting that Schuster actually described a form
latifolia Schuster does not cite any specimens
of this form, but only that it is cultivated in
Garten Verschaffelt. We have been unable to
locate any specimens of this form that were
seen by Schuster, and if one did exist at B it was
destroyed. Therefore, all source material
remains enigmatic.
Acknowledgments

VERACRUZ: Papantla, M. Vzquez-Torres 4568 (CIB),
Allen 19702141 (XAL); Tiguatln, J. Rees 1617 (XAL).
The rst author thanks the Red Latinoamericana de Botnica for the award of a
334
BRITTONIA
Ph. D. fellowship (RLB-06-D2; Systematics

Program, Instituto de Ecologa, A.C., Xalapa,
Mexico) and the Mellon Foundation for a
stipend for a six months residence at the New
York Botanical Garden during 2002. This
research was supported partially by CONACyT-SEMARNAT grant No. 2002-CO1
0183 to AV and NSF Grants BSR-8607049
and EF-0629817 to DWS. The authors thank
Francisco Vergara Silva, Jorge Gonzlez
Astorga, and Victoria Sosa for their comments on a previous version of this manuscript, and Pablo Carrillo-Reyes, Eduardo
Ruiz for their comments on the diagnostic
key to species of Zamia, Carlos Iglesias for
assistance and guidance in the eld, and
Edmundo Saavedra for the illustrations of
Zamia spartea. We thank the Curators and
staffs of the herbaria mentioned for making
their collections available for study, as well as
the staff of the Jardn Botnico Fco. J.
Clavjiero of the Instituto de Ecologa, A.C.
for access to the living specimens of the
Mexican National Cycad Collection. Finally,
the authors would like to thank Roy Osborne
for his salient discussions and diligence in
reviewing this manuscript throughout the
preparation of this work.
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Taxonomic revision of Zamia in Mega-Mexico

FERNANDO NICOLALDE-MOREJN1, ANDREW P. VOVIDES1,

Departamento de Biologa Evolutiva, Instituto de Ecologa, A. C. km 2.5 Antigua Carretera a

According to Stevenson (1992), Zamiaceae

Central America that is oristically similar to

Brittonia, 61(4), 2009, pp. 301335

ISSUED: 1 December 2009

Zamia is the widest distributed genus of the

Materials and methods

NICOLALDE ET AL.: ZAMIA IN MEGA-MEXICO

JFC 1100 sputter coater. Observations were

notable, with branches reaching up to 80 cm

FIG. 1. Trichomes. AB. Zamia furfuracea. CD. Z. katzeriana. EF. Z. polymorpha.

the cone, when mature (Schutzman et al.,

ogy introduced by Mundry and Sttzel (2003)

NICOLALDE ET AL.: ZAMIA IN MEGA-MEXICO

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

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

NICOLALDE ET AL.: ZAMIA IN MEGA-MEXICO

Olivares, 1996). In contrast, all Ceratozamia

Zamia: rst, Norstogs hypothesis (Norstog,

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

Seventy-ve percent of the species in this

Guatemala; (c) Z. loddigesii, ranging along

NICOLALDE ET AL.: ZAMIA IN MEGA-MEXICO

sessile, papyraceous to coriaceous, linear,

hexagonal, 0.30.55 cm long, fertile abaxial

Key to the species of Zamia in Mega-Mexico

18. Leaets lanceolate to oblanceolate, not imbricate, petiolearmed with small

Zamia cremnophila Vovides, Schutzman &

Stem hypogeous, generally unbranched, 5

when immature, red at maturity, 22.8 cm long,

Zamia scheri Miq. in Lem., Hort. Vanhoutt. 1:

NICOLALDE ET AL.: ZAMIA IN MEGA-MEXICO

FIG. 5. Distribution of Zamia cremnophila ( ), Z. furfuracea ( ), Z. inermis (), Z. katzeriana ( ), Z. lacandona

Stem subterranean, dichotomously branching

to ovoid, up to 9 cm long, 4.5 cm in diam, brown

with papyraceous but serrulate leaets and

Zamia furfuracea L. f. in Aiton, Hortus

Stem hypogeous, becoming epigeous with

usually 24, erect, cylindrical, up to 17 cm

NICOLALDE ET AL.: ZAMIA IN MEGA-MEXICO

Zamia herrerae Caldern & Standl., Proc.

be found in secondary growth forests and

Stem hypogeous, bifurcate in adult plants,

FIG. 6. Distribution of Zamia scheri (), Z. herrerae ( ), Z. loddigesii ( ), Z. oreillyi ( ), Z. paucijuga ( ), Z.

cate, 0.70.9 cm high, 1.11.3 cm wide. Seeds

Nicolalde-Morejn & Vovides 1415 (XAL), 1416 (XAL),

Stem hypogeous, unbranched, up to 25 cm

NICOLALDE ET AL.: ZAMIA IN MEGA-MEXICO

FIG. 7. Zamia scheri A. Habit. B. Leaets. (From Stevenson 566, NY).

emerging, dark green when mature; petiole

NICOLALDE ET AL.: ZAMIA IN MEGA-MEXICO

the two populations currently under study is

Zamia lacandona Schutzman & Vovides,

pairs, sessile, coriaceous, lanceolate, opposite

Additional specimens examined.MEXICO.

Zamia loddigesii Miq., Tijdschr. Natuurl.

1028b (lectotype, designated by Stevenson &

Stem hypogeous, branching dichotomously

NICOLALDE ET AL.: ZAMIA IN MEGA-MEXICO

states of Tamaulipas, Hidalgo, Veracruz,

Additional specimens examined.MEXICO.