"CIRCUNSCRIPCIN DE LAS ESPECIES DEL COMPLEJO Zamia katzeriana

(ZAMIACEACE-CYCADALES)"





TESIS QUE PRESENTA EDISON FERNANDO NICOLALDE MOREJN PARA
OBTENER EL GRADO DE MAESTRO EN CIENCIAS EN


SISTEMTICA



Xalapa, Veracruz; Mxico 2005


2




MAESTRA EN CIENCIAS
(SISTEMTICA)

Aprobacin del documento final de tesis de grado:

"Circunscripcin de las especies del complejo Zamia katzeriana (Zamiaceae-
Cycadales)"



Nombre
Director de Tesis Dr. Andrew Peter Vovides Papalouka .
Dr. Dennis W. Stevenson .
Dra. Victoria Sosa Ortega .
J urado Dr. Miguel Angel Prez Farrera .
Dr. Gonzalo Castillo Campos .


km 2.5 Carretera antigua a Coatepec No. 351 Congregacin El Haya, A.P. 63 Xalapa 91070, Veracruz,
Mxico. Tel. (228) 842 18 00 Fax (228) 818 78 09
3

AGRADECIMIENTOS

Este trabajo fue posible gracias a una beca de Maestra otorgada por la Red
Latinoamericana de Botnica (RLB-03-M2), y de un apoyo parcial del proyecto
CONACyT-SEMARNAT, No. 2002-CO1-0183 proyecto del segundo autor.

Agradezco a Andrs Vovides por el apoyo durante toda mi estancia en Mxico, a Dennis
Stevenson por sus sugerencias, a Victoria Sosa por la orientacin durante el desarrollo de
ste trabajo, a Miguel ngel Prez Farrera por su apoyo logstico en el trabajo de campo y
anlisis de los resultados, a Gonzalo Castillo-Campos por sus comentarios al documento.

Mi gratitud a J orge Gonzles-Astorga, Carlos Iglesias-Delfn por soportar mis dilemas
nomenclaturales, a Mario Miguel Ojeda, J ess Hernndez por su apoyo y revisin del
diseo estadstico y por supuesto a mi hija Zamia y mi esposa Rub, a Fernando Vaz de
Mello y Pablo Carillo-Reyes por todas la discusiones taxonmicas planteadas al respecto.








4







DECLARACIN

Excepto cuando es explcitamente indicado en el texto, el trabajo de investigacin
contenido en esta tesis fue efectuada por el Ing. Edison Fernando Nicolalde Morejn como
estudiante de la carrera de Maestra en Ciencias (Sistemtica) entre septiembre de 2003 y
noviembre de 2005, bajo la supervisin del Dr. Andrew Peter Vovides Papalouka.

Las investigaciones aqu reportadas en esta tesis no han sido utilizadas
anteriormente para obtener otros grados acadmicos, ni sern utilizados para tales fines en
el futuro.

Candidato: Ing. E. Fernando Nicolalde Morejn .
Director de tesis: Dr. Andrew Peter Vovides Papalouka .


km 2.5 Carretera antigua a Coatepec No. 351 Congregacin El Haya, A.P. 63 Xalapa 91070, Veracruz,
Mxico. Tel. (228) 842 18 00 Fax (228) 818 78 09
5
NDICE
I. ndice de Tablas y Figuras..6
II. Resumen.7
III. Introduccin8
IV. Species Circumscription in the Zamia katzeriana complex (Zamiaceae-Cycadales)
10
Abstrac...11
Resumen12
1. Introduction...13
2. Materials and Methods..14
2.1 Populations selection.14
2.2 Character selection.15
2.3 Analysis..16
3. Results...17
3.1 Distribution and habitat..17
3.2 Univariate analysis.18
3.3 Principal components analysis (CPA)...19
3.4 Discriminant analysis.19
4. Discussion and conclusions...20
5. Taxonomic treatment.22
6. Acknowledgments.24
7. References.24
V. Discusin y Conclusiones.46
VI. Bibliografa....48
6

NDICE DE TABLAS Y FIGURAS

Table I. Populations selected from the entire area of distribution of Zamia katzeriana
complex29
Table II. Morphological characters selected30
Table III. Qualitative morphological characters..31
Table IV. Principal components analysis.33
Table V. Component weightings......34
Table VI. Discriminant function..35
Table VII. Summary of the discriminant analysis from populations in the Z. katzeriana
complex.36

Figure legends
Figure 1. Zamia katzeriana holotype..38
Figure 2. Zamia verschaffeltii holotype 39
Figure 3. Species distribution of the Zamia katzeriana complex in south east Mexico..40
Figure 4. Plot of analysis using the date matrix of qualitative and quantitative from all
populations and all individuals..41
Figure 5. Scatter plot of derived functions scores produced by stepwise discriminant
analysis of 12 morphometric ratios from populations in the Zamia katzeriana complex42

Appendix 1. Specimens examined..43

7
RESUMEN

En este artculo se caracteriza la variacin morfolgica de las especies de un complejo de
Zamia spp. que presentan fololos anchos y coriceos con distribucin en el sur-este de
Mxico, la mayora de reciente descripcin. El complejo es denominado Zamia katzeriana,
por una coleccin histrica de este nombre, y consiste de Zamia cremnophila, Z.
lacandona, Z. splendens y Z. purpurea. Zamia splendens a partir de la dcada de los ' 90 fue
propuesta en sinonimia bajo dos colecciones histricas Zamia katzeriana y Zamia
verschaffeltii, de las cuales se desconoce con exactitud su lugar de procedencia, excepto
que fueron colectadas en Mxico por colectores alemanes del siglo XVIII. La informacin
en especial sobre el holotipo incompleto de Zamia verschaffeltii es muy ambigua y la
posible localidad tipo de nombre "Socorro" mencionada por Schuster es imprecisa, lo cual
ha generado confusiones taxonmicas. La caracterizacin de la variacin morfomtrica
incluy a todas las poblaciones (11) e individuos (115) actualmente conocidas para el
complejo en todo su rango de distribucin en Mxico. El anlisis discriminante incluyendo
a las colecciones histricas, determina que Zamia verschaffeltii no esta relacionado
morfomtricamente con ninguna especie del complejo y Zamia katzeriana est
estrechamente correlacionada con Zamia splendens. Finalmente, se presenta un revisin
taxonmica para el complejo.





8
INTRODUCCIN

Mxico representa la regin de mayor diversidad de Ccadas del neotrpico, con
aproximadamente 50 especies distribuidas en tres gneros, Ceratozamia (23 spp) y Dioon
(13) mayormente distribuidos en Mxico y Zamia con 15 especies de las 59 actualmente
conocidas para el gnero.

Zamia es el gnero mas diverso de las ccadas americanas, y su distribucin comprende
todo el neotropico. Este gnero evidencia variaciones morfolgicas y ecolgicas muy
grandes, razn por la cual ha sido motivo de atencin de muchos botnicos desde el siglo
XVII; contrariamente su taxonoma se presenta muy inestable, en especial de taxa descritos
entre los siglos XVIII y XIX.

Varias especies de Zamia han sido descritas para Mxico en las dos ultimas dcadas, en
especial desde reas poco exploradas como el sur-este mexicano, entre las cuales podemos
citar a: Zamia cremnophila, Z. lacandona, Z. purpurea y Z. splendens, siendo el grupo de
especies de nuestro particular inters, por compartir una misma rea biogeogrfica y un
juego de caracteres morfolgicos en especial vegetativos que dificultan su reconocimiento.
Esto ha sido motivo para que estudios taxonmicos de estas entidades sean planteados
como un conjunto de especies a los cuales se ha denominado complejo.

No obstante, bajo un contexto de la problemtica nomenclatural que ha enfrentado el
gnero durante todo su historia taxonmica, las especies antes mencionadas no son la
excepcin a pesar de su reciente descripcin; tal es el caso de Zamia splendens que ha sido
9
sinonimizada varias veces desde la dcada de los 90 del siglo pasado hasta la actualidad,
bajo el nombre de un taxon el cual presumiblemente fue colectado en el sur-este de Mxico
a mediados del siglo XIX y del cual actualmente se dispone de un pobre e incompleto
ejemplar tipo nomenclatural; el cual, de corroborar su validez biolgica y nomenclatural
prevalecera sobre cualquier especie de reciente descripcin.

Aunque, el gnero Zamia y en especial las Ccadas en general son consideradas especies
carismticas por toda la historia evolutiva que representan y adems por estar en peligro de
extincin, poco nfasis se ha dado a este grupo desde el punto de vista nomenclatural. Por
lo tanto, la meta de esta investigacin es circunscribir las especies que para este caso hemos
denominado complejo, considerando en el anlisis tanto a las colecciones histricas
relacionadas con el grupo de estudio, como nuevas exploraciones y colectas con el intento
de aclarar la procedencia de dichas colecciones. Sin duda, estas decisiones contribuirn
para la continuacin de estudios de tipo filogentico y de conservacin.










10

Species circumscription in the Zamia katzerianacomplex (Zamiaceae-Cycadales)

FERNANDO NICOLALDE-MOREJ N, ANDREW P. VOVIDES, DENNIS W.
STEVENSON, VICTORIA SOSA

Nicolalde-Morejn, F., A. P. Vovides, V. Sosa (Instituto de Ecologa A.C., Km 2.5
Carretera Antigua a Coatepec No 351, Congregacin El Haya, Xalapa, Veracruz, C.P.
91070, Mxico,e-mail: nicolalde@posgrado.ecologia.edu.mx) & D. W. Stevenson (The
New York Botanical Garden, Bronx, New York, 10458-5126. U.S.A., e-mail:
dws@nybg.org). Species circumscription in the Zamia katzeriana complex (Zamiaceae-
Cycadales). The complex is denominated the Zamia katzeriana species complex in
reference to an historic collection of this name, distributed within south eastern Mexico.

Key words: Holotype, morphometric characterization, discriminant analysis, Mexico,
Zamia.







11
ABSTRACT

The morphological variation of recently described species within a complex of Zamia spp.
that present wide and coriaceous leaflets distributed within south eastern Mexico is
analysed. The complex is denominated the Zamia katzeriana species complex in reference
to an historic collection of this name, and consists of Zamia cremnophila, Z. lacandona, Z.
splendens and Z. purpurea. During the 1990s Z. splendens was proposed under synonymy
of two historical collections; Zamia katzeriana and Zamia verschaffeltii, either of which
precise locality is unknown other than collected in Mexico by German collectors during the
XVIII century. Information especially of the incomplete Z. verschaffeltii holotype is very
ambiguous and the possible type locality named Socorro mentioned by Schuster is
imprecise, thus generating taxonomic confusion. Morphometric characterization and
discriminant analysis of the contemporary and historic collections (Z. katzeriana and Z.
verschaffeltii) also including all the known populations (11) and individuals (115) of the
complex throughout its range has shown that Z. verschaffeltii is not morphometrically
related to any of the species in the complex but Z. katzeriana is closely related to Z.
splendens. Finally a taxonomic revision is presented for the complex.

Key words: Zamia, Mexico, holotype, morphometric characterization, discriminant
analysis.




12
RESUMEN

En este artculo se caracteriza la variacin morfolgica de las especies de un complejo de
Zamia spp. que presentan fololos anchos y coriceos con distribucin en el sureste de
Mxico, la mayora de reciente descripcin. El complejo es denominado Zamia katzeriana,
por una coleccin histrica de este nombre, y consiste de; Zamia cremnophila, Z.
lacandona, Z. splendens y Z. purpurea. Zamia splendens a partir de la dcada de los ' 90 fue
propuesta en sinonimia bajo dos colecciones histricas Zamia katzeriana y Zamia
verschaffeltii, de las cuales se desconoce con exactitud su lugar de procedencia, excepto
que fueron colectadas en Mxico por colectores alemanes del siglo XVIII. La informacin
en especial sobre el holotipo incompleto de Zamia verschaffeltii es muy ambigua y la
posible localidad tipo de nombre "Socorro" mencionada por Schuster es imprecisa, lo cual
ha generado confusiones taxonmicas. La caracterizacin de la variacin morfomtrica
incluy a todas las poblaciones (11) e individuos (115) actualmente conocidas para el
complejo en todo su rango de distribucin en Mxico. El anlisis discriminante incluyendo
a las colecciones histricas, determina que Zamia verschaffelttii no esta relacionado
morfomtricamente con ninguna especie del complejo y Zamia katzeriana est
estrechamente correlacionada con Zamia splendens. Finalmente, se presenta un revisin
taxonmica para el complejo.





13
There are 59 known species in Zamia has (Hill et al., 2004) and is the most widespread
cycad genus of the Neotropics, ranging between latitudes 30 N and 18 S. It occurs in
Georgia and Florida in the USA, the Greater Antilles, Mexico, Central America, Colombia,
Venezuela, Ecuador, Peru, Brazil and Bolivia (Balduzii et al., 1982; Sabato, 1990; Norstog
& Nicholls, 1997). Zamia also has the greatest morphological, ecological and cytological
variation within the cycads (Vovides, 1983; Moretti & Sabato, 1984; Moretti, 1990;
Vovides & Olivares, 1996).

The last complete taxonomic treatment (study) for the neotropical species of Zamia was by
Schuster (1932); nevertheless, posterior works (Sabato, 1990; Stevenson, 1990, 1991, 2001,
2004; Norstog & Nicholls, 1997) and the typification of valid names (Stevenson & Sabato,
1986) have thrown evidence on nomenclatural and taxonomical errors of this treatment
mainly due to insufficient field work and a paucity of collections.

Various local treatments on the neotropical Zamiaceae have since appeared (see Vovides et
al., 1983; Stevenson, 1991, 1993a, 1999, 2001, 2004). However none of these treatments
have dealt with the problem of the identity of Zamia katzeriana (Fig. 1), Zamia
verschaffeltii and Z. splendens, mainly because the existing information on the type locality
especially that of Z. verschaffeltii reported by Schuster (1932) as well as the description and
incomplete sterile holotype are very ambiguous (Fig. 2).

The binomial Z. verschaffeltii has been in use for over 15 years (Stevenson et al., 1990;
1993b; 1995; Hill et al., 1998; Osborne, 1999; Hill et al., 2004a; Hill et al., 2004 b;
Osborne et al., 2005;), but in no contemporary species descriptions within the complex
14
(Vovides et al., 1983; Schutzman, 1984; Vovides et al., 1988; Schutzman & Vovides, 1998)
has a possible relation with Z. katzeriana and Z. verschaffeltii with any other species been
discussed. However, since the publication of the world cycad list by Hill et al. (1998) and
considering publication date priorities, Zamia katzeriana and Zamia splendens are included
as synonyms under Z. verschaffeltii, which status has been upheld by the Cycad Specialist
Group (Hill et al., 2004a; Hill et al., 2004b; Stevenson et al., 2004, Osborne et al., 2005 in
press). The main polemic has been centered on the synonymy of Z. splendens under Z.
verschaffeltii as discussed and refuted by Schutzman (2004) whose argument is based
solely on the ambiguity of the Z. verschaffeltii holotype and type locality.

Because of the lack of a revision that corroborates the taxonomic and nomenclatural status
of these entities, the goal of this research has been aimed at defining the identities and
relationships of Z. katzeriana, Zamia verschaffeltii and Z. splendens. In doing so the study
also included Zamia cremnophila Vovides, Schutzman & Dehgan, Z. purpurea Vovides,
J .D. Rees & Vzq. Torres, and Z. lacandona Schutzman & Vovides. The latter species have
been included in the analysis beacuse these taxa form a complex of species based on
morphological similarities and also share the same biogeographical distribution in
southeastern Mexico (A. Vovides, 2004 pers. comm.) (Fig. 3), especially in the arc
florisitic refuge of Wendt (1987).

Materials and Methods

POPULATION SELECTION
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The populations chosen for the study correspond to the Zamia katzeriana/splendens
complex suggested by Vovides (2004 pers. com.). The inclusion of Zamia variegata in the
analysis suggested by Stevenson (2004 pers. com.) with the premise that it forms part of
this complex, even though from a different geographical distribution. We justify the
inclusion of Z. paucijuga because a locality by the name of Socorro exists in Oaxaca,
which forms part of the geographical distribution of this species (and the possible type
locality for Zamia verschaffeltii, Schuster 1932).

A total of 115 individuals from 11 known populations of the Z. katzeriana complex was
studied (Table I), which includes the total number of populations and individuals therein
currently known for Mexico. Also, specimens held in the living cycad collection at the
J ardn Botnico Fco. J avier Clavijero, Instituto de Ecologa, A. C., were examined as well
as herbarium specimens from: B, CHIP, ECOSUR, ENCB, HEM-UNICACH, IBUG,
MEXU, K, XAL, LE, NY. U.

CHARACTER SELECTION

Twelve morphological characters were selected (Table II). Distance measurements were
made manually by using calipers or a ruler. These variables represent a suite of
characteristics that have been used for diagnosing species within Zamia. Furthermore,
consideration was taken into account that the specimens represented in herbaria are usually
sterile, and also that taxonomic treatments have placed major emphasis on foliar characters
(Miquel, 1861; De Candolle, 1868; Schuster, 1932; Standley, 1923; Eckenwalder 1980;
Vovides, 1983; Newell 1986; Schutzman, 1998; Stevenson, 1993, 2001, 2004). Also, 28
16
qualitative morphological characters were coded as binary state (presence/absence) and
multistate from foliar and reproductive structures as well as habit (Table III).

ANALYSIS

A univariate exploratory analysis was performed with the aim of assessing the variation of
individuals within and between species, using a students t-test. A univariate analysis was
performed using Statistica for Windows, version 6.0

A principal components analysis (PCA) was done to identify which variables and the
percentage of each that are contributing to the variation within the species complex
(eigenvectors and eigenvalues from the correlation matrix after linear standarization of the
original data). The analyses were based on two different data sets: 1) Quantitative
characters, 2) both quantitative and qualitative characters. Correlation matrixes were
generated for matrix 1, to evaluate the variance of the new factors extracted successively
and to explain the variance percentage of each component. A principal components analysis
(PCOA) was done on matrix 2 to evaluate grouping patterns of the species defined a priori,
a scenario that could not be observed with matrix 1. The factors that explained most of the
variation were extracted from the correlation matrix as well as the contribution from each
variable. All quantitative variables were log (base 10) transformed because they were not
normally distributed. All analyses were done using Statistica for Windows v. 6.0.

Finally, to explore and corroborate the position of Zamia katzeriana and Z. verschaffeltii,
(where only historical information on origin and vegetative characteristics defining them as
17
taxonomic entities exists) a discriminant analysis was applied where all the variables
including those from Miquel (1869) and Retting (1896) and the existing holotypes were
considered as just one more case in the analysis (Figs. 1, 2). The final graphic was obtained
using Statigraphic Plus v. 2.0 then edited in Photoshop v. 6 for Macintosh.

Results

DISTRIBUTION AND HABITAT

Zamia katzeriana (=Z. splendens) is the most widely distributed species of the complex at
an altitudinal range between 200 and 700 m as well as presenting the highest range of leaf
and leaflet measurements. It is present in the states of Chiapas, Tabasco (Vovides et al.,
2003) and Veracruz, with the latter locality being a new record for the species. Agricultural
expansion and road building have decimated populations, especially in the areas close to
the type locality of Z. splendens Schutzman (1984).

Zamia cremnophila represents a micro-endemic for the state of Tabasco, especially on the
rocky limestone outcrops (mogotes) of southern Tabasco that have steep slopes and
efficient drainage. Zamia lacandona is from remnants of tropical rain forest of northern
Chiapas and occurs on more or less plane clayey substrates with little humus. Altitudinal
ranges for this species fluctuate from 85 to 270 m. Zamia purpurea is known only from the
Uxpanapa area that is common to the states of Veracruz and Oaxaca on deep clayey soils
with little humus. Finally, Z. variegata, represented by few records of this species in
18
Mexico, and is found on limestone outcrops and humus-rich soils in the Lacandona rain
forest, Chiapas.

UNIVARIATE ANALYSIS

Zamia lacandona shows the longest leaflets of the three populations evaluated and leaflet
mid-region and apex also show high variation between individuals of the same population
and between populations of the same species. With respect to these variables, Z.
cremnophila and Z. purpurea are homogenous between and within populations, whereas
Zamia katzeriana (splendens) and Z. variegata, show slight variation between their
populations.

Leaflet width (up to 12 cm) at base, midway, and leaf apex separate Z. katzeriana from the
rest of the species in the complex, values that are maintained constant between populations
2 and 3 of this species. Zamia lacandona, Z. cremnophila, Z. purpurea and Z. variegata
show little variation for this variable, and their populations are very homogeneous, with the
exception of one individual of Z. variagata with a leaflet width of up to 9 cm. Z.
paucijuga shows narrower leaflets.

Leaf length separates Z. cremnophila (up to 170 cm long) from the rest of the species,
where Z. katzeriana and Z. lacandona have individuals with leaf length up to 180 cm, and a
third group such as Z. purpurea and Z. paucijuga with shorter leaves (up to 80 cm).


19
PRINCIPAL COMPONENTS ANALYSIS (PCA)

According to the correlations matrix, component 1, represented by the character suite in the
leaves of basal, midway and apical leaflet lengths, as well as leaflet articulation width
explain 36.9% of the total variation. Component 2, represented by number of leaflets,
width of basal, and median leaflets of the leaves contribute 27.2%. Finally component 3,
represented by number of leaves, leaf length and petiole length contribute 14.6%.
Collectively, the three components explain 78.6% of the observed morphological variation
of the complex (Tables IV, V). The combined matrix analysis (qualitative and quantitative)
presents a better grouping resolution (Figure 4), which permits us to easily characterize the
species included in this study through component 1 and 2 which explain 55% of the total
variation and that have the following qualitative characteristics for component 1: cataphyll
persistence, petiole type, adult plant leaflet shape, type of megastrobilus peduncle,
microstrobilus color and peduncle type; for component 2: cataphyll apex shape, leaf
orientation with respect to a vertical axis, petiole color in adult plants, leaflet arrangement
along the rachis, seed sarcotesta color when ripe and habit.

DISCRIMINANT ANALYSIS

Figure3 shows the configuration of the groups derived from the discriminant analysis,
supported by four discriminant functions that contribute significantly (p <0.01) for the
discrimination of three groups (Tables VI, VII). Four of the five species form one large
group, in which overlap is observed according to the variables analyzed, and these species
represent the Z. katzeriana complex. Zamia paucijuga is clearly separated from this
20
complex and Z. variegata also forms an isolated group from the rest of the species
analyzed. The prediction places Zamia verschaffelttii amongst individuals of Z. paucijuga,
and Z. katzeriana amongst populations of Z. splendens (Fig. 4).

Discussion and Conclusions

Throughout the Z. splendens and Z. verschaffeltii polemic, (Hill et al., 2004, Osborne et al.,
2005; Schutzman, 2004), there is not one instance of discussion concerning the position of
Zamia katzeriana under synonymy of Z. verschaffeltii (Hill et al., 1998, 2004a, 2004b) and
its relationship with Z. splendens (Schutzman, 1984; Schutzman, 2004).

According to our morphological studies and discriminant analysis (Fig. 4), Z. katzeriana
shows very close morphometric relationships to Z. splendens, especially to the San
Fernando population of Chiapas, whose morphometric parameters are constant and
consistent with the attributes of the holotype and original description of Z. katzeriana (=
Ceratozamia katzeriana Regel). Furthermore, the qualitative characters observed in the
holotype (see Figure 1) such as; conspicuous marginal teeth of the leaflets, cuneate leaf
base, brilliant cuticle on adaxial leaflet surface a unique attribute to this species of the
genus in Mexico and throughout the neotropics.

Based on the evidence presented and invoking the publication date criterion we conclude
that Zamia katazeriana Regel (Retting) has nomenclatural preference of valid species over
Zamia splendens Schutzman.

21
Socorro, a locality mentioned by Schuster (1932) for the Miquelian Z. verschaffeltii type
opens up a possibility to find an origin for the species; however eight localities with this
name are known in Mexico, only two of which are in cycad habitat, the other six being in
northern Mexico, in arid zones, where no cycads have been reported. The two probable
localities are in southeastern Mexico; La Ruta del Socorro and El Socorro in southern
Veracruz and Tabasco respectively.

In the case of the Veracruz locality, no Zamia spp. were found, the whole area and
surrounding regions converted into vast sugar cane plantations. However 20 km west of
this locality in the vicinity of the city of Tuxtepec, Oaxaca, only plants affin to Z. sylvatica
Chamberlain of the Zamia loddigesii Miq. complex were located. In the case of El
Socorro, Tabasco, in the vicinity of the small town of J alapa, all the adjacent areas have
been converted into immense pasturelands, no remaining relict forest was found nor any
cycads in the scarce secondary vegetation of the region. At El Socorro ranch, a cultivated Z.
loddigesii was found growing in the ranchers garden which we suspect came from the
Macuspana region to the west of El Socorro, a known locality for this species.

We can confidently say that no Zamia spp was 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 under study. Also no individual or population of Zamia
studied conforms to the original description nor the Z. verschaffeltii type. We are therefore
in accord to the standpoint of Schutzman (2004), and conclude that there is no synonymy
relation between Z. verschaffeltii and Z. splendens. Furthermore, on being unable to find
22
another record of Z. verschaffeltii since its publication in 1870, we propose that this taxon
be considered extinct.

Taxonomic Treatment

Zamia katzeriana (Regel) Retting, Gartenflora 45: 148-149. 1896.
Type: ex Horto Katzer., Regel s.n. (Lectotype: LE)

Ceratozamia katzeriana Regel, Acta Horti Petrop. 4(4): 298. 1876.
Type. ex Horto Katzer., Regel s.n. (Lectotype: LE)
Zamia splendens Schutzman, Phytologia 55(5): 299-303. Fig. 1. 1984
Type. Cultivated in Fairchild Tropical Garden, Miami, accession no.
FTG 76-1046, J. Watson s.n. (Holotype: NY; Isotype, FLAS, FTG, MEXU)

Dioecious plant up to 1.6 m tall; trunk subterranean up to 25 cm long, 7 cm in diameter.
Cataphylls chartaceous, semidecidious, base triangular apex aristate, 1.4 cm wide, 5.3 cm
long, with yellowish indument. Leaves 1-2 (3), adpressed, 49-220 cm long, 35-58 cm
wide, juvenile leaves bright pink; petiole terete with simple prickles; rachis 35-75 cm long,
inermis, with 3-7 leaflet pairs. Leaflets coriaceaous, oblong-lanceolate, opposite to
subopposite 18-35 cm long, 4.5-12 cm wide midway along leaf, venation inconspicuous,
adaxial surface with brilliantly shining cuticle, sessile, base cuneate, apex accuminate,
asymmetric, margin denticulate along distal half, subrevolute; articulation 0.6-1.6 cm wide.
Microstrobili 1-5, conical, 3.9 cm long, 1.1 cm in diameter, brown tomentulose, apex
accute; microsporophyll cuneiforme, distal end hexagonal-truncate; peduncle 3.8 cm long,
23
descending, yellowish. Megastrobilus usually solitary, 8-12 cm long, 4.5-6 cm in diameter,
elliptic, brown, tomentulose, apex aristate up to 0.8 cm; megasporophylls 2.6X1.5 cm,
cuneate-peltate, distal end hexagonal-truncate; peduncle 11-14 cm long, descending,
tomentose. Seeds obovoid, 1.2 cm long, 1.8 cm in diameter, sarcotesta pink when
immature, red upon maturity. Chromosome number 2n =16.

Distribution and habitat - The widest geographically distributed species of the
complex, occurring in Chiapas, Veracruz and Tabasco, and between altitude ranges of 200
to 700 m. However, the populations are poorly conserved in their natural state.

Representative especimens examined: MEXICO. VERACRUZ: Las Choapas,
Rubn Martnez & N. Martnez M. 825 (UNICACH); F. Nicolalde et al. 1436, 1437 (XAL).
TABASCO: Macuspana, A. Vovides et al. 1344, 1345 (XAL). Teapa, Fco. Hdez-Najarro
622 (CHIP); Miguel A. Prez F. sn (XAL); Miguel A. Prez F. 899 (MEXU, UNICACH);
T. Walters sn (FTG accession 12-2, XAL). CHIAPAS: San Fernando, A. Vovides et al.
1266 (XAL); E. Palacios E. 383 (CHIP); F. Nicolalde & Miguel A. Prez F. 1420 (XAL);
M. A. Prez F. sn (XAL); T. Walters sn (FTG accession 23-2, XAL). Malpaso, A. Gmez-
Pompa 705 (MEXU); F. Nicolalde et al 1453, 1454, 1455, 1456, 1457, 1458, 1459, 1460
(XAL). Ocozocoautla, Miguel A. Prez F. 29 (CHIP, MEXU). Tila, A. Vovides et al. 1340,
1343, 1341 (XAL).

Diagnostic characters - This species has a subterranean trunk, adpressed leaves
bright pink when emergent, leaflets up to 12 cm wide with very shiny cuticles and dentate
24
along the distal half of the leaflet; micro-and megastrobili descendent. This species has the
widest and highest leaflet variation of the complex.

Acknowledgements

This research was supported by a grant (RLB-03-M2) from the Red Latinoamericana de
Botnica (RLB-03-M2) and partially by CONACyT-SEMARNAT grant No. 2002-CO1-
0183 awarded to the second author. The author expresses thanks to J orge Gonzles-
Astorga, Mario Miguel Ojeda, Miguel A. Prez-Farrera, Carlos Iglesias-Delfin, and J ess
Hernndez, Gonzalo Castillo-Campos for their help on statstic design and revision of the
manuscript and Nayeli Martnez for all the logistic help for field work.

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Num. 2 - J une 2004. 8-11.
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(ZAMIACEAE, CYCADALES) from southern Mexico. Bot. Gaz. 149(3): 347-360.
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Chiapas, Mxico. Novon 8, 441-446
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------------------1993a. The Zamiaceae in Panam with comments on
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Cromosome numbers and Karyotypes. American J ournal of Botany. 70: 1002-1006
Vovides, A. P., J. D. Rees & M. Vzquez-Torres. 1983. Zamiaceae, Flora de
Veracruz. Fascculo 26. INIREB, Xalapa, Veracruz.
Vovides, A. P & M. Olivares. 1996. Karyotipe polymorphism in the cycad Zamia
loddigesii (Zamiaceae) of the Yucatan Peninsula, Mxico. Botanical J ournal
of the Linnean Society. 120: 77-83.
28
Vovides, A. P., M. A. Prez-Farrera., C. Iglesias., S. Avendao & Salas-Morales S.
2003. New Cycad (Zamiaceae) reports from Chiapas, Oaxaca and Tabasco, Mxico.
Rhodora 105: 71-76.
Wendt, T. 1987. Las selvas de Uxpanapa, Veracruz-Oaxaca, Mxico: evidencia de
refugios florsticos Cenozoicos. Anales del Instituto de Biologa UNAM (Serie
Botnica) 58: 29-54


















29

TABLE I
POPULATIONS SELECTED FROM THE ENTIRE AREA OF DISTRIBUTION OF
Zamia katzeriana COMPLEX.
Species State No. of
populations
No of
Individuals
Z. cremnophila Tabasco 2 20
Z. lacandona Chiapas 2 20
=Z. splendens Chiapas,Veracruz 3 32
Z. purpurea Oaxaca, Veracruz 2 20
Zamia katzeriana Mxico Holotype and
description
1
Z. verschaffeltii Mxico Holotype and
description
1
*Zamia paucijuga Oaxaca 1 12
*Zamia variegata Chiapas 1 11
* These species do not conform to the Z. katzeriana species complex






30

TABLE II
MORPHOLOGICAL CHARACTERS SELECTED

Character Code
Number of leaves per plant NL
Leaf length LL
Petiole length PL
Number of leaflets per leaf NLF
Length of leaflets at leaf base LLB
Length of leaflets at leaf
midway
LLM
Length of leaflets at leaf apex LLA
Width of leaflets at leaf base WLB
Width of leaflets at leaf midway WLM
Width of leaflets at leaf apex WFA
Distance between leaflets DL
Articulation width AW




















31
TABLE III
QUALITATIVE MORPHOLOGICAL CHARACTERS

Character State of character
1 - Cataphyll apex: (0) aristate; (1) acuminate
2 - Cataphyll consistency: (0) chartaceous; (1) membranaceous
3 - Cataphyll persistence: (0) semi-deciduous; (1) deciduous
4 - Leaf orientation: (0) adpressed; (1) ascending; (2)
descending
5 - Petiole type: (0) terete; (1) subterete
6 - Petiole color in adult plants: (0) green; (1) brown; (2) purple
7 - Presence of prickles on rachis: (0) absence; (1) presence
8 - Leaflet texture: (0) coriaceous; (1) chartaceous
9 - Leaflet form in adult leaves: (0) oblong-elliptic; (1) elliptic; (2)
Linear-lanceolate
10 - Venation: (0) conspicuos; (1) inconspicuos
11 - Leaflet base: (0) cuneate; (1) attenuate; (2) cuneate to
attenuate
12 - Leaflet apex: (0) acuminate; (1) acute
13 - Presence of sinuous margin: (0) present; (1) absent
14 - Leaflet arangement on rachis: (0) imbricate; (1) non-imbricate
15 - Megastrobilus shape: (0) ellipsoid; (1) conical; (2) cylindrical
16 - Peduncle habit (megastrobilus): (0) descending; (1) erect
32
17 - Megastrobilus color: (0) brown; (1) purple; (2) green; (3)
yellow

18 - Distal face of megasporophyll: (0) truncate; (1) scutiform; (2)
protuberant.
19 - Megastrobilus apex: (0) aristate; (1) acute; (2) apiculate;
(3) mucronate.
20 - Megastrobilus shape: (0) cone; (1) conical; (2) conical-
cylindrical; (3) cylindrical
21 - Microstrobilus Color: (0) creamy; (1) purple; (2) brown
22 - Microstrobilus apex: (0) mucronate; (1) acute; (2) truncate.
23 - Peduncle habit (microstrobilus): (0) descending; (1) erect; (2) decumbent
24 - Sarcotesta color (ripe seeds): (0) red; (1) orange.
25 - Adaxial cuticle: (0) lustrous; (1) non-lustrous.
26 - Leaflet variegation: (0) presence; (1) absence
27 - Bulbose petiole base: (1) presence; (0) absence
28 Habit: (0) hipogeal; (1) semi-hipogeal.







33
TABLE IV
PRINCIPAL COMPONENTS ANALYSIS
Component
Number
Eigenvalue Percent of Variance Cumulative Percentage
1 4.42557 36.880 36.880
2 3.25819 27.152 64.031
3 1.7489 14.574 78.606
4 0.787914 6.566 85.171
5 0.476508 3.971 89.142
6 0.390987 3.258 92.401
7 0.24894 2.075 94.475
8 0.238251 1.985 96.461
9 0.213003 1.775 98.236
10 0.117933 0.983 99.218
11 0.059461 0.496 99.714
12 0.0343366 0.286 100.000







34
TABLE V
TABLE OF COMPONENT WEIGHTS
Character Component 1 Component 2 Component 3
NH -0.0752502 -0.164739 -0.441456
LH 0.304574 0.108935 -0.484207
LP 0.246457 0.0197849 -0.553604
NF 0.100931 0.487297 -0.0823356
LFB 0.369359 0.19868 0.158082
LFM 0.359071 0.238206 0.152202
LFA 0.341607 -0.407671 0.265383
AFB 0.275796 -0.422145 0.140114
AFM 0.283815 -0.389967 0.104798
AFA 0.286783 -0.389967 0.166342
IF 0.302971 -0.197849 -0.275905
AA 0.343425 0.189332 0.0664224








35
TABLE VI
DISCRIMINANT FUNCTION
Discriminant
Function
Eigenvalue Relative
Percentage
Canonical
Correlation
1 7.40869 54.75 0.93866
2 3.51754 25.99 0.88241
3 1.33662 9.88 0.75633
4 0.659817 4.88 0.63049
5 0.340681 2.52 0.50409
6 0.173926 1.29 0.38491
7 0.0625471 0.46 0.24262
8 0.0270593 0.20 0.16232
9 0.00501395 0.04 0.07063
10 0.0000590585 0.00 0.00768









36
TABLE VII
SUMMARY OF THE DISCRIMINANT ANALYSIS RESULTS OF THE ANALYSIS
FROM POPULATIONS IN THE Z. KATZERIANA COMPLEX
Functions
Derived
Wilks Lambda Chi-Square DF P-Value
1 0.003932 576.0152 110 0.0000
2 0.0330629 354.5716 90 0.0000
3 0.149363 197.7430 72 0.0000
4 0.349005 109.4775 56 0.0000
5 0.579285 56.7799 42 0.0635
6 0.776636 26.2894 30 0.6603
7 0.911714 9.6126 20 0.9747
8 0.968739 3.3031 12 0.9930
9 0.994952 0.5263 6 0.9975
10 0.999941 0.0061 2 0.9969





]


37

Figure legends:

Figure 1. Zamia katzeriana holotype
Figure 2. Zamia verschaffeltii holotype
Figure 3. Species distribution of the Zamia katzeriana complex in south east Mexico.
Figure 4. Plot of analysis using the date matrix of qualitative and quantitative from all
populations and all individuals.
Figure 5. Scatter plot of derived functions scores produced by stepwise discriminant
analysis of 12 morphometric ratios from populations in the Zamia katzeriana complex














38
Figure 1


39
Figure 2



40
Figure 3











41
Figure 4

L1
K1
C1
P1
P1
PA
Va1
P2
Va1
L2
K2
C2
P2
P2
K3
-8 -6 -4 -2 0 2 4 6 8 10
Fact or 1: 33.26%
Simbologa: Zamia cremnophila (C1, C2); Z. katzeriana (K1, K2, K3); Z. lacandona (L1, L2);
Z. paucijuga (PA); Z. variegata (Va)
-8
-6
-4
-2
0
2
4
6
8
10
F
a
c
t
o
r

2
:

2
0
.
9
6
%










42
Figure 5













43
Appendix 1. Specimens examined

Zamia cremnophila Vovides, Schutzman & Dehgan

MEXICO. TABASCO: M. A. Magaa & S. Zamudio 343 (MEXU, MO); Teapa, Miguel A.
Prez F. 900 (MEXU, UNICACH).

Zamia katzeriana (Regel)Retting

MEXICO. Regel s.n. (LE); CHIAPAS: J. B. Watson 1870 (NY). San Fernando. A. Vovides
et al. 1266 (XAL); E. Palacios E. 383 (CHIP); F. Nicolalde & Miguel A. Prez F. 1420
(XAL); M. A. Prez F. sn (XAL); T. Walters sn (FTG accession 23-2, XAL). Malpaso, A.
Gmez-Pompa 705 (MEXU); F. Nicolalde et al 1453, 1454, 1455, 1456, 1457, 1458, 1459,
1460 (XAL). Ocozocoautla, Miguel A. Prez F. 29 (CHIP, MEXU). Tila, A. Vovides et al.
1340, 1343, 1341 (XAL). TABASCO: Macuspana, A. Vovides et al. 1344, 1345 (XAL).
Teapa, Fco. Hdez-Najarro 622 (CHIP); Miguel A. Prez F. sn (XAL); Miguel A. Prez F.
899 (MEXU, UNICACH); T. Walters sn (FTG accession 12-2, XAL). VERACRUZ: Las
Choapas, Rubn Martnez & N. Martnez M. 825 (UNICACH); F. Nicolalde et al. 1436,
1437 (XAL).

Zamia lacandona Schutzman & Vovides

MEXICO. CHIAPAS: Palenque, B. Schutzman 510, 511, 512, 513, 514, 515, 516, 517,
518, 519, 520 (XAL); F. Nicolalde & N. Martnez 1418 (XAL); Miguel A. Prez F. 890
44
(MEXU, UNICACH); T. Walters sn (FTG accession 14-2, XAL). San J ernimo Tulija, J.
Chavelas et al. ES=315 (ENCB, MEXU); B. Schutzman 521, 522, 524, 523, 525 (XAL).

Zamia paucijuga Wieland

MEXICO. GUERRERO: J ose Azueta, A. Vovides et al. 1426 (XAL); Petatlan, A. Vovides
et al. 1427, 1428, 1429, 1430, 1431, 1432, 1433, 1434 (XAL); Unin de Isidro Montes De
Oca, A. Vovides et al. 1416, 1417, 1418, 1420, 1421 (XAL); Zihuatanejo, G. Castillo C. et
al. 6633 (XAL). JALISCO: El Arenal, G. Castillo C. et al. 9822 (XAL); Cabo Corrientes,
G. Castillo C. et al. 10147, 11733, 10280, 10466 (XAL); Cuautitln, J. A. Prez de la Rosa
1039, 1040, 1518 (IBUG, XAL); Nayarit, A. P. Vovides et al. 1487, 1488, 1489, 1490,
1491, 1493 (XAL); San Sebastin, F. Nicolalde et al. 1422, 1423, 1424, 1425, 1426, 1427,
1429 (XAL); J. A. Prez de la Rosa 1084, 1097, 1098 (IBUG, XAL). OAXACA: Loxicha,
F. Nicolalde et al. 1422, 1423, 1424, 1425, 1426, 1427, 1429 (XAL). Pochutla, B.
Shutzman 544, 543, 545, 546, 547, 548, 550, 551, 552, 553, 554, 555, 556, 557, 558, 560,
561, 562, 563, 565 (XAL); Puerto Escondido, T Walters sn (FTG accession 7-14, XAL).

Zamia purpurea Vovides, J .D. Rees & Vzq

MEXICO. OAXACA: Santa Mara Chimalapa, A. Snchez A. et al. 40 (B, MEXU); Santa
Mara Lachixio, M. Cern et al. 266 (XAL). San J uan Guichicori, T. Walters sn (FTG
accession 10-1, XAL). VERACRUZ: A. P. Vovides 743 (XAL); J. Rees et al. 1654 (IEB);
Hidalgotitln, J. I. Calzada 8374 (XAL-MEXU); J ess Carranza, M. Vzquez et al. V-2532
(CHAPA, XAL).
45

Zamia variegata Warszewicz

MEXICO. CHIAPAS: Ocosingo, G. Castillo C. et al. 3884, 3885 (XAL); Margaritas, F.
Nicolalde et al. 1443, 1444. 1445, 1446, 1447, 1448, 1449, 1450, 1451, 1452 (XAL)

Zamia verschaffeltii Miq.

MEXICO. Miquel s. n. (U)















46
DISCUSIN Y CONCLUSIONES

De acuerdo a nuestros anlisis morfolgicos y discriminantes (Fig. 5), Z. katzeriana
muestra estrecha relacin con Z. splendens, especialmente con la poblacin localizada en
San Fernando-Chiapas, cuyos parmetros morfomtricos son constantes y consistentes con
los atributos que presenta el holotipo y descripcin original de Z. katzeriana (=Ceratozamia
katzeriana Regel). Adems, caracteres cualitativos observados en el holotipo (ver figura 1),
tales como: dientes marginales conspicuos en los fololos, base de la hoja cuneada, cutcula
brillante sobre la base adaxial de los fololos como nico atributo para esta especie del
gnero en Mxico y todo el neotropico confirman su relacin con Z. splendens.

Con base en la evidencia presentado y por criterios de prioridad de publicacin, nosotros
concluimos que Zamia katazeriana Regel (Retting) tiene preferencia nomenclatural de
especie valida sobre Zamia splendens Schutzman.

Socorro una localidad mencionada por Schuster (1932) como localidad tipo para la
especie Miquelina Zamia verschaffeltii, abri la posibilidad para encontrar un origen para
esta especie; sin embargo, ocho localidades con este nombre fueron registradas para
Mxico, de las cuales solamente dos representan un hbitat potencial para Ccadas, las otras
seis estn distribuidas en el norte de Mxico, en zonas ridas, donde ninguna ccada ha sido
reportada. Las dos probables localidades estn en el sureste de Mxico; La Ruta del
Socorro y El Socorro en el sur de Veracruz y Tabasco.

47
En el caso de la localidad de Veracruz, ninguna especie de Zamia fue encontrada, debido a
que toda la regin y zonas aledaas fueron convertidas en plantaciones de caa de azcar.
Sin embargo, a 20 km al oeste de esta localidad en la reas cercanas a la ciudad de
Tuxtepec, Oaxaca, se localizaron plantas afines a Z. sylvatica Chamberlain, especie que
pertenece al complejo Zamia loddigessi Miq. En el caso de El Socorro, Tabasco, en la
reas cercanas al pueblo de J alapa, no fue posible encontrar ni un solo relicto de bosque y
tampoco especie de Zamia alguna, ya que toda la regin fue convertida en inmensas
pasturas. Sin embargo, en el jardn del rancho El Socorro, una planta cultivada de Zamia
loddigessi fue encontrada, la cual sospechamos viene desde la regin de Macuspana
ubicada hacia el oeste del El Socorro, una localidad conocida para esta especie.

Nosotros confidencialmente podemos decir que ninguna especie de Zamia fue encontrada
cerca de las dos localidades de nombre Socorro, y en ninguna otra localidad de este
nombre dentro del rango de distribucin del complejo de especies en estudio. Tambin
ningn individuo o poblacin de Zamia estudiadas conforme a la descripcin original y el
tipo de Z. verschaffeltii. Por lo tanto, nosotros estamos de acuerdo con el punto de vista
planteado por Schutzman (2004), y concluimos que entre Z. verschaffeltii y Z. splendens no
existe ninguna relacin de sinonimia. Sin embargo, al no disponer de ningn otro registro
para Z. verschaffeltii desde su publicacin en 1870, nosotros proponemos que este taxon
debe ser considerado extinto.




48
BIBLIOGRAFA

Miquel, F. A. W. 1870. Z. verschaffeltii sp. Nov. Archives Neerlandaises des Sciences
Exactes et Naturales vol. 1 p. 458
Regel, E. 1876a. Cycadearum generum specierumque revisio. Acta Horti petro. 4(4): 273-
320
Regel, E. 1876b. Cycadearum. Acta Horti petro. Tomus IV, Fasciculus II. 298.
Rettig .1896. Die Cycadeen des Botanischen Gartens in Petersburgo. Gartenflora 45: 149-
149
Schuster, J. 1932. Cycadaceae. In. Engler. Das Pflanzenreich 99 (IV. 1): 1-168.
Schutzman, B. 2004. The History of Zamia splendens. The Cycad Newsletter. Vol. 27
Num. 2 - J une 2004. 8-11.