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Key words: Species boundaries, Palmae, Arecaceae, AFLP, Chamaedorea alternans, Chamaedorea tepejilote, endemics,
conservation, Los Tuxtlas, Mexico.
INTROD UCTION
Over the past 15 years, phylogenetics has become the
primary focus for much of systematic botany, and results
from these studies have prompted major advancements
towards our understanding of higher level relationships
(e.g. Chase et al., 1993; Soltis et al., 1999; Qiu et al.,
2000; Soltis and Soltis, 2004). Progress in higher level
phylogenetics has been made in Arecaceae (Palmae), and
many relationships among subtribes and genera are now
robustly supported (Asmussen et al., 2000; Baker et al.,
2000a,b; Lewis and Doyle, 2001; Hahn, 2002a,b; Lewis
and Doyle, 2002; Gunn, 2004; Dransfield et al., 2005;
Roncal et al., 2005).
Despite these gains, many issues at lower taxonomic
levels continue to hinder complete understanding of palm
systematics. Inadequate understanding of both the limits
between closely related species and relationships among
them have been attributed to a growing crisis in the field
of taxonomy that directly impinges on the development
of justified recommendations in conservation (e.g. Tautz,
2002, 2003; Gower and Wilkinson, 2003). In fact, the
limited progress toward the understanding of species limits
in most plant groups may be accredited to both recent focus
on phylogenetic research as well as the scale of the
problem (Scotland, 2003). In-depth evaluation of hundreds
of thousands of species is far more demanding than broad
*
The Author 2006. Published by Oxford University Press on behalf of the Annals of Botany Company. All rights reserved.
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Background and Aims The establishment of justified recommendations in conservation biology requires robust
taxonomic treatments for the group(s) being considered. Controversial or poorly developed taxonomies can have a
negative impact on conservation assessments. One example of a taxonomically difficult and controversial species
complex that is important in conservation involves two species of Mexican palms, Chamaedorea tepejilote and
C. alternans. The goal of this study was to investigate whether C. alternans and C. tepejilote are genetically distinct
within the Los Tuxtlas region of Veracruz.
Methods Individuals corresponding to the morphology of C. alternans and C. tepejilote were collected from
sympatric and allopatric regions within the Los Tuxtlas Biological Station. Eighty-eight samples were genotyped
using amplified fragment length polymorphism (AFLP) markers. Cluster and ordination analyses were used to
investigate patterns of differentiation.
Key Results UPGMA and PCO analyses of AFLP profiles recovered two divergent clusters corresponding to
morphologically defined C. tepejilote and C. alternans. No intermediate genotypes were observed and five of the
45 loci were fixed for either the presence or absence between the species. The patterns of divergence observed do
not identify a role for sympatric speciation.
Conclusions The observed patterns of differentiation support the recognition of C. alternans as distinct from
C. tepejilote. A suite of vegetative and reproductive morphological features can be used to help distinguish these taxa
in the field, but they can be difficult to differentiate from preserved material. Previous treatments of the variation
found within the complex (C. alternans and C. tepejilote) as intraspecific variance is unjustified. Given that the
Los Tuxtlas region has suffered from historical and ongoing deforestation and that Chamaedorea includes numerous
endangered taxa, retaining conservation status for C. alternans serves to help safeguard individuals of the species
as well as a region and larger taxonomic group all under considerable threat from human activities.
756
MATERIA LS A ND METHODS
Materials and sampling
DNA samples from leaf tissue collected from populations of both putative taxa were analysed using AFLP
markers. Forty-four samples of each potential species
were selected to represent the geographic distribution
across which all samples were collected. Plant material was
ground using liquid nitrogen, and total genomic DNA was
extracted using a DNeasy Plant Mini Kit (Qiagen). DNA
concentrations were estimated by comparison with a
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9210
Area
enlarged
1840
9205
757
9200
PUNTA ORGANO
RANCHERIA
MONTE PIO
1840
1835
72
73 71 69 67
VOLCAN SAN
MARTIN TUXTLA
BARRA DE
SONTECOMOPAN
1835
LA PALMA
SONTECOMOPAN
1830
1830
LOS AMANTES
SANTIAGO
TUXTLA
LAGUNA ENCANTADA
SINUAPAN
SAN ANDRES
TUXTLA
1825
CATEMACO
Km
10
9215
1825
20
9210
9205
9200
F I G . 1. Map of the Los Tuxtlas Biological Station, adapted from Lot-Helgueras (1976).
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ESTACION DE
BIOLOGIA TROPICAL
LOS TUXTLAS
AFLP fragments*
Fixed presence fragments*
Coloration of leaf sheath
Venation in leaf sheath
Inflorescences per node
Life form
Habitat
Chamaedorea
tepejilote
Chamaedorea
alternans
39
3
Green
Vague
Singular
Multiple stem
Rocky substrate
35
2
White
Distinct
Multiple
Solitary
Rich, developed soil
Chamaedorea tepejilote
RESULTS
072
06
048
036
024
012
Eight accessions per species were rerun from the preselective amplification stage to test for reproducibility
of AFLP fingerprints. In a replicate UPGMA analysis,
including both independently derived profiles for the same
sample, 14 of the 16 grouped with their duplicate. Two
accessions failed to group with their replicate but each
remained within their respective UPGMA cluster.
DISCUSSION
Genetic differentiation
Accessions representing the two divergent clusters recovered in both UPGMA and PCO analyses (Figs 2 and 3)
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Chamaedorea alternans
758
759
Axis 1
26
21
15
10
26
21
15
10
05
05
10
15
21
Axis 2
26
05
10
15
21
26
F I G . 3. Principal coordinates analysis diagram based on Euclidean distance. Chamaedorea alternans is indicated by triangles and C. tepejilote
by squares.
Using data derived from field and herbarium observations, Hodel (1992) concluded that C. alternans represents
a distinct taxon. Chamaedorea alternans was distinguished
from C. tepejilote primarily by differentiation in leaf
sheath colouring and number of inflorescences per node.
Chamaedorea alternans has leaf sheaths that are whitish
and exhibit distinct green venation, whereas C. tepejilote
has green leaf sheaths with no obvious venation. Also,
C. alternans has multiple inflorescences per node, whereas
C. tepejilote has singular inflorescences per node
(Fig. 4). In addition to these more persistent features,
Hodel (1992) also used ephemeral floral characteristics,
including the number and orientation of branches in
pistillate rachillae and the spacing of pistillate flowers, to
help differentiate these taxa.
More recently, Henderson et al. (1995) addressed the
status of C. alternans using observations from available
herbarium specimens. These authors concluded that
specimens of putative C. alternans were insufficiently
distinct from those of C. tepejilote to justify recognition
of the former. Subsequently, Henderson et al. treated all
representatives of this group as C. tepejilote, which has
nomenclatural priority over C. alternans.
Both the genetic data and field-based morphological
observations generated in this study confirmed the utility
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05
760
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ACKNOWLEDGEMENTS
This research represents a partial fulfilment of the
requirements for the degree of Master of Science in
biology at NMSU and was supported by the International
Palm Society, the NMSU Biology Departement and the
Loustaunau Memorial Fellowship. The authors thank
M. Murphy Thomas and Tom Parchman for providing
constructive advice, Richard Spellenberg, Brook Milligan
and Michael Fay for editorial support, John Dransfield
and Scott Zona for their encouragement with the manuscript, and Geovanni Romero for technical assistance.
The efforts of two anonymous reviewers greatly improved
the manuscript. We are also grateful to research staff at
Estacion de Biologa Los Tuxtlas (UNAM), herbaria at
UNAM, Los Tuxtlas and NYBG, and to Barbara Ayala
and Argelia Cuenca Navarro for their assistance with the
voucher specimens.
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