Blackwell Publishing LtdOxford, UKBOJBotanical Journal of the Linnean Society0024-4074The Linnean Society of London, 2006?

2006
151?
103111
Review Article

MORPHOMETRICS IN PALM SYSTEMATICS

A. HENDERSON

Botanical Journal of the Linnean Society, 2006, 151, 103–111.

The Palms
Guest edited by William J. Baker and Scott Zona

Traditional morphometrics in plant systematics and its

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role in palm systematics
ANDREW HENDERSON*
New York Botanical Garden, Bronx, NY 10458, USA

Received April 2005; accepted for publication November 2005

A review is given of the role of traditional morphometrics in plant systematics. The three most commonly used tech-
niques of data analysis – Cluster Analysis, Principal Component Analysis and Discriminant Analysis – are dis-
cussed. The kinds of data that can be taken from palm specimens and the problems of using specimens as data
sources are outlined. Published systematic studies of palms using traditional morphometrics are reviewed. More
recent studies indicate that: hybrid zones between species may be common; infraspecific diversity is greater than
previously suspected; there may be more than double the currently accepted number of species; and our current
knowledge of morphological variation in palms is superficial. A procedure for scientific systematics is given, which
incorporates traditional morphometric methods. © 2006 The Linnean Society of London, Botanical Journal of the
Linnean Society, 2006, 151, 103–111.

ADDITIONAL KEYWORDS: Arecaceae – Cluster Analysis – Discriminant Analysis – geometric morphomet-

rics – numerical taxonomy – Principal Component Analysis – Palmae.

INTRODUCTION are size and shape variables (i.e. distances or angles

between homologous landmarks) and also include
Morphometrics can be defined as the quantitative
qualitative variables, usually taken from herbarium
analysis of biological form. It has been widely used in
specimens. I emphasize that my perspective is that of
a variety of disciplines, including systematics. The
a herbarium systematist and this bias means I am
field has developed rapidly over the last 20 years, to
interested in both size and shape variables. Other
the extent that we now distinguish between tradi-
morphometricians consider that the field concerns
tional morphometrics (e.g. Marcus, 1990) and the
only the analysis of shape (e.g. Rohlf, 1990).
more recent geometric morphometrics (e.g. Rohlf &
I use the term variable for any qualitative (binary
Marcus, 1993; Adams, Rohlf & Slice, 2004). Geometric
or multistate) or quantitative (continuous, meristic)
morphometrics, in which information about the rela-
attribute. Although I distinguish between qualitative
tive spatial arrangement of landmarks is preserved, is
and quantitative variables, the distinction is not
not widely used in plant systematics (Jensen, 2003)
always clear (Stevens, 1991; Thiele, 1993). Qualitative
and has not yet found its way into palm systematics. It
variables can also be referred to as characters or
will not be discussed further here.
traits, in the sense of the Phylogenetic Species Con-
In this paper I shall be concerned solely with tradi-
cept (PSC) of Nixon & Wheeler (1990). Characters are
tional morphometrics as used by plant systematists,
qualitative variables that are found in all comparable
and with the descriptive, as opposed to the phyloge-
individuals within a terminal lineage (i.e. species);
netic, component of systematics. The data employed
traits are qualitative variables that are not distri-
buted universally amongst comparable individuals
*E-mail: ahenderson@nybg.org within such a lineage. I return to this topic later.

© 2006 The Linnean Society of London, Botanical Journal of the Linnean Society, 2006, 151, 103–111 103
104 A. HENDERSON
The methods of analysis in traditional morphomet-
rics are primarily multivariate statistics. Although
the terms morphometrics and multivariate statistics
are sometimes used interchangeably – and some
authors speak of multivariate morphometrics – it is
possible to analyse morphometric data without multi-
variate statistical methods (Rohlf, 1990). I give a brief
review of traditional morphometrics in plant systema-
tics, and of some of the most commonly used methods
of analysis. I then concentrate on palm systematics
and discuss the kinds of data used, and give examples
of traditional morphometric analyses used to address
systematic problems in palms. I conclude by outlining
what I take to be a scientific methodology for descrip-
tive systematics, and show how morphometrics is an
integral part of this method.
HISTORICAL OVERVIEW OF TRADITIONAL
MORPHOMETRICS IN PLANT SYSTEMATICS
Some of the earliest papers in plant systematics using
morphometrics were associated with the school of
numerical taxonomy. The development of this school
in the late 1950s and its expanding influence in the
1960s and 1970s (e.g. Sneath & Sokal, 1973) led to an
increase in the use of morphometrics and multivariate
statistics. This coincided with the increased availabil-
ity of computers capable of rapid analysis of large
datasets. However, most of the effort in numerical tax-
onomy, and the source of its conflict with other schools
of systematics, concerned the classification of taxa (i.e.
phylogeny), rather than the descriptive component of
systematics. The unfortunate legacy of numerical tax-
onomy is that many researchers associate morphomet-
ric methods with the demise of that school of
systematics. Similarly, many associate the term phe-
netic with numerical taxonomy, and phenetics or
numerical phenetics both refer to numerical taxon-
omy. I prefer therefore not to use the term phenetic.
In the last 30 years, independently of numerical tax-
onomy, there have been numerous studies using mor-
phometrics, approached from a number of different
perspectives. Commonly, these methods are used in
the study of species complexes (e.g. Chandler & Crisp,
1998; Naczi, Reznicek & Ford, 1998; Raulings & Ladi-
ges, 2001; Gengler-Nowak, 2002) and hybrids (e.g.
Murrell, 1994; Campbell & Wright, 1996; Marhold
et al., 2002). Recently, researchers have combined
molecular and morphological data (e.g. Hansen, Elven
& Brochman, 2000; Lihová et al., 2004). In most cases,
the methods used in these studies have solved
problems that proved intractable using the methods
of traditional herbarium systematics. Traditional
monographs and revisions, published in such series as
Flora Neotropica and Systematic Botany Monographs ,
very seldom employ morphometric methods.
© 2006 The Linnean Society of London, Botanical Journal of the Linnean Society, 2006, 151, 103–111
MOST COMMONLY USED MORPHOMETRIC
METHODS
There are two important discussions of morphometric
methods as applied to botanical systematics: those of
Pimentel (1981) and Crisp & Weston (1993). More gen-
eral reviews are provided by Marcus (1990) and James
& McCulloch (1990). An excellent and detailed review
of the field of multivariate statistics, from an ecologi-
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cal perspective, is that of Legendre & Legendre (1998).
The computer program NTSYSpc (Rohlf, 2000) is
designed specifically for morphometric analysis of sys-
tematic data.
Here I discuss morphometric techniques, at least as
far as systematics is concerned, under three headings,
depending on the data used and the aims of the study.
(1) Cluster Analysis of qualitative data with no a pri-
ori knowledge of specimen groupings. (2) Ordination of
quantitative data (sometimes with qualitative data)
with no a priori knowledge of specimen groupings. (3)
Discriminant Analysis of quantitative data with a pri-
ori knowledge of specimen groupings. The first two are
exploratory analyses (i.e. they are not necessarily
inferential but merely look for patterns in the data),
while the third is confirmatory, with inferential anal-
ysis. First, I discuss some of the assumptions of these
methods, and then give a brief outline of them as
applied in plant systematics.
Difficult problems for systematists using multivari-
ate statistics are the underlying assumptions of the
methods, particularly if they are being used inferen-
tially. The most important requirement for statistical
inference is that the sample be random. Although
some authors insist on this (e.g. Marcus, 1990), sam-
ples of herbarium specimens cannot be random in the
strict sense – ‘a sample collected from a population in
such a manner that every individual in the population
has the same probability of being sampled’ (Pimentel,
1979). A second requirement for statistical inference is
that data be distributed in a multivariate normal fash-
ion. This again is a difficult requirement to meet. Mul-
tivariate normality means that each and all variables
combined are normally distributed (Tabachnik &
Fidell, 2001). This is not readily tested, although some
multivariate procedures are relatively insensitive to
deviations from normality (Tabachnik & Fidell, 2001).
Tabachnik and Fidell give discussions of assumptions
for each multivariate procedure.
Generally, authors of systematic papers using mul-
tivariate statistical methods have ignored the require-
ments of random sampling and other assumptions.
Perhaps the problem is solved by using only noninfer-
ential methods. Pielou (1984), for example, makes the
distinction between interpreting the data at hand
using cluster analysis and ordination, and using mul-
tivariate statistics based on inference. She considers

the latter activity to be a separate field of enquiry (see
also James & McCulloch, 1990). On the other hand,
Tabachnik & Fidell (2001) place much less emphasis
on the importance of random sampling. They write,
‘Use of inferential and descriptive statistics is rarely
an either-or proposition’.
CLUSTER ANALYSIS
Cluster Analysis (CA) is an exploratory tool for classi-
fying objects. It is an example of a Q-mode type of
analysis, in which the association amongst objects
(specimens) is being assessed (Legendre & Legendre,
1998). There are no statistical assumptions about the
data. The most common procedure is for similar
objects to be placed in groups and these in more inclu-
sive groups, in a hierarchical manner. Results are usu-
ally presented in the form of trees or dendrograms.
The greatest development of CA in systematics was
associated with the school of numerical taxonomy (e.g.
Sneath & Sokal, 1973), and there are numerous dif-
ferent procedures available.
The two preliminary decisions in CA concern the
choice of an association coefficient and of a clustering
algorithm. Association matrices for this kind of anal-
ysis are produced either by similarity or distance coef-
ficients. The simplest similarity coefficient is the
simple matching coefficient. This can be used only for
qualitative data (binary or multistate) and CA works
better with these kinds of data (Crisp & Weston, 1993;
James & McCulloch, 1990). There are many different
clustering algorithms available. The most commonly
used algorithm in systematics is the hierarchical,
agglomerative algorithm using averages – UPGMA
(unweighted pair-group method using arithmetic
averages). Examples of the use of CA in plant system-
atics are provided by Binns, Baum & Arnesen (2002),
Crisp & Weston (1993), Gengler-Nowak (2002).
In contrast to cluster analysis, R-mode analysis
assesses the association amongst variables, and
begins with matrices produced by coefficients of
dependence amongst variables. Principal Component
Analysis (PCA) is an R-mode type of analysis.
PRINCIPAL COMPONENT ANALYSIS
PCA is usually used as an exploratory tool in system-
atics. It is a method for rotating the axes of a coordi-
nate system such that the first axis (first principal
component) passes through the greatest dimension of
the swarm of data points, and thus accounts for the
greatest amount of variance of any possible axis. The
second principal component, orthogonal to the first,
accounts for the greatest amount of residual variance,
and so on. There are as many components as original
variables, and these components are linear combina-
© 2006 The Linnean Society of London, Botanical Journal of the Linnean Society, 2006, 151, 103–111
MORPHOMETRICS IN PALM SYSTEMATICS 105
tions of the original variables. Rotation of axes is rigid,
so that the data points retain their positions relative
to one another. Most of the variance is usually sum-
marized by the first few components, and PCA thus
reduces a larger number of variables to fewer vari-
ables, which are often easier to interpret. PCA is thus
described as a dimension reducing method. Scores of
each specimen on the principal components, usually
the first two, can be plotted on bivariate scattergrams,
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allowing visualization of the relative positions of the
specimens. Some authors (e.g. James & McCulloch,
1990) consider that PCA should not be used for mul-
tiple samples – a restriction which would entirely
eliminate its usefulness in systematics. This opinion is
not shared by others (e.g. Humphries et al., 1981).
Examples of the use of PCA in plant systematics are
provided by Boyd (2002), Chandler & Crisp (1998),
Crisp & Weston (1993) and Naczi et al. (1998).
If a dataset contains both quantitative and qualita-
tive variables, and is thus unsuitable for analysis with
coefficients of dependence as in PCA, Principal Coor-
dinates Analysis (PCoA) can be used. PCoA begins
with any kind of distance matrix amongst specimens
(or a similarity matrix transformed to a distance
matrix) and is thus a Q-mode type of analysis. The dis-
tance matrix is often based on Gower’s coefficient
(Legendre & Legendre, 1998). PCoA extracts eigenval-
ues and eigenvectors from the distance matrix. Dis-
tances amongst specimens are maximized on the first
principal coordinate, and then on the second coordi-
nate, and so on. PCoA thus provides a geometrical rep-
resentation of the distances amongst specimens, and it
is the visualization of specimens in two- or three-
dimensions that is its main purpose.
DISCRIMINANT ANALYSIS
Also known as Discriminant Function Analysis or
Canonical Variate Analysis, Discriminant Analysis
(DA) comprises a group of methods rather than a sin-
gle procedure (Pimentel, 1979). The kinds of data used
are the same as for PCA, quantitative variables, but
the specimens are identified a priori to groups (i.e.
taxa). The first stage in DA is usually a multivariate
analysis of variance (MANOVA), which tests the
hypothesis that group centroids are the same. If the
MANOVA supports the alternate hypothesis, that
group centroids are significantly different, then DA
proceeds with two operations – classification and dis-
crimination. Some authors (e.g. James & McCulloch,
1990) regard the initial, inferential stage, MANOVA,
as a separate procedure, so that DA itself becomes a
noninferential method.
For classification, DA produces classification, or
identification functions that are used to determine to
which group specimens belong. Results of this may be
106 A. HENDERSON
presented in the form of a classification matrix in
which each specimen is classified according to the clas-
sification functions, either correctly according to the
original grouping, or into another group. The percent-
age of correct classifications is given and this gives an
indication of the validity of the original grouping.
These functions can also be used for identification, and
can predict group membership of an unidentified spec-
imen (assuming it belongs to one of the groups).
For discrimination, DA finds discriminant functions
(i.e. linear combinations of variables) that best
discriminate amongst the predefined groups, by
maximizing the differences amongst groups while
minimizing variation within groups. Discriminant
functions can be used to assess the relative impor-
tance of the original variables for discriminating
amongst groups.
DA is well suited to systematics. The first stage
(MANOVA) may not seem at first sight to be of much
interest, because the groups have been identified a
priori on some grounds. However, MANOVA of groups
which have been delimited by some other dataset, e.g.
qualitative data, can be tested by DA using an inde-
pendent dataset in the form of quantitative data. If
these groups represent species, then the initial
MANOVA of DA is a test of species as hypotheses (see
later). DA can be used to show which variables are the
most discriminatory, to classify specimens, and to
identify unknown specimens. Examples of the use of
DA in plant systematics are provided by Battaglia &
Patterson (2001), Binns et al. (2002), Boyd (2002) and
Murrell (1994).
TRADITIONAL MORPHOMETRICS IN PALM
SYSTEMATICS
In this section I first review the kinds of data used in
palm morphometrics, and some of the problems asso-
ciated with their collection. I then review all papers in
which traditional morphometrics has been used in
palm systematics.
SOURCES OF DATA
The data used in traditional morphometric studies of
palms are morphological or anatomical, taken from
herbarium specimens. Less often data are taken
directly from living plants. There are several problems
associated with using specimens as a data source.
First, palms are tropical plants, and there are still
numerous gaps in the collection coverage of tropical
countries. Even in small, relatively well-collected
countries such as Panama, there is uneven coverage.
These kinds of gaps and the problems they cause are
magnified many times in countries such as Brazil,
where enormous areas of the Amazon region have
© 2006 The Linnean Society of London, Botanical Journal of the Linnean Society, 2006, 151, 103–111
never been collected for any plants, let alone palms
(Henderson, 1995). The consequences of uneven collec-
tion density are that distribution gaps may be artefac-
tual, leading to erroneous conclusions.
Secondly, palms are woody plants and are often
large and/or spiny and do not fit into the general col-
lector’s main purpose – to collect as many specimens
as possible. Consequently there are relatively few
palm collections in herbaria, and those that do exist
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are often fragmentary or incomplete. Poorly collected,
pressed and mounted specimens are often useless as
data sources. For example, collectors routinely cut off
the base of the inflorescence, making it impossible to
see the important variables of peduncle length and
bract number or type of insertion. Leaves are often
folded and pressed in such a way that it is impossible
to score such variables as number, angles, lengths and
widths of leaflets. In general, the larger the size of the
palm, the more incomplete the specimen. The conse-
quences of poor specimens are missing data. These are
the bane of multivariate analysis.
Even when complete organs are present on the spec-
imen, there may still be problems in their measure-
ment. Many organs are distorted as a result of
pressing and drying. Some organs continue to change
in size as they age. For example, petioles continue to
elongate after the leaf opens (Chazdon, 1991) and
rachillae thicken as the fruits develop. Such variables
should be used with some caution. The extent of onto-
genetic allometry in palms (at least beyond the juve-
nile stage), and the extent to which this may confound
systematic studies is not yet clear. Do younger or
shorter adult palms have smaller organs than older or
taller adults of the same species? Nor have the poten-
tial size differences between sexes of dioecious species
been investigated. The various methods to remove the
effects of size in morphometric studies have been
reviewed by Humphries et al. (1981), but these have
not been attempted in palm studies. Indeed, some
would be difficult to apply because we have no mea-
sure of overall size (e.g. weight, volume or length) of
palms, nor a reasonable surrogate.
Despite these problems, one can usually recover a
reasonable amount of data for analytical purposes
using simple tools such as a protractor, digital cali-
pers and a ruler. Typically, 20–40 variables can be
measured or counted from herbarium specimens, and
there may be several hundred specimens in the sam-
ple. Some variables can be taken from specimen
labels, such as stem height or stem diameter,
although these are seldom measured accurately in
the field.
The use of leaf anatomical data circumvents the
problem of missing data, because virtually all speci-
mens have leaves. The drawback is that it requires
much more time and labour to make anatomical sec-

tions and extract the data, and such data have seldom
been used.
SYSTEMATIC STUDIES OF PALMS USING
MORPHOMETRICS
The first morphometric study of a group of palms was
that of Madulid (1981). This concerned variation in
leaf morphology in Calamus javensis Blume, and the
purpose was to find morphologically based groups
within this complex species. Madulid scored 125 spec-
imens for 35 leaf variables. Almost half of these were
quantitative but were scored as multistate qualitative
variables. Madulid used CA and PCA to analyse the
data, but found few discernible groupings.
Nauman & Sanders (1991) carried out a study of the
classification of species of Coccothrinax. Based prima-
rily on a literature survey, rather than on specimens,
they found 22 variables. All of these, including quan-
titative ones, were scored as qualitative (binary or
multistate). Nauman and Sanders used PCoA and CA
to analyse the data, and found three main clusters of
species.
Pinheiro (1997) studied generic boundaries in the
subtribe Attaleinae. He used 87 qualitative and quan-
titative, leaf anatomical variables from 75 herbarium
specimens, representing 30 species. Also included
were a number of putative hybrids. He used PCA and
CA to analyse the data. Although Pinheiro found sev-
eral reasonably well-defined groups of species, he con-
sidered that extensive hybridization made generic
resolution difficult. However, the observed variation
appears typical of closely related genera and may not
necessarily be indicative of hybridization. Pinheiro’s
study is remarkable in that he was the first to use leaf
anatomy and was able to score many variables and
produce a dataset free of missing data.
Rustiami (1999) analysed species classification in
section Piptospatha of Daemonorops. She used 58
specimens to represent 11 previously recognized spe-
cies. She scored 39 vegetative and reproductive vari-
ables, both qualitative and quantitative. She used CA
and PCA to analyse the data, and the result was a
reduction in the number of species in the section from
11 to five.
Mogea (1999) scored 51 morphological, qualitative
variables from 22 species of Arenga. He used CA to
produce a classification of species, and this was con-
sidered to represent a phylogeny of the genus. At an
arbitrarily chosen level of similarity, four groups of
species were recognized.
Bayton (2001) analysed the Calamus hollrungii
Becc. species complex. Using 40 vegetative variables
scored from 137 specimens, he tested an informal clas-
sification that recognized seven species within the
complex. Bayton used PCoA, but got inconclusive
© 2006 The Linnean Society of London, Botanical Journal of the Linnean Society, 2006, 151, 103–111
MORPHOMETRICS IN PALM SYSTEMATICS 107
results. One of the problems Bayton encountered was
that leaf sheath spines provided a suite of six qualita-
tive variables, but spines were lacking on some
specimens. Bayton discussed the problem of scoring
such inapplicable variables. However, Bayton’s use of
Strong & Lipscomb’s (1999) coding regimes is not
appropriate. Strong and Lipscomb’s study is orien-
tated to phylogenetic analysis and variable coding,
using such programs as PAUP and HENNIG86.
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Clearly, in a morphometric study these scoring con-
ventions and programs are not applicable. A better
approach to the problem of inapplicable variables is to
treat them in terms of characters and traits. Based on
Bayton’s results, leaf sheath armature seems more
likely to be a trait, rather than a character, and thus
should not be used to delimit species. I return to this
topic later.
In four of the six studies discussed above, the aim
was to classify species (Nauman & Sanders, 1991;
Mogea, 1999; Pinheiro, 1997; Rustiami, 1999), and in
the other two the purpose was to delimit taxa within a
species complex (Madulid, 1981; Bayton, 2001). How-
ever, all six studies were infused with the philosophy
of numerical taxonomy and all suffer because of that
to a greater or lesser extent. Procedures that were
commonly used in numerical taxonomy would not be
used today. For example, quantitative variables were
scored as qualitative, qualitative variables were used
in PCA, and CA was used to classify species.
Borchsenius’s (1999) paper was something of a turn-
ing point in palm systematics, because it was the first
to apply morphometric methods free from the influ-
ence of numerical systematics. Borchsenius studied
variation within the Geonoma cuneata H. Wendl. ex
Spruce species complex. He measured 12 quantitative
variables from stems and leaves from 105 individual
living plants at one site in western Ecuador. At this
site he found four different varieties of G. cuneata.
Using PCA and DA, he could clearly separate the four
varieties. However, differences between the varieties
broke down when he included plants from other sites
in western Ecuador in the analysis. Borchsenius con-
cluded that the current varietal classification was not
applicable in western Ecuador, much less throughout
the total range of G. cuneata, from western Ecuador to
Nicaragua.
Another species complex in Geonoma, the Amazo-
nian G. stricta (Poit.) Kunth, was studied by Hender-
son & Martins (2002). They measured 19 quantitative
variables from 238 herbarium specimens and analy-
sed the data with PCA, CA and DA. Although their
approach was different from that of Borchsenius
(1999), i.e. they used herbarium specimens as a data
source across the range of the species, as opposed to
Borchsenius who used living plants from a limited
part of the range, their conclusions were similar. The
108 A. HENDERSON
current varietal classification of G. stricta was unreal-
istic, and one was probably not possible based on the
data and methods employed. The main problem was
found to be in variation in leaf size and shape.
Loo et al. (2001) carried out an analysis of the vari-
ation within Licuala glabra Griff. in Peninsular
Malaysia. They studied 74 herbarium specimens and
scored 43 vegetative and reproductive variables, both
qualitative and quantitative. They used PCoA with
qualitative and quantitative variables, and PCA with
quantitative variables. Results showed three clear
groupings of specimens, and these corresponded with
certain mountain ranges in the region. Like that of
Borchsenius (1999), this paper is important in palm
systematics because of its use of appropriate method-
ology and conclusive results. Loo et al. (1999) used
molecular data to analyse three populations of
L. glabra, and used both CA and PCA.
Kahn & Gluchy (2002) measured 4393 pistillate
flowers from 135 living plants of Astrocaryum
urostachys Burret in Amazonian Ecuador. Using
univariate statistics, they found that pistillate
flower morphology varied quantitatively but not
qualitatively.
In 2002, Henderson and coworkers began a series of
morphometric studies on palm genera, mostly from
Mesoamerica. Henderson & Ferreira (2002) studied
variation between and within the two species of Syn-
echanthus. They used 23 qualitative and quantitative
variables from 355 herbarium specimens, and ana-
lysed the data with CA, PCA and DA. They found that
qualitative but not quantitative variables clearly sep-
arated the two species. Within S. warscewiczianus H.
Wendl., there were populations of very small plants
from isolated mountains in central Panama. Within
S. fibrosus (H. Wendl.) H. Wendl., there were three
separate populations, one in Mexico, a second in Bel-
ize, Guatemala, Honduras and Nicaragua, and a third
in Costa Rica. PCA showed that S. warscewiczianus
varied mostly in size, and regression showed that
some of this variation was associated with elevation.
In contrast, S. fibrosus varied more in shape and
showed no association with elevation.
Henderson (2002) analysed 22 qualitative and
quantitative variables from 476 specimens of Rein-
hardtia, using CA, PCA and DA. In marked contrast to
other studies of palms, PCA of quantitative variables
clearly separated the six species. Using regression,
Henderson found clinal variation in some variables
with elevation. One species, R. gracilis (H. Wendl.)
Burret, exhibited considerable variation and seven
different populations were distinguished.
The two studies discussed above, on Synechanthus
and Reinhardtia, lacked any systematic content in
that they only discussed variation discernible by
morphometric analysis of data from specimens. The
© 2006 The Linnean Society of London, Botanical Journal of the Linnean Society, 2006, 151, 103–111
absence of systematic conclusion can be considered a
shortcoming of these studies. In Henderson (2004) and
Henderson (2005a), multivariate analysis of specimen
data was combined with delimitation of taxa.
In Hyospathe (Henderson, 2004), 31 variables from
428 specimens were scored. CA was used to divide
qualitative variables into characters or traits. Charac-
ters only were used to delimit species, and six species
were recognized, based on groups of specimens with
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unique combinations of character states. This is an
application of the Phylogenetic Species Concept (PSC;
Nixon & Wheeler, 1990). Variation within each species
was then analysed using PCA and DA. One species
was found to be widespread and complex, and was
divided into six subspecies. Henderson hypothesized
the existence of a hybrid zone between two species
along eastern Andean slopes.
Henderson (2005a) analysed variation within
Calyptrogyne, for which 35 variables, both qualitative
and quantitative, were scored from 563 specimens.
CA, PCA and DA were used to analyse the data. As a
result of the analysis, the number of recognized spe-
cies increased from eight to 18, these with 13 subspe-
cies. Regression showed evidence of clinal variation of
some variables with longitude, and there was also evi-
dence of the existence of a hybrid zone between two
species in Costa Rica.
The four papers on Synechanthus, Reinhardtia,
Calyptrogyne and Hyospathe show the development of
my own thinking on methods in herbarium systemat-
ics. A shortcoming of the Synechanthus and Reinhard-
tia papers is that they have no systematic basis: they
are merely analyses of morphological variation. In the
Calyptrogyne study there is a systematic basis, but
characters are not distinguished from traits, and spec-
imen groups were delimited before a species concept
was applied. In Hyospathe, qualitative variables were
divided into characters and traits, and the PSC
applied.
CONCLUSIONS FROM RECENT STUDIES
A review of these recent papers using morphometric
methods on palms leads to several conclusions. These
derive from the complexity of variation uncovered by
morphometric analysis, a complexity that is not
revealed, or is glossed over, in traditional systematic
studies.
1 Hybrid zones may be common phenomena in palms.
Henderson (2004, 2005a) postulated the existence of
zones of intermediacy between two species in both
Hyospathe and Calyptrogyne. In studying Geonoma,
I have found evidence of a hybrid zone between
G. deversa (Poit.) Kunth and G. leptospadix Trail.
Hybrid zones will have important implications
 MORPHOMETRICS IN PALM SYSTEMATICS 109

for systematics, but will need to be investigated
further using ecological and genetic data and
methodologies.
2 Subspecific variation in many species of palms is
greater than previously suspected, and is seldom
adequately documented by traditional methods.
Species such as Synechanthus warscewiczianus and
Reinhardtia gracilis, as well as many species of
Geonoma, are widespread and exhibit marked
species concept (Henderson, 2005b). This should have
diversity. Recently I have put forward what I
consider to be a scientific method for herbarium
systematics, at least its descriptive component
(Henderson, 2005b). This method involves several
separate yet sequential stages, two of which involve
morphometric methods. I give a brief review of this
method here.
The first stage in a systematic study is to choose a

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morphological variation, and this often correlates
an appropriate theoretical background and an appli-
with geographical disjunction. In such situations,
cable discovery operation – and these two parts of the
subspecies can be recognized. I think it is useful to
concept are closely linked. Herbarium systematists
distinguish here between these kind of variable,
are therefore constrained in their choice of concept. By
or polytypic, species and polymorphic, or ochlo-,
taking into account theoretical background, discovery
species (Cronk, 1998). Variation in the former can
operation and data (i.e. primarily morphological data
readily be understood using morphometric methods.
taken from specimens), I consider that the PSC is the
Variation in the latter is not so easily resolved.
most appropriate for herbarium systematics. Phyloge-
Ochlospecies exhibit great variation that does not
netic species are: ‘the smallest aggregation of popula-
correlate with geography, and are not amenable to
tions . . . diagnosable by a unique combination of
systematic treatment (Cronk, 1998). There are sev-
character states in comparable individuals’ (Nixon &
eral examples from Geonoma, where ochlospecies
Wheeler, 1990).
have been termed species complexes (G. cuneata:
The next stage is to assemble a sample of specimens
Borchsenius, 1999; G. stricta: Henderson & Mar-
and to state an initial hypothesis: that a certain
tins, 2002). However, I think that ochlospecies may
number of groups are present in the sample. Data are
be relatively uncommon in palms; most species com-
gathered from the specimens and the matrix is
plexes are polytypic and may be understood using
constructed. Qualitative attributes are divided into
morphometric analysis. I have found examples of
characters and traits using either Population Aggre-
polytypic species in Geonoma, e.g. G. interrupta
gation Analysis (Davis & Nixon, 1992) if applicable, or
(Ruiz & Pav.) Mart.and G. maxima (Poit.) Kunth.
CA and the sequential removal of suspected traits
3 Morphometric studies suggest the number of spe-
from the analysis (e.g. Henderson, 2004). Characters
cies of palms may be underestimated. The number
alone are used to produce groups of specimens with
of species recognized in Hyospathe has increased
unique combinations of character states. At this stage
from two to six, these with six subspecies (Hender-
the initial hypothesis may be accepted or rejected, and
son, 2004). In Calyptrogyne the number has
in the latter case a new hypothesis proposed. This may
increased from eight to 18, these with 13 subspecies
be in the form of a null hypothesis – for example, that
(Henderson, 2005a). Similar increases are found in
group centroids are different – and such a hypothesis
Geonoma (A. Henderson, unpubl. data). I estimate,
can be tested inferentially using DA of quantitative
from this admittedly small sample, that there may
data. The PSC is applied to these groups. Quantitative
be double the currently accepted number of 2300
and geographical variation within species may then be
species of palms.
analysed and subspecies recognized. It is this stage at
4 Following directly from this, our knowledge of spe-
which morphometrics is most useful. Descriptions of
cies-level variation in general is poor. The founda-
taxa are given in the form of summary statistics of the
tion of our systematic knowledge of palms at all
sample. Ranges, means or medians, confidence inter-
levels rests on our knowledge of species, and yet we
vals or coefficients of variation, and sample size, taken
have hardly scratched the surface when it comes to
directly from the data matrix, contain informative,
understanding morphological variation in palms. I
unambiguous statements about the sample. For
consider the main impediment to understanding
qualitative data in descriptions, all that is needed
species-level variation is not so much the data
are the relevant states of the characters. Finally,
source, i.e. specimens, but rather flawed methodol-
names are applied to species according to the rules of
ogy. Although there are limitations on the knowl-
nomenclature.
edge that can be gained from herbarium specimens,
it is apparent that when we use morphometric
methods we recover far more information than we
FUTURE PROSPECTS
do with traditional methods. Analysis of this infor-
mation, combined with a scientific systematic Considering the total number of systematic papers on
method, gives a much improved estimate of species palms published over the last 30 years, the role of mor-

© 2006 The Linnean Society of London, Botanical Journal of the Linnean Society, 2006, 151, 103–111
110 A. HENDERSON
phometric analysis has been minor. I consider this is
due in part to the legacy of numerical taxonomy. As
shown above, the few palm papers based on this philo-
sophical background have not been overly successful.
Nevertheless, traditional systematics has not, in my
opinion, been entirely successful either. Palm system-
atists have, for the most part, ignored the problems of
species concepts and species delimitation, and pro-
duced monographs and revisions without any recourse
to a scientific methodology. The result has been wildly
different estimates of the numbers of species of palms
and an unstable systematics that changes with every
revision. Traditional herbarium systematics of palms
is untenable because of the lack of stated species con-
cepts and of a scientific methodology. Recent system-
atic revisions employing an explicit methodology give
a scientific estimate of species diversity.
I think that if we can move toward a more scientific
systematics of palms, as outlined above, incorporating
morphometric methods, then we will have a more sta-
ble systematics. We will also have a sound basis on
which to study other problems, such as subspecific
variation, hybrids and hybrid zones, species com-
plexes and biogeographical patterns. Furthermore, we
should complete descriptive systematic studies before
we attempt phylogenetic analysis, although these
things are often carried out the wrong way round
(Wheeler, 2004). Geometric morphometric methods
should also be applied to systematic problems in
palms. Geographic Information Systems (GIS) tech-
nology will become increasingly important in palm
systematics, and this will be especially informative
with morphometric datasets. We will begin to under-
stand biogeographical patterns and the association
between morphological variables and environmental
variables. A group of researchers at the University of
Aarhus is already active in this field (e.g. Borchsenius
& Skov, 1997; Skov & Borchsenius, 1997).
ACKNOWLEDGEMENTS
I thank the organizers of this conference for the oppor-
tunity to present this paper.
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