Cladistics

Cladistics 32 (2016) 573–576

10.1111/cla.12144

What is a cladogram and what is not?

Andrew V.Z. Brower*
Evolution and Ecology Group, Department of Biology, Middle Tennessee State University, Murfreesboro TN 37132, USA
Accepted 20 October 2015

Abstract

The origins and meanings of “cladogram” are reviewed. Traditionally, “cladogram” has been defined as a graphical represen-
tation of an empirical hypothesis of relationships among taxa, based on evidence from synapomorphies alone. Disturbingly,
numerous recent authors treat “cladogram” as synonymous with “dendrogram” and do not appreciate the particular method-
ological connotations of the former term. This is lamented.
© The Willi Hennig Society 2015.

One would hope, 50 years after the publication of
Phylogenetic Systematics (Hennig, 1966) that cladistics
would have achieved clear and unambiguous terminol-
ogy. And perhaps, within our own modest circle
of like-minded researchers, we have done so. However,
it seems that in the broader world of “phylogenetics”,
concepts that were once precisely defined have become
vague and obfuscated. As part of an ongoing
campaign for semantic clarity (cf. Brower, 2010,
2012, 2015; Brower and de Pinna, 2012, 2014), I offer
the following observations regarding the term
“cladogram”.
Hennig (1966, p. 196) said, “Our contention is gener-
ally true, that ‘family tree’ and ‘written fixation of the
system’ correspond exactly only when the family tree
clearly shows recognized or presumed sister-group rela-
tionships, and also makes clear which groups are
undoubtedly monophyletic, and which are doubtfully
so . . . . By no means all of the family trees scattered
through the literature satisfy these criteria.” Although
Hennig’s book was full of cladograms, he did not
describe them as such. Rather, it was evolutionary tax-
onomist Mayr (1965), and pheneticists Camin and
Sokal (1965) who independently proposed the term.
Camin and Sokal said, “We suggest the term cladogram
*Corresponding author:
E-mail address: abrower@mtsu.edu
to distinguish a cladistic dendrogram from a phenetic
one, which might be called a phenogram.” Mayr (1965)
did not provide a concise definition, but in the glossary
of his systematics textbook (Mayr, 1969), he defined a
cladogram as “a dendrogram based on the principles of
cladism; a strictly genealogical dendrogram in which
rates of evolutionary divergence are ignored.”1
In the 1970s, cladists (e.g. Platnick, 1977) recog-
nized that a single cladogram reflects multiple possi-
ble narratives of evolutionary change. For example,
the cladogram (A(B, C)) interpreted from an evolu-
tionary perspective implies either that A is the sister
taxon of B and C, that A is the ancestor of B and
C, or that A, B and C are three stages of a single
1
As these quotes indicate, multiple terms for different kinds of
tree diagram exist. The most general of these is dendrogram, which
refers to any tree-like diagram. A synonym in casual parlance is
“tree”. Phenograms are results of phenetic analyses; phylograms are
trees that indicate something about relative branch length (these can
be parsimony trees with branch lengths drawn proportional to the
number of apomorphies hypothesized to have undergone transfor-
mation on particular branches under some optimization scheme, or
likelihood trees, which use a model to accomplish the same end).
Chronograms are tree diagrams that illustrate inferred absolute ages
of divergence of sister taxa. There does not seem to be a term for
trees inferred by Bayesian methods, which are a posteriori
model-based truth claims about phylogenetic divergence. I propose
“credograms”.

© The Willi Hennig Society 2015

574 A.V.Z. Brower / Cladistics 32 (2016) 573–576
lineage changing through time. These observations
highlight the conceptual distinction between clado-
grams as graphical summaries of empirical evidence
of sister group relations, and phylogenetic trees as
evolutionary scenarios. Hennig (1966, p. 194) was
aware of this: “the task of the phylogenetic system is
not to present the result of evolution, but only to
present the phylogenetic relationships of species and
species groups on the basis of the temporal sequence
of origin of sister groups.”
Three important post-Hennigian books about
cladistics were published in 1980/81. Eldredge and
Cracraft (1980) did not provide a succinct definition
of cladogram, but offered the following considera-
tions: “Cladograms, as defined and discussed here,
are specific kinds of hypotheses about the history of
life. They are hypotheses about pattern” (p. 20);
“Cladograms depict nested sets of synapomorphies,
thereby defining monophyletic groups and simultane-
ously presenting a hypothesis of relationships among
the taxa” (p. 113); “. . . cladograms are hypotheses
about the structure of that history, that is, not specif-
ically about the history itself, but about the structure
of the relationships of the organisms as expressed in
their patterns of shared evolutionary novelties.”
(p. 212); and (notably), “. . . cladograms, in them-
selves, are not phylogenies, but rather hypotheses
about the pattern of nested evolutionary novelties.”
(p. 21). Wiley (1981, p. 97) defined cladogram as “a
branching diagram of entities where the branching is
based on the inferred historical connections between
the entities as evidenced by synapomorphies”, and
also distinguished cladograms from phylogenetic trees.
Nelson and Platnick (1981, p. 14) succinctly said,
“phyletic trees depict aspects of evolutionary genealo-
gies” while “cladograms depict structural elements of
knowledge”.
In more recent systematics texts, the following defi-
nitions of cladogram are found:
“. . . a pattern of character distributions in the natu-
ral hierarchy. In a phylogenetic tree, the nodes, instead
of representing the defining characters, are ancestors,
the branch points speciation events, and the lines
actual lineages of descent with modification. By con-
trast, a cladogram is a general expression of empirical
evidence; of organisms and their characters consistent
with a variety of phylogenetic trees.” (Humphries and
Parenti, 1986, p. 24).
“The cladogram specifies sister-group relationships
on the basis of homologies shared at the correct level
of inclusiveness (synapomorphies).” (Rieppel, 1988,
p. 150).
“branching diagram[s] depicting the genealogical
relationships between species or other suitable terminal
taxa.” (Minelli, 1993, p. 10). (Minelli also noted that
“The differences between one tree-like diagram and
another are not simply matters of aesthetics. Several
conceptually different tree-like diagrams are possible
and one should check carefully, whether these are phy-
logenetic trees, cladograms, or something else.”
(p. 32)).
“a dendrogram (tree diagram) specifically depicting
a phylogenetic hypothesis and therefore based on
synapomorphies. A cladogram generally only indicates
the branching pattern of the evolutionary history.”
(Quicke, 1993, p. 263).
“a branching diagram specifying hierarchical rela-
tionships among taxa based on homologies (synapo-
morphies). A cladogram includes no connotation of
ancestry and has no implied time axis.” (Kitching
et al., 1998, p. 202).
“an evolutionary tree that has no information on
branch lengths” (p. 24) and “simply shows relative
recency of common ancestry” (p. 10). (Page and
Holmes, 1998)
“a depiction of hierarchic relationships among taxa
in the form of a treelike diagram, which shows relative
recency of relationship and on which character-state
transformations may be mapped, but without the con-
notation of the amount of difference or time since
divergence.” (Schuh and Brower, 2009, p. 259).
“a parsimony tree where the weight of the edges is
not relevant” (Wiley and Lieberman, 2011, p. 104).
“a representation of nested sister-group relationships.
. . . Cladograms make no statements about character
change, ancestors, or the evolutionary process—they
are simply nested sets.” (Wheeler, 2012, p. 28).
The salient features of (almost) all of these defini-
tions are that a cladogram is a special sort of dendro-
gram, depicting an empirically supported hypothesis of
branching order that implies relative recency of com-
mon ancestry, as evident from the presence of shared,
derived character states (synapomorphies), and which
does not take into account degree of similarity or dif-
ference, branch length or absolute time. So, what’s the
problem?
Some authors do not appreciate that a cladogram is
not just any old dendrogram: W€agele (2005), for
example, explicitly conflated cladograms with dendro-
grams, phylogenetic trees and tree graphs. Parenti and
Ebach (2009, p. 252) also defined cladogram extremely
broadly: “any branching diagram, graph or written
statement that depicts the relationship between three
or more taxa.” And the UC Berkeley site “Under-
standing Evolution” (http://evolution.berkeley.edu/
evolibrary/article/0_0_0/phylogenetics_02, accessed 1
September 2015) says, “What’s the difference between
a phylogeny, an evolutionary tree, a phylogenetic tree,
and a cladogram? For general purposes, not much.
This site, along with many biologists, use these terms
interchangeably—all of them essentially mean a tree
structure that represents the evolutionary relationships
 A.V.Z. Brower / Cladistics 32 (2016) 573–576
within a group of organisms.”2 Others reject the termi-
nology outright: Ax (1987, p. 46) said, “A ‘cladogram’
is supposed to represent an abstract synapomorphy
scheme—each branching (whose nature is not further
specified) leads to unities characterized by evolutionary
novelties unique to them alone. A ‘phylogenetic tree,’
on the other hand, specifies that the branching points
of the diagram are species splits, and it thus portrays
the historic happenings which have caused the phylo-
genetic relationships between the unities displayed.
The distinction between cladogram and phylogenetic
tree may or may not be useful . . . . it is irrelevant what
notions different authors may connect with the terms
cladogram and phylogenetic tree. I shall remain consis-
tent and use the expression ‘diagram of phylogenetic
relationship’ as an unambiguous term that cannot be
misunderstood.” But, of course, it can . . .
Others are just as confused. Baum and Offner (2008)
defined cladogram as “either a general term for a tree
diagram, or a particular style of tree diagram in which
neither the amount of change nor time is depicted.”
The former definition makes the same error as W€agele;
the implication in the latter definition is that it is the
manner in which it is drawn, and not the evidence
upon which it is based, that makes a cladogram a
cladogram. Baum and Smith (2013, p. 441) defined
cladogram as “a tree diagram that communicates the
topology (the pattern of branching) but not the branch
lengths” (still neglecting synapomorphy, no doubt
because to Baum, “tree thinking” “does not necessarily
entail knowing how phylogenies are inferred by prac-
ticing systematists” (Baum et al., 2005)), and added,
“A cladogram provides all the information relevant to
determining the degree of evolutionary relatedness.”
Ernst Mayr would certainly not have agreed with that,
and I imagine that most people who advocate model-
based methods (like Baum does) would also argue that
taking account of anagenesis (=“branch length”) is a
critical component of modern phylogenetics.
Williams and Ebach (2008) offered historiographical
comments regarding the origins and evolution of
cladogram concepts that are parallel to these. They
said “speaking with one voice—or so it seemed—the
‘cladists’ stated clearly that a cladogram is a synapo-
morphy scheme. . .” (p. 5), an idea that seems to sup-
port the empirically based definitions above. In
support of this, they cited Schlee (1969), Farris (1979),
Nelson (1979), Schuh and Farris (1981) and Patterson
(1982). But then Williams and Ebach rejected that
2
This is evidently also the prevailing wisdom at Wikipedia
(https://en.wikipedia.org/wiki/Cladogram, accessed 30 August 2015),
which asserted “Algorithms for cladograms include least squares,
neighbor-joining, parsimony, maximum likelihood, and Bayesian
inference.” Efforts to correct that page were stymied by its self-
appointed gatekeepers, providing the inspiration for this manuscript.
575
consensus: “As far as we can establish, the terms
‘cladistic’ and ‘cladogram’ were first given a cladistic
treatment by Nelson (1978, 1979); Nelson and Plat-
nick, 1981): ‘. . . a cladogram . . . is merely a branching
diagram that summarizes general knowledge about the
kinds and relationships of organisms.’ (Nelson, 1978,
p. 108). ‘Cladogram’ can be understood as a general
term for any kind of branching diagram or hierarchi-
cal classification.” (Williams and Ebach, 2008, p. 5).
This definition conflates cladograms, phenograms,
phylograms and even, it would seem, fanciful tree dia-
grams with no empirical basis at all. Williams and
Ebach’s argument is lengthy and subtle, and the reader
is encouraged to read their book to gain a full appreci-
ation. Nevertheless, it is hard to see the utility of a
definition of “cladogram” that eschews synapomorphy.
Last, it is notable (but perhaps not surprising) that
three widely cited foundational texts of algorithmic or
model-based phylogenetics, Swofford et al. (1996), Nei
and Kumar (2000) and Felsenstein (2003), do not
define and barely mention cladograms at all (but then
they are also not interested in other old fashioned
notions such as “homology”, either).
In the post-cladistic Black Box Age of barcode
K2P phenograms, unrooted haplotype networks and
heuristically inaccessible model-based tree diagrams, I
think it is important that the word “cladogram”
retains its historical meaning and not be coopted as a
generic term for trees produced by other, less philo-
sophically sound approaches. Just as the French
claim that not all sparkling wine is Champagne, so
cladists need to assert that not all dendrograms are
cladograms: cladograms represent the results of
cladistic analyses.
Acknowledgements
I thank Peter Coxhead for provocation. Research in
my lab is supported by a collaborative grant, Dimen-
sions US-Biota-S~ao Paulo: Assembly and evolution of
the Amazon biota and its environment: an integrated
approach, supported by the US National Science
Foundation (NSF), National Aeronautics and Space
Administration (NASA) and the Fundacß~ao de Amparo
a Pesquisa do Estado de S~ao Paulo (FAPESP).
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