UNCORRECTED PROOF
© The Norwegian Academy of Science and Letters • Zoologica Scripta, 00, 0, January 2005, pp000–000 1
Z S C 1 9 6 Operator: Zhou rong Dispatch: 21.02.05 PE: Julian Staddon
Journal Name Manuscript No. Proofreader: Ni xiuqiang No. of Pages: 3 Copy-editor: Jonathan Lewis
Traditional (‘Linnaean’) definitions of taxon names ‘have
types and no circumscriptions’ (Moore 1998: 565), and the
contents of taxa thus defined are inherently unstable. For
instance, the Linnaean taxon Hominidae contains the type
species Homo sapiens and all other species that ‘deserve’ to be
in the same family as the type. However, different people will
have different interpretations of just how much divergence or
variability is permitted within the Linnaean rank of ‘family’
(for instance), resulting in endless and inconclusive argu-
ments over the limits of this family, even if there is complete
agreement over phylogenetic relationships (e.g. Cantino
et al. 1997; Laurin & Anderson 2004).
In contrast, phylogenetic definitions tie taxon names to
clades that are precisely circumscribed through a definition
and phylogenetic hypothesis, and these well-defined taxon
boundaries improve taxonomic stability (de Queiroz &
Gauthier 1992; Cantino & de Queiroz 2003). However, since
the boundaries of named taxa are defined with respect to an
inferred phylogeny, instability in taxon composition within this
system can occur with changes in hypothesized phylogenetic
relationships. If the composition of taxa with phylogenetic-
ally defined names is to be maximally stable, the boundaries
of named taxa should be buffered against phylogenetic
uncertainty.
The choice of reference taxa can greatly influence the sta-
bility of phylogenetic taxonomies. The three most common
types of phylogenetic definitions use one or more reference
taxa or ‘specifiers’ (analogous to Linnaean ‘types’) to delimit
named clades. For instance, ‘the least inclusive clade contain-
ing species A and species B’ (node-based), ‘the most inclusive
clade containing species A but not species B’ (stem-based),
and ‘the most inclusive clade exhibiting character state X
synapomorphic (‘homologous’) with that in species A’
(apomorphy-based).
Here, building upon previous suggestions (e.g. Schander
& Thollesson 1995; Lee 1998; Sereno 1999; Cantino & de
Queiroz 2003), I suggest how such instability can be mini-
mized through choosing phylogenetically stable reference
taxa. In particular, it is possible to quantify taxon stability
using simple (but seldom-used) indices, and then choose only
the most stable terminals to use as reference taxa in phylo-
genetic definitions. Using this approach enables one to apply
names only to stable clades in the best-supported ‘backbone’
subtree within a phylogeny. Unstable taxa can float in and out
of these named clades causing only minimal nomenclatural
disruption.
The example discussed below employs node-based defini-
tions and morphological data, but the conclusions apply to all
types of phylogenetic definitions and all types of character
data. Also, while example definitions below use one or two
reference taxa, the suggestions apply to definitions that use
more reference taxa (e.g. Cantino et al . 1997; Joyce et al . 2004).
Consider the data set in Table 1 and the resultant phylogeny
(Fig. 1). As will be clear, there is a strong signal (congruent
characters 1–10) suggesting relationships among taxa A to E
as follows: A(B(C(D(EF)))). The relationships of taxon W are
less certain: the most parsimonious tree of 18 steps places it
deeply nested within the ingroup as the sister to taxon F, on
the basis of six characters (7–12). However, there are five con-
Table 1 Example data matrix containing a highly unstable, ‘wildcard’
taxon (W). O = outgroup.
itap0 Taxon Characters
O 0 0 0 0 0 0 0 0 0 0 0 0 0
A 1 1 0 0 0 0 0 0 0 0 0 0 1
B 1 1 1 1 0 0 0 0 0 0 0 0 0
C 1 1 1 1 1 1 0 0 0 0 0 0 0
D 1 1 1 1 1 1 1 1 0 0 0 0 0
E 1 1 1 1 1 1 1 1 1 1 0 0 0
F 1 1 1 1 1 1 1 1 1 1 1 1 0
W 1 1 0 0 0 0 1 1 1 1 1 1 1
Blackwell Publishing, Ltd.
Point of View
Choosing reference taxa in phylogenetic nomenclature
MICHAEL S. Y. LEE
Lee M. S. Y. (2005). Choosing reference taxa in phylogenetic nomenclature — Zoologica
Scripta, 00, 000–000.
Michael S. Y. Lee, School of Earth and Environmental Sciences, University of Adelaide, SA 5005,
Australia; and Earth Sciences Section South Australian Museum, North Tce, Adelaide 5000,
Australia. E-mail: lee.mike@saugov.sa.gov.au
Accepted: ?????? 2000
doi:10.1111/j.1463-6409.2005.00196.x
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