The Botanical Review 69(1): 59-78

The Linnaean System and Its 250-Year Persistence

RANDALL T. SCHUH

Division o f lnvertebrate Zoology

American Museum o f Natural History
New York, N Y 10024, U.S.A.

1. Abstract . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 59
II. Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 60
III. The Linnaean System . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 60
A. Hierarchy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 60
B. Concepts . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 61
C. Stability . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 62
D. Simplicity and Continuity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 64
IV. Criticisms of the Linnaean System of Nomenclature . . . . . . . . . . . . . . . . . . . . . . . . . . 65
A. Phylogenetic Nomenclature . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 65
B. Definition of Names, Definitions of Taxa, and Essentialism? . . . . . . . . . . . . . . . . 65
1. General . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 65
2. Karl Popper and Phylogenetic Nomenclature . . . . . . . . . . . . . . . . . . . . . . . . . . . 69
C. Concepts and the Evolutionization of Taxonomy . . . . . . . . . . . . . . . . . . . . . . . . . . 70
D. Stability . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 71
1. The PhyloCode . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 71
2. Names, Ranks, and the Representation of Hierarchy . . . . . . . . . . . . . . . . . . . . . 73
E. A philosophical perspective . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 75
V. Conclusions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 76
VI. Acknowledgments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 77
VII. Literature Cited . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 77

I. Abstract
The Linnaean system o f nomenclature has been used and adapted by biologists over a
period o f almost 250 years. Under the current system o f codes, it is now applied to more than
2 million species o f organisms. Inherent in the Linnaean system is the indication o f hierarchi-
cal relationships. The Linnaean system has been justified primarily on the basis o f stability.
Stability can be assessed on at least two grounds: the absolute stability of names, irrespective
o f taxonomic concept; and the stability of names under changing concepts. Recent arguments
have invoked conformity to phylogenetic methods as the primary basis for choice of nomen-
clatural systems, but even here stability o f names as they relate to monophyletic groups is
stated as the ultimate objective. The idea of absolute stability as the primary justification for
nomenclatural methods was wrong from the start. The reasons are several. First, taxa are con-
cepts, no matter the frequency o f assertions to the contrary; as such, they are subject to change

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60 THE BOTANICAL REVIEW

at all levels and always will be, with the consequence that to some degree the names we use to
refer to them will also be subject to change. Second, even if the true nature of all taxa could be
agreed upon, the goal would require that we discover them all and correctly recognize them for
what they are. Much of biology is far from that goal at the species level and even further for
supraspecific taxa. Nomenclature serves as a tool for biology. Absolute stability of taxonomic
concepts--and nomenclature--would hinder scientific progress rather than promote it. It can
been demonstrated that the scientific goals of systematists are far from achieved. Thus, the
goal of absolute nomenclatural stability is illusory and misguided. The primary strength of the
Linnaean system is its ability to portray hierarchical relationships; stability is secondary. No
single system of nomenclature can ever possess all desirable attributes: i.e., convey informa-
tion on hierarchical relationships, provide absolute stability in the names portraying those
relationships, and provide simplicity and continuity in communicating the identities of the taxa
and their relationships. Aside from myriad practical problems involved in its implementation,
it must be concluded that "phylogenetic nomenclature" would not provide a more stable and
effective system for communicating information on biological classifications than does the
Linnaean system.

I1. Introduction
Biological nomenclature is a communication system. It provides a way for systematists and
others to talk about taxa. As such, names have meaning only when those who use them jointly
understand the concepts attached to them. Those concepts may refer to the attributes of individual
taxa, such as species, as well as connote information about relationships among taxa. The effective-
ness of a system of nomenclature, then, might best be judged on its ability to transmit such types of
information. Whereas there is a substantial body of literature investigating the ability of hierarchical
systems to transmit taxonomic information--much of it embodied in the debates among cladists,
pbeneticists, and evolutionarytaxonomists--there seems to be little in the way of what I would view
as objective analysis of the attributes of systems of nomenclature.
There is the widely pandered idea that biological nomenclature---especiallyas attributed to
Linnaeus--is a mess. Those who promote this point of view argue that if we could only make
nomenclature stable, not only would communication about taxa proceed in a more harmonious
fashion but the taxonomic enterprise itself would be disencumbered of the Linnaean nomen-
clatural albatross that has long hung around its neck. They further argue that the existing
nomenclatural approach--what I will call "Linnaean"--lacks an evolutionary foundation. Ac-
cording to this view of systematics, we have methods appropriate for discovering lineage rela-
tionships, but the methods of naming those lineages are not themselves properly evolutionary.
In the following pages I will describe what I believe to be some salient aspects that have allowed
the Linnaean system to serve biologists well for nearly 250 years. I will then outline some of the
criticisms of the Linnaean system and describe why I believe that most of those arguments are
specious and misguided. Nixon and Carpenter (2000) published what appears to have been the most
extensive rebuttal of arguments for "phylogenetic nomenclature" before the appearance of the ar-
ticles in this issue of the Botanical Review. For the most part I will not repeat subjects treated by
them but, rather, offer what I believe to be new or extended criticisms.

III. The Linnaean System

A. HIERARCHY
The system of nomenclature currently used in biology can be traced back to the works of
Linnaeus. Hierarchical relationships oftaxa are conveyed--at least in part--by the names them-
 LINNAEAN SYSTEM 61

selves. In the original works of Linnaeus himself, examination of the taxon names allowed only
for direct recognition of taxa at the level of genus and species. It was nearly 100 years before a
system of standardized endings was developed and consistently applied to names in multiple
ranks above the level of genus. The incorporation of additional formal hierarchical levels al-
lowed for the description of increasingly more complex classifications. The adoption of stan-
dardized endings for most of the ranks greatly eased communication.
What was lacking in the Linnaean system of nomenclature until early in the nineteenth
century was a consistent system for determining how a name should be applied--the fixation
of concept, what we now refer to as "typification." The method chosen in all branches of
biology was that ofnomenclatural types. With the exception of some of the most highly ranked
taxa, under this system groupings above the species-group level derive their names--and con-
sequently concepts--from genera. The types of genera are species, the latter being the only
grouping level in which specimens serve as types. The practice of recognizing nomenclatural
types was first introduced in the Stricklandian Code, prepared in the 1840s, and relies on the
concept of priority. Typification was codified in its essentially modem form in the early twen-
tieth century in what we recognize today as the International Codes of Botanical and Zoologi-
cal Nomenclature. The system of nomenclatural types exists in contrast to what Moore (1998)
and others have called "conceptual types." The conceptual system would allow the concept
applied to names to become the personal choice of individual investigators. Whereas it might
at times produce a system of names with more obvious concepts, it would undoubtedly do so at
the expense of continuity in the use of names.
Under the Linnaean system, as currently applied, information on hierarchical relationships
is conveyed directly. At the least inclusive level, species are nested within genera. Because
species names comprise a combination of generic and species epithets, the placement--and
therefore relationships---ofspecies are self-evident from the use of their names. At more inclu-
sive levels the endings of the taxon names connote relative positions in the hierarchical system,
up to a given point, depending on whether we are using the botanical or zoological code. Thus,
within the limits of the number of levels recognized in the hierarchy, the system directly con-
veys information on group membership and on group relationships.

B. CONCEPTS
Most taxon names in biology have little or no meaning in and of themselves. Although the
hierarchical system of Linnaean nomenclature can be quickly learned--and therefore the rank
order and inclusion/exclusion relationships of names comprehended--information on the mean-
ings of the names is still dependent on knowledge of biology. Thus the method for applying
names is largely disconnected from biological issues, excepting the issue of hierarchical rela-
tionship; the names themselves become meaningful only when the concepts attached to them
are understood.
Taxon concepts exist in the form of diagnoses. This was as true in the works of Linnaeus
and his contemporaries as it is today. The justification for the concepts--i.e., whether the
groupings they connote represent works of divine creation or the products of organic evolu-
t i o n - i s what has changed over time. Furthermore, the fidelity with which the concepts (diag-
noses) recognized at lower levels nest within those used at higher levels has varied depending
on the quality of the investigation and the grouping methods applied. The scientific utility of
concepts is very closely tied to the latter point. Under the cladistic paradigm, the concepts for
paraphyletic groups are flawed and must be revised. Therefore, the composition of such groups
will of necessity change, and the application of names will sometimes change as well when
only monophyletic groups are eventually recognized.
62 THE BOTANICAL REVIEW

Taxonomic concepts are theories. Like all theories in science, theories about taxa are sub-
ject to testing and modification or rejection. Testing of taxonomic theories is a desirable prac-
tice. Each of us might wish to protect our own favored theories, especially those of our own
authorship. Nonetheless, if those theories are not maximally informative and predictive, they
should be replaced. This is a scientific goal for which we should strive. The issue of what
constitutes a taxonomic theory has been greatly clarified under the application of cladistic
methods. Prior to ascendancy and refinement of the methods articulated by Willi Hennig (1966),
the concept of a taxonomic theory was vague at best, and the validity of a taxonomic theory
often rested on power of assertion and strength of reputation of its author. Those "tests by
authority" have now been replaced by the application of parsimony as a way of judging char-
acter distributions. The change in taxonomic methodology--whether viewed as incremental or
revolutionary~oes not, however, fit well with one of the major tenets of nomenclature: the
concept of stability (see Gaffney, 1979; Dominguez and Wheeler, 1997).

C. STABILITY
A major tenet of nearly all systems of nomenclature ever proposed is stability. For example,
the most recent International Code of Zoological Nomenclature (ICZN, 1999) indicates in its
preamble that one of the objects of the code is "to promote stability and universality in the
scientific names of animals." The International Code of Botanical Nomenclature (Greuter et
al., 2000) notes that the code "aims at the provision of a stable method of naming taxonomic
groups," a statement which on its face places emphasis on the stability of the method rather
than of the names themselves.
It may seem odd that the codes speak of establishing a stable nomenclature (or at least
having a stable method of applying names), whereas much of what is in them has to do with the
management of a system in which synonymy, homonymy, and recombination--all mediated
under the concept of priority--are paramount concerns. So, by their very nature, the codes are
designed not so much to create a stable nomenclature but possibly better said, "an orderly one
which minimizes changes in names as they are applied to taxonomic concepts."
The issue of whether nomenclature lacks stability because of the system of naming itself, or
whether instability is the result of the taxonomy to which names are applied, can be explored by
examining some case studies. First we might examine the issue of catalogs or indices. In botany the
oldest and best known of these would be the IndexKewensis. Many comparable compilations exist
in zoology for vertebrates and nonvertebrates alike, although none with the scope of the Index
Kewensis, and few with the history. What these types of compilations offer is an organized presen-
tation of taxon names in a hierarchical format, in the best cases providing a detailed history of their
usage and where to find information about the taxa to which they refer. They may give the impres-
sion, especially when newly prepared, that taxonomy is a neat and tidy enterprise and that if we just
had catalogs (checklists) for all groups, we would have an accurate assessment of numerical and
hierarchical aspects of biological diversity. In a recently published article, Edwards et al. (2000:
2313) commented, with regard to the preparation of Web-based lists of world scope, that "many
species have been named more than once by different scientists, and, as our Dlowledge of species
has changed over the years, species have been merged together or split into several new species.
Thus there are many (perhaps three times as many) more scientific names than there are valid
species. The scientific community has begun several international projects to sort out the nomencla-
tural confusion that has resulted."
On one hand, the statement could be interpreted to mean that if taxonomy had been better
done there would be no synonymy or change of species concepts. On the other hand, it strongly
 LINNAEAN SYSTEM 63

Table I
Changes in taxonomy of Plagiognathus Fieber, 1858 (Insecta: Hemiptera: Miridae)
Total Valid Valid Species New
valid Nearctic Palearctic Generic Species New New incertae combina-
Year species species species synonymssynonyms species genera sedis tions
1923 57 39 18 0 - 20 - 0 -
1995 101 77 24 2 - - - 0 -
2001 110 86 24 3 30 (new) 23 1 16 26

suggests that once the synonyms and varied combinations have been sorted out through the
cataloging process, we would have only to add heretofore undescribed taxa to the classifica-
tion, and an accurate map of biological diversity would be at hand--that taxonomy could be
frozen in time.
Reality, in my view, could be no farther from the truth. My own experiences at cataloging
and producing revisions, comparable experiences of my colleagues at the American Museum
of Natural History, and my examination of the literature indicate that although catalogs serve
as indispensable tools for improving the organization of our knowledge of biological diversity,
in the end they provide only a summary of the state of the art at any given point in time.
Whether the existing taxonomy is good or bad will be determined by examination of the origi-
nal literature, subsequent revisionary work, and additional specimen materials, processes greatly
facilitated by a well-prepared catalog but for which there is no substitute. Now for the ex-
amples.
1. I recently completed a revision of a Holarctic group of bugs, the genus Plagiognathus
Fieber, 1858 (Schuh, 2001). The most recent catalog for the group had been published just six
years earlier (Schuh, 1995). Table I provides information on taxonomic changes resulting
from the revision. The existing taxonomy, although appearing orderly enough in the catalog,
dealt with a group that at once was paraphyletic with numerous species placed in other genera,
had a large number of undescribed species, had many named taxa that ultimately were treated
as junior synonyms, and included many species that did not belong to the group, some of these
being moved to other genera and the remainder placed as incertae sedis in lieu of additional
revisionary work. All of these changes were the result of interpretation of taxonomic concepts.
In only one case was a name changed for strictly nomenclatural reasons, because of a case of
secondary homonymy.
2. As a second example, let us consider the taxonomy of birds at the species level. Accord-
ing to Haffer (1997) the number of bird species reached nearly 19,000 early in the twentieth
century. By the 1940s the number of species had been reduced to 86 l 6. The numbers have now
begun to rise again, with somewhere between 9000 and 10,000 species being recognized.
These changes have almost nothing to do with the vagaries of nomenclature but, rather, result
from the application of varying concepts of species. The reduction in species numbers between
1909 and 1946 came about with the rise of population biology and the application of the
methods of the "new systematics." More recent increases in numbers of species are not so
much the result of the description of new species--because, according to Haffer's figures, only
153 taxa were described as new species over a period of about 70 years. Rather, the changes in
numbers are largely the result of applying a phylogenetic species concept, wherein minimally
diagnosable entities are recognized as valid species. Again, these changes are not the result of
64 THE BOTANICAL REVIEW

the system of nomenclature but altered views of what represents the most appropriate taxo-
nomic practice.
3. Within botany, up until about 20 years ago, the prevailing view divided seed plants
into Gymnospermae and Angiospermae, and within the latter into Dicotyledoneae and Mono-
cotyledoneae. This view is no longer considered valid, with both the Gymnospermae and
the Dicotyledoneae now being recognized as paraphyletic. These and other dramatic changes
in seed-plant classification have resulted primarily from the application ofcladistic methods,
as for example in the work of Bremer (1985) and others; more recently, additional data from
DNA sequences have substantially influenced plant classification, as reflected, for ex-
ample, in the work of Chase et al. (1993). The methods of nomenclature are irrelevant to
these changes; under any definition of the term, the higher-level taxonomy of green plants--
and seed plants in particular---over the last 20 years would be interpreted as anything but
stable.
From this limited sample, we must conclude that taxonomy is by its very nature not a stable
enterprise, and consequently neither can be the nomenclature that is tied to it. One might
venture that one day all taxa will be described and their diagnoses agreed upon. This eventual-
ity is, however, a long time in the future, if it is ever to be achieved. Although a significant
proportion of the species of macroscopic organisms have been described, a substantial number
still remain undiscovered and/or undescribed. As the examples of bugs and birds given above
clearly show, however, just because one person says the species in a group are now recognized
and adequately diagnosed, another worker my have a quite different opinion. Possibly more
important, studies of taxonomy at the supraspecific level often lag far behind what exists at the
species level. A completely dichotomous hierarchical system would require n - 1 higher taxa,
where n is the number of species. Nowhere near that many higher-taxon concepts exist. In even
the best-known groups the goal of achieving such a system is decades away. Furthermore,
many of the higher-taxon concepts now applied are flawed, as clearly evidenced by the ex-
ample of seed plants given above. The classifications (taxonomy) of even those higher taxa for
which the greater majority of species have been described--e.g., birds and angiosperms--is in
most cases far from phylogenetic. This being the case, it does not take much thought to realize
that classifications for the vast majority of taxa are obviously going to require substantial
additional work before they can be labeled as phylogenetic and therefore containing maximum
scientific value, let alone having achieved a modicum of completeness in terms of descriptive
work. For these reasons it is not reasonable to think of nomenclature as stable in an absolute
sense. It is much preferable to think of the methods of applying names as being stable. This
conclusion presupposes, of course, that names cannot be perfectly stable if the taxonomy to
which they are attached is not stable. Not everybody adheres to this point of view, a subject
that 1 will return to later in the article.

D. SIMPLICITY AND CONTINUITY

The Linnaean system provides a relatively simple and effective system of communicating
hierarchical information. It is not the only possible system capable of communicating such
information, but it now has a nearly 250-year history of doing so. It is that continuity which
provides one of the strongest arguments for the persistence of the Linnaean approach and also
for maintaining it. With at least 2 million names for biological taxa now in existence, any
efforts to replace Linnaean nomenclature should be very well founded indeed. Otherwise,
massive confusion will result, and much of the existing literature might be rendered essentially
useless.
 LlNNAEAN SYSTEM 65

IV. Criticisms of the Linnaean System of Nomenclature

A. PHYLOGENETICNOMENCLATURE
The main detractors of the Linnaean system are those who advocate what I will refer to as
"phylogenetic nomenclature." Under phylogenetic nomenclature, methods of diagnosis, typi-
fication, and indication of hierarchical placement differ from those in the Linnaean system.
Although not all details of methods for "phylogenetic nomenclature" are well worked out and
the system is still little applied, the general approach can nonetheless be articulated; an exten-
sive review of the literature can be found in Bryant and Cantino (2002). A brief description
will be of help for understanding the remainder of this article.
Definitions." Its proponents argue that in phylogenetic nomenclature it is taxa that are de-
fined, not names. Accordingly, definitions might be of three types: node based, stem based, or
apomorphy based.
The choice as to which form of definition is most desirable and will produce optimal results
under the broader philosophy of phylogenetic nomenclature is not clear from the literature.
Indeed, Sereno (1999: 334) has argued that apomorphy-based definitions should be avoided in
favor &node-based or stem-based definitions because of drawbacks inherent in the first, whereas
Lee (2001: 177) has advocated the use of apomorphy-based definitions because they will be
more stable than the other two types. What is clear is that most of the literature is organized
around the concept of node-based definitions; furthermore, Kojima (2003) has persuasively
argued that even apomorphy-based definitions are node based. For these reasons, the remain-
der of this article will treat phylogenetic nomenclature as if all definitions within the system
were node based. Furthermore, if the phylogenetic nomenclaturalistswere to adopt apomorphy-
based definitions, many of the arguments for their system would seemingly evaporate. Typical
node-based definitions, then, might read as follows: "Lepidosauria is the most recent common
ancestor of Sphenodon and squamates and all of its descendants" or "Mammalia is the most
recent common ancestor of montremes and therians and all of their descendants."
Typification: As I noted earlier, in the Linnaean system the application of names is con-
trolled by the designation of nomenclatural types. In phylogenetic nomenclature taxa are cir-
cumscribed by indicating in their definition two or more taxa--specifiers--that are descended
from a node; in the case of apomorphy-based definitions one of those specifiers would be a
character. For this reason Nixon and Carpenter (2000) have referred to phylogenetic nomen-
clature as the "node-pointing system."
Hierarchical position." In an effort to provide stability of names, indications of rank are
removed from names. One &the more widely distributed proposals would have all supraspecific
names terminating in "-ina," simply to indicate that the name was coined under the system of
phylogenetic nomenclature.

B. DEFINITIONSOF NAMES, DEFINITIONSOF TAXA,AND ESSENTIALISM?

1. General
Some authors have referred to the methods of phylogenetic nomenclature as the "phyloge-
netic systematic" approach (e.g., Kron, 1997) or as "phylogenetic taxonomy" (de Queiroz and
Gauthier, 1990; Lee, 2001). I am not interested in arguing the exact meanings of terms; none-
theless, as Nixon and Carpenter (2000) pointed out, the term "phylogenetic systematics" has
long been used for the methods articulated by Willi Hennig; appropriation of the term for an
activity with a totally different purpose does little but obscure the actual nature of the argu-
66 THE BOTANICAL REVIEW

ments being made. Furthermore, some authors, such as Lee, (2001: 175) in his use of the term
"phylogenetic taxonomy," seem to confuse the recognition of groups with the naming of them.
The term "phylogenetic systematics" is best applied to the methods used to recognize mono-
phyletic groups on the basis of synapomorphic characters; it should not be applied to conven-
tions developed for assigning names to those groups. A similar comment could be made with
regard to the use of the term "taxonomy" when referring to methods that deal solely with the
process of applying names. Such usage will at best be confusing and at worst convey an im-
pression that is just plain false. Worse yet is the frequently conveyed connotation that natural
(monophyletic) groups can be recognized only in that system (of phylogenetic nomenclature)
and that the existence of such groups will be obscured, or their recognition impossible, using
any other approach.
All criticisms of Linnaean nomenclature by phylogenetic nomenclaturalists devolve from
arguments about definitions; more specifically, the definitions of taxon names. What is being
criticized is the use of the attributes of organisms to "define" taxon names. The phylogenetic
nomenclaturalists believe this process to be essentialistic--to convey an immutable essence.
This view, as espoused by de Queiroz and Gauthier (1990), de Queiroz (1994, 1997), and
others, was adopted from works such as those of Popper (1964, 1966), Hull (1965), and Ghiselin
(1984) rather than having been invented by the advocates of phylogenetic nomenclature; fur-
thermore, the opinions of these authors are based on the application of classical ideas in the
field of philosophy to a limited sampling of relevant literature, and in the case of Hull and
Popper, by individuals with no personal experience in the field of taxonomy.
The alternative to the Linnaean system, as proposed by the advocates of phylogenetic no-
menclature, is an approach in which taxa (lineages) are defined on the basis of their relation-
ships to one another, under the model indicated above for Lepidosauria and Mammalia. Such
definitions are said to achieve a level of acceptabilitybecause they are not attached to names, do not
treat taxa as classes, and therefore do not connote an "essence." Thus, under this system of defini-
tions, phylogenetic nomenclature purportedly turns taxonomy into a truly evolution-based enter-
prise. The substance of this line of reasoning may appear as little more than obfuscation, and I
for one would contend that it is, a point of view that I will now explain in some detail.
De Queiroz's writings are littered with disclaimers indicating that he is not labeling the
majority of practicing taxonomists as essentialists. For example, "Most contemporary biolo-
gists no longer hold the view that eternal, immutable essences are the underlying bases of taxa,
or even that all members of a taxon have certain invariant characters by which they can be
distinguished" (de Queiroz, 1994: 501). Yet he makes it abundantly clear that he considers
their thinking tainted with essentialism when he notes that "It]hey have nonetheless continued
to formulate taxonomic definitions as lists of organismal traits or characters" (p. 501), that
"[t]reating lists of organismal traits, i.e., descriptions of individual organisms, as the defining
formulas of taxon names implies that taxa are abstract categories (kinds), the members of which
belong to those categories for the reason that they share certain (defining, essential) characters. This
view is very much in keeping with--and probably has been inherited directly from--the metaphys-
ics of essentialism, in which individual organisms are viewed as the concrete manifestations of an
abstract, ideal organism (essence)" (p. 501), and that "elements of the Aristotelean form of defi-
nition have persisted in modern biological taxonomy in that the names of taxa continue to be
treated as if they are defined by lists of organismal traits (see the section titled 'Definition' in
numerous recent taxonomic papers)" (de Queiroz and Gauthier, 1990: 308).
Nixon and Carpenter (2000) have labeled de Queiroz's presentations as condescending, a
point of view with which I wholeheartedly agree. But then, de Queiroz is not the first person to
have labeled working taxonomists as essentialists and not the first to use the term as one of
 LINNAEAN SYSTEM 67

derogation and wrongheaded thinking. Ernst Mayr (see 1988 for a summary) argued over a
period of many years that taxonomy must be rid of essentialism, to be substituted in his case
with "population thinking." Although Mayr was attacking working taxonomists as essential-
ists, he was also complaining about what he viewed as the defects of Hennig's then recently
introduced methods and the fact that they would undoubtedly reduce the Ayes to a status lower
than "class" (in a phylogenetic classification), the rank to which birds were truly deserving in
his view.
Mayr thus seems to have proceeded de Queiroz in providing arguments that produce an
obvious paradox: in this case that essentialism is a scientific scourge and at the same time
that the appropriate ranks for given taxa are somehow preordained. The phylogenetic no-
menclaturalists have simply avoided the paradox of their argument rather than addressing it.
They are certainly correct that taxa in the Linnaean system taxa have character-based defini-
tions (diagnoses). But then, taxon recognition in the cladistic system--the one from which a
phylogenetic taxonomy is supposed to spring--is also character based. The issue here, then, is
not the difference between character-based definitions and lineage-based definitions, as the
proponents of phylogenetic nomenclature would have us believe. It is, rather, their particular
construal-~)r misconstrual--of the term "definition" as widely used in the taxonomic litera-
ture.
De Queiroz (1997: 127), in a relatively recent form of his ongoing manifesto promoting
phylogenetic nomenclature, has argued that there are two primary kinds of taxonomic and
nomenclatural systems: Linnaean and phylogenetic. He observed that taxonomic systems can
be both Linnaean and phylogenetic, whereas nomenclatural systems "must be one or the other."
The statement suggests that the Linnaean system of nomenclature cannot be used in a phyloge-
netic context, an idea that will strike most practicing taxonomists as absurd.
De Queiroz offers no simple or direct statement about why the Linnaean hierarchy cannot
be interpreted in a phylogenetic context. His reasons, however, appear to be based almost
totally on his concept of the definition of a taxon name. He stated that, although the codes of
nomenclature are elaborate, "their method of definition is implicit rather than explicit" (1997:
133). De Queiroz ventured that the nature of the definition (of the term "definition") must be
inferred from the codes. His inference is that the definition of the name Asteraceae is some-
thing like "the taxon including the genus Aster that is assigned to the category Family." He
further states that the Zoological Code refers to definitions of names, whereas the Botanical
Code more appropriately speaks of "descriptions." According to de Queiroz, the latter term
(description) refers to taxa rather than names (p. 133) and, in the Linnaean system, definitions
are dependent on the Linnaean hierarchy, so categorical assignments play a critical role in the
application of taxon names (p. 134). On the other hand, according to de Queiroz, in a "phylo-
genetic" system of nomenclature, definitions are independent of the Linnaean hierarchy, so
categorical assignments play no role in the application of taxon names (p. 135).
Stuessy (2000) recently stepped forward to counter de Queiroz's reasoning concerning defi-
nitions. He argued that taxon names are not defined--that lists of diagnostic attributes do not
form a definition of a name--but rather that the names serve as labels for the taxa and that the
lists of attributes are circumscriptions of the taxa. In line with the view of taxa as concepts, as
expressed above, I believe that Stuessy has an accurate perception of what is actually taking
place in taxonomic practice. Nonetheless, Stuessy's comments were summarily dismissed by
de Queiroz (2000) as having been presented without any evidence to support his conclusions.
To bolster this assertion, de Queiroz simply gave a citation of his own work and requoted his
definition of Asteraceae as given above. This can hardly be viewed as anything but authori-
tarianism, because de Queiroz has no argument himself, as can be seen in this quotation: "Per-
68 THE BOTANICAL REVIEW

haps the reason Stuessy thinks that taxon names are not defined is that definitions under the
traditional system are taken for granted. That is to say, the bacteriological, botanical, and
zoological codes do not explain how taxon names are defined. . . . Nevertheless, taxon names
have implicit definitions of the kind described above [for the Asteraceae]" (de Queiroz, 2000:
533-534). One wonders how it is that de Queiroz knows the true and correct method of defin-
ing taxon names, if the works that codify the use of those names themselves make no explicit
mention of the nature of such definitions.
I agree with Stuessy that taxa are concepts and that character-based diagnoses are the defi-
nitions for the taxa to which those names apply. De Queiroz's perspective on essentialism
aside, such a view makes the direct--and I would claim necessary--connection between the
taxa, the means by which they are recognized, and the names by which we refer to them. Under
de Queiroz's model, one wonders how we are to make the connection between taxa and names.
The definitions of names, of necessity, make no mention of attributes. The taxa must remain
nameless, otherwise their names would be character based, and we would have returned to
what de Queiroz believes are entities with essences.
I have regarded (Schuh, 2000), and still regard, nearly all of the arguments put forth by de
Queiroz and those who adhere to his line of reasoning as specious, for the following reasons.
First, taxonomists have long used the terms "definition," "description," and "diagnosis" as
being virtually synonymous. Certainly the term "definition" is not restricted to the meanings of
names in the modem literature--as suggested by Hull and endlessly reasserted by de Queiroz--
but is broadly applied in the sense of "taxon concepts"; such usage is widespread in both
botany and zoology and occurs in the phylogenetic, taxonomic, and nomenclatural literature.
As such, "definition," "diagnosis," and "description" all have the intended meaning of listings
or summaries of the attributes of taxa, the first two terms usually referring to a more restricted
list of attributes than the last.
De Queiroz falsely assumes that definition is (always) used in the sense that he suggests,
but he cites no explicit examples from the taxonomic or nomenclatural literature that would
allow him to arrive at such a conclusion. Indeed, he says that the meaning of definition, as
applied to taxa, must be inferred. One is forced to conclude that his inference is simply wrong.
Names are used as connotations of those diagnostic attributes, which in sum are frequently
referred to as definitions oftaxa. Such an inference (fact?) would seem obvious, for de Queiroz
himself has noted that the Botanical and Zoological Codes use "definition" and "diagnosis" in
an interchangeable way, as do many works in the taxonomic literature; but he prefers to view
this as an apparent lapsus, or the result of confusion, concerning what the "correct" term should
actually be. The real issues, despite many pages of explication to the contrary on the part of de
Queiroz, are his slavish adherence to a meaning of the term "definition," which few biologists
but de Queiroz himself accept, and his belief that "defining" taxa by a list of attributes con-
notes for those taxa some essential and immutable properties.
The definition of Asteraceae (and similarly for any other taxon)---contraryto the view of de
Queiroz--would certainly be treated by the bulk of practicing taxonomists as "the set of at-
tributes shared by Aster and other plant taxa grouped together and placed at that categorical
level." Although the name Asteraceae may indicate hierarchical level, the level of placement
has no fixed reality but only indicates the relative hierarchical relationship of Asteraceae to
other plant taxa at more and less inclusive levels. Yhe continued insistence by de Queiroz, and
others infatuated with the idea of phylogenetic nomenclature, that placement of a taxon at
some level in the Linnaean hierarchy confers some "essential" quality to that taxon is simply
misguided. Asteraceae can maintain its status as monophyletic, while retaining its position in
the hierarchy of relationships at a level other than family. Nothing about the assignment to
 LINNAEAN SYSTEM 69

family rank is fixed. Indeed, by changing only the ending of the word, the relative hierarchi-
cal placement of the clade within the phylogenetic system for flowering plants could be
changed without altering the diagnosis for the taxon itself, or its composition. The assign-
ment of the "-aceae" ending has everything to do with communication of information about
relative rank and nothing to do with Aristotelian definitions or ascription of essential properties.

2. Karl Popper and Phylogenetic Nomenclature

The works of Karl Popper were cited by de Queiroz (1994) as a source of inspiration for his
writings. Popper's works have also been accorded approval by many authors, including myself
(Schuh, 2000), concerning the conception and testing of taxonomic theories in the cladistic
paradigm. The reader may wonder how Popper's writings can figure as a positive influence in
the writings of de Queiroz and me, when ! have little good to say about the works of de Queiroz
on the subject of phylogenetic nomenclature. 1 will explain.
In two of his works, Popper (1964, 1966) discussed at length what he believed to be the
scientific status of the social sciences. These writings necessarily diverge to some degree from
others of his that deal almost strictly with philosophy (or, if you will, epistemology) as it might
be applied in the natural sciences, as for example The Logic ~?fScientific Discovery (1968).
Nonetheless, de Queiroz (1994) has found in the aforementioned works statements that he
believes support his cause; namely, Popper's arguments concerning "methodological nominal-
ism" and "methodological essentialism." By way of paraphrase, methodological nominalism
has everything to do with identifying concepts or ideas, describing them, and if need be, at-
taching terminology to them. Methodological essentialism is much more strongly oriented
toward producing unbending definitions for terms representing favored concepts, irrespective
of their scientific interest; the approach is strongly associated by Popper with the Aristotelian
view of scientific thought, a view Popper treats most harshly.
It should not be thought unusual to find such distinctions in the works of Karl Popper. His
disdain for fomaal definitions is well known, as is his advocacy (indeed, maybe even penchant)
for conveying ideas through context rather than through formal definition. Indeed, The Poverty
of Historicism (1964) includes no explicit definition of the term "historicism," and the reader is
near the end of the work before the historicists are actually identified and labeled as such.
De Queiroz (1994) would have us believe that (most) practicing taxonomists are essential-
ists because he equates the methods of taxonomists with what Popper labels as "methodologi-
cal essentialism." Even though de Queiroz allows that the distinction between nominalism and
essentialism is a relative one, he nonetheless pursues the line of thought that the nominalistic
perspective--his--represents an important step in the development of a more broadly scien-
tific approach to taxonomy in the form of"phylogenetic nomenclature."
Under de Queiroz's construal of methodological essentialism, taxonomists are, for example,
"associated with a seemingly endless and strictly verbal debate about the meaning of the name
Mammalia" (de Queiroz, 1994: 499); one would suppose that just about any taxon name could
be substituted for Mammalia if essentialistic beliefs are as widely held among taxonomists as
de Queiroz would have us believe. In de Queiroz's view, it would appear that taxonomists dote
on the choice of the names and then spend all of their time devising concrete (essentialistic,
character-based) definitions for them, rather than concerning themselves with more productive
aspects of scientific inquiry, such as discovering and describing new species, recognizing mono-
phyletic groups, and testing existing taxonomic theories. Practicing taxonomists would cer-
tainly question whether de Queiroz has ever taken a serious and broad-based look at the
descriptive and revisionary literature.
70 THE BOTANICAL REVIEW

There is no mistaking that Popper, in his writings, showed little tolerance for the Aristotelean
approach, by which he meant positing what is true about the world (the essences), providing
names for those truths, and producing definitions for those names. Popper does not speak of
whether taxonomists---or biologists more broadly--are essentialists. But in the mind of de
Queiroz, the masses of practicing systematists are guilty by association, because they are known
to "define" taxon names, and those definitions consist of lists of traits. De Queiroz agrees with
Popper that methodological essentialism stifles scientific progress and concludes that practic-
ing taxonomists must therefore be stifling progress in the science of systematic biology be-
cause of his prior----erroneous--assessment of their methods as essentialistic.
The arguments of de Queiroz might be found worthy if his major premise had a leg to stand
on. But it does not. Practicing taxonomists know full well what they are doing, and it has
nothing to do with essentialism, in the sense used by de Queiroz, or any of those from whom he
has adopted his ideas. Rather, taxonomic definitions (or diagnoses, if you prefer) are hypoth-
eses about entities in nature. Those hypotheses are subject to test and rejection or revision. It is
in this area that the philosophy of taxonomic practice has been substantially clarified since the
last round of essentialist attacks by Mayr, a development that de Queiroz and his like-minded
colleagues are so happy to overlook in their quest to provide what they believe will be a central
role for evolutionary theory in taxonomic and nomenclatural practice.

C. CONCEPTSAND THE EVOLUTIONIZATIONOF TAXONOMY

Many authors have asserted that the publication by Charles Darwin of On the Origin of
Species and the subsequent rise of the evolutionary paradigm revolutionized the methods of
taxonomy. In the view of de Queiroz and his fellow phylogenetic nomenclaturalists, this ap-
parently only leaves nomenclature to be revolutionized. In my view--and here I am not alone
or expressing an original interpretation--nothing could be farther from the truth. Darwin may
have offered a way of viewing classifications as reflective of genealogy and argued explicitly
that such should be the case, but he certainly did not propose a method for constructing classi-
fications that contain only monophyletic groups (e.g., Padian, 1999; Schuh, 2000). This is not
to say that many taxa recognized between the time of Linnaeus and the advent of cladistics
were not monophyletic--we need only mention groups such as Angiospermae and Aves. None-
theless, explicit methods for recognizing such groups--those defined on the basis of synapo-
morphies--became clearly formulated and widely accepted only upon publication of the works
of Willi Hennig (e.g., 1966) in English in the mid-1960s. No amount of argumentation over
essentialism, definitions, or other subjects cloaked in philosophical jargon can change that
fact.
De Queiroz (1997, and elsewhere) is fond of the phrase "evolutionization of taxonomy."
The phrase includes, of course, the wishful adoption of his much-promoted system ofphyloge-
netic nomenclature. What is rarely mentioned in his writings is the fact that the evolutionization
of taxonomy (the acceptance of Darwinian theory by most taxonomists) had little effect on
taxonomic practice (see Patterson, 1987; Schuh, 2000). The justification for much of taxo-
nomic practice has no doubt changed markedly since the time of Linnaeus and his contempo-
raries and particularly since the publication of On the Origin of Species in 1859. However, the
recovery of hierarchical structure in nature has been the driving force within taxonomy from
Linnaeus to the present. Although Hennig may not have been the first---or the only--person to
appreciate the idea that grouping by "synapomorphy" was the only way to construct maxi-
mally efficient and predictive diagnoses of taxa in a hierarchical classification, his writings
were certainly the ones that catalyzed the idea among biologists worldwide. The consistent and
 LINNAEAN SYSTEM 71

widespread application of those methods has revolutionized certain aspects of taxonomic prac-
tice in a way not seen since the time of Linnaeus. Nixon and Carpenter (2000: 300-301) have
a slightly different take on this argument, but their comments--like mine--are not favorable
with respect to the views of de Queiroz, as can be deduced from the fact that their discussion of
this subject is placed under the heading "revisionist history."
De Queiroz, like some others, seems to have lost sight of (or possibly never read) the
argument that the concept of evolutionary relationships among organisms gains strength from
an independent taxonomy (e.g., Brady, 1985). Nor do he and his co-authors give much space
to discussion of the source of information for phylogenetic relationships. If the practice of
taxonomy has been strengthened in the past 35 years, it is not because of "evolutionization"
but, rather, from "objectification." That objectification has to do largely with the way taxa are
diagnosed; i.e., by apomorphies.
De Queiroz and his followers (e.g., Bryant and Cantino, 2002) would have us believe that
their approach to producing taxon definitions--"node-pointing system" (Nixon and Carpen-
ter, 2000)--will evolutionize nomenclature, because names in this system are defined on the
basis of clade (lineage) relationships rather than by lists of diagnostic characters. One has to
wonder about the relevance of the continued discussion of definitions, when in reality the
clades that de Queiroz would wish to recognize receive their character-based diagnoses from
cladistic analyses.

D. STABILITY

1. The PhyloCode
Stability under the codes of nomenclature has to a certain extent been engineered. As I
pointed out above, the idea of stability will of necessity be transitory. We could, of course,
devise a system of biological nomenclature that would be perfectly stable, but in the end it
would be almost certainly be useless scientifically. The authors of the PhyloCode (Cantino and
de Queiroz, 2000) argue that reassignment of a taxon to a higher or lower rank can cause an
undesirable cascade of name changes, a situation that would be obviated within a system of
phylogenetic nomenclature. Phylogenetic nomenclature would, of course, also remove much
of the ease with which we are able to communicate concerning relationships among taxa.
When dealing with typified taxa, shifts in categorical ranks in the Linnaean system will be no
greater, and often much less, than would be the case under the node-pointing system, as shown
by Nixon and Carpenter (2000). The proponents of phylogenetic nomenclature further argue
that:
When the PhyloCode is extended to species, it will improve nomenclatural
s t a b i l i t y . . , by removing the linkage [of a species name] to a genus name. A
major source of instability in species names under the preexisting c o d e s [ , ] . . .
revision of generic limits, will thereby be eliminated. There will, of course, be a
consequent absence of hierarchical information in species names governed by
the PhyloCode; specifically, one will not be able to infer phylogenetic relation-
ship from these names in the way that one can infer genus assignment from spe-
cies names governed by the preexisting codes. However, an underlying principle
of the PhyloCode is that the primary purpose of a taxon name is to provide a
means of referring unambiguously to a taxon, not to indicate its relationships.
From this perspective, the loss of nomenclatural stability of species names under
the preexisting codes is too high a price to pay for incorporating taxonomic infor-
72 THE BOTANICAL REVIEW

mation (genus assignment) into the names. Moreover, although such information
will not be built into species names under the PhyloCode, phylogenetic position
can easily be indicated by associating the species name with the names of one or
more clades to which it belongs. (Cantino & de Queiroz, 2000)
This statement offers a truly tortured piece of logic. Phylogenetic nomenclature would
appear by definition to be designed to convey information on phylogenetic relationships. The
name suggests as much, as does virtually all of the published discussion promoting such a
system. Furthermore, in its role as part of the "evolutionization of taxonomy," phylogenetic
nomenclature could hardly have a more important objective. Yet we are told that absolute
stability of names is more important than being able to directly communicate relationships
among species through the system of nomenclature. In the end, if producing stability is sub-
lime in the PhyloCode, indicating phylogenetic affinity is ridiculous. According to the above-
quoted statement, after we have discarded the association between generic and specific epithets
in an attempt to promote stability in the application of scientific names, we can easily commu-
nicate this information by reassociating those names, presumably in the form of a uninomial.
Thus the Phylocode proposes the reinvention an inferior version of the system that it just
discarded. Furthermore, it presupposes a fixity for species concepts and lineage identities that
has little to do with reality, as the examples described early in this article were designed to
illustrate.
To date, few published works have addressed the issue of how to deal with species under
the system of phylogenetic nomenclature. In one such work (Cantino et al., 1999) the authors
offered 13 proposals on how species names might be dealt with. In addition to the argument
that stability of species names is paramount, the paper is based on what I will call the"preemp-
tive assertion" that the system of naming species must be changed because "Linnaean binomial
nomenclature is logically incompatible with the phylogenetic nomenclature of de Queiroz and
Gauthier." No doubt all working taxonomists have found aspects of species-group nomencla-
ture under the codes tedious at times. I count myself as one of those. However, I am not
encouraged by the arguments of Cantino et al., and those presented in the PhyloCode, for the
following reasons. [Lee (2001) has also taken the "preemptive" approach to argumentation,
suggesting that the work of Nixon and Carpenter (2000) actually advocated the methods of
phylogenetic nomenclature (or phylogenetic taxonomy as he prefers to call it). Lee's argument
is based on the following erroneous syllogism, whose major premise is clearly false: Any
taxon definition involving the mention of apomorphies is apomorphy-based in the sense of
phylogenetic nomenclature; the taxon definition advocated by Nixon and Carpenter (2000)
involves the use of apomorphies; therefore, Nixon and Carpenter advocate the methods of
phylogenetic nomenclature. In actuality, nothing could be farther from the truth concerning
the conclusions of Nixon and Carpenter.]
First, assertions concerning the "logical incompatibility" of Linnaean nomenclature with
phylogenetic nomenclature are irrelevant; this is the "preemptive assertion." What is relevant
is whether the Linnaean system can accurately associate names with taxa and whether it can
represent hierarchical relationships of monophyletic groups. The answer is that it unequivo-
cally can.
Second, absolute stability is not necessarily a desirable goal. Species, like taxa at higher
levels, are concepts. To imply that fixity of species names will somehow promote fixity of
taxonomy at the species level (or any other level, for that matter) is a fatuous idea. It will more
likely simply impede the discovery of concepts in need of test and revision (Gaffney, 1979;
Dominguez and Wheeler, 1997).
 LINNAEAN SYSTEM 73

Third, adding "some" stability at the expense of losing a relatively simple method of com-
municating information on relationships is a price not worth paying. Most of the proposals by
Cantino et al. (1999) would adopt existing binomials for the names of species in the system of
phylogenetic nomenclature. As such, nothing novel is offered by the move to phylogenetic
nomenclature, except the claim by Cantino et al. that the names will not have ostensible defini-
tions and will therefore have lost their "logical incompatibility" with phylogenetic nomencla-
ture. Those names would remain the same in perpetuity. Thus, any revisions in concepts at the
supraspecific level would complicate our ability to communicate directly information about
relationships of species.
Fourth, with at least 2 million available species names already in existence, one wonders
just how phylogenetic nomenclature will ultimately communicate about the relationships of
species and distinguish among the taxa vulgarisl-vulgarislO00. Homonymy, synonymy, and
recombination may be messy issues for the taxonomist. Yet, because the classification of large
groups is seldom the work of a single individual, and because different taxonomists with equally
credible credentials and methods may reach different conclusions, taxonomy will never be-
come an absolutely stable enterprise. The idea that species names can let alone should--
remain absolutely stable is simply misguided and unscientific.

2. Names, Ranks, and the Representation of Hierarchy

Nixon and Carpenter (2000) showed that the node-pointing system is no more stable than
the Linnaean system. They based their observations on some of the examples favored for the
promotion ofphylogenetic nomenclature. I will offer a separate set of observations, also using
examples from the work of de Queiroz (1997).
De Queiroz provided an example to justify his assertion that, under the Linnaean system,
names can change under different ranking schemes and that the same names can refer to differ-
ent taxa. His figure demonstrating these effects is here reproduced as Figure 1. Types are
represented by asterisks. The differences between Scheme One and Scheme Two indeed show
what de Queiroz asserts concerning the Linnaean system: that, in fact, the (endings of the)
names are different because the ranks are different, What he views as irrelevant is that the
hierarchical relationships among the taxa are identical in both diagrams. Under a system in
which the "definition" of a taxon name is its diagnosis, exactly the same information on inclu-
sion/exclusion relationships is transmitted by both schemes. De Queiroz would have us be-
lieve that the different schemes cause confusion about the information being transmitted, but
this could only be the case if one were to accept his concept of definition for the taxa in
question.
Figure 2 offers de Queiroz's justification of phylogenetic nomenclature from the point of
view that taxon names are unaffected by rank (categorical assignment). Certainly the example
offers evidence for his point. But information on inclusion/exclusion relationships is lost!
Stability of names is the only criterion of interest to de Queiroz, but it can hardly be the only
relevant criterion in evaluating a system of biological nomenclature. Thus this example cannot
be viewed as anything but weak.
De Queiroz offers additional examples, in which the theories ofphylogenetic relationships
are changed and, therefore, the concepts applied to at least some of the included taxa are
changed. He argues that because concept is tied to rank in the Linnaean system, change in rank
produces change in meaning of taxon name. De Queiroz would have us believe in this case that
phylogenetic nomenclature will be less confusing than the Linnaean approach because the
names remain stable even though the taxon concepts have changed. This can hardly be the
74 THE BOTANICAL REVIEW

Fig. 1. Two taxonomicschemes showingchanges in nameswith changes in rank under the Linnaean
system (see the text for further explanation).Source: de Queiroz, 1997.

Fig. 2. Two taxonomic schemes showing unchangingnames with changes in rank under phyloge-
netic nomenclature(see the text for further explanation).Source: de Queiroz, 1997.

case. Nixon and Carpenter (2000) and Carpenter (2003) have shown that the Linnaean system
will be more stable for typified taxa when taxon concepts change.
All of these arguments are essentially irrelevant for the evaluation of stability. The only
relevant example offered by de Queiroz (1997) does not make a case for phylogenetic nomen-
clature on the basis of stability. Neither does it show well the difference between the two
systems with regard to inclusion/exclusionrelationships: namely, under the de Queiroz model
no information on inclusion/exclusion relationships will be found in the names themselves,
whereas the Linnaean system serves this function inherently, and well. On the basis of de
Queiroz's own examples, I can only conclude that phylogenetic nomenclature will certainly be
no more stable than the Linnaean system and that its methods for depicting even the most basic
of inclusion/exclusionrelationships will be substantially more cumbersome than those of the
Linnaean system.
 L1NNAEAN SYSTEM 75

E. A PHILOSOPHICALPERSPECTIVE
Elsewhere in the present issue Keller et al. (2003) argue for the grounding of phylogenetic
nomenclature in the philosophical view of species as individuals. Although there are aspects of
their presentation with which I do not agree, we are certainly in accord on the view that phylo-
genetic nomenclature is an ill-founded and illogical replacement for the Linnaean system.
There are also additional published arguments for why phylogenetic nomenclature should
be preferred from a philosophical point of view. One of these, by Ereshefsky (2001), was
published in book tbrm. This might, in and of itself, be though to confer some special weight.
The title, The Poverty of the Linnaean Hierarchy, would seemingly add credence to such
thinking. Between the covers of Ereshefsky's work one finds many ideas that mainstream
biologists might find perplexing, if not downright disturbing. For the purposes of the present
discussion, I will restrict my comments primarily to points dealing with the issue of biological
nomenclature.
Ereshefsky is what I would call relatively cagey in most of his presentation, weaving ideas
together in such a way that it is difficult at times to determine his position. Our first clues
undoubtedly come from his presumptuous title. Like those promoting phylogenetic nomencla-
ture, and like his mentors David Hull and Eliot Sober, Ereshefsky argues that nomenclature
must fit within an appropriate theoretical context. Not surprisingly, this context is "evolution-
ism." Ereshefsky's view, stated in his own words, sounds remarkably like what I quoted above
from de Queiroz.
A defender of the Linnaean system would, of course agree that the vast major-
ity of Linnaeus's theoretical [essentialist and creationist] assumptions are obso-
lete. Nevertheless, she could argue that we can continue to use various aspects of
the Linnaean system without adopting his outdated assumptions . . . . Still, the
Linnaean system does carry unwanted theoretical baggage. There may be a con-
sensus among biologists that taxa lack essences but the assumption that the Lin-
naean categories have essences is widely held. Taxa in a single category are often
considered comparable, and taxa in different categories are considered impor-
tantly different. (Ereshefsky, 2001: 283)
Ereshefsky does not indicate who the offending parties might be or just what they have said
and done that allows him to discern their essentialist leanings. As I argued above, such claims
are largely empty and represent little more than futile efforts to justify a position based on
philosophical preconception.
Ereshefsky further argues for what he describes as the practical advantages ofphylogenetic
nomenclature, as opposed to what he views as the practical limitations of the Linnaean system.
Among these is stability. In this regard Ereshefsky notes that because "positions and ranks of
taxa must be incorporated into their names" we are forced to accept "unwanted instability
during revision because the names of the taxa must be altered." According to Ereshefsky, the
post-Linnaean system avoids such instability because "the ranks and positions of taxa are not
indicated by their names, so taxon names remain constant during revision" (2001: 285).
Ereshefsky seems to admit to the necessity of studies in which taxa are reviewed and revised
(taxonomically). He seems to overlook the fact that such studies will frequently force us to
accept revised concepts. How nomenclature can remain entirely stable in the face of revised
concepts has not been satisfactorily explained by de Queiroz, and it is certainly not explained
by Ereshefsky. The latter's assertion that stability of nomenclature is a desirable practical
objective seems to rest on the assumption that "[t]wo biologists can disagree on the placement
of a taxon yet refer to the taxon with the same name" (2001: 285). The absolute stability of
76 THE BOTANICAL REVIEW

names seems to be the only achievement realized under this kind of thinking, an empty objec-
tive from the scientific point of view, as Gaffney (1979) argued more than 20 years ago,
Ereshefsky further states that Linnaean ranks do not reflect true divisions in nature and that
"biologists should be freed from the theoretically empty task of having to assign higher Lin-
naean ranks to taxa." With regard to species he argues that taxa typically thought of as species
are incomparable entities. As he puts it, "It is doubtful that a feature exists that distinguishes
species taxa from all other types of taxa" (2001: 285). After all of the disparaging remarks
about essences, essentialists, and essentialism by Ereshefsky and others, these remarks come
as close to describing an essentialist perspective as any one might find, no matter what the
theoretical or philosophical persuasion of the author.
Ereshefsky's arguments, in my view, are carefully tuned to conform to his preconceived
views on how evolutionary biology, and systematic biology more particularly, should func-
tion. Like many others, he seems happy enough to attempt to construct a new orthodoxy. His
viewpoint rejects the continuity of observations from at least the time of Linnaeus to the present,
that there does seem to be a hierarchical structure in nature and that it is the effective descrip-
tion of that hierarchy that has preoccupied systematic biologists. I can only conclude that
Ereshefsky's claims concerning the "poverty" of the Linnaean hierarchy are empty, in large
part because he misunderstands the very system he is trying to reform.

V. Conclusions
De Queiroz (1997) argued that the current codes of nomenclature promote "nomenclatural
clarity, universality, and stability" (p. 137) but that they do so in an inappropriate theoretical
context. That inappropriate context rests on de Queiroz's ideas concerning the definition of
names and his belief that the only thing that is "clear, universal, and stable [in the Linnaean
system] is the association between a name and one of the Linnaean taxonomic categories"
(p. 137). He argued that the clear, universal, and stable association within the system ofphylo-
genetic nomenclature will be the "association between a taxon name and a clade or monophyl-
etic group of species" (p. 138). As 1 discussed above under "Definitions," I see little evidence
in support of de Queiroz's point of view. Indeed, the characterization of the phylogenetic
nomenclature by Nixon and Carpenter (2000) as one of "metaphysical correctness" would
seem to be entirely accurate.
The context in which biological classifications must function is a hierarchical one. In this
sense the Linnaean system has served biology effectively for nearly 250 years. Although
some workers may have misconstrued the use of the "family" category to mean that place-
ment of a taxon at that level conveys some particular (essential) properties, most investiga-
tors using the system in this day and age are well aware that"family" and all other categorical
ranks are used to denote a scheme of nested hierarchical relationships, not to convey infor-
mation on some additional aspect of reality over and above the characters by which the taxa
are diagnosed.
In claiming that the "theoretical context" of hierarchical Linnaean nomenclature is flawed,
de Queiroz, Ereshefsky, and others have produced arguments that are little more than distrac-
tions. The important issue is the portrayal of hierarchical relationships of taxa. The Linnaean
system has done that for nearly 250 years and should not be abandoned simply to bring nomen-
clature into conformity with flawed metaphysics. If changes are to be made, they should be to
repair deficiencies in the existing codes of nomenclature, along the lines suggested by Nixon
(2003).
 LINNAEAN SYSTEM 77

VI. A c k n o w l e d g m e n t s

I thank Jerry Davis for inviting me to contribute to this discussion and for his belief in the
unity o f the taxonomic enterprise, Dennis Stevenson for his efforts in helping me locate the
botanical literature and for discussing botanical nomenclature, and Andrew Brower for a one-
line comment that inspired the title o f this article and a portion o f its content. Pamela Beresford,
James Carpenter, and Jun-ichi Kojima offered helpful comments on the manuscript. The ar-
ticle is dedicated to the late Ron Brady, in recognition o f his arguments concerning the inde-
pendence of systematics as a field of enquiry.

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