Cladistic Taxonomy

Prévia do material em texto

1995 POLNTS OF VIEWSSTEMATIC BIOLOG 
schoois ot taxonomy (Fig. 1) are as foliows. 
Aristotelian taxonomy proceeds by logical subdivision, in which each member of a Cladistic Taxonomy, Phylogenetic Systematics, and 
Evolutionary Ranking 
pair of taxa is characterized, respectively, by the presence and absence of a chosen 
N Dt feature. All present attempts to derive a 
taxonom from an identitication key reflect 
this process. In Linnaean taxonomy, there 
is an explicit attempt to distinguish the im- 
portant (stable, essential) properties from 
the unimportant (variable, nonessential) 
properties. In traditional taxonomy, groups 
are reconstructed based on perceived sim-
ilarities and ditterences among taxa. Qui- 
narian taxonomists (eg., MacLeay, 1819) 
looked for idealistic groupings in circles of
five, supposed to reveal the harmonious 
MARTIN L. CHRISTOFFERSEN 
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5S050-900. joào Pessoa. Paraiba Bras1l
A 
The anaent discipline of biological tax 
onomr has been veT slow to incorporate phylogenetic representations (Gritfiths,
maior shifts in worid views throughout the 
millennia. imperiious to the derision of 
saentists trom the more giamorous fields
0 research. marnr taxonomists today sim- onvmy tied to Linnaean categories, 
and re- 
pi take tor graried secular traditions of dundant taxon names (Christoftersen, 
describing and naming the diversity of na- 
Te Ther mav persist stoically for a life- conflict with evolutionary approaches. 
The 
time in such a sel:-appointed descriptive codes thus help perpetuate essential1stic
role, avoiding theor, philosophy, and ex taxonomic traditions rather than promote 
pianation. Some of these taxonomists may Conceptual evolutionary
nnovations.
rated by the codes, simply conflict with 
1974a, 1974b, 1976). For exampie, manda- 
tor categories, the names o: genera as 
parts of binomiais, typification and syn- 
FiGURE 1. A possible phyiogenv o: intellectual lin- 
eages in biological taxonorn. A-M. sequence of an-
cestor-descendant lineages; N monophyletic taxon of 
multiple lineages or single lineage o 11nterbreedng 
populations. A = Aristotelian taxonomy; B = Linnae- 
an (essentialistic, typolog1cal) taxonomy, C = tradi- 
tional taxonomy; D = extinct qunarnan taxonomy; E 
conventional taxonomy; F= 0mnispeciive tcompi- 
latory, practical, utilitarian) taxononir, G = natural Od LdXOnomy nvolves interpretation of 
taxonomy; H = phenetic (numencal taxonomy (phe similarities and differences as reflecting 
netics);= orthodox taxonomy,! = evoiunonary degrees of phylogenetic relationships. Om-
(Mavrian, Simpsonian, sVntheic, siTICTetistic. gradis- 
tic, eclectic) taxonomy (phyletics), K = Hennigian tax- 
onomy, L phylogernetic taxonOny iphylogenetic cia 
distics), M=ciadisti: taonomy patterr. 
methodoiogica., transtormed ). = phyiogeneuc svs 
tematics (pnvlogenetics 
and lawful relations of numbers as evi- 
dence of the rational plan of creation. In-
Corporating evolutionary theory, conven- 
1987; de Queiroz and Gauthier, 1992) all 
nispective taxonomists explicity reject arny 
connections between a practical and utili- 
tarian taxonomy and the processes 
thought to be responsible for biological di-
versitt eS. Biackweider, 1964). In natural 
taxonom, groups are supposed to be dis- 
COvered in nature rather than fabricated in 
venture intuitive classitications tor their 
CONCETUAL LINEAGES
named groups bu: will often delegate to 
otners the task 0: derivVing evolutionar 
meanings rom the: propOsals. 
EvolunOnarv concepts have still not pen 
etrated tne core 0: nomenclature. The in- tlicting conceptual systems. Following 
ternationa. coaes (internationai Commis- Hull's (1984) and Mishler's (198:) analog 
Sion on Loological Nomnenclature, 1985; 
Lnternat1onal Botanica Congress, 1988; in- research groups, rather
than being defined 
ternational Associaion of Microbial Soci- by the presence of Some necessary and sut 
eties, 1992 are totaly couched in a tradi- 
ficient set of shared ideas, are viewed as 
tional Linnaean ramewOrk developed 
more than 100 vea:s before the widespread 
acceptarnce oi evOiutionary ideas. Purport- 
relations with other lineages. I propose a 
ediy not to interIere with taxonomic free- 
dom, the codesprovide no rules for the svstems of biological taxonomr (Fig.
1). Fo: 
Tecógnition and derinition of taxa and cat- 
egores et the available rules significantly 
Constrain taxonomic practice by tving tax- (individual spokespersons) 
on the phylo-
ot names to categorical ranks. This struc-
ture interteres wjtn the codes own avowed who provided a 
numerical cladistic studv
goals of providing expiicit, universal, and of phylogeneticists 
and a review' of prei 
stablenames tor taxonomy because a taxon ous single-character classifications
of the 
ame must be changed every time it is three main contemporary schools of tax 
placed in a different category (de Queiroz onom. 
and Gauthier, 1994). Furthermore, several 
taxonomic conventions, variously incorpo- atics, and classification have been advocat- 
laxonomists, like other segments of the 
scientific community, are presently divid- 
ed into several tactions that support con- 
1970; Griffiths, 1974a; Wile, 1981: de the mind of the taxonomist. Phenetic tax- 
Queiroz, 1988). When these denominations onomists attempt to quantify data and es-
become interehangeably qualined by such tablish groups by overall similarity. In or-
terms as traditional, numerical, phenetic, 
evolutionary, cladistic, and phylogenetic used to access taxa (similarity, diversity, 
(Fig. 1), their meanings become fmulti�ari- Size of gaps, ecology, behavior, etc.). In 
ous, extensively Overlapping. and sutti- evolutionary taxonomy, species are rede-
Ciently confusing So as to lose much of fined as evolutionary units (MayT, 1942, 
their heuristic and interpretative value íct. Simpson, 1961), and phenetic, Patristic, Charig, 1982; Hill and Crane. 1982; de and cladistic data are combined into a Sin- 
Queiroz and Donoghue, 1090b: Nixon and gle taxonomic system (e.g., May, 1981; 
Wheeler, 1990). lerms and concepts are 
necessarily context bound and shouid be 
allowed to change through time. I provide granted a central role at all levels in the 
here successively more inciusive defini- taxonomic hierarchy. in the last two de-
tions for the basic concepts of taxonomy, 
systematics, and Classification. I hope my 
assignment of several commonly used Mishler, 1987; de Queiroz and Donoghue, 
qualitying terms to distinct conceptual 1990a) (Fig. 1). Pattern cladists are empir systems (Fig. 1) wll reduce ambiguity, 
even if total agreement is not possible. 
Brief haracterizaions ot the sereral a taxonomy (Brady, 1985; Nelson, 1985. 
with biological phylogenies, tnese difterent 
thodox taxonomy, multiple criteria are 
intellectual lineages, with strong historicai 
Conerence, social! cohesion, and adversaria. 
tentative phvlogeny tor these intellectua. 
those that would have prererred to see 
characters (shared ideas) and Characters 
Stuessy, 1987, 1990). In Hernigian taxono-
my, the priniple of common descent is genetic tree, I deier to Carpenter (1987: 
cades, a phiosophical split has developed 
within phylogenetic systematics e.g 
Different concepts of taxonomy, system-
ICists that avoid all assumpthons and pre- 
Conceptions about process in constructing ed by various authors (e.g., Simpson, 196. 
Blackwelder, 1967; Mayr, 1969; Nelsor mal endii nrufpb bitnet. 
SYSTEMATIC BIOLOG VOL. 1995 POINTS OF VIEW 
42 
avoiding the time axis and the production practices. Cladistic taxonomy mav be iden-tified with what has been called the taxic 
Rieppel, 1988b). Phyiogenetic taxonomists (Hennig. 1966:6). Characters are instanta 
are evoiutionary theorists that deduce the neous morphologies (de Queiroz, 1985: 
most useful taxonomic concepts and meth- 296) that are compared among "specimens 
ods from general evolutionary processes at similarstages in their life history" (Wi- 
(de Queiroz, 1985). Notwithstanding this ley, 1981:119). Character generalities are es- 
basic difference in general outlook, manv tablished by the ontogenetic method (Nel-
phyiogeneticists stil promote social cohe- son, 1978; Nelson and Platnick, 1981; 
Patterson. 1982, 1983) and the outgroup 
sights, meetings, and journal. In analogy method (Watrous and Wheeler 1981; Far- 
with the evoiutionary species concept, ris, 1982 Maddison et al, 1984). Nelson 
these two factions may still be considered (1985) calied ontogeny a direct method 
interbreeding populations. Below, I try to and outgroup analysis an indirect method
deemphasize the individuality of cladistic for the reconstruction of the taxonomic hi 
of truncated hierarchies. 
The notion of actual ancestT is incom- 
patible with a logically stringent pattern Eldredge, 1979; Rieppel, 1988a). Pattern 
analysis. In the initial version ot cladistics adistics is a logical and intermallv consis- 
(Nelson, 1971), taxa were defined by tent approach in which evolutionary theo-
shared haracters, and the lines in the 
cladogram indicated relative recency of Viewed as an 'after-the-fact generalization 
common ancestry berween pairs of taxa. In 
this way, the notion of hypothetical com- 
mon ancestors became associated with ed a strict independence of "that aspect of 
cladograms. This procedure, however, still systematics concerned with the identifica- 
produces a temporalized version of a clado- 
gram resembling a phyiogenetic tree, thus 
blurring the concerptual gap between cla- 
dists and phylogenetic taxonomists. 
In the more radical version of trans- 
formed cladism (Platnick, 1979, 1982, 
1985), pattern cladism (Patterson, 1982), or 
methodological cladism (Hill and Crane, 
1982), evolution is not considered a neces- 
sary prerequisite for the practice of taxon- 
omy. The same model may be called a syrn-
apomorphy scheme, where the connecting methods in gerneral and the use of cladistic 
lines in the scheme indicate no more than methods in particular d0 not seem to re- 
shared sets of character generalities. Such quire any assumption about evolution. 
a synapomorphy scheme, or strictly atem-
poral and acausal cladogram, represents a reconstruct evolutionary history, then this 
hierarchy of static homologies obtained assumption will dictate which cladistic 
from downward ciassification br logical methods are useful" (de Queiroz, 1985: 
subdivision (Platrnick, 1977; Patterson, 
1983; Rieppel, 198ba). One of the implica-
tions of this approach is that all taxa are 
treated as classes that are defined by char- 
acters (Beatty, 19821. Platnick (1977, 1985) iples, and practice of identifving (discov- and Patterson (1978) tried to avoid tnis es- ering) systems, .e., of ordering the diver- sentialistic connotation of cladistics by im- sity of organisms (parts) into more general plying that taxa are only recognized by svstems of taxa (wholes) according to the 
apomorphic characters. According to this most general causal processes. Under this 
Conception, characters would be diagnostic definition, systematics differs from taxon- 
rather than defining. This approach, how- omy basically in not attempting to divorce 
ever, leaves cladistic taxa without defini- the practice of building a general biologi- tions. If cladistic theory corresponds to the 
notion that there is order in nature, it is of how general biological processes (in 
agnostic about what taxa represent for bi- contrast to particular biological mecha- 
ology. 
Pattern cladistics represents the most patterns of diversity. Feedback loops be- 
empirical, operational, and objective stage 
attained by Hennigians. it represents the considered fundamental for a constant re- 
logical consequence of striving after the 
most formal cladistic principles and of 
adopting the least theory-laden taxonomic 
approach to systematics (Stanley, 1975; 
ry, rather than a unifying concept, is 
sion by sharing the same Hennigian in- 
to be deduced from taxonomy. Brady (1985:113) is one philosopher that support 
tion of the empirical patterns" from " the- 
ories of process." But is it necessary for 
cladists to use methodological essentialism 
to construct both cladograms and taxono-
taxonomy and attempt to incorporate its erarchy. 
most fundamental contribution-rigorous, in cladistic taxonomy, only groups based 
operational, cladistic analysis-as a basic on synapomorphies (shared derived char-
procedure within phylogenetic systemat- acters) are recognized as monophyletic 
taxa and named. Because ancestral lin- 
eages are not recognizable by positive cla- 
distic data and because ancestor-descen-
mies? 
1CS. 
PHYLOGENETIC SYSTEMATICSs 
CLADISTIC TAXONOMY The theory of evolution was developed 
partly as a generalization from taxonomy 
(Nelson, 1978, 1985; Patterson, 1983; de 
Queiroz, 1985). Thus, the use of taxonomic 
I define taxonomr as the practice of rec- 
ognizing, naming. and ordering taxa into hierarchical pattern, only nested sets of 
a system of words consistent with any monophvietic taxa are admitted in the tax- 
kind of relationships among taxa that the onomy Taxa become operationally diag 
investigator has discovered in nature. Un- nosed by uniquely held characters, and 
der this definition. taxonomy becomes the species become simply the smailest ot 
most basic actiiit in biology, deaiing ex- these Ciusters (Neison, 19895). Br associ 
clusively with tne aiscovery, orderng, and ating species names with 
the jeast inciu- 
communicatior. o patterns of biological sive monophvietic groups, it Wil always 
aant lineages in principle do not form a 
Howeve, "if systematics is an attempt to 
taxa. 
be posiDie to discover Smaller groups n 
Cladistic taxonomy uses a cladogram the future with the retinement of data (e-g. 
(sensu Nelson anc Platnick, 1981) as the molecuiar data). Thus, species will not be 
graphical model for constructing a biolog- 
ical system. A dadogram is a predom 
nantlr bifurcairng. asymmetric, nontrun- recognized. 
Furthermore, some organisms 
cate derndrogram. with no defined vertical or popuiations will 
not belong to any 
and horizonta. axes. A ciadogram is thus monopnt"ietic group because ontologically 
a more genera. Starement than a phvloge- ther are direct 
ancestors or belong to lin 
netic tree, because it does not attempt to eages in which apomorphies 
have not yet 
depict actual ancestors (de Queiroz, 1988: 
250; Rieppel, 1990:187). A cladogram cor- The logic of cladistic taxonomy requires 
responds to a ser of possible trees rather 
than to a singie tree (Platnick, 1985; cf. Wi- 
iey, 1981). Cladograms are constructed by fixed characters ("character" 
sensu Nixon 
maximizing the congruence of several
types of derived characters, usually mor- ual organisms logically become the 
small- 
phological or moiecular. Character congru- est terminal entities for cladistic analysis 
ence is accessed by the methodological (Vrana and Wheeler, 1992). Monophyletic 
prinaple of parsimony (Farris, 1982). The 
basic elements o taxonomy are semapho- ducing what Colless (1977) called a "cor 
ronts, or organisms "during a certain, the- nucopia of categories." Fossil taxa are treat 
oretically inhnitely small, period of lite ed in the same wav as extant taxa, thus 
281). Phyiogenetic systematics is the at-
tempt to deduce evolutionary history from 
the axiom of evolution. comparabie units and will tend to be much 
more numerous than those conventionally define systematics as the theory, prin-
developed. 
no d1stinction between species and mono-
phvletic taxa nor between variable and cal system írom our theoretical knowledge 
and Wheeler's [1990] "attribute"). Individ- 
nisms) are supposed to affect the resultant 
tween patterm and process information are 
taxa are not usually ranked to avoid pro- 
shaping of the best system tor bioiogy 
This view is similar to the concept of Mayr 
(1969) and approaches what Nelson (1970)S!STEMATIC BIO-O3 OL 1995 PO!NTS OF VIEN 
445 
called "comparative bioiog, in that sys- netic tree or a temporaiized cladogram, i. 
tematics must deal with the transtormation which ancestor-descendant lineages ma 
of form through space and thme, thus also be given names. Usually, nested sets o 
integratng intormation on pattern and multiple lineages are also given names and 
process trom ontogeny, paleontologY, and all taxa may be further ranked into a series 
biogeographi. Svstematics is thus both of categories. Of course, there seems to be 
broader in scope than taxonomy and nec- no logical requirement for naming both 
essarilv more theory laden. It integrates a nestec sets and ancestor-descendant se 
ranstormational approach (Eldredge, quences in the same svstem. in a new phy- 
1979; Rieppel. 1988a) with the predomi- logenetic system proposed by Papavero et 
nantly taxic perspective of taxonomy. Riep-
pel (1988a:170) treated patterm and process phylogenetic tree are given names. lt is 
analyses as "different ways of seeing" in- 
compatibie with whiie complementary 
each other. The world may appear static or 
dVnamic, discontinuous or continuous, hi- 
erarchical or linear, as revealed by the taxic system, how to name taxa, and whether or 
versus transtormational approach. These not to rank taxa have led to several ap- 
approaches are complementary in that a proaches within phviogenetic systematics. 
serial or linear hierarchv mav be transiated 
into a subordinated hierarchy by the spec-
incation of inclusive taxa, whereas the sub- 
wise noted that only characters that are 
fixed in terminal lineages produce branch- 
ing patterns. 
The implication for cladistic analy'sis is ties seem to exist at the same hierarchical that there are sensible limits to the appli- 
cation of cladistic methods. If these bound- namically as sequences of individual on- aries are trespassed, cladistic methods wl togenies to be cormpared among organ- still produce a hierarchical pattern oI cnar 1Sms, ontogenetic polarities are seen to 
acters, however poorly resolved, even belong to one hierarchical level (i.e., thev 
when these are known not to form hier-
archical relationships in nature. Bryant ary poiariies are seen to belong to a dis- (1992:259) noted that "'as an inductive inct hierarchical level (i.e., they exist be- 
summary of the data set, a cladogram nas tween characters). Failure to make these 
no separate empirical content; it provides erarchical distinctions may lead to errors 
only a consensus of the hierarchical infor- in cladistic analyses. Ontogenetic polari- 
mation in the data matrix." 
Thus, allelic polymorphisms, ontogenet- alterations in developmental timing (het ic transformations, and populational vari- erochrony; see Gould, 1977), do not nec- 
able characters must be correctly interpret essarily coincide with phylogenetic polar ed in a phylogenetic context. Amorim et al. ities of ontogenies. The sequence of (1993) showed that it is not possible to particular ontogenetic transformations 
transpose the phylogenetic concept of syn- within an organism (character adjacency apomorphy to the lower hierarchical levels in Wheeler's [1990) nomenclature) may be 
without further resolution. Two evolution- irrelevant to cladistic analysis. The critical 
ary events are actualy involved: (1) the elements from ontogeny to be used in cla- 
molecular process of apomorphic modih- 
cation of a preexisting gene at a given io 
cus and (2) the populational process of ple- 
siomorphic alieie elimination at the same 
in phyiogenetic systematics. When static 
morphologies are treated as characters, both ontogenetic and evoiutionary polari- 
level. But when characters are viewed dy- 
exist within characters) whereas evolution- 
al. (1992), for exampie, oniy the lines of the 
also not necessary to rank taxa within the 
Svstem. Such possibilities for choice as to 
which particular phylogenv-based dendro 
gram to use as a model for the biological ties of instantaneous characters, because of 
Cladistic Analysis 
Basic methods of phvlogenetic taxono-
ordinative hierardhy may be changed to a my are the same as those described for Cla- 
serial arrangement of forms by specifving distic taxonomy, but several additional re 
actual ancestors. Phvlogenetic systematics quirements and quaiincations are necessar 
involves integration of these two world In phvlogenetic taxonomy, not all kinds 0 
iews by recognition of two ontological characters empiricaly observed in sema 
phoronts as instantaneous morpnologies 
Snoula be usea in the cladistic analysis 
Phyiogenetic methods are based on the 
premise that there exists a nested hierar-
which are discontinuous taxa ranked hi- dhical pattern of reiationships. It is thus 
not appropriate to apply cladistic methods 
to entities that are not expected to be re 
kands o taxa: species, which are continu- 
ous strings o ancestor-descendant poppu- 
1ations ranked serially (the transtormation- 
a. approach). and monophyletic taxa, 
distic analysis are the relative generalities 
ot different life cvcles of organisms. P'attern 
cladists. br avoiding evolutionary assump-
tions. mai produce taxonomies that are 
named the shared presence of not oniv evolutionarily neutral but are at 
locus. 
apomorphic alleles suntrep:y and the odds with evolutionary thinking" (Beatty, shared absence ot plesiomorphic or apo- 1982:33). An important consequence tor morphic alleles synapous. Because these phylogenetic systematics 1s that the onto-
events occur at ditferent times and may be genetic method of determining character separated by one or more cladogenetic polaritv, like the paleontological method, events, allelic polvmorphisms will be in- becomes a special case ot the outgroupherited by difterent species. Ihis concep- method (de Queiroz, 1985.293). In other 
tual resolution should have important con- words. the two methods are not indepen-sequences for the pars1mont concept as dent (F1nk, 1982; Kluge. 1985), the distinc- applied to allele matrices. lt aiso prOV 1des 
a plausible bas1s tor the intuiive concept tion methods is nonex1stent (Wheeler of ""underlying synapomorph1" (Saether 1990), and ontogeny stands to phylogeny 1979). Many characters no interpreted as as a part-whole relationship (Brooks and homoplasies between taxa mar be found Wiley. 1985). 
to correspond to independeni events o 
synapousy of the plesiomorphic alele. 
Likewise, independent events of sVnapou- 
sy ot the apomorphic alele mav account of data for the same taxa (e g., Doyle, 1992) for many cases ot multiple reversais in 
cladograms. 
De Queiroz (1985) claritied how onto- the estimates of phvlogeny (Kim, 5 genetic transtormations must be handled 335). However, molecular phvlogenies ma 
erarchically (the taxic approach) (Rieppel, 
1988a:105). 
The preterred modern graphic models lated hierarchically. Characters are associ 
or basing biological systems within phv- ated with difterent patterns of evolutionar 
Ogenetic svstematics are ramifying, asym- reiationsnips at 
difrerent huerarchical lev- 
metric denarograms (phylogenetic trees or els. For example, mutations 
at the molecu- 
ciadograms. which mav be viewed as lar level may proauce ontogenetic changes somewhat stvlized and simplified deriva- in organisms. variable characters in popu-
tions ot traditional phylogenetic trees. lations, ifixed characters in species, and 
here are two wavs of applying names to transtormation series in monophvletic 
Such dendrograms (Griffiths, 1974a, 1974b): groups. Only fixed characters and trans- 
(1) the naming o: time-extended lineages tormation series Droduce branching pat 
of ancestor-descendant populations (e.g., terns of genealogical relationships. This 
taxa A-M in Fig. 1) and (2) the naming of view corresponds closely to Hennig 
nested sets of these lineages (e.g., taxon N (1966:31, hg. 6) corntrasts among ontoge 
in Fig. 1) In cladistic taxonomy, an ahis netic (within organisms). tokogenet 
torical cladogramserves as a model for (within populations), and phylogenetic (be 
representing relationships, and oniy nest- tween species) descent systems, in whic 
ed sets are given names. In phylogenetic only the phvlogenetic system produces ni- 
taxonomy, the model becomes a phyloge erarchies. Nixon and Wheeler (1990) like 
tion berween direct and ndirect polariza-
One o the main probie 
porary cladistic analvsis reiers to explor- 
ing methods for conmbining different kinds 
of contem- 
AgTeement among trees estimated by dif- 
ferent methods lends greater credibilitv to 
VOL 1995 POINTS OF VIEW SYSTEMATIC BIOL OG) 447 
that have chosen to emphasize monophyiy Dreeding that may serve to d1stinguish at the lowest levels in phvlogenetic taxon- 
omy (e.g., Rosen, 1979; Hill and Crane, asexual organisms. Cladistic taxonomy has 
1982; Donoghue, 1985; Mishler and Bran- surrace structure (descriptive adequacv) 
don, 1987) are obliged to exclude at least but lacks deep structure (explanatory ad- 
some organisms (viz., those that are ances equacy) (BrookS and Wiley, 1985). A SVs- 
tors and those that have not yet developed tematic technique justified only in terms of 
apomorphies) from their ontological spe- surtace structure has not demonstrated its 
cies concepts. Epistemologically, an artifi relevance to studies of dharacters of the liv- 
Cial dist1nction must then be made be- 
disagree with each 
other and with those gram will ther. be hypothesized to form 
generated from morphologY (Gariand et homologies. Homologies 
shouid be viewed 
al, 1993:289). With the number of molec- as relationships among historicai individ- 
ular analyses incTeasing at a greater rate uals, a perspective that combines 
the taxic 
than that of morphological analyses (San- and the transformational views of homol- 
derson et al, 1995) and with over 100 avail- ogy (McKitrick, 1994:2). Genetically and 
able methods o' pnyiogenetic estimation developmentally individualized ertities 
(Huelsenbeck and Hillis, 1993:247), there is form intraorganismal homologies. Because 
cleariy a problem in identifving those con- 
ditions under which particular methods form reticuiated patterns, onlt interorgan 
and models períorm well or poorly. Opti-
maity criteria such as parsimony are use 
ful, but they must be recognized for what 
thev are: toois that help direct perception 
towards realit (Knight et al., 1993:357). De 
Queiroz (1993) noted that at the higher tax- 
onomic levels disCordances between gene cladogram (sensu Rieppel, 1988a) as 
and species trees due to ancestral poly- graphical model for constructing a biolog- 
morphisms are vanishingly small, and ical system. A phylogeny is 
a predomi 
consequently a combined approach to data nantly bifurcating, asymmetric, and trun 
analysis is usualiy preferable at suprage- cate dendrogram, with time as its vertical 
neric levels. At the species and population axis (Griffiths, 1974a, 1974b). It is only by 
levels, however, much disparity occurs be- the addition of the time dimension and 
or 
tween pnylogenetic patterms based on mor- the notation of ancestry 
that the static ho- 
phology and those based on molecular mologies of cladistic analysis 
can be inter- 
data (e.g., DeSalle and Grimaldi, 1993; preted as evolutionary 
novelties character- 
Sage et al., 1993; Patton and Smith, 1994). izing monophrletic 
taxa (Rieppel, 1988a: 
De Queiroz (1993) suggested that consen- 138). All taxa are 
defined in terms of ge 
sus methods ot aata analysis are more ap- nealogical reiationships 
rather than o 
propriate at these lower levels. However, 
shared attributes (de Queiroz and Gau- 
gene trees, organismal ontogenies, 
and thier, 1990; de Queiroz, 1992). "Distin- 
taxon phviogenies belong to three distinct 
hierarcical leveis, which show part-whole 
relationships among each other. Further- 
more, disagreements among analyses at 
these different leveis seem to be the rule from genealogical relationships 
among 
rather than the exception. Does it make their parts. Descriptions 
and diagnoses are 
sense to provide consensus trees for these epistemological 
statements about how we 
Contlicting data sets? I tend to agree with 
Bull et al. (1993) that when difterent his- 
ciusters of sexual organisms from clones of 
intraorganismal historical phenomena mar 
ing worid that it classifies. Rather than in- 
sisting on a dichotomy between surtace isma! homoiogies will revea! hierarchical 
relationships. 
tween real ciadospecies and provisional 
metaspecies (Donoghue, 1985; Ardhibald, and deep structure, pattern and process, 
1994), some of which may represent actual 
ancestors. 
organisms and groups, cladistics and phy- 
logenetic taxonomy, these part-whole re- Phuiogenetic Taxonomy 
It may be tempting to try to avoid the lationships may be more profitably viewed 
species problem entirely by using only or- 
ganisms as terminal entities for cladistic 
analysis (Vrana and Wheeler, 1992). Under 
this approach, taxa are viewed as relation- vides an explanation tor pattern (Rieppel, 
ships among organisms rather than among 
groups. Neison (1989a, 1989b) concluded
that a basic taxonomic unit does not exist 
and that there is no empirical difference ly and ancestry is to recognize two distinct 
between species and other taxa. The prob-
lem with this view is that it ignores the 
empirical observation that "character in- Under this approach, there is still a tension 
congruence atfects lower leveis of the ge- between those who propose to use inter- 
nealogica! hierarchy to a greater degree breeding (e.g., Hennig, 1966; Gould, 1986; 
than higher ievels" (Rieppel, 1989:57). The Ridley, 1989) and those who reject all pro- 
pars1mony metnod provides a cladogram cess defnitions in tavor of strictiy morpho- 
in which there is maximum congTuence logical pattern detinitions (e.g, Wheeler among character generalities 
this method does not specify the amount My positiorn is that no distinct biological of character incongruence that would re- entity would exist between the organism fute the initial hypotheses of hierarchy and and the monophyletic taxon if it were not 
monophvly. Cladistic methods cannot pro- for the biologica! process of interbreedin8 vide a rigid test for the expectation o hi- lnterbreeding is the most general processerarchical order in nature. Something more known to maintain a reticulated pattern than observations ot organisms is required anmong organisms through time. Conse tor an understanding of character gener quently, interbreeding is of primary inter- alities at more incusive leveis. A covering est to phylogenetic systematics (de Quei-
theory, involving assumptions about bio- roz and Donoghue, 1988) and should 
logical processes such as tokogenesis and continue to be the basis of species con- 
interbreeding, appears to be necessary. cepts. My theoretical (ontological) species con- 
Otherwise, the recurrent character incon- cept defines a species as a single lineage of 
gruences between different empirical data ancestor-descendant sexual populations, sets cannot be accounted tor. Doubts have genetically integrated by historically contingent consequently been raised as to the adequa- events of interbreeding. This concept is clos-
cy of the cladistic approach to phylogeny est to what has been called the evoiution- 
reconstruchon at low taxonomic levels (Ar- 
nold, 1981). Many phylogeneticists have re- Because there are empirical limitations to jected the very' process assumption (inter- 
Phvlogenetic taxonomy uses a phyloge- 
ny (sensu Hennig, 1966) or a temporalized as complementary. Under such an ap- 
proach to systematics, pattern provides 
process with a direction, and process pro- 
1988a:451). 
The only alternative to the problem of 
logical incompatibility between monophy- 
ontological entities in phylogenetic sys- 
tematics: monophyletic taxa and species. 
gushing descriptions shouid not 
be con- 
fused with definitions" (Beatty, 1982:27). 
Definitions are ontologicalstatements 
about the existence of entities that result 
However, and Nixon, 1990; Davis and Nixon, 1992). 
recognize the parts of those 
entities (de 
Queiroz and Gauthier, 1990:307). 
in phylogenetic taxonomy, 
ancestor-de 
tories are detected by different data anal-
yses, it may be best to keep the datä sets scendant relationships 
are recognized in 
separate The next logical step wouid be to 
addition to the sister-group relationships 
Identity the causes of heterogeneity and re- of 
cladistics because ancestors belong on 
interpret the results accordingiy. 
If cladistic analysis is to avoid tautology, time-extended lineages 
of ancestor-de- 
homology cannot be resolved only by em- scendant populations are 
conveniently
pirical observation. Homology is distinct named as species. 
Under this procedure, 
from synapomorphy (de Queiroz, 1985: however, the notion of monophyly 1S n 
280). Similarities must first be analyzed by compatible with the notion of ancestry 
be 
parsimony criteria and summarized on a cause monophyletic species cannot be an 
cladogram. synapomorphies on this clado cestors (Rieppel, 1988a:157). Those authors 
tologically to our evolutionary models. 
The 
ary species concept (see Wiley, 1981:25) 
the possibility of recognizing evolutionary 
450 SST -TIC BIOLOT 1995 
POINTS OF VIEW 51 
tionary ranking. Transiormed ciad1stics, 
when divested from evoiutionary theory, 
detailed biogeographic reconstructions are becomes a strictly methodological actvitv 
still not possible for before the Jurassic. In It will not produce the best taxonomy be- 
cause gTOups known to #orm reticulated 
in the reconstruction of global tectonic pro- 
cesses in the Paleozoic (eg, Young. 1986), 
TABLE 1 Pnviogenetic taxonomy of Olbogastridae based on Aristotelian essentiaiistic iogic 
With the subsequent addition ot more ca ptera. bibionomorpha), with biogeographical labeis 
egories to the system, these ranks have be- 
come quite arbitrary, represent1ng no mOre 
than an empty formalism that has lost 1ts
theoretical idealistic foundation. After the 
added to each taxon name. Biogeographical labeis are 
based on an area ciadogram for the history of inter 
cont1nental reiationships. Composite labels indicate 
the distibutions ot successively' subordinated ances- 
tral speaes (ether extinct or not sampled), the last 
term indicat1ng the recent representatives (AtOr = 
AITO-UTIental, Au = Northern Australia, CAn Cir- 
these cases, Amorim (1994) proposed pro- 
visional labels based only on minimum patterns in nature will be nevertheiess 
age classes (for exampie, TR', TR, TR, forced into a hierarchy. When erolution is 
etc., for successively older fossils in the made the basis of taxonomy, a spiit be- 
tween pattern cladists and phvlogenetic 
taxonomists is no longer justified. Under 
es determined by paleontological data evolutionary assumptions, the most useful 
Could be graduallr substituted tor age of methods developed by ciadists can be se 
penetrating analvsis of Griffiths (1974a 
1974b. 1976) on the difticulties oi recond: cumantarctc, G = Gondwana, GTe = lemperate 
ing the Linnaean categories with the phy- Gondwana GTTropical Gondwana NoAuN 
logenetic system. many phviogenetiists 
have abandoned further attempts to Cias-
sir taxa into ranks. 
Historical classification corresponds to a convention of Christoffersen (1989) for indicating re- 
Hennig s (1966) original proposal ot cor- dundant. phylogenetically 
uninformahve taxa (adapt- 
relating absolute ages of ongn o: taxa 
ec Tom Amonm, 1992.288, 
with age classes delimited by convention 
rom the geologic time scale. Subsequent 
diffhculties in implementing such a scheme 
in practice ied Hennig (1969, 1981) and 
some followers (e.g, Løvtrup, 1977; Wil 
mann, 1989) to adopt an alternative nu- 
I riassic and similarly for successively old- 
er time periods). These minimal age 
class- 
Australian; NoAus Neotropicai-Northern 
Neotropica:-Southerm Australian; NoN = Northerm
Neotropica., NoS = Southern Neotropical 
Pangaea. The square brackets in the taxonomy follow lected and used for the construction of a 
origin of the taxon and then for a biogeo-
graphical category if the necessary geolog-
Ical evidence ever becomes available. Even 
phylogernetic taxornomy. At the higher lev- 
els of systematic generalization, all phylo 
though provisional age classes will always genetic nVpotheses become retrospective 
remain partially arbitrary and subjective narratives. In these cases, there is basic 
(Craske and jefferies, 1989), they are still of agreement between pattern cladists and 
phylogenetic taxonomists on how to rec 
Ranking according to Amorim's (1992) ognize and name monophyletic taxa. Con- 
proposal is based on biogeography and ceptual and methodological disagreements 
stratigraphy. Such extrinsic data are best at the lower taxonomic levels result be 
kept separate from the intrinsic data of cause gene trees, organismnal ontogenies, 
and taxon phylogenies may produce in 
the earth, however, is causally related to congruent histories tor each of these hier-
archical ievels. My erolutionary species 
concept iS ontologically based on inter- 
ical data sets and their representation side breeding and epistemoiogically recog 
nized br diagnostic characters. Codes o 
nomenciature that do nor tie taxon names 
which particular evolutionary mechanisms to categorical ranks are required. Such a 
release irom Linnaean conventions would 
Olbiogastridae P-G 
new subtamily GTe-CAn-NoAuS-NoS 
Olbiogastrinae GIrT 
new tribe [new genus 1] AfOr 
Olbiogastrini NoAuN 
new genus 2 AuN 
Olbiogaster NoN 
some heuristic value 
mericlature. But none of the alternatives to 
the subjective Linnaean categories has won 
widespread use. 
These ditficulties notwithstanding. he formation (characters). 
Griffiths (1974a, 
demand for evolutionar1 ranking 1s Sili 
warranted because ranking adds retnea- cause o the noncomparability among 
dif- 
ble intormaior. content to the bi0:0gica erent 
characters, we are still very far trom 
SVstem. T like the broader 
connotatior o: being able to measure overall
ditterences 
the term evolutionary, as intended by tne among 
taxa. 
evolutionary taxonomists, for transmiting 
the idea of added information. 
Howeve.
am here advocat1ng Hennigian pr1napies 
to attain evolutionary ranking. In this re- 
spect, theorists of the evolutionar 
school 
such as Mayr (1969, 1974, 1981 and Mayr 
and Ashlock (1991) have misunderstood 
does not interfere with the names 
of taxa 
phylogenetic taxonomy. The evolution of 
the evolution of its biota (Rosen, 1978). 
1974b, 1976) showed compellingir that be- Congruence between these largely empir- 
by side in a general reference system 
should provide the best hypothesis against 
Amorim (1992) proposed a method of 
sV'stematization n which a 
label corre- 
sponding to the biogeographical compo- 
nent of each taxonomic level is added 
to 
may be tested. 
In phylogenetic systematics, only mono- immediately promote stability of taxon 
phyletic taxa with extant representatives names. Furthermore, this action ouid 
are ranked into age classes or biogeo- permit ranking to proceed gradually and 
graphical categories. Species, which are independently of taxonomy. Ranks based 
time-extended lineages logicall distinct on the sy'stematization o: biogeographica 
from monophyietic taxa, and taxa com- components add extrinsic evolutionar in- 
posed exclusiveir of fossils, which form formation on the age o origin and distri 
truncated phvlogenies and have unstable 
positions in the system (Griffiths, 1976), and mav provide the most useful general
should not be formaliy ranked. Fossils may reference iramework for comparative biol 
simpiy be sequenced or indented in the ogy. 
system of extant taxa, with such designa 
tions as monopBylhumm (Lauterbach, 1989) for 
monophyletic taxa and plesicrm (Patterson 
and Rosen, 1977) tor the remaining taxa. 
the nameof a taxon (Table 1). One of the 
great advantages of this proposal 
is that it 
tne dist1nction between 
classincation and in the preexisting ciassification 
nor must 
systematization. The svstems structure 
ot 
real phylogenies is ciadistic (branching.. ir 
respective ot what principie for classiry1ng 
form the basis tor biogeographical catego- 
taxa is adopted (Griffiths, 197-a). Furiher 
more, Mayr and Ashlock's (1091) concept 
cates taxa that have the same age of orign 
ot evolutionary' ranking is not erolutionary 
and that have ancestral species living n 
at all because it 1s based on overall 
s1rm- the same geographicai area. Ihus, infor- 
larity of included taxa rather than on any 
explicit evolutionary criterion. 
Cladistic areas may be gradually associated with in 
and phenetic data cannot be successively 
formation on the phvlogenetic history or 
Combined into a single taxonomic system 
taxa into a single biological system. One 
(cf. MavT. 1981; Stuessy, 19S7) because limitation 
of this proposal reters to taxa 
these data are obtained from differen: and originating before the fragmentation or 
Contlicting analyses of the same basic in- Pangaea. Although progress 
is being made 
the Linnaean categories be abandoned 
im- 
mediatel. The ne label may' 
nevertheless
bution o: the ancestral species of a clade 
ries. Ihe same biogeographical 
label indi-
ACKNOWLEDGMENTS
A researdh scholarsh1p from Conselho Nacional de 
Desenvolimento Cientifico e Tecnoiógico during the 
last 15 vears has permitted me to acciumulate a com 
prehensive collection of literature even though I work 
in relative isolation 2.500 km awav irom the nearest 
provisiones libranes. Kesearchers oni the 
whole rave 
been partcularl" generous r senaing me reprints. 
Editor Michaei Miivamoto has been courteous and 
mation on the history of biogeographical
CONCLUSIONS
I have argued for a three-steP procedure 
in phylogenetic systematics: cladistic anal-
ysis, phylogenetic taxonomy, and evolu- 
:STEMATIk B.c.OG OL 44 1995 PO:TS OF VIEV 53 
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MAYR, E 1942. Systemat1cs and the origin ot species 
from the viewpoint ot a zoologist Columbia Unuv. 
Press, New York. 
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ham Grifiths, Oliver Kieppel, Kevin ae ueiro2, and 
Dalton Amonm, through tneir insighttu. purications 
have done much to focus mt jdeas in taionomy Iwo CHRISTOFFERSE . L 1989 Phvlogent and classifi-
anonmous rererees änc one assoaate eaito: maae 
helpful suggestions on earier verSIONs 0: manu-
script. Such joint efforts aid much 
to improve tne final OLLESS, D H 19,.. A cornucopia of categories. Svst. 
quaiir ot this manusct let the whoe process 
made me very conscious that I can blame oniv mvself 
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