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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 nEto a SISt£MT ITA t Ecoioga. uniersuzae keaETa. ia Paraiva 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 new ram:ies for the Crangonoidea and Alpheoidea Custacea. Decapoda. Car1dea). Cladist1cs 3:348- DISON 1984 Outgroup anaiysis anc parsimony Syst. Zool. 33:83-103 MAYR, E 1942. Systemat1cs and the origin ot species from the viewpoint ot a zoologist Columbia Unuv. Press, New York. constructive in his handiing of the manu>T:)I. Gra 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 tor al the errors and hybnd results tiha: stil remain. Tweeri ontogeni and phv.ogerni Paleobioiogv E.25 264 GARLAND, T., JR., A. K. DiCKERVMAV, C. M. JAVIS, AND A JONES 1995 Pnviogenetic anay'sis Ot cOar- ance by comDuter Simulatio SVSt. B10. 2:265-292. GouLD. S. I 1977. Ontogent and phylogen. 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