CONCLUSIONS AND SUMMARY
As recently stated by Gingerich
and Uhen (1998, p. 3), "Evolution is first and foremost a history of
ancestors and their sometimes-divergent descendants." This is a Darwinian
generality that has profound consequences for estimating the evolutionary
history of all organisms, but one that is often ignored by systematists who
practice a taxogram approach.
The obvious question is, what are the guidelines
or parameters that circumscribe and inform transformational or polarity
determinations of characters and the relationship of taxa beyond a sometimes
compelling fossil record of successive fossils samples of the same kinds of
organisms? As briefly outline in this paper, the Darwinian context of lineage
transformation sets the boundaries within which the contingent attributes of
different lineages at all time levels, or stages of the same lineages, may be
recognized and weighted as being either more probably homologous or homoplasious.
Practitioners of mathematical linearity as a substitute for biological and
paleontological analysis—and who, as a rule, do not order and polarize the
characters they use without algorithms—eschew the need for a conceptually
interdisciplinary and Darwinian context for character analysis. Parsimony-based
cladists customarily rely on diverse data bases without any causal input into
the selection of these traits and employ algorithm-driven congruence to order
and polarize characters. They claim to follow such linear practices in order to
fulfill the need for notions of "consistency" and 'scientific' (=
Popperian) respectability for the testing of what is in fact a highly
idiosyncratic and mosaic manifestation of lineage choreographies of evolving
species and speciation events, phenomena that do not correspond with
paleontological species taxa. They hold, for all intents and purposes, that a
tested disembodied "genealogy" of taxa can be attained without an
understanding of the factors responsible for the properties of organisms.
The following major points are a brief summary of
some aspects of these issues, which are discussed in the body of the paper in
some detail.
- What phylogeny (but not classification) is has bearing on how we should try
to recover it. Descent with ancestrally constrained (but also facilitated)
adaptive (and other) modifications, largely through natural and sexual
selection, contains a host of post-Darwinian advances in evolutionary biology
that relate to this fundamentally Darwinian view of the evolutionary process.
This complex array of precepts is both the theoretical foundation of a Darwinian
phylogenetic analysis and the basis for any comprehensive theory of function (in
a broad sense) and structure. Meaningful functional (in a full Darwinian sense)
and phylogenetically oriented biology is the study of the results of selection
and other attributes, and of the process of their acquisition.
- Fundamentally structuralist approaches consider the goals of adaptational
and other evolutionary analysis to be unattainable for fossils and extant
organisms. Yet adaptations or sexual-selection- related paraphernalia in extinct
species are often better understood than their phylogeny. Such attributes may
form the framework of an analysis. A functional and adaptational (ecologically
utilitarian) assessment of traits in both extant and extinct organisms (whenever
possible) is necessary in order to reliably establish polarities of homologous
features. Such practice enables students of phylogeny to cull convergent
attributes from a body of characteristic features. The Darwinian method,
contrary to misconstrued characterizations that it permits only the use of
adaptive characters in phylogenetics (e.g., Schuh
2000), informs of the incidental and phyletically relevant attributes in
assessing the adaptive solutions. The homologously shared and derived, or
transformationally homologous, features so obtained are the taxonomic properties
against which lineage and taxon phylogeny hypotheses may be tested.
- A Darwinian evolutionary explanation involves both the causal and
historically mediated components of a particular transformation, an evolutionary
becoming. Its macrotaxonomic expression is constrained by heuristics, which are
not directly relevant to phylogenetics. A sharp divide between functional and
evolutionary explanations, distinction are more practice-based than
theory-based. These should be replaced by a less dichotomous, less hierarchic,
and far more interrelated or "temporally-looped" set of theories and
concepts regarding the relationship between the evolution of function and
biorole of features. Functional analysis (broad, Darwinian sense) and the
biostratigraphical record are the valid bases of transformational analysis of
organismal attributes in phylogenetics (Bock
1981). The results of the transformational analyses of features and the
subsequent understanding of the relationships of lineages, independent of
parsimony taxograms, are prerequisites for a meaningfully tested taxon
phylogeny. Both adaptational and phylogenetic analyses are inferential about
specific events in the past. Mapping the results of the "causal role
function" approach of Lauder and colleagues onto taxograms, a structuralist
perspective that is now labeled by its practitioners as the "new
adaptationism," does not advance the cause of phylogenetic reconstruction.
- In spite of the obvious data-based limits of the past that were previously
noted, there is no reason to reject either specific historical narrative
explanations, or their rigorous tests, dealing with adaptations, or phylogenetic
estimates of character transformations. These activities are the core of
comparative biology and macroevolutionary science, and the testing of
phylogenetic hypotheses of lineages and taxa fundamentally relies on them (Figure
1). Organisms are adapted (= "fit," in a fully Darwinian sense)
and an understanding of selection-produced attributes (e.g., mechanical
adaptations or dimorphisms) can have a significant influence on choosing
features against which phylogenetic estimates can be tested. Thus, functional
understanding is essential in any estimation of phylogeny. Model-based analysis
of attributes of selected extant species is a cornerstone of this activity.
- Taxograms, bereft of specific taxic derivation (origins), do not reflect the
lineage-based reality of phylogenetics. Axiomatized attempts to reflect the
history of organisms as taxograms, node-based holophyla, without paraphyla is an
illusory practice. It is contradicted by the continuity of all lineages in the
evolutionary process, and the necessity to make arbitrary demarcations when
delineating taxa.
In closing, I note that the attempt at the
expurgation of the fully Darwinian (= adaptive) context of evolutionary change
from both conceptual and operational methods has been largely driven by a belief
that analysis of patterns independent of tested evolutionary theory will yield a
theory of evolution of such generality, and that this theory will replace an
expanding Modern Synthesis (see Eldredge
and Cracraft, 1980; Grande and
Rieppel, 1994; Rieppel
and Grande 1994). Beyond claims that Darwin has been reinvented (Eldredge
1995; and in this process, not coincidentally, omitted from phylogenetics)
and the rise of a hardened taxic paradigm of parsimony-based cladistics, nothing
of the sort has materialized thus far.
