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3 The New York Years (1931–1953)

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324 10 Systematics and Classification<br />

logenetic analysis and phylogenetic classification; the term “cladistic” was coined<br />

by Mayr). Taxa are based on the joint possession of derived characters (synapomorphies):<br />

“Hennig deserves great credit for having fully developed the principles<br />

of cladistic analysis, especially the clear recognition of the importance of synapomorphies<br />

for the construction of branching sequences.” This step of the analysis<br />

is important for the cladist, the evolutionary systematist, and the historical biogeographer.<br />

<strong>The</strong>re is no argument over the value of cladistic analysis and the final<br />

product, the cladistic diagram or branching pattern for evolutionary studies. Arguments<br />

arise over the construction of a classification from the cladogram (Mayr<br />

1974g). Hennig (l.c.) proposed simply to translate the cladogram into a hierarchical<br />

classification reflecting only the recency of common descent (“branching point”),<br />

whereas evolutionary classification as recommended by Mayr, Simpson and others<br />

reflects common descent and evolutionary divergence. Mayr’s objections to<br />

cladistic classifications may be summarized as follows (Mayr and Bock 2002h):<br />

(1) Relationship means to the cladist only kinship in a strictly genealogical sense<br />

rather than inferred amount of shared genotype or gene content;<br />

(2) Monophyly was, since Haeckel, a distinctly “retrospective” term referring to<br />

the nearest relatives inferred to have descended from the same stem species.<br />

Hennig added the following qualification: “[…] and which includes all species<br />

descended from this stem species.” Monophyly became a “prospective” criterion<br />

and groups which do not include all its descendent species became “paraphyletic.”<br />

Ashlock (1972) discussed how different Hennigian phylogeny is<br />

from Haeckelian phylogeny universally employed by traditional taxonomists.<br />

<strong>The</strong> term monophyly refers to Haeckelian phylogeny and Ashlock introduced<br />

the term holophyly for Hennigian phylogeny.<br />

(3) <strong>The</strong> neglect of the dual nature of evolutionary change: All phylogenetic splits<br />

have equal weight for the cladist, just as all characters have equal weight for the<br />

pheneticist. Autapomorphies or anagenetic invasion of new adaptive zones,<br />

the existence of minor or major “grades,” and mosaic evolution are all ignored<br />

in the construction of a Hennigian ordering system. <strong>The</strong> cladist acts as if he<br />

assumed that all lineages diverge in an equivalent manner and that genealogical<br />

distance corresponds to genetic distance. <strong>The</strong> reptiles, a “paraphyletic” group,<br />

represent a well-defined “grade” between the amphibian level and that of the<br />

derivates of reptiles: birds and mammals.<br />

(4) A purely formalistic species definition: A cladistic species is simply the distance<br />

between two branching points on a phylogenetic tree of related species. This<br />

is both unbiological and unrealistic. If one “branch” has not changed, it is<br />

the stem species which has continued and from which another species has<br />

branched (or “budded”) off. Also, a simple dichotomy into two daughter<br />

species is not the rule, as shown by superspecies often consisting of more than<br />

two allospecies.<br />

(5) <strong>The</strong>modeoforiginofhighertaxa:<strong>The</strong>evolutionofanewhighertaxonis<br />

correlated with the invasion of new adaptive zones by existing species and<br />

has nothing to do with a splitting event (speciation). It is the later process

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