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Evolution__3rd_Edition

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490 PART 4 / <strong>Evolution</strong> and Diversity<br />

Summary<br />

1 There are two main principles, and three main<br />

schools, of biological classification: phenetic and phylogenetic<br />

principles, and phenetic, cladistic, and evolutionary<br />

schools. The schools differ in how (if at all)<br />

they represent evolution in classification.<br />

2 Phenetic classification ignores evolutionary relations<br />

and classifies species by their similarity in appearance;<br />

cladism ignores phenetic relations and classifies<br />

species by their recency of common ancestry; evolutionary<br />

taxonomy includes both phenetic and phylogenetic<br />

relationships.<br />

3 Phylogenetic inference is uncertain and phenetic<br />

classification has the advantage that it is not subject to<br />

revision when new phylogenetic discoveries are made.<br />

4 Phenetic classification is ambiguous because there is<br />

more than one way of measuring phenetic similarity<br />

and the different measures can disagree.<br />

Further reading<br />

5 Cladism is unambiguous because there is only one<br />

phylogenetic tree of all living things.<br />

6 The cladistic philosophy specifies the pattern of<br />

groups within groups in a classification, but not the<br />

ranking of those groups. The large number of nodes in<br />

a fully resolved phylogeny can be difficult to represent<br />

in a Linnaean classification.<br />

7 <strong>Evolution</strong>ary taxonomy avoids some of the extraordinary<br />

properties of cladism. But it suffers from the<br />

ambiguity of phenetic taxonomy, and its argument for<br />

excluding one kind of phenetic relation (convergence)<br />

works equally well against the kind of phenetic relation<br />

(differential divergence) that it includes.<br />

8 Living things show a diverging, tree-like pattern of<br />

relationships. Darwin explained this by his “principle<br />

of divergence”: that competition is stronger between<br />

more similar forms, forcing them to evolve apart.<br />

Schuh (2000) is a recent book about biological systematics. I have previously discussed<br />

many of the points discussed in this chapter, at an introductory level and at greater<br />

length, in Ridley (1986). Sneath & Sokal (1973) is the standard work on numerical<br />

taxonomy; Sokal (1966) is a clear introduction. Hennig (1966) is the classic work on<br />

cladism a but he is not an easy read! Wiley et al. (1991) and Kitching et al. (1998) are<br />

two texts on cladism. Mayr (1976, 1981, and Mayr & Ashlock 1991), Dobzhansky<br />

(1970), and Simpson (1961b) are key works by key evolutionary taxonomists. Mayr &<br />

Diamond (2001) classify the birds of Melanesia according to traditional “Mayrian”<br />

principles.<br />

Bryant & Cantino (2002) review criticisms of phylogenetic nomenclature. Science<br />

March 23, 2001 contains a newspiece about phylocode. Benton (2000b) defends the<br />

traditional (cladistic) practice. Ereshefsky (2001) is more critical. References to the<br />

primary sources can be traced through these sources. This controversial topic can be<br />

followed in occasional papers in the journals Systematic Biology and Cladistics. See<br />

also the website www.ohio.edu/phylocode.<br />

On the principle of divergence, Darwin (1859) is the obvious source. Bolnick (2001)<br />

did an experiment showing how evolutionary divergence, and adaptation to exploit<br />

new resources, is more likely in the presence of competition.<br />

..

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