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Evolution__3rd_Edition

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..<br />

Protein and DNA sequences have<br />

been transcribed in recent decades<br />

Molecules are used in phylogenetic<br />

inference ...<br />

CHAPTER 15 / The Reconstruction of Phylogeny 437<br />

phylogeny c, and 20 to other idiosyncratic arrangements. We then study the characters,<br />

identify the homoplasies and ancestral homologies, and discard them. Maybe 30 of the<br />

100 initial characters remain. If all 30 point to the same phylogeny, our job is done. But<br />

in practice, 20 of the derived homologies may support phylogeny a, six support b, and<br />

four support c. The reason is probably that some of the characters we think are derived<br />

homologies are in fact homoplasies or ancestral homologies.<br />

We have four options when faced with conflict in the evidence: we can scrutinize,<br />

and rescrutinize, the contradictory results to test their reliability; we can suspend judgment;<br />

we can collect more evidence; or we can infer that the phylogeny supported by<br />

the most evidence is the correct one. If 20 of the 30 characters support phylogeny a,<br />

then we could infer a is the correct answer. The viability of the four options will vary<br />

from problem to problem.<br />

15.8 Molecular sequences are becoming increasingly<br />

important in phylogenetic inference, and they have<br />

distinct properties<br />

The sequences of proteins and DNA are both used in phylogenetic inference. Proteins<br />

blazed the trail. The first protein to have its amino acid sequence worked out was<br />

insulin, which was sequenced by Sanger in 1954. Protein sequencing became an automated<br />

process through the 1960s, and the sequences of some proteins, such as<br />

cytochrome c and hemoglobin, became available in enough species for large-scale phylogenies<br />

to be inferred. DNA sequences followed on, about 20 years later. It was Sanger<br />

again who sequenced the first decent-sized sequence of DNA, in this case the whole<br />

genome (containing 5,375 bases) of the bacteriophage φX164, in 1977. DNA sequencing<br />

since then has expanded, almost explosively, and most current molecular phylogenetic<br />

work is concerned with DNA sequences. Many of the methods and concepts of<br />

molecular phylogenetics were established for proteins, however, and here we consider<br />

the two kinds of molecules together.<br />

The deep logic of phylogenetic inference is identical for molecular and morphological<br />

characters, but the two have distinct properties and the methods and concepts<br />

used for each can appear very different. The homology/homoplasy distinction, in<br />

particular, differs for the two. When confronted by apparently conflicting homologies<br />

for morphological characters (like the wings of birds and bats), the first thing to do is to<br />

re-examine the organs, and their embryology, in detail to see whether their similarity<br />

really is fundamental, or superficial and homoplasious. Homology is a powerful concept<br />

for morphological organs such as wings. Wings are complex in structure and can<br />

take on an almost infinite variety of shapes; they have an embryonic development and<br />

morphological relations with the rest of the body. If the information in the structure<br />

and development of a wing in two species is the same, those wings are highly likely to<br />

evolved from a common ancestor who had similar wings.<br />

The homology/homoplasy distinction is much less powerful for molecular evidence.<br />

Suppose a nucleotide is identical in two species. <strong>Evolution</strong>ary changes take place<br />

among a very limited set of alternatives (the four bases A, C, G, and T) and it is fairly<br />

probable that the same informational state could independently evolve in the two

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