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Evolution__3rd_Edition

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Summary<br />

1 Phylogenies show the ancestral relations between<br />

species. For each species, a phylogeny shows which<br />

other species (or group of species) it shares its most<br />

recent common ancestor with.<br />

2 Phylogenetic relations are inferred using the shared<br />

characters of species. The characters can be morphological,<br />

in living and fossil species, or molecular.<br />

3 When different characters imply the same phylogeny,<br />

phylogenetic inference is easy; when they do not, other<br />

methods are needed to unravel the disagreement.<br />

4 Theoretical arguments suggest that some kinds<br />

of shared characters indicate phylogenetic relations<br />

reliably, whereas others do not. Homoplasies and<br />

ancestral homologies do not reliably indicate phylogenetic<br />

groups. Derived homologies do. Phylogenetic<br />

inference should be based on derived homologies.<br />

5 Techniques exist to distinguish homologies from<br />

homoplasies, and ancestral homologies from derived<br />

homologies.<br />

6 Character polarities can be inferred, with varying<br />

degrees of certainty, by outgroup comparison, the<br />

fossil record, and other criteria, including paralog<br />

rooting.<br />

7 For molecular characters, the kinds of character<br />

analysis used with morphology are often inapplicable.<br />

Further reading<br />

CHAPTER 15 / The Reconstruction of Phylogeny 467<br />

Phylogenies are usually inferred by statistical techniques.<br />

The three main classes of techniques are distance<br />

methods, parsimony, and maximum likelihood.<br />

8 Distance methods group species according to their<br />

molecular similarity. With parsimony, the best estimate<br />

of the tree for a group of species is the tree that<br />

requires the fewest evolutionary changes. With maximum<br />

likelihood, the best estimate of the tree is the<br />

one that is most probable, given a model of sequence<br />

evolution.<br />

9 Molecular evidence is often used to infer the phylogeny<br />

in the form of an unrooted tree. An unrooted tree<br />

specifies the branching relations among species, but<br />

not the direction of evolution.<br />

10 Molecular phylogenetics is used in medical<br />

research to identify the source of emerging diseases.<br />

11 Molecular phylogenetic inference encounters problems<br />

with sequence alignment, the large number of<br />

possible trees, multiple hits, inadequate data, unequal<br />

rates of evolution, and confusions between paralogous<br />

and orthologous genes.<br />

12 The unrooted tree of the Hawaiian fruitflies is the<br />

most firmly established phylogeny of any large group<br />

of species. It has been reconstructed using chromosomal<br />

inversions.<br />

Felsenstein (2003) is an authoritative book about phylogenetic inference. However, for<br />

most of the further reading it makes sense to distinguish between cladistic references, in<br />

which rooted trees are inferred using character polarities, often with morphological<br />

evidence, and molecular references, in which unrooted trees are inferred by some statistical<br />

method.<br />

Cladistics. For the cladistic method, see Wiley et al. (1991) and Kitching et al. (1998);<br />

references can be traced from these sources. Kemp (1999) concentrates on fossils, but<br />

introduce cladistics generally. Hennig (1966) is the classic reference. Sober (1989) is a<br />

philosophical discussion of most of the main methods, and he has also edited an<br />

anthology (Sober 1994) that reprints a number of relevant papers. Another philosophical<br />

question, not covered in this chapter, is whether phylogenetic reconstruction

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