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Evolution__3rd_Edition

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

Homologies are recognized by ...<br />

. . . structural similarity ...<br />

. . . relations between parts ...<br />

. . . embryonic development ...<br />

. . . and other criteria<br />

The phenetic similarity between the wolves is homoplasious and is not due to a close<br />

phylogenetic relationship.<br />

15.4 Homologies can be distinguished from homoplasies by<br />

several criteria<br />

Homoplasies do not indicate phylogenetic relationships, and the first task is to recognize<br />

homoplasies, as opposed to homologies. How can we recognize them? The crude<br />

answer is that homologies are identifiably the same character in two species, but homoplasies<br />

differ in some way that suggests the character has evolved independently in the<br />

species that possess it. Research therefore starts with a character that shows some similarity<br />

in two (or more) species, and then examines the character in detail to find out<br />

whether it really is the same in all the species.<br />

First, if a character is homologous it is likely to have the same fundamental structure.<br />

The wings of birds and bats, for example, are superficially similar; but they are constructed<br />

from different materials and supported by different limb digits (Figure 15.3c).<br />

The differences suggest that the bird and bat wings are homoplasies, and evolved independently<br />

from a common ancestor that lacked wings.<br />

Second, homologies usually have the same relations to surrounding characters.<br />

Homologous bones, for example, are usually connected in a similar way with their surrounding<br />

bones.<br />

Third, the character is likely to have the same embryonic development in different<br />

groups. A character that looks similar in the adult forms, but develops by a different<br />

series of stages, is unlikely to be homologous. One example, which we meet again in<br />

Chapter 16, is the relation between a barnacle, a mollusk such as a limpet, and a crab<br />

(Figure 16.1, p. 474). At least superficially, the adult form of a barnacle is more like a<br />

limpet than a crab. The relations of barnacles had been uncertain for centuries until<br />

John Vaughan Thompson discovered their larvae in 1830. The barnacle larva is very<br />

like the larva of several groups of Crustacea, and unlike those of mollusks. Barnacles<br />

therefore share a more recent common ancestor with crabs than with limpets. The<br />

similarities between the adult barnacle and limpet, such as their hard external armour,<br />

attachment to rocks, and feeding through a hole in the shell, are all homoplasious.<br />

Finally, some other criteria can sometimes be useful. Convergence is caused by natural<br />

selection, when organisms in different evolutionary lineages face similar functional<br />

requirements (such as flying in birds and bats). We have grounds for suspecting that a<br />

shared morphological structure may be homoplasious when the species that share it<br />

clearly need it for their way of life.<br />

The criteria in this section are not the only ones that can be used to distinguish<br />

homologies from homoplasies. However, the criteria discussed here do illustrate that<br />

we have techniques to analyze characters shared between species to distinguish homologies<br />

from homoplasies. A homology can be recognized as a character that has fundamentally<br />

the same structure, relations with surrounding parts, and development, in a<br />

set of species. Once the homologies are (often tentatively) identified, they can be retained<br />

in the list of evidence used to infer the phylogeny. The homoplasies are discarded.<br />

..

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