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Principios de Taxonomia

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74j 4 What are Traits in Taxonomy?<br />

protecting bone plates (instead of scales). The absence or additional presence of<br />

particular fins, as well as significant differences in jaw shape, body shape, tooth<br />

structure, the <strong>de</strong>fensive spines and in body coloration is also striking.<br />

Most striking are the two forms of Sticklebacks in many North American lakes.<br />

One lives in the open <strong>de</strong>eper water, and the other lives on the bottom in shallow<br />

areas. The <strong>de</strong>ep-water form exhibits a pronounced, spined ventral fin, similar to the<br />

original sea dweller. The shallow water form lacks both the fin and spine<br />

(Prud homme et al., 2006).<br />

All of these forms are evolutionarily very young. They evolved after the last ice age,<br />

and some forms evolved several times in<strong>de</strong>pen<strong>de</strong>ntly from each other. In captivity,<br />

these phenotypically very different fish can be crossed with each other artificially. Of<br />

course, this does not say anything about their species status, but it has an important<br />

practical consequence for the scientist. The genetic cross-compatibility allows for the<br />

i<strong>de</strong>ntification of genes involved in the reduction of the pelvic fin. Surprisingly, the<br />

remarkable anatomic alterations apparently trace back to only a very small number of<br />

genes that function as the <strong>de</strong>velopmental control genes (Kingsley, 2009).<br />

The presence or absence of the ventral spine in the Stickleback essentially traces<br />

back to the expression of only a single gene. This gene is named Pitxl and is (as<br />

usual) controlled by various enhancers. One particular enhancer is responsible for<br />

the expression of the Pitxl gene in the ventral fin, the ventral fin enhancer. In other<br />

body parts, Pitxl co<strong>de</strong>s for the same protein, but it affects the <strong>de</strong>velopment of other<br />

body structures. Failure of the ventral fin enhancer only causes the failure to express<br />

the gene in the ventral fin area and thus specifically affects only <strong>de</strong>velopment of the<br />

one spiny ventral fin. The other enhancers remain intact and continue function.<br />

The Stickleback is an exceedingly instructive example for typologically oriented<br />

taxonomists. It shows that the reshaping of entire body structures can trace back to a<br />

single mutation.<br />

4.5<br />

What is the Relevance of Differences in Traits between Two Species?<br />

Two traits can appear different or perform different functions but nevertheless be<br />

enco<strong>de</strong>d by the same gene. What then are two different traits if they are based on the<br />

same gene? If the structural genes hardly differ between two species, what is it that<br />

differs if two species have different traits? If one species possesses a certain trait and<br />

another does not, we assume intuitively that the first species has acquired this trait at<br />

a later time. However, traits can also be lost. The control of the structural genes via<br />

enhancer switches makes apparent that anatomical and morphological alterations<br />

during evolution can be based on the repeated switching on and off of structural<br />

genes that otherwise do not change during the speciation processes.<br />

What is a newly evolved trait if its coding gene is not new? This consi<strong>de</strong>ration<br />

questions the concept of Willi Hennig s apomorphy (Hennig, 1966) (Chapter 7).<br />

Hennig s cladistic taxonomy <strong>de</strong>fines the origin of two daughter species from an<br />

ancestor species (stem species) by the fact the two new daughter species must possess

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