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Principios de Taxonomia

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7.9 The Concepts of Monophyly and Paraphyly cannot be Applied to Speciesj207<br />

foun<strong>de</strong>r of cladistic taxonomy, avoi<strong>de</strong>d applying the cladistic unit of a monophylum to<br />

the category of species and restricted the application of the monophyly concept to<br />

higher taxa (Hull, 1997; Ereshefsky, 1999). In contrast, Hennig s successors have<br />

exten<strong>de</strong>d the concept of monophyly to the level of species (Ax, 1995; <strong>de</strong> Queiroz, 1999;<br />

<strong>de</strong> Queiroz and Donoghue 1988; Mishler and Donoghue, 1994).<br />

The problem in this case is that the concepts of monophyly and paraphyly are tied<br />

to bifurcations (Figure 7.5). Without bifurcations, there is no mono- and no paraphyly.<br />

However, biparental species as a gene-flow community do not represent a<br />

bifurcating system. Within a unit in which reproductive connections between<br />

organisms exist, there can be no bifurcations. A bifurcation within a species always<br />

splits the species into two separate gene flow communities, thus representing the<br />

origin of two new species.<br />

Accordingly, the terms monophyly and paraphyly cannot be applied to biparental<br />

species because there is no bifurcation within such species. Mono- or paraphyla must<br />

be groups that inclu<strong>de</strong> distinct branches. The logic of the paraphylum always requires<br />

at least two bifurcations, which lead to three branches with dissimilar kinship<br />

relationships that would then be expressed in the paraphylum (Figure 7.5).<br />

Two branches alone cannot form a paraphylum; and a gene-flow community that<br />

always is a group of organisms without any separated branches, never can be a monoor<br />

paraphylum. Bifurcations only exist at higher levels of taxonomic hierarchies<br />

above the species level.<br />

This is explained by the following example (Figure 7.11). On the Canary Islands of<br />

Tenerife and Gran Canaria, there is an en<strong>de</strong>mic species of finch that only occurs on<br />

these two islands, the Blue Chaffinch (Fringilla tey<strong>de</strong>a). The Blue Chaffinch is a sister<br />

species of the Common Chaffinch (Fringilla coelebs). It is thought that the Blue<br />

Chaffinch separated from the continental Common Chaffinch only a relatively short<br />

time ago evolutionarily.<br />

Let us assume that the stem species of the contemporary Common Chaffinch<br />

and Blue Chaffinch inhabited a wi<strong>de</strong> geographical range from northwest Africa<br />

through Europe to West Asia, similar to the distribution of the contemporary<br />

Common Chaffinch. Let us also assume that during that time, different races<br />

arose that remained connected to each other via continuing gene flow (open<br />

circles within the stem species in Figure 7.11). When population e then reached<br />

the Canary Islands, the gene flow cohesion with the continental populations<br />

ceased due to allopatry.<br />

Two sister species arose: the recent Common Chaffinch and the Blue Chaffinch.<br />

It does not make sense to view the Common Chaffinch as a paraphylum, as its current<br />

races, a through d, are connected by gene flow and, thus, belong to a single gene-flow<br />

community. Gene flow cohesion exclu<strong>de</strong>s the possibility of a more recent common<br />

ancestor or a less recent common ancestor of populations a through e because all<br />

populations are connected and do not consist of a system of separated branches.<br />

Therefore, the Common Chaffinch cannot be a paraphyletic species. The concepts of<br />

monophyly and paraphyly cannot be applied to species as long as species are<br />

un<strong>de</strong>rstood as gene-flow communities.

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