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Ornithology, Evolution, and Philosophy 123

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218 5 Biological Species <strong>and</strong> Speciation—Mayr’s First Synthesis<br />

Fig.5.5. Stages of geographical (allopatric) speciation (from Mayr 1942e, Fig. 16)<br />

found which were difficult to classify (see Table 4.2 on p. 177). However, such “intermediate”<br />

forms illustrated for Mayr the process of speciation graphically. Strictly<br />

speaking it is not the subspecies but the geographically isolated population which<br />

may reach species status. However, little was known about the genetic basis of the<br />

speciation process itself which both Dobzhansky (1937) <strong>and</strong> Mayr (1942e) felt certain<br />

occurred as a continuation of microevolution. In a superspecies the member<br />

species are not ecologically compatible, for which reason the respective species<br />

populations, if their ranges abut, compete along the contact zones <strong>and</strong> exclude<br />

each other geographically without (or only rarely) hybridizing. The closely related<br />

members of a species group may be sympatric in parts of their ranges. In ring<br />

species the overlapping populations are so strongly differentiated that they no<br />

longer hybridize <strong>and</strong> would be considered species if their direct intergradation<br />

through the “ring” of subspecies were to be interrupted (an example are the Asian<br />

greenish warblers Phylloscopus trochiloides, Irwin et al. 2005). According to the<br />

“Wallace-Dobzhansky model” isolating mechanisms of neospecies originate due<br />

to selection forces in sympatry, whereas according to the “Darwin-Muller-Mayr<br />

model” isolating mechanisms arise as incidental, pleiotropic by-products of divergence<br />

in allopatry (but see also under sympatric speciation, p. 223). David Lack<br />

(1944, 1949, 1971) added important data on ecological aspects of the speciation<br />

process, as acknowledged by Mayr (1982d: 274).<br />

Besides the “dumbbell” model of geographic speciation, Mayr (1942e) developed<br />

an alternative speciation model: a new species originates from jump dispersal<br />

of a few individuals forming a small strongly isolated peripheral founder population<br />

(founder principle, 1942e: 237; 1954c; 1963b: 539, 554; peripatric speciation,<br />

1982l). This principle relates the reduced variability of small founder populations<br />

not to accidental gene loss, but to the fact that the entire population was started<br />

by a single pair, a few individuals or even by one fertilized female which contained<br />

only a fraction of the gene pool of the parent population. These “founders” carried

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