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Ornithology, Evolution, and Philosophy 123

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A Modern Unified Theory of <strong>Evolution</strong> 225<br />

(not published until 1941k) <strong>and</strong>, second, at a symposium on speciation organized<br />

by Dobzhansky (Mayr 1940c).<br />

Mayr (1954e) asked the question whether speciation in marine organisms differed<br />

from speciation in l<strong>and</strong> animals (birds, mammals, <strong>and</strong> butterflies). At the<br />

time he was working with West Indian Cerion snails (see p. 331) <strong>and</strong> therefore<br />

selected the West Indian shallow water echinoids for study. Each genus consists of<br />

one or several groups of allopatric species, some of which should be considered as<br />

subspecies. Allopatric populations show every grade of distinctness, ranging from<br />

slight difference through subspecific to specific rank. This pattern corresponds to<br />

that of l<strong>and</strong> animals. This is also suggested for deep sea organisms (crinoids) <strong>and</strong><br />

pelagic organisms (Scyphomedusae). Dispersal barriers which caused speciation<br />

in warm water echinoids are the eastern Pacific, the Isthmus of Panama <strong>and</strong> the<br />

cold waters of southwestern Africa. There was no evidence suggesting any other<br />

mode of speciation in sexually reproducing marine organisms than geographic<br />

speciation. Palumbi <strong>and</strong> Lessios (2005) largely confirmed Mayr’s interpretation of<br />

geographic speciation in sea-urchins using DNA sequence data. They also showed<br />

that the origin of reproductive isolation depends not only on genome-wide steady<br />

accumulation of substitutions but also on the rate of evolution of gamete recognition<br />

proteins.<br />

Sibling species were found among the colorful birds of southeastern Asia (p. 161)<br />

<strong>and</strong> the polymorphic Californian king snake proved to be one biological species<br />

only (p. 226). Particularly interesting cases of geographical speciation in birds<br />

include the Black Bulbul (Hypsipetes madagascariensis; p. 160), trillers (Lalage),<br />

fantails (Rhipidura), Reef Heron (Demigretta sacra), drongos (Dicruridae, p. 158),<br />

Halcyon kingfishers, Australian tree runners (Neositta; p. 179), wheatears (Oenanthe,<br />

p. 226), paradise magpies (Astrapia, p. 155), <strong>and</strong> honeyeaters (Melidectes).<br />

When experimentation with birds turned out as logistically too difficult, Mayr<br />

studied isolating mechanisms in Drosophila (p. 228). He also applied the principles<br />

of new systematics to paleanthropology (p. 330). Numerous additional examples of<br />

geographical variation <strong>and</strong> speciation from peripheral isolates may be found in his<br />

major books on species <strong>and</strong> evolution (1942e, 1963b, Mayr <strong>and</strong> Diamond 2001g).<br />

The number of generic <strong>and</strong> family revisions, faunistic papers or check-list entries<br />

requiring decisions on the species status of geographically isolated populations<br />

totals about 60.<br />

Mayr (1946a) estimated the number of bird species in the world at 8,616 species<br />

<strong>and</strong> Bock <strong>and</strong> Farr<strong>and</strong> (1980) at 9,021. This increase in the number of species<br />

taxa was caused in part by the discovery of genuine new species <strong>and</strong> in part<br />

by reinterpreting the rank of numerous allopatric taxa as species rather than<br />

subspecies. In a study of a local flora, Mayr (1992a) tested the validity of the<br />

biological species concept for plants concluding that the great majority of taxa<br />

(93.5%) form biological species; only 6.44% pose difficulties by apomixis <strong>and</strong> by<br />

autopolyploidy; Whittemore (1993) published a critical discussion of these results<br />

from a botanical point of view.

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