In The Origin of Species (1859) Darwin challenged many of the most deeply-held beliefs of the Western world. Arguing for a material, not divine, origin of species, he showed that new species are achieved by "natural selection." The Origin communicates the enthusiasm of original thinking in an open, descriptive style, and Darwin's emphasis on the value of diversity speaks more strongly now than ever. As well as a stimulating introduction and detailed notes, this edition offers a register of the many (...) writers referred to by Darwin in the text. (shrink)

Interspecific comparisons reveal remarkable diversity in patterns of gene expression, even among closely related species. Combinatorial regulatory mechanisms could facilitate the evolution of this diversity. However, the high degree of interdependence characteristic of combinatorial networks would represent a major constraint on evolution and might generate many features that have no direct adaptive value.

In 1979, Lewontin and I borrowed the archi- tectural term “spandrel” (using the pendentives of San Marco in Venice as an example) to designate the class of forms and spaces that arise as necessary byproducts of another decision in design, and not as adaptations for direct utility in them- selves. This proposal has generated a large literature featur- ing two critiques: (i) the terminological claim that the span- drels of San Marco are not true spandrels at all and (ii) the (...) conceptual claim that they are adaptations and not byprod- ucts. The features of the San Marco pendentives that we explicitly defined as spandrel-properties—their necessary number (four) and shape (roughly triangular)—are inevitable architectural byproducts, whatever the structural attributes of the pendentives themselves. The term spandrel may be extended from its particular architectural use for two- dimensional byproducts to the generality of “spaces left over,” a definition that properly includes the San Marco pendentives. Evolutionary biology needs such an explicit term for features arising as byproducts, rather than adaptations, whatever their subsequent exaptive utility. The concept of biological span- drels—including the examples here given of masculinized genitalia in female hyenas, exaptive use of an umbilicus as a brooding chamber by snails, the shoulder hump of the giant Irish deer, and several key features of human mentality— anchors the critique of overreliance upon adaptive scenarios in evolutionary explanation. Causes of historical origin must always be separated from current utilities; their conf lation has seriously hampered the evolutionary analysis of form in the history of life. (shrink)