Structural Constraints, Spandrels, and Exaptation 1183
education—at the ceremony for his receipt of an honorary degree as Doctor in Civil
Law). In 1969, the Whole Earth Catalog, the commercial bible of the "green"
movement in America, called his major work "a paradigm classic." Few people can
list such diverse distinctions in their compendium of honors. But then, few people
have displayed so wide a range of talent. D'Arcy Wentworth Thompson (1860-1948),
Professor of Natural History at the Scottish universities of Dundee and St. Andrews,
translated Aristotle's Historia animalium, wrote glossaries of Greek birds and fishes,
compiled statistics for the Fishery Board of Scotland and contributed the article on
pycnogonids to the Cambridge Natural History.
But D'Arcy Thompson's current reputation rests almost entirely upon a book of a
thousand pages, revered by artists and architects as well as by engineers and
biologists—the "paradigm classic," On Growth and Form (1917, 2nd edition, 1942).
P. B. Medawar (1967, p. 232) lauded this volume as "beyond comparison the finest
work of literature in all the annals of science that have been recorded in the English
tongue." G. Evelyn Hutchinson (1948, p. 579) regarded Growth and Form as "one of
the very few books on a scientific matter written in this century which will, one may
be confident, last as long as our too fragile culture."
Although I have studied D'Arcy Thompson's wonderful book throughout my
career (see Gould, 1971b, for my first, and in retrospect embarrassingly puerile,
publication in a journal of the humanities), I originally made a major error in siting
him within the history of biology. All intellectuals love a courageous loner, and I had
been beguiled by D'Arcy Thompson's seemingly anachronistic peculiarities—his
flowery, sometimes overblown, but often soaring and powerful, Victorian prose; his
expertise at fully professional levels in Latin and Greek; even his lifelong residence
in an outlying region that, in my false mental geography, might well have been
located above the Arctic Circle. In retrospect, I had unthinkingly conflated my sense
of his intellectual distance from conventional thought with an assumption about
physical isolation as well. When I finally visited the University of St. Andrews (for
the humbling experience of receiving an honorary degree in D'Arcy Thompson's own
bailiwick), I recognized its proximity to Edinburgh, and its easy access by rail. (As a
further confirmation of St. Andrews's central location within the contemporary world,
I began to write this section on the very day that Tiger Woods won the British Open
on the world's original, and still most famous, golf course of St. Andrews.)
I had therefore viewed D'Arcy Thompson as the ultimate man out of time—a
Greek geometer and classical scholar, a Victorian prose stylist at the dawn of
modernism's lean and cynical attitude (for the first edition of Growth and Form
appeared in 1917 in the midst of Word War I, while another and even more
destructive war greeted the second edition of 1942; many historians have noted that,
in a meaningfully ideological, rather than an arbitrarily calendrical, reckoning, the
20th century really begins with World War I and the end of illusions about progress
and the benevolent hegemony of European control).
I shall not try to rob D'Arcy Thompson of his genuine singularities, but