An earlier version of this collection of essays appeared as a special issue of the J. of social and biological structures (Academic Press) in July 1989. Annotation copyright Book News, Inc. Portland, Or.
The editors explain in the introductory chapter of this 1989 collection, “How does change occur in the biological world? Does it happen gradually, as classical evolutionary theory has long assumed, or does it occur by rapid transformations of structure and behavior?... These are the questions posed by a ‘new’ conceptualization of the evolutionary process… common referred to as punctuated equilibrium. And these are the questions this book addresses. Punctuated equilibrium theory makes two contentions: that evolutionary change… occurs in rapid bursts over (geologically) short periods of time, and that there is relative stasis after the punctuational burst… [Niles] Eldredge and Stephen Jay Gould in 1972 unleashed a controversy that still roils the scientific journals and has divided evolutionary theorists into often angrily competing camps. It seems only sensible… to attempt an assessment of its implications for the neo-Darwinian synthesis. In structuring this volume, we had three goals. First, we wanted to identify the component elements of punctuated equilibrium… Our second objective was to assess the present status of punctuated equilibrium among evolutionary theorists… Third, we sought to weigh the scientific implications of punctuated equilibrium.” (Pg. 1-2)
Ernst Mayr noted in his essay, “I believe I was the first author to develop a detailed model of the connection between speciation, evolutionary rates, and macroevolution… my new theory of the importance of peripatric speciation in macroevolution is now widely recognized… it is most curious that the theory was completely ignored by paleontologists until brought to light by Eldredge and Gould… The major novelty of my theory was its claim that the most rapid evolutionary change does not occur in widespread, populous species… but in small founder populations. This conclusion was based on empirical observations… I had found again and again that the most aberrant population of a species… occurred at… the most isolated peripheral location… such a population would have unique opportunities to enter new niches and to select novel adaptive pathways.” (Pg. 25)
He continues, “In refutation of [Richard] Goldschmidt’s claims I demonstrated… that geographic variation in isolated populations could indeed account for evolutionary innovations. Such populations have a very different evolutionary potential than contiguously distributed, clinically varying populations in a continental species… one can defend a moderate form of punctuationism… without having to adopt Goldschmidt’s theory of system mutations.” (Pg. 30)
Gould explains, “Niles Eldredge and I were graduate students in paleontology together… we were primarily interested in evolution… We began our professional lives with a commitment to engage in the empirical study of evolution as illustrated by the fossil record… Now imagine the frustration of two … ambitious young men captivated with evolution… and faced with the following situation. The traditional wisdom of the profession held … that the fossil record … had long been restrictively defined as ‘insensibly graded fossil sequences’---and such hardly existed… In other words, we were told that our primary data base contained virtually no examples of the phenomenon we wished to study… The record, we were told, was so imperfect that truly insensible fossil transitions left no trace in the rocks… Niles and I had one advantage … We had been well trained in the details of modern evolutionary theory. (I also had a fascination for history and philosophy of science. I had been strongly influenced by the then radical view of [Thomas] S. Kuhn, N.R. Hanson, and [Paul] Feyerabend on the hold that theory imposes upon perception… Eventually we (primarily Niles) recognized that the standard theory of speciation… would not, in fact, yield insensibly graded fossil sequences when extrapolated into geological time, but would produce just what we see: geologically unresolvable appearance followed by stasis. For if species almost always arise in small populations isolated at the periphery … and in a period of time… effectively instantaneous in the geological world…then the workings of speciation should be recorded in the fossil record as stasis and abrupt appearance.” (Pg. 55-56)
He goes on, “I continue to have one cardinal difficulty with the attribution of stasis to stabilizing selection---the scales are all wrong… I fully admit that my alternative also relies on gut feelings and, in part, on negative evidence---but I find the logic compelling enough to win a hearing for this view… the profoundness of temporal depth of stasis are trying to tell us that change is ACTIVELY PREVENTED, rather than … merely suppressed because n adaptive advantage would accrue… stasis is primarily an active feature of organisms and populations, maintained by evolved genetic and developmental coherences…” (Pg. 64)
He clarifies, “colleagues have often criticized punctuated equilibrium … Most egregious in this regard has been the long-standing and lamentable confusion of punctuated equilibrium with Richard Goldschmidt’s saltationism---though we have continually, painstakingly, and clearly distinguished the two… I do take an interest in Goldschmidt’s ideas… if only to redress a balance upset by previous derision among the orthodox… My interest in Goldschmidt springs largely from another source in my career—that leading to ‘Ontogeny and Phylogeny; (1977)---and I do assert an intellectual’s birthright to multiple subjects of concern!” (Pg. 70)
Steven Stanley notes, “Speciation by way of very small populations may be more common than has generally been believed. Not to be overlooked here is the new evidence from studies of mitochondrial DNA that all modern humans have descended from a single female.” (Pg. 97)
Niles Eldredge observes, “The notion of punctuated equilibria arose in an attempt to bring evolutionary theory more closely in line with general patterns of evolutionary history that are manifest in the fossil record… evolutionary biology since Darwin’s time has unquestionably seen evolution as a matter of slow, steady, and progressive transformation of the adaptive features of organisms. Punctuated equilibria… sought an explanation for the overlooked phenomenon of marked stability… in which adaptive evolutionary change seems to be concentrated in (relatively) brief episodes, ‘punctuating’ vastly longer intervals when little or no change occurs.” (Pg. 105)
Antoni Hoffman, however, argues, “The concept of macromutations are the main mechanism of speciation appears today implausible, to say the least. Although there is no logical way to disprove such a mechanism’s potential operation in nature, the genetic data suggest that point mutations with large phenotypic effects generally are adaptively disadvantageous, and whenever they are not, they may not bring about speciation… The strong version of punctuated equilibrium and its macromutationary implication are thus indefensible; in fact, no one seems willing to advocate it any longer.” (Pg. 128)
Susan Cachell points out, “I also note that mosaic evolution is a major problem for adherents of punctuated equilibria. If different morphological or behavioral characters evolve at different rates in the same organism, that organism will be a mosaic of characters, some primitive and some derived. This concatenation of traits … represents the operation of mosaic evolution. Hence there is no coherent transformation of all or most characters in evolution. Traits may vary in rate of evolution even within the same functional system… or one functional system, made up of an intricate complex of traits.” (Pg. 196-197)
This book will be “must reading” for anyone interested in punctuated equilibrium, or current evolutionary theory.
Punctuated equilibrium is a theory of evolutionary change originally championed by Niles Eldredge and Stephen Jay Gould. It posits stasis in evolutionary change over long periods of time--punctuated by a rapid (in evolutionary terms) change. This volume features essays by key figures in the debate over the validity of this perspective--including by Eldredge and Gould themselves. There is also a magisterial essay by one of the deans of neo-Darwinian theory--Ernst Mayr. There is a second part to the book, where implications of punctuated equilibrium theory for the social sciences is discussed.
I co-edited this volume; I'm pleased to say that many of the major figures in the debate over punctuated equilibrium theory consented to write chapters for this book.