Microevolution and Macroevolution

Author:

Ayala Francisco J.

Publisher

Elsevier

Reference15 articles.

1. Gould's language is combative in the extreme. He speaks of a “preemptive strike” (p. 31) against his enemies, their “destruction” (p. 33), their “jealousy” (p. 1021), and how ultimately “we won” (p. 1022). Depressingly frequent throughout the book are such words as “battle,” “conflict,” “retreat,” “victory,” and the like.

2. Gould's rich language and elegant metaphors are marred by redundancy, long elaboration, and repetition. The metaphor of a tree with three branches is redundant with that of a coral with three branches, reproduced from a 1670 engraving, and with that of a tripod supported by its three legs. These three metaphors are repeated and elaborated at length. The 1670 coral engraving is reproduced at nearly full-page size twice, identically on pages 18 and 97. Milan's Duomo, introduced as an architectural structure that acquired ornamental and other features centuries after it was built, serves as a metaphor for later elaborations of the fundamental Darwinian logic. The first time this metaphor appears is belabored over five pages (pp. 2–6) and illustrated with two photographs of the cathedral. These two photos, the duplicated coral engraving, plus another 1670 engraving representing two human figures with shells, are the only illustrations for the first 182 pages of the book. The following two paragraphs may serve as examples of Gould's literary style, eloquence, and prolixity: “The specific form of the image—its central metaphorical content, if you will—plays an important role in channeling or misdirecting our thoughts, and therefore also requires careful consideration. In the text of this book, I speak most often of a ‘tripod’ since central Darwinian logic embodies three major propositions that I have always visualized as supports—perhaps because I have never been utterly confident about this entire project, and I needed some pictorial encouragement to keep me going for 20years. (And I much prefer tripods, which can hold up elegant objects, to buttresses, which may fly as they preserve great Gothic buildings, but which more often shore up crumbling edifices. Moreover, the image of a tripod suits my major claim particularly well—for I have argued, just above, that we should define the ‘essence’ of a theory by an absolutely minimal set of truly necessary propositions. No structure, either of human building or of abstract form, captures this principle better than a tripod, based on its absolute minimum of three points for fully stable support in the dimensional world of our physical experience.)” (p. 15).“Galton's Polyhedron, the metaphor and model devised by Darwin's brilliant and eccentric cousin Francis Galton, and then fruitfully used by many evolutionary critics of Darwinism, including St. George Mivart, W.K. Brooks, Hugo de Vries, and Richard Goldschmidt, clearly expresses the two great, and both logically and historically conjoined, themes of formalist (or structuralist, or internalist, in other terminologies) challenges to functionalist (or adaptationist, or externalist) theories in the Darwinian tradition. This model of evolution by facet-flipping to limited possibilities of adjacent planes in inherited structure stresses the two themes—channels set by internal constraint, and evolutionary transition by discontinuous saltation—that structuralist alternatives tend to embrace and that pure Darwinism must combat as challenges to basic components of its essential logic (for channels direct the pathways of evolutionary change from the inside, albeit in potentially positive and adaptive ways, even though some external force, like natural selection, may be required as an initiating impulse; whereas saltational change violates the Darwinian requirement for selection's creativity by vesting the scope and direction of change in the nature and magnitude of internal jumps, and not in sequences of adaptive accumulation mediated by natural selection at each step” (p. 66).

3. A religious architecture metaphor for replacement would have been the splendid Gothic cathedral of Leon, Spain. The 14th-century wealth and exuberant ambition of the citizens of the then capital of Castile moved them to demolish the preexisting 12th-century Romanesque cathedral and build a much larger and taller Gothic cathedral on the same location. This replacement of Romanesque by Gothic cathedrals was common through 14th- and 15th-century Christendom. (How much wiser were the citizens of Salamanca in Spain, who left the Romanesque cathedral standing and built a new Gothic cathedral next to it—which, admirable architectural masterpiece that it is, is nevertheless aesthetically surpassed in my judgment by the earlier Romanesque monument!)

4. Eldredge, N., 1971. The allopatric model and phylogeny in Paleozoic invertebrates. Evolution 25, 156–167; Eldredge, N., Gould, S.J. 1972. Punctuated equilibria: an alternative to phyletic gradualism. In: Schopf, T.J.M. (Ed.), Models in Paleobiology. Freeman, Cooper, Co., pp. 82–115.

5. Gould refers to me, with kind words, as supporting this claim of macroevolutionary autonomy and quotes me at length: “I have particularly appreciated the fairness of severe critics who generally oppose punctuated equilibrium, but who freely acknowledge its legitimacy as a potentially important proposition with interesting implications, and as a testable notion that must be adjudicated in its own macroevolutionary realm. Ayala (1983) has been especially clear and gracious on this point.” “If macroevolutionary theory were deducible from microevolutionary principles, it would be possible to decide between competing macroevolutionary models simply by examining the logical implications of microevolutionary theory. But the theory of population genetics is compatible with both punctualism and gradualism; and, hence, logically it entails neither. Whether the tempo and mode of evolution occur predominantly according to the model of punctuated equilibria or according to the model of phyletic gradualism is an issue to be decided by studying macroevolutionary patterns, not by inference from microevolutionary processes. In other words, macroevolutionary theories are not reducible (at least at the present state of knowledge) to microevolution. Hence, macroevolution and microevolution are decoupled in the sense (which is epistemologically most important) that macroevolution is an autonomous field of study that must develop and test its own theories” (Ayala, 1983, as quoted by Gould, 2002, p. 1023).To avoid misunderstanding, I summarize here issues that I have discussed at some length in the paper cited by Gould (p. 1023) and elsewhere. Macroevolution and microevolution are not decoupled in two senses: identity at the level of events and compatibility of theories. First, the populations in which macroevolutionary patterns are observed are the same populations that evolve at the microevolutionary level. Second, macroevolutionary phenomena can be accounted for as the result of known microevolutionary processes. That is, the theory of PE (as well as the theory of phyletic gradualism) is consistent with the theory of population genetics. Indeed, any theory of macroevolution that is correct must be compatible with the theory of population genetics, to the extent that this is a well-established theory. The decoupling asserted by me in the quotation cited above by Gould, as well as later in his book, concerns epistemology: the logical autonomy of theories.

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