Few of Stephen Jay Gould’s accomplishments in evolutionary biology have received more attention than his hierarchical theory of evolution, which postulates a causal discontinuity between micro- and macroevolutionary events. But Gould’s hierarchical theory was his second attempt to supply a theoretical framework for macroevolutionary studies—and one he did not inaugurate until the mid-1970s. In this paper, I examine Gould’s first attempt: a proposed fusion of theoretical morphology, multivariate biometry and the experimental study of adaptation in fossils. This early “macroevolutionary synthesis” was predicated on the notion that parallelism and convergence dominate the history of higher taxa, and moreover, that they can be explained in terms of adaptation leading to mechanical improvement. In this paper, I explore the origins and contents of Gould’s first macroevolutionary synthesis, as well as the reasons for its downfall. In addition, I consider how various developments during the mid-1970s led Gould to identify hierarchy and constraint as the leading themes of macroevolutionary studies—and adaptation as a macroevolutionary red herring.
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Parallelism and convergence are processes resulting in the evolution of similar traits or character states in independent lineages. In the case of parallel evolution, the lineages under consideration must possess a recent common ancestor—for instance, they may be species in a genus, or (according to an older conception) genera within a family. By contrast, convergent evolution occurs when the lineages in question do not share a recent common ancestor.
Norman Newell was a celebrated invertebrate paleontologist and pioneer of the quantitative study of mass extinctions, who spent the majority of his career at Columbia University and the affiliated American Museum of Natural History (see Sepkoski 2012, Ch. 2). Along with John Imbrie (also an invertebrate paleontologist at the AMNH), Newell was an important influence on Gould’s early career, in particular, his decision to apply quantitative methods to problems in invertebrate paleontology (see Princehouse 2009; Eldredge 2013).
Gould also published a single study in theoretical morphology, a simulation of the spiral morphology of a group of Ordovician organisms known as receptaculids (Gould and Katz 1975).
To use the terminology of Gould (1971b), Raup’s studies of coiled shell morphology are primarily concerned with the “how” of form: that is, with reducing complex morphologies to a small set of factors that can generate them during growth. However, as Gould (1970a, p. 92) observes, “Raup has extended [his] methods far beyond the simple insight that a complicated form can be produced by a few simple instructions,” and is actively investigating the “why” of form as well (see especially Raup 1967, pp. 54–65). And it is the latter issue, “the explanation of form in terms of adaptation,” which constitutes “the fundamental problem of evolutionary paleontology” (Gould 1967, p. 385).
See, for instance, Gould (1966b), (1967), (1968), (1969), (1970c), (1972), (1973a), (1973b), (1974), (1975), Gould and Garwood (1969), Gould and Johnson (1972), Pilbeam and Gould (1974). Several of these projects are discussed in Roger D.K. Thomas’s essay: “Gould’s odyssey” (2009), which also considers Gould’s morphological researches after 1975.
As I mentioned above, it was John Imbrie who is most responsible for Gould pursuit of multivariate analyses of form and geographical variation during his early career (Eldredge 2013, p. 8). Recruited to the AMNH by Newell, Imbrie was the foremost paleontological expert on multivariate statistics during Gould’s graduate years, and a keen advocate of factor analysis. For an excellent account of Imbrie’s “statistical paleontology,” see Sepkoski (2012) (pp. 83–91).
These individuals commanded the most space in the text, and earned the greatest number of citations. Equal in citation number to Seilacher, but featured less prominently in the text, are the vertebrate paleontologists George Simpson and Björn Kurtén.
Since Gould (1970a) defines “history” as “directional change through time” (p. 108), and since (on the very next page) he equates “history” with “biological progress” (p. 109), “change without history” means something like “directionless change,” or change unaccompanied by mechanical improvement.
George Simpson made this very claim in a 1961 review paper, stating: “The study of fossil vertebrates elucidates the general principles of evolutionary biology” (p. 1679, emphasis added). This statement seems innocent enough, until one considers the state of invertebrate paleontology during the first half of the twentieth century (see Rainger 2001). “[It] is almost as if invertebrate paleontology [is] in bondage to geology,” the gastropod specialist J. Brookes Knight complained in a presidential address to The Paleontological Society (Knight 1947, p. 282). The average invertebrate paleontologist is “not a paleontologist at all…He is…a stratigraphical or ‘soft rock’ geologist,” and therefore ill-equipped to investigate evolutionary problems. Edwin Colbert, a vertebrate paleontologist, carried this criticism even further: “[It] is not only a question of lack of interest in the subject, for it is a fact that most of our contemporary geologists are not even competent to take more than a superficial interest in evolutionary problems” (Colbert 1947, p. 289). To be sure, the situation had improved somewhat by 1963, when Gould entered graduate school. But it was far from rectified, even in the elite centers of paleontological training (see Sepkoski 2012, pp. 55ff; Eldredge 2013, p. 6).
Bobb Schaeffer was a longtime curator of fossil fishes at the American Museum of Natural History, and Gould’s teacher at Columbia.
By “biological improvement,” Gould means mechanical improvement of the engineering type that occurs in a constant physical environment.
As Gould observes in “Evolutionary paleontology,” natural experiments afford an opportunity to study adaptation in situ without limiting ourselves to “modern manipulation” (Gould 1970a, p. 89). (Natural experiments are empirical studies in which the experimental and control conditions are determined by nature, not by human intervention.) Here he cites the work of Adolph Seilacher, who examined patterns of boring on fossilized belemnite shells, which he assumed to be “adjusted to the normal, head-on movement of the belemnite” (Seilacher 1968, p. 279). This led Seilacher to conclude that “the streamlining of the rostrum is related only to occasional backward escapes ([an] ‘emergency adaptation’).” Gould’s 1970 study of parallel evolution in Bermudian microgastropods also utilized a natural experiment, which leveraged historical fluctuations in the availability of calcium carbonate (an important mineral in gastropod shell construction) to test a number of adaptive hypotheses relating to shell morphology (Gould 1970c).
Problems of Relative Growth is dedicated to D’Arcy Thompson, with whom Huxley maintained an active correspondence. Interestingly, the long epigram that follows Huxley’s dedication—a Thompsonian meditation on the correlation of parts—is quoted at length in Gould’s most influential paper, “Punctuated equilibria: an alternative to phyletic gradualism.” .
Here it bears mentioning that grades, in Huxley’s view, are stable units of anagenetic advance (Huxley 1957, p. 454). Grades are therefore attained only when anagenesis is arrested; that is, when progress is consolidated by “stasigenetic,” or stability inducing, evolutionary processes.
Notice that while this process will tend to produce monophyletic grades “near the level of the taxon actually arising” (e.g., orders, classes), it should not be assumed that these would qualify as monophyletic at lower taxonomic levels (e.g., species, genera). While anagenesis followed by cladogenesis sometimes produces monophyletic grades at the species level, this outcome requires that no other lineages independently achieve the same organizational level. And this Huxley and others presumed to be rare (see, e.g., Simpson 1953, p. 348; Mayr 1963, p. 609).
Neoteny refers to the slowing of development, leading to the sexual maturity of an animal while it is still in a juvenile (or even larval) state.
A traditional assessment of the evolutionary significance of neoteny appealed to the ability of neotenic species to escape from ecological specialization—the putative enemy of diversification.
Notice that the theory of life history tactics does not demand that adaptations constitute “superior a priori designs for living in [an] environment,” at least if superiority is cashed out in engineering terms. Indeed, certain changes may confer fitness by decreasing the mechanical efficiency of a structure, like the loss of eyes in cavefishes.
Elsewhere, Gould seems to back off this statement, suggesting that size-imposed characters “merely [provide] the same efficiency for a primary adaptation of altered size,” and therefore do not constitute biological improvements (Gould 1970a, p. 110). Nonetheless, their importance for transspecific evolution is manifest: “the expanded potential for further progress…conferred upon organisms bearing [size-required adaptations] is a true and most significant evolutionary advance” (Gould 1966a, p. 591). If size-required adaptations do not constitute biological improvements, yet they set the stage for potential biological improvements in the future.
I owe this insight to Alan Love.
While Gould made superficially similar claims in Ontogeny and Phylogeny, his emphasis fell on the capacity of natural selection (acting on the life history parameter of “difficult transitions” between “fundamentally different designs in the origin of taxa,” not on positive channeling (Gould 1977a, p. 338). This phrasing betrays a concern for the original problematic of macroevolutionary studies: the origin of higher taxa or organizational types (e.g., Schaeffer 1965). By contrast, Gould’s later interest in the ability of constraints to “impart a preferred direction to evolutionary change not based on natural selection” is responsive to a new problematic: the causation of statistical trends within large taxa (e.g., Stanley 1975, p. 648; Gould 1982a, p. 385).
Sepkoski’s Rereading the Fossil Record contains an excellent account of this period in the history of paleobiology (see especially chapter 5, on the origins of “Punctuated equilibria,” and chapter 7, on the introduction of stochastic models). Further useful information can be found in chapters 11 and 16 of The Paleobiological Revolution (written by Todd Grantham and John Huss, respectively), and in Raup’s synopsis of the MBL project (Raup 1977).
In 1974, Gould collaborated with David Raup on a series of stochastic simulations of morphological evolution—a project with distinct bearings on the problem of the causation of phyletic trends (Raup and Gould 1974). The results suggested that “trends in morphology” (“[even those] of outstanding duration and unreversed direction”) could occur in the absence of deterministic causes like directional natural selection (Raup and Gould 1974, p. 314). While Gould was wary of assigning too great a significance to these results (ibid, 321), yet he later enlisted them in his campaign for a hierarchically expanded evolutionary theory (e.g., Gould 2002, p. 741).
In their celebrated discussion of trends, Eldredge and Gould (1972, pp. 108–112) failed to consider that the differential success of species may owe to irreducible species-level properties, although they did recognize that certain species will outlast others in virtue of the superior adaptedness of their members. Years later, Gould would admit that this was indeed a major limitation of their earlier discussion (Gould 1982a, p. 101, 2002, p. 731), and one he did not fully appreciate until papers by Stanley (1975) and Gilinsky (1981).
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I would like to thank Alan Love, Emilie Snell-Rood, Mark Borrello, Ruth Shaw and Staffan Müller-Wille for their keen editorial insights during the writing process. Dr. Love read the manuscript several times and provided invaluable feedback on its organization and scope. In addition, I would like to thank Niles Eldredge, Roger D.K. Thomas and Richard Lewontin for their generous correspondence during various stages of this project. Last but not least, I owe a debt of gratitude to the participants of the 2015 MBL-ASU History of Biology Seminar (“Perspectives on Stephen Jay Gould”), and especially to the seminar organizers, John Beatty and David Sepkoski, for inviting me to participate.
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Dresow, M.W. Before hierarchy: the rise and fall of Stephen Jay Gould’s first macroevolutionary synthesis. HPLS 39, 6 (2017). https://doi.org/10.1007/s40656-017-0133-6
- Stephen Jay Gould