Abstract
The adaptive landscape is an important diagrammatic concept that was conceived in population genetics. During the Modern Synthesis, in the first half of the twentieth century, the landscape imagery was used to represent evolution on a large scale, aiding in the construction of a common language for a new evolutionary biology. Not only historic adaptive landscapes by Dobzhansky, Simpson, and others are a record of how macroevolution was thought of in those decades; they stimulate reflection on “combination spaces” that underlie them. In fact, any landscape diagram is the three-dimensional transposition of a multidimensional space of combinations of genes, morphological traits, or other kinds of variables. This is an important and enduring general point of awareness: The diagram displays some aspects of the considered space while hiding others, exposing the author and the user to incomplete understanding and to conflating different spaces. Today, macroevolution is studied as a multifarious exploration of spaces of possibilities of all different sorts, interconnected in complex ways: genotype spaces, molecular spaces, morphospaces, geographical spaces, ecological spaces, and genealogical spaces. Actual macroevolutionary stories and outcomes are a subset of the universes of possible combinations—of genes, nucleotides, morphological traits, and environmental variables. Visualizations of macroevolution are a challenge of showing both distinction and correlation between spaces of possibilities.
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Notes
- 1.
When did the MS begin? Provine, finding no trace of origin in any of the major works of the MS, cited Thomas Kuhn saying that “the actual origins of a scientific field generally will not be found in the major books that embody the fundamental beliefs of the field” (Provine 1980a: 400–401). He even expressed the feeling of evolutionary synthesis having “been a part of biology for a long time, almost since Darwin” (Ivi: 400).
- 2.
A dictionary definition of metaphor states that “a metaphor is defined as a figure of speech concisely comparing two things, saying that one is the other,” while its etymology contains the idea of transfer. The English “metaphor” derives from the sixteenth century’s old French métaphore, from the Latin metaphora “carrying over.” In Greek metaphorá (μεταφορά) “transfer,” from metaphero (μεταφέρω) means “to carry over,” “to transfer”: from meta (μετά) “between” + phero (φέρω), “to bear,” “to carry.” With this etymology, metaphor seems just the right word to qualify the adaptive landscape in the MS as seen à la Ernst Mayr.
- 3.
Ironically, for personal and arbitrary reasons, Wright was absent from Mayr’s list of bridge builders (Provine 1986).
- 4.
In some versions of the landscape, including Wright’s (1932), the vertical dimension also depends on absolute, intrinsic values, such as “harmony” of the genetic combination (see also Bokma, this volume). As we see below, in Dawkins’s landscape, the vertical elevations of the surface are largely determined by some intrinsic and absolute measure such as “complexity” or “perfection.”
- 5.
“The relative capacity of a given genotype to transmit their genes to the gene pool of the following generations constitutes the adaptive value, or the Darwinian fitness, of that genotype. The adaptive value is, then, a statistical concept which epitomizes the reproductive efficiency of a genotype in a certain environment” (chapter “Toward a Natural Philosophy of Macroevolution”, “Selection”: 78).
- 6.
See Stigall, this volume, for updated evidence on the nonadaptive nature of the matching between climate and speciation in this evolutionary radiation.
- 7.
By framing the concept of “design” in an evolutionary view, and by taking just the eye evolution example, Dawkins harked back to a historical case in the development of Darwin’s theory of natural selection: The eye had been a classical example by which natural theologists built the “argument from design,” advocating the need for an intelligent designer to account for the most “adaptively complex” structures. William Paley, one of Darwin’s intellectual interlocutors, was the oft-cited champion of this school of thought. He had defended the argument from design in his influential book, Natural Theology: “…there is precisely the same proof that the eye was made for vision, as there is that the telescope was made for assisting it. They are made upon the same principles; both being adjusted to the laws by which the transmission and refraction of rays of light are regulated” (Paley 1828: 17). Darwin had looked at the eye with some worry while he was developing the theory of natural selection, and perceiving that the theory would require the demonstration of gradual implementation of complex structures. In his Notebook C (1838), he had written: “We never may be able to trace the steps by which the organization of the eye, passed from simpler stage to more perfect, preserving its relations. The wonderful power of adaptation given to organization. This really perhaps greatest difficulty to whole theory.” And again, in the Origin of Species, Darwin had declared: “To suppose that the eye with all its inimitable contrivances for adjusting the focus to different distances, for admitting different amounts of light, and for the correction of spherical and chromatic aberration, could have been formed by natural selection, seems, I freely confess, absurd in the highest degree […]. If it could be demonstrated that any complex organ existed, which could not possibly have been formed by numerous, successive, slight modifications, my theory would absolutely break down” (Darwin 1859: 143, 146). The main concern of Dawkins’s book was, in fact, a new clarification of the classic “problem of design.” Mount Improbable answered the problem by giving a visual shape to the classical argument of “chance and necessity” (cf. Monod 1970). Darwinism was defended, for Dawkins, by demonstrating the existence and viability of several intermediate forms (Elsdon-Baker 2009).
- 8.
- 9.
Kaplan (2008), for example, explains: “The problem […] is that this compression misrepresents the distances between most of the genotypes—accurate representations of distance cannot survive the packing of many dimensions into a few” (630). Pigliucci (2008) writes: “there is no metric that allows one to ‘pack’ genotypes side-by-side” (593). Sewall Wright was actually in agreement with this 'impossible packing' objection: “The two dimensions […] are a very inadequate representation of such a field” (Wright 1932: 356–357), and a surface picture “cannot accurately represent relations that are multidimensional” (Wright 1988: 116). But, while he would probably have considered it a major objection for a geometrical model, he shielded the surface picture by declaring the latter “useless for mathematical purposes” (Ibidem). The surface is not to be considered a point-by-point map of the combination space: It captures and displays some general features of the space. It is, indeed, a metaphor. Kaplan (2008), like others, accepts Wright’s defense of metaphor in general, but then, he criticizes this particular metaphor as poor and misleading with respect to the complexity of the space it represents. Other authors (e.g., Gavrilets 1997, 2004) accept Wright’s idea of metaphors and the clarity of this particular metaphor, but think that its messages are wrong, i.e., that the genotype space is not like Wright thought. For further discussion of what I call the “impossible packing objection,” see Serrelli (2011).
- 10.
Neo-Darwinian population genetics strongly held recombination as the major genetic evolutionary mechanism. For Ernst Mayr (e.g., 1980), the underemphasis of mutation by Wright and others in the first decades of the twentieth century could be straightforwardly explained by the fact that macromutations were studied by authors such as Hugo DeVries, who considered it as the non-Darwinian mechanism for the origin of species. The redefinition of mutation was, for Mayr, achieved later by the second phase of the MS.
- 11.
Dobzhansky recognizes that mutation works together with recombination, so that even the right mutations may fail to prevent extinction “if the requisite constellations of these elements do not appear in time” (Dobzhansky 1937: 277).
- 12.
In the second phase of the MS, mutation was reintegrated into population genetics and began to be seen as a major mechanism for the origin of evolutionary novelties. Those were the years in which Dobzhansky worked and wrote. But the absence (almost “eclipse”?) of recombination in those later works was, for Ernst Mayr, an “astonishing” fact: “It would be decidedly whiggish to suppress the fact that even [many biologists] slighted recombination in the 1930s. Dobzhansky, who later did much to establish the evolutionary importance of recombination, hardly referred to it in the first edition of his book (1937) […]. Recombination was, of course, well known since 1900 as one of the basic Mendelian processes and described in every genetics textbook. Remarkably, only a few evolutionary geneticists used it as a source of material for selection” (Mayr 1980: 23).
- 13.
Attributing the term “niche” to Wright is slightly anachronistic. I use niche in the “evolutionary” sense (sensu Odling-Smee et al. 2003), as the set of selective pressures acting on a population. See below for further analysis of ecological niches.
- 14.
This answers, in passing, an often expressed doubt on Wright’s landscape (e.g., Provine 1986).
- 15.
An indirect link between trees and landscapes is the following. Phylogenetics today is carried out with the aid of computer programs that, given a dataset of genetic sequences and other characters, explore the space of possible trees and determine which tree is the most likely under the given data, as well as how much we can be confident in it being the solution (see Wiley and Lieberman 2011). The process of discovering the tree is often imagined as a series of blind “robot walks” on the landscape of all possible trees, with the most likely trees on top of likelihood peaks, surrounded by similar trees that are good, but not quite!
- 16.
Some examples of this flourishing literature are O’Hara (1992, 1998), Baum (2005), Gregory (2008a), Omland et al. (2008), Thanukos (2009, 2010), McLennan (2010), Meisel (2010), Halverson (2011), Torrens and Barahona (2012). Perhaps similar discourses could be made on network thinking, concerning for example network drawing choices (that are also available as options in computer programs), cognitive tendencies in reading them, biases in recognizing modules and hierarchical levels in networks (Papin et al. 2004). But, unlike the tree-thinking “story,” this one has not been written yet.
- 17.
Hofmann and Mountjoy studied a geologic formation called the “Miette Group,” in British Columbia, Canada. Within it, they focused on the Byng carbonate platform, a recent part of the Upper Miette Group. The Byng formation within the Upper Miette context is a case of spatial and geographical heterogeneity. Furthermore, parts of the Byng Formation are “dolomitized.” Dolomitization is a chemical process by which calcium is replaced by magnesium, obliterating fossil shells. Other parts of the Byng formation are predominantly limestone. In the beds of limestone made by accumulations of biogenic carbonate—that is, shells—millimeter-scale tiny animals such as Cloudina and Namacalathus are found. These fossil shelly organisms are from the Neoproterozoic, a very ancient period—from 1,000 to 541 million years ago—that preceded the so-called Cambrian explosion. The Cambrian explosion is the relatively rapid and abundant appearance of animal phyla, including all those surviving today (except Bryozoa, which appeared later). See “Cambrian fauna” in Fig. 7a.
- 18.
Cloudina and Namacalathus are part of the worldwide Ediacaran biota, the earliest known complex of multicellular organisms. Most Ediacaran life forms are known by indirect traces (“trace fossils”), but Cloudina and Namacalathus are particular for their skeleton which is liable to fossilization. Cloudina has global distribution, while Namacalathus is rare. The Cloudina–Namacalathus association studied by Hofmann and Mountjoy (2001) in British Columbia was first found in Namibia, in the “Nama sequence.” Today, Canada and Namibia are antipodal, but, back to 1 million years ago when rocks formed, their latitude was very different, and so, was the global configuration of landmasses as we see in Fig. 11. Hofman and Mountjoy argue that finding a new instance of the Nama sequence constitutes a widening of the known geographic range of such assemblage, making it “cosmopolitan” and enhancing expectations that the assemblage will be found at least in other areas that, in the Neoproterozoic, were between current Canada and current Namibia.
- 19.
See the Paleomap Project by Chrisptopher R. Scotese (http://www.scotese.com/). Some interesting works with GIS in paleontology have been carried out by Lieberman (Rode and Lieberman 2004; Hendricks et al. 2008; Abe and Lieberman 2009, 2012; Lieberman 2012; Myers and Lieberman 2011) and Jablonski (e.g., 2008). GIS is routinely used to study current biodiversity. GIS techniques are also used the context of “geophylogenies,” i.e., integrations between geographical and genealogical data (Kidd and Liu 2008; Kidd 2010) that we shall see below.
- 20.
- 21.
From left, Rangeomorphs are “fractally quilted with frondlets arranged to form a repetitive, self-similar pattern;” Erniettomorphs “have biserially quilted tubes that are alternately arranged along a midline,” and they are not bilaterally symmetric; Bilateral forms are characterized by “anterior–posterior differentiation with a differentiated ‘head’ region;” Discoidal forms are characterized by “concentric and sometimes radial features;” taxa such as Charniodiscus and Palaeopascichnus are morphologically unique intermediates between more abundant morphologies; Triradial forms are characterized by triradial symmetry or consist of three spiral arms; Tetraradial forms are rare, Pentaradial forms are characterized by a five-fold symmetry, Octoradial forms include “one species which consists of eight spiral arms tightly wrapped into a disk”.
- 22.
The importance of this pioneering study can hardly be overestimated. Later, many “mass extinctions” beyond the Big Five have been revealed and studied in the fossil record, and knowledge in this field is constantly evolving.
- 23.
Hutchinson distinguished the fundamental niche—the maximum inhabitable hypervolume in the absence of biotic constraints—from the realized niche, a smaller hypervolume occupied by the species when under competition, predation, parasitism, and other constraints.
- 24.
A hypothetical sequence space accommodating whole genomes would be absurdly high dimensional (a human sequence space, for instance, would have 2.9 billion dimensions); not to mention the inconceivable fitness effect of every nucleotide in the sequence and of any single nucleotide substitution.
- 25.
For another example of a two-dimensional measure extracted from the variability in a multidimensional space, see Fig. 13.
- 26.
A notion of landscapes tightly related to dynamical systems is extremely interesting (see Fusco et al. 2014). Here, a dynamic vector is associated with each and every combination of the considered variables, so that the space—so to speak—fully includes the rules of its own exploration. The idea diverges somehow from all the presentation made in this chapter, which relates landscapes to static combination spaces whose exploration mechanisms exceed the combinations. As a matter of fact, macroevolution studies often employ these kinds of spaces (e.g., morphospaces, ecological niches) and connect them in different ways. They are rarely able to define dynamical systems.
- 27.
“Range size heritability” is a strong concept that has been also tied to the issue of species selection. In an interesting study on carnivores, Machac et al. (2011) clarify that range size heritability patterns are accounted for by phylogeny only in part. The patterns actually emerge as a consequence of the interplay between evolutionary and geographic constraints: Geographic constraints (e.g., temperatures, precipitations) are the most proximate factor shaping species’ ranges, but related species are sensitive to shared sets of geographic constraints.
- 28.
Stigall, this volume exemplifies the methodology of integrating biogeography and phylogeny at the macroevolutionary scale. When a daughter species occupies an area different or additional to the ancestral distribution, the speciation mode of that branching is classified as dispersal: The new species must have originated as a migrating subpopulation. When a daughter species occupies a subset of the ancestral range, speciation mode is considered vicariance: The ancestral species must have been passively fragmented by ecological barriers.
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Serrelli, E. (2015). Visualizing Macroevolution: From Adaptive Landscapes to Compositions of Multiple Spaces. In: Serrelli, E., Gontier, N. (eds) Macroevolution. Interdisciplinary Evolution Research, vol 2. Springer, Cham. https://doi.org/10.1007/978-3-319-15045-1_4
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