Macroevolution refers (most of the time, in practice) to evolutionarypatterns and processes above the species level. It is usuallycontrasted with microevolution, or evolutionary change withinpopulations. This customary way of drawing the macro/micro distinctionis not perfect, however, because species sometimes consist of multiplepopulations. Some evolutionary processes, such as the spread of atrait from one population to another, might count as within-speciesprocesses but not within-population processes.Population genetics, which emerged during the modern synthesis of the early- tomid-twentieth century, explains within-population microevolutionarychange in terms ofnatural selection,genetic drift, mutation, and migration.

One question that looms over philosophical work on macroevolutionarytheory is how macroevolution and microevolution are related. One view,which is closely associated with the modern synthesis, is thatmacroevolutionary patterns are fully explicable in terms ofmicroevolutionary processes. On this view, macroevolution is“nothing but” successive rounds of microevolution (aformulation due to Grantham 2007). Stephen Jay Gould pejorativelyreferred to this kind of view as “extrapolationism” (Gould2002). This issue links up with more general philosophical questionsabout the potential reduction of higher-level biological phenomena tolower levels (Oppenheimer & Putnam 1958; Fodor 1974; Kitcher 1984;Sarkar 1992, Rosenberg 1997).

During the paleobiological revolution of the 1970s and 1980s, a numberof paleontologists sought to establish paleontology’s status asan evolutionary discipline, with something to say about evolutionarytheory. This shift was reflected in the decision to use the term‘paleobiology’ (Sepkoski & Ruse 2009). For some ofthese scientists of the 1970s and 1980s, the disciplinary autonomy ofpaleobiology was closely linked to ideas about macroevolution. AsSteven Stanley (1975) put it, their thought was that macroevolution is“uncoupled” from microevolution, and thatmacroevolutionary patterns sometimes require explanation in terms ofmacro-level processes. One early move in this direction was theconstruction of the MBL model (Gould, Raup, Sepkoski, Schopf, andSimberloff 1977), a computer simulation of macroevolutionary processesthat explicitly ignored microevolutionary factors such as naturalselection (Huss 2009, Sepkoski 2012). Today, this approach issometimes known as the “hierarchical expansion” ofevolutionary theory, and it remains the focus of quite a bit oftheoretical work (see, e.g., Eldredge, Pievani, Serrelli, & Temkin2016). See also Lieberman and Eldredge (2024) for retrospectivereflections on this period of macroevolutionary theorizing. Dresow(2017, 2019) explores Gould’s move toward hierarchical thinking in the1970s.

Since the 1970s, theorizing about macroevolution has focused onseveral important ideas: punctuated equilibria, species selection,hierarchical theory, historical contingency, passive vs. drivenevolutionary trends, major evolutionary transitions, and more recentlythe zero-force evolutionary law (orZFEL) defended by McSheaand Brandon (2010). One philosophical challenge is to understand therelationships among these ideas; another is to work out what theymight mean for the relationship between macro- and microevolution.

The fossil record is the chief source of empirical evidence concerningmacroevolutionary patterns, and so macroevolutionary theory is closelyassociated with paleontology. In addition to questions ofmacroevolutionary theoryper se, there are alsoepistemological questions about how paleontologists infer pattern fromprocess, and how they correct for biases in the fossil data (Bokulich2021). Erwin (2024) reflects on changes in the methods and researchquestions that paleobiologists have brought to the study of the fossilrecord. In addition, the distinction between macroevolution andmicroevolution sometimes mirrors the distinction between paleontology,with its focus on the fossil record, and neontology, with its focus onobservable, extant populations. For this reason, metaphysicalquestions about the relationship between macro- and microevolution aresometimes difficult to disentangle from epistemological questionsabout what sorts of things can be inferred from different evidencebases.

The extent of various evidence bases may also be worth considering, assome have recently argued that “the vast proliferation of data,knowledge, and theory since the advent of the Modern Synthesis”(Kearney, Lieberman, and Strotz 2024) undermines any prior reason fortreating micro-and macroevolution as distinct domains, and thatcontinuing to do so thwarts potential progress on integrated researchquestions. See Hautmann (2020) for a recent discussion of three waysto define macroevolution, only one of which (purportedly) conclusivelydistinguishes macroevolution from microevolution. See Liow, Uyeda, andHunt (2023) for an introduction to current practice incross-disciplinary macroevolutionary modeling—utilizingphylogenetic comparative methods (PCMs) and (e.g.) birth-death orOrnstein-Uhlenbeck (OU) models but also fossils, ecology, evo-devo,mechanics, and other data sources as (e.g.) constraints, priors ondiversification, or generators of likelihood surfaces.

Epistemically speaking, it is worth noting that introducing more anddifferent types of data does not necessarily lead to a reduction incompetition amongst various explanatory candidates inmacroevolutionary biology. For instance, what has been termed“the nonidentifiability problem” (Louca and Pennell 2020,2021) in macroevolutionary modeling implies that, in certain types ofcases at least, “we cannot identify the true history of lineagediversification but perhaps we can identify all of the histories thatare consistent with our data” (Pennell 2023). Since adding datafrom different sources can increase rather than decrease the numberof consistent histories which serve as explanatory candidates,interdisciplinary approaches to attempts to estimate rates ofdiversification may complicate rather than resolve the explanatorylandscape (Kopperud, Magee, and Höhna 2023; Pennell 2023). Thecurrent study of macroevolution is an exciting area in which thegrowth of “big data,” the evolution of modeling practice,and the advent of interdisciplinary approaches are all impacting boththe selection of problems as well as previously domain-specific waysof addressing them—with exciting, attendant implications forepistemology and philosophy of science.

• 1. Punctuated Equilibria
• 2. Species Selection
• 3. Hierarchical Theory
• 4. Theory Accommodation and Change
• 5. Issues of Rhetoric and Risk
• 6. Historical Contingency
• 7. Passive vs. Driven Trends
• 8. The Zero-Force Evolutionary Law
• 9. Major Evolutionary Transitions
• 10. The (Dis)unity of Macroevolutionary Theory
• Bibliography
• Academic Tools
• Other Internet Resources
• Related Entries

1. Punctuated Equilibria

The classic paper on punctuated equilibria (PE) was co-authored byNiles Eldredge and Stephen Jay Gould (1972), although Eldredge (1971)was originally responsible for the idea. Their (1972) presentation ofPE draws heavily on themes from Thomas Kuhn’s (1962) work, asthey present PE as a new paradigm to rival traditional Darwiniangradualism (Turner 2011b: chapter 2). Far from challenging the modernsynthesis in that early work, they present PE as a straightforwardconsequence of allopatric speciation (speciation driven by isolatinggeographic change). Eldredge and Gould reasoned that if speciationwere typically allopatric, then it would tend to happen fairly rapidlyin geological terms, and new species would be geographically distantfrom parent species. Consequently, in the fossil record, one shouldnot expect to see very many cases where a fossil series documents aclear and gradual transition from one species to another. Eldredge andGould also argued that most morphological change occurs in(geologically) rapid spurts during speciation events and that newspecies, once they arise, typically exhibit stasis.

When Eldredge and Gould (1972) introduced punctuated equilibria, thereceived view of expected morphological change via evolution was thatof phyletic gradualism. This is a notion that can be traced all theway from Darwin’sOrigin of Species to, for instance,Simpson (1970). In accepting phyletic gradualism, paleontologists hadlong had to accept that the incompleteness of the fossil record wassignificantly responsible for the lack of (ideally expected)geological documentation of gradual transformation in the majority offossilized speciation events. What Eldredge and Gould uniquelyrealized, however, was that the expectation of phyletic gradualismclashed not only with what the fossil record actually showed, but alsowith what the dominant model of (allopatric) speciation predicted. Sothey developed PE: an alternative view of morphological evolution (andstasis) intended to synthesize what allopatric speciation projectedwith what the fossil record depicted.

In theOrigin of Species, Darwin had attributed the lack offossil series exhibiting gradual morphological transition to theincompleteness of the fossil record. He thought of speciation andmorphological evolution as gradual processes (though he didacknowledge variation in evolutionary rates), and thought that theyoften happen under the geological radar. As he put it: “Naturemay almost be said to have guarded against the frequent discovery ofher transitional or linking forms” (Darwin 1859 [1964: 292]).Eldredge and Gould were, in contrast, proposing that the fossil recordis more complete than it seems, and that we should take the rapidappearance of new species more or less at face value. As theyeventually put it, in designing the theory of punctuated equilibriumthey were “hoping to validate our profession’s primarydata as signal rather than void” (Gould & Eldredge 1993:223). One epistemological issue in play here is whether the absence ofevidence—in this case, the absence of evidence of gradualevolutionary change—is really evidence of absence (Sober 2009;Currie & Turner 2017).

The ensuing debate about punctuated equilibria proved to be complexand contentious. Gould, in particular, made increasingly bold andcontroversial claims on behalf of PE during the 1980s, and he tendedto see PE as the opening wedge for a more ambitious critique of themodern synthesis. Notoriously, Gould also flirted with non-Darwinianaccounts of speciation, such as Richard Goldschmidt’s (1940)idea that new species arise from “hopeful monsters” (Gould1977). The stasis claim of PE was also controversial, especially sinceGould took it to mean that cumulative, directional natural selectionis a less significant factor in evolution than many had thought. Thereasoning was that if directional natural selection were the dominantdriver of evolutionary change, then you would not expect to see suchclear patterns of stasis in the fossil record. PE elicited a greatdeal of opposition from biologists and philosophers who were morecommitted to a selectionist picture of evolution (Dawkins 1986;Dennett 1995; and see Sterelny 2001a for a useful guide to thecontroversy).

In the meantime, PE also inspired new empirical research inpaleontology, with scientists doing statistical analysis of largefossil samples to try to assess the relative importance of stasis vs.gradual directional change. This research continues to get moresophisticated, and as it happens, PE has fared reasonably well,empirically (J. Jackson & Cheetham 1999; Hunt 2007; Uyeda, Hansen,Arnold, & Pienaar 2011).

One other philosophical issue raised by PE concerns the explanation ofevolutionary stasis (Turner 2017). Eldredge and Gould never reallyoffered a very clear account of the mechanism(s) that could maintainstasis, a fact that critics often seized upon (e.g., Coyne &Charlesworth 1996). They did propose that species might be homeostaticsystems. To make matters worse, there is a mismatch between the fossilevidence (where stasis is common) and studies of extant populationsthat show lots of directional evolutionary change (Price, Grant,Gibbs, & Boag 1984; Endler 1986; Lenski & Travisano 1994;Harshman & Hoffmann 2000), a puzzle sometimes called the“paradox of stasis” (Hendry 2007). Many evolutionarybiologists (e.g. Estes and Arnold 2007) are quick to attribute stasisat larger scales to well-known processes such as stabilizing selectionand/or habitat tracking, but Kaplan (2009) raises some criticalquestions about that move. And Sterelny (2001b) has pointed out thatcases of “coordinated stasis” in the fossil record posespecial explanatory challenges. Coordinated stasis is a pattern wherea whole ecological assemblage seems to persist without change formillions of years. Stasis, the idea at the heart of PE, remains anissue of considerable theoretical and empirical interest inpaleontology (Lidgard & Hopkins 2015; Lidgard & Love2018).

2. Species Selection

If punctuated equilibria (PE) was the first innovative idea aboutmacroevolution to emerge during the 1970s, species selection followedclose on its heels. Just how the two ideas are related is itself atopic of some philosophical interest (Turner 2010). According toEldredge and Gould (1972), driven ordirectional change isonly part of the theoretical story about the evolution of morphology;it is not, perhaps, even a geologically common narrative. Directionalselection is selection that changes a population, pushing itsdistribution of traits from one pattern of trait expression toanother. More typical, according the Eldredge and Gould, are cases ofeithernon-directional change or evenstasis. Likedirectional selection, nondirectional selection also changes apopulation, but not towards any particular distribution oftraits—it just moves a population from one pattern to another.Stabilizing selection is selection that keeps a population the same,nudging it back to its trait distribution pattern whenever it startsto stray away. PE as originally formulated favors stabilizingselection, suggesting that both individuals and species are inherentlystable homeostatic systems—systems that are “amazinglywell-buffered to resist change and maintain stability in the face ofdisturbing influences” (Eldredge & Gould 1972: 114). On thisview, whatever the mechanisms that “resist change and maintainstability” are, they are in conflict with and must be overcomeby directional natural selection in order for natural selection tosuccessfully push for or “force” directional change.Whether natural selection is properly thought of as a force is itselfan interesting philosophical issue (Sober 1984; Walsh 2000; Matthen& Ariew 2002; Walsh, Lewens, & Ariew 2002; Bouchard &Rosenberg 2004; Stephens 2004; Brandon 2006; Millstein 2006; Filler2009). Setting that question aside, however, allows us to consider thesuggestion that species might be homeostatic systems which“resist change by self-regulation” (Eldredge & Gould1972: 114)—as this is a suggestion rife with implications formacroevolutionary theory.

At first, the idea that species are homeostatic systems may have beena way of addressing critics of PE, who wondered what sort ofbiological mechanisms could maintain stasis over long stretches ofgeological time. However, it is a short step from thinking of speciesas homeostatic systems to thinking of them as the sort of thing thatcould be a unit of selection. Eldredge and Gould thought thatvariation introduced amongst organisms is typically isolated withinthe population and constrained by stabilizing selection, unlessgeographic separation creates the conditions for rapid evolutionarychange. During allopatric speciation events, the adaptive forcesnegate the stabilizing ones effectively enough to disrupt the stasisof the species and cause the formation of a new lineage. Eventuallythough, the stabilizing forces (re)assert themselves upon the emergentspecies and (re)establish stasis. Most biologists think of stabilizingselection as an ordinary microevolutionary process (Schmalhausen1949); however, the suggestion that a species is a homeostatic systemmakes it look like some form of stabilizing selection might beoperating (in some sense) as a mechanism that maintains equilibrium atthe level of populations.

When modeling how these interactions generate macroevolutionarypatterns, PE presents the dominant (i.e., equilibrium) state as one ofpopulation-level morphological stasis—a state in whichspeciation attempts often occur but generally fail to take. Theseperiods of stasis are occasionally interrupted (i.e., punctuated) byperiods of population-level morphological change—periods ofsuccessful speciation. This bifurcated model (depicting states of bothstasis and change) is one in which species behave a little bit likeorganisms: speciation begins to look like reproduction, and extinctionbegins to look like death. Crucial to this picture is the claim thatonce a new species comes into existence, it mostly exhibits stasis.The analogy is by no means perfect (Havstad 2016 in Other InternetResources), but it did open up a new field of theoretical exploration.What if the differential speciation and extinction of whole lineagesis a lot like the differential survival and reproduction ofindividuals in a population? PE may not logically imply speciesselection, but it does point in that direction by suggesting thatspecies resemble organisms in relevant ways—for instance, byhaving comparable integration and cohesion, enough for populations toblock directional change in favor of stasis.

Initially, Eldredge and Gould (1972) were quite explicit that they didnot see their model as entailing a new type of selection. Theystraightforwardly wrote “we postulate no ‘new’ typeof selection” (Eldredge & Gould 1972: 112). But in 1975, thepaleontologist and evolutionary theorist Steven M. Stanley dubbedEldredge and Gould’s population-level selection process a formof species selection, and by 1977 the inventors of PE concurred. Theywrote that they nonetheless understood species selection asrepresenting “no more than the operation of natural selection athigher levels” (Gould & Eldredge 1977: 139), but still, thisinterpretation presents a challenge to one typical way ofinstantiating the dominant model of natural selection.

Regarding the dominant model: classically and summarily described (seeGodfrey-Smith 2007), evolution occurs when individuals in a populationaccumulate a change to the degree that this change becomes a featurein the population. On this view, natural selection requires variation,fitness, and heritability (Lewontin 1970; though see Brandon 1990).When all three of these things coincide within populations, and changeresultantly accumulates, evolution occurs. Proponents of speciesselection argue that these three conditions are met at the specieslevel.

But on a typical way of instantiating this rather abstract model ofnatural selection, one can identify what plays the role of theindividuals in this model (organisms) and what plays the role of thepopulations (species). This popular view treats organisms as the unitsof selection and species as the units of evolution. Species evolve,owing to the differential survival and reproduction of organisms. Thisemphasis on organism-level selection and species-level evolution isanother which dates back to Darwin’sOrigin of Species,although Darwin also countenanced family selection (Darwin 1859),sexual selection (Darwin 1871) and group selection (Stauffer 1975).Some early proponents of the modern synthesis (e.g., Mayr 1942, 2001)also shared this basic picture in which selection acts on organisms,while species evolve. To propose that selection occurs at the level ofspecies as opposed to that of organisms, is to suppose that not onlycan organisms act as the units of selection but that species cantoo.

So, if the proper way to understand natural selection casts organismsas the units of selection, and species as the unit of evolution, thenGould and Eldredge’s model of PE challenges that view.Alternatively, if the proper way to understand natural selectionmerely names biological individuals as the units of selection andbiological populations as the unit of evolution, and species can bebiological individuals too, then Gould and Eldredge’s model ofPE is not really a challenge to the model of natural selection, it isjust a new instance of it. This is because the notion ofbiological individual is more expansive than that of biological organism; biologicalindividuals include but are not limited to biological organisms.Biofilms, coral reefs, insect colonies, obligate symbionts,siphonophores, slime molds, and viruses are just a few of the manybiological entities that we might want to consider as individuals,although they are not organisms (see Buss 1983; J. Wilson 1999; Clarke2010, 2013; and R.A. Wilson & Barker 2013, among others). Ifspecies are themselves individuals (Ghiselin 1974; Hull 1976), thenthe traditional evolutionary theorist may be able to accommodatespecies-level selection—in this sense, it is just another flavorof individual-level selection. Note that is it is very easy for thisdispute to become an instance of what the philosopher John Beatty(1997) calls a “relative significance” debate—one inwhich proponents move from arguing about whether something is possible(species acting as the unit of selection) to whether that thing issignificant (as opposed to a mere oddity).

Many of these issues—such as how to understand stabilizingmechanisms and selection, what biological individuals are, whetherspecies are individuals, and whether species-level selection isreducible to individual-level selection—are ongoing. In thedecades since Eldredge and Gould first outlined the theory ofpunctuated equilibria, various biological theorists have attempted toprovide definitive and irreducible examples of species-levelselection—ones that cannot be recreated at lower levels such asthat of the organism. Species-level traits like range size (where aspecies may be wide-ranging even though its organisms are not; seeJablonski 1987) and rate of speciation (where a species may be slow to“reproduce” although its organisms are not; see Vrba 1987)are especially promising candidates. In addition to the issue ofproviding definitive examples of irreducible species selection, thereis also the issue of providing a satisfactory definition of theconcept itself. The biologist Elisabeth Vrba (1984) posits that allproposed definitions of species selection ought to identify causes, berestricted enough for testing, and employ a concept of selection thatis consistent with how selection is understood to act at other levelsas well. See Vrba (1984), Cracraft (1985), Grantham (1995), Jablonski(2008), and Turner (2011b: chapters 4 and 5) for further discussion ofspecies selection.

3. Hierarchical Theory

It is worth noting that several discussions of species selectionpre-date Eldredge and Gould’s (1972) proposal (e.g., Fisher1929, Wright 1956; Lewontin 1970). However, it is the ensuingdiscussion of stabilizing selection prompted by PE—incombination with a contemporaneous uptick in discussion of kinselection, group selection, and altruism (Haldane 1932; Wright 1945;Wynne-Edwards 1962; Hamilton 1963; Maynard Smith 1964; Boorman &Levitt 1973; E.O. Wilson 1973; Levin & Kilmer 1974; D.S. Wilson1975; Wade 1978)—which transforms these macroevolutionary sparksinto an ongoing, hierarchical conflagration (see Gould 1980b, 1982,2002; Vrba & Eldredge 1984; Cracraft 1985; Eldredge 1985; Grantham1995; and Eldredge, Pievani, Serrelli, & Temkin 2016, amongothers). Indeed, some theorists see the paleobiological revolution ascontributing to a “hierarchical expansion” of evolutionarytheory.

Evolutionary theorists often distinguish between the unit of selection(Williams 1966; Franklin & Lewontin 1970; Lewontin 1970) and theunit of evolution (Williams 1966; Hull 1978; Ereshefsky 1991).Expanding the category of what can act as a unit of selection leadsdirectly into a corresponding expansion of what can act as a unit ofevolution. This is how a hierarchical theory of evolution istheoretically generated: when theoreticians consider not justorganisms or individuals as theunit of selection but also species and (kin) groups, and genes (for instance), they arealso and correspondingly considering not just species or populationsas the unit of evolution but also clades, and cultures, and cells orperhaps organisms (respectively). Each proposed unit of selectionincurs another, higher-level candidate for the unit of evolution.Hierarchical evolutionary theory is just the name for theories ofevolution that admit of different units of selection and evolutionthan that of (merely and respectively) organisms or individuals andspecies or populations. Hierarchical theorists see evolutionaryprocesses (especially selection processes) unfolding simultaneously atdifferent biological levels and temporal scales.

Hierarchical thinking about evolution is perhaps best illustrated withan example. It has already been noted that one way of explainingmorphological stasis is stabilizing selection. Stabilizing selectionis a well-understood Darwinian mechanism that operates on organisms ina population. Here, organisms are the units of selection while speciesare the units of evolution. But Sheldon (1996) introduced anotherpotential explanation of stasis, which he called the “plusça change” model. Sheldon observed that in times ofenvironmental turmoil, ecological generalists usually have lowerextinction risk, while ecological specialists are more prone toextinction. Perhaps surprisingly, then, a pattern of stasis in thefossil record might be due to the persistence of ecologicalgeneralists during times of environmental upheaval. This model appealsto species selection, or to the differential extinction andpersistence of whole species, rather than to stabilizing selectionprocesses occurring within those species. Hierarchical theorists holdthat evolutionary patterns (like stasis) can be generated andsustained by processes operating at different biological levels.

Altogether, hierarchical evolutionary theory is potentially inclusiveof macroevolutionary processes (such as species selection), along withparadigmatic microevolutionary processes (such as organism-levelselection), sub-microevolutionary processes (such as genic selection),and even super-macroevolutionary processes (like clade selection).Lewontin (1970) considers the possibility of the following acting asunits of selection: molecules, cell organelles, cells themselves,gametes, individuals (this is the term he uses), kin groups,populations, species, communities of species, and phyla. Williams(1966) and Dawkins (1976) are often credited with developing thenotion of genic selectionism. Debate over genic selectionism—atheory within which genes are both the units of heredity and the unitsof selection—has led to further philosophical discussionregarding what counts as a replicator as opposed to an interactor(Hull 1980; Brandon 1988, 1990; Lloyd 1988; Griffiths & Gray 1994,1997; Sterelny 1996; Sterelny, Smith, & Dickison 1996;Godfrey-Smith 2000; Griesemer 2000; Nanay 2002). Various biologists(e.g., Mayr 1963; Gould 1980a; Lewontin 1991) have ardently disputedthe claim that genes act as direct targets of selection. Philosophersof biology have taken positions both pro (e.g., Sterelny & Kitcher1988; Waters 1991) and con (e.g., Sober & D.S. Wilson 1994; Lloyd2005) on the topic of genic selectionism.

Philosophical discussions of genic selection, clade selection, otherunits of selection, and the hierarchical theory of evolution areextensive and ongoing (in addition to works already cited, see Wimsatt1980; Kitcher, Sterelny, & Waters 1990; Sober 1990; Godfrey-Smith& Lewontin 1993; Stanford 2001; Glennan 2002; R.A. Wilson 2003;Walsh 2004; Waters 2005; and Okasha 2006, among others). Anotherinteresting, relatively recent, and somewhat related development isthe proposal of a further, even-more-extended evolutionary synthesis,or EES (Jablonka & Lamb 2005; Pigliucci & Müller 2010;Laland et al. 2014; though see Wray et al. 2014). Proponents of EEScontend that phenomena such as developmental bias (Arthur 2004), nicheconstruction (Odling-Smee, Laland, and Feldman 2003), phenotypicplasticity (West-Eberhard 2003), and non-genetic inheritance processeslike social learning and cultural transmission (Hoppitt & Laland2013) challenge evolutionary theory as standardly understood.Opponents contend that the now-traditional theoretical frameworkconstructed via the modern synthesis can accommodate all thesephenomena and more. See Pigliucci and Finkelman (2014) for adiscussion of the philosophical implications of the EES debate.

Whether these phenomena challenge the modern synthesis or not, factorssuch as developmental bias—understood as the manner in which“developmental systems impose uneven probability distributionson the phenotypes accessible from a given evolutionary startingpoint” (Jablonski 2020)—provide candidate means of linking(for instance) micro-level phenotypes with macroevolutionary trends,and vice versa. For instance, a recent study—one focused on theevolution of teeth in extant and extinct primates—contrasted theexplanatory power of various comparative models in quantitativegenetics, attempting to link microevolutionary processes withhigher-level patterns of macroevolutionary divergence. Study resultssuggested that “biologically informed morphospaces alongsidequantitative genetics models allow a seamless transition between themicro- and macroscales, whereas biologically uninformed spaces donot” (Machado et alia 2023). Work like this suggests that fossildata, developmental studies, and other biological and paleontologicalevidence bases may, when better integrated with macroevolutionarymodels, help to reconcile previously discordant trends andobservations. Methodical attempts to bridge the micro and the macro,whether reductive in their approach or otherwise, echo throughout thehistory of this scientific domain and continue unabated in its presentmoment (e.g., Rolland et alia 2023; Schluter 2024; Tsuboi et alia2024; Voje, Saito-Kato, and Spanbauer 2024).

4. Theory Accommodation and Change

One meta-scientific issue raised by these macroevolutionary disputesis just how much a scientific theory can flex and change before iteither becomes a different theory altogether, or it becomes a badscientific theory (via too much of what Popper [1974] would callad hoc revisionism).

As an illustrative example, consider punctuated equilibrium, which hascertainty been articulated in many and varying ways. The two initialpublications by Eldredge (1971) and Eldredge and Gould (1972) areundoubtedly the founding ones, but there are other crucial moments inthe development of the theory (e.g., Gould & Eldredge 1977, 1983,1986, 1993; Eldredge & Gould 1988; Gould 2002). At times theshifting nature of PE has been a target of criticism (e.g., Levinton1986; Coyne & Charlesworth 1997; see Eckhardt 1986 for aparticularly entertaining visit from “the shade of FrancisGalton”).

Another dynamic theoretical dimension of PE, hierarchical theory, andthe like is how radically these scientific notions areperceived—even by their own creators. In 1982, for instance,Gould first asked and then answered:

What would a fully elaborated, hierarchically based evolutionarytheory be called? It would neither be Darwinism, as usuallyunderstood, nor a smoothly continuous extension of Darwinism, for itviolates directly the fundamental reductionist tradition embodied inDarwin’s focus on organisms as units of selection. (1982:386)

But in 2002, Gould wrote

I do believe that the Darwinian framework, and not just thefoundation, persists in the emerging structure of a more adequateevolutionary theory. (2002:3)

He devoted much of the extensive efforts of hisStructure ofEvolutionary Theory to defending an interpretation of Darwinianevolutionary theory as resting on a revisable “tripod ofnecessary support” (Gould 2002: 586)—one whose threecritical supports were challenged by proposed expansions toevolutionary theory, but which nonetheless could be buttressed viaintegration with these challenges, rather than replacement by them. Inshort, the later Gould argues that Darwinian evolution traditionallyemphasizes organismic selection, adaptationism, and extrapolationism;that punctuated equilibria, stabilizing selection, and speciesselection challenge the first of these supports, developmental andother constraints challenge the second, and major evolutionarytransitions challenge the third; but that all three of thesechallenges can be accommodated by a revised but still Darwinianconception of evolutionary theory.

Note that efforts to retain (or reject) the Darwinian label show up inother contexts as well—for instance, in discussions ofadaptationism (e.g., Gould & Lewontin 1979) and the neutral theoryof molecular evolution (Dietrich 1994). Darwin himself declared that“He who rejects these views [his] on the nature of thegeological record, will rightly reject my whole theory” (1859[1964: 342]). Whether or not Darwin’s emphatic claim is correctis still an open question. Additional philosophical resources ontheoretical and conceptual change in science include but are by nomeans limited to Kuhn (1962), LaPorte (2004), and M. Wilson(2006).

5. Issues of Rhetoric and Risk

From October 16^th to 19^th of 1980, a conferenceon the subject of macroevolution was held at the Field Museum ofNatural History in Chicago, Illinois. Shortly thereafter—in theNovember 21, 1980 issue ofScience—an exciting reporton some events of the conference was published (Lewin 1980). Thereport was entitled “Evolutionary Theory under Fire”, andit prompted a cascade of responses in the Letters section ofScience—an exchange of scientific correspondence thatcontinued for almost two decades. Whatever the motives behind theheadlining of the report, it is empirically evident that the chosentitle was inflammatory.

The February 21, 1981 issue ofScience included no fewer than5 separate contributions (in order of printing: Futuyma et al. 1981;Templeton & Giddings 1981; Carson 1981; Olson 1981; Armstrong& Drummond 1981). The February 4, 1983 issue ofSciencecontained a letter by critics of PE (Schopf & Hoffman 1983) and areply (Gould 1983). In the March 11 issue of that same year both priorparties were accused of prolonging a fruitless discussion of a“philosophically intractable” and dubious“pseudoquestion” (Grant 1983: 1170). In April, onecorrespondent wrote in agreement with Grant (Schoch 1983) and anotherchided him for failing “to explore and appreciate the latestblooming in the desert of dogma” (Maderson 1983: 360). To put itmildly, and purely descriptively: many of the responses elicited bythe original report were impassioned and emotive.

In the February 14, 1986 issue ofScience, the originalreporter on the macroevolutionary conference fanned the flames withanother piece, one entitled “Punctuated Equilibrium is Now OldHat” (Lewin 1986). In March, Jeffrey Levinton registered anardent objection to Lewin’s characterization of the theory aswidely accepted, as well as to PE in general (Levinton 1986). InApril, Gould responded (Gould 1986), and in May, Robert Eckhardtresponded to Gould’s response (Eckhard 1986). Throughout theseexchanges and related ones, many interesting meta-scientific chargeswere laid and lobbed. Questions of “where is theevidence?” as well as accusations of dogma, religionism,revisionism, sloganeering, and vagueness abounded.

So did references to creationism, specifically. The very first batchof letters in response to Lewin’s initial piece included oneobjecting strongly to the choice of title (Armstrong & Drummond1981). The correspondents wrote that “this article isundoubtedly destined to enter the out-of-context arsenal that hasbecome a mainstay of recent creationist literature” (Armstrong& Drummond 1981: 774). Incidentally, the correspondents werecorrect about this (e.g., Meyer 1994; Parker 2006).

And in the March 10, 1995 issue, a different journalist forScience published another piece, this one entitled “DidDarwin Get It All Right?” (Kerr 1995). By May 5, a scientist hadwritten in to protest the use of such an inflammatory headline(McInerney 1995), referencing Lewin’s original piece (from1980). McInerney wrote that Lewin’s earlier piece had“provided a gold mine for subsequent creationistpropaganda” (1995: 624). He bemoaned the fact that another suchheadline had just been published byScience, and recommendedthat “a journal representing some 130,000 scientists be a bitmore judicious in its choice of headlines” (1995: 624).

These references to a wider social context—the one in which suchardent macroevolutionary debate is being held—raise interestingissues that have perhaps been somewhat neglected by philosophers ofscience (though not by those working onthe social dimensions of scientific knowledge). Do the scientists involved in this dispute, and the scientificjournalists reporting on it, have a responsibility to consider the(unintended yet foreseeable and potentially avoidable) uses to whichtheir headlines might be put (by creationists and other parties)? Somescientific correspondents evidently think they do. And if suchresponsibility exists, does it fall upon the shoulders of thescientistqua scientist—or upon the shoulders of thescientistqua citizen, or scientistqua educator?Philosophers focusing on the relation between science and value havedone some relevant work on questions like these, usually whilefocusing on other scientific domains—work which could be appliedto macroevolutionary instances and issues (e.g., Rudner 1953; Hempel1960; Longino 1990; Douglas 2000, 2009; Kourany 2010).

These are not the only issues of provocative linguistic expressionawaiting further exploration by philosophers of macroevolution.Questions of what it means to sit “at the high table”(Maynard Smith 1984; Sepkoski 2014) or to “replay life’stape” (Gould 1989; Beatty 2006; Sepkoski 2016) are perhaps morefamiliar to philosophers of biology, but still worth exploring.

6. Historical Contingency

Stephen Jay Gould (1989) famously argued that evolutionary history iscontingent. George Gaylord Simpson (1963) foreshadowed that claim whenhe suggested that paleontology is a distinctively historical sciencethat seeks to understand “configurational change”. Gouldclaimed that if we could rewind the tape of history to some point inthe deep past and play it back again, the outcome would probably bedifferent.

Gould’s thinking about contingency has had significant impactsin both philosophy and biology. In philosophy, for instance, JohnBeatty (1995) argued that the contingency of evolutionary historyimplies that there are no distinctively biological laws, an argumentthat served as a touchstone for much of the subsequent debate aboutlaws in biology (Brandon 1997; Mitchell 1997, 2000, 2002; Sober 1997;Giere 1999; Shapiro 2000; Woodward 2000, 2001, 2004; Elgin 2003, 2006;and Hamilton 2007; for important precursors to Beatty 1995, see Smart1963; Ruse 1970; Cartwright 1983; van Fraassen 1989; and Rosenberg1994). More recently, Beatty (2016) has argued that the contingency ofevolutionary history is closely connected to narrative modes ofexplanation. His suggestion is that narrative explanations areespecially well suited to historically contingent series of events.McConwell (2017) argues that the contingency of evolutionary historyexplains why we should be pluralists about biological individuality.In biology, on the other hand, Gould’s argument aboutcontingency, particularly his thought experiment concerning rewindingand playing back the tape of history, helped to inspire research inlong-term experimental evolution (e.g., Travisano, Mangold, Bennett,& Lenski 1995; Blount, Lenski, and Losos 2017). In addition tofueling interest in understanding the Cambrian explosion,Gould’s work also instigated something of a backlash from otherscientists convinced that selection-driven convergence is the hallmarkof evolutionary history (e.g., Conway Morris 2003). This debateconcerning contingency and convergence sometimes also has theologicalovertones. Conway Morris has at least hinted that he seesconvergentism about evolution as cohering especially well with theism.This contrasts with Gould’s well-known view that science andreligion should be regarded as “non-overlappingmagisteria” (Gould 1997). Scientists have also published popularbooks exploring the theme of contingency (De Quieroz 2014; Losos2018).

There is some question about just what Gould meant by the term‘contingency’. Some philosophers read him as meaning thatdownstream outcomes are sensitive to small changes in upstreamconditions (Ben-Menahem 1997; Inkpen & Turner 2012). On thisreading, contingency comes in degrees, and in contrast with historicalinsensitivity to changes in upstream conditions, which wecould think of as historical convergence or robustness. When, forexample, Sterelny (1995, crediting F. Jackson & Pettit 1992) talksabout “robust process explanations”, he means explanationsthat show how an outcome would have resulted from a wide range ofupstream conditions.

Beatty (2006), however, has shown that there are two different sensesof ‘contingency’ in play in Gould’s work. Inaddition to what Beatty calls contingency as causaldependence—basically, sensitivity to initialconditions—there is a second form of contingency that Beattyinitially called contingency as unpredictability, but now callscontingencyper se (Beatty 2016). These two senses ofcontingency correspond with two versions of the famous thoughtexperiment that Gould (1989) deployed. Sometimes, Gould imaginesrewinding the tape of history, tweaking an upstream variable, and thenplaying the tape back. On other occasions, he talks about playing thetape back from the same initial conditions. Beatty (2016) thinks thatboth senses of ‘contingency’ are important, and he takesit that the second sense—contingencyper se—mustcommit us to some sort of causal indeterminism. On the other hand,Turner (2011a) has tried to give an account of this second sense ofcontingency that is neutral with respect to determinism. Hissuggestion is that what Gould really cared about was random orunbiased macroevolutionary sorting. Processes such as coin tosses, orrandom genetic drift, can be random or unbiased (in a sense) withoutviolating causal determinism. One way to think about this is byadopting a frequentist conception of probability: the outcome of acoin toss could be causally determined by small-scale physicalinfluences, but the outcome is still random or unbiased in the sensethat over a long series of trials, the ratio of heads to tails willapproximate 50:50. This account is closely allied withSepkoski’s (2016) suggestion that Gould’s metaphor ofreplaying the tape of history was rooted in his work on simulations ofmacroevolution in the 1970s. Turner (2015) has also argued that thereis a connection between contingency, thus understood, and the notionof a passive evolutionary trend.

McConwell and Currie (2017) have added further complexity to thediscussion by arguing that there is an important difference betweenabstract,source-independent accounts of contingency andsource-dependent accounts. The notion of sensitivity toinitial conditions is a good example of what they mean by asource-independent account: it gives an abstract characterization ofhistorical or biological processes without saying what the source ofcontingency is. Does the source have something to do with geneticdrift? With mutational ordering? Or something else entirely? McConwelland Currie argue that in order to establish the autonomy ofmacroevolutionary theory, a source-dependent account is required.

Some commentators have noted that Gould’s defense ofevolutionary contingency sits in tension with his arguments, in the1970s and early 1980s, for a more “nomothetic”paleobiology (e.g., Raup & Gould 1974). A more nomotheticpaleobiology would seem to place the emphasis on laws andpredictability. However, contingency is thought by some to mean thatthere are no distinctively biological laws (Beatty 1995). Haufe (2015)addresses this puzzle by arguing that Gould was interested indeveloping a “nomothetic” science of macroevolutioninvolving generations at large spatial and (especially) temporalscales, but that this is compatible with saying that contingencyreigns at other scales. One other larger theme that may help to unifysome of Gould’s thinking is his skepticism about the power ofnatural selection, which is the classic example of amicro-evolutionary process (see also Gould & Lewontin 1979). Oneway to limit the importance of natural selection is to insist on theautonomy of macroevolutionary theory, and to show that naturalselection is insufficient to explain certain larger scale patterns.Another way is to insist that evolutionary history is contingent,since one would expect selection to produce convergent outcomes.

Thus, philosophers have done a lot of work on challenging conceptualquestions about the various senses of ‘contingency’ andtheir relationship to other macroevolutionary ideas. One problemlurking in the background of this discussion is empirical: how canscientists even tell whether evolutionary history is contingent or(say) convergent? Conway Morris (2003) assembles a whole catalog ofexamples of evolutionary convergence in an effort to providecounterevidence against Gould’s contingency thesis. Oneimportant issue here is to find a principled way of distinguishingcases of convergent evolution (where unrelated lineages subject tosimilar selection pressures evolve similar traits) from parallelevolution (where lineages with similar traits to start with followsimilar evolutionary trajectories) (Pearce 2012). Another issue isthat one can multiply convergences by giving coarser-graineddescriptions of the traits in question—for example, by countingagriculture in humans and some ant species as the same trait (Sterelny2005). Powell & Mariscal (2015) argue that not all of the putativeexamples of convergence really tell against the contingency thesis,and they try to specify more clearly what would actually count asevidence against contingency.

In developing his initial claim about evolutionary contingency, Gould(1989) made much of the Burgess Shale fauna, exquisite Cambrianfossils from western Canada first described by the Americanpaleontologist Charles Doolittle Walcott in the early 1900s. Gould waswriting in the wake of what some call the “firstreclassification” of the Burgess fossils, when work by Britishpaleontologist Harry Blackmore Whittington and students Derek Briggsand Simon Conway Morris had placed many of the strange forms in theirown phyla, with the idea that each phylum represented a unique bodyplan. Most of those Burgess phyla went extinct, but some (including alikely ancestor of chordates) persisted. But what if that sorting hadhappened differently? Since 1989, scientists Graham Budd, AllisonDaley, and others using cladistic methods of classification (whichwere slow to be adopted by paleontologists) have reclassified theBurgess fauna yet again, placing many of the strangest ones in stemgroups of existing clades (such as arthropods). So one furtherphilosophical question is whether and to what extent this change intaxonomic practice matters to the argument that Gould tried to makeabout contingency (Brysse 2008).

There is a closely related debate about the status of phyla. Somecladists see phyla as being nothing terribly special. Like any highertaxa, phyla must be monophyletic, but from a cladistic perspectivethere is nothing about the taxonomic rank of phylum that has anyparticular importance to macroevolutionary theory. Others have arguedthat lower ranks like order and class are evolutionarily meaningful ina way that phyla are not (e.g., Holman 1989). Gould (1989), however,drawing upon German paleontological tradition, saw phyla ascorresponding to stable morphological body plans (orBaupläne). This connects closely with his argument aboutcontingency: if different Cambrian phyla had persisted, downstreamevolutionary history would have involved modification of totallydifferent body plans. Some scientists have recently tried to revivethe idea that there is something special about the role of phyla inmacroevolution (Levin, et al. 2016; though see Hejnol & Dunn2016).

Historical contingency is a counterfactual notion, and although thisissue has not gotten as much attention as it deserves, there is anascent philosophical literature on historical counterfactuals (Tucker2004: 227ff; Nolan 2013; Radick 2016; Zhao 2017 in Other InternetResources; Zhao 2023). The debate about historical contingency can beconstrued as a disagreement about the truth of various historicalcounterfactuals. Gould claimed that if things in the Cambrian had beenslightly different, there would be no vertebrates today, let alonehumans, while other convergentists claim that humanlike cognitiveabilities, language, tool use, and sociality would have evolved evenif other things had been different in the past—for example, ifthe non-avian dinosaurs had not gone extinct.

More recently, philosophers and historians interested in contingencyhave begun to turn their attention to the various ways in whichcontingency intersects with normative questions. For example, Perez(2024) explores connections between Stephen Jay Gould’s interest incontingency and his political commitments. McConwell (2023) looks athow George G. Simpson and Stephen Jay Gould both thought about thenormative consequences of their evolutionary ideas, includingcontingency. Although interest in contingency owes much topaleontologists’ theorizing about macroevolution, contingency has alsofound its way into more popular discussions about history and thesocial sciences (e.g. Klaas 2024).

7. Passive vs. Driven Trends

Paleontologists have long been interested in documenting andexplaining larger-scale evolutionary trends in the fossil record(McShea 1998). Perhaps the classic example of a macroevolutionarytrend, and one that paleontologists have studied extensively, isCope’s rule of size increase. Paleontologists routinelydistinguish within-lineage trends (e.g., average size increase withina single lineage) from among-lineage trends (e.g., increase in theaverage size of mammals over the Cenozoic). The latter aredistinctively macro-level patterns. While it might be tempting tothink that trends such as evolutionary size increase are generallyattributable to natural selection, Stanley (1973) pointed out thatthese trends could be merely statistical phenomena. Suppose that aclade, such as mammals, starts out at or near a fixed lower boundaryon body size. And suppose that evolutionary size increases anddecreases are equally probable. As the clade evolves, the mean bodysize might do a “random walk” away from the fixedboundary—an idea that Gould (1996) later popularized. Manyscientists also think of this as a passive diffusion model ofevolutionary change.

The notion of a passive trend is closely related to other themes ofmacroevolutionary theory. Gould, in particular, was enthusiastic aboutthe possibility that Cope’s rule might be a passive trend. Ifthat were the case, then it would mean that evolutionary size increaseis not attributable to natural selection. What’s more, if weexplain size increase by supposing it to be a passive trend, then theexplanation need not reference any micro-level causes. The fixedboundary in the state space could be maintained by developmentalconstraints—another favorite theme of Gould’s. Andfinally, a trend resulting from a random walk away from a fixedboundary would not seem to be progressive at all. To the extent thatlarger-scale evolutionary trends are passive, chance looks like a moresignificant factor in evolutionary history, and selection looks like aless significant one.

McShea (1994) gave one widely-cited formulation of the distinctionbetween passive and driven trends, and also suggested severalapproaches for determining, empirically, whether a trend is passive ordriven. One such approach is the stable minimum test: track, say, thebody size of the smallest members of the clade. If that increasessteadily over time, it is a signal that the trend is driven.Paleobiologists now routinely try to determine whether trends arepassive or driven, though this work can sometimes encounterunderdetermination issues (Turner 2009). Body size has received a lotof empirical attention, in part because it is easy to estimate fromfragmentary fossil remains, and partly for the simple reason that itis a trait that every animal has.

In addition to its role in the development of macroevolutionary theoryand its influence on scientific research, the passive/drivendistinction raises a number of philosophical issues. As noted already,the notion of a passive trend links up with a traditional set ofquestions about the role of chance or randomness in evolution(Millstein 2000). Indeed, the passive/driven distinction looks a lotlike the distinction between random genetic drift and selection inpopulation genetics. Drift could be construed as a passive trend ingene frequencies. Grantham (1999) argues that the notion of a passivetrend points toward a certain sort of explanatory pluralism: a trendthat is best explained as a stochastic phenomenon at the macro-levelcould receive a more deterministic explanation at a lower level, andthose explanations need not conflict. Turner (2014) suggests that thenotion of a passive trend poses a challenge to popular interventionistideas about causal explanation. Explaining a trend by showing that itis passive could turn out to be a non-causal explanatory strategy.

One issue that remains under-explored, philosophically, is how thedistinction between passive vs. driven larger-scale trends mightintersect with philosophical disagreements about whether neo-Darwinianevolutionary theory is best understood as a causal theory (a viewgiven its canonical formulation by Sober 1984) or a statistical theory(Matthen and Ariew 2002; Walsh, Lewens, and Ariew 2002). There may besome question about whether it is best to understand thepassive/driven distinction as a distinction between trends generatedby different causal mechanisms, vs. a distinction between differentsorts of statistical phenomena.

8. The Zero-Force Evolutionary Law

Many paleobiologists have assumed that driven trends must be driven bynatural selection. For example, if body size increase is driven,surely that means that larger body size has conferred some survival orreproductive advantage (Hone & Benton 2005). McShea (2005) andMcShea and Brandon (2010) have forcefully challenged that assumption.McShea (2005) argues that increase in structural complexity is adriven trend, but one that need not be driven by selection. That isbecause increasing structural complexity might just be the zero-forcecondition for evolving systems, an idea that McShea and Brandon (2010)develop in great detail. McShea and Brandon’s zero-forceevolutionary law—orZFEL—has elicited criticalresponses from other philosophers (Barrett, et al. 2012). Whereastraditional evolutionary thinkers, from Darwin on down, assumed thatnatural selection would have to explain complexity increase, McSheaand Brandon see complexity increase as the default expectation, andthey invoke selection to explain deviations from that defaultexpectation—such as cases of eye loss in cave-dwelling species.They treat eye loss in cave dwellers as a case where natural selectionfavors a reduction in structural complexity. Alternatively,O’Malley, Wideman, & Ruiz-Trillo (2016) consider thepotentially wide-ranging and macroevolutionary role of simplificationin conferring “adaptive” advantage.

McShea and Brandon also argue that theZFEL has the virtue ofgiving a unified account of both micro- and macro-evolutionaryphenomena. They invoke theZFEL to explain a wide variety ofdifferent patterns, ranging from complexity increase in the history ofanimal life, as represented by the fossil record, to increasinggenomic heterogeneity over time. Whatever one thinks about their claimthat theZFEL is “biology’s first law”,their project raises a question worth exploring: is it reasonable tothink that explanations of macroevolutionary patterns should takefundamentally the same form as explanations of patterns at vastlydifferent spatiotemporal scales? How much does scale matter?

TheZFEL also raises some fundamental philosophical questionsabout the structure of evolutionary theory. How, for example, shouldwe determine what counts as the default expectation, or zero-forcecondition, for an evolutionary system (Gouvêa 2015)? Is thereany way to distinguish empirically between theories that disagreeabout the default expectation? Is there even a fact of the matterabout what the default expectation should be? These questions aboutdefault expectations for evolutionary systems may link up with largerquestions about the function of null hypotheses in scientificreasoning. These questions about default assumptions inmacroevolutionary theory also prompt comparison with questions about,for instance, neutral theory in molecular evolution (Kimura 1983) andin community ecology (Hubbell 2006). For relevant philosophicaldiscussion, see e.g., Dietrich (1994); Bausman and Halina (2018).

9. Major Evolutionary Transitions

If one way to think about the larger picture of the history of life onEarth is to focus on evolutionary trends (size increase, complexityincrease, etc.), another approach is to focus on major transitionalevents in the history of life (Maynard Smith & Szathmáry1995; Szathmáry & Maynard Smith 1995). The point ofdeparture for most work on major transitions has been Szathmáryand Maynard Smith’s (1995: 228) list of game-changingalterations in evolutionary history:

• Replicating molecules to populations of molecules incompartments
• Unlinked replicators to chromosomes
• RNA as gene and enzyme to DNA and protein (genetic code)
• Prokaryotes to eukaryotes
• Asexual clones to sexual populations
• Protists to animals, plants, and fungi (cell differentiation)
• Solitary individuals to colonies (non-reproductive castes)
• Primate societies to human societies (language)

One of Maynard Smith and Szathmáry’s insights is that thevery mechanisms of evolution—the way evolution works—havechanged over the course of evolutionary history (Calcott &Sterelny 2011).

One philosophical concern about the theory of major transitions isthat the original list was something of a mixed bag (McShea &Simpson 2011). For example, some of the transitions, like theevolution of eukaryotes and the evolution of multicellularity, seem toinvolve jumps in part/whole complexity. But others, like the evolutionof sexual reproduction, do not. So it is not clear at all that theseare transitions of the same kind. Indeed, Maynard Smith andSzathmáry suggest that the transitions have three features: (1)changes in the way that information is stored and transmitted; (2) newdivision of labor; and (3) individuals that could once reproduceindependently can no longer do so after the transition. Criterion (3),in particular, links the issue of major transitions to the largerdebate about the nature of biological individuality. One potentialproblem, though, is that not all the transitions on the original listsatisfy all three criteria. For example, the shift from primatesocieties to human societies clearly involves (1) and (2), but not(3). To complicate things further, Szathmáry (2015) revisitsthe original list of transitions and proposes some changes.

Related to this worry that the major transitions do not comprise aunified type of evolutionary event, some theorists (e.g.,O’Malley 2014) argue that other significant events should becount as major transitions. O’Malley & Powell (2016) thinkthat the biological oxygenation of Earth is a major transition, andCurrie (2019) makes the case for treating mass extinctions as majortransitions. Another potential candidate for major evolutionarytransition is the Angiosperm Terrestrial Revolution, a shift indomination from gymnosperms to angiosperms across many terrestrialecosystems. This shift corresponded with a surge in insectdiversification, as might be expected given the relationship—onewith both metabolic and reproductive import—between pollinatorsand flowering plants. Recent discussion of this transition has beenthe locus for another iteration of the philosophically significantdebate about what can be inferred from correlation or co-location inthe fossil record about potential causal relationships or drivers ofmajor evolutionary change (Jouault, Condamine, Legendre, &Perrichot 2024a, 2024b; Vermeij 2024).

A second philosophical concern is that Maynard Smith andSzathmáry’s original idea looks a bit teleological, andin a way that likely raises the hackles of some scientists andphilosophers (e.g., Dawkins 1986; Dennett 1995). The transitions inSzathmáry and Maynard Smith’s (1995) list all seem likenecessary steps on the way to the evolution of humans, and cynically,one might wonder if their relation to the later appearance of humansis what makes the transitions “major”. Relatedly, onemight wonder if the idea of major transitions comes a little too closeto reviving pre-evolutionary thinking about the great chain of being.This also connects with traditional questions about evolutionaryprogress (Ruse 1997). While some of the concepts surveyed in thisentry—especially passive trends and historicalcontingency—might seem like threats to the belief inevolutionary progress, the theory of major transitions coheres wellwith progressivist views about evolution.

In spite of these potential concerns, however, the major transitionsreally are historical events that pose distinct explanatorychallenges. Arguably, our picture of macroevolution will not beentirely complete without tackling some of those challenges.

It is noteworthy that neither Maynard Smith nor Szathmáry arepaleontologists, and so there may be a bit of a disconnect betweentheir original proposal and the way that many paleontologists havecome to think about major transitions. Several of the transitions fromthe original list must have occurred way back in the deep past; thefossil record is not likely to give us much insight into the origin ofeukaryotes, for instance. Molecular tools may be able to shed somelight on macroevolutionary queries about the deep past, however(O’Malley 2016). And paleontologists sometimes take a muchbroader (and looser) view of major transitions, such that any majorevolutionary novelty would count as a major transition. For instance,the evolution of placentation in mammals, or the evolution of feathersin archosaurs, might count as major transitions on this broader view.Currie (2019) draws a distinction between theory-driven approaches tomajor transitions and phenomena-driven approaches. Paleontologiststend to fall in the latter camp, since they usually begin by thinkingabout major morphological or phenotypic changes that show up in thefossil record.

10. The (Dis)unity of Macroevolutionary Theory

One significant and remaining challenge in the philosophy ofmacroevolution is to work out how the various ideas aboutmacroevolution surveyed above may or may not fit together. Whatexactly is the relationship between punctuated equilibria and speciesselection? Or between PE and the major transitions? Or between speciesselection and historical contingency? Or between contingency and themajor transitions? Do (some, all of) these ideas weave together toform a unified picture of evolution at large scales? Or do theyrepresent something more like a heterogeneous grab bag of scientificideas that are more or less useful in different contexts? A furtherquestion is how the relative (dis)unity of macroevolutionary theorymight bear on the longstanding question of how macroevolution isrelated to microevolution. One recent proposal linking micro- andmacroevolution suggests a positive scaling relationship betweenevolutionary divergence and evolvability, likely resulting fromgenetic constraints limiting trait fluctuation (Holstad et alia 2024).Evolvability is generally understood as propensity to evolve, and itis an attribute which can be found at any level within the biologicalhierarchy (Jablonski 2022). There is a growing body of work on thistopic; see Hansen et alia (2022) for an introduction to the field.There is also a growing body of work on the macroevolutionarysignificance of ecological interactions, exploring the ability ofbiotic interactions to influence evolution at the macro level; seeHembry and Weber (2020) for an introduction to work in this field.Work on this topic introduces the possibility of an integrated,mezzo-incorporated approach to evolutionary trends, one whichconsiders biotic and other ecological factors in addition to the usualmicro (organism) and macro (species) foci.

Finally, the investigation of macroevolution challenges the purporteddistinction between the experimental and historical sciences (Sober1993; Cleland 2001). Investigation in this domain is historical andoften emphasizes the deep past, to be sure, but it also involvestheoretical and prospective modeling, making predictions about whatthe models will or will not find, making predictions about whatpaleontologists will or will not find, running bottle and other kindsof experiments, simulations, and more. The scientific study ofmacroevolution is a philosophically rich and challenging field withmany deposits to mine.

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