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Transactions of the Royal Society of South Australia (2013), 137(2): 102–148

ORGANIC EVOLUTION IN DEEP TIME: CHARLES DARWIN

AND THE FOSSIL RECORD

bRIAN MCGOWRAN

School of Earth and Environmental Sciences

The University of Adelaide, Adelaide SA 5005, Australia

brian.mcgowran@adelaide.edu.au

Abstract

The heart and soul of geology are to be found in rock relationships and earth history. The fossil record was central and critical to geology emerging as geohistory, the first historical science, from the speculative geotheories of the 18th Century. The key figure was Cuvier. In the process of shaping geohistory, Cuvier and palaeontology produced the second historical science, namely biohistory, including faunal and floral succession in deep time; and biohistory and geohistory have been intertwined for two centuries.

lamarck kept alive the venerable theory of organic change, but evolution as heuristic scientific theory was stumbling. Even so, the fossil-based geological time scale was constructed in the six decades between Cuvier nailing bioextinction and Darwin nailing biospeciation. by about 1830, French molluscan palaeontology was building Tertiary stratigraphic succession, correlation and age determination in the palaeontological synthesis, i.e., biostratigraphy. Palaeontology revealed ancient and exotic life in a deep-time panorama of succession punctuated by extinctions and demanding explanation, but it contributed little to theory of evolutionary processes.

Darwin’s world was lyell’s gradualist world and his appreciation of environmental change as an evolutionary forcing factor lessened as competition came to dominate his thinking. Darwin’s Darwinism comprised five theories. Two were historical theories (the world and its biospecies change in deep time; and common descent in branching evolution produces the tree of life) and both were widely accepted by the generation after Darwin. The other three were causal or nomothetic theories (respectively speciation; gradual change not saltational; and variational change by natural and sexual selection) and they were accepted only in the 20th Century. For its importance to our culture, Darwin’s historicist worldview, in the face of entrenched, ahistorical opinion as to what science really is, outweighs disputes about the importance of selection.

As stratigraphy and palaeontology went global and highly successful on most criteria, their evolutionary direction went ‘anti-Darwinian’ in the later 19th Century, towards such theories as orthogenesis, saltationism and a resurgent ‘Neo-lamarckism’, mostly in Hyatt, Cope and Osborn in North America. This cluster of trends culminated a second time in the 1930s–1940s, in the macromutational typostrophism of the German synthesis, dominated by Schindewolf.

Meanwhile there was a thin red line of Darwinian palaeontology down those decades from the 1860s to the 1920s. When population genetics emerged from decades of its own anti-Darwinism the Modern Synthesis was forged between natural history, genetics and palaeontology, the latter especially embodied by Simpson’s macroevolution. Darwin’s three causal theories came into their own in the 1930s–1950s in completing the Darwinian Revolution—or, as I prefer, installing the Darwinian Restoration. However, the Restoration was dominated by variational evolution, whilst practitioners of embryology and morphology, in the transformational mode of evolution, felt excluded.

The roots of modern palaeobiology are firmly in the Darwinian Restoration and Simpsonian palaeontology and macroevolution. Modern palaeobiology (i) is thoroughly Darwinian in its historicism and variational evolution but (ii) is beyond Darwin in becoming pervasively hierarchical whilst (iii) reconciling with elements of the German Synthesis through collaboration with developmental genetics in evo-devo. Also (iv) we have gone beyond Darwin (and Simpson) primarily in the rise of micropalaeontology with its untold millions of specimens and in enormous progress in chronologically resolving and reconstructing bioevents and environmental shifts in

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the geological past. And (v) the tree of life is underlain by an anastomosing web of life. Deep-time palaeobiology becomes more autonomous as major soluble problems arise from the fossil record, utterly beyond the reach of shallow-time neontology.

Securely embedded in thriving research programmes recovering the recorded history of life on earth, Darwinism lives!

Introduction

Tyrants and popes, populists and thought police have always realized that it is essential to own the historical narrative of the tribe, the society and the culture, but it is one of the greatest triumphs of the Western cultural tradition to have expanded that history back beyond the civilizations and into human prehistory, into biohistory and into geohistory in the deepest geological past of this planet. Darwin’s pre-eminence in that enterprise is beyond challenge. However, it is under-appreciated that Darwin inherited the first historical science, the still quite young geology, and that geology was intricately intertwined from the outset with another recently christened and historical science, biology. Darwin’s first scientific publications were geological and the awards made by his peers were for his geology, not his biology, but the accolades are not significant here. What is important is the historicist way of thinking—historicism—that he could absorb so thoroughly only as a geologist-insider and go on to exploit, uniquely, in his biology.

Figure 1. Four figures pivotal to any narrative of evolution and the fossil record—of geohistory and biohistory. (From left) Chrétien Frédéric Dagobert Cuvier (‘Georges’) (1769–1832) (Muséum d’Histoire Naturelle) established historical science qua science in the form of geohistory and biohistory, advances easily outweighing his more frequently mentioned catastrophist and anti-evolutionist stances (the inherently unstable taxa in the evolutionary theory of the times would have rendered impossible a meaningful fossil succession and biostratigraphy). Rudwick (see text) linked Charles lyell (1797–1875) (University of Adelaide library) with Cuvier as accomplishing Tertiary biostratigraphy and the palaeontological revolution, less for the threadbare mantra of uniformitarianism, ‘the present is the key to the past’. For Charles Robert Darwin (1809–1882) (Cambridge University library), all the exquisite design and adaptation in the organic world attributed to a Designer was explained instead by historicist adaptation and natural selection, the latter meanwhile explaining the fossil record and extinction, a body of knowledge entirely refractory to all scenarios for creationist Design. George Gaylord Simpson (1902–1984) (US Army Major in 1944; léo F. laporte) put palaeontology in the Darwinian restoration, reconciling it with variational evolution and rejecting goal-directedness and other excrescences of transformational evolution.

It was an anti-Darwinist and neo-lamarckian, so help me, who wrote this on ‘the conception of the organism as an historical being’:

We have seen this thought expressed with the utmost clearness by Darwin himself. In his eyes the structure and activity of the living being were a heritage from a remote past, the organism was a living record of the achievements

of its whole ancestral line. What a light this conception threw upon all biology! (Russell, 1915, p. 308). This essay discusses the place of fossils and palaeontology in the realization of biohistory and organic evolution. What did palaeontology do for Darwin and what did Darwin do for palaeontology? The short answers are that palaeontology bequeathed him the first truly historical science and his first personal

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experience of same, it supplied him a deep-time geological time scale and it gave him his invaluable historicism; and in return palaeontology received its comprehensive intellectual frame and context for all subsequent research programs.

The half-century or so of earth-and-life science before Darwin embarked on the Beagle in 1831 has received less scrutiny than has the ensuing Darwinian half-century. However, there has been substantial recent advance capped in two magnificent volumes by M.J.S. Rudwick (2005, 2008). The first, Bursting the limits of time: The reconstruction of geohistory in the age of revolution (blT), demonstrates the crucial innovations of Georges Cuvier during a time of great intellectual ferment and progress in the earth and life sciences. The second volume, Worlds before Adam: the reconstruction of geohistory in the age of reform (WbA), does likewise for Charles lyell (somewhat to its author’s surprise and, for that matter, to mine) as the sciences progressed. The volumes together give us an unprecedented portrait of the milieu that Darwin’s generation inherited, the rise of earth and life history named felicitously by Rudwick the Cuviero-Lyellian Revolution. (A reviewer preferred the simpler but too restrictive Geohistory revolution.) (Fig. 1.)

Table 1. Fossil record and evolution: seven advances in two centuries’ progress

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1980s–2000sVII

1980s–2000sVI

1960s–1970sV

1930s–1940sIV

Geohistory and biohistory highly integrated and multi- disciplinary in deep-time biospheric evolution Molecular clock, developmental genetics,

phyletic reconstruction, speciation Oceanfloor spreading and plate tectonics

Darwinian Restoration (Darwinian Revolution II) The Modern Synthesis: reconciling palaeontology, field biology, and population genetics

(Anti-selectionism discarded as dead and rotten science)

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Two centuries’ worth of progress: seven major advances and turning points (“revolutions”) in earth and life history and pertaining to their development. Strands of evolution-denial (and deep-time-denial) are labelled simply as dead science (Kitcher, 2009). See discussion in text.

Accepting Ernst Mayr’s argument that the Darwinian Revolution has two parts in separate centuries, we have a convenient structure for this essay—the Cuviero-lyellian Revolution, the Darwinian Revolution Mark I and the Darwinian Revolution Mark II, commonly known as the Modern Synthesis or the Synthetic Theory but better called the Darwinian Restoration (Table 1, Fig. 2, 3). Recent triumphalism distinguishes the 1970s-on as The Paleobiological Revolution (Sepkoski and Ruse, 2009), which this essay finds unconvincing. I note too that Hodge (2005) objects to the exuberant use of the terms ‘evolution’ and ‘revolution’ and that Cain (2009) would abandon the term ‘the evolutionary synthesis’.

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Two flow charts (Fig. 2, 3) reconstruct the geological and palaeontological strands of this narrative of historical science. Rudwick’s central theme is the rise and flowering of historicism, extending back to the advent of Cuvier the importance of that theme long advocated in organic evolution and Darwinism. Cuvier not only was the first great historical geologist and biologist and founder of vertebrate palaeontology; he also adopted the word ‘geology’ when satisfied that it was free of the 18th century genre ‘geotheory’ where De luc had first used it. Where Rudwick used ‘geohistory’ (whilst making it abundantly clear that fossils were the driving force behind its origins and subsequent exuberant flourishing), this essay uses ‘geohistory’ and ‘biohistory’ (Fig. 2).

Figure 2. The first of two charts centred on the fossil record and together comprising a visual summary of this essay. This one highlights outstanding names, titles and themes of evolution. biohistory and organic evolution through time (running upwards and unscaled) since the Enlightenment are depicted to contrast two modes of evolution, variational to the left and transformational to the right, all passing through the sphinctral unifying force of On the Origin of Species... only to diverge again through most of the next hundred years. In Natural History as distinct from Natural Philosophy (see Fig. 3) I depict streams represented by Cuvier and Goethe plus von baer; lamarck is set apart as the only major figure not an agnostic about or an uncompromising opponent of the evolution theory. lamarck had affinity with the transformational anatomy and embryology of the idealistic morphologists and von baer and Owen, but the essential difference was that he was an accomplished systematist: in pioneering Cenozoic molluscan systematics he was antecedent to the French palaeontologists Deshayes and d’Orbigny who established biostratigraphy. Somewhat likewise, Haeckel had a lot to say about transformation and recapitulation, but more significant was his immediate grasp of Darwin’s variational theory and his own vast contribution to radiolarian and animal systematics (Gliboff, 2008; Richards, 2008). Sources are in the text. The central objective of this chart is to contrast down the decades the two cultures of variational evolution and transformational evolution: on the left, populational and biodiversity, and all teleological theories of direction, internal drive and goal-direction rejected; on the right, embryology and morphology, and those teleological theories tending to recur. This is a deeper cleavage than are the traditional evolution/antievolution or catastrophism/uniformitarianism.

(Geohistory and biohistory have sustained close ties for more than two centuries and the ties will only strengthen as we come to appreciate the role of the biosphere not merely as a passive reactor but as an active player in environmental change (e.g., Kasting, 2010). I like ‘biogeohistory’ for that reason and indeed chapter eight of McGowran (2005) is entitled ‘On biostratigraphy and biogeohistory’. However, adverse reception of the term convinces me reluctantly to revert to ‘geohistory and biohistory’ as used elegantly by Simpson (1970).)

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Figure 3. The second half of the visual summary of this essay: palaeontology and the fossil record pertaining to biohistory and geohistory, compiled from many sources cited in the text. The rake structure is intended to convey the notion of innovation and expansion, but some inputs from geology and neontology are omitted. The narrative proceeds up-chart but there is no linear time scale. The palaeontological synthesis is the core of the Cuviero-lyellian Revolution (Rudwick, 2005, 2008). The crosses mark ‘extinctions’ of top-down geotheories some of which, however, are capable of lazarus-comebacks. Thus lyell’s cyclism resurrects Hutton’s ahistorical steady-statism; and idealistic morphology and transformational evolution reappeared after Cuvier, after Darwin, and after the Restoration. The five Darwinian theories are after Mayr (1982 and references); the two historical theories were accepted within a decade or so but acceptance of the three causal theories was deferred, to be fulfilled only in the Restoration. The ‘German Synthesis’ is sensu Princehouse (2009), not sensu Reif (see text). Autonomous Palaeobiology has also been called the Paleobiological Revolution (Sepkoski & Ruse, 2009); certainly the label marks an exuberant efflorescence with roots in the Restoration, in fusion with molecular and developmental genetics (evo-devo), and in the rise of micropalaeontology and palaeoceanography providing the context of environmental shifts. Ghiselin (2004–2006) would, reasonably, add the new ontology of species-as-individual. ‘Autonomous’, because the uniformitarian notion that palaeontology is merely equivalent to watered-down neontology (i.e., modern and ahistorical biology) has been an exhausted cliché long since.

Evolution and evolutionism

In the 18th century and much of the 19th century ‘evolution’ referred to what is now ‘ontogeny’ i.e. the development of an individual organism, not the emerging of an inclusive or collective entity such as a species or higher taxon. It is necessary to distinguish three modern meanings of the word ‘evolution’ as in the three principle theories (or theory-groups) of evolutionism: Saltational, Transformational, and Variational (Table 2). The analogy of individual development with taxic evolution has been remarkably persistent and highly significant over almost two centuries. If ‘Darwinian’ refers to the tree of life generated by variational evolution, then ‘anti-Darwinian’ refers, not to the long-irrelevant and now-cynical denial of evolution, but to authentically scientific challenges from the transformational and saltationist strands. The most commonly used term for these is ‘(neo)lamarckism’, popularly meaning the inheriting of use and disuse rather than lamarck’s theory of evolution (below and Fig. 4).

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ORGANIC EVOlUTION IN DEEP TIME: CHARlES DARWIN AND THE FOSSIl RECORD It is also useful to distinguish the three aspects of organic evolution, namely:

(i) Did evolution actually happen? Since the passing of Darwin’s generation this question about the biosphere has been a non-question, therefore of no scientific interest. It is dead science (Fig. 1). Accepting that deep time in earth history means billions of years has taken longer, as too has accepting the highly mobile and reworking of the earth’s crust by internal heating.

(ii) What has been the course of evolution? Reconstructing biohistory on the template of geohistory comprises a series of research programs in disciplines ranging from geology and palaeontology to biosystematics to molecular genetics.

(iii) How does evolution happen? This too is a series of research programs with much the same range but different emphases.

Numerous familiar dualisms refer to query #(ii) vis-à-vis query #(iii), such as ultimate/proximate, functional/historical, contingent (or configurational)/immanent, or pattern/process. Note too the recurring styles of thought, such as catastrophist/uniformitarian and gradualist/punctuationalist. Useful rhetorical and pigeonholing devices as they may be, these polarities too easily tend to obfuscate instead of clarifying. I have the same opinion about the widespread use of ‘lamarckian’ vs ‘Darwinian’, but I confess to liking ‘time’s arrow’ versus ‘time’s cycle’ (Gould, 1987). A useful if pernicious dualism was expressed in the late 18thC as ‘natural history’/‘natural philosophy’ (Fig. 2, 3).

What is ‘Darwinism’? Meanings range from extremely broad and unhelpful, such as anything to do with organic change at geological time scales, to the narrow and more useful variational evolution mostly by natural selection. This was the position of one of Darwin’s most comprehending defenders and most articulate promoters through the somewhat dismal ‘anti-Darwinist’ years, Ernst Haeckel: ‘... the Theory of Selection, or breeding, might be justly called Darwinism, being that portion of the Theory of Development which shows us in what way and why the different species of organisms have developed from those simplest primary forms’ (1876, p. 150; emphasis his).

I would add though that Darwinism should evoke some amalgam of variational evolution with a thoroughgoing historicism. Anti-Darwinian in evolutionary or more broadly scientific discourse has tended to mean one or more of three things (none uncontested). (i) One is to challenge the extrapolating of intraspecific processes (microevolution) seamlessly to supra-specific levels (macroevolution). (ii) In a second meaning, structure or morphology (and in the later 20th century, evolutionary developmental biology, or evo-devo) challenge the dominance of variational change including population genetics. (iii) Third is promoting punctuational change over gradual change, be it biological saltation or abrupt environmental shift.

History and historicism

Palaeontology and its discipline biostratigraphy are profoundly historical sciences (McGowran, 2005, Ch. 8). As Simpson (e.g., 1959b) put it, the cosmos is broadly twofold: it has the immanent characteristics inherent in the nature of matter-energy, the stuff of the physical sciences and most experimental biology, and its other characteristic is configurational, concerning structure and organization (‘contingent’ is not synonymous with configurational but has displaced that term). Within biology, Mayr (1958) long distinguished between ahistorical (functional) and historical biology and railed against the mindset for which, for example, a frog in a swamp and a fossil frog in a mudstone are natural history whilst a frog in a laboratory blender is real science. All palaeontologists are familiar with the roll-call of the physicists: John Herschel, for whom natural selection was the law of the higgledy-piggledy; the intellectual conservatism and arrogance of William Thomson (lord Kelvin) who granted the geologists less and less time (down to perhaps 20 Myrs) for the age of the earth as he himself aged; Ernest Rutherford, for whom all historical science was stamp-collecting; Fred Hoyle, whose notion of natural selection was as a tornado assembling an airliner in a junkyard; luis Alvarez, lead author of the bolide hypothesis of mass extinction based on the Iridium spike at the end of the Mesozoic Era, with his notorious contempt for the palaeontology that discovered then focused the biology and chronology of the problem in the first place. We would like to think that we have moved on, so that it

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Table 2. The three principle theories of evolutionism

Theories of evolutionism: Saltational, transformational and variational evolution, indebted to levins and lewontin (1985).

‘now seems natural’ (Sober, 2009) to recognize that science is of two types. Nomothetic science aims to discover laws and uses historical-type data as a means to that end. Historical science aims to reconstruct particular or specific histories and uses information about laws as a means to that end. Nor is it sufficient to say that physics and chemistry are nomothetic and palaeontology and geology are historical. Indeed, Gould (1980) promoted palaeontology as nomothetic science.

At any rate, evolutionary biology is historical science! Grasping this point at fundamental or philosophical levels entailed contrasting pre-Darwinian with Darwinian change. For Mayr (1964, 1982, 1988) it was the shift from essentialism, the Platonic Idea, where variation was trivial, species were natural kinds or classes with their own essences and essentially similar to, say, the chemical elements, to the more Aristotelian

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populational thinking where Darwin showed variation to be all-important. In his customarily crystallizing and confronting way Mayr lumped philosophers, theologians and physical scientists as Platonic essentialists, the reactionaries whose overthrow by the fact and theory of evolution made Darwin such a successful intellectual revolutionary (Mayr, 1983). Pre-Darwinian systematists were typologists, post-Darwinian systematists were populationists. Hodge and Radick (2009) attributed prime responsibility to the American pragmatist philosopher John Dewey for this portrait of 2000 years of Platonic-Aristotelian doctrine being overthrown by Darwin, freeing us from the ‘tyranny of absolutes’ (the one true picture of the world, the one true morality, the ultimate nature of things)—a portrait that they aimed to ‘correct out of existence’. Winsor (2003), levit and Meister (2005) and Wilkins (2009) argued that the pre-Darwinians were not a solidly essentialist bloc; even so, much of the stark contrast in mindset remains.

The German palaeontologist H.G. bronn in the 1850–1860s strongly promoted the study of life as a proper intellectual pursuit—Wissenschaft—distinct both from natural theology and descriptive natural history (Gliboff, 2008). This is important, for Gliboff has shown that bronn was as historicist as Darwin himself in ascribing organic change to environmental change, not to the internal laws, or the developmental plans, or the archetypes of the transformationists (indeed, Gliboff suggested that bronn was the more evolutionary, in that his earth was evolving, whilst Darwin was still in some thrall to lyell’s steady-statism).

Ghiselin (1969) wrote what for me is the most significant book about Darwin, meanwhile launching the ontological thesis (1971, 1974, 1997), now (in his words) the ‘new evolutionary ontology’ (2002, 2005a, b), that the biospecies is an individual, not a class, a thesis also owing much to Hull (1988, 1989) and resonating with Mayr’s distinguishing the species taxon from the species category. (Perhaps ‘historical entity’ is a little more apposite than ‘individual’ (Sober, 2009).) The species is a real chunk of a real genealogical lineage awaiting discovery, not a convenient accounting procedure or logical trick. Darwin did more than establish the reality of evolution once and for all—he showed us a new way of thinking—thinking historically, thinking like an evolutionist. In Ghiselin’s words (2005a, p. 127) he established:

The priority of the concrete over the abstract. So far as metaphysics goes, it was an intellectual revolution of the first magnitude. Much as Copernicus had moved the sun to the center of the physical world, Darwin moved concrete particular things to the center of the metaphysical world. Individuals, not classes, were the ultimate reality: The individual organisms that struggle for existence and also the individual species that speciate and are transformed through time. In taxonomy, this change in perspective has often been explicated in terms of the rejection of essentialism, and that is surely a part of it. Taxonomic groups are individuals and individuals have neither the essences nor the intellectual baggage that goes with them. Their component organisms are also individuals and the fact that it is they that differ from one another and reproduce differentially means that they play a crucial role in evolutionary processes. Variation is not deviation from a norm, but a reflection of the underlying causality.

Historicism acquired a bad name when Popper (1957) shifted the meaning pejoratively to grand-scale, teleological theorizing (McGowran, 2005, Ch. 8). However, there should be much more historicism in epistemology including science within itself and within broader contexts (Kadvany, 2001; Munz, 2004; Carroll, 2004; Kitcher, 1993, 2009).

The Cuviero-Lyellian Revolution

There is no historicism in [The Reverend Dr William Paley’s] Moral and Political Philosophy and no geology in his Natural History ... ; and the two books are good illustrations that a sense of history was as uncharacteristic of utilitarian political philosophy as a sense of evolution was of eighteenth-century natural philosophy.

(Gillispie, 1951).

How was it not seen that the birth of the theory of the earth is due to fossils alone; and that without them we would perhaps never have dreamt that there had been successive epochs, and a series of different operations, in the formation of the globe?

Georges Cuvier, 1812, Preliminary Discourse on the

Revolutions of the Globe, in Rudwick (1997, p. 205).

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Palaeontology is an ancient proto-subject but its flourishing was delayed for centuries. Fossils inform us about three matters—ancient environments (hence the possibility of reconstructing environmental change), the geological ages of strata (hence the possibility of any deep-time history at all), and the pathways of the history of life on earth (hence the possibility of a testable genealogy). As for major environmental change in the deep past, leonardo and numerous if unknown others puzzled over the ‘[marine] fossils on the mountain top’, many meters above and many kilometres inland from the modern sea. Did the sea fall, or did the earth rise, or both? Such matters are covered in a beautifully told account of sedimentary geology in the Mediterranean region (Fischer & Garrison, 2009).

As for the age of rocks, this possibility was enunciated by the 17th century scientist Robert Hooke (Jardine, 2003): However trivial a thing a rotten shell may appear to some, yet these monuments of nature are more certain tokens of antiquity than coins or medals ... and though it must be granted that it is very difficult to read them and to raise a chronology out of them, and to state the intervals of the time wherein such or such catastrophes and mutations

have happened, yet it is not impossible... (Hooke, 1668, 1705 as quoted in lyell, 1832; my emphasis). Hooke also referred to the shells known from high in the mountains, perhaps raised by earthquakes; he considered the question of organisms going extinct, even at different times and places; and the discovery of giant ammonites made it ‘necessary to suppose that England once lay under the sea within the torrid zone’. There was more than enough here to stimulate research programs in earth and life history, and had that happened it might well have been the beginnings of historical geology (Oldroyd, 1996). but the necessary research did not happen; nor did the geotheories of the Enlightenment, meaning the speculative, top-down theories of the earth also known as cosmogenies, trigger a recognizably modern geology. Instead, we have to look to the early describers of stratal succession in the field, particularly Giovanni Arduino (1714–1795) and his celebrated 1758 cross-section in the Vicentian region in Italy (Vai, 2007), and to the economic imperatives of minerals exploration and mining operations for the rise of geognosy (see below).

Late Eighteenth Century view of the earth

Enlightenment knowledge was twofold, literary and humanist-philosophical, the latter concerning the natural world including human nature (Fig. 3). Natural world enquiry in turn deconstructs into two categories, natural history, concerning the ordering and arranging of real-world diversity (organic and inorganic: mineral species as well as plant and animal species), and natural philosophy, investigating regularities and their causes and seeking natural laws.

Natural History pertaining to the earth included the materials genre mineralogy (including fossils), the spatial discipline physical geography, and geognosy, the analysis of rock structures and rock formations in three dimensions (Rudwick, 2005). biological enquiries fell more or less naturally into two streams, organic structure (morphology) and its individual development (embryology) on the one hand and describing and ordering biodiversity, i.e. systematics and taxonomy, on the other.

The Natural Philosophy discipline was earth physics, seeking the processes, the causal origins of the phenomena and entities of the natural history disciplines, from earthquakes and volcanism to mineral veins, from mountain building to the accumulation and consolidation of fossils. The category antecedent to earth physics was conjectural geotheory (largely synonymous with the above-mentioned cosmogeny) referring to speculative, a priori and deductive theories of the earth—hypothetico-deductive theory preceded field evidence. The best- known geotheorists of the late 18th century were the Frenchman De luc and the Scot James Hutton. Hutton, celebrated down the generations of Anglophonic students for his ‘rock cycle’ (erosion, sedimentation, deformation and metamorphism, igneous intrusion), was strictly steady-statist and tacitly eternalist, well known for summoning up a conjecture about the earth and scouring Scotland for evidence to support it (e.g. granites as igneous intrusions). He was fundamentally ahistorical in his outlook (e.g., Hallam, 1989; Gould, 1987) but he appreciated the biosphere: for example, the presence of modern-looking marine shells high in the mountains gave him the crustal uplift necessary to keep the rock cycle operating. De luc in contrast was strongly directional and the most geohistorically minded of the 18th century geotheorists and he coined the word geology (see below).

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by the end of the 18th century geognostic savants were comfortable with the fundamentals of rock relationships in the field—faults, folds, igneous rocks displaying intrusive and crosscutting configurations, angular unconformities, and stratigraphic superposition and the beginnings of what would be termed stratigraphic succession. The English geognost William Smith was demonstrating the power of fossils in characterizing and identifying the newly appreciated entity, the stratigraphic formation (laudan, 1987) (meanwhile giving us the word ‘stratigraphy’). Progress was stimulated by economic and industrial demands—engineering works, groundwater and springs, minerals and coal exploration and exploitation— and the objective was ‘structural’ (a much broader term then than now), solving the patterns of rock types in the field and in the subsurface. Just as the deep space of the heavens (laplace, William Herschel) was commonplace, so too was the notion of a vast time scale.

From Geognosy and Geotheory to Geohistory

Rudwick (2005) has sustained the consistent distinction between (the preceding) structural and temporal but ahistorical geognosy plus the deductive, also ahistorical geotheory, and (the succeeding) evidence-based geohistory. Geognosy was strong on the configurations above/below and higher/lower, whereas geohistory’s strength resided preeminently in its configuration of younger/older. Several conjectural geotheories were popular in the 1780s–1790s, whereas attempts to reconstruct the past from its discovered concrete remains were rare and not popular, but De luc’s geohistorical mindset radically undermined the genre of geotheory by beginning its transformation into geohistory.

That change of mindset from geotheory to geohistory developed for one reason above all others, the study of fossils and especially the distribution of fossil species in space and time.

but historicism arose in a pertinent cultural context:

Oldroyd (1979) made a strong case for the emergence of historical geology as an integral part of a major intellectual shift at the end of the 18th century. Scholars became more interested in the human past, the prehistorical past and the geological past, in contrast to the philosophers and scientists of the immediately preceding Enlightenment, who were thoroughly ahistorical in their concern with how the universe and its components, from atom to organism,

fit together and function ...(McGowran, 2005). The disciplines linking biogeohistory with human history—prehistory and prehistoric archaeology—arose at this time when prehistory as a respectable, evidence-based discipline replaced the legendary prehistory of earlier times. Prehistory slotted neatly, time-wise, between the emerging geohistory and human history, itself stimulated by such excavations as Pompeii and Herculaneum. In due course Christian Thomsen in Copenhagen from 1815 onwards demonstrated archaeological succession and the three ages of man, the Stone, bronze, and Iron Ages (Daniel, 1962).

Taxonomy influenced the shift from mineralogy-dominated earth science to strata-dominated (laudan 1987, 1989). Attempts to employ linnaean principles in mineralogy collapsed and attention moved from minerals to rocks (the vast bulk of which were composed of only a small number of the known minerals), and A.G. Werner replaced mineralogy with mode and time of formation as the essential characteristics of rocks. Hence the rise of the ‘formation’, as the basic systematic unit, explained by Cuvier and brongniart as ‘a group of beds of the same or different nature, but formed at the same epoch’ (laudan, 1989). Hence too a new emphasis on superposition— position in the succession of strata—as the prime criterion for identification. Formations, individuated, historical entities, catered for those who asked chronological questions (and in due course built the geological time scale). An ahistorical rock and rock-type classification was available to those asking causal questions.

Evolutionism was in the air

If historicism was flourishing and the notion of earth and life history was rising, then surely evolution in the modern sense of the word could not be far away—and indeed there were subsequently claimed ‘forerunners of Darwin’ (Glass et al., 1959). but only lamarck stands out as a visionary evolutionist with strong claims to be the founder of the theory that life on earth has evolved through time (Fig. 4). Thus, in his Philosophie zoologique... : ‘...after a long succession of generations these individuals, originally belonging to one species, become at length transformed into a new species distinct from the first’ (in Mayr, 1972).

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Time

Figure 4. Models of evolution. A Darwinian clade is shown in the traditional or gradualist mode and the punctuated/stasis mode. In lamarck’s alternative scenario there is not the clear sense of literal relatedness in a tree of life, more of analogy with ontogeny. lamarck took the ancient, single linear pattern—the ‘scala naturae’, the Great Chain of being—and reinterpreted it as an inherent pattern followed by organisms in the course of their evolution. Since species are inherently unstable or do not exist in this theory, there are no evolutionary criteria for taxonomy and no entities with which to construct a biostratigraphic succession. See also Tables 1 and 2.

In his sixth decade lamarck was transmuted by administrative decree in the Paris Museum from distinguished botanist to tyro invertebrate zoologist and palaeontologist (e.g., burkhardt, 1970, Mayr, 1972, Corsi, 1988, 2005). Clearly an accomplished naturalist and biotaxonomist, he also had a strong geological background and it so happened that he acquired an entirely new worldview, namely evolutionism, meaning organic change in a gradually but constantly changing world, at that time, the turn of the 18th-19th Centuries. He embarked upon a major monographic study of the rich fossil molluscan assemblages around Paris (Rudwick, 2005, Fig. 7.15)—faunas preserved well enough to be compared and contrasted with analogues among living species. He explained the similarities and differences as due to slow transmutation through geological time—but with no extinction. Mayr (1982) attributed lamarck’s dramatic conversion to his new insights into the molluscan temporal record. Corsi (1988) concluded more cautiously that no single factor brought on the conversion, pointing out that he ‘seemed distinctly reluctant’ to defend his theory with palaeontological arguments and never discussed the importance of the fossils for his theory of transformism; nor are fossils prominent in ‘lamarck’s great change of mind’, 1794–1802 (Hodge, 2008).

lamarck took the classical reconstruction, the scala naturae or Great Chain of being, the concept handed down from Aristotle and the Mediaeval scholastics of a linear progression from the humblest mud, through the plants to the lower animals and so on up to the Supreme being (lovejoy, 1936), and reinterpreted it as an inherent pattern followed by organisms in the course of their evolution (e.g., Simpson, 1959). Since species are inherently unstable or even do not exist in this theory, there are no evolutionary criteria upon which to base a taxonomy and no stable units, bioentities, upon which to found a biostratigraphy—and accordingly a dismally sparse legacy of a great natural scientist. lamarck’s theory of evolution did not impress the two peers who most mattered (and who destroyed the ‘scala naturae’), Georges Cuvier and the embryologist-morphologist Carl von baer, but his evolutionism did survive in the culture of natural history, in a low-profile and marginal way to be sure, but importantly.

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ORGANIC EVOlUTION IN DEEP TIME: CHARlES DARWIN AND THE FOSSIl RECORD Advent of Georges Cuvier

Cuvier founded vertebrate palaeontology, founded comparative invertebrate morphology (in parallel with lamarck) and founded geology as true geohistory (with Alexandre brongniart). He also established three of the basic principles of the history of life:

(i) The pattern of that history is multiple, not linear..

This principle marked the final and successful overthrow of the ‘scala naturae’ Animal taxonomy and classification, chaotic and stagnant if not downright regressive in the 17th-18th century (including the linnaean classification) was spectacularly invigorated by Cuvier’s 1795 Memoir on the classification of the animals named Worms, in which linnaeus’ catch-all Vermes were sorted into no less than six classes. later he classified all animals into four phyla (‘embranchements’), Radiata, Mollusca, Articulata, and Vertebrata, initiating modern comparative invertebrate and vertebrate zoology (Cole, 1944). Carl von baer also rejected the ‘scala naturae’ in recognizing four types of animal, congruent with Cuvier’s embranchements, and issued four laws of development. (von baer and Cuvier were the most uncompromising opponents of evolution theories (e.g., Russell, 1916) and von baer carried this denial forward to include Darwinian natural selection, but he seems not to have been entirely ahistorical or against all notion of evolution (Gliboff, 2008).)

(ii) When ancient strata and their fossils are put in the sequence of time, the fossil faunas are found to have changed markedly in the course of geological time.

For giving close, systematic attention to the distribution of fossils in distinguishing strata—i.e., fulfilling the numerous antecedent suggestions that this would be useful—credit is widely given to the geognost stratigrapher William Smith in England. The Parisians Cuvier and his long-time colleague Alexandre brongniart took such work well beyond Smith’s, by reconstructing an alternation of marine and freshwater environments on the molluscan assemblages and thence a ‘geohistory’ of the Paris region:

They reconstructed a complex story in which the seas had alternated in the deep past with freshwater lakes or lagoons: it was a geohistory as unpredictable and contingent as the turbulent politics they had both lived

through in the past two decades.(Rudwick, 2005, p. 648). (iii) Many or most of the species found as fossils became sequentially extinct.

The fossil faunas of vertebrate animals in the Paris basin were separated abruptly by ‘revolutions’ and in overall aspect were directional towards the present (Fig. 5). by then Cuvier had reinforced the fact of extinction of terrestrial and marine vertebrates beyond argument and it was uncontroversial that trilobites, ammonites and belemnites had gone extinct in ancient seas.

The Tertiary System: Correlation and age determination and synthesis

Stimulated by this work in the Paris basin, the stratigraphers of Europe advanced Tertiary studies on two fronts in the 1820s—(i) more detailed and more accurate description of the local successions of strata and meticulous locating of their contained fossils, and (ii) comparing and ‘correlating’ the various scattered sections in the other sedimentary basins in Europe. The Tertiary record contrasted with the more widespread and continuous, Secondary Chalk in three ways. The first was the more scattered distribution of that record, making correlation all the more urgent since tracing is much less feasible (Fig. 6). The second was that the earth was known already to be cooling towards the present (in modern language and concept: the greenhouse earth was transforming into the icehouse earth) although the notion of the ice ages had to await the 1840s. Cooling forced more equator-to-pole differentiation of floras and faunas and correspondingly more difficulty in using fossils for correlation. The third point is to the Tertiary’s advantage—the faunas and floras were becoming more modern (i.e., like the living) through time, which was powerful evidence that the geohistory and biohistory were directional.

Alexandre brongniart pioneered this enterprise of deconstructing and reconstructing the Tertiary with his extensive travelling, visiting, fieldwork and collecting, less difficult after the Napoleonic wars. The main outcome for our developing insights into earth and life history was the utterly preeminent role of fossils as distributed in space and time. Fossils were the primary evidence (i) for correlation and relative age 113

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Figure 5. The vertebrate faunal succession in the Paris basin, the evidence-based, bottom-up guts of biohistory and geohistory overthrowing the old, speculative, top-down geotheories. The succession is punctuated by turnover or ‘revolutions’ (a stronger term then than now), implying extinctions and restocking from some vague source (not organic evolution).

determination and (ii) for determining ancient environments, from the small and local scale (e.g., distinguishing marine from nonmarine environments) to the large and geographic scale (e.g., the configuration of continents and oceans or clues to global climatic change).

This power of fossils brings with it the problem of competing variables—how do we discriminate the age signal from the environmental signal in a fossil assemblage? (e.g., Rudwick, 2008). The French geologist and palaeontologist Prévost was the first to see this problem clearly and do something about it. The Italian brocchi found that faunas in northern Italy were more similar to those far away in the Paris basin than to those just across the Po Plain in the Sub-Apennines, his explanation being biogeographic, namely a climatic gradient. Prévost proposed instead that the difference was not environmental but in age, the Sub-Apennine faunas being the younger, like the faunas he studied in the Vienna basin, both having more species in common with the modern seas than did lamarck’s faunas from the Paris basin. Prévost also produced in 1827 a superb and ‘astonishingly modern-looking’ reconstruction of Parisian stratigraphy in the next step beyond Cuvier and brongniart (Rudwick, 2008, Fig. 10.5). The strata of the Paris district were regarded as the central reference point and, as Charles lyell discovered in the later 1820s, the conchologists and stratigraphers kept Paris in its leading position as established in the 1790s. The most significant in a glittering array was Gérard-Paul Deshayes, successor to lamarck as the world’s leading conchologist. Deshayes distinguished three natural assemblages of significance in chronological subdivision and correlation of the Tertiary.

Consider Figure 6, sketched by lyell to illustrate the conclusions of the French conchologists as they distinguished successional assemblages in the 1810–1820s. The Tertiary beds of Paris (d) could be compared in their faunas with beds in the london and Hampshire basins in one direction and, by a mixture of stratal superposition and the faunal contents of the strata, with the loire district in the other, where it underlay a distinctly younger fauna (e) which was also brocchi’s in the Sub Apennines, where it underlay in turn a fauna younger still (f), so that the identification of fauna (f) in Hampshire gave the English

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Figure 6. Top: synclines, dips exaggerated, showing the Green Sand and Chalk succeeded by the Tertiaries under london and

Paris (lyell, 1871). lower left: the Parisian faunas had to be traced to Touraine to be shown to be below superpositionally therefore older than the Miocene, thence to the Piedmont and the Pliocene, ditto (right), giving thereby a sense of the missing fossil record in southern England. Sketches by lyell (1833) redrawn in berry (1987, Figs 10 and 11), and used with permission of Wiley- blackwell, Oxford. This was difficult biostratigraphy relying on the superb molluscan palaeontology by Deshayes and colleagues in France, Italy, Germany and Austria.

palaeontologists some notion of what was missing, the ‘hiatus’, in their succession of Tertiary faunas. lyell from about 1829 adopted and actively supported (including financially) this splendid conchological research program in Paris and employed Deshayes’ faunal succession to erect Eocene, Miocene, Pliocene and in due course Pleistocene as the proportion of still-living species present increased. (The Oligocene and Palaeocene divisions came later.) It was the enormous efforts by Deshayes in handling more than 40,000 specimens of molluscs, living and fossil, that enabled the quantitative estimates in the percentage method (Rudwick, 1982) and pioneered the fossil-based geological time scale—interpreting faunas assembled in stratigraphic order.

The palaeontological synthesis and the fossil-based geological timescale: Fossil succession without genealogical evolution

by about 1830 ... the spectacular success of three or four decades of research on fossils had transformed Cuvier’s early demonstration of a single recent organic revolution into a palaeontological synthesis of very wide scope and explanatory power. The geological time-scale was firmly established as almost unimaginably lengthy by the standards of human history, yet documented by an immensely thick succession of slowly deposited strata. The successive formations of strata, and even in some cases individual strata, were clearly characterised by distinctive assemblages of fossil species, which enabled them to be identified and correlated over very wide areas. This correlation proved that in its broader outlines the history of life had been the same in all parts of the world.

(Rudwick, 1972, p. 156; emphasis added). For a puzzled Herbert (1985) this statement about the situation in 1830 was a good summary of the british scene in about 1840, i.e. with a decade’s lag. However, the pioneering ‘biostratigraphic’ work on the Tertiary was centred on Paris by the heirs of Cuvier and lamarck. Some of the divisions of the modern geological timescale date back to the 1820s (and the Tertiary and Quaternary are Neptunian hangovers from the previous century and regrettably are still with us), and documenting the succession of life in the fossil record expanded rapidly after the Tertiary triumphs, both in the geographic sense (due not least to European imperialist expansion) and in the geochronic sense, back down the geological column to the oldest known fossils. John Phillips proposed the fossil-based, three-part succession of Palaeozoic, Mesozoic and Cainozoic Series (now Eras/Erathems). The detailed list of ammonites characterizing the Mesozoic stratigraphic succession (Fig. 7) is a sample of Phillips’ summary. The unscaled curve in Fig. 8, with an overall expansion interrupted by two strong constrictions, strongly anticipates modern diversity curves based on much more data and numerical age control, as has been observed (e.g., Rudwick, 1972, 2008).

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Figure 7. Two figures from Phillips’ (1860) Life on Earth. left: the classical three-part ammonoid succession in the sutural pattern (the junction of the chamber partition with the inner surface of the outer shell of these extinct cephalopods). Although his book exemplifies the triumphs of the palaeontological synthesis, the sequence goniatite→ceratite→ammonite was not to Phillips a phyletic lineage. Right: the ammonoid zonation of the Mesozoic still displaying the style of Phillips’ uncle, the legendary William Smith, in matching the lithological units with their characteristic species.

Clearly stratigraphy and stratigraphic palaeontology achieved impressive levels of sophistication in the 1830s–1850s. lithostratigraphy, biostratigraphy, facies: ‘modern stratigraphy rests securely on these three basic achievements of the human mind’ (Teichert, 1958), and all were well established then. The all-important biostratigraphy itself developed on three pedestals, namely (i) the recognition of successional assemblages of fossils in successional strata; (ii) the successful testing and confirmation of that succession in other localities and other regions; and (iii) the perception that similarity among assemblages of fossils indicates similarity in geological age (McGowran, 2005).

The discriminating of successional fossil assemblages, as in Deshayes’ great achievement, proceeded in two directions concurrently in European geohistory and biohistory. The French palaeontologist and explorer Alcide d’Orbigny assembled a monumental palaeontological succession of 27 marine stages, based on the stratigraphic distributions of ~1800 species (Vénec-Peyré, 2004). The recurring overturns in species’ composition were wholesale, representing mass extinctions. This was extreme catastrophism; even so, many of d’Orbigny’s stages survived in the time scale. In another direction the succession of fossil assemblages gave way in some degree to compiling taxon ranges through strata and, by interpretation, through time, hence presenting boundary criteria for recognizing the biozones of Albert Oppel in the 1850s. To Oppel:

... is due not the credit for the inception of the zonal idea, but for a very great refinement in its use, and, most important of all, for emancipating the zones from the thralls both of local facies, lithological and palaeontological, and of cataclysmic annihilations, thus giving them an enormous extension and transferring them from mere local records of succession to correlation-planes of much wider (theoretically universal) application

(Arkell, 1933; in McGowran, 2005). Thus, we have here in the mid-19th century a full-blown science of stratigraphy and palaeontology and their fusion in biostratigraphy, producing a geological time scale capable of ordering and arranging strata and events in a global geohistory and biohistory based on millions of rigorously studied fossil specimens. An example is shown in Fig. 9. biostratigraphy, the palaeontological synthesis, happened in the six decades between Cuvier’s demonstration of extinction and Darwin’s demonstration of evolution, it all depended on the record on ancient life, and it was supported by very little accepted theory on the extinction of species and by no consensual theory at all on the origin of species. And biostratigraphy’s brilliant practitioners steadfastly refused to accept Darwin’s theory.

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Figure 8. Phillips’ geological time scale with the Palaeozoic, Mesozoic and Cainozoic ‘series of organic affinities’ (now eras), also showing the remnants of the lithology-based scale. Using stratal thickness as a proxy for time, four decades before radiometric age determinations became feasible, Phillips (1860) produced a ‘remarkably modern looking’ diversity curve (Rudwick, 1972). The pronounced contractions at the major boundaries were explained away by lyell and Darwin as artefacts of sampling an highly incomplete stratigraphic record, but are now known to be due to mass extinctions.

Reverent silence on the mysteries of mysteries

Our simple question for the architects of the palaeontological synthesis is: Was not the evolution of life blindingly obvious? It has a simple answer (‘no’) and a complex answer. The complete dominance of palaeontological evidence in all the research that mattered relied upon species extinct and extant being stable and reliable entities. lamarck’s transformism, with no extinction and no concept of stable species, attracted naturalists and interested citizens, thereby keeping evolutionism alive in the culture, but it offered little to the actual practitioners of taxonomy and biostratigraphy and geohistory whose currency simply had to be stable and trustworthy; more than that, for Cuvier, lamarckian transmutation was a real threat to progress in biohistory and geohistory—for how could a fossil record of unstable or even nonexistent species ever deliver a stable biostratigraphy?

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progress in fossil-based research. Two of the three principle theories of evolutionism were ruled out (Table 2) and the third, the theory of variational evolution, had not been assembled.

The early-mid-19th century reality was that biostratigraphic correlation and age determination could drive geohistory to reach around the globe without resolving these questions of origin and extinction. Chronicling, ordering and correlating could proceed scientifically—as theories of pattern—in the absence of theories of causation.

Jumping ahead for a moment, certainly there were three particularly refractory problems with the fossil record as to species’ origins and evolution, as Darwin stated candidly (1859, p. 310):

... [i] our not finding in the successive formations infinitely numerous transitional links between the many species which now exist or have existed; [ii] the sudden manner in which whole groups of species appear in our European formations; [iii] the almost entire absence, as at present known, of fossiliferous formations beneath the Silurian strata... .

Accordingly, he acknowledged the forbidding reality about the leading discoverers of the stratigraphic and palaeontological succession, namely that:

... all the most eminent palaeontologists, namely Cuvier, Owen, Agassiz, barrande, Falconer, E. Forbes, &c., and all our greatest geologists, as lyell, Murchison, Sedgwick, &c., have unanimously, often vehemently, maintained the immutability of species.

As examples we can cite the reactions to the Origin of eminent and thoroughly historicist minds remaining resistant to Darwin’s evolution: Heinrich Georg bronn (1800–1862), Richard Owen (1804–1892), louis Agassiz and John Phillips.

The German palaeontologist bronn was very similar to Darwin, with whom he was in contact, in seeking a scientific history of life on earth and drawing on all kinds of evidence from palaeontology to artificial breeding to achieve it (Gliboff, 2007, 2008). For bronn, German biohistory was stranded in a swamp of 118

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ORGANIC EVOlUTION IN DEEP TIME: CHARlES DARWIN AND THE FOSSIl RECORD

idealistic morphology and Naturphilosophie and he saw in Darwin’s approach a promising new way forward. And yet, he could not accept the theory of evolution by speciation; his review of the Origin (in Hull, 1973) was mired in such matters as spontaneous generation and the origin of life.

The british comparative anatomist and vertebrate palaeontologist Owen was surely not antievolutionary but had a personal problem: ‘he found himself in the uncomfortable position of wanting to censure Darwin for enunciating his theory of evolution while simultaneously claiming priority for himself’ (Hull, 1973, p. 215). Agassiz’ student Alpheus Hyatt (1897, p. 212) had this to say about his mentor, half a century later:

One wonders, as he reads, how any man holding such views could have held his mind closed to the conclusion that animals were evolved from simpler or more primitive forms. The effect of theoretical preconceptions in closing the mind to the reception of new ideas never had a stronger illustration. louis Agassiz, in 1849, had all the facts that would have placed him in the history of science on the same line with lamarck and Darwin.

John Phillips quoted the concluding paragraphs of On the Origin of Species with their famous ‘entangled bank’ and ‘There is grandeur in this view of life’, but forcefully rejected the ‘Darwinian hypothesis’, adding that Sedgwick did so too. Here is Phillips’ own resonant final paragraph in Life on the Earth concluding an impressive survey of the fossil record through geological time:

It may be thought that, while professing to keep to the old and safe method of reasoning on known causes and ascertained effects, we deviate from this principle in regard to the origin of life, and introduce an unknown cause for phenomena not understood, by calling to our aid an act of ‘creation’. be it so, let the word stand for a confession of our ignorance of the way in which the governing mind has in this case acted upon matter; we are equally ignorant in every other instance which brings us face to face with the idea of forces not manifested in acts. We see the stream of life flowing onward in a determined course, in harmony with the recognized forces of nature, and yielding a great amount of enjoyment, and a wonderful diversity of beautiful and instructive phenomena, in which MIND speaks to mind. life through many long periods has been manifested in a countless host of varying structures, all circumscribed by one general plan, each appointed to a definite place, and limited to an appointed duration. On the whole the earth has been thus more and more covered by the associated life of plants and animals, filling all habitable space with beings capable of enjoying their own existence or ministering to the enjoyment of others; till finally, after long preparation, a being was created capable of the wonderful power of measuring and weighing all the world of matter and space which surrounds him, of treasuring up the past history of all the forms of life, and of considering his own relation to the whole. When he surveys this vast and co-ordinated system, and inquires into its history and origin, can he be at a loss to decide whether it be a work of Divine thought and wisdom, or the fortunate offspring of a few atoms of matter, warmed by the anima mundi, a spark of electricity, or an accidental ray of sunshine?

The Darwinian Revolution Mark I

The verdict of the most distinguished palaeontologist since Cuvier:

Darwin was one of history’s towering geniuses and ranks with the greatest heroes of man’s intellectual progress. He deserves this place first of all because he finally and definitely established evolution as a fact, no longer a speculation or an alternative hypothesis for scientific investigation. His second greatest achievement was correct identification of a major element in the rise of adaptation: natural selection.(Simpson, 1949 (1967), p. 268.)

Darwin was a geologist

‘... the Darwinian revolution did not arise out of morphology and taxonomy ... It arose out of biogeography and ecology. The discoverer was a geologist with extensive knowledge of natural history’ (Ghiselin, 1980). Darwin became a superb geohistorical thinker and deeply involved in palaeontology (Herbert, 2005; Herbert and Norman, 2009).

Rock relationships and earth history—geognosy and geohistory—are the heart and soul of geology. Darwin grew to maturity collecting beetles and shooting partridges in a family worldview seasoned in the geotheory of his grandfather Erasmus. As an undergraduate in Edinburgh he was influenced by Robert

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Grant, a distinguished marine biologist and transformational evolutionist, Robert Jameson, mineralogist and the last of the Neptunists of Werner’s school in Freiberg, and presumably also the charismatic chemist Thomas Hope, Plutonist and follower of James Hutton (Secord, 1991). In Cambridge were the geologist and botanist Henslow and the divine and geologist Sedgwick, who took Darwin on a field-mapping trip in Wales. The young man’s lively and curious mind was well-stocked indeed in the natural history disciplines and he joined the Beagle at the time that Cuvier’s geohistory and biohistory were in full flower and lyell’s was bursting forth. And by the first landfall in early 1832 he was identifying himself as a geologist.

Which brings us to the changing place of lyell in the Darwin narrative, from highly influential geological mentor to loyal friend and ally and, at last, to reluctant acceptor of organic evolution. Rudwick (1990, 2008) described how the young lyell in the mid-1820s clearly endorsed a strongly directed geohistory including a progressive trend in the fossil record but then, steeped in natural theology as he was, took fright at the indignity of man being descended from animals, as implied by lamarckian transformism. He assessed the best way as a geologist to undermine lamarck to be to abjure all directionality in geohistory and biohistory, which course brought him inevitably into a Huttonian worldview or geotheory of steady state and eternal cycling. Even the goal of the acclaimed Tertiary studies based on Deshayes’ molluscan labours was not merely to erect robust divisions of Tertiary strata for a Tertiary timescale. The longer game was to place the Tertiary in a grand geological cycle, its workings to be understood by known and observable natural causes and to be calibrated by a biological clock founded in an unchanging ahistorical law, namely that species enter the scene, exist, and depart in a metronomic pattern (Rudwick, 1982). Time’s cycle quite dominates time’s arrow (Gould, 1987) and evidence for the latter is to be explained away as failure of the record.

lyell’s peers in the 1830s acknowledged his comprehensive views and outstanding powers of synthesis, his insistence on the rigorous use of causes observable in action and his Tertiary studies; Principles killed off the Noachian Deluge once and for all. The peers did not accept his steady-state view of geohistory and biohistory including his notions of species (Herbert, 1985). Speaking of those peers: ‘Contemporary leaders of the science such as Sedgwick and Murchison filled notebooks with strike and dip symbols, sections and other information about strata, only occasionally interspersing these jottings with more reflective passages. Darwin was highly speculative, with causal explanation as his central goal’ (Secord, 1991). Secord found it extraordinary that Darwin aligned himself with lyell, ‘whose controversial program of reform had almost no followers in England’, but surely this was a fertile meeting of minds and a shrewd move with little extraordinary about it.

Fossils, biogeography and genealogical inference

Richard Owen (1860) was impressed by the close correspondence between the modern mammalian faunas and the Pliocene-Pleistocene faunas on the Europaeo-Asiatic land mass. He found that Darwin’s collection of South American fossil mammals were as distinct from the Europaeo-Asiatic forms ‘as they are closely allied to the peculiarly South American existing genera’. Again on his first-hand knowledge, he found that ‘the ossiferous caves of Australia’ confirmed the ‘law’ that ‘with extinct as with existing Mammalia, particular forms were assigned to particular provinces, and that the same forms were restricted to the same provinces at a former geological period as they are at the present day.’ The extinct giant Megatherium was a placental close to the living sloth in South America; the extinct giant Diprotodon and others were marsupials close to the living forms in Australia. Thus, on the evidence of living and fossil mammals there was a very strong regional signature implying discrete centres of origin, not only now but preceding a putative Deluge. Darwin derived from this pattern his ‘law of succession of types’.

Darwin could add this biogeographic configuration to others, as summarized schematically (Fig. 10). On the long-appreciated, often exquisite adaptations of organisms to their respective environments one might predict close relationships among the tropical biotas and likewise among the oceanic island biotas and the temperate- continental biotas respectively. Not so! It was quite the other way around. The centres-of-origin notion was reinforced and no evidence was stronger than this biogeography and palaeontology in building Darwin’s theory of evolution (e.g., Ghiselin, 1969; Herbert and Norman, 2009).

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Figure 10. A schematic rendering of southern biogeography with the fourth dimension. Adaptations in an evolution-denying mode would predict similarities within the realms of tropical, temperate and oceanic-island habitats respectively. Several biogeographers observed the patterns that falsified that prediction, a falsification compounded by adding in the Pleistocene megafaunas (as they were later called) observed acutely by Owen, but only Darwin made the jump from geographic similarity to evolutionary centers of origin. As he explained in a letter to Haeckel, 8 October, 1864 (Haeckel, 1876, p. 134):

In South America three classes of fact were brought strongly before my mind. Firstly, the manner in which closely allied species replace species in going southward. Secondly, the close affinity of the species inhabiting the islands near South America to those proper to the continent. This struck me profoundly, especially the difference of the species in the adjoining islets in the Galapagos Archipelago. Thirdly, the relation of the living Edentata and Rodentia to the extinct species. I shall never forget my astonishment when I dug out a gigantic piece of armour like that of the living armadillo.

The five Darwinian theories

Darwin referred repeatedly to ‘my theory’ and Ghiselin (1969) more than anyone has emphasized the unity, the strong coherence in the Origin, of evidence drawn from everywhere. Some refer to a two-part theory (e.g., Sober, 2009) comprising the tree of life, whereby all organisms now living trace back to a common ancestor, and natural selection, being an important cause of the similarities and differences in the biota. Kitcher (2009) refers to Darwin’s three doctrines as pervasive variation, relation by descent, and modification mainly by natural selection. Mayr (1985, 2004) sorted ‘Darwinism’ into five theories—evolutionism and tree of life (two pattern or historical theories), and speciation, gradualism and natural selection (three causal theories). I find this deconstruction to be helpful (Fig. 3) as does Kutschera (2009), although for others it underplays the unity of his thinking and the coherence of his system (Reif et al., 2000; Ghiselin, 2002).

First, evolutionism, the evolution of life on an ever-changing planet, was not original to Darwin, but certainly original was his powerful and meticulous assembling of a wide-ranging case to prove that evolution happened. Second, the tree of life was not original iconography (Archibald, 2009) but this tree, the branching transmutation of species, was compelling. Next: change through time is one thing but the generation of diversity, the emergence of two or more species from a common parent is quite another, and Darwin came to realize that a theory of speciation (the word itself is much younger) was essential. The principle is ecological, coming out of economics

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(Adam Smith) via agronomy, and arguably is the keystone of the Origin (Kohn, 2009, who has coined ‘divergence selection’). Fourth: also seen as essential was calming down Cuvier’s and d’Orbigny’s revolutions and anticipating the catastrophism seen by many in the notoriously disjunctive stratigraphic and fossil record (not to speak of biblical creation or extinction in the Noachian deluge), and he strove to establish the case for gradual not saltational change through deep time (the latter also anachronic as a mid-20th century appellation). The fifth theory, natural selection, has been the most-discussed component of this comprehensive super-theory of evolution.

Certainly there were critics (Hull, 1973). To repeat, palaeontologists and geologists from Cuvier onwards rejected evolution as an explanation of the fossil record of organic succession. All british geologists rejected Darwin’ theories, as did von baer, Agassiz and bronn. Even so, Darwin’s two historical or pattern theories came to be accepted within a decade or so, after which the evolution of life as a fact was a non-issue for virtually every competent and professional biologist and, more tardily, geologist; ‘Darwinism’ was widely accepted (Ellegard, 1958) and ‘Darwinian impacts’ (Oldroyd, 1980) were far-reaching. but it was not a clearcut example of Planck’s principle whereby younger scientists accept new scientific ideas with greater alacrity than their elders (Hull et al., 1978). At any rate, the three causal theories were another matter (see below).

What of the fossil record? ‘Darwin’s dilemma ... was that he both needed palaeontology and was embarrassed by it’ (Sepkoski and Ruse, 2009). The embarrassment was for the discontinuity and the need was for strong biological evidence from deep time. Examples of species-by-species lineages were not forthcoming, as has been remarked many times. but the grand sweep of the fossil record as a whole and in innumerable details was a different matter and ‘dilemma’ is a vast overstatement. The fossil-biogeography configuration was especially compelling (above). Then there were the broad trends from the Palaeozoic to the Recent—the more ancient a form is, the more it differs from living species. In detail, as had been and was being worked out for biostratigraphic reasons, fossils in adjoining stages or formations have much more in common than either has with assemblages more removed, up or down the succession. Perhaps, with ‘grand natural system’, Darwin called it best. The great radiations (as they were later named) such as the Palaeozoic trilobites or the Mesozoic ammonites could be placed within higher taxa already discriminated, even though the fossil groups themselves were extinct.

Genealogical relationship explained taxonomic groupings just as it explained the patterns in the fossil record, in both cases compiled mostly by non- or anti-evolutionists. A critically significant item in the Darwinian narrative is his project on the barnacles, carried out 1846–1854, i.e., between producing an unpublished outline of his evolutionary theories and being goaded into renewed action to publish by receiving Wallace’s manuscript on natural selection. Challenged by Hooker to accrue his own biosystematic credentials, he commenced the project as a tyro and emerged as a full-blown comparative morphologist and pioneering evolutionary or genealogy-based taxonomist (Ghiselin & Jaffe, 1973; Padian, 1999; Ghiselin, 2004).

Palaeontology presented Darwin with a massive compendium of the succession and history of life on earth and the insights of half a century’s wrestling with the discipline of geohistory. Admittedly, the opening chapters were missing. In return, palaeontology received a comprehensive explanation of that document, a great boosting of that discipline and likewise in a new and overlapping discipline, evolutionary taxonomy. but what of the three causal theories of speciation, gradualism and natural selection?

Eclipse of Darwinism

There is a very deep background, a long history to ‘evolution’, to the notion of change in the organic world, or to the idea that a system or a grand structural plan based on similarities—systematic affinities—might be due to blood relationship (Osborn, 1929; Russell, 1916). leibniz and Kant were among those to whom such thoughts occurred. However, the leading embryologists and comparative morphologists became stranded at the level of homology and affinity, one level short of literal genealogy and descent, Goethe being a prominent example (Cole, 1944). Cole portrayed Goethe as inferring that two of his studies, one an admirable exercise in comparative anatomy and the other an exploitation of The Idea or unitary whole, confirmed that idea. Thus Goethe was a forerunner of Darwin only in the sense that he was a leader in formulating a doctrine of homologies, an essential precondition of an evolution-theory, as Darwin demonstrated. Russell’s view (1916, p. 213) was like Cole’s, but stronger:

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It is a remarkable fact that morphology took but a very little part in the formation of evolution-theory. When one remembers what powerful arguments for evolution can be drawn from such facts as the unity of plan and composition and parallelism, one is astonished to find that it was not the morphologists at all who founded the theory of evolution.

Russell also described how it was for the morphologists of the old school, the classical tradition—they accepted evolution and Darwin’s laying the foundations of evolutionary morphology, but Darwinism still was not synonymous with evolution.

That detaching of the accepted fact of evolution from the not-accepted theory of natural selection characterized most of the work in the transformational disciplines for decades after the Origin. In Europe, as the Origin progressed through six editions, palaeontology, which had already unearthed one specimen transitional between groups, the Neanderthal humanoid skull, revealed the most spectacular of all, the Jurassic Archaeopteryx, part-reptile, part-bird; it was developing a plausible pattern of horse evolution; and Mesozoic and Cenozoic lineages were being reconstructed from successions of shelly marine fossils. but the major advances were occurring in North America where fossil faunal and floral discoveries were spectacular, where evolution as a fact was not contested but its patterns and modes were, and vigorously. North American vertebrate palaeontology was a well-defined branch of post-Darwinian morphology (Rainger, 1981, 1985, 1986). beyond doing the systematics of the fossils avalanching out of the west, its practitioners sought to identify ancestral and intermediate forms, to define morphogenetic changes through time, and to construct phylogenetic trees. This (pre-Darwinian) tradition of palaeontology coupled with morphology and embryology was reinvigorated among the vertebrates by E.D. Cope (1840–1897) and to a lesser extent O.C. Marsh (1832–1899), subsequently H.F. Osborn (1857–1935) and, in the marine invertebrates, by A. Hyatt (1838–1902). Cope was one of several to happen upon what became known as Haeckel’s biogenetic law: ontogeny recapitulates phylogeny, which ‘law’ retained influence well into the next century (Russell, 1916; Gould, 1977). (For Hyatt (1897), ‘The so called Haeckelian “law of biogenesis” is really Agassiz’s law of embryological recapitulation restated in the terms of evolution.’) Cope asserted that Darwin neglected the origin of those variations which could not be accounted for by natural selection and which arose from a growth force, dormant in the tissues until actualized by, especially, use/disuse and effort, in turn directed by a something he called unconscious choice, the originator of the fittest. This ‘causal efficacy of the reactions of organisms to environmental conditions’ (Rainger, 1981) became influential in the late 19th century. but it is not clear to me if or how Cope’s theorizing affected his own palaeontological achievement, which Rainger (1981, p. 138) summarizes as very impressive indeed, thus:

... Cope defined in detail the changes in the bones of the feet of fossil and recent mammals. Thus by the mid 1870s Cope had defined the ancestral form of a major division of the mammals, had established the principal structural changes that had occurred from the ancestral to the more modern forms, and had provided tables and illustrations of the patterns of descent defined by those changes in structure. In short, Cope’s work fulfilled the major objectives of a science of morphology.

However, in the 1880s and 1890s he used more explicitly the doctrine of inherited use/disuse to explain observed morphological changes in the successions of fossils. ‘Kinetogenesis’, the effects of parts moving against other parts, was elevated to the status of a theory explaining the origin and evolution of changes in vertebrate structure. Cope’s book The Primary Factors of Organic Evolution (Cope, 1896) profoundly influenced American biology at the turn of the century (Rainger, 1981). The theory came to be known as ‘Neo-lamarckian’, which is peculiar because it omitted the generally perfecting tendency in evolution, central to lamarck’s theory, and committed heavily to the inheriting of direct environmental impact on organic structure, explicitly denied by lamarck, whilst having use/disuse in common with lamarck (e.g., Simpson, 1949 (1967)).

Hyatt (especially 1897) expanded his views on perceived parallels between embryonic growth, the structural gradation among living forms and the geological succession of extinct forms, for which the ontogeny- preserving, character-rich shells of the extinct cephalopods, the ammonites, are ideal material. Hyatt identified in ontogenetic acceleration both progressive and retrogressive evolutionary change. From the latter he developed in the 1890s his theory of old age or racial senescence, whereby individual or ontogenetic aging markedly paralleled the stages of evolutionary decline and extinction (e.g., Rainger, 1981; Gould, 2002).

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During this time evolution itself and the tree of life were not in doubt amongst the informed, but the recovery of Darwinism from the Neolamarckian-Neodarwinian polarizing had to await the awakening of population genetics and a turnaround in palaeontological thought in the 1920s–1930s (below). Meanwhile, palaeontology in the German-speaking lands in Europe lurched still further in the orthogenetic direction. The following section draws heavily on very fine discussion by W-E Reif (1983, 1986, 2003; Reif et al., 2000).

Typostrophism and the German Synthesis

Darwin’s evolution rapidly supplanted the remnants of Naturphilosophie among German biologists generally but much less widely among the palaeontologists, who were more geological and for whom it was not a burning question whether their taxa had evolved or been created (Reif, 1986, 1993). Virtually all fossil workers accepted descent from a common ancestor but very few had much enthusiasm for natural selection. There were outstanding exceptions. Wladimir Kowalevsky (1842–1883) 1874 achieved in the 1870s an evolutionary history of the horse and other ungulates which was ‘one of the first and greatest triumphs of evolutionary palaeontology’ (Simpson, 1944, p. 103; see also Reif, 1986).

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In the last two decades of the 19thC, virtually all palaeontologists in Germany (as indeed in western Europe and North America) failed to find the pattern of gradual improvement deduced from Darwin’s theory. The particularly able Wilhelm Waagen (1841–1900), close contemporary of Hyatt and also an ammonite specialist, exemplified those. It was Waagen who distinguished intraspecifically between contemporaneous morphs and successional mutants, or mutations, before the geneticists apprehended that word, and he came close to the notions of ecophenotypes and index taxa.

The German palaeontological culture developed a strong self-consciousness and sense of autonomy. Even if microevolution as observed by biologists in living populations was controlled by natural selection, macroevolution documented by the fossil record could only be explained by specific macroevolutionary processes—by the fossil record itself (Reif, 1993). It is too easy to characterize this as reactionary—anti- Darwinian, anti-biological, anti-selection, anti-uniformitarian. Consider Othenio Abel (1875–1946), namer of ‘palaeobiology’ in 1929, long before its uptake by the Americans (Reif, 1980; Kutschera, 2007). Adaptation by evolution was more interesting and important to Abel than to his biostratigraphic colleagues. lamarckian mechanisms tended to be:

[but] minor corrections in the flow of the evolutionary stream, which had its own internal driving force and hence a momentum independent of environmental factors. Thus a lineage could evolve beyond the adaptive realm and acquire hypertrophied organs and other deleterious characters that led inexorably to extinction.

(Reif, 1993, p 437).

Abel’s law of biological inertia embraced four subsidiary generalizations: 1, Haacke’s 1893 law of orthogenesis, the internal driving force; 2, Rosa’s 1899 law of continuous reduction of variability, hence adaptive versatility in the origin of a higher taxon; 3, Cope’s 1893 law of the unspecialized stem group, so for Abel a new higher taxon arrives on the stage with a new reservoir of variability; and 4, Dollo’s law of the irreversibility of evolution—the high momentum of evolution does not allow any reversals.

The school of typostrophism culminated in ‘the triumph of the anti-Darwinian, anti-uniformitarian, orthogenetic, and saltationist theories that dominated [German palaeontology in] a large part of this century’ (Reif, 1993). based in orthogenesis as an orderly process of a lineage unfolding, typostrophism added these: saltationism, whereby macroevolutionary novelties are produced in jumps; cyclism, whereby evolutionary history proceeded in cycles analogous to the life cycle of an organism; and idealistic morphology and the morphological type. The leaders were Karl beurlen and Otto Schindewolf, invertebrate palaeontologists who independently derived macroevolutionary conclusions from ammonites and corals—well-skeletonized and well-fossilized organisms preserving ontogenetic development in their skeletons. The necessary genetics requiring macromutations came from the geneticist Richard Goldschmidt, culminating in his Material Basis of Evolution, published in 1940.

The typostrophist manifesto was from beurlen in 1932 (in Reif, 1993):

It is a very general rule that the pathway of evolution within a taxon—irrespective of whether it is a unit of higher or lower rank—is cyclic. Evolution starts with a first phase of rich saltation and explosive creation of forms. It goes on in a phase of orthogenetic continuity that is directional and purposive and does not produce new types of forms. Finally, a phase of degeneration and disintegration of forms leads to extinction.

Schindewolf published in 1936 Paläontologie, Entwicklungslehre, und Genetik, apparently the first serious attempt to synthesize palaeontology with genetics. His enormous influence on German palaeontology for several decades was reinforced by the 1950 Grundfragen der Paläontologie, very belatedly issued in English (Schindewolf, 1993). The two major theories were typostrophism and proterogenesis, the former being the core, ‘which I oppose to Darwinism, for the latter, inadmissibly, in my opinion, has simply applied the mechanisms responsible for the formation for race and species to the entire evolutionary process’ (p. 215). In proterogenesis new features arise suddenly by transformation early in ontogeny and move forward, appearing in the adult stage in due course (Fig. 12). Schindewolf displayed (his Fig. 3.73) a schematic typostrophic cycle in three parts: (i) typogenesis, many groups arising vigorously, saltationally producing new structural organizations; (ii) typostasis, gradual, lengthy orthogenetic phase, nonadaptive; and (iii) typolysis, the types lose their consistent morphological identity while producing all kinds of ‘degenerative offshoots’

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Figure 12. Schindewolf’s ammonite proterogenesis in reconstructed lineages, Callovian-Oxfordian (Middle-late Jurassic), drawn in Schindewolf’s style as if dissected to expose the juvenile whorls. In each lineage the change in form was initiated in the juvenile shell and proceeded in due course into the mature shell. From Ziegler (1972, Fig. 123), courtesy of E.Schweizerbart’sche Verlagsbuchhandlung (www.schweizerbart.de).

leading to extinction. As soon as the class type appears in the typogenetic period it splits abruptly into the subtypes of its orders, which in turn split into members of suborders, then families ... And again Schindewolf stakes his territory (p. 202, italics his): ‘The real problem in evolution thus shifts from the “origin of species”, which until now has been in the forefront, to the understanding and explanation of the far-reaching, unmediated differences between types.’ Princehouse (2009) has dubbed this ‘beautifully integrated’ synthesis the German Synthesis. (Note: the name ‘German Synthesis’ was used previously by Reif et al. (2000) for quite another movement in Germany, namely the German contribution to the Synthetic Theory centred on the zoologist bernhard Rensch (1947–1972).)

For some, the above gives too much space to the anti-Darwin Schindewolf and the German synthesis. Thus, Conway Morris’ review (1994) of Schindewolf (1993) is titled ‘wonderfully, gloriously wrong’.

Darwinian Restoration: the Synthetic Theory of evolution

I quote from the blurb of the triumphalist The Paleobiological Revolution: Essays on the growth of modern paleontology (Sepkoski and Ruse, 2009):

Paleontology has long had a troubled relationship with evolutionary biology. Suffering from a reputation as a second-tier science and conjuring images of fossil collectors and amateurs who dig up bones, paleontology was marginalized even by Darwin himself, who worried that incompleteness in the fossil record would be used against his theory of evolution. but with the establishment of the modern synthesis in the 1940s and the pioneering work of George Gaylord Simpson, Ernst Mayr, and Theodosius Dobzhansky, as well as the subsequent efforts of Stephen Jay Gould, David Raup, and James Valentine, paleontology became embedded in biology and emerged as paleobiology, a first-rate discipline central to evolutionary studies.

The ‘Synthetic Theory’ or the ‘Modern Synthesis’ is prominent in any account of Darwin and the fossil record. Recognized at the genetics level were the importance of mutation, (individual) selection, recombination, isolation and drift, collectively driving two phenomena, namely (i) speciation (predominantly allopatric) and evolution which was gradual (meaning not saltational and not to imply one-speed), and (ii) macroevolution, meaning the origins of higher taxa, radiations (bursts of diversification), and adaptive shifts into new environmental realms.

Rejected at the genetics level and at neontological timescales were macromutations, or the systemic mutations of Goldschmidt (1940), and the ‘soft’ or ‘lamarckian’ inheritance of acquired characteristics. Rejected at higher levels and palaeontological timescales were all such notions as internal evolutionary drives and

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orthogenesis, inherent progressive trends, goal-oriented evolution or finalism, cyclical evolution analogous to ontogeny (i.e., youth/maturity/old age and racial senescence), and overshooting by lineages into nonadaptive evolution. Also ostracised were archetypes, Baupläne and saltations, or jumps. Clearly, palaeontology, in the writings of Cope and Hyatt, culminating in the ebullient writings of Henry Fairfield Osborn (1934), was grossly over-represented in this compendium of rejects and discards. Morphology, claimed by some to lead the way in demonstrating the fact of evolution in the years after the Origin (e.g., Davis, 1949), continued in its uneasy relationship with evolutionary biology, again failing to assimilate Darwinism (Simpson, 1959; Ghiselin, 1980, 2006).

Tempo and Mode in Evolution (Simpson, 1944) preeminently brought palaeontology back to Darwinian evolution and vice versa, but a thin Darwinian red line did traverse the anti-Darwinian decades. The outstanding statement for Darwin in those dark times was Kellogg’s Darwinism Today (1907). One outstanding European example of a selectionist and gradualist was the above-mentioned Kowalevsky; another was Melchior Neumayr (1845–1890) in Vienna (Reif, 1986). In North America Rainger (1986) argued cogently that William Diller Matthew (1871–1930), ‘great paleomammalogist and hero’ to Simpson (1984, his own words), did provide:

a new means for a young vertebrate paleontologist to approach the study of evolution and the fossil record. In contrast to the work of Osborn, lull, and others, Matthew’s researches offered a means for interpreting the fossil record in accordance with selection theory, a means for recognizing the importance of genetics on the understanding of the fossil record, and a new means for understanding the variations among fossil remains that would have important consequences for systematics. In short, Matthew’s work provided a new conception of the fossil record

upon which Simpson would build.(Rainger, 1986, p. 470). Simpson needed in addition a grasp of statistical analysis, in which he became more than competent (Simpson and Roe, 1939), and an insight into the new population genetics: he absorbed Dobzhansky’s (1937) Genetics and the Origin of Species, the work that more than any other and ‘both by field and laboratory studies established the recognition that natural selection can and does produce adaptive evolution’ (Simpson, 1984, p. xxii). He did not have access at the time to Mayr’s (1942) Systematics and the Origin of Species. The thesis of Tempo and Mode:

... in briefest form, is that the history of life, as indicated by the available fossil record, is consistent with the evolutionary processes of genetic mutation and variation, guided toward adaptation of populations by natural selection, and furthermore that this approach can substantially enhance evolutionary theory, especially in such matters as rates of evolution, modes of adaptation, and histories of taxa, particularly at superspecific levels ...

(Simpson, 1976, p.5). Simpson’s impact on palaeontology resembled more than slightly Darwin’s impact on biology—a powerful and pervasive influence ranging from mastering the palaeontology and neontology of the mammals, through taxonomy to macroevolution to biological education, from producing in Tempo one of the few truly key documents of the synthesis to four more decades as major public advocate of Darwin and of the synthesis (1964, 1970) (Cain, 1992). Simpson presented his threefold view of micro-, macro- and mega-evolution as the basis for the three modes, speciation, phyletic evolution and quantum evolution (Fig. 13, 14).

As early as 1925 he worked on fossil mammals at the three levels of taxonomy (as defined later by Mayr), alpha, defining numerous new species, beta, their phylogenetic classification, and gamma, their broader biological meaning (see also Simpson, 1951) (laporte, 1994). In due course he produced the magisterial classification of mammals with a powerful statement on his taxonomic philosophy (1945) which became a book (1961) ranking in my opinion with Tempo. laporte traced Simpson’s developing concepts of species, from type to population, statistical treatment of the taxon, then from taxonomic to ecological concept, yielding a highly plausible scenario of small and localized populations breaking through at very high evolutionary rates, all consistent with the neontological scenarios of population genetics and natural history.

It was said that Simpson did a splendid job in Tempo and Mode (a) in demonstrating that evolution and population genetics could be reconciled (with smooth extrapolation) with evolution and the fossil record, and

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italics) surely was justified in observing (four

decades ago and it still holds): ‘Modern evolutionary theory is closer to the original Darwinian formulation today than it has ever been’. In the interests of open communication, though, here is Ghiselin (2001), somewhat biliously, on those referred to by Futuyma (1988) as the ‘old masters’:

The notion that “the” Synthesis was somehow complete at one time or another in its history implies that the participants were aiming at some culminating event, like the Resurrection of Christ. The canonical texts are being treated as if they were The Gospel according to Saint Doby, The Gospel according to Saint Ernst, The Gospel according to Saint G.G., The Gospel according to Saint Julian, The Gospel according to Saint bernhard, and The Gospel according to Saint ledyard. Scientists are explorers, not prophets. For them to display themselves otherwise is as dishonest as it is misleading.

Towards the Palaeobiological Revolution?

Critical assertions about the Synthesis

Its non-acceptance by Schindewolf and the typostrophists is obvious (Fig. 3), but there were also criticisms from others in palaeontology and morphology, especially Olson (1959); still others recur in the modern literature on evo-devo (laubichler & Maienschein, 2007). One discontentment with the Synthesis was of virtually ignoring the fact of extinction (below). Gould (1980, 1983, 2002) asserted that the Synthesis ‘hardened’, meaning making adaptation and natural selection all-important. In parallel was the theory that microevolutionary processes could be extrapolated entirely into the macroevolutionary realm, such as in the origin of higher categories, thereby downgrading any autonomy of macroevolution and ignoring the claim

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(e.g., Olsen, 1959, p. 524) that morphology ‘had provided the greatest single source of data in the formulation and development of the theory of evolution’. So far as palaeontology was concerned, much of this had to do with Gould’s reiterating that Simpson lost his nerve between (1944) and (1953) on quantum evolution, watering down that bold and exciting theory to the point of blandness, although for Cain (2003) Gould went too far. Ghiselin’s (2002, p. 269) gripe with the Synthesis was with the shortcomings of the taxonomy: thus Willi Hennig, founder of cladism, wished to participate by combating idealistic morphology but was ignored, and the ‘failure of the Synthetic Theory to come up with more than just a superficial and inadequate treatment of phylogenetics and to reflect it adequately in higher- level classification suggests that it was less of a synthesis than has been claimed’. Again, Mayr (1988) accused palaeontologists generally of ignoring the ‘horizontal’ dimension of his biological species concept and attending only to the ‘vertical’ dimension, emphasizing lineage change (anagenesis) over splitting (cladogenesis). This is a somewhat cryptic criticism in view of laporte’s analysis (above), but Simpson did favour anagenesis as occurring in 90% of evolutionary shift (whereas the reverse is now more accepted in these cladogenetic times).

Simpson (1976, 1984; in Mayr and Provine, 1980; in Cain, 2009) would have none of this, especially of the implication that palaeontology at its own time scales and on its own terms had nothing to contribute to Darwinian evolution. Nor would I accept another Gouldism: in his Foreword to the English translation of Schindewolf’s book, Gould (1993) would have us believe that the high priests of an orthodoxy need, in their Orwellian endeavours to ignore, silence or distort dissent, designated ‘whipping boys’. For the Darwinian orthodoxy in the form of the Modern Synthesis,

the boys were Goldschmidt and Schindewolf. Rereading Simpson and Mayr in this perspective hardly supports such overblown rhetoric. Even so, the ironic wheel of history keeps turning and now turns up this, under the rubric of sociology: ‘Ritual patricide: why Stephen Jay Gould assassinated George Gaylord Simpson’ (Cain, 2009).

The expansion of palaeobiology

Figure 3 merely lists eight subjects of modern evolutionary research of special concern to palaeontology. Molecular biology has produced the ‘molecular clock’ and invigorated genealogical reconstruction (Ayala, 2009). That morphology as ontogeny-phylogeny never sat comfortably in the modern synthesis is well known (e.g., Riedl, 1983; Ghiselin, 1980(1998), 2006; laubichler and Niklas, 2009); in evolutionary developmental biology (evo-devo), developmental genetics has infused new life into these problems (laubichler, 2007).

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Mass extinction became a research program unto itself (Hallam, 2004). Jablonski (2009) outlines and exhaustively references these and other items in a menu for palaeontology in the twenty-first century.

Progress in later Phanerozoic biogeohistory

Simpson’s accomplishment in hands-on palaeontology/zoology was in the mammals of the later Mesozoic and especially of the Cenozoic Era. He never lost sight of the environmental and ecological dimensions of evolution. We should remember though that our understanding of Cenozoic geohistory accelerated during the decades of the Synthesis and of the expansion of palaeobiology. These developments were driven by as well as affecting palaeontology—the world of fossils was not simply divided into competent and worthy if essential systematics on the one hand, and dynamic palaeobiological science on the other. but it might easily be inferred from numerous passages in the palaeobiological manifesto (Sepkoski and Ruse, 2009) that the relevance of the environment and its profound shifts is seriously downplayed. To compensate, the following list is slanted strongly towards micropalaeontological advances revealing the environmental template upon which deep-time evolution has happened. Indeed, the expansion of palaeobiology has been paralleled by the expansion of biostratigraphy, much of which I have discussed elsewhere (McGowran, 2005).

biostratigraphy drove micropalaeontology early in the 20th century (in geological mapping and economic drilling accompanying European imperial expansion). Progress accelerated with the fossil planktonic foraminifera in the 1930s and post-war oceanography, when the arsenal enlarged to comprise phytoplankton (diatoms, dinocysts and coccoliths) and zooplankton (foraminifera and radiolarians) (Riedel, 1973; lipps, 1993; lazarus, 2011). biostratigraphy was a driver, from onshore and offshore drilling to the advent of deep- ocean drilling in the late 1960s. Specimen numbers and sampling precision and refinement were unprecedented in palaeontology. Whereas the role of palaeontology in large-scale evolutionary research was defined narrowly thanks to a false belief, tracing back to Darwin and his early followers, that the fossil record is woefully incomplete (Stanley, 1979), marine micropalaeontology removes such strictures.

(ii) Correlation and age determination: the expansion of biostratigraphy

Planktonic foraminiferal phylozones were identified transhemispherically by speciational and extinctional bioevents. These bioevents could be (a) cross-correlated with the other microfossil successions (bolli et al., 1985), (b) tied to the rapidly evolving geomagnetic polarity reversal time scale (berggren et al., 1995), and

(c) tied to sequence stratigraphy (McGowran, 2005), cyclostratigraphy and astrochronology (lourens et al., 2004; Strasser et al., 2006), all (d) under rigorous and refined numerical calibration. Chronicling biohistory and macroevolution are under increasingly rigorous discipline. For example: we now accept the sheer speed of far-reaching, inter-ocean, paleoceanographic shifts which are seen in biostratigraphic markers (both phyletic and ecological) holding true across the huge geographic ranges of microplanktonic populations, between oceans and sometimes across tens of degrees’ latitudes. but how true is true? In Neogene nannofossil astrobiochronology, Raffi et al. (2006) consider a biohorizon to be isochronous when the available estimates of its age are constrained within a single orbital cycle (~20-100 kiloyears). Eighteen distinct and well-defined bioevents, demonstrably isosochronous worldwide, were identified and another seven biohorizons were deemed to be reasonably isochronous.

Cyclostratigraphy and astrochronology now provide a superb Neogene example of environmental impact on evolution (van Dam et al., 2006). Turnover cycles in densely sampled lineages of rodents closely match low- frequency modulations of Milankovitch oscillations—obliquity nodes and eccentricity minima which in turn 130

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are associated with icesheet expansion, cooling and precipitation changes. The authors infer that ‘long-period astronomical climate forcing [at 106 years’ scale] is a major determinant of species turnover in small mammals and probably other groups as well’.

(iii)Cenozoic environmental trajectory and greenhouse-to-icehouse transition Plate tectonics and continental drift transformed geography and biogeography ocean Tethys and opening the Southern Ocean initiated circum-Antarctic circulation,

and atmospheric circulation and CO2 levels and triggered long-term climatic change. Tracking

and their feedbacks with the biosphere employs elegant templates based on isotopes of carbon, strontium and others, sampled from single shells of foraminifera and bulk open-oceanic carbonate. have narratives for global temperature changes, fertility changes and carbon dioxide changes (Zachos 2001, 2008; Pagani et al., 2005; Thomas, 2008). The phrase ‘greenhouse-to-icehouse’ invokes a stately change and indeed it is a broad trend over the past 100 million years, but at all more refined chronological the record is strongly punctuated or episodic, as we realized already in the 1970s (e.g., berggren & Couvering, 1984) with much cruder data. McGowran (2005, Fig. 6.12) plotted eight well-corroborated environmental punctuations of Cenozoic geohistory—four sudden chills, two warmings and transgressions, one hyperthermal and one bolide impact—all shown to have biotic impacts and none particularly characteristic of the Cenozoic alone; the pre-Cenozoic fossil record is acquiring an environmental template too.(Hallam, 1994). Closing thealtered global oceanicthese changesoxygen,Thus weet al.,globallevelsvan

(iv) Mass extinction

Glaessner (1937) demonstrated the extinction of the planktonic foraminifera at the end of the Maastrichtian Age and Cretaceous Period and bramlette and Martini (1964) demonstrated the even more spectacular demise of the calcareous phyto/nannoplankton at the same horizon, already known for extinctions among the marine reptiles, ammonites and rudistid bivalves. It took careful integrating with geomagnetic stratigraphy and correlations between the pelagic, neritic and terrestrial realms to actually confirm that the dinosaurs disappeared at the same time, work that was still in progress (berggren et al., 1985) when the bolide hypothesis arrived (Alvarez et al., 1980) with the rigorous cyclostratigraphic confirmation of synchronicity still to come (Herbert et al., 1995). For Hull (1988), finding this mass extinction by counting higher taxa in the library (Sepkoski, 1981) was unexpected. Raup (Sepkoski & Raup, 2009, p. 468) observed : ‘What little Darwin said about extinction was dead wrong, absolutely dead wrong. It was all competitive replacement’, and the Synthesis virtually ignored extinction: ‘Rather like a demographer ignoring death’. Neither Hull nor Raup noted that the demise of the ammonites had been known since the 1930s to be part of a geologically sudden event, thanks to the precision of micropalaeontology; nor was the compilation by the anti-evolutionist Phillips (Fig. 8) accorded its due in the modern frenzy of mass extinction research.

(v)Phenotypes and genotypes and cryptic speciations After Glaessner’s (1937) phyletic reconstruction reconstructions became commonplace from the 1950s on, evolutionary study of the group by berggren (1969). Kennett pioneered modern-type taxic-evolutionary studies and some

(Kucera & Malmgren, 1998) were cited by Gould (2002). Evolutionary foraminiferal higher taxa and their relationships (e.g., Pawlowski, 2000). The with well-defined phenotypic species, are being revealed as species clusters. Darling and Wade (2008), 19 morphospecies had been searched for their small RNA (rRNA) genotypes/subtypes and the 19 morphospecies yielded 54 genotypes.

sorting may underestimate cryptic species by a factor of ~3. Moreover, the planktonic acknowledged by Gould to display phenotypic evidence for gradualism, in this integrated case evidence for punctuation and stasis. These cryptic clusters within morphospecies are the planktonic species proposed by De Vargas et al. (2003), who perceive significance in the very conservative nature morphological species or phenotype, which actually is a monophyletic cluster of sibling species which several of million years ago (on molecular clock calculations) and still display only slight, subtle, far obvious morphological divergence between the ecotypes/genotypes after all that time. They propose

this distinctive evolutionary mode characterizes most marine planktonic taxa, metazoans as well as protists. It is a powerful addition to the case for stasis in punctuated speciation (next section).of late Cretaceous planktonic foraminifera, suchthe most noteworthy outcome being a Simpsonian(Wei & Kennett, 1983; Stanley et al., 1988)of these and other microplanktonic studiesgenetics has radically challengedNeogene-Holocene planktonics,As of the major review bysubunit (SSU) ribosomalThus morphospeciesforaminifera,present strongsuper-of thesplitfromthat

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(vi) Punctuated speciation

This is a specific hypothesis about speciation whereby most evolutionary change is located in punctuated cladogenesis, followed by pronounced stasis: thus Gould (2002) in prolonged defence of perhaps the most- discussed of all palaeontological hypotheses excepting extraterrestrial mass extinction. PE is not the same as Simpson’s quantum evolution and it was not anticipated by anyone back to Darwin (Rhodes, 1983) whilst at the same time not being anti-Darwinian in any of the transformationist senses mentioned above. The outstanding post-Simpson contribution to palaeontology-based macroevolution was Stanley’s (1979) Macroevolution: pattern and process, writtern in punctuationist mode. Mayr (1992) came to accept PE in due course. Prothero (2009b) reported that PE has been widely accepted among palaeontologists for several years and that well- established stasis still lacks acceptable explanation.

(vii) Hierarchy and scale

... experimental biology in general and genetics in particular have the grave defect that they cannot reproduce the vast and complex horizontal extent of the natural environment and, particularly, the immense span of time in which population changes really occur. They may reveal what happens to a hundred rats in the course of ten years under fixed and simple conditions, but not what happened to a billion rats in the course of ten million years under the fluctuating conditions of earth history. Obviously, the latter problem is much more important.

(Simpson, 1944, p. xxix). Three kinds of hierarchy in the realm of historical biology are the linnaean taxonomic hierarchy, the evolutionary ‘extension of Darwinism’, and the deep-time ecological hierarchy. The linnaean system brought order into the seeming chaos of organic diversity and Darwinian organic evolution and evolutionary taxonomy showed why it did (Simpson, 1961; Mayr, 1969). There is a real genealogical tree ‘out there’, comprising real entities or individuals produced by splitting (speciation) and terminating (extinction) and growing in real lineages or clades with a real sequence of branchings, awaiting discovery and reconstruction. This is the ‘what’ of evolution.

Another biological hierarchy addresses the extension of Darwinism, in its narrow sense of evolution by natural selection within populations of individuals (microevolution), upwards to specific and supra-specific levels (macroevolution). The question is whether upwards extrapolation suffices to explain higher-level effects. In the long history of debate over evolution by natural selection, one view asserts that intrapopulational selection and microevolution is sufficient to extrapolate to macroevolutionary levels. In this view, hierarchy is merely a useful tool for ordering and classifying. In contrast is Gould’s (2002) exposition of evolutionary hierarchy (Table 2) in radical denial of extrapolation and in radical affirmation of its opposite, emergence. In this view the organic world is tiered nonfractally, a different process operating at each level (of the upper three) in the hierarchy. Hierarchical thinking now has a strong and solid presence in macroevolutionary thinking (Jablonski, 2007).

Also hierarchical is ecology and environment. So long as palaeoecology was regarded as but a deep-time extension of modern ecology, on the other hand, it took quite some time to find its own voice as being strongly autonomous, and capable of posing and answering questions that are beyond the time-frames and therefore beyond the competence of (neo)ecology. The hierarchical structure of ecosystems is the key to understanding their origins and development in deep time (Miller, 2008). Eldredge (2008) tied the ecological hierarchy rather tightly to the evolutionary hierarchy in his sloshing bucket theory of evolution—environmental-biotic perturbations begin at low levels and the interactions gear up through both hierarchies until we attain mass extinctions at the level of the biosphere. ‘[T]he larger the environmental jolt, the bigger the environmental reaction, an inherently hierarchical approach’ (Miller, 2008).

Thus these biological hierarchies are more than a mere organizing of data and its retrieval and communication (itself essential). They are natural in so far as they are based on discoverable patterns in nature; they are the locus and impetus for developing deep insights and theoretical advances into the evolution of the biosphere. Extrapolation versus emergence is far from settled and the hierarchies are significant players.

(viii) Taxonomy, classification, systematics

Classification is the ordering and arranging of organic diversity, the study of which is systematics, and giving the overarching philosophy and rationale is taxonomy (Simpson, 1961; Mayr & Ashlock, 1991; Mayr and bock, 2002). The New systematics (Huxley, 1940) arose as intrinsic to the population thinking at the heart

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of the Modern Synthesis. Three schools of macrotaxonomy demanded attention in that period (Mayr & Ashlock, 1991). Traditional (Darwinian) Evolutionary taxonomy, thought through by Simpson (1945, 1961), goes back to Darwin (1859, Ch XIII) who saw the need to balance genealogy (descent) with divergence (similarity). (The claim (Mayr & bock, 2002, with references) that Darwin sought that balance is disputed (Padian, 1999, 2004; Ghiselin, 2004).) Phenetics (Sokal & Sneath, 1963), gives primacy to similarity. Cladistics (Hennig, 1966) gives primacy to the branching points of descent—of reconstructing the clade-splits into the right order as signalled by inferred specializations, or evolutionary novelties.

Figure 15. The essence of cladism (Prothero, 2009a, Fig. 5.4, 5.5) as opposed to evolutionary classification, known to cladists as ‘gradism’. Each diagram, left and right, shows an uncontested phyletic succession. Above, a traditional evolutionary classification is shown for each diagram, emphasizing divergence as grades (the birds from all reptiles, the humans from apes) and permitting paraphyly. below is shown for each the strictly monophyletic succession in which paraphyly is forbidden. Redrawn with permission of D.R. Prothero.

Prothero (2009a) demonstrated the core of cladistics with his customary clarity (Fig. 15). The cladists attacked the influential and probably dominant evolutionary taxonomy of the 1960s (especially Simpson, 1961; also Mayr, 1969) on their paraphyly and on tolerance of some polyphyly in their taxa, justified in their intent to balance divergence with genealogy. For the evolutionary taxonomists the marked divergence of birds and hominids from their nearest respective relatives (and birds have diversified greatly as well) should be expressed in their classification. but for the cladists that leaves the Reptilia and Pongidae as nothing-taxa—Reptilia are not-mammals and not-birds; Pongidae are not-monkeys and not-hominids; such taxonomy is not rigorous, not consistent, not falsifiable (the cladists discovered Popper in the 1970s: Hull, 1999) and not scientific.

Hennig’s disciples claimed too that palaeontology was not the only keeper of the deep-time perspective and that other disciplines could reveal phylogenetic patterns (Williams & Ebach, 2004, 2008). A paper by Zangerl (1948, p. 358), one of the morphologists discontented with the Synthesis, included this:

The widely held opinion that paleontology alone among the morphological sciences could claim to have a voice in the discussion of evolutionary problems, because of the ‘historic’ nature of the materials, is a distinct fallacy.

Reforming this allegedly stunted outlook entailed (Williams & Ebach, 2004):

replacing the central role palaeontology once played with biogeography, adding a spatial dimension to the concept of phylogeny. This approach to phylogeny replaces the old “transformationist” view with the cladistic view, the latter dependent on discovering relationships among taxa. Numerical phylogenetic methods are inherently “transformationist” and have replaced stratigraphy as the key to phylogenetic relationships. Numerical methods in systematics and biogeography are inherently transformational and suffer the same problems as the old palaeontology ... computer algorithms have replaced the older palaeontological method (stratigraphy) and nucleotides have replaced fossils as the sure and certain guide to the course of phylogeny ...’

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The turbulence of those taxonomic times (Hull, 1988) passed micropalaeontology by (McGowran, 2005, Ch. 4). Foraminiferal systematics tended to be typological, top-down, pre-Darwinian, with Glaessner (1945, 1955) the most prominent of the few exceptions. The rise of detailed phyletic reconstructions of the planktonic foraminifera exposed the rampant convergence among rather few shell forms and brought about evolutionary classifications (McGowran, 1968, 1971). In the numerous areas of palaeontology where cladistics has taken hold the gaps in lineages can be great and ghost lineages are common. Cladistic methods in planktonic-foraminiferology integrate molecular with phenotypic data in the Holocene and extrapolate the inferred outcomes back into the earlier Neogene (Darling & Wade, 2008). Cladistics has been advocated among the extinct clades, but taxonomic precepts in recent definitive monographs on the Paleocene (Olsson et al., 1999) and Eocene (Pearson et al., 2006) planktonics are stratophenetic—meaning that temporal ranges can be employed as data. The difference is that the phenotypic fossil record of the planktonic foraminifera is much more complete than are even the iconic examples of evolutionary palaeontology such as the horse family (McGowran, 2005), and much more so again than are the records of fish and plants that triggered the ‘cladistics revolution’ and the ‘reform’ of palaeontology. A new stratophenetic reconstruction of the macroperforate planktonic foraminifera throughout the entire Cenozoic (Aze et al., 2011) takes another big step towards a complete family tree. by the standards of chrono-resolution and completeness considered normal in micropalaeontology, all other data bases of fossils, even the fossil records used to construct the patterns of horse evolution in Fig. 14, are meagre.

(ix) Chronofaunas and community in deep time

The chronofauna was christened by Olsen (1952, 1983) who recognized the persistence through geological time of community types. Networks of species were held together through local variants of the overall environment; those species can be replaced by others in more or less the same role without destroying the structure of the chronofauna. There is stability and co-adaptation whilst evolution proceeds. Studying the patterns of ecological organization and change through long periods of time, i.e., the anatomy of chronofaunas, is the discipline of evolutionary palaeoecology (Wing et al., 1992). Very similar to chronofaunas are the ecological- evolutionary units (EEUs) of boucot (2009), whose central concept was the community group, a stable association of genera whose content changes more in the uncommon species. Community groups typically take 1–2 Myrs to become established and are fixed for millions of years, sometimes lingering longer. Also similar, from a somewhat different standpoint, is coordinated stasis (brett and baird, 1995; brett et al., 1996; brett, 2012), in which punctuation occurs collectively in both ecological structures and clades. Yet again, the ‘turnover pulse’ (Vrba 1985, 1995) assumes the phenotypic conservatism and concentration of phenotypic change in punctuated speciation, with the important addition of synchronism among lineages.

Referring to community groups but referrable to the others is boucot’s exuberant ‘reconciling Darwin with d’Orbigny’, reconciling the gradual in hierarchy with the punctuated, instead of the usual opposing of the two worldviews ((vii), above, and Table 2).

Chronofaunas were conceived amongst Permo-Triassic terrestrial vertebrate faunas and caught on in the Cenozoic North American vertebrate culture (Webb, 1984), and the turnover pulse was based on the African Neogene vertebrate record. EEUs and coordinated stasis come from Palaeozoic neritic invertebrate faunas. These matters and the next were reviewed by DiMichele et al. (2004) and McGowran (2005, Ch. 6).

(x) Environmental impact on evolution?

It has often been noted that Darwin moved away from interactions between organism and physical environment in his later thinking on evolutionary dynamics and moved closer to organism/organism interactions, culminating in Raup’s accusation ‘It was all competitive replacement’ (above). but the attention paid in recent years to mass extinctions has tended to be at the expense of testing for lower-level punctuations and their correlations (or not) with the well-established environmental punctuations or caesuras, especially in the late Phanerozoic stratigraphic and geohistorical record.

lieberman and Melott (2012) recounted the history of the debate about whether periodic astronomical phenomena shaped large-scale patterns in the history of life; they concluded that there is some evidence that periodicity may be real and they discuss the possible role of our solar system motion in the galaxy.

Prothero (2004) cast a very sceptical eye over the three most popular extrinsic environmental events (climatic shifts, ET impacts, volcanic events) in relation to the North American land mammal record, itself in a rigorous

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framework of correlation and age determination. He found no appreciable effect. The most spectacular non- effect was at the end of the Eocene and glaciation Oi-1—the first major Antarctic glaciation heralding the Neogene icehouse state—where abundant regional evidence confirmed profound environmental shifts but the land-mammal communities did not notice (Prothero and Heaton, 1996). If evolutionary change is not extrinsic it must be intrinsic, Prothero concluded, but the how and why were unknown.

Figure 16. Successions of chronofaunas in the three environmental realms from the Eocene into the Oligocene—the most critical transition from the greenhouse world of the Cretaceous and Palaeogene to the icehouse world of the Neogene and today. Excess nutrient caused the first crisis in planktonic foraminifera in the open ocean, coevally with a similar crisis in two successions in the neritic and coevally with the chronofaunal turnover in terrestrial mammals. The reverse happened at the second crisis which was caused by the onset of modern-type Antarctic glaciation (event Oi-1). The central point is that long-lived (14–15 million years) and deep-time ‘community’ entities turned over in concert in all realms. See McGowran (2009, 2012) and Kamikuri and Wade (2012) for further discussion and sources.

However, extrinsic influence on organic evolution is suggested by another approach—by correlations between the three environmental realms. Inspect the successions spanning the Eocene/Oligocene boundary (Fig. 16). long-lived chronofaunas in the three realms turn over at the Middle/late Eocene boundary. The dominant photosymbiotic planktonic foraminifera in the pelagic realm went extinct coevally with numerous radiolarians (in a quite different nutritional regime), and in a very likely cause-and-effect, the similarly long-lived deep- ocean benthic foraminiferal fauna lurched toward turnover. Almost certainly coevally the flourishing photosymbiotic benthic foraminifera in the Tethyan neritic went extinct, being replaced by algae. This was a wholesale shift towards eutrophy in the global ocean! (McGowran, 2009). The chronofaunal turnover in the North American mammals within the Duchesnean Age was similarly profound (and acknowledged by Prothero). These major caesuras in three realms were coeval. How the marine evidence relates to the terrestrial is unclear—yet the correlations are persuasive enough to indicate that Prothero abandoned extrinsic factors as terrestrial drivers prematurely.

For Alvarez (2009) it was because of the Cretaceous/Paleogene catastrophic, extraterrestrial impact (Alvarez et al., 1980) that he could announce: ‘After a century and a half, the uncompromising uniformitarian

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gradualism of lyell was dead’. However, the shift in worldview away from gradualism and into strongly episodic geohistory and biohistory was well underway by 1980 (McGowran, 2005, Ch. 8). Meanwhile, Keller (2012) has argued for many years on biostratigraphic grounds that the biotic and environmental effects of this large impact have been vastly overestimated—and likewise for all other large impacts.

General discussion and summary

The bare outlines of two-centuries’ saga of palaeontology in earth and life history can be sketched with two recurring themes and three caesuras. The themes on the one hand are population and ecology, and variation and inheritance (transmission), and on the other, morphology and embryology (development). From the former emerged variational evolution; from the latter, the transformational and saltational modes of evolution. The first caesura was realizing the possibility of geohistory and biohistory and accordingly establishing a scientific enquiry into the fossil record. There were two key figures, Cuvier and lamarck. The second caesura was the Darwinian revolution, without which nothing in biology makes sense, in Dobzhansky’s famous observation, and named after the perpetrator, crucially a geologist turned biologist. The third caesura was the Darwinian restoration and the key figure in palaeontology and macroevolution figure was Simpson.

Also recurring were environmental change vis-à-vis organic evolution and a uniformitarian/catastrophic dualism used frequently as shorthand in both.

Geohistory and biohistory took hold as Cuvier overthrew the deductivist, speculative, top-down geotheories of the 18th century with hard-won, bottom-up stratigraphic and morphological evidence supporting his own theories. In the span of Cuvier’s own scientific lifetime there were established the reality of entirely vanished communities in the neritic and terrestrial environmental realms (if not yet in the pelagic), the reality of organic extinction, and the basic principles of biostratigraphy making possible a fossil-based geological time scale controlling full-blown geohistory and biohistory. This was the palaeontological synthesis of ~1830, rich in evidence and hypothesis, proceeding circum-globally with all scientific rigour and becoming one of the— perhaps the—most impressive scientific achievement of the earlier 19th Century. (The widely-acclaimed historical study, The Age of Wonder (Holmes, 2008), neglected the saga of geohistory and biohistory; clearly the author is a physicalist more interested in mechanics and technology.)

There were casualties (Fig. 3). The ahistorical, steady-state or cyclical mindset of the 18th century disappeared along with the rock-based ‘Neptunian’ time scale; even though lyell tried to resurrect steady-state eternalism. The scala naturae or Great Chain of being went too. Out of Goethe’s idealistic morphology and sundry nature-philosophy, romanticism, typology and transcendentalism there emerged anti-evolutionist morphology and embryology and the anatomist and palaeontologist Owen, the last major practitioner of that school. All of this was without evolution. Those people were groping upwards in search of the fourth level of science, explanation, to cap the three levels of descriptions, comparative studies, and generalizations as to relationships (Simpson, 1959).

Palaeontology and natural history revealed the richness of ancient and modern diversity and morphology, and built the geological time scale, without an overwhelming need to answer two fundamental, explanatory questions: whence all this diversity, and how come there has been so much and such incessant change? The options were lamarck’s evolution, spontaneous generation, biblical creationism, and reverent silence, none satisfactory. lamarck deserves all credit for what he achieved as a systematist but although he was the major evolutionist of his time his transformationalism did not flourish. Russell (1916) observed that opposition by the two strongest evolution-deniers, von baer and Cuvier, was not so much to organic derivation and transmutation as such as to the lack of tangible and compelling evidence. but it must be added that lamarck’s transformationalism implied taxa that would have been too unstable to support either a biostratigraphic succession and correlation or an evolutionary taxonomy.

‘Much of Darwin’s genius was of ... infusing evolutionary significance into observations and generalizations already established in a different context’ [in this case reinterpreting the work of the idealistic morphologists and embryologists] (Simpson, 1960, p. 173n). Darwin achieved the above-mentioned fourth level of science, almost single-handed and almost in one leap (Simpson, 1959).

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Although a parade of Darwin’s peers in palaeontology and geology could not accept evolution, their successors could—but in the fact of evolution and in the excitement of discovery and phylogenetic reconstruction, not in natural selection as the mechanism. Again, the transformationalists were discontented—their disciplines supported and were reinvigorated by evolution but did not contribute to the process itself. The high-profile palaeontologists Cope and Hyatt ‘were but the tip of an iceberg of non- and anti-Darwinian thinking moored firmly across the flood-tide of evolutionism’ (bowler, 2005). From evolutionary palaeontology, post-Darwin, emerged a lamarckian structuralism, orthogenesis and saltationism as formal or structural constraints on development and mysterious internal drives came to overshadow function and adaptation. As morphology and embryology decayed into the 20th century (laubichler & Maienschein, 2007) palaeontology produced the idiosyncratic theories of evolution and biostratigraphy of the North American School, and the German Synthesis emerged.

Meanwhile, the theory of variational evolution of A.R. Wallace and Darwin was grasped by Fritz Müller (1869), Ernst Haeckel (e.g. 1876), Vernon Kellogg (1907) and very few others in the decades before the Restoration. The Darwinian Restoration occurred when palaeontology, most prominently through Simpson, presented a panorama of macroevolution which no longer supported the anti-Darwinian theories of process and development through mysterious internal forces. Meanwhile and concomitantly, inheritance and mutational genetics gave way to population genetics, which fused with natural history and biogeography in a theory of microevolution and speciation.

This essay has favoured heavily the ‘internalist’ view of science, mostly avoiding its professional, cultural and political and sociological, ‘externalist’ aspects (e.g., Junker, 1996). It supports the ‘presentist’ habit of interpreting the past to some degree in terms of the present rather than entirely on its own terms (e.g., Hull, 1989; Mayr, 1982). More importantly here, palaeontology is flanked by and interacts with biology on one side and geology on the other, the interactions rising and falling down the decades and the geological, deep-time factor being neglected by shallow-time, neontological theorists. Cuvier’s catastrophism expressed as revolutions (Fig. 5) was bolstered by Elie de beaumont’s tectonic theory (Rudwick, 2005) and was amplified in d’Orbigny’s erecting of stages. Darwin’s physical worldview owed a lot to lyell’s gradualism. The contrasting theories of Schindewolf and Simpson have under-acknowledged contexts. Schindewolf had studied with the tectonic geologist Hans Stille (1876–1966), well-known for his episodic, punctuated world view expressed in global synchronous orogenic cycles. Simpson (1952) was part of the (1940s–1960s) powerful consensus among opinion leaders in stratigraphy, tectonics, palaeontology and evolutionary biology that the record in the rocks is episodic preservationally but gradualistic in its message, well accommodated in lyellian uniformitarianism and unsympathetic to Stille’s tectonism and its successor, Umbgrove’s (1947) Pulse of the Earth. The world does not change in so stately a manner as it did when Simpson was writing magisterially; the worldview has shifted (McGowran, 2005, Ch. 8). Plate tectonic shifts and bolide impacts are only part of our mindset, which is that environmental change through thresholds and feedbacks is always episodic.

From the palaeontological standpoint, Gould (2002 and numerous earlier writings) devoted much effort to revising Darwinism by attacking (what he perceived as) the excessively extrapolationist and adaptationist aspects of the Darwinian Restoration, micro-to-macroevolution, and to pressing the case for punctuated speciation and hierarchical expansion (Erwin, 2002), as did Eldredge’s (1985) Unfinished synthesis. Evolution is an expanded synthesis (Kutschera & Niklas, 2004). The ‘revolution in palaeobiology’ has been presented as an ‘incredible ascendance’ over the Simpsonian palaeontology of the Restoration (Sepkoski & Ruse, 2009). There are expectations of a Simpson-Schindewolf rapprochement (Princehouse, 2009), and clear if minority disagreement with the theory of punctuated equilibrium (boucot, 2009). Advances in Darwin’s most oppressive blank, Precambrian palaeontology (Schopf, 2009; brasier, 2009), are accompanied by assaults on the universal Tree of life itself, such as that lateral gene transfer among prokaryotic lineages seems to exclude prokaryotes from the Tree (Doolittle, 2010). Emphasized are the influences on evolutionary palaeontology of developmental genetics, molecular taxonomy/phylogeny, and cladistics.

There is no overarching theory signifying a new paradigm in palaeobiology. I see the rise of palaeobiology as a series of pronounced but natural developments within the Darwinian Restoration. This would have been clearer had attention been paid to an advance as spectacular as any: namely, reconstructing micropalaeontologically

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the palaeoenvironmental panorama in the pelagic realm through deep time. but I agree strongly with Jackson and Erwin (2006) that Simpson’s great synthesis of genetics, evolution and palaeontology in Tempo and Mode formed the core of modern palaeontology.

Even so, this statement by one of those authors holds true, too:

[Population genetics] has established evolutionary biology as a far more robust discipline than was the case earlier, but the Modern Synthesis is a curiously ahistorical view of a historical discipline. beyond the fact that it provides little insight into how form evolves (something we know now a great deal about from comparative studies of molecular developmental biology), the Modern Synthesis is silent (and indeed probably antagonistic) to issues such as whether the nature of variation upon which selection can act has changed systematically over time, whether the relative significance of selection, mutation and genetic drift (the principle [sic] drivers of evolution) has changed over time, or how the changing structure of ecological relationships has altered evolutionary ooportunities through time.

(Erwin, 2011, p. 35; emphasis added). Darwinism in palaeontology is extended hierarchically (Table 3). long-lived standoffs on deep-time lineage, community and causation are resolved or resolvable here. Gould (2002) embraced the ontological species- as-historical-entity and extended it down to the gene and up to the clade. Three tiers are occupied by, in ascending order, gradualist anagenesis, punctuated cladogenesis, and coordinated or concurrent, sometimes mass extinction. boucot (2009) summarized his notion of the ecological-evolutionary unit (EEU), of which there are about 40 in half a billion years, bounded by accelerated taxic turnovers (‘d’Orbignyan biostratigraphy’). Each EEU contains several biofacies to be reconstructed as community groups within which individual species’ turnovers give biozonations (‘Oppelian biostratigraphy’). This is evolutionary palaeoecology, beyond the reach of and not predicted by the ecology of shallow-time neontology.

Table 3. Hierarchy of historical entities

IIIOrganism- individual

IICell- individual

IGene- individual

Hierarchical expansion of Darwinism in palaeontology. Gould (2002) argued for non-fractal and non-extratrapolational hierarchy in the individuated organic world, Darwinian individuals (i.e., historical entities) being produced by tiered evolutionary processes at their respective ranks. Speciational and transformational evolution are both variational, i.e., ‘Darwinian’, but hierarchical (Mayr, 1992). Community evolution in deep time, with d’Orbignyan and Oppelian biostratigraphy in hierarchical relationship, is from boucot (2009). Scale-dependent causation models, Court Jester and Red Queen, from benton (2009).

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Again, we have had since the 19th century the opposition of two families of models for evolutionary causation, one lot going back to Darwin’s competition and the other to environmental ‘catastrophe’. benton (2009) resolves the contest of the Red Queen (biotic causation) versus the Court Jester (abiotic causation) temporally, in this way: the Red Queen operates locally at short time scales and the Court Jester operates regionally and globally at longer time scales. However, using the unparalleled phylogenetic record of the planktonic foraminifera for the entire Cenozoic Era (Aze et al., 2011), Ezard et al. (2011) showed that macroevolutionary dynamics depended on the interaction between species’ ecology and changing climate— that is, neither the Court Jester nor the Red Queen hypothesis is dominant. Studies such as this herald the use in macroevolution and palaeobiology of the deep-sea microfossil record, richer and more complete by orders of magnitude than the record available to Darwin or the Darwinian Restoration.

Darwinism was restored, but as a thriving group of research programs, of course Darwinism has gone beyond Darwin!

Acknowledgements

This paper began as a lecture to the Royal Society of SA and the SA Museum celebrating the two Darwin centenaries in 2009. I thank Oliver Mayo and Ole Wiebkin, Editor and Guest Editor respectively, for their most positive reception and their efforts in planning and bringing to fruition this special volume. lisi McGowran helped greatly with the figures. The manuscript was been read critically and encouragingly by bill berggren, Ole Wiebkin, Oliver Mayo, John Cann, Qianyu li, and an anonymous referee. As ever, Jennifer Thurmer’s expertise and patience turned this manuscript into a publication.

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