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Organic Evolution in Deep Time: Charles Darwin and the Fossil Record
Brian McGowrana
aSchool of Earth and Environmental Sciences The University of Adelaide, Adelaide SA 5005, Australia
Published online: 13 Oct 2014.
To cite this article: Brian McGowran (2013) Organic Evolution in Deep Time: Charles Darwin and the Fossil Record, Transactions of the Royal Society of South Australia, 137:2,
To link to this article: http://dx.doi.org/10.1080/03721426.2013.10887188
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Transactions of the Royal Society of South Australia (2013), 137(2):
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
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
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
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
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ORGANIC EVOlUTION IN DEEP TIME: CHARlES DARWIN AND THE FOSSIl RECORD
the geological past. And (v) the tree of life is underlain by an anastomosing web of life.
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
Figure 1. Four figures pivotal to any narrative of evolution and the fossil
It was an
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
The
Table 1. Fossil record and evolution: seven advances in two centuries’ progress
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Geohistory and biohistory highly integrated and multi- disciplinary in
phyletic reconstruction, speciation Oceanfloor spreading and plate tectonics
Darwinian Restoration (Darwinian Revolution II) The Modern Synthesis: reconciling palaeontology, field biology, and population genetics
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1910s |
III |
Radiometric calibration of earth history |
1870s |
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(Novelty creationism discarded as dead and rotten science) |
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II |
Darwinian Revolution I |
Organic evolution and universal historicism: |
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Evolution in deep time triumphs; natural selection on hold |
|
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(Genesis creationism (young earth) discarded as dead and rotten science) |
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I |
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The Palaeontological Synthesis: fossils and rock relationships |
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yield geohistory and biohistory and geological time scale |
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lamarck’s theory: evolutionism survives but theory fails |
Two centuries’ worth of progress: seven major advances and turning points (“revolutions”) in earth and life history and pertaining to their development. Strands of
Accepting Ernst Mayr’s argument that the Darwinian Revolution has two parts in separate centuries, we have a convenient structure for this
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ORGANIC EVOlUTION IN DEEP TIME: CHARlES DARWIN AND THE FOSSIl RECORD
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
(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
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
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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
(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)
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
I would add though that Darwinism should evoke some amalgam of variational evolution with a thoroughgoing historicism.
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
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Table 2. The three principle theories of evolutionism
Saltational evolution |
Changes take place through the origin of new types. The mutation theories of |
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19thC palaeontologists (and of 20thC geneticists who took over the word) belong |
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here. Schindewolf, the last macrosaltationist palaeontologist of major significance, |
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advocated that higher taxa emerged virtually instantaneously. Of very similar |
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views were bateson, the most influential advocate for Mendelian genetics, and |
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Goldschmidt, whose systemic mutations produced the notorious, postulated, |
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hopeful monsters. |
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(i) A given object changes through time, transforming without losing its identity, |
Transformational |
analogous to ontogeny (e.g. the changes from egg to old age in an organism). Internal |
forces dominate external; individual elements are the subjects in that changes within |
|
evolution |
themselves produce the evolution of evolution. There is recurring reference to |
(The organism |
unfolding, predetermination, preconditions, and development. before Darwin, all |
theories of historical change including organic evolution were transformational. |
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as subject of |
(ii) lamarck, the founder of the theory of organic evolution, was transformational in |
the evolutionary |
that theory, but stands apart from the transformationalist morphologists and |
process; internal |
embryologists in being a systematist concerned with variation, taxa and biodiversity. |
forces dominate) |
(iii) The evolution of cosmological entities, of magmas and continents, and of all other |
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geological and environmental configurations are transformational. All human- |
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historical and social changes identified as evolution are transformational. Prime |
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examples |
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theories of the psyche (Freud) and the body (Piaget). |
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(iv) Subsequent theories such as orthogenesis and finalism, along with all theories |
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ignoring or rejecting the reality of species as historical entities, are |
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transformational. Also transformational were the ‘eclipse of Darwinism’ in the |
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later 19th and early 20thC, the decline of evolutionary morphology and recovery |
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by |
Variational |
(i) In variational evolution there is an alternation of: (a) lavish production of internal |
variation through mutation and recombination, and (b) a new generation consisting |
|
evolution |
of a limited number of survivors of external pressures. This occurs only in organic |
(The organism |
evolution including the origin of new species. In that developmental pathways are |
the consequence of history, not its cause, variational evolution is quite different |
|
as object of |
from the continuing sense of transformational evolution. Haeckel’s theory of |
the evolutionary |
recapitulation looks strongly transformational, but it is variational and Darwinian, |
process; external |
for it held that ontogeny was the trace of the evolutionary past, was in no sense |
forces dominate) |
the image of its future, and was merely another target for selection. |
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(ii) If ‘Darwinism’ has an identifiable core, the core comprises two components. |
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The origin of species or speciational evolution is variational. The other Darwinian |
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component, descent with modification and |
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However, in an hierarchical expansion of Darwinism both are variational, the latter |
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at |
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
At any rate, evolutionary biology is historical science! Grasping this point at fundamental or philosophical levels entailed contrasting
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populational thinking where Darwin showed variation to be
The German palaeontologist H.G. bronn in the
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
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
The
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
(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
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,
Late Eighteenth Century view of the earth
Enlightenment knowledge was twofold, literary and
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
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by the end of the 18th century geognostic savants were comfortable with the fundamentals of rock relationships in the
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)
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
Taxonomy influenced the shift from
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
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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
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
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
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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
(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
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
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
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
Consider Figure 6, sketched by lyell to illustrate the conclusions of the French conchologists as they distinguished successional assemblages in the
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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
The palaeontological synthesis and the
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
(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
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Figure 7. Two figures from Phillips’ (1860) Life on Earth. left: the classical
Clearly stratigraphy and stratigraphic palaeontology achieved impressive levels of sophistication in the
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
... 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
(Arkell, 1933; in McGowran, 2005). Thus, we have here in the
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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
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
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Cuvier in turn established successional |
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and naturalistic extinctions but nothing |
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of value on origins. lyell, anti- |
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evolutionary and reverting spiritually |
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to Huttonian geotheory and a steady- |
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state, cyclical model of the earth, |
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could do no better than resurrect the |
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1814 suggestion of brocchi of the Sub |
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Apennines that species in analogy with |
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individuals had |
intrinsically |
limited |
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lifespans between their origin and their |
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extinction (Rudwick, 2008). |
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Mayr (1982, 1988) steadfastly saw the |
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19th century species problem in |
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Platonic and essentialistic terms. The |
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Platonic |
world |
comprises |
natural |
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kinds, discontinuous types, essences. |
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To move from one kind to another |
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Figure 9. Three Early Jurassic ammonites occur consistently in the same |
requires a pronounced jump, a |
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saltation. If species are natural kinds |
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stratigraphic order across western Europe, implying that they diagnose |
then saltations are required to achieve |
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successive slices of geological time. These are empirical observations, |
the origins of species, but there were no |
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susceptible to testing, corroboration and refinement without the benefit of |
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theories of speciation, extinction, or biogeography. This is a modern textbook |
acceptable naturalistic mechanisms on |
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example (Ziegler, 1972, Fig. 123), courtesy of E.Schweizerbart’sche |
offer. Also, for Cuvier especially but |
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Verlagsbuchhandlung (www.schweizerbart.de), but it exemplifies |
for others too, that a species might |
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century biostratigraphy. |
transmute implies a threat to species’ |
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stability, |
in turn |
a threat to |
orderly |
progress in
The
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
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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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
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
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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
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
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
Fossils, biogeography and genealogical inference
Richard Owen (1860) was impressed by the close correspondence between the modern mammalian faunas and the
Darwin could add this biogeographic configuration to others, as summarized schematically (Fig. 10). On the
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Figure 10. A schematic rendering of southern biogeography with the fourth dimension. Adaptations in an
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
First, evolutionism, the evolution of life on an
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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
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
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
Genealogical relationship explained taxonomic groupings just as it explained the patterns in the fossil record, in both cases compiled mostly by non- or
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
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It is a remarkable fact that morphology took but a very little part in the formation of
Russell also described how it was for the morphologists of the old school, the classical
That detaching of the accepted fact of evolution from the
... 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
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,
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Just as ‘the old man returns to second childhood |
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in mind and body’, so does the evolutionary |
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lineage rise (epacme), expand or flower (acme), |
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and contract (paracme); ‘[in] fact, there is no end |
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to the homological and analogical similarities and |
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parallelisms of ontogeny and phylogeny |
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wherever both are found complete’ (Hyatt, 1897). |
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Schuchert (1915, p. 904), a student of Hyatt, |
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stated in a famous textbook, ‘the ammonids were |
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making their last ineffectual stand. Old age was |
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upon them and their doom was foreshadowed in |
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the uncoiling, the unnatural twisting of the shells |
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... and the straight baculites ...The genus |
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Heteroceras [Fig. 11 herein, top] displays the |
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extreme of irregular growth’. |
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Hyatt’s theorizing was an extreme example of |
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orthogenesis, whose meanings ranged from |
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schemes, some |
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notions of finalism and vitalism. Cope included |
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saltations, meaning evolution by large jumps |
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(Table 1). This assortment of Neolamarckism, |
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saltation, linking of ontogeny to phylogeny, and |
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orthogenesis has in common a strong internal |
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drive, strongly rejecting natural selection and |
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culminating in a strongly |
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the |
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palaeontologists in Europe and North America |
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rejected Darwin’s understanding of either the |
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process or the pattern of evolution (Rainger, |
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1986; Reif, 1986). bowler (1988, 2005) analysed |
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this episode under such headings as the eclipse |
Figure 11. A ‘typical’, |
of Darwinism and the |
As for invertebrate palaeontology, more |
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lower left. The others represent various departures from ‘typical’, |
entwined in its biostratigraphy and biofacies with |
departures attributed during the |
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German Synthesis as being due to the lineages entering old age |
geohistory than with biohistory, Darwinism |
(‘racial senescence’) and lurching into inadaptedness then extinction. |
made very little impact on large tracts of that |
Drawings mostly from Schuchert (1915, Pl. 37 & Fig. 495). |
enterprise (bretsky, 1979). |
During this time evolution itself and the tree of life were not in doubt amongst the informed, but the recovery of Darwinism from the
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
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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
The German palaeontological culture developed a strong
[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
The school of typostrophism culminated in ‘the triumph of the
The typostrophist manifesto was from beurlen in 1932 (in Reif, 1993):
It is a very general rule that the pathway of evolution within a
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,
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
For some, the above gives too much space to the
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
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
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,
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
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
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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(b) in cleaning away all the theories and notions |
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accumulated in macroevolution, listed as rejected |
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in a preceding paragraph. Simpson agreed that |
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showing that palaeontology was not contradictory |
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to the genetics of that time ‘was one of my aims, |
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but I do not see how anyone who has really read |
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this book could fail to understand that it was not |
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my only or even my main aim. My main aim was |
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to explore and in a way to exploit the fact that |
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paleontology is the only |
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time, “tempo”, is inherent in it. Thus the aim of this |
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book, which I think it accomplished, was to bring |
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this dimension squarely, methodically, into the |
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study of evolutionary theory’ (1984, p. xxii; my |
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italics). It is noteworthy that Tempo and Mode was |
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widely and favourably reviewed with the |
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astounding exception of Simpson’s own discipline, |
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palaeontology, which produced but the one |
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mention discovered by laporte (1983). In that |
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review Jepsen (1946, p. 538) opined that the |
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contributions to interpretations of fact and |
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epistemology, such as rates of evolution, |
31): speciation or cladogenesis, phyletic evolution or anagenesis, |
explanation of gaps as more than sampling failure, |
and emphasis on adaptation as a continual theme. |
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Figure 13. The three modes of evolution (Simpson, 1944, Fig. |
For subscribers to the Synthetic Theory, Darwin’s |
and quantum evolution. The bell curves signify the |
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variation in each population. Simpson estimated that anagenesis |
three causal theories (Fig. 3), namely origin of |
was considerably more common than cladogenesis; modern |
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estimates tend to the reverse. In quantum evolution small, |
species, gradualism and natural selection, were |
peripheral, maladaptive populations almost always go extinct, but |
now fulfilled, completing the Darwinian |
the occasional exception breaks through to a new adaptive zone, |
Revolution, although I prefer Ghiselin’s (2005b) |
or way of life. Used with kind permission of Joan Simpson burns. |
Darwinian Restoration. Hull (1973, p. 77, my |
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
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ORGANIC EVOlUTION IN DEEP TIME: CHARlES DARWIN AND THE FOSSIl RECORD
(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
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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
Progress in later Phanerozoic biogeohistory
Simpson’s accomplishment in
(i) The rise and rise of micropalaeontology |
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Darwin’s biota, living and fossil, comprised invertebrates, vertebrates, and plants. Skeletonized microfossils |
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(phytoprotists and zooprotists) were missing. English micropalaeontologists’ |
using d’Orbigny’s catastrophism |
to denigrate his systematics misled Darwin into dismissing the |
evolutionary potential of the zooprotistan |
foraminifera, thereby impeding their study for decades (lipps, 1981, |
also quoted in McGowran, 2005, 2012). |
Systematic study of the global oceanic microbiotas flourished in the later |
19th century with the monographing |
of the most prominent groups, the foraminifera and the radiolarians. |
Apropos of the latter, the huge output |
by Haeckel was blighted by his careless attitude to collections and |
types, cursory descriptions and a highly |
arrtificial, |
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
(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)
(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
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
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ORGANIC EVOlUTION IN DEEP TIME: CHARlES DARWIN AND THE FOSSIl RECORD
are associated with icesheet expansion, cooling and precipitation changes. The authors infer that
(iii)Cenozoic environmental trajectory and
and atmospheric circulation and CO2 levels and triggered
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
(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
(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
(Kucera & Malmgren, 1998) were cited by Gould (2002). Evolutionary foraminiferal higher taxa and their relationships (e.g., Pawlowski, 2000). The with
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
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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
(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
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
Also hierarchical is ecology and environment. So long as palaeoecology was regarded as but a
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
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
Hennig’s disciples claimed too that palaeontology was not the only keeper of the
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,
(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
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
(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
lieberman and Melott (2012) recounted the history of the debate about whether periodic astronomical phenomena shaped
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
Figure 16. Successions of chronofaunas in the three environmental realms from the Eocene into the
However, extrinsic influence on organic evolution is suggested by another
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
General discussion and summary
The bare outlines of
Also recurring were environmental change
Geohistory and biohistory took hold as Cuvier overthrew the deductivist, speculative,
There were casualties (Fig. 3). The ahistorical,
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
‘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
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ORGANIC EVOlUTION IN DEEP TIME: CHARlES DARWIN AND THE FOSSIl RECORD
Although a parade of Darwin’s peers in palaeontology and geology could not accept evolution, their successors
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
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,
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,
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).
Table 3. Hierarchy of historical entities
|
|
|
|
|
|
|
level |
Darwinian |
Organism to Clade: |
beyond ecology’s reach: |
|
||
|
Individual |
|
of Historical Entities |
Community Evolution in |
causation models of |
|
|
|
|
|
Deep Time |
Evolution |
|
VI |
Clade- |
Third |
‘Mass’ extinction |
|
Court Jester (abiotic |
|
|
individual |
Tier |
between clades derails |
Unit turnover and |
causation: unpredictable |
|
|
|
|
punctuated speciation |
d’Orbignyan biostratigraphy |
changes in |
|
V |
Species- |
Second |
Punctuated and cladogenetic |
[~40 EEUs in Phanerozoic] |
physical environment) |
|
speciation undoes anagenesis; |
|
dominates Red Queen |
||||
|
individual |
Tier |
differential success within clades |
[regional and global, |
||
IV |
Deme- |
First |
[Speciational evolution] |
evolution and Oppelian |
long time scales] |
|
Anagenesis within populations |
biostratigraphy |
Red Queen (biotic causation |
||||
|
individual |
Tier |
in ecological time |
[several ecologically distinct |
||
|
|
|
[Transformational evolution; |
CGs in each EEU] |
especially competition) |
|
|
|
|
phyletic gradualism] |
|
dominates Court Jester |
|
|
|
|
|
|
[local, short time scales] |
IIIOrganism- individual
IICell- individual
IGene- individual
Hierarchical expansion of Darwinism in palaeontology. Gould (2002) argued for
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ORGANIC EVOlUTION IN DEEP TIME: CHARlES DARWIN AND THE FOSSIl RECORD
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
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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