Evolution Through Punctuated Equilibrium: History Of An Idea

Palaeontologist Niles Eldredge explains how one of the most famous papers on paleontology and evolution came to be published: 

Steve was determined to be a part of Tom’s plan to do a GSA symposium and publish a book of essays on this new-fangled concept of “paleobiology.” Tom had a list of topics and was shopping around for speakers to be assigned to each one. When Steve saw the list, he told me that he had first wanted “morphology”—but that was already assigned to Dave Raup. So he opted instead for “phylogeny”—but that had been grabbed up by Mike Ghiselin. That left only “speciation,” the last of the evolutionarily imbued topics on Tom’s list, as yet unassigned. Steve called me up, explained the situation, and said he had settled for speciation—but could not think of anything much to say about it beyond the manuscript I had written and recently submitted to Evolution—there of course being no Paleobiology as yet. “The Allopatric Model and Phylogeny in Paleozoic Invertebrates”—a distinctly un-Gouldian, plodding, if accurate, title (Eldredge 1971). Without Ralph Gordon Johnson in the editorial chair of Evolution at that time, I doubt that that early paper would have been accepted. As it was, it was likely to have gone relatively unnoticed—had not Tom come along, Steve grabbing “Speciation”—and Steve asking if we could coauthor the paper along the basic lines of my first effort. He was stuck with “speciation,” and couldn’t think of anything much to say beyond what I had said in the Allopatric Model manuscript.

This passage is from an article by Niles Eldredge titled Reflections On Punctuated Equilibria, published in a recent issue of Paleobiology. The 1972 paper he refers to, coauthored with Stephen Jay Gould, was, Punctuated equilibria: an alternative to phyletic gradualism. It marked the beginnings of a long debate on how to interpret the patterns of morphological change observed in the fossil record. Do species remain in stasis, showing little morphological change through much of their existence as Eldredge and Gould argued? Do periods of rapid morphological change coincide with the origin of new species (speciation)? Supporters hailed it as a revolutionary work. Critics called it 'evolution by jerks', a jibe aimed not just at the patterns of change.

Dr. Eldredge provides a very insightful look at the history of this idea including some fascinating snippets on Darwin's thinking about divergence and species origins. For Darwin, change accumulates incrementally over long passages of time. Divergence via natural selection can give rise to descendant varieties even without geographic isolation of a population. Later thinking has given more importance to exogenous factors like climate change in causing habitat fragmentation and reproductive isolation. Populations gets geographically isolated first, and then diverge from the ancestral species either through natural selection or random genetic drift. Eldredge and Gould applied this idea to the fossil record and emphasized that the sudden appearance of new fossil species is a manifestation of long periods of stability interrupted by episodes of isolation and geologically rapid shifts in morphology (allopatric speciation).

There is a lot to take in and think about the long term patterns of change preserved in the fossil record. But it is enriching reading. The article is open access.

Links: Birth Of Species, Children's Book, Rise Of Oxygen

 Some geology and evolution material for your perusal.

1) I came across this well crafted documentary by Niles Eldredge and Stefano Dominici on the history of the development of ideas on the birth of species. It is a paleontologist's perspective with fossils being given the centre stage. A great many personalities who contributed to the early thinking on the origin of species are featured. Among the prominent ones who influenced Darwin were Lamarck, Cuvier, Giambattista Brocchi, and John Herschel to name a few. Completely left out of this film is Alfred Wallace. 

It does center around Darwin, and, later towards the end, on Niles Eldredge's work on Punctuated Equilibrium, which he published in collaboration with Stephen Jay Gould. I have often wondered whether there was any tension between Eldredge and Gould regarding proprietorship over Punctuated Equilibrium given Gould's very bombastic advocacy of this idea. The last section of this film is revealing! 

Beautifully compiled. Do watch. 

 

If  you are unable to access the embedded video, view it at this permanent link - The Birth of Species. 

2) Zircon (zirconium silicate) is a remarkable mineral. Born in the cauldron of magma chambers, it is henceforth virtually indestructible unless it is melted down again. This makes it a witness to geological processes affecting and shaping terrains over hundreds of millions of  years. What a story it has to tell us. And that is precisely what geochronologist Matthew Fox has done. He has written a children's book titled Jane's Geological Adventure, which follows Jane, the zircon grain, from her birth in a magma chamber to a life lasting 400 million years. Alka Tripathy-Lang reviews the book.

Meet Jane, the Zircon Grain—Geochronology’s New Mascot.

3) Elizabeth Pennisi writes about a new paper published in Nature Geosciences on the link between the increase in the length of the day and the rise of atmospheric oxygen on early earth. 

‘Totally new’ idea suggests longer days on early Earth set stage for complex life.

Punctuated Equilibrium Is About Small Subtle Changes

The theory of punctuated equilibrium (PE) put forth by paleontologists Niles Eldredge and Stephen Jay Gould proposes that morphological evolution speeds up during lineage splitting events when new species form. In practice, a paleontologist sampling fossil species in a sedimentary section will find that a species over its lifetime does not show any trend in morphological changes. That species eventually goes extinct at a particular interval. Fossils of the inferred descendant species appear abruptly in the sedimentary bed above. A fossil population showing a mix of traits diagnostic of both species is not generally seen. 

Eldredge and Gould argued that this pattern of change observed in fossil species shows that the history of a species is really characterized by long periods of no change, eventually interrupted by rapid evolution to a new species.

A reassessment of one textbook example in bryozoans suggests no evidence of such a punctuated mode. 

A write up in Phys.org describes the new work. 

Revisiting a Landmark Study System: No Evidence for a Punctuated Mode of Evolution in Metrarabdotos - Kjetil Lysne Voje, Emanuela Di Martino, and Arthur Porto

I am not going into the work itself. There are well documented examples of the punctuated equilibrium mode of evolution while others as in this case may see revision from time to time. 

Instead, I wanted to comment on one of my pet peeves about this topic which is the common use of the phrase "major evolutionary change" to describe the actual evolutionary changes.

This idea which never seems to go way that PE involves big or major changes has caused a lot of misunderstanding about the theory.  As the authors of PE took pains to point out, the amount of morphological difference between ancestor and descendant species is subtle, often taking excruciating examination to recognize. 

Here is Niles Eldredge describing his work on Devonian trilobites.

I measured some 50 different lengths and widths- length of the head, distance between the eyes, height of the eyes, length of the tail and so on -on hundreds of specimens. This was tedious. Each specimen had to be cleaned to at least well enough to make all the anatomical landmarks visible. Each had to be mounted on a block of wood, stuck to a blob of plastilene, with the tops of the eyes (a flat surface) in a horizontal plane, perpendicular to my line of sight down the barrel of the microscope. There was a little scale inside the right eyepiece reticle from which to read off the various measurements. After a day of measuring, and until I got used to the microscope, I would see double from the bus window on my way home. 

 from Time Frames: The Evolution of Punctuated Equilibria.

Trilobites have compound eyes, made up of columns of lenses. All this detailed work led to the recognition that a new species had 17 columns of lenses, instead of 18 in the ancestral species found in slightly older strata!

This point about small changes was missed by many observers. Creationists conflated PE with a theory of macro-mutation or saltation, a sudden origin of big morphological change as for example seen in the transition from reptiles to mammals. They claimed that Darwin with his insistence on gradual accumulation of small changes had been wrong all along! 

Not so. The morphological differences observed involved small changes between the ancestor and descendant species.  It takes an expert to recognize this shift in form. 

The real significance of PE is not about the amount of change, but the pattern. PE proposes that very little morphological divergence takes place during the lifetime of a species. It is only when a sub-population of that species gets isolated that it may experience relatively rapid evolution. This genetic isolation results in the formation of a new species. The ancestral species may persist in the main parts of its range, while one of its peripheral populations has evolved into a new species. Biologists term this process of branching or lineage splitting as cladogenesis. 

PE is about explaining these rhythms in the history of lineages. Long periods of stasis (little or no change in form), punctuated by relatively rapid transition to a new form. This transition, though rapid in geologic time,  may take place over hundred of generations and thousand of years.

Why then isn't this transition to a new form preserved in the rock record as a fossil population with a mix of characters? The answer Eldredge and Gould prefer is that reproductive isolation takes place in outlier regions of a species range. Sediments deposited in these isolated basins will be archiving the incremental evolution of a new form, just as Darwin envisaged,  via a mixed or transitional population. However, these outliers containing transitional populations have less chance of getting preserved in the rock record.  The sedimentary sections that paleontologists examine are generally from the main part of the basin. This is not where the change to a new species has taken place.  The sudden appearance of an inferred descendant species in this strata is really recording the migration of the descendant from an isolated part of the species range where it evolved, into the central range of its extinct ancestor. 

Biologists have named this process of formation of a new species through reproductive isolation in a geographically distant area as allopatric speciation. Eldridge and Gould's theory of punctuated equilibrium invokes allopatric speciation and migration to explain the abrupt appearance of new species in the fossil record. In terms of the mechanisms of change, it is not an alternative to Darwinism as is often portrayed. Both the authors accept that even during the period of 'rapid' evolution, changes accumulate through natural selection or genetic drift incrementally across generations. But the vagaries of preservation means that sediments which record this transition are rarely available for study.