Friday, June 12, 2020

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 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.

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