The punctuated equilibrium model (top) consists of morphological stability followed by rare bursts of evolutionary change via rapid cladogenesis. It is contrasted (below) to phyletic gradualism, the more gradual, continuous model of evolution.
Punctuated equilibrium (also called punctuated equilibria) is a theory in evolutionary biology which proposes that once species appear in the fossil record the population will become stable, showing little evolutionary change for most of its geological history. This state of little or no morphological change is called stasis. When significant evolutionary change occurs, the theory proposes that it is generally restricted to rare and geologically rapid events of branching speciation called cladogenesis.
Cladogenesis is the process by which a species splits into two distinct
species, rather than one species gradually transforming into another.
Punctuated equilibrium is commonly contrasted against phyletic gradualism, the idea that evolution generally occurs uniformly and by the steady and gradual transformation of whole lineages (called anagenesis). In this view, evolution is seen as generally smooth and continuous.
In 1972, paleontologists Niles Eldredge and Stephen Jay Gould published a landmark paper developing their theory and called it punctuated equilibria. Their paper built upon Ernst Mayr's model of geographic speciation, I. Michael Lerner's theories of developmental and genetic homeostasis, and their own empirical research. Eldredge and Gould proposed that the degree of gradualism commonly attributed to Charles Darwin is virtually nonexistent in the fossil record, and that stasis dominates the history of most fossil species.
History
Punctuated equilibrium originated as a logical consequence of Ernst Mayr's concept of genetic revolutions by allopatric and especially peripatric speciation
as applied to the fossil record. Although the sudden appearance of
species and its relationship to speciation was proposed and identified
by Mayr in 1954, historians of science generally recognize the 1972 Eldredge and Gould paper as the basis of the new paleobiological research program.
Punctuated equilibrium differs from Mayr's ideas mainly in that
Eldredge and Gould placed considerably greater emphasis on stasis,
whereas Mayr was concerned with explaining the morphological discontinuity (or "sudden jumps") found in the fossil record.
Mayr later complimented Eldredge and Gould's paper, stating that
evolutionary stasis had been "unexpected by most evolutionary
biologists" and that punctuated equilibrium "had a major impact on
paleontology and evolutionary biology."
A year before their 1972 Eldredge and Gould paper, Niles Eldredge published a paper in the journal Evolution which suggested that gradual evolution was seldom seen in the fossil record and argued that Ernst Mayr's standard mechanism of allopatric speciation might suggest a possible resolution.
The Eldredge and Gould paper was presented at the Annual Meeting of the Geological Society of America in 1971. The symposium focused its attention on how modern microevolutionary
studies could revitalize various aspects of paleontology and
macroevolution. Tom Schopf, who organized that year's meeting, assigned
Gould the topic of speciation. Gould recalls that "Eldredge's 1971
publication [on Paleozoic trilobites]
had presented the only new and interesting ideas on the paleontological
implications of the subject—so I asked Schopf if we could present the
paper jointly."
According to Gould "the ideas came mostly from Niles, with yours truly
acting as a sounding board and eventual scribe. I coined the term punctuated equilibrium and wrote most of our 1972 paper, but Niles is the proper first author in our pairing of Eldredge and Gould." In his book Time Frames
Eldredge recalls that after much discussion the pair "each wrote
roughly half. Some of the parts that would seem obviously the work of
one of us were actually first penned by the other—I remember for
example, writing the section on Gould's snails. Other parts are harder
to reconstruct. Gould edited the entire manuscript for better
consistency. We sent it in, and Schopf reacted strongly against it—thus
signaling the tenor of the reaction it has engendered, though for
shifting reasons, down to the present day."
John Wilkins and Gareth Nelson have argued that French architect Pierre Trémaux proposed an "anticipation of the theory of punctuated equilibrium of Gould and Eldredge."
Evidence from the fossil record
The fossil record includes well documented examples of both phyletic gradualism and punctuational evolution. As such, much debate persists over the prominence of stasis in the fossil record. Before punctuated equilibrium, most evolutionists considered stasis to be rare or unimportant. The paleontologist George Gaylord Simpson, for example, believed that phyletic gradual evolution (called horotely in his terminology) comprised 90% of evolution. More modern studies, including a meta-analysis
examining 58 published studies on speciation patterns in the fossil
record showed that 71% of species exhibited stasis, and 63% were
associated with punctuated patterns of evolutionary change. According to Michael Benton, "it seems clear then that stasis is common, and that had not been predicted from modern genetic studies." A paramount example of evolutionary stasis is the fern Osmunda claytoniana.
Based on paleontological evidence it has remained unchanged, even at
the level of fossilized nuclei and chromosomes, for at least 180 million
years.
Theoretical mechanisms
Punctuational change
When Eldredge and Gould published their 1972 paper, allopatric speciation was considered the "standard" model of speciation. This model was popularized by Ernst Mayr in his 1954 paper "Change of genetic environment and evolution," and his classic volume Animal Species and Evolution (1963).
Allopatric speciation suggests that species with large central
populations are stabilized by their large volume and the process of gene flow. New and even beneficial mutations
are diluted by the population's large size and are unable to reach
fixation, due to such factors as constantly changing environments.
If this is the case, then the transformation of whole lineages should
be rare, as the fossil record indicates. Smaller populations on the
other hand, which are isolated from the parental stock, are decoupled
from the homogenizing effects of gene flow. In addition, pressure from natural selection
is especially intense, as peripheral isolated populations exist at the
outer edges of ecological tolerance. If most evolution happens in these
rare instances of allopatric speciation then evidence of gradual
evolution in the fossil record should be rare. This hypothesis was alluded to by Mayr in the closing paragraph of his 1954 paper:
Rapidly evolving peripherally isolated populations may be the place of origin of many evolutionary novelties. Their isolation and comparatively small size may explain phenomena of rapid evolution and lack of documentation in the fossil record, hitherto puzzling to the palaeontologist.
Although punctuated equilibrium generally applies to sexually reproducing organisms, some biologists have applied the model to non-sexual species like viruses,
which cannot be stabilized by conventional gene flow. As time went on
biologists like Gould moved away from wedding punctuated equilibrium to
allopatric speciation, particularly as evidence accumulated in support
of other modes of speciation. Gould, for example, was particularly attracted to Douglas Futuyma's work on the importance of reproductive isolating mechanisms.
Stasis
Many hypotheses have been proposed to explain the putative causes of stasis. Gould was initially attracted to I. Michael Lerner's theories of developmental and genetic homeostasis. However this hypothesis was rejected over time, as evidence accumulated against it. Other plausible mechanisms which have been suggested include: habitat tracking, stabilizing selection, the Stenseth-Maynard Smith stability hypothesis, constraints imposed by the nature of subdivided populations, normalizing clade selection, and koinophilia.
Evidence for the existence of stasis has also been corroborated from the genetics of sibling species,
species which are morphologically indistinguishable, but whose proteins
have diverged sufficiently to suggest they have been separated for
millions of years.
According to Gould, "stasis may emerge as the theory's most important contribution to evolutionary science." Philosopher Kim Sterelny
in clarifying the meaning of stasis adds, "In claiming that species
typically undergo no further evolutionary change once speciation is
complete, they are not claiming that there is no change at all between
one generation and the next. Lineages do change. But the change between
generations does not accumulate. Instead, over time, the species wobbles
about its phenotypic mean. Jonathan Weiner's The Beak of the Finch describes this very process."
Hierarchical evolution
Punctuated equilibrium has also been cited as contributing to the hypothesis that species are Darwinian individuals, and not just classes, thereby providing a stronger framework for a hierarchical theory of evolution.
Common misconceptions
Much
confusion has arisen over what proponents of punctuated equilibrium
actually argued, what mechanisms they advocated, how fast the
punctuations were, what taxonomic scale their theory applied to, how
revolutionary their claims were intended to be, and how punctuated
equilibrium related to other ideas like saltationism, quantum evolution, and mass extinction.
Saltationism
Alternative explanations for the punctuated pattern of evolution observed in the fossil record. Both macromutation
and relatively rapid episodes of gradual evolution could give the
appearance of instantaneous change, since 10,000 years seldom registers
in the geological record.
The punctuational nature of punctuated equilibrium has engendered
perhaps the most confusion over Eldredge and Gould's theory. Gould's
sympathetic treatment of Richard Goldschmidt, the controversial geneticist who advocated the idea of "hopeful monsters," led some biologists to conclude that Gould's punctuations were occurring in single-generation jumps. This interpretation has frequently been used by creationists to characterize the weakness of the paleontological record, and to portray contemporary evolutionary biology as advancing neo-saltationism.
In an often quoted remark, Gould stated, "Since we proposed punctuated
equilibria to explain trends, it is infuriating to be quoted again and
again by creationists—whether through design or stupidity, I do not
know—as admitting that the fossil record includes no transitional forms. Transitional forms are generally lacking at the species level, but they are abundant between larger groups."
Although there exist some debate over how long the punctuations last,
supporters of punctuated equilibrium generally place the figure between
50,000 and 100,000 years.
Quantum evolution
Quantum evolution was a controversial hypothesis advanced by Columbia University paleontologist George Gaylord Simpson, who was regarded by Gould as "the greatest and most biologically astute paleontologist of the twentieth century."
Simpson's conjecture was that according to the geological record, on
very rare occasions evolution would proceed very rapidly to form
entirely new families, orders, and classes of organisms.
This hypothesis differs from punctuated equilibrium in several
respects. First, punctuated equilibrium was more modest in scope, in
that it was addressing evolution specifically at the species level. Simpson's idea was principally concerned with evolution at higher taxonomic groups. Second, Eldredge and Gould relied upon a different mechanism. Where Simpson relied upon a synergistic interaction between genetic drift and a shift in the adaptive fitness landscape,
Eldredge and Gould relied upon ordinary speciation, particularly Ernst
Mayr's concept of allopatric speciation. Lastly, and perhaps most
significantly, quantum evolution took no position on the issue of
stasis. Although Simpson acknowledged the existence of stasis in what he
called the bradytelic mode, he considered it (along with rapid
evolution) to be unimportant in the larger scope of evolution. In his Major Features of Evolution
Simpson stated, "Evolutionary change is so nearly the universal rule
that a state of motion is, figuratively, normal in evolving populations.
The state of rest, as in bradytely, is the exception and it seems that
some restraint or force must be required to maintain it." Despite such
differences between the two models, earlier critiques—from such eminent
commentators as Sewall Wright as well as Simpson himself—have argued that punctuated equilibrium is little more than quantum evolution relabeled.
Multiple meanings of gradualism
Punctuated equilibrium is often portrayed to oppose the concept of gradualism, when it is actually a form of gradualism.
This is because even though evolutionary change appears instantaneous
between geological sedimentary layers, change is still occurring
incrementally, with no great change from one generation to the next. To
this end, Gould later commented that "Most of our paleontological colleagues missed this insight because they had not studied evolutionary theory and either did not know about allopatric speciation or had not considered its translation to geological time. Our evolutionary colleagues also failed to grasp the implication(s), primarily because they did not think at geological scales".
Richard Dawkins dedicated a chapter in The Blind Watchmaker to correcting, in his view, the wide confusion regarding rates of change. His first point is to argue that phyletic gradualism
— understood in the sense that evolution proceeds at a single uniform
rate of speed, called "constant speedism" by Dawkins — is a "caricature
of Darwinism" and "does not really exist".
His second argument, which follows from the first, is that once the
caricature of "constant speedism" is dismissed, we are left with one
logical alternative, which Dawkins terms "variable speedism". Variable
speedism may also be distinguished one of two ways: "discrete variable speedism" and "continuously variable
speedism". Eldredge and Gould, proposing that evolution jumps between
stability and relative rapidity, are described as "discrete variable
speedists", and "in this respect they are genuinely radical."
They assert that evolution generally proceeds in bursts, or not at all.
"Continuously variable speedists", on the other hand, advance that
"evolutionary rates fluctuate continuously from very fast to very slow
and stop, with all intermediates. They see no particular reason to
emphasize certain speeds more than others. In particular, stasis, to
them, is just an extreme case of ultra-slow evolution. To a
punctuationist, there is something very special about stasis." Dawkins therefore commits himself here to an empirical claim about the geological record, in contrast to his earlier claim that "The paleontological evidence can be argued about, and I am not qualified to judge it." It is this particular commitment that Eldredge and Gould have aimed to overturn.
Criticism
Richard Dawkins
regards the apparent gaps represented in the fossil record to document
migratory events rather than evolutionary events. According to Dawkins,
evolution certainly occurred but "probably gradually" elsewhere.
However, the punctuational equilibrium model may still be inferred from
both the observation of stasis and examples of rapid and episodic
speciation events documented in the fossil record.
Dawkins also emphasizes that punctuated equilibrium has been "oversold by some journalists", but partly due to Eldredge and Gould's "later writings". Dawkins contends that the hypothesis "does not deserve a particularly large measure of publicity".
It is a "minor gloss," an "interesting but minor wrinkle on the surface
of neo-Darwinian theory," and "lies firmly within the neo-Darwinian
synthesis".
In his book Darwin's Dangerous Idea, philosopher Daniel Dennett
is especially critical of Gould's presentation of punctuated
equilibrium. Dennett argues that Gould alternated between revolutionary
and conservative claims, and that each time Gould made a revolutionary
statement—or appeared to do so—he was criticized, and thus retreated to a
traditional neo-Darwinian position. Gould responded to Dennett's claims in The New York Review of Books, and in his technical volume The Structure of Evolutionary Theory.
English professor Heidi Scott argues that Gould's talent for
writing vivid prose, his use of metaphor, and his success in building a
popular audience of nonspecialist readers altered the "climate of
specialized scientific discourse" favorably in his promotion of
punctuated equilibrium.
While Gould is celebrated for the color and energy of his prose, as
well as his interdisciplinary knowledge, critics such as Scott, Richard
Dawkins, and Daniel Dennett have concerns that the theory has gained
undeserved credence among non-scientists because of Gould's rhetorical
skills.
Philosopher John Lyne and biologist Henry Howe believed punctuated
equilibrium's success has much more to do with the nature of the
geological record than the nature of Gould's rhetoric. They state, a
"re-analysis of existing fossil data has shown, to the increasing
satisfaction of the paleontological community, that Eldredge and Gould
were correct in identifying periods of evolutionary stasis which are
interrupted by much shorter periods of evolutionary change."
Some critics jokingly referred to the theory of punctuated equilibrium as "evolution by jerks",
which reportedly prompted punctuationists to describe phyletic gradualism as "evolution by creeps."
Darwin's theory
The sudden appearance of most species in the geologic record and the lack of evidence of substantial gradual change in most species—from their initial appearance until their extinction—has long been noted, including by Charles Darwin who appealed to the imperfection of the record as the favored explanation. When presenting his ideas against the prevailing influences of catastrophism and progressive creationism, which envisaged species being supernaturally created at intervals, Darwin needed to forcefully stress the gradual nature of evolution in accordance with the gradualism promoted by his friend Charles Lyell. He privately expressed concern, noting in the margin of his 1844 Essay, "Better begin with this: If species really, after catastrophes, created in showers world over, my theory false."
It is often incorrectly assumed that he insisted that the rate of
change must be constant, or nearly so, but even the first edition of On the Origin of Species
states that "Species of different genera and classes have not changed
at the same rate, or in the same degree. In the oldest tertiary beds a
few living shells may still be found in the midst of a multitude of
extinct forms... The Silurian Lingula differs but little from the living species of this genus". Lingula is among the few brachiopods surviving today but also known from fossils over 500 million years old. In the fourth edition (1866) of On the Origin of Species
Darwin wrote that "the periods during which species have undergone
modification, though long as measured in years, have probably been short
in comparison with the periods during which they retain the same form." Thus punctuationism in general is consistent with Darwin's conception of evolution.
According to early versions of punctuated equilibrium,
"peripheral isolates" are considered to be of critical importance for
speciation. However, Darwin wrote, "I can by no means agree ...
that immigration and isolation are necessary elements.... Although
isolation is of great importance in the production of new species, on
the whole I am inclined to believe that largeness of area is still more
important, especially for the production of species which shall prove
capable of enduring for a long period, and of spreading widely."
The importance of isolation in forming species had played a significant part in Darwin's early thinking, as shown in his Essay of 1844. But by the time he wrote the Origin he had downplayed its importance. He explained the reasons for his revised view as follows:
Throughout a great and open area, not only will there be a greater chance of favourable variations, arising from the large number of individuals of the same species there supported, but the conditions of life are much more complex from the large number of already existing species; and if some of these species become modified and improved, others will have to be improved in a corresponding degree, or they will be exterminated. Each new form, also, as soon as it has been improved, will be able to spread over the open and continuous area, and will thus come into competition with many other forms ... the new forms produced on large areas, which have already been victorious over many competitors, will be those that will spread most widely, and will give rise to the greatest number of new varieties and species. They will thus play a more important role in the changing history of the organic world.
Thus punctuated equilibrium is incongruous with some of Darwin's
ideas regarding the specific mechanisms of evolution, but generally
accords with Darwin's theory of evolution by natural selection.
Supplemental modes of rapid evolution
Recent work in developmental biology has identified dynamical and physical mechanisms of tissue morphogenesis
that may underlie abrupt morphological transitions during evolution.
Consequently, consideration of mechanisms of phylogenetic change that
have been found in reality to be non-gradual is increasingly common in
the field of evolutionary developmental biology,
particularly in studies of the origin of morphological novelty. A
description of such mechanisms can be found in the multi-authored volume
Origination of Organismal Form (MIT Press; 2003).
Language change
In linguistics, R. M. W. Dixon has proposed a punctuated equilibrium model for language histories, with reference particularly to the prehistory of the indigenous languages of Australia and his objections to the proposed Pama–Nyungan
language family there. Although his model has raised considerable
interest, it does not command majority support within linguistics.
Separately, recent work using computational phylogenetic methods claims to show that punctuational bursts play an important factor when languages split from one another, accounting for anywhere from 10 to 33% of the total divergence in vocabulary.
Mythology
Punctuational evolution has been argued to explain changes in folktales and mythology over time.
