Punctuated Equilibrium

Punctuated Equilibrium Definition

Punctuated equilibrium is a theory that states that evolution occurs primarily through short bursts of intense speciation, followed by lengthy periods of stasis or equilibrium. It postulates that nearly 99% of a species’ existence on earth is spent in stasis. So, if a species appears in fossil records for about 10 million years, it is likely that speciation occurred over the span of less than 100,000 years. Once complete however, there is little, if any, morphological change.

The theory also provides an explanation for the absence of intermediate forms in fossil records, where new species seem to appear from ancestral forms abruptly and ultimately disappear without experiencing any apparent morphological change during their existence.

While this was a shift from the idea that all new species arose due to continuous, gradual and incremental changes, the founders of this theory have also conceded that other modes of evolution could co-exist.

Features of Punctuated Equilibrium

One of the cornerstones of this hypothesis is that reproductive isolation is necessary for the formation of new species. This implies that the fossil record at any one place is unlikely to record the process of speciation because new species can evolve only from small, isolated populations. Therefore, variations will be seen only in fossils of the same age arising from different geographical locations.

Punctuated equilibrium postulates that genetic and morphological changes that bestow a survival advantage will be amplified quickly in small populations. The rapid pace of evolution in these isolated groups is also stated as the reason why there is no fossil record of evolution, and new species seem to appear abruptly.

It also predicts that while intermediates will be rare in the evolution of single species, they will be seen among larger groups. For example, while, Australopithecus afarensis, is the precursor of modern humans, there are no fossils showing a gradual change in the cranial capacity or body size of the Australopithecus. However, there are other species such as Homo habilis and Homo erectus that show the transition from Australopithecus towards modern man in terms of cranial capacity and body size.

Another important feature of this hypothesis is the explanation given for extended periods of stasis. It implies that the average morphology of a species is under a homogenizing influence. Interbreeding populations are said to appear static because, in the absence of active selection pressure, any changes are diluted among the large number of individuals. A number of explanations have been given for this phenomenon observed in fossil record. These include the effect of gene flow, assertions that the morphology of a species is under ‘homeostatic’ pressure, and koinophilia or the rejection of mates with unusual attributes.

Examples of Punctuated Equilibria

Support for punctuated equilibrium is seen in fossil records, and the impact of reproductive isolation has been observed by biologists, systematists and taxonomists across the world. Given the fact that this is a theory of evolution, its predictions cannot be directly tested. While fossil record can provide support for the theory, some indications need to arise from the living world. For example, animals living in similar environments but experiencing reproductive isolation must become incapable of interbreeding, indicating the emergence of a new species.

Reproductive Isolation among Kingfishers

The study of kingfishers in Papua New Guinea showed the deep impact of reproductive isolation on speciation. There are three subspecies that reside on the mainland, where the environment can vary wildly from humid, dense rain forest, to monsoon forests with extended dry seasons. These subspecies can not only interbreed, but are nearly indistinguishable from each other. However, on islands a few hundred kilometers away, even when the environment is similar to the nearest part of the mainland, the kingfishers are markedly different. More species have been found on these smaller islands than in mainland – a landmass spanning nearly 300,000 square miles. Similar observations have been made for birds and reptiles and invertebrates across the world, where geographical separation has led to the emergence of new species, while large continuous tracts with varying conditions maintain homogeneous populations.

Land Snails of Bermuda

About 300,000 years ago, Poecilozonites bermudensis, an air-breathing land snail, colonized the island of Bermuda, possibly carried on driftwood from North America. The fossils of these snails constitute the large majority of Bermuda’s land fossils and, until recently, one species continued to survive on the island. The earliest populations of this snail had two stocks, with distinct color banding patterns. When these became extinct, a derivative from a peripheral population that was evolving on a separate island became dominant. Fossil samples taken from six different geological times and from various geographical locations points to the repeated evolution of species from peripherally isolated populations that ultimately led to the formation of the land snail that remained morphologically static till it was observed in the 1950s.

Gradualism vs Punctuated Equilibrium

Punctuated equilibrium is often pitted against phyletic gradualism as competing theories of evolution. Both of these hypothesize about the rate of emergence of species. Gradualism places importance on the slow appearance of new characters in interbreeding subspecies that, over time, lead to the evolution of a new species from ancestral forms.

However, this is not supported by fossil data, where new species seem to appear suddenly. Punctuated equilibrium tries to explain these fossil ‘gaps’ or the absence of intermediate forms, but stating that they exist for very short periods of time, when speciation occurs intensely in an isolated population.

The criticism of punctuated equilibrium focuses on the possibility that fossil records may simply be incomplete and intermediate forms may still be found in regions where fossils are abundant and well-preserved. In addition, critics point to the fact that there is no evidence that an external homogenizing influence keeps interbreeding populations in stasis.

Related Biology Terms

  • Allopatric speciation – Speciation that occurs after a population splits into two groups that are reproductively isolated from each other.
  • Koinophilia – Phenomenon where individuals with unusual features are not preferred for sexual reproduction.
  • Peripatric speciation – Speciation in a small, isolated, peripheral population.
  • Phyletic Gradualism – A model that theorizes that speciation is gradual, incremental and slow.
  • Saltation – Sudden change that occurs over the span of a single generation.


1. Which of these is a major feature of punctuated equilibrium?
A. Rapid increase in reproductive capacity
B. Detailed fossil record of intermediates
C. Long periods of stasis with no morphological changes
D. Mutation

Answer to Question #1
C is correct. Punctuated equilibrium states that short bursts of intense speciation are interspersed with long periods of equilibrium or stasis.

2. How does punctuated equilibrium explain the lack of intermediates in the fossil record?
A. Fossil record is incomplete
B. Environmental change destroys fossils
C. Speciation is too rapid to leave behind a fossil record
D. All of the above

Answer to Question #2
C is correct. Speciation is too rapid to leave behind a fossil record. Punctuated equilibrium specifically argues against the idea of the fossil record being incomplete or destroyed.

3. Gradualism and punctuated equilibrium are often considered as opposing theories of evolution.
A. True
B. False

Answer to Question #3
True. Though the founders of punctuated equilibrium concede that the two could co-exist as evolutionary processes.