What Are Spandrels?

Spandrels are byproducts of evolution. They arise due to the evolution of one trait, but this unintended trait may not have any function.

Take a look at this picture:

A dusk view of south side of Constantine's Arch, standing at between the Colosseum, right, and the Roman Forum, left( Sean Xu)S

The arch of Constantine, Rome (Photo Credit: Sean Xu/ Shutterstock)

That is the Arch of Constantine in Rome, a type of triumphal arch that is situated between the Colosseum and Palatine Hill. A triumphal arch served as a gateway for the victorious Roman army when it came back home after winning a war.

This particular one was erected to celebrate the victory of Constantine I’s victory over the tyrant ruler Maxentius in the Battle of Milvian Bridge in 312 AD. On it are beautiful collages of the past Roman emperors who had ruled before Constantine. It is easy to imagine Constantine walking with his chest filled with pride as he walked through the arch, the emperors of yore congratulating him on his supreme victory. 

Arch of Constantine, a massive triumphal arch built to celebrate Emperor Constantine located front the Coliseum in Rome( Jose HERNANDEZ Camera 51)S

A close-up of the arch relief on the south face of the arch of Constantine, Rome. these decorative parts of the arch are byproducts of making the arch shape. (Photo Credit: Jose HERNANDEZ Camera 51/ Shutterstock)

If we removed the decorative elements on the corners of the arch, the arch is still structurally fine (although now we probably have an angry emperor on our hands). The triangular portion just above the arch could be called a byproduct.

These parts of a building (most commonly seen in the construction of arches or domes) have a name—spandrels. As formally defined by the Oxford Dictionary, a spandrel is “the almost triangular space between one side of the outer curve of an arch, a wall, and the ceiling or framework.”

The makers of the arch probably didn’t design the empty triangles on the corners specifically for Constantine’s happiness. How could you allow Constantine to enter through a plain-faced arch? How embarrassing! 

Spandrels are seen everywhere in architecture, but spandrels don’t only exist in man-made structures; they are found in nature as well!

In 1979, two evolutionary biologists (among various other titles), Stephen Jay Gould and Richard Lewontin, published a paper titled “The Spandrels of San Marco and the Panglossian Paradigm: A Critique of the Adaptationist Programme”. 

What are spandrels in biology?

Gould and Lewontin defined a biological spandrel as a byproduct of evolutionary adaptation. They are necessary ‘leftovers’ created because some other trait had evolved. This means that the spandrel isn’t an adaptation to anything in the environment. Instead, it is a secondary trait that arose from the development of another primary trait.

The easiest spandrel to visualize is the human chin. On hypothesis about why humans are the only animals that have a chin is that it is merely a byproduct of the growth of different parts of the jaw.

Side view of the teeth and jaws.

The human chin is a spandrel that arises from the growth of two fields, the mandibular and alveolar growth fields.  (Photo Credit: H. V. Carter/Wikimedia Commons)

The development of the jaw was an adaptation to the kind of food humans used to consume during our good old prehistoric days. The chin is a secondary trait that developed because of the first jaw adaptation. This secondary trait isn’t an adaptation to any specific environmental conditions. The evolution of the chin, in isolation from the jaw’s evolution, would mean very little to the animal’s survival.   

This proposal of a trait that served no adaptive purpose was a critique of the thought that every trait is an adaptation and has been selected for through natural selection.

The Panglossian Adaptationist

Dr Pangloss, in Voltaire’s novel, Candide, is one of the most optimistic fictional characters ever created. His ‘all for the best attitude’ is on clear display in quotes like, “Things cannot be other than they are… Everything is made for the best purpose. Our noses were made to carry spectacles, so we have spectacles. Legs were clearly intended for breeches, and we wear them.” Gould and Lewontin’s ‘spandrels’ paper critiques this Panglossian optimistic adaptationist view of evolution. 

Adaptationists argue that every trait must have some evolutionary function or else natural selection wouldn’t have selected for the trait to persist. They argue that the environment weeding out the less fit individuals is the biggest pressure and driving force behind evolution, keeping only the functionally important traits around. 

On the flip side, Gould and Lewontin argue for a more pluralist approach to evolution.

Pluralism in evolution refers to considering multiple factors that may have affected a trait. In popular science, natural selection is synonymous with evolution, but that isn’t the whole story. There are three other major mechanisms of (micro)evolution—genetic drift, mutations and gene flow. Pluralists argue that adaptationism often attributes a purpose to a trait because it must, not because the evidence leads to it. 

Gould and Lewontin weren’t the first to start the debate between pluralism and adaptationism. The first rift began when the theory of evolution itself was being born, between Charles Darwin, and Alfred Russel Wallace and August Weissmann.

Darwin held a pluralistic view of evolution advocating other mechanisms of evolution also be considered along with natural selection. Wallace and Weissmann, on the other hand,  took up the adaptationist mantle. Over the years, any other prominent evolutionists have helped to further shape and influence adaptationist and pluralist views. The debate between the two has reached a stalemate, with many having accepted the spandrels argument. 

Implications of spandrels

Adaptationism, according to Gould and Lewontin, not only showed an incomplete history of certain traits but could also lead to misleading and counterproductive results. If one looks at the human chin like an adaptationist, one would conclude that the chin evolved because humans found George Clooney-esque chins sexy.

Thus, people with chins found mates more often than those who didn’t. Subsequently, those humans who had sexier chins were more successful and therefore thrived. While chins can be a metric for attractiveness, this purpose of the chin came after its appearance in humans, argued Gould. Gould referred to the something useless attaining a purpose as ‘exaptation’ of function.

Spandrels have a wonderful ability to co-opt certain traits that appear to modern scientists as a primary trait. Consider architectural spandrels. They are used to display complex imagery and art that serves to elevate the aesthetic value of the building. Does that mean the architects created the spandrel solely to decorate? Obviously not. That sounds illogical. Gould argues a similar case for biological spandrels. Think of evolution as an architect putting a non-adaptive trait to use. 

There are many other examples. The hollow space in the shells of gastropods (snails) that arises as a consequence of the shell spiraling inwards. This space is used by some gastropods to store their eggs (a co-opted trait) but is left hollow in others. 

great Irish deer

The extinct great Irish deer has a shoulder hump caused by the elongation of the neural spines in the vertebra, in order to better manage the weight of the head with its giant horns. The hump is a secondary, non-adaptive trait—a spandrel. (Photo Credit: Hutchinson/Wikimedia Commons)

The hump of the great Irish deer is another spandrel. It arose due to the elongation of the vertebrate so that the horned head of the deer could be supported. The hump, through evolutionary time, became patterned and co-opted for sexual selection. 

If one assumes that the hump of the deer was an adaptation, it would mean messing up the historical origin of the trait. Doing so might lead to errors in predicting when the trait appeared, or lead to ignoring other important interactions (environment, viral, other animals, etc.). 

The examples we have considered thus far have touched upon physical traits, but what about behavioral traits? This is where the debate gets murky. Phenomena like war, language and art were considered by Gould as spandrels of a large human brain. Sure, they help humans get along and become the dominant species on the planet, but they might not have started out with that function as its inherent purpose.


  1. University Of Washington
  2. PNAS
  3. Stanford University
  4. UCLA
The short URL of the present article is: http://sciabc.us/M9Rw5
Help us make this article better

Salama has a degree in Life Science and Biochemistry from St. Xavier’s College, Mumbai. She enjoys being in the water much more than being on land. She is passionate about science and wants to declutter science from its jargon to make people understand its beauty.

Science ABC YouTube Videos

  1. Why Do We Dance To Music?Why Do We Dance To Music?
  2. What is Quantum Entanglement: Explained in Simple WordsWhat is Quantum Entanglement: Explained in Simple Words
  3. Can We Harness Electricity From Lightning?Can We Harness Electricity From Lightning?
  4. Are Giant Insects Larger Than Humans Possible?Are Giant Insects Larger Than Humans Possible?
  5. What are Glial Cells: Definition, Types, Functions of Glial Cells | Role in PsychologyWhat are Glial Cells: Definition, Types, Functions of Glial Cells | Role in Psychology
  6. Why Don't They Have Parachutes For Passengers In Commercial Planes?Why Don't They Have Parachutes For Passengers In Commercial Planes?
  7. Methusaleh: The oldest tree in the world | What's the mystery of trees' immortality?Methusaleh: The oldest tree in the world | What's the mystery of trees' immortality?
  8. 7 Scientifically Inaccurate Things They Show in Movies: Most Common Movie Mistakes and Myths7 Scientifically Inaccurate Things They Show in Movies: Most Common Movie Mistakes and Myths