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# The spandrels of San Marco and the Panglossian paradigm: a critique of the adaptationist programme

## Details

**Authors** Gould, Stephen Jay and Lewontin, Richard C.

**Year** 1979

**Publisher** Proceedings of the Royal Society B: Biological Sciences, 205(1161), 581–598

**Kind of work** article

**Discipline** Evolutionary Biology

**Secondary disciplines** Philosophy

**Applied** false

[Read it at the publisher](https://doi.org/10.1098/rspb.1979.0086) 
10.1098/rspb.1979.0086

## In authors' words

### What they set out to do (purpose)

To criticize the adaptationist programme in evolutionary biology.

### How they did it (methods)

Theoretical critique using the architectural spandrel as an analogy.

### What they found (results)

The authors argue that the adaptationist programme divides organisms into separate traits and supplies an adaptive story for each, confuses current utility with origin, fails to test alternatives, and overlooks nonadaptive explanations such as random fixation, developmental correlation and byproduct structures. They call for analyzing organisms as integrated wholes shaped by inherited body plans and developmental constraints, and support a pluralistic approach to the agents of change.

## Commentary

### In short

A critique of explaining every trait of an organism as a separate adaptation, arguing that organisms are integrated wholes whose structure and development limit and shape what selection can do.

### Response

The argument is built on the part-whole relation: atomizing an organism into traits misrepresents a whole whose parts constrain one another. It also relies on distinctions, notably between the origin of a feature and its current use and between adaptations and byproducts, and on relationships such as developmental correlation. It is a conceptual critique, not a formal or empirical test. Findings are drawn from the abstract only.

**Patterns it shows** D, S, R

**Added** 2026-10-26

**How to cite this** Gould, Stephen Jay and Lewontin, Richard C. (1979). The spandrels of San Marco and the Panglossian paradigm: a critique of the adaptationist programme. Proceedings of the Royal Society B: Biological Sciences, 205(1161), 581–598.