A quantitative test of the predicted relationship between countershading and lighting environment

Am Nat. 2012 Dec;180(6):762-76. doi: 10.1086/668011. Epub 2012 Oct 25.


Countershading, a vertical luminance gradient from a dark back to a light belly, is perhaps the most common coloration phenotype in the animal kingdom. Why? We investigated whether countershading functions as self-shadow concealment (SSC) in ruminants. We calculated "optimal" countershading for SSC by measuring illumination falling onto a model ruminant as a function of time of day and lighting environment. Calibrated images of 114 species of ruminant were compared to the countershading model, and phylogenetic analyses were used to find the best predictors of coats' countershading characteristics. In many species, countershading was close to the model's prediction of "optimal" countershading for SSC. Stronger countershading was associated with increased use of open lighting environments, living closer to the equator, and small body size. Abrupt transitions from dark to light tones were more common in open lighting environments but unassociated with group size or antipredator behavior. Though the SSC hypothesis prediction for stronger countershading in diurnal species was not supported and noncountershaded or reverse-countershaded species were unexpectedly common, this basic pattern of associations is explained only by the SSC hypothesis. Despite extreme variation in lighting conditions, many terrestrial animals still find protection from predation by compensating for their own shadows.

Publication types

  • Research Support, Non-U.S. Gov't

MeSH terms

  • Animals
  • Biological Evolution
  • Body Size
  • Cytochromes b / genetics
  • Environment*
  • Food Chain
  • Light
  • Motor Activity
  • Phenotype
  • Photography
  • Pigmentation*
  • Ruminants / classification
  • Ruminants / genetics
  • Ruminants / physiology*


  • Cytochromes b