small and large species may be present in canines. The small red fox (Vulpes vulpes) has vertical slit pupils whereas their large relatives, the gray wolf (Canis lupus lupus) and domestic dogs (Canis lupus familiaris) have round pupils. Evolution and adaptation One explanation for the evolution of slit pupils is that they can exclude light more effectively than a circular pupil. This would explain why slit pupils tend to be found in the eyes of animals with a crepuscular or nocturnal lifestyle that need to protect their eyes during daylight. Constriction of a circular pupil (by a ring-shaped muscle) is less complete than closure of a slit pupil, which uses two additional muscles that laterally compress the pupil. For example, the cat's slit pupil can change the light intensity on the retina 135-fold compared to 10-fold in humans. However, this explanation does not account for circular pupils that can be closed to a very small size (e.g., 0.5 mm in the tarsier) and the rectangular pupils of many ungulates which do not close to a narrow slit in bright light. An alternative explanation is that a partially constricted circular pupil shades the peripheral zones of the lens which would lead to poorly focused images at relevant wavelengths. The vertical slit pupil allows for use of all wavelengths across the full diameter of the lens, even in bright light. It has also been suggested that in ambush predators such as some snakes, vertical slit pupils may aid in camouflage, breaking up the circular outline of the eye. Activity pattern and behavior In a study of Australian snakes, pupil shapes correlated both with diel activity times and with foraging behavior. Most snake species with vertical pupils were nocturnal and also ambush foragers, and most snakes with circular pupils were diurnal and active foragers. Overall, foraging behaviour predicted pupil shape accurately in more cases than did diel time of activity, because many active-foraging snakes with circular pupils were not diurnal. It has been suggested that there may be a similar link between foraging behaviour and pupil shape amongst the felidae and canidae discussed above. A 2015 study confirmed the hypothesis that elongated pupils have increased dynamic range, and furthered the correlations with diel activity. However it noted that other hypotheses could not explain the orientation of the pupils. They showed that vertical pupils enable ambush predators to optimise their depth perception, and horizontal