It’s been said that the eyes are the windows to the soul. They allow us to communicate feelings across a room, direct the attention of others, and express emotion better than words ever could. The importance of eye contact in non-human species is well known—we’ve all heard that you shouldn’t stare a bear or angry dog in the eyes—but we don’t know a whole lot about how gaze is used between individuals of the same species. Japanese researchers took on this topic in a recent PLOS ONE article, focusing specifically on how eye contact and communication is affected by eye visibility and facial patterning around the eyes of canids.
Their research observed 25 canid species, comparing variations in facial pattern and coloring to observations about their social behavior and evolutionary history. They found that canines may use facial markers to either highlight or de-emphasize their eyes. Species with more distinguishable eyes tended to live and hunt in groups, where gaze-communication facilitates the teamwork that is necessary to bring down large prey and stay safe. Those with camouflaged eyes were more likely to live alone or in pairs, where communication with other members of their species may not be needed in the same way.
Using photos of each species, the authors analyzed the contrast between five areas of the canine face: pupil, iris, eyelid margin, coat around the eyes, and facial area including the eyes, as shown in the figure above. They measured contrast assuming red-green colorblindness of the observer (fun fact: canids cannot see the full spectrum of color). Species were then grouped according to the visibility of their eyes, described in the figure below:
- Group A contained species with easily visible pupils and eye placement
- Group B contained species with camouflaged pupils but clearly defined eye placement
- Group C contained species with fully camouflaged eyes and pupils
The authors found the strongest correlation between eye visibility and living and hunting behavior. More species in Group A, like gray wolves, live and hunt in packs, whereas more species in Groups B and C, like the fennec fox and bush dog, live and hunt alone or in pairs. Species in Group A also spend significantly more time in “gazing postures,” with their sight and body directed at another animal, an action that accentuates their focused attention to other members of the group. The genetic similarity between species was not as useful in explaining these differences, with A-type faces found in 8 of 10 wolf-like species, and in 3 of 10 red fox-like species. The authors suggest that A-type markings developed independently once these groups had evolutionarily split.
Lighter iris coloring is thought to be an adaptation to ultraviolet light in many species, similar to variations in human skin pigmentation. To determine whether this adaptation could explain the variation seen in canid iris color, the researchers compared the eye coloring of three wolf subspecies from Group A originating from arctic, temperate, and subtropical regions, to see if any differences in their lighter coloring could be attributed to geographical origin. They found that iris color did not vary significantly between the subspecies, suggesting that it may have developed to facilitate communication and not as an adaptation to specific geographical locations.
When the authors reviewed social behaviors, they found a number of social species with B- and C-type faces, the groups normally found alone or in pairs. These species are known to use acoustic or other visual signals, like a howl or the flash of a white tail, to communicate with their comrades. This allows them to skirt one possible disadvantage of gaze-communication: when prey can also identify and follow a gaze, and realize they’ve been targeted.
Gaze communication may be an important tool for other canids, including our own companions, domestic dogs. Previous studies have shown that domestic dogs are more likely to make direct eye contact with humans than wolves raised in the same setting. This could mean that after thousands of years of cohabitation, dogs see us in socially useful ways that wolves never will. Luckily for us, that means we get to see this.
Citation: Ueda S, Kumagai G, Otaki Y, Yamaguchi S, Kohshima S (2014) A Comparison of Facial Color Pattern and Gazing Behavior in Canid Species Suggests Gaze Communication in Gray Wolves (Canis lupus). PLoS ONE 9(6): e98217. doi:10.1371/journal.pone.0098217
Images 2 and 3: Figures 1 and 2 from the article