Preliminary evidence that eye appearance in parrots (Psittaciformes) co-varies with latitude and altitude.

External eye appearance in avian taxa has been proposed to be driven by social and ecological functions. Recent research in primates suggests, instead, that, photoprotective functions are important drivers of external eye appearance. Using similar methods, we examined the variation in external eye appearance of 132 parrot species (Psittaciformes) in relation to their ecology and sociality. Breeding systems, flock size and sexual dimorphism, as well as species' latitude and maximum living altitude, and estimated UV-B incidence in species' ranges were used to explore the contribution of social and ecological factors in driving external eye appearance. We measured the hue and brightness of visible parts of the eye and the difference in measurements of brightness between adjacent parts of the eye. We found no link between social variables and our measurements. We did, however, find a negative association between the brightness of the inner part of the iris and latitude and altitude. Darker inner irises were more prevalent farther away from the equator and for those species living at higher altitudes. We found no link between UV-B and brightness measurements of the iris, or tissue surrounding the eye. We speculate that these results are consistent with an adaptation for visual functions. While preliminary, these results suggest that external eye appearance in parrots is influenced by ecological, but not social factors.

No immediate attentional bias towards or choice bias for male secondary sexual characteristics in Bornean orang-utans (Pongo pygmaeus).

Primate faces provide information about a range of variant and invariant traits, including some that are relevant for mate choice. For example, faces of males may convey information about their health or genetic quality through symmetry or facial masculinity. Because perceiving and processing such information may have bearing on the reproductive success of an individual, cognitive systems are expected to be sensitive to facial cues of mate quality. However, few studies have investigated this topic in non-human primate species. Orang-utans are an interesting species to test mate-relevant cognitive biases, because they are characterised by male bimaturism: some adult males are fully developed and bear conspicuous flanges on the side of their face, while other males look relatively similar to females. Here, we describe two non-invasive computerised experiments with Bornean orang-utans (Pongo pygmaeus), testing (i) immediate attention towards large flanges and symmetrical faces using a dot-probe task (N = 3 individuals; 2F) and (ii) choice bias for pictures of flanged males over unflanged males using a preference test (N = 6 individuals; 4F). In contrast with our expectations, we found no immediate attentional bias towards either large flanges or symmetrical faces. In addition, individuals did not show a choice bias for stimuli of flanged males. We did find exploratory evidence for a colour bias and energy efficiency trade-offs in the preference task. We discuss our null results and exploratory results in the context of the evolutionary history of Bornean orang-utans, and provide suggestions for a more biocentric approach to the study of orang-utan cognition.

Is there no beauty in sexually dimorphic eyes? Facial attractiveness and White Europeans ocular morphology-Brief communication.

The link between human ocular morphology and attractiveness, especially in the context of its potential adaptive function, is an underexplored area of research. In our study, we examined the association between facial attractiveness and three sexually dimorphic measures of ocular morphology in White Europeans: the sclera size index, width-to-height ratio, and relative iris luminance. Sixty participants (30 women) assessed the attractiveness of the opposite-sex photographs of 50 men and 50 women. Our results show that in both men and women, none of the three measures was linked to the opposite sex ratings of facial attractiveness. We conclude that those ocular morphology measures may play a limited role in human mate preferences.

Individual attractiveness preferences differentially modulate immediate and voluntary attention.

Physical attractiveness plays a crucial role in mate choice for both men and women. This is reflected in visual attention: people immediately attend towards and look longer at attractive faces, especially when they are motivated to find a partner. However, previous studies did not incorporate real-life dating decisions. Here, we aimed to combine attentional tasks with individual attractiveness ratings and a real-life mate choice context, namely a speed-dating paradigm. We investigated whether heterosexual non-committed young adults showed biases in immediate and voluntary attention towards attractive faces and preferred dating partners. In line with previous research, we found considerable individual differences in individual attractiveness preferences. Furthermore, our results showed that men had a bias towards attractive faces and preferred dating partners in the immediate attention task, while results for women were mixed. In the voluntary attention task, however, both men and women had an attentional bias towards attractive faces and preferred dating partners. Our results suggest that individual attractiveness preferences are good predictors of especially voluntary attention. We discuss these findings from an evolutionary perspective and suggest directions for future research.

The adaptive significance of human scleral brightness: an experimental study.

Homogeneously depigmented sclerae have long been proposed to be uniquely human-an adaptation to enable cooperative behaviour by facilitating interpersonal coordination through gaze following. However, recent evidence has shown that deeply pigmented sclerae also afford gaze following if surrounding a bright iris. Furthermore, while current scleral depigmentation is clearly adaptive in modern humans, it is less clear how the evolutionarily intermediate stages of scleral pigmentation may have been adaptive. In sum, it is unclear why scleral depigmentation became the norm in humans, while not so in sister species like chimpanzees, or why some extant species display intermediate degrees of pigmentation (as our ancestors presumably did at some point). We created realistic facial images of 20 individually distinct hominins with diverse facial morphologies, each face in the (i) humanlike bright sclera and (ii) generalised apelike dark sclera version. Participants in two online studies rated the bright-sclera hominins as younger, healthier, more attractive and trustworthy, but less aggressive than the dark-sclera hominins. Our results support the idea that the appearance of more depigmented sclerae promoted perceived traits that fostered trust, increasing fitness for those individuals and resulting in depigmentation as a fixed trait in extant humans.

Ecological factors are likely drivers of eye shape and colour pattern variations across anthropoid primates.

External eye appearance across primate species is diverse in shape and colouration, yet we still lack an explanation for the drivers of such diversity. Here we quantify substantial interspecific variation in eye shape and colouration across 77 primate species representing all extant genera of anthropoid primates. We reassess a series of hypotheses aiming to explain ocular variation in horizontal elongation and in colouration across species. Heavier body weight and terrestrial locomotion are associated with elongated eye outlines. Species living closer to the equator present more pigmented conjunctivae, suggesting photoprotective functions. Irises become bluer in species living further away from the equator, adding to existing literature supporting a circadian clock function for bluer irises. These results shift the current focus from communicative, to ecological factors in driving variation in external eye appearance in anthropoid primates. They also highlight the possibility that similar ecological factors contributed to selection for blue eyes in ancestral human populations living in northern latitudes.

Scleral pigmentation leads to conspicuous, not cryptic, eye morphology in chimpanzees.

Gaze following has been argued to be uniquely human, facilitated by our depigmented, white sclera [M. Tomasello, B. Hare, H. Lehmann, J. Call, J. Hum. Evol. 52, 314-320 (2007)]-the pale area around the colored iris-and to underpin human-specific behaviors such as language. Today, we know that great apes show diverse patterns of scleral coloration [J. A. Mayhew, J. C. Gómez, Am. J. Primatol. 77, 869-877 (2015); J. O. Perea García, T. Grenzner, G. Hesková, P. Mitkidis, Commun. Integr. Biol. 10, e1264545 (2016)]. We compare scleral coloration and its relative contrast with the iris in bonobos, chimpanzees, and humans. Like humans, bonobos' sclerae are lighter relative to the color of their irises; chimpanzee sclerae are darker than their irises. The relative contrast between the sclera and iris in all 3 species is comparable, suggesting a perceptual mechanism to explain recent evidence that nonhuman great apes also rely on gaze as a social cue.

Humans do not perceive conspecifics with a greater exposed sclera as more trustworthy: a preliminary cross-ethnic study of the function of the overexposed human sclera.

Understanding the adaptive function of the unique morphology of the human eye, in particular its overexposed white sclera, may have profound implications for the fields of evolutionary behavioural science, and specifically the areas of human interaction and social cognition. Existing hypotheses, such as the cooperative eye hypothesis, have attracted a lot of attention but remain untested. Here, we: (i) analysed variation in the visible sclera size in humans from different ethnic backgrounds and (ii) examined whether intraspecific variation of exposed sclera size is related to trust. We used 596 facial photographs of men and women, assessed for perceived trustworthiness, from four different self-declared racial backgrounds. The size of the exposed sclera was measured as the ratio between the width of the exposed eyeball and the diameter of the iris (sclera size index, SSI). The SSI did not differ in the four examined races and was sexually monomorphic except for Whites, where males had a larger SSI than females. In general, the association between the SSI and trustworthiness was statistically insignificant. An inverted U-shaped link was found only in White women, yet the strength of the effect of interaction between sex and race was very small. Our results did not provide evidence for the link between exposed sclera size and trustworthiness. We conclude that further investigation is necessary in order to properly assess the hypotheses relating to the socially relevant functions of overexposed sclera.

Not everything is blue or brown: Quantification of ocular coloration in psychological research beyond dichotomous categorizations.

The notion that phenomenologically observable differences in the human eye are correlated with behavioral tendencies (other than gaze-following) has been addressed poorly in the psychological literature. Most notably, the proposed correlations are based on an arbitrary categorization in discrete categories of the continuous variability across various traits that could be contributing to individual eye morphologies. We review the relevant literature and assume a view of human eyes as sign stimuli, identifying the relative contrast between the iridal and scleral areas as the main contributor to the strength of the signal. Based on this view, we present a new method for the precise quantification of the relative luminosity of the iris (RLI) and briefly discuss its potential applications in psychological research.