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Self-Domestication and the Emergence of Behavioral Modernity in Homo sapiens

Claire Brandes

Department of Anthropology, University of Georgia

ANTH 4100: Evolution and Human Behavior

Dr. Bram Tucker

April 29, 2021

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Self-Domestication and the Emergence of Behavioral Modernity in Homo sapiens

Understanding the origins of widespread behavioral modernity, the category of behaviors

characteristic of modern humans, remains a challenge within the field of evolutionary

anthropology. Cieri et al. define behavioral modernity as having the capacity to engage in

symbolic and abstract thought. Archaeologists attribute these behaviors to a population through

the discovery of prehistoric artifacts like clothing and jewelry, art, and musical instruments.

While there is some evidence for the presence of behavioral modernity in the Middle Stone Age,

these behaviors remained inconsistent until the Late Stone Age, around 50,000 years ago. Within

this proposed timeline, there remains a near 150,000 year gap between the development of the

anatomical structure and the behavior of modern humans (Cieri et al., 2014).

Two broad theories aim to explain the mechanisms which triggered the emergence of

behavioral modernity within this time period. Some scholars posit that changes in cognitive

abilities generated the capacity for behavioral modernity. Archaeological evidence from

Neanderthal sites provide evidence that these hominids also had the capacity to engage in

symbolic behaviors. Neanderthals presumably went extinct around 50,000 years ago, suggesting

that cognitive advancements cannot account for the prevalence of behavioral modernity after this

point in time. Another theory attributes cultural, ecological, and demographic factors to the rise

of behavioral modernity (Cieri et al., 2014).

One cultural factor hypothesized to have given rise to behavioral modernity is the

phenomenon of self-domestication. Self-domestication occurs through the course of selection

against aggression and for prosociality. A group of traits associated with domesticates called

domestication syndrome, can be used to diagnose self-domestication in a species (Cieri et al.,

2014). A non-exhaustive list of these traits includes reduced aggression, lower stress response,
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reduced cranial capacity, feminized (i.e., short and wide) faces, and depigmentation (Hare,

2017). The self-domestication theory holds that through increases in social tolerance, humans

were able to utilize preexisting cognitive abilities in a new way, ultimately contributing to the

spread of behavioral modernity (Cieri et al., 2014).

This essay is organized as follows: first, I will explore domestication in nonhuman

animals; second, I consider evidence for the self-domestication hypothesis in humans, consisting

of morphological, physiological, and genetic data; finally, I discuss the implications of the

self-domestication process as it relates to the emergence of modern human behavior. I conclude

by summarizing my thesis that the self-domestication phenomenon is both applicable to humans

and a plausible explanation for the origins of behavioral modernity in our species.

Comparisons of various domesticates and their wild counterparts serve to create a

foundation upon which I will later discuss the applicability of self-domestication to humans. By

comparing domesticated to analogous wild species (e.g., dogs to wolves) we can begin to

determine which characteristics may be a product of the domestication process. Besides

behavior, the most easily observable traits that indicate a species has been domesticated are their

distinct morphological traits. An experimental group of silver-foxes selected for non-aggression

toward humans over forty-five generations displayed high prosociality as well as domestication

syndrome (Hare, 2017). While these foxes were only bred for tameness, they developed many

characteristics as a side-effect of this process, apparent when compared to the control population.

A majority of foxes exhibited piebald coats, reduced snouts, feminized faces and floppy ears, all

established effects of domestication syndrome. Domesticated dogs also exhibit many

morphological differences in comparison to wolves (Cieri et al., 2014). Relative to wolves, dogs

have a reduced cranial capacity, feminized faces, and decreased sexual dimorphism in canine
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size. Guinea pigs, in comparison to wild cavies, also display reduced cranial capacity (Cieri et

al., 2014). To summarize, the most pronounced and common morphological characteristics

amongst domesticates include feminization of the face, reduced cranial capacity, and reduced

sexual dimorphism. Later I will explore physiological and genomic indicators of domestication

syndrome.

Distinct differences between chimpanzees and bonobos provide evidence of

self-domestication in a primate closely related to humans. Compared to chimpanzees, the only

other species within the genus Pan, bonobos are considerably more peaceful and prosocial. Male

chimps frequently engage in aggressive behaviors directed toward male rivals and females, while

male bonobos rarely display aggressive physical contact and are known to form strong alliances

with their mothers and other females. Chimpanzees are extremely territorial and inter-group

conflicts regularly occur, oftentimes resulting in fatalities. While bonobos have also been

observed to be territorial, inter-group hostility is severely muted compared to chimps, and these

interactions have never been fatal. Bonobos sometimes even engage in positive social behaviors

with members of other groups. Bonobos frequently participate in play, as well as non-conceptive

sexual behavior as a way to relieve social tension (Hare et al., 2012). Hare et al. posit that

ancestral stable ecological conditions allowed female bonobos to form coalitions with the ability

to selectively breed with non-aggressive males, leading to the evolution of social tolerance in this

species (Hare et al., 2012).

Although generally very similar in appearance, bonobos exhibit several parallel

morphologies to domesticates that chimpanzees do not. As opposed to chimpanzees, bonobos

have a reduced cranial capacity, feminized faces, loss of pigmentation in their tail-tuft and lips,

and low sexual dimorphism in canine size. A hallmark of domesticate physiology is a reduction
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in stress response, which bonobos, but not chimps, share. Chimpanzees are hyper-androgenized

in utero, meaning they are exposed to higher levels of male reproductive hormones (e.g.,

testosterone) than bonobos during gestation (Hare et al., 2012). The ratio of the lengths of the

index and ring fingers, which I refer to as the 2D4D ratio, is used as an indicator of fetal

exposure to testosterone; lower ratios correspond to lower levels of exposure (Cieri et al., 2014).

Analysis of bonobos and chimpanzee cortisol levels, a stress hormone, reveal that bonobos

experience stress when they anticipate social conflict, while chimps experience a surge in

testosterone, preparing them to act aggressively (Hare et al., 2012). I will use our closest living

relatives, the bonobos, as a comparative model throughout this paper.

Morphological data derived from the hominin fossil record provides evidence for

domestication syndrome in humans. Reduced aggression and lower reactivity to androgens is

evident in feminization of the craniofacial region. The self-domestication hypothesis holds that

we should observe increases in craniofacial feminization in humans over time, characterized by

reduced brow ridges and shorter faces (Cieri et al., 2014). In an analysis of over 1,000 human

skulls from three temporal categories (Middle Stone Age, Late Stone Age, and Holocene), Cieri

et al. found a statistically significant reduction in mean brow ridge projection across the sample

groups. Facial length, measured by the length from nasion to prosthion, shortened considerably

over time (Cieri et al., 2014). Homo neanderthalensis, a now extinct, but close relative of Homo

sapiens, had faces significantly masculinized compared to the oldest modern humans, as

observed through the fossil record (Hare, 2017).

Analysis of human physiology serves as further evidence of domestication syndrome in

humans. As mentioned previously, the craniofacial feminization of human skulls can be used as a

proxy measure of reduced reactivity to androgens over time. The 2D4D ratio of humans
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compared to Neanderthals and other species of the genus Homo reveals low levels of androgen

exposure throughout gestation, resulting in lower aggression. Reduction in brain size observed

during Holocene late human evolution suggests an increase in serotonin levels, which has been

observed in several animal domesticates (Hare, 2017).

Data collected from a genomic analysis of animal domesticates and humans shows an

overlap in alleles that supports self-domestication in humans. Theofanopoulou et al. isolated

genes believed to be correlated with self-domestication in humans by identifying a list of genes

positively selected for in Homo sapiens compared to Neanderthals and Denosovans, resulting in

742 genes. These 742 genes were compared to 691 genes associated with positive selection in

animal domesticates compared to their wild counterparts, yielding an overlap of 41 genes.

Theofanopoulou et al. ran statistical tests that determined the likelihood of observing this number

of genes in common between the two groups was small enough to be considered highly

significant. Five genes were found to be under selection in modern humans as well as multiple

domesticated species, two of which the researchers deemed especially significant to theory of

self-domestication. BRAF was found present in cats, horses, and humans; this gene is part of the

ERK/MAPK pathway, which plays a role in plasticity, memory and learning, and is thought to be

involved in domestication. Genes involved in glutamate metabolism, like GRIK3 (present in cats,

horses, and humans), is hypothesized to play a role in reducing fear responses in domesticated

animals compared to wild animals (Theofanopoulou et al, 2017).

Given that humans have undergone a process of self-domestication, behavioral modernity

may have risen from a brand of social intelligence unique to domesticates. Experiments with

domesticated dogs show that these animals are uniquely skilled at understanding human social

cues (Hare & Tomasello, 2005). Data derived from a variety of experiments conducted by Hare
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and Tomasello with dogs in which humans attempt to communicate the location of hidden food

through the use of pointing, nodding, or gazing toward the target demonstrated that dogs are

highly skilled at these tasks. Dogs are even able to outperform chimps on these types of tasks.

Wolves raised by humans do not share these abilities, while dogs across a spectrum of exposure

to humans do, suggesting that these skills result from domestication. Silver foxes selected for

tameness also exhibited capabilities in using human social cues on par with dogs. Both species

were able to outperform great apes on these types of tasks Chimps hold the necessary cognitive

skills to carry out these tasks, but lack the social tolerance to successfully engage in this type of

cooperative communication (Hare & Tomasello, 2005). While bonobos also perform poorly in

this area, eye tracking data has revealed that they are sensitive to human gaze direction and focus

on the eyes of experimenters in order to orient themselves while chimpanzees do not. In general,

bonobos also show greater cooperation than chimpanzees when it comes to things like

food-sharing (Hare, 2017). Human infants are able to successfully complete these

communicative tasks as early as 14 months of age, in tandem with language acquisition (Hare &

Tomasello, 2005).

The development of more cooperative temperaments in humans through

self-domestication may have played a role in the evolution of complex social cognition that

affected our social structure and capacity for language (Benitez-Burraco & Progovac, 2020). A

hallmark of behavioral modernity is the capacity for language. Human demographics began to

shift during the Late Stone Age in the form of high population densities, leading to higher rates

of interaction among individuals (Cieri et al., 2014). Benitez-Burraco and Progovac (2020)

propose a mutually-reinforcing feedback loop between the forces of self-domestication and

advances in human language. As language complexity increased, verbal communication came to

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replace physical aggression, supporting cooperation and the development of culture

(Benitez-Burraco & Progovac, 2020). Wrangham (2019) proposes that the development of

languages reinforced the self-domestication process through his “language-based conspiracy”

hypothesis. In order to establish social rules that would lead to selection against aggression, the

development of language is essential. Linguistic capability allowed for the establishment of

coalitions that could punish or kill alpha males for the aggressive bullying (Wrangham, 2019).

This theory complements the idea that self-domestication in bonobos occurred through sexual

selection by female coalitions (Hare et al., 2012). The development of language promoted

proactive, or premeditated, forms of aggression over reactive, or impulsive aggression, which is

exemplified in present-day culture (Benitez-Burraco & Progovac, 2020; Wrangham, 2019).

Analyses of non-human domesticates provide a list of characteristics associated with the

domestication process, that, when compared to humans provide strong evidence for

domestication syndrome in Homo sapiens. Behavioral, morphological, physiological, and genetic

data support my hypothesis that humans have self-domesticated, and furthermore, that this

process shaped modern behavior. It is plausible that the development of language interacted with

the domestication process in a mutually-reinforcing way. Through the study of evolutionary

processes, we arrive at conclusions upon which other academic disciplines such as psychology,

linguistics, or biology can grow, contributing to a more holistic understanding of humanity.

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References cited:

Benitez-Burraco, A., & Progovac, L. (2020). A four-stage model for language evolution under
the effects of human self-domestication. Language & Communication, 73, 1-17.
https://doi.org/10.1016/j.langcom.2020.03.002

Cieri, R. L., Churchill, S. E., Franciscus, R. G., Tan, J., & Hare, B. (2014). Craniofacial
feminization, social tolerance, and the origins of behavioral modernity. Current
Anthropology, 55(4), 419-443. 10.1086/677209

Hare, B. (2017). Survival of the Friendliest: Homo sapiens Evolved via Selection for
Prosociality. Annual Review of Psychology, 68, 155-186.
10.1146/annurev-psych-010416-044201

Hare, B., & Tomasello, M. (2005). Human-like social skills in dogs? Trends in Cognitive
Sciences, 9(9), 439-444. 10.1016/j.tics.2005.07.003

Hare, B., Wobber, V., & Wrangham, R. (2012). THe self-domestication hypothesis: evolution of
bonobo psychology is due to selection against aggression. Animal Behaviour, 83,
573-585. 10.1016/j.anbehav.2011.12.007

Theofanopoulou, C., Gastaldon, S., O'Rourke, T., Samuels, B. D., Messner, A., Martins, P. T.,
Delogu, F., Alamri, S., & Boeckx, C. (2017). Self-domestication in Homo sapiens:
insights from comparative genomics. PloS one, 12(10), e0185306.
https://doi.org/10.1371/journal. pone.0185306

Wrangham, R. W. (2019). Hypothesis for the evolution of reduced aggression in the context of
human self-domestication. Frontiers in Psychology, 10, 1914. 10.3389/fpsyg.2019.01914