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. 2026 Jan 21;21(1):e0336161. doi: 10.1371/journal.pone.0336161

Gatherer ancestry associated with national happiness

Matthew Frederick Basilico 1,2,3,4,*
Editor: Ran Barkai5
PMCID: PMC12822993  PMID: 41563945

Abstract

Efforts to improve human emotional wellbeing through economic growth have seen varied success. One interpretation of the lack of wellbeing returns to economic growth is that humans may have been more emotionally suited to patterns of life in pre-agricultural societies. This study examines the hypothesis, dating to Rousseau, that descendants of hunter-gatherer societies have higher levels of subjective wellbeing. It utilizes data from 1265 small scale societies in the Murdock Ethnographic Atlas to construct a country-level measure of gatherer ancestry. Average country-level happiness and life satisfaction were derived from the World Values Survey which covered 104 countries from 1981–2014. Gatherer ancestry was significantly associated with happiness, controlling for contemporary income per capita (beta = 13.58; standard error = 3.0, R2 = 11.8%, p < 0.01). Results were robust to an extensive list of historical and contemporary controls. The findings are consistent with the hypothesis that gatherer lifestyle organization may hold insights for human emotional wellbeing.

Introduction

The relationship between human economic organization and societal emotional wellbeing continues to hold many puzzles. The well-known Easterlin Paradox is the documentation of a pattern that country-level economic growth and happiness are not meaningfully correlated in the long-run since data became available in the 1960s, a relationship that is of increasing concern given the consequences of economic growth for the environment [1,2]. Income and emotional wellbeing are also tenuously related for middle to high income individuals in the United States and Europe [3,4]. Emotional distress among young people has appeared at new levels in the United States despite historically unprecedented average levels of consumption and effective material living standards [5,6]. The notion that individual income improvements should lead to wellbeing improvements remains central to economic modelling and policymaking [7,8].

Other prominent social theorists, such as Jean-Jacques Rousseau, have hypothesized that hunter-gatherer social organization may hold benefits for human emotional wellbeing despite income limitations. Rousseau hypothesized that these benefits are eroded in societies with a higher degree of division of labor and thus inequality [9]. Contemporary evolutionary psychologists, such as Randolph Nesse, Glenn Geher and Nikhil Chaudhary, have similarly hypothesized that high prevalence rates of emotional distress, anxiety and depression may be the result of a “mismatch” between human emotional systems which evolved in the environment of small-scale societies and contemporary human environments (e.g., typically hierarchical, non-subsistence, including agricultural and industrial organization) [1015]. The work of these theorists leads to the hypothesis that wellbeing levels may be higher on average in small-scale hunter-gatherer societies than in contemporary industrialized societies, despite the higher consumption levels of the latter. The hypothesis has been given empirical support by several recent studies. For example, Miñarro and colleagues (2021) find higher levels of subjective wellbeing in a sample of minimally-monetized societies compared to high income countries [16]. Reyes-Garcia and colleagues (2021) find high levels of subjective wellbeing in a sample of 474 adults from 3 small-scale societies (Baka, Punan and Tsimane’), while Galbraith and colleagues (2024) similarly find high levels of life satisfaction in a sample of 19 small-scale societies [16,17]. Fedurek and colleagues (2023) find that status does not correlate with physiologic stress levels in a sample of Hadza hunter-gatherer men, in contrast to consistent findings in high-income societies of a correlation between stress and status [18]. Gurven and colleagues (2025) examine patterns of subjective wellbeing across the lifecycle in three small-scale societies, discovering departures from the typical age-patterns observed in high-income country samples [19]. The fields of Positive Evolutionary Psychology and Evolutionary Psychiatry have expanded rapidly over the last decade around the insight that human affect may be naturally attuned to particular historical small-scale societies [10,12,2024].

The present study seeks to enhance our understanding of how wellbeing levels in small-scale societies may relate to contemporary, country-level subjective wellbeing. It investigates the hypothesis that societies with a higher levels of ancestry from hunter-gatherer societies may be associated different rates of subjective wellbeing. Recent innovations in historical economics have allowed scholars to link features of ancestral societies to modern-day populations, and investigate relationships between historical economic organization and modern-day beliefs [2527]. One of the initial works in this space examined how ancestral cultures that used plough-based agriculture had a higher gender-based segregation of labor, and modern-day populations with a higher fraction of descendants from plough-based societies have more unequal gender norms today [28]. Since this paper, over a dozen works in historical economics have uncovered mechanisms of cultural persistence, linking features of pre-industrial societies to contemporary economic and cultural outcomes [26,27]. While it has been well-utilized in several influential economics papers, the approach does have important limitations, such as assigning country-level cultural ancestry based on language group, reducing important sources of variation [29,30]. Nevertheless, the approach makes possible a degree of mapping from the features of ancestral societies to help characterize the influence on contemporary populations.

The Ethnographic Atlas tables, compiled in 1967 by Anthropologist George Peter Murdock, “incorporate nearly 50,000 distinct items of information” based upon earliest available ethnographic sources [31]. The data in the Ethnographic Atlas reflects understanding of human social organization for several generations up to the 20th century. Although most humans before the neolithic revolution (circa 10,000 BCE) are believed to have lived in small-scale societies, Henrich and colleagues have noted that the substantial heterogeneity of human societal organization over the past several millennia, including pastoral, agricultural and industrial practices, has led to important variation in cultural norms [3234]. Hence, we are able to ask the question: is a higher fraction of recent ancestral descendants from hunter-gatherer societies associated with contemporary average wellbeing, controlling for income? As noted above, several investigations have directly sampled wellbeing levels in hunter- or gatherer-based contemporary societies and typically found higher levels of subjective wellbeing than industrialized country samples, although these studies are naturally limited by the remaining prevalence of this form of social organization [3537]. The accumulating body of evidence in historical economics suggests, if they existed, some of these wellbeing advantages could transmit to modern day populations through channels of cultural persistence [38,39].

The present study investigates this question using contemporary tools of historical economics, including linking the Murdock Ethnographic Atlas to countries, and utilizing cross-country subjective wellbeing data available from the World Values Survey and economic controls available from the World Bank Group’s World Development Indicators.

Methods

This study examines how the variance in measured average population-level happiness varies with the fraction of ancestry from hunter-predominant, gather-predominant, or agricultural-predominant societies. We follow the methodology utilized in Alesina, Giuilano 2013 which linked the Murdock Ethnographic Atlas data to country-level populations using the steps outlined below [28].

The Ethnographic Atlas was constructed in 1967 by George Peter Murdock, coding data on 1265 small scale societies utilizing earliest-available ethnographic evidence. The dataset includes over 100 variables, including coding economic organization features such as predominant mode of food production, levels of political hierarchy, and kinship practices [40]. Within the ethnographic atlas, variable v42 indicates the predominant mode of food production in society. Here, “Gathering contributes most” (103 of 1265 ethnic groups) is coded as Gathering Predominant, “Hunting contributes most” is coded as Hunting Predominant (75 of 1265 ethnic groups). “Fishing contributes most” is coded as Fishing Predominant (114 of 1265 ethnic groups). For comparison, the majority of societies are coded as “Intensive agriculture contributes most” (270 groups), “Extensive agriculture contributes most” (475 groups), “Agriculture contributes most, type unknown” (86 groups), or “Pastoralism contributes most” (77 groups). Remaining observations are either more than one equal sources (64 groups), or missing observation (1 group). The ethnographic atlas data is then linked to the Ethnologue: Languages of the World dataset, which maps the prevalence of 7612 languages globally. Each language in the Ethnologue is linked to an observed society in the Ethnographic atlas. Finally, the Ethnologue is linked to modern nation states by the percentage of the population speaking the language coded. Following Giuilano and Nunn 2018, each variable is then rescaled to adjust for missing values. Namely, v42 group 1 denotes missing values (1 of 1265 ethnic groups). The resulting country-level estimate of “Gathering Predominant Ancestry” is thus the fraction of contemporary ancestry estimated to have descended from Gathering predominant societies (v42). Country-level “Hunting Predominant Ancestry” is estimated in a similar fashion, and the two fractions are summed to create the variable “Hunting or Gathering Predominant Ancestry.” This technique was pioneered by Alesina, Giuiliano and Nunn 2013, and a mapping to country-level data is available from Giuiliano and Nunn 2018 [26,28]. The result of this mapping is an estimate of the country-level fraction of ancestral dependence on gathering or hunting. A similar procedure is used to create the ancestral fractions of control variables used from the ethnographic atlas.

Outcome data on happiness and life satisfaction at the country level comes from the World Values Survey (WVS) Wave 1–6 Key Aggregates, available online through worldvaluessurvey.org. Waves 1–6 covered 108 countries at least once from 1981–2014. Happiness was recorded in each survey using a four-point scale, according to the question, “taking all things together, would you say you are: very happy, rather happy, not very happy, not at all happy.” Life satisfaction is recorded in each survey using a scale from 0 to 10, indicating the degree to which respondents “are satisfied with their life as a whole.” Answers are rescaled from 0 to 1.0, utilizing fractions for intermediate responses [41]. Average happiness and life satisfaction are reported at the country-level for the most recent wave available in the dataset.

To estimate the relationship between the fraction of ancestral population from hunter and gatherer populations and subjective wellbeing, ordinary least squares (OLS) regression with heteroskedasticity robust standard errors was conducted using Stata 18 (StataCorp 2024) [42]. Our baseline specification includes contemporary and historical controls as well as continent fixed effects. Contemporary controls include GDP per capita, using 2019 estimates available publicly from the World Bank Group’s World Development Indicators [43]. Historical controls include observables for key dimensions of societal variation available in the Murdock Atlas: levels of political hierarchy, presence of polygyny, patrilineal descent, and kinship score. Kinship score is an aggregate scale created from four Murdock Atlas variables reflecting degree of kinship tightness, including nuclear family, post-wedding residence, unilineal descent, segmented clans. The scale was developed by economist Benjamin Enke is publicly available from his 2019 paper Kinship, Cooperation and the Evolution of Moral Systems [30]. The linkage of these ancestral characteristics to the contemporary country-level was accomplished through the same procedure for the ancestral gathering variable outlined above. The practice of using a baseline specification with contemporary GDP per capita and selected historical controls was established by Alesina et al 2013 and has become standard practice across over 20 papers in the historical economics literature [27,28]. Coefficient estimates with and without the presence of these controls are presented in the results.

Results

Ethnographic Atlas and World Values Survey outcome data were matched on n = 102 countries. World Bank data for contemporary controls was matched on n = 100 of the full sample, and a full set of Ethnographic controls, including the Enke Kinship Score, was matched for n = 97 countries. For n = 102 countries with World Values Survey and Ethnographic Atlas observables, average ancestral gatherer fraction was 0.048% (range: 0.0 to 1.26%, std. dev. 0.18%), average hunter ancestry was 0.044% (range: 0.0 to 2.1%, std. dev. 0.24%), average hunter or gatherer ancestry was 0.092% (range: 0.0 to 2.3%, std. dev. 0.31%).

In our baseline specification, the fraction of the population with ancestry from gathering-predominant societies is positively associated with country-level happiness, controlling for contemporary income per capita (beta = 13.58; Standard Error (SE) = 3.0, R2 = 11.8%, p < 0.01; Table 1, Column 2). A one standard deviation increase (0.4%) in the fraction of gathering predominant accounts for a 0.05 point increase in average country-level happiness, equivalent to roughly 55% of the standard deviation (0.091 points) of average happiness rates between countries. The fraction of population ancestry from gatherer-predominant populations is positively associated with country-level happiness with or without including controls for GDP per capita, historical controls, or continent fixed effects (Table 2; Columns 2–5).

Table 1. Gathering Predominant Ancestry Fraction and Country-Level Happiness.

(1) (2) (3) (4) (5)
Happiness
Gathering Predominant Ancestry 13.86*** 13.58*** 10.61*** 13.16*** 11.53***
(2.503) (2.997) (1.883) (3.635) (2.534)
Log GDP Per Capita 2019 (WDI) 0.00751** 0.0136*** 0.0143*** 0.0184***
(0.00314) (0.00411) (0.00437) (0.00417)
Average Level of Political Hierarchy 0.0232 0.0250*
(0.0143) (0.0131)
Average Settlement Complexity 0.00244 0.00489
(0.00704) (0.00808)
Patrilineal Descent 0.0224 0.0239
(0.0340) (0.0277)
Matrilineal Descent −0.125 −0.0803
(0.0778) (0.0861)
Polygynous 0.00873 0.0657
(0.0482) (0.0660)
Plough Use −0.170*** −0.168***
(0.0324) (0.0340)
Kinship Score −0.0452 −0.0446
(0.0484) (0.0398)
Constant 0.697*** 0.558*** 0.470*** 0.502*** 0.373***
(0.00910) (0.0624) (0.0668) (0.125) (0.0994)
Observations 102 100 100 97 97
R-squared 0.081 0.118 0.247 0.361 0.488
Continent Fixed Effects Yes Yes

Table 1. Results from OLS regression of WVS average country-level Happiness on Gathering Predominant Ancestry (Column 1) as well as including contemporary and historical controls (Columns 2-5). Robust standard errors in parentheses. *** p < 0.01, ** p < 0.05, * p < 0.1.

Table 2. Hunting Predominant Ancestry Fraction and Country-Level Happiness.

(1) (2) (3) (4) (5)
Happiness
Hunting Predominant Ancestry 4.138** 3.754*** −1.028 2.767* −1.716
(1.685) (1.202) (1.692) (1.498) (1.685)
Log GDP Per Capita 2019 (WDI) 0.00737** 0.0137*** 0.0138*** 0.0178***
(0.00310) (0.00410) (0.00440) (0.00410)
Average Level of Political Hierarchy 0.0147 0.0230
(0.0161) (0.0145)
Average Settlement Complexity 0.00361 0.00596
(0.00715) (0.00817)
Patrilineal Descent 0.00812 0.0128
(0.0347) (0.0280)
Matrilineal Descent −0.147* −0.0909
(0.0823) (0.0908)
Polygynous 0.0209 0.0836
(0.0446) (0.0636)
Plough Use −0.160*** −0.159***
(0.0338) (0.0335)
Kinship Score −0.0348 −0.0494
(0.0497) (0.0400)
Constant 0.702*** 0.564*** 0.472*** 0.523*** 0.388***
(0.00918) (0.0618) (0.0666) (0.127) (0.129)
Observations 102 100 100 97 97
R-squared 0.013 0.059 0.211 0.304 0.449
Continent Fixed Effects Yes Yes

Table 2. Results from OLS regression of WVS average country-level Happiness on Hunting Predominant Ancestry (Column 1) as well as including contemporary and historical controls (Columns 2-5). Robust standard errors in parentheses. *** p < 0.01, ** p < 0.05, * p < 0.1.

The fraction of population with ancestry from hunter-predominant societies is positively associated with country-level happiness, controlling for contemporary income per capita (beta = 3.75, SE = 1.2, R2 = 5.9%, p < 0.01; Table 2, Column 2). The coefficient is approximately 27% as large as the estimated coefficient on gathering-predominant ancestry. However, when including controls for continent fixed effects and historical controls, the results are mostly insignificant (Table 2, Columns 3–5).

The fraction of ancestry from either hunting or gathering-predominant societies was positively associated with country-level happiness, controlling for contemporary income per capita (beta = 6.62, SE = 2.5, R2 = 10.1%, p < 0.01; Table 3, Column 2). Results were robust to inclusion of historical controls (Table 3, Column 2) and were no longer significant when including continent fixed effects (Table 3, Columns 3, 5).

Table 3. Hunting or Gathering Predominant Ancestry Fraction and Country-Level Happiness.

(1) (2) (3) (4) (5)
Happiness
Hunting or Gathering Predominant Ancestry 7.225*** 6.618*** 3.056 5.882** 2.790
(2.671) (2.469) (2.307) (2.735) (2.788)
Log GDP Per Capita 2019 (WDI) 0.00721** 0.0134*** 0.0144*** 0.0178***
(0.00311) (0.00410) (0.00437) (0.00409)
Average Level of Political Hierarchy 0.0158 0.0211
(0.0148) (0.0140)
Average Settlement Complexity 0.00262 0.00549
(0.00708) (0.00815)
Patrilineal Descent 0.0122 0.0126
(0.0338) (0.0273)
Matrilineal Descent −0.135 −0.0927
(0.0805) (0.0883)
Polygynous 0.0159 0.0778
(0.0450) (0.0642)
Plough Use −0.159*** −0.159***
(0.0326) (0.0333)
Kinship Score −0.0275 −0.0410
(0.0489) (0.0401)
Constant 0.697*** 0.563*** 0.475*** 0.506*** 0.390***
(0.00928) (0.0619) (0.0665) (0.125) (0.127)
Observations 102 100 100 97 97
R-squared 0.064 0.101 0.219 0.338 0.454
Continent Fixed Effects Yes Yes

Table 3. Results from OLS regression of WVS average country-level Happiness on Hunting or Gathering Predominant Ancestry (Column 1) as well as including contemporary and historical controls (Columns 2-5). Robust standard errors in parentheses. *** p < 0.01, ** p < 0.05, * p < 0.1.

Results for life satisfaction were qualitatively similar to the results on happiness. The fraction of the population with ancestry from gathering-predominant societies is positively associated with country-level life satisfaction, controlling for contemporary income per capita (beta = 7.74; Standard Error (SE) = 4.5, R2 = 33.0%, p < 0.1; Table 4, Column 2). The fraction of population ancestry from gatherer-predominant populations is positively associated with country-level life satisfaction with or without including controls for GDP per capita, historical controls, or continent fixed effects (Table 4; Columns 2–5). Results for life satisfaction and hunting, as well as for hunting or gathering ancestry, were qualitatively similar to those for happiness, and are reported in Supporting Information Tables S1 and S2 in S1 File. Inclusion of Fishing predominant ancestry with hunting and gathering ancestry did not change results qualitatively, as reported in Supporting Information Tables S3-S5 in S1 File.

Table 4. Gathering Predominant Ancestry Fraction and Country-Level Life Satisfaction.

(1) (2) (3) (4) (5)
Life Satisfaction
Gathering Predominant Ancestry 8.444** 7.748* 4.722* 10.67** 9.070***
(4.199) (4.498) (2.773) (4.889) (3.016)
Log GDP Per Capita 2019 (WDI) 0.0227*** 0.0249*** 0.0257*** 0.0293***
(0.00310) (0.00437) (0.00444) (0.00454)
Average Level of Political Hierarchy 0.0446*** 0.0421***
(0.0147) (0.0154)
Average Settlement Complexity −0.00420 −0.00488
(0.00715) (0.00727)
Patrilineal Descent 0.0933*** 0.108***
(0.0290) (0.0287)
Matrilineal Descent 0.0110 0.0990*
(0.0630) (0.0538)
Polygynous −0.0823 −0.00868
(0.0585) (0.0537)
Plough Use −0.168*** −0.152***
(0.0338) (0.0339)
Kinship Score −0.141*** −0.120***
(0.0404) (0.0391)
Constant 0.639*** 0.219*** 0.165** 0.245** 0.104
(0.0109) (0.0617) (0.0708) (0.111) (0.104)
Observations 102 100 100 97 97
R-squared 0.022 0.330 0.480 0.492 0.634
Continent Fixed Effects Yes Yes

Table 4. Results from OLS regression of WVS average country-level Life Satisfaction on Gathering Predominant Ancestry (Column 1) as well as including contemporary and historical controls (Columns 2-5). Robust standard errors in parentheses. *** p < 0.01, ** p < 0.05, * p < 0.1.

Discussion

There appears to be a statistically significant association between the fraction of ancestry from gatherer-predominant societies and contemporary average country-level happiness. The relationship is robust to a set of commonly used contemporary and historical controls, as well as continent fixed effects. The results also suggest a smaller effect for the influence on hunter-predominant ancestry, although these results were not fully robust to the inclusion of historical controls and continent effects. The findings are also notable given the limited quantitative contribution of hunting and gathering predominant societies, accounting for 178 of 1265 societies in the Ethnographic Atlas.

There are several potential explanations for these results. One important explanation is that the finding could be due to omitted variable bias, a threat to any study using associative techniques. Although the Gatherer results are robust to an extensive list of controls, those including hunting are less robust with inclusion of continent controls. It is also notable that the baseline fractions of hunter and gatherer ancestry in most countries are quite small. However, it is possible that the country-level cultural influences of this ancestry extend beyond the estimated quantitative fraction. While we caution against any causal interpretation of these results, several features of the analysis support a real association between these variables. These factors include the historical nature of the predictor of interest, with ancestral fraction contributions set in time well before the estimation of contemporary happiness rates. Additionally, the statistically and economically significant positive relationship remained strong across specifications, including uncontrolled regression as well as OLS regression with rich sets of historical controls, contemporary GDP.

The possibility of a relationship between gatherer ancestry and societal happiness is consistent with studies of wellbeing in contemporary hunter-gatherer societies [10,44,45]. Franckowiak and colleagues (2021) find subjective happiness among the Hadza compares favorably to a sample of Polish individuals, while Reyes-Garcia et al 2021 find notably high and seasonally stable levels of wellbeing among their study populations from three small-scale societies [16,37]. Theorists from Thoreau to Weber have posited that detachment from nature has had negative implications for the human experience, such as separation from natural cycles of life [46,47]. The growing body of evidence for nature exposure and social rhythm therapy in various psychiatric disorders attests to the salience of this concept to human affective wellbeing [48,49]. Another potential mechanism, the argument of Rousseau’s famous 1761 discourse on the origin of inequality, is that human hierarchies are responsible for a large fraction of human unhappiness, and emotional wellbeing may have been higher in ancestral hunter-gatherer societies due to reduced class stratification [9]. This mechanism could be driven by either material inequality or formation of status hierarchy, a distinction beyond the scope of the analysis in this paper but indicated for future study [50,51]. Of note, average ancestral level of political hierarchy and average ancestral settlement complexity were included as controls in the specification, and were not significant.

There are several limitations to this study. One important limitation is the use of the Ethnologue approach to linking ancestral societal features to modern populations. In the approach, a country’s ancestral representation in the Murdock Atlas is estimated using the Ethnologue language atlas. With the prevalence of language as the basis of mapping societies in the Murdock atlas, there is much additional ancestral richness and heterogeneity that is not incorporated in this approach. This critique has been noted in several key papers in historical economics, which nevertheless utilize the Ethnologue approach as the most effective known method for linking the Murdock Atlas to modern nation states [26,2830,52]. Another important limitation is the associative nature of the findings as discussed above, and caution should be used in applying a causal interpretation. Additionally, the World Values Survey data covers only 104 countries, omitting variation from dozens of countries globally [41]. Further investigation could link micro-level analyses of wellbeing to the Ethnographic Atlas, or identify ecological instruments for gathering ancestry to improve identification.

Conclusion

Contemporary techniques and data availability in historical economics have indicated a positive, economically significant relationship between the fraction of ancestry from gatherer societies and contemporary average subjective wellbeing. While additional work is necessary to confirm this relationship, the study finds that ancestral social organization may contribute meaningfully to modern population-level wellbeing, and that these relationships can be characterized using techniques from historical economics. Given the limitations of economic growth in improving happiness, and negative externalities including global warming, gatherer ancestry may hold wisdom for population wellbeing worth further exploration.

Supporting information

S1 File. Supporting Information.

Tables S3-S5 are included in the Supporting Information file.

(DOCX)

pone.0336161.s001.docx (31.2KB, docx)

Acknowledgments

The author acknowledges helpful feedback from presentation attendees at the American Psychiatric Association Annual Meetings 2025.

Data Availability

All data are publically available with sources references in the manuscript. In addition, the data underlying the results presented in the study and replication code are available from: https://github.com/mfbasilico/GatheringHappiness.

Funding Statement

The author(s) received no specific funding for this work.

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Decision Letter 0

Ran Barkai

8 Sep 2025

Dear Dr. Basilico,

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[Note: HTML markup is below. Please do not edit.]

Reviewers' comments:

Reviewer's Responses to Questions

Comments to the Author

1. Is the manuscript technically sound, and do the data support the conclusions?

Reviewer #1: Yes

Reviewer #2: Partly

**********

2. Has the statistical analysis been performed appropriately and rigorously? -->?>

Reviewer #1: I Don't Know

Reviewer #2: Yes

**********

3. Have the authors made all data underlying the findings in their manuscript fully available??>

The PLOS Data policy

Reviewer #1: Yes

Reviewer #2: No

**********

4. Is the manuscript presented in an intelligible fashion and written in standard English??>

Reviewer #1: Yes

Reviewer #2: Yes

**********

Reviewer #1: This is a very significant article, and with a few additions and some clearer wording, should definitely be published

The paper is about two related, but distinct, hypothesized phenomena. The first is that there exist “links between hunter-gatherer economic organization and emotional wellbeing”. The second is that these links may be passed on to subsequent generations that have transitioned to non-hunter-gatherer economies “through transgenerational cultural transmission”. From the author’s conclusion, it is clear that the second of these phenomena is what the paper focusses on and hopes to demonstrate. This is confirmed in the first sentence in the methodology section, which states: “This study examines how the variance in measured average population-level happiness varies with the fraction of ancestry from hunter-predominant, gather-predominant, or agricultural-predominant societies.”

However, the author needs to signal that these are two rather different phenomena, particularly in terms of what might account for them. In the case of the first hypothesis, the author briefly suggests possible explanations: that “forms of economic organization more closely tied to rhythms of natural environmental cycles contained advantages for emotional wellbeing”, and also that emotional wellbeing is related to the relative lack of inequality and “a lack of class stratification” among hunter-gatherers.

Where the author refers to the relationship between happiness and low levels of inequality, some reference is needed to the distinction between material inequality and inequality based on the existence of a status hierarchy, as these may not coincide.

In the case of the second hypothesis, the author leaves several issues only briefly examined, and in particular when. why and how the cultural transmission of the ancestral benefits for hunter-gatherer emotional well-being persist or do not persist in descendant generations. Much depends on the author’s reliance on a single 2018 source, Giuliano P, Nunn N. Ancestral Characteristics of Modern Populations. Economic History of Developing Regions. Part of the issue here is in the method to distinguish between a population with an ancestry from hunter-gatherer societies, and those without such an ancestry. This distinction is made on the basis of language, also implying that this is the mode of cultural transmission. According to Google Scholar, the Giuliano and Nunn article has been cited no less than 117 times. Given its key place in the author’s argument, I would have expected some discussion of any issues raised that are relevant to his hypothesis in this subsequent literature.

In addition, I found that in some places the wording was either a bit awkward, ungrammatical, or difficult to understand. As examples, I note the following:

“and appears to have intellectual the culture of Anglo-American analytic philosophy, including the notion of preference optimization dating to Jeremy Bentham.”

“Social theorists including Freud, Durkheim and Rousseau identified potential links between hunter-gatherer economic organization and emotional wellbeing that would not be well captured by a utility from consumption framework.”

“caution should be used in applying a causal.”

Reviewer #2: This article combines a previously-published method linking the geographical distribution of ethnic ancestry, via language groups and pre-industrial ethnographic records, with the World Values Survey life satisfaction scores. The main finding is a statistically-significant, positive relationship between the estimated fraction of Gatherer ancestry and life satisfaction, at the national level. I think this is an interesting and important finding, worthy of publication. However I think the article is not sufficiently developed at present, and there are a number of aspects that need to be strengthened so that the analyses are performed to a high technical standard and the conclusions adequately supported.

1. I think it’s important to expand the discussion of the ethnologue approach. I know the linking to the ethnography is not the new contribution of this work, but I was surprised that the original references for the method did not seem to address this aspect. Is the United States linked, in this analysis, to an English heritage because that’s the predominant language for the country? If so, how does this affect countries that strongly enforced national languages in the 19th century, like France? A bit of discussion about this seems pertinent - there might be systematic underestimates of ancestry of groups that were coerced to speak a dominant language rather than maintaining their traditional language. I think it’s quite important for the conclusions and needs to be discussed, particularly given that prior research has shown that immigrants tend to shift their life satisfaction toward their new country, even within a given lifetime (e.g. Helliwell et al., International Migration and World Happiness, World Happiness Report 2018).

2. It’s not clear to me that the binary distinctions of ‘gathering predominant’ vs. ‘hunting predominant’ is the best way to analyze the atlas data. Forager lifeways were diverse, and usually involved a combination of hunting, fishing and gathering in variable proportions (see e.g. Kelly, 2013, Lifeways of Hunter Gatherers). It would therefore be surprising that the hunting-predominant societies were not also doing a lot of gathering (and most were probably fishing too). I see that the Atlas includes a column that actually assesses all forms of subsistence - could this be used instead of the simple binary distinctions?

3. I feel the paper needs some more substantive thought into exactly how the ancestral populations may be connected to average national life satisfaction. I presume that the fractions of modern populations that are linked to either gathering or hunting are mostly very small? (A map would be very helpful, to show the % of each ancestry by country.) If so, how would it be that these small populations manage to exert significant influence on the entire national populations? There is some literature on South American societies that might be useful for this, such as regarding the Buen Vivir idea (Villalba, Buen Vivir vs. Development: a paradigm shift in the Andes? Third World Quarterly, 2014). Related to this, it also might be helpful to discuss the fact that really this is getting at populations with recent hunting and gathering ancestry, since all humans are descended 100% from hunters and gatherers if one goes back far enough in time.

4. I think it would also be worthwhile comparing with the Gallup world poll results. There is some important nuance between how people answer the Cantril ladder vs. life satisfaction questions (Nilsson, The Cantril Ladder elicits thoughts about power and wealth, Scientific Reports 2024 https://doi.org/10.1038/s41598-024-52939-y), so I don’t think it would undermine the results here if the Gallup results were not similarly correlated with hunter-gatherer ancestry. Whatever it shows, it would provide useful additional insight.

5. There are also some additional references that may prove helpful. The works by Minarro (Miñarro et al., Happy without money: Minimally monetized societies can exhibit high subjective well-being. PLoS ONE 2021 https://doi.org/10.1371/journal.pone.0244569) and Galbraith (Galbraith et al., High life satisfaction reported among small- scale societies with low incomes, PNAS 2024 https://doi.org/10.1073/pnas.2311703121) seem particularly relevant, and would tend to support the findings here with a more direct, though more locally-focused method.

6. The discussion states that ‘findings are also notable the limited quantitative contribution of hunting and gathering predominant societies, accounting for 180 of 1265 societies in the Ethnographic Atlas.’ So what were the other 1085 societies doing for subsistence?

7. The figure does not seem appropriate. This figure simply shows the previously reported WVS life satisfaction score by country, without adding anything related to the work at present. I think it would be far more useful to include a figure of the hunter and gatherer ancestry fractions by country, or similar. Perhaps the three could be shown as adjacent panels - each of the two ancestries, and the life satisfaction?

Again I think this is potentially very valuable work and hope that these comments can help to strengthen it.

**********

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Reviewer #1: Yes:  Adrian Tanner

Reviewer #2: No

**********

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PLoS One. 2026 Jan 21;21(1):e0336161. doi: 10.1371/journal.pone.0336161.r002

Author response to Decision Letter 1


19 Oct 2025

This text has been uploaded seperately as part of the rebusmission, with color-coding to indicate query responses.

1. Please ensure that your manuscript meets PLOS ONE's style requirements, including those for file naming. The PLOS ONE style templates can be found at

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https://journals.plos.org/plosone/s/file?id=ba62/PLOSOne_formatting_sample_title_authors_affiliations.pdf

I have reviewed the style requirements, including file naming, and to my understanding this draft is consistent with the requirements. I am very happy to make any additional adjustments if indicated.

2. We note that your Data Availability Statement is currently as follows: [All relevant data are within the manuscript and its Supporting Information files.]

Please confirm at this time whether or not your submission contains all raw data required to replicate the results of your study. Authors must share the “minimal data set” for their submission. PLOS defines the minimal data set to consist of the data required to replicate all study findings reported in the article, as well as related metadata and methods (https://journals.plos.org/plosone/s/data-availability#loc-minimal-data-set-definition).

For example, authors should submit the following data:

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- The values used to build graphs;

- The points extracted from images for analysis.

Authors do not need to submit their entire data set if only a portion of the data was used in the reported study.

All the data used in this study is publicaly available, from the four following data sources: the World Bank World Development Indicators, the World Values Survey, the supporting files of Giuliano and Nunn 2018 and supporting files of Enke 2019.

However, I have now also taken the additional step of posting the merged dataset and STATA code for production of all tables in the analyses in github: https://github.com/mfbasilico/GatheringHappiness. The posted code reproduces all tables.

If your submission does not contain these data, please either upload them as Supporting Information files or deposit them to a stable, public repository and provide us with the relevant URLs, DOIs, or accession numbers. For a list of recommended repositories, please see https://journals.plos.org/plosone/s/recommended-repositories.

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We require you to either (1) present written permission from the copyright holder to publish these figures specifically under the CC BY 4.0 license, or (2) remove the figures from your submission:

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In the figure caption of the copyrighted figure, please include the following text: “Reprinted from [ref] under a CC BY license, with permission from [name of publisher], original copyright [original copyright year].”

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The following resources for replacing copyrighted map figures may be helpful:

USGS National Map Viewer (public domain): http://viewer.nationalmap.gov/viewer/

The Gateway to Astronaut Photography of Earth (public domain): http://eol.jsc.nasa.gov/sseop/clickmap/

Maps at the CIA (public domain): https://www.cia.gov/library/publications/the-world-factbook/index.html and https://www.cia.gov/library/publications/cia-maps-publications/index.html

NASA Earth Observatory (public domain): http://earthobservatory.nasa.gov/

Landsat: http://landsat.visibleearth.nasa.gov/

USGS EROS (Earth Resources Observatory and Science (EROS) Center) (public domain): http://eros.usgs.gov/#

Natural Earth (public domain): http://www.naturalearthdata.com/

Thank you very much for this guidance. After reflection including Reviewer 2 Comment #7 as well as copyright limitations, I am in agreement that the figure does not add sufficiently to the analysis, and that these results are available elsewhere. I have included an additional table in the manuscript from the helpful suggestion regarding Life Satisfaction, and instead of a map I have included summary statistics within the text on the prevalence and variance of hunter and gatherer ancestry using the Ethnologue approach as suggested by this comment.

4. If the reviewer comments include a recommendation to cite specific previously published works, please review and evaluate these publications to determine whether they are relevant and should be cited. There is no requirement to cite these works unless the editor has indicated otherwise.

Thank you very much for this comment. The updated manuscript includes an expanded literature review which touches on additional themes mentioned in the reviewers and editors, including wellbeing in small-scale societies, limitations to the Ethnologue approach, and the citations suggested by Reviewer 2 in Comment #5.

[Note: HTML markup is below. Please do not edit.]

Reviewers' comments:

Reviewer's Responses to Questions

Comments to the Author

1. Is the manuscript technically sound, and do the data support the conclusions?

The manuscript must describe a technically sound piece of scientific research with data that supports the conclusions. Experiments must have been conducted rigorously, with appropriate controls, replication, and sample sizes. The conclusions must be drawn appropriately based on the data presented.

Reviewer #1: Yes

Reviewer #2: Partly

________________________________________

2. Has the statistical analysis been performed appropriately and rigorously?

Reviewer #1: I Don't Know

Reviewer #2: Yes

As mentioned above, I have included data and STATA code used for each of the tables publicly on github. Of note, the standard errors in columns 4 & 5 of Tables 1-3 have been updated to heteroskedasticity-robust as originally indicated in the methods section.

________________________________________

3. Have the authors made all data underlying the findings in their manuscript fully available?

The PLOS Data policy requires authors to make all data underlying the findings described in their manuscript fully available without restriction, with rare exception (please refer to the Data Availability Statement in the manuscript PDF file). The data should be provided as part of the manuscript or its supporting information, or deposited to a public repository. For example, in addition to summary statistics, the data points behind means, medians and variance measures should be available. If there are restrictions on publicly sharing data—e.g. participant privacy or use of data from a third party—those must be specified.

Reviewer #1: Yes

Reviewer #2: No

As mentioned above, all data used in the analysis is available from 4 publicly available datasources (World Bank WDI, World Values Survey, Enke 2019 and Giuliano-Nunn 2018). I have taken the additional step of posting merged dataset with all data elements used in the analysis, and code for the tables.

________________________________________

4. Is the manuscript presented in an intelligible fashion and written in standard English?

PLOS ONE does not copyedit accepted manuscripts, so the language in submitted articles must be clear, correct, and unambiguous. Any typographical or grammatical errors should be corrected at revision, so please note any specific errors here.

Reviewer #1: Yes

Reviewer #2: Yes

________________________________________

5. Review Comments to the Author

Please use the space provided to explain your answers to the questions above. You may also include additional comments for the author, including concerns about dual publication, research ethics, or publication ethics. (Please upload your review as an attachment if it exceeds 20,000 characters)

Reviewer #1: This is a very significant article, and with a few additions and some clearer wording, should definitely be published

The paper is about two related, but distinct, hypothesized phenomena. The first is that there exist “links between hunter-gatherer economic organization and emotional wellbeing”. The second is that these links may be passed on to subsequent generations that have transitioned to non-hunter-gatherer economies “through transgenerational cultural transmission”. From the author’s conclusion, it is clear that the second of these phenomena is what the paper focusses on and hopes to demonstrate. This is confirmed in the first sentence in the methodology section, which states: “This study examines how the variance in measured average population-level happiness varies with the fraction of ancestry from hunter-predominant, gather-predominant, or agricultural-predominant societies.”

However, the author needs to signal that these are two rather different phenomena, particularly in terms of what might account for them. In the case of the first hypothesis, the author briefly suggests possible explanations: that “forms of economic organization more closely tied to rhythms of natural environmental cycles contained advantages for emotional wellbeing”, and also that emotional wellbeing is related to the relative lack of inequality and “a lack of class stratification” among hunter-gatherers.

Where the author refers to the relationship between happiness and low levels of inequality, some reference is needed to the distinction between material inequality and inequality based on the existence of a status hierarchy, as these may not coincide.

Thank you very much for this thoughtful comment. I have provided additional discussion on the distinction between material inequality and status hierarchy, and how each could be relevant to the results of the analysis.

In the case of the second hypothesis, the author leaves several issues only briefly examined, and in particular when. why and how the cultural transmission of the ancestral benefits for hunter-gatherer emotional well-being persist or do not persist in descendant generations. Much depends on the author’s reliance on a single 2018 source, Giuliano P, Nunn N. Ancestral Characteristics of Modern Populations. Economic History of Developing Regions. Part of the issue here is in the method to distinguish between a population with an ancestry from hunter-gatherer societies, and those without such an ancestry. This distinction is made on the basis of language, also implying that this is the mode of cultural transmission. According to Google Scholar, the Giuliano and Nunn article has been cited no less than 117 times. Given its key place in the author’s argument, I would have expected some discussion of any issues raised that are relevant to his hypothesis in this subsequent literature.

Thank you very much for this important comment. I have expanded the discussion of critiques of the Ethnologue approach within the Introduction and Discussion sections, within the section on study limitations. This approach has been used in dozens of papers within the historical economics literature; however, as you note there are important limitations, notably the assignment of a language group to an observation in the ethnographic atlas, and the use of language as the sole connection to ancestry in contemporary nation-states.

In addition, I found that in some places the wording was either a bit awkward, ungrammatical, or difficult to understand. As examples, I note the following:

“and appears to have intellectual the culture of Anglo-American analytic philosophy, including the notion of preference optimization dating to Jeremy Bentham.”

“Social theorists including Freud, Durkheim and Rousseau identified potential links between hunter-gatherer economic organization and emotional wellbeing that would not be well captured by a utility from consumption framework.”

“caution should be used in applying a causal.”

Thank you very much for these helpful examples. I edited each of areas mentioned to use clearer wording. I have also reviewed the manuscript for similar instances where the language could be clearer and adjusted accordingly.

Reviewer #2: This article combines a previously-published method linking the geographical distribution of ethnic ancestry, via language groups and pre-industrial ethnographic records, with the World Values Survey life satisfaction scores. The main finding is a statistically-significant, positive relationship between the estimated fraction of Gatherer ancestry and life satisfaction, at the national level. I think this is an interesting and important finding, worthy of publication. However I think the article is not sufficiently developed at present, and there are a number of aspects that need to be strengthened so that the analyses are performed to a high technical standard and the conclusions adequately supported.

1. I think it’s important to expand the discussion of the ethnologue approach. I know the linking to the ethnography is not the new contribution of this work, but I was surprised that the original references for the method did not seem to address this aspect. Is the United States linked, in this analysis, to an English heritage because that’s the predominant language for the country? If so, how does this affect countries that strongly enforced national languages in the 19th century, like France? A bit of discussion about this seems pertinent - there might be systematic underestimates of ancestry of groups that were coerced to speak a dominant language rather than maintaining their traditional language. I think it’s quite important for the conclusions and needs to be discussed, particularly given that prior research has shown that immigrants tend to shift their life satisfaction toward their new country, even within a giv

Attachment

Submitted filename: Response to Reviewers.pdf

pone.0336161.s003.pdf (294.7KB, pdf)

Decision Letter 1

Ran Barkai

21 Oct 2025

Gatherer Ancestry Associated with National Happiness

PONE-D-25-39711R1

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Acceptance letter

Ran Barkai

PONE-D-25-39711R1

PLOS ONE

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Associated Data

    This section collects any data citations, data availability statements, or supplementary materials included in this article.

    Supplementary Materials

    S1 File. Supporting Information.

    Tables S3-S5 are included in the Supporting Information file.

    (DOCX)

    pone.0336161.s001.docx (31.2KB, docx)
    Attachment

    Submitted filename: Response to Reviewers.pdf

    pone.0336161.s003.pdf (294.7KB, pdf)

    Data Availability Statement

    All data are publically available with sources references in the manuscript. In addition, the data underlying the results presented in the study and replication code are available from: https://github.com/mfbasilico/GatheringHappiness.


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