Hunter-Gatherers and the Origins of Religion
Hunter-Gatherers and the Origins of Religion
Hervey C. Peoples 0 1 2
Pavel Duda 0 1 2
Frank W. Marlowe 0 1 2
0 Department of Zoology, University of South Bohemia , České Budějovice , Czech Republic
1 P. O. Box 8850, Longboat Key, FL 34228 , USA
2 Department of Archaeology and Anthropology, University of Cambridge , Cambridge , UK
Recent studies of the evolution of religion have revealed the cognitive underpinnings of belief in supernatural agents, the role of ritual in promoting cooperation, and the contribution of morally punishing high gods to the growth and stabilization of human society. The universality of religion across human society points to a deep evolutionary past. However, specific traits of nascent religiosity, and the sequence in which they emerged, have remained unknown. Here we reconstruct the evolution of religious beliefs and behaviors in early modern humans using a global sample of hunter-gatherers and seven traits describing hunter-gatherer religiosity: animism, belief in an afterlife, shamanism, ancestor worship, high gods, and worship of ancestors or high gods who are active in human affairs. We reconstruct ancestral character states using a time-calibrated supertree based on published phylogenetic trees and linguistic classification and then test for correlated evolution between the characters and for the direction of cultural change. Results indicate that the oldest trait of religion, present in the most recent common ancestor of present-day hunter-gatherers, was animism, in agreement with long-standing beliefs about the fundamental role of this trait. Belief in an afterlife emerged, followed by shamanism and ancestor worship. Ancestor spirits or high gods who are active in human affairs were absent in early humans, suggesting a deep history for the egalitarian nature of hunter-gatherer societies. There is a significant positive relationship between most characters investigated, but the trait “high gods” stands apart, suggesting that belief in a single creator deity can emerge in a society regardless of other aspects of its religion.
Religion is unique to humans. Belief in supernatural agents and the entailed religious
practices occur in virtually all human cultures (Brown 1991; Johnson 2005; Murdock
1965; Murdock and White 1980). The universality of religion across human societies
(Brown 1991) suggests a deep evolutionary past. The human ability to create complex
culture (“cultural capacity”) may be older than the first split in the modern human
lineage (Lind et al. 2013), and some aspects of religion may have been present before
the appearance of anatomically modern humans (Rossano 2006). Religion has
generally been assumed to have emerged among anatomically modern humans in Africa
during the Upper Paleolithic, and to have played a vital role in the subsequent
out-ofAfrica expansion (Balme et al. 2009; Rossano 2009a).
Explanations for a natural emergence of religion have been debated for hundreds, if
not thousands, of years (Martin and Wiebe 2012; Wiebe 2008). That debate has now
turned to empirical approaches and testable hypotheses, many of them grounded in the
framework of evolutionary theory (Alcorta and Sosis 2005; Atran and Norenzayan
2004; Barrett and Lanman 2008; Boyer and Bergstrom 2008; Irons 2001; Rossano
2006; Tremlin 2006). During the past decade, evolutionary psychologists have
identified and described the activity of cognitive biases that enable us to accept the
counterintuitive concepts and beliefs of religion (Atran and Henrich 2010; Barrett and Lanman
2008; Tremlin 2006). Research into the dynamics of religion has revealed the nature of
ritual behavior to promote high levels of cooperation (Atran and Henrich 2010; Fischer
et al. 2013; Sosis and Ruffle 2003; Xygalatas et al. 2013). The contribution of belief in
morally punishing high gods to enhancing prosociality and ensuring growth and
stabilization of society has also been demonstrated (Norenzayan and Shariff 2008;
Norenzayan et al. 2016; Peoples and Marlowe 2012).
This research has often focused on characteristics of the large prosocial religions that
have emerged during the 10,000–12,000 years since the advent of agriculture
(Matthews 2012; Norenzayan 2013). Little attention has been paid to the religion of
hunter-gatherers whose religious beliefs and behaviors have been evolving during the
vast majority of human history (Lee and DeVore 1968). Despite established
speculations about various beliefs and behaviors that may represent an original form of
religion, specific traits of nascent religiosity, and the sequence in which they emerged,
have remained unknown.
There have been as many attempts to define religion as to explain its origins.
Broadly defined, religion is a set of beliefs and behaviors based on a shared worldview
that separates the sacred, or supernatural, from the profane (Durkheim 1965 ). In
this study of its origins, we view religion as a biocultural adaptation (Alcorta and Sosis
2005; Harris and McNamara 2008; Sanderson and Roberts 2008).
How old are religious concepts and their engendered behaviors? Ritual behavior is
widespread among humans today, operating in a variety of social environments, both
religious and secular (Brown 1991; Cronk 2005; Spencer 1870). Behavior reminiscent
of ritual can be seen in many animals, including the ecstatic “rain dances” and
directional drumming of chimpanzees (Goodall 1986; Nishida et al. 1999), which
along with bonobos are our closest extant evolutionary kin. It is highly likely that
archaic hominins would have exhibited ritualistic behavior in some form, but evidence
for nascent religiosity remains difficult to infer from the archaeological record (Rossano
Whether early hominins held religious beliefs prior to the emergence of language is
unknown. We should not dismiss the possible presence of non-linguistic religious
thought and sentiment among early members of the genus Homo. However, a case
can be made that transmission of religious concepts from one individual to another
requires complex mental imaging, and a capacity for symbolic thought and
communication that might include ritual, dancing, singing, gestures, art and ornamentation, as
well as language (Deacon and Cashman 2010; Mithen 1998). Some genetic and
anatomical changes enabling speech and language can be traced to the most recent
common ancestor of Neanderthals and anatomically modern humans about half a
million years ago (Barney et al. 2012; Dediu and Levinson 2013; Krause et al.
2007). It has been suggested that rudimentary language could be older than symbolic
thought (Barnard 2012).
The archaeological record documents the presence of artifacts and technology dating
from ca. 400 kya that would probably have required a level of symbolic communication
close to that of language (Zilhão 2007). Early evidence for the processing and use of
red ochre, often considered a marker for symbolic behavior, dates to the Middle
Pleistocene >400 kya in Africa (Barham 2002) and to >284 kya (Deino and
McBrearty 2002) in the presence of Levallois blade technology.
Finds at Pinnacle Point in southern Africa (Marean et al. 2007) demonstrate the use
and processing of pigment among anatomically modern humans as early as
165,000 years ago (McDougall et al. 2005). Ochre nodules bearing engraved abstract
patterns and perforated shell beads found at Blombos Cave in South Africa, dating to
75,000–100,000 years ago (d’Errico et al. 2005; Henshilwood et al. 2009), suggest
symbolically mediated behavior. These and other similar finds lend substantial support
to the theory of progressive development of symbolic behavior and complex imagery
along with the evolution of modern human anatomy (Barnard 2012; Conard 2010;
Deino and McBrearty 2002; d’Errico and Stringer 2011; McBrearty and Brooks 2000;
Zilhão 2007). Pleistocene hunter-gatherers would most likely have possessed both the
cognitive and communicative skills to share religious beliefs and practices prior to
dispersal out of Africa more than 60 kya (Fu et al. 2013; Henn et al. 2012; Johansson
2011; Lind et al. 2013). Although present-day hunter-gatherers are not direct analogues
of those early societies and may not be direct, unbroken descendants of ancestral
hunter-gatherers, they can provide a window onto traits selected for in the Pleistocene
(Marlowe 2005), including traits of early religion.
The uniqueness of “natural” religions of hunter-gatherers, and likely those of our
Paleolithic ancestors, cannot be overemphasized when compared with the “world”
religions that have emerged along with the advent of agriculture. Many
huntergatherer societies have little or no concept of religion per se, though a religious
dimension often permeates normal activities and is continuous with daily life (Lee
1989). Hunter-gatherer religions are seldom religions of protest or evangelism
(Woodburn 1997). Instead, each society focuses on maintaining its unique beliefs and
culture, along with a sense of self-worth and the general health and well-being of the
group (Woodburn 1997, 2005). Simple egalitarian hunter-gatherer groups generally
hold fewer religious beliefs and participate in less ritual (Marlowe 2010) than more
complex groups. But hunter-gatherers do have religion, embodied in sacred healing
dances and rituals marking life events. Although there is considerable variation in
specific religious traits among hunter-gatherer societies, a cross-cultural view reveals
underlying similarities in cosmology, ritual, and belief (Rossano 2007). These often
include gods and spirits with limited powers who are typically not omniscient and
usually lack concern for morality and human affairs (Marlowe 2010; Norenzayan et al.
2016; Peoples and Marlowe 2012; Swanson 1960; Woodburn 1997), as Marshall
The concept of sin as an offence against the gods is vague among the !Kung.
Man’s wrong-doing against man is not left to #Gao!na’s punishment nor is it
considered to be his concern. Man corrects or avenges such wrong-doings
himself in his social context. (Marshall 1962:245)
What were the specific traits of early religion? How did traits of nascent
religiosity evolve and interact over time? Recently phylogenetic comparative
methods have been increasingly applied to the study of the evolution of
material and non-material culture (Mace and Holden 2005; Mace and Jordan
2011), including religion (Matthews 2012; Watts et al. 2015). Reconstructing
ancestral character states on phylogenies based on genetic or linguistic data has
proven valuable in revealing the history of various sociocultural phenomena
(Currie et al. 2010; Opie et al. 2014; Walker et al. 2011, 2012). Although
religious beliefs are regarded as one of those cultural traits that are historically
labile and prone to cultural borrowing (Guglielmino et al. 1995), cross-cultural
research suggests that religion (and mythology) can be surprisingly stable
across time and space, and shared religious beliefs can be indicative of deep
ancestry (Berezkin 2008; Blust 2013). The use of phylogenetic methods is
important for understanding not only the origins of religious traits, but also
the behavioral systems that emerged from them that have determined patterns of
social constraint and have impacted believers and non-believers alike.
Importantly, homology is not limited to morphology and its genetic and/or
developmental underpinnings. Behavior, which is often evolutionarily conserved, is also a
proper subject of homology relations and can be used in phylogenetic reconstruction
(Hall 2013; Powell and Shea 2014; Rendall and Di Fiore 2007). A behavioral
homology need not have a particular structural basis. We do not argue for homology of the
particular religious beliefs (e.g., different afterlife beliefs across hunter-gatherer
societies that possess this trait) but for the homology of the fundamental religious concepts
(e.g., the concept of afterlife itself) and their continuity. Even characters that cannot be
hypothesized as strictly homologous among sampled cultures can be analyzed because
they can represent non-homologous psychological-behavioral responses to identical
selective pressures (see Murdock 1965 for similar reasoning in anthropology).
In this study we investigate early evolution of religion by reconstructing ancestral
states for seven characters describing religious beliefs and behaviors in a global sample
of 33 hunter-gatherer societies (Fig. 1). Using a time-calibrated supertree based on
published genetic and linguistic phylogenetic trees, and linguistic classification as a
proxy for population history, we reconstruct ancestral character states and test for
correlated evolution between the characters and for the direction of cultural change.
Fig. 1 Geographic locations of the 33 hunter-gatherer societies in the study sample
Phylogenetic reconstruction of ancestral character states is a two-step process. It
requires a set of characters (data matrix) with known character states (e.g., present or
absent) based on traits of interest, and a phylogenetic tree that represents the historical
relationships between the populations under investigation.
We used primary ethnographic sources to create a data matrix of characters describing
religiosity in a sample of 33 hunter-gatherer societies. The study sample consists of 28
hunter-gatherer societies selected from the Standard Cross-Cultural Sample (SCCS)
(Murdock and White 1980), and five additional hunter-gatherer societies selected from
both the Ethnographic Atlas (EA) (Murdock 1967) and original ethnographic
descriptions of the societies.
We used ratings of other researchers to obtain a subsample of 28 hunter-gatherer
societies from the SCCS. The variables used to define the sample were SCCS variable
1: Intercommunity Trade as Food Source; variable 3: Agriculture Contribution to Local
Food Supply; variable 5: Animal Husbandry Contribution to Food Supply (Murdock
and Morrow 1970); and variable 858: Subsistence Type–Ecological Classification
(coded by D. White after Paige and Paige 1981). We defined our sample of
huntergatherers as follows: contribution to local food supply less than 10% agriculture (SCCS
v3), less than 10% animal husbandry (SCCS v5), and trade accounting for less than
50% and no more than any single local source (SCCS v1). We excluded mounted
hunters (SCCS v858 = 5, mounted hunting) to more accurately represent
preagricultural hunter-gatherer societies. We excluded eight SCCS hunter-gatherer
societies from the sample either because of the lack of phylogenetic information (Ingalik,
Micmac, E. Pomo, Yokuts [Lake], Paiute North, Klamath, and Kutenai) or because
their main source of subsistence stated in the EA does not classify them as
huntergatherers (Shavante depend 16–25% on agriculture). We added five additional societies
chosen from the EA and original ethnographic sources in order to balance the
geographic distribution of the sample. These additional societies are the Aka and G/wi
(Africa), Agta (Philippines) and Walbiri (Australia) hunter-gatherers, and the Sandawe
(Africa), who are now mainly herders and farmers but share a deep genetic and
historical relationship with the Hadza (Tishkoff et al. 2007). Figure 1 shows the
geographic locations of the 33 hunter-gatherer societies in the study.
Trait Definition and Character Matrix Construction
Original coding of data in all 33 sample societies for the traits of animism (Tylor 1871),
belief in an afterlife (Bering 2006), shamanism (Eliade 1964; Winkelman 1990), and
ancestor worship (Sheils 1975; Spencer 1870; Steadman et al. 1996; Swanson1960)
was based on principal ethnographic source descriptions (White 1989) and additional
ethnologies referenced in the character matrix.
Tylor defined animism as a general belief in the “animation of all nature” (1871:258)
and fundamental to religion. Animism includes a “belief in personal souls” (1871:260)
as well as “a sense of spiritual beings. . . inhabiting trees and rocks and waterfalls”
(1871:260). We define animism as the belief that all “natural” things, such as plants,
animals, and even such phenomena as thunder, have intentionality (or a vital force) and
can have influence on human lives. Animism is coded as present or absent in each
society, based on assessments of principal ethnographers. Belief in an afterlife is
defined as belief in survival of the individual personality beyond death (Bering 2006)
and is coded as either present or absent.
A global definition of shamanism remains contentious (Sidky 2010). We define
shamanism as the presence in a society of a “shaman” (male or female), a socially
recognized part-time ritual intercessor, healer, and problem solver (Sidky 2010;
Winkelman 1990). Shamans often use their power over spirit helpers during
performances involving altered states of consciousness (Eliade 1964; Winkelman 2010) to
benefit individuals and the group as a whole (Eliade 1964; Winkelman 1990, 2010). We
view shamans as a general category of individuals often found in hunter-gatherer
societies who mediate between the earthly and spirit worlds to promote cohesion and
physical and mental well-being in the society (Eliade 1964; Sidky 2010; Winkelman
1990, 2010). Shamanism is coded as present or absent.
Ancestor worship is defined as belief that the spirits of dead kin remain active in
another realm where they may influence the living, and can be influenced by the living
(Sheils 1975; Spencer 1870; Steadman et al. 1996; Swanson 1960). Ancestor worship
is coded as four states in the SCCS: a belief in ancestor spirits can be (1) absent in a
society or the spirits can be (2) present but inactive in human affairs. In other societies,
ancestor spirits are believed to be active in human affairs but (3) may not or (4) may be
influenced by humans through prayer and sacrifice (Sheils 1975; Spencer 1870;
Steadman et al. 1996; Swanson 1960). “High gods” is EA variable 34 and SCCS
variable 238, defined by Swanson (1960) as single, all-powerful creator deities who
may be active in human affairs and supportive of human morality. The variable is coded
as four states. It differentiates between societies in which a creator deity is (1) absent,
(2) present but inactive in human affairs, (3) active in human affairs but does not
support a moral agenda, or (4) active and morally punishing. The SCCS provides
coding for high gods in 28 of the 33 societies in our sample. Original coding in the
additional five societies, based on principal ethnographic sources, completed the coding
for all 33 societies.
Based on the five traits of hunter-gatherer religion described above, we created a set
of seven characters of hunter-gatherer religiosity. The character matrix was coded as
follows: animism (absent, present); belief in an afterlife (absent, present); shamanism
(absent, present); ancestor worship (absent, present); high gods (absent, present). Two
additional traits, active ancestor worship (absent, present) and active high gods (absent,
present), were derived from the basic traits of ancestor worship and high gods. These
two additional characters recognize those societies that not only hold a belief in
ancestor spirits or high gods but also believe that those spirits or high gods are active
in human affairs. “Active ancestor worship” denotes the absence or presence of
ancestral spirits actively meddling in human affairs, who may or may not be influenced
through prayer or sacrifice (Sheils 1975; Spencer 1870; Steadman et al. 1996; Swanson
1960). Similarly, “active high gods” identifies societies that believe in a high god that is
also active in human affairs, and may or may not be morally punishing (Swanson
The resulting character matrix has 33 terminals (hunter-gatherer populations) and 7
characters (Fig. 2). (Character matrix and coding references are provided in the ESM,
Tables A1a and b.)
tre Belief in an Afterlife
lg Ancestor Worship (total)
teh High Gods (total)
tun Active Ancestor Worship
Number of societies with character (n=33)
Fig. 2 Distribution of the seven characters describing hunter-gatherer religiosity in the study sample
Phylogenetic Supertree Inference
We used a dataset described in Duda and Zrzavý (2016) to generate a phylogeny of 33
hunter-gatherer populations using the Matrix Representation with Parsimony
(hereinafter MRP) supertree method (Baum 1992; Ragan 1992). The dataset is based on 301
genetic and linguistic source trees from 199 studies published in journals, edited
volumes, and books from 1990 to ca. 2014. It also includes a character set based on
language classifications of Greenberg (1987), Ruhlen (1991), and Greenberg and
Ruhlen (2007) on the level of linguistic stocks and families.
Data from White (2009) and Ethnologue (Lewis et al. 2013) were used to match the
hunter-gatherer populations in the study sample with the populations present in the
Populations absent from or underrepresented in the supertree dataset were replaced
by either a more inclusive population group or by a genetically closely related
population that was used as a proxy for the hunter-gatherer population in question.
Positions of four North American populations (Kaska, Eyak, Twana, and Yurok) were
based solely on linguistic classification (ESM Table A2).
In order to overcome the problem of the lack of genetic data and the conflicting
signals caused by recent genetic admixture and language shifts in some hunter-gatherer
populations in the study sample, the characters based on linguistic classifications were
up-weighted by a factor of 100 to serve as a topological constraint or “scaffold.” This
scaffold tree constrains the topology for a subset of populations for which linguistic
affiliation can be determined (i.e., those scored for characters). (Details on supertree
dataset manipulations are given in the Methods section of the ESM.)
The linguistic scaffold tree included 20 phylogenetically informative characters for
the 33 populations in the study sample. Note that this linguistic scaffold implies
relatively few internal groupings (clades based on linguistic classification), particularly
among the Old World hunter-gatherers (see ESM Figure A1).
Semi-rooted coding (sensu Bininda-Emonds et al. 2005) was employed. The
supertree was rooted by an “all-0” hypothetical outgroup that preserves the rooting
information for rooted source trees; for unrooted source trees the hypothetical outgroup
was scored as “?”.
The complete dataset included 974 taxa + outgroup. The analysis was performed in
TNT 1.1 (Goloboff et al. 2008) under “new technology search” with search level 10
using sectorial, ratchet, and tree fusing searches, obtaining trees from a 10,000-replicate
random addition sequence, treating gaps as missing data and all character changes as
equal and non-additive. The recovered trees were then subjected to additional branch
swapping with up to 10,000 trees held during each step. The resulting supertree is a
semi-strict consensus tree reduced to 33 hunter-gatherer populations + outgroup. The
topology of this supertree is fully resolved (ESM Figure A2).
Time-Calibrating the Supertree
Time-calibrated branch lengths were obtained from published time estimates of
divergence events and colonization events in human population history based on molecular,
linguistic, and archaeological data (ESM Table A3b). These estimates were used as
time constraints on the nodal ages of the supertree. In order to test the robustness of
reconstructions of ancestral character states and given the considerable variance in
molecular-based time estimates of divergence dates and discrepancies between
estimates based on molecular, linguistic, and archaeological data, two sets of time
estimates were used. (Two sets of divergence dates for time-calibrating the supertree are
given in ESM Table A3a.)
The first set of dates assumes a greater time depth of the supertree and consists
mostly of molecular-based estimates of divergence dates. The second set of divergence
dates is based on molecular-based estimates, archaeological data, and glottochronology
and is close to the minimum time estimates. We emphasize the results based on the
second set of dates since these represent the time when the populations in our sample
last shared close cultural contact, which arguably suits our analyses better than the
estimates of deeper, molecular-based divergence dates between the populations in
The age of the nodes for which time estimates were available was fixed using the
Node Age Constraint tool in Mesquite 3.02 (Maddison and Maddison 2015). The
timecalibrated supertree was inferred using the combination of Enforce Minimum Node
Age Constraints and Arbitrarily Ultrametricize functions in Mesquite 3.02 (Maddison
and Maddison 2015).
Phylogenetic Reconstruction of Ancestral Character States
The set of characters was mapped onto the tree topology. All character states in the
outgroup were scored as 0, absent (i.e., plesiomorphic character states for all
characters). Maximum parsimony and maximum likelihood reconstruction of ancestral
character states were performed in Mesquite 3.02 (Maddison and Maddison 2015). The
Markov k-state 1 parameter model (Mk1) that assumes an equal rate of change between
all character states (Lewis 2001) was used for maximum likelihood reconstruction. An
asymmetric likelihood ratio test (Pagel 1999b) indicated that the asymmetric
twoparameter model does not offer a significant improvement over the Mk1 model for
any of the seven characters in question, thus validating the Mk1 model. Each character
was mapped onto a set of topologies using the Trace Character History function.
The statistical support for the ancestral state reconstructions was determined using a
likelihood decision threshold of T = 2, indicating support at least 7.4 times greater for
the character state in question than for the alternative character state(s) (Schluter et al.
Testing for Correlated Character Evolution
In order to test hypotheses about temporal ordering of character state changes and
coevolution of traits we used Pagel’s test for correlated discrete character evolution
(Pagel 1994, 1999a) implemented in the Pagel94 module in Mesquite 3.02 (Maddison
and Maddison 2015). This method uses a continuous-time Markov model to infer
character changes along each branch of a phylogenetic tree in order to establish the
most likely temporal ordering and direction of evolutionary change and the most
probable evolutionary pathway between two discrete binary characters. Evolutionary
change in each character along the tree branches is modelled as a Markov process, in
which the likelihood of character change is dependent on its current character state.
Two models are fitted: an independent, four-parameter model (Li) in which evolution in
each character is independent of the state of the other character, and a dependent,
eightparameter model (Ld) in which the probability of change in one trait is dependent on the
state of the other trait. (For example, the probability of a culture gaining shamanism can
differ between cultures with a belief in an afterlife and cultures without one.) A
likelihood ratio (LR) is used to compare the log likelihoods of the independent and
dependent models. The advantage of this method is that its use is not conditioned on the
ability to unequivocally reconstruct ancestral character states (Nunn 2011).
The supertree with the preferred set of divergence dates (i.e., shallower divergences)
was used for correlation analysis. The probability that a model of dependent evolution
fits the data significantly better than the model of independent evolution was estimated
with a likelihood-ratio test involving 1000 Monte Carlo simulations. A likelihood-ratio
test generates a null distribution of likelihood ratios, against which the significance of
the observed LR is tested. For each simulation, maximum-likelihood estimates of
model parameters were optimized using 500 iterations. If the dependent model fits
the data significantly better than the independent model, this indicates that the state of
one character affects the probability of change in the other, and that the two characters
The resulting supertree topology (see ESM Figure A2) indicates a deep split between
sub-Saharan African and non-African hunter-gatherers. Within Africa, South African
Khoisan who speak !Kung and G/wi are more closely related to Central African
Pygmies than to click-speaking East African Hadza and Sandawe. Outside Africa,
the Vedda of Sri Lanka is the deepest-rooting lineage, followed by Andaman Islanders
and Australian Aboriginals, presumably the remnants of early out-of-Africa expansion
via the “southern route” (Macaulay et al. 2005). A large clade follows, consisting of
two groups: hunter-gatherers of East Asia and those in Beringia and America. Within
East Asia, two sister groups appear: Southeastern (Negritos of Malaysia and Philippines
and Badjau Tawi) and Northeastern (the “Paleo-Asiatic” peoples). The
BeringianAmerican clade consists of related Eskimo-Aleut and Na-Dene speakers, and the
Amerind speakers of North and South America as a sister group of the two.
The reconstructions of ancestral states for religious beliefs and behaviors show
several consistent patterns using maximum parsimony and maximum likelihood
methods, and topologies with two alternative sets of divergence dates. The presence
of animistic concepts in the religions of all sample societies (Fig. 2) is in accord with
Tylor’s (1871) theory that animism is fundamental to religion. The presence of animism
in the last common ancestor (LCA) of present-day hunter-gatherers is significantly
supported (proportional likelihood = 0.99, p < 0.05*).1 Reconstructions of ancestral
states for the six remaining characters describing religion in our hunter-gatherer sample
are shown in Fig. 3. (Reconstructions of ancestral states for each node based on
maximum parsimony and maximum likelihood using two sets of divergence dates
are given in ESM Table A4.)
1 Asterisk (*) indicates significant result p ≤ 0.05
Maximum likelihood reconstructions of ancestral states for six characters describing hunter-gatherer
religiosity. a Belief in an Afterlife. b Shamanism. c Ancestor Worship. d Active Ancestor Worship. e High
Gods. f Active High Gods. The scale indicates time depth in kya. (see ESM Table A4 for details)
Belief in an afterlife and shamanism are present among 79% of sample societies (Fig. 2)
and have similar, although not identical, distribution across societies (Fig. 3a, b). These
characters are less common among African hunter-gatherers. The reconstructed ancestral
state in the deepest node for belief in an afterlife and shamanism is equivocal (proportional
likelihood = 0.5 and 0.56, respectively; ESM
Table A4a) according to maximum
likelihood. Maximum parsimony favors the absence of shamanism. We cannot determine
whether belief in an afterlife and shamanic practices were present in the LCA of
presentday hunter-gatherers. Among present-day African hunter-gatherers (the deepest-rooting
clades) the !Kung and G/wi hold a belief in an afterlife, whereas Hadza, Mbuti, and Aka
either never acquired it or had the trait and then lost it (Fig. 3a). The !Kung and G/wi have
shamans, but Hadza, Sandawe, Aka, and Mbuti do not (Fig. 3b). The presence of healers
among the Mbuti suggests that the Mbuti may once have had shamans, but they lost the
trait at some point (Winkelman 1990).
Ancestor worship is present in 45% and active ancestor worship in 24% of sample
societies (Fig. 2). The ancestral state of “ancestor worship” is equivocal (proportional
likelihood = 0.5; ESM Table A4b). Ancestral presence of active ancestor worship is
somewhat less likely. However, this is not significantly supported (proportional likelihood
= 0.43429948; ESM Table A4b) on the topology with shallower divergence dates. In
contrast, its ancestral absence is significantly supported (0.10739427*; ESM Table A4b)
on the topology with deeper divergence dates. Maximum parsimony reconstruction favors
the absence of active ancestor worship (ESM Table A4b). These results suggest ancestor
worship could have been present among ancestral hunter-gatherers, but probably not the
Only 39% of sample societies have the trait “high gods,” and even fewer (15%) have
active high gods (Fig. 2). The equivocal results based on maximum likelihood and
absence according to maximum parsimony for the ancestral state of “high gods”
suggests possible presence of belief in a single creator deity among ancestral
huntergatherers, albeit one that is not active in human affairs (proportional likelihood = 0.05*)
(see ESM Table A4b for details).
The consistency and retention indices (CI, RI) calculated for each character and the
whole character matrix quantifies the degree of character “fit” on the tree. CI (with values
from 0 to 1) measures the amount of homoplasy on a tree; RI (also 0–1) measures the
degree to which shared derived character states are exhibited on a tree. The resulting CI
and RI values for the whole character matrix are low (0.17 and 0.31, respectively),
indicating that most characters are highly labile. The characters displaying the worst fit
on phylogeny are ancestor worship and high gods (CI = 0.1), and the reconstructed
ancestral states for both are equivocal (Fig. 3c, e; ESM Table A4b).
The results of Pagel’s test for correlated evolution indicate a significant positive
relationship between most traits investigated. The dependent models for the evolution
of selected pairs of characters showing support for correlated evolution are shown in
Fig. 4. (The results of Pagel’s test for correlated evolution, including transition rates for
independent and dependent model of character change, log likelihood values for each
model, and p values, are given in ESM Table A5.)
Belief in an afterlife and shamanism emerge in the presence of the fundamental trait
of animism. Once these two traits are gained, they are unlikely to be lost (Fig. 4a).
Coevolution of belief in an afterlife and shamanism is significantly supported
(Li = −35.71, Ld = −28.75, LR = 6.96, p = 0.00*) (Fig. 4c). The transitional probabilities
indicate that belief in an afterlife evolves more likely in the absence of shamanism than
shamanism would evolve in the absence of a belief in an afterlife. This suggests that belief in
an afterlife is likely to have emerged first from the base of animistic beliefs, and later
shamanism evolved in the presence of belief in an afterlife (Fig. 4c). It also indicates that
shamanism is likely to be lost in the absence of belief in an afterlife (Fig. 4c; ESM Table A5).
Fig. 4 Transitions between character states for selected pairs of characters showing significantly higher
likelihood of the dependent model of evolution, indicating that these traits coevolve. Widths of arrows are
proportional to rates of change (see ESM Table A5 for details)
There is significant support for coevolution of belief in an afterlife and ancestor
worship (Li = −40.48, Ld = 34.66, LR = 5.82, p = 0.003*) (Fig. 4d). Belief in an
afterlife evolves prior to ancestor worship, and its presence stimulates the subsequent
evolution of ancestor worship. Ancestor worship is unlikely to be lost in the presence of
belief in an afterlife (Fig. 4d).
There is also significant support for coevolution of both shamanism and ancestor
worship (Li = −40.48, Ld = −36.60, LR = 3.88, p = 0.01*) and shamanism and active
ancestor worship (Li = −34.74, Ld = −28.72, LR = 6.02, 0.001*). Shamanism seems to
have a deep history and continuity, whereas ancestor worship, although it could have
evolved very early in the history of modern humans, is a highly labile trait (Fig. 3c).
Active ancestor worship probably appeared later (Fig. 3d). Ancestor worship without
shamanism seems to be an unstable cultural state that results either in a loss of
worshipful relationship with dead kin or in the appearance of the shaman. Ancestor
worship with shamanism, on the other hand, appears to be a stable cultural state, rarely
lost once achieved, and the same is seen for active ancestor worship with shamanism
(Fig. 4e, f).
There is no support for coevolution of any pair of characters that includes high gods
and active high gods, with the obvious exception of the two traits themselves
(Li = −33.58, Ld = −27.93, LR = 5.65, p = 0.001). Surprisingly, not even belief in an
afterlife shows any correlation with high gods. Belief in an afterlife evolved prior to
high gods, as is evident from reconstruction of ancestral states and the transition rates
based on Pagel’s test for correlated evolution. But these pairs of characters do not
coevolve: in other words, the probability of change in one is not affected by the state of
the other (see ESM Table A5 for details).
Our results reflect Tylor’s (1871) belief that animism was the earliest and most basic
trait of religion because it enables humans to think in terms of supernatural beings or
spirits. Animism is not a religion or philosophy, but a feature of human mentality, a
byproduct of cognitive processes that enable social intelligence, among other capabilities.
It is a widespread way of thinking among hunter-gatherers (Bird-David 1999; Charlton
2007; Klingensmith 1953; Piaget 1929). Animistic thought is a natural by-product of
the human capacity for intentionality or “theory of mind mechanism” (Dunbar 2003).
This innate cognitive trait allows us to attribute a vital force to animate and inanimate
elements in the environment (Piaget 1929; Tylor 1871). Once that vital force is
assumed, attribution of other human characteristics will follow. Animistic beliefs are
generally adaptive in the environments that prevail in hunter-gatherer societies
(BirdDavid 1999; Charlton 2002). Animistic thinking would have been present in early
hominins, certainly earlier than language (Coward 2015; Dunbar 2003).
It can be inferred from the analyses, or indeed from the universality of animism, that
the presence of animistic belief predates the emergence of belief in an afterlife. Once
animistic thought is prevalent in a society, interest in the whereabouts of spirits of the
dead could reasonably lead to the concept of an unseen realm where the individual
personality of the deceased lives on. The afterlife might be a rewarding
continuation of life on earth, or a realm of eternal punishment for those who
break social norms. Belief in an afterlife may have generated a sense of “being
watched” by the spirits of the dead, prompting archaic forms of social norms
(Bering 2006) actualized in the role of the shaman.
Shamanism significantly correlates with belief in an afterlife, which emerged first.
Shamanism then evolved in the presence of belief in a realm of spirits of the dead. If
belief in an afterlife is lost, shamanism is also likely to be lost. The single exception to
this in our sample is the Slave, who have shamanism without belief in an afterlife.
Although shamanism has been described as the universal religion of Paleolithic
huntergatherers (Eliade 1964; Winkelman 1990), it is not a religion per se, but a complex of
beliefs and behaviors that focus on communication with the ancestral spirits, as well as
the general world of spirits in the realm of the afterlife. Shamans are healers, ritual
leaders, and influential members of society whose keen insight and success in solving
social problems (Rossano 2007; Winkelman 1990) can lead to wealth, power, and
access to mates. Shamanism acts as a mechanism to reinforce social norms,
encouraging group cooperation through ritual and social bonding, and calming anxiety during
times of resource stress (Hayden 1987; Rossano 2007; Winkelman 1990; Winkelman
2010). Shamans, as Vitebsky (2000) puts it, are both spiritual leaders and social
workers. It would be reasonable to argue that shamans, who draw their power from
communication with the world of spirits, would have initially emerged in strongly
animistic societies that believed in an afterlife. Communication with omniscient and
perhaps judgmental spirits of known deceased, including ancestors, would have been a
useful tool in the work of the shaman.
As humans migrated out of Africa more than 60 kya (Henn et al. 2012; Macaulay
et al. 2005), the shaman’s curing skills and group rituals would have enhanced survival
through physical and emotional healing, enforcement of group norms, and resource
management. At the time of the rapid population dispersal of AMH out of Africa along
the southern route into Wallacea and Sahul, the physical stress of travel and encounters
with unfamiliar cultures in areas already occupied by other hominins would have
driven the need for use of both material and non-material culture, including religion,
to negotiate identities and relationships among and between groups (Coward 2015).
Evidence for more complex information exchange systems, planning depth and
authority, and increased symbolization appears in the archaeological record of Wallacea
and Australia prior to 40 kya (Balme et al. 2009). These suggestions are in line with the
elevated likelihood support of the ancestral presence of shamanism in the deepest
outof-Africa nodes (Fig. 2b). Our results support Rossano’s (2009a) hypothesis that
shamanism preceded and is more basic than ancestor worship, although the presence
of shamanism in the LCA of present-day hunter-gatherers is not supported.
Despite established speculation by Spencer (1870) and Tylor (1871) that universal
ancestor worship was the rudimentary beginning of religion, our analysis shows that
worship of dead kin is neither widespread among hunter-gatherers nor the oldest trait of
religion. Fewer than half of the societies in our sample believe that dead kin can
influence the living (Fig. 2). In many hunter-gatherer societies the concept of ancestor
spirits is absent, or present but they are inactive in human affairs (Sheils 1975; Swanson
1960). For example, among the! Kung there is a general fear of active spirits of “the
dead,” who are often the ghosts of recently deceased kin. But the concept of having a
worshipful relationship with their own ancestors is absent (Marshall 1962). Greater
likelihood of the presence of active ancestor worship has been linked to societies with
unilineal descent where important decisions are made by the kin group (Sheils 1975;
Swanson 1960). Ancestor worship is an important source of social control that
strengthens cohesion among kin and maintains lineal control of power and property
(Sheils 1975; Steadman et al. 1996; Swanson 1960), particularly in the more complex
hunter-gatherer societies. In contrast, immediate-return hunter-gatherer societies
(Woodburn 1982) seldom recognize dead ancestors who may intervene in their lives.
The social structure of these societies does not usually consist of strong kin ties, and
individuals do not depend on help from close kin, living or dead (Barnard and
The minimum requirement for veneration of dead ancestors is animism and belief in
the survival of the personal identity beyond death. In our analyses, ancestor worship is
significantly positively related with belief in an afterlife and shamanism. Belief in an
afterlife evolves prior to shamanism and ancestor worship. There is significant support
for coevolution of shamanism with ancestor worship and active ancestor worship.
Belief in an afterlife with shamanism appears to be a stable cultural state, rarely lost
once achieved. Ancestor worship is also less likely to be lost in the presence of belief in
an afterlife with shamanism.
This is not to say that the reduction of complexity of religious beliefs and behaviors
cannot occur in simple hunter-gatherers. The presence of belief in an afterlife and
shamanism is significantly supported in the LCA of Beringian-American as well as
North and South American hunter-gatherers (Fig. 3a,b; ESM Table A4), suggesting that
the absence of these traits among the Siriono in Bolivia and Botocudo in southern
Brazil is due to independent secondary losses. In the Siriono, this loss was probably
part of a substantial decrease in cultural complexity during the expansion of Tupi
language speakers across lowland South America (Walker et al. 2012).
It can be argued that those societies under higher resource stress, encountering
difficulties with resource extraction that demands cooperative effort, would benefit
most from the shaman’s skills (Hayden 1987; Rossano 2007; Winkelman 1990, 2010).
This is supported by the prevalence of shamanism among hunter-gatherer societies of
Eurasia, and corresponding support for the presence of shamanism among their
ancestors (Fig. 2b). In “Paleo-Asiatic” peoples (Ainu, Gilyak, and Yukaghir) and in
EskimoAleut peoples of the circumpolar region, the presence of shamanism combines with
active ancestor worship. The presence of both traits in the LCAs of these groups is
significantly supported (Fig. 2d). High gods were not the first supernatural entities to
monitor morality (Geertz 2014). The power and leadership of the shaman was often
based on reaffirming traditional social behavior that was presumed to have been carried
out by the ancestors, and still desired and monitored by punishing ancestral spirits, even
in those societies where spirits of dead kin were not considered a part of the religion
(Steadman and Palmer 2008).
Belief in high gods appears to be a rather “stand-alone” phenomenon in the evolution of
hunter-gatherer religion. Prior studies have shown that among the four modes of
subsistence (hunter-gatherers, pastoralists, horticulturalists, and agriculturalists) hunter-gatherers
are least likely to adopt morally punishing active high gods, if any high gods at all (Botero
et al. 2014; Norenzayan 2013; Peoples and Marlowe 2012; Swanson 1960). This pattern is
reflected in the distribution and the reconstructed evolution of high gods and active high
gods in our sample (Fig. 2; Fig. 3e, f). Early egalitarian hunter-gatherers would rarely have
acknowledged an active high god (Norenzayan 2013; Peoples and Marlowe 2012) and
would be the least likely to accept or benefit from the supernatural meddling and social
constraints of deities who would be seen as “high rulers” (Peoples and Marlowe 2012).
The leaders of complex hunter-gatherer societies whose subsistence relies on collective
effort should be more likely to benefit from the coercive power of a punishing high god.
Our analysis does not support the prevalence of either type of high god among ancestral
hunter-gatherers, and the evolution of high gods does not correlate with any of the other
traits of hunter-gatherer religion, including ancestor worship.
In a study by Guglielmino et al. (1995) analyzing the transmission pattern of cultural
traits in sub-Saharan Africa, high gods (Swanson 1960) was among a group of traits
(taboos, ritual mutilation, premarital norms, etc.) that were consistently the least
correlated with either language or ecology, suggesting they evolve rapidly and are
prone to cultural borrowing. On the other hand, according to a more recent study by
Currie and Mace (2014), high gods are among those cultural traits that evolve at
relatively slow rates in Bantu and Austronesian societies. Our results are consistent
with Guglielmino et al. (1995) since high gods (and ancestor worship) show the worst
fit on phylogeny (CI = 0.1) of all studied characters. This suggests that the presence of
high gods and some other traits related to religion and ritual are influenced by more
socioculturally oriented factors, and it lends support to the idea that these types of traits
may be more labile. Such traits would be more readily gained or lost as the adaptively
relevant sociopolitical environment changes (Irons 1998). To some extent this finding
may explain the independent pattern of emergence of high gods in our study.
Ancestral spirits and local gods with limited powers of supernatural monitoring may
have come relatively easily to the minds of early human hunter-gatherers. These types
of supernatural entities operate in a different realm from omniscient and powerful
creator gods (high gods), who have been shown to be related to a culture of some type
of control or decision-making structure (Peoples and Marlowe 2012; Radin 1937;
Swanson 1960). The absence of belief in active gods and spirits in the LCA of
present-day hunter-gatherers, according to the reconstructions, indicates a deep
evolutionary past for the egalitarian ethos of most simple hunter-gatherer societies, whose
small mobile populations of self-sufficient individuals make collective action problems
less of an issue. Those societies would be the least likely to accept or benefit from the
personal restraints of active ancestors or active high gods.
In this study we used a suite of phylogenetic comparative methods to investigate the
early evolution of religion. We reconstructed ancestral states for seven characters
describing religious beliefs and behaviors in a global sample of 33 hunter-gatherer
societies and tested for correlated evolution between these characters and for the
direction of cultural change.
Our results indicate that the oldest trait of religion, shared by the most recent
common ancestor of present-day hunter-gatherers, was animism. This supports
longstanding beliefs about the antiquity and fundamental role of this component of human
mentality, which enables people to attribute intent and lifelike qualities to inanimate
objects and would have prompted belief in beings or forces in an unseen realm of
spirits. Reconstructions are equivocal on whether or not the religion of the LCA of
present-day hunter-gatherers included belief in an afterlife, shamanism, ancestor
worship, and the concept of a single creator deity, or a high god. Belief in either ancestral
spirits or creator deities who remain active in human affairs was not present in ancestral
hunter-gatherer societies, according to the reconstructions. This may be indicative of a
deep past for the egalitarian nature of hunter-gatherer societies, to whom high gods
would appear to be rulers (Peoples and Marlowe 2012).
The majority of traits of religion we investigated exhibit a correlated pattern of
character change on phylogeny. The results suggest that belief in an afterlife,
shamanism, and ancestor worship evolve in concerted fashion as an integrated system of
beliefs and practices. However, neither high gods nor active high gods exhibit
correlated evolution with the rest of the religious traits, including ancestor worship, despite
Spencer’s and Tylor’s suggestions.
This is in line with a variety of evidence from other studies (Botero et al. 2014;
Norenzayan 2013; Peoples and Marlowe 2012; Radin 1937; Swanson 1960) suggesting
that if a society acquires belief in an omniscient and potentially morally punishing
creator deity, it does so regardless of other aspects of its religion but more as a reflection
of its social and political structure.
Acknowledgments We thank Toomas Kivisild and Nathaniel Dominy for critical discussion; Olaf
BinindaEmonds and Jan Zrzavý for discussion of the methods; and Pavel Flegontov for critical comments on an earlier
version of the manuscript. Pavel Duda was supported by the Grant Agency of the University of South
Bohemia (042/2013/P; 140/2013/P).
Open Access This article is distributed under the terms of the Creative Commons Attribution 4.0 International
License (http://creativecommons.org/licenses/by/4.0/), which permits unrestricted use, distribution, and
reproduction in any medium, provided you give appropriate credit to the original author(s) and the source, provide a
link to the Creative Commons license, and indicate if changes were made.
Hervey Peoples (http://humanquestion.com/) is a PhD student in the Division of Biological Anthropology,
Department of Archaeology and Anthropology at the University of Cambridge, UK. Her research interests
include the evolution of religious behavior and the behavioral ecology of contemporary religious beliefs. She
is the author of The Human Question: What People Believe about Evolution, Human Origins, and the
Beginning of Life (Red Lion, 2003)
Pavel Duda is a PhD student in the Department of Zoology, Faculty of Science, University of South Bohemia,
České Budĕjovice, Czech Republic. His research interests include behavioral evolution of primates, including
humans, cultural evolution, and human population history
Frank W. Marlowe is a retired lecturer in the Biological Anthropology Division, Department of Archaeology
and Anthropology at the University of Cambridge, UK. His research focuses on the behavioral ecology of
mating systems and cooperation, especially among foragers. He has worked with the Hadza since 1995 and is
the author of The Hadza: Hunter-Gatherers of Tanzania (University of California Press, 2010)
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