A worldwide view of matriliny: using
cross-cultural analyses to shed light on
royalsocietypublishing.org/journal/rstb
Research
Cite this article: Surowiec A, Snyder KT,
Creanza N. 2019 A worldwide view of
matriliny: using cross-cultural analyses to shed
light on human kinship systems. Phil.
Trans. R. Soc. B 374: 20180077.
Downloaded from https://royalsocietypublishing.org/ on 07 April 2023
http://dx.doi.org/10.1098/rstb.2018.0077
Accepted: 31 May 2019
One contribution of 17 to a theme issue ‘The
evolution of female-biased kinship in humans
and other mammals’
Subject Areas:
behaviour, computational biology, evolution
Keywords:
post-marital residence, evolution of behaviour,
cultural evolution, matrilineal descent
Author for correspondence:
Nicole Creanza
e-mail: nicole.creanza@vanderbilt.edu
Electronic supplementary material is available
online at https://doi.org/10.6084/m9.figshare.
c.4532573.
human kinship systems
Alexandra Surowiec, Kate T. Snyder and Nicole Creanza
Department of Biological Sciences, Vanderbilt University, Nashville, TN 37240, USA
NC, 0000-0001-8821-7383
Although matriliny and matrilocality are relatively rare in contemporary
human populations, these female-based descent and residence systems are
present in different cultural contexts and across the globe. Previous research
has generated numerous hypotheses about which cultural traits are associ-
ated with the stability or loss of matrilineal descent. In addition, several
studies have examined matrilineal descent with phylogenetic analyses; how-
ever, the use of language phylogenies has restricted these analyses to
comparisons within a single language family, often confined to a single con-
tinent. Cross-cultural comparisons are particularly informative when they
account for the relationships between widely distributed populations, as
opposed to treating each population as an independent sample or focusing
on a single region. Here, we study the evolution of descent systems on a
worldwide scale. First, we test for significant associations between matriliny
and numerous cultural traits that have been theoretically associated with its
stability or loss, such as subsistence strategy, animal domestication, mating
system, residence pattern, wealth transfer and property succession. In
addition, by combining genetic and linguistic information to build a
global supertree that includes 16 matrilineal populations, we also perform
phylogenetically controlled analyses to assess the patterns of correlated evol-
ution between descent and other traits: for example, does a change in
subsistence strategy generally predict a shift in the rules of descent, or do
these transitions happen independently? These analyses enable a worldwide
perspective on the pattern and process of the evolution of matriliny and
matrilocality.
This article is part of the theme issue ‘The evolution of female-biased
kinship in humans and other mammals’.
1. Introduction
In matrilineal systems, descent is traced along female lines [1]. Matrilineal kin-
ship organization occurs relatively infrequently in human populations [2,3],
and much research by anthropologists and ethnographers has discussed the
rarity and apparent instability of matriliny [2 –14]. Some of this work has
focused on whether matrilineal systems are inherently unstable from the male
perspective, since men in these cultures do not belong to the same kin group
as their biological children; instead, with matrilineal kinship organization, a
man is a member of his own mother’s kin group while his children are members
of their mother’s (his wife’s) kin group. Therefore, in matrilineal systems, men
are expected to invest resources in their sister’s children instead of their own,
which appears to violate the evolutionary predictions of inclusive fitness that
individuals will prefer to invest resources in their closest kin [6,9]. This has
been dubbed the ‘matrilineal puzzle’. With inclusive fitness and kinship
theory in mind, the evolutionary puzzle of matriliny could be resolved by
paternity uncertainty: a man might be uncertain that his wife’s children are
his own, but, given that he and his sister share a mother, he is certainly related
to his sister’s children [15], providing an explanation for inheritance from a
& 2019 The Author(s) Published by the Royal Society. All rights reserved.
 mother’s brother to his sister’s son. However, the level of
paternity uncertainty required for this explanation of matri-
liny appears to be much higher than is observed in real
human populations [16], although more recent theoretical
approaches have shown that mother’s brother–sister’s son
inheritance can also be stable in the context of high paternity
certainty if certain restrictive assumptions are relaxed [8,17].
Although matriliny is relatively rare across human popu-
lations, matrilineal populations are distributed around the
world (figure 1). Previous studies of matriliny have often
focused on one geographical region or a single language
family, and relatively few analyses have accounted for phylo-
genetic relationships between studied populations [14,19]. In a
key phylogenetically controlled study, Holden and Mace
assembled data on rules of descent (binarized as ‘matriliny’
versus ‘patriliny’) and on the presence or absence of cattle
across 68 Bantu- and Bantoid-speaking populations [14]. Using
a phylogeny of the Bantu language family to control for ancestral
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relationships between populations, this study was able to
demonstrate a directional evolutionary association in which
the spread of cattle domestication led previously matrilineal
populations to transition to patriliny, supporting the adage
that ‘the cow is the enemy of matriliny’ [12].
Anthropological research on matriliny and matrilocality
in different parts of the world has generally predicted that
female-based descent and residence patterns are becoming
increasingly rare [6,9,12,20]. Accompanying this general pre-
diction, however, has been a wide variety of proposed traits
that have been predicted to influence matrilineal descent,
either reinforcing it or destabilizing it [9]. In addition to
cattle as mentioned above [14], large domesticated animals,
in general, have been predicted to destabilize matriliny [12].
Other hypotheses hinge on subsistence strategy; for example,
horticulture has been predicted to be positively associated
with matriliny, whereas plough agriculture has been linked
to the loss of matriliny [12]. Still other authors predict that
the transition from matriliny to patriliny is associated with
factors related to inheritable resources, such as increased
wealth, property and inequality [2,7], but that matriliny can
be fostered by daughter-biased investment of resources
[12,13]. Related to the inclusive fitness interpretation of the
matrilineal puzzle, some authors predict that matriliny is
associated with cultural traits that would reduce paternity
certainty; for example, external warfare, which requires
men to be absent for long periods, is thought to promote
matrilocal residence [21,22], and high divorce rates are
hypothesized to promote or stabilize matriliny [10,11]. In
addition, populations with matrilocal or other female-based
post-marital residence systems are often also matrilineal; in
these cases, it has been hypothesized that matrilocality pre-
cedes and promotes the transition to matriliny [2–4]. We
summarize these hypotheses in table 1.
Previous evolutionary studies of descent and post-marital
residence have primarily been conducted within language
families [14,25–28], which limits their ability to draw
conclusions about the evolution of descent and residence
patterns on a larger scale, both because different language-
family trees cannot be intuitively merged and because
language groups demonstrate their own dynamics of
change [29]. To date, there has not been a global evolutionary
analysis to detect which cultural traits appear to maintain,
generate or destabilize matriliny. Here, we evaluate hypoth-
esized relationships between cultural traits and female-
based descent and residence patterns on a global scale. We 2
first assess these associations across 1291 populations in the
royalsocietypublishing.org/journal/rstb
Ethnographic Atlas; we then account for the non-indepen-
dence of these populations (Galton’s problem [30,31]) in
two ways. First, we repeat our analyses in the Standard
Cross-Cultural Sample [32], which is a subset of populations
whose cultures are relatively independent from one another.
Second, we construct a worldwide phylogeny that is a com-
posite of numerous genetic and linguistic phylogenies
merged with an existing supertree of human populations
[33] and use this phylogeny to perform analyses that account
for the ancestral relationships between populations [31]. With
these phylogenetically controlled analyses, we aim to assess
the likelihood that traits in the Ethnographic Atlas show cor-
Phil. Trans. R. Soc. B 374: 20180077
related evolution with matrilineal descent and matrilocal
residence patterns, enabling us to both test and generate
hypotheses about the evolutionary dynamics of kinship
system organization.
2. Methods
(a) Constructing the D-PLACE dataset
We gathered ethnographic data from D-PLACE [18], an online
resource that stores digitized data from the Ethnographic Atlas
[34] (data downloaded from github.com/D-PLACE/dplace-
data), and compiled it into a matrix of 94 cultural categories
from 1291 populations. For 76 of these categories, we were able
to reclassify the cultural data into binary traits for analysis. For
example, from ‘Settlement patterns [EA030],’ we could encode
sedentary (coded as 1) or non-sedentary (coded as 0). In some
cases, we could not have coded a single binary trait for a given
cultural category without excluding a significant amount of
information from the analysis. In these cases, we created multiple
binary traits from one category in the Ethnographic Atlas. For
example, ‘Descent: major type [EA043]’ was binarized twice,
once according to the presence or absence of matrilineal descent
and then according to the presence or absence of patrilineal des-
cent (table 2). For 17 of the cultural categories in D-PLACE, there
was not an intuitive way to binarize the data, so those categories
were excluded from the analysis. Additionally, for the ‘Age
or occupational specialization: gathering [‘EA061]’ and ‘Age or
occupational specialization: agriculture [‘EA065]’ only one of
the populations exhibited age or occupational specialization, so
those categories were also excluded from the analysis. Data
were not available for all cultural traits for each population;
missing values for a population were coded as ‘NA’. We classi-
fied D-PLACE data from 76 categories into 126 binarized traits;
for a full list of these binarized traits, see electronic supplemen-
tary material, Dataset S1.
(b) Tests for association between descent/residence
systems and other cultural traits
We next determined whether cultural traits significantly
co-occurred with certain descent or residence systems more
than would be expected by chance, based on the frequency of
each trait across 1291 populations in the Ethnographic Atlas
[34]. We tested for significant associations between each of the
126 binary traits with each of our focal traits: ‘matrilineal des-
cent,’ ‘patrilineal descent,’ ‘matrilocal residence’ or ‘patrilocal
residence’. For each of these comparisons, we made a contin-
gency table (as in table 3). If all expected counts for the 2  2
contingency table were five or more populations, we performed
a x2 test; otherwise, we performed a Fisher’s exact test since the

patrilineal
duolateral
matrilineal
bilateral
ambilineal
mixed
quasi-lineages
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Figure 1. Map of major descent types in worldwide populations, adapted from D-PLACE [18]. Out of 1291 populations in the Ethnographic Atlas, there are 160
matrilineal populations, 590 patrilineal populations, 362 bilateral populations, 52 duolateral populations, 50 populations with mixed descent, 48 ambilineal popu-
lations and 12 populations with quasi-lineages. 17 populations are missing data and are not included.
x2 test is unreliable with small expected numbers in any one
category.
We then conducted a similar set of tests, but only included
the subset of populations that are in the Standard Cross-
Cultural Sample; this sample of 186 populations was chosen in
an effort to survey populations that are as independent from
one another as possible [32]. For each set of tests, we performed
a Holm–Bonferroni correction for multiple hypothesis testing.
(c) Compiling a genetic dataset
We downloaded mtDNA sequences from the National Center for
Biotechnology Information (NCBI) database using the following
search terms for each population in D-PLACE: (1) ‘(Population)
mitochondrial genome’ and (2) ‘(Population) AND human[or-
ganism] AND mitochondrion[title] AND genome[title]’, where
‘(Population)’ was the name or alternative name of a human
population. Additional sequences were obtained from Kivisild
et al. [35], the Human Mitochondrial Genome Database [36]
and the 1000 Genomes Project [37]. The genetic data generally
included metadata with the population name and geographical
sampling location for each sequence; when this was absent
(such as when the population name in the NCBI search term
occurred in the title of the study but not in the sequence meta-
data) we checked the citation given in NCBI and its electronic
supplementary material to assign a population name and
sampling location to the sequence.
(d) Matching the genetic data to D-PLACE population
names
For each mitochondrial genome, we first determined whether the
population name given in NCBI was identical to a population in
D-PLACE; if so, we considered the mtDNA and the ethno-
graphic data to correspond to the same population. To validate
these cases, we checked whether the D-PLACE population
location and the mtDNA sampling location were from the
same geographical region. If populations could not be associated
in this way, we examined the languages spoken by each popu-
lation. D-PLACE provides a standardized language code (ISO
639-3) for most populations in the downloadable data tables,
and we had previously matched many of the sequenced
3
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Phil. Trans. R. Soc. B 374: 20180077
populations to their spoken language using the Ruhlen database
of phoneme inventories from Creanza et al. [38]. If a population
in D-PLACE had an identical ISO code to a sequenced popu-
lation, we matched them and checked the validity of the match
with the geographical locations of each population. For popu-
lations that were still unmatched, we checked alternative
population, language and dialect names given in Ethnologue
[39] for the sequenced populations to attempt to match an
alternative name to a the D-PLACE population. For populations
that were matched by population name and location, we also
verified the match with language information whenever
possible. In sum, we collected 4210 mitochondrial genome
sequences from a total of 193 populations, 165 of which were
successfully matched to populations listed in D-PLACE
(electronic supplementary material, Dataset S2).
(e) Making the supertree
In order to test our hypothesis using a global phylogeny of
D-PLACE populations, we built a supertree: a single tree
assembled from a set of smaller phylogenies. This method
enables us to combine trees that were constructed using different
data types. Our supertree was constructed by combining eight
continent-level mtDNA phylogenies, 12 linguistic phylogenies
from D-PLACE and a previously constructed supertree from
[33]. This previously published supertree, which has 186 popu-
lations and four non-human outgroups, was generated by
merging phylogenies constructed from several types of genetic
and linguistic data. We included in subsequent analyses those
populations from D-PLACE that we could associate with genetic
data, enabling us to estimate their genetic distance from one
another, and we incorporated both genetic and linguistic
phylogenies from these populations into the published supertree.
For each population included in this study, we acquired full
mitochondrial DNA sequences from at least one individual. Most
matrilineal populations that have been genotyped had only
mtDNA available, but mitochondrial DNA alone is not well
suited to building a large-scale phylogeny of populations;
mitochondrial phylogenies are generally constructed at the hap-
logroup level [35,40]. Accordingly, we assigned each of the
genotyped populations to a region based on the geographical
location of that population and the United Nations Geoscheme
to make smaller, continent-level mtDNA phylogenies within
 Downloaded from https://royalsocietypublishing.org/ on 07 April 2023

Table 1. Existing hypotheses linking matriliny to other cultural factors. The ‘SCCS p-value’ columns give the significance of the association between each trait and matriliny/matrilocality in the Standard Cross-Cultural Sample
(x2 analysis). The ‘BayesTraits p-value’ columns give the significance of the BayesTraits analysis of correlated evolution between each trait and matriliny/matrilocality, using the supertree. Bold red text indicates values that were
significant after Holm-Bonferroni correction. * indicates that only one economic specialization (metalworking) was statistically significant.

matriliny matrilocality

related D-PLACE category tested BayesTraits SCCS x 2 BayesTraits

hypothesis ref. here SCCS x 2 p-value p-value p-value p-value

domestic factors: marriage, family, residence

‘matrilineal descent groups have probably developed independently. . . on the basis of prior [2–4] matrilocal residence 6.1 3 10214 6.7 3 10210 n.a. n.a.
matrilocality’
the rule of exogamous marriage is difficult to reconcile with matriliny (matrilineal puzzle) [6] exogamy 0.05991 0.35762 0.00772 0.03189
matriliny is incompatible with a focus on nuclear family [7] nuclear family 0.80878 1 0.30207 0.03631
polygyny/polyandry associated with matrilineal inheritance [8] polygamy 0.99909 0.61088 0.15830 0.30649
communal breeding promotes matrilineal system [23] (no equivalent variable)
high frequencies of divorce are associated with matriliny [10,11] (no equivalent variable)
low paternity certainty promotes matriliny [2,12] (no equivalent variable)
economic factors: wealth transfer, inequality, economic specializations
matriliny can arise from daughter-biased investment by parents and/or grandparents [13] matrilineal inheritance of movable 6.9 3 1027 3.0 3 10215 1.2 3 1026 2.3 3 10210
property
matrilineal inheritable of real property 6.9 3 1028 4.4 3 10212 7.7 3 1027 7.3 3 1029
matriliny is vulnerable with increasing wealth [2] (no equivalent variable)
‘economic changes brought about by contact with Western industrial nations’ disrupt matrilineal [5] (no equivalent variable)
systems
economic differentiation (presumably within a population) is incompatible with matriliny [7] economic specializations none sig. none sig. 0.00022* none sig.
class 0.05130 0.74781 0.02660 0.00125
caste 0.00755 0.02065 0.03970 0.00096
matriliny is incompatible with inequality: ‘power, property and prestige’ [2] class 0.0513 0.74781 0.02660 0.00125
caste 0.00755 0.02065 0.03970 0.00096
slavery 0.53279 0.00012 0.14228 0.00377
warfare promotes matrilocal residence [21,22] (no equivalent variable)
(Continued.)
4

Phil. Trans. R. Soc. B 374: 20180077 royalsocietypublishing.org/journal/rstb

 the following regions: North-Central America (13 populations), 5
South America (12), Central-South Asia (17), Europe (31),

9.2 3 1025
2.1 3 1026

3.4 3 1026

1.9 3 1027
BayesTraits
North-East-Southeast Asia (50), West Asia (12), Africa (47) and

royalsocietypublishing.org/journal/rstb
0.00011
p-value

0.18108

0.00030
0.00016

0.01898
0.00045
0.02705
matrilocality Oceania (11). To construct these genetic trees, we aligned the
genetic data using MAFFT [41] and constructed a pairwise gen-
etic distance matrix with the Kimura 2-parameter distance
metric. The distance matrix of the individual sequences was
then averaged within populations, resulting in a population-
level pairwise distance matrix that quantified the average genetic
SCCS x 2

0.00279

0.00103

0.00024
p-value

0.00206
0.19944

0.06572
0.02041
0.00389

0.32645
0.01612
distance between the populations being studied. We then con-
structed continent-level neighbour-joining trees from these

1
distance matrices to merge with the published supertree, as
described below and in the electronic supplementary material.
We included data from 12 linguistic phylogenies available for
download on D-PLACE, which had been pruned to only societies
BayesTraits

Phil. Trans. R. Soc. B 374: 20180077

present in the D-PLACE database [42 –52]. Each linguistic phylo-
0.00043
p-value

0.00191
0.02905
0.81965
0.02038

0.53215
0.08807
0.00311
0.02418
0.08403
0.51263
geny used in the construction of the supertree included at least
matriliny

one population from our genetic dataset.

We added our additional genetic and linguistic phylogenies
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to the published supertree using TNT (Tree analysis using New

SCCS x p-value

Technology) [53] following the methods in [33], resulting in a

phylogeny with 290 human populations and four non-human
2

0.02110
0.10952

0.09594
0.22234
0.46334
0.06909
0.00311
0.02418
0.13718
0.75392 outgroups. Detailed methods are included in the electronic sup-
plementary material. Then, we trimmed this large phylogeny
1

(which included populations from the previously published

supertree not included in this study) to include only the 165
populations for which we had ethnographic and genetic data.
related D-PLACE category tested

Branch lengths were necessary for our evolutionary analyses,

but the supertree methods do not produce a tree with branch
large domesticated animals

lengths, so we used mtDNA genetic distance data to add

the presence of fishing

branch lengths to the supertree (electronic supplementary

extensive agriculture
intensive agriculture

cattle domestication

material). The result was a global phylogeny of the 165 popu-

plough agriculture

animal husbandry

leather working

lations that included branch lengths. Finally, we rescaled the

boat building
pastoralism

phylogeny to have a mean branch length of 0.1, based on the rec-

milking

ommendation in the documentation for the BayesTraits package

here

used in subsequent analyses, which states that this scaling pre-

vents the predicted evolutionary rates from ‘becoming small
[or] hard to estimate’ [54].
[14,24]
[12]

[19]
ref.

(f ) Testing for correlated evolution

We next tested for correlated evolution between binary traits
coded from the Ethnographic Atlas [34]. If two cultural traits
matrilineal societies are less likely to have ploughs or large domesticated animals, more likely to

are evolving independently from one another, then the tran-

sitions between the presence and absence of one trait should
not depend on the other trait. In contrast, if the evolution of
the two traits is correlated, then transitions in one trait are signifi-
cattle domestication destabilizes matriliny (the cow is the enemy of matriliny)

cantly dependent on the state of the other trait. For example,

increased reef density promotes matriliny (the fish is the friend of matriliny)

Holden and Mace found that Bantu populations with matriliny

were significantly more likely to transition to patriliny when
cattle were present compared to when they were absent [14].
To test this type of prediction on a worldwide scale, we used
the function ‘Discrete’ in the program ‘BayesTraits (V2)’ via the
subsistence factors: food sources, agriculture, domestication

R package ‘BayesTraitsWrapper V1’ (btw v. 1.0). We tested

each of the traits ‘matrilineal descent,’ ‘patrilineal descent,’
‘matrilocal residence’ or ‘patrilocal residence’ in combination
with each of 124 other cultural traits using each of the four
trees that contained at least 30 populations (global supertree,
and the Bantu, Austronesian and Indo-European language
trees) (4  124 comparisons for each of four trees). For each com-
bination of factors, we performed 20 runs of Discrete using the
maximum-likelihood method to generate independent and
dependent models of correlated evolution. We performed a like-
lihood ratio test using lrtest ( package: btw v1.0) between each
Table 1. (Continued.)

be horticultural

pair of models to obtain a p-value indicating whether or not

hypothesis

the dependent-evolution model fit the data substantially better

than the independent-evolution model. The reported p-values
are the medians of the p-values from these 20 runs. We obtained
the final values for transition rates between states by calculating
 Table 2. Binary descent and post-marital residence traits. The binary categories of descent as classified for our analyses.
D-PLACE Category from our
Ethnographic Atlas categorization
descent: major type [EA043] matrilineal
non-matrilineal
patrilineal
non-patrilineal
marital residence with kin: matrilocal
prevailing pattern [EA012]
non-matrilocal
patrilocal
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non-patrilocal
Table 3. Contingency table of female-based descent and female-based
post-marital residence patterns. Here, the matrilocal category also includes
other ethnographic classifications indicating female-based residence
patterns, such as avunculocal and uxorilocal residence.
matrilineal non-matrilineal
matrilocal 110 88
non-matrilocal 47 1008
total 157 1096
the means and 95% confidence intervals of the computed rates
from the 20 runs of the dependent model for each trait combi-
nation. We performed a Holm– Bonferroni correction for
multiple hypothesis testing to determine the threshold for signifi-
cance for each test. Two traits (male-biased house construction
and female-biased house construction) did not converge in
BayesTraits and were omitted from these results.
3. Results
(a) Tests of cultural trait association
It has been hypothesized that the evolution of matrilineal des-
cent correlates with female-based post-marital residence;
specifically, that matrilocal residence facilitates the transition
to matrilineal descent in a population [2 –4]. In line with this
hypothesis, we found that matrilineal descent often co-
occurred with matrilocal residence in D-PLACE populations,
with 70% of matrilineal populations also having matrilocal
residence, even though both characteristics are relatively rare
across human populations (table 3). We tested this association
with a x2 test and found that it would be extremely unlikely for
this degree of co-occurrence between matriliny and matrilocal-
ity to occur by chance ( p-value 2.8  10287).
We repeated this process for each of the 126 binary cul-
tural traits that we coded from D-PLACE to determine
whether they were significantly associated with matrilineal
descent, patrilineal descent, matrilocal residence or patrilocal
residence (electronic supplementary material, table S1 and
6
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D-PLACE type
matrilineal
patrilineal, bilateral, duolateral, quasi-lineages, ambilineal, mixed
patrilineal
matrilineal, bilateral, duolateral, quasi-lineages, ambilineal, mixed
matrilocal, avunculocal, uxorilocal, ambilocal with marked preponderance of uxorilocal
practice
patrilocal, virilocal, ambilocal with marked preponderance of virilocal practice, ambilocal,
Phil. Trans. R. Soc. B 374: 20180077
optionally uxorilocal or avunculocal, optionally patrilocal (or virilocal) or avunculocal,
neolocal, non-establishment of common household
patrilocal, virilocal, ambilocal with marked preponderance of virilocal practice
matrilocal, avunculocal, uxorilocal, ambilocal with marked preponderance of uxorilocal
practice, ambilocal, optionally uxorilocal or avunculocal, optionally patrilocal
(or virilocal) or avunculocal, neolocal, non-establishment of common household
see table S2 for full results). We conducted these x2 tests on
both the 1291 populations in the Ethnographic Atlas and, to
correct for Galton’s problem, on the Standard Cross-Cultural
Sample of 186 populations (table 1; electronic supplementary
material, tables S1 and S2). Broadly speaking, we found that
total matrilineal descent was positively associated primarily with
other female-based aspects of the kinship system (matrilineal
198 inheritance of real and movable property, matrilocal resi-
1055 dence and female-biased hereditary political succession)
and negatively associated with their male-based counterparts
1253
(table 1; electronic supplementary material, table S1). In
addition, when we considered the full Ethnographic Atlas
dataset (1291 populations), matrilineal descent was positively
associated with extensive agriculture and female-biased
participation in agriculture, and negatively associated with
cows, large domestic animals and milking. However, these
associations were not significant for the 186 populations in
the Standard Cross-Cultural Sample (table 1; electronic
supplementary material, table S1).
We conducted a similar analysis to identify cultural traits
that were significantly associated with matrilocal residence.
The same cultural traits that were associated with matriliny
in the Standard Cross-Cultural Sample were also associated
with matrilocal residence: matrilocal residence was signifi-
cantly positively associated with several female-biased
patterns (e.g. inheritance of real/movable property and her-
editary political succession) and negatively associated with
their male-biased counterparts (table 1; electronic supplemen-
tary material, table S1). In addition, matrilocal residence was
negatively associated with several subsistence-related traits,
such as intensive agriculture, plough cultivation, milking
and specialized metal working.
(b) Tests for correlated evolution
In addition, we used phylogenetically controlled analyses to
shed new light on existing hypotheses about the cultural fac-
tors that have been proposed to either foster or destabilize
female-based descent and residence patterns (table 1). For
these analyses, we used four phylogenies—the global super-
tree and Bantu, Austronesian and Indo-European language
 trees (figure 2)—and we tested whether each of the binarized
cultural traits from D-PLACE showed correlated evolution
with matrilineal descent, patrilineal descent, matrilocal resi-
dence or patrilocal residence (electronic supplementary
material, tables S1, S3, S4 and figures S1 –S7). These tests
allowed us to assess whether these kinship and residence
traits were evolving independently from other cultural traits
or whether the transition rates of one were significantly
dependent on the state of the other trait (see electronic sup-
plementary material, table S4 for full results). For example,
under the prediction that matrilocality facilitates the emer-
gence of matriliny, we might expect to see an elevated rate
of transition from no matrilineal descent to matrilineal
descent in the presence of matrilocal residence.
Broadly, we found support for the hypothesis that matri-
liny and matrilocal residence show correlated evolution,
corroborating the results of the association test (tables 1 and
3; electronic supplementary material, table S1). However,
Downloaded from https://royalsocietypublishing.org/ on 07 April 2023
our global phylogenetic analysis suggests a different direc-
tionality from previously hypothesized. Murdock suggested
that populations that were already matrilocal would be
more likely to develop matrilineal descent systems [2,3].
Contrary to this prediction, our results indicated that
matrilocal populations have a very low transition rate from
non-matrilineal to matrilineal systems (figure 3a). Instead,
it appears more likely that populations transition first to
matriliny and then subsequently to matrilocality.
Daughter-biased investment has also been hypothesized to
foster matriliny [13]. While we did not have a cultural variable
to directly test the daughter-biased investment hypothesis, we
examined maternal inheritance of both real and movable
property as a proxy. We found that both of these inheritance
patterns form a stable state in combination with matriliny:
matrilineal descent and matrilineal inheritance were stable
together in the sense that there were low rates of evolutionary
transitions away from this combination, but one trait without
the other was not stable (figure 3b; electronic supplementary
material, table S3). In addition, corroborating the results of
the association tests above, we found that both matrilineal
descent and matrilocal residence showed correlated
evolution with other sex-biased aspects of the kinship
system (matrilineal/patrilineal inheritance of real and movable
property, matrilocal/patrilocal residence and female/male-
biased hereditary political succession; figure 3c and table 1;
electronic supplementary material, tables S1, S3 and S4).
Other cultural aspects of family structure have been puta-
tively associated with female-based descent and residence:
polygamy is hypothesized to associate with matrilocality
and matrilineal inheritance, exogamy is thought to destabi-
lize matriliny and nuclear family structure is hypothesized
to be incompatible with matriliny [6–8]. We did not find sig-
nificant evidence for correlated evolution between matriliny
and any of these factors in the global analysis, although we
found support for correlated evolution between matriliny
and the presence of both matrilineal and patrilineal exoga-
mous groups (electronic supplementary material, tables S1,
S3 and S4). In particular, matrilineal descent is stable in com-
bination with matrilineal exogamous groups but not stable in
combination with patrilineal exogamous groups.
Various forms of economic differentiation have also been
hypothesized to be incompatible with matriliny [2,5,7].
Although in the x2 analysis we found that matrilocal residence
and metalworking co-occurred less often than would be
expected by chance in the Standard Cross-Cultural Sample, 7
we did not find evidence for correlated evolution between matri-
royalsocietypublishing.org/journal/rstb
liny and any sex-biased economic specializations, including
metalworking (table 1; electronic supplementary material,
table S1). Matriliny is also hypothesized to be incompatible
with economic inequality—’power, property and prestige’ [2].
We tested several traits related to economic inequality and
observed evidence for correlated evolution between matriliny
and the existence of slavery in a population, with slavery increas-
ing the rates of transition between modes of descent. However,
we did not find correlated evolution between matriliny and caste
or class systems across global populations. Interestingly, class
systems showed significant correlated evolution
with matriliny and matrilocal residence only for the Bantu popu-
Phil. Trans. R. Soc. B 374: 20180077
lations. In these populations, class systems were evolutionarily
unstable in combination with both matriliny and matrilocal resi-
dence, with populations quickly transitioning away from class
systems or away from female-based descent or residence.
Various aspects of subsistence strategy have been frequently
hypothesized to associate with matriliny; in particular, the
intensification of agricultural practices and the domestication
of cows and other large animals have been predicted to destabi-
lize matriliny and promote the transition to patriliny [12,14,24].
In support of this hypothesis, we found significant evidence of
correlated evolution between matriliny and extensive agricul-
ture—the practice of farming over a large land area with
relatively small labour investment and low yields (figure 3d).
This type of agriculture system appeared to stabilize matriliny,
as matriliny was lost more rapidly when extensive agriculture
was absent. However, across numerous tests, we found more
statistical support for this suite of hypotheses when considering
matrilocal residence instead of matrilineal descent. For example,
matrilocal residence was more stable in the presence of female-
biased participation in agricultural practices (electronic sup-
plementary material, tables S3 and S4), and matrilocal
residence was never gained in the presence of intensive agricul-
ture (figure 4). In addition, the combination of matrilocal
residence with plough agriculture was not stable, and popu-
lations with this combination appeared to quickly lose either
ploughs or matrilocality. Moreover, populations almost never
transitioned to matrilocality once they acquired plough agricul-
ture (figure 4). We found a similar pattern for other agricultural
traits: matrilocal residence was not stable in combination with
large domestic animals, animal husbandry or milking
(figure 4 and table 1; electronic supplementary material,
tables S3 and S4). However, matrilocal residence was also
gained and lost rapidly without these associated traits, so the
absence of agricultural and domestication practices did not
fully stabilize post-marital matrilocal residence (figure 4).
The spread of cattle specifically has been hypothesized to
destabilize matriliny in Bantu populations [14,24]. We did not
observe correlated evolution between cows and matriliny on
any scale. We did, however, observe significant correlated
evolution between cows and matrilocal residence, but only
in the Bantu lineage; a matrilocal society was more likely to
adopt cows than a non-matrilocal one, but then matrilocal
residence was quickly lost in the presence of cows (electronic
supplementary material, tables S1 and S4).
4. Discussion
The long-term dynamics of matrilineal descent and matrilocal
residence, including the factors contributing to their stability
 Tswana
global supertree Ambo Bantu linguistic tree 8
Shona Kom
Turkana Widikum
Nama Bamum
Tsonga

royalsocietypublishing.org/journal/rstb
Sotho Bali Nyonga
Naro Bamileke
Ju/’hoan North Tiv
Luhya Bafia
Chewa Fang
Bisa Kundu
Mende
Lozi Kpe
Plateau Tonga Puku
Masa Duala
Lala
Yoruba Sanga
Bemba Ngumba
Zulu Kota
Xhosa Mpongwe
Bergdama
Herero Mitsogho
Ndebele Ndaka Babali
Kikuyu Plains Bira
matrilineal Ila
Luvale Bira
Chokwe Mbuti
Mossi Kumu
non-matrilineal Mambwe Ngombe
Masai
Djafun Budja
Afar Ngala
Yao Kuba
Gurage Ekonda
Amhara
Wolof Mongo
Tuareg Tetela
Algerians Dzing

Phil. Trans. R. Soc. B 374: 20180077

Egyptian Songo
Gibe
Bedouin Yanzi
Austronesian linguistic tree Druze
Armenian
Bunda
Sakata
Syrian
Marshallese Abkhazian Yombe
Bikinians Lebanese Suku
Ponapeans Kurd Yaka
Nomoians Turkish Bakongo
Latvian
Downloaded from https://royalsocietypublishing.org/ on 07 April 2023

Carolinians Sundi
Portuguese Pende
Trukese Greek
Lamotrek
Kumyk Mbala
French Canadian Luwa
Ifaluk Romanian Ndembu
Kosraeans Kalmyk
Czech Herero
Makin Spanish Mbundu
Rotumans Basque Luvale
Tikopia Bulgarian
Dutch Chokwe
Ontong Java North Ossetian Luba
Kapingamarangi Russian Songye
Tokelau Saami Kaonde
Ellice Byelorussians
Ukranian Garanganze
Raroians Chuvash Songola
Mangaians Hungarian Buye
Tongarevans Kazan tatar Tonga
Tahitians Estonian
Irish Lamba
Hawaiians Chechen Lala
Easter Islanders Lithuanian Bemba
Mangarevans Adygei Shila
Iranian
Marquesans Yusufzai Gusii
Futunans Sinhalese Toro
Rennell Islanders Balochi Nyoro
Sherpa
Niueans Punjabi Haya
Tongans Bengali Zinza
American Samoans Khasi Kerewe
Mbau Fijians Sindhi Vugusu
Tamil
Lifu Uzbek Gisu
Tannese Turkmen Soga
Eromangans Kazakh Ganda
Burusho
Seniang Hazara Rundi
Mota Udihe Ha
Ulawans Nganasan Bashi
Lau Cham Safwa
Tagbanua
Kwaio Moken Nyakyusa
Simboese Atayal Fipa
Choiseulese Balinese Pimbwe
Ami Turu
Kombe Puyuma
Aua Paiwan Sukuma
Lakalai Vietnamese Sumbwa
Burmese Bende
Manam Sugbuanon
Wogeo Merina Pokomo
Mekeo Malay Digo
Javanese Giriama
Motu Ifugao
Trobriands Thai Chagga
Dahuni Igorot Meru
Toraja Kikuyu
Bwaidoga Bunun
Dobuans Minangkabau Kamba
Molima Semang Sonjo
Waropen Kwaio Pare
Tikopia Ngulu
Midobi Onotong Java Zigula
Rotinese Nauru
Ambonese Simbo Luguru
Kei Hawaiian Kwere
Lakalai Kaguru
Tanimbarese Maori Gogo
Ili−Mandiri American Samoans
Tongans Rangi
Kodi Ellice Sangu
Sumbanese Yakut Bena
Palauans Mansi
Mongola Hehe
Chamorro Hezhen Ngoni
Toradja Negidal Nyasa
Macassarese Tu Makonde
Toba Batak Selkup
Japanese Yao
Malays Korean Shona
Minangkabau Buryat Tsonga
Iban Miao Ndebele
Javanese Khmer
Evenk Zulu
Cham Ket Xhosa
Moken Yukaghir Tswana
Merina Aleut
Shuswap Lozi
Badjau Tawi−Tawi Tlingit Venda
Kalinga Nivkh Makua
Ifugao Koryak
Chukchi Kunda
Sagada Tlicho Chewa
Tao Eastern Ojibwa Tumbuka
Bilaan Tsimshian
Yaqui
Subanun Mapuche
Manobo Pima
Sugbuanon Warao
Bisayan Yucatec Maya
Aymara
Tagbanua Wayuu
Hanunoo Embera
Paiwan Brazilian
Guna
Bunun Cherokee
Atayal Mixtec
Puyuma Zapotec
Ami Wichi
Guarani
Yekuana

Figure 2. The evolution of matrilineal and non-matrilineal lineages across multiple phylogenies. The tips of each tree show whether populations were coded as
matrilineal (green) or non-matrilineal (white). At each node, the probability that the common ancestor was matrilineal is represented by the fraction of the bar that
is green; a white bar indicates a small probability that the common ancestor at that node was matrilineal. The Indo-European tree does not include any matrilineal
populations and is not shown. For a discussion of the estimated ancestral states, see the electronic supplementary material.
 9
(a) matrilineal descent and matrilocal (b) matrilineal descent and matrilineal
residence; p < 1 × 10–6 inheritance (real); p < 1 × 10–6

royalsocietypublishing.org/journal/rstb
no matrilineal 0.59 no matrilineal no matrilineal 0.25 no matrilineal
descent descent descent descent
no matrilocal matrilocal no matrilineal matrilineal
residence residence inheritance inheritance
20.13 35.64

189.49 7.16 11.54 0.33 46.34 1.09 4.54 21.75

matrilineal 187.44 matrilineal matrilineal 31.04 matrilineal

descent descent descent descent

Phil. Trans. R. Soc. B 374: 20180077

no matrilocal matrilocal no matrilineal matrilineal
residence residence inheritance 3.91 inheritance
59.63

(c) matrilineal descent and matrilineal (d) matrilineal descent and extensive
Downloaded from https://royalsocietypublishing.org/ on 07 April 2023

hereditary political succession; p < 1 × 10–6 agriculture; p = 4.3 × 10–4

no matrilineal 0 no matrilineal no matrilineal 0.91 no matrilineal
descent descent descent descent
no matrilineal matrilineal no extensive extensive
succession 0 succession agriculture 3.51 agriculture

26.14 1.34 1.06 2.12 32.48 2.1 4.18 1.45

matrilineal 12.11 matrilineal matrilineal 1.27 matrilineal

descent descent descent descent
no matrilineal matrilineal no extensive extensive
succession 4.15 succession agriculture 0 agriculture

Figure 3. The evolution of matrilineal descent is correlated with matrilocal residence, matrilineal inheritance and hereditary political succession, and extensive agri-
culture across a global supertree. We tested for correlated evolution between pairs of binary cultural traits, assessing whether the two traits were evolving independently
or whether the transition rates of one trait depended significantly on the state of the other trait. (a) Matrilineal descent was highly unstable without matrilocal resi-
dence. (b) Matrilineal descent was significantly correlated with matrilineal inheritance of real property, such that the presence of this trait was associated with an
increased rate of transition towards matrilineal descent. (c) Matrilineal populations without matrilineal hereditary political succession were unstable, and tended
to lose matrilineal descent. (d ) Matrilineal descent was lost more quickly in the absence of extensive agriculture. The predicted rate of transition between each
state is indicated by the numbers adjacent to each arrow, and the arrow thickness is scaled to the rate. Arrows with a rate of zero are indicated in black.

and loss, have been a focus of research since the late nine-
teenth century but only rarely studied in the context of the
evolutionary relationships between populations [2–15,22].
Here, we take a worldwide view of the evolution of matrili-
neal descent and matrilocal residence patterns across
globally distributed populations, and we study the inter-
actions between cultural traits while accounting for the
cultural non-independence of related populations. In particu-
lar, we test multiple hypotheses that have been proposed in
the literature that pinpoint cultural traits that might either
stabilize or destabilize patterns of matrilineal descent and
matrilocal residence (table 1).
Taken together, our results indicate that only a subset of
these hypotheses are supported by worldwide analyses to
detect cultural trait associations or correlated evolution. For
matriliny in particular, our significant findings pointed to
associations between matrilineal descent and other patterns of
cultural inheritance through the female line, such as female-
biased hereditary political succession, matrilocal residence
and matrilineal inheritance of real and movable property
(table 1; electronic supplementary material, table S1). Our
results lend support to the hypothesis that daughter-biased
investment (using a proxy of wealth transmission through the
maternal line) fosters matriliny [13], but we also note that matri-
liny might in turn also foster maternal inheritance of wealth
(figure 3). In addition, these quantitative results suggest that
female-biased cultural patterns of descent, inheritance and
residence might be parts of a suite of traits or a broader cultural
norm of female-based transmission rather than independently
evolving cultural traits. Our results also shed new light on
Murdock’s hypothesis that matriliny evolves on the basis of
prior matrilocality [2–4]: in contrast to Murdock’s prediction,
our evolutionary analysis indicates that it might not be uncom-
mon for matriliny to develop first, prompting a transition to
matrilocality (figure 3).
There is an intuition from previous literature that matriliny
is a potentially unstable form of kinship system organization;
interestingly, our results appear to support this hypothesis in
the sense that most of our observed associations were negative
(with the exceptions of the female-based inheritance patterns
mentioned above), with matriliny and matrilocality occurring
less often with other cultural traits than would be expected
 (a) (b) 10
matrilocal residence and matrilocal residence and
large domestic animals; p = 2 × 10–6 intensive agriculture; p = 1 × 10–5

royalsocietypublishing.org/journal/rstb
no matrilocal 1.35 no matrilocal no matrilocal 1.54 no matrilocal
residence residence residence residence
no large large domestic no intensive intensive
domestic animals animals agriculture agriculture
0.42 1.26

40.62 19.15 6.82 0.58 24.54 8.04 4.61 0

matrilocal 1.48 matrilocal matrilocal 1.6 matrilocal

residence residence residence residence

Phil. Trans. R. Soc. B 374: 20180077

no large large domestic no intensive intensive
domestic animals animals agriculture agriculture
5.18 6.94

(c) matrilocal residence and (d) matrilocal residence and

Downloaded from https://royalsocietypublishing.org/ on 07 April 2023

plough cultivation; p = 9.2 × 10–6 milking; p = 1.9 × 10–7

no matrilocal 0.65 no matrilocal no matrilocal 1.36 no matrilocal

residence residence residence residence
no plough plough
cultivation cultivation no milking milking
1.05 0

26.77 7.06 5.5 0.35 13.99 4.36 8.9 1.04

0 0.01
matrilocal matrilocal matrilocal matrilocal
residence residence residence residence
no plough plough no milking milking
cultivation 6.87 cultivation 24.07

Figure 4. The evolution of matrilocal residence is correlated with the presence of large domestic animals, intensive agriculture, plough cultivation and milking on a
global scale. As above, we tested for correlated evolution between pairs of binary cultural traits to determine if the rate of transition of one trait was significantly
dependent on the state of the other trait. The presence of large domestic animals (a), intensive agriculture (b), plough cultivation (c) and milking (d ) all reduced the
rate of transition between types of residence system. Once gained, these traits seemed to preclude the evolution of female-based systems. Arrows are labelled with
predicted transition rates (also indicated by thickness).

by chance (electronic supplementary material, table S1, blue
cells). In particular, a number of hypotheses have linked the
loss of matriliny to subsistence and economic factors, particu-
larly shifts from horticulture and relative economic equality
to intensifying agriculture, animal domestication and increas-
ing economic inequality [2,12]. In line with these hypotheses,
we found a stabilizing effect in which extensive agriculture
(as opposed to intensive agriculture) appeared to reduce the
rate of loss of matriliny (figure 3). However, we found no sig-
nificant association or correlated evolution between matriliny
and of the subsistence or economic factors hypothesized to
destabilize it [2,12,14,24]. Instead, we found significant associ-
ations between these factors and matrilocality: for example, the
presence of intensive agriculture, animal husbandry, milking
or plough agriculture in a population all appeared to be
unstable in combination with matrilocal residence. Many of
these same trends were similar for matriliny but were not stat-
istically significant; this might indicate that matrilocality has a
stronger pattern of being destabilized by intensifying agricul-
ture or animal domestication than matriliny does, or it could
simply suggest that we have more statistical power to detect
this phenomenon in our larger sample of matrilocal
populations.
Another related hypothesis predicts that the ‘cow is the
enemy of matriliny’ [14,24]. In testing this hypothesis, we
found a negative association between cows and matriliny,
and between cows and matrilocality, across the full dataset of
1291 populations. However, these associations were no longer
significant when we accounted for the non-independence of
populations, either by using the subset of populations in the
Standard Cross-Cultural Sample or by conducting phylogeneti-
cally controlled analyses (table 1, electronic supplementary
material, table S1). Earlier studies tested the specific hypothesis
that the domestication of cattle was correlated with a loss of
matriliny in Bantu populations [14,24]. We repeated these ana-
lyses with a Bantu language phylogeny that included more
populations (101 compared to 68 in [14]). In this Bantu subset
of populations, we found significant evidence for correlated
evolution between cows and matrilocality ( p ¼ 3.1  1025),
but the observed correlated evolution between cows and matri-
liny was not significant when we corrected for multiple
hypotheses ( p ¼ 0.00028). The significant association that we
observe between cows and post-marital residence patterns
could be an example of a lineage-specific evolutionary trend
[29] that is particularly salient in Bantu populations. Alterna-
tively, since we observe significant associations between
 matrilocality and other cultural traits that are related to domes- significant evidence of correlated evolution with other sex- 11
tication but not limited to cattle, such as animal husbandry, biased inheritance patterns and with traits related to intensive

large animal domestication and milking, on a worldwide
scale, it is possible that framing the hypothesis as the ‘cow is
the enemy of matriliny’ limits our ability to detect similar pro-
cesses that are occurring in societies that have domesticated
other large animals besides cattle.
There are several caveats to this and any cross-cultural
analysis. Any subset of populations is likely to be not fully
representative of the diversity of worldwide human popu-
lations. Our two approaches for addressing Galton’s
problem (that related populations are not independent
samples) make different sets of assumptions: using the Stan-
dard Cross-Cultural Sample makes the assumption that
royalsocietypublishing.org/journal/rstb
agriculture and animal domestication (electronic supplemen-
tary material, table S1). However, we found fewer significant
results with the Austronesian tree, with matrilineal descent
correlated with matrilineal inheritance of property, residence
and political succession but no economic- or subsistence-
related cultural traits. This result warrants further investigation
in the context of hypotheses that are tailored to this region; for
example, the ‘fish is the friend of matriliny’ hypothesis hinges
not just on the existence of fishing in a population but also on
reef density [19], which might correspond to the relative effort
investment required to catch fish. This nuanced view of fishing
is not captured in the Ethnographic Atlas, demonstrating a

Phil. Trans. R. Soc. B 374: 20180077

choosing populations from different regions produces a
subset of populations that are functionally independent
from one another, whereas conducting phylogenetically con-
trolled analyses of cultural traits makes the assumption that
Downloaded from https://royalsocietypublishing.org/ on 07 April 2023
limitation of this type of large-scale comparative analysis.
In sum, on a worldwide scale, we find support for several
of the previous hypotheses linking the evolution of female-
based descent to other cultural factors, and our global

those cultural traits are transmitted in a way that is well rep-
resented by the phylogeny (i.e. vertically transmitted). We
constructed a global phylogeny from several regional
language- and genetic-based phylogenies to maximize the
number of matrilineal populations that could be studied in
a phylogenetic context. However, the matrilineal populations
on this global phylogeny are spread out across the tree,
making it difficult to evaluate the evolutionary patterns that
might occur among closely related populations that show evi-
dence of recent gains and losses of matriliny, such as those
clades with multiple matrilineal and non-matrilineal popu-
lations seen on the Bantu and Austronesian trees (figure 2).
This underscores the need to genotype populations more
broadly, with cultural diversity in mind.
In addition, the results of our evolutionary analyses were
generally similar between the global and Bantu trees, with
matrilineal descent and matrilocal residence showing
analysis adds to the discussion of these hypotheses by
proposing the direction of these associations across many cul-
tures and by hypothesizing whether they apply to descent,
post-marital residence, or both. Taken together, our analyses
suggest that complex and nuanced factors contribute to the
evolution of matriliny and matrilocality, and both are best
studied in the rich cultural context of human populations.
Data accessibility. We include all data and code analysed in this article as
electronic supplementary material.
Authors’ contributions. N.C. conceived of the experiment. A.S. collected
data. A.S., K.T.S. and N.C. designed research, analysed data, made
figures and wrote the manuscript.
Competing interests. We declare we have no competing interests.
Funding. Funding was provided by Vanderbilt University to A.S.,
K.T.S. and N.C., and by the Vanderbilt University Summer Research
Program to A.S.

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