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Advances in Complex Systems

Vol. 15, Nos. 1 & 2 (2012) 1150006 (20 pages)
c World Scientic Publishing Company


DOI: 10.1142/S0219525911003372

THE EVOLUTION OF ETHNOLINGUISTIC DIVERSITY

THOMAS E. CURRIE,,, and RUTH MACE,

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Evolutionary

Ecology Research Group,

Department of Anthropology,
University College London,
London, United Kingdom

Evolutionary

Cognitive Science Research Centre,

University of Tokyo, Japan

AHRC

Centre for the Evolution of Cultural Diversity,

University College London,
London, United Kingdom
t.currie@ucl.ac.uk
Received 7 January 2011
Revised 24 February 2011
Published 6 January 2012

Humans divide themselves up into groups based on a shared cultural identity and common descent. Culturally inherited dierences in dress, language, and institutions are
often used as symbolic markers of the boundaries of these ethnic groups. Relatively little is known about the function of such ethnic groups, and why ethnic diversity is high
in some regions yet lower in others. In this paper, we demonstrate how investigating the
spatial distribution of ethnolinguistic groups can reveal the factors that aect the origin
and maintenance of human ethnic group diversity. Here we describe the use of a Geographic Information System to construct a large database that integrates information
about languages with a number of environmental, ecological, and ethnographic variables.
Using these data on the spatial distribution of ethnolinguistic groups, we employ a hierarchical linear modeling approach to test a variety of hypotheses concerning the function
of such groups. Despite revealing intriguing spatial patterns such as the latitudinal gradient in ethnolinguistic diversity, previous analyses suggested that the direct eects of
environmental variables on the distribution of ethnolinguistic groups were in fact quite
small. Here we show that the strength of the relationship between ethnolinguistic area
and environmental variables is stronger in societies whose primary mode of subsistence
is foraging. We then go on to demonstrate this same nding using the estimated native
distributions of ethnolinguistic group in the Americas and Australia. In particular, Net
Primary Productivity is shown to be a good predictor of the area covered by ethnolinguistic groups in foragers but not in agriculturalists. This provides support for the
idea that the factors aecting ethnic diversity have changed in a systematic way with
changes in subsistence strategies and social organization. We highlight future avenues
for spatially explicit investigations of the evolution of ethnic diversity, and suggest that
the evolutionary ecological approach adopted here may provide important insights into
processes aecting ethnic diversity in the modern world.
Keywords: Ethnic diversity; ethnolinguistic groups; cultural evolution; Niche Construction; political complexity.
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1. Introduction
Despite being a relatively homogenous species genetically, humans are extraordinarily adept at nding dierences between themselves and dividing themselves up
into separate ethnic groups [48]. While this tendency has created a wonderful array
of distinct cultures and ways of life, it has also led to some of the darker aspects of
human history as groups from one ethnic group have sought to subjugate or destroy
those from another. Understanding the function of ethnic groups, and what factors
cause ethnic diversity to be high in some regions yet lower in others is therefore
of vital importance. However, relatively little is known about these questions. Here
we demonstrate how examining the geographical distribution of ethnic groups can
help shed light on the factors that aect the origin and evolution of such groups
[39, 48, 17].
1.1. Identifying and classifying ethnic groups
While the terms ethnic and ethnicity are widely used today these concepts
only become widely used in the social sciences relatively recently (especially in
comparison with concepts such as race or class) [32, 21]. This rise is often
linked to the breakdown of colonialism in Africa and Asia in the 1960s, and gained
yet further prominence with the collapse of Communist regimes in the early 1990s
[29]. Despite the prominence of ethnicity, the term is often used in a variety of
contexts with dierent meanings, and is often not explicitly dened [12, 21]. Here
we dene ethnic groups as groups of individuals that share a common, self-ascribed
identity based on the belief in common descent, and/or a shared culture, distinct
in some respect from those of other groups. This belief is culturally inherited from
one generation to the next. Symbolic markers of these groups such as dierences in
dress, language, and institutions are themselves culturally inherited. It is important
to stress that what is important in terms of the ways individuals and groups interact
with one another is the belief in cultural uniqueness or common descent rather than
the objective reality of such beliefs. We can also dene ethnogenesisa as the process
by which ethnic groups are formed, while ethnopathosis refers to the extinction of
ethnic groups, which may involve either the physical death of all members of the
group, or the incorporation of at least some of them into another group.
In practice, identifying ethnic groups is not an easy task. Ethnic groups often
dene themselves in relation to other groups and dierent markers may be more or
less important in dierent contexts. However, this diculty should not dissuade us
a It should be noted that [40] employs a dierent denition of ethnogenesis, one that incorporates
the idea that the formation of human cultural groups always involves reticulate, or rhizotic process
rather being a purely splitting or cladistic process. However, this denition is rather confusing as
it privileges one particular mode of group formation. The terms ethnogenesis and ethnopathosis
as we and others (e.g. [32, 58]) employ them are more directly analogous to the terms speciation
and extinction which are used in the biological literature [15], and denote phenomena that could
potentially come about via a variety of processes.

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The Evolution of Ethnolinguistic Diversity

from attempting to identify ethnic groups in a systematic manner using a consistent

criterion in order to understand what factors aect their formation and maintenance. The relational nature of ethnic groups is something that is held in common
with concepts of biological species, and delimiting species boundaries is by no means
straightforward [15]. However, it would be hard to argue that species thinking
has not been of enormous value to biologists in cataloguing and explaining the massive variation seen in living organisms. Perhaps the most widespread ethnic marker
is dierences in spoken language, with the critical development period in which a
language can be learnt making it particularly suitable for this task [44, 53]. Here,
we examine the diversity of ethnic groups that are based on dierences in language.
This focus on language has a number of advantages. Language has received more
attention over a greater portion of the world, and for a longer time, than probably
any other cultural group marker. Consistent criteria, such as mutual intelligibility,
have been employed in order to identify and classify distinct varieties of speech,
and are open to testing and evaluation [20]. Importantly, the classicatory system
employed in this study, the Ethnologue [24], explicitly uses group identity in its
determination of distinct languages and does not rely solely on more narrow linguistic criteria. Again this has parallels with the identication of biological species
where the degree of genetic similarity by itself is not necessarily the most pertinent factor in delimiting species boundaries [15]. It should also be emphasized that
although multilingualism is undoubtedly a common feature in many societies here
we are interested only in the primary languages of groups, i.e. those associated
with group identity, and not secondary languages or widespread trade languages
such Tok Pisin. In this paper, we therefore use the term ethnolinguistic group.
It is also important to stress at what level we are discussing the diversity of such
groups, as ethnicity, and variation in general, is often a nested phenomenon [57], e.g.
a person born in London may describe themselves as a Londoner, English, British,
European, or Western depending on the situation. Figure 1 shows two dimensions
of ethnolinguistic diversity: (1) the level of individual ethnolinguistic groups, (2) the
higher taxonomic grouping of language family. In any particular region these two
dimensions of diversity may overlap (e.g. the island of New Guinea is highly diverse
both in the number of languages and number of language families (Fig. 1(d)),
while eastern China has relatively few languages, and all come from a single family (Fig. 1(a))), or they may be discordant (e.g. the Pacic Island of Vanuatu is
extraordinarily diverse in number of languages, yet all belong to the Austronesian
language family (Fig. 1(c)), while the middle east does not have many languages
yet the languages that are present belong to a variety of dierent language families
(Fig. 1(b)). It should also be pointed out that higher levels of language diversity
may or may not be relevant in terms of a groups own ethnic (or meta-ethnic)
identity, and often the classications are based on the work of linguists (e.g. Hindi
and German are both classied as Indo-European languages, yet few speakers of
either language would feel anity with those of the other language based on this).
The factors aecting diversity at these dierent levels may also be very dierent.
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T. E. Currie and R. Mace

Fig. 1. (Color online) The diversity of cultural groups can be assessed on a number of dierent
levels. The thick black lines delimit distinct ethnolinguistic groups, while separate colors within
each panel indicate the dierent, higher-level groupings of language families. Language family
classications are taken from the Ethnologue [24]. Note the scale bars (in km). In this paper, we
examine the diversity at the lower-level of distinct ethnolinguistic groups.

For example, researchers such as Jared Diamond, Peter Bellwood, and Colin Renfrew have argued that the distribution of certain language families is related to
the expansion of populations fuelled by some competitive advantage (most commonly agriculture) [18, 52, 10]. Under this hypothesis, regions that have relatively
few language families are those that have undergone such an expansion during the
last 10,000 years or so, while those that have larger numbers of language families
indicate longer periods of settlement with relatively less population replacement.
In this paper, we address what factors explain the diversity at the lower level of
individual ethnolinguistic groups, dened as those speaking distinct languages.
1.2. The global distribution of ethnolinguistic groups
Examining how ethnic groups are distributed in space may provide clues as to what
factors have been important in shaping the diversity of dierent regions. Figure 2
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Fig. 2. Ethnolinguistic diversity and genetic diversity exhibit dierent global patterns. The current distribution of ethnolinguistic groups does not
reect time since dispersal from Africa. Kernel density map of ethnolinguistic groups was created in ArcGIS (data sources: Old World [24], Americas
and Australia [42]). Genetic diversity data (autosomal short tandem repeats) were taken from [61]. Keys for Genetic Diversity and Ethnolinguistic
Density represent upper bound of each category, with Ethnolinguistic Density being in units of ln languages per km2 . Data on Ethnolinguistic diversity
for the scatter plot was calculated as the density of languages (number of languages divided by land area) within in a 500 km radius of each of the 49
focal societies for which genetic data was available (only regions of land contributed to this density calculation). The dotted regression line indicates
this relationship is not statistically signicant.

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T. E. Currie and R. Mace

shows the global distribution of ethnolinguistic diversity. The red regions represent the areas of highest diversity (in terms of number of groups per unit area),
while the yellow regions are those of lowest diversity. It can be seen that broadly
speaking there is a latitudinal gradient in ethnolinguistic diversity, with diversity
increasing as one moves from the poles towards the equator [17]. Interestingly, this
parallels well-known patterns of biological species diversity and biologists have long
been interested in explaining these patterns [60]. This non-random distribution of
groups suggests some ecological factors may be important in shaping ethnic diversity. Interestingly, this latitudinal pattern of ethnolinguistic diversity is very dierent to the global pattern of genetic diversity. Generally speaking, genetic diversity
of populations decreases as with increasing distance from Africa (see Fig. 2), most
likely due to serial founder eects following the migration of modern human populations out from their place of origin [50]. This suggests that dierent processes have
shaped these two dierent aspects of human diversity, and that the distribution of
ethnolinguistic groups cannot simply put down to the length of human occupation
of these areas (e.g. Vanuatu has a huge number of languages on a few small islands,
yet was rst inhabited only around 3,500 years ago).
The association between cultural group diversity and latitude or other environmental factors has led to a number of hypotheses concerning how this diversity is
determined. The topography hypothesis [55, 11, 23] argues that physical barriers,
such as mountain ranges, divide populations with ethnic groups forming in a manner
analogous to allopatric speciation. This idea reects quite a passive view of ethnic
group formation, implying that regions would be less diverse if such impediments
were not in place. However, while some regions such as the Highlands of New Guinea
and the Himalayas are both mountainous and ethnically diverse other regions are
either mountainous but not particularly diverse (e.g. the Altai mountains of Central
Asia), or diverse without such obvious physical barriers (e.g. Aboriginal northern
Australia). The ethnographic record also shows that people often maintain ethnic
or other culturally dened boundaries despite prolonged contact with other groups,
and the movement of individuals across these boundaries [3]. The environmental
productivity hypothesis can be traced back to Birdsells pioneering work on Australian aboriginal groups [8], and argues that distributions of ethnic groups are
related to the productivity of a region [17]. This argument is based on the idea that
membership of a group, while providing certain benets, such as more eective
defense or increased hunting capacity, also involves some costs, such as the potential for altruistic acts to go unreciprocated. Therefore the group should be no larger
than that needed to be meet subsistence and reproductive needs (ethnic groups are
usually endogamous and therefore need to be large enough to avoid the deleterious
eects of inbreeding). Birdsell argued that for Aboriginal tribal groups this number
was around 500 individuals. More productive regions will be able to support higher
population densities and therefore will contain more ethnic groups. In support of
this idea Birdsell found a strong correlation between the area covered by Aboriginal
tribal groups and amount of rainfall. The environmental risk hypothesis [30, 43],
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The Evolution of Ethnolinguistic Diversity

on the other hand, argues that the risk of failing to meet subsistence requirements
is the most important factor in determining the area over which a group is spread.
In more variable environments social networks need to spread over larger areas to
provide a buer against temporary shortfalls in production, making more environmentally risky regions lower in ethnic diversity. In an analysis using country-level
data, Nettle [43] found a negative relationship between the number of languages
and an estimate of environmental risk (mean growing season: the number of months
in a year in which the climate is conducive to growing crops).
While these spatial patterns and correlations might suggest that the direct
eects of environmental factors have had a strong inuence in shaping ethnolinguistic diversity previous studies have suered from a number of methodological
issues (see Methodological Considerations below). In a previous study [17], we
addressed these problems and found that in fact the direct eects of environmental
variables on the present-day distribution of ethnolinguistic groups were relatively
weak, and that in particular the productivity and topography hypotheses received
no support in our analysis. Instead, we found that the mobility of groups aected
the area over which they were spread (more mobile subsistence strategies such
as Pastoralism were associated with more widespread ethnolinguistic groups), and
that the degree of political complexity exhibited by a society was the biggest single
predictor of the area covered by their ethnolinguistic group, suggesting that regions
which are lower in ethnolinguistic diversity are those in which more complex societies have arisen and expanded, with those regions being generally away from the
tropics.
These results suggest that the factors aecting ethnogenesis and ethnopathosis
have altered over time. Currie and Mace [17] argued that the relationship between
the environment and ethnic group distributions may have been stronger prior to
the development of agriculture and increasingly complex forms of political organization over the last 12,000 years. While it is dicult to ascertain the distribution
of ethnic groups in the remote past [34, 59], this hypothesis can be tested by examining whether the relationship between environmental factors and ethnolinguistic
diversity vary systematically across societies with dierent subsistence strategies.
From a niche construction perspective [45], we would predict that societies that rely
primarily on foraging and pastoral (animal herding) subsistence strategies would be
more strongly aected by direct climatic and environmental factors than agriculturalists who can often modify their environments in ways that buer themselves
from such direct eects. For example, the construction of large-scale irrigation systems means that farmers can control the supply of water to their crops, and raise
productivity without having to rely on direct rainfall. To test these predictions, we
rst use the database of ethnolinguistic groups described in [17]. We then go on to
take advantage of a kind of natural experiment in history [19] by examining the
distribution of ethnolinguistic groups in the Americas and Australia at the time of
rst contact with European societies, a period when foraging societies were more
abundant.
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2. Methods and Results

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2.1. Methodological considerations

In our previous paper [17], we argued that there were several problems with existing studies that attempted to explain large-scale patterns of cultural group diversity. Firstly, many studies have only provided descriptions of patterns rather than
testing specic hypotheses [39, 13, 41], or have only assessed single environmental
predictors of a preferred hypothesis in isolation [43, 22, 55]. As many environmental
variables vary with latitude, many of these correlations are likely to be spurious.
Secondly, many studies were conducted at a coarse scale of analysis, usually at the
level of countries, which masks a lot of potentially informative variation. To illustrate, Nettle [43] used the relationship between mean growing season (MGS) and
number of languages per country to argue for the Ecological Risk hypotheses, while
Fincher and Thornhill [22] use the correlation between number of pathogens and
number of languages per country to argue that it is transmissible disease that is
the most important factor aecting ethnic diversity. Neither study attempts to control for other potentially important environmental variables. In fact if both MGS
and number of pathogens are entered in to a regression model with number of languages as the dependent variable (controlling for country area) then the relationship
between pathogens and number of languages is no longer statistically signicant.b
We include this here not as a conclusive test of either the ecological risk or the
pathogen hypothesis but merely to highlight the potential problems with existing
studies. In our previous paper [17] and here, we have addressed these issues by
taking ethnolinguistic groups themselves as the unit of analysis. We examine the
association between the area covered by these groups and a number of dierent
ecological or cultural variables in line with the predictions of dierent hypotheses,
such as those described above. Finally, the units of analysis (in our case individual ethnolinguistic groups) cannot be considered as independent for the purposes
of statistical analysis due their historical relatedness [38]. We therefore employ a
Hierarchical Linear Modeling (HLM) approach [51] that controls for this problem.
In these analyses, individual ethnolinguistic groups are nested within the language
family to which they belong, with language family included as a random eect.
2.2. Ethnolinguistic groups and subsistence strategy
in the old world
In this study, we use the global ethnolinguistic database described in [17], based
on a digital map showing the geographical distribution of entries in the Ethnologue
[24], a comprehensive catalogue of the worlds languages.c Figure 1 shows examples
b Multiple

regression with number of languages as the dependent variable, MGS is a signicant predictor ( = 0.41, p < 0.001), while number of pathogens is not a statistically signicant predictor
( = 0.20, p = 0.056), size of country is included as control variable. Data taken from [43, 28].
c Digital language maps are produced by Global Mapping International (http://www.gmi.org).
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Table 1. HLM parameter estimates showing the relationship between ethnolinguistic area
(log10 km2 ) and various environmental variables. (LAT: Absolute Latitude, NPP: Net Primary
Productivity, MGS: Mean Growing Season, PREC: Mean annual precipitation, TEMP: Mean
annual temperature).

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Environmental predictors
Subsistence strategy

LAT

Foraging
Pastoralism
Agriculture

0.589
0.617
0.319

NPP
0.397
0.567
0.038 (ns)

MGS

PREC

TEMP

0.585
0.558
0.173

0.488
0.699
0.249

0.543
0.010 (ns)
0.157

of the language polygons that make up this map. Briey, this database integrates
information about the geographical extent of ethnolinguistic groups, with environmental, ecological, and ethnographic data. Here we test how the strength of the
relationship between various environmental variables and the area covered by an
ethnolinguistic group varies across dierent subsistence strategies. Societies are classied according their primary mode of subsistence using the available ethnographic
data from the Ethnographic Atlas [25] (Foraging [Hunting, Gathering, and Fishing],
n = 28; Pastoralism, n = 55; Agriculture, n = 498).
Table 1 shows the parameter estimates of the correlation between the area covered by an ethnolinguistic group and several (mean-centered) environmental variables in a series of HLMs. There is a positive relationship between ethnolinguistic
area and absolute latitude across all subsistence strategies. However, as latitude is
only a proxy for some other variable, and to test the hypotheses discussed above, we
assess the climatic variables annual precipitation and temperature, and also examine a measure of environmental productivity (Net Primary Productivity: a measure
of the net amount of plant biomass converted from solar energy during photosynthesis), and a measure of environmental risk (Mean Growing Season: the number
of months in a year in which the mean temperature is above 6 C and the total
precipitation in millimeters is more than twice the mean temperature in centigrade
[43]). As predicted, the strongest correlations between the climatic variables and
group area are found in the foraging and pastoral societies while the correlations are
generally weaker for those societies that practise agriculture (see Fig. 3). An interesting exception to this general pattern is the non-signicant relationship between
area and temperature in pastoralists. This may reect the fact that pastoralists
tend to inhabit quite arid environments, and therefore the most pertinent variable
aecting survival will be the amount of rainfall.

2.3. Ethnolinguistic groups and subsistence strategy in the

Americas and Australia
Although the results from the old world sample are in line with the predictions of
the niche construction hypothesis, we only have a relatively small sample of foraging
populations to consider, which prevents more rigorous multivariate analyses from
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ethnolinguistic area

ethnolinguistic area

6
6

2
0.00

0.02

0.04

0.06

0.08

0.10

0.00

0.02

0.04

0.06

0.08

0.10

NPP

(a)

(b)
7

ethnolinguistic area

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NPP

6
5
4
3
2
0.00

0.02

0.04

0.06

0.08

NPP

(c)
Fig. 3. The relationship between the area covered by an ethnolinguistic group (log10 km2 ) and
Net Primary Productivity varies according to the groups subsistence strategy. Agriculturalists
(a) show no relationship, while the negative correlation between these variables for both Pastoralists (b) and Foragers (c) is reasonably strong.

being performed. We therefore examine the distribution of ethnolinguistic groups

in Australia and the Americas. At the time of rst European contact with these
regions, Australia was populated solely by foragers, while large numbers of foragers
were also still present in the Americas. The Ethnologue and associated maps only
show the current distribution of native languages in this region, which are often
heavily circumscribed compared to their original distributions. We therefore had
to construct a new ethnolinguistic database using maps describing the estimated
geographical distribution of languages at the time of rst contact with European
societies [42]. These maps are undoubtedly only an approximation of the true distribution of native groups, and were constructed from sources reecting an extended
period (and therefore do not represent a single narrow time slice). However, such
sources of error are only likely to introduce noise into our data and it seems unlikely
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The Evolution of Ethnolinguistic Diversity

Fig. 4. Where only point data were shown on the maps ethnolinguistic polygons were created
using the Thiessen algorithm, which works by placing a line perpendicular to the midpoint between
two points. In this example from North Queensland, Australia the curved lines indicate where the
maps already indicated the boundaries of dierent linguistic groups in some places.

that they have introduced a systematic bias with respect to the hypotheses being
tested in this paper. These maps were scanned and geo-referenced in the Geographic
Information System ArcGIS v9.1 (using multiple reference points and a third-order
polynomial algorithm) in order to bring all maps into the same coordinate and
projection system. Polygons were then created by drawing around each individual language. In some regions, single languages were shown only as a number. In
this case, the language was represented as a point and the geographical extent of
that language was estimated using Thiessen polygons (see Fig. 4). The area of these
polygons was then calculated under a Robinson projection. Multiple languages were
sometimes indicated with only at a single point with a range of numbers, in such
cases the area for the entire polygon was calculated and divided by the number of
languages indicated (in such cases only one entry was used in the statistical analyses). The resulting language polygons were overlaid onto high resolution maps of
various environmental variables and a number of values were calculated for each
language (e.g. mean annual temperature) [17]. Although agriculture was invented
independently in the Americas, its range was still somewhat limited before European contact. In order to categorize the subsistence strategy of the ethnolinguistic
groups in the Americas, we took the estimated boundary of native agriculture in the
Americas as delimited by [5] (Fig. 5). We can assign the societies living outside this
region to a foraging mode of subsistence, the societies that fall inside this zone are
categorized as agricultural (although it should be noted that some societies within
this zone may also have subsisted predominantly via foraging).
Table 2 shows the HLM parameter estimates of several environmental predictors of the area covered by an ethnolinguistic group (again groups are modeled as
being nested within language families). The results support the ndings of the rst
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ethnolinguistic area

5
4
3
2
1
0.00 0.02 0.04 0.06 0.08 0.10

ethnolinguistic area

ethnolinguistic area

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NPP

2
0.00 0.01 0.02 0.03 0.04 0.05 0.06

0.00

NPP

0.02

0.04

0.06

0.08

NPP

Fig. 5. The strength of the relationship between ethnolinguistic area and NPP varies according
to the subsistence strategy employed by native societies of the Americas and Australia. The area
covered by agricultural ethnolinguistic groups in the Americas (A) which show no relationship
with NPP, while the extent of ethnolinguistic groups of foraging societies of the Americas (B)
and Australia (C) are negatively related to NPP. The strength of this relationship is increased
further in Australia if societies found in the eastern temperate forests of Australia (empty triangles) are removed from the analysis (Regression lines: Solid: all societies; Long-dashed: Australian
non-eastern temperate forests; Short-dashed: Australian eastern temperate forests). Boundary of
prehistoric agriculture adapted from [5].

Table 2. HLM parameter estimates indicating the strength of relationship between ethnolinguistic area and various environmental predictors. Ethnolinguistic group area is generally better
predicted by the environmental variables in the foraging regions of the Americas and Australia,
than in the agricultural region of the Americas.

Australia
Americas
Foraging
Agriculture

Latitude

NPP

MGS

PREC

TEMP

0.298

0.238

0.165

0.390

0.259

0.352
0.080

0.531
0.004 (ns)

0.358
0.045 (ns)

0.392
0.040 (ns)

0.409
0.011 (ns)

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Table 3. HLMs showing the relationship of NPP and MGS with ethnolinguistic area in the
same model. AIC scores show the t of these models in relation to a null model containing only
the grouping variable language family.
Americas

AIC Null model

AIC Alternative model

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Intercept
NPP
MGS
Note:

# Excluding

Australia#

Foraging

Agriculture

506.053
343.592

504.957
398.640

981.132
981.274

Parameter
estimate
4.150
25.048
0.003

p
<0.001
<0.001
0.838

Parameter
estimate
5.120
36.185
< 0.001

p
<0.001
<0.001
0.992

Parameter
estimate
4.154
2.553
0.029

p
<0.001
0.193
0.050

cases from eastern temperate forest zone (see Fig. 5).

analysis: the societies in Australia and the Foraging regions of the Americas, show
stronger correlations with environmental factors than the agricultural societies of
the Americas. We also ran HLMs with both NPP and MGS as predictors in order to
test between the productivity and environmental risk hypotheses. The results show
that in both the forager zones NPP is a signicant predictor of ethnolinguistic
area, while in the agricultural zone of the Americas NPP remains non-signicant.
By including both NPP and MGS in the same model, MGS is on the borderline of
statistical signicance (at p = 0.05) in the agricultural zone. However, we should be
wary about over-interpreting this result as the overall AIC score for this alternative
model is only marginally better than the null model. Furthermore, MGS is a weak,
non-signicant predictor of ethnolinguistic area if it is the only co-variate included
in the HLM (Table 2).
3. Discussion
Our results lend support to the hypothesis that the factors shaping the diversity
and distribution of ethnic groups have changed over the course of human evolutionary history. Based on these ndings, and those from previous studies, we tentatively
suggest the following temporal sequence. For the majority of our history, foraging
(of one form or another) was the only subsistence strategy employed. Therefore,
the most important factor determining the distribution of ethnic groups during
this time would have been environmental productivity, determined primarily via
climatic variables such as temperature and precipitation. With the advent of agriculture after the end of the last ice age, the direct eects of climate on ethnic
diversity were reduced. While animal herding is also a Holocene development, pastoralists are less able to modify their environment than agriculturalists and our
results suggest that the area covered by their ethnolinguistic groups appears still
to be determined substantially by the direct eects of climate. Finally, the development of agriculture eventually led to the evolution of larger-scale political groups,
with more complex social and political institutions that were able to integrate and
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T. E. Currie and R. Mace

coordinate the actions of larger numbers of people. Such groups were also able to
out-compete less complex groups and as they expanded they displaced, replaced
or assimilated existing ethnic groups, leading to an overall reduction in diversity.
This process has further severed the link between the geographical extent of ethnic
groups and the direct eects of environmental variables.
There is some evidence that ecological risk might become a more important
factor shaping ethnolinguistic diversity for agriculturalists. This may reect the fact
that while the societies practicing foraging and pastoralism can respond to shortfalls
in production through their greater mobility, the more sedentary agriculturalists
may have to rely to a greater extent on buering such risks through their social
networks as envisaged by the ecological risk hypothesis. However, we note that
ethnolinguistic group area in the agricultural zone of the Americas showed only
a weak relationship with MGS at best, while MGS explains only small amount
of variation in the extent of ethnolinguistic groups in the agricultural societies of
the Old World. For agriculturalists, productivity may still be an important factor
determining the distribution of ethnic groups, however, the productivity of any
particular region came to depend more on labour inputs into farming, the kinds of
crops grown and the process of articial selection to create more desirable cultivars.
Furthermore, dierent crops require dierent climatic conditions for optimal growth
with some vegetative crops doing best in warm, wet environments such as taro while
others such as sweet potato do better in drier conditions [36]. Grain crops, such as
rice, need a seasonal climate with appropriate annual patterns of temperature and
rain to allow the grain to dry once harvested [4].
While our results are consistent with the niche construction hypothesis it is
important to point out that a number of other factors tend co-vary with the presence
of agriculture, and may also decrease the link between the area of an ethnic group
and environmental variables. For example, the presence of large-scale, expansive
polities mentioned previously, or trade of products including staple crops. It should
also be noted that foragers also conduct important niche construction activities
(e.g. the use of controlled burning as a resource management strategy in Australian
Aborigines [7], or have a resource bases that are less linked directly to terrestrial
environmental variables (e.g. Salmon in the Northwest Coast of America [1]). These
factors may also aect the relationship between the climatic variables considered
here and the area covered by ethnic groups. In order to test the relative importance
of these dierent processes, future work will attempt to integrate such pertinent
ethnographic information with information about the native distribution of ethnic
groups in the Americas and Australia in the manner that we have previously done
for the old world [17].
While environmental factors such as Net Primary Productivity were shown
to be better predictors of the area covered by ethnolinguistic groups for foragers
than for foragers than for agriculturalists, productivity more broadly dened may
still be a key factor underlying the propensity for human to split themselves o
into distinct ethnic groups in the modern world. Interestingly, major metropolises
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The Evolution of Ethnolinguistic Diversity

such as New York, or London, which can be thought of as being very productive
economically, are well known for containing many dierent ethnic communities. For
example, it is estimated that more than 300 languages are spoken by schoolchildren in London [2]. [46] found that economic productivity of US cities correlates
positively with the number of languages spoken. This suggests that the evolutionary ecological perspective we have employed in this paper to examine may also be
fruitfully applied in future work to explain the patterns and processes of ethnic
dierentiation in contemporary, urban settings. It will also be important for such
an approach to incorporate the impact that national or regional policies on such
factors as trade, immigration, and the assimilation of dierent groups, in shaping
ethnic diversity in these settings.
Our results demonstrate that the distribution of human cultural groups varies
systematically with a number of social and environmental factors. As such, our
approach may have important implications for archaeologists who wish to study
the ethnicity of past populations, and may provide an independent lens through
which to view this aspect of the past. Interpretations about the ethnic aliations
of societies represented in the archaeological record have a long and controversial history, and there is a vigorous debate about to what extent ethnicity can be
deduced from material remains [34, 54, 59, 32]. One possibility is that the expected
number of ethnic groups of any particular region could be estimated based on estimates of productivity, and information about such things as population density and
social complexity. Such estimates could then be tallied with hypothesized archaeological markers of ethnic groups. A similar idea about using ecological data as a
window into the past has been proposed by Binford [6], particularly with respect to
predicting the distribution of hunter-gatherer groups. His approach was explicitly
inductive and attempted to make predictions about the past based on statistical
associations within his particular dataset without reference to an underlying theory as to why those associations existed. However, such a strategy runs the risk of
nding associations that cannot be generalized to other situations. We argue that is
better to make predictions about the possible past distribution of groups or cultural
traits using the kind of data we have analyzed in this paper on the basis of specic
hypotheses that postulate a link between these phenomena.
In this paper, we have used a Geographical Information System to integrate
data from a number of dierent sources in order to statistically test a number
of hypotheses about the evolution of ethnic diversity. The use of spatially explicit
mathematical models or computer simulations could provide further tests of these or
other hypotheses, and would represent an important complement to this approach.
Such techniques have previously been employed to address related questions about
biological species diversity [49], human genetic diversity [23], and other aspects of
cultural evolution (see other papers in this volume). Hypotheses involving the relationship between environmental variables and ethnic diversity, such as the productivity and ecological risk hypotheses, are based on equilibrium models [13, 23, 43].
However, if the sequence described above is correct then, on a global scale, such
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T. E. Currie and R. Mace

equilibria have been systematically disrupted at least twice. First, in the transition
from foraging to agricultural modes of subsistence [43], and secondly in the development of more complex forms of political organization [17]. Estimates of high recent
rates of language loss [56] suggest that we have not yet settled down to a new stable equilibrium level of ethnolinguistic diversty. Furthermore, the development and
spread of other cultural innovations may cause disruption to equilibrium that may
be witnessed on a more local or regional scale and over shorter time scales, with
some practices creating either positive or negative feedback that may have consequences for the distribution of ethnic groups. The extent to which these dynamic,
non-equilibrium processes have shaped the evolution of human ethnic diversity will
be modeled and tested using a complex systems framework in future work. Simulations could look explicitly at changes in distributions over time, or could test
how well alternative evolutionary models explain the current distribution of ethnolinguistic groups as indicated in the kind of maps we have examined in this paper.
Kandler et al. [35] have recently used such spatially explicit models to examine the
replacement of Celtic languages by English in Scotland.
Another future direction in which a spatial information can be used to test
hypotheses about the formation of ethnic groups that we are currently persuing
involves the spatially explicit use of cultural phylogenetics [37, 27, 16]. This kind
of cultural phylogeographic thinking has been employed in previous studies that
have applied phylogenetic methods to culturally inherited data, particularly those
that have examined large-scale migrations of human populations (e.g. [31, 26, 14, 9].
However, these studies have not considered explicitly how geographically associated
factors may aect cultural group diversication. One particular advantage of such
an approach is that it will allow us to explore how ethnic groups have diversied over
time and can be used to examine how rates of ethnogenesis and ethnopathosis vary
according to movements into new environments, or the adoption of new cultural
practices or social institutions (Fig. 6). This kind of cultural phylogenetic approach
will also be used to assess the relationship between ethnic diversication and other
aspects of cultural diversity. For example, Pagel et al. [47] used language phylogenies
to infer that the rate of change of lexical items increases following the divergence
of two languages, a phenomenom they interpreted as evidence that language was
being used as a marker of group identity. This approach will be adapted to examine
whether other ethnic group markers also change in this punctuated manner.
In conclusion, we have shown that the strength of the relationship between ethnolinguistic area and environmental variables is stronger in societies whose primary
mode of subsistence is foraging. This nding was replicated using the estimated
native distributions of ethnolinguistic groups in the Americas and Australia. In
particular, Net Primary Productivity is shown to be a good predictor of the area
covered by ethnolinguistic groups in foragers but not in agriculturalists. This provides support for the idea that the factors aecting ethnic diversity have changed
in a systematic way with changes in subsistence strategies and social organization.
In this paper, we have demonstrated how examining geographic and environmental
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The Evolution of Ethnolinguistic Diversity

Fig. 6. (Color online) The use of spatially explicit cultural phylogenies can be used to test
hypotheses about ethnic group formation and how the rates of ethnogenesis and ethnopathosis
may be aected by ecological variables. This hypothetical example shows a phylogeny mapped on
to a background representing an environment that exhibits a cline (red to white) in a particular
variable. A number of dierent ethnic groups (large circles) have formed after spreading out and
diverging from a common ancestor in the homeland A. The phylogeny shows that the net rate of
ethnogenesis has been larger in the redder areas.

information can shed light on the processes that have shaped human ethnolinguistic
diversity. The use of such a spatially explicit framework to understand the evolution of biological diversity is common in the natural sciences, and it is hoped that
such an approach will prove fruitful in our attempts to understand the evolution of
human ethnic groups and other aspects of cultural diversity.

Acknowledgments
The authors are supported by a European Research Council grant. Currie was also
supported at various stages of this research by an ESRC/NERC interdisciplinary
studentship, a Japan Society for the Promotion of Science Post-doctoral fellowship,
and the AHRC Centre for the Evolution of Cultural Diversity. This paper reects
only the authors views, the European Union is not liable for any use that may be
made of the information contained therein. We thank the editors James Steele and
Anne Kandler and two anonymous reviewers for their constructive comments on an
earlier draft of this paper.

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