Journal of Archaeological Science 120 (2020) 105168

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Journal of Archaeological Science

journal homepage: http://www.elsevier.com/locate/jas

The global ecology of human population density and interpreting changes

in paleo-population density
Jacob Freeman a, b, *, Erick Robinson a, Noelle G. Beckman b, c, Darcy Bird d, Jacopo A. Baggio e, f,
John M. Anderies g
a
Anthropology Program, Utah State University, Logan, UT, 84322, USA
b
Ecology Center, Utah State University, Logan, UT, 84322, USA
c
Department of Biology, Utah State University, Logan, UT, 84322, USA
d
Department of Anthropology, Washington State University, Pullman, WA, 99164, USA
e
School of Politics, Security and International Affairs, University of Central Florida, Orlando, FL, USA
f
Sustainable Coastal Systems Cluster, National Center for Integrated Coastal Research, University of Central Florida, Orlando, FL, USA
g
School of Sustainability, Arizona State University, Tempe, AZ, USA

A R T I C L E I N F O A B S T R A C T

Keywords: Explaining variation in human population density constitutes a basic research problem in human ecology and
Human ecology archaeological science. To contribute to this basic research problem, we build a graphic model and conduct a
Radiocarbon global analysis of the effects of ecological variables, controlling for technological differences, on human popu­
Climate change
lation density. Our results indicate that human population density displays a consistent relationship with
Human population ecology
Macroecology
ecological variables across productive technologies. Human population density peaks in high productivity en­
vironments with moderate levels of species richness and moderate to low levels of pathogens among hunter-
gatherer, subsistence agricultural, and industrial societies. Population density is lower in low productivity en­
vironments as well as environments with high productivity, high species richness and high pathogen loads. The
productive technologies associated with agriculture and industrial production impact human population density
more than ecological variables, and shifts toward agriculture or industrial production may weaken the effect of
net primary productivity on population density. These results illustrate the nature of the relationship between
population density and ecological variables, which are partly driven by climate, and the results provide a basis
for hypotheses that researchers can use to analyze the potential effects of climate change on material record
estimates of human paleo-population density.

1. Introduction us to make a very practical point about the analysis of paleodemo­

Explaining variation in the global distribution of human population
density has been a research question in archaeology and human ecology
for decades (e.g., Tallavaara et al., 2018; Freeman, 2016; Freeman and
Anderies, 2015; Kelly, 2013; Hamilton et al., 2007; Luck, 2007a, b;
Binford, 2001, 1983; Hassan, 1981). In this paper, we make two con­
tributions to this research question. First, to increase our understanding
of the global scale variation in human population density, we evaluate a
model that describes the effects of productive technology and three key
ecological variables on human population density: the productivity of an
ecosystem, the species richness of an ecosystem, and the pathogen load
of an ecosystem (from a human’s perspective). Second, our work allows
graphic proxies: we can use the results of our study (and similar studies)
to create more informed hypotheses about when climate variables
should and should not strongly correlate with estimated changes in
paleo-population density. To this end, we use our results to propose
hypotheses that researchers can use to guide their investigation of po­
tential relationships between paleoclimate and paleodemographic
proxies. In the remainder of this paper, we propose a graphic model of
ecological constraints on human population density, evaluate the model,
and develop a set of hypotheses for analyzing
paleoclimate-paleo-population density proxies.

* Corresponding author. Anthropology Program, Utah State University, Logan, UT, 84322, USA.
E-mail address: jacob.freeman@usu.edu (J. Freeman).

https://doi.org/10.1016/j.jas.2020.105168
Received 12 August 2019; Received in revised form 22 April 2020; Accepted 25 May 2020
Available online 6 June 2020
0305-4403/© 2020 Elsevier Ltd. All rights reserved.
J. Freeman et al.
1.1. An ecological model of human population density
The inspiration for our paper comes from the explosion of research
that uses radiocarbon ages to develop time-series of relative human
population density (population size in a constant area) over the last 15
years (e.g., Robinson et al., 2019; Freeman et al., 2018a; Bevan et al.,
2017; Gayo et al., 2015; Kelly et al., 2013; Shennan et al., 2013),
referred to as the ‘dates as data’ approach. While this approach has been
critiqued, our purpose is not to defend against or add to such critiques.
Rather, we want to address a separate research problem that this corpus
of research raises. Suppose that the ‘dates as data’ approach provides a
reliable proxy for changes in human population density. Would our
understanding of the factors that affect human population density be
sufficient to interpret associations (or lack thereof) between paleo­
climate proxies and such datasets? We propose that the answer to this
question is: No (e.g., Robinson et al., 2019; Tallavaara et al., 2018;
Freeman et al., 2018b; Cromb�e and Robinson, 2014). As Tallavaara et al.
(2018) argue, we cannot understand the impacts of paleoclimate change
on paleo-human population density without understanding how
ecological variables affect human population density in the first place.
Holding technology and social organization equal, climate does not
directly impact human populations. Rather, climate affects variables
such as the net primary productivity, species richness, and the pathogen
load of ecosystems (Kreft and Jetz, 2007). It is these attributes of eco­
systems that impact the population density of humans and mammals in
general (Santini et al., 2018; Luck, 2007a).
Therefore, a key research problem in the analysis of paleoclimate and
paleodemography is to understand the chain of relationships between
climate and ecosystem attributes on the one hand and ecosystem attri­
butes and human population densities on the other hand. The relation­
ship between climate and ecological variables has been the subject of
research in ecology for decades and is fairly well developed (Kreft and
Jetz, 2007; Odum, 1997; Rosenzweig, 1968). However, the relationships
between ecosystem attributes and human population density needs
more description and explanation (Tallavaara et al., 2018). To help
build a knowledge base of the relationship between ecosystem attributes
and human population densities, we build on previous research to pro­
pose and evaluate a graphic model that may explain global scale vari­
ation in human population density.
Studying hunter-gatherers, Tallavaara et al. (2018, p. 1232) propose
a qualitative model to explain global variation in hunter-gatherer pop­
ulation density. They propose that: (1) The main ecosystem attribute
that affects population density is potential net primary productivity
(NPP). NPP refers to the amount of biomass produced per unit space per
unit time (Odum, 1997). This variable, thus, estimates the amount of
potential energy available for secondary consumers in a given area. As
NPP increases, the availability of food (energy) increases and, thus, so
should population density. (2) Studies of biodiversity indicate that as the
species richness of ecosystems increases, so does the stability and mean
of ecosystem productivity (Wang et al., 2019; Mazancourt et al., 2013;
Cardinale et al., 2012). Thus, more species rich ecosystems, at large
scales, should be more productive and less risky for human foragers
because foragers can take advantage of species that serve similar needs
but have asynchronous responses to climate drivers. (3) Pathogen load
should affect population density by increasing morbidity and mortality,
which creates incentives for foragers to spread out to avoid the disease
transmission that causes such morbidity and mortality.
Tallavaara and colleagues’ analysis of approximately 300 hunter-
gatherer societies broadly supports the above model. They conclude
that, despite much social and technological variation, hunter-gatherer
populations do not transcend the ecological constraints that affect the
population densities of other mammals. While important work, the
above model needs more formalization and testing for three reasons.
First, Tallavaara and colleagues’ analysis indicates that ecosystem
attributes interact in their effect on human population density (see, for
example, Figure S1 Tallavaara et al., 2018). For example, in more
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Journal of Archaeological Science 120 (2020) 105168
species rich environments, a one unit increase in NPP results in a larger
increase in population density than in species poor environments. This
might occur because increases in NPP result in more energy available for
human consumers, and higher species richness decrease the variability
of NPP and important nutrients. Thus, increases in species richness
might amplify the positive effect of NPP on population density because
foragers in more species rich environments experience shortfalls of re­
sources and nutrients less often. Understanding such interactions ben­
efits from developing a more formal model of the effects of ecosystem
attributes on human population densities.
Second, throughout the Holocene human societies experienced
drastic changes in technology. One of the most important examples of
such technological change occurred with the adoption of agriculture.
Conventional wisdom states that the adoption of agriculture increased
the carrying capacity of human societies (Freeman, 2016; Glassow,
1978). If the effect of technological change is large relative to changes in
ecosystem attributes, then these technological changes might be driving
observed shifts in population density despite changes in climate that
affect ecosystem attributes. Interpreting changes in the density of
human populations over time requires understanding the relative effects
of technological change vs. ecological attributes on population density.
Finally, it is unclear whether population density displays similar
relationships with NPP, species richness, and pathogen load among
agricultural and industrial societies as it does among hunter-gatherers.
Do ecosystem attributes constrain the population density of agricul­
tural and industrial societies in the same way as they do human for­
agers? If ecosystem attributes constrain human population densities in
the same way across different productive technologies, this would sug­
gest that a general underlying process affects human population den­
sities. If the relationships between ecological attributes and population
density vary systematically with technology, then this would suggest
that shifts in climate that impact ecosystem attributes would have
different consequences for human populations, depending on their
productive technology. Investigating the above potentialities can help
clarify when we should and should not expect associations between
material record estimates of population density, paleoclimate proxies,
and material traces of productive technology.
1.2. A graphic model
Fig. 1 builds on Tallavaara and colleagues’ qualitative model and
provides a set of hypotheses to guide an empirical analysis of global
scale variation in hunter-gatherer, agricultural and industrial society
population density. Fig. 1A describes the marginal benefits and costs of
higher NPP and higher population density, respectively, and Fig. 1B
summarizes the expectations for how population density varies with
NPP, holding the effects of species richness and pathogen load equal, for
two different productive technologies.
The red curves on Fig. 1A describe the marginal benefits of increases
in NPP for human consumers. These curves capture the idea that as NPP
increases, the availability of energy in an ecosystem that humans can
consume increases, and thus the ecosystem can support more people per
unit area. The red curves also capture the idea that the marginal benefits
of NPP display diminishing returns. This means that as energy becomes
more available, humans experience diminishing benefits. Benefits
diminish because, even though more energy is available as NPP in­
creases, not enough time exists in the sleep/wake cycle or efficient
enough infrastructure does not exist to exploit all of the extra available
energy. As a starting hypothesis, we propose that the relationships dis­
played by the red curves holds across types of productive technology.
This may be the case as, ultimately, agricultural societies (subsistence
and industrial) depend on energy from the sun and water to grow crops,
just as hunter-gatherers depend on these climate inputs for the growth of
wild resources.
Second, we propose that the marginal benefits of increases in NPP
vary with species richness. In low NPP environments, high species
J. Freeman et al.
Fig. 1. A-Graphic model of the marginal benefits of higher NPP and the mar­
ginal costs of higher population density. The dashed red curve illustrates the
marginal benefits of NPP holding species richness equal at a moderate level and
the solid red curve holding species richness equal at a high level. The dashed
blue curve illustrates the marginal costs of higher population density holding
pathogen load equal at a moderate level and the solid blue curve at a high level
of pathogens. For all values of NPP that a red curve is above a blue curve, an
increase in NPP will result in an increase in population density. For all values of
NPP for which a blue curve is above a given red curve, increases in NPP will
result in decreases in population density. B-Expected relationships between
population density and NPP for two different combinations of species richness
and pathogen load and for two different productive technologies. Dashed
curves ¼ hunter-gatherers; Solid curves ¼ subsistence agriculturalists. (For
interpretation of the references to color in this figure legend, the reader is
referred to the Web version of this article.)
richness leads to more stable productivity and amplifies the benefits of
increases in NPP. However, high species richness may also result in more
steep diminishing returns. The basic idea behind this postulate is that all
individuals face an exploration-exploitation tradeoff. The more re­
sources used, the more that time is spread among resources leading to
less efficiency in the production of food. This amplifies the saturation
effect of more energy (NPP) caused by not enough time or efficient
enough infrastructure to exploit all of the energy available in high NPP
settings. This is captured by the difference between the moderate di­
versity and high diversity red curves in Fig. 1A. Steeper diminishing
returns occur in the high species richness (diversity) environment due to
the impact of allocating time across many resources. Again, as a starting
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Journal of Archaeological Science 120 (2020) 105168
hypothesis we expect this dynamic to occur across types of productive
technology. Indeed, species richness in agricultural societies mimics the
underlying species richness of ecosystems (Freeman, 2012) and could
lead to an exploration–exploitation tradeoff that generates steeper
diminishing returns.
Finally, the blue curves in Fig. 1A describe the marginal costs of
higher population density. The costs of higher population density
include increased disease transmission and potentially more conflict as
individuals encounter each other more often. These costs increase at an
accelerating rate as human population density increases. Note the two
blue curves. These represent different levels of pathogen load: A mod­
erate pathogen load curve and a high pathogen load curve. These two
curves capture the idea that the marginal costs of living at higher pop­
ulation density will increase faster when pathogens are more abundant
than when less abundant in ecosystems.
The key insight of the above model is that global scale variation in
human population density, holding productive technology equal, should
result from an interaction between NPP, species richness, and pathogen
load. As long as the red curves in Fig. 1A are higher than the blue curves,
population density increases as NPP increases, though at a diminishing
rate. Once a given red curve crosses a blue curve, population density
should decline as the costs of the next unit of increase in population
density outweigh the benefits of the next unit increase in NPP.
Fig. 1B summarizes a set of guiding empirical expectations that
follow from the model. The black dashed curves in Fig. 1B illustrate a
less productive technological base and the solid curves a more produc­
tive technological base. The more humped curves illustrates how pop­
ulation density should increase with NPP, holding species richness and
pathogen abundance equal at high levels. The flatter curves illustrates
how population density should increase with NPP holding species
richness and pathogen abundance equal at more moderate levels. The
key here is that as NPP increases from very low levels, population
density should, initially, be higher in species rich and pathogen abun­
dant environments than in moderate species richness and pathogen
abundance environments. This is because the distance between the high
diversity red curve and high pathogen blue curve (the net benefits of
higher NPP) is greater than the distance between the moderate diversity
red curve and moderate pathogen load blue curve in Fig. 1A.
However, as NPP continues to increase, the distance between the
moderate diversity red curve and moderate pathogen blue curve actu­
ally becomes greater than the distance between the high diversity red
curve and high pathogen blue curve in Fig. 1A. This means that at a high
enough NPP, population density in moderate richness and moderate
pathogen abundance environments exceeds the population density of
high species richness, high pathogen abundance environments. Indeed,
the high diversity red curve and high pathogen blue curve actually cross
each other at high values of NPP, leading to a decline in population
density in such high NPP environments.
Note, the marginal benefits of NPP, alone, are always positive or
neutral in the model. The negative relationship between NPP and pop­
ulation density displayed by the humped curves in Fig. 1B results from
the interaction of NPP with species richness and pathogen load. The
moderate species richness and pathogen load red and blue curves in
Fig. 1A never cross; thus, the moderate diversity and pathogen load
curves in Fig. 1B never decline. In such environments, the benefits of
higher NPP always outweigh the costs of higher population density,
though the rate of population density increase does decline.
In sum, the effect of NPP on human population density depends on
context. At low levels of NPP, higher species richness decreases variance
in the supply of food, and this has benefits that far outweigh the costs of
aggregating into more dense populations. However, at high levels of
species richness, the benefits of more stable NPP do not outweigh the
costs of aggregating into higher population densities in terms of disease
transmission and fights. Thus, population density may decline due to the
interaction between species richness, pathogen load and NPP. As a
starting point, we propose that this process holds across multiple types
J. Freeman et al.
of productive technology systems. The solid curves in Fig. 1B describe
the proposed effect of productive technology in addition to ecological
attributes. Here, the intercepts of the solid curves are higher than the
dashed black curves, but the forms of the curves remain exactly the
same. This quantifies the proposition that the dynamics proposed in
Fig. 1A operate across types of productive technologies. Shifts to agri­
cultural or industrial economies simply augment the energy available
along NPP gradients, leading to a higher population density at a given
level of NPP, on average, for an industrial society vs. an agricultural
society vs. a hunter-gatherer society.
2. Data and methods
To analyze the global distribution of human population density, we
use three global data sets on human population density. We selected
these data sets because (1) all three data sets attempt to provide a global
sample of hunter-gatherer, subsistence agriculturalist/pastoralist or in­
dustrial societies, and (2) the data sets were collected independently and
without knowledge of our specific research question when assembled.
The first data set was published by Binford (2001) and contains infor­
mation on the population density of 339 ethnographically recorded
hunter-gatherer societies from around the world. The second data set
includes 86 societies primarily dependent upon agriculture and pasto­
ralism for their subsistence needs, previously published to study the
latitude-domesticated plant species richness gradient (Freeman, 2012)
and population-territory scaling (Freeman, 2016). The third data set
consists of population, land area, and energy consumption data
compiled from the World Bank (TWB, 2016) and International Energy
Agency (IEA, 2016) since 1973 to study population size–energy scaling
(Freeman et al., 2018b). We simply calculated the mean population
density of each country since 1973. The data are available for 117
countries. Together, these three data sets provide information on the
global distribution of human population densities among
hunter-gatherer, subsistence agricultural and industrial societies.
The resulting dataset consists of 529 societies (329 hunter-gatherer,
86 subsistence agricultural and 114 modern countries). Ten hunter-
gatherer societies were removed due to our inability to accurately
calculate ecological parameters as a consequence of the societies living
on coasts or islands. We decided to remove rather than impute the
missing values as this does not change our results. The sample of hunter-
gatherer societies overlaps with Tallavaara et al. (2018). The
hunter-gatherer and subsistence agricultural societies territories are
defined by ethnolinguistic units and the territory of modern countries is
defined by modern boarders. This means that several countries have
very large territories. The consequences is that our calculation of
average NPP, species richness, and pathogen load (discussed below)
masks a high degree of environmental variation in several cases. This
fact actually decreases the power of the industrial data set to detect the
effects of ecological attributes on population density; nonetheless, we
still detect such effects as noted in the results below.
For comparability, to estimate NPP and species richness, we follow
the methods developed by Tallavaara and colleagues (Tallavaara et al.,
2018, 2017). We estimate NPP from the climatological parameters of
temperature and rainfall using the Miami model. Temperature and
rainfall data were downloaded from WorldClim.org (Hijmans et al.,
2005), and the underlying projection of the climate data, resolution and
extent of the climate rasters was used as a template to create species
richness rasters (Tallavaara et al., 2017, pp. 1–4). Species richness was
estimated by combining mammal, bird and vascular plant richness.
Specifically, bird and mammal richness data were obtained from Biod
iversityMapping.org website (Jenkins et al., 2013; Tallavaara et al.,
2017). Native vascular plant richness was modeled from climate pa­
rameters following Kreft and Jetz (2007), and then interpolated using
inverse weighted distance to the grid size of the data on bird and
mammal richness (Tallavaara et al., 2017, 4–7). To link NPP and species
richness data to each society, we used the center of each
4
Journal of Archaeological Science 120 (2020) 105168
ethnographically recorded societies’ territory as the point to extract NPP
and species richness. We used a mean value for the modern nation states.
Finally, we estimated pathogen load at a global scale by using esti­
mates of pathogen diversity and the incidence of pathogens in modern
populations. Pathogen load is a composite index of both pathogen
abundance and ease of transmission. The pathogen load index is calcu­
lated using the average z scores from the prevalence of ten different
pathogens (spirochetes, leprosy, plague, schistosomes, leishmaniasis,
trypanosomes, typhus, filariae, dengue, and malaria) (Tallavaara et al.,
2018; Low, 1990). The pathogen stress index was interpolated to
generate a global raster, again scaled to the resolution of the climate
rasters used to project NPP and species richness (Tallavaara et al., 2017).
Fig. 2 compares the estimated NPP and species richness of each of the
three productive technology groups in our data set. We constructed
Fig. 2 to inspect potential biases between productive technology groups
in terms of the terrestrial ecosystems that they inhabit. The curves are
visual aids that help divide the graph into spaces where combinations of
NPP and species richness are common (between the curves) and
extremely rare (outside the curves) on earth. The key point is that the
productive technology types have distributions that overlap in the main
linear scatter, suggesting pretty equal representation of NPP-species
richness environments on earth across the productive technology
groups. The only minor exceptions occur in the upper left quadrant and
lower right quadrant of the graph. In the upper left quadrant, agricul­
tural and contemporary countries dominate these rare ecosystem types.
In the lower right quadrant, hunter-gatherers are more frequent in the
more rare ecosystem combination of high productivity but low species
richness.
2.1. Methods
The expectations illustrated by Fig. 1B guide our analysis of global
human population densities. To evaluate these expectations, we conduct
a multiple regression analysis and specify interaction effects between
the three ecological variables of NPP, species richness, and pathogen
load. A multiple regression recognizes that numerous explanatory var­
iables may impact a single outcome variable. In this case, the model
sketched above indicates that NPP, species richness, pathogen stress,
Fig. 2. Scatter plot of NPP (x-axis) and species richness (y-axis); with all
documented societies in this study coded as hunter-gatherer, subsistence agri­
culturalists/pastoralist, and industrial societies. The figure creates a two
dimensional representation of the range of ecosystems that human societies
occupy on earth. The upper and lower curved boundary lines mark combina­
tions of NPP and species richness that are rare among terrestrial ecosystems
on earth.
J. Freeman et al.
and productive technology should all contribute to explaining variation
in the outcome variable of human population density. Interaction effects
refer to how two or more explanatory variables may work in concert,
rather than in an additive way, to explain variation in an outcome
variable. For instance, if the relationships between NPP, species richness
and pathogen load were simply additive, then we would only postulate
one curve on Fig. 1B, and, by controlling for species richness and
pathogen abundance in a regression, the remaining variation in popu­
lation density would fit the postulated relationship between NPP and
population density well. However, the model described above proposes
that NPP, species richness, and pathogens interact. This means that
changes in species richness and pathogens modify the relationship be­
tween NPP and population density (as one can see in the difference
between the moderate and high diversity and pathogen curves in
Fig. 1B).
Here, we use general least squares and linear mixed effects multiple
regression models to assess the effects of ecological variables and pro­
ductive technology on human population density at a global scale. We fit
linear mixed effects models because such models are appropriate in
cases where variation in an outcome variable may be caused by differ­
ences between groups. For example, how often parents read to their
children might affect a child’s reading performance in school, but per­
formance might also vary between schools due to different school en­
vironments. A mixed effects model allows one to model the variation
explained by parents reading to their children (the “within effect”) and
variation explained by differences between school environments (the
“between effect”). Here, we expect ecological variables to affect human
population density across all types of productive technology (the within
effect), but we also expect density to vary between groups of societies
with different productive technologies (the between effect of hunter-
gatherer, agricultural, industrial technology groups).
We fit three main regression models. First, we fit a general least
squares model fit by maximum likelihood that does not model differ­
ences in population density between productive technology groups. This
is a “null” model that allows us to understand if allowing population
densities to vary between groups improves the performance (variation in
population density explained at a given level of model complexity) of a
regression model. Specifically,
lnD ¼ β1 þ α1 NPP þ α2 BIO þ α3 PAT þ α4 NPP : BIO : PAT þ ε
where D is the population density of a society; NPP is net primary pro­
ductivity; BIO is species richness; PAT is pathogen load; and NPP : BIO :
PAT denotes the interaction of these three variables.
Second, we introduce a random intercept using a mixed effects
regression model fit using maximum likelihood.
lnDij ¼ β1 þ α1 NPPij þ α2 BIOij þ α3 PATij þ α4 NPPij : BIOij : PATij þ vi0 þ εij
where D is the population density of the ith society in productive tech­
nology group j; NPP is net primary productivity; BIO is species richness;
PAT is pathogen load and NPP : BIO : PAT denotes the interaction of
these three variables. The term vi0 is random variation of productive
technology groups around the fixed intercept of β1 . This random vari­
able captures the possibility that productive technologies may cause
variation in human population density. The ε term is the residual error
not explained by the other variables in the regression model.
Third, we introduce a random slope into Eq. (2) and fit a regression
using maximum likelihood.
lnDij ¼ β1 þ α1 NPPij þ α2 BIOij þ α3 PATij þ α4 NPPij : BIOij PATij
þ vi0 þ vi1 NPPij þ εij
where all variables are the same as in Eq. (2) above. However, in this
regression, we add the term vi1 NPPij . This term allows for the possibility
that the slope of the relationship between NPP and the log of population
Journal of Archaeological Science 120 (2020) 105168
density may vary by productive technology groups.
Our methodology compares the fits of the above regression models
and selects the best fitting model as a representation of the data, in
particular, whether population density varies significantly between
productive groups as we expect. Table 1 compares the fit of a general
least squares regression model (Eq. (1) above, denoted glsm in the table)
with a random intercept (Eq. (2), TimeRI) and random intercept and
slope model (Eq. (3), TimeRS). The table provides fit statistics and uses
the log likelihood ratio (L.Ratio) to test the null hypotheses that TimeRI
does not explain more variation in human population density than
model glsm, and that TimeRS does not explain more variation in human
population density than model TimeRI. In this case, we can reject the
successive null hypotheses at p< 0:05, indicating that allowing the in­
tercepts to vary randomly as in Eq. (2) explains more variation in pop­
ulation density than Eq. (1). Further, Eq. (3), which allows both slopes
and intercepts to vary randomly explains more variation in human
population density than Eq. (2) (TimeRI). In short, human population
density varies between productive technology groups, and the slope of
the main effect of NPP on population density varies between productive
technology groups.
Hence, in our main results below, we report the results from Eq. (3).
To evaluate the effect of the interaction of NPP, species richness, and
pathogens, we use effect plots and contour plots. These plots graph the
relationship between NPP and population density holding species rich­
ness and pathogens equal at particular values (as in Fig. 1B). This allows
us to observe if species richness and pathogens modify the relationship
between NPP and population density consistent with the expectations of
the model. Note that in all regression models above, we used z-scores to
center all variables around their mean to avoid potential coefficient
biases introduced by colinearity. We ran all regression models in R using
the nlme and effects packages. All data and scripts are available at: htt
ps://github.com/people3k/JAS2020Ethnographic-Population-Density.
3. Results
To begin, we provide a summary of our results and then describe the
details of the outcomes of our analysis. Our results, in general, are
consistent with the model presented in Fig. 1. (1) As NPP increases,
human population density increases; however, species richness and
(1)
pathogen load significantly modify the relationship between NPP and
human population density. Specifically, there is an optimal population
density environment for humans, holding productive technology equal,
that is defined by high NPP, moderate species richness and moderate to
low pathogen loads. (2) Differences in productive technology have a
much larger impact on potential population density than changes in
ecosystem attributes. (3) Productive technology significantly increases
the population density of human societies and weakens the underlying
(2) positive, fixed effect of NPP on human population density.
3.1. The effects of ecological attributes and productive technology on
population density
Table 2 illustrates the fixed coefficients (within effects) from the best
fitting regression model in Table 1 (TimeRS, Eq. (3)) associated with
individual ecological attributes, the interaction of NPP, species richness
and pathogen load and the random effects associated with productive
technology groups. As predicted, NPP has a positive and significant fixed
effect on human population density. This means, on average, across all
productive technology groups a one unit increase in NPP results in a
0.809 unit increase in the log of human population density. Species
(3) richness (BIO) and pathogen load (PAT) also have positive fixed effects,
though these effects are not statistically significant. More importantly
given the hypothesis developed in conjunction with Fig. 1, NPP, species
richness and pathogen load have a significant fixed interaction effect on
human population density. Species richness and pathogen load modify
the relationship between NPP and population density as depicted in
5
J. Freeman et al. Journal of Archaeological Science 120 (2020) 105168

Table 1
Table comparing the fits of alternative regression models. The mixed effects models that allow population density to vary randomly between productive technology
groups fit the data better than a general least squares model that does not allow population density to vary between technology groups.
Model df AIC BIC logLik Test L.Ratio p-value

glsm 6.00 2501.93 2527.56 1244.97

TimeRI 7.00 1973.19 2003.09 979.60 1 vs 2 530.74 <0.001
TimeRS 9.00 1964.47 2002.90 973.23 2 vs 3 12.73 <0.001

Fig. 3.
Table 2
Fig. 3 illustrates how species richness and pathogen load modify the
The fixed and random effects coefficients in the TimeRS regression model dis­
predicted effect of NPP on the log of population density. First, in very
played in Table 1 and Eq. (3). Standard errors reported in parenthesis and *** ¼
significant at the 99% level. low pathogen load environments (Panel A), groups living environments
with high species richness and low NPP have the lowest population
Fixed Coefficients
densities; however, as NPP increases there is a NPP threshold at 0.2
Predictors Coefficients (se) (about 650 g/m/yr) above which groups living in high species richness
NPP 0.809***
environments are predicted to have the highest population densities.
(0.18)
BIO 0.03 Above the 0.2 threshold, species richness amplifies the positive effect
(0.12) of NPP on population density (purple line has a steeper slope and is
PAT 0.164 above all other lines).
(0.103) Second, as pathogen load increases (panel A to B to C to D), the
NPP : BIO : PAT 0.224***
amplifying effect of species richness on the positive relationship be­
(0.072)
Constant 0.73 tween NPP and population density weakens. In fact, in high pathogen
Random Effects load environments (Panels C and D), low species richness and high NPP
θ2Intercept 2.52 environments have the highest predicted population densities. In both
θ2NPP 0.25 Panels C and D, below a threshold of 0.25 (1330 g/m/yr), high species
Model Fit richness increases population density at a given level of NPP, but above
N 529 this threshold, high species richness decreases population density at a
AIC 1964.47
given value of NPP. These results are consistent with our model and
Log Likelihood 973.23
hypothesis that high species richness diminishes the marginal benefits of
NPP and a greater pathogen load increases the marginal costs of high
population density in high NPP settings, actually resulting in a decline in
population density.
Fig. 4A provides an additional way to view the predicted effects of
NPP and species richness on population density by holding productive
technology equal. Two observations: First, the slope of the relationship
between NPP and population density decreases from hunter-gather to
agricultural to industrial societies. This means that a one unit increase in
NPP results in a less steep increase in population density among indus­
trial and agricultural societies than among hunter-gatherer societies.
The implication is that NPP has a positive fixed effect across all pro­
ductive technology groups, but industrial societies and agricultural so­
cieties, in a sense, are less constrained by NPP than hunter-gatherer
societies. Second, note that below a NPP threshold of 0.25 (1330 g/m/
yr), groups who live in high species richness environments fall above
each respective regression line. Conversely, above this threshold, groups
who live in high species richness environments fall below each respec­
tive regression line. This is consistent with our model and the results
presented in Fig. 3. The effect of NPP is modified by species richness and
pathogen stress, with high species richness contributing to declines in
Fig. 3. Three way effect plot of the interaction effects of NPP, species richness population density at high levels of NPP.
and pathogens on population density. The plot holds pathogen load equal in a Fig. 4B illustrates NPP on the x-axis, predicted population density on
series of panels (A,B,C,D) and allows species richness to vary within each panel the y-axis and contours that represent levels of pathogen load. Observe
while graphing the relationship between NPP and population density. Hori­ an interesting and complex pattern. Between standardized NPP values of
zontal lines indicate inflection points of NPP where each linear curve meets in a 1.2 and 0.9 (300 and 1650 g=m2 r), holding productive technology
given plot. All ecological variables are standardized. A-Low pathogen load, one
equal, groups who live in high pathogen load environments actually
standard deviation below the mean pathogen value; B-Mean pathogen load; C-
have higher population densities than groups who live in lower path­
one standard deviation above the mean pathogen load; D 1.5 standard de­
viations above the mean pathogen load. In each panel, purple curve ¼ 1.5 ogen load environments. One can see this pattern by noting that the red
standard deviations above mean species richness; blue curves ¼ 1 standard and dark orange data points fall above each respective regression line
deviation above mean species richness; green curves-mean species richness; and within the NPP window of 1.2 to 0.9. Consistent with Fig. 3, at mod­
red curves ¼ 1 standard deviation below mean species richness. (For inter­ erate levels of NPP, the benefits of higher species richness seem to
pretation of the references to color in this figure legend, the reader is referred to outweigh the negative effects of higher pathogen load. Conversely,
the Web version of this article.) above the NPP threshold of 0.9, higher pathogen load societies typically
fall below each respective regression line. Interestingly, below the NPP
threshold of 1.2 (very low productivity environments), societies
among agricultural and industrial societies with high pathogen loads

6
J. Freeman et al. Journal of Archaeological Science 120 (2020) 105168

human population densities occur in setting with high NPP, moderate

levels of species richness and moderate to low pathogen loads. At lower
levels of NPP, higher species richness increases population density, and
at high levels of NPP, higher levels of species richness lead to lower
population densities. This may occur because of the combined effects of
steeper diminishing returns from more NPP in high species richness
environments and the increasing costs of higher population density due
to the abundance of pathogens in such environments.
However, not all of the patterns that we observed were consistent
with our model. Most importantly, the slope of the fixed effect of NPP on
population density decreases from hunter-gatherers to agriculturalists to
industrial societies. This suggests that while NPP does lead to higher
population density, regardless of productive technology, NPP con­
straints population density less among agricultural and industrial soci­
eties than among hunter-gatherers. This could make sense. Agriculture
is, after all, an attempt to augment ‘wild’ NPP by controlling the
dispersal of plants and animals and the flow of water and nutrients to
increase productivity. Industrial economies depend upon fossil plant
and animal remains that further augment the energy available for pro­
duction and transportation purposes. In short, technological evolution
may not fully emancipate humans from the constraints of current levels
of NPP, but technology and infrastructure do weaken the constraints of
NPP on population density.

4.1. Insights for paleoclimate–paleo-population analysis

The motivation for our work comes from recent research that com­
pares radiocarbon time-series across different climate, ecosystem, and
technological contexts. Much of this recent work has focused on how
prehistoric human populations were impacted by climate changes.
Climate changes do not directly impact human populations, but rather,
indirectly impact human populations through their effects on different
Fig. 4. Depiction of NPP on the x-axis, predicted population density from the ecosystem attributes. For research on human paleodemography across
TimeRS regression model (Eq. (3)) on the y-axis, and contours that represent different climatic contexts to advance, it is important to build middle
level of species richness in graph A and pathogen load in graph B. The color of range theory for how different ecosystem attributes affect human pop­
the points also indicates the level of species richness (A) or pathogen load (B)
ulations. We must be able to predict how paleoclimate changes might
for a particular society, and the regression lines are the best fit for each
have differently impacted the ecosystem attributes that then affected
respective productive technology group (hunter-gatherer, agricultural, indus­
trial). (For interpretation of the references to color in this figure legend, the
human populations throughout the Holocene. We can use the results of
reader is referred to the Web version of this article.) our study to create more informed hypotheses about when climate
variables should and should not associate with estimated changes in
paleo-population density. To illustrate, we provide two brief examples.
clearly fall below the regression line, while high pathogen load cases are
First, many archaeologists use large data sets of radiocarbon to
not seen among hunter-gatherers. The higher population densities
develop summed probability distributions or kernel density estimates of
generated by more productive technologies in very low productivity
paleo-population density over time. Once these proxies are constructed,
environments may also carry a pathogen penalty in such environments.
researchers often attempt to identify associations between paleoeco­
logical or paleoclimatological records and such paleo-population esti­
4. Discussion
mates. This type of research takes place without the context provided by
the finding of our paper: changes in productive technology have a larger
The goal of this paper has been to make two interrelated contribu­
effect than ecosystem attributes on population density and ecological
tions to the science of archaeology and human ecology. First, we have
factors that affect population density interact with each other. These
built and evaluated a graphic model that may explain global scale
results allow us to develop a set of hypotheses about the strength of
variation in human population densities. Following previous research,
association between paleoclimate/ecological and paleo-population
the model allows us to hypothesize that human population density
density fluctuation in any given region.
varies between three well defined productive technology groups
(hunter-gatherer, agricultural, and industrial) and, within each respec­
(1) If productive technology changes over the period of study in
tive group, net primary productivity (NPP), species richness and path­
question (e.g., the sequence displays the process of increasing use
ogen load interact to affect global scale variation in human population
of domesticated plants), then we should expect population den­
density.
sity to increase for most time-steps in a series from t to tþ1 to t þ
Our results partly support the model presented in Fig. 1. In partic­
i, just as a function of increasing productive capacity. This should
ular, hunter-gatherers have lower population densities than agricultural
weaken the cross association between fluctuation in paleo­
societies, and agricultural societies have lower population densities than
climatological (as well as paleoecological) variables and a given
industrial societies. Though there is some overlap in population den­
paleo-population estimate. In short, social-technological changes
sities, societies do form distinct productive technology groups. Within
that increase the carrying capacity of a system should result in
each productive technology group, higher NPP results in higher popu­
weak cross association between paleoclimate and paleo-
lation densities; however, species richness and pathogen load modify the
population time-series. Thus, weak associations that do not
relationship between NPP and human population density. The highest
meet an arbitrary level of significance should be expected–not

7
J. Freeman et al.
interpreted as a negative result–when a system is undergoing
increased productive capacity. Such lack of association is a
consequence of social-ecological system dynamics.
(2) The association should be strongest between paleoclimate
records and paleo-population estimates when social-
technological systems are near equilibrium (change very lit­
tle over some interval of a time-series). In this case, one
should choose carefully their paleoclimate proxies and take
into consideration that NPP is likely the main driver of pop­
ulation density, but this relationship is modified in complex
ways by species richness and pathogen stress.
In sum, to more productively investigate associations between
paleoclimate and paleo-population time-series: first, split the paleo-
population time-series into social-technological phases of productivity
expansion, stasis and perhaps contraction. Next, expect the strongest
associations between paleoclimate records and paleo-population records
during periods of relative social-technological stasis. Finally, consider
using multiple paleoclimate/ecological proxies to attempt to capture
interactions between NPP, species richness, and pathogen load.
Obtaining these measures from paleoenvironmental records will require
statistical techniques and modeling, such as rarefaction analysis (Birks
and Line, 1992; Odgaard, 1999) and biomization methods (Prentice
et al., 1996).
Second, an emerging issue in the interpretation of radiocarbon time-
series is the extent to which human population densities respond to
changes in climate over large geographic areas. For example, Robinson
et al., 2019 have recently developed a method that reconstructs paleo­
demography from radiocarbon time series according to different pale­
oclimate zones developed from downscaled and transient paleoclimate
models over the whole western US. This method attempts to move
beyond approaches that reconstruct paleodemography according to
arbitrary modern or cultural historical criteria, in favor of paleodemo­
graphic reconstructions that consider the climatic contexts and changes
in which different prehistoric populations lived. This new method for
reconstructing paleodemography according to transient paleoclimate
zones promises to build a more uniformly comparative approach for
research on the impacts of climate change on human populations.
However, this methodology still does not allow the ranking of climate
zones as more or less productive for human populations. We have a good
sense of how climate impacts ecosystems, and our work builds the link
between ecosystems and human population densities. One can, thus, use
our results to rank climate zones in terms of their expected ecological
attributes and the distance of these attributes from the ecological space
with the highest population densities observed in the ethnographic re­
cord, regardless of social-technological organization. This would allow
researchers to observe how climate-ecological space on a landscape
fills-up with people. The zones modeled as most close to the highest
population density ecosystems observed ethnographically should fill-up
with populations first, leap frogging less suitable zones, and so on.
5. Conclusion
Archaeologists and paleoecologists have long collaborated to inves­
tigate the effects of climate on human societies and populations (e.g.,
Barton et al., 2018; Naudinot and Kelly, 2017; Kelly et al., 2013; Dalfes
et al., 2013; Van de Noort, 2011; Kerr et al., 2009; McIntosh et al., 2000;
Dean, 1994). The ubiquity of radiocarbon data for Holocene archae­
ology means that radiocarbon time series provide the most widespread
data for research on the impacts of paleoclimate changes on Holocene
human populations. However, one of the challenges associated with
studying the effects of climate on human populations is that climate
indirectly affects human populations through the direct effects of tem­
perature and water on the productivity of resources. Moreover, social
institutions and technologies also change the potential density of re­
sources in an environment. Hence, interpreting the effects of climate on
8
Journal of Archaeological Science 120 (2020) 105168
changes in human population densities requires understanding the in­
teractions between climate, productivity, technology and population
density. As Tallavaara et al. (2018) argue, research into the relationships
between ecological attributes, which are controlled to some degree by
climate, and human population densities provides a critical frame of
reference for interpreting changes in human population densities over
time. Our study illustrates a remarkably consistent relationship between
human population density and ecological attributes at a global scale.
Future work that uses radiocarbon time series to investigate the impacts
of paleoclimate change on Holocene human populations must consider
the relative impacts of ecosystem attributes and productive technology
in the specific populations under investigation. These considerations
will improve the ability of researchers to investigate how climate
changes impact human populations, and provide a foundation for
comparative studies of human population ecology using
paleo-population proxies.
Declaration of competing interest
The authors have no conflicts of interest to declare.
Acknowledgments
This study was undertaken by PEOPLE 3000, a working group of the
Past Global Changes (PAGES) project, which in turn received support
from the Swiss Academy of Sciences and the Chinese Academy of Sci­
ences.The authors also wish to thank Miikka Tallavaara and two anon­
ymous reviewers for helpful comments on an earlier draft of the paper.
Appendix A. Supplementary data
Supplementary data to this article can be found online at https://doi.
org/10.1016/j.jas.2020.105168.
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