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The biogeography of human diversity in life history strategy.

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Evolutionary Behavioral Sciences
The Biogeography of Human Diversity in Life History
Strategy
Aurelio José Figueredo, Steven C. Hertler, and Mateo Peñaherrera-Aguirre
Online First Publication, April 20, 2020. http://dx.doi.org/10.1037/ebs0000198

CITATION
Figueredo, A. J., Hertler, S. C., & Peñaherrera-Aguirre, M. (2020, April 20). The Biogeography of
Human Diversity in Life History Strategy . Evolutionary Behavioral Sciences. Advance online
publication. http://dx.doi.org/10.1037/ebs0000198
 Evolutionary Behavioral Sciences
© 2020 American Psychological Association 2020, Vol. 2, No. 999, 000
ISSN: 2330-2925 http://dx.doi.org/10.1037/ebs0000198

The Biogeography of Human Diversity in Life History Strategy

Aurelio José Figueredo Steven C. Hertler

University of Arizona College of New Rochelle

Mateo Peñaherrera-Aguirre
University of Arizona
This article is intended solely for the personal use of the individual user and is not to be disseminated broadly.
This document is copyrighted by the American Psychological Association or one of its allied publishers.

Controversial theories have been advanced (e.g., Rushton, 1985, 2000) relating “race”
to human life history strategy: (a) different human populations (“races”) evolved in
different physical and community ecologies; (b) these ecologies should at least partially
determine the selective pressures shaping the evolution of human life history strategies
in different parts of the world; ergo (c) different human populations (“races”) should be
associated with different modal life history strategies. Although the argument seems
plausible in its stark logical form, there were several limitations in operationalization:
(a) the traditional “Big Three Races” used (Caucasoid, Mongoloid, and Negroid) do not
correspond very closely to the five or six major population clusters identified by
modern human genetics; (b) these “races” are neither discrete nor mutually exclusive,
having many zones of overlap and interbreeding, making geographical boundaries
fuzzy and imprecise; (c) fixing the “race” issue will still not directly address the
fundamental premise of the theory that human life history strategy is largely determined
by ecological factors. We therefore divided a sample of 141 national polities into
zoogeographical regions instead of conventionally defined “races.” We only used
regions for this analysis that were still inhabited mostly by the aboriginal populations
that existed there prior to the 15th century AD. Although obtained by different
procedures than those used originally by Rushton, these produced results that were
surprisingly convergent with the basic premise underlying the original hypotheses.

Public Significance Statement

This study found that zoogeographic regions explain much of the observed diversity
among national polities in several basic dimensions of the physical and community
ecology as well as in human life history strategies.

Keywords: life history strategy, zoogeographical regions, human biodiversity, races,

genetic clusters

Differential-K Theory (Rushton, 1985, 2000)
predicts that different modal life history strate-
gies should have evolved in the different ances-
X Aurelio José Figueredo, Department of Psychology,
University of Arizona; Steven C. Hertler, Department of
Psychology, College of New Rochelle; Mateo Peñaherrera-
Aguirre, Department of Psychology, University of Arizona.
Correspondence concerning this article should be addressed
to Aurelio José Figueredo, Department of Psychology, Uni-
versity of Arizona, 1503 East University Boulevard, P.O. Box
210068, Tucson, AZ 85721. E-mail: ajf@email.arizona.edu
1
tral ecologies by the major continental “races”,
by virtue of the varying selective pressures ex-
erted in different regions of the world by factors
such as climate. Differential-K Theory was thus
intended primarily as a test of evolutionary eco-
logical theory and only secondarily as a descrip-
tive theory of race differences; “race” was
merely used as a convenient proxy for ancestral
ecological differences that were hypothesized
as truly causal (J. P. Rushton, personal commu-
nication, November 11, 2011) and not as the
original intellectual focus of the hypothesis. The
theory was found to have a rough qualitative fit
 2 FIGUEREDO, HERTLER, AND PEÑAHERRERA-AGUIRRE
to the data, in spite of many anomalies in the
details, and this incomplete correspondence was
probably attributable to the inadequacy of the
conventional tripartite racial categorization, in
terms of which the most of the primary data
utilized had already been collected, with respect
to the results of modern molecular genetics in
identifying five to seven discriminable genetic
clusters in the human population that do not fit
very neatly into the traditional categorization.
This article is intended solely for the personal use of the individual user and is not to be disseminated broadly.
Most of the tests performed were secondary
This document is copyrighted by the American Psychological Association or one of its allied publishers.
data analyses, so it was not possible to address
the fundamental taxonomic problems.
Nevertheless, a signal was identifiable
through all the noise that reproduced the same
rank order of life history speed among the major
continental races across a wide variety of indi-
cator variables. This suggests that a meaningful
pattern might be found in the human biodiver-
sity of life history strategy if the various meth-
odological problems that plagued the original
work could somehow be circumvented. The
present study attempts to do just that by entirely
avoiding the use of “race”, however defined, as
a proxy for ancestral ecology and referring di-
rectly to the zoogeographical regions of origin
of contemporary human populations as the pri-
mary predictive factor. This has the benefit of
constituting a more direct test of the original
evolutionary ecological hypothesis and remain-
ing safely agnostic on the politically and scien-
tifically controversial topic of racial categoriza-
tions. Many critics have made valid points
against some of Rushton’s methods but at the
cost of missing the central point of the entire
scientific exercise. The present study seeks to
refocus us on the original objectives of Differ-
ential-K Theory as an evolutionary ecological
prediction. Further, the present study avoids
definitional distractions by relying on well-
accepted biodemographic indicators of life his-
tory strategy, such as fertility, longevity, and
mortality, and does not venture into the various
psychosocial indicators to which Rushton ex-
tended the logic of life history theory
(Figueredo, Cabeza de Baca, & Woodley,
2013).
It is also important to note that the described
rankings of fast-to-slow life history speeds do
not in any way imply some kind of normative
hierarchy or bogus evolutionary progression
from more primitive to more advanced. Evolu-
tionary ecological thinking would instead pre-
dict that the evolved life history speeds within
each zoogeographical region are an adaptation
to that specific ecology and thus the most ob-
jectively viable therein, by virtue of natural
selection, independent of any cultural value
judgments that some might misguidedly make.
Thus, we consider it self-evident that no single
life history strategy is universally “better” or
more adaptive than another but instead is con-
tingent in fitness upon the nature of the ambient
ecology in which it has evolved. No group or
individual should therefore take any umbrage at
these quantitative rankings as if they repre-
sented some kind of scala naturae of subjective
preference. Anyone who does has entirely
missed the point of evolutionary ecological the-
ory.
Race as Red Herring
Starting in the 1980s, John Phillipe Rushton
analyzed human biodiversity through the lens of
life history evolution, a longstanding biological
theory originally postulated to explain cross-
species variation in longevity, pace of living,
and the allocation of bioenergetic resources ei-
ther to survival and somatic maintenance on one
hand or mating effort and reproductive output
on the other hand. Within the context of life
history evolution, Rushton was interested in hu-
man intergroup differences as they evolved
within geographically disparate ecologies. For
instance, Rushton was an early advocate of
what became known as cold winters theory. He
additionally discussed migration out of Africa
into Eurasia as imposing cognitively demanding
environmental exigencies. The seasonal varia-
tion of temperate zones into which humans mi-
grated, Rushton theorized, was predictable,
which contrasts with the “sudden droughts and
devastating viral, bacterial, and parasitic dis-
eases” common to Africa (Rushton, 2000, p.
231). To demonstrate such variation required
data points spanning a host of life history traits
from populations disparately distributed across
the world. Naturally, Rushton therefore looked
to mine previously assembled data, much of
which was found in past anthropological studies
and in other databases employing three broad
racial categories: Mongoloids, Negroids, and
Caucasoids.
The capacity for global racial categorization
came after the age of exploration: Trading
 BIOGEOGRAPHY OF HUMAN LIFE HISTORY STRATEGY
routes, shipping lanes, crusades, and conquests
were among the many mechanisms of cross-
cultural contact between far flung human pop-
ulations. Swedish systematizer Carl Linnaeus
(1758), for instance, parsed between Homo
Americanus, Europeanus, Asiaticus, and Africa-
nus within the context of erecting the modern
foundations of taxonomy. Georges-Louis
Leclerc, Comte de Buffon, 18th century French
naturalist, published The Varieties of the Hu-
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man Species (1807), which more finely discrim-
This document is copyrighted by the American Psychological Association or one of its allied publishers.
inated between populations, including classifi-
cations such as Muscovite that corresponded
with national territories and subnational groups
such as the Ostiacks. Blumenbach (1825) later
subdivided humanity among Caucasian, Mon-
golian, Malayan, Ethiopian, and American rac-
es. Subsequently, Georges Cuvier (1833), born
in France during the latter half of the 18th
century, gave preference to a simpler tripartite
classification into Caucasian, Mongolian, and
Ethiopian. Thereafter in the mid-19th century,
Joseph Arthur, Comte de Gobineau, in an Essay
on the Inequality of the Human Races (1853)
also employed a tripartite scheme, labeling
these same three groupings black, white and
yellow races. William Z. Ripley’s The Races of
Europe, produced in 1899, delineated the Teu-
tonic, Mediterranean, and Alpine races, which
was followed by Madison Grant’s (1916) Cau-
casoid, Negroid, and Mongoloid classifications.
Interestingly, at least when viewing this small
sampling of theorists, lumpers predominated
over splitters as time went on, such that one sees
four and five divisions routinely being replaced
by three, which then, by intellectual inertia or
otherwise, continued on into early anthropolog-
ical formulations. Carleton Coon’s Caucasoid,
Congoid, Capoid, Mongoloid, and Australoid,
in this light, served as a bellwether for the
coming transition back to finer distinctions
(Coon & Hunt, 1966). Nevertheless, as Brian
Siegel discusses, a tripartite division was used
by early anthropologists that captured much
variation but awkwardly incorporated or other-
wise failed to incorporate disparate peoples. For
many decades of the 20th century, a small num-
ber of racial categories, often three, were used
by those collecting data on human racial diver-
sity, including anthropologists; with the same
often holding for research samples compiled by
the United States Census Bureau1 and the
United States Army.2 Of note is the observation
3
that categories only multiplied into finer group-
ings in later decades, after Rushton had com-
piled the research that undergirded his Differ-
ential K Theory, which was first published in
1985.
Intensive genetic surveys of the kind initi-
ated by Cavalli-Sforza (1997) have demon-
strated the existence of extant regional ge-
netic clusters, which are found across vast
geographic areas. Within these areas, large as
they are unto themselves, one finds gradients
of linear genetic variation. However, there is
a concomitant trend wherein geographic bar-
riers impart genetic discontinuities. It is from
mountains, rivers, seas, and oceans, among
other geographic barriers that add to distance
to restrict gene flow, that the realities of racial
categories rest. Without these barriers, one
would only have uninterrupted continuous
variation, as seen within regions.
With these barriers, one sees the justification
for categorization not into three broad groups,
but into the following seven groupings: Africa,
Europe, Middle East, Central/South Asia, East
Asia, Oceania, and America. Again, within each
of these clusters are clinal gradients of varia-
tion, but between them are sufficient genetic
discontinuities to support the continuance of
some degree of categorical classification
(Rosenberg et al., 2005). Linnaeus and other
early typologists came closest to this genetically
informed sevenfold classificatory scheme, but
even these were not sufficiently nuanced to
comport with the reality of variation within our
cosmopolitan species, spread as it is across a
vast a geographically varied earth.
In an age becoming increasingly racially con-
scious and politically correct, Rushton (1988),
following the “Big Three” categorical classifi-
cations, found himself reporting that the rate of
dizygotic twins differs by race, such that Mon-
goloids have one quarter the rate of twins as
compared to Negroids, with Caucasoids falling
in the middle. In addition to rates of twinning,
Rushton (1988) also describes race differences
in the rates of fetal growth, motor milestones,
sexual maturation, and age of menarche.
1
https://www.pewsocialtrends.org/interactives/multi
racial-timeline/.
2
https://www.npr.org/sections/thetwo-way/2014/11/07/
362243042/army-drops-use-of-term-negro-in-document.
 4 FIGUEREDO, HERTLER, AND PEÑAHERRERA-AGUIRRE
Rushton additionally described life history
driven differences in monogamy and mating
alongside differences in the onset of menarche
and fertility. Rushton himself, for all that he
remonstrated, never publicly explained that his
decision to use racial categories was in effect a
matter of convenience sampling, akin to how
many studies use conveniently accessible col-
lege students as stand ins for the population at
large. Though he never said as much in pub-
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lished form or in a public forum, Rushton was
This document is copyrighted by the American Psychological Association or one of its allied publishers.
perfectly aware that there may exist approxi-
mately five or six broad genetic clusters, de-
pending on the method used to identify them
(Cavalli-Sforza, 1997; Cavalli-Sforza, Menozzi,
& Piazza, 1994; Jobling, Hurles, & Tyler-
Smith, 2004; Rosenberg et al., 2002), many of
which are collapsed and combined across the
ill-informed tripartite racial divisions around
which most official data collection efforts had
been organized. From the outside, some critics
saw race categorization as the end of Rushton’s
reasoning and research, missing altogether that
his was a study of how ecological determinants,
as they varied geographically, created life his-
tory diversity among humans. The fateful,
though possibly unavoidable, decision to allow
racial categories to serve as proxies for the
evolved effects of continuously distributed eco-
logical variation blunted the thrust of his life
history research program and subsumed his later
years in bitter controversy.
Rushton, and to a lesser extent the application
of life history theory to human biodiversity,
became mired in a defense against critics.
Rushton’s energies were channeled in three
paths. First he protested against attacks on his
academic freedom, insisting that the “ultimate
aim of science is causally to explain the world
around us . . .” Rushton continued, writing of
the “chilling effect of self-imposed censorship,”
which pervaded “our own academic institutions,
and indeed even our own minds” (Rushton,
1999, p. 220). Second, he politicized attacks by
squarely identifying the ideological quarter
from which those attacks proceeded. “The po-
litical left,” Rushton wrote, “poses a more seri-
ous and immediate threat to the advance of
evolutionary science than does religious funda-
mentalism because fundamentalism has no
clout in research universities whatsoever.” Far
and away, however, the main tributary carrying
away Rushton’s energies was his defense of race,
which he may have felt pinioned into mounting. In
doing so, he entrenched, ostensibly confirming the
centrality of race in the eyes of critics. Rushton
(2000) asserted that “race is a biological concept”
(p. 96). He continued, writing:
Races are recognized by a combination of geographic,
ecological, and morphological factors and gene fre-
quencies of biochemical components. However, races
merge with each other through intermediate forms,
while members of one race can and do interbreed with
members of other races.
Thereafter we can intuit his tacit recognition
that, of course, racial categories did not amount
to immutable Platonic types:
Within each race, several varieties or minor races have
been proposed, although there is no agreed upon num-
ber. Mostly for political reasons, a majority of inves-
tigators avoid the use of the term race as much as
possible and use, for the major human races, the word
‘population’ and for the minor races, the phrase ‘ethnic
group’ (Rushton, 2000, p. 96).
Feeling himself uniquely exposed, Rushton
continued, lamenting that deliberate withhold-
ing of evidence had become commonplace
among evolutionary scientists writing about
race. Rushton held out the following authors as
examples: Stephen J. Gould, author of the re-
vised and expanded edition of The Mismeasure
of Man (1996), Jared Diamond, author of Guns,
Germs, and Steel (1997), and Christopher
Stringer, coauthor with Robin McKie of African
Exodus (1996). Rushton was not without his
champions, few and far between though they
were. Glayde Whitney, in reviewing Rushton’s
work, wrote of the “remarkable resistance to
racial science in our times,” which he compared
to the inquisition in Rome during the Renais-
sance. He thereafter likened Darwin, Galton,
and Rushton to Copernicus, Kepler, and Gali-
leo; all being men who generated ideas that
encountered stalwart resistance from their re-
spective quarters. L. S. Gottfredson defended
Rushton’s Race, Evolution, and Behavior: A
Life History Perspective even as it was deemed
by others “racist trash.” Yet, those defending
Rushton, like Rushton himself, followed the
lead of critics, and assembled battle lines across
the issue of race.
One exception is a review by Figueredo,
Cabeza de Baca, and Woodley (2013), which
pointedly regretted that Rushton’s “work on the
psychometrics of LH strategy was overshad-
owed by the controversy surrounding his pre-
 BIOGEOGRAPHY OF HUMAN LIFE HISTORY STRATEGY
diction of race differences in LH strategy as
adaptations to ancestral ecological conditions in
the different regions of the world.” With the
field of social biogeography only lately mitigat-
ing the loss, the spectacle of race long sup-
pressed a better understanding of ecological re-
gions, along with the various selective regimes
they impart and the evolved differences they are
capable of creating.
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Zoogeographic Regions as Alternative
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Approach
Since the 19th-century naturalists noticed
that florae and faunae varied across environ-
ments in different parts of the world (Sclater,
1858; Wallace, 1876), the interdisciplinary field
of biogeography emerged to address how re-
gional differences influenced the geographical
distribution of species and species assemblages
around the world (Morrone, 2015). This field
employed a hierarchical partitioning of geo-
graphical areas into a multilevel system com-
prised of realms or kingdoms, regions, domin-
ions, provinces, and districts (Morrone, 2015).
Each category captured unique ecological as-
pects, from broad general patterns at the level of
the kingdom to specific conditions correspond-
ing to each district. Within the field of bioge-
ography, one of its disciplines, zoogeography,
specialized in generating geographic maps ac-
cording to the variation and distribution of fau-
nae (Sclater, 1858; Wallace, 1876).
Hence, zoogeography provides a powerful
framework to test predictions regarding trait
evolution in response to regional variation.
Each region was hypothesized to pose unique
ecological challenges, from differences in tem-
perature, humidity, and altitude to the presence
of prey, predators, parasites, and competitors
(Wallace, 1876). Hence, species– environment
matching influenced the distribution of animals,
with geographical barriers limiting the migra-
tion of non-native species lacking the necessary
traits to establish into a new environment (Wal-
lace, 1876). These classifications capture cova-
riation among taxa in response to the local con-
ditions whereby, rather than viewing the local
ecology operating independently on each native
species, environmental pressures generate a cas-
cade of effects among entire symbiotic species
assemblages (Hertler, Figueredo, Peñaherrera-
Aguirre, Fernandes, Woodley of Menie, 2018).
5
While early physical atlases relied solely on
indicators of physical ecology such as latitude
and longitude (Johnston, 1850), subsequent
zoogeographic examinations considered species
variation as a critical marker for regional cate-
gorization (Sclater, 1858; Wallace, 1876).
Sclater, for example (1858), compared the pres-
ence of avian species across land areas, allow-
ing the author to identify six zoogeographic
regions. The Palearctic region included Europe,
North Asia, and North Africa. The Ethiopian
region comprised Arabia, Madagascar and the
remaining African areas south of the Sahara.
The Indian region involved Southern Asia and
the Indian Archipelago. The Australian region
contained Australia, New Zealand, and the Pa-
cific Isles. The Nearctic region included North
America and extend south to central Mexico,
and the Neotropical ranged from southern Mex-
ico to the southern areas of South America
(Sclater, 1858).
Sclater’s contributions influenced other natu-
ralists who proceeded to examine whether this
classification also extended to other clades.
While works such as Günther’s zoogeographic
reconstructions are worth mentioning (1858), it
was Alfred Wallace (1876) who, by adopting
Sclater’s classification, was able to generalize
this system to multiple animal families. Wallace
(1876) also described the presence of four sub-
regions within each major region, providing ad-
ditional information on the degree of intrare-
gional variation, as well as the presence of
transition zones between these areas.
Though over the years some critics have chal-
lenges some of the boundaries specified by
Sclater-Wallace system, recent examinations
support their predictions. Morrone (2015), for
example, reviewed the biogeographic, zoogeo-
graphic, and phytogeographic literature pub-
lished between the 1850s to the 2010s. The
author’s hierarchical classification consisted of
three kingdoms: the Holarctic, the Holotropic,
and the Austral. The Holarctic kingdom con-
tained Europe, North Africa, North Asia, North
America, and Greenland (i.e. Nearctic and Pale-
arctic). The Holotropic kingdom included the
Neotropic, the Ethiopian, and the Oriental re-
gions. Morrone also concluded the presence of
four regions comprising the Austral kingdom:
Andean, Cape, Australian, and the Antarctic.
The Andean region extends from the Andes
highlands the southern areas of South America
 6 FIGUEREDO, HERTLER, AND PEÑAHERRERA-AGUIRRE
(Morrone, 2015). The Cape region includes the
southern parts of South Africa (Morrone, 2015).
New Zealand, New Guinea, New Caledonia,
and Australia comprise the Australian region
(Sclater, 1858; Morrone, 2015). Finally, the last
area corresponded to Antarctica (Morrone,
2015). Like Wallace (1876), Morrone (2015)
also suggested the presence of transition zones.
The Mexican zone extends from southern North
America to central Meso America and joins the
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Nearctic with the Neotropical region. The Sa-
This document is copyrighted by the American Psychological Association or one of its allied publishers.
hara-Arabian zone corresponding to the Ara-
bian Peninsula and parts of North Africa di-
vided the Palearctic with the Ethiopian region.
The Indo-Malayan region acted as a border be-
tween the Australian and the Oriental region.
The Chinese zone divides the Oriental from the
Palearctic region.
In tandem with these publications, phylogeog-
raphers have also examined species’ distribution
after accounting for phylogenic effects. Holt et al.
(2013) proposed a zoogeographic classification
following the distribution of over 21,000 species,
accounting for the phylogenetic proximity be-
tween the species in the dataset. The analyses
revealed 11 zoogeographic realms, which could,
in turn, be subcategorized into 20 regions. The
classification included the Palearctic, the Sino-
Japanese, the Saharo-Arabian, the Afrotropical,
the Madagascan, the Oriental, the Oceanian, the
Australian, the Nearctic, the Panamanian, and the
Neotropical realms.
While the work of Holt et al., supported
Sclater’s and Wallace’s predictions, the analy-
ses also differed in some respects. For instance,
The Saharo-Arabian realm extended beyond
North Africa and acted as an intermediate zone
between the Afrotropical and the Sino-Japanese
realms (Holt et al., 2013). Similarly, the authors
discovered that insular regions east of Borneo
and Bali, rather than being part of the Australian
biogeographical region, clustered with other is-
lands in the Oceanian realm. Although Holt and
colleagues estimated the presence of 11 realms,
the classification of zoogeographic regions var-
ied depending on the taxa taken into consider-
ation. Hence, while analyses restricted to mam-
malian data detected 34 regions, examinations
based on the distribution of either avian or am-
phibian taxa discovered fewer areas (Holt et al.,
2013).
As a species, behavioral modern humans
were characterized for their widespread coloni-
zation on numerous environments around the
globe. This cosmopolitan status exposed human
communities to an array of unique environmen-
tal pressures leading to trait variation (Cochran
& Harpending, 2009). Social biogeography, a
branch of human sociobiology, emerged as a
discipline destined to examine the influence of
physical and community ecology (e.g., mean
annual temperature, mean annual precipitation,
altitude, latitude and longitude, parasite burden)
on interpopulation differences in life history as
well as in social, political, and economic out-
comes (e.g., Figueredo et al., 2017; Peñaher-
rera-Aguirre et al., 2019). Since biogeographic
and zoogeographic classifications are predicted
to capture differences in physical ecology, these
regions should also adequately detect variations
between human populations. Similarly, since
each region features a unique interplay of sym-
biotic interactions between species assem-
blages, zoogeographic regions are predicted to
detect differences in parasite diversity.
It could be argued that differences between
human populations are exclusively the product
of human behavioral flexibility. However,
rather than considering these abilities as con-
trary to physical and community ecology, social
biogeography predicts cultural and cognitive
ecology react to their respective environmental
conditions, often increasing underlying popula-
tion differences. Moreover, it is worth noting
that while behaviorally modern humans are
characterized for their reliance on cultural niche
construction (Odling-Smee, Laland, & Feld-
man, 2013), several aspects of physical ecology
are less susceptible to human interference, such
as altitude, temperature, and precipitation, ex-
cept for the construction of shelters. Current
biogeographic examinations have demonstrated
the additive and nonadditive influence of vari-
ous dimensions of physical, community, and
cognitive ecology on human variation (e.g.,
Figueredo et al., 2017; Peñaherrera-Aguirre et
al., 2019).
The Present Study
We address the various concerns regarding
race-based schemes of life history variation
by using the updated zoogeographical regions
(Holt et al., 2013) based on data from over
21,000 species of terrestrial vertebrate to es-
tablish ecologically informed geographical
 BIOGEOGRAPHY OF HUMAN LIFE HISTORY STRATEGY
areas within which humans (and other spe-
cies) can be expected to evolve and develop
different life history adaptations. By dividing
modern humans roughly into such regional
ecotypes, instead of conventionally defined
“races”, we can more directly test the basic
premise of Rushton’s hypothesis while avoid-
ing much of the unnecessary controversies
surrounding the definitions and utilities of
such historically received but antiquated ra-
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cial concepts.
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Method
Sample
National polities. One hundred forty-one
recognized national polities located in Asia, Af-
rica, and Europe were selected for study based
on the availability of sufficiently complete in-
formation on all the variables modeled. Na-
tional polities in the Americas and Australasia
were not sampled as they had recently experi-
enced massive invasions and “colonizations”
from other zoogeographic zones by human and
nonhuman animals and plants. Such invasions
were deemed to have disrupted the evolutionary
relations among physical, community, and so-
cial– ecological parameters that would be other-
wise expected by life history theory, producing
a certain degree of mismatch between the pop-
ulation phenotypic characteristics and their cur-
rent local ecologies. For example, the skin pig-
mentation of European Americans currently
living in desert regions (such as the American
Southwest) remains extremely low, making
them vulnerable to extremely high melanoma
rates as compared to aboriginal peoples (Cen-
ters for Disease Control and Prevention, 2016).
In contrast, the current genetic structures of
indigenous populations in Africa, Asia, and Eu-
rope have generally remained quite similar to
those existing several millennia ago in the same
or neighboring regions (e.g., Basu et al., 2003;
Hanihara, 1991; Leslie et al., 2015; Martínez-
Cruz et al., 2012; Piazza, Cappello, Olivetti, &
Rendine, 1988; Qi et al., 2013), better preserv-
ing the evolutionarily predicted ecological rela-
tions matching phenotypes to their current en-
vironments. Thus, we only used regions for this
analysis that were still inhabited mostly by the
aboriginal populations that existed there prior to
the 15th century AD.
7
Zoogeographic regions. The selected na-
tional polities were then assigned to the updated
zoogeographical regions published by Holt and
colleagues (2013), based on data from over 21
thousand species of terrestrial vertebrate, to es-
tablish ecologically informed geographical ar-
eas within which humans and other species can
be expected to have evolved and developed
different life history adaptations. In certain
cases, these assignments were decided based on
maximum overlap between the areas delimited
by political and zoogeographic boundaries. The
zoogeographical regions selected for analysis
were therefore as follows: Afro-Tropical (AT),
Madagascan, (MA), Oceanian (OC), Orienta-
l(OR), Palearctic (PA), Saharo-Arabian (SA),
and Sino-Japanese (SJ).
Measures
Physical ecology. The annual mean precip-
itations and temperatures for the selected na-
tional polities were obtained from Climate
Charts (2015), with some of the missing data
imputed for mean annual temperatures obtained
from CantyMedia (2015).
Slow life history strategy. This common
factor was indicated by a unit-weighted com-
posite of the following four biodemographic
indicators: (a) infant mortality (Central Intelli-
gence Agency, 2015) as an indicator of age-
specific environmental harshness (see Ellis,
Figueredo, Brumbach, & Schlomer, 2009); (b)
life expectancy (Central Intelligence Agency,
2015) as an inverse indicator of morbidity and
mortality partially reflecting bioenergetic in-
vestments in somatic effort (e.g., Charnov,
1991; Figueredo, Vásquez, Brumbach, &
Schneider, 2004; Figueredo, Vásquez,
Brumbach, & Schneider, 2007; van Schaik &
Isler, 2012; Williams, 1957); (c) birth rate (Cen-
tral Intelligence Agency, 2015) as a measure of
fertility and thus indirectly of the tradeoff be-
tween mating and parental effort central to life
history strategy (MacArthur & Wilson, 1967);
(d) teenage birth rate (or adolescent fertility
rate; World Bank, 2019), operationalized as the
number of births per 1,000 women aged 15–19,
representing early investments in mating effort
and thus related to life history speed (van
Schaik & Isler, 2012; Stearns, 1992).
Parasite burden. Both infant mortality and
life expectancy were statistically adjusted for
 8 FIGUEREDO, HERTLER, AND PEÑAHERRERA-AGUIRRE
the specific effects of parasite burden prior to
aggregation; we obtained per capita disability
adjusted life years (DALY) from the World
Health Organization (2015) and historical infec-
tious disease prevalences from Murray and
Schaller (2010) as indicators of human parasite
burden, using a logarithmic transformation of
their unit-weighted composite for each national
polity.
Population. Populations for each national
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polity within the sample were obtained from
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the United Nations Department of Economic
and Social Affairs, Population Division’s
(2011) World Population Prospects: The
2010 Revision, supplemented by official na-
tional statistics published in the United Na-
tions (2010) Demographic Yearbook, 2009-
2010, and data compiled by the Secretariat of
the Pacific Community (2010) SPC Statistics
and Demography Programme. Population
densities for each national polity within the
sample were obtained from the World Bank
Web Site (2019).
Statistical Analyses
All statistical analyses were performed using
SAS 9.4; the unit-weighted factor structures
were estimated using PROC CORR, and the
analyses of variance were done using PROC
GLM, with each national polity weighted by its
population. Multilevel models (MLMs) were
performed using PROC MIXED.
Results
The Measurement Model
The theoretically expected positive manifold
was found among the four biodemographic in-
dicators of life history strategy. Both infant
Table 1
Bivariate Correlations Among Slow Life History Strategy Indicators
Teen pregnancy
Teen pregnancy 1.00
Birth rate .88ⴱ
Infant mortality .65ⴱ
Life expectancy ⫺.70ⴱ
Note. Adjusted for natural logarithm of parasite burden (LPB) above the diagonal; not
adjusted for natural logarithm of parasite burden (LPB) below the diagonal.
ⴱ
p ⬍ .0001.
mortality and life expectancy were statistically
adjusted by exporting the regression residuals
for the logarithmic effects of human parasite
burden. This was done to control statistically for
any cross-cultural variation with respect to his-
torically recent advances in medical technology
and public health interventions that were not
present in the ancestral environments and to
which current human populations have presum-
ably not had very much time to adapt geneti-
cally by selection. This statistical adjustment
had relatively negligible effects upon the mag-
nitudes of the parameters within the correlation
matrix, as shown in Table 1.
Table 2 shows the unit-weighted factor load-
ings of each of the indicators from its latent
common factor, the slow life history composite
(see Table 2), operationalized as part-whole
correlations, each serving as convergent validity
coefficients among the indicators (Gorsuch,
1983). These results are shown in the factor
structure table for both the adjusted and nonad-
justed forms of the variables. Again, the adjust-
ment had relatively negligible effects upon the
magnitudes of the parameters, but we decided to
use the more conservative (adjusted) estimates
in all subsequent analyses.
The Structural Models
Four basic parameters of the physical and
community ecology were used to examine the
predictive validity of the zoogeographic re-
gions: (a) average annual temperatures, (b) av-
erage annual precipitation, (c) the natural loga-
rithms of human parasite burden, and (d) the
population densities. While this list of relevant
parameters is by no means exhaustive, it sum-
marizes many of the ecological selective pres-
sures that have been proposed to account for life
history evolution and thus for the diversity in
Birth rate Infant mortality Life expectancy
.88ⴱ .72ⴱ ⫺.76ⴱ
1.00 .83ⴱ ⫺.82ⴱ
.73ⴱ 1.00 ⫺.86ⴱ
⫺.71ⴱ ⫺.78ⴱ 1.00
 BIOGEOGRAPHY OF HUMAN LIFE HISTORY STRATEGY 9

Table 2 accounted for 39.0%, F(6, 133) ⫽ 14.17, p ⬍

Slow Life History (SLH) Factor .0001, of the variance across national polities;
Unit-Weighted Loadings for the natural logarithm of human parasite bur-
SLH (Not LPB- SLH (LPB- den, the assigned zoogeographic regions ac-
Adjusted) Adjusted) counted for 69.1%, F(6, 133) ⫽ 49.66, p ⬍
Teen pregnancy ⫺.92ⴱ ⫺.90ⴱ
.0001, of the variance across national polities;
Birth rate ⫺.95ⴱ ⫺.92ⴱ for the population density, the assigned zoogeo-
Infant mortality ⫺.88ⴱ ⫺.88ⴱ graphic regions accounted for 21.1%, F(6,
Life expectancy .91ⴱ .89ⴱ 133) ⫽ 5.92, p ⬍ .0001, of the variance across
Note. Not adjusted for natural logarithm of parasite bur- national polities. These preliminary analyses in-
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den (LPB) in column 1; adjusted for natural logarithm of dicated that the assigned zoogeographic regions
This document is copyrighted by the American Psychological Association or one of its allied publishers.

parasite burden (LPB) in column 2; bivariate correlation
between unit-weighted factor scores for LPB-adjusted SLH
and non-LPB-adjusted SLH is .98ⴱ.
ⴱ
p ⬍ .0001.
life history strategies among human populations
(e.g., Rushton, 1985, 2000). The bivariate cor-
relations among these ecological parameters as
well as with LPB-adjusted slow life history
strategy are shown in Table 3.
Five analyses of variance were performed to
determine how well the assigned zoogeographic
regions explained the variance among the na-
tional polities nested within them. The first four
of these were done mostly as “manipulation
checks” to see how well the zoogeographic re-
gions partitioned the cross-national variance in
basic parameters of physical and community
ecology; afterward, the fifth of these analyses of
variance was done on life history strategy itself
to test the main hypothesis of the present study.
All analyses of variance were weighted by the
population of each national polity.
For average annual temperature, the assigned
zoogeographic regions accounted for 86.5%,
F(6, 133) ⫽ 142.58, p ⬍ .0001, of the variance
across national polities; for average annual pre-
cipitation, the assigned zoogeographic regions
were doing a reasonably good job of partition-
ing the variance among national polities in these
basic dimensions of the physical and commu-
nity ecology.
Finally, for our unit-weighted factor scale of
slow life history strategy, the assigned zoogeo-
graphic regions accounted for 78.6%, F(6,
133) ⫽ 81.59, p ⬍ .0001, of the variance across
national polities; when tested individually,
dummy variable analyses also revealed that
each of the zoogeographical regions, excepting
only the Madagascan (MA) and Oceanian (OC),
was significantly different in life history strat-
egy from the ultimately ancestral Afro-Tropical
(AT) region, indicating subsequent adaptive ra-
diation in life history strategies. The distribution
of these means is presented graphically in Fig-
ure 1.
We also performed hierarchical general lin-
ear models to determine the relative incremental
validities of the four continuous ecological pa-
rameters with respect to the zoogeographical
regions. As with the previous analyses of vari-
ance, all hierarchical general linear models were
weighted by the population of each national
polity. When entering the four continuous eco-
logical parameters hierarchically prior to the
categorical zoogeographic regions, all four were

Table 3
Bivariate Correlations Among Slow Life History Strategy Predictors
Average annual Average annual Logarithm Population SLH (LPB-
temperature precipitation parasite burden density Adjusted)
Average annual temperature 1.00 .47ⴱ .52ⴱ .27ⴱ ⫺.54ⴱ
Average annual precipitation .47ⴱ 1.00 .16 .34ⴱ ⫺.08
Logarithm of parasite burden .52ⴱ .16 1.00 .16 ⫺.21ⴱ
Population density .27ⴱ .34ⴱ .16 1.00 .09
SLH (LPB-adjusted) ⫺.61ⴱ ⫺.11 ⫺.39ⴱ .06 1.00
Note. Adjusted for natural logarithm of parasite burden (LPB) above the diagonal; not adjusted for natural logarithm of
parasite burden (LPB) below the diagonal.
ⴱ
p ⬍ .05.
 10 FIGUEREDO, HERTLER, AND PEÑAHERRERA-AGUIRRE
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This document is copyrighted by the American Psychological Association or one of its allied publishers.
Figure 1. Distribution of slow life history across zoogeographical regions: AT ⫽ Afro-
Tropical; MA ⫽ Madagascan; OC ⫽ Oceanian; OR ⫽ Oriental; PA ⫽ Palearctic; SA ⫽
Saharo-Arabian; SJ ⫽ Sino-Japanese.
found statistically significant and collectively
accounted for 38.8%, F(4, 129) ⫽ 20.29, p ⬍
.0001, of the variance, leaving zoogeographic
regions accounting for the remaining 40.8%,
F(6, 129) ⫽ 42.86, p ⬍ .0001, of the systematic
variance in the model. When instead entering
zoogeographic regions hierarchically prior to
the four continuous ecological parameters, how-
ever, none of them were found statistically sig-
nificant. Collectively, they only raised the over-
all squared multiple correlation of the model to
79.6%, F(10, 129) ⫽ 50.19, p ⬍ .0001, of the
variance, only adding a paltry and statistically
nonsignificant 1% to the total. This indicated
that the zoogeographical regions did a good job
of summarizing the continuous ecological pa-
rameters of each biologically defined geo-
graphic area, precisely as they were intended to
do.
We believe that these results, although ob-
tained by procedures quite different from those
used originally by Rushton (1985) and subse-
quent works, nevertheless produced results that
were strikingly convergent with the basic prem-
ise underlying the original hypotheses. On the
other hand, these results are significantly differ-
ent from those obtainable with the way that the
hypothesis was originally operationalized. To
evaluate this assertion formally and quantita-
tively as an alternative hypothesis to ours, we
did our best to reconstruct the original tripartite
racial categories with our data on zoogeo-
graphic regions, understanding that this is only
a very rough fit. This approximate reconstruc-
tion required inappropriately lumping the in-
habitants of the Afro-Tropical and Madagascan
regions into the so-called “Negroid” category,
inappropriately lumping those of the Palearctic
and Saharo-Arabian regions into the so-called
“Caucasoid” category, and inappropriately
lumping those of the Oceanian, Oriental, and
Sino-Japanese regions into the so-called “Mon-
goloid” category. These reconstructions were
based on which category within the tripartite
“racial” classification the majority of the popu-
lation within each zoogeographic region could
be assigned. One can discern by simple visual
inspection of Figure 1 what such overinclusive
categorizations would do to the balance among
between-groups and within-groups variance,
but we went ahead and did the analysis anyway
to make this point more precisely. The differ-
ence between the relative explanatory power of
the alternative models, meaning the loss en-
 BIOGEOGRAPHY OF HUMAN LIFE HISTORY STRATEGY
tailed by using the reconstructed “racial” cate-
gories rather than the zoogeographic regions,
amounts to 16.5%, F(4, 133) ⫽ 25.79, p ⬍
.0001, of the variance across national polities,
which is a substantial and statistically signifi-
cant proportion of the total.
In addition, we ran some multilevel models
(MLMs) to evaluate the possible effects of any
spatially autogregressive effects by constructing
a simple ordinal variable representing the se-
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quential distance “Out-of-Africa” for each na-
This document is copyrighted by the American Psychological Association or one of its allied publishers.
tional polity, estimating the serial autocorrela-
tions of each national polity with each spatially
adjacent one within that ordered sequence. This
model is not designed to control statistically for
the effects of cultural diffusion, which would
presumably require estimating network autocor-
relations, but instead models the possible phy-
logenetic effects of common human origins and
subsequent outward migrations (see Hertler et
al., 2018). We limited the autocorrelational
analysis to this simplified sequential model be-
cause we strongly doubt that the highly herita-
ble biodemographic population parameters used
in the present analyses would be very highly
susceptible to hypothetically alternative (mean-
ing nonselective) processes such as cultural dif-
fusion across national boundaries.
MLM residuals were thus exported for LPB-
adjusted slow life history strategy and used for
subsequent general linear modeling. These
MLM residuals had been statistically adjusted
for the ordinal distance Out-of-Africa (OOA) as
well as of any single-lagged heterogeneous au-
toregressive serial dependencies (ARH1)
among successive (meaning spatially contigu-
ous) data prior to regression modeling, thus
circumventing this potential problem as a threat
to the validity of correlational analysis. Using
the studentized MLM residuals of OOA, the
assigned zoogeographic regions accounted for
62.6%, F(6, 133) ⫽ 37.11, p ⬍ .0001, of the
variance across national polities in OOA-
adjusted slow life history strategy. In contrast,
the reconstructed “racial” categories only ac-
counted for 16.8%, F(2, 137) ⫽ 13.79, p ⬍
.0001, of the variance across national polities in
OOA-adjusted slow life history strategy, which
once again constituted a statistically significant
loss of explanatory power, F(4, 133) ⫽ 40.76,
p ⬍ .0001, with respect to the zoogeographic
regions.
11
Discussion
The purpose of this study was to investigate
the original principle behind Rushton’s (1985,
2000) hypothesis regarding the ecology of hu-
man biodiversity in life history strategy. After
considering the various limitations of historical
race-based partitionings of human populations,
we decided to circumvent the problematical na-
ture of racial classifications entirely by instead
applying a classificatory scheme based on the
most recently updated (Holt et al., 2013) map-
ping of zoogeographical regions of origin, be-
lieving this idea to be the most faithful to the
original theoretical motivation behind
Rushton’s evolutionary theory of the diversifi-
cation of human life history strategies.
Having accomplished this reconceptualiza-
tion of the problem, we found that the great
preponderance of the variance among contem-
porary national polities in aggregate life history
speeds can be explained with reference to zoo-
geographical regions. This was done without
any reference to or use of racial classifications,
indicating empirical support for the originally
hypothesized causal principle of ancestral ecol-
ogy. The pattern of results matches quite closely
what one would expect from a purely ecological
theory of life history variation, as well as the
progression of ancestral human migrations “Out
of Africa”, without the many imprecisions in-
troduced by obsolete racial categorizations.
Nevertheless, statistically controlling for the re-
constructed “Out of Africa” sequence, as well
as the serial spatial autocorrelations possibly
produced by those successive population expan-
sions, produced a reduction of at most 16% in
the variance explained by the zoogeographical
regions, leaving nearly 63%. This suggests that:
(a) neither phylogenetic inertia (Huber, 1939;
but see Shanahan, 2011) nor “Isolation by Dis-
tance” (Ishida, 2009; but see Slatkin, 1993)
among the contiguous areas that were colonized
successively by human “Out of Africa” migra-
tions were sufficient to account for our reported
pattern of results in human life history biodi-
versity, and (b) the preponderance of the vari-
ance among contemporary national polities is
instead best explained by divergent evolution
produced as a result of differential ecological
selective pressures among zoogeographical re-
gions. It should be noted, however, that Thorn-
hill and Fincher (2013) have argued quite con-
 12 FIGUEREDO, HERTLER, AND PEÑAHERRERA-AGUIRRE
vincingly that, as contiguous geographic areas
tend to be similar in ecology, such statistical
adjustments might inadvertently absorb the ef-
fects of current continuities in evolutionarily
relevant ecological parameters among adjacent
ecosystems. Spatial contiguity might thus pro-
duce similarities among adjacent populations
that are attributable to current adaptations to
similar environments by means of continuing
selection, thus statistically adjusting for spatial
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autocorrelations might inappropriately attenuate
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our estimates of the magnitude of the effects of
ecological selective pressures.
We therefore offer this reanalysis as one way
to disentangle the science from the unnecessary
ideological baggage historically surrounding
human biogeographic variation in life history in
the hope that we can circumvent the intellectual
barriers that have needlessly plagued this area
of study in the past.
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Appendix

Table of Life History Indicator Traits by Country (Central Intelligence Agency, 2015;
World Bank, 2019)

Country Population Density Teen pregnancy Birth rate Infant mortality Life expectancy
Afghanistan 33397058 57 68.79 5.27 71.7 59.68
Albania 3227373 105 20.68 1.74 13.8 77.44
Algeria 36485828 18 10.42 2.91 22.4 74.32
Angola 20162517 25 154.47 6.25 104.1 51.46
Armenia 3108972 104 23.98 1.58 14.6 74.45
Austria 8428915 107 7.21 1.44 3.3 80.94
Azerbaijan 9421233 120 52.61 2.00 31.2 70.62
Bahrain 1359485 2017 13.49 2.10 6.3 76.41
Bangladesh 152408774 1240 84.41 2.25 35.3 70.86
Belarus 9527498 47 17.98 1.62 4.0 71.97
Belgium 10787788 377 5.11 1.79 3.5 80.39
Benin 9351838 102 88.15 4.93 68.5 59.12
Bhutan 750443 20 22.08 2.15 30.8 68.72
Bosnia and 3744235 65 10.43 1.28 5.9 76.12
Botswana 2053237 4 31.72 2.88 38.2 64.22
Brunei Darussalam 412892 81 10.94 1.91 8.1 78.25
Bulgaria 7397873 65 40.29 1.50 10.5 74.31
Burkina Faso 17481984 72 106.54 5.69 65.4 57.88
Burundi 8749387 435 27.40 6.12 59.5 55.79
Cabo Verde 505335 135 74.74 2.38 22.4 72.83
Cambodia 14478320 92 49.90 2.74 30.7 67.33
Cameroon 20468943 53 108.83 4.86 62.4 54.59

(Appendix continues)
 BIOGEOGRAPHY OF HUMAN LIFE HISTORY STRATEGY 15

Appendix (continued)
Country Population Density Teen pregnancy Birth rate Infant mortality Life expectancy
Central African Rep 4575586 7 105.79 4.45 97.7 49.11
Chad 11830573 12 164.52 6.37 90.2 50.78
China 1353600687 148 6.50 1.55 11.5 75.39
Comoros 773344 447 67.17 4.63 59.8 62.58
Congo 4233063 15 114.09 4.96 37.6 60.92
Cote d’Ivoire 20594615 79 133.39 5.12 72.8 50.86
Croatia 4387376 73 9.37 1.51 4.1 76.92
Cyprus 1129166 129 4.70 1.46 2.8 79.76
Czech Rep 10565678 138 10.21 1.45 3.2 78.08
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Democratic 69575394 37 125.24 6.20 80.5 57.85

This document is copyrighted by the American Psychological Association or one of its allied publishers.

Denmark 5592738 138 4.19 1.73 3.2 80.05

Djibouti 922708 41 19.42 3.33 58.9 61.30
Egypt 83958369 99 50.96 3.31 22.6 70.73
Equatorial 740471 47 157.85 5.01 74.3 56.89
Eritrea 5580862 32 53.49 4.44 37.0 62.64
Estonia 1339762 30 13.33 1.56 2.9 76.33
Ethiopia 86538534 109 64.86 4.64 46.2 62.79
Fiji 875822 48 43.73 2.62 20.0 69.74
Finland 5402627 18 6.90 1.80 2.2 80.63
France 63457777 122 8.80 2.01 3.5 81.97
Gabon 1563873 8 98.48 4.01 39.7 63.28
Gambia 1824777 225 81.94 5.78 50.1 59.78
Georgia 4304363 65 47.05 1.82 12.8 74.35
Germany 81990837 237 6.84 1.38 3.3 80.89
Ghana 25545939 131 67.64 4.24 47.0 60.98
Greece 11418878 83 7.49 1.35 3.9 80.63
Guinea 10480710 51 137.41 5.18 66.8 57.64
Guinea-Bissau 1579632 67 87.21 4.97 67.4 54.50
Hungary 9949589 108 19.81 1.34 5.4 75.06
Iceland 328290 4 7.32 2.04 1.7 82.92
India 1258350971 455 24.54 2.51 42.6 67.29
Indonesia 244769110 148 47.99 2.50 25.3 68.52
Iran 75611798 50 25.72 1.74 15.1 74.80
Iraq 33703068 89 79.80 4.64 28.7 68.95
Ireland 4579498 70 10.14 2.01 3.4 80.90
Israel 7694670 411 9.68 3.05 3.4 81.70
Italy 60964145 205 6.16 1.43 3.2 82.24
Japan 126434653 347 4.16 1.41 2.2 83.10
Jordan 6457260 112 23.29 3.51 16.8 73.75
Kazakhstan 16381297 7 28.42 2.62 16.0 69.61
Kenya 42749418 90 81.79 4.48 39.2 60.27
Kuwait 2891553 232 9.40 2.21 46.6 65.60
Kyrgyzstan 5448085 33 38.81 3.20 22.9 70.00
Lao PDR 6373934 31 63.29 3.14 55.4 65.25
Latvia 2234572 31 13.83 1.44 7.6 73.78
Lebanon 4291719 669 12.18 1.67 7.9 78.91
Lesotho 2216850 69 89.71 3.25 72.3 48.84
Liberia 4244684 50 128.76 4.87 59.4 60.20
Libya 6469497 4 5.73 2.54 12.9 71.65
Lithuania 3292454 45 11.39 1.60 4.5 73.86
Luxembourg 523362 250 5.44 1.57 1.6 81.39
Macedonia 2066785 83 16.57 1.50 6.5 75.03

(Appendix continues)
 16 FIGUEREDO, HERTLER, AND PEÑAHERRERA-AGUIRRE

Appendix (continued)
Country Population Density Teen pregnancy Birth rate Infant mortality Life expectancy
Madagascar 21928518 45 111.68 4.53 39.3 64.25
Malawi 15882815 192 141.01 5.32 50.2 60.05
Malaysia 29321798 96 13.36 1.97 6.6 74.42
Maldives 324313 1719 6.50 2.18 9.2 76.46
Mali 16318897 16 171.08 6.40 79.2 57.10
Malta 419212 1511 16.80 1.43 5.5 80.75
Mauritania 3622961 4 80.50 4.72 68.1 62.56
Mauritius 1313803 623 26.93 1.54 12.8 73.86
Moldova 3519266 124 22.70 1.27 14.3 70.77
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Mongolia 2844081 2 24.23 2.64 22.1 68.61

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Montenegro 632796 46 12.10 1.73 5.3 75.71

Morocco 32598536 81 31.68 2.55 26.4 73.36
Mozambique 24475186 38 138.95 5.47 63.6 54.21
Myanmar 48724387 82 29.04 2.28 43.2 65.43
Namibia 2364433 3 75.00 3.59 35.5 63.88
Nepal 31011137 196 62.08 2.38 33.1 68.82
Netherlands 16714228 511 4.08 1.72 3.5 81.10
Niger 16644339 18 194.01 7.64 61.6 60.07
Nigeria 166629383 215 109.27 5.76 76.2 52.11
North Korea 24553672 212 0.29 2.00 22.7 69.49
Norway 4960482 15 5.68 1.85 2.3 81.45
Oman 2904037 16 7.85 2.86 10.0 76.59
Pakistan 179951140 275 37.69 3.74 70.6 65.72
Papua New Guinea 7170112 19 53.44 3.87 48.0 62.30
Philippines 96471461 358 59.86 3.05 23.9 68.01
Poland 38317090 124 13.03 1.33 4.6 76.75
Portugal 10699333 112 9.91 1.28 3.1 80.37
Qatar 1938754 240 10.19 2.06 7.3 78.23
Romania 21387517 85 33.72 1.52 10.9 74.41
Russian Federation 142703181 9 22.53 1.70 9.3 70.37
Rwanda 11271786 499 26.78 4.14 37.1 62.80
Saudi Arabia 28705133 16 8.29 2.87 13.8 74.02
Senegal 13107945 82 74.87 5.16 44.1 65.32
Serbia 9846582 80 19.31 1.45 6.2 74.84
Sierra Leon 6126450 106 115.58 4.87 97.9 49.75
Singapore 5256278 7953 3.72 1.29 2.2 82.00
Slovakia 5480332 113 22.15 6.5 76.11
Slovenia 2040057 103 4.31 1.58 2.4 80.12
Solomon Islands 566481 23 47.83 4.10 25.5 67.51
Somalia 9797445 24 102.19 6.67 92.6 54.69
South Africa 50738255 48 44.42 2.41 36.8 56.10
South Korea 48588326 530 1.67 1.30 3.3 81.21
South Sudan 11062113 18 65.25 5.20 66.1 54.73
Spain 46771596 94 8.68 1.32 3.7 82.43
Sri Lanka 21223550 346 14.83 2.14 8.9 74.53
Sudan 45722083 24 67.20 4.49 51.1 62.83
Swaziland 1220408 66 78.53 3.41 51.5 48.85
Sweden 9495392 25 5.27 1.91 2.4 81.70
Switzerland 7733709 216 3.09 1.52 3.7 82.70
Syrian Arab Rep 21117690 92 39.53 3.03 12.6 70.79
Tajikistan 7078755 66 36.92 3.53 42.1 69.17
Tanzania 47656367 64 116.59 5.29 38.8 63.52

(Appendix continues)
 BIOGEOGRAPHY OF HUMAN LIFE HISTORY STRATEGY 17

Appendix (continued)
Country Population Density Teen pregnancy Birth rate Infant mortality Life expectancy
Thailand 69892142 136 51.79 1.53 11.6 74.07
Timor-Leste 1187194 85 45.62 5.30 49.3 67.80
Togo 6283092 145 89.62 4.73 56.5 58.55
Tunisia 10704948 74 7.64 2.20 13.6 74.00
Turkey 74508771 107 26.93 2.10 14.2 74.64
Turkmenistan 5169660 12 24.83 2.35 47.6 65.31
Uganda 35620977 213 110.53 5.96 42.5 57.10
Ukraine 44940268 77 24.66 1.53 9.1 70.94
United Arab Emirates 8105873 136 28.23 1.82 6.7 77.02
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United King 62798099 275 13.55 1.92 4.1 80.90

This document is copyrighted by the American Psychological Association or one of its allied publishers.

Uzbekistan 28077486 77 16.72 2.30 37.2 68.10

Viet Nam 89730274 308 29.04 1.96 18.8 75.32
Yemen 25569263 54 61.82 4.42 38.4 63.33
Zambia 13883577 23 85.97 5.51 49.0 58.36
Zimbabwe 13013678 37 105.82 4.63 55.7 12.64

Received July 15, 2019

Revision received February 4, 2020
Accepted February 14, 2020 䡲

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