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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
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.
Differential-K Theory (Rushton, 1985, 2000) tral ecologies by the major continental “races”,
predicts that different modal life history strate- by virtue of the varying selective pressures ex-
gies should have evolved in the different ances- 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-
X Aurelio José Figueredo, Department of Psychology, tive theory of race differences; “race” was
University of Arizona; Steven C. Hertler, Department of merely used as a convenient proxy for ancestral
Psychology, College of New Rochelle; Mateo Peñaherrera- ecological differences that were hypothesized
Aguirre, Department of Psychology, University of Arizona. as truly causal (J. P. Rushton, personal commu-
Correspondence concerning this article should be addressed
to Aurelio José Figueredo, Department of Psychology, Uni-
nication, November 11, 2011) and not as the
versity of Arizona, 1503 East University Boulevard, P.O. Box original intellectual focus of the hypothesis. The
210068, Tucson, AZ 85721. E-mail: ajf@email.arizona.edu theory was found to have a rough qualitative fit
1
2 FIGUEREDO, HERTLER, AND PEÑAHERRERA-AGUIRRE
to the data, in spite of many anomalies in the dict that the evolved life history speeds within
details, and this incomplete correspondence was each zoogeographical region are an adaptation
probably attributable to the inadequacy of the to that specific ecology and thus the most ob-
conventional tripartite racial categorization, in jectively viable therein, by virtue of natural
terms of which the most of the primary data selection, independent of any cultural value
utilized had already been collected, with respect judgments that some might misguidedly make.
to the results of modern molecular genetics in Thus, we consider it self-evident that no single
identifying five to seven discriminable genetic life history strategy is universally “better” or
clusters in the human population that do not fit more adaptive than another but instead is con-
very neatly into the traditional categorization. tingent in fitness upon the nature of the ambient
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 ecology in which it has evolved. No group or
This document is copyrighted by the American Psychological Association or one of its allied publishers.
data analyses, so it was not possible to address individual should therefore take any umbrage at
the fundamental taxonomic problems. these quantitative rankings as if they repre-
Nevertheless, a signal was identifiable sented some kind of scala naturae of subjective
through all the noise that reproduced the same preference. Anyone who does has entirely
rank order of life history speed among the major missed the point of evolutionary ecological the-
continental races across a wide variety of indi- ory.
cator variables. This suggests that a meaningful
pattern might be found in the human biodiver- Race as Red Herring
sity of life history strategy if the various meth-
odological problems that plagued the original Starting in the 1980s, John Phillipe Rushton
work could somehow be circumvented. The analyzed human biodiversity through the lens of
present study attempts to do just that by entirely life history evolution, a longstanding biological
avoiding the use of “race”, however defined, as theory originally postulated to explain cross-
a proxy for ancestral ecology and referring di- species variation in longevity, pace of living,
rectly to the zoogeographical regions of origin and the allocation of bioenergetic resources ei-
of contemporary human populations as the pri- ther to survival and somatic maintenance on one
mary predictive factor. This has the benefit of hand or mating effort and reproductive output
constituting a more direct test of the original on the other hand. Within the context of life
evolutionary ecological hypothesis and remain- history evolution, Rushton was interested in hu-
ing safely agnostic on the politically and scien- man intergroup differences as they evolved
tifically controversial topic of racial categoriza- within geographically disparate ecologies. For
tions. Many critics have made valid points instance, Rushton was an early advocate of
against some of Rushton’s methods but at the what became known as cold winters theory. He
cost of missing the central point of the entire additionally discussed migration out of Africa
scientific exercise. The present study seeks to into Eurasia as imposing cognitively demanding
refocus us on the original objectives of Differ- environmental exigencies. The seasonal varia-
ential-K Theory as an evolutionary ecological tion of temperate zones into which humans mi-
prediction. Further, the present study avoids grated, Rushton theorized, was predictable,
definitional distractions by relying on well- which contrasts with the “sudden droughts and
accepted biodemographic indicators of life his- devastating viral, bacterial, and parasitic dis-
tory strategy, such as fertility, longevity, and eases” common to Africa (Rushton, 2000, p.
mortality, and does not venture into the various 231). To demonstrate such variation required
psychosocial indicators to which Rushton ex- data points spanning a host of life history traits
tended the logic of life history theory from populations disparately distributed across
(Figueredo, Cabeza de Baca, & Woodley, the world. Naturally, Rushton therefore looked
2013). to mine previously assembled data, much of
It is also important to note that the described which was found in past anthropological studies
rankings of fast-to-slow life history speeds do and in other databases employing three broad
not in any way imply some kind of normative racial categories: Mongoloids, Negroids, and
hierarchy or bogus evolutionary progression Caucasoids.
from more primitive to more advanced. Evolu- The capacity for global racial categorization
tionary ecological thinking would instead pre- came after the age of exploration: Trading
BIOGEOGRAPHY OF HUMAN LIFE HISTORY STRATEGY 3
routes, shipping lanes, crusades, and conquests that categories only multiplied into finer group-
were among the many mechanisms of cross- ings in later decades, after Rushton had com-
cultural contact between far flung human pop- piled the research that undergirded his Differ-
ulations. Swedish systematizer Carl Linnaeus ential K Theory, which was first published in
(1758), for instance, parsed between Homo 1985.
Americanus, Europeanus, Asiaticus, and Africa- Intensive genetic surveys of the kind initi-
nus within the context of erecting the modern ated by Cavalli-Sforza (1997) have demon-
foundations of taxonomy. Georges-Louis strated the existence of extant regional ge-
Leclerc, Comte de Buffon, 18th century French netic clusters, which are found across vast
naturalist, published The Varieties of the Hu- geographic areas. Within these areas, large as
This article is intended solely for the personal use of the individual user and is not to be disseminated broadly.
man Species (1807), which more finely discrim- they are unto themselves, one finds gradients
This document is copyrighted by the American Psychological Association or one of its allied publishers.
inated between populations, including classifi- of linear genetic variation. However, there is
cations such as Muscovite that corresponded a concomitant trend wherein geographic bar-
with national territories and subnational groups riers impart genetic discontinuities. It is from
such as the Ostiacks. Blumenbach (1825) later mountains, rivers, seas, and oceans, among
subdivided humanity among Caucasian, Mon- other geographic barriers that add to distance
golian, Malayan, Ethiopian, and American rac- to restrict gene flow, that the realities of racial
es. Subsequently, Georges Cuvier (1833), born categories rest. Without these barriers, one
in France during the latter half of the 18th would only have uninterrupted continuous
century, gave preference to a simpler tripartite variation, as seen within regions.
classification into Caucasian, Mongolian, and With these barriers, one sees the justification
Ethiopian. Thereafter in the mid-19th century, for categorization not into three broad groups,
Joseph Arthur, Comte de Gobineau, in an Essay
but into the following seven groupings: Africa,
on the Inequality of the Human Races (1853)
Europe, Middle East, Central/South Asia, East
also employed a tripartite scheme, labeling
Asia, Oceania, and America. Again, within each
these same three groupings black, white and
of these clusters are clinal gradients of varia-
yellow races. William Z. Ripley’s The Races of
tion, but between them are sufficient genetic
Europe, produced in 1899, delineated the Teu-
tonic, Mediterranean, and Alpine races, which discontinuities to support the continuance of
was followed by Madison Grant’s (1916) Cau- some degree of categorical classification
casoid, Negroid, and Mongoloid classifications. (Rosenberg et al., 2005). Linnaeus and other
Interestingly, at least when viewing this small early typologists came closest to this genetically
sampling of theorists, lumpers predominated informed sevenfold classificatory scheme, but
over splitters as time went on, such that one sees even these were not sufficiently nuanced to
four and five divisions routinely being replaced comport with the reality of variation within our
by three, which then, by intellectual inertia or cosmopolitan species, spread as it is across a
otherwise, continued on into early anthropolog- vast a geographically varied earth.
ical formulations. Carleton Coon’s Caucasoid, In an age becoming increasingly racially con-
Congoid, Capoid, Mongoloid, and Australoid, scious and politically correct, Rushton (1988),
in this light, served as a bellwether for the following the “Big Three” categorical classifi-
coming transition back to finer distinctions cations, found himself reporting that the rate of
(Coon & Hunt, 1966). Nevertheless, as Brian dizygotic twins differs by race, such that Mon-
Siegel discusses, a tripartite division was used goloids have one quarter the rate of twins as
by early anthropologists that captured much compared to Negroids, with Caucasoids falling
variation but awkwardly incorporated or other- in the middle. In addition to rates of twinning,
wise failed to incorporate disparate peoples. For Rushton (1988) also describes race differences
many decades of the 20th century, a small num- in the rates of fetal growth, motor milestones,
ber of racial categories, often three, were used sexual maturation, and age of menarche.
by those collecting data on human racial diver-
sity, including anthropologists; with the same 1
https://www.pewsocialtrends.org/interactives/multi
often holding for research samples compiled by racial-timeline/.
the United States Census Bureau1 and the 2
https://www.npr.org/sections/thetwo-way/2014/11/07/
United States Army.2 Of note is the observation 362243042/army-drops-use-of-term-negro-in-document.
4 FIGUEREDO, HERTLER, AND PEÑAHERRERA-AGUIRRE
Rushton additionally described life history which he may have felt pinioned into mounting. In
driven differences in monogamy and mating doing so, he entrenched, ostensibly confirming the
alongside differences in the onset of menarche centrality of race in the eyes of critics. Rushton
and fertility. Rushton himself, for all that he (2000) asserted that “race is a biological concept”
remonstrated, never publicly explained that his (p. 96). He continued, writing:
decision to use racial categories was in effect a Races are recognized by a combination of geographic,
matter of convenience sampling, akin to how ecological, and morphological factors and gene fre-
many studies use conveniently accessible col- quencies of biochemical components. However, races
lege students as stand ins for the population at merge with each other through intermediate forms,
large. Though he never said as much in pub- while members of one race can and do interbreed with
members of other races.
This article is intended solely for the personal use of the individual user and is not to be disseminated broadly.
perfectly aware that there may exist approxi- Thereafter we can intuit his tacit recognition
mately five or six broad genetic clusters, de- that, of course, racial categories did not amount
pending on the method used to identify them to immutable Platonic types:
(Cavalli-Sforza, 1997; Cavalli-Sforza, Menozzi,
Within each race, several varieties or minor races have
& Piazza, 1994; Jobling, Hurles, & Tyler- been proposed, although there is no agreed upon num-
Smith, 2004; Rosenberg et al., 2002), many of ber. Mostly for political reasons, a majority of inves-
which are collapsed and combined across the tigators avoid the use of the term race as much as
ill-informed tripartite racial divisions around possible and use, for the major human races, the word
‘population’ and for the minor races, the phrase ‘ethnic
which most official data collection efforts had group’ (Rushton, 2000, p. 96).
been organized. From the outside, some critics
saw race categorization as the end of Rushton’s Feeling himself uniquely exposed, Rushton
reasoning and research, missing altogether that continued, lamenting that deliberate withhold-
his was a study of how ecological determinants, ing of evidence had become commonplace
as they varied geographically, created life his- among evolutionary scientists writing about
tory diversity among humans. The fateful, race. Rushton held out the following authors as
though possibly unavoidable, decision to allow examples: Stephen J. Gould, author of the re-
racial categories to serve as proxies for the vised and expanded edition of The Mismeasure
evolved effects of continuously distributed eco- of Man (1996), Jared Diamond, author of Guns,
logical variation blunted the thrust of his life Germs, and Steel (1997), and Christopher
history research program and subsumed his later Stringer, coauthor with Robin McKie of African
years in bitter controversy. Exodus (1996). Rushton was not without his
Rushton, and to a lesser extent the application champions, few and far between though they
of life history theory to human biodiversity, were. Glayde Whitney, in reviewing Rushton’s
became mired in a defense against critics. work, wrote of the “remarkable resistance to
Rushton’s energies were channeled in three racial science in our times,” which he compared
paths. First he protested against attacks on his to the inquisition in Rome during the Renais-
academic freedom, insisting that the “ultimate sance. He thereafter likened Darwin, Galton,
aim of science is causally to explain the world and Rushton to Copernicus, Kepler, and Gali-
around us . . .” Rushton continued, writing of leo; all being men who generated ideas that
the “chilling effect of self-imposed censorship,” encountered stalwart resistance from their re-
which pervaded “our own academic institutions, spective quarters. L. S. Gottfredson defended
and indeed even our own minds” (Rushton, Rushton’s Race, Evolution, and Behavior: A
1999, p. 220). Second, he politicized attacks by Life History Perspective even as it was deemed
squarely identifying the ideological quarter by others “racist trash.” Yet, those defending
from which those attacks proceeded. “The po- Rushton, like Rushton himself, followed the
litical left,” Rushton wrote, “poses a more seri- lead of critics, and assembled battle lines across
ous and immediate threat to the advance of the issue of race.
evolutionary science than does religious funda- One exception is a review by Figueredo,
mentalism because fundamentalism has no Cabeza de Baca, and Woodley (2013), which
clout in research universities whatsoever.” Far pointedly regretted that Rushton’s “work on the
and away, however, the main tributary carrying psychometrics of LH strategy was overshad-
away Rushton’s energies was his defense of race, owed by the controversy surrounding his pre-
BIOGEOGRAPHY OF HUMAN LIFE HISTORY STRATEGY 5
diction of race differences in LH strategy as While early physical atlases relied solely on
adaptations to ancestral ecological conditions in indicators of physical ecology such as latitude
the different regions of the world.” With the and longitude (Johnston, 1850), subsequent
field of social biogeography only lately mitigat- zoogeographic examinations considered species
ing the loss, the spectacle of race long sup- variation as a critical marker for regional cate-
pressed a better understanding of ecological re- gorization (Sclater, 1858; Wallace, 1876).
gions, along with the various selective regimes Sclater, for example (1858), compared the pres-
they impart and the evolved differences they are ence of avian species across land areas, allow-
capable of creating. ing the author to identify six zoogeographic
regions. The Palearctic region included Europe,
This article is intended solely for the personal use of the individual user and is not to be disseminated broadly.
Zoogeographic Regions as Alternative North Asia, and North Africa. The Ethiopian
This document is copyrighted by the American Psychological Association or one of its allied publishers.
(Morrone, 2015). The Cape region includes the zation on numerous environments around the
southern parts of South Africa (Morrone, 2015). globe. This cosmopolitan status exposed human
New Zealand, New Guinea, New Caledonia, communities to an array of unique environmen-
and Australia comprise the Australian region tal pressures leading to trait variation (Cochran
(Sclater, 1858; Morrone, 2015). Finally, the last & Harpending, 2009). Social biogeography, a
area corresponded to Antarctica (Morrone, branch of human sociobiology, emerged as a
2015). Like Wallace (1876), Morrone (2015) discipline destined to examine the influence of
also suggested the presence of transition zones. physical and community ecology (e.g., mean
The Mexican zone extends from southern North annual temperature, mean annual precipitation,
America to central Meso America and joins the altitude, latitude and longitude, parasite burden)
This article is intended solely for the personal use of the individual user and is not to be disseminated broadly.
Nearctic with the Neotropical region. The Sa- on interpopulation differences in life history as
This document is copyrighted by the American Psychological Association or one of its allied publishers.
hara-Arabian zone corresponding to the Ara- well as in social, political, and economic out-
bian Peninsula and parts of North Africa di- comes (e.g., Figueredo et al., 2017; Peñaher-
vided the Palearctic with the Ethiopian region. rera-Aguirre et al., 2019). Since biogeographic
The Indo-Malayan region acted as a border be- and zoogeographic classifications are predicted
tween the Australian and the Oriental region. to capture differences in physical ecology, these
The Chinese zone divides the Oriental from the regions should also adequately detect variations
Palearctic region. between human populations. Similarly, since
In tandem with these publications, phylogeog- each region features a unique interplay of sym-
raphers have also examined species’ distribution biotic interactions between species assem-
after accounting for phylogenic effects. Holt et al. blages, zoogeographic regions are predicted to
(2013) proposed a zoogeographic classification detect differences in parasite diversity.
following the distribution of over 21,000 species, It could be argued that differences between
accounting for the phylogenetic proximity be- human populations are exclusively the product
tween the species in the dataset. The analyses of human behavioral flexibility. However,
revealed 11 zoogeographic realms, which could, rather than considering these abilities as con-
in turn, be subcategorized into 20 regions. The trary to physical and community ecology, social
classification included the Palearctic, the Sino- biogeography predicts cultural and cognitive
Japanese, the Saharo-Arabian, the Afrotropical, ecology react to their respective environmental
the Madagascan, the Oriental, the Oceanian, the conditions, often increasing underlying popula-
Australian, the Nearctic, the Panamanian, and the tion differences. Moreover, it is worth noting
Neotropical realms. that while behaviorally modern humans are
While the work of Holt et al., supported characterized for their reliance on cultural niche
Sclater’s and Wallace’s predictions, the analy- construction (Odling-Smee, Laland, & Feld-
ses also differed in some respects. For instance, man, 2013), several aspects of physical ecology
The Saharo-Arabian realm extended beyond are less susceptible to human interference, such
North Africa and acted as an intermediate zone as altitude, temperature, and precipitation, ex-
between the Afrotropical and the Sino-Japanese cept for the construction of shelters. Current
realms (Holt et al., 2013). Similarly, the authors biogeographic examinations have demonstrated
discovered that insular regions east of Borneo the additive and nonadditive influence of vari-
and Bali, rather than being part of the Australian ous dimensions of physical, community, and
biogeographical region, clustered with other is- cognitive ecology on human variation (e.g.,
lands in the Oceanian realm. Although Holt and Figueredo et al., 2017; Peñaherrera-Aguirre et
colleagues estimated the presence of 11 realms, al., 2019).
the classification of zoogeographic regions var-
ied depending on the taxa taken into consider- The Present Study
ation. Hence, while analyses restricted to mam-
malian data detected 34 regions, examinations We address the various concerns regarding
based on the distribution of either avian or am- race-based schemes of life history variation
phibian taxa discovered fewer areas (Holt et al., by using the updated zoogeographical regions
2013). (Holt et al., 2013) based on data from over
As a species, behavioral modern humans 21,000 species of terrestrial vertebrate to es-
were characterized for their widespread coloni- tablish ecologically informed geographical
BIOGEOGRAPHY OF HUMAN LIFE HISTORY STRATEGY 7
areas within which humans (and other spe- Zoogeographic regions. The selected na-
cies) can be expected to evolve and develop tional polities were then assigned to the updated
different life history adaptations. By dividing zoogeographical regions published by Holt and
modern humans roughly into such regional colleagues (2013), based on data from over 21
ecotypes, instead of conventionally defined thousand species of terrestrial vertebrate, to es-
“races”, we can more directly test the basic tablish ecologically informed geographical ar-
premise of Rushton’s hypothesis while avoid- eas within which humans and other species can
ing much of the unnecessary controversies be expected to have evolved and developed
surrounding the definitions and utilities of different life history adaptations. In certain
such historically received but antiquated ra- cases, these assignments were decided based on
This article is intended solely for the personal use of the individual user and is not to be disseminated broadly.
the specific effects of parasite burden prior to mortality and life expectancy were statistically
aggregation; we obtained per capita disability adjusted by exporting the regression residuals
adjusted life years (DALY) from the World for the logarithmic effects of human parasite
Health Organization (2015) and historical infec- burden. This was done to control statistically for
tious disease prevalences from Murray and any cross-cultural variation with respect to his-
Schaller (2010) as indicators of human parasite torically recent advances in medical technology
burden, using a logarithmic transformation of and public health interventions that were not
their unit-weighted composite for each national present in the ancestral environments and to
polity. which current human populations have presum-
Population. Populations for each national ably not had very much time to adapt geneti-
This article is intended solely for the personal use of the individual user and is not to be disseminated broadly.
polity within the sample were obtained from cally by selection. This statistical adjustment
This document is copyrighted by the American Psychological Association or one of its allied publishers.
the United Nations Department of Economic had relatively negligible effects upon the mag-
and Social Affairs, Population Division’s nitudes of the parameters within the correlation
(2011) World Population Prospects: The matrix, as shown in Table 1.
2010 Revision, supplemented by official na- Table 2 shows the unit-weighted factor load-
tional statistics published in the United Na- ings of each of the indicators from its latent
tions (2010) Demographic Yearbook, 2009- common factor, the slow life history composite
2010, and data compiled by the Secretariat of (see Table 2), operationalized as part-whole
the Pacific Community (2010) SPC Statistics correlations, each serving as convergent validity
and Demography Programme. Population coefficients among the indicators (Gorsuch,
densities for each national polity within the 1983). These results are shown in the factor
sample were obtained from the World Bank structure table for both the adjusted and nonad-
Web Site (2019). justed forms of the variables. Again, the adjust-
ment had relatively negligible effects upon the
Statistical Analyses magnitudes of the parameters, but we decided to
use the more conservative (adjusted) estimates
All statistical analyses were performed using in all subsequent analyses.
SAS 9.4; the unit-weighted factor structures
were estimated using PROC CORR, and the The Structural Models
analyses of variance were done using PROC
GLM, with each national polity weighted by its Four basic parameters of the physical and
population. Multilevel models (MLMs) were community ecology were used to examine the
performed using PROC MIXED. predictive validity of the zoogeographic re-
gions: (a) average annual temperatures, (b) av-
Results erage annual precipitation, (c) the natural loga-
rithms of human parasite burden, and (d) the
The Measurement Model population densities. While this list of relevant
parameters is by no means exhaustive, it sum-
The theoretically expected positive manifold marizes many of the ecological selective pres-
was found among the four biodemographic in- sures that have been proposed to account for life
dicators of life history strategy. Both infant history evolution and thus for the diversity in
Table 1
Bivariate Correlations Among Slow Life History Strategy Indicators
Teen pregnancy Birth rate Infant mortality Life expectancy
Teen pregnancy 1.00 .88ⴱ .72ⴱ ⫺.76ⴱ
Birth rate .88ⴱ 1.00 .83ⴱ ⫺.82ⴱ
Infant mortality .65ⴱ .73ⴱ 1.00 ⫺.86ⴱ
Life expectancy ⫺.70ⴱ ⫺.71ⴱ ⫺.78ⴱ 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 ⬍ .0001.
BIOGEOGRAPHY OF HUMAN LIFE HISTORY STRATEGY 9
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 were doing a reasonably good job of partition-
between unit-weighted factor scores for LPB-adjusted SLH ing the variance among national polities in these
and non-LPB-adjusted SLH is .98ⴱ.
ⴱ
p ⬍ .0001. basic dimensions of the physical and commu-
nity ecology.
Finally, for our unit-weighted factor scale of
life history strategies among human populations slow life history strategy, the assigned zoogeo-
(e.g., Rushton, 1985, 2000). The bivariate cor- graphic regions accounted for 78.6%, F(6,
relations among these ecological parameters as 133) ⫽ 81.59, p ⬍ .0001, of the variance across
well as with LPB-adjusted slow life history national polities; when tested individually,
strategy are shown in Table 3. dummy variable analyses also revealed that
Five analyses of variance were performed to each of the zoogeographical regions, excepting
determine how well the assigned zoogeographic only the Madagascan (MA) and Oceanian (OC),
regions explained the variance among the na- was significantly different in life history strat-
tional polities nested within them. The first four egy from the ultimately ancestral Afro-Tropical
of these were done mostly as “manipulation (AT) region, indicating subsequent adaptive ra-
checks” to see how well the zoogeographic re- diation in life history strategies. The distribution
gions partitioned the cross-national variance in of these means is presented graphically in Fig-
basic parameters of physical and community ure 1.
ecology; afterward, the fifth of these analyses of We also performed hierarchical general lin-
variance was done on life history strategy itself ear models to determine the relative incremental
to test the main hypothesis of the present study. validities of the four continuous ecological pa-
All analyses of variance were weighted by the rameters with respect to the zoogeographical
population of each national polity. regions. As with the previous analyses of vari-
For average annual temperature, the assigned ance, all hierarchical general linear models were
zoogeographic regions accounted for 86.5%, weighted by the population of each national
F(6, 133) ⫽ 142.58, p ⬍ .0001, of the variance polity. When entering the four continuous eco-
across national polities; for average annual pre- logical parameters hierarchically prior to the
cipitation, the assigned zoogeographic regions 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
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.
quential distance “Out-of-Africa” for each na- applying a classificatory scheme based on the
This document is copyrighted by the American Psychological Association or one of its allied publishers.
tional polity, estimating the serial autocorrela- most recently updated (Holt et al., 2013) map-
tions of each national polity with each spatially ping of zoogeographical regions of origin, be-
adjacent one within that ordered sequence. This lieving this idea to be the most faithful to the
model is not designed to control statistically for original theoretical motivation behind
the effects of cultural diffusion, which would Rushton’s evolutionary theory of the diversifi-
presumably require estimating network autocor- cation of human life history strategies.
relations, but instead models the possible phy- Having accomplished this reconceptualiza-
logenetic effects of common human origins and tion of the problem, we found that the great
subsequent outward migrations (see Hertler et preponderance of the variance among contem-
porary national polities in aggregate life history
al., 2018). We limited the autocorrelational
speeds can be explained with reference to zoo-
analysis to this simplified sequential model be-
geographical regions. This was done without
cause we strongly doubt that the highly herita-
any reference to or use of racial classifications,
ble biodemographic population parameters used
indicating empirical support for the originally
in the present analyses would be very highly
hypothesized causal principle of ancestral ecol-
susceptible to hypothetically alternative (mean- ogy. The pattern of results matches quite closely
ing nonselective) processes such as cultural dif- what one would expect from a purely ecological
fusion across national boundaries. theory of life history variation, as well as the
MLM residuals were thus exported for LPB- progression of ancestral human migrations “Out
adjusted slow life history strategy and used for of Africa”, without the many imprecisions in-
subsequent general linear modeling. These troduced by obsolete racial categorizations.
MLM residuals had been statistically adjusted Nevertheless, statistically controlling for the re-
for the ordinal distance Out-of-Africa (OOA) as constructed “Out of Africa” sequence, as well
well as of any single-lagged heterogeneous au- as the serial spatial autocorrelations possibly
toregressive serial dependencies (ARH1) produced by those successive population expan-
among successive (meaning spatially contigu- sions, produced a reduction of at most 16% in
ous) data prior to regression modeling, thus the variance explained by the zoogeographical
circumventing this potential problem as a threat regions, leaving nearly 63%. This suggests that:
to the validity of correlational analysis. Using (a) neither phylogenetic inertia (Huber, 1939;
the studentized MLM residuals of OOA, the but see Shanahan, 2011) nor “Isolation by Dis-
assigned zoogeographic regions accounted for tance” (Ishida, 2009; but see Slatkin, 1993)
62.6%, F(6, 133) ⫽ 37.11, p ⬍ .0001, of the among the contiguous areas that were colonized
variance across national polities in OOA- successively by human “Out of Africa” migra-
adjusted slow life history strategy. In contrast, tions were sufficient to account for our reported
the reconstructed “racial” categories only ac- pattern of results in human life history biodi-
counted for 16.8%, F(2, 137) ⫽ 13.79, p ⬍ versity, and (b) the preponderance of the vari-
.0001, of the variance across national polities in ance among contemporary national polities is
OOA-adjusted slow life history strategy, which instead best explained by divergent evolution
once again constituted a statistically significant produced as a result of differential ecological
loss of explanatory power, F(4, 133) ⫽ 40.76, selective pressures among zoogeographical re-
p ⬍ .0001, with respect to the zoogeographic gions. It should be noted, however, that Thorn-
regions. hill and Fincher (2013) have argued quite con-
12 FIGUEREDO, HERTLER, AND PEÑAHERRERA-AGUIRRE
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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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(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
This article is intended solely for the personal use of the individual user and is not to be disseminated broadly.
(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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