Original Paper
Brain Behav Evol 2023;98:93–106 Received: May 26, 2022
DOI: 10.1159/000528710
Climate Change Influences Brain Size in
Humans
Jeff Morgan Stibel  
Natural History Museum, Los Angeles, CA, USA
Keywords
Climate change · Brain size · Encephalization · Human
evolution
Abstract
Brain size evolution in hominins constitutes a crucial evolu-
tionary trend, yet the underlying mechanisms behind those
changes are not well understood. Here, climate change is
considered as an environmental factor using multiple paleo-
climate records testing temperature, humidity, and precipi-
tation against changes to brain size in 298 Homo specimens
over the past fifty thousand years. Across regional and glob-
al paleoclimate records, brain size in Homo averaged signifi-
cantly lower during periods of climate warming as compared
to cooler periods. Geological epochs displayed similar pat-
terns, with Holocene warming periods comprising signifi-
cantly smaller brained individuals as compared to those liv-
ing during glacial periods at the end of the Late Pleistocene.
Testing spatiotemporal patterns, the adaptive response ap-
pears to have started roughly fifteen thousand years ago and
may persist into modern times. To a smaller degree, humid-
ity and precipitation levels were also predictive of brain size,
with arid periods associated with greater brain size in Homo.
The findings suggest an adaptive response to climate change
in human brain size that is driven by natural selection in re-
sponse to environmental stress. © 2022 The Author(s).
Published by S. Karger AG, Basel
Karger@karger.com © 2022 The Author(s)
www.karger.com/bbe Published by S. Karger AG, Basel
This is an Open Access article licensed under the Creative Commons
Attribution-NonCommercial-4.0 International License (CC BY-NC)
(http://www.karger.com/Services/OpenAccessLicense), applicable to
the online version of the article only. Usage and distribution for com-
mercial purposes requires written permission.
Accepted: December 12, 2022
Published online: December 27, 2022
Introduction
Understanding evolutionary brain size changes within
a genus is critical to determining how different species
adapt to broader environmental and cultural changes.
This is particularly true for humans, given the reliance on
increases in cognitive capacity over the past two million
years. Yet we know surprisingly little about what causes
adaptive changes to the human brain [Dunbar and Shultz,
2017].
Increasing brain size has been one of the more consis-
tent by-products of hominin evolution [Pilbeam and
Gould, 1974; Beals et al., 1984; Ruff et al., 1997; McHenry
and Coffing, 2000; Rightmire, 2004; Shultz et al., 2012;
Antón et al., 2014; Du et al., 2018; González-Forero and
Gardner, 2018]. Early Homo benefited from large increas-
es in relative and absolute brain size [Pilbeam and Gould,
1974; McHenry and Coffing, 2000; Antón et al., 2014;
Püschel et al., 2021; Gingerich, 2022]. This was followed
by new species within the genus, including H. erectus, H.
heidelbergensis, and H. neanderthalensis, all of whose spe-
ciation events were marked in part by increased brain size
and encephalization [Ruff et al., 1997; McHenry and
Coffing, 2000; Antón et al., 2014]. H. sapiens evolved even
greater brain size and encephalization around their spe-
ciation event roughly 300,000 years ago [Rightmire, 2004;
Antón et al., 2014; Hublin et al., 2015; Du et al., 2018].
Correspondence to:
Jeff Morgan Stibel, jeff @ bryantstibel.com
 While the evolutionarily benefits of Homo brain
growth are well known – increased cognitive capacity,
tool use, language acquisition, sociality [Jerison, 1973;
Dunbar, 2003; Shultz et al., 2012; Antón et al., 2014;
González-Forero and Gardner, 2018] – the reasons why
the human brain increased over time are not well under-
stood [Dunbar and Shultz, 2017]. Numerous hypotheses
have been proposed to explain Homo brain size increases,
including environmental and ecological changes [Antón
et al., 2014; González-Forero and Gardner, 2018], diet
and food availability [Aiello and Wheeler, 1995; Antón et
al., 2014], competitive [Bailey and Geary, 2009] and co-
operative [Dunbar, 2003; Antón et al., 2014] dynamics
within the genus as well as with other animals [González-
Forero and Gardner, 2018], and as a general response to
greater cognitive demands [Dunbar and Shultz, 2017]. It
is likely that some combination of each of these factors,
in addition to other variables, played a role in driving
brain size increases in Homo.
With respect to environmental factors, variability in
precipitation levels [Will et al., 2021], net primary pro-
duction [Stewart, 2014; Will et al., 2021], climate pulses
[Shultz and Maslin, 2013], and climate variability [Ash
and Gallup Jr, 2007; Antón et al., 2014] have all been pro-
posed to impact brain size to some extent. However, rig-
orous testing of these hypotheses has not been conducted,
and as such, the mechanisms driving brain size changes
are not fully understood. Prior studies have relied on data
records that do not account for geochronological errors
that are prevalent in older fossils and do not take into ac-
count the spatiotemporal lag associated with adaptive re-
sponses to environmental variables [Sabeti et al., 2006;
Dehasque et al., 2020; Evershed et al., 2022]. More impor-
tantly, no study to date has controlled for body size and
encephalization, factors known to contribute to brain size
variability [Jerison, 1973]. As a result, studies testing evo-
lutionary brain size changes [Ash and Gallup Jr, 2007;
Bailey and Geary, 2009; Shultz and Maslin, 2013; Stewart,
2014; Will et al., 2021] have either been inconclusive [Bai-
ley and Geary, 2009; Will et al., 2021] or contested [Shultz
et al., 2012].
At greater issue is that almost all prior studies have
generally assumed brain size has persistently increased
throughout time. However, despite general trends of in-
creasing brain size in Homo across time, Homo brain size
has not been on a persistent upward march. Periods dur-
ing the early Middle Pleistocene (roughly 875,000 years
ago) [Ruff et al., 1997] and the Holocene (roughly 12,000
years ago to present) [Henneberg, 1988; Henneberg and
Steyn, 1993; DeSilva et al., 2021; Stibel, 2021] both reflected
94 Brain Behav Evol 2023;98:93–106
DOI: 10.1159/000528710
deceleration in brain size, and modern evidence shows
a decrease in both brain size and encephalization [Stibel,
2021]. For humans, there is a clear adaptive advantage
to larger brains, and speciation events have repeatedly
yielded lineages with greater relative brain size; increas-
es in cognitive ability have consistently outweighed the
costs of expensive brain tissue. As such, it is less clear
why the brain would shrink during certain evolutionary
periods, leading to hypotheses that the cognitive advan-
tage that comes with larger brains has a breakpoint
where it may no longer be evolutionarily beneficial
[DeSilva et al., 2021]. There are exceptions, however, to
the brain size advantage: brains are metabolically expen-
sive and produce high levels of heat relative to their mass
[Martin, 1981; Aiello and Wheeler, 1995; Hublin et al.,
2015; Dunbar and Shultz, 2017]. Environmental chang-
es, as a result, can be a key determinant of the net value
of greater brain size. This is particularly true with re-
gards to changing climates, as climate has been shown
to directly impact metabolic and thermoregulatory sys-
tems in animals [Bergmann, 1847; Beals et al., 1984;
Sheridan and Bickford, 2011; Kasabova and Holliday,
2015; Pomeroy et al., 2021; Will et al., 2021]. Given re-
cent global warming trends, it is critical to understand
the impact of climate change, if any, on human brain
size and ultimately human behavior.
Materials and Methods
The challenge with testing Homo brain size changes is a lack of
comparable sample data. Few intact skeletons exist, and brain size
estimates are difficult to ascertain when derived from cranial ca-
pacity leveraging fragmentary skeletal remains. What measure-
ments are available tend to be sparse and come from small samples
without sufficient information about the specimen. In addition,
precise carbon-14 dating is not available beyond roughly 50,000
years which makes it difficult to ascertain precise geochronology
of older fossils. Prior studies [Ash and Gallup Jr, 2007; Bailey and
Geary, 2009; Shultz and Maslin, 2013; Stewart, 2014; Will et al.,
2021] suffered as a result of either insufficient sample breadth or
an inability to control for dating errors, body size, taxon, and en-
cephalization, all of which have been shown to impact brain size
[Jerison, 1973]. The present study utilized a relatively large sample
(n = 298) which allowed for controls of each of the above con-
founds. Given that body size is highly correlated with brain size in
Homo [Pilbeam and Gould, 1974; Beals et al., 1984; Ruff et al., 1997;
McHenry and Coffing, 2000; Grabowski, 2016; Stibel, 2021], lati-
tude and sex – two strong correlates of body size – were used as
proxies for body size and served to control for encephalization dif-
ferences [Jerison, 1973; Beals et al., 1984; Ruff et al., 1997; Ruff et
al., 2018]. Fossils were also limited to the past 50,000 years (50 kyr)
to control for geochronology errors and because it is a period of
relative stasis in brain/body isometry [Ruff et al., 1997; McHenry
Stibel
and Coffing, 2000; Stibel, 2021]. While limiting the study to the
past 50 kyr presents some difficulties in analyzing longer term
trends, there is a particular advantage in that the climatic period
can be broken down into two dramatic temperature periods before
and after the last interglacial, roughly 17 kyr BP.
Ten studies that documented cranial capacity (cm3) in Homo
over the past 50 kyr BP were used to compile cranial data (see Da-
taset 1 for sources of measurements and dating). Cranial measure-
ments were obtained from meta-analyses, published sources, and
relevant updated data [Holloway, 1980; Holloway, 1981; Beals et
al., 1984; Brown, 1992; Henneberg and Steyn, 1993; Ruff et al.,
1997; De Miguel and Henneberg, 2001; Hawks and Wolfpoff, 2001;
Manjunath, 2002; Lordkipanidze et al., 2013]. Incomplete mea-
sures, unusable skull fragment data, and juvenile specimens were
discarded. Where multiple measurements were available for a par-
ticular skull, they were aggregated to provide an average cranial
capacity estimate. In total, 373 independent cranial capacity mea-
surements were utilized across 298 skulls. Brain size estimates were
derived from cranial capacity measurements, as cranial volume has
been shown to highly correlate with brain size in both modern and
prehistoric humans [Pilbeam and Gould, 1974; Beals et al., 1984;
McHenry and Coffing, 2000]. Cranial capacity was converted to
brain size using a formula derived from a least-squares regression
(r2 = 0.995, LSR) of 27 primate species (brain mass = 1.147 x cra-
nial capacity0.976) [Ruff et al., 1997]. Where precise dates or car-
bon-14 dating were available (n = 266), that source was used; oth-
erwise, the average was taken across all date estimates (n = 32).
Specifics on the type of measure, chronometric date, sex, taxon,
geography, formula derivations, and all sources are available in
Dataset S1 and Stibel, 2022.
Brain size data were compared to four climate records. The pri-
mary dataset utilized paleotemperatures derived from the Euro-
pean Project for Ice Coring in Antarctica (EPICA) at Dome C [Jou-
zel et al., 2007] (online suppl. Dataset S1; for all online suppl. ma-
terial, see www.karger.com/doi/10.1159/000528710). Records
from the ice core at EPICA Dome C provide precise 100-year sur-
face temperatures that have been found to correlate closely with
stacked climate data across 58 different regional sites, including
regions near the fossils used in this study [Lisiecki and Raymo,
2005; Jouzel et al., 2007]. Brain size estimates were compared to
coeval temperatures as well as to past temperatures at intervals dat-
ing back 5 kyr, 10 kyr, and 15 kyr from the fossil date to control for
the uncertainty surrounding the delayed adaptive response of the
spatiotemporal interaction between the two variables. To test for
evolutionary effects and control for the accuracy of carbon dating
which is not reliable within 100-year periods [Wright, 2017], the
temperature data were also cataloged for the present study into
increments of 5,000, 10,000, and 15,000 years (online suppl. Data-
set S1).
The ice core at EPICA Dome C provides precise surface tem-
perature measurements dating back 809,950 years [Jouzel et al.,
2007]. Temperature records at EPICA Dome C were measured as
a function of differences in past temperatures (∆T) relative to the
most recent 100-year temperature average. Temperature change
was calculated against a modern mean temperature as follows:
“The surface temperature change, ∆Ts (estimated with respect to
the mean value calculated over the last millennium), (was) calcu-
lated using the present-day spatial slope of 6.04‰/°C after correc-
tion for the change in the isotopic composition of the ocean (and)
a slight correction linked with changes in the altitude of the ice
Climate Change and Human Brain Size
sheet as calculated from the glaciological model used to derive the
timescale” [Jouzel et al., 2007]. Average temperature change (mean
∆T) across the EPICA Dome C records as compared to the last
millennium was calculated at −5.30°C.
The global climate records also offer insight into regional dif-
ferences. The climate records produced from Dome C in Antarc-
tica have been shown to correlate closely with stacked benthic δ18O
climate data derived from oceanic sediment cores across 58 differ-
ent regional sites, including sites located near the fossil remains
used in this study [Lisiecki and Raymo, 2005; Jouzel et al., 2007].
The stacked regional data displayed strong correlations (average r2
of 0.88 after alignment) with the Antarctic average [Jouzel et al.,
2007], suggesting relative consistency of the ice sheet record and
stacked regional averages. An additional measure using regional
surface temperatures derived from a sediment core from Lake Ma-
lawi in East Africa [Johnson et al., 2016] (see below for more de-
tails) was also compared to the EPICA Dome C record and was
found to be highly correlated during the periods used herein over
the past 50 kyr (LSR, r2 = 0.78, p < 0.0001, ANOVA).
Surface temperatures (°C) derived from sediment samples re-
covered from Lake Malawi (10°–14° S in eastern Africa) over the
past 1.3 million years [Johnson et al., 2016] were used as proxies
for regional temperatures in Africa (online suppl. Dataset S1). The
Lake Malawi record was shown to correlate with past changes in
atmospheric carbon dioxide and, when compared to average rain-
fall estimates, demonstrated a relationship between warmer, wet-
ter periods and cooler, drier cycles [Johnson et al., 2016].
Grain-size analyses of siliciclastic marine sediments from the
coast of Mauritania (core GeoB7920; 2,278-meter water depth)
were utilized to derive a humidity index (mm) for Northwest Af-
rica starting at 500 years ago and going through the entire 50 kyr
BP period (online suppl. Dataset S1) [Tjallingii et al., 2008]. To
derive relative changes in humidity, log ratios of three core end-
members from the marine sediment were utilized (log (hemi-pe-
lagic mud)/(coarse aeolian dust + fine aeolian dust)), whereas he-
mi-pelagic mud provided a measure of river runoff and aeolian
dust records, which served as a proxy for subaerial erosion and
continental vegetation cover [Tjallingii et al., 2008].
Pollen sequences (mm) were used to derive a time series of
mean annual precipitation level estimates by calculating the differ-
ence between the reconstructed absolute value of the pollen counts
and modern precipitation values (online suppl. Dataset S1) [Bon-
nefille and Chalie, 2000]. Four African pollen sequences were de-
rived from two sources dating back to 40 kyr BP: three samples
from the equatorial mountains in the Burundi highlands at 1,850–
2,240 m of elevation and a fourth sample from the South Tangan-
yika basin at 770 m of elevation (the fourth sample was derived
from a prior record provided by the same researcher and recon-
structed with a new synthesis in conformance with updated meth-
odology) [Bonnefille and Chalie, 2000]. Six additional sequences
were available but not utilized as there were no records with match-
ing time periods to available fossil remains.
Paleoclimate records were provided over 100-year averages or
greater, whereas fossil records were dated to the year, as best esti-
mated by geochronologists without regard for variance. The tem-
perature record was rounded down to match the estimated age of
each skeletal remain. Given the considerable variability in fossil
dating, efforts were made to ensure that the results accounted for
potential geochronological errors. Homo records in the sample
were limited to the past 50,000 years, where precise records or
Brain Behav Evol 2023;98:93–106 95
DOI: 10.1159/000528710
radiometric carbon dating accounted for the majority of date esti-
mates (266 of 298 specimens). Where exact dates or radiometric
dating were not available, multiple estimates were used and aver-
aged. Additionally, the fossils were clustered into 100-year, 5,000-
year, 10,000-year, and 15,000-year groupings, the latter of which
can account for dating errors up to 15,000 years. The full dataset is
available in the online supplementary Data and Stibel, 2022.
Results
A significant relationship was found between temper-
atures at EPICA Dome C and brain size estimates over the
past 50 kyr (Fig. 1a) (p < 0.0001, ANOVA), which held
constant after controlling for geography, sex, and taxon
(p < 0.0001, ANCOVA). Despite the correlation, it is im-
portant to note that the climate and brain size records do
not appear to correspond temporally (Fig. 1b–c). Brain
size changes appear to take place thousands of years after
changes to climate, and this is particularly pronounced
after the last glacial maximum, approximately 17 kyr.
While acclamation unfolds within a single generation and
natural selection can happen in as short as a few succes-
sive generations, species level adaptation often takes
many successive generations [Sabeti et al., 2006; Evershed
et al., 2022]. The time series trends demonstrate what ap-
pears to be a pronounced delay that may have been driv-
en by evolutionary adaption (Fig. 1d–f, online suppl. S1).
To test for evolutionary effects, brain size changes
were compared to prior generational climate periods as
well as across temperatures averaged over extended peri-
ods of time. Brain size estimates were first modeled
against 100-year average temperatures dating back 5 kyr,
10 kyr, and 15 kyr (online suppl. Fig. S2). A second set of
models was derived for brain size estimates as compared
to temperatures averaged across 5 kyr, 10 kyr, and 15 kyr
(online suppl. Fig. S2). Across each model, brain size re-
mained significantly correlated with global average tem-
peratures (all tests, p < 0.0001, ANOVA, online suppl.
Table S1). Inclusion of sex and latitude as covariates in-
creased the explanatory power of the models to approxi-
mately 40% (all tests, p < 0.0001, ANCOVA, online suppl.
Table S2), with the 10 kyr spatiotemporal lag showing the
highest predictive power at roughly 42% (online suppl.
Fig. S2).
To control for geochronological errors and to increase
statistical power, the Homo sample was next divided into
groups of individuals living during periods that were
above and below average temperatures. To account for a
bias toward warmer climatic periods in the skeletal sam-
ple, the data were compared using the average temperatures
96 Brain Behav Evol 2023;98:93–106
DOI: 10.1159/000528710
derived solely from the 298 periods where skeletal re-
mains were available (mean ∆T of −1.07°C). Across 100-
year mean average global temperatures over the past
50,000 years [Jouzel et al., 2007], brain size in Homo spec-
imens living during cooler than average climatic periods
was found to be significantly larger than that of those liv-
ing during warmer periods (p < 0.0001, t test). Homo
brain size during cooler periods averaged 1,426.31 g ±
137.30 (mean ± standard deviation) (n = 65) as compared
to 1,280.89 g ± 141.67 (n = 233) for warmer periods, or a
roughly 10.74% difference (Fig. 2a; Table 1).
To control for evolutionary effects, the brain size data
were also grouped using data aggregated into 10,000-year
increments (while 10 kyr increments were chosen for rep-
resentation in the study, tests using temperatures aver-
aged over 5-kyr and 15-kyr periods yielded directionally
similar results). Similar to the 100-year data, Homo brain
size was found to be significantly larger during periods of
global cooling (1,421.69 g ± 136.56 [n = 70]) as compared
to warmer cycles (1,279.12 g ± 141.77 [n = 228]), or a
roughly 10.57% difference (p < 0.0001, t test) (Fig. 2b;
Table 1).
Body size in both modern and prehistoric Homo tends
to be larger in high latitude (above 30° North) geogra-
phies [Beals et al., 1984; Ruff et al., 1997]. Given that body
size tends to correlate highly with brain size [Pilbeam and
Gould, 1974; Beals et al., 1984; Ruff et al., 1997; McHenry
and Coffing, 2000; Grabowski, 2016; Stibel, 2021], the sig-
nificant changes found in brain size over time could have
been a result of Homo dispersal across latitudinal clines.
Despite a bias toward high-latitude Homo in the sample
(220 of 298), controlling for body size across clines above
and below 30ºN latitude did not materially change the
results, and significant differences persisted in brain size
across climatic periods (p < 0.0001, ANCOVA, across
100- and 10,000-year periods). During cooler average
temperatures, larger bodied Homo brain size was 8.14%
larger (p < 0.0001, t test) and smaller bodied Homo brain
size was 19.87% larger (p = 0.0001, t test) than during
warmer than average climatic cycles (Fig. 3a, b; Table 1).
Similar to the ecogeographic patterns, sexual dimor-
phism is present in Homo, with males on average having
larger body sizes than females [Ruff et al., 1997; Ruff et al.,
2018]. Despite a bias toward males in the sample (167 of
257 sexed specimens), controlling for body size through
sex did not change the relationship between climate
change and brain size (p < 0.0001, ANCOVA, across 100-
and 10,000-year periods). Using sex as an intermediate
control, larger bodied Homo averaged 8.40% larger brain
size (p < 0.0001, t test) during cooler average temperatures
Stibel
 a b

c
d

e f

Fig. 1. Relationship between Homo brain size (g) and global aver-
age temperatures (changes in Antarctic surface temperature [°C]
relative to the mean for the last millennium). a Across the past
50,000 years, brain size demonstrated an inverse relationship with
climate change (LSR, rg = −0.362; p < 0.0001; Durbin-Watson,
1.224). b, c However, the linear relationship between brain size and
climate change is confounded by differences across time series
(online suppl. Fig. S1) and cubic trends. d–f As the climate record
moves from present day back temporally, the time series (online
suppl. Fig. S1) and cubic trends more closely align with brain size
trends over time. Black lines represent linear trends; red lines rep-
resent cubic trends; gray lines indicate 95% confidence intervals
around the observations; and dotted gray lines represent the con-
fidence intervals for each model.

Climate Change and Human Brain Size Brain Behav Evol 2023;98:93–106 97
DOI: 10.1159/000528710
 a b

Fig. 2. Scatter plot of Homo brain size (g) during periods of colder and warmer global average temperatures (changes
in Antarctic surface temperature [°C] relative to the mean for the last millennium) across 100-year and 10,000-year
periods. Homo brain size was significantly larger during cooler average temperatures as compared to warmer tempera-
tures across 100-year (a) and 10,000-year (b) periods. Diamonds represent the 298 brain mass estimates across cooler
(blue) and warmer (red) than average temperatures; black lines denote mean brain mass for each period.

as compared to warmer periods (smaller bodied Homo
was directionally consistent – 4.89% greater brain size
during cooler than average periods – but not significant,
possibly due to limited sample size, with only 11 cooler
than average specimens) (Fig. 3a, b; Table 1).
Because the sample consisted of modern and archaic
Homo, robusticity differences across species could also
have influenced the results. Anatomically modern H. sa-
piens (AM Homo) in particular tend to be significantly
more gracile with less skeletal and body mass than both
earlier hominins (Middle Pleistocene and archaic Homo)
and more recent archaic H. sapiens (i.e., Neanderthals)
[Hawks and Wolfpoff, 2001; Harvati et al., 2006; Hublin,
2009], which could account for some of the effect of cli-
mate on brain size given the estimates rely on skeletal
measurements. As compared to the archaic fossils in the
sample, brain size was 9.96% smaller for AM Homo on
average (p < 0.005, t test). To control for robusticity dif-
ferences, the impact of climate change was tested exclud-
ing the archaic Homo fossils, which eliminated 9 of the
298 specimens. Across AM Homo, the differences in brain
size during colder and warmer periods persisted (p <
0.0001, t test). During colder average periods (∆Ts below
−0.86°C), brain size across AM Homo averaged roughly
11.02% larger than during warmer than average periods,
or roughly 1,422.03 g ± 142.52 (n = 56) as compared to
1,280.89 g ± 141.67 (n = 233). The results held direction-
ally within 10,000-year periods and after controlling for
body size (all results significant [p < 0.01, t test], except
for small bodied sexed specimens, which were direction-
ally similar but not significant) (Fig. 3a, b; Table 1).
The results present a general pattern of changing
brain size in Homo that is correlated with climate change
as temperatures increase and decrease. On closer in-
spection, the effects of climate change on brain size ap-
pear more pronounced nearest the mean and do not ac-
celerate as temperatures grow more extreme. While sig-
nificant differences were found for moderate temperature
ranges (interquartile range, p < 0.0001, t test), there were
no significant differences found in brain size as temper-
atures became more extreme (all tests, p > 0.50, t tests).
As with the broader Homo sample, AM Homo brain
size also appeared more sensitive to moderate tempera-
ture changes with an adaptive response that occurred in

98 Brain Behav Evol 2023;98:93–106 Stibel

DOI: 10.1159/000528710
 Climate Change and Human Brain Size
Table 1. Homo brain mass during above and below average climatic periods

Homo sample Temporal range Climate record Brain mass (g)

(kyr BP) (region [mean climate*])
colder/arid climatic warmer/wetter climatic differencea
periods (mean ± SD [n]) periods (mean ± SD [n])

All Homo 50–present 100-year Antarctic surface temperatures (−1.07°C) 1,426.31 g±137.30 (65) 1,280.89 g±141.67 (233) 10.74%
All Homo 50–present 10,000-year Antarctic surface temperatures (−1.90°C) 1,421.69 g±136.56 (70) 1,279.12 g±141.77 (228) 10.57%
All Homo, Holocene versus Pleistocene 50–present 100-year Antarctic surface temperatures (−0.50°C) 1,426.96 g±139.28 (63) 1,281.95 g±141.58 (235) 10.71%
All Homo, latitude, large bodied 50–present 100-year Antarctic surface temperatures (−1.39°C) 1,426.85±136.14 (53) 1,319.49±127.89 (167) 7.82%
All Homo, latitude, small bodied 45–present 100-year Antarctic surface temperatures (−0.19°C) 1,388.45±145.36 (21) 1,158.28±108.43 (57) 18.08%
All Homo, sex, large bodied 50–present 100-year Antarctic surface temperatures (−0.23°C) 1,427.65±124.73 (42) 1,316.99±143.76 (125) 8.06%

DOI: 10.1159/000528710
All Homo, sex, small bodied 31–present 100-year Antarctic surface temperatures (−0.44°C) 1,258.54±121.75 (11) 1,199.76±113.10 (79) 4.78%a
All Homo 50–present E. African lake temperatures (25.75°C) 1,366.86 g±151.09 (94) 1,287.61 g±147.40 (204) 5.97%

Brain Behav Evol 2023;98:93–106

All Homo 50–0.55 African marine sediment humidity levels (−0.30 mm) 1,409.84 g±148.79 (45) 1,337.32 g±137.84 (72) 5.28%
All Homo 40–present African pollen sequence precipitation levels (−94.74 mm) 1,319.79±164.58 (190) 1,284.57±121.18 (99) 2.70%a
AM Homo 45–present 100-year Antarctic surface temperatures (−0.86°C) 1,422.03 g±142.52 (56) 1,280.89 g±141.67 (233) 10.44%
AM Homo 45–present 10,000-year Antarctic surface temperatures (−1.73°C) 1,417.98±141.11 (61) 1,279.12±141.77 (228) 10.23%
AM Homo 45–present E. African lake temperatures (23.19°C) 1,413.36 g±156.34 (40) 1,325.27±131.75 (68) 6.43%
AM Homo 45–0.55 African marine sediment humidity levels (−0.30 mm) 1,413.36±156.34 (40) 1,325.27±131.75 (68) 6.43%
AM Homo 38–present African pollen sequence precipitation levels (−95.20 mm) 1,319.79±164.58 (190) 1,283.91±121.62 (98) 2.76%a

period); E. African lake temperatures (ºC) measured using sedimentary core records from Lake Malawi GLAD7-MAL05-1 (43°N, 31°W, 378 m depth); African humidity index (mm) was derived from
* Antarctic surface temperatures (ºC) measured as change in temperature at EPICA Dome C (−75.10ºN, 123.35ºW, 3,189.45 m depth) relative to current millennium (most recent 100-year

grain-size analyses of siliciclastic marine sediments and hemi-pelagic mud from the coast of Mauritania (core GeoB7920; 2,278-m water depth); Annual mean precipitation level estimates (mm)
were calculated from the difference between the reconstructed absolute value of four African pollen sequences between 2°28′ to 3°42′ S, at 1,850–2,240 m of elevation and modern precipitation
values. Mean climates were derived from each of the four climate records by averaging the paleoclimates associated with a given specimen across each sample (see Methods). a All differences
significant p < 0.01 (t test), except sex, small bodied (p = 0.11) and precipitation (p = 0.061, All Homo; p = 0.058, AM Homo).

99
 a b
Fig. 3. Differences in Homo brain size (g) during periods of cooler
and warmer global average temperatures (changes in Antarctic sur-
face temperature [°C] relative to the mean for the last millennium)
across 100-year periods. a Homo brain size is significantly higher
during periods of cooler average temperatures across the past 50,000
years. Differences persist across high and low latitudes and across
Fig. 4. Relationship between brain size (g) and global climate change
(changes in Antarctic surface temperature [°C] relative to the mean
for the last millennium) across 100-year increments during extreme
temperature periods. “All Homo” (n = 298) represents the average
estimated brain size across the entire 50 kyr BP period using the data
from Fig. 2a. To either side of “All Homo” is the mean across colder
(n = 65) and warmer (n = 233) average temperatures, respectively (as
reported in Table 1). The sample was then broken down into quar-
tiles and colder and warmer than average temperature periods were
100 Brain Behav Evol 2023;98:93–106
DOI: 10.1159/000528710
both sexes when used as controls for body size, as well as when lim-
ited to AM Homo over the past 50,000 years (b). All comparisons
were significant (p < 0.001, t tests) except for the smaller bodied
sexed sample (p ≥ 0.11, t test). Blue represents cooler than average
temperatures; red represents warmer than average temperatures; er-
ror bars represent the standard error of each mean estimate.
compared against one another. Significant differences were found
between the sample average (“All Homo”) and all other groups with-
in the chart (all tests, p < 0.02, t tests), as well as across all warmer
and colder groups (all tests, p < 0.0001, t tests), except the warmest
quartile with respect to the sample average, which was directionally
similar but not significant (p = 0.08, t test); no significant differences
were found within warmer or colder average periods (all tests, p >
0.50, t tests). Error bars represent the standard error of each mean
estimate.
Stibel
Fig. 5. Changes to estimated brain size (g) across global climate
change (changes in Antarctic surface temperature [°C] relative to the
mean for the last millennium) across 100-year increments during the
Holocene and late Late Pleistocene. “All Homo” (n = 298) represents
the average estimated brain size across the entire 50 kyr BP period
using the data from Fig. 2a. To either side of “All Homo” is the mean
estimated brain size across the late Late Pleistocene (50‒12 kyr BP,
n = 63) and Holocene (12 kyr BP–present, n = 235), respectively. The
sample was then broken down into colder and warmer than average
advance of extreme temperatures (interquartile range,
p < 0.001, t test; extreme temperatures, all tests p > 0.50,
t test) (Fig. 4).
Temporal periods produced similar patterns of chang-
es to brain size. Testing temporal periods within the past
50,000 years presents a unique opportunity, as the time
period includes the last interglacial that produced consis-
tently colder average temperatures up until the end of the
Late Pleistocene (mean ∆T of −6.96°C) and the Holocene
(mean ∆T of 0.51°C), which has produced significantly
warmer temperatures on average throughout present day
(p < 0.0001, t test). The Holocene is a period noted for
declining brain size generally, and numerous hypotheses
have been proposed to explain the temporal decline [Hen-
neberg, 1988; Henneberg and Steyn, 1993; DeSilva et al.,
2021; Stibel, 2021], but global warming has not been di-
rectly tested against Homo brain size declines. Testing the
two epochs independently produced effects consistent
with the interquartile results. Whereas brain size in Homo
during the late Late Pleistocene (50–12 kyr BP) was
significantly larger than during the Holocene (12 kyr
Climate Change and Human Brain Size
temperature periods within each epoch and compared against one
another. Significant differences were found between the sample aver-
age (“All Homo”) and all other groups within the chart (all tests, p <
0.04, t tests), as well as between all Pleistocene and Holocene groups
(all tests, p < 0.0001, t tests) except the warmer Holocene quartile
with respect to the sample average, which was directionally similar
but not significant (p = 0.08, t test); no significant differences were
found within the Pleistocene or Holocene (all tests, p > 0.50, t tests).
Error bars represent the standard error of each mean estimate.
BP-present) (p < 0.0001, t test), there were no signifi-
cant differences between colder and warmer periods
within each epoch (all results p > 0.50, t test), suggest-
ing that most of the changes in brain size in Homo hap-
pened as a result of the deglaciation period. This was
further represented by the temporal trends in brain
size and climate change (Fig. 1, online suppl. S1). Pleis-
tocene brain size averaged 1,426.96 g ± 139.28 (n = 63)
as compared to 1,281.95 g ± 141.58 (n = 235) during
the Holocene, a roughly 10.71% difference (Fig. 5).
Because Homo dispersal originated out of Africa, three
regional African climate records were also tested against
the brain size sample: sedimentary records, marine sedi-
ment layers, and pollen sequences (online suppl. Dataset
S1). Sedimentary records have been shown to be predic-
tive of regional temperatures [Johnson et al., 2016], and
the sample utilized was highly correlated with the EPICA
Dome C climate record (LSR, r2 = 0.78, p < 0.0001, ANO-
VA); marine sediment layers have been shown to be pre-
dictive of humidity levels and vegetation cover [Tjallingii
et al., 2008]; and pollen sequences have been shown to be
Brain Behav Evol 2023;98:93–106 101
DOI: 10.1159/000528710
predictive of precipitation levels as well as estimated rain-
fall and water availability [Bonnefille and Chalie, 2000]
(See Methods for details). Similar to the global tempera-
ture results, colder regional temperatures were predictive
of larger brain size in humans; in addition, more arid con-
ditions (lower average rainfall and humidity) were found
to be predictive of larger brain size in humans (all tests, p
< 0.05, ANCOVA controlling for sex and latitude across
Homo and AM Homo). Differences in Homo brain size
were roughly 6.15% for regional temperature (p < 0.0001,
t test), 5.28% for humidity (p < 0.002, t test), and 2.74%
for precipitation levels (nonsignificant, p = 0.061, t test)
(Table 1).
For each of the above regional climate records, the
tests were also performed limiting the Homo sample to
the continent of Africa (n = 26) and sub-Saharan Africa
(n = 23) (there were no archaic specimens, so the entire
sample consisted of AM Homo). In all cases, the patterns
were directionally consistent with the overall findings,
but no significant results were produced through group
testing (all results, ≥0.06, t test), possibly as a result of
the low sample size after limiting to a single geography.
Modeling the African temperature record, humidity es-
timates, and precipitation levels against African Homo
and sub-Saharan African Homo accounted for roughly
42% (LSR, r2 = 0.421, p < 0.01, ANOVA) and 44% (LSR,
r2 = 0.439, p < 0.01, ANOVA) of the variability in brain
size, respectively.
Discussion
The reasons for hominin brain size adaptions are com-
plex and not well understood. Climate has frequently
been proposed as a driver of brain size evolution [Potts,
1996; Ash and Gallup Jr, 2007; Shultz and Maslin, 2013;
Antón et al., 2014; Stewart, 2014; Will et al., 2021], but
testing has not produced conclusive results [Ash and Gal-
lup Jr, 2007; Bailey and Geary, 2009; Shultz et al., 2012;
Shultz and Maslin, 2013; Stewart, 2014; Will et al., 2021].
By systematically controlling for encephalization, body
size, and taxon, the present study was able to isolate evo-
lutionary changes to the brain and systematically test the
impact of climate change. The results suggest that climate
change is predictive of Homo brain size, and certain evo-
lutionary changes to the brain may be a response to envi-
ronmental stress.
Adaptive responses to natural selection are difficult
to unpack given the spatiotemporal delay between the
underlying cause and effect [Sabeti et al., 2006; Burger
102 Brain Behav Evol 2023;98:93–106
DOI: 10.1159/000528710
et al., 2020; Dehasque et al., 2020; Evershed et al., 2022].
While macroevolutionary adaptations can take millions
of years, some modern adaptive responses in Homo,
such as lactose tolerance, have evolved more quickly
[Gerbault et al., 2011; Burger et al., 2020; Evershed et
al., 2022]. Given the data within this study can only pro-
vide correlational support for spatiotemporal relation-
ships, future studies will be needed to confirm any as-
sumptions regarding the responsiveness of brain size to
climate change. However, the results of the current
study demonstrate what may be an evolutionary adap-
tation to environmental stress in human brain size be-
ginning roughly 15,000 years ago and persisting through
present day.
The primary variable tested herein was global tem-
perature change. There is a strong basis for temperature
differences impacting brain size but that has not previ-
ously been demonstrated evolutionarily. To maintain
thermoregulatory balance in response to different tem-
peratures, physical size is believed to be adapted to re-
gional geographies given that smaller sizes dissipate
heat better than larger volumes with less relative surface
area [Bergmann, 1847; Beals et al., 1984; Sheridan and
Bickford, 2011; Kasabova and Holliday, 2015]. This is
true for human body size overall and brain size [Beals et
al., 1984]. Temperature-driven climate change has also
been shown to impact body size evolutionarily [Will et
al., 2021; Pomeroy et al., 2021; Stibel, 2021]; however,
the effect has not previously been demonstrated for
brain size. The present study demonstrates broad brain
size changes in response to changing temperatures using
both a global paleoclimate record, as well as a regional
climate record from Africa. The impact of climate
change on human brain size appears to be similar to how
geographic temperature differences affect body size in
response to thermal stress: global warming tends to fa-
vor smaller brain size, whereas global cooling favors
larger size.
In addition to temperature changes, humidity and
precipitation were also associated with brain size adapta-
tions, although this effect was less pronounced. In both
cases, aridity was predictive of greater brain size and
wetter periods were predictive of smaller size. As with
thermal stress producing larger brains in cooler cli-
mates, it appears as if environmental stress related to de-
creased water availability may produce larger brain size
in Homo. Some caution should be considered in inter-
preting these results in at least two respects: first, climate
records that measure water production tend to be high-
ly localized, and the data used herein were limited to
Stibel
regions in Africa; and second, at least with respect to
certain African climates, arid conditions tend to corre-
late with cooler climates [Johnson et al., 2016], such that
the impact of aridity may be mediated by the larger effect
of temperature on brain size.
Environmental hypotheses that have been proposed
for brain size increases can broadly be summarized into
two groups: those that assume hospitable environmental
conditions (wetter, fertile, and more consistent climates)
increase food availability which allows for larger brain
size [Stewart, 2014; Will et al., 2021] and those that as-
sume climate stressors (cold or variable temperatures and
arid environments) produce brain size increases as a cop-
ing response to more extreme conditions [Potts, 1996;
Ash and Gallup Jr, 2007; Antón et al., 2014; Will et al.,
2021]. With respect to increasing brain size, the results
herein generally support the latter hypothesis and are in-
congruent with the first. There was no relationship found
between hospitable environments and greater brain size.
Rather, there was evidence that colder, more arid periods
correspond to larger brain size.
In its response to temperature changes and water avail-
ability, the human brain appears to be reacting to envi-
ronmental stress as opposed to another adaptive mecha-
nism. However, when taking into account brain size de-
clines, climate stability appears to be the more critical
environmental variable underlying brain size changes.
While past hypotheses have focused on brain size increas-
es, the present data demonstrate a more general response
to climatic factors wherein brain size modulates depend-
ing on the environmental conditions. Brain size was larg-
er during periods of colder and more arid environments
and was smaller when temperatures were warmer and
wet. This was found irrespective of time period, and the
latest Holocene warming cycle was predictive of brain siz-
es in Homo that were smaller than prior periods. The
changes in brain size were proximate to the mean average
temperatures, and the general trend toward larger or
smaller brain size did not accelerate as climates grew
more extreme, suggesting a pattern of changing brain size
limited to transitionary periods as climates become more
extreme. However, some caution should be taken when
considering more granular trends given fossil dating con-
straints and spatiotemporal delays, so it is best to inter-
pret the data as demonstrative of a broader effect across
warming and cooling cycles until more precise methods
are available.
These findings have other taxonomic and phylogenic
implications. Specifically, some variation in brain size
within the genus Homo may be attributable to climate in
Climate Change and Human Brain Size
addition to broader evolutionary causes and should be
considered when interpreting observed morphological
variation. Speciation events have often been marked in
part by changes to brain size in hominins, and those deci-
sions may be worth reconsidering in the context of the
present results. Cranial morphology, brain mass, and en-
cephalization in particular have been considered as fac-
tors for classification within the genus Homo or as criteria
for demarcating a new species [Wood and Collard, 1999;
Harvati et al., 2006; Hublin, 2009; Hublin et al., 2015], but
these variables may be affected by climate change. By way
of example, there has been considerable debate as to
whether H. neanderthalensis should be considered a sub-
species of H. sapiens, in part because of their large brain
and body mass [Hawks and Wolfpoff, 2001; Harvati et al.,
2006; Hublin, 2009; Hublin et al., 2015; VanSickle et al.,
2020]. Neanderthal remains have almost exclusively been
found in high-latitude regions and from cooler climatic
periods, both of which are consistent with bigger body
and brain sizes [Hawks and Wolfpoff, 2001; Harvati et al.,
2006; Hublin, 2009]. The differences in brain and body
size may be more a function of climate than taxon, and
the present results may offer some additional guidance on
whether H. neanderthalensis should remain independent
of the H. sapiens clade. At a minimum, it may be worth
considering whether climatic periods should act as a con-
trol when considering morphological differences within
the genus.
The effects of climate change on brain size may have
been intermediated by body size adaptations, as similar
trends have been found for Homo body size changes in
response to climate change [Will et al., 2021; Pomeroy et
al., 2021; Stibel, 2021]. Fossil records have shown that in-
vertebrates, burrowing insects, and mammals all shrunk
during past periods of global warming [Sheridan and
Bickford, 2011; Peralta-Maraver and Rezende, 2021] and
periods of global cooling have been shown to drive body
size increases in many organisms [Millien et al., 2006].
These results are consistent with studies that have shown
body size changes in response to climate change in Homo
[Will et al., 2021; Pomeroy et al., 2021; Stibel, 2021]. It is
not clear from the present study whether brain size was
specifically selected for or against during periods of
changing climate or whether phenotypic changes to the
brain were a result of selective pressures acting on body
size that caused brain size to drift alongside the body. Re-
gression models have shown that selection on the brain
or the body independently in organisms can impact both
variables and produce a correlated response [e.g., Lande,
1979; Atchley, 1984; Riska and Atchley, 1985]. For hominins,
Brain Behav Evol 2023;98:93–106 103
DOI: 10.1159/000528710
at least one study has demonstrated a phenotypic link to Deary, 2015; Sniekers et al., 2017; Okbay et al., 2016; Sti-
brain size acting as the underlying variable under selec- bel, 2021]. Even a slight reduction in brain size across
tion in both body and brain size changes during prehis- extant humans could materially impact our physiology
toric periods [Grabowski, 2016]. In that case, brain size in a manner that is not fully understood.
increases were determined to be the primary phenotype
under selection, driving both brain and body size chang-
es. However, the study did not contemplate decreases in Acknowledgements
size or climate as an intermediating variable. It is not clear
as a result, whether the body, the brain, or a combination The author would like to thank Chris Ruff, Tim White, Robin
Dunbar, and Jeremy DeSilva for a critical review of the manuscript;
of the two are directly under selection in response to cli- Dan Dennett and Phil Lieberman for helpful commentary on spe-
mate change as opposed to one trait drifting through plei- cific areas; Chris Ruff, Jean Jouzel, and Peter Brown for providing
otropy as a result of the other. Given the human brain’s expanded datasets; Lindsey Long and Caitlin Mason for assistance
high thermal output and sensitivity to extreme heat [Falk, with data collection, tabulation, and standardization; and Zack
1990], it may be that climate disproportionately effects Stokes for assistance with mathematical equations and statistical
modeling.
brain size, but more data and analyses will be required to
determine the interrelationship between brain and body
size.
The underlying drivers of brain size changes are not Statement of Ethics
fully clear. While the results demonstrate a relationship An ethics statement was not required for this study type; no
between climate change and Homo brain size, climate human or animal subjects or materials were used.
appears to account for only a small amount of the varia-
tion in brain size evolutionarily. Brain size adaptations
are complex and are likely driven by other factors that Conflict of Interest Statement
affect the ecosystem (such as predation or other com-
petitive dynamics), by an indirect consequence of a The author was employed by Bryant Stibel at the time of the
changing climate (such as vegetation levels and net pri- publication and Bryant Stibel had no role in the study design; col-
lection, analysis, and interpretation of data; writing of the report;
mary production), or by means unrelated to climate or decision to publish.
(such as culture and technology). Environmental hospi-
tability is a complex formula that includes many vari-
ables influencing physiological adaptation. While there
Funding Sources
is some evidence to support temperature as the primary
climatic influence on Homo brain size, more work will No funding was acquired for this work.
be needed to determine whether the impact of climate
change on Homo physiology is a result specifically of
temperature changes or an indirect effect from other el- Author Contributions
ements of a changing environment.
To the extent that temperature changes directly influ- Jeff Stibel was responsible for the results and write-up of the
ence brain size, the present period of accelerated warm- manuscript.
ing could lead to increased evolutionary pressure on the
human brain. The earth is currently warmer than it has
been at any time in the past 125,000 years, with temper- Data Availability Statement
ature increases of more than 1.1°C during the past cen-
The data supporting the findings of this study are available in
tury and projections of an increase of at least 1.5°C by the study by Stibel [2022] and within the manuscript and its sup-
the end of the current century [Tollefson, 2021]. While plementary information files (online suppl. Data 1). Further inqui-
this may be a sampling or dating error, there is some ge- ries can be directed to the corresponding author.
netic evidence to suggest that brain size, general cogni-
tive ability, and educational attainment are all being se-
lected against in modern human populations [Reynolds
et al., 1984; Bouchard Jr et al., 1990; Deary et al., 2007;
Rietveld et al., 2013; Rietveld et al., 2014; Plomin and

104 Brain Behav Evol 2023;98:93–106 Stibel

DOI: 10.1159/000528710
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DOI: 10.1159/000528710