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This paper presents an investigation of the specic nature of environmental inuences on the evolution of human skin coloration. It builds on previous research (Jablonski and Chaplin, 2000) in which
a clear causal relationship between average annual
ultraviolet radiation (UVR) and skin reectance was
established.
PREVIOUS RESEARCH ON GLOBAL
DISTRIBUTION OF ENVIRONMENTAL FACTORS
Few workers have looked systematically at the
environmental factors identied as possible inuences on skin color. Most have used latitude as a
surrogate for UVR because it has a strong correlation with skin color, with an r-value of around 0.8
(Relethford, 1997; Roberts, 1977; Roberts and
Kahlon, 1976; Tasa et al., 1985). Latitude is a poor
surrogate for UVR because other environmental
variables are correlated with it. Further, the correlation between UVR and latitude will be different
between the hemispheres due to orbital effects on
solar intensity (Chaplin and Jablonski, 1998;
Relethford, 1997). The rst use of direct measurements of UVR collected using a satellite (Jablonski
and Chaplin, 2000) showed the highest correlation
to annual average UVR and skin reectance at rvalues above 0.95, but no other environmental factors were taken into account.
Walter (1958) was the rst to use an estimate of
UVR to demonstrate a correlation between UVR and
skin color. He obtained his estimate by calculating
UVR at the Equator and correcting for latitude,
293
Third are hypotheses that involve non-UVR agencies, such as forest cover, altitude, temperature, and
precipitation. These variables have been considered
in theories of skin coloration and were included in
this study. It was suggested that the intensity of
melanin pigmentation is related to the need for camouage in forest environments (Cowles, 1959). A
number of theories revolve around thermoregulation (Baker, 1958; Hamilton and Heppner, 1967;
Razi et al., 1980) or the prevention of cold injury
(Candler and Ivey, 1997; Post et al., 1975; Steegmann, 1967; Taylor, 1992). Precipitation was included because Glogers rule states that warmblooded animals living in warm and humid places
are more heavily pigmented than those in cool, dry
areas. Precipitation may also be indicative of the
need for shelter-seeking and cultural modication of
the UVR regime, leading to pleiotropic decline in
skin coloration (Brace, 1963; Daniels, 1964; Deol,
1975). This led to formulation of the hypothesis that
variables other than direct UVR are responsible for
skin reectance and are only correlated to UVR or
latitude by solar-driven processes such as vegetation, temperature, and precipitation.
MATERIALS
Skin reflectance data
The skin reectance data set used in this study
was essentially the same as that of Jablonski and
Chaplin (2000), but with a few new data points
added (see Appendix). The database comprises samples for reectance at 425 nm (blue lter), 545 nm
(green lter), and 685 nm (red lter), at a site on the
upper inner arm.
Environmental data
The UVR data used in this study represent the
relative daily areal exposure to the ultraviolet minimum erythemal dose (UVMED), i.e., the amount of
UVR effective in causing a barely perceptible reddening of light skin. The values for UVMED used in
this study were derived from readings taken by the
NASA total ozone mapping spectrometer (TOMS)
that was own aboard the Nimbus-7 satellite between 1978 1993 (Herman and Celarier, 1996). In
measuring UVR strength, all attenuating inuences
(humidity, rainfall, temperature, or elevation) were
already discounted, so that any effect that these
variables have, as modeled in this study, was the
result of their own action rather than how they were
affecting UVR transmittance. The treatment of
TOMS data was described in Jablonski and Chaplin
(2000). From the NASA TOMS data set, the following UVMED readings were derived: annual mean
for maximum, minimum, and range, and seasonal
average for winter, spring, summer, and autumn.
Data for global environmental variables were
taken from the unied data set, ArcAtlas: Our Earth
(ESRI, 1996). These data were provided at scales of
25 million to 1, and 10 million to 1, for selected
294
G. CHAPLIN
Nonlinear investigation
Further, graphical techniques were run to investigate the relationships as identied in the nal
regression model. The nonlinear techniques consisted of local regression, (loess) regression, and
spline tting. Smoothing splines that minimize a
penalized residual sum of squares was done using
four degrees of freedom (d.f.), because this will not
force oversmoothing (S-Plus, 2000).
Analysis of categorical variables
To investigate categorical variables (relief and
land-cover type), boxplot techniques were used, and
an analysis of variance was performed. The latter
looked for variation that could be further explained
by inclusion of these categorical variables.
Prediction from final regression model
The nal regression model was used to predict
skin reectance for each color lter. These predicted
skin colors were then mapped. The colors of the map
were produced from the multiple regression formulae for each color lter. This map was, in effect, an
intersection of all the input maps. The legend was
created by assigning a color to each unique combination derived from the values of the red, green, and
blue (RGB) lters, to derive a RGB color value using
the GIS programming language. These colors are
realistic approximations of the true color of skin,
within the limits of the color space of the particular
medium in which it is viewed.
The large number of features and color combinations meant that this map needed generalization for
visualization of any trends in predicted skin color. It
was generalized as follows: cells were rst dissolved
until they had unique combinations of red, green,
and blue values. Then the lowest and highest values
in the distribution tail, that had only a few cells
each, were unioned into a single shape. Finally,
shapes that were almost identical in color were
merged with their neighbors to further reduce the
number of shapes. By this method, the number of
shapes was reduced to just 17 multipart shapes. The
generalization sought to reach a balance between
equal area representation and equal interval representation. The colors in the legend are also the actual skin colors as dened above. This map was
suitable for visualization techniques (Cleveland,
1993), but not for statistical analysis.
RESULTS
Exploratory data analysis
Outliers that were detected were examined, and
where their explanation could be understood in
terms of migration, genetic admixture, or incorrect
placement on the map, they were removed from
further analysis. These consisted of very few cases.
The rst were the Ammassalimiut (Inuit from
Greenland) and Yemeni Jews; these were removed
295
296
G. CHAPLIN
TABLE 1. Pearson correlation coefficients, for skin reflectance measures to environmental variables1
d.f.
UVR Average
UVR Autumn
UVR Winter
UVR Minimum
UVR Spring
UVR Summer
UVR Maximum
UVR Range
Latitude
Annual precipitation
Summer precipitation
Winter precipitation
Average solar radiation
Snow
Frost-free days
Average temperature,
annual
Average temperature,
summer
Average temperature,
winter
Raw by cell
for Red Filter
E685.609
Trimmed
by cell for
Red Filter
E685.609
Melanin
reectance by
cell, including
estimates
Melanin
reectance
grouped by
population
Tissue
reectance
grouped by
population
Blood
reectance
grouped by
population
3,706.0
0.66***
0.76***
0.73***
0.76***
0.18***
0.31***
0.16***
0.54***
0.75***
0.16***
0.05***
0.26***
0.27***
0.35***
0.36***
0.42***
3,674.0
0.69***
0.78***
0.75***
0.78***
0.19***
0.33***
0.16***
0.56***
0.76***
0.16***
0.05**
0.26***
0.30***
0.35***
0.38***
0.43***
4,355.0
0.66***
0.76***
0.72***
0.76***
0.17***
0.33***
0.16***
0.56***
0.75***
0.15***
0.05**
0.24***
0.30***
0.31***
0.36***
0.41***
96.0
0.82***
0.84***
0.83***
0.82***
0.65***
0.66***
0.63***
(0.13)
0.83***
0.23***
0.32***
0.35***
0.63***
0.46***
0.59***
0.67***
70.0
0.89***
0.86***
0.86***
0.82***
0.79***
0.78***
0.79***
(0.14)
0.92***
0.28***
0.38***
0.46***
0.78***
0.48***
0.77***
0.82***
70.0
0.90***
0.92***
0.93***
0.90***
0.75***
0.73***
0.74***
(0.02)
0.93***
0.29**
0.40***
0.44***
0.71***
0.54***
0.74***
0.84***
0.09***
0.09***
0.08***
0.38***
0.61***
0.55***
0.55***
0.55***
0.53***
0.71***
0.74***
0.82***
Note differences in degrees of freedom of different samples due to differing numbers of missing values. Three left-hand columns show
effect, on correlation in data, of having raw data trimmed by having ve outliers removed and for data converted from different
reectometers. Three right hand columns are for data summarized by population. All correlations are very highly signicant, except
UVR Range.
* P 0.05.
** P 0.01.
*** P 0.0001.
Latitude, UVR Autumn, UVR Winter, UVR Minimum, and UVR Average were all much more highly
correlated to skin reection measures than were
UVR Maximum, UVR Summer, UVR Spring, and all
other environmental factors.
Principal components analysis
Principal components analyses were performed on
both the trimmed by cells and the trimmed by
population data sets. The results were comparable.
The diagnostic metrics for the more signicant
components, for the grouped by population PCA,
are given in Table 2. The biplot of this PCA (Fig. 1)
represented both the original variables and the
transformed observations on the rst two principal
components axes. The original variables were represented as arrows that graphically indicated the proportion of the original variance explained by the rst
two principal components.
The rst component comprised the major UVR
measures and also was negatively correlated to skin
reectance variables, so that an increase in UVR
covaries with a decrease in reection, which means
a darker skin color. The direction of the arrow
showed the relative loadings of the variables on PCA
components 1 and 2 (S-Plus, 2000). In Figure 1, the
arrows span much of the data spread, showing that
the two rst components accounted for much of the
variation (cumulative proportion 0.821). The temperature measure arrows were all shorter than
those for other variables, indicating that their covariance was nested within the variation explained
by the UVR measures.
The variables UVR Winter (UV Win), UVR Autumn (UVAut), UVR Minimum (UV Min), and Winter Temperature (Win Tmp) were highly collinear.
These four variables showed the greatest negative
covariance with skin reectance measures in the
biplot graph (Fig. 1 and Table 2). These four were
offset by the number of days with snow on the
ground (Snow) and by Winter Precipitation
(Winavpcp), which was inclusive of snow. That is,
these two variables positively covaried with skin
reectance, with which they grouped.
Another group of variables consisted of Total Average Solar Radiation, UVR Maximum, UVR Summer, and Summer Temperature, that lay to the left
of the second component axis, and that thereby also
correlated with darker skin colors.
The variables Frost-Free Days, UVR Average, and
Annual Temperature, which were averaging measures of the other variables, lay between the groups
from maximum and minimum measures, as would
be expected.
The loadings showed that most of the variance
(65%) was accounted for by the interaction of skin
reectance and snow to solar processes. The next
component (17%) showed an axis whereby solar processes were offset by increasing precipitation, with
increases in UVR range being associated with
297
3.61
0.65
0.65
0.24
0.26
0.26
0.25
0.25
0.24
0.47
0.27
0.45
0.39
0.15
0.25
0.18
0.25
0.26
0.21
0.24
0.26
0.26
0.24
1.84
0.17
0.82
0.17
0.15
0.19
0.20
0.21
0.39
0.45
0.44
0.22
0.21
0.22
0.17
0.14
0.16
1.08
0.06
0.88
0.91
0.04
0.92
0.19
0.12
0.13
0.15
0.14
0.20
0.10
0.32
0.44
0.34
0.63
0.24
0.23
0.22
0.29
0.20
0.20
0.25
0.23
0.16
0.11
298
G. CHAPLIN
TABLE 3. Table of diagnostic statistics of
multiple regression equations
Value
Standard
error
t-value
Intercept
41.7231
2.4860
16.7834
0.0001
UVAUT
0.0457
0.0080
5.7156
0.0001
UVMAX
0.0365
0.0072
5.0912
0.0001
SUMAVPCP
0.0014
0.0004
3.2372
0.0019
WINAVPCP
0.0006
0.0011
0.5947
0.5540
Residual standard error: 4.396 on 67 degrees of freedom
Multiple r2: 0.8327
F-statistic: 83.39 on 4 and 67 degrees of freedom, P 0.0001
Outliers: Japan Hidakka, Eastern Nepal
Green Blood Reectance UVR Autumn UVR Maximum
Summer Precipitation Winter Precipitation
Coefcients
Value
Standard
error
t-value
Intercept
43.5006
2.2443
19.3825
0.0001
UVAUT
0.0758
0.0071
10.7087
0.0001
UVMAX
0.0156
0.0065
2.4072
0.0188
SUMAVPCP
0.0011
0.0004
2.9080
0.0049
WINAVPCP
0.0018
0.0009
1.8976
0.0621
Residual standard error: 3.977 on 67 degrees of freedom
Multiple r2: 0.8872
F-statistic: 131.7 on 4 and 67 degrees of freedom, p 0.0001
Outliers: Eastern Nepal, Yemen
Red Melanin Reectance UVR Autumn UVR Maximum
Summer Precipitation Winter Precipitation
Coefcients
Value
Standard
error
t-value
Intercept
63.0801
3.8472
16.3963
0.0001
UVAUT
0.1014
0.0114
8.9182
0.0001
UVMAX
0.0035
0.0111
0.3186
0.7507
SUMAVPCP
0.0001
0.0006
0.2310
0.8178
WINAVPCP
0.0028
0.0014
1.9205
0.0579
Residual standard error: 7.342 on 92 degrees of freedom
2
Multiple r : 0.7283
F-statistic: 61.66 on 4 and 92 degrees of freedom, P 0.0001
Outliers: Australia Darwin, Cambodia, Namibia, and Yemen
299
Fig. 3. Map of skin color reectance predicted from multiple regression. Map is generalized to reduce number of polygons.
300
G. CHAPLIN
values and much variation about the line. The darkest people seemed to be those who have inhabited
their homelands for very long periods of time, e.g.,
Australian Aborigines and Ethiopians. As these populations are those that seem to be resident the longest, it can be concluded that the nal increments of
skin darkening appear to have required longer to
evolve. The darkest populations have similar reectance values, so that there may be a point where it is
not biologically possible to darken further, and approaching that point will slow the rate of adaptation.
The prediction that there is reduced skin reectance in the Tissue (blue) Reectance range due to
thickening of the stratum corneum caused by UVR
is supported. Tissue (blue) Reectance was slightly
more correlated with UVR Maximum than were the
other reectance measures, a nding that helps to
explain the thickened stratum corneum found in
African populations, although a thickened stratum
corneum would provide little photo-protection, as
can be seen in people with pigment disorders (Robins, 1991).
The second set of hypotheses, concerning theories
of metabolic processes relating direct solar radiation
to skin reectance, nds the strongest support. The
main original nding of this study was that the
evolution of skin color reectance could be almost
fully modeled as a linear effect of UVR in the autumn alone (UVR Autumn with the absorption peak
of Blood (green) Reectance; r 0.927, d.f. 70, P
0.0001). This is also true for all measures of the
period of lower UVR (those with an increased proportion of longer-wavelength UVR): UVR Autumn,
UVR Minimum, and UVR Winter. These measures
were all more important at explaining skin reectance variation than were the peak UVR measures.
This very strong nding gives considerable support
to the nutrient hypothesis. Folate photolysis leading
to folate deciency was proposed as an agent decreasing tness and promoting skin darkening in
lower latitudes near the Equator out to beyond the
tropics (Jablonski, 1992, 1999; Jablonski and Chaplin, 2000). Hypovitaminosis D was found to be the
agent in higher latitudes where there are prolonged
periods without vitamin-D-promoting UVR (UVB)
(Holick, 1995; Jablonski and Chaplin, 2000; Loomis,
1967). High levels of wintertime UVA without UVB,
as found in middle latitudes, would destroy both
vitamin D and folate by photolysis.
This extremely well-tted linear model explains
why latitude was such a good surrogate for UVR,
because Latitude and UVR Autumn were extremely
highly correlated (r 0.967, d.f. 102, P 0.0001),
as was Latitude and UVR Average (r 0. 937, d.f.
102, P 0.0001), but not Latitude and UVR Summer (r 0.767, d.f. 102, P 0.0001). Further, this
indicates that latitude is still useful for human ecological modeling.
Hypotheses about variables that are only correlated to UVR by latitude or through solar-driven
processes such as vegetation, temperature, and pre-
301
302
G. CHAPLIN
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skin colour. Ann Hum Biol 3:1122.
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Walter H. 1958. Der Zusammenhang von Hautfarbenverteilung und Intensitat der ultravioletten Strahlung. Homo 9:1
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Webb AR, Holick MF. 1988. The role of sunlight in the cutaneous
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Ref.
Various
Huachipaeri
Mexico
Shipibo
Slovakia
United Kingdom
United Kingdom & Northern Ireland
United States, Florida
United States, Oklahoma
United States, Western
1
2
2
2
3
4
4
5
5
6
Tissue
Blood
12.8
19.3
15.75
28.8
42.3
42.3
19.1
27.1
21.4
36.9
45.6
45.6
19.1
33.3
Melanin
How mapped
48.6
48.1
53.2
62.5
69
68.9
47.7
50.4
Various
Estimated from atlas
Estimated as sierra region
Estimated from atlas
Western region of Slovakia
Estimated from atlas
Estimated from atlas
Estimated from atlas
Estimated from atlas
Estimated from atlas
References (Ref.) for data: 1, Jablonski and Chaplin, 2000; 2, Ebbinghaus, 1966; 3, Sefcakova, 1986; 4, Little and Wolff, 1981; 5,
Pollitzer et al., 1970; 6, Pawson and Petrakis, 1975. Full references in Chaplin (2001).