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Nutrition Research 31 (2011) 429 – 435

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Diets of modern hunter-gatherers vary substantially in their carbohydrate

content depending on ecoenvironments: results from an
ethnographic analysis
Alexander Ströhle⁎, Andreas Hahn
Nutrition Physiology and Human Nutrition Unit, Institute of Food Science and Human Nutrition, Leibniz University of Hannover,
Hannover D-30167, Germany
Received 18 March 2011; revised 30 April 2011; accepted 2 May 2011

Abstract

In the past, attempts have been made to estimate the carbohydrate contents of preagricultural
human diets. Those estimations have primarily been based on interpretations of ethnographic data of
modern hunter-gatherers. In this study, it was hypothesized that diets of modern hunter-gatherers
vary in their carbohydrate content depending on ecoenvironments. Thus, using data of plant-to-
animal subsistence ratios, we calculated the carbohydrate intake (percentage of the total energy) in
229 hunter-gatherer diets throughout the world and determined how differences in ecological
environments altered carbohydrate intake. We found a wide range of carbohydrate intake (≈3%-50%
of the total energy intake; median and mode, 16%-22% of the total energy). Hunter-gatherer diets
were characterized by an identical carbohydrate intake (30%-35% of the total energy) over a wide
range of latitude intervals (11°-40° north or south of the equator). However, with increasing latitude
intervals from 41° to greater than 60°, carbohydrate intake decreased markedly from approximately
equal to 20% to 9% or less of the total energy. Hunter-gatherers living in desert and tropical
grasslands consumed the most carbohydrates (≈29%-34% of the total energy). Diets of hunter-
gatherers living in northern areas (tundra and northern coniferous forest) contained a very low
carbohydrate content (≤15% of the total energy). In conclusion, diets of hunter-gatherers showed
substantial variation in their carbohydrate content. Independent of the local environment, however,
the range of energy intake from carbohydrates in the diets of most hunter-gatherer societies was
markedly different (lower) from the amounts currently recommended for healthy humans.
© 2011 Elsevier Inc. All rights reserved.
Keywords: Evolutionary ecology; Carbohydrate intake; Hunter-gatherers; Paleolithic diet; Ethnographic Atlas
Abbreviations: P:A energy subsistence ratios; plant-to-animal energy subsistence ratios.

1. Introduction research activities of paleoanthropologists has been the study

of the dietary habits of our prehistoric ancestors who lived
Because diet represents a fundamental environmental
during the Plio-Pleistocene period (from ≈5 million years
factor in evolution via natural selection, knowledge about the
ago to ≈12 000 years ago) [2,13-28]. Since the publication
ancestral diet of Homo sapiens is important to a proper
of Eaton and Konner's [29] locus classicus “Paleolithic
understanding of the paleobiology and evolutionary ecology
Nutrition” and Williams and Nesse's [30] “The Dawn of
of our species [1-12]. Accordingly, one of the fundamental
Darwinian Medicine,” the diets of our ancestors who lived in
the Paleolithic period (“Old Stone Age,” the archaeological
⁎ Corresponding author. Tel.: +49 511 762 5093; fax: +49 511 762 5729. era that began ≈2.6 million years ago) have also become an
E-mail address: alexander.stroehle@lw.uni-hannover.de (A. Ströhle). important field of discourse in nutrition and nutritional
0271-5317/$ – see front matter © 2011 Elsevier Inc. All rights reserved.
doi:10.1016/j.nutres.2011.05.003
430 A. Ströhle, A. Hahn / Nutrition Research 31 (2011) 429–435

medicine [31-41]. Indeed, some researchers expect that activities could also include the collection of small fauna (eg,
knowledge about the dietary patterns of these Paleolithic invertebrates, insects, and eggs). Because of the similarity in
hunter-gatherers can serve as models for health maintenance the mean energy density of animal (7.24 kJ/g) and plant
and disease prevention recommendations [42,43]. foods (6.99 kJ/g) consumed by hunter-gatherers, we also
Because absolute and relative carbohydrate intakes affect followed the assumption of former studies [31,47,50] that the
human health in many ways [44-46], the carbohydrate P:A subsistence ratio based on weight was identical to the
content (as percentage of energy) of preagricultural human P:A ratio based on energy.
diets is of considerable interest. Therefore, various attempts
2.2. Computation of the carbohydrate intake (percentage of
have been undertaken to estimate the carbohydrate content
energy) in the current model
of such diets [29,42,47-49]. Those estimations have been
based on ethnographic records of historically studied The estimations of the carbohydrate intakes were based
hunter-gatherer populations. In the most recent ethnographic on a previous model [47]. According to that model, the
compilation of plant-to-animal (P:A) subsistence patterns in macronutrient composition (percentage of energy) of
hunter-gatherer populations throughout the world, Cordain hunter-gatherer diets could be computed with the
et al [47] estimated that the most plausible percentage of following equation:
total carbohydrate energy ranged from 22% to 40%. The
subsistence patterns of modern hunter-gatherers, however, AðWa × XÞ þ PðWp × XÞ¼ D; ð1Þ
have shown wide ranges, which were dependent on latitude
where A and P denote the mean energy content of wild-
and ecological environments [31,32,47,50,51]. In this study,
animal foods (7.24 kJ/g) and wild plants (6.99 kJ/g),
it was hypothesized that diets of modern hunter-gatherers
respectively. In addition, Wa and Wp represent the assumed
showed substantial variation in their carbohydrate content
proportions (weight percentage) of animal and plant foods,
depending on ecoenvironments. Thus, we aimed at
respectively, X denotes the total mass (in grams) required to
exploring how latitude and variable ecological environ-
provide any given amount of food energy, and D represents
ments alter dietary carbohydrate intake (expressed as
daily energy intake (in kilojoules per day).
percentage of total energy intake) in 229 environmentally
Because the current work only focused on carbohydrate
diverse, modern hunter-gatherer populations throughout the
intake and animal foods consumed by hunter-gatherers
world and presented an update of the original estimations by
contain negligible amounts of carbohydrates, the following
Cordain et al [47].
equation was used to compute carbohydrate intake in hunter-
gatherer diets:
2. Methods and materials
Ctotal ¼ Wp × Cmean × 100; ð2Þ
Taking recent ethnographic data of P:A energy subsis-
where Ctotal denotes the total energy percentage from
tence ratios of 229 worldwide forager populations [52,53],
carbohydrates and Cmean represents the mean carbohydrate
we computed dietary carbohydrate intakes (percentage of
value (62% of the total energy), which was derived from a
energy) (a) of the entire sample population as well as (b)
large wild-plant–food database consisting of 829 plant
subgroups based on latitude. Furthermore, we calculated (c)
species [55]. Wp represents the assumed proportion of plant
likely dietary carbohydrate intake by primary ecological
foods consumed by weight (weight percentage).
environments in hunter-gatherer societies (n = 63) where
data were documented. The procedures were carried out as
detailed below. Table 1
Nutritional characteristics of plant foods implemented in the wild-plant–
2.1. Ethnographic data food database used in this work
Plant food Individual Energy Carbohydrate
The original data of P:A energy subsistence ratios in
type plants (n) (kJ/100 g) content (g/100 g)
relation to latitude and local ecological conditions that were
Fruit 317 397 ± 241 21 ± 14
used in this study came from a revised analysis [52] of the
Tubers 86 406 ± 213 22 ± 12
Ethnographic Atlas [53], a verified [54] cross-cultural Other seeds 74 1238 ± 531 45 ± 22
index that summarized various ethnographic data of 1267 Nuts 74 1280 ± 838 30 ± 20
world societies, which was carried out by Cordain et al [47]. Roots 51 392 ± 363 17 ± 13
In their analysis, Cordain et al [47] identified 229 societies Acacia seeds 55 1472 ± 266 48 ± 19
Bulbs 30 677 ± 368 38 ± 23
whose economic subsistence depended exclusively on
Leaves 28 256 ± 198 12 ± 10
hunting (including trapping and fowling), gathering (wild Flowers 16 354 ± 184 15 ±7
plants and small land fauna), and fishing (including shell Miscellaneous 14 382 ± 357 15 ± 21
fishing). In accordance with our previous analyses [31,50], Dried fruit 7 1215 ± 316 59 ± 22
the present study assumed that gathered food included only Gums 2 996 ± 566 84
plant foods, whereas Murdock [53] stated that gathering Data taken from Brand Miller and Holt [55].
 A. Ströhle, A. Hahn / Nutrition Research 31 (2011) 429–435 431

Table 1 shows that there was substantial variation in both Table 3

energy density and carbohydrate content among the plant Effects of latitude on carbohydrate intake (% of energy) for 229
hunter-gatherer diets
food categories in the database used to determine Cmean ,
which affected the calculation of Ctotal. As Cordain et al [47] Degrees from Absolute Relative Carbohydrate intake
the equator frequency frequency (percentage of
pointed out in a previous report, “studies of hunter-gatherers
(latitude) (no. of societies) (percentage energy/day)
have shown that the various categories of plant foods were of societies)
not randomly gathered, but were collected with a general
0-10 6 2.6 22-28
prioritization that maximized the rate of energy capture 11-20 15 6.6 29-34
relative to energy expenditure; this pattern of gathering is 21-30 11 4.8 29-34
predicted by the ‘optimal foraging theory’” [56-58]. 31-40 64 27.9 29-34
Therefore, a realistic plant-food database should maintain a 41-50 75 32.8 16-22
51-60 38 16.6 10-15
weighting that is generally predicted by the optimal foraging
N60 20 8.7 ≤9
theory [57,58]. The following percentages (percentage of the
total number of food items) of plant food were used in the
present study: fruits, 41%; seeds, 17%; tubers, 11%; nuts, in relation to latitude. According to our calculations, forager
10%; roots and bulbs, 12%; dried fruit, flowers, gums, and diets were characterized by an identical and moderate
miscellaneous plant parts, 9%. Based on this weighting, the carbohydrate intake (30%-35% of the total energy) over a
plant food carbohydrate content was assumed to be 62% of wide range of latitude intervals (11° to 40° north or south of
the total energy [47]. the equator). However, with increasing latitude intervals
Using the Ethnographic Atlas P:A energy subsistence from 41° to greater than 60° north or south of the equator,
ratios in section 2.1 and the algorithm in section 2.2, we were carbohydrate intake decreased markedly from approximately
able to calculate the carbohydrate intakes (percentage of equal to 20% to 9% or less of the total energy.
energy) in all hunter-gatherer diets.
3.3. Calculations of carbohydrate intake in relation to local
3. Results ecological conditions
3.1. Intercultural variation in carbohydrate intake In addition to computing the frequency distributions of
carbohydrate intake in relation to latitude, we calculated
Using data generated from Equation 2, Table 2 shows the
carbohydrate intake in relation to primary living environ-
frequency distribution of carbohydrate intake in the 229
ments in a subsample of 63 societies for whom appropriate
hunter-gatherer societies. Table 2 shows that there was a wide
data were available. The results of this analysis are
range of interval values for carbohydrate intake (median and
summarized in Table 4. Our findings illustrated that the
mode, 16%-22% of the total energy). As P:A ratios increased
diets of hunter-gatherers living in northern areas (tundra and
from 5:95 to 85:15, carbohydrate intake increased from 3% of
northern coniferous forest) were characterized by a very low
the total energy to 53% of the total energy.
carbohydrate content (≤15% of the total energy). There have
3.2. Calculations of the carbohydrate intake in relation only been 2 other ecological environments—temperate
to latitude grassland and tropical rain forest—that have been character-
ized by societies consuming similarly low amounts of
Table 3 summarizes the findings of the computations of carbohydrates (≈20% of the total energy). The largest
carbohydrate intake for each of the 229 hunter-gatherer diets amount of carbohydrates, 29% to 34% of the total energy,
was consumed by hunter-gatherers living in desert grasslands
Table 2 and tropical grasslands.
Frequency distribution of the calculated carbohydrate intake of the diets of
229 hunter-gatherer societies
P:A ratio Absolute Relative frequency Carbohydrate intake 4. Discussion
by class frequency (percentage of (percentage of
Because dietary intake of carbohydrates has considerable
interval (%) (no. of societies) societies) energy/d)
effects on human health [44-46], intercultural variations in
≤5:≥95 11 4.8 ≤3
the carbohydrate content of hunter-gatherer diets are very
6-15:85-94 35 15.3 4-9
16-25:75-84 42 18.3 10-15 interesting. Therefore, we estimated the likely carbohydrate
26-35:65-74 45 19.7 16-22 intake of 229 modern hunter-gatherer populations living at
36-45:55-64 35 15.3 23-28 different latitudes in diverse ecological environments.
46-55:45-54 30 13.1 29-34
56-65:35-44 23 10.0 35-40 4.1. Calculation of carbohydrate intake in the entire group
66-75:25-34 6 2.6 41-46
76-85:15-24 2 0.9 47-53 According to the wide ranges of P:A subsistence patterns
Median 16-22 (gathered plant food, 0%-5% to 76%-85%; animal food,
Mode 16-22
16%-25% to 86%-100%), our analysis of the diets consumed
432 A. Ströhle, A. Hahn / Nutrition Research 31 (2011) 429–435

by hunter-gatherer societies showed that there was a large Table 5

variation in the carbohydrate content of the different diets,
which ranged from approximately 3% of the total energy to
approximately 50% of the total energy. Interestingly, most
hunter-gatherer diets (approximately 85%) were character-
ized by a relatively low carbohydrate intake (b35% of the
total energy), which reflected the high reliance on animal-
based foods of most hunter-gatherer societies. For example,
of the 229 hunter-gatherer populations examined, Cordain
et al [47] found that the median subsistence dependence on
animal food was 66% to 75%. Similarly, an analysis of a
recent ethnographic compilation of P:A subsistence patterns
showed that most members of historically studied hunter-
gatherer societies consumed high amounts of animal-source
food (median P:A ratio, 35:65) [51]. If we interpreted the
range of percentages of energy consumed as carbohydrates
in the diets of hunter-gatherer societies from the perspective
of contemporary levels (see Table 5), 73% (n = 168) of diets
would be classified as “low-carb,” and only approximately
26% (n = 59) would be classified as “moderate-carb” [59].
Therefore, it is not surprising that the range of percentages of
energy from carbohydrates in the diets of most hunter-
gatherer societies is markedly different from the amounts
recommended for healthy individuals (see Table 6). With
respect to human health, however, there has been increasing
evidence from observational and clinical studies suggesting
that diets with restricted carbohydrates do not cause harm per
se. Interestingly, carbohydrate-restricted diets have been
shown to have favorable effects on blood lipids [63],
coronary heart disease risk [64], and body weight [59].
Because many hunter-gatherer societies consumed low
levels of carbohydrates, this fact could explain why these
populations were relatively free of many chronic and
nutrition-related degenerative diseases, such as obesity,
type 2 diabetes mellitus, and coronary heart disease [32,65].
We must take into account, however, that there was a
variety of less Westernized humans worldwide (eg, horti-
culturalists and simple agriculturalists) who ingested high-
carbohydrate diets (approximately 70% of the total energy)
without having those diseases [32,66]. In this context,
Definitions for low-carbohydrate diets [59]
Absolute carbohydrate Relative carbohydrate
content (g/d) content (percentage
of energy/d)
Low-carbohydrate- b50 b10
ketogenic diet
Low-carbohydrate 50-b130 10-b25
diet
Moderate ≥130 26-45
carbohydrate
diet
Ludwig and Jenkins [67] may have been correct when they
stated that “humans can probably do well over the long term
by consuming diets that vary widely in macronutrients, as
long as adequate attention is paid to nutrient quality.”
Indeed, with respect to carbohydrate quality, all traditional
diets mentioned above differ from modern Western diets.
Except for honey—available seasonally and consumed in
high amounts (yearly average up to 14%-18% of total daily
energy) by some hunter-gatherers (eg, the Hadza and the
Ache) [68,69]—the plant component of the diets of hunter-
gatherers [70], horticulturalists [71,72], and simple agricul-
turalists [73] has been based on minimally processed plant
foods with low glycemic index (GI) values and high levels
of dietary fiber [55,71-73]. In contrast, foods characterized
by a high GI, such as refined sugars and grains, which
currently supply approximately 40% of the total energy in a
typical US diet [74], have rarely been consumed by non-
Westernized individuals. There has been increasing evi-
dence suggesting that high-fiber, low-GI diets are associated
with a lower risk of coronary heart disease and type 2
diabetes [75]. Therefore, the quality of carbohydrates rather
than the amount of carbohydrates may partly explain the
fact that hunter-gatherers were relatively free of nutrition-
related degenerative diseases. This interpretation is
in accordance with findings from a meta-analysis of 37
prospective observational studies, which showed that “there
were more positive associations of greater magnitude

Table 4
Effect of different ecological environments on the ratio of plant-food energy intake to animal-food energy intake (P:A ratio) and the corresponding dietary
carbohydrate intake (percentage of energy) for 63 hunter-gatherer diets
Characterization of the P:A ratio by Absolute frequency Relative frequency Carbohydrate intake
ecological environments class interval (%) (no. of societies) (percentage of societies) (percentage of energy/d)
Tundra, northern areas 6-15:85-94 6 9.5 3-9
Northern coniferous forest 16-25:75-84 14 22.2 10-15
Temperate forest, 36-45:55-64 6 9.5 23-28
mostly mountainous
Desert grasses and shrubs 46-55:45-54 11 17.5 29-34
Temperate grassland 26-35:65-74 11 17.5 16-22
Subtropical bush 36-45:55-64 2 3.2 23-28
Subtropical rainforest 36-45:55-64 4 6.3 23-28
Tropical grassland 46-55:45-54 4 6.3 29-34
Monsoon forest 36-45:55-64 2 3.2 23-28
Tropical rainforest 26-35:65-74 3 4.8 16-22
 A. Ströhle, A. Hahn / Nutrition Research 31 (2011) 429–435
Table 6
Nutrition recommendations with respect to dietary carbohydrate intake a
NNR D-A-CH
Carbohydrate intake 55 (50-60) 50
(% of the total energy)
NNR indicates Nordic Nutrition Recommendations; D-A-CH, German Nutrition Society; HCN, Health Council of the Netherlands; WHO, World Health
Organization; and FAO, Food and Agriculture Organization; IOM, Institute of Medicine of the National Academy USA (Food and Nutrition Board); AMDR,
acceptable macronutrient distribution.
a
Composed according to Pavlovic et al [60], Spaaij and Pijls [61], and DGE et al [62].
between GI and chronic disease than between GL
(glycemic load) and chronic disease” [75].
4.2. Calculation of carbohydrate intake varying with
latitude and local ecology
Because latitude has been shown to affect P:A subsistence
ratios [47,50], we calculated the carbohydrate intake of
hunter-gatherers based on latitude. To our knowledge, we
were the first to use this approach. In general, we found no
clear relationship between carbohydrate intake and different
ecoenvironments worldwide. In fact, forager diets were
characterized by an identical and moderate carbohydrate
intake (in the range of 30%-35% of the total energy) over a
wide range of latitude intervals (11°-40° north or south of the
equator). Our evaluation of carbohydrate intake as it related
to the ecological niche of a subsample of 63 hunter-gatherer
societies showed that only hunter-gatherers living in desert
grasslands, shrubs, and tropical grasslands consumed a
moderate-carb diet (29%-34% of the total energy). Further-
more, a more detailed analysis of the ethnographic data
showed that the threshold above which the diets of the 229
hunter-gatherers became “low-carb,” emerged at greater than
40° north and south of the equator. This finding reflected that
hunter-gatherers who lived in higher latitudes ingested
animal-based diets (P:A ratio, b26:N65), which was mainly
a consequence of temperature effects on primary biomass
(a measure of the productivity of a given habitat) [51].
From the view of evolutionary ecology, this finding was
of special interest because it is known that historically
studied hunter-gatherers do not represent “living fossils”
[76] of our hominid ancestors who lived throughout the
Paleolithic period with different climatic and ecological
conditions. Because the evolution to anatomically modern
humans took place in East Africa (latitude of approximately
0°) 100 000 to 200 000 years ago [77], it seems plausible
that, on an average, a moderate-carb diet of approximately
35% of the total energy was the norm of our early ancestors
who lived in the African savannah. This interpretation was
in accordance with estimations made by other investigators.
For example, Eaton [43] and Eaton et al [42] suggested that
those African ancestors obtained approximately 35% to
40% of their dietary energy from carbohydrates.
Recently, in their sensitive and impressing analysis,
Kuipers et al [78] estimated that an East African Paleolithic
diet results in a moderate-carb intake of approximately 40% of
433
HCN Euro diet WHO/FAO IOM (AMDR)
N40 N55 55-75 45-65
the total energy (median). Interestingly, the estimated
medians of the carbohydrate contents of Paleolithic hunter-
gatherers diets were identical for multiple foraging strategies
(meat-based selective savannah diet, 40% energy; fish/meat-
based selective savannah/aquatic diet, 40% energy; fish-based
nonselective aquatic diet, 39% energy) [78]. The calculated
moderate-carb intake of approximately 40% energy in the
models of Kuipers et al [78] is higher but comparable with the
estimated intake of approximately 35% of the total energy in
the model presented in our work. In contrast, the switch to a
low-carb diet (b25% of the total energy) seems to have taken
place late in human evolution (ie, between 46 000 and 7 000
years ago) when our ancestors settled in higher latitude
environments, such as subartic Eurasia, northern Europe,
North America, Greenland, and Siberia [79,80].
4.3. Limitations of the analysis
As we have stated elsewhere [31,47], the most important
limitation of this ethnographic analysis was that ethnograph-
ic data are subjective in nature. Therefore, the assigned P:A
subsistence economy ratios and the carbohydrate content of
hunter-gatherer diets were approximations rather than
precise measurements [81]. However, a compilation of
more exact quantitative dietary studies, which were carried
out on some of the world's hunter-gatherer societies [9,82],
showed a high similarity to the ethnographic data analyzed in
the present study.
A further limitation of our analysis was the assumption
that gathered food would only include plant foods. Therefore,
the calculations presented here might have overestimated the
carbohydrate content of the diets of modern hunter-gatherers.
It should also pointed out that the ethnographically
described P:A ratios of modern hunter-gatherer diets may
represent “a biased sample” [51] of the diets consumed by
our ancient ancestors who lived in the Pleistocene. For
example, many modern hunter-gatherers “have been greatly
influenced by their association with more powerful agricul-
tural societies. It has even been suggested that Holocene
foragers represent a new niche that appeared only with the
climatic changes and faunal depletion at the end of the last
major glaciation […]. The greatest obstacle to using foragers
as analogs of our ancient ancestors is that virtually all
foragers in the ethnographic record have complex technol-
ogy compared to premodern hominins” [51]. Because of this,
the carbohydrate intake of modern hunter-gatherers, as
434 A. Ströhle, A. Hahn / Nutrition Research 31 (2011) 429–435
calculated in our study, cannot with certainty be considered
representative of the carbohydrate intake of the members of
Homo sapiens throughout the Pleistocene.
In summary, our data suggested that most diets of hunter-
gatherers were relatively low in carbohydrates. In accordance
with our hypothesis, there was a wide range of carbohydrate
intake across various hunter-gatherer populations, and
latitude and ecological environments were important factors
that influenced the carbohydrate content of the diets. For this
reason, there is no universal carbohydrate intake that is
representative of hunter-gatherer diets. Interestingly, the
energy derived from carbohydrates in the diets of most
hunter-gatherer societies was markedly different (lower) than
the amounts recommended for healthy individuals.
Acknowledgment
The authors wish to thank Claus Leitzmann and Anthony
Sebastian for their comments and helpful review during the
writing of this article. There was no financial support, and
none of the authors has a conflict of interest.
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