Viewpoint: The economics of huntergatherer societies and the evolution of

human characteristics
Arthur J. Robson

Department of Economics, Simon Fraser

University
Hillard S. Kaplan Department of Anthropology, University of
New Mexico

Abstract. We argue for attention to the evolutionary origins of economic behaviour.

Going beyond this, we argue that the economy of hunting and gathering was the
context in which evolution shaped human characteristics that underlie modern economic behaviour. We first reconsider the basic biological question of why aging occurs
at all. We then illustrate the usefulness of considering foraging economics by asking why
it is biologically advantageous for humans to live long after their reproductive career is
over. Further, we argue that foraging economics would have led to the simultaneous
exaggeration of intelligence and of longevity that is characteristic of humans. JEL
classification: A12, J10
Leconomie des societes de chasseurs-cueilleurs et levolution des traits humains. Les
auteurs sugge`rent quon doit porter plus dattention aux origines evolutionnaires du
comportement economique. Poussant un pas plus loin, ils sugge`rent que leconomie
des chasseurs-cueilleurs a ete le contexte ou` levolution a conforme les traits humains
qui sous-tendent le comportement economique moderne. Dabord, ils reexaminent la
question biologique de base a` savoir pourquoi il y a vieillissement. Ensuite, ils illustrent
lutilite dune economie des activites de cueillette, chasse et engrangement en posant la
question a` savoir pourquoi il est biologiquement avantageux pour les humains de bien
vivre apre`s que leur periode de reproduction active est terminee. Finalement, ils sugge`rent que ce type deconomie aurait entra ne simultanement une surestimation de
lintelligence et de la longevite deux traits des humains.

Robson was supported by the Social Sciences Research Council of Canada and the
Canada Research Chairs program; Kaplan by the National Science Foundation and the
National Institutes on Aging. Email: robson@sfu.ca
Canadian Journal of Economics / Revue canadienne dEconomique, Vol. 39, No. 2
May / mai 2006. Printed in Canada / Imprime au Canada

0008-4085 / 06 / 375398 /

Canadian Economics Association

376 A.J. Robson and H.S. Kaplan

1. Introduction
Serious consideration of the evolutionary origins of human economic behaviour
is likely to pay handsome dividends. Our evolutionary history is inherently
fascinating and logically must underpin relevant human characteristics.
Recently, however, a number of empirical regularities have been found that
cast doubt on the validity of conventional economic theory. These regularities
have inspired a bewildering variety of conflicting alternative theories, many of
which are limited in scope and ad hoc. We believe, then, that the most
important potential contribution from considering the evolutionary origins of
economic behaviour is that it can provide a disciplined way of winnowing
down and unifying the current array of models of economic behaviour. An
evolutionary perspective holds out the hope of modifying and extending
economic theory while retaining its appealing generality and consistency.
Progress can be made by considering in the abstract how biological
evolution shaped attitudes to risk, for example (see, e.g., Robson 1996a, b).
However, an evolutionary approach can be sharpened by considering the
circumstances prevailing during our evolutionary history. For most of the
two million years since the advent of the genus Homo, that is, we have hunted
and gathered. This foraging lifestyle put an unusually heavy emphasis on
intelligence and social organization. We hunted, for example, using a myriad
of techniques that were customized to a corresponding myriad of situations,
and many of these techniques involved cooperative behaviour. The cognitive
complexity of economic production in foraging societies entailed a dramatic
initial increase in the rate of output as individuals age, owing to the effects of
learning-by-doing, in particular. It is then not surprising that foraging societies
are also characterized by huge intergenerational transfers from adults to
children. Thus, we take foraging economies with such characteristics as the context
in which natural selection operated to form modern human demographic and
economic characteristics.
A thesis of this paper is therefore that the usual implicit causal relationship
between economics and biology should be partially reversed. That is, when
economists think of biological evolution (which admittedly is not very often), it
might be as providing the effectively exogenous backdrop against which modern
economic phenomena play out. This paper argues instead that the economics
of archaic societies was the crucible in which biological evolution formed
human demographic and economic characteristics. That is, if we peek behind
the backdrop, there is not only evolutionary biology, but economics as well.
2. The evolved life history of a skill-intensive and social species: an overview
In many ways, we human beings are ideal biological organisms to study. We
live and have lived in a diverse array of environments; we know our ages and
our reproductive history; we can report directly on relevant phenomena; and

Economics of hunter-gatherer societies

377

we leave historical records. For example, we certainly know much more about
aging in humans than we do about aging in any other mammal (with the
possible exception of laboratory rats and mice).
The processes underlying human aging stand as testaments to the forces
shaping life span evolution among living creatures in general. More specifically, research with the few remaining people that still practise a traditional
hunting and gathering lifestyle is providing critical information about the life
histories of our ancestors and the selection pressures that acted on them.
Humans lived as hunter-gatherers for most of our 2 million years of evolutionary history. Modern hunter-gatherers, who are affected by global socioeconomic forces, are not living replicas of our stone age past. Yet, in spite of
their variable historical, ecological, and political conditions, foraging peoples
exhibit remarkable similarities, especially in the realms of development and
aging. This suggests that natural selection has produced a characteristic life
history for our species, a life history that is quite distinct from that of our closest
living primate relatives.
When we compare humans with other creatures, our brains and attendant
mental abilities stand out. The human brain is about three times as large as
that of a chimpanzee, our closest living relative and arguably the most intelligent
non-human species. Any attempt to account for our unique evolutionary path
will also have to explain why human brains became so large and why we
became such good learners.
In addition to our large brain size, humans are also distinctive because of
our very long lives. Comparing the life span of people living in remote societies
without Western medicine to that of wild chimpanzees is particularly revealing.
Overall, such humans have an average life span that is about twice that of
chimpanzees in the wild. In greater detail: the risk of dying is high during
infancy for both humans and chimpanzees, but then decreases rapidly. For
chimpanzees the lowest point about 3% per year is reached at about age 13
(the age of first reproduction for females) and increases sharply after that. In
contrast, mortality among people still living as hunter-gatherers drops to a
much lower point about 0.5% per year and remains low without much
increase between about 15 and 40 years of age. Human mortality rates then rise
slowly, followed by a rapid jump only in the 60s and 70s.
As a result, about 60% of hunter-gatherer children survive to adulthood,
compared with 35% of chimpanzees. Chimpanzees also have a much shorter
adult life span than humans. From the age when they first reproduce,
chimpanzees live an additional 15 years, on average, compared with 38 more
years among human foragers. It is important to note that very few female
chimpanzees even reach a post-reproductive phase, whereas women spend more
than a third of their adult life in such a phase. Fewer than 10% of chimpanzees
live to age 40, but more than 15% of hunter-gatherers make it to age 70.
Thus, delayed aging and long adult life spans appear to be evolved characteristics of our species. Adult mortality risks are remarkably uniform across

378 A.J. Robson and H.S. Kaplan

human societies without access to western medicine. Although the ancestors of
the aboriginal peoples of South America, Africa, and New Guinea diverged
more than 10,000 years ago, the risk of dying at each age is very similar from
society to society, and not that different from that in Europe prior to modern
medicine.
It helps to examine our primate ancestors in order to understand the
evolution of our larger brains and longer lives. As the primate order evolved
and different species radiated across the worlds warmer areas, there was a
series of four grade shifts that increased encephalization brain size relative
to body size and increased longevity.
The first grade shift began about 60 million years ago with the evolution of
prosimians, who probably lived longer because living in trees afforded them
more safety. The second grade shift came about 35 million years ago with the
evolution of the monkey lineage. It involved a huge increase in both brain size
and life span. The development of the ape lineage represented today by
chimpanzees, gorillas, and orangutans was the third major grade shift.
Apes can live almost twice as long as most monkeys and have much bigger
brains, even after adjusting for their larger body size. The fourth grade shift
occurred with the divergence of the hominid line, particularly the advent of the
genus Homo about 2 million years ago.
This coevolution of brains and life spans makes sense when one considers
that natural selection depends on both costs and benefits. Brains are very
costly: humans use about 65% of base metabolism for the maintenance and
growth of the brain during the first year of life. Brains resulting from natural
selection must provide benefits that pay for their costs. If the benefits provided
by the brain were outweighed by its cost, individuals with smaller brains would
leave more descendants than those with larger brains, and the average brain
size of the population would shrink.
Since learning transforms present experiences into better future performance,
we can see brain development especially development of the cerebral cortex
(which expanded disproportionately during primate evolution) as an investment
in the future. During primate evolution, brain size appeared to increase as the
learning required by the feeding niche grew more intense. The advent of
monkeys, for example, involved a shift from the prosimians smell- and
hearing-based insect eating to a reorganized sensory system with binocular,
colour vision able to find many different plant foods, captured by dexterous
hands and manipulated through hand-eye coordination. Because monitoring
and exploiting the fruits and leaves of different trees demanded more learning, brain size increased.
During the third shift, apes adopted a diet that emphasized ripe fruits, requiring
even more environmental monitoring and more complex extractive techniques.
These higher prerequisites for learning mean apes take longer than monkeys to
become competent foragers. For example, even though chimpanzees can
provide most of their own caloric needs by age five, juveniles under age

Economics of hunter-gatherer societies

379

seven are unable to utilize and make tools for termite fishing and therefore still
rely upon their mothers for protein, even after weaning. Orangutans also
depend upon their mothers for about seven years. Today, as conservationminded scientists attempt to reintroduce captive-born orangutans into the
forest, they are discovering how much learning must take place before these
creatures are able to survive on their own.
During our evolutionary history, humans pushed learning-intensive feeding
techniques to the extreme. Evidence from a wide variety of sources shows that
humans became specialized in the consumption of calorie-dense, low-fibre
foods rich in protein and fat. Although there is considerable variation across
societies depending on their ecology, modern foragers all differ considerably in
diet from chimpanzees. The majority of forager diets is meat, accounting for
about 60% of calories. In contrast, chimpanzees obtain only about 2% of their
food energy from hunted foods.
The next most important food category for foragers is resources such as
insects, roots, nuts, seeds, and difficult-to-extract plant parts such as palm fibre
or growing shoots. These resources tend to be embedded in a protective
context, such as underground or in hard shells. Such extracted foods make
up about 30% of the forager diet, as opposed to 3% among chimpanzees. In
contrast to such difficult-to-acquire resources, collected foods such as fruits,
leaves, flowers, and other easily accessible plant parts form, on average, 95%
of the chimpanzee diet but only 8% of the human forager diet. The data thus
suggest that humans specialize in rare but nutrient-dense resource packages
such as meat, roots, or nuts whereas chimpanzees specialize in easily attainable
but less nutritionally dense plant parts.
Owing to the learning-intensive nature of their feeding niche, hunter-gatherers
take even longer than apes to become competent foragers. Whereas chimpanzees
can meet most of their caloric needs by age five, human foragers produce fewer
calories than they consume for close to twenty years! In fact, the total calories that
human parents must provide for their offspring each year increases from birth to
about age 14, as children grow but remain unproductive.
Yet however burdensome child rearing may sometimes seem to huntergatherer parents, this long period of dependence pays off in the long run.
Human adults are much more productive than chimpanzees. Net production
(the surplus after ones own food consumption is taken into account) climaxes
at about 1,750 calories per day for human adults, compared with 250 for adult
female chimpanzees, although human foragers generally do not reach that level
of productivity until about age 45.
Between ages 20 and 40, human hunters go through the equivalent of
graduate school and a period of on-the-job training. This is because humans,
unlike other predators, rely more on knowledge than on physical prowess. For
example, in one group of South American hunter-gatherers, the Ache, men
know that the paca (a large rodent) lives in burrows with a main entrance and
up to seven hidden escape hatches. When jaguars attempt to dig a paca out, it

380 A.J. Robson and H.S. Kaplan

can run away unscathed through one of its alternative exits. But rarely does it
fool an Ache hunter, who calls the other hunters to the hole while quietly and
systematically searching for each escape route. When the other hunters arrive,
a younger, less-skilled hunter is given the task of ramming a large log into the
main entrance. Simultaneously, each of the remaining hunters dives on top of
his respective escape hatch while his hands form a noose, allowing the fleeing
pacas head through his open arms only to grab its throat and suffocate it.
Feeding niches that demand such high levels of learning and cooperation
should select not only for bigger brains, but also for increased longevity. This is
because the brain costs a great deal early in life, providing benefits only later in
life. To illustrate the importance of longevity to a learning-intensive feeding
niche, consider what would happen if a human forager group suffered the same
mortality rates that chimpanzees do. Humans depend on their parents for a
long time, creating a large calorie deficit that they only gradually pay back
during their highly productive middle adulthood years. However, less than
10% of chimpanzees live to the age at which human productivity peaks. With a
chimpanzee life span, a human forager group would then necessarily experience
a precipitous decline in numbers.
The flows of food and other services both within and among families
that support this long period of dependence are also particular to humans.
Unlike our ape relatives, such as chimpanzees, who give females the entire
burden of feeding and caring for infants, humans cooperate in this activity,
practising a sexual division of labour, a practice that appears to have ancient
roots. Although the details vary from society to society, all existing huntergatherers and peoples who depend on a mix of foraging and farming worldwide collaborate in raising children. Typically, the male contribution to this
joint endeavour focuses on food supply, mainly from hunting but with some
gathering as well, while women participate in a wider mix of activities such as
gathering, food preparation, and direct child care.
Why is a division of labour so fundamental to the human way of life?
After all, women are physically capable of being adept hunters and sometimes do so when it is necessary. However, since traditional human hunting
is so learning intensive, it pays to hunt only if one spends many years doing
so. If women were to hunt only when they were not pregnant or nursing,
they might get less food from hunting than from gathering because of their
lack of experience.
Males probably became the primary provider of food when hunted animals
became an important part of human diets. The protein and fat from meat
along with the carbohydrate obtained from plant foods created a balanced
diet. Although the proportion of food provided by men and women varies
across hunter-gatherer societies, men provide, on average, about twice as many
calories and seven times as much protein as women. After taking into account
their own consumption, women supply only 3% of the calories to offspring
while men provide the remaining 97%.

Economics of hunter-gatherer societies

381

This assistance from men enables women to focus their energy on providing
high-quality child care, resulting in almost double the survival rate for human
children compared with that of chimpanzee offspring. In addition to behaviour, womens physiology is consistent with an evolutionary history of extensive male parental investment. Unlike other primates, human females lower
their metabolic rates during pregnancy and store fat, which is permitted only
by receiving provisions. Human female foragers also tend to work less during
lactation, unlike female chimpanzees, who heighten mortality risk by working
more in this circumstance.
This extensive cooperation between hunter-gatherer men and women makes
evolutionary sense only once the reproductive performance of spouses is
linked. Even though divorce is common in many hunter-gatherer societies,
marriages stabilize once children are born. The long period of dependence on
parents means that, at any one time, most parents are raising several dependent
offspring of different ages. This puts pressure on couples to stay together.
Those who divorce and remarry while raising children frequently argue with
new spouses over the division of resources among their joint children.
Avoiding those conflicts is an additional incentive for a couple to stay together
and have all or most of their children together.
In fact, although men are physiologically capable of reproducing throughout their lives, most men undergo behavioral menopause. Among Ache
hunter-gatherers, for example, if a couple had at least two children together,
the womans last birth was the same as her husbands last child in 90% of
cases.
Indeed, the phenomenon of menopause, whether physiological or behavioural, is puzzling from a biological perspective. That is, how could humans
have evolved to outlive their biological usefulness? One suggestion is that
menopause evolved because women would leave more genetic descendants by
helping as a grandmother than by continuing to reproduce. As women age, the
reasoning goes, their pregnancies are less likely to succeed and they are more
likely to die in childbirth. Indeed, among foragers, there are no golden
retirement years; both grandmothers and grandfathers spend significant time
helping to raise children. Although older people switch to less physically
demanding and more knowledge-intensive activities as they age, they still
work very hard until death.
This paper examines some of the issues arising here in greater detail, utilizing our published and unpublished work, as follows. Given the apparent
advantages of longevity, we begin by re-examining a basic question applicable
to a wide range of species, but to humans in particular: Why do we age at all?
We then ask questions that focus more on the human case. These questions
also illustrate how the economics of hunting and gathering was the context in
which biological evolution shaped human demography: Why is fertility concentrated towards the middle of life? Why do women live after menopause?
Finally, we come back, in particular, to the major themes of this overview:

382 A.J. Robson and H.S. Kaplan

Why are we so smart? Why do we live so long? Why might these last two
attributes be linked together?

3. Why do we age? Why is fertility concentrated towards the middle of life?

From a biological point of view, aging refers to an eventual increase in
mortality. But why does aging occur? How can it be evolutionarily optimal
for us to be programmed to first invest heavily in physical growth (while also
learning productive skills), only to subsequently decay and senesce? It is
important to appreciate in this connection that recent empirical findings
imply that aging in this sense may not be inevitable.1
Robson and Kaplan (2005) show this issue can be addressed by applying
economic analysis of intertemporal allocation of scarce resources. Organisms
grow in order to produce more energy that is then available for additional
growth, maintenance, and reproduction. Natural selection chooses the life
history with the maximum rate of population increase.2
There are two important related problems in evolutionary theories of aging
(Key references: Charlesworth 2000; Hamilton 1966; Kirkwood 1990;
Medawar 1952; Rose 1991; Williams 1957). First, existing evolutionary theories
fail to provide an integrated model explaining both the typical decrease in
mortality with age during the first phase of life and the subsequent typical rise
in mortality during the second phase. Second, this existing formal theory does
not explain the extended periods of survival after reproduction has ceased, as
observed in humans (and in a few other species).
Inspired by the available data for hunter gatherers on mortality, body
weight, and output, as sketched in figure 1, we suggest as an answer to the
first problem that (a) growth in the quantity of somatic tissue helps to drive the
increasing productivity during the first phase of life and this results in an
endogenous decline in mortality, and (b) Deterioration in the functional efficiency of somatic tissue drives the decline in productivity later in life, and this,
in turn, drives an endogenous increase in mortality.
The second problem is: Why do we survive after fertility has fallen to zero?
This essentially asks: Why do we not age faster? The existence of this phenomenon in humans is demonstrated by the data on hunter-gatherer fertility and
mortality in figure 1. The economic explanation for this phenomenon that we
adopt relies upon the huge intergenerational transfers that are also observed
in these societies. (Data on cumulative expected net energy production for
hunter-gatherers in figure 2 below demonstrate the importance of these intergenerational transfers.)
1 For example, aging is negligible in some species of tree that live for several millennia (Finch
1998). It is particularly revealing that humans age at about half the rate of chimpanzees
(Kaplan et al. 2000), despite the close evolutionary relationship of the two species.
2 Stearns (1992) presents the evolutionary theory of life history.

Economics of hunter-gatherer societies

0.14

0.13

0.9

0.12

0.8

0.11

0.7

0.1
0.09

0.6

0.08

0.5
0.4
0.3

Production

0.07

Weight

0.06

Female Fertility

0.05

Mortality

0.04

Mortality

Production, Weight and Fertility

383

0.03

0.2

0.02
0.1

0.01

0
0

10

15

20

25

30

35

40

45

50

55

60

0
65

Age
FIGURE 1 Standardized production, standardized weight, standardized female fertility, and
mortality, all by age
SOURCES: The data on production and mortality are derived from Kaplan et al. (2000) and
represent the average production and age-specific mortality of males and females by age among
three foraging groups: the Ache of Paraguay, the Hiwi of Venezuela, and the Hadza of Tanzania.
Data on weight are derived from unpublished measures of Ache males and females made by
Kaplan, along with coworkers Kim Hill, A. Magdalena Hurtado, and K. Hawkes. Data on fertility
are based upon the Ache of Paraguay from Hill and Hurtado (1996), those on the !Kung of
Botswana from Howell (1979)

Our theory hypothesizes that each individuals somatic capital is characterized by both its quantity and its quality. The quantity of capital is the number
of cells. There is initial investment in quantity, so cell number increases
irreversibly up to some age, but it is thereafter constant, capturing the determinate growth pattern of humans. Cell quality, as measured by productive
efficiency, is also capable of improvement by energetic investment. Without
such investment, however, cell quality depreciates over time.
Our approach to somatic quality is based on what is now known about the
biochemistry of cellular aging. Research shows that metabolic activity in the
cell inevitably produces various harmful biochemical by-products, including
reactive oxygen species, or ROS, for example. These by-products are implicated in aging; For example, ROS is implicated in some serious neurological
disease. (See Love and Jenner 1999). However, such aging is not inevitable.
That is, it is possible to remove such by-products and hence reduce or even
prevent aging. Consider, for example, the nematode worm C. elegans, one of
biologys favorite experimental organisms. This has a genetic variant in which

384 A.J. Robson and H.S. Kaplan

particular relatively large molecules are produced within each cell, which then
bind to the small harmful by-product molecules. The entire molecular combination can then be purged from the cell. The interesting fact is that this genetic
variant, although typically present, is not common in the wild population (See
McElwee et al. 2004). The issue then must be: Is somatic quality maintenance
too expensive to be worthwhile?
We propose a new theory of aging in which such quality maintenance is
expected to be too costly, as follows. Human beings, and many other species,
are characterized by a segregation of the germ line and the somatic line. That
is, the cells that form the body of each individual are separate from the sex cells
of that individual the eggs for females or the sperm for males. Thus, for
example, mutations arising from the replication of somatic cells during the
growth of the individual are not transmitted to offspring via the sex cells.
In our model, the cost of investment in quality depends on the quantity of
cells, because each cell is subject to deterioration and has its own maintenance
costs. It may then be evolutionarily optimal to generate a high level of initial
quality, but then to let it fall with age. This is because the quality of the
relatively small number of cells in the germ line can be maintained cheaply,
while the quality of the large soma achieved after growth would be much more
expensive to maintain.
Indeed, we assume, as a reasonable limiting case, that the quality of the
germ line can be maintained costlessly. Thus the plans for a new individual can
be maintained without cost, implying that the initial quality of such a new
individual is high. It is relative to this alternative of producing a small but
high-quality new individual that the cost of maintaining the quality of a fullgrown individual is too high.
Our model also then explains why mortality rates are U-shaped, a common
feature of a wide variety of species. That is, we explain not only why mortality
rates eventually rise, or why aging occurs, but why mortality rates initially fall
during infancy and childhood. In our model, optimal investments in mortality
reduction respond to the overall value of remaining life. As expected net future
production increases as a result of somatic growth, the optimal investment in
mortality reduction then increases and mortality falls. As expected net future
production declines later in life, owing to decreased somatic quality, optimal
investment in mortality reduction decreases and mortality rises.

3.1. A two-period example

It is impossible to illustrate all of the important theoretical issues here in a twoperiod example. Merely to obtain U-shaped mortality, for example, requires at
least four periods. Indeed, to obtain aging as an optimal outcome requires an
unlimited number of periods, since to impose a terminal date guarantees aging
near that date. A key aspect of the model, however, concerns the dependence
of the cost of quality maintenance on the quantity of somatic capital. That

Economics of hunter-gatherer societies

385

quality maintenance is thus discouraged by the optimal accumulation of

somatic capital can be illustrated by a two-period example as follows.
Suppose that gross energy output of an individual is given by
FK; Q aK  bK2 eQ  fQ2 ; where a; b; e; f > 0;
K is the quantity of somatic capital and Q is its quality. An individual lives two
periods and is endowed with somatic capital with initial quantity K0 and initial
quality Q0. Suppose the energy cost of producing each individual is C0, a
constant, so that the reproductive technology is constant returns to scale.
(This simplifying assumption cannot be made in the general model below if
this is to yield a determinate pattern of fertility.) Suppose that individuals have
fixed probability p of surviving from age 0 to age 1. This abstracts from an
important property of the general model that mortality is endogenous.
At t 0, the individual invests v0  0 in the quantity of somatic capital and
w0  0 in the quality of this capital. The quantity of capital at t 1 is then
K1 K0 v0; the quality at t 1 is Q1 Q0  Q0 w0, for some  2 (0, 1).
There is no depreciation in the quantity of somatic capital, in line with its interpretation as the number of cells; but quality tends to depreciate, in line with its
interpretation discussed above. The energy cost of v0 is v0; that of w0 is K1w0.
The latter cost function builds in a greater cost of quality maintenance for a larger
soma, where the most appropriate soma is that from the next period, K1. (This
issue does not arise in the general continuous time formulation sketched below.)
Consider, now, a large population in steady-state growth at rate r, say,
so that the age distribution is constant. The ratio of old to young is then
(p(s)/(1 r)). That is, there are fewer old individuals because there is some
probability of death in the interim, but also because there were fewer in the
birth cohort one period ago, given a positive growth rate.
What population growth rates are then economically feasible in the long
run? It is assumed that the energy flow is food, for example, which cannot be
stored but is also not wasted. Since the steady state is then feasible if and only
if it is possible for the adults to cover the deficits of the children, it follows that
C0 F0  K0 I0 w0  I0

pFK0 v0 ; Q0  Q0 w0
;
1r

where F0 F(K0, Q0) is fixed. Thus, the growth rate of the population is
1r

pFK0 v0 ; Q0  Q0 w0
where K0 I0 w0 I0 C0  F0
K0 I0 w0 I0 C0  F0
> 0;

which follows, for example, if C0 > F0.

The most evolutionarily successful type must then maximize this expression
for 1 r over choice of v0 and w0. For simplicity, suppose, furthermore, that
this maximized growth rate is zero. Indeed, humans must have had essentially

386 A.J. Robson and H.S. Kaplan

a zero growth rate over the 2 million years of our evolutionary history.
(However, it is easy to allow a positive growth rate.)
Assuming an interior solution for v0, the first-order conditions are then
FQ K1 ; Q1
FK K1 ; Q1

K1


and
;
FK1 ; Q1
v0 C0  F0
FK1 ; Q1
v0 C0  F0
with equality if w0 > 0. Equivalently,
pFK K1 ; Q1 a  2bK1  and pFQ K1 ; Q1 e  2fQ1  K1 ;
with equality if w0 > 0. It follows that
K1

a
a
; so that; if 
> e; then w0 0:
2b
2b

That is, the optimal strategy is to increase the quantity of somatic capital, even
though such an increase renders the maintenance of quality suboptimal. It is
not that quality is forced to decline; it is a prediction of the model. For that
matter, it may well be feasible to maintain quality, although a type that did so
would experience a a lower growth rate.
Indeed, if investment in quantity v0 is simply constrained to be zero, and K0
and Q0 are small enough, then it will be optimal to invest in quality, w0. That is,
if v0 0; but e > K0 and

e  K0
> Q0  Q0 ;
2f

then the optimal

Q1

e  K0
Q0  Q0 w0 > Q0  Q0 :
2f

From an economic point of view, the form of the cost of investment in quality
implies that the quantity and quality of somatic capital are essentially substitutes: an increase in the quantity of somatic capital reduces the attractiveness
of investing in its quality. A key source of the evident asymmetry between
quantity and quality is that initial quality is relatively high but initial quantity
is low.
3.2. A sketch of a general theory
Consider now a sketch of the general model of Robson and Kaplan (2005).
Greater generality is required in order to obtain U-shaped mortality as well as
a fertility profile that is at first constant at zero, then rises, then falls, finally
becoming constant at zero again.
A key aspect of this general model is a non-linear cost of fertility. This
implies that the optimal time profile of fertility is determinate and has a
realistic shape. Suppose, then, that the gross energy production rate of an
organism is given by the concave function G(K, Q). Again, K is the somatic

Economics of hunter-gatherer societies

387

capital of the individual and Q is the quality of this somatic capital. Fertility is
now s and the energy flow net of the cost of fertility is given by the concave
function F(G, s). This function is increasing in G, decreasing in s and
FGs(G, s) < 0, so that reproduction is cheaper, at the margin, the greater the
amount of gross energy available. For simplicity, individuals here reproduce
parthenogenetically, by means of virgin birth.
The rate of augmentation of the capital stock at age t is denoted by v, where
dK
v 2 0; v:
dt
The quantity of somatic capital does not depreciate, as is consistent with the
data. The energy cost of increasing the capital stock at rate v is v.
Also for simplicity, investment in capital is confined to an initial range of
ages. That is, there exists t*  0 such that
v v; for all t 2 0; t*
v 0; for all t > t*:
Thus, the choice of the investment profile of the quantity of somatic capital
reduces to the choice of t*  0.
The evolution of the quality of the somatic stock, Q, is given by
dQ
w  Q:
dt
Here,  is the rate of decline of quality of the stock in the absence of discretionary maintenance. The contribution of discretionary maintenance to offsetting or reversing this decline is w, and the energy cost of this is wd(K),
where d is increasing in K. That is, a given reduction in the rate of decline of
quality is more energetically costly, the greater the quantity of the somatic
stock involved. This reflects the energy economics of removing metabolic
by-products, for example.
Another major component of the model that was absent in the example is
endogenous mortality. If  is the rate of mortality and p is the probability of
survival, then these are connected by the identity
1 dp
:
p dt
However, mortality is subject to natural selection, where the energetic cost of
mortality  is e(), a decreasing and convex function. Consider the operation
of the immune system, for example. The larger the number of antibodies of a
given type, the better protected one is against the corresponding disease; the
wider the spectrum of types of antibody, the greater the variety of diseases to
which one has immunity. But the larger the army of antibodies maintained, the
greater the metabolic cost.

388 A.J. Robson and H.S. Kaplan

Suppose, now, that the population is in a steady-state growth equilibrium,
with growth rate r. The Euler-Lotka equation, a foundation of demography,
must hold (see, e.g., Charlesworth 1994, chap. 1). That is,
Z 1
ert psdt 1:
0

In addition, it follows from the above that the energy flow surplus for an
individual at age t is
F  v  wd  e:
This surplus is not constrained to be non-negative. Rather, intergenerational
transfers can be made freely, a reasonable abstraction for the human case.3
With such transfers, but given that energy in the form of food cannot be
stored, economic feasibility requires that the total energy excess generated by
the old covers the total energy deficits of the young, at any given point in time,
so that the steady-state budget balance condition is
Z 1
ert pF  v  wd  edt  0:
0

This, then, is a new foundation of our approach, a foundation based on

economics rather than demography.
This is an idealization of hunter-gatherer society that is, despite its
possibly having only 2030 people, it is assumed that the steady-state age
structure has been attained. Individuals in each group of hunter-gatherers are
also assumed to be related to one another. This provides the basis for an
answer to the question: Why do the current old not simply renege on the
obligation to make large transfers to the current young, even if these old
individuals themselves were once the beneficiaries of such transfers? That is,
from a biological point of view, individuals have a strong incentive to transfer
resources to the young if these individuals are close relatives genetically. The
proximate emotional mechanism driving such behaviour is parental love.
The basic evolutionary problem is then to maximize the growth rate r
subject to the differential equations governing K, Q, and p, the Euler-Lotka
equation, and the budget balance constraint.
The main result of Robson and Kaplan (2005) is that it may be optimal to
invest in somatic capital during an initial growth phase. Nevertheless, at the
same time it may be optimal never to counteract the decline in the quality of
somatic capital. Furthermore, fertility is hump-shaped and is zero at the
beginning and again at the end of life. Finally, mortality is U-shaped with a
minimum before the end of the growth phase, so that mortality has the shape
seen in the majority of species.

3 Kaplan and Hill (1985) provide evidence on food sharing and examine its causal basis.

Economics of hunter-gatherer societies

389

The present theory can be shown to imply that mortality is governed mostly
by narrow biological considerations, based on fertility, when young. At the
other end of life, however, where fertility is eventually zero, mortality is
determined by purely economic considerations, based on expected net energy
production. It is therefore evolutionarily optimal to live beyond the age at
which fertility falls to zero, because expected net energy production remains
positive there. The model then accounts for life beyond menopause.
It is worth emphasizing that it is generally feasible to maintain a value of life
that is constant and hence mortality that is also constant. However, this is not
optimal, and a type that did this would not maximize the population growth
rate. We grow old because the marginal cost of not growing old, of keeping
mortality constant, exceeds the marginal benefit. This is in the light of the
option of using the pristine germ line blueprints to create another individual,
and is despite the enormous biological and economic costs that are incurred in
raising such an individual.
At the same time, the Robson and Kaplan model can be generalized so that
it pays to eventually arrest the decline in the quality of somatic capital. That is,
since the marginal product of quality rises as quality falls, eventually it may be
optimal to invest appropriately in somatic quality. This would produce a
plateau in output and hence in mortality. Although such an individual would
not live forever, since her mortality would remain positive, she would not age
beyond a certain point, in the present biological sense. This is of interest, since
it has been claimed that such a mortality plateau exists in the human data in
advanced old age.4

3.3. A note on the literature

Our model is best seen as a more attractive formulation of Kirkwoods disposable soma theory of aging (see, e.g., Kirkwood 1990, 1999). Kirkwood
argues that repair of somatic tissue must be optimized by natural selection. At
some point, greater returns will be obtained through reproduction than
through repair. Perhaps, then, optimal repair is less than complete, and the
soma deteriorates with age, ultimately being replaced by descendants. The
segregation of the somatic and germ lines permits the degradation of the
former.
However, there are key differences in the detailed explanations of aging
between Kirkwoods model and ours. In Kirkwoods model, in contrast to
ours, it is simply impossible to make the mortality rate fall over time and
prohibitively expensive at the margin to keep it constant.5 This assumption of
4 See Vaupel (1997). Indeed, there is sketchy evidence for a decrease in mortality in advanced
old age. The obvious alternative explanation for such a phenomenon is that individuals are
not homogeneous. Hence, since individuals with higher mortality rates tend to be winnowed
out of the population first, this reduces the mortality rates of the remainder.
5 Sozou and Seymour (2004), show that relaxing this assumption implies that constant
mortality may be optimal.

390 A.J. Robson and H.S. Kaplan

Kirkwood guarantees aging without explaining it. His approach also obviously
fails to account for an initial phase of life with falling mortality.
Some of the variables in the above model can be reinterpreted in a more
conventional economic fashion. For example, expenditures on mortality reduction could be interpreted as discretionary investment in health capital, rather
than as genetically controlled metabolic expenditures on the immune system.
This would make the current model more comparable to the work of
Grossman (1972) and Ehrlich and Chuma (1990), for example. In contrast to
their assumption that the depreciation rate for health capital is an exogenously
given increasing function of time, however, the current model explains why
mortality eventually falls without presupposing this effect.

4. Why are we so smart? Why do we live so long?

A detailed look at data on hunter-gatherer production over time (as in figure 1,
above) raises an issue that has not been addressed yet. Output continues to rise
dramatically in the teens and twenties, well after somatic growth has ended. In
Kaplan and Robson (2002) and Robson and Kaplan (2003) we argue that this
is due to on-the-job-training to increasing skill in performing mentally
demanding tasks but that the acquisition of such skill is possible only because
a large brain has already been created.6 The brain is a key component of
somatic capital. Indeed, since the lag between the end of brain growth and
peak output is dramatic as much as 2530 years investment in the brain is
significantly more roundabout than is investment in the body in general.
What circumstances promoted such an exaggerated brain? Why are we as
smart as we are? Perhaps the advent of the African savanna and the subsequent
flourishing of large herbivores created a niche for a species that relied on
intelligence and tight social organization to hunt effectively (see, e.g.,
Jurmain et al. 2000). Human hunting is certainly cognitively demanding,
applying techniques that are varied and creative. The wide diversity of behaviour would be difficult for evolution to program directly; rather, it entails the
evolution of a large brain with the ability to learn general rules and to solve
particular problems. Such a strategy produces a life history with the dramatic
intertemporal trade-offs in figure 1.
This steeply rising human productivity curve, which directly reflects the
learning-by-doing needed to become an effective forager, is only possible
given the physical growth of the brain early in life. The dynamic trade-off
here begs for some interpretation of the brain as embodied or somatic capital.
It is worth recalling that, during the first year of life, up to 65% of basal
metabolism is devoted to the maintenance and growth of the brain (Holliday
1978).
6 See also Kaplan, Lancaster, and Robson (2003). Reader and Laland (2002) provide evidence
of the crucial link between brain size and intelligence when primate species are compared.

Economics of hunter-gatherer societies

391

Kaplan and Robson (2002) and Robson and Kaplan (2003) focus upon a
more specific and accessible question than that concerning the origin of the
intelligence of humans, namely: Why might the extreme intelligence and longevity of humans be expected to go together?
Such an association holds quite generally in a cross-section sense (Allman,
McLaughlin, and Hakeem 1993). Multiple regression with data on primates,
for example, shows that brain weight has a statistically significant effect on
longevity, even when body weight is also included as an explanatory variable
and the effect of body weight on brain weight is allowed for. Humans have a
brain that is roughly three times as big as that of chimpanzees, in particular,
and live about twice as long, despite being only slightly heavier.
Furthermore, this association between intelligence and longevity is a salient
feature of our evolutionary history. This history, in the first place, has featured
dramatic increases in brain size, with no similar clear trend in body size. The
encephalization quotient, EQ, a measure of brain size relative to body size, has
increased dramatically.7
In the second place, the evidence concerning the longevity of our evolutionary ancestors is necessarily less direct. However, evidence on the sequence
of eruption of teeth, for example, suggests that Australopithecus had a life span
comparable to that of a modern chimpanzee, much less, then, than that of
modern humans; intermediate species have intermediate life spans (Smith,
1991). Thus, human evolutionary history involves a large increase in brain
size and an apparent substantial concurrent increase in longevity.
A key part of an explanation of this association is that investment in the
brain is a more roundabout biological strategy than investment in the body.
This is illustrated by the productivity profile in figure 1. That is, the costs of a
large brain are even more concentrated towards the start of life than is the cost
of physical growth overall, but the returns as hunting productivity, for example, do not peak until about age 35, well beyond the age of physical maturity.
Using both productivity and mortality data, it is possible to make even
more vivid the resulting huge intertemporal trade-off in hunter-gatherer life
history. Figure 2 plots cumulative production net of consumption: transfers,
that is, weighted by the probability of survival, as a function of age, for all
available hunter-gatherer societies. It then takes almost until age 50 to break
even and the ultimate payback is relatively small.
This also demonstrates the essential way in which intelligence and mortality
are intertwined. That is, if this calculation is performed using human net transfers
but chimpanzee mortality, the curve remains in substantial deficit. Given
that chimpanzee mortality approximates the mortality of human evolutionary
ancestors, this ancestral demography would have had to change to support the
emergence of human intelligence and the associated economic life history.
7 See Martin (1981). Jerison (1973) provides evidence concerning the evolution of intelligence in
a variety of species.

392 A.J. Robson and H.S. Kaplan

4.1. A two-period example of neural capital
The theoretical model of Robson and Kaplan (2003) can be illustrated by
means of the following two-period example.
Consider a species in which individuals live, at most, for two periods. At

t 0, gross energy output is F()
< 0, so that the child here must rely on
transfers from adults. The role of the parameter  is to create a shift to more
challenging environment in which there are greater short-run costs (but longrun benefits), so F0 () < 0. Nature can invest at t 0 in a somatic stock, K,
where this is interpreted as the brain. At t 0, Nature can also divert energy s
to increase the probability of survival to t 1, p(s), say, where p0 (s) > 0,
p00 (s) < 0. For example, the range and/or the depth of immune function can
be increased, at a metabolic cost. At t 1, gross energy output is F(K, ) > 0,
so adults generate a surplus, which can be used to foster children. In addition,
production satisfies FK(K, ) > 0, FKK(K, ) < 0, F(K, ) > 0, and
FK(K, )  0.
Net energy flow at t 0 is F  K  s < 0, whereas at t 1, it is
F(K, ) > 0. In a large population in steady-state growth at rate r, the proportion of old to young is ( p(s)/(1 r)). If the energy flow is food, for example,
and so is non-storable but also is not wasted, the growth rate r is then given by
ps
FK;  0:
F  K  s
1r
The steady state is feasible if and only if the adults cover the deficits of the
children. Hence, the growth rate is
1r

psFK; 
;

K s  F

where it will also be assumed that r* 0.

An increase in the parameter  reflects the advent of a more learningintensive environment. Further, suppose that  maintains the zero growth
rate, implicitly capturing the ultimate effect of environmental limits. It can
then be shown that @s*/@ > 0 and @K*/@ > 0. That is, a shift towards such
an environment causes a simultaneous increase in both intelligence and longevity, as in the human data.

4.2. A sketch of a general continuous time model

Consider, now, the more general model of Robson and Kaplan (2003). Again,
it is necessary to go beyond the two-period case in order to capture U-shaped
mortality and to be sure that the above comparative statics results are not an
artefact of that simple case.
In this more general model, K is still neural capital. The gross energy output
of the individual at age t is then given by the production function F(K, t). This
function is increasing and concave in K and depends on t as follows. First,

Economics of hunter-gatherer societies

393

2,000,000
1,000,000
0

Net Calories

1,000,000
2,000,000
3,000,000
4,000,000
Human Prod, Human Surv.
Human Prod., Chimp. Surv.
Chimp. Prod. Chimp. Surv

5,000,000
6,000,000
7,000,000
0

10

15

20

25

30

35

40

45

50

55

60

65

Age
FIGURE 2 Cumulative expected energy production net of consumption, by age
SOURCE: This is adapted from Kaplan et al. (2001), to which reference should be made for data
sources.

there is an initial range of ages, t, over which F(K, t) rises. The key factor
leading to rising productivity is learning-by-doing. Thus, a large brain, K,
induces high levels of F(K, t), which may only become evident at ages t long
after brain growth itself has ended. Second, senescence is captured in a terminal range of ages over which F(K, t) declines. That is, as justified by the
analysis of the previous section, aging is now built in.
Output can be used as investment, v, which augments the capital stock as
dK
v; where v 2 0; v:
dt
The upper bound on investment v represents a simple specification of an
increasing cost of investment, which is plausible in the current setting. For
analytic simplicity, neural investment is now constrained to occur in a block of
time at the beginning of life. That is, for some t  0,

v > 0
 2 0; t
v
:
0
 t
In addition, there is assumed to be a constant initial energy cost C0  K0 > 0
for each individual. In terms of the model of the previous section, this

394 A.J. Robson and H.S. Kaplan

assumption is equivalent to assuming that the cost of fertility is linear and
independent of the level of gross energy, G. That is, it is equivalent to specifying that F(G, s) G  s. This simplifies the problem, but means that the
pattern of fertility is indeterminate.
Essentially, as in the previous model, some of the energy output, s, reduces
the rate of mortality, (s). The probability of survival to age t, p(t), say, satisfies
dpt
pts where p0 1:
dt
The net output is y F(K)  v  s.
There is an essentially unique optimal path for s, determined by the condi0
~
tion that J(t)
(s) 1, where
Z 1
~ 1
Jt
pFK;   v  sd:
pt t
In this model, it can be shown that the maximization of the steady state growth
rate, together with the requirement that this maximum growth rate be zero, can
be achieved by maximizing the undiscounted lifetime expected net output of
energy excluding the cost C0:
Z 1
~
J0
pFK; t  v  sdt;
0

where, in addition, the maximum value of J~ is equal to C0. Indeed, given that
optimal neural investment ends before output peaks, optimal mortality  is
U-shaped, with a minimum that is no later than peak output.

4.3. Comparative statics

Consider, now, the effect of a more challenging environment, modelled by an
increase in a parameter  introduced into the production function. Suppose the
increase in  decreases output right after brain growth ends, but eventually
increases output. This represents the advent of a more cognitively demanding
foraging activity. Learning this activity results in a substantial ultimate
increase in productivity, but this is predicated on investment in a large brain
in the first place. In addition, such on-the-job-training may also necessitate
sacrificing an occupation that is more productive at first. Suppose, however,
that the overall effect of the increase in  is either to raise or to hold constant
total expected energy surplus, and also that increasing  does not lower the
marginal product of capital. It can then be shown that such a shift raises K and
reduces mortality at every age, thus increasing longevity.
Alternatively, it can also be shown that a more favourable environment, in
the form of an exogenous reduction in mortality, also induces such shifts in K
and mortality. That is, if  parameterizes an additive shift in , reducing  also
has these two effects.

Economics of hunter-gatherer societies

395

These comparative static results imply that human intelligence and life
expectancy might have co-evolved as follows. Consider the circumstances
when a drier climate in Africa precipitated the replacement of the African
rain forests by savanna, several million years ago. The result was to foster
greater numbers of herbivores, for example, as well as energy intensive plant
resources. The skill intensity needed to exploit this new foraging niche plausibly raised the productivity of the brain. Greater expenditure on mortality
reduction then became worthwhile and longevity rose. In addition, greater
investment in the brain itself also paid off.
The skills, weapons, and social organization in early humans required for
hunting must simultaneously have reduced predation on humans. This reduction in exogenous mortality caused an additional decrease in endogenous
mortality. The additional reduction in overall mortality then induced an
further increase in investment in the brain. These two effects from increasing
productivity and from reducing extrinsic mortality therefore reinforced one
another.
These comparative statics results also illuminate the differences that presently exist between humans and our closest living relatives, the chimpanzees,
as discussed in the introduction. That is, after humans evolved to fill a niche on
the African savanna characterized by somewhat higher skill intensity and lower
mortality, perhaps we continued to be positioned to fill a series of niches with
yet higher skill intensities and lower mortalities, while chimpanzees remained
in a niche with relatively low skill intensity and relatively high mortality. The
above comparative statics results are then consistent with our greater intelligence our brains are about three times as large as those of chimpanzees and
our greater longevity, since we live about twice as long.
Our approach, more generally, illuminates the evolution of species with low
rates of mortality and low rates of senescence. That is, we show how the
productivity of capital affects human life history evolution. Circumstances
where capital is highly productive favour reduced senescence not only in
humans, but also in diverse non-human species. For example, the extreme
longevity of queens in social insect colonies may reflect the productivity of
various forms of capital not only somatic capital, but also non-somatic
physical capital, the hive or nest, for example.
5. Conclusions
In this paper, we have advocated consideration of the evolutionary origins of
human demographic and economic characteristics. Going beyond this, we have
illustrated how the economics of hunting and gathering societies was the anvil
on which natural selection formed these innate characteristics. We have considered the shaping of the basic human characteristics of mortality, fertility,
and intelligence. In particular, we have explained why our mortality profile is
U-shaped, but why we also live well beyond the cessation of fertility. We have

396 A.J. Robson and H.S. Kaplan

also explained why human intelligence and longevity has increased in a lockstep fashion.
Indeed, the remarkable similarities evident in figure 1, for example, between
foraging and modern economies are to be expected under our approach. That
is, our innate capacities and preferences were adapted by natural selection to
the foraging economy. These capacities and preferences then make it optimal
for us to reproduce key features of the foraging economy within the modern
economy.
There are a number of additional topics that we intend to investigate. In the
introduction, we discussed the sexual division of labour in foraging societies. It
would be interesting to capture the forces inducing this division in a formal
model. Perhaps, for example, a relatively small difference between the sexes
based on their biological roles in reproduction would be amplified by the
demands of the foraging economy. Such a model might ultimately help to
address the question: Why do women now live longer than men?
Another important topic concerns the formation of the rate of time preference as a function of age. Here, the existence of intergenerational transfers
has implications for the rate of time preference of older individuals. The
prospect of imminent death of the individual would have less dramatic effects
on time preference when the resources involved in the intertemporal trade-off
accrue substantially to offspring.
A final ambitious aim is to consider how strategic behaviour has been
shaped by the economics of hunting and gathering. This concerns the extent
that strategic behaviour is not completely rational. Perhaps, for example,
humans are conditionally cooperative willing to anticipate cooperation at
first, and therefore themselves to be cooperative initially, despite also energetically punishing defection. Such a propensity might have arisen readily in
small groups of hunter-gatherers, in which each individual knew all other
members, being indeed genetically related to many of them.8
The pioneers of modern economic theory, such as Paul Samuelson, enjoyed
tremendous success in applying relatively simple mathematics to provide convincing explanations of basic economic phenomena. However, the very success
of these early pioneers means that such possibilities that still exist within
conventional economics are now few and far between. On the other hand,
there is a vast, untouched expanse of territory awaiting exploration in the
adjoining hinterlands of economics, biology, and anthropology. This is a
golden opportunity to apply simple optimization models to explain important
phenomena in a convincing way.

8 See Seabright (2004) for vivid descriptions of such phenomena.

Economics of hunter-gatherer societies

397

References
Allman, John, Todd McLaughlin, and Atiya Hakeem (1993) Brain weight and life-span
in primate species, Proceedings of the National Academy of Sciences of the USA 90,
11822
Charlesworth, Brian (1994) Evolution in Age-Structured Populations (Cambridge:
Cambridge University Press)
(2000) Fisher, Medawar, Hamilton and the evolution of aging, Genetics 156, 92731
Ehrlich, Isaac, and Hiroyuki Chuma (1990) A model of the demand for longevity and
the value of life extension, Journal of Political Economy 98, 76182
Finch, Caleb E. (1998) Variations in senescence and longevity include the possibility of
negligible senescence, Journal of Gerontology: Biological Sciences 53A, B235B239
Grossman, Michael (1972) On the concept of health capital and the demand for health,
Journal of Political Economy 80, 22355
Hamilton, William D. (1966) The moulding of senescence by natural selection, Journal
of Theoretical Biology 12, 1245
Hill, Kim, and Anna Magdalena Hurtado (1996) Ache Life History: The Ecology and
Demography of a Foraging People (Hawthorne, NY: Aldine)
Holliday, Malcolm A. (1978) Body composition and energy needs during growth, in
Human Growth, ed. F. Falker and J. Tanner (New York: Plenum)
Howell, N. (1979) Demography of the Dobe !Kung (New York: Academic Press)
Jerison, Harry J. (1973) Evolution of the Brain and Intelligence (New York: Academic
Press)
Jurmain, Robert, Harry Nelson, Lynn Kilgore, and Wenda Trevathan (2000)
Introduction to Physical Anthropology (Belmont, CA: Wadsworth)
Kaplan, Hillard S., and Kim Hill (1985) Food-sharing among Ache foragers: tests of
explanatory hypotheses, Current Anthropology 26, 22345
Kaplan, Hillard S., and Arthur J. Robson (2002) The emergence of humans: the
coevolution of intelligence and longevity with intergenerational transfers,
Proceedings of the National Academy of Sciences 99, 10221226
Kaplan, Hillard S., Kim Hill, A. Magdalena Hurtado, and Jane B. Lancaster (2001)
The embodied capital theory of human evolution, in Reproductive Ecology and
Human Evolution, ed. P. Ellison (Hawthorne, NY: Aldine de Gruyter)
Kaplan, Hillard S., Kim Hill, Jane B. Lancaster, and Anna Magdalena Hurtado (2000)
A theory of human life history evolution: diet, intelligence, and longevity,
Evolutionary Anthropology 9, 15685
Kaplan, Hillard S., Jane B. Lancaster, and Arthur J. Robson (2003) Embodied capital
and the evolutionary economics of the human lifespan, Population and Development
Review 29, Supplement, 15282
Kirkwood, Thomas B.L. (1990) The disposable soma theory of aging, in Genetic
Effects on Aging II, ed. D. Harrison (Caldwell, N.J.: Telford Press)
(1999) Time of Our Lives (New York: Oxford University Press)
Love, Seth, and Peter Jenner (1999) Oxidative stress in neurological disease, Brain
Pathology 9, 556; introduction to a symposium available at http://www.brainpath.
medsch.ucla.edu/brainpath/abstracts/vol9/901/901toc.html
Martin, M.J., S.B. Hulley, W.S. Browner, L.H. Kuller, and D. Wentworth (1986)
Serum cholesterol, blood pressure, and mortality: implications from a cohort of
361,662 men, Lancet 8513, 9336
Martin, Robert D. (1981) Relative brain size and basal metabolic rate in terrestrial
vertebrates, Nature 293, 5760
McElwee, Joshua. J., Eugene Schuster, Eric Blanc, James H. Thomas, and David Gems
(2004) Shared transcriptional signature in Caenorhabditis elegans dauer larvae and

398 A.J. Robson and H.S. Kaplan

long-lived daf-2 mutants implicates detoxification system in longevity assurance,
Journal of Biological Chemistry 279, 4453343
Medawar, Peter B. (1952) An Unsolved Problem in Biology (London: Lewis)
Reader, Simon M., and Kevin N. Laland (2002) Social intelligence, innovation, and
enhanced brain size in primates, Proceedings of the National Academy of Sciences of
the USA 99, 443641
Robson, Arthur J. (1996a) The evolution of attitudes to risk: lottery tickets and relative
wealth, Games and Economic Behavior 14, 190207
(1996b) A biological basis for expected and non-expected utility, Journal of
Economic Theory 68, 397424
Robson, Arthus J., and Hillard S. Kaplan (2003) The evolution of human longevity
and intelligence in hunter-gatherer economies, American Economic Review 93,
15069
(2005) Why we grow large and then grow old: economics, biology, and mortality,
Simon Fraser University Working Paper
Rose, Michael (1991) The Evolutionary Biology of Aging (Oxford: Oxford University
Press)
Seabright, Paul (2004) The Company of Strangers (Princeton, NJ: Princeton University
Press)
Smith, B. Holly (1991) Dental development and the evolution of life history in hominidae, American Journal of Physical Anthropology 86, 15774
Sozou, Peter and Rob Seymour (2004) To age or not to age, Proceedings of the Royal
Society: Biological Sciences 271, 45763
Stearns, Stephen C. (1992) The Evolution of Life Histories (Oxford: Oxford University
Press)
Vaupel, James W. (1997) Trajectories of mortality at advanced ages, in Between Zeus
and Salmon: The Biodemography of Longevity, ed. K. Wachter, and C. Finch
(Washington, DC: National Academy Press)
Williams, George C. (1957) Pleiotropy, natural selection, and the evolution of
senescence, Evolution 11, 398411