Human Behavioral Ecology Online

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Human Behavioral Ecology Bobbi Low
• LAST MODIFIED: 26 May 2016
• DOI: 10.1093/obo/9780199941728-0076
• Online at: http://www.oxfordbibliographies.com/view/document/obo-
9780199941728/obo-9780199941728-0076.xml
Introduction
Human behavioral ecology (HBE) asks how environmental conditions—physical, biotic, and
social—shape human lifetimes and behavior. Along with evolutionary anthropology,
evolutionary psychology, and dual-inheritance theory it is one of four major evolutionary
approaches to the study of human behavior. Based in evolutionary theory, behavioral ecology
tests hypotheses designed to understand the ultimate causation of the patterns we see—the
selective pressures that shape these patterns. It seeks to understand the actual function of
behaviors. Behavioral ecology employs selectionist logic (in any given environment, what
strategy is fitness maximizing?), decision rules to understand conditional strategies and
facultative traits, and optimization techniques. It begins with simple models and employs what
we call the “phenotypic gambit”—a simplifying assumption that allows us to examine behaviors
without regard to the particulars of inheritance. Analyses may be ethnographic studies of one
society, comparative (e.g., cross-cultural), or experimental (game theoretic), or may employ
modeling. Of course, in modern human groups, this gambit may or may not be effective; we
change environments quite rapidly, but our behavior is extraordinarily flexible. Thus, great care
is important in defining terms and making inferences. There is considerable overlap between
HBE and evolutionary anthropology; a writer’s choice of terms appears to reflect his or her
academic background. Dual-inheritance theory also overlaps greatly because it seeks to
understand the roles cultural transmission has in shaping our evolution and behavior. The related
field of evolutionary psychology caught journalists’ eyes, and the term has come to be widely
used, even when it is not appropriate. Many evolutionary psychologists are concerned primarily
with mental models and the forces that shape them, but journalists sometimes call papers on life
history pacing, for example, “evolutionary psychology,” although they clearly are not. In short,
interesting papers can be found in all these categories—which in fact can overlap greatly.
General Overviews
A number of works reflect the various approaches to HBE, ranging from overviews for a broad
audience to more-detailed summaries and analyses of different approaches for specialists. Cronk
1991 and Borgerhoff Mulder 1991 are perhaps the earliest works to use the term “human
behavioral ecology” in print; they are classic introductions. Smith 2000, an important
contribution, delineates the sometimes-subtle differences among behavioral ecology and other
approaches to studying adaptive responses of humans. Winterhalder and Smith 2000 is a
coherent assessment of evolutionary (or behavioral) ecology from its beginnings, with excellent
coverage. Low 2015 takes a life history approach to sex differences in traditional, historical, and
(less deeply) modern Western societies. Nettle, et al. 2013 is a forward-looking summary; Brown
and Richerson 2014 gives a nice analysis of modern controversies and tracks developments in
cultural-evolution theory. Laland and Brown 2011 not only tracks the development of theory, but
it treats early and nontrivial political attacks. A fine resource for anyone mastering early-21st-
century papers is Kermyt Anderson’s online compendium of behavioral ecology papers from
2000 through 2013 (Human Behavioral Ecology Bibliography).
• Borgerhoff Mulder, M. 1991. Human behavioural ecology. In Behavioural ecology: An

evolutionary approach. 3d ed. Edited by J. R. Krebs and N. B. Davies, 69–98. London:
Blackwell Scientific.
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This overview analyzes human patterns in the context of broad behavioral ecology. It
may have been the first time a standard work on behavioral ecology written for biologists
included a review of work on HBE. The volume was the third of a series of four edited
volumes on current issues in behavioral ecology.
Find this resource:
• Brown, G. R., and P. J. Richerson. 2014. Applying evolutionary theory to human

behaviour: Past differences and current debates. Journal of Bioeconomics 16.2: 105–128.
DOI: 10.1007/s10818-013-9166-4Save Citation »Export Citation »E-mail Citation »
This modern comparison of the fields considering human behavior has excellent
summaries; it also clarifies and tracks early-21st-century advances in cultural evolution.
Find this resource:
• Cronk, L. 1991. Human behavioral ecology. Annual Review of Anthropology 20:25–53.
DOI: 10.1146/annurev.an.20.100191.000325Save Citation »Export Citation »E-mail

Citation »
This cogent summary of the basics of HBE is one of the first scholarly treatments of the
subject. Though much has been learned in the ensuing time, this continues to be a very
nice introduction.
Find this resource:
• Human Behavioral Ecology Bibliography (HBEB).

This is not, strictly speaking, an introduction to HBE; rather, it is a comprehensive
collection of HBE papers from 2000 through 2013. A labor of love by Kermyt Anderson,
it is a wonderful tool to uncover papers in HBE.
Find this resource:
• Laland, K. N., and G. R. Brown. 2011. Sense and nonsense: Evolutionary perspectives on
human behaviour. 2d ed. Oxford: Oxford Univ. Press.
This is a balanced treatment of evolutionary approaches to human behavior, and a good

summary of early political attacks. Its last chapter compares the approaches, using the
broader “cultural evolution” rather than “evolutionary anthropology,” to make clear the
concept that selective forces are still active in the early 21st century.
Find this resource:
• Low, B. S. 2015. Why sex matters: A Darwinian look at human behavior. Rev. ed.
Princeton, NJ: Princeton Univ. Press.
DOI: 10.1515/9781400852352Save Citation »Export Citation »E-mail Citation »
This examines life history strategies and sex differences; original edition was published
in 2000. Covers basics of selection (chapters 1, 2), life history (6, 7, 8), mating effort (2,
4, 5), cooperation and conflicts (9, 10), and warfare (13, 14). The final chapter (15)
examines how our evolutionary past—the way we evolved to get and use resources—
influences modern conservation issues.
Find this resource:
• Nettle, D., M. A. Gibson, D. W. Lawson, and R. Sear. 2013. Human behavioral ecology:
Current research and future prospects. Behavioral Ecology 24.5: 1031–1040.
DOI: 10.1093/beheco/ars222Save Citation »Export Citation »E-mail Citation »
This assessment of the strengths of a behavioral-ecological approach emphasizes more-

recent papers, including those on modern industrial populations. It summarizes the
growth of HBE in a number of major journals. Commentaries follow the paper.
Find this resource:
• Smith, E. A. 2000. Three styles in the evolutionary analysis of human behavior. In

Adaptation and human behavior: An anthropological perspective. Edited by L. Cronk, N.
Chagnon, and W. Irons, 27–46. Evolutionary Foundations of Human Behavior. New
York: Aldine de Gruyter.
Smith compares behavioral ecology, evolutionary psychology, and dual-inheritance

theory in their approaches and assumptions, making clear the advantages and pitfalls of
each. The boundaries can be fuzzy. This is a must-read for anyone hoping to understand
the similarities and divergences (and arguments) among these closely related fields.
Find this resource:
• Winterhalder, B., and E. A. Smith. 2000. Analyzing adaptive strategies: Human

behavioral ecology at twenty-five. Evolutionary Anthropology: Issues, News, and
Reviews 9.2: 51–72.
This collection is a unified, coherent overview of advances in selectionist thinking from

the mid-1970s to 2000. It covers theoretical foundations, closest kin, resources, work and
space, and reproduction and social relations.
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Journals
Papers on behavioral ecology appear in a number of journals, at least two of which are specific to
HBE, evolutionary anthropology, dual-inheritance theory, and evolutionary psychology.
Evolution and Human Behavior is the official journal of the Human Behavior and Evolution
Society and, along with Human Nature, is exclusively dedicated to papers focused on
evolutionary analyses of human behavior and demography. Evolutionary Anthropology is an
important source of current reviews. Current Anthropology includes many papers on behavioral
ecology, as do Cross-Cultural Research and American Journal of Human Biology. Behavioral
Ecology and Behavioral Ecology and Sociobiology often include papers on humans. A high-
quality, broad-ranging journal that often features HBE is Proceedings of the Royal Society B:
Biological Sciences. Students should note that when accessing journals online, using your
institution’s library links may give you free access to papers.
• American Journal of Human Biology.
An interdisciplinary journal of original research, theoretical articles, and occasional

reviews. HBE papers appear frequently.
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• Behavioral Ecology.
This is the official journal of the International Society for Behavioral Ecology; it
occasionally publishes papers on HBE.
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• Behavioral Ecology and Sociobiology.
This journal publishes HBE articles as well as papers on interactions between humans
and other species.
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• Cross-Cultural Research.
This journal publishes articles that span the social sciences, and it often includes HBE
papers.
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• Current Anthropology.
This journal is wide ranging, covering anthropological scholarship in many areas, from
evolution to musicology; it also includes papers on some nonhuman primates.
Find this resource:
• Evolution and Human Behavior.
This is the official journal of the Human Behavior and Evolution Society. It publishes
theoretical and empirical papers in behavioral ecology, evolutionary anthropology, and
evolutionary psychology.
Find this resource:
• Evolutionary Anthropology: Issues, News, and Reviews.

An authoritative review journal that focuses on issues of current interest in biological

anthropology, paleoanthropology, archaeology, functional morphology, social biology,
and bone biology—including dentition and osteology—as well as human biology,
genetics, and ecology.
Find this resource:
• Human Nature.
This specializes in evolutionary approaches to human behavior. It includes papers in

HBE, along with evolutionary anthropology and evolutionary psychology papers. An
excellent source.
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• Proceedings of the Royal Society B: Biological Sciences.
Dedicated to rapid publication of relatively short, high-quality papers. It aims to cover the
breadth of biology; most papers are on organismal biology. HBE papers that set human
data into the context of, for example, other primates’ behavioral ecology, appear
routinely.
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Textbooks
Currently, texts on behavioral ecology seldom integrate human data; an exception is Danchin, et
al. 2008. Boyd and Silk 2014, a text on evolutionary anthropology, is excellent. Some good
general texts on animal behavior have chapters or information on HBE (e.g., Dugatkin 2004,
Alcock 2013), and there are focused edited collections such as Chagnon and Irons 1979; Betzig
1997; Cronk, et al. 2000; and Low 2015 (the latter cited under General Overviews) that have
been used for classes, from undergraduate lecture courses to graduate seminars.
• Alcock, J. 2013. Animal behavior: An evolutionary approach. 10th ed. Sunderland, MA:
Sinauer.
Chapter 14, the last chapter, is on the evolution of human behavior. The book is highly
readable, with a wealth of examples.
Find this resource:
• Betzig, L., ed. 1997. Human nature: A critical reader. New York: Oxford Univ. Press.
This collection pairs classic analyses with each author’s retrospective about advances
since the time of the original publication, and thoughts about what the author might do
differently. An engaging exploration.
Find this resource:
• Boyd, R., and J. B. Silk. 2014. How humans evolved. 7th ed. New York: Norton.
This text on evolutionary anthropology could easily be titled “Human behavioral

ecology.” Robert Boyd and Joan Silk are evolutionary anthropologists with important
additional specialties that inform their work: Silk is a primatologist, and Boyd works on
the evolution of cooperation.
Find this resource:
• Chagnon, N. A., and W. Irons, eds. 1979. Evolutionary biology and human social
behavior: An anthropological perspective. North Scituate, MA: Duxbury.
This is the first edited collection of seminal papers in HBE, containing several classic
papers.
Find this resource:
• Cronk, L., N. Chagnon, and W. Irons, eds. 2000. Adaptation and human behavior: An
anthropological perspective. Evolutionary Foundations of Human Behavior. New York:
Aldine de Gruyter.
This book focuses on advances in human behavior ecology in the twenty years since the
publication of Chagnon and Irons 1979.
Find this resource:
• Danchin, É., L.-A. Giraldeau, and F. Cézilly. 2008. Behavioural ecology. New York:
Oxford Univ. Press.
This text incorporates data on a number of human traits, in the context of behavioral
ecology: infanticide, lactose intolerance, sex ratios, etc.
Find this resource:
• Dugatkin, L. A. 2004. Principles of animal behavior. New York: Norton.
This work is lavishly illustrated and very accessible. It has information on human
behavior integrated throughout the text: on aggression, alliances, family stability,
infanticide, homicide, polygyny, social learning and cultural transmission, and more.
Find this resource:
Human Lifetimes
Life history theory seeks evolutionary explanations for observed patterns in lifetimes, in the
context of how environmental selective pressures shape both life history and behavior. Major
trade-offs include those between current and future reproduction and, in any reproductive bout,
between quantity and quality of offspring. Such analyses help us understand variations in, for
example, age at first reproduction, length of gestation and infant dependency, growth rates,
population growth rates, and more. Stearns 1992 and Roff 1992 are general treatments of life
history evolution; each discusses human life history traits in comparison with other vertebrates.
More-specific focus on humans is provided in Hill and Kaplan 1999 and Mace 2000. Hill and
Hurtado 1996 is perhaps the best demographic and ecological study of a traditional society.
Bribiescas, et al. 2012 analyzes male life histories and, in that context, sex differences. Human
life histories are unusual in a number of ways; Kuzawa and Bragg 2012 discusses our
extraordinary plasticity, Kaplan, et al. 2000 treats the evolutionary impacts of our diets, and
Hawkes and Paine 2006 discusses longevity and women’s early reproductive senescence, which
the authors argue may also have helped create our unusual lifetimes.
• Bribiescas, R. G., P. T. Ellison, and P. B. Gray. 2012. Male life history, reproductive
effort, and the evolution of the genus Homo: New directions and perspectives. In Special
issue: Human biology and the origins of Homo. Current Anthropology 53.S6: S424–
S435.
Human males’ life histories are strikingly different from those of other hominid males,
and these differences are crucial to human life history evolution in general. For example,
male paternal care influences a suite of traits, such as large newborn size after a
predicted-length gestation, altriciality, early weaning, large brain, and more.
Find this resource:
• Hawkes, K., and R. R. Paine, eds. 2006. The evolution of human life history. School of
American Research Advanced Seminar. Santa Fe, NM: School of American Research
Press.
This edited collection explores the potential driving force of grandmothering in extending
human life spans (and creating the associated long post-reproductive period in women’s
lives), first proposed by Hawkes (also see Parental Effort).
Find this resource:
• Hill, K., and A. M. Hurtado. 1996. Ache life history: The ecology and demography of a
foraging people. Foundations of Human Behavior. New York: Aldine de Gruyter.
This remains perhaps the best and most thorough work integrating life history and
ecology of a human society.
Find this resource:
• Hill, K., and H. Kaplan. 1999. Life history traits in humans: Theory and empirical
studies. Annual Review of Anthropology 28:397–430.
DOI: 10.1146/annurev.anthro.28.1.397Save Citation »Export Citation »E-mail Citation »
A classic, this retains importance.
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• Kaplan, H., K. Hill, J. Lancaster, and A. M. Hurtado. 2000. A theory of human life
history evolution: Diet, intelligence, and longevity. Evolutionary Anthropology: Issues,
News, and Reviews 9.4: 156–185.
This paper suggests that important life history traits of humans—very large brain, long
life, extended juvenile dependence, and reproductive assistance by husbands, brothers,
and other family (“cooperative breeding” and “grandmothering”)—are possible because
early humans switched to calorie-dense food requiring considerable skill to obtain.
Find this resource:

• Kuzawa, C. W., and J. M. Bragg. 2012. Plasticity in human life history strategy:
Implications for contemporary human variation and the evolution of genus Homo. In
Special issue: Human biology and the origins of Homo. Current Anthropology 53.S6:
S369–S382.
Humans’ long development time, extensive allocare, and early weaning are characteristic
of our life history. Nonetheless, traits of human life history show extensive within- and
between-population variation. Explores how such phenotypic and developmental
plasticity may have arisen: phenotypic plasticity may have made rapid changes possible.
Find this resource:
• Mace, R. 2000. Evolutionary ecology of human life history. Animal Behaviour 59.1: 1–
10.
DOI: 10.1006/anbe.1999.1287Save Citation »Export Citation »E-mail Citation »
A succinct, well-written summary. Useful for seminars needing a concise overview.
Find this resource:
• Roff, D. A. 1992. The evolution of life histories: Theory and analysis. New York:
Chapman & Hall.
Roff’s classic is very well written and contains interesting analyses of important aspects
of human life histories in a comparative context.
Find this resource:
• Stearns, S. C. 1992. The evolution of life histories. New York: Oxford Univ. Press.
This classic treats some human data in a comparative context, setting human life history
into the same selective framework as other species.
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Somatic Effort
To succeed, any organism, including humans, must successfully survive and reproduce. Somatic
effort encompasses any investment into one’s own soma (essentially any part of the body not
dedicated to reproduction): growth, homeostatic regulation, foraging, and avoiding death or
disability from, for example, predators and pathogens. The two major areas of inquiry within
HBE are growth, attachment, anddevelopment; and optimal foraging.
Growth, Attachment, and Development
In HBE, much work in somatic effort has focused on child development (Hrdy 2005, Hewlett
and Lamb 2005, Hagen and Hammerstein 2005), health (Crespi 2011), and play and learning
(Lancy, et al. 2010). These represent areas in which what we learn about our evolutionary past
and life history constraints may have direct applications for health and development.
• Crespi, B. 2011. The evolutionary biology of child health. Proceedings of the Royal
Society B: Biological Sciences 278.1711: 1441–1449.
DOI: 10.1098/rspb.2010.2627Save Citation »Export Citation »E-mail Citation »
Human genetic and epigenetic studies indicate that juvenile growth, development, and
behavior are mediated by imprinted genes and resulting genomic conflicts. Cancers, the
primary cause of noninfectious childhood mortality, mirror child growth rates from birth
to adolescence.
Find this resource:
• Hagen, E. H., and P. Hammerstein. 2005. Evolutionary biology and the strategic view of
ontogeny: Genetic strategies provide robustness and flexibility in the life course.
Research in Human Development 2.1–2: 83–97.
Much developmental work is difficult to set into an evolutionary context, yet

environmental conditions clearly influence the success of development strategies. This
uses game-theoretic approaches to synthesize an evolutionary argument that
flexible/conditional strategies are founded on genetically encoded strategies shared by
most of a population.
Find this resource:
• Hewlett, B. S., and M. E. Lamb, eds. 2005. Hunter-gatherer childhoods: Evolutionary,

developmental, and cultural perspectives. Evolutionary Foundations of Human Behavior.
New Brunswick, NJ: Aldine Transaction.
This edited collection provides cross-cultural ethnographic information on childhood.

The chapters vary in emphasis and degree of theoretical underpinnings: some chapters are
not evolutionary or behavioral ecological, instead focus on developmental or cultural
aspects of childhood.
Find this resource:
• Hrdy, S. B. 2005. Evolutionary context of human development: The cooperative breeding

model. In Attachment and bonding: A new synthesis. Edited by C. S. Carter, L. Ahnert,
K. E. Grossmann, et al., 9–32. Dahlem Workshop Reports 92. Cambridge, MA: MIT.
This is a clear exploration of the Pleistocene context that likely led to cooperative
breeding, an unusual pattern for an ape, and one that probably allowed for development
of further human traits that resulted in Homo being so successful competitively.
Find this resource:
• Lancy, D. F., J. Bock, and S. Gaskins, eds. 2010. The anthropology of learning in
childhood. Walnut Creek, CA: AltaMira.
This edited volume provides an excellent overview of the behavioral ecology of learning
cross-culturally. It examines cultural, social, and ecological influences; learning mores as
well as skills; adult-child interactions in learning; and some aspects of learning in novel
modern societies.
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Optimal Foraging
Optimal foraging is an important focus in behavioral ecology. Humans are diverse in their
foraging techniques, main sources of calories, and cultural arrangements, so that this is a broad
field, from analyses of nonhuman prey choice to complex and coordinated efforts by humans
(Winterhalder and Smith 1981, Smith 1991, Belovsky 1988). Analyses of optimality in
nonhuman foraging are closely related (Stephens and Krebs 1986). Some studies of optimal
foraging have begun to recognize that studies of humans can contribute to theory (Stephens, et
al. 2007). Exciting early-21st-century work demonstrates the utility of behavioral ecology in
analyzing prehistoric patterns (Bird and O’Connell 2006, Garvey and Bettinger 2014) and
understanding more fully the changes arising during the Neolithic revolution (Kennett and
Winterhalder 2006).
• Belovsky, G. E. 1988. An optimal foraging-based model of hunter-gatherer population

dynamics. Journal of Anthropological Archaeology 7.4: 329–372.
DOI: 10.1016/0278-4165(88)90002-5Save Citation »Export Citation »E-mail Citation »

Belovsky used linear programming to determine optimal diet for a number of hunter-
gatherer populations. He built on earlier work on generalist herbivores and tested models
against empirical data. Because his models included the demands of dependent
nonforagers, they supersede earlier models that considered foragers as independent
individuals.
Find this resource:
• Bird, D. W., and J. F. O’Connell. 2006. Behavioral ecology and archaeology. Journal of
Archaeological Research 14.2: 143–188.
This paper is concise, a fine example of how behavioral-ecological logic and inference
can be useful and offer insights in fields not previously associated with behavioral
ecology.
Find this resource:
• Garvey, R., and R. L. Bettinger. 2014. Adaptive and ecological approaches to the study
of hunter-gatherers. In The Oxford handbook of the archaeology and anthropology of
hunter-gatherers. Edited by V. Cummings, P. Jordan, and M. Zvelebil, 69–91. Oxford
Handbooks. Oxford: Oxford Univ. Press.
This is a comprehensive and well-written comparison of how adaptive approaches to

understanding human behavior compare to anthropological approaches. It documents
how anthropology has experienced a paradigm shift over time, moving from largely
descriptive pieces to more hypothesis-testing work (e.g., evolutionary anthropology).
Find this resource:
• Kennett, D. J., and B. Winterhalder, eds. 2006. Behavioral ecology and the transition to
agriculture. Origins of Human Behavior and Culture 1. Berkeley: Univ. of California
Press.
This volume may be the first in which archaeologists and ethnographers use behavioral-
ecological approaches to understand what was involved in the Neolithic revolution, as
hunter-gatherers settled down and began growing crops. Advances in theory are covered
and empirical data are given their due.
Find this resource:

• Smith, E. A. 1991. Inujjuamiut foraging strategies: Evolutionary ecology of an Arctic
hunting economy. Foundations of Human Behavior. New York: Aldine de Gruyter.
This is an early classic study; it is both detailed and broad ranging, a prime example of a
well-done study on human foraging.
Find this resource:
• Stephens, D. W., J. S. Brown, and R. C. Ydenberg, eds. 2007. Foraging: Behavior and
ecology. Chicago: Univ. of Chicago Press.
This broad and well-edited work is a comprehensive review of advances in foraging

theory for the previous two decades or more. It highlights the breadth of foraging theory
and includes chapters on human foraging.
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• Stephens, D. W., and J. R. Krebs. 1986. Foraging theory. Monographs in Behavior and
Ecology. Princeton, NJ: Princeton Univ. Press.
This classic is the biological foundation for much of the literature on human optimal
foraging.
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• Winterhalder, B., and E. A. Smith, eds. 1981. Hunter-gatherer foraging strategies:

Ethnographic and archeological analyses. Prehistoric Archeology and Ecology. Chicago:
Univ. of Chicago Press.
This is a classic collection of analyses of optimal foraging for a number of hunter-

gatherers, focusing primarily on adult optimal-foraging strategies.
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Reproductive Effort
Reproductive effort encompasses calories spent and risks taken to achieve matings and to
produce offspring that survive to adulthood: mating effort and parental effort (Geary 2010;
Betzig, et al. 1988; Ellison 2003; Burger, et al. 2010). In general, the behaviors that make an
individual successful in one of these endeavors are costly in terms of the other: what helps a man
gain status or resources is antithetical to careful parenting, for example, because the most-
effective competitive behaviors carry survival risks (e.g., see Low 2015, cited under General
Overviews). Thus, the reproductive interests of males and females often diverge. Males’
reproductive return for effort in polygynous species is typically characterized by considerable
mating effort, with a very large fixed cost even to enter the mating competition at all. When
males are highly parental, they typically have relatively low testosterone (Alvergne, et al. 2009).
This appears to be true for humans, although humans are unusual among primates in having
considerable male parental care (see Parental Effort) as well. Female primates, including
humans, typically focus effort on parental duties. These characteristic sex differences in spending
reproductive effort arise partly from disruptive selection in the evolution of anisogamy (Parker,
et al. 1972) and partly from a cytoplasmic “war of all against all” among gametes in species in
which gametes combine (Hurst 1992). Sociocultural factors can also obviously influence
optimal-mating strategies (Schacht and Borgerhoff Mulder 2015).
• Alvergne, A., C. Faurie, and M. Raymond. 2009. Variation in testosterone levels and
male reproductive effort: Insights from a polygynous human population. Hormones and
Behavior 56.5: 491–497.
DOI: 10.1016/j.yhbeh.2009.07.013Save Citation »Export Citation »E-mail Citation »
This is one of the first papers to find that male testosterone levels covaried with men’s
allocation of reproductive effort differentially to mating versus parental effort. Married
fathers had lower testosterone levels than unmarried non-fathers. Among fathers, more
paternal men had lower testosterone than others; polygynous fathers had higher
testosterone.
Find this resource:
• Betzig, L., M. Borgerhoff Mulder, and P. Turke, eds. 1988. Human reproductive
behaviour: A Darwinian perspective. Cambridge, UK: Cambridge Univ. Press.
This edited collection is an anthropological tour through a variety of human societies,

exploring selective aspects of human mating and parental strategies in a variety of
conditions and for different societies. Topics primarily consider behavioral-ecological
analyses of mating and parenting.
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• Burger, O., R. Walker, and M. J. Hamilton. 2010. Lifetime reproductive effort in humans.
Proceedings of the Royal Society B: Biological Sciences 277.1682: 773–777.

If one measures the total amount of metabolized energy diverted to reproduction over
lifetimes, using standard measures of reproductive effort, human values converge on the
theoretically predicted value of 1.4. Human lifetime reproductive effort is not an outlier:
demographic values are traded off in a predictable and constrained manner.
Find this resource:
• Ellison, P. T. 2003. Energetics and reproductive effort. American Journal of Human

Biology 15.3: 342–351.
DOI: 10.1002/ajhb.10152Save Citation »Export Citation »E-mail Citation »
Natural selection favors optimal allocations of effort, and trade-offs exist. As a result,
human reproductive physiology exhibits traits that can be viewed as mechanisms to help
optimize the allocation of effort to different tasks. This paper reviews a number of such
trade-offs.
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• Geary, D. C. 2010. Male, female: The evolution of human sex differences. 2d ed.
Washington, DC: American Psychological Association.
DOI: 10.1037/12072-000Save Citation »Export Citation »E-mail Citation »
This is a comprehensive review of sexual selection; in particular it asks how sexual

selection shapes many aspects of human life even today. Geary moves from the evolution
of basic sex differences through how these differences play out in interactions with
environmental conditions.
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• Hurst, L. D. 1992. Intragenomic conflict as an evolutionary force. Proceedings of the

Royal Society B: Biological Sciences 248.1322: 135–140.
This foundational paper explains why humans (and other species producing gametes that
combine) face special conflicts. These conflicts result in anisogamy: the gametes we call
sperm profit from losing cytoplasm before combining with eggs. Laurence Hurst’s
explanations and those of Geoff Parker and colleagues (Parker, et al. 1972) are not
alternative hypotheses; most scholars think both operate.
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• Parker, G. A., R. R. Baker, and V. G. F. Smith. 1972. The origin and evolution of gamete
dimorphism and the male-female phenomenon. Journal of Theoretical Biology 36.3:
529–553.
This seminal paper modeled disruptive selection resulting in anisogamy. Two tasks must
be accomplished in successful reproduction: meeting other gametes and producing well-
invested zygotes. Intermediate-sized gametes lose to small gametes in the first task, and
to large gametes in the second task. The result: gametes of quite different size and
mobility.
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• Schacht, R., and M. Borgerhoff Mulder. 2015. Sex ratio effects on reproductive strategies
in humans. Royal Society Open Science 2.1: 140402.
DOI: 10.1098/rsos.140402Save Citation »Export Citation »E-mail Citation »
The evolution of anisogamy does not dictate mating strategies, of course. Fisher 1958
(cited under Mating Effort, Mate Choice, and Resource versus Reproductive Value) notes
that the relative availability of each sex as mates shaped the relative value of each sex in
a mating market. Here, that is demonstrated in humans. Schacht and Borgerhoff Mulder
found that men’s mating strategies (degree of sociosexuality) varied predictably with the
adult sex ratio—the mating market.
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Mating Effort, Mate Choice, and Resource versus Reproductive Value
Differences in male and female patterns reach their height at sexual maturity (Gray and Garcia
2013; Conroy-Beam, et al. 2015; Thornhill and Gangestad 2008). Like most other mammals,
human males typically expend more mating effort than do females; men show off their prowess
(Hawkes and Bliege Bird 2002) and take risks (e.g., Wang, et al. 2009) as they compete. As a
result, male and female survivorship curves differ because, typically, men’s risk taking affects
survivorship more than women’s (e.g., Kruger and Nesse 2006), and male mortality is higher
than female mortality after sexual maturity. Further, what each sex seeks in the other differs: in
most societies, a woman’s mate value is her reproductive value (defined in Fisher 1958), while a
man’s mate value is typically resource value of some sort (e.g., wealth, hunting prowess, status).
These lead to robust differences in qualities each sex seeks in a mate, even when careful
multivariate analyses are used. There are variations cross-culturally, but assortative mating
(Zietsch, et al. 2011) is common, and the pattern is clear and strong.
• Conroy-Beam, D., D. M. Buss, M. N. Pham, and T. K. Shackelford. 2015. How sexually

dimorphic are human mate preferences? Personality and Social Psychology Bulletin 41.8:
1082–1093.
The general pattern in mate choice is that men seek women of high reproductive value
(young) while women (or their families) seek men of high resource value (wealthy). This
study uses sophisticated statistical methods, finding that the overlap between male
preferences and female preferences is only about 22 percent: mate preferences are
robustly dimorphic.
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• Fisher, R. A. 1958. The genetical theory of natural selection. Dover Books on Science
S466. New York: Dover.
Fisher defined the important variable of reproductive value and calculated it for
Australian women (pp. 27–30): the number of daughters (sometimes now calculated as
children) a woman will have in the rest of her life, given the prevailing age-specific
fertility and mortality schedules. First published in 1930 (Oxford: Clarendon); most
recently published in 2013 (Miami, FL: HardPress).
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• Gray, P. B., and J. R. Garcia. 2013. Evolution and human sexual behavior. Cambridge,
MA: Harvard Univ. Press.
This is a well-written and thorough book that ranges from the evolution of sex differences
to cross-cultural variation in sexuality, adolescent sexual development, and developing
sexual behavior in cultural contexts.
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• Hawkes, K., and R. Bliege Bird. 2002. Showing off, handicap signaling, and the
evolution of men’s work. Evolutionary Anthropology: Issues, News, and Reviews 11.2:
58–67.
DOI: 10.1002/evan.20005Save Citation »Export Citation »E-mail Citation »
In many societies, men expend primarily mating effort. Successful effort must be
unbluffably expensive, demonstrating ability. This argues that the widespread male
competition of hunting large game may have evolved as an expensive signal of male
quality; men often ignore safer, less flashy foraging choices.
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• Kruger, D. J., and R. M. Nesse. 2006. An evolutionary life-history framework for
understanding sex differences in human mortality rates. Human Nature 17.1: 74–97.
DOI: 10.1007/s12110-006-1021-zSave Citation »Export Citation »E-mail Citation »
Because mating effort typically involves unbluffably expensive behavior to advertise

men’s abilities, it often involves risky behavior. Mating effort in humans is primarily a
male endeavor, so sex-differential survivorship curves are the norm: men die at a higher
rate than women throughout much of their lives. Modern transnational data are analyzed.
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• Thornhill, R., and S. W. Gangestad. 2008. The evolutionary biology of human female
sexuality. New York: Oxford Univ. Press.
Although the generalization is that females specialize in parental effort rather than mating
effort, women do compete for top males in societies in which men’s status benefits their
families, in societies in which men give significant parental investment, or both. This
book takes an evolutionary look at women’s sexuality and competition.
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• Wang, X. T., D. J. Kruger, and A. Wilke. 2009. Life history variables and risk-taking
propensity. Evolution and Human Behavior 30.2: 77–84.
DOI: 10.1016/j.evolhumbehav.2008.09.006Save Citation »Export Citation »E-mail

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Risk taking is not a singular phenomenon. In mating competition, for example, men may
take physical or social risks. This paper defines five domains of risk taking in the context
of between-group competition, within-group competition, mating and resource
acquisition, and fertility and reproduction.
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• Zietsch, B. P., K. J. H. Verweij, A. C. Heath, and N. G. Martin. 2011. Variation in human

mate choice: Simultaneously investigating heritability, parental influence, sexual
imprinting, and assortative mating. American Naturalist 177.5: 605–616.
This multivariate analysis of mate choice of twins found that the mate choice was
primarily assortative in initial choice, not through convergence. Natal family environment
also mattered. Height, body mass index, age, education, income, social attitudes, and
religiosity had no discernable independent impact.
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Mating and Marriage
Perhaps only the dunnock (Prunella modularis), a small, sparrow-like European bird, displays as
many different mating systems within the species as do humans: polygyny, social monogamy,
polyandry, polygynandry, and perhaps genetic monogamy. Marriage systems are clearly not
identical to mating systems, and marriage typically involves culturally defined transactions
(Huber, et al. 2011) and even dispersal (Towner 2002). The culturally defined system is typically
not the genetic system—which is why, for example, we distinguish social monogamy from
genetic monogamy (e.g., Reichart and Boesch 2003, Strassmann 2003). Nonetheless, both are
influenced by conflicts of interest between men and women. Typically, in polygynous systems
(which comprise most societies of the world, although not the majority of people), men’s
resources are important in obtaining wives: thus the importance of resource value for men. The
importance of women’s reproductive value is clearly seen in brideprice societies, in which
typically a younger woman commands a higher brideprice than others (e.g., Monique Borgerhoff
Mulder’s study of Kipsigis bridewealth payments in Betzig, et al. 1988, cited under Reproductive
Effort, pp. 65–82). Polygynous systems are the norm cross-culturally, and polyandry is quite rare
(Starkweather and Hames 2012, Smith 1998). Polyandry is often associated with agriculture on
poor land: when land is poor, and one man’s efforts may not be sufficient to raise the children of
one woman, fraternal polyandry can occur. Beyond this broad distinction, we are discovering
new and important subtle aspects to marriage rules cross-culturally; even the demographic
transition has important selective relationships to life history theory (Shenk 2009). A special
issue of Human Nature (Borgerhoff Mulder and Bliege Bird 2009) focuses on gendered
inequalities and has a number of papers on marriage disparities.
• Borgerhoff Mulder, M., and R. Bliege Bird, eds. 2009. Special issue: Gendered
inequalities in evolutionary perspective. Human Nature 20.2.
This issue, guest-edited by Borgerhoff Mulder and Bliege Bird, is dedicated entirely to
gendered inequalities and includes several papers on marriage.
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• Huber, B. R., W. F. Danaher, and W. L. Breedlove. 2011. New cross-cultural

perspectives on marriage transactions. Cross-Cultural Research 45.4: 339–375.
This is a nuanced approach to reanalyzing marriage patterns cross-culturally, adding

transactions (e.g., groom service, groom wealth) that have previously not been clearly
identified. Most societies have two or more kinds of transactions, although most result in
a transfer of wealth from the groom’s family to the bride’s.
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• Reichart, U. H., and C. Boesch, eds. 2003. Monogamy: Mating strategies and
partnerships in birds, humans and other mammals. Cambridge, UK: Cambridge Univ.
Press.
This edited collection makes the distinction between genetic and social monogamy; we
expect genetic monogamy only when males cannot profit reproductively from deserting
the female to seek additional matings. Chapters cover pair bonding in birds and
mammals, including humans.
Find this resource:
• Shenk, M. K. 2009. Testing three evolutionary models of the demographic transition:

Patterns of fertility and age at marriage in urban South India. In Special issue: Trade-offs
in female life histories: Integrating evolutionary frameworks. American Journal of
Human Biology 21.4: 501–511.
This paper examines the 20th-century demographic transition in India, testing three
models: the costs of children (with modernization, parents must invest more to ensure
success), effects of mortality (which typically decreases with transitions), and cultural
norms of imitating high-status families. All have significant impact, but the child-
cost/investment model appears strongest.
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• Smith, E. A. 1998. Is Tibetan polyandry adaptive? Methodological and metatheoretical

analyses. Human Nature 9.3: 225–261.
This is an examination of classic Tibetan polyandry. In agricultural systems, when one

man’s work cannot support a family, brothers may marry a single woman (fraternal
polyandry); typically, all children are counted as children of the elder brother. The system
kept land in the lineage, rather than having it dispersed.
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• Starkweather, K. E., and R. Hames. 2012. A survey of non-classical polyandry. Human
Nature 23.2: 149–172.
DOI: 10.1007/s12110-012-9144-xSave Citation »Export Citation »E-mail Citation »
Because polyandry is so rare in human societies, it is often difficult to study.

Starkweather and Hames identified fifty-three societies beyond those typically
represented, to analyze demographic and economic characteristics. They identified
characteristics beyond classical Himalayan and Marquesan systems, important in
understanding polyandry.
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• Strassmann, B. I. 2003. Social monogamy in a human society: Marriage and reproductive

success among the Dogon. In Monogamy: Mating strategies and partnerships in birds,
humans and other mammals. Edited by U. H. Reichart and C. Boesch, 177–189.
Cambridge, UK: Cambridge Univ. Press.
DOI: 10.1017/CBO9781139087247.012Save Citation »Export Citation »E-mail Citation

»
This paper on the Dogon of Mali documents classical social monogamy. The mating
system is biologically polygynous; male reproductive success (RS) varies more than
female RS. For each additional wife, a man fathered 1.8 extra offspring; in contrast,
monogamously married women averaged one more child than polygynously married
women.
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• Towner, M. C. 2002. Linking dispersal and marriage in humans: Life history data from
Oakham, Massachusetts, USA (1750–1850). Evolution and Human Behavior 23.5: 337–
357.
DOI: 10.1016/S1090-5138(02)00094-6Save Citation »Export Citation »E-mail Citation »
Dispersal in many primates appears linked to inbreeding avoidance, but human natal
dispersal may be more complex. This study found that resource availability mattered
(thus, men were less likely to disperse than women), as did competition for mates.
Inbreeding avoidance was less important.
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Parental Effort
Trade-offs are rife in parental care: size versus number of offspring, versus care expended
(evenly or differentially) on offspring. Parental investment, defined in Trivers 1972, is offspring
specific; parental effort (Low 1978) includes both offspring-specific and more general,
generalizable investment. Because of its zero-sum nature, parental investment considers trade-
offs affecting offspring. Lawson and Mace 2009 and Lawson and Mace 2011 explore parental
trade-offs and optimal family size. Parental effort is useful for examining cases with multiple
offspring when non-zero-sum efforts are examined (e.g., repeatedly used shelter). Parental care is
delicately balanced (Hagen, et al. 2006) and can vary across societies, with differential
occurrence of male parental care, for example. Maternal strategies are beautifully documented in
Hrdy 1999. Typically, because of our polygynous evolutionary history, raising a successful son
is more costly than raising a daughter; however, in some populations (often low-status groups in
multicultural societies; see Cronk 2004 and Neill 2011), parents may bias investment into
daughters, for example. In a variety of lowland South American cultures, the concept of partible
paternity, the assignment of fatherhood to more than one man, arises (Beckerman and Valentine
2002). Perhaps the variation in cultural beliefs and conditions relates to how parental-effort
trade-offs are solved.
• Beckerman, S., and P. Valentine, eds. 2002. Cultures of multiple fathers: The theory and
practice of partible paternity in lowland South America. Gainesville: Univ. Press of
Florida.
In a number of South American traditional societies, there exist a variety of cultural

beliefs about paternity arising not from one man’s sperm and one woman’s egg, but
rather contributions from several men to one woman’s egg. The distribution of these
beliefs suggests that they are culturally labile. In some of these societies, children deemed
to have multiple fathers are at an advantage.
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• Cronk, L. 2004. From Mukogodo to Maasai: Ethnicity and cultural change in Kenya.
Westview Case Studies in Anthropology. Boulder, CO: Westview.
Parental-effort biases can be optimization strategies. Cronk found that low-status

Mukogodo co-residing with Maasai neighbors biased their parental investment toward
daughters (e.g., better weight for height): a daughter might be able to “marry up” to a
Maasai man. The result of such hypergyny yielded better-invested grandchildren, and
cattle as brideprice for the bride’s family.
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• Hagen, E. H., H. C. Barrett, and M. E. Price. 2006. Do human parents face a quantity-
quality tradeoff? Evidence from a Shuar community. American Journal of Physical
Anthropology 130.3: 405–418.
DOI: 10.1002/ajpa.20272Save Citation »Export Citation »E-mail Citation »
Charles Darwin suggested that parents face a quantity-quality trade-off. However, direct
evidence is largely lacking. These authors test the relationship in a Shuar community in
Ecuador, using anthropometric data and the household ratio of consumers to producers
(CP). There was a clear negative relationship: children in smaller families received more
investment.
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• Hrdy, S. B. 1999. Mother Nature: A history of mothers, infants, and natural selection.
New York: Pantheon.
A classic: a thorough and accessible treatment of selection on maternal strategies,

allomothering, and more. Hrdy debunks several popular myths (e.g., pregnancy as a rosy
interlude). She covers not only maternity but allomothering, and she treats the interests of
(and conflicts of interest among) mother, fetus, and more.
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• Lawson, D. W., and R. Mace. 2009. Trade-offs in modern parenting: A longitudinal study
of sibling competition for parental care. Evolution and Human Behavior 30.3: 170–183.

Citation »
Because many forms of parental effort are true parental investment—what is given to one
offspring cannot be also given to another—siblings are in competition for finite parental
resources. The form such competition takes will vary across the life span.
Find this resource:
• Lawson, D. W., and R. Mace. 2011. Parental investment and the optimization of human
family size. Philosophical Transactions of the Royal Society B: Biological Sciences
366.1563: 333–343.
DOI: 10.1098/rstb.2010.0297Save Citation »Export Citation »E-mail Citation »
Ideas of quantity-quality trade-offs and optimal family size in humans have been
subjected to careful testing only since the late 20th century; most have relied on
correlational data. The authors find that a range of studies confirm that parents balance
family size against offspring success. Evidence of optimization is less clear.
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• Low, B. S. 1978. Environmental uncertainty and the parental strategies of marsupials and
placentals. American Naturalist 112.983: 197–213.
Draws the distinction between true parental investment (Trivers 1972), a zero-sum
allocation (what is invested on one offspring is not available for others), and more
generalizable allocation that can profit multiple offspring, perhaps related to R. A.
Fisher’s undefined concept of parental expenditure (see Fisher 1958, cited under Mating
Effort, Mate Choice, and Resource versus Reproductive Value).
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• Neill, D. B. 2011. Urbanization and daughter-biased parental investment in Fiji. Human

Nature 22.1: 139–155.
This paper documents parental-effort biases in a modern society. Even in the early 21st
century, parental effort and investment biases that result in greater net reproductive
success are not simply cultural relicts of a distant past or oddities of modern societies but
may reflect actual selective advantages.
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• Trivers, R. L. 1972. Parental investment and sexual selection. In Sexual selection and the
descent of man, 1871–1971. Edited by B. Campbell, 136–179. Chicago: Aldine de
Gruyter.
This classic paper defines parental investment as care given to one offspring that
otherwise could be devoted to different offspring, thus defining important trade-offs in
parental care.
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Senescence
Senescence, the failure of body systems to repair or regenerate, is relatively rare in wild
populations of nonhuman organisms. Early explanations (e.g., Comfort 1956) were not general.
They involved naive group selection (e.g., death mechanisms to clear the way for new
individuals) or selective irrelevance and failed to predict the timing of onset of senescence as
well as rate differences across species. The classical theory (Williams 1957, Hamilton 1966)
helps explain the general process. Natural selection ordinarily should work to delay or eliminate
senescence. Selection “nibbles” away at the strongest deleterious impacts, beginning at peak
reproductive value. But it is difficult for selection to discriminate between an allele that greatly
improves fitness in an organism’s early years (yet imposes deleterious effects later in life) and an
allele that has only good early effects—so deleterious effects accumulate toward the end of life.
Interestingly, the pattern of senescence and death in humans may not have changed greatly since
hunter-gatherer societies (Gurven and Kaplan 2007). More recently, Baudisch and Vaupel 2012
have suggested that broader consideration of trade-offs is important in understanding senescence.
Senescence is of particular interest in humans because, for example, human female reproductive
capacity senesces much earlier than other systems (menopause). Similar loss of fertility by older
females is seen only in a few other species, such as elephants, horses, and odontocete whales. As
modern women delay childbearing, some experience difficulty in carrying out a successful
pregnancy, so our interest is not idle.
• Baudisch, A., and J. W. Vaupel. 2012. Getting to the root of aging. Science 338.6107:
618–619.
DOI: 10.1126/science.1226467Save Citation »Export Citation »E-mail Citation »
Following Hamilton 1966 and Williams 1957 on the relative potency of selection at
various life history stages, organisms with fast life histories should show relatively early
senescence. This challenges that argument. Given finite resources for competing needs,
an organism makes difficult choices at every moment of life. Important comparative
analyses are suggested.
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• Comfort, A. 1956. Ageing: The biology of senescence. New York: Holt, Rinehart &
Winston.
This older work exemplifies earlier “explanations” of senescence: wearing out, toxin
accumulation, making way for younger individuals. These early theories failed to explain
why senescence began at different ages, and why it followed different paces in different
species, for example. Third edition published as recently as 1979 (New York: Elsevier).
Find this resource:
• Gurven, M., and H. Kaplan. 2007. Hunter-gatherer longevity: Cross-cultural perspectives.

Population and Development Review 33.2: 321–365.
DOI: 10.1111/j.1728-4457.2007.00171.xSave Citation »Export Citation »E-mail Citation

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This examines survivorship cross-culturally: post-reproductive longevity is a robust

feature of humans—two-thirds of mature individuals survive to grandparenthood. Until
about age forty, mortality rates increased by only about 1 percent per year; after forty,
mortality rates doubled in six to nine years. Even in hunter-gatherers, the modal age at
death is seventy-two (range sixty-eight to seventy-eight) years.
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• Hamilton, W. D. 1966. The moulding of senescence by natural selection. Journal of

Theoretical Biology 12.1: 12–45.
Furthering George Williams’s analysis (Williams 1957), Hamilton noted that any allele
with age-specific impacts is pleiotropic; thus, even if alleles with positive early-life
effects had late-in-life deleterious effects, they would be favored: natural selection could
not “discriminate” because alleles that affected performance late in life (1) affected fewer
individuals and (2) affected far less of those individuals’ reproduction. He noted that
natural selection should “nibble” away at deleterious impacts most strongly around the
age of first reproduction.
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• Williams, G. C. 1957. Pleiotropy, natural selection, and the evolution of senescence.

Evolution 11.4: 398–411.
Williams, in this classic early analysis, recognized that alleles affecting individuals at
different points in their life history would be under differing selective pressures. He
found existing literature on senescence inadequate, in that it failed to predict age at onset
and pace; he derived testable hypotheses for the evolution of senescence patterns and
moved the theory forward significantly.
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Inclusive Fitness, Cooperative Breeding, and Grandparental

Investment
Hamilton 1964, a seminal paper, made the point that in terms of genes passed on to future
generations, it paid individuals to adjust the help or harm they offered others in response to the
likely proportion of shared genes. William Hamilton’s work was occasionally misconstrued,
leading Richard Dawkins (Dawkins 1979) to write about such misconceptions. Degree of
relatedness, as well as reciprocity, matters greatly in raising children. Human infants are
expensive, and humans differ from their closest kin in degree and kind of parental and nepotistic
care. In other hominids, females are almost entirely single parents. In human societies, others
besides the mother may contribute food, care, protection, teaching, and more, to a degree not
seen in other hominids. Sometimes even nonrelatives such as wet nurses have provided such care
(Hrdy 1992). Such “cooperative breeding” means that human infants can be heavier at birth than
one would predict after a gestation period that is just what one would predict, and can be weaned
earlier and lighter than expected, giving humans a demographic advantage (Kramer 2010).
Different outcomes in Scelza and Silk 2014 and Hrdy 1992 suggest interesting comparative work
to be done. A special case of cooperative breeding, grandparental investment (Hawkes, et al.
1998), has been proposed as an influence on the evolution of menopause, although this remains a
controversy in the field (Strassmann and Garrard 2011).
• Dawkins, R. 1979. Twelve misunderstandings of kin selection. Zeitschrift für

Tierpsychologie 51.2: 184–200.
Many misconceptions arose from Hamilton 1964, a classic paper; it was because
Hamilton used honeybees as an example (and sisters are on average 75 percent alike
genetically rather than 50 percent, as in mammals). Dawkins created this useful piece to
clarify such points relevant to kin selection. It is important for students, but it remains
relevant to some modern research.
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• Hamilton, W. D. 1964. The genetical evolution of social behaviour I, II. Journal of

Theoretical Biology 7.1: 1–52.
This seminar paper introduced the concept of kin selection: behaviors that would be
simple competition when directed at a competitor would be spite if directed toward
relatives; conversely, help that would be genetically costly altruism when directed to a
stranger would be profitable when extended to a relative. The degree of help or harm
varies with degree of relatedness: behavior should evolve when br – c > 0; that is, when
the benefit to the recipient (b) times its degree of relatedness (r) to the doer, minus the
cost (c) to the doer is greater than zero (genetically profitable). A huge literature has
followed.
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• Hawkes, K., J. F. O’Connell, N. G. Blurton Jones, H. Alvarez, and E. L. Charnov. 1998.

Grandmothering, menopause, and the evolution of human life histories. Proceedings of
the National Academy of Sciences of the United States of America 95.3: 1336–1339.
DOI: 10.1073/pnas.95.3.1336Save Citation »Export Citation »E-mail Citation »
Curiously, human female reproductive function senesces long before other systems
degenerate. This fact has sparked considerable work; empirical studies (to date) have
mixed results. Grandparents (especially grandmothers, and particularly maternal
grandmothers) may enhance lineage success, but there are many confounding variables,
so no clear consensus yet exists.
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• Hrdy, S. B. 1992. Fitness tradeoffs in the history and evolution of delegated mothering
with special reference to wet-nursing, abandonment, and infanticide. Ethology and
Sociobiology 13.5–6: 409–442.
DOI: 10.1016/0162-3095(92)90011-RSave Citation »Export Citation »E-mail Citation »
Hrdy reviewed delegated mothering in evolutionary context, analyzing allocare in 18th-

century Paris. Wealthy women could afford wet nurses; they had short interbirth intervals
(IBIs) and low infant mortality (but with health problems). Poor women had long IBIs,
low fertility, and high infant mortality, but wet nurses fared worst of all.
Find this resource:
• Kramer, K. L. 2010. Cooperative breeding and its significance to the demographic

success of humans. Annual Review of Anthropology 39:417–436.
DOI: 10.1146/annurev.anthro.012809.105054Save Citation »Export Citation »E-mail

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Cooperative breeding may strongly influence life histories, both in terms of delayed
maturation or breeding and in terms of shortened IBIs and heightened fertility of breeding
individuals. This paper pulls together data on the helping roles and resulting impacts of
different categories of individuals in a number of societies.
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• Scelza, B. A., and J. B. Silk. 2014. Fosterage as a system of dispersed cooperative

breeding: Evidence from the Himba. Human Nature 25.4: 448–464.
Most studies of cooperative breeding focus on supplemental help from others. This paper
proposes that fostering can operate as a form of dispersed cooperative breeding,
enhancing women’s ability to concentrate their investment on a subset of their children.
Find this resource:
• Strassmann, B. I., and W. M. Garrard. 2011. Alternatives to the grandmother hypothesis:

A meta-analysis of the association between grandparental and grandchild survival in
patrilineal populations. Human Nature 22.1–2: 201–222.
This meta-analysis of seventeen studies of cooperative breeding tests predictions from

five hypotheses: confidence of paternity, grandmothering, helpful kin proximity,
senescence of grandparents, and local resource competition. Results favored the last
hypothesis (local resource competition), but there are confounding variables such as
marital residence and wealth.
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Inheritance
In other hominids, a parent’s status may influence the success of offspring. In humans, not only
parental status, but material goods, may help. Thus, inheritance is a special form of parental
investment. Parents in essentially all societies try to skew resources to favor their own children
(Borgerhoff Mulder, et al. 2009; Aldenderfer 2010). In polygynous societies, inheritance tends to
be preferentially directed toward sons. One study (on 19th-century Sweden) appeared to be an
exception in which parents did not favor their own offspring. Further analysis, however, showed
that within-family favoritism, especially in inheritance of land, obscured the pattern: oldest living
adult sons inherited (see chapter 8, “Sex, Resources, and Fertility in Transition,” in Low 2015,
cited under General Overviews, pp. 116–134). Hooper, et al. 2015 integrates inclusive-fitness
theory. Thus, in all societies examined so far, parents attempt to skew resources in favor of their
own offspring. This bias is reflected in the fact that stepparents tend to invest less in their
stepchildren than in their own (Anderson, et al. 2007) and sometimes even constitute a danger to
their stepchildren (see Daly and Wilson 1988, cited under Conflict and Competition).
• Aldenderfer, M., ed. 2010. Special section: Intergenerational wealth transmission and
inequality in premodern societies. Current Anthropology 51.1.
Devoted to specific papers on societies that flow from the summary in Borgerhoff
Mulder, et al. 2009, focusing on societies using different resource bases. Papers report
inheritance, and the inequality that follows in so many cases, in hunter-gatherers,
pastoralists, horticulturalists, and agrarian societies. Commentary and replies are
included.
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• Anderson, K. G., H. Kaplan, and J. B. Lancaster. 2007. Confidence of paternity, divorce,

and investment in children by Albuquerque men. Evolution and Human Behavior 28.1:
1–10.

Citation »
This is an excellent, theoretically driven empirical paper examining the role of genetic
relatedness in terms of paternal investment, from educational finding to inheritance.
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• Borgerhoff Mulder, M., S. Bowles, T. Hertz, et al. 2009. The intergenerational

transmission of wealth and the dynamics of wealth in pre-modern societies. Science
326.5953: 682–688.
DOI: 10.1126/science.1178336Save Citation »Export Citation »E-mail Citation »
This paper examines how parents strive to skew resource distribution in favor of their
own children in a wide array of societies, from hunter-gatherers and agricultural societies
to postindustrial nations; the authors used the same analytics for all. The results are
powerful in the uniformity of parents’ attention to giving their offspring every advantage,
in a variety of ecologically appropriate ways.
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• Hooper, P. L., M. Gurven, J. Winking, and H. S. Kaplan. 2015. Inclusive fitness and
differential productivity across the life course determine intergenerational transfers in a
small-scale human society. Proceedings of the Royal Society B: Biological Sciences
282.1803: 20142808.
This paper analyzes intergenerational transfers, integrating inclusive-fitness theory. The

authors predict that patterns of intergenerational transfers will track the interactions of
biological kinship with age-specific resource production.
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Cooperation
Cooperation has long been a puzzle. Clearly, no organism should evolve to cooperate in ways
that are genetically costly, yet often fitness costs and benefits are not readily apparent. Further, a
number of major papers have redefined terms in confusing ways, mostly in the use of the terms
“altruistic” and “altruism.” The term “altruism” is properly restricted to behavior that produces a
net genetic loss of fitness for the actor. Helping kin and helping reciprocators are relatively easy
to understand; the possibility for cooperation among kin is obvious, and both direct and indirect
reciprocity appears to be important in the evolution of cooperation. West, et al. 2011 argues that,
for example, Trivers 1971 and other major papers such as Gintis, et al. 2005 use some terms
wrongly or confusingly. Humans do cooperate, sometimes at large scales, and this complicates
our understanding because it is clear that the more competition within a group, the more difficult
it is for members of that group to compete against other groups (Barker, et al. 2012). Strategies
to reduce one’s own cost do not disappear, and within-group versus between-group payoffs must
be balanced. Humans are probably prosocial—predisposed to cooperate (e.g., Gintis, et al. 2005),
probably as a result of the selective pressures of other human groups. Certainly humans behave
so as to generate norms and punish defectors (Kurzban, et al. 2015). We even cooperate in
setting up large-scale rules—the law (Jones and Goldsmith 2005), and across quite diverse
societies we display quite uniform concepts of justice (Robinson, et al. 2007). Here is a dynamic
area of inquiry in which clear definitions and inference are needed.
• Barker, J. L., P. Barclay, and H. K. Reeve. 2012. Within-group competition reduces

cooperation and payoffs in human groups. Behavioral Ecology 23.4: 735–741.
DOI: 10.1093/beheco/ars020Save Citation »Export Citation »E-mail Citation »
When individuals within a group cooperate to produce excess resources, these may be
monopolized by competitive individuals—but most studies focus on noncontestable
public goods. Using game theory and empirical data, this study finds that within-group
competition decreases cooperation and reduces individual payoffs.
Find this resource:
• Gintis, H., S. Bowles, R. Boyd, and E. Fehr, eds. 2005. Moral sentiments and material
interests: On the foundations of cooperation in economic life. Economic Learning and
Social Evolution 6. Cambridge, MA: MIT.
This is an interesting read, and an important stride in the development of our

understanding of cooperation. A caveat is that in using a social science perspective to
understand cooperation from human morality (and the psychology behind it) to the
emergence of institutions, some misconceptions exist (laid out in West, et al. 2011).
Find this resource:
• Jones, O. D., and T. H. Goldsmith. 2005. Law and behavioral biology. Columbia Law
Review 105.2: 405–502.
A pioneering evolutionary approach to the law, examining the selective significance of a

group’s agreed-upon rules. A core argument is that law uses legal rules to change
behavior, and that changing behavior effectively (and efficiently) requires sound models
of human behavior—and that such models require an evolutionary perspective. Available
to read online for free via registration.
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• Kurzban, R., M. N. Burton-Chellew, and S. A. West. 2015. The evolution of altruism in
humans. Annual Review of Psychology 66:575–599.
DOI: 10.1146/annurev-psych-010814-015355Save Citation »Export Citation »E-mail

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This is an analysis of the evolution of fitness-costly behaviors, testing several alternative

hypotheses. An excellent guide to new methodologies and a review of still-problematic
areas, it emphasizes kinship, reciprocity, indirect reciprocity, punishment, and morality.
Find this resource:
• Robinson, P. H., R. Kurzban, and O. D. Jones. 2007. The origins of shared intuitions of
justice. Vanderbilt Law Review 60.6: 1633–1688.
This explores why intuitions of justice (or fairness) appear almost invariant across quite
different societies. There is consistency across animal studies, brain studies, and patterns
of moral development. An important constant is that we focus on getting our fair share,
not necessarily ensuring we give enough to help others.
Find this resource:
• Trivers, R. L. 1971. The evolution of reciprocal altruism. Quarterly Review of Biology

46.1: 35–57.
This is the classic paper on reciprocity. As West, et al. 2011 notes, however, there is a
confusing misuse of the term “altruism” common in much literature on cooperation:
using the term simply to imply short-term phenotypic costs. The term “altruism” today
specifies a loss of genetic fitness. Available to read online for free via registration.
Find this resource:
• West, S. A., C. El Mouden, and A. Gardner. 2011. Sixteen common misconceptions

about the evolution of cooperation in humans. Evolution and Human Behavior 32.4: 231–
262.

Citation »
This paper is essential in assessing the wealth of literature on cooperation and “altruism.”
In studies of cooperation, definitions have frequently been developed independently,
resulting in confusion. The authors identify misconceptions (e.g., “altruism” as all sorts
of behaviors, some of which are genetically profitable) with examples.
Find this resource:
Conflict and Competition

Conflict is rife at many levels, from parent-offspring (Trivers 1974; see also Hrdy 1999 in
Parental Effort) to allelic battles of mother’s and father’s genes within the fetus, to family
members and other within-group competitors, to reproductive competitors (Daly and Wilson
1988), to informal intergroup competition, to war between nation-states. Haig 2004 (and
citations therein) develops the concept of genomic conflict: paternal and maternal alleles in a
fetus have different interests and are thus in conflict. Individuals within families also have
conflicting interests (e.g., Mace 2013); in fact, Charles Darwin noted that one’s closest relatives,
because they were more similar than nonrelatives, could be a fierce source of competition. The
term “mating effort” really concerns conflicting goals of competitors within a population, the
escalation of which may lead to homicide (Daly and Wilson 1988). Free riders within a group are
also liable to be punished (Henrich and Boyd 2001). Of course, intergroup conflict is a major
phenomenon. Finally, much cooperation exists among members within groups to facilitate
intergroup conflict. There is evidence that humans have, over time, developed a variety of
strategies that result in lowering death rates from intergroup conflicts (Pinker 2011).
• Daly, M., and M. Wilson. 1988. Homicide. Foundations of Human Behavior. New York:
Transaction.
This classic evolutionary approach to the study of killing one another ranges from
traditional to modern societies. Homicide frequencies vary with the demographics of the
perpetrator: highest for young adult males, usually due to sexual selection. Penalties for
homicide also may vary with the victim’s demographics. Nongenetic children (e.g.,
stepchildren) also are at risk not only for reduced investment, but even homicide.
Find this resource:
• Haig, D. 2004. Genomic imprinting and kinship: How good is the evidence? Annual
Review of Genetics 38:553–585.
DOI: 10.1146/annurev.genet.37.110801.142741Save Citation »Export Citation »E-mail

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This is an excellent summary of Haig’s theory of genetic imprinting, introduced in 1992:

parent-specific gene expression at a locus evolves because a gene’s level of expression in
one individual has fitness impacts on other individuals. At any locus in a fetus, paternally
derived alleles have “interests” (maximizing what the fetus gets from the mother)
different from maternally derived alleles (which trade off maternal investment and
maternal condition).
Find this resource:
• Henrich, J., and R. Boyd. 2001. Why people punish defectors: Weak conformist
transmission can stabilize costly enforcement of norms in cooperative dilemmas. Journal
of Theoretical Biology 208.1: 79–89.
DOI: 10.1006/jtbi.2000.2202Save Citation »Export Citation »E-mail Citation »
This suggests that norms of punishment can establish cooperation. However, because
different selective forces can be involved in the establishment and maintenance of a trait,
West, et al. 2011 (cited under Cooperation) argues that it can be difficult for cooperation
to spread initially (because defecting in the early stages is an evolutionarily stable
strategy [ESS]). An area of intense interest.
Find this resource:
• Mace, R. 2013. Cooperation and conflict between women in the family. Evolutionary
Anthropology: Issues, News, and Reviews 22.5: 251–258.
This examines female conflicts of interest within families, and it models situations to
predict; for example, when cooperation among co-wives is reproductively profitable even
if wives’ “shares” are not equal.
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• Pinker, S. 2011. The better angels of our nature: Why violence has declined. New York:
Viking.
This accessible work reiterates and extends Daly and Wilson’s points about who kills
whom in traditional societies. Comparing traditional and modern societies suggests that
the development of large heterogeneous societies leads to imposition of larger-scale
formal rules and institutions; this in turn leads to decreased interpersonal violence and
lower death rates.
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• Trivers, R. L. 1974. Parent-offspring conflict. American Zoologist 14.1: 249–264.
DOI: 10.1093/icb/14.1.249Save Citation »Export Citation »E-mail Citation »

This is the seminal original description and explanation of the fact that an offspring’s
interests do not align fully with the mother’s interests, leading, for example, to
phenomena such as offspring (in normally silent species) screaming during the weaning
process.
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Cultural Transmission (Dual-Inheritance Theory)

Although humans represent an extreme in cultural transmission, it occurs in many species, from
the learning of song in some birds, to the optimality of foraging in ground squirrels, to tool use in
dolphins. The idea that ideas and behaviors could be culturally transmitted was introduced in
Dawkins 1976: the meme, the cultural homologue of the gene, could be copied perfectly or with
what amount to transcription errors. In humans, cultural transmission is particularly important,
often in ways that affect fitness (Henrich and McElreath 2003; Henrich, et al. 2008; Boyd, et al.
2011). Cultural-transmission theory integrates the problems of cooperation and conflict.
Although cultural transmission exists in other species, its influence in humans may be reaching
novel levels (e.g., social media, Google as a primary source of information). Cultural
transmission exhibits the three traits required for evolution by natural selection: variation,
heritability, and fitness effects. But one cannot assume that mechanisms and patterns are simply
parallel to genetic evolution (Henrich, et al. 2008). It differs from genetic transmission in
important ways: genetic transmission can only be downward, from parents to children. Cultural
transmission can be down, up (children to parents), oblique (unrelated adults such as teachers to
children; children to adults), and horizontal (among members of a cohort). Cultural transmission
is more malleable and can change more rapidly than biological transmission; innovations can
spread quickly. Cultural transmission of norms certainly can affect life history patterns (Huber,
et al. 2010). An accessible summary, useful for classes, is Prinz 2011.
• Boyd, R., P. J. Richerson, and J. Henrich. 2011. The cultural niche: Why social learning
is essential for human adaptation. Proceedings of the National Academy of Sciences of
the United States of America 108.S2: 10918–10925.
DOI: 10.1073/pnas.1100290108Save Citation »Export Citation »E-mail Citation »
While humans have more “computing power” than other animals, no individual is smart
enough to master all the information necessary to survive in any single habitat. Even in
the simplest societies, cultural transmission is essential for human success.
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• Dawkins, Richard. 1976. The selfish gene. Oxford: Oxford Univ. Press.
Parallel to genes, which are strands of DNA that are copied from individual to individual,
memes are- culturally transmitted ideas or practices; Dawkins defined a meme as a
cultural unit of replication. Like genes, memes can be copied perfectly or imperfectly.
Dawkins’s definition fostered subsequent development in studies of cultural evolution.
The 30th-anniversary edition was published in 2006.
Find this resource:
• Henrich, J., R. Boyd, and P. J. Richerson. 2008. Five misunderstandings about cultural
evolution. Human Nature 19.2: 119–137.
The authors suggest that in early-21st-century debates about the characteristics and
importance of cultural evolution, widespread misconceptions exist. They discuss five: the
nature of mental representations, the requirement of replicators, whether only content-
dependent psychological biases matter, whether one can assess “cultural fitness” from
cultural transmission, and whether sources of variation must be random. They propose an
evolutionary science of culture.
Find this resource:
• Henrich, J., and R. McElreath. 2003. The evolution of cultural evolution. Evolutionary
Anthropology: Issues, News, and Reviews 12.3: 123–135.
Human behavior exhibits remarkable phenotypic plasticity; it is likely that basically the
same genetic endowment has given rise to, for example, subsistence strategies from
Arctic hunting to tropical horticulture to desert pastoralism. Ecologically and socially
appropriate strategies are transmitted by social learning; they have accumulated over
generations. The authors examine how (and when) culturally evolved adaptations arise.
Find this resource:
• Huber, S., F. L. Bookstein, and M. Fieder. 2010. Socioeconomic status, education, and
reproduction in modern women: An evolutionary perspective. American Journal of
Human Biology 22.5: 578–587.
Cultural transmission of women’s roles correlates, in many Western societies, with

education and income. For modern Western women, there can be a negative association
between their own status and realized fertility, as modern women delay their
reproduction. This paper examines the impacts of a woman’s own (versus her husband’s)
income and education (see also Extrinsic Mortality and Reproductive Timing).
Find this resource:

• Prinz, J. 2011. Culture and cognitive science. In The Stanford encyclopedia of
philosophy. Edited by E. N. Zalta. Stanford, CA: Stanford Univ.
This broad discussion of culture and cultural transmission is easily accessible and is a
good overview. It essentially summarizes the Boyd-Richerson arguments.
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Environmental Extremeness, Unpredictability, and Risk

Taking
Unpredictability of important environmental variables clearly raises an organism’s cost of
responding optimally. When rainfall, for example, fluctuates irregularly both in timing and
amount, an organism may have little reliable information about when to initiate important
activities. One of the earliest ecological analyses was by Robert Colwell (Colwell 1974), who
developed a homologue of species diversity indices: high species diversity generates
unpredictability in sampling. Colwell partitioned predictability into that arising from constancy
(low variance) and that due to contingency (covariance with another factor; e.g., seasonality).
The concept of unpredictability has a checkered history; it has often been confused with
extremeness (“harshness”), although the selective pressures are quite different (Ellis, et al. 2009)
and may interact. When an important resource is rare, with predictable timing (e.g., very harsh
winters), contingent response strategies are not difficult to find. On the other hand,
unpredictability (which can be spatial, temporal, or both) raises an organism’s costs.
Unpredictability can be analyzed with Fourier analyses (temporal) and Colwell’s predictability
measures. Human strategies in the face of environmental uncertainties typically aim at risk
reduction, both in terms of strategies such as storage and reciprocity (Cashdan 1990;
Winterhalder, et al. 1999; Bliege Bird, et al. 2002) and in personal life history events (Hill, et al.
1997; Leslie and Winterhalder 2002; Quinlan 2006). Cashdan and Downes 2012 contains a series
of relevant papers.
• Bliege Bird, R., D. W. Bird, E. A. Smith, and G. C. Kushnick. 2002. Risk and reciprocity
in Meriam food sharing. Evolution and Human Behavior 23.4: 297–321.
DOI: 10.1016/S1090-5138(02)00098-3Save Citation »Export Citation »E-mail Citation »
The authors examine the “risk reduction reciprocity” hypothesis: reciprocal sharing can
reduce the risk of failure. In Meriam foragers, high-risk hunting kills were shared more
than low-risk kills from the same foraging patch, but risk alone was not a good predictor
of sharing.
Find this resource:

• Cashdan, E., ed. 1990. Risk and uncertainty in tribal and peasant economies. Boulder,
CO: Westview.
This collection contains some important early papers; although it can be difficult to find,
not having been reprinted, it clearly sparked further research on risk and unpredictability
at a number of levels.
Find this resource:
• Cashdan, Elizabeth, and Stephen M. Downes, eds. 2012. Special issue: Evolution of
human aggression. Human Nature 24.1.
This collection contains papers on coping with environmental risk and uncertainty at
individual and group levels. It considers pathogen stress, social-network predictors of
perceived stress, violence, father absence, and more.
Find this resource:
• Colwell, R. K. 1974. Predictability, constancy, and contingency of periodic phenomena.

Ecology 55.5: 1148–1153.
This classic adaptation of Shannon-Weiner diversity index calculations (higher diversity

= lower sampling predictability) partitions the predictability of any phenomenon into that
due to constancy (no information is gained about better-worse timing of activities) and
contingency (degree of co-occurrence with some other variable; e.g., diurnal or seasonal
rainfall patterns).
Find this resource:
• Ellis, B. J., A. J. Figueredo, B. H. Brumbach, and G. L. Schlomer. 2009. Fundamental

dimensions of environmental risk: The impact of harsh versus unpredictable
environments on the evolution and development of life history strategies. Human Nature
20.2: 204–268.
Argues that harshness and unpredictability are major determinants of life history patterns
for humans, reprising some chapters of Cashdan 1990. Impacts will depend on population
densities, levels of intraspecific competition, resource scarcity, and the extent to which
environmental fluctuations fluctuate over short versus long timescales.
Find this resource:
• Hill, E. M., L. T. Ross, and B. S. Low. 1997. The role of future unpredictability in human
risk-taking. Human Nature 8.4: 287–325.
DOI: 10.1007/BF02913037Save Citation »Export Citation »E-mail Citation »
This paper examines, among late-20th-century community college students, individuals’

propensity to take risks as a function of their perceived future uncertainties.
Unpredictability beliefs remained a strong predictor after accounting for standard
predictors such as sex and temperament.
Find this resource:
• Leslie, P., and B. Winterhalder. 2002. Demographic consequences of unpredictability in

fertility outcomes. American Journal of Human Biology 14.2: 168–183.
Both unpredictability (sensu Colwell’s work) and variance (even if predictably timed)
that affect survivorship also will affect fertility patterns. Risk-sensitive (i.e., variance-
sensitive) adaptive behaviors produce significantly different family sizes compared to
those under low-variance and predictable conditions. This connects specific social and
economic conditions directly to birth spacing and completed fertility, with long-term
implications.
Find this resource:
• Quinlan, R. J. 2006. Human parental effort and environmental risk. Proceedings of the
Royal Society B: Biological Sciences 274.1606: 121–125.
Quinlan examined measures of parental effort cross-culturally as parents responded both

to extremeness and unpredictability. Maternal care was negatively associated with
warfare and with famine; it showed a quadratic relationship to pathogen intensity.
Paternal investment showed an inverse relationship with pathogen intensity.
Find this resource:
• Winterhalder, B., F. Lu, and B. Tucker. 1999. Risk-sensitive adaptive tactics: Models and
evidence from subsistence studies in biology and anthropology. Journal of
Archaeological Research 7.4: 301–348.

Risk sensitivity in foraging is important when outcomes are uncertain, and the
consequences for fitness are nonlinear. Here, a general model of risk sensitivity, and a
review of risk-sensitive adaptive tactics both in nonhumans and traditional human
societies.
Find this resource:
How Modern Environments Shape (and Reshape) Human

Life History and Behavior
Modern environments are, in selective terms, novel in many ways. Some selective forces are
almost certainly similar across populations, locations, and time: the centrality of family (e.g., kin
selection) and friends (reciprocity), and the influence of life expectancy on reproductive patterns.
Other forces are changing dramatically as humans alter the environment. Teasing apart these
categories represents an important contribution that behavioral ecologists could make.
Points of View
Behavioral ecologists and evolutionary anthropologists tend to focus on “what behavior is

adaptive?” in particular circumstances, seeking to match particular behaviors to specific
environmental conditions. In contrast, many evolutionary psychologists (Barkow, et al. 1992) are
concerned with defining what constitutes an “adaptation.” Typically, they restrict the term
“adaptation” to mean specific genes selected for in the “environment of evolutionary
adaptedness” (EEA), usually assumed to be hunting and gathering in the Pleistocene. Thus,
running away from a fast-approaching truck can never be an adaptation (because there were no
trucks in the Pleistocene), although it may be adaptive in the behavioral ecologist’s sense of
improving fitness. Behavioral ecologists and evolutionary anthropologists (e.g., Irons 1998) are
likely to argue that environments were hardly uniform around the world in the Pleistocene. As a
result, some traits (e.g., valuing kin) may have remained the same across many environments,
while others will have varied in specifics: perhaps the mental rule is not “run away from a tiger”
but “run away from anything large that is approaching fast.” Those apparently small differences
assume greater importance when studying modern populations. Humans, with their considerable
phenotypic plasticity and cultural transmission, may well show adaptive responses whether we
can define particular genes for specific behaviors or not. The result has been a lively debate.
• Barkow, J. H., L. Cosmides, and J. Tooby, eds. 1992. The adapted mind: Evolutionary
psychology and the generation of culture. Oxford: Oxford Univ. Press.
This is the classic paper restricting the meaning of “adaptation” to behaviors responding
in a domain-specific way to Pleistocene conditions. In the early 21st century it looks
somewhat problematic because we see adaptive genetic change occurring (see Selection
and Genetic Responses in Modern Populations).
Find this resource:
• Irons, W. 1998. Adaptively relevant environments versus the environment of

evolutionary adaptedness. Evolutionary Anthropology: Issues, News, and Reviews 6.6:
194–204.
DOI: 10.1002/(SICI)1520-6505(1998)6:6<194::AID-EVAN2>3.0.CO;2-BSave
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Irons coined the term “adaptively relevant environment” (ARE) to distinguish between
traits likely to be universal (e.g., valuing kin) and those for which specifics may shift
across environments (running away from tigers or from fast-approaching dangerous
things in general, including, for example, trucks). Behavioral ecologists, concerned with
whether a trait enhances or depresses fitness—and unable to specify particular causal
genes—frequently prefer ARE.
Find this resource:
Extrinsic Mortality and Reproductive Timing
Extrinsic mortality affects patterns of parental effort (e.g., when adults are safer than juveniles
and can transfer that safety) and patterns of fertility (e.g., age at first birth is reduced when life is
nasty, brutish, and short). It arises from conditions not related to an organism’s behavior; in
humans this can result in many factors. Thus in Wilson and Daly 1997, drive-by shootings were
the major source of extrinsic mortality. In fact, any intense mortality risk affects maturation and
reproduction, even today (e.g., Nettle 2010). Modern women’s lives, though variable, are mostly
so different from our ancestors’ that women’s health may be at risk (Jasienska 2013). High
extrinsic mortality favors a “fast” life history: early reproduction, and high fertility with
attendant lower per capita investment (Anderson 2010; Low, et al. 2008). To complicate matters
further, the important relevant conditions are in the early 21st century unlikely to be at
equilibrium (Low, et al. 2013).
• Anderson, K. G. 2010. Life expectancy and the timing of life history events in developing
countries. Human Nature 21.2: 103–123.
This study tests the extrinsic-mortality age at first-birth relationship by using two detailed
samples: cross-sectional analysis of sixty-two countries, and multiple cross-sectional data
from forty-eight countries. Interestingly, the predictions are not supported in sub-Saharan
Africa. Implications for understanding HIV are considered.
Find this resource:
• Jasienska, G. 2013. The fragile wisdom: An evolutionary view on women’s biology and
health. Cambridge, MA: Harvard Univ. Press.
Focusing on women’s life histories (and more), the author concentrates on the ways that
early-21st-century evolutionary novelty affects women’s life histories and health. For
example, selection has, in the past, favored alleles that enhance fertility, not those that
reduce disease risk (and especially not to protect from relatively new diseases).
Find this resource:
• Low, B. S., A. Hazel, N. Parker, and K. B. Welch. 2008. Influences on women’s

reproductive lives: Unexpected ecological underpinnings. Cross-Cultural Research 42.3:
201–219.
This paper explores life expectancy and age at first birth in more than 170 modern
societies. The relationship is nonlinear, and it is strongest in nations deemed to have high
human development scores (UN Development Programme, or UNDP) and weakest in
countries with low scores. Life expectancy at birth is also strongly correlated with age-
specific mortality: countries with lower life expectancies have earlier ages at first
reproduction, and higher fertility at all ages, than others.
Find this resource:
• Low, B. S., N. Parker, A. Hazel, and K. B. Welch. 2013. Life expectancy, fertility, and
women’s lives: A life-history perspective. Cross-Cultural Research 47.2: 198–225.
Life expectancy at birth changed from 1955 to 2005 in most societies. Patterns differ for
countries categorized as high, moderate, or low in the UNDP Human Development
Index. This highlights the difficulties arising from assumptions that underlie many
analyses: that any population will adequately represent the species, and that variables are
in equilibrium and not changing.
Find this resource:
• Nettle, D. 2010. Dying young and living fast: Variation in life history across English
neighborhoods. Behavioral Ecology 21.2: 387–395.
DOI: 10.1093/beheco/arp202Save Citation »Export Citation »E-mail Citation »
Nettle’s study finds that when extrinsic mortality is high, not only is age at first birth
shifted earlier, but breastfeeding, a major maternal investment, is decreased.
Find this resource:

• Wilson, M., and M. Daly. 1997. Life expectancy, economic inequality, homicide, and
reproductive timing in Chicago neighbourhoods. British Medical Journal 314.7089:
1271–1274.
DOI: 10.1136/bmj.314.7089.1271Save Citation »Export Citation »E-mail Citation »
This study examined the impact of extrinsic (usually violence-related) mortality and age
at first birth in three hundred Chicago neighborhoods. It is one of the first papers to test
the well-established generalization that high extrinsic mortality leads to early
reproduction, and relatively high fertility, in modern human populations.
Find this resource:
Selection and Genetic Responses in Modern Populations
Sometimes one can discover the actual genetics of a trait; since Crow 1958, a seminal work,
analyses continue to advance. We know that humans experienced genetic change over the last
few thousand years (Pritchard 2012). Today, there is a small but growing literature showing that
selection does operate in modern populations, and that genetic changes result (Losos, et al. 2013;
Milot, et al. 2011; Stearns, et al. 2010). Our modern responses to health risks are (perhaps
slowly) coming to deal with host behavior and biology (Nesse and Williams 1996),
distinguishing, for example, “symptoms” that are the result of host manipulation by a pathogen
(e.g., sneezing, coughing) from those that are in fact adaptive (e.g., fever). Thus, the argument
over whether there exist actual adaptations (genetic changes produced by selection) in modern
humans may be over. As genetic analyses continue to develop, we are able to see ongoing
genetic change as the result of selection.
• Crow, J. F. 1958. Some possibilities for measuring selection intensities in man. Human
Biology 30.1: 1–13.
This modestly titled piece is the foundation of all following papers. Amazingly insightful
and prescient.
Find this resource:
• Losos, J. B., D. A. Baum, D. J. Futuyma, et al. 2013. The Princeton guide to evolution.
Princeton Reference. Princeton, NJ: Princeton Univ. Press.
Section 8 (“Evolution and Modern Society”) discusses a variety of topics examining how
natural selection operates in modern societies.
Find this resource:

• Milot, E., F. M. Mayer, D. H. Nussey, M. Boisvert, F. Pelletier, and D. Réale. 2011.
Evidence for evolution in response to natural selection in a contemporary human
population. Proceedings of the National Academy of Sciences of the United States of
America 108.41: 17040–17045.
DOI: 10.1073/pnas.1104210108Save Citation »Export Citation »E-mail Citation »
This study examines variation in age at first birth (AFB) in a preindustrial French
Canadian population. AFB was highly heritable and correlated to fitness: earlier births
resulted in higher lineage success.
Find this resource:
• Nesse, R. M., and G. C. Williams. 1996. Why we get sick: The new science of Darwinian
medicine. New York: Vintage.
A groundbreaking book. It examines selective pressures exerted by pathogens, and host

responses. The authors also argue something not widely appreciated at the time: that
persistent use of antibiotics, in the long run, merely selects for antibiotic-resistant
pathogens.
Find this resource:
• Pritchard, J. K. 2012. How we are evolving. In Special issue: What makes us human.
Scientific American 22.1: 98–105.
This reprises and updates an earlier article in the magazine (Scientific American 303.4
[2010]: 40–47) by the same author, giving examples of human populations changing
genetically, in response to changing environmental conditions, in the past few thousand
years. Excellent for undergraduates.
Find this resource:
• Stearns, S. C., S. G. Byars, D. R. Govindaraju, and D. Ewbank. 2010. Measuring

selection in contemporary human populations. Nature Reviews Genetics 11.9: 611–622.
A relatively up-to-date review of studies that measure actual genetic changes, to estimate
the impact of selection on modern populations.
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Conservation: How Behavioral Ecology Predicts Patterns
Issues of sustainability and rapid environmental change are in the headlines in the early 21st
century. Humans, like all other organisms, evolved to get and use resources efficiently, and the
combination of our prosociality and high intelligence means that we are so efficient at
maximizing in the short term that sustainability may be elusive (e.g., Marean 2015). In small-
scale societies, hunting for optimal returns was always profitable (Hames 2007, Alvard 1994,
Smith and Wishnie 2000). In many cases, indigenous technology was relatively inefficient and
markets were largely lacking; widespread extirpation was rare. We have a long history of
directional selection: more has always been better in fitness terms. In the early 21st century,
human technology is so efficient at extracting resources that conservation is increasingly
important (Borgerhoff Mulder and Coppolillo 2005). Levi, et al. 2011 shows that “prey profiles”
(how many of which prey are taken by hunters) are not a good predictor of impacts on prey
populations for Neotropical hunters; integrating spatial information greatly improves prediction.
Primary and secondary impacts of our resource use have the potential to create ecological and
social problems, from the impacts of climate change on human habitation and supporting
resources such as food to intranecine pitched battles over controlling valuable and localized
resources such as diamonds, tin, cooper, and more. We are therefore interested in patterns of
resource competition and control, as well as any evidence of deliberate conservation. An
evolutionary approach helps us understand why convincing ourselves to “do the right thing” is
not easy, but some scholars are hopeful (e.g., Wilson 2002).
• Alvard, M. S. 1994. Conservation by native peoples: Prey choice in a depleted habitat.

Human Nature 5.2: 127–154.
DOI: 10.1007/BF02692158Save Citation »Export Citation »E-mail Citation »
Alvard explicitly tested the alternative hypotheses that Amazonian hunters were (1)
optimal foragers (even when optimality meant, for example, killing an individual from an
endangered species) or (2) conservators.
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• Borgerhoff Mulder, M., and P. Coppolillo. 2005. Conservation: Linking ecology,

economics, and culture. Princeton, NJ: Princeton Univ. Press.
This book is an evolutionarily informed approach to conservation, with broad, wide-

ranging coverage: from basic definitions to policy aspects, and from preindustrial
societies to modern issues. Relevant basic ecology and conservation biology are also set
into context.
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• Griskevicius, V., S. M. Cantú, and M. van Vugt. 2012. The evolutionary bases for
sustainable behavior: Implications for marketing, policy, and social entrepreneurship.
Journal of Public Policy & Marketing 31.1: 115–128.
DOI: 10.1509/jppm.11.040Save Citation »Export Citation »E-mail Citation »
This is one of the first application of human behavioral-ecological principles to

sustainable behavior in the context of marketing and policy. In their table 1, the authors
identify principles and then compare well-matched, and poorly matched, strategies.
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• Hames, R. 2007. The ecologically noble savage debate. Annual Review of Anthropology
36:177–190.
DOI: 10.1146/annurev.anthro.35.081705.123321Save Citation »Export Citation »E-mail

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Hames identifies two different research threads underlying the debate. The first is a
relatively classical analysis of the ethnography of resource use. The second is broader
(and more politically charged): definitions of “ecological nobility,” ideology, and the use
of the noble-savage concept as a political tool.
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• Levi, T., F. Lu, D. W. Yu, and M. Mangel. 2011. The behaviour and diet breadth of
central-place foragers: An application to human hunters and Neotropical game
management. Evolutionary Ecology Research 13.2: 171–185.
An important advance. When opportunity costs, spatial structure, and finite time horizons
are included in Neotropical optimal-foraging models, it is clear that using prey profiles
does not reflect impact on the prey population. Spatial information shows that prey
profiles do not change substantially even when high-value game species may be locally
extirpated.
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• Marean, C. W. 2015. The most invasive species of all. Scientific American 313.2: 32–39.
This is an easily accessible article, suitable for nonspecialists and undergraduate courses.
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• Smith, E. A., and M. Wishnie. 2000. Conservation and subsistence in small-scale
societies. Annual Review of Anthropology 29:493–524.
DOI: 10.1146/annurev.anthro.29.1.493Save Citation »Export Citation »E-mail Citation »
The authors review theory and data from multiple fields and assess predictions about
sustainable use of resources. They offer an excellent definition of conservation: to meet
the definition, “any action or practice must not only prevent or mitigate resource
overharvesting or environmental damage, it must also be designed to do so.”
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• Wilson, E. O. 2002. The future of life. New York: Knopf.
Within behavioral ecology, Wilson is best known for his 1975 classic Sociobiology: The
New Synthesis (Cambridge, MA: Belknap). More recently he has written broadly about
human nature, genetics, and culture. Here he lays out modern conservation problems and
suggests, more in hope than analysis, how to solve them.
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• Borgerhoff Mulder, M. 1991. Human behavioural ecology. In Behavioural ecology: An

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• Brown, G. R., and P. J. Richerson. 2014. Applying evolutionary theory to human

DOI: 10.1007/s10818-013-9166-4Save Citation »Export Citation »E-mail Citation »

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• Cronk, L. 1991. Human behavioral ecology. Annual Review of Anthropology 20:25–53.

DOI: 10.1146/annurev.an.20.100191.000325Save Citation »Export Citation »E-mail

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• Human Behavioral Ecology Bibliography (HBEB).

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This is a balanced treatment of evolutionary approaches to human behavior, and a good

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This assessment of the strengths of a behavioral-ecological approach emphasizes more-

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• Smith, E. A. 2000. Three styles in the evolutionary analysis of human behavior. In

Smith compares behavioral ecology, evolutionary psychology, and dual-inheritance

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• Winterhalder, B., and E. A. Smith. 2000. Analyzing adaptive strategies: Human

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This collection is a unified, coherent overview of advances in selectionist thinking from

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• American Journal of Human Biology.

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An interdisciplinary journal of original research, theoretical articles, and occasional

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• Behavioral Ecology and Sociobiology.

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• Evolution and Human Behavior.

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• Evolutionary Anthropology: Issues, News, and Reviews.

An authoritative review journal that focuses on issues of current interest in biological

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This specializes in evolutionary approaches to human behavior. It includes papers in

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• Proceedings of the Royal Society B: Biological Sciences.

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This text on evolutionary anthropology could easily be titled “Human behavioral

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