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Introduction
Compared with our closest living relativesthe extant great
apesthe sole surviving member of the genus Homo possesses
a suite of features that make us quite distinct, including unusually large brains, obligate bipedality, a reliance on the production and use of tools, and a strikingly different life history.
Generally, patterns of mammalian growth, development, and
life history are thought of as lying along a spectrum somewhere between two end points that are colloquially referred
to as live fast, die young and live slow, die old. Modern
human life history incorporates elements of both schedules:
long gestation periods, altricial offspring, enlarged brains,
slow maturation rates, increased life span, and protracted periods of offspring dependence are suggestive of a live slow
strategy, whereas relatively early weaning, short interbirth intervals, and the ability to overlap births (resulting in the presence of multiple offspring) are suggestive of a live fast schedule. Of interest to paleoanthropologists is whether this
modern human life history package evolved as a single developmental module or accumulated in a mosaic fashion
(with different attributes appearing at different points in
time). Furthermore, it is of great interest to understand when
Gary T. Schwartz is Associate Professor of Anthropology at the
Institute of Human Origins at Arizona State University (900 South
Cady Mall, Tempe, Arizona 85287-2402, U.S.A. [garys.iho@asu.edu]).
This paper was submitted 12 XII 11, accepted 28 VI 12, and
electronically published 20 XI 12.
this transition occurred and thus determine whether the human life history package appeared as part of a suite of fundamental adaptations at the base of the hominin clade;
whether it evolved somewhat later, perhaps tied to the reorganization of the cranium and postcranium that characterized the earliest members of the genus Homo; or whether
it appeared even later still, perhaps in the last hundred thousand years or so.
The last decade has been marked by a tremendous amount
of research into reconstructing the pattern of growth, development, and life history of extant great apes, australopiths,
and early to later members of the genus Homo. Novel analytical techniques, imaging modalities, and hard-fought observational data from naturalistic studies of great apes can
now be synthesized to paint a broad view of the evolution of
life history throughout the course of the human story. The
goal of this paper is to review the current state of knowledge
of the evolution of human life history within the comparative
context of what we know about these attributes in populations
of extant hominoids and fossil hominins. These data will be
evaluated within the light of what is known about primate
life history, ecology, diet, and so forth, and will be used to
help suggest future avenues of inquiry into studies of hominin
life history.
2012 by The Wenner-Gren Foundation for Anthropological Research. All rights reserved. 0011-3204/2012/53S6-0011$10.00. DOI: 10.1086/667591
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striations and Retzius lines in the enamel and the corresponding von Ebner and Andresen lines in dentine (Dean 1987,
2006; Schwartz and Dean 2000; Smith 2006, 2008). Careful
counts of these lines reveal the time taken to form a tooth,
including the tooth crown and however much root had
formed at the time of death, thereby providing a detailed
chronology of dental development that can, under the right
circumstances, also yield precise ages for key events such as
molar emergence (e.g., Beynon, Dean, and Reid 1991; Dean
et al. 2001; Dirks 1998; Kelley and Schwartz 2010; Smith,
Reid, and Sirianni 2006; Smith et al. 2010a, 2010b).
Data on molar emergence ages in hominins, while rare, are
becoming more accessible, especially with the application of
new, noninvasive imaging modalities that allow access to the
internal dental growth record (e.g., Smith et al. 2007b, 2007c,
2010b). While it is critically important to establish a growing
database on molar emergence ages in key hominin taxa, it is
also necessary to chart more fully dental developmental variation in extant hominoids and to interpret that variation in
light of a particular species population ecology, demography,
and life history. For now, good population data on molar
emergence ages exist only for P. troglodytes (Anemone, Mooney, and Siegel 1996; Conroy and Mahoney 1991; Kuykendall,
Mahoney, and Conroy 1992; Nissen and Reisen 1945, 1964;
Reid et al. 1998; Schultz 1940; Smith et al. 2007a, 2010a;
Zihlman, Bolter, and Boesch 2004; though see Dirks 2003 and
Dirks and Bowman 2007 for individual hylobatids; and Beynon, Dean, and Reid 1991; Winkler, Schwartz, and Swindler
1991; and Kelley and Schwartz 2010; and Willoughby 1978
for individual gorillas and orangutans). There is also a growing awareness that emergence ages for captive primate populations may be slightly advanced compared with wild populations (e.g., Kelley and Schwartz 2010; Smith and Boesch
2011; Zihlman, Bolter, and Boesch 2004), suggesting at the
very least some caution in relying on databases derived exclusively from captive colonies. Expanding our knowledge of
dental developmental variation across natural fertility populations of modern humans is another key element (e.g., Liversidge 2003) and will ultimately allow more fine-scale tests
of how population-level variation in aspects of dental development (e.g., M1 emergence age) relates to that for various
life history attributes.
Early Homo
It has generally been viewed that the origin of the genus Homo
was characterized by a trend away from bipedal apelike forms
to obligate terrestrial bipeds who were endowed with much
larger brains and the capacity to manufacture and use stone
tool technology and who also exhibited a shift in dietary and
foraging adaptations. Fossil representatives of the genus Homo
were first described by Leakey, Tobias, and Napier (1964) from
material derived from Bed I, Olduvai Gorge, Tanzania, and
were dated to 1.81.7 Ma. These authors viewed the material
as distinct from australopith material given its possession of
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Table 1. Comparative life history and M1 emergence age (years) in extant great apes and humans
Variable
Age at first reproduction
Interbirth interval
Survivorshipe
Age at M1 emergence
Cranial capacity (cm3)
Gorilla
Pan
Pongo
Homo
10.1a (51.3)
4.3
20.6f (38.1)
3.8 (65.5)
484
14.3b (72.6)
5.8b
29.7 (54.9)
4.0b (69.0)
383
15.7c (79.7)
6.9c
43.0 (79.5)
4.6c (79.3)
379
19.7
3.4d
54.1
5.8
1,293
Note. Numbers in parentheses indicate the percentage relative to the values in modern humans. References for life
history, survivorship, and M1 emergence data are reported in Kelley and Schwartz (2010).
a
Value for mountain gorilla Gorilla gorilla beringei, which is likely to be earlier than in Gorilla gorilla gorilla.
b
Values for Pan troglodytes verus only (Ta Forest, Ivory Coast).
c
Values for Pongo pygmaeus pygmaeus (i.e., orangutans from Borneo) only.
d
Interbirth interval is anomalously low in modern humans compared with other anthropoid species. This life
history variable is included for between-ape comparisons only, and percentages of human values were not calculated.
e
Expected age at death at age 15 years based on empirically derived survivorship curves.
f
Average of female (24.8 years, or 45.8%) and male (16.4 years, or 30.3%) values. Courtesy of Anne Bronikowski
and the Dian Fossey Gorilla Fund International.
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Table 2. Estimated ages (years) at death and M1 emergence in great apes, australopiths,
and Homo
Species (specimen)
Great apes:
Pongo pygmaeus pygmaeus
Gorilla gorilla beringei
Pan troglodytes verus
Australopiths:
Australopithecus afarensis (LH 2)
Australopithecus africanus (Sts 24)
A. africanus (Taung 1)
Paranthropus robustus (SK 62)
P. robustus (SK 63)
Paranthropus boisei (KNM-ER 1820)
Homo:
Homo erectus (Sangiran S7-37)
Homo ergaster (KNM-WT 15000)
Homo neanderthalensis (La Chaise)
Homo sapiens (Global)
Age at M1 emergence
4.6
3.8
4.0
3.25
3.30
3.733.93
3.353.48
3.154.23
2.53.1
8.38.8
2.9*
2.9*
3.33.5*
3.83.9*
2.93.2*
2.73.3*
4.4*
4.5*
6.7*
5.8 (4.77.1)
species groups). Taken together, they suggest a dental developmental profile that was more modern apelike than modern
humanlike, providing some tantalizing evidence that neither
species likely possessed an extended period of childhood dependence (Dean 1995; Dean et al. 2001).
Later species of Homo. Given the extraordinarily complete and
well-preserved juvenile specimen KNM-WT 15000, we perhaps know more about overall growth and development, and
thus life history, in African H. erectus than any other early
hominin taxon. Based on the lack of fusion of the distal elbow
joint and the acetabulum in particular, KNM-WT 15000 was
given an age of death at ca. 13 years, or the early part of the
adolescent stage of growth (i.e., postpubertal; Ruff and Walker
1993). Interestingly, the mostly completed dentition (26 permanent teeth had emerged, all but the M3s and maxillary
canines) suggested an age at death of 10.2 years. This discord
of almost 3 years suggests a somewhat unique developmental
trajectory for H. erectus (Smith 2004). Equally interesting is
that based on a chimpanzee developmental standard, the state
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Future Directions
Discoveries of new infant and subadult fossils along with
advances in noninvasive imaging and analytical methods are
providing opportunities to probe further the fossil record of
human growth and development. For instance, it is likely that
the age at which important life history events, such as weaning, will be assessable directly from the fossil record. The
timing of weaning is a key life event for both mother and
offspring, and analyses of life history variation as it relates to
weaning across Primates provide one example of the selective
basis of these sorts of developmental dissociations. Within
Malagasy prosimians, selection has acted to accelerate weaning
and dental development but has delayed the age at first reproduction (Godfrey et al. 2001; Richard et al. 2002; Schwartz
et al. 2002). It has been suggested that some australopiths
show rapid deciduous tooth wear, which was taken as evidence
suggestive of relatively early weaning (Aiello, Montgomery,
and Dean 1991; Dean 2006, 2010). It may now be possible
to retrieve direct evidence for reconstructing weaning age and
shifts in energy provisioning for offspring through the evolution of Homo. Across Primates, weaning is closely tied in
time to the emergence of M1. However, human life history
is characterized by relatively early weaning followed by a prolonged period of postweaning dependency. The advancement
of weaning age throughout human evolution coupled with
rapid and early brain growth implies a shift in how the rising
energetic demands of offspring are met: initially, energetic
costs are subsidized completely by the mother but then by
members of the social group through the provisioning of
weanlings (Humphrey 2010). This pattern of high maternal
investment and alloparenting behavior is important because
it is a clear determinant of birth spacing. Such a stratagem
has also been suggested to characterize the earliest members
of Homo (Aiello and Wells 2002). Recently, models have been
proposed to establish the precise age at which organisms were
weaned by accessing the isotopic record, in particular, strontium : calcium ratios (Sr/Ca) preserved within dental hard
tissues (e.g., Humphrey et al. 2008a, 2008b). Charting shifts
in this ratio throughout the developmental period associated
with early tooth tissue formation is one exciting way of reconstructing infant diet as well as tracking dietary transitions
throughout early life. If early Homo and later-occurring archaic Homo populations were indeed characterized by relatively early weaning, then analyses of the isotopic chemistry
throughout enamel development hold the potential to verify
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Figure 2. Bivariate plot of bite point positions for all teeth and masticatory muscle positions relative to the temporomandibular
joint (TMJ; y-axis) versus age (x-axis) in a cross-sectional ontogenetic series of modern humans (Nubian archaeological sample
housed at University of Colorado, Boulder). Bite points are measured as the distance of each tooth from the TMJ, measured in the
occlusal plane, and are illustrated by the two vertical arrows (left) indicating bite points for the dm1 and dm2. Masticatory muscle
position (for the superficial and deep masseters, temporalis, and medial pterygoid muscles) is defined as the point where each
muscles resultant force crosses the occlusal plane relative to the position of the TMJ. Primary masticatory adductor position and
orientation are based on a series of 2-D and 3-D linear and angular measurements and, taken together, capture what has been
termed masticatory system configuration (see Spencer 1995, 1999). Note the consistent position of the dm2 (small white sphere)
and the permanent M1, M2, and M3 (large white spheres) anterior to the masticatory muscles (i.e., above the white dotted line) at
the time of emergence. Also note the differing rates of anterior growth of the dental arcade and masticatory muscles (as indicated
by the slopes of second-order polynomials); space for emerging molars is a product of these different growth rates. A color version
of this figure appears in the online edition of Current Anthropology.
this with direct evidence for the age at weaning from the fossil
record itself.
While continued probing of the fossil record to establish
more precise demographic and maturational profiles holds
the potential to yield key details in the evolution of human
life history, another interesting way forward is to attempt to
understand the processes that lie behind the slightly dissimilar
patterns in dental development, and thus inferred life history
profiles, among hominins. A host of studies have advanced
our understanding of how dental developmental variation
intersects with primate life history variation, but surprisingly
little is known about the precise mechanism that governs,
modulates, regulates, constrains, and so forth, the timing of
molar eruption and as such the underlying processes that
regulate these temporal events are largely unknown.
Recently, it was postulated that a set of biomechanical constraints regulates masticatory system configuration throughout ontogeny and therefore modulates the position and ultimately the timing of emerging molars within developing
faces (Spencer and Schwartz 2008). Based on an ontogenetic
sample of modern human crania, it seems that successive
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Acknowledgments
I would like to thank Leslie Aiello and Susan Anton for their
invitation to participate in the Wenner-Gren workshop, all of
the conference participants for their stimulating and thoughtful discussions throughout the conference, and the two anonymous reviewers for their input and constructive comments
on this manuscript. The ideas and work laid out here are the
results of many discussions and ongoing collaborations, especially with Chris Dean, Jay Kelley, Tanya Smith, Debbie
Guatelli-Steinberg, Laurie Godfrey, Wendy Dirks, Bill Kimbel,
Mark Spencer, Terry Ritzman, Kierstin Catlett, and Halszka
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