AMERICAN JOURNAL OF PHYSICAL ANTHROPOLOGY 145:337–347 (2011)
Perspectives
Using the Life History Model to Set the Stage(s) of
Growth and Senescence in Bioarchaeology and
Paleodemography
Mirjana Roksandic1* and Stephanie D. Armstrong2
1
Department of Anthropology, University of Winnipeg, Winnipeg, Manitoba, Canada, R3B 2E9
2
Department of Anthropology, University of Manitoba, Winnipeg, Manitoba, Canada, R3T 2N2
KEY WORDS skeletal biology; paleodemography; life history theory; bioarchaeology
ABSTRACT Paleodemography, the study of demo-
graphic parameters of past human populations, relies on
assumptions including biological uniformitarianism, sta-
tionary populations, and the ability to determine point age
estimates from skeletal material. These assumptions have
been widely criticized in the literature and various solu-
tions have been proposed. The majority of these solutions
rely on statistical modeling, and have not seen widespread
application. Most bioarchaeologists recognize that our abil-
ity to assess chronological age is inherently limited, and
have instead resorted to large, qualitative, age categories.
However, there has been little attempt in the literature to
systematize and define the stages of development and age-
ing used in bioarchaeology. We propose that stages should
be based in the human life history pattern, and their skele-
tal markers should have easily defined and clear endpoints.
As a study of demographic parameters of past human
populations, paleodemography relies on three major
assumptions: biological uniformitarianism, stationary
populations, and the ability to determine point age esti-
mates from skeletal material. Despite numerous
criticisms raised on each of these assumptions (Petersen,
1975; Bocquet-Appel and Masset, 1982, 1985, 1996; Sat-
tenspiel and Harpending, 1983; Horowitz et al., 1988),
and a number of unsatisfactory attempts to address these
criticisms (Van Gerven and Armelagos, 1982; Buikstra
and Konigsberg, 1985; Greene et al., 1986; Milner et al.,
1989; Konigsberg and Frankenberg, 1992, 1994), there
seems to be a reluctance in the field of paleodemography
to let go of these assumptions and examine different
approaches to overcoming these difficulties. As currently
conceptualized, the construction of mortality profiles is
based on methods developed for modern human popula-
tions, founded on cohort statistics. A cohort is defined as
a group that shares a common experiential trait, such as
birth year. Relationships such as this can rarely be deter-
mined in an archaeological context. This has been recog-
nized as problematic since the very onset of paleodemo-
graphic research by Angel (1969), who recognized that it
cannot be assumed that the individuals have cohort rela-
tionships, nor can it be assumed that the sample popula-
tion is unbiased. Furthermore, paleodemography com-
monly uses 5- or 10-year age categories (Lancaster, 1990)
which stem from the application of model life tables
C 2011
V WILEY-LISS, INC.
In addition to a standard five-stage developmental model
based on the human life history pattern, current among
human biologists, we suggest divisions within the adult
stage that recognize the specific nature of skeletal samples.
We therefore propose the following eight stages recogniz-
able in human skeletal development and senescence:
infancy, early childhood, late childhood, adolescence, young
adulthood, full adulthood, mature adulthood, and senile
adulthood. Striving toward a better prediction of chronolog-
ical ages will remain important and could eventually help
us understand to what extent past societies differed in the
timing of these life stages. Furthermore, paleodemogra-
phers should try to develop methods that rely on the type
of age information accessible from the skeletal material,
which uses life stages, rather than point age estimates. Am
J Phys Anthropol 145:337–347, 2011. V 2011 Wiley-Liss, Inc.
C
derived from modern human populations. This is prob-
lematic not only because we are unable to determine
point age estimates, but because these categories have no
underlying biological meaning insofar as these categories
group individuals of different life stages together based
on chronological age, with little correlation to biological
age. In this article we propose to work within the biologi-
cal reality of the skeletal material and redefine age
categories to suit this reality. This would likely mean
avoiding cohort statistics and the assumption of biological
uniformitarianism by using only biological age groups,
independent of chronological age. Furthermore, the size
and the structure of the population at risk are not
known; it is commonly assumed that the excavated
(analyzed) population is an unbiased sample of the
Grant sponsor: SSHRC; Grant number: 861-2009-0071; Grant
sponsor: NSERC-USRA.
*Correspondence to: Mirjana Roksandic, Department of Anthro-
pology, University of Winnipeg, 515 Portage Avenue, Winnipeg,
Manitoba, Canada, R3B 2E9. E-mail: m.roksandic@uwinnipeg.ca
Received 29 June 2010; accepted 10 January 2011
DOI 10.1002/ajpa.21508
Published online 5 April 2011 in Wiley Online Library
(wileyonlinelibrary.com).
338 M. ROKSANDIC AND S.D. ARMSTRONG
population at risk. This assumption is not always war-
ranted, and even though a fairly robust method for the
examination of bias in buried populations has been pro-
posed (Jackes, 1993; Jackes and Meiklejohn, 2008), it is
rarely examined or reported.
Bioarchaeologists are increasingly recognizing the
problems within the field and—as we show below—they
are avoiding both point age estimates and 5-year catego-
ries. Increasingly, they are using biologically meaningful
categories that are not dependent on precise determina-
tion of chronological age. In this article, we draw atten-
tion to the changes in the field of bioarchaeology that
are the result of the earlier criticisms of paleodemogra-
phy, and recognize the increasing use of age categories
rather than point age estimates and 5- and 10-year age
groups derived from point age estimates. These age cate-
gories are, however, never properly defined in relation to
a sound underlying principle or a theoretical concept:
their meaning is assumed rather than discussed in any
of the papers. We propose that life history theory pro-
vides the best underlying model for constructing age cat-
egories. In that context, we examine the biology of
human growth and development using a life-history per-
spective. The age categories derived from life history are
matched to stages of skeletal and dental development in
non-adults. These developmental markers are examined
against primate models, to stress their universality and
applicability across populations. We further discuss skel-
etally recognizable and meaningful stages within adult-
hood that are critically important for paleodemographic
and paleopathological studies. While it is evident that
these groups will not in and of themselves solve the
problems inherent in paleodemographic research, we
hope that the article will prompt paleodemographers to
build their models on thus redefined, readily available
information that, limited as it is, could potentially reveal
important aspects of past human groups based on sub-
stantiated biological processes. In itself—even if mortal-
ity/survivorship profiles could not be reconstructed due
to cemetery biases and problems inherent to paleode-
mography (Petersen, 1975)—analyzing the population in
terms of relative representation of different stages could
advance our understanding of past human experiences
of life. Furthermore, while we deliberately avoided set-
ting chronological boundaries to these life stages, they
carry intrinsic age information that could be used to
build mortality profiles, provided development of proper
paleodemographic techniques. It is also important to
stress that finding better methods of point age estima-
tion is still a very important goal. Better methods in
age-at-death estimation would allow us to examine at
which chronological ages past populations experienced
the transition from one life-history stage to another.
Issues and criticisms
The problems associated with using life tables in pale-
odemography were recognized at the very onset of paleo-
demography as a discipline. In his theoretical article,
Angel (1969) outlined three unsubstantiated assump-
tions that are made in constructing life tables from
cemeteries: ‘‘. . . that the cemetery represents a single
generation cohort, that death rates are even at all ages
after infancy and hence directly reflected in cemetery
age frequencies, and that the population is virtually sta-
ble biologically and socially over the period of cemetery
American Journal of Physical Anthropology
use,’’ (Angel, 1969: p 428). Despite Angel’s warnings, life
tables—a tool used to calculate life expectancy and the
probability of survival at a particular age—went on to be
widely used throughout the field starting with Acsádi
and Nemeskéri (1970) and Angel himself (1971). Acsádi
and Nemeskéri (1970) provided the first standard text
on determining demographic parameters from ancient
populations by using life tables, followed by a number of
case studies (e.g., Blakely, 1971; McKinley, 1971; Ben-
nett, 1973; Owsley and Bass, 1979; Piontek and Henne-
berg, 1981, etc.). These case studies used abridged life
tables that grouped populations into 5-year age intervals
as advocated by Ubelaker (1978), who deemed these
intervals large enough to eliminate most of the probable
errors in age determination.
Following the initial enthusiasm for paleodemography,
the field underwent its first substantial criticism by Boc-
quet-Appel and Masset (1982). In their ‘‘Farewell to Pale-
odemography’’ the authors argued that age determination
was inaccurate and that age-at-death profiles tended to
mirror the reference samples from whence they were
derived (Bocquet-Appel and Masset, 1982). This latter
criticism was addressed by many in favor of using more
appropriate reference samples (Van Gerven and Armela-
gos, 1982; Buikstra and Konigsberg, 1985; Greene et al.,
1986), though more recently it has been recognized that
reference population mimicking remains a valid concern
(Buikstra, 1997).
The problem of inaccuracy in age determination is one
that continues to plague paleodemography. This issue
has been thoroughly discussed in the anthropological lit-
erature (Murray and Murray, 1991; Kvaal et al., 1994;
Ley et al., 1994; Aykroyd et al., 1999; Nagar and Hersh-
kovitz, 2004). There is a definite tendency of archaeologi-
cal populations to show a mortality peak between 30 and
45 years of age and an absence of older individuals—a
pattern not seen in historical hunter-gatherer and sub-
sistence populations. Rather than reflect a reality of
archaeological populations, this pattern shows the bias
in skeletal age determination methods, and reflects ref-
erence populations that rarely contain individuals older
than 60 years of age (see Fig. 1 of Aykroyd et al., 1999;
Chamberlain, 2006).
In addition to the practical issues of age determination,
there are further problems with the assumption of biolog-
ical uniformitarianism. This assumption implies that
age-related changes in humans are the same throughout
time and space. It has been noted that this is only par-
tially correct—though the pattern of age-related changes
may be the same, the rate at which these changes occur
varies between males and females, and across popula-
tions (Hoppa, 2002). Cultural practices can also affect
individual ageing patterns (Gowland, 2006).
The most problematic assumption of paleodemography,
rarely addressed by paleodemographers, is the existence
of cohort relationships in a sample population. Cohorts
are studied for the very reason that each cohort is differ-
ent and that ‘‘. . . successive cohorts do something other
than merely repeat the patterns of behavior of their pred-
ecessors’’ (Weeks, 2005: p 321). It is also noted that
though successive cohorts are expected to be similar in
some ways, they can differ in others (namely size, compo-
sition, and age-specific patterns of survival or longevity),
which is precisely why they are studied (Weeks, 2005).
Understanding these changes is the goal of paleodemog-
raphy, but by their reliance on artificial cohorts and
assumption of ‘stationary populations’ paleodemographers
 STAGES OF GROWTH AND SENESCENCE
Fig. 1. Graph of age categories used in American Journal of
Physical Anthropology, in two periods: the pre-criticism period
between 1978 and 1982 at the top and the recent 2005 and 2009
period at the bottom. It should be noted that the category
‘‘Other numerical’’ includes many studies using only non-adult
specimens, which generally used small age categories due to the
relative ease of determining age in subadults. The majority of
articles in 2005–2009 employing 5-year age categories were self-
defined paleodemographic studies.
deny that any change occurs. Furthermore, although a
cohort can be any delineated population that experiences
a particular life event in a given period of time, paleode-
mography uses a population as if it were a birth cohort,
which is by definition one that is born in the same
defined period of time (usually a year or a decade), in
order to examine death by age (Glenn, 1977). Ultimately,
it is inappropriate to treat an archaeological population
as a birth cohort when it is unknown when the individu-
als were born, or when the period of time in which the
cohort as a whole was born is large, as is often the case
with archaeological populations. The result of this method
is that it attempts to take into account only the effects of
age on life expectancy, while neglecting other important
and inseparable effects, such as cohort effects (the effects
due to being part of a particular birth cohort) and period
effects (effects due to the period of time in which that
individual or group lived) (Glenn, 1977).
Archaeological populations do not allow us to account
for cohort and period effects, and as a result, we can
only estimate a very crude age-specific death rate (i.e.,
number of individuals who died within a particular age
range over the total population) to measure mortality
among archaeological populations, with the knowledge
that no cohort relationship exists between the individu-
als who perished at any particular age (Lancaster, 1990).
Crude death rates may be as precise as paleodemogra-
phy is capable of achieving, but even they might be a
lofty goal, given that the structure of the population at
risk is hard to ascertain. While the dead are never an
unbiased sample of a living population, as Jackes (2011)
points out, ‘‘an unbiased cemetery sample. . . closely
339
approximates this ‘normal’ U-shaped mortality by age
curve.’’ Jackes further proposes a relatively robust test
of bias that is based on comparing fertility estimates
derived from the Juvenile: Adult ratio (juvenility index
of Bocquet-Appel and Masset, 1977) on one hand, and
her total fertility estimate based on the Mean Childhood
Mortality on the other (Jackes, 2011). Consequently,
archaeological populations should be tested for bias, and
those that can be recognized as biased should be
excluded from any further paleodemographic study.
Current approaches
The response to the criticisms regarding inaccuracies in
age determination and in the underlying age distribution
in sample populations has focused on statistical methods
that could address these inaccuracies such as: hazard
models (Gage, 1988); maximum likelihood models (Paine,
1989); the iterated-age-length key method (Konigsberg
and Frankenberg, 1992); the use of Bayes’ theorem (Ayk-
royd et al., 1999); and weight functions (Müller et al.,
2002). Most recently, the ‘‘Rostock manifesto’’ proposed
that the Bayesian approach should be the method of
choice, but that the age distribution (f (a)) of the target
population must be established before osteologists are able
to determine the probability of a person from that target
population dying at a particular age, which again depends
on statistical modeling to determine how the probability of
death varies with age (Hoppa and Vaupel, 2002).
This somewhat circular logic and heavy reliance on
statistics makes practical application difficult, as
reflected in the relative absence of case studies that
have emerged from the manifesto. Furthermore, the
probabilities are calculated on the basis of the same poor
age indicators (for a recent attempt to apply this model
see DeWitte, 2009, although the author uses additional
age information provided by documentary evidence).
Because of the practical and theoretical complexities of
using the statistical methods suggested over the past
few years, a number of more recent case studies (Steyn,
2003; Castro et al., 2004; Mosothwane and Steyn, 2004;
Nagaoka and Hirata, 2007), although recognizing the
problems with the life table method, continue to use the
traditional paleodemographic approach.
Consequently, a substantial rift separates theoretical
statistical paleodemography and bioarchaeological
attempts to understand past human populations. This is
no less true with regards to age categories. As previously
mentioned, 5-year intervals have been commonly used in
paleodemographic reconstructions. Used for simplicity of
analysis for examining birth cohorts in modern demogra-
phy (Weinstein and Pillai, 2001), 5-year intervals have
been used in paleodemography due to the difficulties in
establishing point age estimates from the individual skel-
etons. Outside of paleodemography, however, very few
bioarchaeologists use 5-year categories, as evidenced by a
survey of articles using age categories in the past 5 years
of the American Journal of Physical Anthropology (see
Fig. 1). This journal being the most relevant biological
anthropology journal, the past 5 years should reflect cur-
rent trends in the use of age categories. It becomes clear
that many researchers use qualitative categories (e.g., ju-
venile, adolescent, young adult, mature adult, etc.) with
or without a defined year range, as opposed to paleode-
mography articles which tend to use strict 5- or 10-year
age categories. This may reflect the urge to use biologi-
cally meaningful age categories, as opposed to simply
American Journal of Physical Anthropology
340 M. ROKSANDIC AND S.D. ARMSTRONG
TABLE 1. Age categories in AJPA 1978–1982, 2005–2009
Age categories
Year 5-year 5-year (%) 10-year 10-year (%)
1982 4 57.14 1 14.29
1981 2 40.00 3 60.00
1980 2 40.00 0 0.00
1979 1 14.29 4 57.14
1978 0 0.00 3 100.00
Total average 9 30.29 11 46.29
2009 2 7.14 7 25.00
2008 1 5.00 4 20.00
2007 1 6.25 4 25.00
2006 4 21.05 6 31.58
2005 1 6.67 5 33.33
Total average 9 9.22 26 26.98
outlining chronological age. When this is compared to the
five years of the journal before the major criticisms of
paleodemography emerged in 1982, it becomes evident
that views on the quality of methods of age determina-
tion have changed—it is no longer assumed that precise
ages can be determined for adults (Table 1).
Despite some positive reviews of our ability to deter-
mine age in skeletal remains (Wright and Yoder, 2003),
determining point age estimates remains an imprecise
endeavor, subject to substantial scrutiny, particularly for
the age assessment of adult specimens. It has already
been noted that methods of age determination provide a
better idea of physiological or biological age rather than
chronological age (Kemkes-Grottenthaler, 2002; Jackes
et al., 2008). In fact, biological age is the only age that is
apparent on skeletal remains; it is usually the bioarch-
aeologist who attempts to convert this into years based
on a correlation between biological and chronological age,
and the assumption of biological uniformitarianism. Yet
it is understood that, in addition to the difficulties of
establishing chronological age from skeletal remains and
the issues of mimicking a reference population, chrono-
logical age is often an irrelevant measurement in other
cultural settings (Gowland, 2006; Bocquet-Appel and
Bacro, 2008). The physical process of growth and senes-
cence, however, is a human universal, though certainly
the importance assigned to any particular maturational
stage, or even the timing of these events, may be affected
by cultural practices, genetics, and environment, among
others. It is well understood that the chronological timing
of biological events can vary wildly between individuals
and populations. Thus, for biological anthropologists it
seems only logical to determine maturational or degener-
ative stage, as opposed to chronological age. The increas-
ing use of descriptive age categories by bioarchaeologists
indicates that this might be the next logical step for pale-
odemographic research as well. Ultimately, it is likely
more informative, even from a paleodemographic per-
spective, to discover mortality patterns by meaningful,
biological life-stage as opposed to chronological age. It is
nevertheless obvious that projecting a population by life
stage will be further hampered by the fact that stages
will be of different lengths. Therefore, establishing better
methods of assigning chronological age remains an im-
portant goal that will ultimately allow us to discuss the
chronological ages at which different life history transi-
tions were experienced in different populations. Since
even methods such as the analysis of tooth cementum
annulations, that claimed high levels of accuracy in a
known age context (Stott et al., 1982; Wittwer-Backofen
American Journal of Physical Anthropology
Other numerical Other numerical (%) Qualitative Qualitative (%)
2 28.57 0 0.00
0 0.00 0 0.00
0 0.00 3 60.00
2 28.57 0 0.00
0 0.00 0 0.00
4 11.43 3 12.00
7 25.00 12 42.86
8 40.00 7 35.00
4 25.00 7 43.75
0 0.00 9 47.37
4 26.67 5 33.33
23 23.33 40 40.46
et al., 2004), have been heavily criticized (Renz and
Radlanski, 2006; Roksandic et al., 2009), we still do not
have a method that will allow us to calibrate these stages
in relation to their chronological age expression in differ-
ent populations. This, however, should be our ultimate
goal. In the meantime, age categories currently used by
bioarchaeologists need to be defined and linked to mean-
ingful biological processes.
Life history model
There are many complicating factors to determining bi-
ological age. As Beall (1984) notes, most maturational
changes finish by the end of the second decade of life and
subsequent degenerative changes are much more vari-
able. As already noted, there has been no discussion on
how to appropriately divide the human lifespan into life
stages, as evidenced by different descriptive categories
used by researchers, not only when dealing with adults,
but also in non-adult skeletons.
The model proposed here uses biologically meaningful
stages, under the overarching framework provided by life
history theory. Life history theory is concerned with the
trade-offs between growth, reproduction and maintenance
(McDade, 2003), and how these interacting factors affect
the evolution of a particular life history pattern. This
ontogenic process and consequent biological compromises
result in a distinct pattern of life stages where each stage
represents the sum of the coordination between behavior
and physiology (Del Guidice et al., 2009); in humans, this
process has led to the pattern characterized by helpless
infants, slow maturation and long childhood dependency,
menopause and long life span (Bogin, 1999). The use of a
model of progressive life stages can circumvent many of
the difficulties that arise when determining the chrono-
logical age of individuals. It requires identifying clear de-
velopmental and degenerative endpoints on the skeleton,
which serve as a proxy for those biological events that
cannot be observed on the skeleton, but through which all
humans must pass (Leidy, 1996). The chronological age at
which these endpoints are achieved may vary, between
individuals and across populations, depending on a num-
ber of factors. Though understanding the variation of age
at these endpoints across populations remains an impor-
tant area of study, the chronological ages at which these
endpoints occur is not the focus of the proposed approach.
The model outlined here is an adaptation of the pat-
tern proposed by Bogin (1990, 1994, 1999, 2001, 2003).
Bogin’s five-stage model includes infancy, childhood,
juvenility or juvenescence, adolescence, and adulthood.
 STAGES OF GROWTH AND SENESCENCE 341
TABLE 2. Skeletal markers of model stages
Skeletal and dental markers
Bogin model Proposed model Starting point Endpoint
Infancy Infancy Full deciduous dentition
Childhood Early Childhood Full deciduous dentition Eruption of first permanent tooth:
First molar
Central incisor
Juvenescence Late Childhood Eruption of first permanent tooth Eruption of permanent canine
Adolescence Adolescence Eruption of permanent canine (sexual Epiphyseal fusion of long bones (cessation of
maturity) longitudinal growth, reproductive
maturity)
Adulthood Young adulthood Epiphyseal fusion of long bones Epiphyseal fusion of medial clavicle
Closure of spheno-occipital synchondrosis,
iliac crest and ischial tuberosity
Full adulthood Epiphyseal fusion of medial clavicle Complete obliteration of all fusion lines on
Closure of spheno-occipital synchondrosis, vertebrae
iliac crest, and ischial tuberosity Closure of superior sacral segments
Degenerative changes begin
Mature adulthood Complete obliteration of all fusion lines on Substantial degenerative changes
vertebrae Final stages of pubic symphysis, iliac
auricular surface, sternal rib end methods
Cranial suture obliteration
Closure of superior sacral segments Substantial intravitam tooth loss, with
Degenerative changes begin resorption
Senile adulthood Severe degenerative changes
Final stages of pubic symphysis, iliac
auricular surface, sternal rib end methods
Cranial suture obliteration
Substantial intravitam tooth loss, with
resorption

Since the term ‘‘juvenile’’ is used by both paleodemogra-
phers and bioarchaeologists (even in very important vol-
umes such as Bocquet-Appel and Masset, 1977; Scheuer
and Black, 2000) to indicate non-adults in general, we
prefer to substitute the terms childhood and juvenility
with early and late childhood, respectively. Each of these
stages is defined by distinct biological events. In addition
to Bogin’s model, which recognizes only one adult stage,
we outline several stages within adulthood that are
based on observable skeletal changes associated with the
final developments of the skeleton and the degenerative
changes that follow the end of skeletal growth (Table 2).
Given the nature of bioarchaeology and the skeletal
material it uses, we will focus mainly on the skeletal and
dental developments, though it is understood that these
developmental stages are also marked by cognitive and
physiological changes. Dentition is of particular impor-
tance in defining stages of growth, as it is less susceptible
to environmental and nutritional changes, and is ‘‘... inte-
grated into the plan of growth and development, timed to
the growth of the skull, maturation of muscles of mastica-
tion, and somatic growth in general’’ (Smith, 1989: p
683). Once full dentition is reached, changes associated
with the post-cranial skeleton become more relevant.
Nonadults (juveniles)
The juvenile (the term is here used as is common in
bioarchaeological practice to denote a non-adult) period
can be divided into infancy, early childhood, late child-
hood, and adolescence.
Infancy can be defined, among other biological and
cognitive developments, as the time between birth and
the eruption of a full set of deciduous teeth at 2–3
years of age (Bogin 2001). This also tends to coincide
with weaning, which is a highly culturally dependent
event in humans, but can nonetheless be roughly corre-
lated to complete eruption of the deciduous dentition,
occurring at about 2.5 years (Smith et al., 1994). After
this time, substantially more non-maternal sources of
food are introduced into the diet, though the exact tim-
ing is, again, culturally dependent.
In primates, infancy is defined as the period from birth
to weaning which coincides with the eruption of the first
permanent molar (Smith, 1991). Even in primates, wean-
ing is a gradual process, and its timing may be affected by
seasonality and the availability of weaning foods (Lee,
1996; Strier, 2007). In humans, the end of infancy also
tends to coincide with weaning, but this occurs years
before the first permanent tooth erupts (Bogin, 1999).
Consequently, humans not only have a relatively short
infancy compared to non-human primates, but they also
experience the unique stage of early childhood between
weaning and the eruption of the first permanent molar.
Early childhood, a time of rapid brain growth and
extensive learning, is recognized as a stage primarily for
its intellectual and social, rather than physical develop-
ment (Mace, 2000). In terms of skeletal biology this
stage corresponds to the time between the full set of de-
ciduous teeth and the eruption of the first permanent
tooth, which tends to be the first molar, but can also be
one of the central incisors (Smith et al., 1994; Bogin,
2001; Del Guidice et al., 2009). Although all deciduous
teeth have emerged, the individual is still dependent on
adults for care and feeding, and they have difficulty
processing an adult diet (Bogin, 2003). At least theoreti-
cally, the eruption of the full deciduous dentition indi-
cates the ability of the child to process food, though
there is variation between different societies and individ-
uals as to the extent of solid food introduced in the diet
and the continued reliance on breastfeeding.
After weaning, nonhuman primates are relatively in-
dependent in comparison to human children, who are
still helpless for a number of years during which the

American Journal of Physical Anthropology

342 M. ROKSANDIC AND S.D. ARMSTRONG
child must continue to be fed by specially prepared food
by care-giving adults (Del Guidice et al., 2009). The fact
that this adult need not be the mother is important as
this reduced dependency for breast milk allows for rela-
tively shorter birth intervals in humans (Bogin, 2003).
Furthermore, it is possible to speculate that the food
processing behavior, particularly thermal processing
(cooking), seen in humans has aided the usefulness of
this stage, given that children cannot process a fully
adult diet and have a high food-energy requirement for
brain growth (Carmody et al., 2010). Childhood may pro-
vide humans with the necessary time for learning and
understanding culture, in accompaniment with the rapid
brain growth characteristic of this stage (Bogin, 1990).
Further proof for the existence of early childhood as a
uniquely human life stage is the difference in growth
profiles. Humans are distinct in that three mathematical
functions are required to plot their growth, due to the
mid-childhood growth spurt which is not seen in other
primates (Bogin, 1990). The mid-childhood growth spurt
is characterized by another event not observable in the
skeleton, the onset of adrenal androgen production, or
adrenarche (Bogin, 1997). The effects of adrenarche are
wide-ranging, from social and cognitive behavior, to so-
matic growth and development, to effects on the neuro-
logical and immune systems (Campbell, 2006).
Following early childhood is late childhood, coinciding
with the time between the eruption of the first perma-
nent tooth and the eruption of the permanent canine.
Late childhood is the time of pre-puberty and relative in-
dependence; it is also when the growth rate is particu-
larly slow. In addition, there are a number of social
developments—late childhood is a time of increased play
and increased sexual differentiation of activities (Del
Guidice et al., 2009). The eruption of the first permanent
tooth at the end of early childhood tends to coincide with
the end of the mid-growth spurt that occurs in human
children, as well as with adrenarche (Bogin, 2001). Adre-
narche seems to be related to gonadarche (puberty), and
it has been proposed that it may ‘‘serve to activate amyg-
dala and hippocampus activity prior to the onset of sex
steroids, reducing fearfulness and anxiety and increasing
social interaction with unfamiliar individuals’’ (Camp-
bell, 2006: p 585).
Adolescence is a time of hormonal changes, secondary
changes (e.g., pubic hair, breast buds, seminal emission,
menarche, etc.), and rapid growth, namely the adolescent
growth spurt (Bogin, 2001). This phase corresponds to
the time between sexual and reproductive maturity,
marked by the eruption of the permanent canines and
skeletal maturity, respectively (Bogin, 2001). Skeletal ma-
turity in this case refers to the end of longitudinal
growth, with complete epiphyseal fusion of the long
bones. To a lesser extent, the end of adolescence may also
be marked by the eruption of the third molar, although
this has become a variable, and consequently, unreliable
event (Smith, 1989). Once adolescence has ended, adult-
hood is achieved and the majority of growth-related
changes are complete.
The characteristic dramatic increase in the rate of skel-
etal growth in adolescent humans is not observed in
nonhuman primates, although chimpanzees may demon-
strate an increase in body mass in this period (Bogin and
Smith, 1996; Kappeler and Pereira, 2003). In females,
sexual maturity is marked by menarche, while reproduc-
tive maturity is marked by the ability to successfully
carry a child to term. The gap between sexual and repro-
American Journal of Physical Anthropology
ductive maturity, the time which is occupied by adoles-
cence in humans, is not unique to humans as it is also
seen in great apes. Unlike humans, in great apes this
gap is relatively short in terms of total lifespan (Bogin,
1999). Similarly, while sexual maturity is also the time of
permanent canine eruption in humans, it is instead
marked by eruption of the third permanent molar in pri-
mates such as Pan and Gorilla, an event which does not
occur in humans until reproductive maturity or later, if
it occurs at all (Dainton and Macho, 1999).
Though perhaps not well-recognized, the timing of hor-
monal changes indicating the onset of puberty in girls
and boys coincides with the eruption of the permanent
canines, and consequently the canines serve as a better
predictor for the onset of adolescence than the eruption
of the second molars. Though he does not discuss the
association between the two events, Bogin (2001) does
separately discuss the timing of canine eruption (p 83)
and onset of puberty (p 87). Similarly, the eruption of
the canines is a less variable predictor of the onset of pu-
berty than menarche, an event which is under strong
environmental pressures and whose variation in timing
is well-documented (Moffitt et al., 1992; Worthman,
1999; Freedman et al., 2002; Chumlea et al., 2003). The
advantages to using the eruption of canines as a proxy
for the onset of sexual maturity and adolescence is that
it is observed in both males and females, and can be
observed in a bioarchaeological context.
The onset of reproductive maturity indicates adulthood
and cessation of growth, as growth and reproduction are
mutually exclusive in terms of life history theory
(Harvey et al., 1985). The evolutionary purpose of ado-
lescence is far more speculative than the distinct evolu-
tionary advantages of childhood. Nonetheless, it has
been proposed that since the timing and type of changes
that occur in girls and boys differ, the evolutionary rea-
sons behind them may differ as well. In girls, menarche
is followed by a few years of sterility—indeed, the female
pelvis is not affected by the adolescent growth spurt and
according to Bogin (1994) does not reach its adult size
until several years after adult stature is attained. Conse-
quently, they appear to be fully grown adults before they
are in fact reproductively mature, which Bogin (1994)
suggests leads adults to encourage these adolescent girls
to participate in adult social interactions, and conse-
quently learn how to be reproductively successful. In
contrast, boys appear more child-like despite the hormo-
nal changes which encourage them to participate in
adult behavior. Consequently, due to their appearance,
they are not perceived to be adults and do not constitute
competition for adult males (Bogin, 1994). Ultimately, it
can be said that, ‘‘Girls best learn their adult social roles
while they are infertile but perceived by adults as
mature, whereas boys best learn their adult social roles
while they are sexually mature but not yet perceived as
such by adults. Without the adolescent growth spurt this
unique style of social and cultural learning could not
occur’’ (Bogin, 1994: p 33).
It should be noted that reproductive maturity cannot
be equated with fertility; while sexual maturity is fol-
lowed by a period of sterility, human females are fertile
before they are reproductively mature (Bogin, 1994). Sim-
ilarly to the onset of menarche, fertility is under strong
environmental pressures. While in well-fed populations,
individuals can be fertile shortly after menarche; this has
not traditionally been the case, and generally females are
not fertile until a few years after the onset of menarche
 STAGES OF GROWTH AND SENESCENCE
(Apter, 2008). Reproductive maturity tends to coincide
with skeletal maturity, which may be explained by the
involvement of gonadal estrogen in both the ovarian cycle
and the regulation of epiphyseal fusion (Kramer and
Greaves, 2010). In addition, the pelvis develops more
slowly than the rest of the skeleton, and is not really
affected by the adolescent growth spurt, coinciding better
with the end of long bone epiphyseal fusion (Moerman,
1982). Consequently, reproductive maturity is attained
when the pelvic basin has completely developed, though
there are secondary centers of ossification in the coxal
bone which continues to develop in adulthood. While the
timing of skeletal changes in adolescent males and
females certainly differs, there are few formalized studies
on both living and known age populations. However, the
overall pattern of these changes seems to follow the same
course, which allows the application of the skeletal end-
points associated with adolescence in this model.
Adults
According to life history theory, adulthood commences
upon attaining reproductive maturity, once the longitudi-
nal growth is complete. The model outlined above is a
useful and well-defined biological model for understand-
ing human growth and development as reflected in hard
tissues. However, once the long bone growth is complete
and the individual reaches reproductive maturity, or
adulthood, the five-stage life history model does not dif-
ferentiate or recognize any other stages besides meno-
pause in women, which cannot be observed in the skele-
ton, and does not apply to men. We could accept this,
and use different estimators for paleodemographic stud-
ies, such as the juvenile/adult ratio, to understand past
populations (Bocquet-Appel and Masset, 1996). Having
only one adult stage, however, would effectively preclude
our ability to build mortality profiles. While these are
not necessarily informative about age-specific mortality,
they could contribute information about fertility. Accord-
ingly, while further stages do not have a firm basis in
life history theory, we include them here because they
are observable in skeletal samples, allowing us to use
skeletal markers of senescence in the same way that the
markers of growth are used: to assess the stage in the
life of an individual. However, we have to acknowledge
that there are no universal or definite biological corre-
lates to these age stages other than progressive changes
in the experience of the quality of life; although they
vary individually in extent and timing, they are experi-
enced by all humans. Furthermore, we cannot associate
these with either chronological age or point age esti-
mates, or any a priori social meaning that the stage of
life could carry. We propose that there are identifiable
changes that occur to the bone once longitudinal growth
is complete, and though these changes may be less
grounded in life history theory than the stages outlined
above, they may still prove useful to bioarchaeology, and
possibly to paleodemography. In terms of bioarchaeologi-
cal studies, if our categories were easily recognized in
any skeletal population, we could use these categories
against disease prevalence data, burial data or any other
relevant marker of the biological or social persona whose
remains we are studying.
We suggest that—provided sufficient preservation—it is
relatively easy to recognize four adult stages: young
adult, fully adult, mature adult, and senile adult. Preser-
vation is obviously an issue for all skeletal collections.
343
Given that the method relies on a limited number of
clearly defined and easily observable age markers, if the
preservation of these elements is not systematically bi-
ased it should allow observation as often as any other
ageing method. For comparison, the pubic symphysis and
the auricular surface, the most commonly used age indi-
cators in bioarchaeology, preserve very poorly in archaeo-
logical populations. For the Iron Gates Gorge Mesolithic
population, the auricular surface was available in 13.1%
(43/328) of individuals, while the pubic symphysis was
available in 3.6% (12/328) (Roksandic et al., 2002). It is
true that in some cases skeletal preservation will pre-
clude us even from distinguishing a juvenile from an
adult. However, if the relevant elements are preserved,
even a poorly preserved skeleton can be assigned a stage.
The beginning of the young adult stage is character-
ized by the attainment of longitudinal growth: all long
bone epiphyses are closed except for the medial clavicle.
However, epiphyseal lines are still visible and a number
of secondary structures remain open: the iliac crest, the
ramal epiphysis (Scheuer and Black, 2000), and the
spheno-occipital synchondrosis. The fusion of the latter
coincides with the epiphyseal fusion of the medial clavi-
cle (Krogman and Is _ can, 1986; Schwartz, 1995). The
young adult stage then ends when the spheno-occipital
synchondrosis is closed, the clavicle is completely fused,
and the iliac crest and ischial tuberosity are joined to
the coxal bone. The clavicle should be used in particular,
as there has been some evidence that the spheno-occipi-
tal synchondrosis may begin to close earlier than previ-
ously thought (Madeline and Elster, 1995). Once this
endpoint has been achieved, the individual enters full
adulthood.
Full adulthood would thus start with the closure of all
secondary centers of ossification in long bones and last
until the obliteration of fusion lines between the cen-
trum and the annular rings of the vertebrae, particu-
larly the thoracic and upper lumbar vertebrae (Albert,
1998). During the full adulthood stage, two superior sac-
ral segments open in young adulthood close (Scheuer
and Black, 2000). Degenerative changes are generally
not present at this stage, but since these changes can be
caused by pathological conditions, which may or may not
be related to the aging process itself, this criterion
should be applied with caution.
The onset of mature adulthood can be associated with
the complete obliteration of the vertebral lines and the
appearance of degenerative changes. However, degenera-
tive changes which occur are not consistent among indi-
viduals or across populations; not only is there variabili-
ty in timing, but the changes themselves may or may
not occur depending on lifestyle, occupation and genet-
ics. Each of the traditionally used age determination
methods carries substantial information on the process
of ageing: definite signs of degenerative changes on ver-
tebrae, the pelvis, the beginning of closure of skull
sutures and dental attrition. Within and between popu-
lations, variation makes these changes unreliable in
terms of point age estimates; however, taken together
and evaluated against each other they generally provide
enough information on the skeletal stage. In terms of
traditional methods of age determination, it may be pos-
sible to use and combine some of the phases outlined in
the Suchey-Brooks pubic symphysis (particularly Phase
3) and Lovejoy et al. auricular surface (Phases 3–6)
methods to indicate a mature adult, though these should
be used very cautiously (Lovejoy et al., 1985; Brooks and
American Journal of Physical Anthropology
344 M. ROKSANDIC AND S.D. ARMSTRONG

Suchey, 1990). It should be noted that the downside to

using these methods is the overlap that occurs between
the phases and the high degree of inter-observer error.
These methods should be used in conjunction with one
another, not individually, and skeletal structures need to
be evaluated against each other.
Senile adult is by far the most difficult stage to deter-
mine, and in many cases—especially with poor preserva-
tion—it may not be possible to distinguish a senile adult
from a mature adult or an individual with a pathological
condition. It can be defined by severe degenerative
changes in the axial skeleton, suture obliteration, and
substantial intravitam tooth loss with resorption. These
changes would include indicators used in traditional
methods of age determination, such as the pubic symphy-
sis, auricular surface of the ilium, and the sternal rib end
_ can and Loth, 1986; Brooks and
(Lovejoy et al., 1985; Is
Suchey, 1990). Rather than using the gradient involved
in these systems, only the final stages (which are tradi-
tionally used to ascribe an age of 501) would be consid-
ered for this model in order to describe a senile adult.
Although in the context of life history theory these
changes could be related to menopause—a time when
endosteal bone deposition more or less stops and resorp-
tion begins—there are several reasons that argue against
this approach. First, it may not be possible to identify
resorption and attribute it to menopause (Bogin and
Smith, 1996), and furthermore, menopause does not
account for changes associated with senility in males, nor
can it be accurately observed in a skeletal remains. In
essence, the senile stage would indicate an individual—
regardless of their actual chronological age—that experi-
ences significant loss of quality of life due to a number of
systemic degenerative changes recognizable in their skel-
eton, that are not limited to a specific pathology.
Our ability to define an adult stage would depend
strongly on the preservation of relevant skeletal areas.
While it is relatively easy to distinguish between young
and fully adult categories as well as between these two
and older individuals, the borderline between mature
and senile adults is more fluid, as it relies on cumulative
loss of quality of life rather than on a set of predefined
features. Therefore, where preservation is partial we
might need to reduce the number of categories to three
easily recognizable ones: young adult, fully adult, and
older adult.
Can the use of the life stage model improve
paleodemographic research?
Criticisms of paleodemography have resulted in a num-
ber of novel approaches in the field that all rely on so-
phisticated modeling. Proper understanding of both
demography and statistical modeling are therefore a nec-
essary prerequisite for paleodemographic research. These
are rarely within the reach of practicing bioarchaeolo-
gists, as few of us are properly trained in demography.
Since bioarchaeologists are responsible for providing the
age-related skeletal information used in statistical
approaches, we need to bridge this rift between theoreti-
cal paleodemography and bioarchaeology. Bioarchaeolo-
gists are increasingly wary of point age estimates, and
are resorting to larger age groups, especially for adult
skeletons. Paleodemographers need to acknowledge this
trend and work with the available data. The proposed
life stages have the benefit of reflecting biological and
skeletal reality within which bioarchaeologists operate.
Fig. 2. Weight functions of life stage model compared with
Suchey-Brooks pubic symphysis phases. The Suchey-Brooks dia-
gram was adapted from Müller et al. (2002), who applied the
examined Suchey-Brooks method on a known age sample. Our
life stage diagram (adapted from Roksandic et al., 2002) was
based on a test performed on the adult Coimbra sample. Note the
relatively uniform distribution of the four peaks in the life stage
model compared to the broad, variable distributions obtained
from the Suchey-Brooks method.
Furthermore, these stages carry substantially more pre-
dictable chronological age information—especially for the
adults—that can be used by demographers to build survi-
vorship curves.
According to Müller et al. (2002), constructing a paleo-
demographic profile by using estimates of age at death of
individual skeletons is not a correct approach. Rather,
one should estimate the probability of assigning skeletons
into a specific osteological age indicator stage. They pro-
posed the use of weight functions to relate age-at-death
to the assigned categories. By using this approach, they
have shown that nonparametric estimates of weight func-
tions of the six categories of the Suchey-Brooks pubic
symphysis method are poor estimators of age when
applied to a population other than the one that was used
as a reference sample for the model (Müller et al., 2002;
Fig. 1).
In 2000, the senior author scored the known age sam-
ple from the University of Coimbra that comprises 504
individuals ranging from 7 to 96 years, of which 462
were adults. ‘‘Weight functions’’ were calculated from
these scores to determine the correlation between age
and adult stage (Roksandic et al., 2002). When compared
with weight functions for the commonly used Suchey-
Brooks pubic symphysis method, these stages show
much higher probabilities of assigning age correctly (see
Fig. 2). It should be stressed here that our model was
not developed on the Coimbra population and thus
avoids the major criticism of Bocquet-Appel and Masset
(1982) that the method would a priori mimic the refer-
ence population.
From the point of view of a bioarchaeologist, it is not
necessary or, ultimately, even informative to assign chro-

American Journal of Physical Anthropology

 STAGES OF GROWTH AND SENESCENCE
nological age points to the beginning and the end of each
stage, as these are expected to vary both within and
between populations. However, as demonstrated by the
weight functions in Figure 2, the probabilities of certain
ages are much higher and more normally distributed than
in (at least one of the) other ageing techniques. The senile
category remains large, reflecting the variability of degen-
erative stages observed in any biological population.
Therefore, the model carries sufficient age information
that can be used to build mortality profiles by paleodemog-
raphers. Using the life stage model to build mortuary pro-
files still faces other criticisms proper to paleodemography
as a field, but it deals with one major issue: the reliance
on poor point age estimates. At the same time, this model
allows bioarchaeologists to discuss their findings by using
the same categories used in paleodemography, increasing
our ability to communicate across the fields.
CONCLUSION
Rather than trying to fit skeletal data into the
demands of point age estimates and cohort statistics,
paleodemography needs to be based in the reality of age-
associated information provided by skeletal assemblages.
We suggest that this can be accomplished by adjusting
the methods of age determination to the biological real-
ity of human growth and senescence. By using life stages
instead of chronological ages, this model can be easily
applied to any archaeological population, to allow for
comparisons between populations, while acknowledging
the inherent limitations of the assumptions made in
paleodemography. The model proposed here, using stages
based in the human life history pattern, systematizes
what has rapidly become a more common practice among
bioarchaeologists. The suggested stages of adulthood,
while not directly comparable to life history stages, use
single defined points of reference to which other skeletal
information can be calibrated within the studied popula-
tion. While admittedly far from perfect, it can serve as a
starting point for a discussion on the pattern of aging
that can be applied across human populations.
ACKNOWLEDGMENTS
The authors acknowledge that the comments by the
reviewers and editors have been instrumental in reconsi-
dering some points of the article; without them the arti-
cle would have been far more rudimentary and incom-
plete than in its current form. They are grateful to Mary
Silcox and Dejana Nikitovic (University of Toronto at
Scarborough) for their comments on the very first draft.
MR thanks James Vaupel of Max Planck Institute for
Demographic Research (Rostock, Germany) for the sup-
port for data collection at Coimbra; Eugenia Cunha, Ter-
esa Ferreira, and the colleagues at the Department of
Anthropology at the University of Coimbra; and Kon-
stantin Arbeev for the statistical analysis.
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