92 EVolutionary Anthropolog\~

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Evolutionary Anthropology 93

32 Dausser J. Colombani J (eds) (1972) Histocompatibility Testing 1972. Copenhagen:

Munksaaard.

33 Te~saki PI (ed) (1980) Histccompotibihty Testing J 980. Los Angeles; UCLA Tissue Typing Laboratorv,

34 Albert ED, Baur MP, MaYT WR (eds) (1984) Histocompatibility Testing 1984. Berlin: Springer-Verlag.

35 Tsuji K, Aizav ... 'a M, Sasazukl T (eds) (1992) HLA ./991, Proceedings of the. Elevmrh 'lnternalf0l1a.l Histocompatibilitv l-vorkslwp and COII- 1erel1Ce., I--blume 1. Oxford: Oxford University Press.

36 Baur MP, Neugebauer M, Albert ED (1984) ~efurencetablesoft\\'o.locusha.plotypefre.:::!uen· ClCS for all MHCmarker loci, In Albert Elj.Banr MP, Mays- WR (eels), Histocompatibility Testing 1984, pp 677-755. Berlin: Springer-Verlag,

37 Williams RC, Chen SN, Gill DK, Lane ITL,

Mcauley JE, Strothman R, Mitral KK (1989) Antigen Society #6 Report (Ell. B49, B\,,-'';O. BN2J, 811, B44. B4:S) In Dupont B (00), 1111- munobioiogyof Hl.A; t0l,wne- 1, Histcconrpaiibility Tasting 1987, pp 133~147T New York:

Springer-Verlag.

38 Wflliams RC, Chen SK:. McAuIcy JE, Richiardi P, D'Alfonso S, Mehra N, Mcf.almon RT. Tardif G, Strothman R, Knowlcr '\Ie, PettitL DJ (1992) Report of the Antigen Society lQ6, E21, B49, Bw50, BN2I, Bl2, B44J B45.ln TsujiK. Aizawa M. Sasazuki T(ecis},HL4.1991 Proceedings of the Ele.vfmth International Histocompatibility Workshop and Conference, H)[. umc l, pp 305-307. Oxford: Oxford University Press,

39 Hildebrand WHo Madrigal jA, Belich MP, Zemmour 1, Ward FE, Williams Re, Parham P (1992) Serologic cross-reactivities poorly reflect allelic relationships in the HLA-B12 and

~---------------------------'

1 GLOSSARY

I Bayes' Theorem~A classic theorem in

~ probability theory first pllblishe1~n 1 ~6?

I bv the Reverend Thomas Bayes, "" which states that the posterior probabilir:;·. is I proportional to the ?ro~uct of the prior ; probability and the likelihood.

\ Conditional probability-The pro b·

ability that an event (E:i) will occur if some previous event (Ej) has rurea?y occurred. U suellv written as p(E.' I EJ} and

l read as "the probability of Ej given Ei" or 'I· "the probability of Ej conditional on 'E,:' Gompertz function-One of the many distribution functions used to represent ! continuous age-at.death distributions. It , is a special case of the Weibl111 distribution (see below) with the shape parameter

equal to one.

Hazards analvsis- The study of haz\ ar'ds functions (instantaneous: con di- 1 tiona! failure rates], In dem?graphy, a

I hazard function is the probability that an individual at a particular age will die \ within an infinitesimally small period.

I, The age-specific probability of d~at,h from traditional life-table analysis IS analogous to the hazard function, al~ thouah the time intervals in a life table are I discr~te, and the approach is nonpara-

I metric. A!; applied in demography, he>. ards analvsis generally approximates the \ continuously distributed variable "age" ·1' with discrete, short time intervals such as.

weeks, months, or years.

Lilcelfhcod- The likelihood of a set of

parameters conditional on the observed \ data, which is usually written as L(B I x),

I is proportional to the probability of obtainine the observed data conditional On fixed ~alucs for the parameters. Thus, i 1.(91 x)~p(x 1 e Because likelihood 5 are

l' used rel.at.ive to one another (in rh.e SCD.se that we talk about certain values of parameters heingrr.cre likely than others). the arbirrarv constant of proportionality is of no immediate concern.

Joint probabilit), density function, or joint decsttv- The probability of cb-

HLA-B21 groups.T Immuncl149:3563-3568. 40 Kawaguchi G, Hildebrand vVH, Hiraiwa M, Karaki S, Nagao 1, Akiyama N, Uchida R, Kashiwase K. Akaza T, Williams ac, Juji 1'; Parham P, 'Iakiguchi M (1992) Tho'O subtypes of HL4~B51 differing by substitution at posilion 171 of the (lL helix. Immunogenetics 37:57-63.

41 Hill AVS, Allsopp CEM, Kwiatkowski D, Anstey NM, 'Iwumasi P, Rowe PA, Bennett S, Brewster D, McMichaeJ Al, Greenwood BM (1990 Common Vilest African HLA antigens are associated with protection from severe malaria. Nature 352:595--600.

42 Tiwari JL, Terasaki PI (1985) HVi and Disease Associations. New York: Springer-Verlag.

© 1<X>4 Wiley-l.s, Inc.

Pa[eodemography: "Not QUIte DeodH

LYLE W. KONIGSBERG AND SUSAN R FRANKENBERG

As Kim Hilll recently noted in Evolutionary Anthropology, humans are unique among the horrmolds with regard to the length of their lives, as well as other elements in the individual life histories. The evolutionary details that modified a basic pongid life bistory into a hominid one remain obscure, but aspects of recent human demographic history are assailable. Study of the last 10,000 years or so is an important part of ongoing anthropological discourse, for demographic changes may be intimately linked to such major developments as agriculture and urbanization.2-{! Whelher demographic changes are antecedents for or consequences of these major developments is a matter of great contention, but at the least we should attempt to document the nature of human demographic changes in the recent past. Allhough this documentation can take different forms, the pr.ncipa: sources are archeological information on past settlement patterns and analyses of prehistoric human skeletal material.

Lyle Konigsberg and Susan Frankenberc are, respectively. Assistant Protessor and Research Assistant Professor in the Department 01 Antnrooolopy at the Umverstty of Tennessee. KnoxvrHe. ln addition to peleooemcprapby, their res-earch interests include nistcnca! demography, U,S. Eastern Woodland ~rcheology. quantitative genetiC analysis, and human skeletal biology. Konlgsbe rg is . an associate ecttor tor the American

Journal of Physical Anthropology, the biological/physical anthropology editor for the Journal of Quantitative Anihrop%gy, and the associate edltcr for physical anthropology for the Tennessee AnthropologiSt,

Key words: Age estlmatlcn, hazards analysis, maximum ~ike!]hood

In a scene [TOm the film "Monrv Pvc thon and the Holy Grail," a mortallv wounded squire to one of King Idthurs knights remarks that he is "not quite dead," This statement is followed by other comments from the squire: "I think I could pull through" and "Actually 1 think I'm all right:' Similar statements might be made concerninz the general health of palecdemography Having sustained some crushing blows from various protagonists (see Box 1), paleodemography has begun to re-emerge as a vital field that can inform us about human evolutionary

histoJ.); and, indeed, is stronger now for haVing weathered the criticism s. Rather than having died at the hands of its critics, paleo demography is, like the wounded squire, "not quite dead." But we should not follow the example of the Black Knight in the same film, who cavalierly dismisses the loss of both arms as "just a flesh wound," We should not belittle the problems that still lie ahead to be solved in paleodemography

raining particular values fur two or mare variables. In the case of age and an indicator. the joint density is the probability that an individual will be it particular age and have a particular value of the indica-

lOfL:.-git regression-A particular lype of nonlinear regression in which the dependent variable is a probability or rate; therefore. Its estimate from the regression must be constrained between zero and one.

Marginal probability density function, or marginal -dellsity-The probahilttv of obtaining particular values of a single variable across all the values of one or-more otner va-iables. In the case of age and aJ1 indicator, the marginal density of age is simply the probability that an individual will be any particular age.

Prior probability-The unconditional probability of an event, hypothesis, OJ" set of parameters. For example, in paleodem ograpby the pr-ior probability that a randomlv selected individual from the reference sample died at a particular age is aiven by the marginal density of age.

-Poste~ior probability-The conditional probaoility of an event, hypotheses. or set of parameters. For example, in pap Iecdemography the posterior probability that a randomly selected individual from the target sample died at a particular ~ge is proportional to the product of the pnor probability of the age with the likeli~oo~ (from the reference sample) that an individual with the observed indicator data would be that particular age.

Rayleigh function-One of many distributionfunctions used to represent continuous aae-at-death distributions. It is a special case of the Weibull disrrtbution (see- below) with the shape parameter equal to two.

- \\ieibull function-A general distribution function used to represent conrinuou s ilge-atrdeath distributions. It is characterized by both shape and scale pnrameters (see Hastings and Peacockf").

THE NATURE OF ARCHEOLOGICAL SI(ELETAL SAMPLES

In a Panglossian world, all past members of a population could be recovered in archeolOgical samples. In the real world, this probably is never the case. Instead, a number of natural and cultural filters conspire to produce archeological skeletal samples that cannot be considered as random samples of all members of a population who died within a certain period.

. Natural filters consist of taphonomic processes, which may differentially affect the recovery of skeletons by age andsex9.IO Cultural filters include a di-

verse arrav of factors that can affect the repres~ntativeness of skeletal samples. Chief among these factors is differential mortuary treatment. Biased sex ratios may arise in skeletal samples because one sex is most likely to enter the archeological record in a place or manner that increases its chances of recovery Similarly, certain age groups may have been treated differently with respect 10 interment. Skeletons of infants frequently are under-represented in cemetery samples,' eitber because the bodies were placed elsewhere or because they are

less likelv to have been preserved. In rare instances, two or more spatially separate mortuary tracts may be recovered that are patterned by age or sex. This was true, for example at the Averbuch site near Nashville, Tennessee. Excavation of the three cemeteries associated with the village(Fig. 1 A) produced 852 skeletons, among which infants up to the age of one year we..""e substantially under-represented, whereas excavation of house floors in the village (Fig. 1 B) produced 35 skeletons, of which 27 were less than one year old." Figure lC shows the two

age-at-death distributions for the Averbuch samples (data from Berryman 12) and how the distribution of one sample appears to complement that of the oth cr. Unfortunately, in many archeological cases, we are left with only one segment of the burial

, population and the knowledge that

I, some people must be missing.

. In addition to the cultural factor> . active at the time of corpse disposal, II many other cultural factors can filter

the archeological record at later times. Indeed. archeological methods themi selves can substantially alter skeletal

I,· samples. For example, archeologists may neglect to locate and recover burial s completely or may choose to

) leave material in situ. Other cultural practices that can have nonrandom ef fects on skeletal samples are rcpatriaI tion and reburial. The effects of the

1 latter can be expected to increase in the ensuing years, given the current political and legal climate sur-round-

ling the excavation and curarion of archeologically reco-vered human skeletal material.

Because of the mvriad factors that can affect and ultimately distort the demographic profile of archeological skeletal samples, biological anthropologi sts who work with this material should be conversant in archeological method and theory, cultural anthropological theory, and such diverse areas as taphonomy and soil science. This directive is hardly novel, emanating as it does from the 'articulation between the "new archeology" of the 1960s and the developing methods of human skeletal biology," Unfortunately, there are no "magic bullets" that can be uniformly applied to remove the biases caused bv biological and cultural fil-

ters. Each case i; unique and needs to be treated as such. However, if we can assume that these myriad filters on the archeological record can, to some extent, be controlled, at least in a statistical sense, then the next task of paleodem ography is to esrablish the ages-at-death of skeletal remains.

DEATH AND THE SINGLE SKELETON

In a typical forensic ar.thropologv setting, the skeletal remains of a single individual are presented to an expert, whose task is to describe the r-emains

94 Evolutionary AnlhropoJogy

ARTICLE5

40 DV 60,'· STRUC1URE4 au 530' ..

170ff 210E S878

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Fig~rfl ~::;lhC~ogrc:p~~ of E=lx.colJctiOfl5::t the A!erbUCh SITe and comparison of oqe-ot-deoth distributions. The site was cxcovotoo in ~977 end 1978 b~ ,,:le.~pc. sment or Arohrcpcloqv 01 lr.e, Ur-lverstv of Tennessee 'lmd~r ccmobcrsce with the Arcnceczccrccr cnc Histortc Preservctlon ACT or 197.:1 ~.ktigCTlOn \1';05 necesscrv because O_f the ImP:=tct of housi~g CCP-STTUCTlon on the stte. which 'lies Iocotec in a suburbcr- subdlvsicn near Nash -II"' _~en~essee __ Av~r::\J~h WC~ a lcrqe ~alls.aced IVllssi.ssipp.!a:l.vllI~ge. the excav~tlon of whlch revecled three cemetery o-ecs wlth stone b~); graves ~n~ ..::2_0'Jm8Sh~ 51,1 L.; .... !urcs, :'\! ;hotag,cp.'" of one- or The ourlcls r-em 0 domestic structure. this burial oootonec the- rernos-s of on indiv-dl.:'CI .. , \:J~lr,!n The- fl~S' r:CIT year O~ l.fe. BJ Photocroof of cemetery). C} Cornporlson of ope-ot-oeorn profiles for skeletons from" AveJblJ~h ~emet~a ~I';~ 852) 0:10 vncce structures (n .:::: 35). Data are TokE-II trom Berrvrncn.tz . e es n -

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Evolutionary Anthropology 95

BoX 1. Assumptions in Age l:stimotion

Regressing age on an indicator to estimate the age-at-death in a forensic case requires the assumption that the forensic sample comes from the same age-atdeath distribution as does the reference sample. We prove this here, although we warn readers that some cumbersome notation is necessary, as is a basic knowledge of probability theory, calculus, and statistics. A number of the terms used here are defined in the glossary. In this proof, we wish to find the continuous distribu~on of age (x) for a single skeleton conditional on the observation that the skeleton has a value of y, for a continuous indicator. Using Bayes' Theorem, this conditional distribution is:

f(y, I x) 9 (x)

f If(y, I x) g(X) 1 dx

e-o

where fry, I x) is the conditional probability from the reference sample that an individual who is age x will have an indicator value of Ji. g(x} is the probability density function for age for the forensic skeleton, and the integration is across age until the maximum attainable age of w. Equation 1 can be expanded to:

f(x, Yi) (x) f(x) 9

p(xl y,)

J[fCX,y,) 9(X)]dX

""" f(x)

where fix} and g(x) are the respective probability density functions for age in the reference sample and for the forensic skeleton, and f(x,y,} is the joint density for age and indicator in the reference sample: In this context, g(x} represents the prior density for age; it is our best guess at the distribution of possible ages from which the individual was selected. This is the prior probability, with our guess made independent of any information aboutthe indicator value.

If we assume that the population from which the forensic SKeleton is derived has the same aqsatdcath distribution as th e reference sample, then g(x) = fix), so that lhese terms cancel in both the numerator and denominator. The integration in the denominator is then lor ajoint density across a marginal density, so x integrates out and the denominator is simply fiJi}. This leaves US wlth p(x1Y;} = fix, y,}/f(y,}, which is equal to f(xIJi}; that is, the distribution of age for a single skeleton equals the distribution from the regression of age on indicator in the reference sample. Thus, use of the regression of age an indicator in a reference sample to esti mate the age-at-death of a forensic case makes the implicit as surnption that the forensic case comes from fhe same aqe-at-ceath distribution as that represented by the reference sample.

If, instead, we assume that the forensic case comes from a uniform age-at· death distribution, then g(x} can be replaced with a constant (such as llw) , which can be factored out of the integral and ih en will cancel with this same constant in the numerator. This leaves the condluonal distribution of age on indicator in the target sample as:

P(XiYi)

Jf(Y;ix)dx

"""

The denominator represents a "normalizing" factor" that makes equation 3 a

true density function. Because f(y. Ix) is the regression of indicator on age,· use of this second method to estimate ages only makes the assumption that ages are uniformly distributed. This is a much weaker assumption. for it really is a

. statement of ignorance. We are sayi~g that without recourse to information from i the indicator we' cannot make any statement about the aqe-at-deam.

(1)

so that they can be matched to a missing individual. In addition ·to stature and "race," age and sex are distinguishing characteristics that limit the number of potential matches .. Even though forensic anthropologists are not inherently interested in the demographic structure of their case loads, researchers will estimate the age-atdeath and sex of remains to aid in identification. Derermining the sex for adult remains is not particularly problematic. In the absence of any external information such as age-at-death or circumstances of death, the odds of a correct determination are 1 :1. Further, the development of new techniques for sexing skeletal material from the presence of Y-chromosome mark. ers14.1.5 may provide near certainty in sex assessment for all individuals, reo gardless of age-at-death. Determination of aze-at-death is an entirelv different ';atter. There is considerabl~ variation, not only in how individual researchers determine age, but in how they choose to report it. Some give a simple paint estimate (e.g., the remains are those of a 47-year-old), some give a range (e.g., the individual was between 45 and 50 years old), and some give a standard error (e.g., the individual was 47 ± 2.59 years old).

First consider how a forensic anthropologist arrives at a point estimate or range of ages for individual remains. Usually, the researcher will use information on one or more age indicators relative to a reference sample of skeletons of known age in order to determine the age of the unknown skeleton. Ifthe indicator is disconrinuOUS, as is the case for cranial suture closure." the development of joint surfaces such as the auricular surface or pubic symphysis, 17.18 or dental for. marion and eruption.l? then an age range can be given for each stage in the sample of known age. This range can then be applied to the skeleton under STudy. For continuous indicators, such as the transparency of dentin in teeth 20.21 or the development of histological features in bone,22- it is usual to regress age on the indicator in the reference sample. then predict the age of the skeleton being studied. In either case, the researcher considers [he distribution of age to be conditional on the indicator information, assuming

(2)

(3)

96 Evolutionary Anthropology

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EvoluTIonory Anthropology 97

that for a fixed value of the indicator there is some distribution of age. Essentially; age is treated as a dependent variable and the indicator as an independent variable, so the causal mechanism is reversed. Regressing age on a continuous indicator in a sample of known age is opposite from the procedure human biologists use to study growth, which is to regress an indicator on age. Further, we can show that regressing age on an indicator to estimate ages in future forensic cases makes the critical, often dangerous assumption that future cases come from the same arre-at-dcath distribution as does the original reference material of lmown age (see Box 1).

Rather than he forced into assuming that a given skeleton comes from a particular age-at-death distribution, we consider the indicators as being conditional on age. We say that age causes growth and development, not that growth and development cause age. Although the mathematics of predicting ages for unidentified skeletons then become more complicated, the verbal description of this process is fairly straightforward. To predict age when indicators are conditional on age. we USE: Bayes' Theorem .. -8 classic theorem in probability theory first published by the Reverend Thomas Bayes" in 1763. In one of its simplest forms, Bayes' Theorem tells us that the posterior probability that an individual is a particular age conditional. on the observed indicator (the left side of the first equation in Box 1) is proportional to the prior probability that the individual is that particular age times the probability that an individual of that particular age would be in the observed indi cater state (i.e., the numerator in the first equation in Box!). The pro bability of an individual being a particular age conditional on the observed indicator is called the posterior probability because it is that obtained after-we have made OUT observation on the;;'dicator. The prior probability of that particular age is a statement about the possible age distribUlion for 2.: particular case made before we have observed the indicator. The probability that an individual at a particular age would be in the observed indicator SLate is called the likelihood of age conditional On indicator. The

Box 2, Controversies in Human Skeletal Biology

Mucha! lhe progress made in human skeletal biology during the past few decades has been the product ot numerous controversies regarding paleodemography. Bocquel-Appel and MasseF' initialed one such controversy in 1982, when they suggested that paleodemography be abandoned because age estima~on was too unreliable. Sattenspiel and Harpending'" started another a year later when they suggested that paleademography had more to say about fertility than about mortality. Both of these articles were met with a volley of critiques, 31--39 which made some valid poi nts but also tended 10 distort some issues. Only with the passage of time have the views surrounding these debates matured, the overall results being quite positive. Error in age estimation is now a widely acknowledged problem in paJeodemography, while the previous overemphasis on mortality interpretations is slowly being replaced with an appreciation for the role offertility. Tothese two·debates, as well as broader debates overthe development of human life history and the role of agricultu re in paleodemography, Wood, Milner, Harpending. and Weiss40 have added a new chaUenge to both paleodemography and paleopatholoqy. These authors have leveled serious and soberi ng crmcrsrns at current Interpretations of pa!eodemography and paleopathology, which human Skeletal biologists are only beginning to address.4'.42 If Wood and co-workers' challenge follows the same course as the previous debates in paleodemography, we can expect human skeletal biology to emerge with a stronger theoretical bent

c

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40 Age (Weeks LMP)

48

FigUI8 2. Age-at-dl;l-oth distributions ln weeks since the lost rneristtuol period (LMP) for o.jcrce co'ectcn of prehistoric and protohlstorlc tete' and neonatal Arkcro 1110iorYi. TtiS giOY b?l'S,n the background gi~le the ag;~at·deoth distribution from Ows'ey and Jantz,28 derived by us,ng published repressions of age _on long bone ~er:gth,97 !h-E.'. whlre bars ln the foreground g_lve the agent-decrlh dlstrlbufton derived by the maximum llkellnooc method. using a teqression of femur ienqth on gsta"iiono1 oge frcrn a large Ultrosound study,29 Forty weeks represent a 1erm mfont. Note th~t the origir.al age distrlbtmon from O~lsley and Jantz oppecrs 10 uocerestmote ages substantially wheel corrrpored to me maximum likelihood method

term likelihood here means that we are considering the distribution of age rather than the distribution of the indicator. For the single skeleton, there is no distrfbution of the indicator; there is only the observed indicator state.

In the forensic setting, age estimation Can be simplified because, in the absence of any outside information, the plio]' distribution for age is flat. In other words, without having some information from the ir.dicators, we have no preconceptions about the ageat-death, If we have some prior kno« .. 11- edge about the age distribution to be expected in forensic cases, then we will not have a flat prior distribution but, for the sake of simplicity we assume that we do not have the prior information. In such a case, the posterior probability that the individual died at a particular age conditional on the indicator is proponional to the probability that thev are in the observed indicate!' stat~ conditional on the particular age. Stated another way, the posterior probability is the likelihood of a particular age conditional on the observed indicator state. This makes age estimation a simple problem; we have only to select the age at which the probability of obtaining the indicator state observed in the uniden-

tified skeleton is the highest.

DEATH AND SKELETAL SAMPLES

solved by u sing a common statistical method known as maximum likelihood estimation. Just as we can write the posterior probability of age conditional on indicator for a single skeleton, we can write tbe joint probability of ages for a group of skeletons. This joint probability is the likelihood of the age-at -death distribution conditional on the observed indicator information from all skeletons in the group.

The maneuvers we have described are admittedly more complicated than the methods of age estimation typically used in paleodemograph), Traditionally, one would assign a point estimate or range for the age-at-death for each skeleton based on the conditional distribution of age on indicators ill a reference sample, then tabulate these deaths into a life table. Unfortunately, this form of age determination almost always leads to biased estimates ofthe ~ge-at-death distribution, as was pointed our first in the fisheries literature" and later in the human paIeodemography literature"·27 (see Box 2). This bias occurs because if we make aze conditional on indicators, then we~make the a priori assumption that skeletons in the archeological

Estimation of the age of a group of skeletons is more complicated because there is little reason to assume a flat prior distribution for age-atdeath. Neither the living population age distribution (i.e., the age "pyramid") nor the structure of mortality is uniform across age in any real population. As a result, the age-at-death distribution is not uniform, The typical undergraduate class project in demography of tabulating ages-at-death from gravestone inscriptions quickly demonstrates the non-uniform nature of age-at-death distributions. Thus, for any single skeleton from an archeological sample, the posterior probability that the individual is a particular age conditional on the observed indicator is proportional to the prior probability that the individual is that particular age times the pro bability that an individual that particular age would be in the observed indicator state. However, we do not know this prior probability. If we knew the prior probabilities of death at particular ages, then we would know the age-at-death distribution, which is precisely what we are trying to estimate. This paradox can be

sample follow the same age-at -death distribu tion as does the reference sample (see Box 1). As a simple example of the bias that can arise, Figure 2 presents the estimated age-ai-death

50

distributions for almost 500 perinatal skeletons originally reponed by Owslev and Jantz" and our analysis of the same sample. Owsley and Jantz used a regression of age in weeks on long bone length to estimate ages. whereas we calculated the age-atdeath structure using maximum likelihood and a regression of femur lenzth on a"e.'" The differences betw';-en the distributions are marked. Owsley and Jantz's study suggests that 88% of the remains are of preterm or term infants (usually defined as 40 weeks postmenstruation); our analysis indicates that only 17% are term or preterm infants. The average age-atdeath in the Owsley and Jantz study (38.5 weeks) is consequently about a month lower than in our study (42 weeks).

BIRTH, DEATH, AND SKELETAL SAMPLES

An additional complication in estimating the age-at-death distribution of a skel etal sample is thatthe age structure represented in a cemetery is affected by both deaths and births. For example, suppose that the infants and children in a population had generally adequate nutrition so that their mortality rate was very low. As a conse-

ffI s:

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'0

e OJ rn

C

~

(J.J 0...

50

Figure 3. Comporison of ope-ct-decta distributions for the Averbuch site- under ditferertt fert1l!i\' rates. The gmy ribbon in ihe background shows the one-ct-oeom dlstributlon assumhlQ that the birth and death totes were equol.(i,e .• popuioticn grolJJth rate was zero). The white ribbon. In the foreground shows the oge-otrde~th dlstributiort a~u-ni:1g thotfermty'haci incre:J~€ci to tbs e~ent that population gmll/th was 1.4% per.yeor, it apjJeO:ffi that the proportion ofirLdivlduas \'~O die ct young ages has mcreosed one the proportion dYH1g ot old ages nos decrecsed in the higheT fertmty population. even though mortolltvhcs remained unchanged

]00 Evol~ti"nory Anthropology

of human life-span, however defined, was a subject for debate even before the written word. Because the literature on Iife-spari is voluminous, we touch only tangentially on the paleodem agraphic evidence,

During the last 20 years, the question of the evolution of life-span has been addressed sporadically In the paleodemographic literature, but the main evidence has been rather indirect. Both Cutler" and Sacher'" argued that the maximum. human life-span has remained fairly.constant for at least the last 100,000 years. Their arguments were based on the empirical relationship between increased brain size and life-span across many modern mammalian taxa, together with the observation that the size of the human brain has not changed since the appearance of our species. More recently Allman et a1.6" have noted.a strong relationship between the brain weight and life-span of hominoids, again suggesting that modem human life-span was attained at the same point that modern brain size was attained.

Smith" has reviewed evidence that the rate of maturation and life-span in Australopithecus were probably more ape-like than human-like. She suggests that the modern human life-span was established only on the order of lens of thousands of years ago. Her research on the Nariokotorne Homo erectus skeletonP which had a pattern of dental emergence that was still apelike in some respects, but was closer to

. that of humans, supports her argument that the pattern of human life history was established by the emergence of Homo sapiens.

D~vid Smith63 summarizes the arguments on this subject, suggesting that the life-span of Australopithecus africanus was probably similar to that of modern apes-that is, about so years-and that life-span gradually doubled by the establishment of modern R01110 sapiens approximately" 100,000 years ago. Estimated life-span for humans now is between 80 to 100 years; the 01 dest verified age is slightly in excess of 120 years.55 The evolutionary causes fOT increases in life-span remain obscure, although methods as diverse as biochemistry, cell biology, and computer simulation of the de-

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Evolutionary Anthropology' 101

FigureS. The Shan~dar5 reconstructed cranium. tor which 0 histo!ogi::::ol ag-e at death estimate can be cclculcted by taking samples from ossocloted postcronlcl remains. Courtesy of and © Erik Tnnkous.

mography of J.R.R. Tolkeins elves= have been applied to the problem.

More direct evidence regarding the evolution of the human life-span comes &om recent research on Keandena] skeletal biology. After examining the bone histology of four Neandertals and com bing the literature for documented cases of senescent Neandertals, Tri nk aus and Thompson" commented on the "extreme rarity and possible absence of Neandertals greater than 40 to 4S years in the fossil record" (p. 128). Loth and Iscan= have discounted this finding, although they base their critique primarily on indirect evidence from the brain size literature. The bone histology work by 'Irinkaus and Thompson, and more recent work by Trinkaus, is crucial to the question of the evolution of the human life span, This corpus of work suggests that extensi on of an ape-like life history to that of a human life history did not occur until tbe advent of fullv modem Homo sapiens sapiens. Bone histological methods for age determination are generally fairly accurate and objective, though in the particular case that Trinkaus and Thompson present, the estimated ages are extrapolated beyond the limits of the reference sample data, and the confidence intervals

are quite wide for the estimated Neandertal ages-at-death (Fig, 5). For example, Shanidar 3, whom Trinkaus and Thompson list as being 4 I years old, has a 95% confidence interval around the estimated age, which extends from 24 to 59 years old.

Aside from the inherent difficulties in trying to determine ages-at-death from fossil remains, there is an additional complication. One problem with the search for senescent Neandertals is that larger samples than are available would be necessary in order' to identify such Individuals,

This raises another general problem with the concept of life-span, which Wood et al.48 have discussed in detail. "Life-span" refers to the extreme value from the distribution of ages-at-deatb and, like a range,68 is highly dependent on sample size. As samples increase in size, the potential for outliers increases. Consequently, small samples are unlikely to produce individuals who died at extremely old ages,

An example of this problem is shown in Figure 6, which summarizes a computer simulation of the maxi. mum observed age across samples of increasing size. For the conditions of this simulation, an upper age limit of 45 or more years is attained only 25% of the time when the sample contains 32 individuals more than 1 S years old. Trinkaus and Thompson note that l52 adult Neandertals are available for study, so the probability that one or mare of these individuals lived more than 45 years (if, indeed, this was possible) should be hirrh. On the other hand, many of the;e individuals are relatively incomplete and would often be missing areas of tbe skeleton which might be informative using traditional macroscopic methods for age determination.

If maximum human life-span has remained unchanged during modem human history (or ifwe lack the abili!) to show that it has changed), then has life expectancy at birth also remained constant? In theory, this should be a relatively easy question to answer=we would only need to !mow the average ages-at-death within paleodemographic samples arrayed across time. Unfortunately; it will be very difficult to document changes in life expectancy at birth in archeological samples. One

0,9

0.&

0.7

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a:. 0.4

0.3

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Age- (Years)

SOl

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1'"1

i:3 30 II

~ JI

20 -\-,---1 ---i-----------;------+-------+---------'

Sampl-e Size (Deaths c v eragc15 vears)

reason for this is the existence of biases in age estimation for paleodemographic samples; another is that fertility asserts a stronger influence than does mortality on the age-atdeath distribution."

The biases introduced by traditional methods of age estimation can produce substantial errors in estimates of the mean age-at -death. Although such biases cannot universally explain the many anomalies noted in paleodemographic life tables, theyalmost. certainly explain a general pattern of too few older individuals in Some archeological samples. As an example, Figure 7 contrasts the age-specific probability of death functions (or hazard rates) for Toqua Mounds A and B" (Fig. 7a) with that for GibsonKlunk (Fig. 7h).7° The age-ac-death distributions from these two sites

40

0.00 ~!-------~-----------,

i

0.051

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~ 1

~ c.oa I.

LJ.. 00021

65

,5

25

as 45

Age·a;t.Death (Years)

5.

.5

Fiqure 6. A simulotion of the effect of sample size- on the vc:lue of me observed maximum age-at-death. The top two ponelsshow,irom lett to right the prcocbntv density function tOT oce-ce-oectb and s.urvivorship slmulcted in this example. Ages-do-death greater than 15 years were slrnuloted uslnc a Rayleigh distr;butlon wlth scale po-orneter equal to 16.9. This produces a Ilte expectancy at age15 of 15 years (i.e .. a mean ope-ct-decrth of 30 years. for those who SurVlV8 to oge 1~ yec:rs). This is the lowest life expectancy from Weiss'98 series of :nodej life tables, The tttlrd (lower) panel shows the effec~ of sample szes on the obse~6:d rncxlrnum oge--{]t-deafh. The three n-es mdicote the peTceilt~e. VOlLJ8S out of 100,000 repllcctlcrs of the simulation at ecch sample sze. fOT exomple. the graph ShDWS -r-cc the 25th percentile vcjue for me .maxlmum agerat-death reaches ~5 veers once the sample ccntolns 32 deaths of those over the age o~ 15 veers. For the conditions of this sijmvlotion. the 1 Olh percenftle volue d09S not reach ;.';5 years across any of the sample sizes shown here, Ibislndlcctes that scruples of more then 50 lndivldua~ would be necesscry in orcer to obto'n a max~murn oge-ctdeath of 45 or more veers QO% of the time.

50

were identified in a previous analysis of 26 paleodemographic life tables [YOm prehistoric and protohistoric North American populations as having the greatest difference between them regarding possible underestimates of adult ages." The ages of the Toqua remains were primarily estimated by using a young reference sample of Korean Vifar dead. As a result, the ages-at-death for adults appear to be quite low. The ages of the Gibson-Klunk remains were primarily estimated bv usine a 2:enerallv older dissection 'room ~sa';;'ple, yi"elding adultages-at-deatb that are quite high as compared to Toqua. Figure 7 was' drawn usinz maximum likelihood estimation of ~ logit regression+-" to fit the hazards across age, with the observed numbers of deaths under one year and between one and five years in

each sample used at the beginning of the curves. The rapid rise in morrality Iorthe Toqua series is almost certainly a result of underaging due to the use of a young reference sample. Taken at face value, the Toqua life table indicates that the average age-at-death for those who survived to at least age 20 years was 29 vears: the comparable figure fo,Gibs~n~Klunk was 42 years. These calculations assume a maximum attainable age of 45 years for Toqua6• and 60 vears for Gibson-Klunk. 73

The influences on the mean age-atdeath can be equally perplexing in attempts to document increased life expectancy at birth. It is reasonable to assume that both fertility and mortali tv rates IDa\' have chanzed during ~odern hum~n history The developing consensus, however, is that dramatic increases in human life

98 Evolutionary Anthropology

ARTlCLE5

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Evolutjon~ry AnthropoloQV 99

tionship between mean age-at-deazli and birth rate.

quence, we would expect the mean age-at-death to be rather high. Allernatively, if we found a high mean ageat-death for a skeletal population, we might infer that rnortahtv among infa;15 and children was 10\;'. To this picture, add a high fertility rate, such that fertility exceeds mortality, so the population consequently gl-OWS through time. If we now examine the age-atdeath distribution assembled from skeletal data, the mean age-at-death will appear lower. Mortality did not change, so how did this shift in the mean occur? The shift is a result of differing cohort sizes. Old individuals in the death sample cam e from a smaller population whereas, because of population growth, young Individualscame from a larger population. For example, if we compare deaths before age one to d eaths between the ages of 50 and 51 years within a one year period, deaths under one year will be over-represented because they com e from a larger birth cohort than do the 50 to 51 -year-old group. At a modest growth rate of 1 .4% per year, the ratio of 50- year-old deaths to .newborn deaths would he only half that expected in a nongrowingpopulation. The effect of increased fertility on the age-at-death distribution is shown in Figure 3, which compares the age-at-death distribution for the Averbuch site assurning a stationary population and assuming a population growing at

1 .4% per year.

Although the effect or fertility on the . age-at-death distribution can be perplexing. we could adjust for this effect if we knew the population growth rate, which equals the crude birth rate minus (he crude death rate, or the change in census size per unit of time.11.3J,43 Hov ... -ever; because growth rates are generally unknown, researchers often assume a zero growth rate. Sattenspiel and Harpending30 showed that when the growth rate is unknown, the effect of fertility On the age-at-death distribution. is actually stronger than is the effect of mortality. Specifically, they suggested that the mean age-at-death is approximately equal to the inverse of the crude birth rate (lib). In our Scenario involving a decrease in the mean age-at-death with positive .population growth, it would thus be more reasonable to assume that fertility increased

than that subadult mortality increased. Subsequently, Horowitz and Armelagos-s claimed to have generated "a different but exact expression" (p. 1 YO) from that of Sattenspiel and Harpending and that. consequently, "a more predictable, 'nackable' relationship exists between age at death and life expectancy" (p. 192). Were this true} age-at-death distributions would be more reflective of mortality than of fertility.

Although the mathematical resolution of these issues is far outside the scope of this paper, we must point out that Sattenspiel and Harpending are correct in their statement that birth rate has a stronger relationship to mean age-at-death than does mortaJit), Sattenspiei and Harpending note in their appendix that themean age-at-

... birth rate has a stronger relationship to mean age-at-death than does mortality,

death is a function of the crude death rate, population growth rate, and the mean age in the living population. They give this relationship in a form that is algebraically identical to the expression ~that Hor;~'itz and Annelagos derive. If, given a population growth rate that is noi known but is not zero, we write the birth and death rates as functions of growth rate, mean age-atdeath, and mean age of the living, We find that the death rate is better estimated (han the birth rate. We also find that better estimation of the death rate is attained onlv when (he mean age-atdeath is more than two times the mean age of the living. However, this is a virtual impossibility: the mean age-atdeath will almost always be less than the mean age of the living. \Ve therefore reject Horowitz and AnneJagos' assertion of a stronger relationship between mean age-at-death and death rate in favor of Sattenspiel and Harpending's statement of a stronger rela

OF HAZARDS AND LIKELIHOODS

Two of the most recent dewlopments in paleodernographv the application of hazard models and the explicit use oflikelihood theory, promise to provide broader insights into the past. Both processes are mathematically rather demanding, but have sirnpIe intuitive appeal; it is at this level that we will consider them.

In hazards analysis, which has been extensively reviewed by Wood et al.,-'" a smooth function, the hazard of death, is fit to the mortalitv data contained within a life table. The primary advantage of hazards analysis is that it provides a smooth summary of the age-at-death data. while al the same time generating a small set of parameters that can be compared across different samples (see, for example, Gage's" study of modern census data and Whittington's" study of pre-hispanic Maya). However; hazards analysis does have a shortcoming, Many different parameters may be fit to the life table data, even though some may not have any ready bio]ogkal Interpreration (see, for example, Wood et al.'s48 discussion of the Gornpertz function) and others may not adequately represent the data.

Intimately related (0 the usc of hazards models is the increasing USe of maximum likeli hood method;r Or estimating paleodemographic parameters. Such methods have considerable antiquity in the statistical literature, as recounted in Appendix 2 to Edwnrds'w book on likelihood, and are often used to fit hazards models. Explicit use of likelihood theory in paleodemography is, however. relatively recent, havinz started only with Paine's" 1 989 pu bli"".: cation of a table-fitting method. This was followed by other applications of maximum likelihood theory to age estimation in paleodernograph)~z7,sl.'52

As mentioned earlier, the concept of likelihood is intimately related to that of probability. In much of statistical theory we try to find the probability that an observed data set could have arisen under some fixed hypothesis about a given parameter. For example, we might ask "What is the probabiiity that we coul d have obtained the ob-

Figure 4. Lcq-likellbood surroce against c~~e deeth rote er-e crude birth rate tor the Ubben slte.53 The log-likelihood is tbe log of the probcbmv of ob-omnn the observed age-at-de-ath distributio!l trom Ubben cDnditional an the crude birth and death totes.

served difference between two means if the tr-ue difference between the means was zero?" In this case, we consider the hypothesis as fixed and the observed data as variable. In the likelihood approach, we similarly deal with the probability that we would have obtained (he observed data under a given hypothesis, but now COnsider the data as fixed and the hypothesis as variable. For example, we could ask "What is the probability of obtaining the observed age-at-death data if the crude death rate was 0.05 and the crude birth rate 0.04?" We can then consider a multitude of alternative hypotheses about what the birth and death rates might have been. When comparing alternative hypotheses, it would be reasonable to choose the hypothesized rates at which the observed data are most likely or, in other words, the parameters at which the probability of obtaining the observed data is highest. We would (hen have found the maximum likelihood estimates of the parameters. Because the probabilities are often very small, and because it simplifies the mathematics, we usually search across likelihoods in a Iogarithmic scale and thus try to find the maximum log-likelih~od. The parameter values at the maximum likelihood on the log scale

will correspond to the parameter values at the maximum likelihood on the original scale. Figure 4 shows an example of a log -likelihood. surface for the Libben site" plotted against crude birth and death rates. The log-likelihoods were calculated as in Paine'" althouzh, rather than usinz model life table~, we used the observ;;d mortality profile for Libben (assuming zero population growth). then adjusted birth and death rates. The "death rates were adjusted by a logit regression, 54

Figure 4 shows that the fertility rate has a greater effect on the age-at-death profile than does the mortality rate. If a likelihood or Jog-likelihood surface is very flat, this tells us that we are having difficulty estimating a parameter. Intuitively, this makes sense-s-a completely flat surface tells us that the data are equally likely to have been generated by different values across the parameter space. Therefore, we want to find the maximum likelihood estimates of parameters (as the highest point in the log-likelihood surface); we also want the likelihood surface to fall rapidly from the maximum in all directions, The surface in Figure 4 looks much like a long east Tennessee mountain ridge, with the ridge running parallel to the crude death rate, Because the ridge runs roughly paral-

Iel to the axis for crude death rate, the peak of the ridge viewed across crude birth rate does not move much. Thus. if we did not kIlOW the growth rate (or; for that matter. crude death rate) we could estimate the crude birth rate. The converse is nottrue. If we look at the ridge across crude death rate, the ridge is fiat at the top. slightly convex in the foreground, and slightly conCave in the background. \~Ie therefore could not interPl"et the age-at-death distribution with respect to changes in the crude . death rate. This, which is precisely the point made by Sattenspiel and Harpending.P is borne out again here by the likelihood analysis.

WHAT DO WE KNOW ABOUT PALEO DEMOGRAPHY?

If our samples, our age estimates, and our demographic reconstructions are also biased, what do we know about past human demography? The answer is that although there is little that we 1o.10W with certainty \there is much that we can infer from the archeological record. Considerable effort in paJeodemographic research has focused myopically on the details of individual sites or small groups of Sites, often attempting to relate small changes in life-table parameters to changing su bsistence or health. These indtvidual case histories are interestinz in their own right, but it is at broader levels of analysis that we Can begin to see paleodemographic generalities.

Three far-reaching questions have emerged within the paleo demographic literature over the y-ears. The first is whether human life-span has increased; the second is whether life expectancy at birth has increased; and the third is whether demographic consequences or antecedents are related to the origins oHood production. An initial problem with the concept of life-span is its definition. Olshansky et aJ .55 define it as "the genetically endowed limit to life for a sin de individual jffree of all exogenou s risk factors" (p .. 635). However; current empirical results make a unitary geneti c basis for longevity highly unlikely.=>? Despite difficulties in defining lire-span, the term is conceptually ingrained in both the popular and scientific presses. We can assume that the limit

I D2 Evokrtlcncry Arrtnropolopv

ARTICLE§:

ARTICLES

Evolutionary Anthropology 1 C3

Toqua

Gibson-Klunk

Indian Knoll

Averbuch

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Hpure 7, Log~ reqresslcn smoothed hazard rctes" 1 tor A) Toquo69 orrd B) GilJs.orrKlullk.70 The two lines shown fer each graph are plus and minus tV10 standard errors around the estirnotes. Toque was 0 Missi:;sipPian village located in eost teroessee: Gibson-klunk wcs a Middle \,\Ioodland mour.c group in west-centrolfllnols. The contrast between these two graphs of hazard totes is not -thought to represent djffer~ng fertilITy or rnortostv. but the different methodoloples used to oetermlne ones-ot-deofn

CrowCreei<

expectancy at birth have occurred beginning only in the late nineteenth century 7':'~76 The increased emphasis on fertility (at the expense of mortalit)') in recent anthropological research, as in the work of Bentley et al.,77.1B makes it unlikely that we will find many ethnographic analogies that can be applied to interpret prehistoric mortality When we tum to the actual prehistoric record, it is clifficult to state definitively that life expectancy at birth has increased with increa~ed sedentism or the adoption of agri culture. In point of fact, Coben and Armelaaos"? arzued thatin most regions of the wo;ld 111r:::an age-atdeath .. which they used interchange. ably with life expectancy at birth, decreased with the adoption of agriculture. Although there is clear documentation of changes in the skeletally derived average age-at -deatl- at various points in the past, Sattenspiel and Harpendings+' insight tells us that these changes must first be int erpreted in terms of fertility, and only secondar.ly in terrns of mortality. Most researchers now allow for onlv modest changes, if any, in rriortali~ prior to the industrial revolution,'

The final question regarding the demographic consequences or antecedents related to the origins of rood production is the most difficult. If, in the past, mortality rates truly had littl c. relationship to agricultural produc-

tion, then we will need to consider fertility as a possibly more important force. Sattenspiel and Harpcnding.c? in their reanalysis of prehistoric and historic European life tables from Acsadi and Nemeskeris'" classic study, found that the mean age-at-death had increased through time. They thus concluded that the birth rate had declined. This finding is difficult to square with any notion that the agricultural transition led to high positive growth rates, either through increased fertility or decreased mortality However, Sattenspiel and Harpending noted that much of what is interpreted as rapid population growth with the advent of agriculture may instead represent an increase in population denshy because of aggregation around agricultural resources. In a later study restricted to North America, Bulkstra and Konigsberg" noted a contrast in life tables of pre-agricultural versus agricultural peoples, Specifically, the pre-agricultural peoples appeared to have mortality hazard estimates that rose more rapidlywith advancing age. As an example of this contrast, Figure 8 shows the hazard models for Indian Knoll (Fig. 8a)," a pre-agricultural Archaic site, and for Averbuch (Fig. Sb), a Late Mississippian agricultural site. The Averbuch "mortality" profile is much flatter against age. Why might this be?

If the data represented in Figure 8

rcallv tell us about mortality. the implicarion is that the agricultural lif estyle began to uncoupl e the risk of death from advancement in age. An extreme case in which risk of death does appear to be unrelated to age comes from the Crow Creek massacre site,8i where the risk of death across age is almost completely flat (Fig. 8e). In this case, however. the uncoupling from age is not the result of a preagricultural versus agricultural SU bsistence mode, but of the fact that the massacre represented a constant risk of death with respect to age.

Returning to the distinction between Indian Knoll and Averbuch, and using the insight from Sattenspie1 and Harpending. we should ask how fertility might affect the apparent "mortality" curves when we assume a zero growth rate. In high-fertility populations, younger deaths win be over-represented in the morta liry profile. leading to a flatter hazard curve (set Fig. 3). Consequently. the fJ atter curves Buikstra and Konigsberg noted for agriculturalists are in keeping with higher fertilitv populations. Again, because of the quality of the data we should be careful about interpreting thi s pattern too far; but in a more controlled setting Buikstra et al.'; provided some evidence for increased fertility with the adoption of full-scale agriculture in the prehistoric Lower Illinois Valley. Johansson and

40

c

Horov • .ritz3-7 also suggested that for a diachronic series from a single site in Central illinois, Dickson Mounds, the decreasing mean age-at-death with the adoption of maize agriculture indicates an increasing birth rate.

An increase in average number of "birth scars" on the pubic bones or at the pre-auricular sulci in adult females would be a useful corollary to the inferred increased fertility rates mentioned earlier. Unfortunately, the relationship between such osseous marks and parity among documented skeletal remains is weak at best,"·" so an analysis of prehistoric skeletal remains would be quite tenuous. More promising are studies that focus on the age of infants at weaning or the nature of the weaning process, which may serve as indirect measure of past fertility. For example, Bullingtons,:"

1 I

: .. : f

0.1 ~ 0.6 t o.e 1

0:' ~

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10

20 30

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..

Fig:.Jre 8. Lcgtt repression smoothed hazard rates for A) ~ndian Knoll. en Arc-hem: cemetery In Kentucky 81; B) /werbuch Cemeteries. from 0 Late Mississ:pplan villoge in centro' Tennessee 12 (see Figure l):C)Cro:·: Creek, the site of 0 tourteenth century massacre of a South Dakota Initlol ~oa~ escerr vi:lage.132 The contrcst between hazar~ rates fO: ,Indian Kno~, cr-d Averbuch is thought ta be 0 product of the hIgher fe:rti11ty r~tes at Aver· buch. Crow Creek controsts wrth 0:1 previous graphs In hovlng a hazard rote that ~5 vrtucuvurvejnted to age. presumably as 0 resclt or moseocre victims dy]ng rcocrcress ot their ages.

so

based on a study of rnicrowear on the deciduous den~ition, has suggested that the shift to azriculurre in the prehistoric Lower minois Valley occurred at the same time as the introduction of a softer weaning diet. This possibility reinforces previous suggestions~l3·,58 that a technological change in ceramic production concident with the development of agriculture in this region may have provided an easy means of cooking weaning gruels made from soft. starchy seeds. More rapid and easier weaning may have l_ed to shorter interbirth intervals, and thus to increased fertility This argument remains quite speculative; the relationship between weaning and fertility is complicated.

The relationship that the adoption or intensification of agriculture has to population pressure; [ertilitv. mortal-

iry, and pathology is obviously also quite complicated. We need to be especially careful of attempting to paint the broad history of humanity with the fine brush strokes provided by regional histories and prehistories_" We should expect that the generalities ",,11 be few and the exceptions great. What does seem to emerge repeatedly from the archeological record is a slight decrease in the~skeletally derived meanage-at-death with the adoption or intensification of agriculture.I" The former dogma was that such a decrease was a signal mark of decreasing quality of life and increasing mortality rates.79,89 This view is now being partially challenged by one that replaces the emphasis on increased monalilY with a'; emphasis on increased fertilin~37,S3,flB,90 In coming years, »se must b~ careful not to subscribe to [he

so

104 Evolutionary Anthrop"logV

wholesale adoption of a new dogma. A challenge to the prevailing view that agriculture necessarily leads to poorer quality of life has also recently been presented?" (see Box 2). In this instance, there seems little chance of a paradigm shift in the near future, for the challenge appears to have been dismissed by a few and ignored by many.

The swirl of activity, uncertainty, and controversy that now surrounds pal eodemography is a sign that the field is undergoing a healthy maturation. "With the relatively new appreciation of the problem s of age estimation, sample bias, and the complexity of relationships among fertility mortality, population growth, and life table analysis, we see pa1eodemography as just beginning to embark on what should be a truly productive phase of research. The addition of analytical techniques such as maximum likelihood estimation and hazards analysis to the traditional palcodcmographic arsenal can only strengthen Our ability to make inferences about past human life. Although we well appreciate the en tical comments directed towards paleodemographic rescarch,26,91-95 we believe that it is time to move bevond the methodological problems ar:d return to answering the interesting larger issues.

ACKNOWLEDGMENTS

We thank Jane Bulkstra, John Fleagle, and Hem"y Harpending for their constructive and copious comments on a previous drafi of this paper. We also thank Richard J am, for providing us with the original Owsley and Jantz data in machine-readable form, for commenting on a previous draft, and for encouraging us even \.Iihen our results deviated from some previous findings. The photographs of the Averbuch sile were provided by the McClung Museum (Fig. IE) and Walter Klippel (Fig. IA), whom we gratefully acknowledge. The Neandertal photograph (Fig. 5) was provjded by Erik Trinkaus, who also kindly provided a galley copy of his article on Neandertal mortality, \"hich will appear (undoubtedly prior to this article) in the Joumal of Archaeological Science. We thank him for his help, and strongly encourage readers of

this article to refer to his Journal of Archaeological Sciel1ce article as the definitive source on t-.:eandertal paleodemography.

REFERENCES

1 Hill K (1993) Life lustorv theory and evolutionary anthropology Evol Anthropol 2~78- 89

2 Dumond DE (197:5) The limitation of human population' A natural history, Science /87:713-711.

3 Cohen MN (989) Health and the Rise o{ Civilization. New Haven: Yale Univer-sitv,

4 Boserup E (1965) ThO'; Conditions or Agriculsural Gro·wtJ-L Chicago: Aldine Press.

5 Boserup E (1990) Economic mid Demographic Relationships itt Development. Baltimore: Johns Hopkins University Press.

6 Hand .. vcrker WP (ed) (1986) Culture and Reproduction: A11 Anthropological Critique of Demographic Transition Theory, Boulder; Westview Press.

7 Coale AJ (974) The history of human population. Sci Am 231:40-51.

8 frisch RE (J 978) Population, food intake, and fertility Science 199-:21-30.

9 Gordon ce. Buikstra JE H981) Soil pH, bone preservation, and sampling bias at mortuary sites. Am Antiq 46:566-57l.

10 Walker PL Johnson JR, Lambert PM (1988) Age and sex biases in the preservation afhuman skeletal remains. Am J Phys Anfhro-

po17tolS3-188. .

11 Moore J A. Swedlund AC, Armelagos GJ (1975} The use of life tables tn paleodemography In Swedlund AC (ed). Population Studies ill Arc/wJeoiDgJ.' and Biologic.al Anthropology: A Symposium, pp 57-70. Washington DC: Society for American Archaeology

12 Berryman HE (1981) Tj1e Averbuch skeleWI series: A study o-f biological and social stress at a Late Mississippian Period Site from Middle Tennessee, Ph,D_ Dissertation, University ofTennessee, Knoxville.

13 Buiksrra JE {]977) Biocultural dimensions of archaeological study: A regional perspective. In Blakey RL (ed). Biocultural Adaptation in Prehistoric .4.merica, pp 67-84. Athen.s GA: University of Georgia Press.

] 4 H.ummel S, Herrmann B (]99 n Y-d:u:'O· mosome-specific D!\A amplifi~d in ancient human bone. Naturv.'issenschaftcn 78:266- 267

15 Stone AC. Milner GR, Paabo S (1994) Sex delermj n;,>;~jOI1 of prt"hislOric human reruain.s usjng DNA analysisc Am J Phys Anthl"opol (Suppl) ) 80 18 S.

16 MasseT C (J 989) Ageestimation on the bas.is of crnnlal sutures. In ]scan MY (ed}, Age ,lL1al"lGers i" the, HUmQJt Skelezo{1, pp 71-103. Springfield: Chades C. Thomas.

17 KatzD, Suchey JM (J 9S6} Age deterrninari01:1 of lhema:!e os pubis. Am J Phys Anthropol 690237-244.

IS ·.M~indl RS, Lovejoy CO (1989) Age chang(.':s. in the pelvi.'i: Implications for pa· leodemogr.aphy_ln lscan MY {eci},Age Mm-ken iH the Human S}~letol1. pp 137-168. Spring-odd: Charles C. Thomas

19 . Saunders SR (1992) Subaduh skeletons and J!Towlh rcla!cd ~tudies. In Saunders S1<.., Katz;~nbt!r£ MA (cds), Sl,e.ie!a.l Siolog)' of Pasl Peoples; Res~wch Methods, pp 1-20. Nt':w"{otk:

John \o\iDc\' & Sons, !nc.

20 Dru.s~iAG (1991}_4ge-related changes in 1'001 n-ans]Jat"enc\' of Leeth in males and females. Am·] Hum 13ioI3:629-637.

21 DrusiniAG, Calhart I, VolpeA(199J)Root dentine transparency: Age determination of 'human teeth using computerized densitomet_ ric analysis. Am J Phys Anthrcpol 85~25-30. 22 StOUi SD (1992) Methods of determinina age at death using bone microstrucmre. I~ Saunders SR, Katzenberg MA (eds). Skeletal Biology of Pas! Peoples: Research Methods, PP 2]-35. New York: John Wiley & Sons, Inc.

23 Box GEP, TIao GC (1.992) Ba.vesian Inter. ence in Suuissical Ana~)'sis. Nev.' York John Wiley & Sons, . 24 Bayes TR(1763) An essay towards solvinn a problem in the doctrine of chances. Ph~ Trans Rov Soc London 53370--418,

25 \.Vest·rheim 5J, Ricker WE: (1978) Bias in using an age-length key to estimate age-Irequcncy distributions. J Fish Res Board Can 350184-189.

'26 Bocquet-Appel Jrp, Massa: C (1982) Farewell to palsodemography J Hum E\10J 11;321- -333.

27 Konigsberg LW, Frankenberg SR (1992} Estimation of ag~ structure in anthropological : demography. Am J Phys Anthropol 89:235-

156,

28 Owsley DW, Jantz RL (J985) Long bone lengths and gestational age distributions of Post-contact Period Arlkara Indian perinatal infant skeletons. Am J Phys Anthropol 6&321- 328_

29. Oman SD, \~.,rax Y (1984) Estimating fetal age by ultrasound measurements: An example of mu l rtvar-i ate calibration. Biometrics 40:947-960.

30 Sattenspiel L, Harpending H (1983) Stable populations and skeletal age. Am Antiq 480489-498.

3i Van Gerveu DP, Armelagos GJ (1983) "Farewell to palccdemography?" Rumors of i1S death have been greatly exaggerated. J Hum Evol12:353-360.

32 Greenc Df., Van Gerven Dl'; Armelagos GJ (1986) Life and death in ancien! populations:

Bones of contention in paleodemogrnphy Hum Evo11:193-207.

33 Buikstra IE, Konigsbe-rg LW (985) Palecdemcgraphy: Critiques and controversies. Am Anthrcpol 87:316-333.

34 Lanphear KM (1989) Testing the value of skeletal samples in demographic research: A campalison with "hal registration .samples. lot} Amhropo14:185-193

35 Mensforth RP (1990) Paleodemography of 1-hcCarlston Annis (Bt·5) Latr..:Archaic skeletal population. Am J Phys AnihrDpo182;81-99 36 Piontek l, Weber A (1990} Controvcrsv oI"J palcodemography. Int J Anlh:ropDI5~71-{3. 37 Johansson SR, Horowitz S (1986) Estimating mortality in .skeleu~l populations: In" fluence of the g:rowth ri3.t(C: on [he jnterpl-et3tian of lewIs and trends dUTing the tTamition to agriculmre_ Am J Phy~ Anth;opol 71~233-250.

. 3:8 Ho:r-owitz S, Armelagos G (1988) On generating birth rates from skeletal populations. Am.J Phys Anthropo) 76189-196

39 Jilckes M (1992) Paleodcrnography: Problems and techniques. In Saunders SR, Kaizcnberg YLA (eds), Skektal BiolugyofPas: Peoples: /Z<:.search Methods. pp 189-224. Ne\\..- York~ .1ohn \Viley & SQn~. Inc

40 \"-'ood JM, lVWner GR, Harpending He, \",1els~ Kil,rl (1992) The osteological paradox:

Problcm~ of inferring prehisLoric. health from skeIeuu s'"'-mples_ CUrT AnLhropo133:343-370. 41 Jades M (993) On paradox and osteol. ogy. Curt AnthroroI34':434--439.

42 Goodman AH (1993) On th~ int-C':rpreta- tian of heallh from skdetaJ remains. Cun- An-

ARTICLES

EvolutionaTY Anthropology 105

thropo134:2S1~288. ..

43 Bennett K {1973) On rhe e~u~<'tUo~ of some demographic charactcnstIc.s ~~v:.n deaths. by age. Am J Phys Anthropo139-~2.)~

~~2_ Wood JW, Holman DJ, weiss KM, Bucha[Jan AV, LeFor B {l992) Hazards models for hum<ln population biology. Ycerbk Phys An, tbropoI35~43-8i

45 GaE!e TB (198.8) Mathematical hazard models -of mortality: An alternative to model life lables.. Am J Phys Anthropol 76:419--441. 46 whittington SL (199]) Detection of significant demographic differences between subpop1l1atjons of orehispanic Mava from Copan, Honduras, by survival analysis. Am J Phys Anthmpo1850167-184.

47 G~l-'C'''' TB (I 991) Causes of death and the cornpo;;ems of mortality Testing the biological mterpretations of a competing hazards model. Am J Hum Bio13~289~300.

48 Wood J\\', 'Weeks SC, Bentley GR, Weiss KM (J 994) Human population biology and the evolution of aetna. In Crews De, Garrutc RM (eds). Biologic-=al Anthropology alld-Agilig.c Perspectives 0/1 Human l'al·iatiun ove~ t?ze L~fe Span, pp 19-75. New York: Oxford University Press.

49 Edwards A'NF (1992) Likelihood. Znd ed, Baltimore: Johns Hopkins University Press. 50 PaineRR(1989) Model life table fitting by maximum likelihood estimation: A procedure to reconstruct paleodemog:raphic ch aracteristics from skeletal age distributions. Am J Phvs Anthropol 79:51-56.

51 $jven CH (1991) On estimating mot-talities from osteological age data. Int.J Anthropol 6097-\10.

52 Skvtthe A, Boldsen JL (1993) A method for ccusrruction of standards for determination of skeletal age at death. Am] Phys Anthropol S 16:182.

53 Lovejoy CO, Meindl RS, Pryzbeck YR, Barton rs, Heiple KG, Kotting D (197i) f'aleodemography of the Libben Site, Ottawa County, Ohio. Science 198:291-293.

54 Brass W (1969) A generation method for projecting death 'rates. In Bechhofer F (ed}. Population GlVwth mw t.hc.Bl"ainDrail1. pp 75- 91 Edinburzh; Edinburzh Universitv Press. 55· Olshanskv·SJ, Cam~BA.Cffii$-elC(1990) In search ofMrethuselah; Estima~ingtheupper limlls 10 human longevjr.y. Science 250:634- 640

56 Pres.on SH (1993} De:mogr'dphic change in the United Stales, 1970~2050_ In Manton KG. Sin~erBH.Suzman R~'1 (edsLForecasling lh~ Health of Ekierly Popuiarions. pp 51-77. New Ymk: Springer-Verlag.

57 \n/eis~ K:\-l: (1993} Genetic Va,-iar{OH and H~/,nun Disei.l.Sc. New York: Cambridge Uni· \reTsit\' Press

58 CutlerRG (1 973) Evolutionofhurnan longevity and the genelic complexity governing agihg rate. Proc Nat Acad Sci 72 :4664-4668 . 59 SacherGA (1975) Matu:ratjon and longev• ity in rel2t1on to crani13.1 capaciLy in hominid evolution. In Tuttk RB (e:d), Primale hmclionai Motphology aHd EvolUliolZ,pp 4] 7--44]. The Hague~. Mouton:

60 AUmanJ, McLaughli1'l J, Hakkem-A (1993) Brain \'v'elghl 13.nd ~ife-spiill in pJ:imale splC:Oes. Proc ~at Acad Sci 90:] 18-122.

61 SmhhBH 0992~·UfehIStorY';;3.nd theevo· lution of human maTUration. Evol Anthropol 1:134-142.

62 SmIth BH (1993) The physio~ogi(.al ageof

KNM-WT ]5000_ln Walker A, Le~eyR(eds), The Nariokotcnu: Homo Erecius Skeleton, pp 195-220. Cambridge MA: Har-vard University Press

63 Smith D'\VE (1993) Human LongevilY· New York: Oxford Universit \' Press.

64 Williams GeJ Taylor FD (1987) Demographic consequences of natural selection. In Woodhead AD, Thompson KH {eds), Evolution oi Longevity ill Animals-, pp 235-245 _ New York Plenum Press .

65 "Irinkaus E, Thompson DD (1987) Femoral diaphyseal historncrphornetric age deter" min at ion s for th e Sharirda'r 3,4.5 and 6 Neandertals and Neandertal .longevity, Am J Phvs AnthroDo172:123-129.

66· Loth SR~ Iscan MY(1994) Morphological Indicators of skeletal aging: Implications for palcodemography and paleogcrcntology, In Crews DE, Garruto RM (eds), Biological .4.11- thropology aHd Aging: Perspectives on Human Variation over the Life SPLU1, pp 394-425. New YOTk: Oxford Universirv Press.

67 Thompson DD (19'79) The core technique in the detennhmtion of age at death in skeletons. J Porens Sci 24;902~91 s.

68 Foote M (1993) Human cranial variabilit)': A methodological comment. Am J·Phys Anthropo1900377-379.

69 Parham S (1982) A bioculrural approach to the skeletal biology of the Dallas people from 'Ioqua. M.A. Thesis, University of Tennessee, Knoxville.

70 Buikatra JE (1976}Hopel.1'ell in the Lo11J~J Illinois \!'UUc.y.- A RegiDnal Appmq.dt 10 the ~!Udy D[ Bwlog,~al Variability and Mortua.ry Actwity Evanston: Northwestern University Archaeological Program, Scientific Papers, No.2.

71 Efron E (1988) Logistic regression, sur"ivai analysis. and the Kaplan-Meier curve. J Am Stat Asscc 83:414--425.

72 Pennington R, Harpending B (1993) r:he Structure oi m~Afrh.:'Q/1 Pastoroliss Community:

Demography, History, cud Ecology of the Ngamiland Herero_ Nev.' York: Clar-endon Press.

73 Asch D (1976) The Middle Woodland POP11- lafiol1 of the Lower Illinois Valley; A. Study in Paleodemographic Me.thods. Evanston: Northwestern Archaeological Program,

74 Benjamin B (1986) 'I'he prospects for mor-tality decline and consequent changes in aee stTucture ofthc population. In Bittles AH, Collins KJ (eds), The Biolog'l' or Humall Agin,g, pp 133-J 54. New York: Cambridge Uni .. 'ersity Pr-e:;s.

75 Weiss KJ\,1 (1981) E\.'olutionary perSpectives On human ~ging. In Amoss PT, Harrell S (eds), Orher Ways of Growing Old, pp 25-58 Stanford~ Stanford Universit\: Press.

76 Wrigley EA (1969) PDp~lation an.d His· TOr-.:'. New York l\,1cGr~w-Hili.

7i Bentlcy GR. Jasienska G, Goldb~rg r (1993) Is thefertiliJ;:Y of agriculrura11s~ higher than that of nonag:ticulmraiists? Curr Anthropa134~778-785.

78 Bentley GR, Goldberg T. Ja.sienska G (1993)'The ferullty of agricultural mld nonagD T}cultur-al traditional socieIies. P-op Stud 47;269-.-281.

79 Cohen MN, Anneb£Cls GJ (1984) Paleo· pathology ilt the: origins of agdculture: Editor's sumIli3tion. In Cohen MK. Annelagos GJ {eds!. Paieopathology aT the Origi,-u; of Agricul· . lUre, pp 385--6-01_ Ne\\' York Academic Press. 80 Acsadl G, Neme;sk~ti J (1970) Hi~;lo1)-' of Ht[maJ1 Life Span and Mm-uiliry. Bud~pe;sc Akad€:miai Kie.d6.

81 Johnston FE, Snow CE. 096]) The reassessment of the age and sex of the Indian Knoll skeletal population: Demographic and methodological aspects. Am J Phys Anthropol 19:137 -244.

82 Zimmerman J, Emerson T, Willey p, Sweglc M, GTCgg J, Gregg. P, White E:, Smith C, Haberman T, Bumsted 11.1 (1981) TJII'; Croll' Creek Sire (39BF 11) Massacre: A Preliminary Report, Omaha: Omaha District. U .S, Army Corps of Engineers.

83 Buikstra IE. Konigsberg L\o\i, Bullington J (19&6) Fertility and the development of agriculture in the prehistoric midwest. Am Antiq 5}~528-546.

84 Holt CA (1978) A re-examination of parturition scars on the human female pelvis. Am J Phys Anthropo149:91-94.

85 Suchey JM. Wiseley DV, Green RF, Noguchi TT (1979) Analysis of dorsal pitting in the os- pubis in an extensive sample of modern American females. Am J Phys Anthropol 5J:517-S40.

86 Bullington J (I 988) Deciduous dental microwear in- Middle Woodland and Mississippian populations from the Lower Illinois River Valley Ph.D. Dissertation, Northwestern University Evanston.

87 Bullington J (199]) Deciduous dental microwear of prehistoric juveniles from the Lower Illinois River vallev Am J Phys Anthropo184059-73 c

88 Konigsberg LW. Buikstra JE, Bullington J (1989) Palecdemographic correlates of fertility: A reply to Corruccini, Brandon, ~d Handler and to Hoiland. Am Antiq 54 ~626--636.

89 Armelagos GJ (1990) Health and disease in prehistoric populations in transition. In Swedlund AC, Annda12:05 GJ (eds), Disease in Populations il11hmsflinH: Al1ti~rnpologicalal1d Epidemiological Perspectives, pp 127-144. New York: Bergin and Garvey.

90 Harris M, Ross EE (l987) Death. Sex, and Fen ilitv: Popula1foH Regula/ion in Preindusrrial and Developing Societies. New York: Columbia University Press.

91 Petersen W (1975) A demographer's view of prehistoric demography. Curr Anthropol J6:227-24S.

92JohanssonSR( I 982)The demographic histor'}' of the native peoples of North America: A sdective bibliography. Y~arbk Phy.s. Anthropol 25:133-152

93 HoweH N (1982) Village c-ornpos.ition im· plied by a pal~odemDgraphic life lable: The Ubben sitc_ Am J Phys Anthropo159:263-269. 94 Bocque,.Appel J-P, Ma"e[ C (1985) faleode:mography: Resurrec;:tion or ghost? J Hum Eml 14;107-111.

95 Bocquet-Appel J.p {1986) Once upon a tim~: Palaeodemograpby- Mitteilungen del' Berliner Geselhchah fUr AnthTopolgie. Ethnologie unci Urgeschjchte 7:117-133.

96 .Hastiof>:s NAJ. Peacock ffi (1913) Sta.tistical Di5tJib;;lion5.. New York; John W'iley & Sons

97 Scheuer lL, Musg:rave JR, Evans SF (1980) The estimation of late fetal ~nd peri· natal a>':':e rrom limb bone length by linear and logarithmjc regre.sslon. Ann Hum Bioli:257- 265.

98 \VeiSf; KM (J 973) Demographic Modds fur /1.1~thmp()logy. "''''as.hington. DC: Society for American Archaeology.

© 199"" WiJey-li:3s., Inc