Using the life history model to set the stage(s) of growth and senescence in bioarchaeology and paleodemography

Paleodemography, the study of demographic parameters of past human populations, relies on assumptions including biological uniformitarianism, stationary populations, and the ability to determine point age estimates from skeletal material. These assumptions have been widely criticized in the literature and various solutions have been proposed. The majority of these solutions rely on statistical modeling, and have not seen widespread application. Most bioarchaeologists recognize that our ability 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 ageing used in bioarchaeology. We propose that stages should be based in the human life history pattern, and their skeletal markers should have easily defined and clear endpoints. 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 recognizable 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 chronological ages will remain important and could eventually help us understand to what extent past societies differed in the timing of these life stages. Furthermore, paleodemographers 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.     

Problems in the aging of skeletal juveniles: Perspectives from maturation assessments of living children

We employ samples of children of known chronological age to demonstrate the significance of random and systematic effects on maturation in both dental and skeletal development. Differences between chronological age for dental age in young healthy Canadian children can be as much as 100% of the actual age of the children. For skeletal development by reference to Greulich-Pyle standards, three samples of known-age children from Mexico document parallel effects: 1) 183 six-year-old children have skeletal-based ages with a 95% confidence interval of 4–8 years; 2) 80% of 217 4.0–4.5-year-old children are underaged by 1–3 years; and 3) 130 children of skeletal age between 39 and 44 months are actually between 4 and 7.4 chronological years of age. The Mexican samples are drawn from a population living under conditions of environmental stress with chronic mild to moderate protein-energy malnutrition and moderate to high levels of infectious disease. These children may parallel those from the past, whose remains are studied by skeletal biologists or paleoanthropologists. Our findings reinforce concerns expressed in extant studies regarding the accuracy of age-at-death reconstructions. © 1996 Wiley-Liss, Inc.     

Life and death in ancient populations: Bones of contention in paleodemography

In a recent articleBocquet-Appel & Masset (1985) renew their criticism of paleodemography and criticizeVan Gerven & Armelagos' (1983) defense of paleodemographic methods. In the present analysis we respond toBocquet-Appel andMasset's criticisms ofVan Gerven andArmelagos and then address the question of whether paleodemographic methods are capable of producing plausible results based on what is known about contemporary human populations. This is done by expanding the life tables previously analysed byVan Gerven and co-workers for the Medieval Christian site of Kulubnarti in Nubia'sBatn el Hajar. The expanded results are then compared to data obtained from contemporary human populations as well as interpreted in light of Nubian culture history.     

Urban and rural differences in mortality and causes of death in historical Poland

The purpose of this paper is to document and interpret urban-rural differences in mortality in the past. To this end, we used data on mortality in Wielkopolska, Poland, in the 19th century and at the beginning of the 20th century. The data on mortality in rural areas (N = 1,173,910 deceased), small towns (N = 573,903 deceased), and Poznań, the capital of the Wielkopolska region (N = 86,352 deceased), were gathered from original Prussian statistical yearbooks (Preussische Statistik). Causes of death were also analyzed (rural areas, N = 449,576 deceased; small towns, N = 238,365 deceased; Poznań, N = 61,512 deceased). Mortality measures such as crude death rate (CDR), infant death rate (IDR), and neonatal and postneonatal death rates were calculated. Life tables were constructed for both stationary and stable population models and measures of the opportunity for natural selection calculated (Crow's index I(m), potential gross reproduction rate R(pot), and biological state index I(bs)). Relative frequencies of leading causes of death were computed. Stratification depending on the place of residence was evident in all mortality measures as well as in the values of the life tables and the measures of the opportunity for natural selection, but it was reverse of what is observed today in developed countries. In Poznań (a large industrial city), the mortality situation was the least favorable. It was caused by large population density, lack of water supply and sewage systems (up to 1896), and bad working conditions. The values of CDR ranged between 26.89-31.46, and IDR between 190.6-280.5. Newborn life expectancy (for a stable population model) was 31.6 years, I(m) = 0.79, R(pot) = 0.85, and I(bs) = 0.47. The most common causes of death were tuberculosis, other diseases of the respiratory and circulatory systems, dysentery and diarrhea, and cancer. These diseases were less common in rural areas, so they had the most favorable values of mortality measures (CDR between 22.87-27.32, IDR between 181.8-219.4, life expectancy of newborn e(0) = 42.12, I(m) = 0.55, R(pot) = 0.93, I(bs) = 0.60). Infectious diseases (other than tuberculosis), frailty at birth, and frailty in old age were the most frequent causes of death in rural areas. Small towns (population <20,000) had a mortality intermediate between city and rural areas.     

Reconciling “stress” and “health” in physical anthropology: What can bioarchaeologists learn from the other subdisciplines?

The concepts of “stress” and “health” are foundational in physical anthropology as guidelines for interpreting human behavior and biocultural adaptation in the past and present. Though related, stress and health are not coterminous, and while the term “health” encompasses some aspects of “stress,” health refers to a more holistic condition beyond just physiological disruption, and is of considerable significance in contributing to anthropologists' understanding of humanity's lived experiences. Bioarchaeological interpretations of human health generally are made from datasets consisting of skeletal markers of stress, markers that result from (chronic) physiological disruption (e.g., porotic hyperostosis; linear enamel hypoplasia). Non-specific indicators of stress may measure episodes of stress and indicate that infection, disease, or nutritional deficiencies were present in a population, but in assessing these markers, bioarchaeologists are not measuring “health” in the same way as are human biologists, medical anthropologists, or primatologists. Rather than continue to diverge on separate (albeit parallel) trajectories, bioarchaeologists are advised to pursue interlinkages with other subfields within physical anthropology toward bridging “stress” and “health.” The papers in this special symposium set include bioarchaeologists, human biologists, molecular anthropologists, and primatologists whose research develops this link between the concepts of “stress” and “health,” encouraging new avenues for bioarchaeologists to consider and reconsider health in past human populations. Am J Phys Anthropol 155:181–185, 2014. © 2014 Wiley Periodicals, Inc.     

Field population biology of citrus whitefly, Dialeurodes citri (Ashmead) (Heteroptera: Aleyrodidae), on oranges in California

Populations of Dialeurodes citri (Ashmead) on oranges were studied in two orange orchards in southern California, USA, to quantify developmental times and mortalities affecting this species. Four populations were followed in detail for three generations over 2 years, by monitoring the development and survival of 200 individuals in each population. Overall, preimaginal mortality among the generation studied varied from 56 to 94%. No evidence of disease or parasitism was present in the populations, and mortality was likely due to generalist predators. Weather records for the period were typical for the region, and periods of rainfall or elevated temperatures did not show any correlation with periods of mortality in the populations. Overall, the populations have high survival rates for whitefly populations, and this high survival is likely a major contributing factor to their pest status in California orchards. Addition of natural enemies to reduce the survival rates of these populations is proposed as the next course of action in suppressing populations below damaging levels.     

Warfare related trauma at Orendorf, a middle Mississippian site in west-central Illinois

Skeletal evidence of nonritual interpersonal trauma in the central Illinois valley is currently limited to the terminal prehistoric period in the region. Sixteen percent of the entire Norris Farms Oneota skeletal sample died violently, presumably because they intruded upon small groups of Mississippians who had not yet abandoned the region. Archaeological evidence of palisades, however, suggests that the region was embroiled in conflict before the Oneota arrived though the skeletal evidence supporting more than ritualized or geographically sporadic cases of scalping or embedded projectiles has been elusive. This study examines the frequency and nature of interpersonal trauma at Orendorf, a Middle Mississippian (AD 1150-1250) site at the northern periphery of the region. Nine percent (N = 25) of all 268 individuals documented at Orendorf suffered warfare-related trauma, including 13 cases of scalping, six instances of decapitation, five individuals with healed cranial blunt force trauma, three projectile point impacts, and eight cases of projectile injuries inferred by the burial context. All of the traumatized individuals were at or above the age of 15 years and males and females were victimized equally. The trauma rate among adults is 16%, which is less than that of the Norris Farms Oneota (34%) but higher than other Mississippian groups in the Southeast. The nature of the injuries is more consistent with attacks by outsiders than codified or ritualized intragroup violence.     

Brief communication: the collection of identified human skeletons housed at the Bocage Museum (National Museum of Natural History), Lisbon, Portugal

A large collection of identified human skeletons curated at the Bocage Museum (National Museum of Natural History, Lisbon, Portugal) has remained in relative anonymity since its collecting protocol was initiated in the 1980s. This collection originates from modern cemetery sources and is comprised of 1,692 skeletons with basic documentary data (age at death, place of birth, occupation, place of residence, and date and cause of death). At present, this information is more readily available for 699 individuals. The remaining 993 are in the process of being fully documented. The skeletons consist largely of Portuguese nationals who lived in the 19th and 20th centuries (1805-1975) in Lisbon. Both sexes are equally represented, and ages at death range from birth to 98 years, including 92 subadults (<20 years old).     

On estimating mortalities from osteological age data

A statistical method (based on the maximum likelihood principle) for estimating mortalities of skeletal populations is presented. The method can be applied both when osteological age groups are not overlapping as well as when they are. The results are presented as point estimates and as confidence intervals around these. The method is applied to two series from the Middle Ages: Westerhus and Helgeandsholmen.     

Deconstructing language by comparative gene expression: from neurobiology to microarray

Language is a defining characteristic of our species that has emerged quite recently on an evolutionary timescale. Understanding the neurobiological substrates and genetic underpinnings of language constitutes a basic challenge for both neuroscience and genetics. The functional localization of language in the brain has been progressively refined over the last century through studies of aphasics and more recently through neuroimaging. Concurrently, structural specializations in these brain regions have been identified by virtue of their lateralization in humans and also through comparisons with homologous brain regions in non-human primate species. Comparative genomics has revealed the genome of our closest living relative, the chimpanzee, to be astonishingly similar to our own. To explore the role that changes in the regulation of gene expression have had in recent human evolution, several groups have used microarrays to compare expression levels for thousands of genes in the brain between humans and chimpanzees. By applying this approach to the increasingly refined peri-sylvian network of brain regions involved in language, it may be possible to discern functionally significant changes in gene expression that are universal among humans but unique to our species, thus casting light on the molecular basis of language in the brain.     

Deconstructing language by comparative gene expression: from neurobiology to microarray

Language is a defining characteristic of our species that has emerged quite recently on an evolutionary timescale. Understanding the neurobiological substrates and genetic underpinnings of language constitutes a basic challenge for both neuroscience and genetics. The functional localization of language in the brain has been progressively refined over the last century through studies of aphasics and more recently through neuroimaging. Concurrently, structural specializations in these brain regions have been identified by virtue of their lateralization in humans and also through comparisons with homologous brain regions in non-human primate species. Comparative genomics has revealed the genome of our closest living relative, the chimpanzee, to be astonishingly similar to our own. To explore the role that changes in the regulation of gene expression have had in recent human evolution, several groups have used microarrays to compare expression levels for thousands of genes in the brain between humans and chimpanzees. By applying this approach to the increasingly refined peri-sylvian network of brain regions involved in language, it may be possible to discern functionally significant changes in gene expression that are universal among humans but unique to our species, thus casting light on the molecular basis of language in the brain.     

Expected life years compared to the general population

For cohorts with long-term follow-up, the number of years lost due to a certain disease yields a measure with a simple and appealing interpretation. Recently, an overview of the methodology used for this goal has been published, and two measures have been proposed. In this work, we consider a third option that may be useful in settings in which the other two are inappropriate. In all three measures, the survival of the given dataset is compared to the expected survival in the general population which is calculated using external mortality tables. We thoroughly analyze the differences between the three measures, their assumptions, interpretation, and the corresponding estimators. The first measure is defined in a competing risk setting and assumes an excess hazard compared to the population, while the other two measures also allow estimation for groups that live better than the general population. In this case, the observed survival of the patients is compared to that in the population. The starting point of this comparison depends on whether the entry into the study is a hazard changing event (e.g., disease diagnosis or the age at which the inclusion criteria were met). Focusing on the newly defined life years difference measure, we study the estimation of the variance and consider the possible challenges (e.g., extrapolation) that occur in practice. We illustrate its use with a dataset of French Olympic athletes. Finally, an efficient R implementation has been developed for all three measures which make this work easily available to subsequent users.     

The utility of osteon shape and circularity for differentiating human and non-human Haversian bone

Distinguishing human from non‐human bone fragments is usually accomplished by observation of gross morphology. When macroscopic analysis is insufficient, histological approaches can be applied. Microscopic features, like plexiform bone or osteon banding, are characteristic of non‐humans. In the absence of such features, distinguishing Haversian bone as either human or non‐human proves problematic. This study proposes a histomorphometric approach for classifying species from Haversian bone. Two variables, osteon area (On.Ar.) and circularity (On.Cr.), are examined. Measurements were collected from three species (deer, dog, human) represented by various skeletal elements; only ribs were available for humans (ribs: deer n = 6, dog n = 6, human n = 26; humeri: deer n = 6, dog n = 6; femora: deer n = 6, dog n = 6). Qualitative analysis comparing human to non‐human On.Ar. demonstrated that human ribs have larger mean On.Ar. (0.036 mm²) than non‐human ribs (deer = 0.017 mm², dog = 0.013 mm²). On.Cr. in the ribs showed minor differences between species (deer = 0.877; dog = 0.885; human = 0.898). Results demonstrated no significant difference across long bone quadrants in long bones. Discriminant analyses run on the means for each sample demonstrated overlap in deer and dog samples, clustering the non‐human and human groups apart from each other. Mean On.Cr. proved a poor criterion (ribs only: 76.3%, pooled elements: 66.1%), while mean On.Ar. proved useful in identifying human from non‐human samples (ribs only: 92.1%, pooled elements: 93.5%). When variables were combined, accuracy increased to 100% correct classification for rib data and 98.4% when considering data from all elements. These results indicate that On.Ar. and On.Cr. are valuable histomorphometric tools for distinguishing human from non‐human Haversian bone. Am J Phys Anthropol, 2012. © 2012 Wiley Periodicals, Inc.     

Marginal attack rate,k-values and density dependence in the analysis of contemporaneous mortality factors

1. Procedures for calculating the marginal attack rates and associated k-values from observed death rates are presented for contemporaneous predators or parasitoids that either do or do not discriminate against previously parasitized hosts.
2. The relationships among previous methods for quantifying contemporaneous mortalities are discussed.
3. Solutions to the problem of calculating marginal rates taken from human survival analysis are given under the assumption of proportional hazards. These calculations are easily extended to three or more contemporaneous sources of mortality.
4. The conditions under which proportional hazards can be expected are discussed.
5. These methods are compared to that of Varley et al., 1973 for assessment of density dependence among contemporaneous predators.

     

Critical periods in the life cycle and the effects of a severe spate vary markedly between four species of elmid beetles in a small stream

1. The chief objectives were: (i) to describe quantitatively the life cycles of four species of Elmidae, Elmis aenea, Esolus parallelepipedus, Oulimnius tuberculatus and Limnius volkmari; (ii) to use life tables to identify critical periods for survival in the life cycle of each species; (iii) to evaluate the immediate and longer‐term effects of a severe spate on densities of the four species. Monthly samples were taken over 63 months at two contrasting sites in a small stream: one in a deep section with macrophytes abundant, and the other in a shallow stony section. 2. There were five larval instars for O. tuberculatus, seven for L. volkmari and six for the other two species. The life cycle of each species took 1 year from egg hatching (chiefly in June for E. aenea and O. tuberculatus, and July for the other species) to pupation in the stream bank and a further year before the adults in the stream matured and laid their eggs. Mature adults were present in most months, but were rare or absent in January and February and attained maximum densities in April for O. tuberculatus and May for the other species. 3. Laboratory experiments provided data on egg hatching and pupation periods and the number of eggs laid per female. Life tables compared maximum numbers per square metre for key life‐stages. Within each species, mortality rates between adjacent life‐stages were fairly constant among six cohorts and between sites, in spite of large differences in numbers. The only exception for all species was the high adult, but not larval, mortality during a severe spate. 4. Standardised life tables, starting with 1000 eggs, identified key life‐stages with the highest mortality, namely the early life‐stages for E. aenea (36% mortality), start of the overwintering period to pupation for O. tuberculatus (41%) and L. volkmari (51%), start of pupation to the maximum number of immature adults for E. parallelepipedus (41%) and between the maximum numbers of immature and mature adults for O. tuberculatus (41%). Therefore, critical periods for survival in the life cycle differed between species, presumably because of their different ecological requirements. Similarly, the effects of the spate on adult mortality, and hence egg production, varied between species, being most severe and long‐term for E. aenea and O. tuberculatus, less severe for E. parallelepipedus and least severe with a rapid recovery for L. volkmari. Possible reasons for these discrepancies are discussed, but more data are required on the food and microhabitat requirements of the elmids before satisfactory explanations can be found.     

Geometric morphometric analysis of mandibular ramus flexure

Many characteristics of the human skeleton can only be assessed morphologically, which may be problematic due to factors such as interobserver error and difficulties with standardization. Flexure of the mandibular ramus is one of these traits, and various researchers found widely differing results using this morphological feature. The aim of this study was to determine whether differences between male and female mandibular rami could be observed using the computerized method of geometric morphometrics, a valuable tool that helps quantify shape differences. Twenty-eight mandibular rami of black females and 43 of black males were photographed in a standard plane and assessed. It was found that the females were more scattered on the graph (more variable in shape), while the males clustered more around the center point where the two axes met (shape more constant). There was, however, considerable overlap between the sexes. Although different tendencies exist between the rami of males (being more flexed) and females (tending to be straight), the extent of these differences is not adequate to predict the sex of a single individual.