Age-at-death estimation in an Italian historical sample: A test of the Suchey-Brooks and transition analysis methods

A growing body of research is demonstrating increased accuracy in aging from a relatively new method, transition analysis. Although transition analysis was developed for paleodemographic research, a majority of subsequent studies have been in the forensic arena, with very little work in bioarchaeological contexts. Using the Suchey‐Brooks pubic symphysis phases, scored on a target sample of historic Italians from the island of Sardinia, we compare accuracy of aging between transition analysis combined with a Bayesian approach and the standard Suchey‐Brooks age ranges. Because of the difficulty in identifying a reasonable informative prior for bioarchaeological samples, we also compared results of both an informative prior and a uniform prior for age estimation. Published ages‐of‐transition for the Terry Collection and Balkan genocide victims were used in conjunction with parameters generated from Gompertz hazard models derived from the priors. The ages‐of‐transition and hazard parameters were utilized to calculate the highest posterior density regions, otherwise known as “coverages” or age ranges, for each Suchey‐Brooks phase. Each prior, along with the parameters, were input into cumulative binomial tests. The results indicate that the Bayesian approach outperformed the Suchey‐Brooks technique alone. The Terry Collection surpassed the Balkans as a reasonable sample from which to derive transition analysis parameters. This discrepancy between populations is due to different within phase age‐at‐death distributions that reflect differences in aging between the populations. These results indicate bioarchaeologists should strive to apply a Bayesian analysis when aging historic and archaeological populations by employing an informative prior. Am J Phys Anthropol 149:259–265, 2012. © 2012 Wiley Periodicals, Inc.     

Senescence in European Middle Pleistocene hominids: The Atapuerca evidence

The mandibular specimens AT-250 and AT-793 (individual IV) and AT-888 (individual XXI) from the Atapuerca Middle Pleistocene sample show a combination of traits interpreted as evidence of senescence. In order to explore age-related mandibular changes relevant to the interpretation of the Atapuerca hominids, the sample of known age and sex from Spitalfields (London) and the Neolithic sample from Abu Hureyra (Syria) were used as a baseline. Results obtained indicate that the occurrence of an enlarged and posteriorly located mental foramen, in association with high alveolar resorption in AT-250 and AT-888, support the hypothesis of senescence in individuals IV and XXI from the Atapuerca sample. The findings suggest that European Middle Pleistocene populations lived long enough to reach senescence. The age at death for Middle Pleistocene hominids and Neanderthals has previously been estimated at around 40–45 years in the oldest specimens. The appearance of senescence processes before this age is indicative of higher rates of morphological aging in the clade which gave rise to Neanderthals. The anatomical significance of ageing features when considered in the context of craniofacial remodelling can help in the understanding of the taxonomic status of these morphological traits.     

REPRODUCTIVE PATTERNS OF THE HAWAIIAN MONK SEAL

We evaluated reproductive patterns of the Hawaiian monk seal ( Monachus schauinslandi ) using a combination of fitted age-specific reproductive curves and analysis of reproductive patterns of individual females. We review the difficulties inherent in the acquisition and modeling of reproductive data with emphasis on the significance of reproductive senescence to populations with dissimilar age/sex compositions. Validation of the fitted reproductive parameters was accomplished by Monte Carlo sampling of parameter distributions to compare the expected number of pups with the observed production. Although the fitted reproductive functions appear to provide an acceptable fit to the raw reproductive data, we found that the fitted curves did a poor job of predicting the actual pup production in individual years because of high variability among years. To further verify, and elaborate on, the patterns in the pooled (multi-seal, and multi-year) rates, we examined attributes of the reproductive performance of individual seals. The attributes included age of primiparity, reproductive rates computed over several age ranges, and the relationship between reproductive performance and seal longevity. Analysis of individual seal patterns reinforced the conclusion that reproductive senescence is operative in monk seal populations.     

Haematological parameters do senesce in the wild: evidence from different populations of a long‐lived mammal

Increasing evidence of senescence has been reported from long‐term studies of wild populations. However, most studies have focused on life‐history traits like survival, reproduction or body mass, generally from a single intensively monitored population. However, variation in the intensity of senescence across populations, and to a lesser extent between sexes, is still poorly understood. In addition, the pattern of age‐specific changes in haematological parameters remains virtually unknown to date for any population of vertebrate living in the wild. Using repeated blood samples collected from known‐aged (2–15 years of age) roe deer (Capreolus capreolus) from two populations facing highly different environmental conditions, we filled the gap. In particular, we investigated age‐specific changes in haematocrit, albumin and creatinine. We reported clear evidence of senescence in all haematological parameters. Moreover, senescence patterns differed between sexes and populations. The rate of senescence was higher in males than in females for haematocrit with no site difference. On the other hand, the rate of senescence in creatinine was higher at Trois Fontaines than at Chizé with no sex difference. Our findings provide a first demonstration of age‐specific declines in haematological parameters in wild populations of large herbivores and show that the process of senescence in vertebrates is not restricted to body mass or fitness components. We also demonstrate that the senescence pattern of haematological parameters is context dependent and varies both between sexes and according to environmental conditions.     

Gene expression during two alternative pathways of ovary development in Pisum sativum: fruit development and ovary senescence

Pea ovaries are induced to enter a fruit development pathway involving physiological and morphological changes by pollination or application of plant growth regulators. In the absence of these stimuli, overies stop growing and enter an alternative pathway of senesecence that leads to their degeneration. We have used two dimensional polyacrylamide gel electrophoresis in search of molecular changes underlying fruit development and ovary senescence at the level of total accumulated proteins, newly synthesized proteins, and translatable, RNA populations. We have found changes in gene expression during the processes of ovary formation and ovary senescence. Stimuli that induce fruit set do not appreciably alter the overall patterns of synthesized proteins or translatable RNAs, indicating that fruit development is apparently a natural continuation of ovary formation. However, ovary senescence is an alternative pathway that involves the presence of new RNA messengers and proteins as well as the disappearance of others. These changes were detected earlier than any morphological or structural changes could be observed in the ovary.     

Reconstructing the evolutionary history of an endangered subspecies across the changing landscape of the Great Central Valley of California

Identifying historic patterns of population genetic diversity and connectivity is a primary challenge in efforts to re‐establish the processes that have generated and maintained genetic variation across natural landscapes. The challenge of reconstructing pattern and process is even greater in highly altered landscapes where population extinctions and dramatic demographic fluctuations in remnant populations may have substantially altered, if not eliminated, historic patterns. Here, we seek to reconstruct historic patterns of diversity and connectivity in an endangered subspecies of woodrat that now occupies only 1–2 remnant locations within the highly altered landscape of the Great Central Valley of California. We examine patterns of diversity and connectivity using 14 microsatellite loci and sequence data from a mitochondrial locus and a nuclear intron. We reconstruct temporal change in habitat availability to establish several historical scenarios that could have led to contemporary patterns of diversity, and use an approximate Bayesian computation approach to test which of these scenarios is most consistent with our observed data. We find that the Central Valley populations harbour unique genetic variation coupled with a history of admixture between two well‐differentiated species of woodrats that are currently restricted to the woodlands flanking the Valley. Our simulations also show that certain commonly used analytical approaches may fail to recover a history of admixture when populations experience severe bottlenecks subsequent to hybridization. Overall our study shows the strength of combining empirical and simulation analyses to recover the history of populations occupying highly altered landscapes.     

Test of the Lamendin aging method on two historic skeletal samples

The Lamendin aging method involves the quantification of root translucency and the attachment position of the periodontal membrane. It was developed using recent medical-examiner specimens, and was tested on modern skeletal samples such as the Terry Collection (Lamendin et al. [1992] J. Forensic Sci. 37:1373-1379; Prince and Ubelaker [2002] J. Forensic Sci. 47:107-116). The method may be one of the most useful for estimating age after the mid-30s. The current study is an evaluation of the Lamendin criteria on two historic skeletal samples from Britain. Both the Christ's Church Spitalfields and St. Bride's Church collections represent documented skeletal samples that were interred in the 18th and 19th centuries. In total, 1,188 teeth from 220 adult individuals were examined from these two collections. The Lamendin method requires measuring total root length (cementoenamel junction to apex), gingival regression (cementoenamel junction to periodontal ligament attachment), and root translucency (root apex to maximum level of root translucency) on the labial surface of single-rooted teeth. Our results indicate that postmortem factors affect the applicability of the Lamendin technique to archaeological and historical samples. In particular, root translucency disappears with time, or is obscured by unknown postmortem taphonomic effects related to the length of interment or postmortem environment. Thirty-five percent of our sample showed no root translucency, indicating that caution is required when applying this method to archaeological or historical remains. The mean error of age estimates for Spitalfields and St. Bride's was higher than in the original study of Lamendin et al. ([1992] J. Forensic Sci. 37: 1373-1379), and higher than in the test by Prince and Ubelaker ([2002] J. Forensic Sci. 47:107-116) of the Lamendin method on the Terry Collection.     

Invasive and Non-Invasive Congeners Show Similar Trait Shifts between Their Same Native and Non-Native Ranges

Differences in morphological or ecological traits expressed by exotic species between their native and non-native ranges are often interpreted as evidence for adaptation to new conditions in the non-native ranges. In turn this adaptation is often hypothesized to contribute to the successful invasion of these species. There is good evidence for rapid evolution by many exotic invasives, but the extent to which these evolutionary changes actually drive invasiveness is unclear. One approach to resolving the relationship between adaptive responses and successful invasion is to compare traits between populations from the native and non-native ranges for both exotic invaders and congeners that are exotic but not invasive. We compared a suite of morphological traits that are commonly tested in the literature in the context of invasion for three very closely related species of Centaurea, all of which are sympatric in the same native and non-native ranges in Europe and North America. Of these, C. solstitialis is highly invasive whereas C. calcitrapa and C. sulphurea are not. For all three species, plants from non-native populations showed similar shifts in key traits that have been identified in other studies as important putative adaptive responses to post-introduction invasion. For example, for all three species plants from populations in non-native ranges were (i) larger and (ii) produced seeds that germinated at higher rates. In fact, the non-invasive C. calcitrapa showed the strongest trait shift between ranges. Centaurea solstitialis was the only species for which plants from the non-native range increased allocation to defensive spines, and allocated proportionally less resources to reproduction, patterns contrary to what would be predicted by theory and other empirical studies to enhance invasion. Our results suggest caution when interpreting the commonly observed increase in size and reproductive capacity as factors that cause exotics to become invaders.     

Age‐related variation and temporal patterns in the survival of a long‐lived scavenger

Although senescence has been described for various fitness components in a wide range of animal species, few studies have studied senescence in long‐lived species, and little is known about its interactions with varying environmental conditions. Using a 32 year capture–mark–recapture dataset on the griffon vulture Gyps fulvus, we examined the demographic patterns of actuarial senescence and the patterns of year‐to‐year variation in survival rates. We found a significant, surprisingly late, decrease of annual survival probabilities from the age of 28 years onward and divided individual lifetimes into to three categories (juvenile, mid‐age and senescent birds). In agreement with the environmental canalization hypothesis, our analyses uncovered 1) higher temporal variation of annual survival probabilities in both the juvenile and senescent age classes compared to the mid‐age class and 2) low sensitivity of the population growth rate to the survival of both the juvenile and senescent age classes. Our results further suggested that the temporal variation in the survival of senescent birds might be related to intra‐annual changes in air temperature amplitudes. Finally, using population dynamics modeling, we revealed contrasting effects of the inclusion of the senescent age class on predicted population growth, depending on how survival rates were modeled. Altogether, our results demonstrate the existence of a class of senescent birds that exhibit distinct demographic properties compared to juvenile and mid‐age classes.     

Evidence for senescence in survival but not in reproduction in a short‐lived passerine

Senescence has been studied since a long time by theoreticians in ecology and evolution, but empirical support in natural population has only recently been accumulating. One of the current challenges is the investigation of senescence of multiple fitness components and the study of differences between sexes. Until now, studies have been more frequently conducted on females than on males and rather in long‐lived than in short‐lived species. To reach a more fundamental understanding of the evolution of senescence, it is critical to investigate age‐specific survival and reproduction performance in both sexes and in a large range of species with contrasting life histories. In this study, we present results on patterns of age‐specific and sex‐specific variation in survival and reproduction in the whinchat Saxicola rubetra, a short‐lived passerine. We compiled individual‐based long‐term datasets from seven populations that were jointly analyzed within a Bayesian modeling framework. We found evidence for senescence in survival with a continuous decline after the age of 1 year, but no evidence of reproductive senescence. Furthermore, we found no clear evidence for sex effects on these patterns. We discuss these results in light of previous studies documenting senescence in short‐lived birds. We note that most of them have been conducted in populations breeding in nest boxes, and we question the potential effect of the nest boxes on the shape of age‐reproductive trajectories.     

Social ageing: exploring the drivers of late-life changes in social behaviour in mammals

Social interactions help group-living organisms cope with socio-environmental challenges and are central to survival and reproductive success. Recent research has shown that social behaviour and relationships can change across the lifespan, a phenomenon referred to as ‘social ageing’. Given the importance of social integration for health and well-being, age-dependent changes in social behaviour can modulate how fitness changes with age and may be an important source of unexplained variation in individual patterns of senescence. However, integrating social behaviour into ageing research requires a deeper understanding of the causes and consequences of age-based changes in social behaviour. Here, we provide an overview of the drivers of late-life changes in sociality. We suggest that explanations for social ageing can be categorized into three groups: changes in sociality that (a) occur as a result of senescence; (b) result from adaptations to ameliorate the negative effects of senescence; and/or (c) result from positive effects of age and demographic changes. Quantifying the relative contribution of these processes to late-life changes in sociality will allow us to move towards a more holistic understanding of how and why these patterns emerge and will provide important insights into the potential for social ageing to delay or accelerate other patterns of senescence.     

Population history of native groups in pre- and postcontact Spanish Florida: aggregation, gene flow, and genetic drift on the Southeastern U.S. Atlantic coast

Evolutionary trends and population history and structure are discussed for a series of late prehistoric and historic-period skeletal samples from the Georgia coast and interior (the Guale). Phenotypic dental measurement data were collected for nine samples from the late prehistoric (AD 1200-1400) and historic (AD 1608-1702) periods and subjected to population genetic and statistical analyses. The primary trends were for an increase in tooth size through time, and for an initial increase in dental variability in the early historic period, followed by a subsequent decline in dental variability in the late historic period. Given the increasing stress levels, evidenced by previous bioarchaeological analyses (Larsen [2001] Bioarchaeology of Spanish Florida, Gainesville: University Press of Florida), an environmental explanation for the increase in tooth size is unlikely. It is proposed that the early historic period witnessed aggregation and gene flow with extraregional populations, possibly African slaves or nonlocal Native American population groups. The late historic period may have experienced significant loss of phenotypic variability due to genetic drift. In both time periods, the evolutionary mechanism increased average tooth size, with independent variance effects.Because microevolutionary trends obscure patterns of gene flow and population ancestry, the data were detrended following Konigsberg ([1990a] Hum. Biol. 62:49-70), and submitted to standard population genetic analyses (Relethford et al. [1997] Hum. Biol. 69:443-465). Analysis of the precontact samples in isolation (Irene Mound, Irene Mortuary, and an aggregate coastal sample) indicated little genetic microdifferentiation (F(ST) = 0.008), limited extralocal gene flow, and a small distinction between interior and coastal samples. The inclusion of the historic data dramatically increased variability levels (F(ST) = 0.019). The analysis of extralocal gene flow indicates that the late mission period experienced significantly less external gene flow, which is consistent with historic models that suggest the social organization of the Guale during this time period may have been significantly altered. Genetic distances also indicate a primary division between inland and coastal precontact samples and a maintenance of biological populations along the coast. In other words, the coastal, early historic, and late historic period samples are phenotypically homogeneous, supporting the notion that the mission populations were drawn from the local population base. The late mission period sample was also, however, more closely related to the interior samples. This may suggest that the late mission period population was an aggregate sample composed of both remnant interior and coastal population groups.     

Genetic variation in time and space: the use of herbarium specimens to reconstruct patterns of genetic variation in the endangered orchid Anacamptis palustris

Habitat alteration, fragmentation and destruction as a consequence of human impact are a global phenomenon. Here we document changes in genetic variation in the marsh orchid Anacamptis palustris as a consequence of such habitat changes. We examined historical specimens that were collected during the nineteenth and early twentieth centuries, prior to the most recent massive habitat changes affecting this species. Sequences of a hypervariable region in the plastid DNA, located in the tRNA^LEU intron, from herbarium vouchers were compared with those from a near-exhaustive survey of the extant A. palustris populations on the Italian peninsula. It was found that private haplotypes and alleles found in small, extant populations were once widespread and more common in historic populations and that alleles, once present in historic populations, have gone extinct. In addition, genetic differentiation among populations has increased over time and haplotype frequencies significantly differ among historic and extant populations. These results document that human induced habitat changes have reduced genetic diversity and increased the importance of random genetic drift in this species. It is concluded that the analysis of herbarium specimens may provide important insights into changes of genetic diversity over time and may be critical for correct inference of the evolutionary history of rare and endangered species.     

Analysis of quantitative methods for rib seriation using the Spitalfields documented skeletal collection

Accurate rib seriation is essential in forensic anthropology and bioarchaeology for determination of minimum numbers of individuals, sequencing trauma patterns to the chest, and identification of central ribs for use in age estimation. We investigate quantitative methods for rib seriation based on three metric variables: superior (anterior) costo-transverse crest height (SCTCH), articular facet of the tubercle-to-angle length (AFTAL), and head-to-articular facet length (HAFL). The sample consists of complete but unseriated sets of ribs from 133 individuals from the documented (known age and sex) and undocumented skeletal collections of Christ Church Spitalfields, London. This research confirms the results of an earlier study (Hoppa and Saunders [1998] J. Forensic. Sci. 43:174-177) and extends it with the application of two new metric traits and further analyses of sex differences. Analyses of variance showed that SCTCH and AFTAL are significantly associated (P < 0.001) with rib number. Tukey tests of pairwise rib comparisons revealed that for two dimensions (SCTCH and AFTAL), the central ribs (3rd-6th) are significantly distinct from each other (P < 0.05). Using simple ranking of either the SCTCH or AFTAL traits, the proportion of correctly identified ribs within +/-1 position was 80%, compared to initial seriation using morphological methods (Dudar [1993] J. Forensic. Sci. 28:788-797; Mann [1993] J. Forensic. Sci. 28:151-155). Significant sex dimorphism was also identified for these two traits. Analysis of the HAFL trait produced somewhat equivocal results, suggesting that this variable is not reliable for rib seriation. The variable SCTCH proves to be the most useful dimension for seriation, and shows that all but the 7th-9th ribs can be distinguished from others in the sequence, with important results for the 4th rib, where ranking allowed identification in 86% of cases, consistent with morphological methods for intact ribs.     

Artificial Occurrence of the Fallow Deer, Dama dama dama (L., 1758), on the Island of Rhodes (Greece): Insight from mtDNA Analysis

We investigated the origins of the fallow deer (Dama dama dama) of Rhodes by both morphological and molecular means. Our results show that these deer have homogeneous phenotypic patterns. All specimens fell within the common colour coat variety typical of the wild form. The Rhodian deer appear to be rather small, especially when compared with specimens from central and northern Europe. We then sequenced the HVR-I of 13 deer from Rhodes and compared these sequences with other 31 samples obtained from different European and Anatolian populations of fallow deer. Out of 44 sequences, 23 haploypes were found. When compared to the Turkish and Italian populations, the population of Rhodes revealed lower values of within population genetic diversity. The fallow deer from Rhodes are characterized by an 80-bp mitochondrial DNA (mtDNA) insertion not found elsewhere. As a consequence, all the deer from Rhodes form a tight cluster, distinct from all other fallow deer populations. This uniqueness makes the conservation and management of the Rhodian population particularly urgent.     

Individual covariation in life-history traits: seeing the trees despite the forest

We investigated the influence of age on survival and breeding rates in a long-lived species Rissa tridactyla using models with individual random effects permitting variation and covariation in fitness components among individuals. Differences in survival or breeding probabilities among individuals are substantial, and there was positive covariation between survival and breeding probability; birds that were more likely to survive were also more likely to breed, given that they survived. The pattern of age-related variation in these rates detected at the individual level differed from that observed at the population level. Our results provided confirmation of what has been suggested by other investigators: within-cohort phenotypic selection can mask senescence. Although this phenomenon has been extensively studied in humans and captive animals, conclusive evidence of the discrepancy between population-level and individual-level patterns of age-related variation in life-history traits is extremely rare in wild animal populations. Evolutionary studies of the influence of age on life-history traits should use approaches differentiating population level from the genuine influence of age: only the latter is relevant to theories of life-history evolution. The development of models permitting access to individual variation in fitness is a promising advance for the study of senescence and evolutionary processes.     

Ancient mitochondrial DNA analyses of Iberian sturgeons

Today's European sturgeons are relics of erstwhile widely distributed populations, diminished mainly by overfishing and habitat changes over the centuries. While extinct European populations in the Baltic and North seas have been identified as Acipenser oxyrhinchus or A. sturio , a clear species determination on the Iberian Peninsula is still lacking. Plans to conserve existing populations and to re-introduce extinct wild populations in European rivers will benefit from information of historic population/genotype composition. In this study, we used techniques involving ancient DNA as well as morphological comparisons based on bony scutes to identify twelve samples from five archaeological sites (650 bc –1500 ad , one sample dated 10.1–11.8 ky) on the Iberian Peninsula. All amplified PCR products of bony scutes (n = 5) had the mitochondrial DNA haplotypes of European sturgeon. Neither mitochondrial haplotypes of A. naccarii nor mitochondrial haplotypes of A. oxyrinchus were found.     

Genetic discontinuity, breeding-system change and population history of Arabis alpina in the Italian Peninsula and adjacent Alps

Arabis alpina is a widespread plant of European arctic and alpine environments and belongs to the same family as Arabidopsis thaliana. It grows in all major mountain ranges within the Italian glacial refugia and populations were sampled over a 1300 km transect from Sicily to the Alps. Diversity was studied in nuclear and chloroplast genome markers, combining phylogeographical and population genetic approaches. Alpine populations had significantly lower levels of nuclear genetic variation compared to those in the Italian Peninsula, and this is associated with a pronounced change in within-population inbreeding. Alpine populations were significantly inbred (F(IS) = 0.553), possibly reflecting a change to the self-incompatibility system during leading edge colonization. The Italian Peninsula populations were approaching Hardy-Weinberg equilibrium (outbreeding, F(IS) = 0.076) and genetic variation was highly structured, consistent with independent local 'refugia within refugia' and the fragmentation of an established population by Quaternary climate oscillations. There is very little evidence of genetic exchange between the Alps and the Italian Peninsula main distribution ranges. The Alps functioned as a glacial sink for A. alpina, while the Italian Peninsula remains a distinct and separate long-term refugium. Comparative analysis indicated that inbreeding populations probably recolonized the Alps twice: (i) during a recent postglacial colonization of the western Alps from a Maritime Alps refugium; and (ii) separately into the central Alps from a source outside the sampling range. The pronounced geographical structure and inbreeding discontinuities are significant for the future development of A. alpina as a model species.     

THE ZOOLOGICAL SOCIETY OF LONDON STAMFORD RAFFLES LECTURE 1996 From Gaia to Noah: human responsibilities in nature

Life-table data of 56 natural populations of mammals were analysed, modelling mortality rate as a three-phase step function of age (the step model ) instead of using the Gompertz model. In the step model, mortality rale is constant in each phase. The phases correspond to juveniles and young and old adults. The age of transition between young and old adults is referred to as the age of senescence. This approach has the advantages that, for the first time, the age of senescence is identified objectively using a robust statistical procedure, and that young adult mortality rates are estimated without bias since no assumption is made about how adult mortality rate changes with age. A further statistical problem solved here that has previously caused difficulty is that of correctly accounting for the different levels of precision in data from different age classes.
Significant changes ( P < 0.05) in adult mortality rate with age were found in 27 out of 56 populations. In 23 of these 27 cases, adult mortality rate increased with age. Juvenile mortality rate differed significantly from young adult mortality rate in 21 cases; in 18 of these the rate for juveniles was higher. These results are discussed in relation to earlier analyses, in particular that of Promislow (1991).