Conceptual issues in hominin taxonomy: Homo heidelbergensis and an ethnobiological reframing of species.

Efforts to name and classify Middle Pleistocene Homo, often referred to as "Homo heidelbergensis" are hampered by confusing patterns of morphology but also by conflicting paleoanthropological ideologies that are embedded in approaches to hominin taxonomy, nomenclature, and the species concept. We deconstruct these issues to show how the field's search for a "real" species relies on strict adherence to pre-Darwinian essentialist naming rules in a post-typological world. We then examine Middle Pleistocene Homo through the framework of ethnobiology, which examines on how Indigenous societies perceive, classify, and name biological organisms. This research reminds us that across human societies, taxonomies function to (1) identify and classify organisms based on consensus pattern recognition and (2) construct a stable nomenclature for effective storage, retrieval and communication of information. Naming Middle Pleistocene Homo as a "real" species cannot be verified with the current data; and separating regional groups into distinct evolutionary lineages creates taxa that are not defined by readily perceptible or universally salient differences. Based on ethnobiological studies of this kind of patterning, referring to these hominins above the level of the species according to their generic category with modifiers (e.g., "European Middle Pleistocene Homo") is consistent with observed human capabilities for cognitive differentiation, is both necessary and sufficient given the current data, and will allow for the most clear communication across ideologies going forward.

Paleodemographic analysis of age at death for a population of Black Sea Scythians: An exploration by using Bayesian methods.

OBJECTIVES: Studies of the demography of past populations involving deterministic life tables can be criticized for ignoring the errors of estimation. Bayesian methods offer an alternative, by focusing on the uncertainty of the estimates, although their results are often sensitive to the choice of prior distributions. The aim of this study is to explore a range of Bayesian methods for estimating age at death for a population of nomadic warriors-Scythians from the Black Sea region. MATERIALS AND METHODS: In total, skeletons of 312 individuals (93 children and 219 adults) from Glinoe (Moldova), dated to the 5th-2nd century BCE, were examined. We unified the age categories corresponding to different aging methods, allowing an application of a probabilistic assessment of the age categorization. A hierarchical Bayesian multinomial-Dirichlet-Dirichlet model was applied, with a hypothetical, subjective reference population, a real reference population, and no reference. RESULTS: Stationary-population life expectancy was estimated as 27.7 years (95% CI: 25.1-30.3) for a newborn (e0 ), and 16.4 years (14.0-19.0) for 20-year-olds (e20 ), although with high uncertainty, and sensitive to the model specification. Slight differences in longevity between different social strata and between the Classical and Late chronological periods were found, although with high estimation errors. A more robust finding, confirming earlier studies, was a high probability of death in young adulthood, which could depend on Scythian lifestyle (conflicts, wars). DISCUSSION: Our study shows a way to overcome some limitations of broad age categorization by using the Bayesian approach with alternative model specifications, allowing to assess the impact of reference populations.

Shipping amphorae and shipping sheep? Livestock mobility in the north-east Iberian peninsula during the Iron Age based on strontium isotopic analyses of sheep and goat tooth enamel.

Animal mobility is a common strategy to overcome scarcity of food and the related over-grazing of pastures. It is also essential to reduce the inbreeding rate of animal populations, which is known to have a negative impact on fertility and productivity. The present paper shows the geographic range of sheep provisioning in different phases of occupation at the Iron Age site of Turó de la Font de la Canya (7th to 3rd centuries BC). Strontium isotope ratios from 34 archaeological sheep and goat enamel, two archaeological bones and 14 modern tree leaves are presented. The isotopic results suggest that sheep and goats consumed at the site were reared locally (within a few kilometres radius) during the whole period of occupation. The paper discusses the isotopic results in light of the socio-political structure of this period, as complex, strongly territorial societies developed during the Iron Age in the north-east Iberian Peninsula.

Who were the miners of Allumiere? A multidisciplinary approach to reconstruct the osteobiography of an Italian worker community.

This research presents an in-depth study of the skeletal remains collected from the archaeological site of Allumiere (15th-16th centuries CE; Rome, Italy). A multidisciplinary approach was used, combining skeletal biology, molecular anthropology and archaeobotany with the aim of reconstructing the osteobiography of the alum miners buried at the site. Since 1460, the area of the Tolfa Mountains was significant for the exploitation of alum which was used for a wide range of purposes in the Middle Ages, ranging from woven production to medical practice. A total of 70 individuals (63 adults and 7 juveniles) were studied. The sex ratio of the community indicated a higher prevalence of males with respect to females. Morphological examination indicated occupational musculoskeletal stress markers, which might reflect the specific phase of alum production that each individual was occupied in. Dietary reconstruction was primarily performed through carbon and nitrogen stable isotope analysis with integration of the results obtained by microscopic, genetic and GC-MS investigations on dental calculus. The diet was omnivorous, indicating a reliance on C3-terrestrial protein and evidence for limited C4 consumption by some individuals. Herbivores, such as sheep and cattle, appear to have contributed to the diet more than pigs and chickens. Consumption of Fagaceae and Poaceae species was predominant; moreover, indicators of Brassicaceae and milk and its derivatives were abundantly recurrent in the population, followed by plant oils and theophylline. Furthermore, the detection of pharmacological alkaloids indicated the knowledge and application of medicinal plants by the community. The novel use of multiple techniques based on cutting-edge technologies has provided a unique window on the lifestyles of individuals from one of the first Italian settlements of alum workers.

Multistep food plant processing at Grotta Paglicci (Southern Italy) around 32,600 cal B.P.

Residue analyses on a grinding tool recovered at Grotta Paglicci sublayer 23A [32,614 ± 429 calibrated (cal) B.P.], Southern Italy, have demonstrated that early modern humans collected and processed various plants. The recording of starch grains attributable to Avena (oat) caryopses expands our information about the food plants used for producing flour in Europe during the Paleolithic and about the origins of a food tradition persisting up to the present in the Mediterranean basin. The quantitative distribution of the starch grains on the surface of the grinding stone furnished information about the tool handling, confirming its use as a pestle-grinder, as suggested by the wear-trace analysis. The particular state of preservation of the starch grains suggests the use of a thermal treatment before grinding, possibly to accelerate drying of the plants, making the following process easier and faster. The study clearly indicates that the exploitation of plant resources was very important for hunter-gatherer populations, to the point that the Early Gravettian inhabitants of Paglicci were able to process food plants and already possessed a wealth of knowledge that was to become widespread after the dawn of agriculture.

Timing of maxillofacial-oral injuries in an individual of the ancient city of Herculaneum (79 AD, Naples, Italy): a case report.

BACKGROUND: This study is based on the skeletal remains of an adult female from the ancient city of Herculaneum (Naples, Italy), who was a victim of the eruption of the nearby Vesuvius Volcano on 24-25 August, AD 79. RESULTS AND DISCUSSION: Examination of the maxillofacial region revealed evidence of unilateral condylar fracture and dislocation, as well as traumatic dental injuries. The injuries observed might have been the consequence of a direct blow to the mental region that was transmitted in a direction that raised the mandible, causing an indirect fracture in the right condylar neck when the condylar head collided directly with the temporal glenoid fossa. This indirect impact also resulted in partial fracture of three dental crowns due to the violent impact of the arches between them, and the sharp impact together of the upper and lower teeth. It is suggested that these injuries were sustained due to an accidental fall when the individual was between 7 and 15 years old, which is supported by the morphology, location and extension of the injuries, and the characterization of the impact. CONCLUSION: These results are an illustration of how dental anthropology and forensic approaches can be applied with great benefit to archaeological skeletal remains.

Odontometric sex estimation on three populations of the Iron Age from Abruzzo region (central-southern Italy).

BACKGROUND: In archaeological contexts, sex identification is a necessary step for a complete reconstruction of the biological profile of the individuals and to know demographic patterns of the population, nutritional stress, diseases, growth and development, and distribution of pathological conditions. METHODS: This study is based on the skeletal remains of 149 individuals from three protohistoric populations in close temporal and geographic proximity in Abruzzo region (central-southern Italy): Opi, Alfedena and Bazzano. It has been possible to develop logistic regression equations based on dental measurements of permanent teeth of adult individuals whose sex had previously been estimated based on pelvic and cranial features. These equations were subsequently applied to the permanent dentition of immature individuals and adult individuals whose sex was estimated as uncertain or unknown in order to estimate their sex. RESULTS: The mandibular canine is the tooth with the greatest sexual dimorphism in adults, followed by both maxillary and mandibular first and second molars, providing a correct assignment of sex ranging from 83.7% and 95.9% of cases, depending on the dimensions used for the construction of these equations. Of the 29 individuals in the target sample (14 adultus, 10 juvenilis and 5 infans), sex estimation was possible for 23 (10 adultus, 8 juvenilis and 5 infans), representing an applicability rate of 79.31% of the individuals. CONCLUSIONS: The results indicate that odontometrics is a useful tool for sex estimation and allows to increase the data to perform more complete paleodemographic studies on archaeological populations.

Bone quality in prehistoric, Cis-Baikal forager femora: a micro-CT analysis of cortical canal microstructure.

Bone quality, a contributor to bone strength, is determined by structural and mechanical properties, which may be analyzed by gross and/or microscopic methods. Variables that contribute to bone quality, such as porosity, can provide insight into the health and lifestyles of people in prehistory. This study tests the ability of microcomputed tomography (µCT) to capture and characterize cortical canal systems in archaeological bone. Seven variables and 71 femora are analyzed to explore bone dynamics in prehistoric foragers from Lake Baikal, Siberia. The results indicate that canal number and canal separation differ significantly (P < 0.05) between age-at-death categories, but only for the pooled and male samples. When merged into a new variable by means of principal components analysis, canal diameter and canal surface to canal volume are also able to discriminate amongst age-at-death categories, as well as between the sexes. However, the overall lack of significant differences between the sexes and amongst age-at-death categories indicates that Baikal forager bone quality (i.e., canal architecture) did not change drastically throughout the lifespan. Interestingly, principal component one identified an untested variable that contributes to canal microstructure variability, and a sexual division of labor may promote divergent trends in canal degree of anisotropy between the sexes. Overall, µCT provides an alternate method for exploring bone quality in archaeological remains, complementing existing methods such as thin-sectioning and gross morphological analyses.

Novel contribution on the diagenetic physicochemical features of bone and teeth minerals, as substrates for ancient DNA typing.

The extraction of DNA from skeletal remains is a major step in archeological or forensic contexts. However, diagenesis of mineralized tissues often compromises this task although bones and teeth may represent preservation niches allowing DNA to persist over a wide timescale. This exceptional persistence is not only explained on the basis of complex organo-mineral interactions through DNA adsorption on apatite crystals composing the mineral part of bones and teeth but is also linked to environmental factors such as low temperatures and/or a dry environment. The preservation of the apatite phase itself, as an adsorption substrate, is another crucial factor susceptible to significantly impact the retrieval of DNA. With the view to bring physicochemical evidence of the preservation or alteration of diagenetic biominerals, we developed here an analytical approach on various skeletal specimens (ranging from ancient archeological samples to recent forensic specimens), allowing us to highlight several diagenetic indices so as to better apprehend the complexity of bone diagenesis. Based on complementary techniques (X-ray diffraction (XRD), Fourier transform infrared (FTIR), calcium and phosphate titrations, SEM-EDX, and gravimetry), we have identified specific indices that allow differentiating 11 biological samples, primarily according to the crystallinity and maturation state of the apatite phase. A good correlation was found between FTIR results from the analysis of the v3(PO4) and v4(PO4) vibrational domains and XRD-based crystallinity features. A maximal amount of information has been sought from this analytical approach, by way of optimized posttreatment of the data (spectral subtraction and enhancement of curve-fitting parameters). The good overall agreement found between all techniques leads to a rather complete picture of the diagenetic changes undergone by these 11 skeletal specimens. Although the heterogeneity and scarcity of the studied samples did not allow us to seek direct correlations with DNA persistence, the physicochemical parameters described in this work permit a fine differentiation of key properties of apatite crystals among post mortem samples. As a perspective, this analytical approach could be extended to more numerous sets of specimens so as to draw statistical relationships between mineral and molecular conservation.

Comparability of multiple data types from the Bering Strait region: cranial and dental metrics and nonmetrics, mtDNA, and Y-chromosome DNA.

Different data types have previously been shown to have the same microevolutionary patterns in worldwide data sets. However, peopling of the New World studies have shown a difference in migration paths and timings using multiple types of data, spurring research to understand why this is the case. This study was designed to test the degree of similarity in evolutionary patterns by using cranial and dental metric and nonmetric data, along with Y-chromosome DNA and mtDNA. The populations used included Inuits from Alaska, Canada, Siberia, Greenland, and the Aleutian Islands. For comparability, the populations used for the cranial and molecular data were from similar geographic regions or had a shared population history. Distance, R and kinship matrices were generated for use in running Mantel tests, PROTEST analyses, and Procrustes analyses. A clear patterning was seen, with the craniometric data being most highly correlated to the mtDNA data and the cranial nonmetric data being most highly correlated with the Y-chromosome data, while the phenotypic data were also linked. This patterning is suggestive of a possible male or female inheritance, or the correlated data types are affected by the same or similar evolutionary forces. The results of this study indicate cranial traits have some degree of heritability. Moreover, combining data types leads to a richer knowledge of biological affinity. This understanding is important for bioarchaeological contexts, in particular, peopling of the New World studies where focusing on reconciling the results from comparing multiple data types is necessary to move forward.

Estimating the age structure of a buried adult population: a new statistical approach applied to archaeological digs in France.

Paleodemographers have developed several methods for estimating the age structure of historical populations in absence of civil registration data. Starting from biological indicators alone, they use a reference population of known sex and age to assess the conditional distribution of the biological indicator given age. However, the small amount of data available and the unstable nature of the related statistical problem mean that most methods are disappointing. Using the most reliable reference data possible, we propose a simple statistical method, integrating the maximum amount of information included in the actual data, which quite significantly improves age estimates for a buried population. Here the method is applied to a French cemetery used from Late Antiquity to the end of the Early Middle Ages.

Validity of child anthropometric measurements in the Special Supplemental Nutrition Program for Women, Infants, and Children.

INTRODUCTION: The Special Supplemental Nutrition Program for Women, Infants, and Children (WIC) serves 50% of infants and 25% of preschool-aged children in the United States and collects height and weight measurements from eligible children every 6 mo, making WIC data a valuable resource for studying childhood growth and obesity. We assessed the accuracy of measurements collected by WIC staff by comparing them to "gold standard" measurements collected by trained research staff. RESULTS: Intraclass correlation coefficients (ICCs) measuring agreement between WIC and research protocol measurements for height, weight, and BMI were 0.96, 0.99, and 0.93, respectively. Although WIC measurements overestimated height by 0.6 cm and weight by 0.05 kg on average, BMI was underestimated by only 0.15 kg/m(2) on average. WIC BMI percentiles classified children as overweight/obese vs. underweight/normal with 86% sensitivity and 92% specificity. DISCUSSION: We conclude that height, weight, and BMI measurements of children aged 2-5 y collected by trained WIC staff are sufficiently accurate for monitoring and research purposes. METHODS: At seven WIC clinics in southern California, 287 children aged 2-5 y measured for height and weight by WIC staff using WIC standard protocol were remeasured by research staff using a research protocol (duplicate measurements with shoes and outerwear removed were taken by trained personnel).

Odontometric sex discrimination in the Herculaneum sample (79 AD, Naples, Italy), with application to juveniles.

Sex determination of subadult skeletal remains with satisfactory accuracy represents one of the most important limitations of archaeological research and forensic practice. Teeth are one of the most durable physical elements of an individual that remain after death, and constitute a potential source of information about the biological sex of that individual. This study was based on the skeletal remains of 117 individuals from the ancient city of Herculaneum (Naples, Italy), victims of the eruption of the nearby volcano Vesuvius on 24/25 August, 79 AD. It has been possible to develop discriminant function formulae based on dental dimensions of adult individuals whose sex had previously been determined based on descriptive osteologic criteria. These formulae were subsequently applied to the permanent dentitions of immature individuals of the same population in order to estimate their sex. The results show that the canine is the tooth with the greatest sex dimorphism in adults, providing percentages of correct assignment of sex between 76.5% and 100% depending on the dimension used. Of the 30 subadult individuals in the target sample, estimation of sex was possible for 22 individuals. Sex assignments matched those determined from descriptive characteristics of the ilia and mandible in 73.33% of the cases. The results provide some optimism that this method may be applicable to juvenile archaeological samples.

Investigation of diachronic dietary patterns on the islands of Ibiza and formentera, Spain: Evidence from carbon and nitrogen stable isotope ratio analysis.

To examine how dietary patterns may have changed in the western Mediterranean through time, stable carbon and nitrogen isotope ratios were measured on extracted bone collagen from fauna (n = 75) and humans (n = 135) spanning four distinct chronological periods: Chalcolithic (c.2100-1600 BC), Punic (6th-2nd/1st century BC), Late Antiquity-Early Byzantine (4th-7th century AD), and Islamic (c.10th-13th century AD) on the islands of Ibiza and Formentera, Spain. The Chalcolithic, Punic, and Late Antiquity-Byzantine societies all showed evidence of a predominately C(3) terrestrial-based diet with a possible input of a small amount of marine and/or C(4) dietary resources. In contrast, the Islamic population on Ibiza had a subsistence strategy that was reliant on a significant amount of C(4) plants and/or animals fed a C(4) diet, likely millet. These results indicate a fairly constant C(3) terrestrial-based diet on the islands of Ibiza and Formentera through time, with a shift to C(4) dietary resources during the Islamic Period. Further research is needed from other Islamic populations in and around the Mediterranean to better understand this unique dietary adaptation. Am J Phys Anthropol 143:512-522, 2010. © 2010 Wiley-Liss, Inc.

Craniometric variation in the Aleutians: integrating morphological, molecular, spatial, and temporal data.

Several hypotheses have been put forward about the origins and evolution of the inhabitants of the Aleutian Islands. Both Hrdlicka [The Aleutian and Commander Islands and Their Inhabitants (Philadelphia: Wistar Institute of Anatomy and Biology, 1945)] and Laughlin ["The Alaska gateway viewed from the Aleutian Islands," in Papers on the Physical Anthropology of the American Indian, W. S. Laughlin, ed. (New York: Viking Fund, 1951), 98-126] analyzed cranial morphology and came to somewhat different conclusions using a typological approach and limited analytical methods. Subsequent investigations using morphological data have not significantly improved our understanding of Aleut prehistory. More recently, radiocarbon dating and mitochondrial DNA analyses have shed light on Aleut genetic variation and changes over time, but better morphological methods using multivariate statistical analysis have not yet been used. We analyzed craniometric data using multivariate procedures and found that Aleuts demonstrate significant changes in cranial morphology over time, and these changes correspond to Hrdlicka's observations but may not necessarily reflect in-migration. The morphological changes were concentrated in the very aspects of morphology that are easily observable and that Hrdlicka most often measured, namely, cranial length, breadth, and height, but they were obscured when craniometric variation as a whole was analyzed. Also, we found that the morphological changes over time were not related to the changes in haplogroup frequencies over time, suggesting that migration into the Aleutians did not play a significant role in producing the morphological changes. However, craniometric variability apparently increases over time, suggesting in-migration, localized selection, and/or greater environmental heterogeneity. Our results contradict Laughlin's observations but may be more in line with his hypothesis of in situ evolutionary changes absent gene flow. In addition to selection, gene flow, and gene drift, however, sociocultural changes must also be considered as a factor in why morphology changed over time.

Biohistorical approaches to "race" in the United States: Biological distances among African Americans, European Americans, and their ancestors.

Folk taxonomies of race are the categorizations used by people in their everyday judgments concerning the persons around them. As cultural traditions, folk taxonomies may shape gene flow so that it is unequal among groups sharing geography. The history of the United States is one of disparate people being brought together from around the globe, and provides a natural experiment for exploring the relationship between culture and gene flow. The biohistories of African Americans and European Americans were compared to examine whether population histories are shaped by culture when geography and language are shared. Dental morphological data were used to indicate phenotypic similarity, allowing diachronic change through United States history to be considered. Samples represented contemporary and historic African Americans and European Americans and their West African and European ancestral populations (N = 1445). Modified Mahalanobis' D(2) and Mean Measure of Divergence statistics examined how biological distances change through time among the samples. Results suggest the social acceptance for mating between descendents of Western Europeans and Eastern and Southern European migrants to the United States produced relatively rapid gene flow between the groups. Although African Americans have been in the United States much longer than most Eastern and Southern Europeans, social barriers have been historically stronger between them and European Americans. These results indicate that gene flow is in part shaped by cultural factors such as folk taxonomies of race, and have implications for understanding contemporary human variation, relationships among prehistoric populations, and forensic anthropology.

Trauma-induced changes in diaphyseal cross-sectional geometry in two elites from Copan, Honduras.

This research utilized biplanar radiographs to estimate cross-sectional biomechanical properties for the skeletal remains of two elite individuals from the Early Classic period (ca. AD 400-600) of Copan, Honduras: K'inich Yax K'uk' Mo' (Hunal Burial 95-2), founder of the Early Classic Dynasty at Copan, and the primary female interment (Burial 37-8) from the Motmot tomb. Both individuals survived severe blunt-force insults to the right forearm. Gross skeletal examination and evaluation of the radiographs for K'inich Yax K'uk' Mo' suggest that these traumas resulted from, at least in part, disuse atrophy of the affected forearm skeletal elements. Gross and radiologic evaluation of the Motmot remains countered the possibility that she suffered from a metabolic bone disease, and confirmed the presence of a well-healed parry fracture of the right ulna. The degree of asymmetry in cross-sectional biomechanical properties reported here for K'inich Yax K'uk' Mo' is likely the secondary result of the described blunt-force trauma. The results obtained for the principal Motmot interment are not as dramatic, but suggest subtle changes to humeral cross-sectional geometry subsequent to trauma.

Trophic level and macronutrient shift effects associated with the weaning process in the Postclassic Maya.

The weaning process was investigated at two Maya sites dominated by Postclassic remains: Marco Gonzalez (100 BC-AD 1350) and San Pedro (1400-AD 1650), Belize. Bone collagen and bioapatite were analyzed from 67 individuals (n < or = 6 years = 15, n > 6 years = 52). Five isotopic measures were used to reconstruct diet and weaning: stable nitrogen- and carbon-isotope ratios in collagen, stable carbon- and oxygen-isotope ratios in bioapatite, and the difference in stable carbon-isotope values of coexisting collagen and bioapatite. Nitrogen-isotope ratios in infant collagen from both sites are distinct from adult females, indicating a trophic level effect. Collagen-to-bioapatite differences in infant bone from both sites are distinct from adult females, indicating a shift in macronutrients. Oxygen-isotope ratios in infant bioapatite from both sites are also distinct from adult females, indicating the consumption of breast milk. Among infants, carbon- and nitrogen-isotope ratios vary, indicating death during different stages in the weaning process. The ethnohistoric and paleopathological literature on the Maya indicate cessation of breast-feeding between ages 3-4 years. Isotopic data from Marco Gonzalez and San Pedro also indicate an average weaning age of 3-4 years. Based on various isotopic indicators, weaning likely began around age 12 months. This data set is not only important for understanding the weaning process during the Postclassic, but also demonstrates the use of collagen-to-bioapatite spacing as an indicator of macronutrient shifts associated with weaning.

Isotopic evidence for age-related variation in diet from Isola Sacra, Italy.

This study examines collagen (N=105) and apatite (N=65) data from an Imperial Roman skeletal sample from the necropolis of Isola Sacra (Rome, Italy). This paper explores correlations between the isotopic composition of bone samples and the inferred age and sex of these individuals (aged 5--45+ years). The collagen of males, and older individuals in general, was significantly enriched in (15)N but not (13)C. Bone carbonate was somewhat depleted in (13)C in some older individuals, suggesting increased consumption of olive oil and possibly wine. Subadults (>5 years) in the sample appear to have consumed an almost exclusively terrestrial diet. This study demonstrates a clear trend in dietary patterns between adult age groups, as well as between adults and children within a population.

Dispersal ghosts in Oceania.

Anthropological genetics helps expand our understanding of human phenotypes in the Pacific, in part because of its focus on gene genealogies to infer past episodes of dispersal and to differentiate these events from adaptations due to long-duration directional selection. Sewall Wright's 1949 seminal paper on population structure emphasized that there were two strong forces that exerted systematic and therefore determinant pressure on the gene pool: recurrent immigration and gene flow. These are important topics to all discussions of human dispersal in any region of the world. Furthermore, Wright listed five unique kinds of events that produced indeterminate or unpredictable changes that could lead to phenotypic and genotypic effects. In this category, he placed unique selective incidents, unique hybridization events, unique reductions in number, swamping by mass immigration, and mutational drive due to an allele always being favored since its origin or introduction. This discussion of human dispersal in the Pacific will touch on these topics, since they provide a second level of complexity in knowing who moved about a region of the world found already settled when rediscovered by colonial explorers during the 16-18th centuries.