Lukas Friedl

During Marine Isotope Stage (MIS) 2, abrupt climate changes created highly variable paleoenvironments inhabited by human populations across the Iberian Peninsula. Pollen and sedimentary analyses from deep-sea cores off Portugal provide records of regional-scale paleoenvironmental responses to the climate shifts that punctuated this period. Archaeological assemblages offer a regional and local-scale understanding of human- environment interactions during this period. One site in particular, Lapa do Picareiro, has yielded a continuous, stratified sedimentary sequence that provides a diachronic record for MIS 2 human occupation and environmental change. Here, we present archaeological data from the Late Gravettian through Solutrean occupations (Levels U-O) in order to show how local-scale, assemblage-level variability may or may not help our understanding of human-environment interactions and culture change during this period when we try to fit them into the traditional techno- typological framework. Lithic artifact assemblages record the technological changes associated with the Last Glacial Maximum (LGM), including the Gravettian-Solutrean transition. Age control is provided by 23 radiocarbon dates from Levels U-O. The spatial distribution of artifacts, animal bones and charcoal concentrations shows a relatively high degree of assemblage integrity. The taphonomic study of the faunal remains informs on local paleoenvironments and human diet choice during the LGM. The combined results are used here to understand human responses to long-term environmental change in central Portugal.

Neandetherthal skeletal remains are the first described and longest-studied modern human ancestors. The history of research on Neanderthals dates back to the discovery of the first specimen in the Feldhoffer cave in 1856. Initially, the debate focused on the taxonomic position of the Neanderthals and their exact anatomical relationship with modern humans. Later research concentrated on many other issues including partial aspects of life, biological and cultural adaptation to glacial conditions in Europe, issues of diet and health, or issues such as paleobiology of Neanderthals (reproductive behavior, ontogeny, mortality). Another important direction is the question of Neanderthal extinction in the period between 30 to 25 thousand years BP. This paper provides an overview of the research directions related to Neanderthals in the context of new discoveries of skeletal remains and the development of new methods of study. The authors use the specific research conducted on the Lapa do Pi...

Genetic species determination has become an indispensable tool in forensics, archaeology, ecology, and food authentication. The available methods are either suited for detecting a single taxon across many samples or for screening a wide range of species across a few samples. Here, we introduce “Species by Proteome INvestigation” (SPIN), a proteomics workflow capable of querying over 150 mammalian species in 7.2 minutes of mass spectrometry (MS) analysis. Streamlined and automated sample preparation by protein aggregation capture, high-speed chromatography and data-independent acquisition, and a confident species inference algorithm facilitate processing hundreds of samples per day. We demonstrate the correct classification of known references, reproducible species identification in degraded Iron-Age material from Scandinavia, and test the limits of our methods with Middle and Upper Palaeolithic bones from Southern European sites with late Neanderthal occupation. While this initial s...

Species determination based on genetic evidence is an indispensable tool in archaeology, forensics, ecology, and food authentication. Most available analytical approaches involve compromises with regard to the number of detectable species, high cost due to low throughput, or a labor-intensive manual process. Here, we introduce “Species by Proteome INvestigation” (SPIN), a shotgun proteomics workflow for analyzing archaeological bone capable of querying over 150 mammalian species by liquid chromatography-tandem mass spectrometry (LC-MS/MS). Rapid peptide chromatography and data-independent acquisition (DIA) with throughput of 200 samples per day reduce expensive MS time, whereas streamlined sample preparation and automated data interpretation save labor costs. We confirm the successful classification of known reference bones, including domestic species and great apes, beyond the taxonomic resolution of the conventional peptide mass fingerprinting (PMF)-based Zooarchaeology by Mass Sp...

Sex assessment of skeletal remains in the context of forensic investigation is one of the most important components when constructing biological profile of the deceased individual since it helps to significantly narrow down the number of potential victims. Therefore, the number of methods suitable to estimate sex should be as wide as possible, especially for cases of highly fragmented remains. This paper offers a classification method for sexing human remains based on an area around foramen magnum and tests other similar discriminatory functions published elsewhere on an independent sample from the circummediterranean region. We provide discriminant and logistic regression functions for several sets of variable combinations derived from head CT images. None of the functions performs reliably enough to be used in the forensic context. The same holds true for other discriminatory functions published in the literature. For most of the functions, the failure rate (its inability to successfully assign sex of an unknown individual) reaches 100%. Thus, despite the fact that foramen magnum is sexually dimorphic in most populations, its use in sexing cranial remains in the forensic context should be limited only to cases in which we know population affinity of unknown skeletal remains and can provide referential data from the same population to estimate sex.

Long-term trends in robusticity of lower limb bones in the genus Homo through the Pleistocene until the present have been proposed, which have been interpreted as a consequence of decreasing levels of mobility and activity patterns, changes in lifestyle, and environmental factors. There has also long been evidence that skeletal strength increases over an individual's lifespan. This increase is caused by continuous bone remodeling that optimizes the structure of a bone to resist mechanical loadings and creates a balance between endosteal resorption and subperiosteal apposition. However, none of the previous studies of temporal trends in robusticity has considered both processes and analyzed how individual age-related robusticity might influence higher-level temporal trends. This paper therefore explores temporal trends in robusticity of lower limb long bones within the genus Homo and considers how individual ages-at-death can confound published evolutionary trends, given the fact that some aspects of relative bone strength tend to increase over individual lifespans. Cross-sectional diaphyseal properties of the midshaft and proximal femur and midshaft tibia of Pleistocene and early Holocene individuals, together with data on age-at-death are used to analyze changes in relative bone strength relative to individuals' ages and evolutionary time. The results show increasing bone strength in adulthood until the fourth decade and then a slight decrease, an observation that conforms to previously published results on recent human populations. However, no significant impact of age-at-death on the trends along an evolutionary trajectory has been detected. The evolutionary trends in femoral and tibial relative strength can be described as fluctuating, probably as a consequence of changing mobility patterns, environmentally and technologically influenced behaviors, and demographic processes. The differences between evolutionary trends published in several studies are explained primarily as a result of different ways of standardizing cross-sectional parameters for size, and differences in sample composition.

Caves represent portions of past landscapes that were, for a long time, used by humans and their ancestors for various purposes and therefore, they have been a focus of researchers for more than two hundred years. Accumulated artifacts, ecofacts, and sediments can be used to reconstruct the environments and lives of the past. Here, we describe current methods applied to excavations of cave sites with emphasis put on their interdisciplinary nature. The case of systematic interdisciplinary research is documented with regard to the Lapa do Picareiro limestone cave, located in central Portugal, approximately 100 km north of Lisbon. The cave sediments have been dated to between 45 and 8 kya BP. A wide range of evidence from the cave (artifacts, faunal and floral remains, sediments, etc.) is used to reconstruct past natural environments (e.g., Bicho et al. 2011; Haws 2012), including fauna (Haws and Valente 2006; Hockett and Haws 2009; Valente 2004), human diet (Hockett and Haws 2009) and...

Adaptation to Late Pleistocene climate change is an oft-cited potential contributor to Neanderthal disappearance in Eurasia. Accordingly, research on Neanderthal behaviourincluding subsistence strategies, mobility, lithic technology, raw material procurement and demographyoften focuses on linking changes observable in the archaeological record to specific phases of climate and environmental change. However, these correspondences are often tenuous because palaeoclimatic and archaeological records are rarely available on the same scale. In Iberia, a critical location for understanding the demise of Neanderthals, some research indicates that Neanderthal populations were unable to recover from environmental degradations known as Heinrich Events, while other studies suggest that enclaves of Neanderthal populations survived for several millennia longer in refugial zones. Here, we present a palaeoenvironmental reconstruction study using analysis of δ 13 C and δ 18 O of herbivore tooth enamel recovered from two Mousterian deposits at Lapa do Picareiro, a site located in Portuguese Estremadura. We then use these data, combined with other site-based palaeoenvironmental indicators, to assess whether central Portugal acted as a refugium during periods of unfavourable climate, and to test whether Neanderthals in Portuguese Estremadura reorganised their mobility strategies after severe climate episodes.

Documenting the first appearance of modern humans in a given region is key to understanding the dispersal process and the replacement or assimilation of indigenous human populations such as the Neanderthals. The Iberian Peninsula was the last refuge of Neanderthal populations as modern humans advanced across Eurasia. Here we present evidence of an early Aurignacian occupation at Lapa do Picareiro in central Portugal. Diagnostic artifacts were found in a sealed stratigraphic layer dated 41.1 to 38.1 ka cal BP, documenting a modern human presence on the western margin of Iberia ∼5,000 years earlier than previously known. The data indicate a rapid modern human dispersal across southern Europe, reaching the westernmost edge where Neanderthals were thought to persist. The results support the notion of a mosaic process of modern human dispersal and replacement of indigenous Neanderthal populations.

The abundant femoral assemblage of Homo naledi found in the Dinaledi Chamber provides a unique opportunity to test hypotheses regarding the taxonomy, locomotion, and loading patterns of this species. Here we describe neck and shaft cross-sectional structure of all the femoral fossils recovered in the Dinaledi Chamber and compare them to a broad sample of fossil hominins, recent humans, and extant apes. Cross-sectional geometric (CSG) properties from the femoral neck (base of neck and midneck) and diaphysis (subtrochanteric region and midshaft) were obtained through CT scans for H. naledi and through CT scans or from the literature for the comparative sample. The comparison of CSG properties of H. naledi and the comparative samples shows that H. naledi femoral neck is quite derived with low superoinferior cortical thickness ratio and high relative cortical area. The neck appears superoinferiorly elongated because of two bony pilasters on its superior surface. Homo naledi femoral shaft shows a relatively thick cortex compared to the other hominins. The subtrochanteric region of the diaphysis is mediolaterally elongated resembling early hominins while the midshaft is anteroposteriorly elongated, indicating high mobility levels. In term of diaphyseal robusticity, the H. naledi femur is more gracile that other hominins and most apes. Homo naledi shows a unique combination of characteristics in its femur that undoubtedly indicate a species committed to terrestrial bipedalism but with a unique loading pattern of the femur possibly consequence of the unique postcranial anatomy of the species.

Variation in humeral morphology among hominoids has long been recognized in relation to both phylogeny and behavior. Here, we use 3D landmark data to analyze humeral shape among hominoids, including hylobatids (n = 37), Pongo (n = 33), Homo (n = 74), Pan (n = 55), and Gorilla (n = 45) to examine the relative influence of phylogenetic history vs. locomotor adaptation on humeral shape. Principal components analysis (PCA) of Procrustes shape data derived from 19 humeral type II or type III landmarks (Bookstein, 1991) for these taxa reveals the following: PC1, which primarily reflects the humeral torsion (or lack thereof) and relative diaphyseal and epiphyseal breadths, separates the relatively narrow-shafted, small articular dimensions and low humeral torsion Hylobates, and to a lesser extent, Pongo, humeri from those of the African hominoids. PC2, which largely contrasts shafts that are posteriorly convex (high PC2 scores) with antero-posteriorly straight humeral shafts (low PC2 scores) separates Homo, who tend to have A-P straighter shafts, from the more A-P bowed humeral shafts of the apes. These shape patterns suggest that the bowed shafts of Pan, Pongo, and Gorilla (and to a lesser extent, hylobatids) are due to the fact that in each of these taxa, the humerus is a weight-bearing bone, whereas the shafts of Homo are freed from locomotion. More subtle behavioral indicators are also elucidated, whereas cluster analyses (minimum spanning tree fit to a principal coordinates [PCO] plot and UPGMA dendrogram) reveal strong phylogenetic signals in the hominoid humerus as well.

Variation in humeral morphology among hominoids has long been recognized in relation to both phylogeny and behavior. Here, we use 3D landmark data to analyze humeral shape among hominoids, including hylobatids (n = 37), Pongo (n = 33), Homo (n = 74), Pan (n = 55), and Gorilla (n = 45) to examine the relative influence of phylogenetic history vs. locomotor adaptation on humeral shape. Principal components analysis (PCA) of Procrustes shape data derived from 19 humeral type II or type III landmarks (Bookstein, 1991) for these taxa reveals the following: PC1, which primarily reflects the humeral torsion (or lack thereof) and relative diaphyseal and epiphyseal breadths, separates the relatively narrow-shafted, small articular dimensions and low humeral torsion Hylobates, and to a lesser extent, Pongo, humeri from those of the African hominoids. PC2, which largely contrasts shafts that are posteriorly convex (high PC2 scores) with antero-posteriorly straight humeral shafts (low PC2 scores) separates Homo, who tend to have A-P straighter shafts, from the more A-P bowed humeral shafts of the apes. These shape patterns suggest that the bowed shafts of Pan, Pongo, and Gorilla (and to a lesser extent, hylobatids) are due to the fact that in each of these taxa, the humerus is a weight-bearing bone, whereas the shafts of Homo are freed from locomotion. More subtle behavioral indicators are also elucidated, whereas cluster analyses (minimum spanning tree fit to a principal coordinates [PCO] plot and UPGMA dendrogram) reveal strong phylogenetic signals in the hominoid humerus as well.