Quinoa, potatoes, and llamas fueled emergent social complexity in the Lake Titicaca Basin of the Andes.

The Lake Titicaca basin was one of the major centers for cultural development in the ancient world. This lacustrine environment is unique in the high, dry Andean altiplano, and its aquatic and terrestrial resources are thought to have contributed to the florescence of complex societies in this region. Nevertheless, it remains unclear to what extent local aquatic resources, particularly fish, and the introduced crop, maize, which can be grown in regions along the lakeshores, contributed to facilitating sustained food production and population growth, which underpinned increasing social political complexity starting in the Formative Period (1400 BCE to 500 CE) and culminating with the Tiwanaku state (500 to 1100 CE). Here, we present direct dietary evidence from stable isotope analysis of human skeletal remains spanning over two millennia, together with faunal and floral reference materials, to reconstruct foodways and ecological interactions in southern Lake Titicaca over time. Bulk stable isotope analysis, coupled with compound-specific amino acid stable isotope analysis, allows better discrimination between resources consumed across aquatic and terrestrial environments. Together, this evidence demonstrates that human diets predominantly relied on C3 plants, particularly quinoa and tubers, along with terrestrial animals, notably domestic camelids. Surprisingly, fish were not a significant source of animal protein, but a slight increase in C4 plant consumption verifies the increasing importance of maize in the Middle Horizon. These results underscore the primary role of local terrestrial food resources in securing a nutritious diet that allowed for sustained population growth, even in the face of documented climate and political change across these periods.

Evolutionary and phylogenetic insights from a nuclear genome sequence of the extinct, giant, "subfossil" koala lemur Megaladapis edwardsi.

No endemic Madagascar animal with body mass >10 kg survived a relatively recent wave of extinction on the island. From morphological and isotopic analyses of skeletal "subfossil" remains we can reconstruct some of the biology and behavioral ecology of giant lemurs (primates; up to ∼160 kg) and other extraordinary Malagasy megafauna that survived into the past millennium. Yet, much about the evolutionary biology of these now-extinct species remains unknown, along with persistent phylogenetic uncertainty in some cases. Thankfully, despite the challenges of DNA preservation in tropical and subtropical environments, technical advances have enabled the recovery of ancient DNA from some Malagasy subfossil specimens. Here, we present a nuclear genome sequence (∼2× coverage) for one of the largest extinct lemurs, the koala lemur Megaladapis edwardsi (∼85 kg). To support the testing of key phylogenetic and evolutionary hypotheses, we also generated high-coverage nuclear genomes for two extant lemurs, Eulemur rufifrons and Lepilemur mustelinus, and we aligned these sequences with previously published genomes for three other extant lemurs and 47 nonlemur vertebrates. Our phylogenetic results confirm that Megaladapis is most closely related to the extant Lemuridae (typified in our analysis by E. rufifrons) to the exclusion of L. mustelinus, which contradicts morphology-based phylogenies. Our evolutionary analyses identified significant convergent evolution between M. edwardsi and an extant folivore (a colobine monkey) and an herbivore (horse) in genes encoding proteins that function in plant toxin biodegradation and nutrient absorption. These results suggest that koala lemurs were highly adapted to a leaf-based diet, which may also explain their convergent craniodental morphology with the small-bodied folivore Lepilemur.

The game of models: Dietary reconstruction in human evolution.

Despite substantial additions to the paleontological record and unanticipated improvements in analytical techniques since the Journal of Human Evolution was first published, consensus on the diet of early hominin species remains elusive. For instance, the notable advances in the analyses of hominin dental microwear and stable isotopes have provided a plethora of data that have in some instances clouded what was once ostensibly a clear picture of dietary differentiation between and within hominin taxa. In the present study, we explore the reasons why the retrodiction of diet in human evolution has proven vexing over the last half century from the perspective of both ecological and functional-mechanical models. Such models continue to be indispensable for paleobiological reconstructions, but they often contain rigid or unstated assumptions about how primary paleontological data, such as fossils and their geological and taphonomic contexts, allow unambiguous insight into the evolutionary processes that produced them. In theoretical discussions of paleobiology, it has long been recognized that a mapping function of morphology to adaptation is not one-to-one, in the sense that a particular trait cannot necessarily be attributed to a specific selective pressure and/or behavior. This article explores how the intrinsic variability within biological systems has often been underappreciated in paleoanthropological research. For instance, to claim that derived anatomical traits represent adaptations related to stereotypical behaviors largely ignores the importance of biological roles (i.e., how anatomical traits function in the environment), a concept that depends on behavioral flexibility for its potency. Similarly, in the paleoecological context, the underrepresentation of variability within the 'edible landscapes' our hominin ancestors occupied has inhibited an adequate appreciation of early hominin dietary flexibility. Incorporating the reality of variation at organismal and ecological scales makes the practice of paleobiological reconstruction more challenging, but in return, allows for a better appreciation of the evolutionary possibilities that were open to early hominins.

The effect of diet and sociopolitical change on physiological stress and behavior in late Roman-Early Byzantine (300-700 AD) and Islamic (902-1,235 AD) populations from Ibiza, Spain.

OBJECTIVES: This study evaluated chronological changes in physiological stress and levels of habitual loading of Ibizan populations from the Late Roman-Early Byzantine (LREB) to the Islamic period (300-1,235 AD) using measures of body size and bone cross-sectional properties to compare Urban LREB, Urban Medieval Islamic, and Rural Medieval Islamic groups. It also explored the effect of diet, modeled using stable isotopes, on physiological stress levels and behavior. MATERIALS AND METHODS: The sample comprised individuals from three archeological populations: Urban Late Roman- Early Byzantine (LREB) (300-700 AD), Medieval Urban Islamic (902-1,235 AD), and Medieval Rural Islamic. Bone lengths, femoral head dimensions, and cross-sectional properties, diaphyseal products and circumferences, were compared to assess differences in body size and habitual loading in 222 adult individuals. Ordinary least squares regression evaluated the correlations between these measures and carbon (δ13 C) and nitrogen (δ15 N) stable isotope ratios in 115 individuals for whom both isotope values and osteological measures are available. RESULTS: The Medieval Rural Islamic group had shorter stature and reduced lower limb cross-sectional properties compared to the two urban groups. Limb shape differs between Urban LREB and Urban Medieval Islamic groups. Measures of body size length were positively correlated with δ13 C values in all individuals and separately in the Urban LREB and Rural Medieval Islamic groups. δ15 N showed a positive correlation with left humerus shape in the Urban LREB sample. CONCLUSIONS: The low stature and cross-sectional properties of the Medieval Rural Islamic group may be an indicator of greater physiological stress, potentially due to poorer diet. Positive correlations between measures of body size and δ13 C values further suggest that greater access to C4 resources improved diet quality. Alternatively, this relationship could indicate greater body size among migrants from areas where individuals consumed more C4 resources.

Proteomic evidence of dietary sources in ancient dental calculus.

Archaeological dental calculus has emerged as a rich source of ancient biomolecules, including proteins. Previous analyses of proteins extracted from ancient dental calculus revealed the presence of the dietary milk protein ß-lactoglobulin, providing direct evidence of dairy consumption in the archaeological record. However, the potential for calculus to preserve other food-related proteins has not yet been systematically explored. Here we analyse shotgun metaproteomic data from 100 archaeological dental calculus samples ranging from the Iron Age to the post-medieval period (eighth century BC to nineteenth century AD) in England, as well as 14 dental calculus samples from contemporary dental patients and recently deceased individuals, to characterize the range and extent of dietary proteins preserved in dental calculus. In addition to milk proteins, we detect proteomic evidence of foodstuffs such as cereals and plant products, as well as the digestive enzyme salivary amylase. We discuss the importance of optimized protein extraction methods, data analysis approaches and authentication strategies in the identification of dietary proteins from archaeological dental calculus. This study demonstrates that proteomic approaches can robustly identify foodstuffs in the archaeological record that are typically under-represented due to their poor macroscopic preservation.

Dental topography and the diet of Homo naledi.

Though late Middle Pleistocene in age, Homo naledi is characterized by a mosaic of Australopithecus-like (e.g., curved fingers, small brains) and Homo-like (e.g., elongated lower limbs) traits, which may suggest it occupied a unique ecological niche. Ecological reconstructions inform on niche occupation, and are particularly successful when using dental material. Tooth shape (via dental topography) and size were quantified for four groups of South African Plio-Pleistocene hominins (specimens of Australopithecus africanus, Paranthropus robustus, H. naledi, and Homo sp.) on relatively unworn M2s to investigate possible ecological differentiation in H. naledi relative to taxa with similar known geographical ranges. H. naledi has smaller, but higher-crowned and more wear resistant teeth than Australopithecus and Paranthropus. These results are found in both lightly and moderately worn teeth. There are no differences in tooth sharpness or complexity. Combined with the high level of dental chipping in H. naledi, this suggests that, relative to Australopithecus and Paranthropus, H. naledi consumed foods with similar fracture mechanics properties but more abrasive particles (e.g., dust, grit), which could be due to a dietary and/or environmental shift(s). The same factors that differentiate H. naledi from Australopithecus and Paranthropus may also differentiate it from Homo sp., which geologically predates it, in the same way. Compared to the great apes, all hominins have sharper teeth, indicating they consumed foods requiring higher shear forces during mastication. Despite some anatomical similarities, H. naledi likely occupied a distinct ecological niche from the South African hominins that predate it.

Dental topography of prosimian premolars predicts diet: A comparison in premolar and molar dietary classification accuracies.

OBJECTIVES: This study tests whether (1) premolar topography of extant "prosimians" (strepsirrhines and tarsiers) successfully predicts diet and (2) whether the combination of molar and premolar topography yields higher classification accuracy than using either tooth position in isolation. MATERIALS AND METHODS: Dental topographic metrics (ariaDNE, relief index, and orientation patch count rotated) were calculated for 118 individual matched-pairs of mandibular fourth premolars (P4) and second molars (M2). The sample represents 7 families and 22 genera. Tooth variables were analyzed in isolation (P4 only; M2 only), together (P4 and M2), and combined (PC1 scores of bivariate principal component analyses of P4 and M2 for each metric). Discriminant function analyses were conducted with and without a measure of size (two-dimensional surface area). RESULTS: When using topography only, "prosimian" P4 shape predicts diet with a success rate that is slightly higher than that of M2 shape. When absolute size is included, premolars and molars perform comparably well. Including both premolar and molar topography (separately or combined) improves classification accuracy for every analysis beyond considering either in isolation. Classification accuracy is highest when premolar and molar topography and size are included. DISCUSSION: Our findings indicate that molar teeth incompletely summarize the functional requirements of oral food breakdown for a given diet, and that the mechanism selecting for premolar form is more varied than what is expressed by molar teeth. Finally, our findings suggest that fossil P4s (in isolation or with the M2) can be used for meaningful dietary reconstruction of extinct primates.

The role of farming and fishing in the rise of social complexity in the Central Andes: a stable isotope perspective.

For many years, the rise of stratified societies along the Central Andean coast, known as the birthplace of Andean civilization, has been closely linked to a marine-oriented economy. This hypothesis has recently been challenged by increasing evidence of plant management and cultivation among Andean populations long before the emergence of complex societies and monumental architecture. The extent to which marine and plant-based economies were integrated and their contributions to early sedentism, population growth, and intra-community stratification, however, remain subjects of ongoing and contentious debate. Using Bayesian Mixing Models we reanalyze the previously published stable isotopes (δ15Ncollagen, δ13Ccollagen, δ13Capatite) values of 572 human individuals from 39 archaeological sites in the Central Andes dated between ca. 7000 BCE and 200 CE to reconstruct dietary regimes in probabilistic terms. Our results reveal that fish, terrestrial fauna, and cultivated plants variably contributed to the diet of prehistoric Andean populations; in coastal and middle valley settlements plant cultivation, not fishing, fueled the development of the earliest complex societies during the Formative Period (from 3000 BCE). Similarly, in the highlands the societies that built ceremonial centers show a plant-based economy. Our findings also show that maize only became a staple food (> 25% dietary contribution) in more recent phases of Andean prehistory, around 500 BCE.

Human dietary δ(15)N intake: representative data for principle food items.

Dietary analysis using δ(15)N values of human remains such as bone and hair is usually based on general principles and limited data sets. Even for modern humans, the direct ascertainment of dietary δ(15)N is difficult and laborious, due to the complexity of metabolism and nitrogen fractionation, differing dietary habits and variation of δ(15)N values of food items. The objective of this study was to summarize contemporary regional experimental and global literature data to ascertain mean representative δ(15)N values for distinct food categories. A comprehensive data set of more than 12,000 analyzed food samples was summarized from the literature. Data originated from studies dealing with (1) authenticity tracing or origin control of food items, and (2) effects of fertilization or nutrition on δ(15)N values of plants or animals. Regional German food δ(15)N values revealed no major differences compared with the mean global values derived from the literature. We found that, in contrast to other food categories, historical faunal remains of pig and poultry are significantly enriched in (15)N compared to modern samples. This difference may be due to modern industrialized breeding practices. In some food categories variations in agricultural and feeding regimens cause significant differences in δ(15)N values that may lead to misinterpretations when only limited information is available.

The histology and growth rate of Nile crocodile (Crocodylus niloticus) claws.

The histology and growth of reptilian and crocodilian claws (ungues) have been extensively studied; however, Nile crocodile (Crocodylus niloticus) claws have not received adequate attention. Furthermore, age estimations for reptilian claws remain unexplored, despite Nile crocodile claws being used in long-term dietary reconstruction studies, assuming certain age-related patterns. In this study, we investigate the histology and growth patterns of Nile crocodile claws, aiming to infer axes for sampling cornified material for radiocarbon dating and establish age estimations for crocodilian claws. Our findings reveal that Nile crocodile claws exhibit growth patterns similar to other reptilians, presenting as modified scutes/scales with an age profile along the sagittal plane. This profile starts at the basal germ matrix and progressively expands in thickness and age dorsoventrally towards the apex or "tip." Consequently, the oldest corneous material is concentrated at the most dorsal point of the claw's apex. To validate previous dietary reconstruction assumptions, we conducted radiocarbon dating on this region of the claw, which supported the idea that retained corneous material in the claws is typically relatively young (5-10 years old) due to abrasion. Our study contributes insights into the histology and growth dynamics of Nile crocodile claws, shedding light on their use in dietary reconstruction studies and emphasizing the significance of considering age-related assumptions in such investigations.

Dental erosion in archaeological human remains: A critical review of literature and proposal of a differential diagnosis protocol.

OBJECTIVE: Although studies of dental wear on archaeological human remains have largely focused on mechanical wear (attrition and abrasion) in the past, chemical wear (erosion) is being increasingly identified as a separate form of wear. This paper aims to review the current state of research and to develop a protocol that may be universally used by biorchaeologists to specifically identify dental erosion. DESIGN AND RESULTS: A critical review of literature has been done in order to highlight the issues related to diagnosis of dental erosion in archaeological human remains. The bodies of work based on the analysis of both modern and archaeological dentitions raise their separate problems. In addition to a need to re-evaluate symptoms of dental erosion, notably dentin 'cupping', it is apparent that no specific protocol is adapted from medical to archaeological sciences. Authors rather rely on tooth wear indices and photographs of modern clinical cases for diagnosis. Furthermore, the diagenetic chemical alternation has rarely been considered as a bias. CONCLUSIONS: Here we suggest a three-step protocol: the primary method is the microscopic identification of dental erosion by SEM, followed by the exclusion of taphonomic aetiology on surrounding bone and soil pH analysis. Archaeologists should also explore possible causative agents of wear using archaeological and historic knowledge about the population being analyzed.