Dietary behaviours of Neanderthals

In this post we present information on the dietary behaviours of Neanderthals extracted from several published papers.

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Carbon and nitrogen isotopic ratios of fossil bone collagen reflect those of the average diet, and can be preserved for tens of thousands of years under favorable conditions. Twelve European Neanderthal bones ranging in age from 100,000 to 32,000 years old have yielded reliable collagen. For this well-preserved collagen, isotopic signatures offer the possibility to reconstruct the dietary habits of Neanderthals. The degree of interpretation of the isotopic results depends on the paleoecological context, especially on the knowledge of the available food resources and their isotopic signatures. Animal bones associated with the studied human remains provide the most reliable source for such information. In addition, isotopic data from animal bones can be retrieved from nearby sites of similar age if they are not present in the hominid site. However, the precision of the interpretation decreases when difference in distance and age between hominids and fauna increases.

Dietary subsistence is a major component of the biology of a human population. Neanderthals became extinct around 30,000 years ago, but their subsistence strategies have attracted a lot of scientific attention. An accurate estimate of their subsistence strategies is crucial to evaluate the cognitive abilities of these hominids. Numerous works, based on different approaches such as zooarcheology and tooth wear patterns have provided convincing evidence for a diet largely oriented towards the meat of large terrestrial herbivores (e.g., Lalueza Fox and Pérez-Pérez, 1993; Gaudzinski, 1996). In this context, the first isotopic study of Neanderthal bone collagen published a decade and a half ago that showed isotopic results close to those of animal predators such as wolves and hyenas did not come as a surprise (Bocherens et al., 1991). Many additional specimens have been analyzed since this first study.

(Source: “Neanderthal Dietary Habits: Review of the Isotopic Evidence”, by Hervé Bocherens, 2009)

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During recent decades, Neanderthal diet has been a major research topic in palaeoanthropology. This has been accelerated by the maturation of different techniques, which have produced a plethora of new information. However, this proliferation of data has led to confusing and contradictory results. Furthermore, most of the ecological dietary studies have been carried out on specimens drawn from different time periods and regions, almost exclusively those characterized by cold, open environmental conditions. Subsistence models based on these fragmentary data have been applied to Neanderthals living in a variety of different regions and environments, even though their dietary strategies may have been as variable as regions they inhabited. In this paper we integrate different dietary approaches (studies of the zooarchaeology, stable isotopes and plant remains) from the central and southeastern Mediterranean coast of Iberia in order to develop a broader and more complex picture of Neanderthal diet in different Mediterranean environmental conditions. Our results suggest that there may have been some minor dietary variation due to climatic or environmental differences, but that Neanderthal diet focussed on large terrestrial game, supplemented by plant foods when these were available.

The dietary behaviours of Neanderthals and the relationship between their diet and their social organization, technological abilities, and even their eventual disappearance, have been hotly debated in the anthropological literature. Examining the diets of Neanderthals requires the synthesis of data that have been collected by numerous researchers over the last 150 years. In many cases, the differences between Neanderthal and early modern human diets have been emphasized, at the expense of cogent discussion of the variation within each group. Early modern humans lived in a wide variety of environmental contexts and likely pursued a variety of behaviours throughout this range of environments. The exclusively Eurasian distribution of Neanderthals did not include as many environments, but neither was it the exclusively cold Arctic climate that many have supposed it to be. A more detailed examination of the Neanderthal range and the effects of shifting global climates reveals that this group would have been exposed to a diverse range of environments (Hoffecker, 2009; MacDonald et al., 2009). Just as modern human dietary behaviour likely varied with environments, so, too, can we predict that Neanderthals had different diets in different environments.

Our understanding of the diets of Neanderthals comes from three main lines of evidence. The first is the study of the human remains themselves, including the isotope signal recorded in their bones, the microwear on their teeth, and how factors of their lifehistory are related to diet. The second line of evidence includes the study of the remains of their food, in the form of animal and plant remains preserved at their sites. The third type of evidence comprises archaeological remains in the forms of tools and material culture and how these are related to food procurement and processing. This variety of dietary information is often confusing and, in places, contradictory. Data from different geographical areas and from different research methods provide different details about Neanderthal diets. However, when all of the data is examined rather coarsely, it appears likely that, generally-speaking, Neanderthals had a narrow diet and focused their attentions on large game, whereas a wider variety of foods characterized the diet of early modern humans. The details of exactly how the diet changed, and why, are still a subject of considerable debate.

(Source: “Neanderthal diets in central and southeastern Mediterranean Iberia”, by  Domingo C. Salazar-García et al., 2013)

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Neanderthals have been commonly depicted as top predators who met their nutritional needs by focusing entirely on meat. This information mostly derives from faunal assemblage analyses and stable isotope studies: methods that tend to underestimate plant consumption and overestimate the intake of animal proteins. Several studies in fact demonstrate that there is a physiological limit to the amount of animal proteins that can be consumed: exceeding these values causes protein toxicity that can be particularly dangerous to pregnant women and newborns. Consequently, to avoid food poisoning from meat-based diets, Neanderthals must have incorporated alternative food sources in their daily diets, including plant materials as well.

Neanderthals are undoubtedly the most studied and best-known group in the human fossil record. Despite that, for more than 100 years since their discovery, research on Neanderthal ecology, subsistence strategies, and diet have received remarkably little attention (Ready, 2010). It is only with the emergence of new archeological disciplines and development of innovative analytical approaches in the 1960s, that scholars began to look at Neanderthal behavior and adaptations to their environment. Methods such as faunal analysis, lithic technology, and taphonomic studies progressively led to a general portrait that defined Neanderthals as a homogenous group, with similar nutritional needs typified by a reliance on the consumption of terrestrial animals. This idea was reinforced by the study of stable isotopes and Neanderthal anatomy. The general robusticity of Neanderthal skeletons, with relatively short limbs and heavy trunks, has been interpreted as adaptation to cold stress environments that follow the ecogeographic principles of Bergmann’s and Allen’s rules (e.g., Trinkaus, 1981; Ruff, 1994; Holliday, 1997). Yet, a number of traits, such as the hyper-barrel-shape of the thorax, the thickened cortical bone and pronounced areas of muscle attachment, the length of the calcaneus and the large nasal piriform aperture, are thought to reflect extremely high levels of physical activity (e.g., Trinkaus, 1986; Franciscus and Churchill, 2002; Weinstein, 2008; Raichlen et al., 2011).

Such activity patterns, together with the physiological stress induced by cold climates, would have imposed significant high-energy demands on Neanderthal bodies (Sorensen and Leonard, 2001; Steegmann et al., 2002; Churchill, 2006): metabolic requirements that would have forced Neanderthals to consume an energy-rich diet. Support for this argument mostly derived from stable isotopic analyses (e.g., Richards and Trinkaus, 2009), which suggested that Neanderthals obtained nearly all of their dietary proteins from animal sources, and occupied the higher level of the food trophic web, similar to the top-level carnivores of that time. The traditional interpretation of Neanderthals as exclusively meat-eaters, has, however, been challenged on several fronts over recent decades. Mounting evidence suggestst hat the nitrogen isotope record may underestimate the consumption of starch/carbohydrate rich plant foods by humans (Ambrose et al., 2003; Hedges and Reynard,2007; Sponheimer and Lee-Thorp, 2007). More importantly, analyses of new archaeological evidence from the Mediterranean coastlines of Southern Europe and the Near East, dental wear studies, and microfossils in calculus, have demonstrated that Neanderthals were exploiting a wide variety of food sources (e.g. Stringeret al., 2008; El Zaatari et al., 2011; Fiorenza et al.,2011a; Henry et al., 2011; Salazar-García et al., 2013; Sistiaga et al., 2014). This unexpected dietary complexity, coupled with a very wide temporal and geographic range, has led many to reconsider the ecology and foraging strategies of Neanderthals.

(Source: “To Meat or Not to Meat? New Perspectives on Neanderthal Ecology”, by Luca Fiorenza et al., 2015)

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Neanderthals have been traditionally considered at the top of the food chain with a diet mostly consisting of animal proteins. New findings challenged this view and suggested that Neanderthals living in areas with more favourable climatic conditions exploited various food sources, including plant materials. In this study, the attention is focused on dental macrowear of Neanderthals from Central Italy, whose diet has been largely unexplored. Three-dimensional digital models of teeth have been examined through occlusal fingerprint analysis (OFA), a method used to understand how wear facets are formed. The results show a close similarity between the specimens of Saccopastore 1 and 2, with a wear pattern that indicates the use of diverse sources of food, but with a predominance of animal proteins. On the other hand, the specimens of Guattari 2 and 3 display a slightly different dental wear from each other, which probably reflects the chronological sequence of the Guattari Cave. It appears that at the end of the marine isotope stage (MIS) 5 the occupants of this cave consumed marginally more plant foods, while during MIS 3 they relied more on animal proteins. Finally, a close look at the Saccopastore maxillary molars reveals the presence of a distinct type of wear that has been previously described in some Neanderthals and early Homo sapiens from Near East, and it provides additional information about the culture and lifestyle of these Pleistocene human populations.

The Middle and Late Pleistocene were characterised by severe climatic fluctuations with alternating phases of glacial and interglacial periods. Such climatic instability produced dramatic environmental changes that shaped the dispersion and evolution of European hominins (Hublin, 1998, 2009; Stewart & Stringer, 2012). The emergence of glacial events determined local extinctions at higher latitudes, forcing human populations to retreat towards refugial environments in southern Europe (Dennell et al., 2011; Jennings et al., 2011).

These areas were characterised by a high biodiversity and elevated habitat heterogeneity, which provided abundant and variable food sources. Although Neanderthals have been traditionally viewed as hunter-gatherers with a diet exclusively
composed of meat, recent studies demonstrated that the populations from Mediterranean and Near Eastern regions relied on broader dietary spectrum (Stiner, 1994; Albert et al., 200; Barton, 2000; Hardi, 2001, 204, 2010; Madella et al., 2002; Stringer et al., 2008; Henry et al., 2010, 2014; El Zaatari et al., 2011; Fiorenza et al., 2011a; Hardy & Moncel, 2011; Salazar-Garci et al., 2013). In this context, the Italian peninsula with its peculiar geography and ecology is of great evolutionary importance and may help us to better understand the role of refugia and climate change during the last 150,000 years. Although Italy is distinguished by an abundant fossil record (with numerous Neanderthal remains), we still know very little about the subsistence strategies adopted by these extinct human populations. This study therefore aims to reconstruct the diet of Neanderthals from Central Italy by looking at their dental wear.

The OFA method is an important tool to understand how dental wear facets are formed.
Because there is a close relationship between masticatory food processes, jaw movements, and attritional wear, it is possible to obtain information about the diets of past human populations through wear facets analysis. Although the Neanderthal sample size of this study is rather small, it was possible to show their flexible dietary habits and the importance of the Mediterranean coastal areas, in terms of natural resources, for the evolutionary dynamics of these Pleistocene humans.

The macrowear patterns of the two Neanderthal specimens from Saccopastore suggest the intake of various food sources but with higher proportions of animal proteins, which is consistent with the presence of cold habitats, such as steppe and grasslands. On the other hand the macrowear patterns of Guattari 2 and 3 reflect their chronological sequence, indicating the use of a broader dietary spectrum during warmer periods and increase of meat consumption at colder MIS stages. Finally, the identification of an unusual type of wear in Saccopastore maxillary molars, which is not related to a normal chewing behavior, permits to extrapolate few information about the cultural habits of these Neanderthals from Central Italy, suggesting the use of posterior teeth as tools for daily task activities.

(Source: “Reconstructing diet and behaviour of Neanderthals from Central Italy through dental macrowear analysis”, by Luca Fiorenza, 2015)

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Here we describe the shotgun-sequencing of ancient DNA from five specimens of Neanderthal calcified dental plaque (calculus) and the characterization of regional differences in Neanderthal ecology. At Spy cave, Belgium, Neanderthal diet was heavily meat based and included woolly rhinoceros and wild sheep (mouflon), characteristic of a steppe environment. In contrast, no meat was detected in the diet of Neanderthals from El Sidrón cave, Spain, and dietary components of mushrooms, pine nuts, and moss reflected forest gathering. Differences in diet were also linked to an overall shift in the oral bacterial community (microbiota) and suggested that meat consumption contributed to substantial variation within Neanderthal microbiota. Evidence for self-medication was detected in an El Sidrón Neanderthal with a dental abscess and a chronic gastrointestinal pathogen (Enterocytozoon bieneusi). Metagenomic data from this individual also contained a nearly complete genome of the archaeal commensal Methanobrevibacter oralis (10.2× depth of coverage)—the oldest draft microbial genome generated to date, at around 48,000 years old. DNA preserved within dental calculus represents a notable source of information about the behaviour and health of ancient hominin specimens, as well as a unique system that is useful for the study of long-term microbial evolution.

(Source: “Neanderthal behaviour, diet, and disease inferred from ancient DNA in dental calculus”, by Laura S. Weyrich et al., 2017)

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Correlating cultural, technological and ecological aspects of both Upper Pleistocene modern humans (UPMHs) and Neandertals provides a useful approach for achieving robust predictions about what makes us human. Here we present ecological information for a period of special relevance in human evolution, the time of replacement of Neandertals by modern humans during the Late Pleistocene in Europe. Using the stable isotopic approach, we shed light on aspects of diet and mobility of the late Neandertals and UPMHs from the cave sites of the Troisième caverne of Goyet and Spy in Belgium. We demonstrate that their diet was essentially similar, relying on the same terrestrial herbivores, whereas mobility strategies indicate considerable differences between Neandertal groups, as well as in comparison to UPMHs. Our results indicate that UPMHs exploited their environment to a greater extent than Neandertals and support the hypothesis that UPMHs had a substantial impact not only on the population dynamics of large mammals but also on the whole structure of the ecosystem since their initial arrival in Europe.

Consuming the same diet at different time periods might lead to different nitrogen isotopic values, which would result in incorrect and misleading conclusions about the dietary strategies of humans.

The Troisième caverne of Goyet site (Belgium) provides a unique opportunity to fill the gap because skeletal remains of both types of humans representing several individuals have been recovered, directly radiocarbon dated, and their taxonomic attribution confirmed by palaeogenetic analysis

The Goyet human remains were associated with a rich faunal assemblage, and both the human and faunal skeletal specimens are biochemically well preserved. These circumstances make this site a key location for the understanding of the ecological behavior of the last Neandertals and UPMHs close to the time of replacement between 45,000 and 35,000 years BP in Europe. Here we applied a Bayesian mixing model (SIAR) 30 using carbon and nitrogen stable isotopic data of bone collagen to determine the relative proportions of different prey species in the diet of the UPMHs (this study) and Neandertals from the Troisième caverne of Goyet.

Given that mobility is a key variable, especially in relation to how interactions within the ecosystem occurred in hunter-gatherer societies we also investigate the mobility history using sulphur isotopic composition in bone collagen within and across two Neandertal groups, one from Spy and the other from Goyet, as well as from the UPMHs from Goyet. In the context of evaluating potential group mobility aspects, it is of special interest that the Goyet Neandertals show features of intensive cannibalism on their highly fragmented bones, this being in contrast to the Spy Neandertals.

The isotopic signatures of the Goyet UPMHs and of the Neandertals are similar, both indicating a purely terrestrial diet and a similar preference for particular terrestrial herbivores.

The zooarchaeological records from Goyet and Spy fully support mammoth hunting episodes with a special preference for younger individuals and possibly their mothers.

Interestingly, based on stable isotopes, the mammoth seems to contribute the major part of the dietary protein of humans in a time range between 50,000 and 30,000 years ago and across wide areas spanning from SW France to the Crimean Peninsula.

Analysing the bulk collagen fraction underestimates the plant protein contribution to the diet, but another approach more sensitive to plant food intake indicates a substantial amount of plant protein in the diet of the Spy Neandertals. This supports rather broader subsistence strategies for late Neandertals than previously considered in a palaeoecological context typical of the MIS 3.

The application of a sulphur isotopic approach appears to be an adequate tool for retrieving information of spatial aspects among humans in Late Pleistocene archeological contexts. A broader dietary spectrum for UPMHs cannot be stated as being a substantial reason for the success of one human type over the other; instead, it seems necessary to investigate further the possibility of different concepts of landscape utilization that could have given UPMHs the edge over Neandertals.

(Source: “Stable isotopes reveal patterns of diet and mobility in the last Neandertals and first modern humans in Europe”, by Christoph Wißing et al., 2019)

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Neanderthals are a Late Pleistocene hominin adapted to cool high-latitudes environments. Popular views on how Neanderthals adapted to these environments have changed over time. While once thought of as a largely scavenging hominin, Neanderthals are now accepted to be competent hunters who sourced a major part of their nutrition from ungulates. Neanderthal diet appears to be highly terrestrial but there are difficulties ruling out a contribution of marine foods in many regions. While the important role of large and medium ungulates in debates about Neanderthal diet has largely been settled, recent discussions about Neanderthal diet have explored the extent of their diet varied, the role of minor foods (plant, small mammal and marine foods) and dietary flexibility.

In this article, diet not only lies at the heart of Neanderthal social organisation, their mobility and their adaptations to European climates, but is also essential for modelling their behavioural modernity and understanding their disappearance and replacement by modern humans by 40 ka (Kuhn and Stiner, 2006; Salazar-Garcíaet al., 2013; Stiner, 2013). Neanderthal diet has been considered to be centred on a more restricted range of resources and existed higher on the food chain than modern humans (O’Connell, 2006; Stiner, 2013; Stiner and Kuhn, 2009). Occupying a higher trophic position may have burdened Neanderthals in the sense they were more highly susceptible to periodic interruptions in food supply.

However, understanding Neanderthal diet and placing them in their ecological context are fraught with challenges. Fossil evidence shows that they predominantly originated in Western Eurasia, but their range shifted over time. During the considerable climatic warming of the last Interglacial Period, Neanderthals expanded deep into central Asia (Hoffecker, 2002). While the northern and southern margin of their range also appears to have varied considerably according to climatic cycles (Hublinand Roebroeks, 2009). Although Eurasia shares some ecological commonalities due to biogeography, the region is topographically heterogeneous, and this is reflected in ecology distribution.

Furthermore, cold glacial and warm interglacial cycles drove vast environmental shifts in these territories between warm/cooland open grassy/closed forested habitats, with corresponding changes in the available animal and plant species (van Andeland Tzedakis, 1996). These environments were complex and dynamic, and some particularly the herbivore-rich cold steppe environments lack a contemporary analogue which serves as a significant challenge to researchers hoping to reconstruct subsistence (Stewart, 2005). This review examines the broad lessons that can be learned from recent multidisciplinary research on Neanderthal diet. It also emphasises on how their diets compared with Upper Palaeolithic modern human diets to highlight alternative hominin adaptations to Eurasian environments, furthering widely those that are already widely held.

Dental micro and macrowear are another line of evidence that can inform us more about Palaeolithic diets. Wear analysis are predicated on the fact that some foods have different mechanical properties that generate contrasting patterns of wear on a tooth’s dental enamel. This approach can be widely applied on Neanderthal teeth because it is non-destructive; however, it has the disadvantage that it only provides a generalised picture of diet, because many foods have similar mechanical properties. Nonetheless, the advantage is that it is a non-destructive approach and thus analysts have been able to conduct very large-scale dental wear studies (El Zaatari et al., 2011, 2016; Fiorenza et al., 2011; Power and Williams, 2018). In this respect, dental wear gives us the broadest picture of Neanderthal diets. With the technique, researchers have been able to detect habitat-driven dietary variation as well as the use of teeth as tools to hold materials. Inferring the total contribution of plants and animals is difficult but generally, these studies infer that cold environment Neanderthals had meat-based diets and that warm-environment Neanderthals had more mixed diets. These works verify that meat and plants were a common feature of Neanderthal diet and probably an important feature in warmer forested parts of their range (El Zaatari et al., 2011; Fiorenza et al., 2015).

We know little about how Neanderthals prepared and processed food. Some argue that starch granule damage known as gelatinisation found in Neanderthal dental calculus could potentially reflect thermal alteration and thus is suggestive of cooking, but it may also reflect nonthermal degradation. Although Neanderthals show evidence of fire use, their use of fire appears to be irregular and it is not yet clear if they regularly cooked foods, or if ever. A number of lightly worn grinding stones have been found on Neanderthal sites, which may reflect low-intensity plant food grinding and mashing, but these artefacts are extremely rare and most are known from older excavations with poor stratigraphic integrity (Power and Williams, 2018).

Neanderthal foraging appears to have been largely stable in the limited areas in time and space where we hold data with the notable exceptions including adoption of marine foods and low but perceptible levels of ungulate predation pressure.

(Source: “Neanderthals and Their Diet”, by Robert C Power, 2019)

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Research-Selection for NovoScriptorium: Maximus E. Niles

 

 

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