Article

Can Carbon-13 in large herbivores reflect the canopy effect in temperate and boreal ecosystems? Evidence from modern and ancient ungulates

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Abstract

Local environmental conditions under dense canopy are known to result in depletion in 13C abundance in plants compared to an open land context. This canopy effect has been observed in tropical as well as in mid-latitude forest ecosystems. However, the impact of the canopy effect on tissue 13C abundance of temperate and boreal forest-dwelling herbivores has not been thoroughly explored. Nevertheless, the canopy effect has been suggested to explain a decrease of about 3‰ in collagen δ13C values in ancient large herbivores from western Europe during the forest expansion of the Late-Glacial–Early Holocene period (ca. 15,000–6000 cal BP). Some papers have considered the 13C decrease in large herbivore as the main result of global change in atmospheric CO2 content. A detailed review of δ13C values of large herbivores (reindeer, red deer, roe deer, and bison) from open and closed environments from high and mid-latitudes confirm that the canopy effect observed in plants is passed on to their consumers. In the Paris Basin, the decline in δ13C values of large herbivores at the Late-Glacial/Early Holocene transition around 10,000 years BP appears to be different according to the considered species, namely red deer, roe deer, and large bovines (bison and aurochs). Moreover, differences in the pattern of decrease in δ13C values are observed in red deer between French northern Alps and French Jura. These differences among species in their isotopic response through time for a given geographical location, and within species from different locations, suggest variance in ecological responses of species that are associated with the relative use of forested habitat. As a result, 13C abundances in collagen can be considered as a direct tracker of the degree of closure of the habitat of ancient herbivores.

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... The "canopy cover effect" is thereby reflected in a 13 C depletion in plants growing under the canopy of dense forest stands compared with those grown in open habitats [e.g., 14]. Since herbivore tissues record the δ 13 C values of their plant food, it is possible to identify the type of environment in which the studied animals fed: closed (forested) or open habitats [e.g., 15]. Plant δ 15 N values can vary according to the same abiotic factors as plant δ 13 C values [16][17][18][19][20] and also differ among groups of plants. ...
... To our best knowledge, there are only a few studies comparing bone collagen isotopic compositions of modern free-ranging animals inhabiting various locations of well-known climatic conditions and habitats, and there are still many questions and hypotheses that need to be verified. Drucker et al. [15] reviewed in detail the δ 13 C values of modern large herbivores from open and closed environments. They linked the δ 13 C variability with the observed "canopy effect," but other environmental factors, which potentially could have had an impact on the stable isotopic composition, were not considered. ...
... Individuals from forested areas, dwelling under the canopy, showed lower δ 13 C values than deer from more open areas. These results are concordant with the results of previous stable isotopic studies, including those one modern free-ranging cervids and bovines [15,47]. The canopy cover effect results from depletion of 13 C abundance in plants growing under a closed canopy, in poorly ventilated, more humid and shaded conditions, compared to those from open habitats [14,48]. ...
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Stable isotope analyses of bone collagen are often used in palaeoecological studies to reveal environmental conditions in the habitats of different herbivore species. However, such studies require valuable reference data, obtained from analyses of modern individuals, in habitats of well-known conditions. In this article, we present the stable carbon and nitrogen isotope composition of bone collagen from modern red deer (N = 242 individuals) dwelling in various habitats (N = 15 study sites) in Europe. We investigated which of the selected climatic and environmental factors affected the δ ¹³ C and δ ¹⁵ N values in bone collagen of the studied specimens. Among all analyzed factors, the percent of forest cover influenced the carbon isotopic composition most significantly, and decreasing forest cover caused an increase in δ ¹³ C values. The δ ¹⁵ N was positively related to the proportion of open area and (only in the coastal areas) negatively related to the distance to the seashore. Using rigorous statistical methods and a large number of samples, we confirmed that δ ¹³ C and δ ¹⁵ N values can be used as a proxy of past habitats of red deer.
... δ 15 N values are indicative of the trophic level of individuals on the food chain (Schoeninger and DeNiro 1984). The mean value for C 3 plants is − 27‰ (Smith and Epstein 1971), with a range from − 20 in dry and open habitats to − 37‰ in shaded forest habitats affected by the canopy effect (Drucker et al. 2008;Kohn 2010). Bone collagen δ 13 C values of herbivores grazing C 3 plants in north-western Europe are around − 21.5‰ (Millard et al. 2013). ...
... The δ 13 C values of cattle from the Lesser Town (ninth to eleventh centuries) point to grazing in an open habitat and not in a closed canopy environment (Drucker et al. 2008;Kohn 2010). The obtained δ 13 C values ( boundaries (− 20.6 + 0.4‰; Berthon et al. 2018). ...
... The δ 13 C values of domestic pigs (Table 3) indicate primary consumption of C 3 plants (Tieszen 1991;Millard et al. 2013) in an open grazing environment (Drucker et al. 2008;Kohn 2010). Results of isotopic measurements for the Lesser Town indicate that the dietary behaviour of domestic pigs was similar to that of the Early Medieval wild boars and domestic pigs from the Great Moravian centre at Mikulčice, situated in the valley floodplain of the Morava River (eighth to ninth centuries; Kovačiková et al. 2020), as well as domestic pigs from Prague Castle (2 nd courtyard) and the Levý Hradec stronghold, dating to the ninth to eleventh centuries (Kaupová et al. 2019). ...
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The continuity of occupation at the settlement of Prague, Czech Republic, from the ninth to fourteenth centuries makes it a good case study for investigating the evolution of animal husbandry practices during the Early and High Middle Ages. Previous archaeozoological work shows that people’s dietary behaviour in Prague underwent a transformation at the beginning of the High Middle Ages (thirteenth century). This change has been attributed to large-scale socioeconomic transformations, which may have affected animal management practices. Stable carbon (δ¹³C) and nitrogen (δ¹⁵N) isotope analysis of the tooth dentine collagen of livestock (cattle, domestic pigs, sheep and goats) from three different areas in Prague (the Lesser Town, Prague Castle and the Old Town) presented in this study provides support for this hypothesis. The range of stable isotope values increased over time, which may, in addition to a farm-household model, indicate a more advanced system of livestock supply–for example, through urban markets–and may also be explained by more diversified breeding conditions in the town and its surroundings. The results show an increase in the δ¹⁵N values in pig diet over time and suggest that their management changed. This difference may have been caused by increased animal protein intake or higher δ¹⁵N value of the dietary sources. It has also been corroborated that the root dentine collagen of High Medieval cattle has slightly higher average δ¹⁵N values when compared to cattle from the earlier period, which could be related to gradual changes in landscape utilization and soil processing, such as fertilization.
... In general, the carbon and nitrogen isotopic signal (δ 13 C and δ 15 N) of a carnivore is controlled by many factors. These include biological factors (choice of prey, spectrum of prey, and isotopic variability of prey) and indirect environmental factors, which are responsible for the isotopic signal in prey (connected to vegetation type, aquatic/terrestrial ecosystem, soil development stage, climate, altitude) 26,27,[49][50][51][52][53] . For domestic carnivores, direct or indirect anthropogenic impacts might also play an important role in shaping isotopic signals. ...
... For instance, several sites from the Baltic region (Kołobrzeg, Puck, Napole, and Visby) stood out from the whole dataset due to their high δ 13 C signal ( Fig. 4; Datasets S1 and S2). Several potential factors could be responsible for such a high δ 13 C signal: low forestation 26 , high elevation 64,65 , or significant contribution of marine food resources 54,55 . Another factor that elevates δ 13 C values might be attributed to the important contribution of C 4 plants (e.g., millet) at the basis of the trophic web. ...
... • Presence of cereal crops, cultivated in open fields, characterized by elevated δ 13 C values 26,27 , and available to synanthropic rodents (such as mice and rats, hunted by cats) either directly from fields, storages in barns or granaries, or from processed products; • Cultivation or import of C 4 cereal crops with high δ 13 C signals (such as millet or maize) 50 www.nature.com/scientificreports/ • Access to fish (and other marine resources), usually characterized by an elevated δ 15 N signal and variable δ 13 C values, depending on fish ecology and physiology (i.e., differences between freshwater and marine fish 54,55 , geography (e.g., the difference between the Baltic Sea and North Sea 47,48 , or between pelagic and littoral waters 53 ). ...
... In general, the carbon and nitrogen isotopic signal (δ 13 C and δ 15 N) of a carnivore is controlled by many factors. These include biological factors (choice of prey, spectrum of prey, and isotopic variability of prey) and indirect environmental factors, which are responsible for the isotopic signal in prey (connected to vegetation type, aquatic/terrestrial ecosystem, soil development stage, climate, altitude) 26,27,[49][50][51][52][53] . For domestic carnivores, direct or indirect anthropogenic impacts might also play an important role in shaping isotopic signals. ...
... For instance, several sites from the Baltic region (Kołobrzeg, Puck, Napole, and Visby) stood out from the whole dataset due to their high δ 13 C signal ( Fig. 4; Datasets S1 and S2). Several potential factors could be responsible for such a high δ 13 C signal: low forestation 26 , high elevation 64,65 , or significant contribution of marine food resources 54,55 . Another factor that elevates δ 13 C values might be attributed to the important contribution of C 4 plants (e.g., millet) at the basis of the trophic web. ...
... • Presence of cereal crops, cultivated in open fields, characterized by elevated δ 13 C values 26,27 , and available to synanthropic rodents (such as mice and rats, hunted by cats) either directly from fields, storages in barns or granaries, or from processed products; • Cultivation or import of C 4 cereal crops with high δ 13 C signals (such as millet or maize) 50 www.nature.com/scientificreports/ • Access to fish (and other marine resources), usually characterized by an elevated δ 15 N signal and variable δ 13 C values, depending on fish ecology and physiology (i.e., differences between freshwater and marine fish 54,55 , geography (e.g., the difference between the Baltic Sea and North Sea 47,48 , or between pelagic and littoral waters 53 ). ...
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The domestic cat is the world's most popular pet and one of the most detrimental predators in terrestrial ecosystems. Effective protection of wildlife biodiversity demands detailed tracking of cat trophic ecology, and stable isotopes serve as a powerful proxy in dietary studies. However, a variable diet can make an isotopic pattern unreadable in opportunistic predators. To evaluate the usefulness of the isotopic method in cat ecology, we measured C and N isotope ratios in hundreds of archaeological cat bones. We determined trends in cat trophic paleoecology in northern Europe by exploiting population-scale patterns in animals from diverse locations. Our dataset shows a high variability of isotopic signals related to the socio-economic and/or geomorphological context. This points toward regularities in isotopic patterns across past cat populations. We provide a generalized guide to interpret the isotopic ecology of cats, emphasizing that regional isotopic baselines have a major impact on the isotopic signal.
... Stable carbon and nitrogen isotope ratios (expressed as d 13 C and d 15 N values, respectively) from analyses of sub-fossils of animals can add a dietary dimension to a paleoecological reconstruction (Drucker et al., 2008;Stevens and Hedges, 2004). d 13 C, along with d 18 O, values can be generated from the analysis of inorganic carbon in bones and teeth (Koch et al., 1997). ...
... However, C 4 plants are exceedingly rare or non-existent in the Arctic (Wooller et al., 2007). Furthermore, a lack of trees and even shrubs in Pleistocene Arctic Alaska, even during some warmer interglacials (Willerslev et al., 2014), also excludes "the canopy effect" caused by the concentration of CO 2 in dense forest (Drucker et al., 2008) and differences between herbs and shrubs (Schwartz-Narbonne et al., 2019). Thus, the main source of variability in d 13 C values we can expect in Arctic vegetation is between wetter and drier environments (Wooller et al., 2007). ...
... The d 13 C and d 15 N values from the collagen sample were À20.0‰ (±0.6) and 4.2‰ (±0.1), respectively. The d 13 C value is consistent with a diet of vegetation consists of plants using C 3 photosynthesis in a relatively open environment (Drucker et al., 2008). The d 15 N value is consistent with values from other bison specimens dated to interstadial conditions, and contrast with stadial conditions, which can have higher values Rabanus-Wallace et al., 2017). ...
Article
Detailed paleoecological evidence from Arctic Alaska’s past megafauna can help reconstruct paleoenvironmental conditions and can illustrate ecological adaptation to varying environments. We examined a rare, largely articulated and almost complete skeleton of a steppe bison (Bison priscus) recently unearthed in Northern Alaska. We used a multi-proxy paleoecological approach to reconstruct the past ecology of an individual representing a key ancient taxon. Radiocarbon dating of horn keratin revealed that the specimen has a finite radiocarbon age ~46,000 ± 1000 cal yr BP, very close to the limit of radiocarbon dating. We also employed Bayesian age modeling of the mitochondrial genome, which estimated an age of ~33,000e87,000 cal yr BP. Our taphonomic investigations show that the bison was scavenged post-mortem and infested by blowflies before burial. Stable carbon and oxygen isotope (d13C and d15N) analyses of sequentially sampled horn keratin reveal a seasonal cycle; furthermore, high d15N values during its first few years of life are consistent with patterns observed in modern bison that undertook dispersal. We compared sequential analyses of tooth enamel for strontium isotope ratios (87Sr/86Sr) to a spatial model of 87Sr/86Sr values providing evidence for dispersal across the landscape. Synthesis of the paleoecological findings indicates the specimen lived during interstadial conditions. Our multi-proxy, paleoecological approach, combining light and heavy isotope ratios along with genetic information, adds to the broader understanding of ancient bison ecology during the Late Pleistocene, indicating that ancient bison adopted different degrees of paleo-mobility according to the prevailing paleoecological conditions and climate.
... Environmental conditions, particularly the availability of water, are reflected in the δ 13 C of plants and the tissues of the herbivores that eat them, in both modern and ancient contexts in western Europe (Drucker et al., 2008). Thus, the δ 13 C values obtained from herbivore tooth enamel mirror plant cover distributions, or the proportion of plants using C3 versus C4 photosynthetic pathways, of the territories in which an individual foraged during the period of tooth mineralisation (e.g. ...
... In temperate settings, the δ 13 C composition of a plant located in open areas can be elevated by 2-5‰ in comparison with those encountered in closed forested environments, even for the same species of plants, confirming that the canopy effect operates in nontropical environments as well (e.g. Drucker et al., 2008). Thus, the δ 13 C composition of tooth enamel from non-migratory herbivores can be a measure of how open (i.e. ...
... Thus, the δ 13 C composition of tooth enamel from non-migratory herbivores can be a measure of how open (i.e. fewer trees due to drier conditions) or closed (i.e. more trees due to wetter conditions) (Drucker et al., 2008) local vegetation systems were. However, interpreting this isotopic data requires an understanding of a herbivore's ecology. ...
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Adaptation to Late Pleistocene climate change is an oft-cited potential contributor to Neanderthal disappearance in Eurasia. Accordingly, research on Neanderthal behaviour-including subsistence strategies, mobility, lithic technology, raw material procurement and demography-often 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.
... δ 13 C (‰, VPDB) and δ 15 N (‰, AIR) values of estimated consumed plants (ECP) were calculated by applying trophic corrections of 5‰ and 3‰, respectively, and ranges of values of diet-to-tissue trophic discrimination are incorporated (3-5‰ for δ 13 C; 2-5‰ for δ 15 N). We used the maximum cut-off value (δ 13 C = −27.5 ‰) commonly used to identify specimens of herbivorous mammals that consume a significant number of plants growing in closed-canopy conditions (Drucker et al., 2003(Drucker et al., , 2008Hofman-Kamińska et al., 2018). Number of samples: n; maximum: Max; minimum: Min; mean values: Mean; standard deviation: σ. ...
... δ 13 C (‰, VPDB) and δ 15 N (‰, AIR) values of estimated consumed plants (ECP) were calculated by applying trophic corrections of 5‰ and 3‰, respectively, and ranges of values of diet-to-tissue trophic discrimination are incorporated (3‰-5‰ for δ 13 C; 2‰-5‰ for δ 15 N). We used the maximum cut-off value (δ 13 C = −27.5 ‰) commonly used to identify specimens of herbivorous mammals that consume a significant number of plants growing in closed-canopy conditions (Drucker et al., 2003(Drucker et al., , 2008Hofman-Kamińska et al., 2018). Number of samples: n; maximum: Max; minimum: Min; mean values: Mean; standard deviation: σ. ...
... The δ 13 C (‰, VPDB) and δ 15 N (‰, AIR) values of estimated consumed plants (ECP) were calculated by applying trophic corrections of 5‰ and 3‰, respectively, and ranges of values of diet-to-tissue trophic discrimination are incorporated (3‰-5‰ for δ 13 C; 2‰-5‰ for δ 15 N). We used the maximum cutoff value (δ 13 C = −27.5 ‰) commonly used to identify specimens of herbivorous mammals that consume a significant number of plants growing in closed-canopy conditions (Drucker et al., 2003(Drucker et al., , 2008Hofman-Kamińska et al., 2018 Dietary ecological traits of extinct mammalian herbivores 9 ...
Article
Stable isotopes are a powerful tool for reconstructing the past. However, environmental factors not previously considered can lead to misinterpretations. Our study presents a novel analysis of the feeding behavior of the megafauna that inhabited the Pilauco ecosystem in south-central Chile during the last glacial termination. We analyzed a suite of modern plant and animal samples from closed-canopy forests to establish an isotopic baseline with which to compare stable isotope results from fossil megafauna. Using the modern samples as a reference, the δ ¹³ C results from the Pilauco megafauna indicate feeding behaviors in forested areas. These results were then calibrated with dental calculus samples and coprolites, which suggest the coexistence of graze- and grass-dominated mixed-feeder diets. The δ ¹⁵ N values found in Pilauco megafauna are not consistent with modern reference data sets or with the low δ ¹⁵ N values of extinct proboscideans from other contemporaneous and nearby sites. Probably, the δ ¹⁵ N values of the Pilauco ecosystem were not primarily affected by climate, but rather by disturbance factors (e.g., grazing effect). Our results indicate that the Pilauco megafauna fed mainly on arboreal vegetation; however, non-isotopic proxies indicate that they were also eating open vegetation (e.g., herbs and grasses).
... This pattern is caused by 1) atmospheric CO 2 under the canopy being 13 C depleted relative to the atmosphere, caused by the uptake of recycled CO 2 respired by 13 C depleted organic matter (Tieszen 1991); 2) increased light intensity associated with increasingly more open foliage improving the efficiency of photosynthesis, favouring the transfer of 13 C (Farquhar et al. 1989;Van der Merwe, Medina 1991). The net result is lower foliage δ 13 C values in canopied environments compared to the foliar values of flora growing in open environments, (c. 1 to 6‰ depending on its density (Drucker et al. 2008;Stevens et al. 2006)). Consequently, animals browsing and grazing under forest canopies exhibit low carbon isotope ratios in their skeletal tissues, with precise δ 13 C values influenced by feeding behaviour and habitat preferences. ...
... Consequently, animals browsing and grazing under forest canopies exhibit low carbon isotope ratios in their skeletal tissues, with precise δ 13 C values influenced by feeding behaviour and habitat preferences. For example, red deer feeding in minimally managed mixed temperate deciduous forest environments in Poland exhibit bone collagen δ 13 C values ranging between −24.2‰ to −21.7‰ whereas roe deer from the same environment had values between −24.8‰ to −23.2‰ (Drucker et al. 2008). The difference in range of values is due to the plasticity of red deer dietary habits in comparison to that of roe deer. ...
... The low δ 13 C values averaging -20.1‰ in cattle from Stubline and their carbon isotopic distinction from all other domestic species, but similarity to deer species, suggests that these animals were provisioned using a feeding system centred in woodlands. Present-day roe deer feeding in minimally managed mixed temperate deciduous forest environments in Poland and France have δ 13 C values ranging between −24.6‰ to −22.2‰ (adjusted for the Suess effect) (Drucker et al. 2008). ...
Chapter
Stable isotopic analysis of animal bone can elucidate how livestock were managed in the past and help define animal-environment interactions. The stable carbon (δ13C) and nitrogen (δ15N) isotopes results from wild and domestic fauna of the Late Neolithic phases of Vinča – Belo Brdo and Stubline (Serbia) suggest livestock were managed using a variety of animal pasturing practices. Domestic cattle appear to have grazed across different environments at Vinča – Belo Brdo, whereas at Stubline they may have been kept mainly within wooded or waterlogged environments. Sheep/goat and domestic pigs consumed a diet enriched in 15N at both sites; this strongly suggests these animals were raised on manured foods, which could have included plant cultivars or graze from pastures repeatedly visited by livestock. The analysis of wild species, such as deer, provides an insight into how man-made activity shaped local vegetation landscapes
... The δ 13 C values of plants vary according to their photosynthesis strategies and environmental conditions (Bender 1971). Thus, the plants of closed areas, present more negative values of δ 13 C, due to the forest depletion of 13 C or 'canopy effect' (van der Merwe and Medina 1991;Drucker et al. 2008;Tornero et al. 2020). ...
... Another noteworthy aspect of El Mirador is the low δ 13 C values (lowered to −12.8‰) observed in some individuals ( M I R _ O v i s _ 0 3 , 0 5 , a n d 0 8 ) b u t , e s p e c i a l l y i n MIR_Ovis_06, coinciding, again, with a time of falling temperatures marked by low δ 18 O values (Fig. 3). In temperate latitudes, some studies have shown that the δ 13 C values in tree species are more negative, especially in thicker forests, due to the so-called 'canopy effect' (van der Merwe and Medina 1991;Drucker et al. 2008). Similar low δ 13 C values to those of El Mirador cave have also been documented in the Early Neolithic sheep of southern France sites of Täi and Gazel . ...
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The beginning of husbandry in the Iberian Peninsula has been documented from the second half of the 6th millennium BCE and was based on the breeding of domestic caprines, mainly sheep. The first evidence of these practices comes from the Mediterranean region, but they quickly expanded inland. Previous studies have reported on the importance of the El Mirador sheepfold cave (Sierra de Atapuerca, Burgos, Spain) for understanding the process of the adoption of a farming economy in this inland region. In this study, we focus on husbandry and sheep management occurring in the 6th and 5th millennium cal BCE occupations of El Mirador cave by means of sequential oxygen (δ¹⁸O) and carbon (δ¹³C) isotope analyses in sheep molar tooth enamel. The results show a rapid adaptation and adoption of domestic sheep in the inland of the Iberian Peninsula. In El Mirador cave, well-organised sheep breeding was developed based on the concentration of births at the end of winter and early spring, for a period of ~2.64 months, which made husbandry more compatible with other economic activities. In 5th millennium levels, autumn births were also documented and may be indicative of the intervention of the shepherd in the reproductive cycle of the sheep and of a progressive adaptation of these animals to the interior of the Peninsula. All these strategies may have been supported by the recurrent occupations of the sheepfold, flocks grazing in the areas surrounding the cave and the possible use of leaf fodder in winter.
... Studies that compare herbivore isotopic compositions in ancient C 3 ecosystems to a plant baseline organized by plant type (e.g., Schwartz-Narbonne et al., 2019;Schwartz-Narbonne et al., 2021) presuppose that type is the most important predictor of a plant's isotopic compositions. An alternative approach is to put equal or greater emphasis on major environmental factors that influence plant isotopic compositions, such as the canopy effect (e.g., Drucker et al., 2008;Hofman-Kami nska et al., 2018) or ecosystem changes (e.g., Drucker et al., 2011;Metcalfe & Longstaffe, 2014). ...
... Lower plant δ 13 C values in EINP closed habitats compared to open habitats (~2‰ on average) is consistent with the well-known canopy effect, in which understory plants have significantly lower δ 13 C values than plants that make up the canopy or emergent layers, or plants that grow in open areas (e.g., Bonafini et al., 2013;Chevillat et al., 2005;Drucker et al., 2008;Van Der Merwe & Medina, 1991). The lower δ 13 C values in EINP leaves relative to seeds/flowers (~1‰ on average) is likewise in agreement with the 1-3‰ difference that has been reported in many other studies (e.g., Badeck et al., 2005;Ghashghaie & Badeck, 2014;Metcalfe & Mead, 2019). ...
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Plant isotopic baselines are critical for accurately reconstructing ancient diets and environments and for using stable isotopes to monitor ecosystem conservation. This study examines the stable carbon and nitrogen isotope compositions ( δ ¹³ C, δ ¹⁵ N) of terrestrial C 3 plants in Elk Island National Park (EINP), Alberta, Canada, with a focus on plants consumed by grazers. EINP is located in a boreal mixed woodland ecozone close to the transition area between historic wood and plains bison habitats, and is currently home to separate herds of wood and plains bison. For this study, 165 C 3 plant samples (grasses, sedges, forbs, shrubs, and horsetail) were collected from three habitat types (open, closed, and wet) during two seasons (summer and fall). There were no statistically significant differences in the δ ¹³ C or δ ¹⁵ N values of grasses, sedges, shrubs and forbs. On the other hand, plant δ ¹³ C and δ ¹⁵ N values varied among habitats and plant parts, and the values increased from summer to fall. These results have several implications for interpreting herbivore tissue isotopic compositions: (1) consuming different proportions of grasses, sedges, shrubs, and forbs might not result in isotopic niche partitioning, (2) feeding in different microhabitats or selecting different parts of the same types of plants could result in isotopic niche partitioning, and (3) seasonal isotopic changes in herbivore tissues could reflect seasonal isotopic changes in dietary plants rather than (or in addition to) changes in animal diet or physiology. In addition, the positively skewed plant δ ¹⁵ N distributions highlight the need for researchers to carefully evaluate the characteristics of their distributions prior to reporting data ( e.g ., means, standard deviations) or applying statistical models ( e.g ., parametric tests that assume normality). Overall, this study reiterates the importance of accessing ecosystem-specific isotopic baselines for addressing research questions in archaeology, paleontology, and ecology.
... During winter, the caprines from both sites exhibited low dietary δ 13 C values between −28.6‰ to −27.1‰, except for individual 7245 from Vinča-Belo Brdo. These values are comparable to ruminants dwelling in deciduous forested environments (−29.5‰ to −25.1‰) [94] and deer sampled from Vinča-Belo brdo (−29.7‰ to −25‰). Low plant foliar δ 13 C values can occur in forest with dense leaf cover caused by the canopy effect, which is caused by 13C depletion of atmospheric CO 2 under the canopy due to uptake of recycled CO 2 respired by 13 C depleted organic matter [95], a decrease in light intensity near the forest floor decreasing efficiency of photosynthesis, discriminating against the transfer of 13 C [83,84]. ...
... Our study benefits from paleoenvironment evidence from the teeth of deer, which are typically forest dwelling. Dietary δ 13 C values for red deer between −29.7‰ and −25.0‰ are similar to the range of values for modern forest dwelling ruminants [94] and mid-Holocene aurochs from Boreal environments [100]. Carbon isotope values from individual 7240 fall within that of the deer dietary range. ...
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Late Neolithic Vinča communities, spread over much of central and northern Balkans during the late sixth to mid-fifth millennium BC and characterised by unusually large and densely population centres, would have required highly organised food production systems. Zooarchaeological analysis indicates that domesticate livestock were herded, but little is known about the seasonal husbandry practices that helped ensure a steady supply of animal products to Vinča farming communities. Here, we present new stable carbon (δ13C) and oxygen (δ18O) isotopic measurements of incremental bioapatite samples from the teeth of domesticated livestock and wild herbivore teeth from two late Neolithic Vinča culture sites: Vinča-Belo brdo and Stubline (Serbia). Our results show a low variation overall within sheep and goats in terms of pasture type that may have been composed of seasonal halophyte plant communities, which have higher δ13C values due to the saline rich growing environments. Cattle feeding strategies were more variable and provided with supplementary forage, such as cut branches or leafy hay, during winter. The sharp distinction in the management of cattle and sheep/goat may be associated with the development of herding strategies that sought to balance livestock feeding behaviours with available forage or, more provocatively, the emergence of household-based control over cattle-an animal that held a central economic and symbolic role in Vinča societies.
... Heat/drought stress is known to lead to δ 13 C values as high as − 24‰ in plants growing in dry environments (Kohn, 2010). Consumption of plants growing in water-rich environments (Lynch, Hamilton and Hedges, 2008) or in dense forests (known as canopy effect), however, leads to δ 13 C values as low as − 22.5‰ in bone collagen (Van der Merwe and Medina, 1991;Drucker et al., 2008). δ 15 N values provide insight into animal diet management strategies, including weaning (Balasse and Tresset, 2002;Gillis et al., 2013) and winter foddering (Makarewicz, 2014). ...
... In Dž-II, open C 3 environments remained the main pasture for the majority of domestic and wild species, indicated by shared carbon isotope ratios across species. Additionally, cattle seem to have grazed seasonally in forests or water-rich environments (Lynch, Hamilton and Hedges, 2008) or were foddered (Van der Merwe and Medina, 1991;Drucker et al., 2008), which led to low δ 13 C values in two cases. Although Džuljunica is located close to water sources, we cannot exclude a forest component in the cattle diet because the diet of one of the roe deer (Capr4112) confirms the presence of forested areas near the site. ...
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Cattle were of great importance for the Neolithic farmers of southeastern Europe, in particular as farming expanded towards the well-watered regions of Džuljunica (ca. 6200–5500 cal. BCE), one of the earliest known Neolithic settlements in northeastern Bulgaria. The clear stratigraphy and the substantial Bos assemblage from Džuljunica Provided us with a great opportunity to investigate the beginning and evolution of cattle husbandry in the northern Balkans through stable isotope and zooarchaeological analyses. The relative abundance of Bos at Džuljunica leaves no doubt about the importance of beef and cattle herding. Mortality profiles suggest a transition in the early phases of the Neolithic from beef-oriented to mixed beef and milk production husbandry, enabled through intensified post-lactation culling. Stable carbon and oxygen isotope analysis of tooth enamel on a limited number of samples provides no evidence for an extended calving season for increasing milk availability or for vertical mobility. Stable carbon and nitrogen isotope values of bone collagen suggest that cattle were kept near the site, where C3 and C4 plants were available in summer, and that they were occasionally foddered with forest resources in the winter. Cattle experience a diachronic reduction in size on a regional scale, possibly due to farmers' choices aimed at more manageable herds consisting of smaller individuals. Restricting intermixing with local aurochs and the arrival of a new type of cattle may also have contributed to this change. Local factors or inter-regional influences may have influenced the ways cattle husbandry evolved at Džuljunica in particular and in northeastern Bulgaria more generally. More data from the region are necessary to flesh out the role of the interplay among environmental factors, local developments, and inter-regional contacts that facilitate the transfer of skills and traditions relating to the changing modes of cattle husbandry.
... Potential seasonal variations in the relative contribution of these different plant resources in sheep diet, as a tracer of mobility between the lowlands and the highlands, may be reconstructed through sequential analysis in enamel (Balasse et al., 2002;Navarrete et al., 2019;Tornero et al., 2020). Moreover, δ 13 C values also allow to characterize the feeding environment in which the animals are fed, distinguishing between plants grown in open and closed environments: the later are characterized by significantly lower stable carbon isotope ratios due to the canopy effect (Bonafini et al., 2013;Drucker et al., 2008;Drucker and Bocherens, 2009;van der Merwe and Medina, 1991). ...
... The δ 13 C values show, except for individuals CH Ovis 5 and 9, a diet consisting of C 3 plants from open environments (Bonafini et al., 2013;Drucker et al., 2008;Noe-Nygaard et al., 2005). Low intra-tooth variations (0.4 to 1.6‰) reflect a small amplitude of variation in diet δ 13 C values on the annual scale and no obvious alternation between different diets over the seasonal cycle. ...
Article
Sheep predominate the Early Neolithic faunal assemblages in the Iberian Peninsula. Their exploitation for meat and milk production made them key to the economy of these early farming societies. Management of sheep breeding season and feeding in the context of the local environment were decisive in obtaining these livestock products. This work focuses on these aspects through stable isotope and dental microwear analyses on sheep teeth from the cave of Chaves (Huesca, Spain). The results show the existence of “out of season” (autumn/early winter) sheep births in the Early Neolithic, contrasting significantly with spring lambing prevailing in Neolithic husbandries elsewhere in Europe and confirming the antiquity of a western Mediterranean characteristic in this regard. Furthermore, little changes in sheep diet throughout the year have been documented, as far as could be evidenced from stable carbon isotope ratios and dental microwear. Only two individuals showed higher variability in diet on a seasonal scale with possible contribution of C4 plants, possibly from grazing in the valley steppes at lower altitudes. Overall the results suggest good adaptation of sheep to the Pyrenean mid-altitude environment and strong zootechnical knowledge of the earliest shepherds in this area.
... Hajnalová 1999, 52). Hodnoty stabilního izotopu uhlíku mohou rovněž přiblížit, zda byla vegetace spásána ve vlhkých a zastíněných lesních biotopech nebo na otevřených a suchých stanovištích (Drucker et al. 2008;Somerville-Froehle-Schoeninger 2018, 93). ...
... Získané hodnoty δ 13 C, které se u obou lokalit neliší (obr. 4), jsou charakteristické pro rostliny původem z oblasti mírného podnebného pásma (C 3 rostliny; Smith-Epstein 1971; Kohn 2010) a otevřené pastevní prostředí s minimálním zastíněním (Drucker et al. 2008), například pole, trvalé travní porosty nebo prosvětlené lesy. Pokles hodnot δ 13 C u vzorků z Roztok, které tuto lokalitu vydělují od zbylých nalezišť (obr. ...
... Considering a +14.1‰ enrichment of 13 C between tooth enamel bioapatite of consumers and their diet [60], all-year-round grazing in open environments would result in δ 13 C values of about -11‰ in bioapatite. Plants growing in closed forests can yield δ 13 C values as low as −31‰ due to recycling of 13 C-depleted CO 2 and reduced photosynthesis due to low light levels, known as the canopy effect [61,62]. Therefore, woodland grazing and foddering would be evident by low δ 13 C values, depending on the degree of canopy density [61]. ...
... Plants growing in closed forests can yield δ 13 C values as low as −31‰ due to recycling of 13 C-depleted CO 2 and reduced photosynthesis due to low light levels, known as the canopy effect [61,62]. Therefore, woodland grazing and foddering would be evident by low δ 13 C values, depending on the degree of canopy density [61]. ...
Article
Schipluiden (3630-3380 cal BC), the earliest known year-round settlement in the Rhine-Meuse Delta in the Netherlands, is a key site for addressing the nature of Neolithic subsistence in the wetlands of northwestern Europe. A preliminary zooarchaeological study suggested that cattle husbandry was a major activity at Schipluiden. In contrast, stable carbon and nitrogen isotope analyses of human remains from the site indicated a marine-oriented diet, implying that the Mesolithic-Neolithic dietary transition continued well into the mid-4 th Millennium BC in this region. Here, we re-investigate the role and nature of cattle husbandry at Neolithic Schipluiden using mortality profiles and stable isotope analysis (δ 18 O, δ 13 C, δ 15 N) of animal bone collagen and tooth enamel. The age-at-death analysis suggests that cattle were managed for both meat and milk production. The δ 18 O and δ 13 C analysis of tooth enamel provide evidence that calving spread over five-and-a-half-months, which would have led to a longer availability of milk throughout the year. Cattle were grazing in open, marshy environments near the site and winter foddering was practiced occasionally. The faunal isotopic data also reveal that the high 15 N in human bone collagen is more likely to signal the consumption of products from cattle that grazed on 15 N-enriched salt marsh plants around the site, rather than a marine-oriented diet. This undermines the previous interpretation of the dietary practices at Schipluiden by showing that human diet in mid-4 th millennium BC Rhine-Meuse area was fully "Neolithic", based primarily on products from domesticates, especially cattle, with some input from wild terrestrial and aquatic resources available in their surroundings , contrary to what has been proposed before. Collating these results demonstrates a high level of investment in cattle husbandry, highlighting the social and economic importance of cattle at the lower Rhine-Meuse Delta during the 4 th millennium BC.
... Light attenuation, high water availability, and poor air ventilation under a dense vegetation cover, such as a closed forest canopy, lead to lower plant δ 13 C values than those observed in open landscapes (e.g., van der Merwe & Medina 1991). This so-called canopy effect has been described for several types of ecosystems, including temperate and boreal forests (e.g., Drucker et al. 2008, Bonafini et al. 2013). Finally, the 13 C abundance of atmospheric CO 2 has changed over time, ranging from ca. −7.0 to −6.4‰ between 115,000 and 11,700 cal BP (for a review, see Eggleston et al. 2016). ...
... The same review concludes with a typical pattern of mammoth and horse δ 13 C < bison and woolly rhino δ 13 C < muskox δ 13 C < reindeer δ 13 C over different time periods and regions of the Mammoth Steppe (Figure 3). The high δ 13 C values of reindeer have been recognized as the result of substantial consumption of lichen as is observed today (e.g., Fizet et al. 1995;Iacumin et al. 2000;Bocherens 2003;Drucker et al. 2003Drucker et al. , 2008Drucker et al. , 2012. For the other herbivorous species, physiological traits (ruminant versus nonruminant) and dietary flexibility (specialist versus generalist) contribute to the final distribution of δ 13 C values (Bocherens 2003. ...
Article
The Mammoth Steppe was the dominant terrestrial biome of the Northern Hemisphere during the late Pleistocene. It encompassed a nonanalog community of animals living in a cold and treeless steppe-tundra landscape. The high diversity of species, including megafauna, could be supported by a productive environment. The carbon-13 and nitrogen-15 abundances in bone collagen confirmed that the coexistence of the large herbivores was facilitated by a pronounced dietary niche partitioning, with some species relatively flexible in the exploitation of browse and graze, while others were more specialized. The isotopic abundances of carbon and nitrogen in carnivores confirm a dietary partitioning, probably based on the size of prey, with an increasingly generalist behavior emerging after the Last Glacial Maximum with notable exceptions. Isotopic investigation reveals dynamic processes of ecological displacement and replacement, shedding new light on the potential niche spectrum of extant species that are now present as relic populations. ▪ The Mammoth Steppe is an extinct nonanalog ecosystem with high productivity and biodiversity despite the cold and dry conditions of the Last Glacial Period. ▪ Stable isotopes reveal that niche partitioning among herbivores and carnivores is a dominant trait of the Mammoth Steppe. ▪ Switches in preferred prey and ecological replacement are observed among carnivores over time, with the few highly specialized predators going extinct. ▪ Warmer and more humid conditions preceding the Holocene impacted large herbivores in most regions of the Mammoth Steppe, driving some of the largest ones to extinction. Expected final online publication date for the Annual Review of Earth and Planetary Sciences, Volume 50 is May 2022. Please see http://www.annualreviews.org/page/journal/pubdates for revised estimates.
... Considering a +14.1‰ enrichment of 13 C between tooth enamel bioapatite of consumers and their diet [60], all-year-round grazing in open environments would result in δ 13 C values of about -11‰ in bioapatite. Plants growing in closed forests can yield δ 13 C values as low as −31‰ due to recycling of 13 C-depleted CO 2 and reduced photosynthesis due to low light levels, known as the canopy effect [61,62]. Therefore, woodland grazing and foddering would be evident by low δ 13 C values, depending on the degree of canopy density [61]. ...
... Plants growing in closed forests can yield δ 13 C values as low as −31‰ due to recycling of 13 C-depleted CO 2 and reduced photosynthesis due to low light levels, known as the canopy effect [61,62]. Therefore, woodland grazing and foddering would be evident by low δ 13 C values, depending on the degree of canopy density [61]. ...
Article
Full-text available
Schipluiden (3630-3380 cal BC), the earliest known year-round settlement in the Rhine-Meuse Delta in the Netherlands, is a key site for addressing the nature of Neolithic subsistence in the wetlands of northwestern Europe. A preliminary zooarchaeological study suggested that cattle husbandry was a major activity at Schipluiden. In contrast, stable carbon and nitrogen isotope analyses of human remains from the site indicated a marine-oriented diet, implying that the Mesolithic-Neolithic dietary transition continued well into the mid-4th Millennium BC in this region. Here, we re-investigate the role and nature of cattle husbandry at Neolithic Schipluiden using mortality profiles and stable isotope analysis (δ18O, δ13C, δ15N) of animal bone collagen and tooth enamel. The age-at-death analysis suggests that cattle were managed for both meat and milk production. The δ18O and δ13C analysis of tooth enamel provide evidence that calving spread over five-and-a-half-months, which would have led to a longer availability of milk throughout the year. Cattle were grazing in open, marshy environments near the site and winter foddering was practiced occasionally. The faunal isotopic data also reveal that the high 15N in human bone collagen is more likely to signal the consumption of products from cattle that grazed on 15N-enriched salt marsh plants around the site, rather than a marine-oriented diet. This undermines the previous interpretation of the dietary practices at Schipluiden by showing that human diet in mid-4th millennium BC Rhine-Meuse area was fully "Neolithic", based primarily on products from domesticates, especially cattle, with some input from wild terrestrial and aquatic resources available in their surroundings, contrary to what has been proposed before. Collating these results demonstrates a high level of investment in cattle husbandry, highlighting the social and economic importance of cattle at the lower Rhine-Meuse Delta during the 4th millennium BC.
... Because the carbon stable isotope compositions (d 13 C) of biological apatite are directly linked to the isotope compositions of the average diet of the animal (d 13 C diet ), measurements of the C-isotope compositions of the bioapatite allow for a reconstruction of the vegetative cover used as a food source (e.g., Koch, 1998;Passey et al., 2005). d 13 C diet values can provide information about the ratio of plants using different photosynthetic pathways (C 3 and C 4 type, O 'Leary, 1988;Farquhar et al., 1989;Martinelli et al., 1991), while in C 3 ecosystems information can be obtained about the vegetation "openness" (van der Merwe and Medina, 1991;Bocherens et al., 1996;Drucker et al., 2008), relative humidity or the Mean Annual Precipitation (MAP, Kohn, 2010) and habitat differences or niche partitioning between animals (e.g., Feranec and MacFadden, 2006). The oxygen isotope composition of mammalian teeth enamel is determined by the isotopic composition of the animal's body water, which, in the case of obligate drinking large herbivores reflects mainly the ingested environmental waters (e.g., Longinelli, 1984;Luz et al., 1984;Bryant et al., 1996;Fricke and O'Neil, 1996;Fricke et al., 1998;Koch, 2007). ...
... Herbs and grasses in open areas tend to have higher d 13 C values, while in forests the limited sunlight, high relative humidity, and the low d 13 C CO2 values related to the decaying organic matter in the soil lowers the d 13 C values of the undergrowth. These latter processes, in general, are collectively known as the canopy effect (van der Merwe and Medina, 1991;Bocherens et al., 1996;Drucker et al., 2008). ...
Article
There is an increasing need for paleoclimate records from continental settings to better understand the climatic changes during critical periods such as the Pliocene and Early Pleistocene. Present data indicates a transition from a warmer than present-day climate to a substantially different cooler climate. This study reviews the oxygen and carbon isotope compositions of mammalian tooth enamel for the Pliocene and Early Pleistocene of South and Central Europe to reconstruct the spatial distribution and temporal changes of the vegetation and oxygen isotope composition of precipitation (δ¹⁸Oppt). In addition to a literature review, this study adds new stable isotope measurements for this period. All δ¹³C values indicate C3 ecosystems and reflect major changes in the water use efficiency and/or in the prevailing humidity. The reconstructed major floral types range from woodland to woodland ‒ mesic grassland in all of the investigated regions. The carbon isotope compositions of fossil mammal teeth demonstrate that the spatial distribution of vegetation was broadly similar to those of the present-day for the Early/Late Pliocene – Early Pleistocene, with the most “closed” vegetation in Central and Northern Italy, while open mesic grassland vegetation covers can be reconstructed for the Iberian Peninsula, Massif Central region (Central France) and the Carpathian Basin. The calculated δ¹⁸Oppt values give a negative temporal shift of about 1–1.5‰ from the Early Pliocene to Late Pliocene – Early Pleistocene in three regions (Iberian Peninsula, Central Italy, Carpathian Basin), potentially representing a 1.5–3.0 °C decrease in mean annual temperatures (MAT) over time. In the Massif Central region and the Carpathian Basin, the δ¹⁸Oppt values are almost the same for the Late Pliocene and Early Pleistocene, while in Northern Italy the values decreased over that period. The δ¹⁸Oppt values are in the range of present-day δ¹⁸Oppt values over the Early Pliocene and somewhat lower than present-day values for the Late Pliocene - Early Pleistocene in most of the regions. Because most other proxies indicate warmer than present-day climate for the Early Pliocene and similar to present-day climate for the Early Pleistocene, the δ¹⁸Oppt values are generally lower than expected, which can be partially explained by local effects.
... Robson et al., 2016) Gazzoni, Goude, Dalmeri, Guerreschi, Mottes, Nicolis, Antonioli & Fontana Both for Mondeval de Sora and Vatte di Zambana, the inter-species difference in values for carbon (Δ 2.3‰ for both sites) and nitrogen (Δ from 0.9 to 1.8‰) distinguish populations that are more related to woodland areas (red deer) from those living mostly in open environments (chamois and ibex, figure 3). This observation could therefore be related to the different ecological areas used and the different dietary preferences of each species (Bocherens et al., 2006;Drucker et al., 2008;Schweiger et al., 2015). Red deer and ibex/chamois probably consumed food of isotopically diverse compositions, i.e. various types of trees or grasses from different habitats as demonstrated by the modern study of Schweiger et al. (2015) indicating low plant biomass for chamois, varied plant biomass for ibex and high plant biomass for red deer foraging areas. ...
... Faunal isotopic compositions reflect the environment and ecology of the animals' home range at the time the animal lived, and analysis of zooarchaeological material allows these records to be directly tied to periods when humans were active within the landscape. Faunal δ 13 C values are largely determined by dietary ecology but also respond to environmental variables such as vegetation density and type, temperature and water availability (Heaton 1999;Stevens and Hedges 2004;Drucker et al. 2008;Kohn 2010). Both faunal δ 15 N and δ 34 S values reflect underlying soil processes related to different aspects of the soil environment (e.g. ...
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Kůlna Cave is the only site in Moravia, Czech Republic, from which large assemblages of both Magdalenian and Epimagdalenian archaeological materials have been excavated from relatively secure stratified deposits. The site therefore offers the unrivalled opportunity to explore the relationship between these two archaeological phases. In this study, we undertake radiocarbon, stable isotope (carbon, nitrogen and sulphur), and ZooMS analysis of the archaeological faunal assemblage to explore the chronological and environmental context of the Magdalenian and Epimagdalenian deposits. Our results show that the Magdalenian and Epimagdalenian deposits can be understood as discrete units from one another, dating to the Late Glacial between c. 15,630 cal. BP and 14,610 cal. BP, and c. 14,140 cal. BP and 12,680 cal. BP, respectively. Stable isotope results (δ13C, δ15N, δ34S) indicate that Magdalenian and Epimagdalenian activity at Kůlna Cave occurred in very different environmental settings. Magdalenian occupation took place within a nutrient-poor landscape that was experiencing rapid changes to environmental moisture, potentially linked to permafrost thaw. In contrast, Epimagdalenian occupation occurred in a relatively stable, temperate environment composed of a mosaic of woodland and grassland habitats. The potential chronological gap between the two phases, and their associations with very different environmental conditions, calls into question whether the Epimagdalenian should be seen as a local, gradual development of the Magdalenian. It also raises the question of whether the gap in occupation at Kůlna Cave could represent a change in settlement dynamics and/or behavioural adaptations to changing environmental conditions. Supplementary information: The online version contains supplementary material available at 10.1007/s12520-020-01254-4.
... The measured isotopic signals are underpinned by dietary specialisation, animal behaviour, and environmental conditions. This approach has enabled the reconstruction of late Pleistocene palaeoenvironmental conditions at a range of archaeological and palaeontological sites (e.g., Sponheimer and Lee Thorpe, 2003;Hedges et al., 2005, Stevens andHedges, 2004;Drucker et al., 2008Drucker et al., , 2011Stevens et al., 2008Stevens et al., , 2014Szpak et al., 2010;Reade et al., 2016Reade et al., , 2020aFabre et al., 2011;Jones et al., 2018Jones et al., , 2019Britton et al., 2019). ...
Article
The position of the Banwell Bone Cave mammal assemblage zone (MAZ) in the mammalian biostratigraphy of the British Isles has been the focus of debate for decades. Dominated by fauna typical of cold environments it was originally linked to the marine oxygen isotope stage (MIS) 4 stadial (ca. 72–59 ka). Subsequently it was argued that the Banwell Bone Cave MAZ more likely relates to the temperate interstadial of MIS 5a (ca. 86–72 ka). It is envisioned that “cold fauna” such as bison and reindeer moved into Britain during stadial MIS 5b (ca. 90 ka) and were subsequently isolated by the rising sea level during MIS 5a. Here we investigate environmental conditions during the Banwell Bone Cave MAZ using bone collagen δ ¹³ C and δ ¹⁵ N and tooth enamel δ ¹⁸ O and δ ¹³ C isotope analysis. We analyse bison and reindeer from the MAZ type-site, Banwell Bone Cave. Our results show unusually high δ ¹⁵ N values, which we ascribe to arid conditions within a temperate environment. Palaeotemperature estimates derived from enamel δ ¹⁸ O indicate warm temperatures, similar to present day. These results confirm that the Banwell Bone Cave MAZ relates to a temperate interstadial and supports its correlation to MIS 5a rather than MIS 4.
... Variation in the isotopic values of the cattle is inferred in the animals of the LBA but becomes more marked during the EIA (Albizuri et al. in press) (see Fig. 9 and increasing standard deviations in Table 5). This variation can be attributed to changes in the environments frequented by the animals, since isotopic signatures vary with factors such as the aridity of the habitat, temperature, height above sea level, and even the degree of tree cover (Drucker et al. 2008;Flohr et al. 2011;Handley et al. 1999;Heaton 1999;Hobson et al. 2003;Männel et al. 2007;Mariotti et al. 1980). Environmental conditions may also have changed over time. ...
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We studied 36 dogs (Canis familiaris) from the Can Roqueta site in the Catalan pre-littoral depression (Barcelona), dated between the Late Bronze Age and the First Iron Age (1300 and 550 cal BC). We used a sample of 27 specimens to analyse the evolution of the dogs’ diet based on the carbon δ13C and nitrogen δ15N isotope composition. The results show a marked human influence in that these natural carnivores display a highly plant-based diet. The offset between canids and herbivorous ungulates does not reach the minimum established for a trophic level, which implies an input of C3 and C4 (millet) cultivated plants. Moreover, the homogeneity in the values indicates that humans prepared their dogs’ food.
... Under a dense forest canopy, the atmospheric and plant-incorporated carbon becomes 12 Cenriched due to low air exchange and low intensity of sunlight (Broadmeadow and Griffiths, 1993;Buchmann et al., 1997;Heaton, 1999;Hiyama et al., 2017). In boreal forests, vascular plants exhibit δ 13 C values between − 27‰ and − 32‰, while in open environments this signal ranges between − 24‰ and − 28‰ (Brooks et al., 1997;Marshall et al., 2007), being reflected by herbivore tissues (Drucker et al., 2008;Bocherens and Drucker, 2013;Hofman-Kamińska et al., 2018). In C 3 -dominated European Pleistocene ecosystems, the δ 13 C C values in enamel of herbivores inhabiting dense forests is expected to be below − 14.5‰ (Domingo et al., 2013). ...
Article
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The wooly rhinoceros (Coelodonta antiquitatis) and forest rhinoceros (Stephanorhinus kirchbergensis), were prominent representatives of the Middle and Late Pleistocene glacial and interglacial faunas of Eurasia. Their diet has traditionally been inferred on functional morphology of the dentition and skull. In rare cases, food remains are preserved in the fossas of the teeth or as gut content. New approaches to infer diet include the study of isotopes and mesowear. Here we apply all four methods to infer the diet of these emblematic rhinoceros' species and compare the food actually taken with the food available, as indicated by independent botanical data from the localities where the rhinoceros' fossils were found: Gorzów Wielkopolski (Eemian) and Starunia (Middle Vistulian) as well as analysis of literature data. We also made inferences on the season of death of these individuals. Our results indicate that the woolly rhino in both Europe and Asia (Siberia) was mainly a grazer, although at different times of the year and depending on the region its diet was also supplemented by leaves of shrubs and trees. According to the results of isotope studies, there were important individual variations. The data show a clear seasonal variation in the isotope composition of this rhino's diet. In contrast, Stephanorhinus kirchbergensis was a browser, though its diet included low-growing vegetation. Its habitat consisted of various types of forests, from riparian to deciduous and mixed forests, and open areas. The diet of this species consisted of selected items of vegetation, also including plants growing near both flowing and standing waters. The food remains from the fossae of the teeth indicated flexible browsing, confirming the previous interpretations based on functional morphology and stable isotopes. Long-term data from mesowear and microwear across a wider range of S. kirchbergensis fossils indicate a more mixed diet with a browsing component. The different diets of both of rhinoceros reflect not only the different habitats, but also climate changes that occurred during the Late Pleistocene.
... δ 18 O values are more positive in warm than in cold regions, and at lower than higher elevations (Mook 2002). Plants from arid zones have δ 18 O values more positive than those in humid zones and animals that inhabit grasslands or savannas show more positive δ 18 O than those living in forests (Feranec and MacFadden 2006;Drucker et al. 2008). Similarly, animals that drink water from lakes, such as horses, will have lower δ 18 O than those that obtain water only from food (Harris and Cerling 2002;Hoppe et al. 2005). ...
Article
The aim of this study is to determine the feeding habits and home range of fossil horses (Equus conversidens and E. mexicanus) from La Presita Blanca, San Luis Potosí, central México. For this purpose, we used carbon, oxygen, and strontium isotope ratios. We compared ⁸⁷Sr/⁸⁶Sr from horses with strontium ratios of La Presita Blanca plants and soils to identify if these animals were local or came from elsewhere. The δ¹³C values indicate that E. conversidens specimen consumed C4 plants and E. mexicanus was a mixed feeder. In this case, carbon isotopic values reflect only the animal’s feeding preferences and not the type of vegetation that existed in La Presita Blanca during the Pleistocene. Furthermore, δ¹⁸O values and strontium isotopic ratios show that E. conversidens was an animal that could be local to the Campo Volcánico de San Luis Potosí whereas E. mexicanus came from another region.
... Th e major cause of variation of carbon isotopes is diff erent isotope fractionation during the photosynthesis of C 3 and C 4 plants (Katzenberg, 2000). Moreover -important in forested environments and less relevant to the dry steppesgrowing conditions under a closed canopy or in open habitats and varying humidity levels cause minor deviations within the C 3 regime (Drucker et al., 2008;Kohn, 2010). Nitrogen isotopes vary primarily due to the trophic-level eff ect that causes an enrichment of 15 N along the food chain. ...
... Th e major cause of variation of carbon isotopes is diff erent isotope fractionation during the photosynthesis of C 3 and C 4 plants (Katzenberg, 2000). Moreover -important in forested environments and less relevant to the dry steppesgrowing conditions under a closed canopy or in open habitats and varying humidity levels cause minor deviations within the C 3 regime (Drucker et al., 2008;Kohn, 2010). Nitrogen isotopes vary primarily due to the trophic-level eff ect that causes an enrichment of 15 N along the food chain. ...
Article
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The flanks of the Caucasus Mountains and the steppe landscape to their north offered highly productive grasslands for Bronze Age herders and their flocks of sheep, goat, and cattle. While the archaeological evidence points to a largely pastoral lifestyle, knowledge regarding the general composition of human diets and their variation across landscapes and during the different phases of the Bronze Age is still restricted. Human and animal skeletal remains from the burial mounds that dominate the archaeological landscape and their stable isotope compositions are major sources of dietary information. Here, we present stable carbon and nitrogen isotope data of bone collagen of 105 human and 50 animal individuals from the 5th millennium BC to the Sarmatian period, with a strong focus on the Bronze Age and its cultural units including Maykop, Yamnaya, Novotitorovskaya, North Caucasian, Catacomb, post-Catacomb and late Bronze Age groups. The samples comprise all inhumations with sufficient bone preservation from five burial mound sites and a flat grave cemetery as well as subsamples from three further sites. They represent the Caucasus Mountains in the south, the piedmont zone and Kuban steppe with humid steppe and forest vegetation to its north, and more arid regions in the Caspian steppe. The stable isotope compositions of the bone collagen of humans and animals varied across the study area and reflect regional diversity in environmental conditions and diets. The data agree with meat, milk, and/or dairy products from domesticated herbivores, especially from sheep and goats having contributed substantially to human diets, as it is common for a largely pastoral economy. This observation is also in correspondence with the faunal remains observed in the graves and offerings of animals in the mound shells. In addition, foodstuffs with elevated carbon and nitrogen isotope values, such as meat of unweaned animals, fish, or plants, also contributed to human diets, especially among communities living in the more arid landscapes. The regional distinction of the animal and human data with few outliers points to mobility radii that were largely concentrated within the environmental zones in which the respective sites are located. In general, dietary variation among the cultural entities as well as regarding age, sex and archaeologically indicated social status is only weakly reflected. There is, however, some indication for a dietary shift during the Early Bronze Age Maykop period.
... Consequently, animals browsing and grazing under heavy forest canopies will exhibit low carbon isotope values in their skeletal tissues. Contemporary wild herbivores feeding in minimally managed mixed deciduous forest in temperate environments in both Poland and France exhibit bone collagen δ 13 C values ranging between − 26 and − 23‰ (Drucker et al. 2008). Forest type also appears to impact floral δ 13 C values. ...
Article
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The plant and animal components of Linearbandkeramik (LBK) subsistence systems were remarkably uniform with cattle, emmer and einkorn wheat providing the primary source of sustenance for Europe's earliest agricultural communities. This apparent homogeneity in plant and animal use has been implicitly understood to indicate corresponding similarity in the types of husbandry practices employed by LBK farmers across the entire distribution of the LBK culture. Here, we examine the results from the stable (δ 13 C/δ 15 N) isotope analysis of animal bone and cereal grains from the site of Vráble-Veľké Lehemby (Slovakia), providing new information about Linearbandkeramik farming practices in the western Carpathians. Moderately high carbon isotope values from animal bone collagen show that all livestock were pastured in open areas with no evidence of forest pasturing, previously associated with LBK settlements in northwestern Europe. High δ 15 N values measured from domesticated cereal grains suggest manuring took place at the site, while 15 N enrichment in bone collagen suggest livestock fed on agricultural by-products and possibly grains. An integrated plant-animal management system was in use at Vráble where livestock grazed on cultivation plots post-harvest. Use of such strategy would have helped fatten animals before the lean winter months while simultaneously fertilising agricultural plots with manure. This study contributes to our growing understanding that although the building blocks of LBK subsistence strategies were remarkably similar, diversity in management strategies existed across central and northwestern Europe.
... This may in part be related to the species sampled. Ibex, the most commonly sampled species, typically inhabit rocky, craggy locations, whereas red deer are more flexible in their habitats, and can inhabit woodland and reflect the canopy effect (Drucker et al., 2008(Drucker et al., , 2011. We could infer that at least some of the deer at the site were predominantly living in open environments. ...
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The article presents evidence about the Middle Palaeolithic and Middle to Upper Palaeolithic transition interval in the karst area of the Danube Gorges in the Lower Danube Basin. We review the extant data and present new evidence from two recently investigated sites found on the Serbian side of the Danube River – Tabula Traiana and Dubočka-Kozja caves. The two sites have yielded layers dating to both the Middle and Upper Palaeolithic and have been investigated by the application of modern standards of excavation and recovery along with a suite of state-of-the-art analytical procedures. The presentation focuses on micromorphological analyses of the caves’ sediments, characterisation of cryptotephra, a suite of new radiometric dates (accelerator mass spectrometry and optically stimulated luminescence) as well as proteomics (zooarchaeology by mass spectrometry) and stable isotope data in discerning patterns of human occupation of these locales over the long term.
... The ingested oxygen mostly comes from the ingested water that is present from rain water that is affected by latitude, longitude, rain quantity, but mainly temperature (Castillo et al., 1985;Dansgaard, 1964). Also, plants from warm and arid regions show a 18 O enrichment due to evapo-transpiration in comparison to those living in cold and humid areas (Drucker et al., 2008;Yakir, 1992). Herbivores inhabiting humid and closed (forest) zones show lower δ 18 O values than those living in arid and open (grassland, savannas or prairies) zones (Ambrose and DeNiro, 1986;Feranec and MacFadden, 2006). ...
... Finally, δ 13 C values between − 6.8‰ and − 1.7‰ would indicate feeding in a mixed C 3 -C 4 grassland and values higher than − 1.7‰ would indicate feeding in a pure C 4 grassland. Based on the variability in the δ 13 C of modern herbivores feeding in the same ecosystem (Drucker et al., 2008), we estimate an uncertainty for these cutoff values of ~± 1-2‰. ...
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To investigate the paleoclimate and paleoenvironment of central Italy during the Middle Pleistocene, we analyzed the carbon and oxygen stable isotope composition of the carbonate component of sequential enamel samples from twenty-four rhinoceros (Stephanorhinus spp.) teeth. The samples come from two key archeological and paleontological sites located in the Molise region: Isernia La Pineta (ILP; ∼600 ka) and Guado San Nicola (GSN; ∼400 ka). Carbon isotope ratios of enamel from both ILP and GSN indicate feeding in a pure C3 ecosystem that consisted predominantly of woodlands and/or mesic C3 grasslands with the possible sparse presence of closed canopy forests at or near ILP ∼600 ka. The average (± 1 σ) enamel δ¹³C for ILP (−13.6 ± 0.6‰) is lower than that for GSN (−12.1 ± 0.4‰), suggesting higher mean annual precipitation and lesser aridity ∼600 vs. ∼400 ka. Average intra-tooth variability in enamel δ¹³C from both sites is low (∼1‰), likely indicating seasonally uniform diets, plant carbon isotope compositions, and precipitation amounts. However, the low intra-tooth variability in enamel δ¹³C may also reflect amplitude attenuation of the isotopic signal of the plants into tooth enamel. With respect to oxygen isotopes, the average (± 1 σ) enamel δ¹⁸O for ILP (24.2 ± 0.7‰) is slightly lower than that for GSN (25.0 ± 0.7‰), likely reflecting the higher elevation of ILP as well as lesser aridity and/or slightly lower mean annual temperature in the region ∼600 vs. ∼400 ka. The δ¹⁸O values of meteoric precipitation (∼-7‰) calculated from enamel compositions are indistinguishable from the modern values, suggesting that mean annual temperatures were broadly similar to today (∼13°C). Both sites exhibit a moderate average intra-tooth variability in enamel δ¹⁸O (∼2‰), likely indicating a temperature seasonality similar or slightly decreased relative to today. The temperate climate and the increase in aridity that occurred sometime between ∼600 and ∼400 ka were potentially important factors for the human colonization of the Italian peninsula and for the cultural and behavioral evolution of the early hominins during the Middle Pleistocene.
... The gazelles presented generally mixed C 3 /C 4 signal in δ 13 C-values (-15.5 ± 1.6 ), which is different from other herbivorous wildlife reported in this study (Figure 3). The relatively positive δ 13 C-values of these gazelles (about 3.5 higher than the deer and slightly higher than the cattle and sheep/goat) demonstrate notable intake of C 4 plants and/or ecological influences such as the canopy effect and watering effect (e.g., Drucker et al., 2008;Wallace et al., 2013). Though some scholars argued that wild C 4 plants in the arid desert constitute the C 4 intake of wild herbivores in Hexi Corridor (Ma et al., 2021), it is impossible to eliminate the potential contribution of C 4 crops in these gazelles' diets. ...
Article
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Animal husbandry has been an indispensable part of human subsistence since the origin of agriculture. Along with the prehistoric cultural exchange, several kinds of major domestic animals diffused and gradually got popularized across the Eurasia. The specific geographic setting makes the Hexi Corridor in northwestern China one of the key regions to converge various types of major crops and livestock, and to witness the deep impact of novel species on local subsistence. Archeological evidence reveals an emergence of early oriental-occidental cultural communication at the opening of the local Bronze Age in Hexi Corridor, resulting in a significant shift of local subsistence. However, due to the lack of more detailed archeological evidence, the initial timing and trajectory of the transformation of livestock feeding patterns remain unclear. In this study, we reported systematic stable isotope and precise radiocarbon dating analyses on faunal remains unearthed from Huoshiliang and Ganggangwa, two Early Bronze Age settlements (ca. 4,000–3,700 BP) in middle Hexi Corridor. Our results show distinct diversification in livestock feeding patterns at ∼3,850 cal BP; in contrast with previous periods, some omnivorous livestock appear to have consumed mainly C3 foodstuff and some herbivorous livestock primarily consumed C4 plants. Combined with published stable isotope data and other archeological findings in the neighboring region, a clearer trajectory of the evolution of livestock feeding patterns has been revealed with diversified strategy amid the transformation during the Early Bronze Age in Hexi Corridor. We argued that the alteration of the local livestock feeding pattern reflects the attempt to achieve more efficient economy and sustainable society, in order to withstand the harsh arid environment in Hexi Corridor.
... Cattle, in particular, possess a wide range of δ 13 C values (− 22‰ to − 18.2‰). For example, one individual (MJ8) with a δ 13 C value of − 22‰ could have grazed in a forested environment like the Moncayo area, where depletion in 13 C occurs due to a 'canopy effect' (Drucker et al. 2008). The individual with the highest δ 13 C values may have been grazing at the river valley area where modern plants have been shown to exhibit higher δ 13 C than in mountain areas in the Ebro basin (Tornero et al. 2018), or, potentially they could have had a small contribution of C 4 to their diet. ...
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This study explores human diet and animal management strategies in multiple Late Chalcolithic and Bronze Age (2550–1200 cal BCE) sites (Moncín, Majaladares, El Estrechuelo and El Balcón) located in Muela de Borja in the Ebro Valley, Northern Iberia, through the application of stable carbon (δ¹³C) and nitrogen (δ¹⁵N) isotope analysis and zooarchaeology by mass spectrometry (ZooMS). Thirty-three animal and nine human bones were analysed across the four sites. ZooMS enabled us to identify a number of fragmented animal bones whose species representation reflected that of the wider zooarchaeological assemblage. The isotopic results complemented by a Bayesian stable isotope mixing model (BSIMM) indicate that all humans had a relatively uniform diet consisting of C3 terrestrial plant and animal resources, despite representing a range of burial sites and a broad chronology. In contrast, animals had a highly variable diet, which suggests the existence of diverse feeding and management strategies within and between species. When comparing data from these sites to other published sites of a similar chronology, we see regional patterns in δ¹³C and δ¹⁵N that reflect environmental differentiation. These results provide a deeper understanding of the diversity of human and animal dietary practices during the Late Chalcolithic and Bronze Age in the Ebro Valley and Northern Iberia.
... The δ 13 C data point to a dependence on domesticated C3-plants for these early farmers, whereas the possibility of prevalent wild plant foods and C4-plant cultivation can be excluded [20,50,51]. The δ 15 N values suggest that this domestic plant-based diet was low in leguminous plants which fix nitrogen directly from the soil [21] and was mixed with considerable amounts of animal-derived protein, most likely meat from domestic cattle, sheep, goats, and pigs [20]. ...
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Dental health is closely linked to an individual’s health and diet. This bioarcheological study presents dental caries and stable isotope data obtained from prehistoric individuals (n = 101) from three Early Neolithic sites (c. 5500-4800 BCE) in central Germany. Dental caries and ante-mortem tooth loss (AMTL) were recorded and related to life history traits such as biological sex and age at death. Further, we correlate evidence on caries to carbon and nitrogen isotope data obtained from 83 individuals to assess the relationship between diet and caries. In 68.3% of the adults, carious lesions were present, with 10.3% of teeth affected. If AMTL is considered, the values increase by about 3%. The prevalence of subadults (18.4%) was significantly lower, with 1.8% carious teeth. The number of carious teeth correlated significantly with age but not sex. The isotopic data indicated an omnivorous terrestrial diet composed of domestic plants and animal derived protein but did not correlate with the prevalence of carious lesions. The combined evidence from caries and isotope analysis suggests a prevalence of starchy foods such as cereals in the diet of these early farmers, which aligns well with observations from other Early Neolithic sites but contrasts to Late Neolithic and Early Bronze Age populations in Germany.
... The range of carbon isotope compositions calculated here (range: À29.1 to À23.6‰) is more typical of a steppe environment. Dense forest vegetation typically has more negative d 13 C (as low as À34‰) due to the influence of soil respiration associated with the canopy effect (Schleser and Jayasekera, 1985;Sternberg et al., 1989;Van der Merwe and Medin, 1989;Drucker et al., 2008;Bonafini et al., 2013). The possibility of a steppe-like environment is confirmed by the pollen data for our study site (Fig. 4). ...
Article
Marine Isotope Stage 3 (MIS 3, at 60–25 ka B.P.) was a short interstadial within the Last Glacial period. A generally warm and wet climatic condition during the MIS 3 period has been suggested by many previous studies. Quantitative climatic reconstruction and studies of seasonality, however, have been seldom attempted given the scarcity of useful proxy indicators. A full understanding the climatic characteristics during this period is hampered by their absence. This is especially the case in the East Asian monsoon region in North China, which is very sensitive to climatic change. To help fill this knowledge gap, we have determined the stable carbon (δ¹³C) and oxygen isotope (δ¹⁸O) compositions of tooth enamel from fossil mammals (Bubalus sp [buffalo] and Cervus elaphus [red deer], dated at ∼33–31 ka B.P.) recovered from Longquan Cave in North China. We use these data to constrain ecology, local air temperature and precipitation amounts during the late MIS 3. The average δ¹³C (VPDB) of tooth enamel (−14.4‰ and −8.8‰, respectively for buffalo and red deer) indicate that C3 plants comprised the main diet of animals living in this area. Pollen assemblages demonstrated that vegetation mainly consisted of grasses and herbs (∼64%) with trees only accounting for about 26%. These results indicate a landscape of open steppe with sparsely distributed trees. The reconstructed climate derived from tooth enamel δ¹⁸O was relatively colder and drier than present-day condition, with mean annual air temperature ranging from 4 to 13 °C and mean annual precipitation ranging from 360 to 670 mm. Moreover, the inferred summer and winter half-year temperatures were respectively 8–24 °C and −2 to 5 °C, which represents either similar or weaker seasonality than at the present time. The inferred cold and dry climate suggested by our study contrasts with the warm and wet conditions previously reconstructed using proxy indicators from bulk soil samples in North China. In comparison to the long-term, time-averaged climatic condition reflected by those soil proxies, the stable isotope compositions of fossil more likely recorded climatic conditions at annual or seasonal time-scale. The inferred annual cold/dry climate at Luanchuan Cave suggested by our data may indicate a short climatic cooling event from Greenland Interstadial (GIS) 5 to its following stadial. Mean annual temperature fluctuated by ∼8 °C during this interstadial-stadial cycle.
... We do not use these ages as definitive, but rather as a way of comparing experience between individuals. In order to graphically present results dentine increment values were corrected for the hair-collagen offset: approximately +1.4‰ for δ 13 C, and +2‰ for δ 15 N (Caut, Angulo, and Courchamp 2009;Drucker et al. 2008). ...
Article
Experiences of childhood in colonial New Zealand are difficult to reconstruct from the historical record alone. Many of those who came to the colony were illiterate, and the Victorian tendency to avoid discussion of pregnancy and breastfeeding practices restricts our understanding of this important period. Bioarchaeological investigation, however, has the potential to illuminate the life stories of these first-generation Pākehā (European) settlers. Here we use isotopic evidence combined with dental pathology from children interred in a historic cemetery from Otago, New Zealand, to examine colonial childhood. We show how weaning practices in the colony differ from those experienced by their emigrant parents, highlight periods of illness likely associated with the weaning process, and bring to light the potential problems caused by maternal Vitamin D deficiency in the colony.
... Group 1b 14 À20.7.8 ± 0.4 (À21.3 to À20.0) 5.5 ± 0.7 (4.1 to 6.9) Group 2a 10 À18.9 ± 0.6 (À19.9 to À18.2) 5.3 ± 0.5 (4.5 to 6.2) Group 2b 12 À17.8 ± 0.7 (À18.9 to À16.6) 6.4 ± 0.5 (5.8 to 7.3) in cooler and wetter areas, δ 13 C values are more negative (Ferrio et al., 2005). Furthermore, areas with denser tree cover show more negative δ 13 C values due to the canopy effect (Drucker et al., 2008). ...
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Can Roqueta has provided a large number of equid and dog assemblages from the Late Bronze Age to the Early Iron Age. An analysis of the stable isotopes δ13C and δ15N in 42 bone samples has made it possible to reconstruct the diet of domestic equids. The results and carpological remains show that the territory was managed for agricultural and pasture use. They indicate the cultivation of two types of highly nutritious and digestible millet, used in combination with other cereals to prepare fodder, as well as the use of natural pastures.
... C3 plants, or woody vegetation and grasses that have a cool growing season, have d 13 C values ranging between À34‰ and À23‰, with a suggested mean of À27‰ (Calvin and Benson, 1948;Hatch et al., 1967;Smith and Epstein, 1971;Ehleringer and Monson, 1993;Kohn and Cerling, 2002). The canopy effect can result in up to 5‰ differences in d 13 C in the same species at one site in pure C3 forest environments between the herbivores feeding in a heavily enclosed wooded environment or on plants from different tree elevation, and the herbivores feeding in more open parkland or grassland environments with higher evapotranspiration (Van der Merwe and Medina, 1991;Bocherens et al., 1999;Feranec and MacFadden, 2006;Drucker et al., 2008;Hofman-Kaminska et al., 2018). ...
... Without direct physical evidence of forest fodder consumption, carbon isotope ratios (δ 13 C) of animal bone and teeth can potentially identify the use of forest environments for pasture and forage [26][27][28][29][30] . This is based on the well-known 'canopy effect' principle 31 (see Methods), where relative negative δ 13 C values of bone collagen, dentine and bioapatite samples may indicate animals feeding under a dense forest canopy. ...
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Stable isotope signatures of domesticates found on archaeology sites provide information about past human behaviour, such as the evolution and adaptation of husbandry strategies. A dynamic phase in cattle husbandry evolution is during the 6th millennium BCE, where the first cattle herders of central Europe spread rapidly through diverse forested ecological niches, where little is known about pasturing strategies. Here we investigate cattle pasturing and foddering practices using a multi-regional dataset of stable isotope values (δ13C and δ18O; compound-specific stable isotopic analysis δ15N-amino acids and δ13C-dairy fats) measured from cattle bone and teeth, and pottery residues from early farming contexts, and palaeoenvironmental information. Our analysis reveals that farmers practiced different pasturing strategies with the intensive use of forested ecosystems in some areas for both graze and seasonal forage. We propose that the diversity of strategies is related to the adaptation of herding to new environments, which had a positive impact on cattle breeding and milk availability for human consumption.
... In contrast to the weaning period, a number of instances of opposing covariance have been identified in the post-weaning period, suggesting the existence of episodes of physiological stress, in some cases preceding death in childhood (MOR2, MOR4, and MOR10). There are also some possible dietary explanations for a rise in δ 15 N values without a corresponding rise in δ 13 C, such as the consumption of freshwater fish or meat from animals raised in heavily forested environments (both often showing comparatively low δ 13 C values) (Drucker et al. 2008;Dufour et al. 1999). However, these subsistence practices have not been documented among northern Iberian late prehistoric communities, and the fact that δ 15 N values in most children rise just prior to death is more plausibly explained by physiological stress. ...
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The relationship between infant and child feeding practices and early mortality is difficult to address in past societies. Here, stable carbon (δ¹³C) and nitrogen (δ¹⁵N) isotope measurements of bulk bone and sequential dentine samples of deciduous second and/or permanent first molars of four younger children, one older child, one late adolescent, and two young adults (n = 8) from Moro de Alins cave, north-eastern Iberia, are used to explore the potential impact of early-life nutrition on mortality in the Bronze Age. Isotope results are compatible with generally short exclusive breastfeeding and standard weaning periods compared to other pre-modern populations. However, there are differences in exclusive breastfeeding mean δ¹³C values and in Δ¹³C trophic shifts between exclusive breastfeeding and immediate post-weaning isotope values for those individuals who survived into adolescence and adulthood and those who did not. While the former seem to be consistent with trophic distances published for modern mother–infant pairs, the latter are above most of them. This may suggest that individuals who consumed similar foods to their mothers or suffered from less physiological stress during or after weaning had greater chances of survival during early childhood and beyond. Post-weaning seems to have been a particularly stressful period of life, where a number of instances of patterns of opposing isotopic covariance compatible with catabolic changes, often preceding death among non-survivors, are detected. This outcome shows the key role of nutritional and/or physiological status in early-life morbidity and mortality among partially and especially fully weaned children from pre-antibiotic, pre-vaccination, and poor sanitation contexts and proposes that adult survival is rooted in early life experiences, in keeping with the developmental origins of health and disease.
... This is unusual, as previous isotope studies of pre-LGM Megaloceros bone collagen from south-west France indicated them occupying a similar niche to red deer (C. elaphus) (Immel et al. 2015), with lower δ 13 C (relative to reindeer) associated with leaf feeding in forested environments (Bocherens 2003, Drucker et al. 2008. These values are therefore likely reflecting the contemporary environmental and ecological suite of this part of France (i.e. a lack of extensive woodland cover) and also serve to confirm variability in Megaloceros dietary niche, which is consistent with the suggestion that this species was a mixed/ opportunistic feeder (Chritz et al. 2009). ...
Thesis
The comprehension of cultural traditions during the Paleolithic based on the bone register is still an undeveloped approach, especially for older periods. During the Middle Paleolithic, the essential relationship of Neandertals to fat, particularly to yellow marrow, motivated our interest in testing the existence of traditional butchery practices through the gestures and techniques used to fracture bones. Analysis of the distribution of percussion marks left on long bones during marrow extraction in several series of the Middle Paleolithic from France (Abri du Maras, Saint-Marcel cave) and Italy (Riparo Tagliente) showed that there was probably standardized practices. In parallel, we carried out a very extensive experiment on the role of intuitiveness in the bone marrow extraction process. Using an original spatial analysis method applied to long bones (GIS), this experiment highlighted specific distribution models that were largely constrained by bone morphology.
... C 4 plants were absent in prehistoric Netherlands [64]. In a forested environment, a combination of deliberated factors cause more depleted δ 13 C values, known as the 'canopy effect' [65,66]. In temperate forests, δ 13 C values lower than -22.5‰ are likely to be derived from a diet composed significantly of closed-canopy vegetation [67]. ...
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Suids (Sus sp.) played a crucial role in the transition to farming in northern Europe and, like in many regions, in the Netherlands pig husbandry became an important subsistence activity at Neolithic sites. Yet little is known about wild boar palaeoecology and hunting in the Late Mesolithic Netherlands with which to contextualize this transition. This paper presents the first multi-proxy analysis of archaeological suid remains in the Netherlands. It explores human-suid interactions at the Swifterbant culture sites of Hardinxveld-Giessendam Polderweg and De Bruin (5450-4250 BC) through biometric analysis, estimation of age-at-death, and stable carbon and nitrogen isotope analysis. The results reveal targeted hunting of adult wild boar in the Late Mesolithic (5450-4850 BC), with a possible shift over time towards more juveniles. The wild boar in this period are demonstrated to be of comparably large size to contemporary northern European populations and exhibiting a wide range of dietary regimes. In the final occupational period (4450-4250 BC), small suids are present, possibly domestic pigs, but there is no evidence of pig management. This study demonstrates that the nature of human-suid interactions varied over time, which may have been connected to changing environmental conditions, human mobility, and wild boar behaviour. This study also contributes the first biometric and dietary baseline for mid-Holocene wild boar in the Netherlands.
... Given that all δ 13 C diet results fall within the range expected for C 3 plants, we interpret variation in δ 13 C diet values among the animals studied to be controlled primarily by the degree of water stress experienced by the plants upon which each species fed, as well as differential consumption of isotopically enriched C 3 plant organs such as tubers. The low δ 13 C diet values of roe deer can be attributed to their characteristic preference for closed forest habitats, reflecting the reduction of plant water stress in forest understories due to decreased light intensity (Bonafini et al., 2013;Drucker et al., 2008;Drucker & Bocherens, 2010;Farquhar et al., 1982;Rodière et al., 1996). Roe deer may have inhabited the lower westward-facing slopes surrounding Üçağızlı I. Here, damp winds and precipitation from the Mediterranean Sea increase environmental moisture. ...
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This paper investigates relationships between intervals of local environmental aridity and site occupation intensity at the Upper Paleolithic cave site of Üçağızlı I (Hatay coast, south-central Turkey) by combining a stable carbon isotope-based paleoenvironmental record with several classes of archaeological evidence. A novel method for synthesizing stable isotope data from multiple ungulate species is used to create an integrated archaeofauna-based paleoenvironmental record. This method increases the temporal resolution of the investigation in the absence of precise chronological control for some sedimentary layers and reveals patterns of habitat segregation among coeval prey taxa in each layer. The method also demonstrates significant variation in the δ13C-diet of ungulates occupying contemporaneous landscapes, reflecting the existence of multiple micro-habitats within the foraging ranges of the Paleolithic occupants. Overall, the degree of environmental aridity does not correlate with measurable changes in land use or site occupation intensity based on archaeological proxies in the Üçağızlı I sequence. One exception is the Ahmarian occupation in layer B1-3 that records the wettest environmental conditions in conjunction with a marked increase in site occupation intensity, increased dietary breadth, and evidence for meat storage practices. These patterns likely signal a reorganization of forager land-use strategies in response to a short-lived interval of especially productive environmental conditions, possibly in conjunction with reduced mobility of local foragers.
Article
The North Sea is considered a unique heritage site that yielded a huge amount of zoological and archaeological data. More than 200 palaeozoological and archaeological fossil bone samples from the North Sea bed are dated by ¹⁴ C. About 2/3 of these dated bones are Pleistocene in age; the majority of the bones are from extinct species (in particular woolly mammoth); about 1/3 of the sample date to the Holocene. The presented dataset is important in its kind, but interpretation is limited because of a lack of context of the finds. The stable isotopes ( ¹³ C, ¹⁵ N) of the dated samples provide additional information on palaeoenvironmental conditions and dietary habits in the past. We present primarily a Groningen list of data; a few fossils dated in other laboratories are included for completeness.
Article
The Late Upper Palaeolithic of Europe, particularly the Last Glacial Maximum (LGM: 26-19 kyr cal BP), was a time of dramatic climatic changes. Fauna, and the humans that preyed on them, were forced to adapt their behaviours in response to climate changes to survive. The Cantabrian Region of northern Spain was continuously inhabited during this period when many other areas of Europe were inhospitable. The site of Las Caldas (Asturias) was repeatedly occupied by hunter-gatherers during the Solutrean (26.1–20.3 kyr cal BP) and Magdalenian (18.5–14.3 kyr cal BP). This paper uses δ¹³C and δ¹⁵N stable isotope analysis of red deer (Cervus elaphus) and Spanish ibex (Capra pyrenaica) remains to reconstruct past environments, ungulate niches and habitats, and to understand how this impacted on their availability as a prey source. Results show that during the Solutrean, cold and potentially wet LGM conditions were experienced and ibex lived in lower altitudes, likely due to reduced vegetation, and even small glaciers on the higher mountain slopes. Improved environmental conditions in the Magdalenian allowed ibex to live in their preferred habitat of higher altitudes. Ameliorated Late Glacial conditions correspond with changing economic hunting strategies at the site. Humans exploited wider ecological zones, over larger distances, which coincides with population growth, and greater connectivity across the Cantabrian region. This research highlights the importance of characterising the complex interactions between humans, animals and environments during the Late Pleistocene and how they responded to rapidly changing environmental conditions.
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Exploring the role of changing climates in human evolution is currently impeded by a scarcity of climatic information at the same temporal scale as the human behaviors documented in archaeological sites. This is mainly caused by high uncertainties in the chronometric dates used to correlate long-term climatic records with archaeological deposits. One solution is to generate climatic data directly from archaeological materials representing human behavior. Here we use oxygen isotope measurements of Bos/Bison tooth enamel to reconstruct summer and winter temperatures in the Late Pleistocene when Neandertals were using the site of La Ferrassie. Our results indicate that, despite the generally cold conditions of the broader period and despite direct evidence for cold features in certain sediments at the site, Neandertals used the site predominantly when climatic conditions were mild, similar to conditions in modern day France. We suggest that due to millennial scale climate variability, the periods of human activity and their climatic characteristics may not be representative of average conditions inferred from chronological correlations with long-term climatic records. These results highlight the importance of using direct routes, such as the high-resolution archives in tooth enamel from anthropogenically accumulated faunal assemblages, to establish climatic conditions at a human scale.
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The Promontory caves (Utah) and Franktown Cave (Colorado) contain high-fidelity records of short-term occupations by groups with material culture connections to the Subarctic/Northern Plains. This research uses Promontory and Franktown bison dung, hair, hide, and bone collagen to establish local baseline carbon isotopic variability and identify leather from a distant source. The ankle wrap of one Promontory Cave 1 moccasin had a δ ¹³ C value that indicates a substantial C 4 component to the animal's diet, unlike the C 3 diets inferred from 171 other Promontory and northern Utah bison samples. We draw on a unique combination of multitissue isotopic analysis, carbon isoscapes, ancient DNA (species and sex identification), tissue turnover rates, archaeological contexts, and bison ecology to show that the high δ ¹³ C value was not likely a result of local plant consumption, bison mobility, or trade. Instead, the bison hide was likely acquired via long-distance travel to/from an area of abundant C 4 grasses far to the south or east. Expansive landscape knowledge gained through long-distance associations would have allowed Promontory caves inhabitants to make well-informed decisions about directions and routes of movement for a territorial shift, which seems to have occurred in the late thirteenth century.
Article
Karst cave investigations in the south of Haida Gwaii have opened a small window on human and paleontological components of the early post-glacial landscape. At three cave locations (K1, Gaadu Din 1 and Gaadu Din 2) our investigations recovered a paleontological record extending from ca. 13,400 to 11,000 years ago and a small number of human artifacts dating from ca. 12,600 to 11,000 years ago. The animal bones recovered are dominated by black and brown bear remains, revealing that these caves were being used for winter dens. Other species present include deer, caribou, and salmon. Domestic dog remains with a direct radiocarbon age of 13,100 years ago are the earliest indicator of human presence from the cave assemblages, and are also the earliest known domestic dog remains reported on in the Americas. Brown bear and deer disappear from the paleontological record at the end of the Pleistocene, but other species persist into the Holocene, most of which continue to thrive on the islands to this day, with the exception of caribou which were extirpated in the early 20th Century. The stone tools that we found are predominately spearpoints and fragments thereof, which were used to hunt denning bears. Additional stone tool types from the Gaadu Din Caves reveal that they were occasionally used by humans as temporary shelters. As sea level was lower than today between 13,400 and 11,000 years ago, the caves provide an alternative target for late Pleistocene archaeological prospection that does not involve subtidal work. Our research demonstrates that karst caves on Haida Gwaii provided ecological and cultural focal points during the early post-glacial period. These caves remained sufficiently stable to preserve the residues of activities including bear denning and bear hunting. With the commencement of the Holocene, the record of animal and human use of the three caves diminishes.
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En este trabajo se obtienen datos antropológicos e isotópicos a partir de los restos óseos de dos individuos de edad tardorromana a efectos de reconstruir sus condiciones de vida y las causas de su muerte. Se trata de dos esqueletos femeninos conservados en el Museo Arqueolóxico de A Coruña datados entre los siglos III a VI dC. Se han aplicado técnicas de antropología forense para determinar patologías y lesiones que puedan haber afectado a los individuos estudiados. Para el estudio de paleodieta, se ha extraído colágeno óseo que se analiza mediante espectrometría de masas de relaciones isotópicas. Las relaciones isotópicas de 13C/12C y 15N/14N obtenidas se comparan con datos procedentes de otras poblaciones contemporáneas. Los perfiles antropológicos confirman que se trata de mujeres adultas jóvenes, de posible ancestralidad caucásica, con signos de hiponutrición e infección crónica y cuyas señales isotópicas del colágeno muestran una importante contribución de los moluscos en su dieta. Estos resultados concuerdan con los datos arqueológicos disponibles, reflejando el progresivo empobrecimiento de la ciudad durante la crisis del Bajo Imperio. Open access. full text here: https://revistas.udc.es/index.php/CADLAXE/article/view/cadlaxe.2015.38.0.3682
Article
Pollen records and macrobotanical remains indicate maize was grown minimally during the Middle Woodland (200 BCE - CE 500) in eastern North America. However, previous isotopic studies have found limited evidence of its consumption by people in archaeobotanical assemblages. Hypothetically, if deer (Odocoileus virginianus Zimmermann) were feeding on maize (Zea mays L.) in gardens prior to its adoption as a human dietary staple, this may indicate a niche construction process which would later lead to shifts in hunting strategies. This construction of garden spaces or niches has implications for decreased mobility and greater investment in garden crops. To evaluate this hypothesis in the Ohio River Valley, we measured stable nitrogen (δ15N) and carbon (δ13C) isotope compositions of bulk collagen in deer bones (N = 25) from archaeological sites spanning the Middle Woodland (200 BCE - CE 400/500) to Late Precontact (CE 1400-1630). Our hypothesis was not confirmed: little (one deer of 19) evidence for maize consumption occurred in the analyzed deer remains. Five deer of 19 did indicate the possibility of feeding in deeply wooded areas, reflecting canopy effect. Dog (Canis lupus familiaris L.) bone samples (N = 2) were also analyzed for these isotopes and one indicates some consumption of maize.
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This chapter provides a discussion based on the data and assumptions made in the previous chapters. In Sect. 5.1, the author discusses the particularities of the environment of France (in the Upper Pleniglacial and Tardiglacial), which was located at the western end of the Mammoth steppe and thus explains the particular development of reindeer populations in this region of Europe. The existence of different ecosystems in distinct regions is then detailed for the period considered in France. The economic system of the societies that lived in France between 28,000 and 14,600 cal BP is discussed in Sect. 5.2 and the author defends, on the basis of the data analysed in the previous chapters, the hypothesis of a single system (reindeer system) based on the exploitation of reindeer (the animal and its antlers), a game that was not very mobile in most regions.KeywordsEcosystemsMammoth steppeZoocenosisAnimal populationsEcological nicheVegetationReindeer systemEconomic systemAnnual nomadic cycle
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Thesis
*INRA Laboratoire de Biogéochimie Isotopique Paris Diffusion du document : INRA Laboratoire de Biogéochimie Isotopique Paris Diplôme : Dr. d'Université
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