The first appearance of the Neolithic Linearbandkeramik (LBK) in Central Germany occurred during ... more The first appearance of the Neolithic Linearbandkeramik (LBK) in Central Germany occurred during the 6 th millennium BC. However, though LBK sites are abundant in the German loess areas, there are only a few studies that reconstruct the diet of these first farmers using biochemical methods. Here we present the largest study undertaken to date on LBK material using stable isotope analysis of carbon and nitrogen to reconstruct human diet and animal husbandry strategies. We analysed the bone collagen of 97 human individuals and 45 associated animals from the sites of Derenburg, Halberstadt and Karsdorf in the Middle Elbe-Saale region of Central Germany. Mean adult human values are-19.9 ±0.4 ‰ for δ 13 C and 8.7 ±0.8‰ for δ 15 N. The δ 13 C values are typical for terrestrial, temperate European regions, whereas the δ 15 N values fall within an expected range for farming societies with a mixed diet consisting of products from domestic animals and plants. The consumption of unfermented dairy products is unlikely as there is direct palaeogenetic evidence of lactose intolerance available for one of the sites. There are no clear indications for dietary differences in sex. Young children under three years of age are enriched in δ 15 N due to breastfeeding indicating that weaning likely occurred around the age of three years. The fauna exhibit mean δ 13 C values of-20.9 ±0.8‰ and mean δ 15 N values of 7.0 ±0.9‰ respectively. Variation in the δ 13 C and δ 15 N in the domestic animals is probably caused by different livestock managements.
The feeding ecology of sympatric great ape species yields valuable information for palaeodietary ... more The feeding ecology of sympatric great ape species yields valuable information for palaeodietary reconstructions in sympatric early hominin species. However, no isotopic references on sympatrically living apes and their feeding ecology are currently available. Here we present the first isotopic study on sympatric great apes, namely western lowland gorillas (Gorilla gorilla gorilla) and central chimpanzees (Pan troglodytes troglodytes) from Loango National Park, Gabon. We successfully analyzed the stable carbon and nitrogen isotope ratios in a selection of food plants (n ¼ 31) and hair samples (n ¼ 30) retrieved from sleeping nests to test whether niche partitioning among sympatric chimpanzees and gorillas is detectable using isotope analysis of hair. Ape hair strands with roots were sectioned into sequential segments (n ¼ 100) to investigate temporal isotopic variation related to seasonal variations in food resources. We found significant d 13 C differences between herbaceous plants and fruits, most likely due to canopy effects. While the d 13 C values of chimpanzees indicate the consumption of fruit, the low d 13 C values in gorilla hair indicate folivory, most likely the consumption of 13 C-depleted herbaceous vegetation. Our isotopic data also confirmed dietary overlap between chimpanzees and gorillas, which varied by season. Gorillas showed significant variation in d 13 C values in response to season due to shifting proportions of herbaceous plants versus fruits. In chimpanzees, significant seasonal variation in d 15 N was likely related to the seasonal availability of fruit species with particularly high d 15 N values. In summary, we found isotopic evidence for niche partitioning and seasonal dietary variation among sympatric great apes at Loango. These findings provide a valuable reference for palaeodietary research on fossil hominins using d 13 C analyses, particularly for studies focusing on sympatric taxa and on temporal isotopic variation within incremental tissues such as tooth enamel.
Stable isotope analysis of carbon and nitrogen in hair provides a versatile tool for reconstructi... more Stable isotope analysis of carbon and nitrogen in hair provides a versatile tool for reconstructing feeding behavior in elusive primate species. Particularly in great apes, researchers can sample long hair completely non-invasively from nests, allowing the investigation of inter-and intra-individual dietary variation. Given its incremental growth pattern, hair records temporal shifts in diet over long periods and allows one to reconstruct seasonal dietary patterns in species that cannot be directly observed. However, as for other sample materials, there are potential drawbacks related to the properties of hair keratin. Here I review some important facts on the nature of primate hair and also introduce new isotopic data from infant bonobo hair to provide methodological recommendations for future sample collection in the field and sample preparation in the laboratory. While these methodological guidelines focus on great apes which can be sampled strictly non-invasively, I also consider applications to other free-ranging primates. The biochemical composition, growth cycle, isotope turnover rate and isotopic fractionation in hair keratin are particularly relevant for data analysis and interpretation. Also, one can microscopically identify infant hairs and analyze them separately to study nursing and weaning behavior in primates. The goal of this article is to encourage primatologists to analyze the stable isotope ratios of hair to assess primate feeding ecology. Am. J. Primatol.
Previous observations have noted that in some species of higher termites the soldier caste lacks ... more Previous observations have noted that in some species of higher termites the soldier caste lacks pigmented particles in its gut and, instead, is fed worker saliva that imparts a whitish coloration to the abdomen. In order to investigate the occurrence of this trait more thoroughly, we surveyed a broad diversity of termite specimens and taxonomic descriptions from the Old World subfamilies Apicotermitinae, Cubitermitinae, Foraminitermitinae, Macrotermitinae, and Termitinae. We identified 38 genera that have this ''white-gutted'' soldier (WGS) trait. No termite soldiers from the New World were found to possess a WGS caste. Externally, the WGS is characterized by a uniformly pale abdomen, hyaline gut, and proportionally smaller body-to-head volume ratio compared with their ''dark-gutted'' soldier (DGS) counterparts found in most termitid genera. The WGS is a fully formed soldier that, unlike soldiers in other higher termite taxa, has a small, narrow, and decompartmentalized digestive tube that lacks particulate food contents. The presumed saliva-nourished WGS have various forms of simplified gut morphologies that have evolved at least six times within the higher termites.
The canopy effect describes vertical variation in the isotope ratios of carbon (δ 13 C), oxygen (... more The canopy effect describes vertical variation in the isotope ratios of carbon (δ 13 C), oxygen (δ 18 O) and partially nitrogen (δ 15 N) within plants throughout a closed canopy forest, and may facilitate the study of canopy feeding niches in arboreal primates. However, the nuanced relationship between leaf height, sunlight exposure and the resulting variation in isotope ratios and leaf mass per area (LMA) has not been documented for an African rainforest. Here, we present δ 13 C, δ 18 O and δ 15 N values of leaves (n = 321) systematically collected from 58 primate food plants throughout the canopy (0.3 to 42 m) in Côte d'Ivoire, West Africa. Besides leaf sample height and light availability, we measured leaf nitrogen and carbon content (%N, %C), as well as LMA (n = 214) to address the plants' vertical resource allocations. We found significant variation in δ 13 C, δ 18 O and δ 15 N, as well as LMA in response to height in combination with light availability and tree species, with low canopy leaves depleted in 13 C, 18 O and 15 N and slightly higher in %N compared to higher canopy strata. While this vertical isotopic variation was not well reflected in the δ 13 C and δ 15 N of arboreal primates from this forest, it did correspond well to primate δ 18 O values.
Stable isotope analysis of carbon and nitrogen in hair provides a versatile tool for reconstructi... more Stable isotope analysis of carbon and nitrogen in hair provides a versatile tool for reconstructing feeding behavior in elusive primate species. Particularly in great apes, researchers can sample long hair completely non-invasively from nests, allowing the investigation of inter-and intra-individual dietary variation. Given its incremental growth pattern, hair records temporal shifts in diet over long periods and allows one to reconstruct seasonal dietary patterns in species that cannot be directly observed. However, as for other sample materials, there are potential drawbacks related to the properties of hair keratin. Here I review some important facts on the nature of primate hair and also introduce new isotopic data from infant bonobo hair to provide methodological recommendations for future sample collection in the field and sample preparation in the laboratory. While these methodological guidelines focus on great apes which can be sampled strictly non-invasively, I also consider applications to other free-ranging primates. The biochemical composition, growth cycle, isotope turnover rate and isotopic fractionation in hair keratin are particularly relevant for data analysis and interpretation. Also, one can microscopically identify infant hairs and analyze them separately to study nursing and weaning behavior in primates. The goal of this article is to encourage primatologists to analyze the stable isotope ratios of hair to assess primate feeding ecology. Am. J. Primatol.
A mounting body of evidence suggests that changes in energetic conditions like prolonged starvati... more A mounting body of evidence suggests that changes in energetic conditions like prolonged starvation can be monitored using stable isotope ratios of tissues such as bone, muscle, hair, and blood. However, it is unclear if urinary stable isotope ratios reflect a variation in energetic condition, especially if these changes in energetic condition are accompanied by shifts in dietary composition. In a feeding experiment conducted on captive bonobos (Pan paniscus), we monitored urinary δ13C, δ15N, total C (carbon), total N (nitrogen), and C/N ratios and compared these results with glucocorticoid levels under gradually changing energy availability and dietary composition. Measurements of daily collected urine samples over a period of 31 days showed that while shifts in urinary isotope signatures of δ13C and δ15N as well as total C were best explained by changes in energy consumption, urinary total N excretion as well as the C/N ratios matched the variation in dietary composition. Furtherm...
Proceedings of the National Academy of Sciences, 2011
In primates, age, sex, and social status can strongly influence access to food resources. InPan, ... more In primates, age, sex, and social status can strongly influence access to food resources. InPan, these criteria are assumed to influence access to vertebrate meat. However, the significance of meat in terms of its role in the nutrition ofPanis still debated. Here we present a study using stable carbon and nitrogen isotope ratios in hair samples from habituated, wild bonobos (Pan paniscus) to explore these issues. Over a period of 5 mo hair samples were collected from fresh bonobo nests at LuiKotale, Democratic Republic of Congo. Hair samples were assigned to known individuals and were of sufficient length to allow the evaluation of isotopic variation over several months. Samples of plant foods and sympatric fauna were also analyzed. The δ13C and δ15N results of the bonobo hair were remarkably homogeneous over time and for the group as a whole. There are no differences in diet between the sexes. Within the group of males, however, there was a positive correlation between dominance st...
The first appearance of the Neolithic Linearbandkeramik (LBK) in Central Germany occurred during ... more The first appearance of the Neolithic Linearbandkeramik (LBK) in Central Germany occurred during the 6th millennium BC. However, though LBK sites are abundant in the German loess areas, there are only a few studies that reconstruct the diet of these first farmers using biochemical methods. Here we present the largest study undertaken to date on LBK material using stable isotope
The first appearance of the Neolithic Linearbandkeramik (LBK) in Central Germany occurred during ... more The first appearance of the Neolithic Linearbandkeramik (LBK) in Central Germany occurred during the 6 th millennium BC. However, though LBK sites are abundant in the German loess areas, there are only a few studies that reconstruct the diet of these first farmers using biochemical methods. Here we present the largest study undertaken to date on LBK material using stable isotope analysis of carbon and nitrogen to reconstruct human diet and animal husbandry strategies. We analysed the bone collagen of 97 human individuals and 45 associated animals from the sites of Derenburg, Halberstadt and Karsdorf in the Middle Elbe-Saale region of Central Germany. Mean adult human values are-19.9 ±0.4 ‰ for δ 13 C and 8.7 ±0.8‰ for δ 15 N. The δ 13 C values are typical for terrestrial, temperate European regions, whereas the δ 15 N values fall within an expected range for farming societies with a mixed diet consisting of products from domestic animals and plants. The consumption of unfermented dairy products is unlikely as there is direct palaeogenetic evidence of lactose intolerance available for one of the sites. There are no clear indications for dietary differences in sex. Young children under three years of age are enriched in δ 15 N due to breastfeeding indicating that weaning likely occurred around the age of three years. The fauna exhibit mean δ 13 C values of-20.9 ±0.8‰ and mean δ 15 N values of 7.0 ±0.9‰ respectively. Variation in the δ 13 C and δ 15 N in the domestic animals is probably caused by different livestock managements.
The feeding ecology of sympatric great ape species yields valuable information for palaeodietary ... more The feeding ecology of sympatric great ape species yields valuable information for palaeodietary reconstructions in sympatric early hominin species. However, no isotopic references on sympatrically living apes and their feeding ecology are currently available. Here we present the first isotopic study on sympatric great apes, namely western lowland gorillas (Gorilla gorilla gorilla) and central chimpanzees (Pan troglodytes troglodytes) from Loango National Park, Gabon. We successfully analyzed the stable carbon and nitrogen isotope ratios in a selection of food plants (n ¼ 31) and hair samples (n ¼ 30) retrieved from sleeping nests to test whether niche partitioning among sympatric chimpanzees and gorillas is detectable using isotope analysis of hair. Ape hair strands with roots were sectioned into sequential segments (n ¼ 100) to investigate temporal isotopic variation related to seasonal variations in food resources. We found significant d 13 C differences between herbaceous plants and fruits, most likely due to canopy effects. While the d 13 C values of chimpanzees indicate the consumption of fruit, the low d 13 C values in gorilla hair indicate folivory, most likely the consumption of 13 C-depleted herbaceous vegetation. Our isotopic data also confirmed dietary overlap between chimpanzees and gorillas, which varied by season. Gorillas showed significant variation in d 13 C values in response to season due to shifting proportions of herbaceous plants versus fruits. In chimpanzees, significant seasonal variation in d 15 N was likely related to the seasonal availability of fruit species with particularly high d 15 N values. In summary, we found isotopic evidence for niche partitioning and seasonal dietary variation among sympatric great apes at Loango. These findings provide a valuable reference for palaeodietary research on fossil hominins using d 13 C analyses, particularly for studies focusing on sympatric taxa and on temporal isotopic variation within incremental tissues such as tooth enamel.
Stable isotope analysis of carbon and nitrogen in hair provides a versatile tool for reconstructi... more Stable isotope analysis of carbon and nitrogen in hair provides a versatile tool for reconstructing feeding behavior in elusive primate species. Particularly in great apes, researchers can sample long hair completely non-invasively from nests, allowing the investigation of inter-and intra-individual dietary variation. Given its incremental growth pattern, hair records temporal shifts in diet over long periods and allows one to reconstruct seasonal dietary patterns in species that cannot be directly observed. However, as for other sample materials, there are potential drawbacks related to the properties of hair keratin. Here I review some important facts on the nature of primate hair and also introduce new isotopic data from infant bonobo hair to provide methodological recommendations for future sample collection in the field and sample preparation in the laboratory. While these methodological guidelines focus on great apes which can be sampled strictly non-invasively, I also consider applications to other free-ranging primates. The biochemical composition, growth cycle, isotope turnover rate and isotopic fractionation in hair keratin are particularly relevant for data analysis and interpretation. Also, one can microscopically identify infant hairs and analyze them separately to study nursing and weaning behavior in primates. The goal of this article is to encourage primatologists to analyze the stable isotope ratios of hair to assess primate feeding ecology. Am. J. Primatol.
Previous observations have noted that in some species of higher termites the soldier caste lacks ... more Previous observations have noted that in some species of higher termites the soldier caste lacks pigmented particles in its gut and, instead, is fed worker saliva that imparts a whitish coloration to the abdomen. In order to investigate the occurrence of this trait more thoroughly, we surveyed a broad diversity of termite specimens and taxonomic descriptions from the Old World subfamilies Apicotermitinae, Cubitermitinae, Foraminitermitinae, Macrotermitinae, and Termitinae. We identified 38 genera that have this ''white-gutted'' soldier (WGS) trait. No termite soldiers from the New World were found to possess a WGS caste. Externally, the WGS is characterized by a uniformly pale abdomen, hyaline gut, and proportionally smaller body-to-head volume ratio compared with their ''dark-gutted'' soldier (DGS) counterparts found in most termitid genera. The WGS is a fully formed soldier that, unlike soldiers in other higher termite taxa, has a small, narrow, and decompartmentalized digestive tube that lacks particulate food contents. The presumed saliva-nourished WGS have various forms of simplified gut morphologies that have evolved at least six times within the higher termites.
The canopy effect describes vertical variation in the isotope ratios of carbon (δ 13 C), oxygen (... more The canopy effect describes vertical variation in the isotope ratios of carbon (δ 13 C), oxygen (δ 18 O) and partially nitrogen (δ 15 N) within plants throughout a closed canopy forest, and may facilitate the study of canopy feeding niches in arboreal primates. However, the nuanced relationship between leaf height, sunlight exposure and the resulting variation in isotope ratios and leaf mass per area (LMA) has not been documented for an African rainforest. Here, we present δ 13 C, δ 18 O and δ 15 N values of leaves (n = 321) systematically collected from 58 primate food plants throughout the canopy (0.3 to 42 m) in Côte d'Ivoire, West Africa. Besides leaf sample height and light availability, we measured leaf nitrogen and carbon content (%N, %C), as well as LMA (n = 214) to address the plants' vertical resource allocations. We found significant variation in δ 13 C, δ 18 O and δ 15 N, as well as LMA in response to height in combination with light availability and tree species, with low canopy leaves depleted in 13 C, 18 O and 15 N and slightly higher in %N compared to higher canopy strata. While this vertical isotopic variation was not well reflected in the δ 13 C and δ 15 N of arboreal primates from this forest, it did correspond well to primate δ 18 O values.
Stable isotope analysis of carbon and nitrogen in hair provides a versatile tool for reconstructi... more Stable isotope analysis of carbon and nitrogen in hair provides a versatile tool for reconstructing feeding behavior in elusive primate species. Particularly in great apes, researchers can sample long hair completely non-invasively from nests, allowing the investigation of inter-and intra-individual dietary variation. Given its incremental growth pattern, hair records temporal shifts in diet over long periods and allows one to reconstruct seasonal dietary patterns in species that cannot be directly observed. However, as for other sample materials, there are potential drawbacks related to the properties of hair keratin. Here I review some important facts on the nature of primate hair and also introduce new isotopic data from infant bonobo hair to provide methodological recommendations for future sample collection in the field and sample preparation in the laboratory. While these methodological guidelines focus on great apes which can be sampled strictly non-invasively, I also consider applications to other free-ranging primates. The biochemical composition, growth cycle, isotope turnover rate and isotopic fractionation in hair keratin are particularly relevant for data analysis and interpretation. Also, one can microscopically identify infant hairs and analyze them separately to study nursing and weaning behavior in primates. The goal of this article is to encourage primatologists to analyze the stable isotope ratios of hair to assess primate feeding ecology. Am. J. Primatol.
A mounting body of evidence suggests that changes in energetic conditions like prolonged starvati... more A mounting body of evidence suggests that changes in energetic conditions like prolonged starvation can be monitored using stable isotope ratios of tissues such as bone, muscle, hair, and blood. However, it is unclear if urinary stable isotope ratios reflect a variation in energetic condition, especially if these changes in energetic condition are accompanied by shifts in dietary composition. In a feeding experiment conducted on captive bonobos (Pan paniscus), we monitored urinary δ13C, δ15N, total C (carbon), total N (nitrogen), and C/N ratios and compared these results with glucocorticoid levels under gradually changing energy availability and dietary composition. Measurements of daily collected urine samples over a period of 31 days showed that while shifts in urinary isotope signatures of δ13C and δ15N as well as total C were best explained by changes in energy consumption, urinary total N excretion as well as the C/N ratios matched the variation in dietary composition. Furtherm...
Proceedings of the National Academy of Sciences, 2011
In primates, age, sex, and social status can strongly influence access to food resources. InPan, ... more In primates, age, sex, and social status can strongly influence access to food resources. InPan, these criteria are assumed to influence access to vertebrate meat. However, the significance of meat in terms of its role in the nutrition ofPanis still debated. Here we present a study using stable carbon and nitrogen isotope ratios in hair samples from habituated, wild bonobos (Pan paniscus) to explore these issues. Over a period of 5 mo hair samples were collected from fresh bonobo nests at LuiKotale, Democratic Republic of Congo. Hair samples were assigned to known individuals and were of sufficient length to allow the evaluation of isotopic variation over several months. Samples of plant foods and sympatric fauna were also analyzed. The δ13C and δ15N results of the bonobo hair were remarkably homogeneous over time and for the group as a whole. There are no differences in diet between the sexes. Within the group of males, however, there was a positive correlation between dominance st...
The first appearance of the Neolithic Linearbandkeramik (LBK) in Central Germany occurred during ... more The first appearance of the Neolithic Linearbandkeramik (LBK) in Central Germany occurred during the 6th millennium BC. However, though LBK sites are abundant in the German loess areas, there are only a few studies that reconstruct the diet of these first farmers using biochemical methods. Here we present the largest study undertaken to date on LBK material using stable isotope
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Papers by Vicky M. Oelze