Subsistence practices, particularly animal husbandry and agriculture, are fundamental to human cultural activities, transforming species and ecosystems and driving population growth. Understanding livestock diets in prehistory provides crucial insights into past human behavior, specifically animal feed management strategies in response to local environments. The sixth millennium BCE in Central Europe saw a dynamic phase in domesticated cattle evolution, coinciding with the rapid expansion of the Linearbandkeramik (LBK) culture. This expansion spanned diverse forested environments with varying densities. Initially, slash-and-burn agriculture was proposed, but evidence of intensive crop cultivation within forests suggests alternative strategies. European woodlands historically served dual roles as shelter and animal feed resources (pasture and supplementary feed, ‘leafy hay’). However, the specific role of forests in LBK animal husbandry and regional adaptations remain unknown. LBK farmers initially settled seasonally flooded marshes and forested steppe before expanding northward, eastward, and westward. Despite occupying varied landscapes, these communities are often portrayed as largely homogenous in terms of their subsistence economies. Cattle were crucial, serving dual purposes of milk and meat production. Herd composition likely fluctuated with spring calving and periodic slaughtering. Consistent feed access is essential for herd health, fertility, and milk production; pasture and fodder management, including supplementary feed like leafy hay, are crucial. Early farmers successfully adapted cultivation practices to new landscapes, making it likely that they also experimented with domesticated animal diets, with forest environments potentially shaping regional husbandry strategies. Direct evidence of forest grazing and leafy hay use comes from waterlogged ruminant dung in late Neolithic Swiss villages. Stable carbon isotope values in animal tissues provide indirect evidence. Plants under dense forest canopies exhibit lower δ¹³C values compared to grasslands (canopy effect). Consumers reflect these values in their tissues. Previous studies hinted at forest resource reliance in LBK cattle diets based on lower bone collagen δ¹³C values compared to sheep. Dual analysis of carbon and oxygen isotopes in incrementally sampled enamel of ruminant molars is valuable for understanding animal diets seasonally. Compound-specific isotope analysis (CSIA) of nitrogen in amino acids (AAs) of dentine distinguishes diets derived from woody or herbaceous plants. CSIA-carbon of dairy lipids in pottery also provides ruminant dietary information. This study uses a large dataset from multiple sources to explore forest resource utilization.

Existing literature highlights the importance of understanding prehistoric livestock diets to reconstruct human behavior and adaptation to diverse environments. Studies on the Linearbandkeramik (LBK) culture have suggested various subsistence strategies, including slash-and-burn agriculture, but evidence points toward more complex interactions with forested landscapes. Previous research using stable isotopes has indicated the potential role of forests in cattle diets, showing lower δ¹³C values in bone collagen compared to sheep. However, the extent of forest use and the regional variability in husbandry practices remained unclear. This study builds on previous research by employing a multi-proxy approach, combining stable carbon and oxygen isotope analysis of multiple tissues (bone collagen, enamel bioapatite, and dairy lipids from pottery) with compound-specific isotope analysis of amino acids in dentine to investigate the nuances of LBK cattle husbandry in diverse forested environments.

This study compiled a large dataset of stable isotope values (δ¹³C and δ¹⁸O) from cattle bone, enamel bioapatite, and dairy lipids from pottery, along with CSIA-δ¹⁵N values from dentine amino acids, from 61 early Central European farming sites associated with the LBK culture. The δ¹³C values were used to infer dietary values after applying established enrichment factors for collagen, lipids, and bioapatite. The dataset was correlated with various environmental parameters, including mean forest cover (MFC), summer and winter precipitation, and temperature data from palaeoclimate reconstructions. Compound-specific isotope analysis (CSIA) of amino acids (AAs) in dentine provided insights into the types of plants consumed (woody vs. herbaceous). The analysis of δ¹⁸O and δ¹³C values from sequentially sampled enamel provided seasonal information about plant consumption and the extent of forest use for supplementary fodder. Statistical analyses such as analysis of variance, Pearson's correlation, and Spearman rank correlation tests were conducted to evaluate the relationships between dietary values and environmental parameters. GIS analyses were used to create interpolation maps visualizing the spatial distribution of cattle diet values and forest cover. The study uses detailed lipid analysis of pottery for dairy lipids, focusing on the C16:0 fatty acid.

The study reveals a wide range of δ¹³C values, indicating diversity in cattle diets. There was no significant difference between δ¹³C values derived from bone collagen, dairy lipids, and bioapatite. A negative correlation was observed between mean forest cover (MFC) and bone collagen δ¹³C values, suggesting greater reliance on forest resources in regions with high forest cover. However, a positive correlation was found between MFC and dairy lipid δ¹³C values, suggesting cattle grazed in forested areas even when open pastures were available. Sites in the western regions showed δ¹³Cdiet values falling within or below the baseline values for forest-dwelling wild herbivores, suggesting significant reliance on forest resources. Analysis of δ¹³Cbioap values from individual teeth revealed seasonal variations in diets, with some individuals showing evidence of winter feeding on leafy hay from forests. This was particularly evident in the Bischoffsheim site, which showed low δ¹⁸O values and low β values (from dentine CSIA-AA δ¹⁵N) in winter, indicating consumption of woody plants. The correlation between δ¹³Cbioap values and MFC suggests either diverse dietary plant sources or high seasonal variability in δ¹³C values within local pastures due to environmental factors. Winter climate factors were more strongly correlated with δ¹³C values than summer factors, suggesting that the availability of winter fodder influenced cattle diets. The findings from Bischoffsheim indicated winter foddering with leafy hay. The practice of supplementing winter diets with nutrient-rich feed has implications for female cattle health and fertility, possibly leading to higher out-of-season births. The availability of leafy hay may also have contributed to higher milk production.

The findings challenge the assumption of homogenous subsistence strategies within the LBK culture, revealing regional and seasonal diversity in cattle husbandry. The significant use of forests for grazing and supplementary foddering, particularly in the northwest regions, highlights the adaptive capacity of early farmers in responding to environmental challenges. The positive correlation between dairy lipid δ¹³C values and MFC, despite the availability of open pastures in some regions, suggests a deliberate choice to utilize forest resources for cattle rearing. This challenges interpretations based solely on bone collagen δ¹³C values. The evidence for winter foddering with leafy hay suggests that forests played a crucial role in ensuring the health, reproduction, and milk productivity of cattle herds. The integration of multiple isotopic proxies, including dairy lipids, provides a more nuanced understanding of animal diets and land-use strategies. The results suggest that forests served as both pasture and fodder source, influencing animal management practices and community economy. The interplay between climate and dietary strategies, particularly winter conditions, warrants further investigation.

This study demonstrates the diverse and adaptive cattle husbandry strategies employed by LBK farmers in Central Europe. The significant use of forested environments for both pasture and supplementary fodder, particularly during winter, challenges previous assumptions of a homogenous LBK economy. This underscores the importance of considering multiple isotopic proxies and environmental factors when interpreting prehistoric subsistence practices. The findings highlight the centrality of cattle to LBK communities and the strategic use of forest resources to optimize animal productivity and community resilience. Future research should focus on refining the temporal and spatial resolution of paleoclimate proxies and exploring the long-term impact of forest use on both animal populations and the transformation of forest ecosystems.

The study acknowledges limitations in the precision of some paleoclimate reconstructions, which could influence the correlation analyses. The species-specific identification of dairy lipids (cattle vs. sheep/goat) from pottery remains challenging, although previous large-scale analyses and slaughter profiles indicate that the lipids mostly originate from cattle. The availability of suitable samples for certain analyses (e.g., cattle teeth for bioapatite analysis) was limited by preservation and excavation factors. Finally, localized exploitation of forest resources might be underrepresented in larger-scale environmental reconstructions.