Abstract
Feasting played an important role in cementing social bonds between prehistoric communities. At Early Neolithic Asiab, western Iran, ceremonial feasting is evidenced by the butchered skulls of nineteen wild boars (Sus scrofa), an animal that was not commonly hunted in the region at the time. Here we use microscopic dental growth patterns to guide geochemical analyses of five wild boar teeth from Asiab and examine the geographical scope from which the animals derived. Our dataset includes 165 stable oxygen isotope values, 107 strontium isotope ratios, and Barium concentration maps. The findings indicate that despite Asiab’s ___location in an environment favourable to wild boars, the animals used for ceremonial feasting originated from a wide geographical catchment, with at least some necessitating transport over substantial distance across mountainous terrain. This deepens our understanding of the effort invested by the participating pre-agricultural communities for celebrating social connectivity across the wider landscape.
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Introduction
Hunting, one of the oldest forms of human–animal interactions, has for a long time held important value to human societies not only as a resource acquisition strategy, but also as a vehicle for provisioning symbolic feasts and rituals1. Prior to the domestication of livestock in southwestern Asia around 10,000 years ago, wild animals were used in activities aimed at creating or maintaining social cohesion, which eventually laid the foundation for reciprocal social networks that were critical for the survival of agricultural communities2,3,4,5,6,7. At Early Neolithic (~9660–9340 cal Before Common Era, BCE) Asiab, Iran, the remains of 19 butchered wild boars (Sus scrofa, representing both males and females of varying ages) were packed neatly together and sealed in a pit inside a circular semi-subterranean building (~20 m in diameter). This building was likely used as a communal structure similar to public buildings commonly found at Early Neolithic sites in the region8. Although wild boars appear in animal iconography at both nearby and more distant Early Neolithic sites (e.g., Sarab, Iran9; Göbekli Tepe, Türkiye10), they were not the most commonly hunted animal in the region at this time11 and the exact wild boar hunting strategies that were developed in the lead up to local pig domestication remains unknown.
The faunal material in the ‘boar pit’ at Asiab (which also included a skull and a mandible of a brown bear and fragments of red deer antler) represents evidence of feasting, most likely a single event rather than multiple events spread over a longer period of time11. Based on an estimate of the dressed weight of the 19 wild boars (~700 kg) and the fact that participants in the feast likely consumed more than an average daily meat intake (around 0.4–1.0 kg/person), the wild boars from the feasting pit may have provided food for 350–1200 adults11. It is possible that some of the meat was preserved for later consumption. This would have required relevant expertise as airdrying meat necessitates a suitable ___location or facility to balance airflow, temperature, and humidity so that the meat does not dry too fast (causing the surface to harden, trap moisture, and spoil) or too slow (causing the meat to get infected by maggots). Because the faunal material appears to have been deposited rapidly in one concentrated event, it is most likely that the killing, butchering, and consumption of the animals happened as part of the same event11.
Given the quantity of meat that the wild boar remains represent, the event either 1) involved a group of people that was larger than the current estimates of the number of people that inhabited Epipaleolithic or Early Neolithic settlements; or 2) a large amount of meat was used for purposes other than consumption, possibly sacrifice11. Both scenarios would have necessitated a large hunting operation to realise. Although Asiab is located on the bank of a river that has long provided a natural habitat for wild boars12,13,14, the nature and geographical scale of the hunting activities that enabled feasting and/or sacrificial activities at Asiab are unknown. The faunal assemblages from Asiab and other contemporary sites in the Zagros Mountains (including Ganj Dareh and Sheikh-e Abad) indicate that Early Neolithic inhabitants in the region employed diverse hunting strategies by focusing on species with distinct habitat requirements, such as wild goat (Capra aegagrus), aurochs (Bos primigenius), and chukar partridge (Alectoris chukar)12,13,14. Here we focus on how these diverse hunting strategies were used to realise large-scale ceremonial feasting.
In this study, we examine the geographical scope of wild boar hunting that underpinned feasting at Early Neolithic Asiab (34°18’1.6”N; 47° 8’6.90”E). Our samples include two second molars (M2) and three third molars (M3) recovered from the Asiab ‘boar pit’. The sample size (n = 5) was limited by the number of teeth that were sufficiently preserved. For each tooth, we map microscopic enamel growth patterns visible on longitudinal sections of the sample and use this information to target high-resolution isotopic and trace element analyses. For a summary of the workflow used in this study, see Fig. 1. After sectioning (stage 1), we used established histological methods15 to reconstruct the timing of development of the wild boars’ teeth (stage 2). In stage 3, we applied the estimated duration of enamel extension (in days) and rate of enamel apposition (in µm/day) to target in-situ analysis of phosphate δ18O values in enamel secreted at ~weekly intervals and later attenuated through enamel maturation. We combined these results with analyses of in-situ 87Sr/86Sr ratios, an assessment of the regional distribution of bedrock 87Sr/86Sr ratios, and trace element (barium, Ba) maps of the teeth ground sections (stage 4), to assess the likelihood that all boars were hunted in close proximity to the site.
For details on each stage, see ‘Methods’.
Our multi-proxy suite of analyses provides a unique opportunity to scrutinise the social role of hunting and assess the degree to which the hunting strategies may have been shaped by worldviews around landscape connections that were at the heart of the ceremonial activities at Asiab. Long-distance animal transport has been identified in later ritual contexts (e.g., at Neolithic henge monuments in Britain16), but this is the first time, to the best of our knowledge, that it has been found in pre-agricultural contexts. The approach of using tooth-specific developmental landmarks to guide high-resolution geochemical analyses provides an improvement over the more commonly used method of hand-drilling tooth enamel powder in ~1 mm intervals17,18,19, which provides averaged and attenuated values of dietary inputs. A central aim of this research is to make the methodology reproducible, and we provide extensive documentation in the ‘Methods’ section, in Supplementary files, and in materials deposited to an online repository to enable others to reproduce every step of the process.
Results
Dental histology: identifying incremental markings in tooth enamel to target high-resolution geochemical analysis
The early developmental histories of the study animals were determined using histological analysis of enamel growth features visible in 50 µm-thick ground sections of the selected teeth (see Supplementary Figs. 1–3 for images showing dental growth features from the analysed samples). The analysis showed that the wild boars’ second molars track enamel secretion over a period close to 6 months (ASB174: 183 days; ASB360: 197 days). The third molars track enamel secretion over a period of 11–12 months (ASB133: 330 days; ASB402: 342 days; ASB449: 310 days + ~30–40 days)—see Table 1 for a summary and Supplementary Data 1 for full details.
The wild boar tooth enamel extension rates (EER) for the analysed enamel segments (see Supplementary Figs. 4–8) were used to guide measurement of δ18O values of enamel in locations that correspond to ~weekly intervals of enamel secretion (see ‘Methods’). Although secreted enamel undergoes a degree of over-printing during the later stage of enamel maturation20, this sampling method provides the highest level of temporal control for estimating changes in δ18O intake from tooth enamel. Using experimental data from modern sheep, Green et al.20 estimate that innermost enamel retains 36–48 % of the likely drinking water δ18O range after it has been attenuated during enamel maturation. For this reason, enamel values cannot be directly compared to δ18O values of environmental water. However, any inter-individual differences in enamel δ18O values can still be used to assess the possibility that individuals spent their crown formation time in distinct locations.
Periods of severe stress during dental development would be identifiable by a disruption of enamel microstructure or the occurrence of hypoplastic enamel defects No such defects were recorded in the studied teeth. This indicates that the wild boars were not exposed to any severe adversity during the period of crown formation of their second or third molars.
Isotopic analysis: assessing the residential origins of Asiab wild boars
Two lines of geochemical evidence were used to assess the residential locations of the study animals during the time of their second and third molar crown formation: (1) in situ analysis of enamel phosphate δ18O values measured using a Sensitive High-Resolution Ion Microprobe (SHRIMP); and (2) in situ enamel 87Sr/86Sr ratios measured using Laser-Ablation Multi Collector Inductively Coupled Plasma Mass Spectrometry (LA-MC-ICP-MS). Both δ18O values and 87Sr/86Sr ratios provide some insight into the ___location where the animals resided during enamel growth. Enamel δ18O values reflect the stable oxygen isotope composition of ingested water, which varies depending on the composition of ___location-specific environmental water, mostly rainfall21. Enamel 87Sr/86Sr ratios reflect the composition of biologically available (‘bioavailable’) strontium obtained through diet. Local 87Sr/86Sr ratios are largely driven by the age of underlying geology, but the actual composition of bioavailable strontium may be influenced by more surficial sources (e.g., surface water, aeolian dust, sea spray, sediments transported during erosion) that may have 87Sr/86Sr ratios distinct to those of local bedrock22. In archaeological studies, ‘non-locality’ is commonly interpreted as falling outside the mean ± 2 standard deviations (2σ) of samples that represent locally bioavailable strontium uptake23,24,25; although the choice of sample type for defining the local 87Sr/86Sr baseline has proven very challenging22,26.
Because each geochemical proxy has its limitations for assessing provenance, the recommended approach is to use a combination of proxies22,27,28. Here, both δ18O values and 87Sr/86Sr ratios point to a difference in residential locations between ASB449 and the other four individuals (ASB133, ASB174, ASB360, and ASB402; together referred to as ASB133–402) (see Fig. 2; δ18O sequences are shown individually in Supplementary Fig. 9). The enamel δ18O values of ASB449 (range: 12.2–17.2‰, mean: 15.2 ± 1.2‰ (1σ), n = 44) are notably lower compared to those of ASB133–402 (range of all four samples: 15.9–22.0‰; mean ASB133: 17.8 ± 0.8‰, n = 36; mean ASB174: 18.0 ± 0.9‰, n = 24; mean ASB360: 19.0 ± 2.0‰, n = 23; mean ASB402: 17.9 ± 1.5‰, n = 38). Similarly, the enamel 87Sr/86Sr ratios of ASB449 (0.7068–0.7071, mean: 0.7071 ± 0.00008 (1σ), n = 22) are lower compared to those of ASB133–402 (range of all four samples: 0.7078–0.7085; mean ASB133: 0.7079 ± 0.00005, n = 23; mean ASB174: 0.7083 ± 0.00007, n = 21; mean ASB360: 0.7081 ± 0.00011, n = 20; mean ASB402: 0.7082 ± 0.00008, n = 21). The mean ± 2σ of 87Sr/86Sr ratios of ASB133–402 is 0.7081 ± 0.0003, and ASB449 (0.7071 ± 0.00016) falls outside of this interval. ASB449 is not distinctive from the other four wild boars in terms of its age at death (see Table 1).
a Stable oxygen isotope values (δ18O, SMOW) of tooth enamel phosphate measured in situ using round (⌀ 21 µm) analytical spots placed at locations corresponding to ~weekly intervals in enamel secretion. Colours denote distinctions between second molars (M2; light grey) and third molars (M3; red or dark grey). Notably, second and third molars start and end formation at different times (Supplementary Fig. 10). The plotting symbols are approximately as wide as the instrument measurement error shown with grey error bars. b Strontium isotope ratios (87Sr/86Sr) of tooth enamel measured using rectangular (450 × 110 µm) laser tracks placed approximately in the direction of crown formation; with error bars indicating ± 1 standard error. For description of the placement of analytical spots, see ‘Methods’.
The four geochemically similar wild boars (ASB133–402) have 87Sr/86Sr ratios close to those of modern plants (0.7085 ± 0.0004, n = 11) measured in the vicinity of the archaeological site of Ali Kosh, located just over 100 km directly southwest of Asiab29. Tissue 87Sr/86Sr ratios can be directly compared to 87Sr/86Sr ratios of bioavailable strontium because strontium does not undergo fractionation during tissue assimilation30,31,32,33,34,35. It was not possible as part of this study to carry out further modern plant measurements closer to Asiab, so instead we interpolated the ratios of local bioavailable strontium to provide a coarse estimate of regional 87Sr/86Sr variability. Looking only at the regional geology, it would seem that the 87Sr/86Sr ratios around Ali Kosh (which sits on older Miocene bedrock) should be higher compared to 87Sr/86Sr ratios around Asiab (which sits on younger Quaternary and Mesozoic bedrock)36. This prediction is reflected in the contour map constructed using 87Sr/86Sr ratios of rock samples from the Georoc database and the plants around Ali Kosh29,37 (see Fig. 3). This bedrock-based prediction, with an error at Asiab around 0.00002, would indicate that ASB449 is more local to Asiab than ASB133–402. The map in Fig. 3 provides an estimate of ~70 km as the minimum distance from Asiab to these non-local 87Sr/86Sr ratios between 0.7078 and 0.7085. However, predictions solely based on bedrock measurements are limited, as they do not reflect possible impacts of biologically available strontium from surficial deposits. In addition, there are limited direct 87Sr/86Sr measurements available for the region around Asiab29,37. We therefore use the map simply to illustrate the approximate ranges of 87Sr/86Sr ratios in the wider region, noting that bioavailable 87Sr/86Sr may have further variability on the local level.
a Regional 87Sr/86Sr ratios were estimated using data from the Georoc database37 and measurements of modern plants from Ali Kosh29 and interpolated to the wider region using the underlying lithology (following Barakat et al.87). b Prediction error on the interpolations. In both panels, the ___location of Asiab is shown with a star and the locations of the 87Sr/86Sr observations underpinning the interpolation are shown in points. Rock measurements from the Georoc database are shown as circles; plant measurements from around Ali Kosh are shown as squares.
Even with these caveats, we can eliminate the possibility that all five wild boars originated from an area close to the site for two reasons. First, the lower 87Sr/86Sr ratios recorded in ASB449 are consistent throughout the entire intra-tooth sequence representing 300+ days of crown formation. This is unlikely to be the result of episodic foraging on an anomalous surficial source (i.e., local 87Sr/86Sr variability not represented in Fig. 3). Second, the difference in δ18O values between ASB449 and ASB133–402 is sufficiently large ((mean of ASB133–402, n = 167) – (mean of ASB449, n = 88) = 2.8‰, 95% CI [2.5, 3.1]) to indicate that the animals spent crown formation time in distinct regions. Altitudes within 100 km of Asiab range from ~1000 masl (Seimarreh Valley) to ~3400 masl (Paraw Mount), so some of the variation in δ18O values can be explained by the ‘altitude effect’, which has a global average value around –0.26‰ per 100 m (at altitudes under 5000 masl)38. A mean difference between 2.5‰ and 3.1‰ may represent an altitudinal difference of several hundred metres.
Based on the results obtained here, ASB449 likely originated from closest to Asiab, while the other four wild boars originated from locations further away. It is important to note that the small sample size (limited by the number of teeth that were sufficiently preserved) is too small to make inferences about the overall proportion of wild boars of geographically distinct origin consumed in the ritual feast at Asiab. Our interpretations here are thus constrained to identifying, rather than quantifying, the phenomenon of distance transport of wild boars.
Isotopic and trace element analysis: assessing foraging patterns of the study animals to evaluate their geographical dispersion
In addition to geographical residence, incremental tooth enamel δ18O values provide insight into the timing of the wild boars’ births, which can help better interpret seasonal dietary patterns that may be indicative of geographical dispersion. Modern wild boars from Northern Hemisphere temperate climates are typically born in the spring season (between March and May, with a peak in April)39. However, a second farrowing season may take place in late summer (between August and September)39,40. Crown formation in second molars starts around day 65–70 after birth, which is in June for boars born in the spring and October/November for boars born in the late summer. Crown formation in third molars starts around day 210 after birth, which is in October of the birth year for boars born in the spring and February/March of the following year for boars born in the late summer15,41 (see Supplementary Fig. 10 for a schematic of the chronology of crown formation of mandibular molars of wild boars).
Tooth enamel δ18O values reflect the composition of body water and by extension the composition of ingested water, which varies seasonally due to the effects of temperature, surface water evaporation, and leaf water evapotranspiration42,43,44. Maximum δ18O values reflect enamel that formed during hot/dry summers and minimum δ18O values reflect enamel that formed during cold/wet winters17,45. Given the timing of molar crown formation described above, wild boars born in the spring should record maximum δ18O values around the start of their M2 crown formation (June) and near-minimum δ18O values around the start of their M3 crown formation (October). Wild boars born in the late summer season should record near-minimum δ18O values around the start of their M2 crown formation (October/November) and increasing δ18O values around the start of their M3 crown formation (February/March). The incremental δ18O values measured in this study (shown together in Fig. 2 and individually in Supplementary Fig. 9) indicate that: (1) both second molars (ASB174, ASB360) represent animals born in the spring (with maximum δ18O values at the start of the recorded sequence), (2) two third molars (ASB133, ASB402) represent animals born in the spring (with near-minimum δ18O values at the start of the recorded sequence), and (3) one third molar (ASB449) represents an animal born in late summer (with near-maximum δ18O values at the start of the recorded sequence).
To probe whether the wild boars exhibit dietary differences that would be indicative of dispersed foraging, we used the season of the animals’ birth to determine the timing of episodic spikes in the intake of dietary barium (Ba). We mapped the trace element composition on the surface of the polished sections using Laser-Ablation Inductively Coupled Plasma Mass Spectrometry (LA–ICP–MS, see ‘Methods’ for details). The resulting images (showing Ba concentrations normalised to Ca concentrations) were interpreted with reference to the ___location and shape of the daily growth lines identified through the histological analysis (Supplementary Figs. 4–8). We distinguished between diagenetic patterns resulting from post-depositional uptake of barium from soil (appearing as indiscriminate spikes close to the enamel surface or the enamel dentine junction) and biogenic bands coinciding with the geometry of the growth patterns.
Biogenic spikes in Ba uptake are visible in all teeth, with two closely spaced spikes appearing in ASB174, ASB360, ASB133, and ASB402, and one spike appearing in ASB449 (Fig. 4). In the case of second molars (ASB174, ASB360), the timing at which the spikes occurred was determined through comparison to a ground section of an experimentally fluorochrome-labelled modern wild boar second molar41. Based on the positions of fluorochrome labels resulting from injections administered on known dates, the two closely spaced Ba spikes in ASB174 formed on postnatal day ~165. A broader zone showing Ba enrichment in ASB360 formed between postnatal days ~86 and ~135. In the case of the third molars (ASB133, ASB402, ASB449), the timing of the Ba spikes was likewise inferred from labelling experiments of modern wild boar and the reconstructed rates of enamel and dentine formation of un-labelled modern wild boar third molar15. In ASB133, a broader zone showing Ba enrichment appears between postnatal days ~330 and ~380. In ASB402, biogenic Ba bands occurred on postnatal days ~310 and ~410 (visible in both enamel and dentine) and, after cessation of enamel formation, on days ~565 and ~605 (in the dentine). In ASB449, a single Ba enrichment band appears at postnatal day ~280.
a–e show trace element maps highlighting the concentration of barium normalised to calcium (Ba/Ca) for the second molars (a ASB174, b ASB360) and the third molars (c ASB133, d ASB402, e ASB449). Blue hues indicate lower Ba concentrations; red hues indicate higher Ba concentrations. Black outlines delineate the enamel regions (see shapes of the analysed enamel flanks in Supplementary Figs. 4–8). The regions inside of the enamel (towards the pulp cavity) show dentine. Arrows indicate visible markings annotated by their approximate timing (in postnatal days). Asterisks at the start of an annotation (*70 in ASB174, *220 in ASB133, *210 in ASB402, *210 in ASB449) indicate the approximate timing (in postnatal days) of the start of crown formation. Asterisks at the end on an annotation (255* in ASB174, 550* in ASB402) indicate the approximate timing of the end of crown elongation. Ba concentrations are higher in dentine because of its higher porosity and lower mineral content, making it susceptible to post-depositional uptake of trace elements from soil. Despite the post-depositional Ba uptake in dentine, visible patterns in (c) and (d) follow the geometry of dentine formation and indicate that they are biogenic. White regions show enamel that is missing as a result of bubbles incorporated during the epoxy casting process (see ‘Methods’).
Given the timing of the wild boars’ birth, the Ba uptake spikes occurred in September (in ASB174), June and August (in ASB360), February and April (in ASB133), February and May (in ASB402) and May (in ASB449) (Table 1). As third molars start formation after the cessation of milk feeding, the Ba enrichment in ASB133, ASB402, and ASB449 is unlikely to be the result of nursing. For the calculation of month of Ba bands, 1 April was used as a starting point for spring births39 and 1 September was used as a starting point for late summer births.
Episodic Ba intake increases may have been the result of (1) the wild boars consuming distinct food items (naturally high in Ba); (2) seasonal movement to locations with distinct underlying geology (contributing to higher Ba levels in the local soils and plants); or (3) more intense rooting behaviour (characteristically exhibited by older wild boars) causing increased uptake of soil particles. While the more intense rooting behaviour may explain the Ba enrichment between postnatal days ~565 and ~605 in ASB402, the early spikes were more likely the result of distinct foraging habits at different times of the year. The diversity in timing of the spikes (see Table 1 for a summary) indicates that the five studied wild boars were not part of the same sounder (i.e., a group of wild boars). Instead, it suggests that they may have derived from sounders that were more geographically dispersed.
Discussion
In this study, we used a multi-proxy suite of geochemical data to assess the geographical dispersion of five wild boars recovered from the ‘boar pit’ at Asiab. The findings can now be used to assess the hunting activities and possible animal transport that underpinned the organisation of large-scale feasting at Asiab during the Early Neolithic. The combination of enamel δ18O values and 87Sr/86Sr ratios indicates that the analysed boars (n = 5, limited by the number of sufficiently preserved samples) are unlikely to have originated from close proximity to Asiab. Using the distribution of bedrock 87Sr/86Sr ratios, it would seem that four out of the five wild boars (ASB133–402) were less local to Asiab than ASB449; although this needs to be confirmed through future mapping of bioavailable strontium in the region. The δ18O values of tooth enamel secreted at approximately weekly intervals, although later attenuated through enamel maturation, indicate that the geochemically-similar ASB133–402 wild boars were born in the spring (i.e., during the normal farrowing season). The remaining and geochemically dissimilar boar, ASB449, was born during a second farrowing season in the late summer. Dietary spikes in the wild boars’ dietary Ba intake were most likely caused by episodic consumption of distinct food items or by short-term seasonal movement to locations with naturally higher Ba abundance. The variable timing of these dietary episodes indicates that the wild boars followed different foraging patterns and were thus more likely dispersed across the local region; as opposed to originating from the same sounder that would be easier to hunt.
When using geochemical data to determine the geographical provenance of animals from the archaeological record, it is important to keep in mind that tooth enamel provides information about the animals’ dietary intake during the time the enamel was forming; not during the season in which they were hunted. Wild boars can travel several tens of kilometres on their own; however, home ranges above ~20 km2 are very rare46,47,48,49,50,51,52. It is therefore unlikely that the wild boars moved on their own over large distances and ended up close to Asiab after their tooth crowns completed formation. Instead, it indicates that the animals were sourced at distant locations and that humans were involved in their transportation to Asiab.
Walking at an average speed between 4–5 km per hour (estimated for a broad demographic of men and women between 20–60 years of age53,54), it would have taken the hunters approximately 14–17.5 h to travel 70 km over a flat terrain. Seventy kilometers is the distance from Asiab to the closest ___location with non-local 87Sr/86Sr ratios of 0.7078 and 0.7085 estimated from the map shown in Fig. 3. Although a map based on bedrock 87Sr/86Sr ratios does not fully represent local variability in bioavailable strontium, it provides one possible scenario for the approximate distance over which the animals may have been transported. Given that the environment around Asiab is not flat, a straight-line distance of 70 km would almost certainly equate to a longer actual travelled distance. Additionally, the weight of the animal carcass would have reduced the hunters’ walking speed. Therefore, at a conservative estimate, this trip could have lasted at least 2 days.
There are several plausible scenarios of how the non-local wild boars may have been transported to Asiab. First, Asiab inhabitants may have travelled to hunt the animals in far-off locations. Second, inhabitants of settlements further away may have travelled to Asiab bringing wild boars hunted in their own home range as a contribution to the communal feast. Third, people may have consumed the wild boars at distant locations and travelled to Asiab with the boar heads as hunting trophies to use for ritual deposition. Each scenario represents a considerable physical effort. This high effort associated with boar hunting and animal transport signals that the symbolic dimension of the feasting activities at Asiab extended beyond cementing social relationships between the participants and into the realm of where the hunting took place. In terms of the value of reciprocity for cementing social bonds, the second scenario seems most likely.
A large body of ethnographic, ethnohistoric, and archaeological literature documents that in both foraging and agricultural societies, hunting activities are not simply optimised towards satisfying nutritional and utilitarian needs of the communities1,55,56,57. Importantly, hunting activities are driven by relational thinking (i.e., the worldviews, beliefs, and mythologies created around human–animal relationships), which dictates how hunters and their communities should interact with wild animals, often in ways aimed at fostering mutual respect and reciprocity1,56,57. Some examples of diverse customs that foster these relationships (see Russell1 for a full discussion) are: placing ceremonial objects on animals to encourage them to return in future hunts58, removing the animals’ feet first during the initial butchering to prevent their spirit from wandering59, and returning the animal bones to forests or rivers as a sign of respect60.
The relational dimension of human–animal interactions can manifest itself in the creation of rituals that are intended to bolster the status quo (e.g., ‘maintenance rituals’ aimed to increase the chances of success during future hunts57,58) and rituals that are intended to challenge the status quo (e.g., use of hunting trophies to display the hunters’ skill and/or boost their social status1). Since Asiab represents a unique archaeological deposit, and since wild boars were not the most commonly hunted wildlife species in the region at this time, it is more likely that the hunting activities were formulated within the latter type of rituals aimed at commemorating or envisioning a special human–animal relationality.
Thus, what we have at Asiab is a special function that celebrated relationalities between people (who formed part of the social network) and relationalities between humans and animals. We cannot determine what the specific beliefs and/or mythologies were that shaped the latter relationalities and whether they were shared or unique to the communities that came together at Asiab. However, it is clear that without the geographical dimension underpinning the hunting activities, the ritual event organised at Asiab would have played out differently. With it, the social gathering brought expression to the worldviews and belief systems that configured human–animal interactions across the wider regional landscape, and the scope of this expression was important enough to warrant considerable effort to realise.
Feasting has long been considered to be a key feature of ceremonial activities in Late Epipalaeolithic and Early Neolithic societies in southwest Asia61,62,63,64,65,66,67,68. It may have been used to cement social bonds among groups at particularly challenging times (e.g., funerary activities) or at certain points during the annual cycle2. Some have argued that feasting was a major driving force behind the adoption of plant cultivation and animal domestication in southwest Asia69,70,71,72. While this view is contested67,73, our findings indicate that 10th millennium BCE groups in the central Zagros were prepared to expend considerable time and energy on attending and/or provisioning feasts and ceremonies early on during the transition to agriculture.
The use of particular animals—in this case wild boar—with specific qualities (e.g., ferocious and dangerous behaviour) to provision such feasts is striking. It is probably not a coincidence that wild boars, which feature regularly in the symbolism and art of Göbekli Tepe and other Pre-Pottery Neolithic A/Pre-Pottery Neolithic B sites in Upper Mesopotamia and southeast Anatolia10,74,75, later became some of the first managed animals. Our direct evidence that wild boars were transported over distances to form part of ceremonial activities at Asiab does not just underscore the high status that these animals held in the symbolism and art of the Early Neolithic societies. It documents the people’s willingness to invest vast amounts of time and energy into celebrating the human–animal dynamic, which eventually laid the foundation for the adoption of livestock management as the primary subsistence strategy a few millennia later.
Methods
Determining animal age-at-death
The ages of the studied individuals were determined using two different methods. The first method involved histological assessment of the crown and root developmental stages using data from Magnell and Carter76 and Emken et al.41. The second method involved assessment of macroscopic dental features of tooth eruption and wear77, an approach that is commonly applied in zooarchaeology. The latter determinations encompass wider age ranges because the criteria are not entirely clear-cut for samples with unfinished crown or root formation.
Sectioning, histology, and analysis of incremental markings
The teeth were initially scanned using a Micro Computed Tomography (µCT) scanner at the National Laboratory for X-Ray Micro Computed Tomography, Australian National University. The resulting 3D models were used to determine the optimal locations of the sectioning planes and for avoiding areas with internal fractures caused by diagenesis. The teeth were cast in an embedding medium (EpoTech 301), and a slab of ~200 µm thickness was removed from the specimens using a water-cooled diamond blade (⌀25 cm) at Griffith University, Australia. These thick sections were further ground and polished to a final thickness of ~50 µm at the University of Hildesheim, Germany, employing a series of silicon carbide paper (grits 400 to 4000), followed by a final polishing step on a motorised rotary polisher with a 3 µm diamond suspension and an 0.3 µm aluminium slurry. The ground sections were cover-slipped using removable Dibutylphthalate Polystyrene Xylene (DPX) mounting medium and viewed and analysed in a Zeiss Axio-Imager2-microscope equipped with an Axiocam 503 colour camera under plain and polarised transmitted light, and by using Differential Interference Contrast (DIC) and phase-contrast illumination.
To reconstruct crown development parameters (tabulated in Supplementary Data 1), stitched images of the crown flanks were produced using a 5x magnification objective. In these images, landmark lines (i.e., clearly traceable incremental markings) were marked down to the enamel dentine junction (EDJ). Between these landmark lines, the number of laminations (i.e., daily incremental markings15) was counted using higher magnifications if necessary. See Supplementary Figs. 1 and 2 for images of daily laminations in ASB402 (lingual and palatal side, respectively) and Supplementary Fig. 3 for images of long-period incremental markings (striae of Retzius) in ASB449 (palatal side). It should be noted that wild boar tooth development varies markedly from tooth development in humans and non-human primates, which is why previously applied sampling strategies for using histological information to target high-resolution isotopic and trace element analyses78,79,80 are not directly transferable to wild boars, or domestic livestock commonly found in archaeological faunal assemblages.
Using this approach, the number of days of crown formation between consecutive landmark lines, the enamel extension rate (EER) along the EDJ, as well as the crown flank extension time were established for all samples. The crown flank extension time covers the period that is needed to form the complete enamel dentin junction in the respective crown flank (i.e., the time needed for enamel extension to reach the later crown root border). As the number of traceable landmark lines varied among the teeth, the number of sections along the EDJ for which the EER could be established also varied (between six and nine; Supplementary Figs. 4–8).
The presence of accentuated incremental markings (lines) was microscopically assessed in the ground sections using the abovementioned methods. Incremental markings were classified as accentuated when they could be traced over ≥75% of the extension of the respective secretory front in the microscopic sections, a criterion originally proposed by Goodman and Rose81 for human enamel.
The two lower second molars (ASB174, ASB360) were analysed on the lingual side. The lower third molar (ASB133) was analysed on the buccal side. The upper third molars (ASB402, ASB449) were analysed on both palatal and buccal sides. The buccal side of ASB402 was used for both isotopic and elemental analyses. With ASB449, the buccal side was used for stable oxygen isotope analyses and the palatal side was used for strontium isotope analyses and trace element analyses. Where the cervical-most portion of the enamel was not preserved (ASB360 lingual, ASB402 buccal), the final length along the EDJ was reconstructed. Where the cuspal-most portion of the enamel was missing due to tooth wear (ASB402 palatal), the length along the EDJ was likewise reconstructed. Where readable incremental markings were missing in the ground sections, the time represented in the respective enamel portion was reconstructed based on the mean daily secretion rate recorded in the corresponding enamel portion with readable incremental markings in other specimens.
Strontium isotope ratio analysis
Strontium isotope ratios (87Sr/86Sr) of tooth enamel were measured using a Laser Ablation Multi-Collector Inductively Coupled Plasma Mass Spectrometer (LA–MC–ICP–MS) at the Geoarchaeology and Archaeometry Research Group facility at Southern Cross University, Australia. The instrument was a Thermo Neptune XT connected to an ESI NWR193 ArF excimer laser ablation system. Enamel from tooth ground sections was ablated using analytical spots (450 × 110 µm) placed in the direction of enamel extension, using dwell time of 35 s, 100 Hz frequency and 70% laser intensity (~5j per cm2). The first spot was placed as close as possible to the dentine horn in the cuspal portion of the sample and the last spot was placed as close as possible to the cervical crown border (the distribution of spot placements is shown in Fig. 1, panel 3). Between 20 and 23 87Sr/86Sr ratios were measured from the sample teeth, depending on the length of the enamel crown. Because this analysis did not take into consideration the daily growth markings and the changing rate of enamel extension (higher in the cuspal region and lower in the cervical region), the temporal dimension of each measurement is not directly comparable to the temporal dimension of the stable oxygen isotope measurements discussed below. However, the general direction of the spots’ placement enables an examination of sub-seasonal changes in the animals’ 87Sr/86Sr intake.
Sample aerosol was carried to the ICP–MS using a mixture of He (640 ml per min) and N2) 6 ml per min). The following isotopes were collected in static mode: 82Kr, 83Kr, 84Sr, 85Rb, 86Sr, 87Sr, 88Sr, and 89Y. Each isotope was collected using cycles of 2.054 s each. Data reduction (including a background subtraction, mass bias, Rb/Sr and Ca argide/dimmer corrections) was performed using Iolite82 with the CaAr–CaCa interference correction (using 82 or 83 as a monitor mass) and nominal mass-bias factors based on 88Sr/86Sr applied to these interferences83. Internal laboratory standards (a seal tooth and a clam shell with a modern seawater 87Sr/86Sr ratio of 0.709284) were analysed four times at the start of each run and four times at the end of each run and used for matrix-matched 87Sr/86Sr correction. All individual measurements are presented in Supplementary Data 3.
Although strontium is present as a trace element in soil and can be taken up by archaeological teeth during burial (to a larger extent by the dentine than enamel due to the lower mineral content of the former85,86), the fact that the majority of the analysed samples provided non-local ratios (i.e., distinct from the extrapolated 87Sr/86Sr ratio of bedrock at the ___location of the archaeological site) indicates that diagenesis did not systematically impact the studied enamel samples.
Map of the distribution of 87Sr/86Sr ratios for the study region
A map of the bioavailable 87Sr/86Sr ratios was constructed using data from the Georoc database37 and interpolated to the wider region using the underlying lithology (following Barakat et al.87). We used external drift kriging to estimate bioavailable 87Sr/86Sr in an area extending approximately 10° in each direction from Asiab (Fig. 3). Kriging is a geostatistical prediction method that linearly predicts a variable (87Sr/86Sr) in a continuous area, given discrete observations and a model for their spatial variation. External drift kriging further incorporates information from a spatially continuous auxiliary variable that is correlated with the target variable—in this case, rock types underlying the study area. Our kriging approach closely followed that of Barakat et al.87. 87Sr/86Sr observations (n = 6130) are from the Georoc database37 and measurements of plants collected in the region (close to the archaeological site of Ali Kosh29; n = 12). Where the datasets contained multiple 87Sr/86Sr values for a single ___location, we calculated the mean value (total unique sites = 645). 87Sr/86Sr predictions were informed by local underlying lithologies obtained from the Global Lithological Map (GLiM88), which is available at a 0.5° spatial resolution globally. We performed the kriging using the gstat89 package in R (R Core Team 2021), with a Bessel variogram model. We made predictions over a continuous 0.05° latitude by 0.05° longitude grid spanning 25.17°N–42°N, 37.16°E–57.1°E (informed by the ___location of the most distal 87Sr/86Sr observations within 10° of Asiab in each direction).
We used the bioavailable 87Sr/86Sr grid to estimate the minimum distance where the non-local wild boars (ASB133–402; combined mean 0.7081 ± 0.0003, 2σ) originated from. Specifically, we calculated the distance between Asiab and all grid cells with predicted 87Sr/86Sr between 0.7078 and 0.7085, assuming that the wild boars originated at or near these locations. We report the distance from Asiab of the closest grid cell with a predicted 87Sr/86Sr value in the relevant range.
Stable oxygen isotope value analysis
Incremental stable oxygen isotope values (δ18O) of tooth enamel phosphate were measured using a Sensitive High-Resolution Ion Microprobe for Stable Isotopes (SHRIMP–SI) at the Research School for Earth Sciences, Australian National University. All ground sections were initially cleaned with petroleum spirit, RBS35 detergent solution, hot water, and deionized water to remove any possible surface contamination. The samples were then placed to dry in a vacuum oven at 60 °C for ≥24 h and subsequently coated with 30 nm-thick layer of high-purity aluminium before being placed in the SHRIMP–SI under high vacuum for at least 12 h prior to analysis.
The SHRIMP–SI measurements were performed with a Cs primary beam of ~1.52 nA. The primary ion optics were configured with a 200 μm Kohler aperture to produce a ~21 μm diameter spot on the surface of the ground section. Electrons were delivered to the target surface with an energy of ~ –1.8 keV (i.e., resultant electron energy at sample surface). The secondary beam maximised its signal on Post–ESA monitor (PESAM) and the source slit width was set at 120 μm. Peak shapes were optimised to their best collector position and focus, yielding peaks with high resolution and low residual. The low mass head and high mass head detectors, equipped with Faraday cups, were used for the simultaneous detection of 16O− and 18O−, respectively. The electrometers measuring 16O− and 18O− were set to 10E10Ω (50 V range) and 10E12Ω (5 V range), respectively. For the oxygen isotope measurements presented here, the collector slit widths for both heads were 300 μm.
Each spot analysis lasted around 6 min and consisted of: (1) 60 s of pre-sputtering, (2) ~100 s of automated steering of secondary ions (i.e., adjustments to the secondary ion beam path in the Y and Z directions within the source chamber, prior to the source slit, so as to maximise secondary ion count rates), (3) ~10 s of automated centring of the secondary ions into the collector slits with magnet control, (4) 120 s of data collection (consisting of 6 scans lasting 20 s each), and (5) electron-induced secondary ion emission (EISIE) counts collected before and after data collection (with each set of measurements consisting of 20 s of count time when the electron gun was on, but the Cs primary beam was turned off). Stage navigation lasted additional ~30 s. Analytical precision was monitored using a laboratory standard (Durango apatite 3, δ18OSMOW = 9.8 ± 0.3‰90), measured after every 10–15 experimental measurements. The standard deviation of Durango δ18O values in each analytical session ranged from 0.1–0.2‰ (1σ). All δ18O values are reported to the Vienna Standard Mean Ocean Water (VSMOW) scale91.
To target measurement of enamel secreted at ~weekly intervals (i.e., 7 days apart), we used the calculated enamel extension rate (EER) for each enamel segment identified during histological analysis15,41 (Supplementary Data 1 and Supplementary Figs. 4–8). The first spot was placed as close as possible to the top of the first growth increment, and the subsequent locations were placed at distances of 7 times the segment-specific EER. Where a new segment started before the week was over, an offset was calculated using the number of days of enamel secretion that took place in the new segment times the EER of the new segment. All spots were placed as close as possible to the EDJ, because enamel here is more highly mineralised than peripheral enamel92,93, and thus provides the best time resolution for the progression of enamel extension. However, in the case of ASB402, bubbles that were incorporated during the epoxy casting process meant that the underlying glass was exposed in a small region of the ground section during polishing, necessitating analysis away from the EDJ in segment B.
After the removal of unreliable measurements (identified using photographs taken with the camera inside the SHRIMP; analytical spots that touched the EDJ and that may have been affected by the δ18O values of dentine were deemed unreliable), a total of 23–24 δ18O values were obtained from the second molars (ASB174, ASB360) and a total of 36–44 δ18O values were obtained from the third molars (ASB133, ASB402, ASB449). All individual measurements (including those that were deemed unreliable) are presented in Supplementary Data 2.
To test whether the accuracy of measurements could be impacted by the presence of the 450 × 110 µm wide laser pits left behind by the LA–MC–ICP–MS during 87Sr/86Sr ratio analysis (described above), we performed an experiment where we monitored the primary beam signal and the secondary extraction steering using spots placed increasingly closer to the laser pits. No obvious impact was observed at distances >30 µm from the laser pits, and analytical spots were able to be placed in all the identified target locations.
To test the reproducibility of ~weekly spaced δ18O sequences, two samples (ASB133 and ASB449) were analysed in duplicate. A first set of analytical spots (series A) was placed in the previously identified target locations and a second set of analytical spots (series B) was placed ~50 µm away from each spot in series A (in cervical direction, where possible). Although measurements from series A and series B represent enamel that was secreted a few days apart, the two sequences yielded very similar results (Supplementary Fig. 11). In both cases, series A was used for the main analysis in this study. For further assessment of the accuracy of sample placement, we placed a series of measurements (n = 4) along a single landmark on ASB360 (landmark no. 4 in Supplementary Fig. 5), situated at increasing distance from the cusp. These measurements represent enamel that formed at the same time and have highly reproducible values: 17.91 ± 0.23 (1σ).
It has been observed in the literature that breastfeeding increases δ18O values of body water of nursing mammals17,94,95. In modern wild boars, weaning is complete by 3–4 months of birth. This means that nursing enrichment can likely only impact the oxygen isotope composition of second molars (which start formation around day ~65–70 of life), but not third molars (which start crown formation around day ~210 of life). As the maximum δ18O values of the second molars (ASB174: 20.9‰; ASB360: 22.0‰) are not considerably higher than the maximum δ18O values of the third molars in the ASB133–402 category (ASB133: 19.9‰; ASB402: 20.8‰), it is unlikely that these teeth were affected by nursing enrichment.
Overall, the δ18O sequences are noisier towards the end (with measurements taken from the cervical half of the crown) than they are at the start (with measurements taken from the cuspal half of the crown). This is the result of two factors15,41. First, enamel maturation in the cervical half of the crown does not follow the same incremental pattern as matrix secretion, resulting in a more ‘blurred’ enamel composition during maturation. Second, the EER is markedly reduced in cervical direction, meaning that formation of the cervical half of the crown covers a larger span of time (specifically, three-quarters of the total crown formation time) compared to the cuspal half of the crown. In consequence, daily growth increments are spaced more closely together in the cervical half of the crown, and therefore, the margin of error for SHRIMP spot placement is much lower here than it is around spots in the cuspal region. In other words, a small deviation in sample placement in the cervical region can place the analytical spot on a distinct growth increment, while a small deviation in sample placement in the cuspal region would likely still place the analytical spot close to the target growth increment. The same challenges are faced to a higher degree when analysing human enamel, which has much lower EER than wild boars. This leads to distinctly noisy sequences in the cervical crown region (see Supplementary Materials in Vaiglova et al.80).
Trace element mapping
Trace element concentration of the polished surface of the ground sections was measured using a Laser Ablation Inductively Coupled Plasma Mass Spectrometer (LA–ICP–MS). Four samples (ASB174, ASB360, ASB402, ASB449) were measured at the Geoarchaeology and Archaeometry Research Group facility, Southern Cross University, Australia, using an ESI NWR193 ArF excimer coupled to an Agilent 7700 ICP-MS. One sample (ASB133) was measured at the Research School for Earth Sciences, Australian National University, using a Coherent ExciStar 193 nm ArF excimer laser with HelEx dual-volume ablation cell coupled to an Agilent 8900 Triple Quadrupole ICP-MS. All measurements followed the same set-up and protocol96. Rastered laser beams were placed along the sample surface in adjacent straight lines using a laser spot size of 40 µm, scan speed of 80 µm per second, laser intensity of 80% (~6j per cm2), and a total integration time of 0.50 s. This resulted in data points that correspond to a pixel size of approximately 40 × 40 µm. A full suite of trace elements was measured (7Li, 25Mg, 31P, 43Ca, 44Ca, 66Zn, 88Sr, 138Ba, 140Ce, 208Pb, 238U), and 138Ba was used for interpretation of the wild boars’ seasonal dietary patterns. International reference materials (NIST610 and NIST612) were used to assess signal drift.
Although the stoichiometric concentration of calcium remains constant across the enamel and the dentine portions of teeth, the density varies within and between the two tissues. This variation in density can have an impact on the ablation and may result in the creation of artificial peaks and troughs during measurement, which can be particularly problematic when working with diagenetically altered material. Additionally, although the ground sections were polished to a smoothness of a few microns, small topographic fluctuations between the enamel and dentine structures may result in energy variation at the surface of the sample, which can translate into slight differences in measured values. To correct for this, each trace element was normalised to 43Ca.
Elemental maps (shown as heat maps in Fig. 4) were created using the interactive R Shiny application shinyImaging (Institute for Exposomic Research at the Icahn School of Medicine at Mount Sinai) and Iolite 4.10.082,97,98. Using shinyImaging, individual laser line csv files for each isotope were transformed into counts per second (cps) matrices, with the number of ablation lines multiplied by the number of ablation spots per ablation line. For each element, the gas blank collected during the first 10 s of each laser scan line was subtracted from the raster stack and all the elements were normalised to 43Ca. Within the rectangular scan regions, the areas which did not contain teeth but were filled with epoxy resin were converted to white colouration (corresponding to no intensity) in order to increase the clarity of the maps. Colour scales were applied using the linear blue-red Lookup Table. Using Iolite, the raw data was imported as a single file accompanied by a laser log file that contained the time stamp for each measurement. The data was divided into selections (sample, background, reference materials) and normalised to 43Ca using NIST610. Images were created from the selections using an Empirical Cumulative Distribution Function (ECDF) colour scale99 with upper and lower limits set to two standard deviations around the mean. The aspect ratio was set by specifying the width and the height of the scan region (in µm) using the XY coordinates of the corners. Lastly, a global mask was used to remove the background noise recorded in the epoxy regions using 43Ca as a comparator.
Adherence to Open Science
This article and its accompanying Supplementary Figs. and Data contain all the raw data that was collected in this project and used in the analysis. To adhere with Open Science principles and provide sufficient information that would allow others to reproduce our methods, we have additionally created a page on the Open Science Framework (OSF) repository that contains: the high-resolution jpeg images of the ground sections before and after isotopic and trace element analyses (the images were too large to include as supplementary materials); the SHRIMP Reproducibility Index, which documents all the decisions that have been made while assessing the reliability of the measured δ18O values and identifies which measurements were not included in the final analysis; and the R and Python code that was used to create the map of underlying bedrock 87Sr/86Sr ratios in the study area. So as to not influence and bias the SHRIMP analysis, some of the histological information (i.e., possible presence of any stress indicators) were kept undisclosed until all isotopic data had been collected; only the EER estimates for each identified enamel segment were used to target the SHRIMP analytical spots. In addition, an independent researcher has checked the internal consistency across raw data files and the Supplementary Data files to make sure that no mistakes were propagated during formatting. The OSF page can be found by searching for this paper’s title on the Open Science Framework (https://osf.io), or using the following link: https://osf.io/6fyeu/.
Reporting summary
Further information on research design is available in the Nature Portfolio Reporting Summary linked to this article.
Data availability
Data generated in this study include the results of histological analysis (Supplementary Data 1), tooth enamel stable oxygen isotope (δ18O) values (Supplementary Data 2), tooth enamel strontium isotope ratios (87Sr/86Sr) (Supplementary Data 3), and trace element maps, which can be reproduced using Supplementary Software 3. All datasets have been uploaded to the Open Science Framework (OSF) repository: https://osf.io/6fyeu/. There are no restrictions on data availability and no accession codes are necessary to access the material.
Code availability
R scripts that were used to make Fig. 2, and Supplementary Figs. 9 and 11 are provided in Supplementary Software 1. R and Python scripts that were used to create Fig. 3 are provided in Supplementary Software 2. Instructions and material for re-creating the trace element maps used in Fig. 4 are included in Supplementary Software 3. All scripts with full documentation have been uploaded to the Open Science Framework (OSF) repository: https://osf.io/6fyeu/.
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Acknowledgements
We thank Miles Cornish from Griffith University (Engineering) for assistance with sectioning the teeth; Levi Beeching from the Australian National University (Materials Physics) for carrying out the µCT scanning; Nicholas Condon from the University of Queensland (Molecular Bioscience) and Bence Paul from Elemental Scientific Lasers LLC for software assistance; Sarah Barakat for sharing R code that helped guide the creation of the map of bioavailable 87Sr/86Sr; Johanna Evans for cross-checking all datafiles; Tanya Smith for access to the Griffith Laboratory for Archaeological Science where the teeth were cast into epoxy; and Jacqueline S. Meier and Duncan Wright for helpful comments and discussion. This study was funded by the New Researcher Grant (Griffith University Arts, Education, and Law Group), the Early Career Researcher Grant Development Award (Griffith Centre for Social and Cultural Research and the Australian Research Centre for Human Evolution), and the Early Career Research Support Award (Society for Archaeological Science). The Asiab excavations were financially supported by the David Foundation Grant. The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript. Permission for the research was granted by the Iranian Centre for Archaeological Research at the Ministry of Cultural Heritage, Tourism, and Handicrafts of the Islamic Republic of Iran.
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Petra Vaiglova: conceptualisation; funding acquisition; sample preparation; data analysis; data interpretation; writing: original draft, review & editing; data curation. Horst Kierdorf: sample preparation; data acquisition; data analysis; data interpretation; writing: original draft, review & editing. Carsten Witzel: sample preparation; data acquisition; data analysis; data interpretation; writing: original draft, review & editing. Georgina Falster: data analysis; data interpretation; writing: original draft, review & editing. Renaud Joannes-Boyau: data acquisition; data analysis; writing: method summary. Yue Wang: sample preparation; data acquisition; writing: method summary. Jiade Wu: sample preparation; data acquisition. Ian Williams: data acquisition. Brett Knowles: data acquisition. Yang Wu: data acquisition. Pernille Bangsgaard: data interpretation; writing: original draft, review & editing. Lisa Yeomans: data interpretation; writing: review & editing. Tobias Richter: conceptualisation; data interpretation; writing: original draft, review & editing. Hojjat Darabi: conceptualisation; data interpretation; writing: review & editing.
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Communications Earth & Environment thanks Selena Vitezović and the other, anonymous, reviewer(s) for their contribution to the peer review of this work. Primary Handling Editor: Aliénor Lavergne. A peer review file is available.
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Vaiglova, P., Kierdorf, H., Witzel, C. et al. Transport of animals underpinned ritual feasting at the onset of the Neolithic in southwestern Asia. Commun Earth Environ 6, 519 (2025). https://doi.org/10.1038/s43247-025-02501-z
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DOI: https://doi.org/10.1038/s43247-025-02501-z
