# Multi-isotope biographies and identities of victims of martial victory celebrations in Neolithic Europe

Teresa Fernández-Crespo, Christophe Snoeck, Javier Ordoño, Philippe Lefranc, Bertrand Perrin, Fanny Chenal, Hélène Barrand-Emam, Rick Schulting, Gwenaëlle Goude

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Teresa Fernández-Crespo, Christophe Snoeck, Javier Ordoño, Philippe Lefranc, Bertrand Perrin, et al. Multi-isotope biographies and identities of victims of martial victory celebrations in Neolithic Europe. Science Advances, 2025, 11 (34), 10.1126/sciadv.adv3162. [https://doi.org/10.1126/sciadv.adv3162](https://doi.org/10.1126/sciadv.adv3162). [hal-05217025](https://HAL.inistre.fr/hal-05217025)

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ANTHROPOLOGY

Multi-isotope biographies and identities of victims of
martial victory celebrations in Neolithic Europe

Teresa Fernández-Crespo<fnref n="1" />,²,³*, Christophe Snoeck⁴, Javier Ordoño⁵, Philippe Lefranc⁶,⁷,
Bertrand Perrin⁷,⁸, Fanny Chenal⁷,⁹, Hélène Barrand-Emam⁷,⁸,
Rick J. Schulting³, Gwenaëlle Goude²

Skeletons showing multiple instances of unhealed trauma and isolated skeletal segments of severed left upper limbs have been documented in the Neolithic sites of Achenheim and Bergheim (northeastern France, approximately 4300 to 4150 cal BCE), providing tantalizing evidence of war-related practices of overkill, mutilation, and/or trophy taking. Here, we conduct an innovative multi-isotope reconstruction of the biographies of the "victims" and other individuals from the region that were given normative funerary treatments (nonvictims). A total of 82 humans are analyzed, together with 53 animals and 35 modern plants to establish regional isotope baselines. Results show statistically significant isotopic differences between victims and nonvictims and suggest that the former were members of invading groups brutally killed, perhaps exposed and deposited in pits—together with trophies in the form of severed upper limbs—by local groups in what might be one of the earliest well-documented instances of martial victory celebrations in prehistoric Europe.

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INTRODUCTION

Collective violence in Mesolithic and Neolithic Europe is known to
have taken many forms. Most well-studied cases have been inter-
preted as massacres of entire small communities, with a relatively
recurrent suggestion of young female abduction [e.g., (1-3)], or,
more rarely, as executions and battle graves of mainly adolescent
and adult males [e.g., (4, 5)]. Haphazard deposition is frequent but
not ubiquitous, as well as taphonomic evidence compatible with the
exposure of corpses for a time [e.g., (5, 6)]. Most cases include sev-
eral instances of perimortem blunt force cranial trauma and often of
unhealed long bone fractures and/or projectile injuries. Occasion-
ally, the dehumanization of the victims by brutal savagery has been
suggested, including the breaking of limbs—especially the distal
segment of the lower limb—and even one case of the right upper
limb having been severed through the humerus (7). In a few cases,
the ritualization of violence has been suggested [e.g., (8, 9)].

This is consistent with the observation that ritualized human kill-
ing and mutilation, despite being described in the archaeological and
ethnographic records of many past societies [e.g., (10, 11)], particu-
larly in highly hierarchical early civilizations, seem to be rare in egali-
tarian and intermediate societies, with the possible exception of those
undergoing crisis [(12); although see (13)], particularly warfare.

The late Middle Neolithic of the Upper Rhine Valley provides
one such context. The region witnessed a rapid substitution of local
Bruebach-Oberbergen cultural traditions by western Bischheim
(BORS, i.e., Bischheim Occidental du Rhin Supérieur) traditions
brought by groups originating from the Paris Basin between ap-
proximately 4295 and 4165 cal BCE [modeled at 95.4% probability;

(14, 15)]. This, together with the appearance of fortifications, large
aggregated settlements, and skeletal evidence for violence, suggests
armed conflict, resembling a successful war of conquest (16), the
consequences of which may have affected the lives of the region's
inhabitants for decades.

In this context, circular pits 124 at Achenheim “Strasse 2, RD 45” and 157 at Bergheim “Saulager” (Alsace region of northeastern France) were used for the haphazard deposition of the remains of a number of individuals (6 and 8, respectively) showing multiple instances of unhealed trauma and evidence of overkill (i.e., the use of unnecessarily excessive violence beyond that required to kill), as well as a number of supernumerary and isolated skeletal segments of severed left upper limbs (4 and 7, respectively) (Fig. 1 and text S1) (16, 17).

The combination of such treatments has no previously known
occurrences in the European Neolithic record and does not fit well
with typical massacres or executions of a captured raiding party at-
tacking the local group, as suggested for Early Neolithic Linear-
bandkeramik (LBK) mass graves [e.g., (1, 2)]. Different scenarios
may be envisaged to explain this evidence. One hypothesis is that
the articulated bodies represent enemies captured alive, taken as
prisoners to the community, and then either killed immediately or
after some time. Meanwhile, the severed upper limbs would repre-
sent war trophies removed at the battle site following the violent
encounter and then taken back to the settlement for perhaps fur-
ther transformation and display. Both the articulated bodies and
the severed limbs would lastly be deposited in the pits [e.g., (16, 18)]
during what may plausibly be interpreted as celebrations of victory
(16, 17). However, victims of such war-like practices can also in-
clude community members fallen in combat and repatriated for
home burial (either as entire bodies or selected body parts) [e.g.,
(19)]. Another alternative which cannot be excluded would be to
understand these deposits as resulting from intragroup collective
punishments or sacrifices of social outcasts (20), where torture—
including mutilation—may have been applied beforehand or as a
part of the process (13, 17).

Reconstructing the identity and life histories of the individuals
represented in the pits is, therefore, essential to interpret both sites

<footnote n="1">¹Departamento de Prehistoria, Arqueología, Antropología social y Ciencias y Técnicas Historiográficas, Universidad de Valladolid, Valladolid, Spain. ²CNRS, Aix Marseille Univ, Minist Culture, LAMPEA, UMR 7269, Aix-en-Provence, France. ³School of Archaeology, University of Oxford, Oxford, UK. ⁴Analytical, Environmental and Geo-Chemistry, Department of Chemistry, Vrije Universiteit Brussel, Brussels, Belgium. ⁵Department of Archaeology and New Technologies, Arkikus, Vitoria-Gasteiz, Spain. ⁶Université de Strasbourg, Strasbourg, France. ⁷UMR 7044 Archimède, Université de Strasbourg, Strasbourg, France. ⁸ANTEA-Archéologie, Habsheim, France. ⁹Inrap Grand Est, Centre de Strasbourg, Strasbourg, France.</footnote>

<footnote>*Corresponding author. Email: teresa.fernandezcrespo@uva.es</footnote>
<figure><img src="image_3.png" /><figcaption>**Fig. 1.** Overhead views of late Middle Neolithic violence-related human mass deposits of the Alsace region, France, analyzed in this study. (A) Pit 157 from Bergheim Saulager (Photo credit: Fanny Chenal, INRAP) and (B) pit 124 from Achenheim Strasse 2, RD 45 (Photo credit: Philippe Lefranc, INRAP).</figcaption></figure>

and the mechanisms by which some individuals were selected for such practices. Here, we conduct an innovative multi-isotope approach to reconstructing aspects of diet, social background, and provenience of individuals with unhealed skeletal trauma and/or unconventionally buried (henceforth interpreted as victims) and individuals that show no such evidence and were given what is taken to be a normative funerary treatment (henceforth interpreted as nonvictims). This division is based on the premise that deathways (particularly those involving violence) and mortuary practices are connected to lifeways and social identities to some degree [e.g., (21)].

On the basis of the available evidence, we hypothesize that individuals whose remains were deposited in pits 124 at Achenheim “Strasse 2, RD 45” and 157 at Bergheim “Saulager” sites (i.e., the victims) were captive foreign enemies. We further hypothesize, and subsequently test, that they differed substantially from nonvictims in at least some, if not all, of the following aspects:

1) Average diet and mobility/landscape use during their last years of life, suggesting their affiliation with potentially distinct communities. This is investigated through bulk bone collagen stable carbon (δ¹³C<sub>bcol</sub>), nitrogen (δ¹⁵N<sub>bcol</sub>), and sulfur (δ³⁴S<sub>bcol</sub>) isotope analyses.

2) Infant and child-rearing practices and physiological aspects such as metabolic processes from birth to adolescence, suggesting potentially different cultural backgrounds and/or lifeways. This is investigated through sequential dentine collagen stable carbon (δ¹³C<sub>dcol</sub>) and nitrogen (δ¹⁵N<sub>dcol</sub>) isotope analyses on first and second permanent molars, which allow for a detailed reconstruction of early life histories due to their different developmental timing (22).

3) Landscapes exploited (including the consumption of plants and water from different sources) and mobility patterns during childhood, identifying them as nonlocal to the sites and possibly the region. This is investigated through sequential dentine collagen sulfur (δ³⁴S<sub>dcol</sub>) and bulk enamel carbon (δ¹³C<sub>cap</sub>), oxygen (δ¹⁸O<sub>c</sub>), and strontium (⁸⁷Sr/⁸⁶Sr) isotope analyses on the same permanent molars.

To this end, 49 bone samples (one per individual) and 31 teeth (16 first and 15 second permanent molars from a selection of 16 individuals) are analyzed, from both victims and nonvictims of

Achenheim “Strasse 2, RD 45” and Bergheim “Saulager.” Another 33 human bone samples (one per individual) from 10 Alsatian sites which also date to the end of the fifth millennium BCE and are attributed to Bruebach-Oberbergen, western Bischheim (BORS), Early Michelsberg, and/or Proto-Munzingen cultural groups—comprising nearly all the human skeletal record available for this spatio-temporal context—are also analyzed (Fig. 2, table S1, and text S1).

Last, 53 faunal bone samples from five broadly coeval sites are analyzed to create a regional δ¹³C, δ¹⁵N, and δ³⁴S baseline, and 35 modern tree samples collected from locations characterized by different geologies to establish a biologically available ⁸⁷Sr/⁸⁶Sr baseline [cf. (23)].

## RESULTS

### Summary of data used for analysis

Human and faunal bone samples (n = 135) provided 129 stable carbon (δ¹³C<sub>bcol</sub>) and nitrogen (δ¹⁵N<sub>bcol</sub>) and 124 stable sulfur (δ³⁴S<sub>bcol</sub>) isotope measurements that met conventional collagen preservation criteria (tables S2 and S3) (see text S2 for details). Most human teeth selected for sequential dentine analysis (29 of 31) provided good collagen. The two teeth that failed due to very poor preservation were from the individual in pit 434 of Achenheim. Sequentially sampled dentine collagen increments (n = 665) provided 555 acceptable measurements for stable carbon (δ¹³C<sub>dcol</sub>) and nitrogen (δ¹⁵N<sub>dcol</sub>) isotope analysis (table S4) and 87 for stable sulfur (δ³⁴S<sub>dcol</sub>) isotope analysis (table S5) (see text S2). All enamel samples were successfully analyzed, providing 31 bioapatite carbon (δ¹³C<sub>cap</sub>), 31 carbonate oxygen (δ¹⁸O<sub>c</sub>), and 31 strontium (⁸⁷Sr/⁸⁶Sr) isotope measurements (table S6).

### General trends in isotopic data

General trends in bulk bone collagen isotopic data from both humans and animals from late Middle Neolithic Alsace and in sequential dentine collagen and bulk enamel isotope data from individuals analyzed at Achenheim and Bergheim are presented in text S3, tables S7 to S10, and figs. S1 to S11.
<figure><img src="image_4.png" /></figure>

<table><thead><tr><th>Years BCE</th><th>Northern Alsace</th><th>Breisgau</th><th>Southern Alsace</th></tr></thead><tbody><tr><td><strong>Copper Age / Late Neolithic</strong></td><td colspan="3">Bell Beaker</td></tr><tr><td><strong>Late Middle Neolithic</strong></td><td colspan="3">Corded Ware</td></tr><tr><td><strong>Middle Neolithic</strong></td><td colspan="3">Horgen</td></tr><tr><td><strong>Early Neolithic</strong></td><td colspan="3">Bischheim rhénan</td></tr><tr><td></td><td colspan="3">Roessen</td></tr><tr><td></td><td colspan="3">Planis-Friedberg<br>Grossgartach</td></tr><tr><td></td><td colspan="3">Hinkelstein</td></tr><tr><td></td><td>?</td><td></td><td>?</td></tr><tr><td></td><td>Limbourg</td><td>N Alsatian LBK</td><td>Hoguette</td><td>Hoguette</td><td>S Alsatian LBK</td></tr></tbody></table>

**Fig. 2. Location and chronocultural attribution of the archaeological sites selected for study.** Colors show attribution to main regional facies of the Alsatian late Middle Neolithic, which are displayed on the timeline on the bottom (after 14: Fig. 2). Legend: 1. Achenheim "Strasse 2, RD 45;" 2. Bergheim "Saulager;" 3. Eckbolsheim "Parc d'Activités;" 4. Entzheim "Sablière Oest;" 5. Entzheim "Les Terres de la Chapelle-Zone d'Activités;" 6. Geispolsheim "Schwobenfeld;" 7. Ittenheim "Complexe sportif;" 8. Kolbsheim "Vogesblick;" 9. Matzenheim "Le Lavoir;" 10. Meistratzheim "Station d'épuration intercommunale;" 11. Ungersheim "Desserte Routière du Bioscope;" 12. Vendenheim "Griesklaus/Hasenacker/Les Portes du Kochersberg." BORS, western Bischheim.
## Research hypothesis testing: Differences in isobiographies and identities between victims and nonvictims

Evaluating differences in isotope biography and identity between potential victims of killing, torture, and/or limb severance and non-victims without evidence of such practices of late Middle Neolithic Alsace is limited by the existence of only two deposits with victims in the two main sites studied. In Achenheim, pit 124 has provided isotopic measurements for eight victims, compared to eight nonvictims from other pits. In Bergheim, pit 157 provided measurements for 11 potential victims, while other pits provided measurements from 20 nonvictims. Comparisons are, therefore, affected by relatively small sample sizes, especially in the case of sequential dentine and bulk enamel isotopic analyses, restricted to individuals that permitted sampling of first and second permanent molars (i.e., four victims and five nonvictims from Achenheim and six victims and one nonvictim from Bergheim).

### Were there differences in average diet and mobility/landscape use between victims and nonvictims during their last years of life?

While bulk bone collagen (δ¹³C_bcol) values show no difference between victims and nonvictims (Fig. 3), δ¹⁵N_bcol and δ³⁴S_bcol values exhibit some differentiation. In the case of Bergheim, two findings must be highlighted. First, there is a statistically significant difference in δ¹⁵N_bcol between adult and adolescent victims and nonvictims, observed whether comparing all individuals ($t = 5.499$, df = 12, $P < 0.001$, $d = 2.950$) or only those that are broadly coeval with the victims ($t = 5.457$, df = 10, $P < 0.001$, $d = 3.254$), with victims showing markedly higher values [average δ¹⁵N_bcol: $11.3 \pm 0.6$ per mil (‰) versus $9.7 \pm 0.5‰$] (Fig. 4). Second, there is significantly greater variation in δ³⁴S_bcol values (Levene's test, $P < 0.001$) among victims compared to nonvictims, again whether analyzing all adults and adolescents (SD: $\pm 2.8$ versus $\pm 1.1$) or only those that are broadly coeval with the victims (SD: $\pm 2.9$ versus $\pm 1.3$). Within this greater variation in victims, there are two other important observations.

First, very few values for victims fall within the range shown by non-victims. Second, there is an unexpectedly clear distinction between δ³⁴S_bcol values obtained from complete or almost complete skeletons and from isolated skeletal segments of severed left upper limbs deposited in pit 157, whether infants and children are included ($t = 3.097$, df = 9, $P = 0.013$, $d = 2.101$) or excluded ($t = 2.851$, df = 5, $P = 0.036$, $d = 2.159$) (Fig. 5). The former show higher mean δ³⁴S_bcol values, excepting individuals 2 and 7, which fall within the range of severed upper limbs.

In the case of Achenheim, there is no differentiation in mean δ¹³C_bcol, δ¹⁵N_bcol, and δ³⁴S_bcol values between victims and nonvictims. However, as at Bergheim, there is a statistically significant difference among victims deposited in pit 124 between δ³⁴S_bcol values obtained from complete or almost complete skeletons and from isolated skeletal segments of severed left upper limbs (all from adults and adolescents, $t = 2.510$, df = 6, $P = 0.046$, $d = 2.612$). In this case, only one of the more complete skeletons, individual 4, falls within the range of severed upper limbs. Falling within this range is also an isolated skeletal segment of a probable severed left humerus found during this study mixed with the fauna recovered from Achenheim pit 123, the inclusion of which further reinforces the difference ($t = 2.553$, df = 7, $P = 0.038$, $d = 2.021$) (Fig. 5).

The findings provide an unexpectedly nuanced answer to the question above: Victims, whether represented by complete skeletons or by isolated severed limbs, differed in the foodstuffs they consumed and/or in the environment from which their foodstuffs originated during their last years of life, not only from those of local nonvictims but also from one another.

### Did victims and nonvictims have different infant and child-rearing practices?

Sequential dentine collagen δ¹³C_dcol and δ¹⁵N_dcol values suggest homogeneous ~exclusive (i.e., as seen isotopically, which does not exclude small contributions from supplementary foods) breastfeeding periods for ~6 months between victims and nonvictims, regardless

<figure><img src="image_5.png" /><figcaption>**Fig. 3.** Boxplots of human bone collagen carbon isotope values (δ¹³C_bcol) of adolescents and adults from Bergheim Saulager and Achenheim Strasse 2, RD 45 sites. Values are sorted in three categories: nonvictims, killed individuals represented by complete or almost complete skeletons, and individuals represented by isolated skeletal segments of severed left upper limbs. BER, Bergheim; ACH, Achenheim; VPDB, Vienna Pee Dee belemnite.</figcaption></figure>
<figure><img src="image_6.png" /><figcaption>**Fig. 4.** Boxplots of human bone collagen nitrogen isotope values (δ¹⁵N<sub>bcol</sub>) of adolescents and adults from Bergheim Saulager and Achenheim Strasse 2, RD 45 sites. BER, Bergheim; ACH, Achenheim.</figcaption></figure>

<figure><img src="image_7.png" /><figcaption>**Fig. 5.** Boxplots of human bone collagen sulfur isotope values (δ³⁴S<sub>bcol</sub>) of adolescents and adults from Bergheim Saulager and Achenheim Strasse 2, RD 45 sites. BER, Bergheim; ACH, Achenheim.</figcaption></figure>

of the funerary phase (figs. S7 and S8). No significant differences are detected between victims and nonvictims in the estimated timing of completion of weaning and the mean δ¹³C<sub>dcol</sub> and δ¹⁵N<sub>dcol</sub> values during ~exclusive breastfeeding, weaning and postweaning periods (tables S8 and S9).

However, it is worth noting the consistent duration of both exclusive breastfeeding and weaning cessation among analyzed Achenheim victims (all males), in contrast to nonvictims who exhibited greater variability (table S8). Another relevant observation is that the isotopic profiles of most individuals analyzed show evidence of instances of opposing covariance and/or marked shifts in δ¹³C<sub>dcol</sub> and δ¹⁵N<sub>dcol</sub> values

(figs. S7 and S8), which may relate to physiological stress and/or mobility. The clearest cases comprise not only victims 1, 4, and 6 from Achenheim and 2 and 7 from Bergheim (including, whether coincidentally or not, the three individuals whose δ³⁴S<sub>bcol</sub> values fall within the range of isolated skeletal segments of severed left upper limbs at both sites) but also some nonvictims, such as those from Achenheim pit 286 and from Bergheim pit 118. Although it is difficult to suggest higher frequencies of different processes between groups due to small sample sizes, the most marked shifts detected in the profiles, particularly well-marked in δ¹³C<sub>dcol</sub> and perhaps related to mobility, are seen among some of the Achenheim victims.
These results indicate a complex answer to the posed question: There are no apparent differences between victims and nonvictims in the timing and mean isotopic values of the weaning and postweaning periods, although the victims' sinusoidal isotopic profiles suggest that they might have experienced greater physiological stress and/or mobility in early life.

**Did victims and nonvictims consume foodstuff or drink water from different sources during their early lives? Did they have different mobility patterns across these landscapes?**

Sequential dentine collagen $\delta^{34}S_{dcol}$ values, which derive from the foods (protein) consumed and, by proxy, the environment exploited, do not show significant differences between victims and nonvictims. However, it is worth noting that individuals showing greater variation in $\delta^{34}S_{dcol}$ along the sequence (e.g., individuals 1 and 6 from Achenheim pit 124 or individual 2 from Bergheim pit 157, the aforementioned outlier) are those showing greater variation in $\delta^{13}C_{dcol}$ and $\delta^{15}N_{dcol}$. Another observation is that while most individuals tend to show a rather stable or slightly negative shift from the M1 crown $\delta^{34}S_{dcol}$ values onward, four individuals show a positive and more marked shift. All these cases correspond to victims (male 1 from Achenheim pit 124 and females 1 and 2 and male 3 from Bergheim pit 157), with one—female 2 from pit 157—being a clear outlier, as previously noted.

Enamel $\delta^{13}C_{ap}$ values, which may help identify differential plant consumption and/or use of distinct landscapes, do not show significant differences between victims and nonvictims. However, it is important to note the large positive shift ($z$-score > 2.0) between the M1 and M2 values of victim 7 from Bergheim pit 157 (Fig. 6 and table S6) which, as mentioned above, shows $\delta^{34}S_{bcol}$ values similar to those of the severed upper limbs deposited directly below him. Individual 7 is also—together with nonvictim individual 1 from pit 343 at Achenheim—an outlier ($z$-score > 2.0) in $\delta^{18}O_c$ values, which mainly reflect differences in water sources used. This individual shows lower M1 values and a larger positive shift between the M1 and M2 $\delta^{18}O_c$ values, compared to most individuals at Bergheim (Fig. 6 and table S6).

Last, intrapopulation comparison of $^{87}$Sr/$^{86}$Sr ratios, which address childhood mobility, reveals significant differences in the extent of M1-M2 shifts between victims and nonvictims in both Achenheim ($U = 1.0, Z = 2.205, P = 0.027, d = 1.731$) and overall ($t = 2.175, df = 14, P = 0.047, d = 1.454$), with victims showing higher mean shifts ($0.0009 \pm 0.0005$) than nonvictims ($0.0003 \pm 0.0002$). The greatest shifts are documented among male victims, including individual 7 from Bergheim pit 157, which again stands out as an outlier ($z$-score > 2.0) because of his more enriched $^{87}$Sr/$^{86}$Sr ratios (Fig. 6 and table S6). To test whether this evidence suggests time spent during infancy and childhood in different geological environments, $^{87}$Sr/$^{86}$Sr results have been analyzed on a case-by-case basis to assess “localness.” Unfortunately, regional variability in $^{87}$Sr/$^{86}$Sr ratios is so high that all individuals' M1 and M2 values fall within the biologically available strontium (BASr) ranges ($2\sigma$) estimated for different walking-distance catchments from each site (tables S11 and S12 and fig. S11).

These results suggest that, while most isotope systems capable of differentiating food and water sources do not unequivocally differentiate the infant and childhood landscapes of victims from nonvictims, they consistently indicate that victims had more mobile and/or changing early life histories as reflected by more frequent and pronounced isotopic shifts.

## DISCUSSION

Isotope data obtained from victims of violence who were subject to non-normative mortuary treatments in Achenheim “Strasse 2, RD 45” and Bergheim “Saulager” sites show meaningful differences from those of nonvictims. In the case of Bergheim, victims show significantly higher $\delta^{15}N_{bcol}$ values compared to coeval nonvictims, which could suggest either higher animal-based protein intake or higher physiological stress levels; in either case, a difference in lifeways can be inferred. The most remarkable distinction at both sites, however, is seen in $\delta^{34}S_{bcol}$ values, with victims showing significantly higher variance compared to nonvictims, whose tightly grouped values are consistent with being local and exploiting a more restricted landscape than the victims. The higher variance among victims is driven by a significant difference in $\delta^{34}S_{bcol}$ values at both sites between isolated severed left upper limbs and complete or almost complete skeletons of victims, with the former showing lower and the latter generally higher mean $\delta^{34}S_{bcol}$ values than the nonvictims with which they share burial location. Considering the geographical component that likely explains the small but significant shift of $\delta^{34}S_{bcol}$ values from northern to southern late Middle Neolithic Alsace (see text S3), this may support differences in the environment exploited, suggesting different origins for victims represented by severed upper limbs and by skeletons, despite having been deposited in the same pits. In view of their demise, it is probable that the identities of these victims can be attributed to socially remote, nonlocal enemies that became trophies or captives during battles or raids and that may have been considered by their captors as not properly “human” and hence warranting such treatment (24).

There are a few exceptions where $\delta^{34}S_{bcol}$ values of skeletons resemble those of the severed upper limbs: individual 4 from Achenheim pit 124 and individuals 2 and 7 from Bergheim pit 157. The skeleton of individual 7, an adult male, was found directly overlying the severed upper limbs and presents the only case of overkill at Bergheim, including several unhealed injuries on the cranium, mandible, left clavicle, left scapula, and left ribs and a complete fracture on the left humerus, with the distal segment of the left upper limb missing. He also shows a substantially different life history than the rest of the skeletons deposited above him in pit 157. The latter show isotopic differences with local nonvictims but do not present skeletal signs of violence and show a demography including diverse age categories and both sexes that differs from that of over-killed individuals (all adolescent or adult males) (17). However, they were deposited in direct contact with the severed upper limbs and individual 7, i.e., if not simultaneously, at least over a very short period of time (hours or days), in haphazard positions, which is in line with the hypothesis that they were also killed but in a manner that left no traces on their skeletons (e.g., strangulation, slitting the throat, abdominal stabbing, and/or evisceration). This leaves open the question as to why apparently nonlocal individuals that were all deposited in the same event were recipients of different types of killing and/or mutilations.

With regard to the potential geographic provenience of victims, the $\delta^{34}S_{bcol}$ values obtained from severed upper limbs analyzed from both Achenheim and Bergheim cluster tightly between 3 and 5‰, with their closest parallel being the range shown by supposedly local nonvictims of Achenheim (Fig. 5). A plausible hypothesis is that they belonged to individuals from northern Alsace, where most sites show generally lower $\delta^{34}S_{bcol}$ and higher $\delta^{13}C_{bcol}$ values (see text S3). By contrast, most complete or almost complete skeletons of victims
<figure><img src="image_8.png" /><figcaption>**Fig. 6.** Summary graph of dentine collagen carbon (δ¹³C<sub>dcol</sub>), nitrogen (δ¹⁵N<sub>dcol</sub>), sulfur (δ³⁴S<sub>dcol</sub>), enamel bioapatite carbon (δ¹³C<sub>ap</sub>), enamel carbonate oxygen (δ¹⁸O<sub>c</sub>), and enamel strontium (⁸⁷Sr/⁸⁶Sr) isotope values for the 16 individuals from Bergheim “Saulager” and Achenheim “Strasse 2, RD 45” sites selected for a life history approach. δ¹³C<sub>dcol</sub> and δ¹⁵N<sub>dcol</sub> data represent the mean of the sequential δ¹³C<sub>dcol</sub> and δ¹⁵N<sub>dcol</sub> values corresponding to formation ages of M1 (0 to 3 years) and M2 crowns (2.5 to 8 years) (22), respectively (cf. table S4).</figcaption></figure>
of both sites show higher $\delta^{34}S_{bcol}$ and lower $\delta^{13}C_{bcol}$ values, which are more compatible with values expected for southern Alsace, in view of the correlation between latitude and both $\delta^{13}C$ and $\delta^{34}S$ along the Alsatian Rhine Valley. This correlation is consistent and also explains the significant differences seen in $\delta^{13}C_{(bcol+dcol+ap)}$ and $\delta^{34}S_{(bcol+dcol)}$ values between Achenheim and Bergheim (text S3). However, it may also be that both groups came from other regions, whether nearby or more distant. Bataille et al.'s (25) probabilistic approach to sulfur isoscapes in western Europe showed that Neolithic individuals from interior regions yielded $\delta^{34}S_{bcol}$ values of ~2 to 8‰, including not only the mid-upper Rhine valley but also neighboring regions like the westernmost mid-upper Paris Basin (from which the invading western Bischheim groups are supposed to have arrived) or the easternmost upper Danube Valley. Crucially, this range covers the $\delta^{34}S_{bcol}$ values provided by both the severed limbs and the complete skeletons of victims. Other studies, including sites from the Middle Neolithic Paris Basin [e.g., (26)], have also shown values consistent with this scenario.

The combined presence of isolated skeletal segments of severed left upper limbs and complete or nearly complete skeletons of victims with different proveniences in the same pits may be plausibly interpreted within the context of a war of conquest (16). The most likely scenario is that the severed upper limbs were trophies taken from the bodies of enemies fallen in battle or raids immediately after death and brought to the village. Heads and hands seem to be the most common human trophies documented in the archaeological record (27–30), although written and ethnographic sources often refer to other body parts, including soft tissues which would not generally preserve, such as scalps, ears, or genitals [e.g., (31, 32)]. Conversely, the individuals represented by complete or almost complete skeletons may have been captured in other violent confrontations and taken alive to the same village to be killed—in many cases, overkilled—in martial celebrations of victory, probably in full public view to reinforce community solidarity and “othering” of the enemy (33). At least in the case of pit 124 at Achenheim, such practices seem to have been celebrated in a central community space (16), which may point to an orchestrated piece of political theater [e.g., (34)]. The fact that a number of the hand bones of those upper limbs are either missing (Achenheim) or scattered on the base of the pit (Bergheim) (17) may be in line with the hypothesis that the upper limbs were in the process of decomposing when deposited, suggesting their exposure {perhaps preserved smoked, dried, or embalmed [cf. (35)]} and curation for some time before being deposited in final closing ceremonies. In this scenario, deposition in the pits of trophies together with enemies killed in rituals of triumph would be seen as an act of both revenge and honor restoration for relatives or allies fallen and/or injured in previous violent encounters. This might explain the special treatment observed for the left upper limbs at both sites, which may be evidence of the cultural affinity of the “perpetrators” at Bergheim and Achenheim, considering that ways of mutilating seem to be culturally specific in many prestate societies (36).

It may be that the bodies of the individuals killed were also considered trophies. This could explain differences in provenience with regard to severed upper limbs. It is both energetically logical and ethnographically documented that the size of trophies taken is usually inversely proportional to the distance to the place whence the trophy was taken. Thus, body segments such as upper limbs are more likely to be taken over mid-range distances and complete bodies—whether alive or dead—over short-range distances [(37), after (38)].

In our case, this might explain the fact that $\delta^{34}S_{bcol}$ values exhibited by isolated severed upper limbs are more divergent from those of supposedly local individuals than those represented by complete or almost complete skeletons of victims (Fig. 5).

An alternative explanation is that severed upper limbs were the result of mutilations intended to humiliate and physically impair enemies, such as those described in classical sources in the context of collective war-related sacrifices (e.g., Appian, *Historia Romana*, Ib.68, Ib.93; Strabo, *Geographia*, III.6; Xenophon, *Hellenica*, II.31). In this scenario, all victims would have been brought alive to the site as captives from different locations and then treated (or punished) differently for unknown reasons (e.g., their origins). Survivors of such mutilations (if any) may have been retained as slaves or even adopted, perhaps by families who lost many members during conflict, as documented ethnographically among some Indigenous eastern North American communities (39). Although this hypothesis appears difficult to reconcile with the available osteological evidence, it may not be coincidental that in both sites severed upper limbs show isotope values consistent with northern Alsace and most skeletons of killed individuals with southern Alsace, which, if actually translated to different identities, might supply a reason for the differential treatment of captives.

While the isotopic biographies of individuals whose upper limbs were severed cannot be further assessed, it is possible to delve deeper into the differences between complete or almost complete skeletons of victims and nonvictims deposited in Achenheim and Bergheim through sequential dentine and bulk enamel isotope analyses. The main differences observed here are consistent with the identification of killed individuals as outsiders. First, victims tend to show greater variation in their sequential $\delta^{13}C_{dcol}$, $\delta^{15}N_{dcol}$ and $\delta^{34}S_{bcol}$ isotope profiles after the cessation of breastfeeding. This greater variation apparently includes a higher number of patterns of opposing covariance between $\delta^{13}C_{dcol}$ and $\delta^{15}N_{dcol}$ values, which are commonly attributed to recurrent periods of malnourishment and/or illness during childhood [e.g., (40)]. Second, they also seem to show more marked shifts between values attributable to mobility or exploitation of varied landscapes (particularly with regard to $\delta^{13}C$), especially in the case of Achenheim pit 124. It is precisely greater mobility that is suggested by higher mean M1-M2 shifts in $^{87}$Sr/$^{86}$Sr ratios among victims—especially males—compared to nonvictims, as well as by the fact that most outliers in bulk enamel isotope systems are victims (e.g., individuals 2 and 7 from Bergheim pit 157).

Of course, this overall greater mobility during childhood does not demonstrate that victims represented by complete or nearly complete skeletons were invaders or migrants—even if the comparison between $^{87}$Sr/$^{86}$Sr values and Alsatian BASr ranges were more diagnostic—nor can it be interpreted as greater mobility in adulthood (41). But considering all the available isotopic evidence, we propose their interpretation as incomers, possibly from western Bischheim (BORS) communities originating from the Paris Basin (42) that had settled in the northernmost part of the Middle Rhine Valley some time before they finally ousted local Bruebach-Oberbergen communities. A number of these individuals—if not all—would have potentially been taken in war, and killed and perhaps displayed in celebrations of victory, in view of the indications of graphic violence performed and their deposition together with severed upper limbs as body trophies. Individuals executed and deposited in the Neolithic LBK mass grave of Halberstadt, in present-day Germany, have also been isotopically characterized as nonlocals. In this case,
trauma was systematically directed at the back of the head, with only four clear postcranial fractures affecting two humeri, one femur and one rib. Given the nature of the evidence, here, the more likely explanation is that victims were members of a raiding party who were captured and subjected to capital punishment. The possibility that they were prisoners taken from afar was dismissed on the basis that only young adult females seem to have been captured alive in LBK massacres (5). Even if the execution of enemies had a standardized performance and was a public event to instill fear of the consequences of transgression as a deterrent (43, 44), in Halberstadt, there is no convincing evidence of further ritualization, whether including extreme and deliberately spectacular violence—usually with an “overkill” element—(e.g., torture spectacle) or the impairment, humiliation, and dehumanization of the defeated enemy—either in this or the next life or both (31)—and their exhibition (33), in contrast to Achenheim and Bergheim.

In this regard, it has been proposed that ritualized killing and trophy deposition in the context of prehistoric and ancient triumphal celebrations may have been understood as a votive offering fulfilling a pact initiated when war leaders sought supernatural (ancestors and “gods”) favor before battle (45). Victims may have addressed spiritual goals (e.g., provided food and slaves for ancestors or gods), legitimized political authority through the display of merciless brutality (19), and paid tribute to glory. The dehumanized image created of the enemy—commonly portrayed as depraved and evil, thus meriting cruelty and retaliation—during protracted conflict would enable perpetrators to kill and maim and the rest of the community to support such actions through moral disengagement (46). The fear of what happens if a demonized enemy is not overcome—or at least contained—would become a justification for perpetuating or even escalating violence. Victories celebrated in the Alsatian sites of Achenheim and Bergheim may be interpreted as pyrrhic, however, as the local Bruebach-Oberbergen culture would be rapidly replaced by western Bischheim (BORS) culture brought by the invading groups.

## MATERIALS AND METHODS

### Materials

Samples were obtained from the Alsatian late Middle Neolithic sites of Achenheim “Strasse 2, RD 45,” Bergheim “Saulager,” Eckbolsheim “Parc d’Activités,” Entzheim “Sablière Oesch,” Entzheim “Les Terres de la Chapelle-Zone d’Activités,” Geispolsheim “Schwobenfeld,” Ittenheim “Complexe sportif,” Kolbsheim “Vogeseblick,” Matzenheim “Le Lavoir,” Meistratzheim “Station d’épuration intercommunale,” Ungersheim “Desserte Routière du Bioscope,” and Vendenheim “Griesklaus/Hasenacker/Les Portes du Kochersberg” (see text S1 for a more detailed background on the sites). At the time of the study, all the archaeological human and faunal samples collected for the research were hosted in Alsatian research and/or curatorial institutions, including Inrap Alsace, the Centre de Conservation et d’Étude d’Alsace, the Service Régional de l’Archéologie, the Musée d’Archéologie de Strasbourg, and the Musée Zoologique de Strasbourg (see text S4 for more details on the provenance of the human remains included in the study). A protocol of measures to diminish the impact of sampling was agreed in advance of the study with the Direction régionale des affaires culturelles Grand Est, the main authority for the curation of Alsatian archaeological collections. Thus, sampling was supervised by local curators and fully documented. All the samples

were taken from skeletal elements that were confidently attributed to specific individuals during the excavations—despite peri- and/or postmortem fragmentation—and maintained as such at the hosting museums. The selection prioritized skeletal elements that had antimeres (i.e., from the opposite side) present, were not expected to have a special display value, and/or already showed postmortem damage on the anatomical areas selected for sampling. In the few instances that this was not possible, samples were taken from areas with lower morphological and visual impact (e.g., interior of mandibles).

In the case of teeth, considering the uniqueness of each specimen, only first and second permanent molars from individuals who preserved at least one antimere of the same tooth in either the mandible or the maxilla or both were selected. After the extraction of the selected teeth and before proceeding with the laboratory treatment and analyses, the crown of each tooth was cast using a Colténe dental casting system to record dental morphology and wear, and each tooth was three-dimensional recorded in full at the laboratory through high-precision photogrammetry. Only one complete half of each tooth was used for analysis (see below), and the other half was returned to the corresponding host institution, permitting future replication of the analyses as well as other approaches.

Plant samples taken from living trees were used to characterize the regional strontium ($^{87}$Sr/$^{86}$Sr) isoscape. They were sampled from different geological formations, targeting diverse sampling locations within each formation (cf. table S11 and figs. S12 and S13) to create a BASr map. Nature reserves were excluded from sampling, and, where possible, trees growing close to farmland were avoided to prevent potential contamination from modern sources (e.g., fertilizers and pesticides) [cf. (47)]. Two samples from different trees were preferably taken from each location to assess variability and identify potential outliers.

### Methods

#### Stable carbon ($\delta^{13}$C), nitrogen ($\delta^{15}$N), and sulfur ($\delta^{34}$S) isotope analysis

Bulk bone collagen was extracted following the protocol in the study of Richards and Hedges (48). Incremental primary dentine collagen microsamples for $\delta^{13}$C and $\delta^{15}$N analysis were sampled from cusp to apex from either the medial or distal side of a demineralized 1.5-mm-wide mid-tooth longitudinal slice following Czermak et al.’s (49) method, using a 1-mm-diameter biopsy punch. Those for $\delta^{34}$S analysis were obtained by dividing the opposite side of the same slice in four main areas (crown, neck, upper root half, and lower root half) using a scalpel, to obtain sufficient material for analysis given the presence of sulfur in only one amino acid (methionine). Age alignment was conducted after the study of AlQahtani et al. (21), following the protocol of Czermak et al. (49). In both cases, after demineralization, samples were sequentially treated with 0.1 M NaOH for 30 min and 0.5 M HCl for 1 hour at room temperature with thorough rinsing with deionized water between and after each reagent [cf. (50)].

Resulting denatured bone collagen samples and nondenatured dentine samples were both freeze-dried before being weighed into tin capsules and loaded into a SERCON 20-22 continuous flow isotope ratio mass spectrometer coupled with an elemental analyzer at the Research Laboratory for Archaeology and the History of Art of the University of Oxford for $\delta^{13}$C and $\delta^{15}$N analysis and into a continuous flow isotope ratio mass coupled with a Europa
Scientific elemental analyzer at Iso-Analytical (Crewe, UK) for δ³⁴S analysis.

Bone collagen samples were all measured in duplicate for δ¹³C and δ¹⁵N, while only one in every five samples was measured in duplicate for δ³⁴S. Dentine collagen samples were not measured in duplicate for δ¹³C and δ¹⁵N due to the relatively low yields resulting from the sampling technique and to the impossibility of precisely replicating a sample elsewhere on the same tooth due to the complexity of primary dentine deposition (51). The larger δ³⁴Sdcol samples provided sufficient yields for duplicate measurements in six cases, resulting in a mean difference of 0.2 ± 0.1‰ between the two runs. This is well within the measurement error, and, moreover, the comparison between runs shows no significant differences (paired t test; *t* = 1.238, *df* = 5, *P* = 0.271).

Collagen quality was assessed using the conventional criteria for bone collagen: %collagen between 0.5 and 22% by weight, %C between 15.3 and 47%, %N between 5.5 and 17.3%, %S between 0.13 and 0.35%, atomic C:N ratio between 2.9 and 3.6, atomic C:S ratio between 300 and 900, and atomic N:S ratio between 100 and 300 (52–55). With regard to dentine collagen, a more conservative C:N range of 3.1 to 3.3 was used. Recent work on fluorescence screening of dentine collagen preservation in human primary dentine suggests that C:N ratios higher than 3.3 in nondenatured dentine samples indicate some degree of collagen decay or degradation, which may result in lower %N (56). Also, only samples weighing ≥0.35 mg were considered reliable, because lower weights may affect mass spectrometer measurements [e.g., (57)].

Stable δ¹³C and δ¹⁵N values were all calibrated to Vienna Pee Dee belemnite and ambient inhalable reservoir, respectively, using cow and seal bone collagen, caffeine, and alanine as control check standards. Separate alanine standards were used for instrument drift correction. Stable δ³⁴S values were calibrated to Vienna Cañon Diablo Troilite using IA-R061 (barium sulphate). IAEA-SO-5, IA-R068, and IA-R069 were used as control check standards. On the basis of Szpak et al.'s (58) formulae, the precision [*u*(*R*(*w*)] of the measurements are determined to be ±0.07‰, ±0.10‰, and ±0.24‰, for δ¹³C, δ¹⁵N, and δ³⁴S, respectively, based on repeated measurements of calibration standards, check standards, in-house laboratory standards, and sample replicates. Accuracy or systematic errors [*u*(*bias*)] are determined to be ±0.14 for δ¹³C, ±0.15 for δ¹⁵N, and ±0.33 for δ³⁴S, based on the difference between the observed and known δ values of the check standards and the long-term SDs of these check standards. The total analytical uncertainties are estimated to be ±0.16 for δ¹³C, ±0.18 for δ¹⁵N, and ±0.41 for δ³⁴S.

### Carbonate carbon (δ¹³C<sub>cap</sub>), oxygen (δ¹⁸O<sub>c</sub>), and strontium (⁸⁷Sr/⁸⁶Sr) analyses

The teeth were cleaned by abrasion with a diamond drill to remove the outer layer to minimize contamination with exogenous strontium. Once cleaned, further drilling was carried out to obtain 50 mg of powder, avoiding the dentine-enamel junction which is more prone to diagenesis. The resulting powder was then further homogenized using a mortar and pestle and pretreated using 0.1 M acetic acid for 30 min and then rinsed three times with Milli-Q following the study of Pellegrini and Snoeck (59). For δ¹³C<sub>cap</sub> and δ¹⁸O<sub>c</sub> analyses, ~1 mg of pretreated powder was measured on a Nu Perspective isotope ratio mass spectrometer with a NuCarb carbonate preparation device at the Vrije Universiteit Brussel (VUB, Brussels, Belgium). Internal matrix matched standards ENF (enamel) and CAB [calcined bone; see (60)] were used alongside Iso-Analytical IA-R022 and

international standards IAEA-603, IAEA-CO8, and NBS18. Repeatability of IA-R022 (*n* = 32) yielded an SD of reproducibility of 0.15 and 0.17‰ for δ¹³C<sub>cap</sub> and δ¹⁸O<sub>c</sub>, respectively. For the matrix matched standards CBA (*n* = 4) and ENF (*n* = 4), reproducibility was better than 0.3‰ for both δ¹³C<sub>cap</sub> and δ¹⁸O<sub>c</sub>. Each sample was measured in duplicate, and the mean value is presented here.

The remaining enamel powder was analyzed for ⁸⁷Sr/⁸⁶Sr. Modern plant samples were also analyzed for ⁸⁷Sr/⁸⁶Sr. They were left to dry naturally at room temperature before being processed using micro-wave digestion following the protocol described in (61). Strontium was extracted from enamel powder and plant samples following the procedure described in (62) and measured on a Nu Plasma 3 multicollector inductively couple plasma spectrometer (PD017 from Nu Instrument) at the VUB.

All the data were corrected for mass fractionation by internal normalization to ⁸⁶Sr/⁸⁸Sr = 0.1194. All sample measurements were normalized externally using a standard bracketing method with the recommended value of ⁸⁷Sr/⁸⁶Sr = 0.710248 for NBS987 (63). The ⁸³Kr and ⁸⁵Rb beam intensities are continuously monitored to prevent any spectral interference; the intensities are always below 0.05 mV, which is negligible compared to the total Sr beam intensity (7 to 8 V). Procedural blanks were considered negligible [total Sr (V) of max 0.02 versus 7 to 8 V for analyses, i.e., ≈0.3%]. For each sample, the ⁸⁷Sr/⁸⁶Sr value was reported with a 2 SE (absolute error value of the individual sample analysis—internal error).

During the course of this study, repeated measurements of NBS987 yielded a ⁸⁷Sr/⁸⁶Sr of 0.710239 ± 0.000028 (2 SD, *n* = 22), which is consistent with the mean value of 0.710252 ± 0.000013 (2 SD, *n* = 88) obtained by thermal ionization mass spectrometry (63). Matrix matched standards National Institute of Standards and Technology (NIST) Standard Reference Material (SRM) 1400 (cow bone ash) and NIST SRM 1515 (apple leaf) were also measured and returned ⁸⁷Sr/⁸⁶Sr values of 0.713112 ± 0.000036 (2 SD, *n* = 5) and 0.713961 ± 0.000027 (2 SD, *n* = 4), which are consistent with those reported in (64).

### Statistical tests

Shapiro-Wilk tests were used to assess whether the data were normally distributed. Levene's tests were used to assess differences in the homogeneity of variances. For two-sample comparisons, Student's *t* tests were used when the data did not depart significantly from a normal distribution; when they did, nonparametric Mann-Whitney *U* tests were used. For more than two groups, one-way analysis of variance (ANOVA) tests were conducted, since all the datasets analyzed were normally distributed with approximately equal variance. Tukey's post hoc tests were used to detect significant differences between sample groups. Two-level mixed-model nested ANOVA tests were used to assess variation in dentine collagen value means among groups (i.e., victims versus nonvictims, between sites) since the comparison involves one measurement variable (δ<sup>xx</sup>X<sub>dcol</sub>) and two nominal variables (individuals and groups), the latter being nested (i.e., forming subgroups within groups) (65). Because of unequal group sample sizes, the Satterthwaite formula was applied (66). This permits the use of modified mean squares at each level, providing a better estimate of the effective degrees of freedom and, therefore, more accurate *P* values. Effect size was assessed using Cohen's *d* (67). This statistic refers to the difference between two sample means expressed in pooled SDs, assuming approximately equal variances. Conventionally, a *d* value of less than 0.2 is seen as trivial, regardless of the outcome of the significance tests. Values ≥0.8 are
considered large. Pearson's r/r² and Spearman's rho coefficients were used to assess correlations as appropriate. Z-scores were calculated to detect the presence of outliers (≥2). A significance level of α = 0.05 and two-tailed testing was used for all statistical tests. Means and SDs are used as measures for central tendency and dispersion throughout the text, unless otherwise stated.

## Supplementary Materials

The PDF file includes:

Supplementary Text S1 to S4

Figs. S1 to S13

Legends for tables S1 to S12

References

**Other Supplementary Material for this manuscript includes the following:**

Tables S1 to S12

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**Acknowledgments:** We thank A. Davadie, from the Direction régionale des affaires culturelles Grand Est, for supporting the project by providing sampling permissions and facilitating access to the collections. We are also grateful to H. Koehler and M. Wunderle (CCE), D. Minnip (Inrap), B. Schnitzler (MAS), and R.-M. Arbogast (MZS) for welcoming, supervising, and helping our research team during sampling at their institutions. We also thank N. Mattielli, W. Debouge, and J. de Jong from the G-TIME laboratory of the Université Libre de Bruxelles for the support with the strontium isotope analyses. **Funding:** This project received funding from the European Union's Horizon 2020 research and innovation programme under the Marie

Skłodowska-Curie grant agreement no. MSCA-IF-790491 to T.F.-C. **Author contributions:** Conceptualization: T.F.-C., P.L., F.C., R.J.S., and G.G. Funding acquisition: T.F.-C. and G.G. Project administration: T.F.-C. and G.G. Resources: C.S., B.P., F.C., H.B.-E., and G.G. Data curation: C.S., F.C., and G.G. Methodology: T.F.-C., P.L., R.J.S., and G.G. Investigation: T.F.-C., C.S., J.O., B.P., H.B.-E., and G.G. Formal analysis: T.F.-C., C.S., J.O., and G.G. Validation: T.F.-C., F.C., and G.G. Writing—original draft: T.F.-C., J.O., R.J.S., and G.G. Writing—review and editing: T.F.-C., C.S., J.O., P.L., B.P., F.C., H.B.-E., R.J.S., and G.G. Visualization: T.F.-C., J.O., F.C., and G.G. Supervision: T.F.-C., P.L., R.J.S., and G.G. **Competing interests:** The authors declare that they have no competing interests. **Data and materials availability:** All data needed to evaluate the conclusions in the paper are present in the paper and/or the Supplementary Materials.

Submitted 13 December 2024

Accepted 27 June 2025

Published 20 August 2025

10.1126/sciadv.adv3162