https://orcid.org/0000-0001-9377-298X
Figures
Abstract
The establishment of the first livestock communities in the eastern Iberian Peninsula during the 6th millennium cal BCE marked a significant transition in the region’s economic system. The research of early animal management practices provides crucial valuable understanding into feeding and pastoral strategies, revealing insights into the social organization of landscapes and their resources. Using stable carbon (δ13C) and nitrogen (δ15N) isotopic analyses of faunal remains this study investigates the interplay between environmental conditions, animal management practices, and dietary habits in Neolithic herds from Mas d’Is and Niuet sites in the Serpis Valley. The landscape of the area, characterized by Mediterranean forests and marshes, provided a variety of plant resources. Although most of the valley is covered by C3 types of plants, there were some clusters of C4 vegetation. The isotopic analyses prove that the local herds can be disaggregated in two groups: some animals with a diet based on C3, some feeding on C4 plants. These results reveal diverse feeding strategies and, by default, spatialized management practices. In addition, the animals presenting a C3 signature, have, as well, strong nitrogen values and marks on the bones that indicate they were part of the agricultural cycle as traction force. This complex organisational strategy, with two herds that show different levels of husbandry intensification, might reflect broader socio-economic systems, where the landscape may have been managed with a mosaic of different property regimes: where marginal areas might have been considered as communal or open-access resources, while more fertile areas nearby settlements, might have been used as crop fields, and individually or household managed. The integration of livestock into agricultural systems and the selective use of landscape resources highlight an adaptive and dynamic approach to animal husbandry in response to environmental and social factors during the Neolithic in the Serpis Valley.
Citation: Navarrete V, Vaccaro I, Escribá P, Grau E, García-Puchol O, Bernabeu Aubán J (2025) Isotopes, herds, and landscape management practices: New insights on early farming communities in the Serpis Valley (Eastern Iberian Peninsula). PLoS One 20(6): e0325137. https://doi.org/10.1371/journal.pone.0325137
Editor: Peter F. Biehl, University of California Santa Cruz, UNITED STATES OF AMERICA
Received: October 23, 2024; Accepted: May 6, 2025; Published: June 27, 2025
Copyright: © 2025 Navarrete et al. This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Data Availability: All relevant data are within the paper and its Supporting Information files.
Funding: The research was supported by the Prometeo 2021/007: NEONETS. A social network approach to understand evolutive dynamics of Neolithic societies (C. 7600-4000 cal. BP), and PID2021-127731NB-C21 EVOLMED. Evolutionary cultural patterns in the contexts of the neolithisation process in the Western Mediterranean. Funding acquisition: Oreto García-Puchol, Joan Bernabeu Aubán. The funders had a role in study design, data collection, decision to publish, and preparation of the manuscript.
Competing interests: The authors have declared that no competing interests exist.
Introduction
The study of domestic fauna biochemistry and morphology remains offers an opportunity to understand long-gone, feeding practices and pastoral management strategies. These strategies, in turn, provide insight into the social organization of the landscape and its resources. The comprehension of key aspects of landscape usage such as herd management, intensity of environmental resources’ use, and settlement patterns contribute to the reconstruction of social practices during the establishment of the first livestock communities in the Iberian Peninsula during the 6th millennium cal BCE. Understanding herd feeding practices is crucial for elucidating the relationship between sedentary societies and their environment, including the territorial distribution of grazing areas, spatial distribution of settlements, and access to distinct pasture areas [1–5]. Recent studies on the Neolithic in the northeast, south and west of the Iberian Peninsula, integrating δ13C and δ15N stable isotope analyses in zooarchaeological research, have revealed a high degree of variability in animal management practices, characterizing the access of flocks to distinct pasture areas and/or crop fields, and the use of fodder derived from crop by-products [6–11]. Nevertheless, this approach has not yet been applied to explaining early husbandry practices in the east of the Iberian Peninsula, one of the pioneering areas in the expansion of the Neolithic in Iberia.
In the archaeological context of the eastern Iberian Peninsula, the introduction of livestock practices inaugurated a distinctive phase that represented a new stage in territorial occupation. During the Neolithic, settlements were relocated from topographically dominant localization to the bottom of the valleys. In addition to these valley settlements, caves and shelters were used for specific economic and symbolic purposes (e.g., sheepfold caves and burials spaces), highlighting a multifaceted approach to land use [12–16]. As the Neolithic progressed, evidenced by archaeological record from the 4th and 3rd millennium BCE, sedentary settlements spread over large territories. Segmented ditches delimited open-air settlements and featured numerous structures, such as siles and pits [17–20]. By the 3rd millennium cal BCE, a new cycle in the socio-ecological dynamics emerged, driven by demographic expansion, intensification of economic practices, and growing social complexity in the east of the Iberian Peninsula [21–25].
The Serpis Valley, is revealed as one of the first areas that present evidence of the spread of the Neolithic in the Iberian Peninsula around the middle of the 6th millennium cal BCE, offering onwards a rich and deep archaeological landscape (Fig 1). Numerous sites have been identified along the Serpis River and surrounding areas, from the early Neolithic to the Bronze Age. Archaeological excavations at sites such as Mas d’Is, Cova de l’Or, Abric de la Falguera, and Benàmer, which from the 6th and 5th millennia cal BCE [14,15,26,27], as well as Jovades, Niuet, and La Vital from the 4th and 3rd millennia cal BCE [17,20,21]; along with others such as Cova de les Cendres, Barranquet, En Pardo, Randero, Missena, Arenal de la Costa and Colata [28–33], have provided archaeological remains that offer a multifarious view of the economy and the environment throughout the Neolithic period.
1-Niuet, 2- Mas d’Is, 3- Benàmer, 4-Cova de l’Or, 5-Jovades, 6-La Vital, 7-Arenal Costa, 8-Colata, 9-Cova Cendres, 10-Missena, 11-Abric de la Falguera, 12-Barranquet, 13-Cova d’en Pardo, 14-Randero. Gaianes’ Lagoon and Serpis River are also indicated.
From the beginning of the Neolithic, the main species of domesticated animals and plants were present in the region [34,35]. Agriculture and livestock breeding were the primary subsistence strategies for communities, with a relatively small contribution from hunting and gathering. The location of the settlements at the bottom of the valleys, surrounded by fertile soils, suggests the development of intensive agriculture with hoe in fields close to the inhabited centres [36–38].
Archaeobotanical data from several sites reveal a broad range of crops with a predominance of cereals. While wheat (Triticum aestivum/durum) and naked barley (Hordeum vulgare var. nudum), were the most common, hulled cereals like einkorn (Triticum monococcum), emmer (Triticum dicoccum) and hulled barley (Hordeum vulgare subsp. vulgare), also played a significant role in the cultivar portfolio [39]. A notable change occurred during the Neolithic was the reduction in the variety of cultivated cereals, that is, a process of relative specialization, which became particularly apparent starting on the 4th millennium cal BCE. This process may have started in the 5th millennium cal BCE, although data for this period is limited [40]. From the 4th millennium cal BCE onwards, both wheat and naked barley, became the only cultivated cereal species [40]. Evidence of woodland clearcutting for larger-scale agriculture, combined with the prevalence of cereal-dominated crop assemblages, indicate that significant transformations were occurring in the economic system [37].
Husbandry practices were characterized by herds composed of caprines (Ovis aries and Capra hircus), cattle (Bos taurus) and pig (Sus domesticus) [35]. The regional archaeozoological record suggests a significant economic change by the end of the 4th millennium cal BCE, marked by a notable increase in cattle remains and, compared to the 6th millennium cal BCE data, a decrease in caprine remains. During the 6th millennium cal BCE, cattle remains were represented in very low percentages at sites such as Cova de les Cendres with a 1% [41], Cova de l’Or with a 2% [42] and Cova la Sarsa with a 5.5% [43]. The limited data available from the 5thmillennium cal BCE prevents direct comparisons of faunal remains between these periods. By the end of the 4th millennium cal BCE, there was a progressive increase in cattle remains in the Serpis Valley, with sites like Jovades with a 15% [44] and Niuet with a 27% [45] showing significant growth. By the 3rd millennium cal BCE, cattle represented over 35% of the domestic species in sites such as La Vital with a 35.5% [46] and Arenal de la Costa with a 36% [44]. Navarrete et al. [45], suggest that the increase in cattle remains in the Serpis Valley could be linked to early occupation of the bottom of valley and the development of the territory according to the open-air settlement structuration, benefiting cattle breeding, as these areas had favourable conditions such as vast plains, access to water, and abundant high-quality herbaceous grasses. These conditions were particularly prevalent in river and lake valleys, offering ample grazing areas for herds. Additionally, cattle could have acquired an additional economic significance due to their role in agricultural work (e.g., animal traction) and their contribution to milk and meat production. Slaughtering patterns and bone pathologies, such as joint injuries in distal extremities, observed at sites like Niuet [45] and Jovades [44], could be indicators of these changes.
These developments in livestock management must be contextualized within the broader ecological setting of the Serpis Valley, whose landscape may have played a key role in shaping agropastoral strategies. During the early Holocene, the Mediterranean landscape was dominated by deciduous forests and conifers, interspersed with open areas like pastures covered by herbaceous plants [47]. In particular, the Serpis Valley featured Mediterranean forests dominated by holm oaks and pines, with riparian zones along the Serpis River characterized by poplar and willow groves, as well as reedbeds [48,49]. Within this valley is the Gaianes Lagoon, a wetland with a rich and biodiverse ecosystem (Fig 1). The vegetation in this area is typical of wetlands and marshes, comprising a wide variety of plant species adapted to flooding and high humidity conditions. The lacustrine landscape includes reeds and rushes, aquatic vegetation like duckweed and pondweeds, marsh plants such as sawgrass and club-rushes, riparian vegetation including willows and alders, halophytic plants like glassworts and sea lavenders, and emergent vegetation such as sea milkwort [50].
By combining carbon (δ13C) and nitrogen (δ15N) stable isotope analysis of bulk collagen with archaeozoological data, the study aims to investigate early husbandry practices in the Serpis Valley, a key area for understanding the spread of the Neolithic in the Iberian Peninsula. This study characterises management strategies by identifying grazing patterns and foddering practices. Particular attention is given to how these practices were influenced by the environmental context of the Serpis Valley, including the availability of fertile valley bottoms, riparian ecosystems, and wetland areas. Additionally, this study contributes to a broader understanding of landscape use and subsistence strategies during the Neolithic. To address these objectives, two Neolithic sites from the Serpis Valley -Mas d’Is and Niuet- have been selected, as they provide a representative sequence for the regional Neolithic occupation.
Archaeological sites
Mas d’Is site
The Mas d’Is is an open-air site located within the municipality of Penàguila (Alicante) at 600 metres above sea level (masl), in the valley bottom of the Serpis Valley. It is located at the head of the Penàguila River, a tributary of the Serpis River, the main watercourse in this region (Fig 2A). The site covers a large area with documented occupations from the early Neolithic to the Bronze Age, including some of the earliest radiocarbon dates connected to the Neolithic in the Iberian Peninsula (Table 1). The first Neolithic occupations are documented at the levels associated with the habitat structures of sector 80 [51] and at the base of the ditch (F4) [26]. F4 presents the most extended stratigraphic sequence within the site, and archaeological research has identified eight archaeological phases (Fig 2A).
A) Above: plan view of the Mas d’Is site, showing the location of ditch F4. Below: stratigraphy of ditch F4. B) Above: plan view of the Niuet site, showing the location of Sector A. Below: stratigraphy of Sector A.
Phases 8 to 5 correspond to the Neolithic IA, IB and IIA in the regional sequence (from c. 5600 to 4850 and to 4550 to 4250 cal BCE [52]). Although Phases 8 and 7, represent the early Neolithic (Impressa and Cardial phases [53]), the scarcity of faunal remains limits their relevance for the present study. Instead, the focus is on Phase 6 and 5, which offer valuable insights into herd management strategies and their socioeconomic implications. Phase 6, associated with the Neolithic IB, dated between 5250-4850 cal BCE [52], is characterized by incised-impressed ceramics. Phase 5 aligns with the regional Neolithic IIA, dated between 4550-4250 cal BCE [52]. Subsequently, various episodes concluded the prehistoric sequence of the site at a time parallel to the Bronze Age [12,51].
The subsistence economy during Phase 6 and 5 appears to have been primarily farming practices, complemented by a broad exploitation of forest resources. Despite the poor preservation of organic matter, the zooarchaeological results show that in both phases domestic species (caprine, pig and cattle) predominated over wild species (deer and rabbit), indicating a husbandry practice. Although the percentage of remains is limited, Phase 6 shows a prevalence of cattle remains, while Phase 5 is dominated by pig remains, which may suggest a change in herd composition strategies. Evidence points to a focus on meat production in both phases, as evidenced by the predominance of adult individuals between 18–24 months old among recovered faunal remains. This age range aligns with optimal meat yield before maintenance cost increase (S1 Table). Archaeobotanical data show the predominance of cereals in both phases. Phase 6 includes hulled barley, wheat and einkorn, and Phase 5 naked barley and wheat [39].
Anthracological analyses indicate a diverse use of forest resources, with the predominance of Quercus (Evergreen Quercus and Quercus sp.) throughout the archaeological sequence. In addition, in Phase 6, remains of Juniperus sp., and in Phase 5 remains of cf. Clematis sp., Coniferae, Juniperus sp., Leguminosae, Pinus halepensis, Pinus sp. and Monocotyledoneae [54] (S2 Table) have been recovered. This diversity in Phase 5 underscores the interaction between environmental resources and farming economies, suggesting the use of surrounding forests for wood, fuel, and possible complementary food sources.
Conventional radiocarbon date along with calibrated radiocarbon ages using OxCal 4.4.4 software of the study phases of the Mas d’Is and Niuet sites (IntCal20.14c [55,56]).
Niuet site
Niuet is an open-air site located on the left bank of the Serpis River, in the valley bottom of the Alcoi Valleys (Serpis Valley), at 340 masl, within the municipality of l’Alqueria d’Asnar (Alicante) (Fig 2B). Situated on a terrace in an interfluve, it is bordered by the Serpis River on one side and the Querola ravine (Barranquet de Vargas) on the other. Archaeological works have identified three sectors: western, eastern and central. Sector A, in the central area, is the richest archaeologically, covering 41m2. Five stratigraphic levels have been documented, with the four lower levels (II to V) corresponding to the filling of a ditch with a V-shaped cross-section, measuring 5.5 m in width and 2.4 m in depth [21] (Fig 2B). All 14C dates were obtained from Sector A, placing the Niuet site sequence between the end of the 4th millennium and the beginning of the 3rd millennium cal BCE. Phase 2, including Phases N3 and N4 (levels: V, IV, III, IId, IIc), span from the end of the 4th millennium to the beginning of the 3rd millennium cal BCE (c. 3300-2900 cal BCE), while Phase 1, includes Phases N1 and N2 (levels: IIb, IIa, I, ES) and corresponds to the beginning of the 3rd millennium cal BCE (c. 2900-2600 cal BCE) [21,45] (Table 1). These chronological sequences provide a framework for examining shifts in economic practices and their implications for herd management and resource exploitation.
The economic practices at the Niuet site reflect a diversified subsistence strategy, where farming was complemented by the exploitation of forest and riparian resources. Livestock consisted of mixed and multipurpose herds, with caprines predominating across all phases analysed. During this period (Phases 1 and 2) cattle becomes increasingly more significant, eventually reaching a third of the herd in both phases. The primary production focus of livestock was meat, but the herds were likely used for multiple purposes, including dairy production and bovine strength for agricultural tasks [45]. This hypothesis is supported by the recovery of part of a pottery recipient with holes that could correspond to a cheese maker [21] and evidence of traction work, possible agricultural work, related pathologies in cattle [45]. Additionally, the analysis of sheep size has yielded new insights into the discussion of animal size reduction at the beginning of the 3rd millennium cal BCE [35,45]. Archaeobotanical data show the presence of cereals such as wheat and leguminous such as grass pea.
Anthracological evidence reflects to the extensive exploitation of forest and riparian vegetation, with Evergreen Quercus dominating throughout the Neolithic sequence, complemented by shrub taxa such as pistacia or leguminosae. Riparian vegetation, including Fraxinus oxyhylla, Salix sp., Populus nigre, Crataegus sp., and Laurus nobilis, reflect the use of nearby water-associated resources [59].
Materials and methods
Sample selection of isotopic analysis
No permits were required for the described study, which complied with all relevant regulations.
A total of 66 faunal samples from Mas d’Is and Niuet sites were selected for stable isotope analysis of carbon and nitrogen in bone collagen. To establish the local δ13C and δ15N isotopic baselines for wild and domestic mammals’ species, we selected a range of herbivores, omnivores and carnivores represented at the archaeological sites. To ensure representation from different individuals, the diaphysis of long bones from the same skeletal element and laterality were analysed for each animal species and stratigraphic level. When long bones were unavailable, mandible and phalanx were selected.
From the F4 of Mas d’Is site, 18 samples were selected: 12 from Phase 6 (5250-4850 cal BCE), and 6 from Phase 5 (4550-4250 cal BCE). The samples include domestic herbivores (Capra hircus Phase 6 n = 2; Bos taurus Phase 6 n = 8, Phase 5 n = 1; Ovis aries Phase 5 n = 1), domestic omnivores (Sus domesticus Phase 5 n = 2) and wild herbivores (Cervus elaphus Phase 6 n = 2, Phase 5 n = 2).
From the Sector A of Niuet site, 48 samples were selected. 8 samples from Phase 2 (c. 3300-2900 cal BCE), and 40 samples from Phase 1 (c. 2900-2600 cal BCE). Domestic herbivores (Capra hircus Phase 2 n = 2, Phase 1 n = 8; Ovis aries Phase 2 n = 2, Phase 1 n = 9; Bos taurus Phase 2 n = 1, Phase 1 n = 13), domestic omnivores (Sus domesticus Phase 2 n = 3, Phase 1 n = 5), wild herbivores (Cervus elaphus Phase 1 n = 2; Oryctolagus cunniculus Phase 1 n = 2), and carnivores (Canis familiaris Phase 1 n = 1) have been included in this study.
The limited number of samples from Phase 5 of Mas d’Is site (n = 6) and from the Phase 2 at Niuet site (n = 8) is due to the poor preservation of organic remains at these sites. Even so, these samples come from adult individuals of animal species and provide valuable information for the analysed periods, contributing to the isotopic baseline of the eastern Iberian Peninsula.
Collagen extraction and δ13C and δ15N stable isotope analysis
Sample selection was conducted at the University of Valencia (Spain). Collagen extraction took place at the HERCULES Laboratory at the University of Évora (Portugal). Samples (ca. 200–300 mg) were cleaned to remove contaminants using a DREMEL tool. Collagen extraction followed the modified method of Longin [60]. The bone samples were demineralized in 10 ml of 0.5 M HCl at 4°C for approximately 14 days. Afterward, the samples were cleaned with ultrapure water until neutral and immersed in 0.125 M NaOH for 20 hours at room temperature. Subsequently, they were gelatinized in 0.01 M HCl at 70°C for 48 hours. To remove impurities, the liquid fraction was filtered using Ezee‑Filter™, and the solubilized collagen was freeze-dried for 48 hours.
Collagen samples (0.3 mg) were analysed using a Thermo Flash 1112 elemental analyser (EA) coupled to a Thermo Delta V Advantage isotope ratio mass spectrometer (IRMS) with a Conflo III interface, at the Institute of Science and Technology of the Autonomous University of Barcelona (ICTA‑UAB). The average analytical error was less than 0.2‰ (1σ), as determined from duplicate analyses of δ13C and δ15N. The international laboratory standard IAEA 600 (caffeine) was used for control. Vienna PeeDee Belemnite (V‑PDB) was used for δ13C, while N2 air (AIR) served as standard for δ15N. A one-way ANOVA (α = 0.05) was used to compare δ13C values among wild and domestic herbivores and omnivores, following verification of normal distribution with the Shapiro–Wilk test (α = 0.05). The null hypothesis of normal distribution was rejected for the δ15N values, so these were analysed using the Kruskal–Wallis test (α = 0.05). All statistical analyses were conducted using PAST 4.11 [61].
δ13C and δ15N isotope analysis of bulk collagen for animal dietary reconstruction
Bulk collagen carbon (13C/12C) and nitrogen (15N/14N) stable isotopes provide valuable insights into early pastoral management strategies by characterizing the contributions of animal and plant macronutrients to animal diets [52–64]. Collagen, the primary protein in bone, reflects an individual’s diet over an extended period of time (approximately 5–15 years old) through its carbon and nitrogen isotope composition [65].
Stable carbon isotopes vary significantly among terrestrial, marine, and estuarine ecosystems, and between plants with different photosynthetic pathways (e.g., C3 and C4 plants) [66–68]. These variations are also influenced by environmental factors such as water availability, ambient temperature, and CO2 concentrations [69–72]. According to a carbon isotope fractionation of ~5‰ between whole plants and consumer’s bone collagen [62] and a correction of ~ + 1.5‰ for the fossil fuel effect in pre-industrial ecosystems [73,74], δ13C values around or above than −19‰ in herbivorous species could indicate the consumption of drought-resistant or dry vegetation, such as shrubs. These plants often have higher δ13C values due to water stress and reduced photosynthetic discrimination against 13C. On the other hand, values greater than −16‰ would suggest the consumption of C4 plants, most common in water saturated environments [72,75].
Nitrogen isotopes derive exclusively from dietary protein, with δ15N values typically indicating the trophic level of the food source due to predictable isotopic fractionations (approximately +3‰ to +6‰) throughout the food web [62,68,76]. Furthermore, increases of bulk collagen δ15N values in humans and animals in agricultural settings can reflect variations in δ15N abundance in plant-soil systems influenced by management practices, such as the use of animal fertilizers [77,78].
Results
Collagen quality
The results of stable isotope analyses of δ13C and δ15N in bulk bone collagen are summarized in Table 2.
At the Mas d’Is site, collagen was successfully extracted from 11 out of 18 samples (61%). Collagen yields ranged from 0.6 to 5%. Phase 5 has the lowest levels of collagen yields. The C% and N% values ranged from 17 to 41%, and from 6 to 14%, respectively, with C:N ratios between 3.3 to 3.6, consistent with the preserved collagen values proposed by DeNiro [79] and Van Klinken [76]. Cut-off values of 13% for C% and 4.8% for N% were applied, as recommended by Ambrose [81].
At the Niuet site, collagen was successfully extracted from 41 out of 48 samples (89%). Collagen yields varied widely, ranging from 1 to 13%, with no significant differences across species or Phases. The C% and N% values ranged from 24 to 43%, and from 8 to 15%, respectively, with C:N ratios between 3.2 to 3.5. These results are consistent with the values proposed by DeNiro [79] and Van Klinken [80], and the cut-off values of 13% for C% and 4.8% for N% were also applied, as recommended by Ambrose [81].
δ13C and δ15N values at the Mas d’Is site
At the Mas d’Is site, the average δ13C value was ‑18.7 ± 1.9‰ and the average δ15N value was 4.6 ± 1.0‰ (n = 11). Group-specific values were as follows: wild herbivores had δ13C x̄ = ‑20.4 ± 0.3‰, δ15N x̄ = 3.7 ± 0.3‰ (n = 3), domestic herbivores had δ13C x̄ = ‑17.8 ± 1.9‰, δ15N x̄ = 5.2 ± 0.8‰ (n = 7) and domestic omnivore had δ13C = ‑19.7‰, δ15N = 3.7‰ (n = 1). From Phase 5, collagen was obtained from deer (δ13C x̄ = −20.3 ± 0.1‰, δ15N x̄ = 3.7 ± 0.4‰, n = 2), pig (−19.7‰, n = 1) and cattle (−16.9‰, n = 1). In Phase 6, collagen was obtained from cattle (δ13C x̄ = −17.6 ± 2.0‰, δ15N x̄ = 5.1 ± 0.8‰, n = 5), goat (−19.9‰) and deer (−20.8‰). Cattle in Phase 6 exhibited significant variability of δ13C values, ranging from −20.5‰ to −14.7‰, documenting a −5.7‰ intra-range variability. When comparing δ15N values, goat showed the highest enrichment of δ15N among domestic herds compared to wild herbivores (2.4‰), followed by cattle in Phase 6 (1.4‰) and in Phase 5 (0.8‰). The pig shows the same δ15N value as average wild herbivores (Fig 3).
(A) Bone collagen δ13C and δ15N values compared to the mean δ13C and δ15N values ±1σ s.d for each taxon from the Mas d’Is site. (B) Box plot δ13C and δ15N values form faunal samples from the Mas d’Is site.
δ13C and δ15N values at the Niuet site
At the Niuet site, the average δ13C value was ‑19.4 ± 1.6‰ (Phase 2, n = 6; Phase 1, n = 35). Wild herbivores, represented only in Phase 1, had an average δ13C value of −20.4 ± 0.7‰ (n = 3). No significant statistical differences were found between domestic herbivores and domestic omnivores across the phases. Domestic herbivores had an average δ13C values of −19.1 ± 1.8‰ (Phase 2 = −20.4‰ to −18.6‰ n = 4; Phase 1 = −20.7‰ to −14.1‰ n = 31), while domestic omnivores averaged −20.4 ± 0.5‰ (Phase 2 = −21.0‰ to −20.2‰, n = 2; Phase 1 = −21.0‰ to −19.8‰, n = 4). In Phase 2, the intra-range specie δ13C variability was noted in goat (−0.7‰) and pig (−0.8‰). In Phase 1, cattle showed the greatest intra-range specie δ13C variability (−6.3‰) compared to goat (−1.6‰), sheep (−1.6‰) and pig (−1.2‰), with significant differences observed between cattle and sheep (р = 0.03).
The average δ15N value for the Niuet site was 4.6 ± 1.0‰ (Phase 2, n = 6; Phase 1, n = 35). Wild herbivores had an average δ15N values of 4.6 ± 0.6‰ (Phase 1, n = 3). For domestic herbivores, the average δ15N value was 5.7 ± 1.5‰ (Phase 2, n = 4; Phase 1, n = 31), and for domestic omnivores, it was 6.3 ± 1.3‰ (Phase 2, n = 2; Phase 1, n = 4). Among domesticates, there were no significant statistical differences in δ15N values. Comparing domestic herds to wild herbivores in Phase 2, pig showed the highest δ15N-enrichment (2.0‰), followed by goat (1.8‰), sheep (1.2‰) and cattle (0.6‰). In Phase 1, pig and sheep showed the highest δ15N-enrichment (1.6‰), followed by cattle (1.3‰), and goats (0.2‰), with cattle reaching a peak δ15N value of 10.9‰. Among caprines, in Phase 2 the δ15N intra-range variability is 1.1‰, specifically 1‰ among goats, there is no data on sheep because there is only one individual analysed. And in Phase 1 the δ15N intra-range variability is 4.3‰, specifically 4.35‰ among sheep and 1.8‰ among goats (Fig 4).
(A) Bone collagen δ13C and δ15N values compared to the mean δ13C and δ15N values ±1σ s.d for each taxon from the Niuet site. (B) Box plot δ13C and δ15N values form faunal samples from the Niuet site.
Discussion
Pig, caprine and cattle herd management strategies in the Serpis Valley
The isotopic variability in δ13C and δ15N values from Mas d’Is and Niuet sites suggests that herds had access to a diverse range of plant resources, reflecting specific feeding strategies employed by Neolithic communities in the Serpis Valley.
In the study area, C3 plants are predominantly expected, with δ13C variations up to 4‰ throughout the seasonal cycle, peaking in summer and dipping in winter depending on water stress [70,72,82–84]. The δ13C values from the faunal remains of Mas d’Is and Niuet sites suggest that majority of the protein in these animals was obtained from a terrestrial food web based either on C3 or C4 plants. C3 values would be characterized by plants from open areas, while the higher δ13C values observed in cattle in Mas d’Is site (−14.7‰) and Niuet site (−14.1‰, −14.3‰ and −14.4‰) indicate the presence of environments dominated by C4 plants [85]. The selective targeting of grazing areas, particularly those adapted to wetland environments, may be the cause of high δ13C values [86]. These aquatic or semi-aquatic C4 species can thrive in environments with variable moisture levels, contributing to the overall plant diversity available to grazing animals [87]. The presence of such plants in the diets of cattle could further explain the elevated δ13C values. In the Serpis Valley, the Cyperaceae family, which includes plants located in wetland and marshes contexts like Juncus sp., Carex sp., Scirpus sp. and Schoenoplectus sp., could contribute to the C4 signal. Cyperaceae plants are adapted to saturated soils and flooded conditions, and while they are not grasses, they play a crucial role in wet ecosystems, and provide important habitats for various species of fauna. These Cyperaceous plants might have been present during the Neolithic along the Serpis River and more abundant in wetland areas like the Gaianes Lagoon, and their recurrent consumption could be reflected in the C4 signal observed in the δ13C values.
Documented evidence of C4 plants in the Iberian Peninsula during the Neolithic is scarce, with evidence primarily found at archaeological sites such as Costamar (Castellón, Spain) and La Vital (Gandia, Spain), both in eastern Iberia [88,89].
Pig management.
Pigs show δ13C values from open C3 environments at both the Mas d’Is and Niuet sites. At the Mas d’Is site, the δ15N value of the single individual analysed is similar to that of wild herbivores, suggesting an herbivorous diet without agricultural surplus contributions. At the Niuet site, pigs exhibit the least variability in δ13C values among the analysed herds, suggesting access to a narrower range of plant resources. The differences in δ15N values are observed between pigs from Phase 2 and Phase 1. In Phase 2, a greater enrichment of δ15N values is observed, possibly indicating a higher intake of protein in the pigs’ diet. This may suggest that pigs had access to protein sources not directly related to plant consumption, such as animal by-products or agricultural by-products. In contrast, pigs from Phase 1 show lower δ15N values, similar to wild herbivores, except for one individual displaying significantly enriched δ15N values (8.5‰), which could be indicative of a distinct dietary strategy or an unusual feeding pattern. The variability in δ15N values could reflect differences in the amount of animal protein in pigs’ diet [90] Pigs, being omnivorous, could have supplemented their plant-based diet with various protein sources, such as worms, insects, small animals, or human food waste. The lower δ15N values, closer to those of wild herbivores, suggest a diet richer in plant products, possibly indicating that pigs were managed in extensive systems in semi-free-range conditions. In contrast, the higher δ15N values in some pigs could be related to an increased intake of protein, possibly from waste disposal sites or access to agricultural by-products [2,6,91]. This interpretation has also been proposed for the Pyrenees [7,11], the Mediterranean areas [6,9] and the Atlantic areas of the Iberian Peninsula [8]. This dietary flexibility would have allowed pigs to take advantage of various food resources in human-dominated landscapes, where waste from daily activities, including animal processing and crop production, provided an accessible source of nutrition. Such enrichment in δ15N values may reflect the early stages of a more intensive pig-rearing strategy, where pigs were increasingly integrated into the livestock economy through the use of human-generated food surpluses and proximity to settlements, perhaps even stabled.
Caprine management.
The variation in δ13C values among the caprine herds at Mas d’Is and Niuet sites suggests a diet of open C3 plants, typical of environments dominated by trees and scrubs. This is a common dietary pattern for animals grazing in open areas, where C3 plants predominate. At Mas d’Is site, the isotopic analysis of a single goat revealed 15N-enriched values (2.4‰) compared to wild animals, which could suggest that this individual had access to fields or was provided with surplus crops as forage. At Niuet site, caprines from chronological Phases 2 and 1 show very similar δ13C isotopic signatures (difference δ13C 0,5‰ between phases), suggesting a relatively stable environment across the 4th millennium and the beginning of the 3rd millennium cal BCE, as also reflected in the anthracological analyses of the site [60]. δ15N values show high variability among caprines, mainly in Phase 1. In Phase 2, goats show more δ15N enriched values than sheep, although the intra-range variability amplitude is 1.1‰. In contrast, in Phase 1, goats show values similar to wild herbivores, unlike sheep, which show an intra-range variability of 4.3‰, and with an 15N-enrichment of 0,7‰ over goats. The δ15N values observed in the goats of Phase 1, point to grazing in areas with minimal human impact, such as soils not used for agriculture or arid areas. Is possible that goats were managed in environments outside cultivated areas or areas less suitable for intensive farming. In addition, the difference with some of the sheep’s diets could be affected by the type of diet of goats, which are browsers, also feeding on branches and bushes. Sheep, in Phase 1, in general, show higher δ15N values than goats. This enrichment could correspond to a more selective feeding, based on different grazing areas, access to fields or on the supply of agricultural surplus. The elevated δ15N values in some sheep samples could also be linked to the physiological needs of neonatal and infantile individuals under 3 months of age, as feeding practices are crucial for the nutritional well-being of the mother, which is essential for the survival of lambs and to ensure sufficient milk production [92]. In Phase 1, the recovery of a cheese maker fragment, a ceramic vessel with small perforations in the walls, provides additional evidence to support the use of milk and the consumption of dairy products at the Niuet site [21]. The caprines feeding practices at Mas d’Is and Niuet sites indicate early forms of controlled herding and the integration of livestock into agricultural activities and environmental management.
Cattle management.
Cattle exhibit significant variability in δ13C values at both sites, suggesting that different individuals consumed plants with different isotopic compositions. This variation may reflect differing animal management practices within the same archaeological site and concerning the same animal species. The δ13C values for the analysed specimens range from −20.5‰ to −16.9‰ and −14.7‰ at Mas d’Is site, and from −20.4‰ to −17.4‰ and from −14.4‰ to −14.1‰ at Niuet site. These results suggest that while some cattle consumed C3 plants, others consumed C4 plants. This feeding strategy appears to have been consistently applied at both sites, regardless of their location and chronology, reflecting a continuous compartmentalized management approach of the landscape throughout the Neolithic sequence in the Serpis Valley. Regarding the δ15N values, cattle from Mas d’Is site show a 15N-enriched by 1.4‰ in Phase 6, and 0.8 ‰ in Phase 5, and Niuet site show a 15N-enriched by 0.6‰ in Phase 2, and 1.3‰ in Phase 1, compared to wild herbivores. The diachronic analysis of the Mas d’Is and Niuet sites shows differences in δ15N values, which could reflect a change in the intensity of cattle management over time. At Mas d’Is site, cattle exhibit δ15N values reaching 6.2‰ with a variability range of 2.1‰. In contrast, at Niuet site, δ15N values reached 10.9‰, with a variability range of 7.5‰. This 15N-enrichment may suggest that some cattle consumed 15N-enriched plants, possibly due to fertilization with manure. However, the impact of animal dung on plant δ15N values is highly variable, depending on factors such as manuring frequency and type of fertilizer used [77,78,93]. High δ15N values could indicate that cattle had access to grazing in manured fields or consumption of agricultural surpluses, such as crop stubble. These practices have been documented in various Neolithic contexts across Europe [93,94] and the Iberian Peninsula [6–9]. Additionally, pathologies documented in the distal extremities at the Niuet site, associated with repetitive mechanical activities like animal traction, could suggest that some cattle were used in agricultural or traction tasks, providing them access to crop fields [45]. The difference in δ15N values between the Mas d’Is and Niuet sites, and the documented cattle bone pathologies in Niuet site, suggest a progressive intensification of cattle management over time. It should be noted that the individuals with C4 isotopic signatures do not show 15N-enriched in Mas d’Is or Niuet sites.
The isotopic signatures from this study suggests a bimodal distribution of the cattle herds over the landscape. On the one hand we have some cattle that had access to pastures in marshes or along riverbanks, such as the Gaianes Lagoon area, where they could consume plants with a C4 isotopic signature. Floodplain pastures offer an abundance of fresh vegetation for much of the year, allowing cattle to optimize the available natural resources. Additionally, cattle can endure prolonged periods in flooded areas due to various physiological adaptations, such as thick skin on their limbs that protects against moisture and infections, and strong hooves that enable them to withstand wet and muddy areas [95]. This access to flood zones by cattle grazing has been documented in several historical and contemporary areas such as the Camargue in France [96], Eastern Europe [97], South Sudan [98], or Australia [99] amongst many others. In the Iberian Peninsula, access by cattle to flooded sites has not yet been documented. Other isotopic data from Neolithic sites available that show a similar pattern of differential C3 and C4 isotopic signatures in cattle belong to the sites of La Vital, located at the mouth of the Serpis River and surrounded by marshes, and the site of Costamar, located on the coastal plain and also surrounded by marshes [88,89]. These results may have indicated cattle access to flood zones as well. The low δ15N values would reinforce this hypothesis, since they are associated with extensive husbandry practices, or even with wild, feral or semidomesticated animals. Regarding the status of animals, biometric analyses from Mas d’Is and Niuet have characterized individuals in their domestic form, but, with the data available we are unable to differentiate feral from hybrid animals [45].
Meanwhile, on the other hand, the rest of the cattle at Mas d’Is and Niuet sites grazed in pasture areas dominated by C3 plants, with similar grazing patterns than pig and caprine in both sites. The 15N-enrichment observed in some of these cattle could indicate varying levels of management intensity, with high δ15N values correlating with intensification of agricultural practices. Studies by García Atiénzar [36], Bernabeu Aubán [100], McClure et al. [38] have indicated that the Neolithic settlements in the Serpis Valley were strategically located in fertile valley bottoms surrounded by rich agricultural soils. These locations suggest the development of intensive agriculture with a hoe in fields close to the inhabited centres [36–38]. Therefore, at Mas d’Is and Niuet sites, specimens with 15N-enrichment C3 might have been household managed, relying on the fields nearby the settlements for access to resources or agricultural by-products.
Husbandry practices in an environmental and socioeconomic context
Isotopic analyses of the Mas d’Is and Niuet sites reveal different strategies in animal management and landscape use strategies during the Neolithic in the Serpis Valley. The results suggest compartmentalized land use, with complementary exploitation of agricultural zones, grazing areas, and lacustrine ecosystems, in response to environmental conditions and socioeconomic transformations.
By the mid-6th millennium cal. BCE, the Serpis Valley saw effective occupation, characterized by valley bottom settlements such as the Mas d’Is site, alongside cave and shelter occupations [36–38]. According to García Atiénzar [36], low demographic density characterized these communities, with concentration of open-air settlements in particular points of the territory and focusing on small-scale cultivation and livestock management. The riverine areas, together with marshes and lakes of the valley, may have played a crucial role in these systems, offering available grazing areas for livestock and influencing settlement and mobility patterns. Furthermore, the potential for communal exploitation of these natural resources, alongside other surrounding environments, cannot be overlooked [36]. By the 4th millennium cal BCE, the process of intensification in the Serpis Valley continued, marked by new settlements in previously unoccupied areas, such as Niuet site. This change marked a transition to an aggregated settlement system with a reduction in cave-pens [40], surrounded by fields of crops and delimited by ditch [36], and the presence of negative structures, interpreted as fodder storage [44,100]. By the end of the 4th millennium cal. BCE, a significant shift occurred, marked by a reduction in the variety of cultivated cereals, indicating a trend towards relative specialization in agricultural practices [40]. Additionally, the intensification of cattle management may be associated with the occupation of the valley bottoms, where favourable conditions such as expansive plains, access to water, and abundant high-quality herbaceous grasses provided ideal grazing environments for herds. These areas, particularly along river and lake valleys, provided abundant grazing opportunities, facilitating the expansion of cattle breeding and its integration into the broader economic system [45]. Isotopic analyses of cattle, specially δ15N-enrichment, have revealed an intensification of management practices, further corroborated by the bone pathologies observed at the Niuet site.
The evolution and intensification of farming practices in the Serpis Valley reflect the adaptability of Neolithic communities to environmental conditions, such as topography, the availability of key resources like water, and the socioeconomic changes [101,102]. There are several historical examples of small agrarian communities in the Middle Ages and early modern period that often divided their land into two blocks: areas close to the villages, easier to work and control, because they are nearby and tend to be the flat and more fertile, tend to become the exclusive property of the family that works each plot (what capitalism has termed private property) [103–106]. Mountain, hilly or marshy areas, hard to access or to defend against interlopers, but with key resources for specific moments of the year (i.e., summer pastures), have often been managed by the entire community as commons [107]. This dynamic reflects a complex interaction between the needs of the community and households, environmental characteristics, and management capabilities, and in some ways, could coevolve through mutually beneficial interactions [99,108].
In the context of Neolithic pastoralism, property theory can offer a valuable framework for understanding territorial management practices. This theory emphasizes two key aspects: a) the characteristics of the natural resources (including their productive cycle, influenced by seasonality, topography, and water availability), and b) the characteristics of the managerial institutions developed by human communities to control these resources, influenced by human population density and competition with other groups [109]. In this framework, property is defined by the capacity to include or exclude users from access to specific resources [108]. Private property implies that an individual or household has the capacity to control and enforce a claim over a specific resource. Common property implies that a group of families exert such a claim. Open access signifies, nobody is interested or has been able to establish control over something, and that everybody has access to a given resource [110,111]. In the Serpis Valley, as isotopic results suggest, marshes or riverine areas may have played a key role in land management during the Neolithic. Marshes, characterized by difficult mobility, potential zoonosis, and/or frequent flooding, have historically been challenging environments for human settlement, making them poor candidates for the development of private property [112,113]. These complexities often led to their management as common property or their transformation into open-access resources [114,115].
Within this context, isotopic data from the Mas d’Is and Niuet sites indicate that such marginal areas also contributed significantly to husbandry practices. These marshes or riverine areas would have provided fresh fodder for much of the year and could support animals that were primarily exploited for meat in the case of cattle. Meanwhile, agricultural areas near settlements offered additional grazing opportunities, benefiting those individuals who were used for traction work. These specimens could be associated with those with C3 signals and higher 15N values. The ethnography of pastoralism identifies, plenty of cases in which animals circulate through all the areas of a given landscape and marshes are used, depending on the latitude, as winter or dry season pastures [116,117]. In other words, the resources are browsed considering the seasonal productivity of each ecological patch and niche. In this situation, however, the C4 signature is so high that the intermittent use of pastures in marshes and dryland, seems unlikely. The exclusive presence of nitrogen in the C3 animals works as well, against this possibility. To fully discard seasonal alternance between C3 and C4 areas a carbon and oxygen isotopic analysis should be implemented. This dual approach to animal management—combining a settlement-based individuals engaged in agricultural work with a more isolated individuals grazing in marshlands—reflects the emergence of a complex system of resource use. By the 4th millennium cal BCE, archaeozoological data suggest an intensification of livestock specialization, with increased production and the accumulation of surpluses, underscoring the growing economic importance of livestock, specially cattle management, and the specialization of these practices.
In addition to the direct use of marshlands or riverine areas, social and economic factors likely shaped how different groups within the Neolithic communities accessed and managed land [101]. Isotopic results reveal two different cattle feeding patterns at each site analysed, suggesting that socioeconomic implications could have influenced access to different grazing areas. Ethnographic examples suggest that differential access to marginal areas could be related to identity markers such as economic status, kinship, age, gender, or ethnicity [118–120].
Alternatively, comparative ethnographic and archaeological analysis suggest that in some inhospitable areas, escaped domestic animals may become feral and adapt to suboptimal environments [121,122]. Neighbouring communities, not necessarily involved in the management of these relatively dangerous environments might not interfere, besides the occasional capture when needed, with this rewilding process. In some cases, these feral animals have become intrinsic parts of the functionality of these environments [123,124] and become an additional source of meat when needed [125]. The C4 cattle individuals could indicate the presence of these semi-wild herds, with little interaction with nitrogen-rich or C3 plants. However, as abovementioned, only the domesticated status has been documented.
The diachronic evolution of land management strategies in the Serpis Valley highlights a gradual shift from small-scale, localized agricultural practices to more complex and integrated systems. The integration of marshes and other difficult-to-access landscapes into the broader land management system illustrates a sophisticated approach to resource use, where communal property regimes and seasonal variations in resource availability played a crucial role. From the 6th to the 3rd millennium cal BCE, the growing complexity of settlement patterns, resource use, and livestock management reflects the increasing integration of agriculture and pastoralism. By the 4th millennium cal BCE, as population densities grew and economic practices intensified, driven by both environmental pressures and social changes, underscores the resilience and adaptability of Neolithic communities in the Serpis Valley, who shaped their landscapes and economies to meet the demands of both subsistence and surplus production, ensuring the sustainability of their agricultural and pastoral practices.
Conclusion
The variability in δ13C and δ15N values at the Mas d’Is and Niuet sites reveals a complex interplay between environmental conditions, feeding strategies, and animal management practices during the Neolithic in the Serpis Valley. The diverse δ13C and δ15N values indicate that cattle had access to a variety of spatially segregated plant resources, suggesting a nuanced approach to livestock management. Notably, the differential presence of C3 and C4 plants in the diets of these animals reflects the availability of different grazing environments, including marshlands, which may have been managed as communal or open access resources. The high nitrogen values in some cattle suggest access to fertilized pastures or agricultural by-products, highlighting the integration of animal husbandry with early agricultural practices. The fact that these higher nitrogen values coincide with C3 specimens seems to provide further evidence of a two-prone approach to cattle breeding: a) a settlement-based herd, part of a process of a relative agricultural intensification, grazing on the fields and pastures nearby the settlements and involved in the agricultural works, and b) a more isolated herd, mostly located and feeding in marshland and, apparently not involved with the larger agricultural system developing in the area. Furthermore, variations in the diets of caprines and pigs point to differing management strategies, with implications for social dynamics and economic organization within these Neolithic communities. The isotopic evidence extracted from the zooarchaeological remains combined with a study of their morphological incidences, supports the idea of a diversified landscape management system in the Serpis Valley, where resource availability and social structure influenced grazing practices and herd use. Furthermore, between the 6th and the beginning of the 5th millennium BCE, there was a significant shift in the strategies for occupying and exploiting territory, which may imply a new approach to animal management. The data become increasingly complete in the 4th and 3rd millennia cal BCE, revealing a certain cattle specialization and a substantial rise in production and surplus accumulated by the household units. This consolidation of husbandry practices in the Neolithic underscores the dynamic relationship between communities and their environments, reflecting an adaptive response to changing economic and social conditions.
Supporting information
S1 Table. Taxonomic representation of animal remains recovered in Phase 5 and Phase 6 of the Mas d’Is site (Penàguila, Alicante).
NISP = Number of Identified Specimens; MNI = Minimum Number of Individuals.
https://doi.org/10.1371/journal.pone.0325137.s001
(DOCX)
S2 Table. Taxonomic representation of charcoals remains in Phase 5 and Phase 6 of the Mas d’Is site (Penàguila, Alicante).
https://doi.org/10.1371/journal.pone.0325137.s002
(DOCX)
Acknowledgments
The authors thank the anonymous reviewers for their valuable comments on this article. The authors thank Dr. Pau Comes (ICTA-UAB) for his help with the IRMS data and Dr. Anne-France Maurer and Dr. Ana Curto (HERCULES Lab) for her help with collagen extraction.
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