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In foreign pastures: Livestock mobility at Hazor and the permeability of Iron Age geopolitical borders in the Southern Levant

  • Cheryl A. Makarewicz ,

    Roles Conceptualization, Data curation, Formal analysis, Funding acquisition, Investigation, Methodology, Project administration, Resources, Visualization, Writing – original draft, Writing – review & editing

    c.makarewicz@ufg.uni-kiel.de

    Affiliations Department of Archaeological Sciences, School of Archaeology and Maritime Cultures, University of Haifa, Israel, Archaeology Stable Isotope Laboratory, Institute for Prehistoric and Protohistoric Archaeology, University of Kiel, Germany

  • Sarah Martini,

    Roles Formal analysis, Investigation, Visualization, Writing – original draft, Writing – review & editing

    Affiliation Department of Anthropology, Yale University, New Haven, Connecticut, United States of America

  • Nimrod Marom,

    Roles Writing – original draft, Writing – review & editing

    Affiliations Department of Archaeological Sciences, School of Archaeology and Maritime Cultures, University of Haifa, Israel, The Leon Recanati Institute for Maritime Studies, School of Archaeology and Maritime Cultures, University of Haifa, Israel, Interdisciplinary Centre for Archaeology and Evolution of Human Behavior (ICArEHB), University of Algarve, Portugal

  • Shlomit Bechar

    Roles Conceptualization, Funding acquisition, Writing – original draft, Writing – review & editing

    Affiliations The Leon Recanati Institute for Maritime Studies, School of Archaeology and Maritime Cultures, University of Haifa, Israel, Department of Archaeology, School of Archaeology and Maritime Cultures, University of Haifa, Israel

Abstract

Southern Levantine societies during the Iron Age II (10th–8th centuries BCE) witnessed the formation of competing territorial polities and urban revival following a period of settlement ruralization and dwindling regional exchange economies associated with the collapse of Late Bronze Age Canaanite city-states around 1200 BCE. Recent work has revealed diverse expressions of complex political organization, state-sponsored cultic activity, and inter-polity conflicts within the region during the Iron Age II, but the impact of regular military confrontations and ensuing territorial reconfiguration on agro-pastoralist economic systems that supported these polities is unknown. Here we explore inter-polity border dynamics between the Israelite and Aramean states during periods of conflict (Iron Age IIA) and concord (Iron Age IIB) by establishing landscape-use strategies involving the most mobile element of Iron Age subsistence and production systems – domesticated sheep and goats, at Tel Hazor located in the Hula Valley, a place of direct contact between Aram and ancient Israel. Multi-stable isotopic analyses of bovid livestock teeth indicate agro-pastoralist herders grazed their animals in well-watered pastures locally near Hazor and also further afield in the Golan Heights. The continuous use of spatially diverse pasturing regimes throughout the Iron Age II suggests household-based agro-pastoralist land use transcended regional political discord. Everyday movement of herders and their flocks to distant pastures was not restricted despite conflict between the military and ruling elites of the Israelite and Aramean states, suggesting that borderlands between states were permeable.

Introduction

Ancient borders and borderlands shaped how states enacted hegemonic power internally and externally, engaged in military actions, participated in exchange networks, and directed their own subsistence production. Such demarcated places may have included spatially diffuse areas claimed by competing states as their own territory, neutral buffer zones between two states, zones of intersection between hinterlands linked to different, cooperating or competing urban cores, or hard boundaries demarcated by physical barriers− either human built or distinctive topographical features, that impeded cross-polity movement and facilitated the exercise of political and military power over the landscape (e.g., the Roman limes) [13]. Despite the role of borders in shaping cross-polity interaction at various scales between households, communities, and states, it remains to be seen the impact, if any, of state borderlands beyond political and military activities. Much attention has been paid to how borders, frontiers, and borderlands defined and influenced political interaction, trade relations, alterity, and the movement of people and goods [1,46], but less so on the subsistence activities carried out by communities and households. There is similarly scant information on how ancient borders and borderlands restricted or encouraged agro-pastoralist production that supported the state apparatus.

In addition to revealing different modes of inter-state relations, isolating mobility dynamics potentially informs on the degree of vertical integration within ancient states. Curtailed movement of people, goods, and activities associated with a particular economic segment or production cascade, for example, would create different social, economic, and political outcomes than controlled movement of elites who exercised authority over a local urban political apparatus (Fig 1). One such group of non-elite individuals whose mobility patterns could be influenced by different border dynamics are herders, whose livelihood depends on moving their livestock to plentiful pastures.

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Fig 1. Schematic illustration of relationships between communities belonging to conflicting polities, contrasting a permeable border model (top) and non-permeable border model (bottom).

(+)/(−) represent cooperative/hostile ideological connections.

https://doi.org/10.1371/journal.pone.0328934.g001

Under conditions of a permeable border, conflict encompasses political violence between established power groups (i.e., ‘elites’), with no percolation of the concept of a ‘dangerous other’ to people distant from ruling and military circles. Livestock and agricultural goods freely moved across borderlands, stopped intermittently by active warfare every few decades. In such circumstances, access to pastoralist infrastructure including seasonal pastures and water sources was conducted independently of the state, regulated instead through inter-community negotiations and traditional tribal arrangements, cross-cutting national boundaries.

When a non-permeable border is in place, quotidian every-day socio-economic interactions across borderlands are strongly curtailed across all sectors of society. Furthermore, non-permeable borders may imply the existence of a vertically integrated territorial society in which conflictual ideologies filter down to the agrarian hinterland to such an extent that much of the flow of goods between hostile territories stops. Bound to ideologies legitimizing the indigenous state, as much as they might be constrained by physical conflict, agro-pastoralists no longer enter subsistence goods and capital into regional exchange networks. Accordingly, access to pastoral resources that are distributed along geographical and environmental clines is curtailed when a non-permeable border is maintained, with state-level political dynamics superseding traditional pasture access agreements. A shift from a permeable to non-permeable borderland could result, for example, in a zooarchaeologically visible shift from mobile sheep pastoralism, which requires transhumant movement between high-quality pastures and predictable access to water, to goat husbandry that does not require high-quality forage and, thus, can be more readily conducted in a circumscribed territory.

The Iron Age II (IAII) of the southern Levant (10th–8th centuries BCE) provides an opportunity to better understand the articulations between ancient border dynamics, landscape use, and livestock pasturing strategies. The Iron Age II saw the emergence and subsequent consolidation of multiple territorial polities (Fig 2), each of which exhibited an unusually strong internal cultural cohesion and ascriptive identity, several hundred years after the collapse of Late Bronze Age Canaanite city-states and palace economies that previously controlled the southern Levant [7]. These various polities, separately governed by the Phoenicians, Neo-Hittites, Philistine, Israelites, and Arameans, appear to have emplaced borders, alluded to in historical texts and thus assumed in archaeological models describing Iron Age II political organization. While biblical narratives and other textual evidence tell of endemic warfare between these Iron Age II polities in the centuries between the fall of the Egyptian empire in the 12th century BCE and the occupation by the Neo-Assyrian Empire in the second half of the 8th century BCE [810], there is no mention in these various records of outright borders or borderlands where political claims on a landscape may have been reduced, negotiable, or relatively unimportant.

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Fig 2. Location of Iron Age II polities in the southern Levant (following ref [11]).

Estimated geographic extent of polities discussed in this text indicated by color shading: Israel = red, Aram = light gray. OpenStreetMap contributors. 2015. Planet dump [SRTM30 Colored Hillshade and Topographic WMS by Terrestris from 2022]. Retrieved from https://planet.openstreetmap.org. The lake vector was made with Natural Earth [ne_10m_lakes]. Free vector and raster map data are available at naturalearthdata.com.

https://doi.org/10.1371/journal.pone.0328934.g002

On-going conflict between the kingdoms of Aram and Israel during the early first millennium BCE shaped regional exchange networks and subsistence production alike. We examine borderland dynamics between these two states through investigation of landscape-use strategies engaged by agro-pastoralist herders, focusing on the diversity and spatial extent of livestock pasturing regimes in contested landscapes further dissected by political borders. Focusing on domesticated sheep and goats that provided households and cities with both subsistence and raw materials that could be transformed into commodities (i.e., wool), we analyzed the strontium (86Sr/87Sr), carbon (δ13C), and oxygen (δ18O) isotopes of incrementally sampled caprine teeth in order to establish the relative extent of mobile herding activities and attendant pasturing strategies, taking advantage of predictable geospatial distribution in bioavailable strontium isotopes, oxygen isotopes in meteoric water, and also carbon isotopes at the floral base of the foodweb across the varied environments of the southern Levant. We specifically investigate if pasturing regimes in the Hula Basin (upper Galilee region) were geospatially concentrated around Israelite urban centers during the 10th–9th centuries BCE (IRIIA) when conflict between the Israelite and Aramean states peaked and was followed by more extensive pasture access during first half of the 8th century BCE (IRIIB) when conflict subsided and the hinterlands were controlled by the Israelites.

We use faunal remains recovered from Tel Hazor, located in the contested Israelite-Aramean border area in the Mediterranean landscapes of the Upper Galilee region, to empirically test two contrasting models for early inter-polity boundaries as they existed during the 10th–8th centuries near Hazor. Under the non-permeable border model, conflict around Hazor in the 10th-9th century BCE (IRIIA) would result in limited access to pastures over a relatively small catchment area suggestive of established borders that served as a bulwark against the movement of people and goods between the Israelite and Aramean states thus hindering movement of livestock across the landscape. Under a permeable border model, livestock mobility would be similar across the early first millennium BCE, suggesting that borders were relatively porous during Aramean and Israelite conflict and also during later Israelite political consolidation. In this case, animals and agricultural goods liberally moved across territories, suggesting that warfare between kingdoms did not have significant effect on household and communities distant from political and military centers. The alleviation of conflict around Hazor in the 8th century BCE would result in expanded access to pastures over a larger catchment area, thus stimulating livestock mobility in order take advantage of more varied and higher-quality pastures.

Tel Hazor in the Iron Age

The borderlands of Israelite Hazor changed hands between Aram and Israel throughout the 10th – 8th BCE centuries. The border between these two states was most probably located around the Yarmouk River, but the Hula Valley, as the area of direct contact between Aram and Israel, was likely also a contentious borderland zone [12]; the upper Jordan Valley was likely part of the Israelite kingdom [10,13]. The Israelite city of Hazor, built as a fortified urban center in the 10th century BCE, functioned both as a border fortification against the Aramean state and central settlement that supported extensive domestic quarters and some administrative buildings [14,15]. Conflict erupted between both kingdoms at the end of the 10th century BCE (IRIIA), after Israel separated from Judea, when the Aramaeans attacked Israel on behalf of Judea (1 Kings, 15:18–20). Although it is unclear how long the Aramaean occupation over the Israelite territories lasted, the establishment of the Omride dynasty in 881 BCE consolidated power in the Israelite state. The Omrides held the highlands of Samaria, the lower and upper Galilee, and a small portion of the Transjordan Plateau through a network of newly constructed palatial architecture placed in various cities; these buildings served to bring together diverse groups while simultaneously broadcasting their own political authority [16].

The new political authority in ancient Israel established in 842 BCE quickly succumbed to Assyrian attackers in 841/840 BCE who also engaged the Aramaean kingdom. The cessation of military activity by the Assyrians around 836 BCE, who then focused their efforts on western campaigns, opened an opportunity for the re-opening of direct conflict by the Aramaeans against the Israelites. The Tel Dan stele, for example, indicates the Aramaean king Hazael conquered the entire northern Jordan Valley [10,13]. At some point during the 9th century BCE, Hazor was transformed into a large administrative center, having doubled in size and domestic buildings replaced by public buildings, which included at least five large storage buildings and two granaries thought to have supplied both the city and residents beyond. Substantial investment in a subterranean water system also took place at this time, providing the city with the means to withstand a long-term siege [14,15,17]. The rapid growth of Hazor might indicate a demographic bulwark established by the Israelite kingdom along the Aramean border in response to heightened political unrest between the two entities or, alternatively, reflect a general increase in the population suggested by shifts in regional settlement patterns (see below, section 2.2.2). The subsequent devastation of the Aramean state by the Assyrians around 803/796 BCE led to a calm period for the Israelite state, as the Arameans were no longer able to exercise autonomous political power in the region [8].

Corresponding with the decline in conflict with the Aramean state, the built environment of Hazor underwent significant changes in the 8th century BCE (IRIIB) when several of the public buildings, including all the granaries and storage buildings, went out of use and were replaced by new domestic buildings and workshops. Storage facilities were identified in some domestic buildings, indicating storage was no longer conducted in public buildings but instead performed within households. Such a change in storage mechanisms at Hazor may indicate a broader change in subsistence modes, probably agricultural activities, engaged by the local inhabitants of the city. Alternatively, this may reflect a shift in the acquisition and (re)distribution of food commodities that coincided with a change in local political structures, a process suggested by the dissolution of administrative buildings and their replacement by domestic buildings and workshops. These changes in architectural construction and use within Hazor were accompanied by considerable settlement shifts in its hinterland regions. The Upper Galilee, the northern Golan Heights and the northern part of the Hula Valley experienced a sharp decrease in the number of settlements during the transition from the 9th century (IRIIA) to the 8th century (IRIIB) BCE [18,19]. Settlement abandonment was particularly pronounced in the mountainous regions of the Galilee, which may have served as a buffer zone between the kingdom of Israel and the Phoenician coast [18]. In contrast, the southern part of the Hula Valley and the central and southern Golan Heights experienced a dramatic increase in settlement density, perhaps reflecting the growth of Hazor [19,20], now wealthier and stronger. More settlements sprung up around the city, hoping to take advantage of economic opportunities that created trade and wealth. It may also be that northern communities relocated from potentially volatile and contested terrains to the more politically stable area surrounding Hazor, which was still under Israelite control, and, by relocating to this area, they were able to seek protection from Hazor.

The shifting political relations between Aram and Israel certainly affected trade and economic relations between and within each kingdom, but the extent to which political conflict, engaged largely by the ruling elite and military circles, was projected onto the hinterland landscapes remains largely unknown. How the economic practices of domestic households, including agro-pastoralist herders who husbanded livestock for their own subsistence and profit, were affected is not documented, nor are shifts in extensive pasturing systems strategies most affected by borderland political dynamics. Hazor’s positioning in the contested landscapes proximal to the Hula Valley makes the settlement ideally suited to examine the permeability dynamics of the Israelite−Aramean border throughout the Iron Age, when political establishments were repeatedly reconfigured by indigenous and foreign influences.

Materials and methods

Isotopic foundations for establishing ancient pasturing regimes

Where herders direct their livestock for pasturing depends on complex intersection of factors, including seasonally driven changes in pasture graze quality and abundance, location of pastoralist infrastructure such as water and shelter [21], pasture access rights [22], and directives from institutionalized authorities [23]. Multi-stable isotopic analyses of ancient animal skeletal tissues provide a robust means to trace livestock dietary change and mobility at high temporal and geospatial resolutions, information that further supplies new insights into the spatial dynamics of pastoral landscape use [24].

Carbon (δ13C) isotopes

The carbon isotope ratios of herbivore tooth bioapatite reflect those of ingested plants, providing a means to directly document seasonality in the carbon isotopic composition of local vegetation communities and thus enabling reconstruction of fodder provisioning and livestock pasturing practices associated with mobile herding regimes [25,26]. The carbon isotope ratios of plants are determined by photosynthetic pathway, aridity, and water availability [27]. C3 and C4 plants globally average −26‰ and −12‰ in δ13C, respectively [28]. C3 plants are sensitive to water stress, closing their stomata under conditions of reduced water availability. This physiological response decreases photosynthesis, leading to a decrease in carbon fixation from CO2, which in turn enriches leaf tissue growth approximately 2 − 5‰ in 13C [29,30]. The carbon isotopic composition of semi-arid environment floral growth shifts seasonally with warmer season growth enriched in 13C relative to cooler wet seasonal growth [31,32]. C3 plants are also sensitive to arboreal cover density. The combination of decomposing leaf litter, recycling of atmospheric CO2, and/or low light levels that slow photosynthesis rates creates a ‘canopy effect’ that can lower understory floral δ13C values up to 5‰ [33,34]. We apply a enamel bioapatite–diet isotope enrichment factor of 14.1 ± 0.5‰ to estimate dietary δ13C values for bovid livestock [35].

Oxygen (δ18O) isotopes

The geospatial distribution of environmental oxygen isotopes is broadly predictable across landscapes, thereby providing a means to examine animal mobility. The oxygen isotopic composition of precipitation is influenced by rainfall amount, temperature, and humidity [36,37]. In semi-arid environments, seasonal variation in temperature, precipitation levels, and aridity produces corresponding seasonal shifts in precipitation oxygen isotope values, with warmer season meteoric waters typically enriched in 18O due to higher temperatures, increased aridity, and reduced rainfall [36]. Precipitation δ18O values are also influenced by altitude with 18O-depleted meteoric waters falling on cooler, high elevation locations less impacted by evaporative fractionation [36]. Meteoric water δ18O values decrease approximately 0.3‰ per 100m increase in elevation [38,39].

Leaf water oxygen isotopes reflect in part the oxygen isotopic composition of meteoric waters [40,41], but are strongly influenced by soil water δ18O values, rooting depth, leaf structure, and evapotranspiration rates [42]. Evaporation at the ground-air interface results in 18O enrichment of upper layer soil waters relative to meteoric water [43]. Fractionation effects are minimal during transport of soil water via non-transpiring plant parts such as roots and stems [31,44]. Substantial 18O enrichment of plant water does occur, however, at evaporation sites in leaves [45,46], from which the 18O-enriched water diffuses throughout the leaf [42].

The δ18O values of herbivore tooth enamel carbonates are influenced by the oxygen isotopic composition of imbibed water which includes open water sources, groundwater, and leaf water [47,48]. Obligate drinkers, animals such as cattle that require substantial quantities of water on a daily basis, exhibit lower δ18O values than non-obligate or semi-obligate drinkers that meet their water needs from 18O-enriched leaf water [49]. Furthermore, herbivores ingesting a higher proportion of grass exhibit higher δ18O values compared to animals consuming higher amounts of browse [25]. Shallow-rooting grasses exhibit pronounced evaporative effects across the leaf surface [50] so that ingestion of 18O-enriched leaf water by sheep and goats would result in higher δ18O values compared to obligate drinking cattle.

Strontium isotopes

Strontium isotopes in biogenic tissues including enamel hydroxyapatite provide a robust means to evaluate animal mobility at local, regional, and transregional scales [5153]. The distribution of bioavailable strontium in soils varies across a defined region, determined by the composition of local weathered bedrock as well as strontium delivered by aeolian dusts and rainfall [54,55]. Strontium isotope ratios (87Sr/86Sr) of different bedrock geologies are defined by substrate age and the original ratio of rubidium and strontium present in bedrock [54]. Strontium isotopes enter the foodweb primarily through plants which take up resident strontium present in soils without isotopic fractionation due to the heavy molecular weight of strontium isotopes [56]. Herbivores thus directly incorporate strontium isotopes derived from graze and, to a lesser extent, water [57]. Unlike bone, which in general does not retain in vivo strontium isotope ratios [58], the highly organized crystalline structure of tooth enamel is highly resistant to diagenetic alteration, providing authentic values associated with the period of enamel mineralization [59].

Southern Levantine environments in the Upper Jordan Valley

Hazor is situated in the Hula Basin (70 m a.s.l.), which constitutes the immediate hinterland and periphery of Hazor (Fig 2). Part of the Upper Jordan Valley (UJV), the Hula Valley covers 180km2. To the west, the UJV is bounded by the Upper Galilee Mountains (1000 + m a.s.l.) composed of Cretaceous limestone and Neogene sedimentary rocks. To the east of the UJV lies the Golan Heights, reaching over 1000 m a.s.l and made up of successive layers of Pliocene and Pleistocene volcanic bedrock [60]. Basalts and pyroclast comprise the geology to the south/southeast of Hazor (north of Lake Kinneret). The distribution of bioavailable strontium in the soils of the Upper Jordan Valley, Golan Heights, and Galilee is determined by the composition of locally weathered bedrock and strontium from aeolian dusts, largely originating from the Sahara, as well as by precipitation moving bioavailable exchangeable strontium from marine deposits and sea spray [61,62].

Differences in bedrock geology, associated soil members, deposition of atmospheric strontium, and mean annual precipitation together produce spatial distinction in bioavailable strontium isotopes (87Sr/86Sr) in the hinterlands surrounding Hazor [63]. The Upper Galilee Mountains are dominated by terra rossa soils derived from the weathering of underlying Cretaceous limestones and dolomites, yielding 87Sr/86Sr ratios of 0.70831 to 0.70883. Rendzina soils, comprised of calcareous soils developed from Cretaceous and Neogene chalks and marl, yield lower strontium isotopes ranging from approximately 0.70790 to 0.70840 [63]; rendzinas occur west and southwest of Hazor, developed over Eocene limestone bedrock (Fig 3). Golan basalts exhibit low strontium isotope values ranging from approximately 0.70448 to 0.70770, with the youngest flows dating to the late Pleistocene yielding the lowest values, and flows dating to the upper Pliocene exhibiting the highest values [63].

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Fig 3. Geological map of the Upper Jordan Valley and Lower Galilee [60].

Jurassic limestones = blue, Cretaceous dolomites and limestones = green, Eocene limestones = orange, Neogene conglomerates = yellow, Pliocene and Pleistocene cover basalts = red. Soil and rock 87Sr/86Sr ratios from ref [120]. Estimated 87Sr/86Sr ratios for Jurassic limestones associated with terra rossa soils (green box 1) (displayed in ref [109]). Bioavailable plant 87Sr/86Sr isotope values are indicated for terra rossa soils (green box 1), rendzinas (blue box 2), and basalts (red box 3 – 6) (original data from ref [63] and also displayed in ref [109]). Dissolved strontium in Banias spring water derived from the Jurassic unit yields a 87Sr/86Sr ratio of 0.7072 [64].

https://doi.org/10.1371/journal.pone.0328934.g003

The Hula Valley is situated within the Lake Kinneret watershed (2570 km2), which includes the lower-elevation topographic drainage of the upper Jordan River that flows directly into Lake Kinneret, as well as an upper watershed defined by the karstic limestone mountain ridges of Mt. Hermon. There is a strong precipitation cline across the southern Levant along an east-west and north-south gradient. Strong orographic effects contribute to the precipitation gradient over the Lake Kinneret watershed, where rainfall amounts range from 400 mm/year for Lake Kinneret to 1200 mm/year for Mount Hermon [65,66]. Precipitation falls during the cooler seasons (November to April) with higher precipitation levels at higher elevations. Mean annual precipitation in the Hula Valley is 500–600 mm/year. Summers are hot and dry with annual temperatures averaging 27ºC in the Hula Valley and 21ºC for the Upper Galilee.

Hazor is situated within a seasonal C3 Mediterranean phytogeographic zone, relatively close to other phytogeographic zones characterized by different vegetation communities. C4 sedges growing around springs and ponds are also present in the Mediterranean environments of the UJV [67]. Higher precipitation levels in the Golan support C3 growth depleted in 13C particularly during the wet season [32]. A drier Mediterranean facies that also supports some Irano-Turanian flora spreads south of Lake Kinneret and along the mid-elevation landscapes of the Rift Valley, while a more arid Saharo-Arabian zone, also containing some Irano-Turanian vegetation including 13C-enriched C4 taxa, is distributed along lower elevations [68]. The very lowest elevations of the Rift Valley averaging −200 to −300 m b.s.l. receive less than 200 mm of rainfall per year, supporting Saharo-Sudanian and Sudan-Zambezian phytogeographic zones that contain a higher abundance of C4 flora [69].

Zooarchaeological methods

Zooarchaeological analyses of faunal remains from Hazor provide a means to evaluate herd management strategies that may articulate with particular pasturing practices and Aramean-Israelite political dynamics. The faunal remains from Area M were recovered from accumulations of urban refuse representing time-averaged secondary discard associated with storage buildings, domestic structures, and also general fill contexts (Table 1). Not unexpectedly, no animal pens have so far been identified inside the built environment. Livestock may have belonged to Hazor households, kept outside the city and managed through kin-based or hired labor, or owned by pastoralists extrinsic to the urban community at Hazor. Regardless if livestock herds were husbanded by urban or rural components of Israelite society, successful caprine husbandry requires regular access by herds to sizeable pastures, and caprines consumed in the city would reflect the isotopic inputs related to pasturing regimes enacted on herds.

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Table 1. Contextual information for mandibular tooth specimens sampled for isotope analyses. O = Ovis aries, C = Capra hircus, OC = Ovis/Capra. ‘n’ indicates number of increments measured for each tooth. Mandibular wear stage determined following ref [76] and age at death estimated using ref [86]. Age cohorts: Age 1 (MWS A-C, 0–12 months; Age 2 = MWS D, 1–2 years; Age 3 = TWS E, 2–3 years; Age 4 = MWS F, 3–4 years; Age 5 = TWS G-I, 4–10 years).

https://doi.org/10.1371/journal.pone.0328934.t001

Faunal remains were collected by hand during the excavation and were identified using the comparative collection of the Laboratory of Archaeozoology at the University of Haifa. Recovery by hand was complemented by extensive wet-sieve sampling, which yielded very few mammal bones that were taxonomically or otherwise informative, suggesting that hand recovery provides as good a sample of the faunal record for medium- and large-sized mammals (cf. ref [70]). Faunal remains are archived in the Israel Antiquities Authority while teeth analyzed for carbon and oxygen stable isotopes as part of this study are archived in the Archaeology Stable Isotope Laboratory (ASIL) at the University of Kiel. To estimate taxonomic and demographic variables of livestock use, we used a focused protocol that records a set of data-rich diagnostic zones [71], (see discussion in [7274]). Epiphyseal fusion [75], tooth eruption and wear [76], and biometrical [77,78] data were collected when possible for all livestock taxa (S1 Table and S2 Table). To check the equivalence of taphonomic deposition scenarios between contexts, data on bone surface modification such as weathering, carnivore gnawing, butchery, and burning marks were noted when observed with the naked eye.

Statistical analyses were carried out in R (R Core Team 2018) and visualized using the ggplot2 package. The association plot was drawn using the “vcd” package v. 1.4–7 [79]. Least-square regression of cosine functions was conducted in Excel using a file published by Pieterse [80].

Selection of teeth and intra-tooth sampling for isotope analysis

Teeth do not remodel after mineralization, preserving a seasonal-scale record of dietary intake and environmental inputs. We selected third mandibular molars for our analyses because these specimens were the most well represented in the assemblage. For sheep and goats, formation of the third mandibular molar begins at approximately 9–10 months and is completed by 20–22 months [81,82]. Crown formation and enamel mineralization rates in goats and sheep appear to be broadly similar [83]. The third mandibular molar exhibits greater temporal variation in crown formation rates between individuals compared to the second mandibular molar, although the amplitude of seasonal isotopic change captured in the tooth crown is similar [84]. For cattle, the third mandibular molar begins formation at approximately 9–10 months in age and is completed by ca. 23–24 months [85]. Continuous wear of hypsodont bovid teeth erases some portion of seasonal isotopic inputs recorded in tooth enamel. Consequently, the amplitude of intra-tooth isotopic change visible in older animals is reduced due to elimination of seasonal peaks and troughs within the sequence (i.e., minimum and maximum isotope values). Mandibular wear stages used to assess relative animal age at death, described above, were used to estimate the extent of tooth structure loss of sampled specimens (Table 1).

All necessary permits were obtained for the described study, which complied with all relevant regulations. Archaeological tooth samples were exported to the Archaeology Stable Isotope Laboratory (ASIL) at the University of Kiel under permit number 14197 issued by the Israel Antiquities Authority. Third mandibular molars (M/3) from domesticated sheep (Ovis aries, n = 2), goats (Capra hircus, n = 3), sheep/goat (Ovis/Capra, n = 2), and cattle (Bos taurus, n =1) were selected from early Iron Age II (IRIIA) contexts dated to approximately the 10th–9th centuries BCE, a period of conflict between the Aramean and Israelite states. This chronological assignment is based on the stratigraphic context of the finds and ceramic typology of pottery sherds recovered from the same stratigraphic context as the faunal remains. M/3’s from five sheep/goat (n = 5, Ovis/Capra) one gazelle (n = 1, Gazella sp.), and one cattle (n = 1, B. taurus) were selected from the late Iron Age II (IRIIB) contexts dated on the basis of stratigraphy and ceramic topology to the 8th century BCE, a period defined by a cessation of conflict between the two polities reflecting the devastation of the Aramean state following the Assyrian conquests. Third mandibular molars were identified as belonging to either sheep or goat based on morphological criteria established by ref [87].

Teeth were cleaned mechanically using a scalpel in order to remove both dental calculus and any adhering sediments. Teeth were then sonicated in distilled water to ensure full removal of contaminating particles and dried overnight at 60oC. A sequence of horizontal bands approximately 1 mm in width were drilled perpendicular to the growth axis from the tooth crown (earliest forming) to the enamel root junction (ERJ; latest forming) on the buccal side of the anterior tooth loph using a diamond-tipped drill bit. Sampling the same tooth loph enhances comparability of isotopic time-series between individuals by reducing the impact of tooth geometry on the shape and amplitude of intra-tooth isotopic change [8890]. Overall, the seasonality of intra-tooth isotopic change recorded in hypsodont teeth is attenuated relative to seasonal change in environmental isotopes due to incremental deposition of enamel matrix at daily scales, successive cross mineralization during subsequent enamel maturation over longer timescales, tooth geometry, and a sampling strategy that cross-cuts successive mineralization fronts. Amelogenesis progresses through multiple, discontinuous stages during which a primary matrix is laid down in successive cone-like structures, subsequently mineralizes, and matures over an extended period of time [9194]. The extended process of enamel maturation involves the deposition of secondary mineralization fronts that run transverse to the long axis of enamel prisms, introducing time-averaging in the measured isotopic sequences [9597]. An exponential decrease in tooth growth rates towards the crown-enamel junction further integrates isotopic inputs [9799].

Enamel powders were soaked in approximately 1 mL of 0.1 M acetic acid for 4 hours to remove diagenetic carbonates [100,101], then rinsed with distilled water and centrifuged at 6000 rpm five times. Enamel powders were then freeze-dried. After treatment, samples were reacted with 100% orthophosphoric acid at 75ºC in an automated cryogenic distillation system (Kiel IV device) interfaced with a ThermoScientific MAT 253 mass spectrometer located in the Leibniz Laboratory for Radiometric Dating and Stable Isotope Research at the University of Kiel. Two international carbonate standards, NBS-19 and IAEA-603, were run daily. Analytical precision was 0.05‰ for carbon isotope ratios and 0.08‰ for oxygen isotope ratios. Three different internal enamel standards measured an analytical precision of 0.10‰ for oxygen isotope ratios and 0.15‰ for carbon isotope ratios (CM1_D; 48 measurements), 0.08‰ for oxygen isotope ratios and 0.14‰ for carbon isotope ratios (ER1_C; 8 measurements), and 0.07‰ for oxygen isotope ratios and 0.09‰ for carbon isotope ratios (ER1_D; 40 measurements). All isotopic values were reported relative to the V-PDB (Vienna Pee Dee Belemnite) standard using NBS-19.

Enamel samples for strontium isotope analyses were removed from portions of the tooth exhibiting high summer season maximum δ18O values and low winter season minimum δ18O values. For strontium isotope analysis, separation chemistry for enamel samples was performed in a class 100 clean lab facility located in the MC-ICP-MS facility in the Department of Geological Sciences, University of Cape Town (UCT). Samples were transferred to Savillex beakers digested in 2–3 mL of 65% HNO3 at 140ºC. After complete dissolution, the samples were dried overnight at 130ºC and re-dissolved in 2 mL 2M HNO3 and loaded into 800 μL Teflon columns containing pre-conditioned Eichrom Sr-spec pure resin (500 = 100 μm). After six washes with 2M HNO3, the Sr was eluted with 3 mL of ultrapure deionized water, dried, and re-dissolved to a volume of 0.2% HNO3. 87Sr/86Sr isotope analyses were performed using a Nu Instruments NuPlasma HR multi-collector ICP-MS instrument housed in the Department of Geological Sciences, University of Cape Town. 87Sr/86Sr data presented were referenced to bracketing analyses of the international strontium isotope standard NIST SRM987 using a 87Sr/86Sr normalizing value of 0.710255. Isobaric interference of 87Rb on 87Sr was corrected using the measured signal for 85Rb and the natural 85Rb/87Rb ratio. The effect of instrumental mass fractionation was corrected using the exponential law and a 86Sr/88Sr value of 0.1194. All samples were run to an internal precision of ± 0.000037 (2SE) or better. Repeat analyses of an in-house carbonate reference material (NM95) analyzed as an unknown along with the enamel samples during processing of samples for this study yielded an 87Sr/86Sr value of 0.708910 ± 0.000030 (n = 3) in agreement with long-term data for this material (0.708911 ± 0.000040 2s; n = 414; over >8 years). Total procedural strontium blanks were typically better than ~250 pg and therefore negligible.

Modelling birth seasonality

The position of maximum δ18O and δ13C values in the tooth crown relative to the ERJ provides information on the seasonality of birth and dietary intake, respectively. However, inter-individual variability in tooth size and geometry impacts the shape of isotopic curves and, in turn, influences the relative positioning of minimum and maximum isotope values in time series [102]. Here, we reduced the impact of inter-individual variation in tooth size to better fit isotopic time-series alignments by normalizing distances of sampled increments using an equation derived from a cosine function developed by ref [103]. Although the M/3 exhibits wider variation in growth rate than the M/2 [102,103], third mandibular molar-based calculations will also separate season of birth between individual animals [84,104]. Previous work comparing oxygen isotope curves modelled for M/2 and M/3 teeth from the same analytical cohort indicates temporal disparities in modelled birth seasonality, with the second molar demonstrating a shorter birthing period of only four months while the third molars demonstrate an extended birth season taking place over six months [84].

A least square regression was used to obtain best-fit values for parameters A, x0, X, and M while the values of p, b, and xB were fixed to 0 and xA was fixed to 106. This four-parameter model was found to be sufficient by ref [103] for modelling the majority of cases. The season of birth was determined by evaluating the position in the tooth crown where the highest (xmax= x0) and lowest (xmin= x0+ X/2) δ18O values were modelled relative to the period of the annual cycle (X). The birth seasonality of individuals 5756 and 5808 from the IRIIA and 5758, 5812, and 5817 from the IRIIB could not be determined due to shortness of isotopic sequence.

Results

Retention of caprine management systems over time at Hazor

Shifts in geopolitical boundaries and border permeability, as well as subsistence and economic dynamics influence the spatial extent of pastoral systems and shape of herd management strategies that entail adjustment of livestock mobility and herd demography. Zooarchaeological results indicate similarity in faunal assemblages representing IRIIA and IRIIB occupation at Hazor. A total number of 737 specimens were identified to genus in the Iron Age II assemblage (Table 2). Sheep (Ovis aries) and goats (Capra hircus), evenly represented with a ratio close to 1:1 (Table 2, Fig 4; identifications based on ref. [105]), were heavily exploited in both periods, followed by cattle (Bos taurus) and equids. The latter included both donkey (Equus asinus) and horse (E. caballus), clearly distinguishable by size (Late IRIIA, 9th century: 13 horse, 14 donkey; IRIIB, 8th century: 13 horse, 16 donkey, see Table 2). Deer (Dama mesopotamica) and gazelle (Gazella cf. gazella) were occasionally hunted. Tooth wear data from sheep/goat specimens indicate focused harvesting of mature adults approximately 4 years in age. Some slaughter of yearlings was also undertaken during the early IRIIA (10th century BCE; Table 1, Fig 5). The predominance of skeletal remains from mature adults is typical of a livestock producer economy and contrasts with an abundance of older juveniles and young adults expected for consumer settlements (e.g., ref [106]). Overall, strong similarity in sheep-to-goat ratio and age-at-death calculated for IRIIA and IRIIB occupation suggests no major change in caprine production goals for subsistence or otherwise between the 10th and the 8th centuries BCE.

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Table 2. Relative frequencies of taxa from Iron Age II Hazor (Area M). Number of Identified Specimens (NISP) values are based on diagnostic zone counts according to ref. [71]. Early IRIIA (10th century BCE), Late IRIIA (9th century BCE), IRIIB (8th century BCE).

https://doi.org/10.1371/journal.pone.0328934.t002

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Fig 4. Chi-square standardized residuals of taxa by period.

Dark purple = significant under representation, dark blue = significant overrepresentation. Total chi-square = 43.97, P < 0.001.

https://doi.org/10.1371/journal.pone.0328934.g004

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Fig 5. Age-at-death distributions for caprines for Iron Age II Hazor analyzed as part of this study based on ref [86] (computed according to mandibular wear stages defined by ref [76]).

Age cohorts: Age 1 (MWS A-C, 0–12 months; Age 2 = MWS D, 1–2 years; Age 3 = TWS E, 2–3 years; Age 4 = MWS F, 3–4 years; Age 5 = TWS G-I, 4–10 years).

https://doi.org/10.1371/journal.pone.0328934.g005

However, a decline in the frequency of caprines from 81% to 53% from the early IRIIA to the IRIIB, accompanied by a concomitant, dramatic increase in the frequency of cattle and equids, rising from 16% in the IRIIA to 35% in IRIIB, suggests an intensification in agriculture, an activity which involved a larger number of traction animals, over large-scale pastoral production (see Table 1 and Fig 4). Notably, this corresponds with a decrease in tree cover between 9th–8th centuries BCE associated with land clearance, possibly for increased agricultural output [107]. A shift in economic strategies linked to food production is also reflected in the architectural changes of the city with large, public storage buildings falling out of use during the IRIIB, replaced by silos placed in households. Despite changes in the type of animal and agricultural production, similarity in caprine management systems with the exploitation of goats and sheep more or less similar across the Iron Age II suggests that caprine mobility patterns were not dictated by household subsistence or economic goals but by landscape accessibility, as defined by polities and borders.

Caprine birth seasonality

Establishing the timing and distribution of animal season of birth is an important husbandry strategy that helps herders coordinate the production of milk and meat throughout the year. Birth seasonality in sheep and goats can be established through analysis of the positioning of summer-season maximum and winter-season minimum oxygen isotope values expressed in mandibular molars [102]. The locations of minima and maxima δ18O values relative to the enamel-root junction (ERJ) are presented in Table 3 and, based on those data, the modelled season of birth is presented in Fig 6 for Late IRIIA and IRIIB animals. For each case, the fit of the model was tested using an ANOVA; all p-values were below 0.05. Model results indicate moderate variation in caprine birth seasonality. An offset of ¼ of a full period (X) is visible between late IRIIA (n = 5) and IRIIB (n = 3) xmax and xmin values. When normalized to the period of the cycle, the position of maximum δ18O values indicates extended birth seasonality at Tel Hazor, stretching over ca 5–6 months (IRIIA: 0.54 year; IRIIB: 0.42 year; Fig 5), and that this period shifted by approximately 3 months (0.24 year) between the earlier and later Iron II occupations.

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Table 3. Best fit (least-squares regression) and cosine function for A (amplitude), x0 (position of δ18Omax value in tooth crown), X (period of cycle), M (mean), xmin = position of lowest δ18O value in tooth crown; p-value refers to ANOVA test on the least-square regression. O = Ovis aries, C = Capra hircus, OC = Ovis/Capra. Absent specimens are those that did not yield clear seasonal minimum or maximum oxygen isotope values required for calculations.

https://doi.org/10.1371/journal.pone.0328934.t003

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Fig 6.  Relative positions of maxima δ18O values measured in intra-tooth oxygen isotope sequences of caprine third molars, normalized to the period of the yearly cycle [period (X0/X)], as determined using the modelling procedure according to ref [103].

https://doi.org/10.1371/journal.pone.0328934.g006

Asynchronicity in modelled sequences likely reflects a combination of loosely staggered birth seasonality and also inter-tooth differences in third mandibular molar tooth formation rates which are more variable than the second mandibular molar more commonly used to establish birth seasonality [84]. Short lambing periods are associated with transhumant herd movement for the production of wool and meat, while extended birth seasons are associated with closely pastured animals for meat and milk. The extended birth season seen at Hazor fits well with production strategies oriented toward milk production for household consumption along with staggered meat production for markets. Extended birth seasonality observed in both the IRIIA IRIIB period could also reflect ad hoc provisioning of Hazor with live animals or meat obtained from multiple herding households employing distinct caprine husbandry strategies for different production goals. Alternatively, the intensification of agricultural activities (suggested by the increased importance of cattle) necessitated an extended season of birth in order to reduce convergence of labor-intensive plowing of fields and planting new crops with equally labor-intensive lambing.

Oxygen (δ18O) and carbon (δ13C) isotope results

The oxygen isotopic composition of the Hazor caprines reflects not only seasonal variation in geospatially sensitive meteoric water δ18O values, but also animal drinking and feeding behaviors that partially overwrite meteoric water oxygen isotope inputs [108]. Bovid livestock from Hazor exhibit moderate seasonal variation in oxygen isotopes expressed in sequentially sampled molars. Mean δ18O values yielded by each tooth specimen range from −3.4 to 2.6‰. Caprines from the IRIIA level range from −5.4 to 1.5‰ in δ18O while caprines from the IRIIB levels range from −4.8 to 3.7‰ (Table 4; S3 Table). Caprines exhibited moderate intra-tooth oxygen isotopic change (Δ18O) ranging from 2.7 to 5.0‰ in the IRIIA (Figs 7 and 8) and 2.4 to 6.1‰ in the IRIIB (Fig 9) and, in general, exhibited sinusoidal variation reflecting seasonal shifts in the oxygen isotope ratios of imbibed water. Cattle display low-amplitude intra-tooth oxygen isotope change, likely reflecting the intake by obligate drinking cattle of groundwater water from wells or springs, sources that yield an integrated isotopic signal derived from paleowaters, seasonal recharge events, recharge from different regional and local aquifers, or open water consistently affected by evaporative processes. The single IRIIA Bos specimen exhibits δ18O values ranging from −3.6 to −2.0‰, while the IRIIB Bos specimen exhibited a somewhat lower amplitude of oxygen isotopic change ranging from −3.6 to −2.9‰. The single gazelle specimen exhibited the highest intra-tooth δ18O values ranging from −1.8 to 3.7‰.

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Table 4. Summary statistics for carbon (δ13C), oxygen (δ18O), and strontium (87Sr/86Sr) isotope values measured from sequentially sampled third mandibular molars from cattle (Bos cf. taurus), goat (C, Capra hircus), sheep (O, Ovis aries), sheep/goat (OC), and gazelle (Gazella sp.). ‘n’ indicates number of increments measured for each tooth, ∆ = indicates range of intra-tooth isotopic change (i.e., difference between minimum and maximum isotope values), asterisk (*) indicates specimens exhibiting a full seasonal cycle in oxygen isotope sequences. Strontium isotope values coincide with maximum and minimum δ18O values expressed within each tooth.

https://doi.org/10.1371/journal.pone.0328934.t004

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Fig 7. Intra-tooth oxygen (δ18O) and carbon (δ13C) isotope sequences measured from a single sheep (Ovis aries) tooth recovered from the Early IRIIA period deposits.

Carbon isotope (δ13C) values = blue, oxygen isotope (δ18O) values = yellow.

https://doi.org/10.1371/journal.pone.0328934.g007

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Fig 8. Intra-tooth oxygen (δ18O) and carbon (δ13C) isotope sequences measured for bovid taxa recovered from Late IRIIA period deposits.

Squares = Bos sp., circles = Capra/Ovis (see Table 1 for detailed taxonomic assignments for caprines). Carbon isotope (δ13C) values = blue, oxygen isotope (δ18O) values = yellow.

https://doi.org/10.1371/journal.pone.0328934.g008

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Fig 9. Intra-tooth oxygen (δ18O) and carbon (δ13C) isotope sequence measured from a single sheep (Ovis aries) tooth recovered from early IRIIB period deposits.

Squares = Bos sp., circles = Capra/Ovis, diamonds = Gazella sp. (see Table 1 for detailed taxonomic assignments for caprines). Carbon isotope (δ13C) values = blue, oxygen isotope (δ18O) values = yellow.

https://doi.org/10.1371/journal.pone.0328934.g009

Caprines from Hazor yield a wide range of intra-tooth δ13C values from −12.2‰ to −10.6‰ observed in the single early IRIIA specimen, −12.3‰ to −5.0‰ for late IRIIA animals, and −13.6‰ to −6.9‰ for IRIIB animals (Fig 10). Caprines from both periods exhibited similarly wide variation in intra-tooth carbon isotopic change (Δ13C) ranging from +1.2‰ to 4.6‰ in the IRIIA (Table 4, Figs 7 and 8) and +1.0‰ to 4.5‰ in the IRIIB (Fig 9); carbon isotope sequences exhibit sinusoidal variation reflecting seasonal changes in dietary intake. The single Bos specimen from the late IRIIA level at Hazor exhibits moderate intra-tooth carbon isotopic variation ranging from −11.8 to −9.0‰ while the Bos specimen from the IRIIB level shows higher δ13C values from −7.3 to −4.5‰. Furthermore, intra-tooth carbon and oxygen isotopic variation exhibits consistent seasonal patterning between individuals. In general, seasonal shifts in intra-tooth carbon isotope change are inversely related to changes in oxygen isotopic composition, such that low wet-season δ18O values coincide with high δ13C values, while high summer season δ18O values correspond with low δ13C values (Figs 79). Other sequences instead exhibit high δ13C values associated with mid-range δ18O values likely representative of a spring season enamel formation (e.g., 5752), whereas another set of sequences show  low δ13C values corresponding to mid-range δ18O values expressed after summer maxima oxygen isotope values consistent with an autumn season formation period (e.g., 5805).

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Fig 10. Range of intra-tooth δ13C values for Hazor caprines (boxes without labels), cattle (B), and gazelle (G).

The periods are marked by color: light blue = Early IRIIA, blue = Late IRIIA, dark blue = IRIIB.

https://doi.org/10.1371/journal.pone.0328934.g010

Strontium isotope results

The range of variation of strontium isotope values.

The strontium isotopic composition of caprine livestock from both the IRIIA and IRIIB indicates these animals were pastured on basalt geologies (Table 4, Fig 11). A high proportion of strontium isotope ratios measured for animals from both periods ranged from approximately 0.7075 to 0.7079, values associated with Pliocene Dalton basalts, Dalwe basalts and cover basalts that cover the western and southeastern Golan, politically associated with Aram during the Iron Age II (Figs 2 and 3). Two IRIIA animals, one sheep (5752) and one cattle (5750) exhibited the very low strontium isotope ratios typically associated Pliocene basalts distributed to the east of the northern Hula Valley (Fig 3). Only one IRIIB cattle individual (5754; yielding radiogenic Sr values of 0.70844 and 0.70850) was grazed on terra rossa soils associated with Cretaceous limestones that covered the Lower Galilee region west and northwest of Hazor including the western edge of the Hula Valley, and also Jurassic limestones to the north-northeast. IRIIA and IRIIB bovids exhibited both low intra-tooth shifts in strontium isotope ratios suggestive of small-scale seasonal moves as well as wide intra-tooth differences in 87Sr/86Sr indicating larger scale movement to different pastures (Fig 11).

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Fig 11. Strontium isotope values for Hazor caprines, cattle, and gazelle.

Open circles coincide with δ18Omax values and black circles coincide with δ18Omin values. ‘B’ indicates cattle (Bos cf. taurus) and ‘G’ indicates gazelle (Gazella cf. gazella); all other values represent caprines. Colored horizontal bands and vertical bars indicate range of bioavailable strontium isotope values based on mean ± 1σ according to ref [63]. Horizontal blue dashed lines indicate the full range of 87Sr/86Sr values expected for rendzinas from refs [63,109].

https://doi.org/10.1371/journal.pone.0328934.g011

Discussion

Caprine pasturing systems unaffected by Iron Age conflicts

Multi-isotope data together suggest that bovid livestock grazed on spatially distinct pastures local to Hazor, within the Upper Jordan Valley, and also on more distant pastures located in the adjacent Golan Heights, an area that included contested landscapes held by Aram. Notably, there is no evidence that livestock pastured on rendzina soils located immediately north of Hazor or to the west in the Lower Galilee nor grazed on pastures supported by the radiogenic terra rossa soils that characterize the much of the Lower Galilee, the western portion of the UJV, or Mt. Hermon except for a single IRIIB cattle specimen (Fig 11). This individual (5754) exhibited the most radiogenic 87Sr/86Sr ratio (mean = 0.70846) of all sampled individuals from Hazor as well the highest dietary δ13C values (range = −21.9 to −19.0‰), indicating the animal regularly ingested water-stressed C3 plants and probably some C4 graze. This strontium value is expected for animals ingesting forage growing on terra rossa soils present along the western flanks of the Jordan Valley [63], which also supports 13C-enriched C4 sedges growing near springs and flowing water [32].

Instead, Hazor pasturing systems were oriented eastward during both the IRIIA and IRIIB, focused on the volcanic landscapes of the Golan Heights where well-watered pastures would have supplied herds with an abundant food supply. Pasture systems also included grazing grounds located to the south of Hazor in the rolling hills that surround the northern end of Lake Kinneret. Bioavailable 87Sr/86Sr ratios across these volcanic landscapes vary according to age with isotopic separation visible between plants growing on Pliocene, early Pleistocene, and late Pleistocene volcanic layers comprised of basalts and pyroclasts; this difference is driven largely by an increased contribution of atmospheric (strontium) deposition in soils that formed on old bedrock relative to younger bedrock [32].

The southern reaches of the UJV and much of the Golan Heights are comprised of Pliocene cover basalts (Pβc, 0.7074–0.7079) and Late Pliocene basalt flows (Dalwe basalts, 0.7077) (Fig 3; Dalwe values from ref. [109]). Both IRIIA (5756, 5757, and 5808) and IRIIB (5758, 5801, 5813, and 5817) caprines yielded strontium isotopes values, indicating animals were pastured in the Golan or to the south of Hazor where cover basalts are also present. Taking advantage of the pronounced elevation and precipitation gradients across the UJV and Golan, pasturing in either of these regions can be more finely resolved based on the oxygen isotope ratios recorded in discrete portions of the tooth enamel structure. The oxygen isotopic composition of meteoric precipitation and groundwater is influenced by altitude [110]. In the UJV and Golan region, there is a −0.26‰ decrease in meteoric water δ18O values per 100 m increase in elevation [111], similar to the global gradient of −0.28‰/100 m [38]. The precipitation ‘amount effect’ further modifies meteoric water oxygen isotope values with a decrease in precipitation levels corresponding with increasing meteoric water δ18O values [37]. Tel Hazor annually receives ca 450 mm of precipitation while the Golan Heights receives from 500 to 800 + mm per year.

We expect winter wet season δ18Omin values visible in intra-tooth sequences measured from archaeological caprines to more closely reflect the oxygen isotopic composition of open or groundwater sources which more closely reflect meteoric water oxygen isotope ratios, particularly under cooler, more humid conditions such as the wet season when evaporative enrichment is slowed. Furthermore, the isotopic composition of groundwater in the Cenomanian-Turanian limestone aquifer in northern Israel reflects meteoric water values in recharge areas [110]. Either one of these water sources would direct the oxygen isotopic composition of caprine body water during the wet season, in contrast to the later spring and summer seasons when 18O-enriched leaf water would also heavily influence bioapatite δ18O values of these semi-obligate drinkers [25,49]. Leaf water δ18O values are strongly influenced by evapotranspiration and air relative humidity [112]; these isotope values shift seasonally with leaf water present in foliar growth pushed out under cooler, more humid conditions during the wet season conditions more similar to meteoric water oxygen isotopes (i.e., depleted in 18O) compared to leaf water in later foliar growth enriched in 18O under more arid and warmer evaporative conditions [31,112].

Obligate-drinking cattle from Hazor, exhibiting an intra-tooth range from −3.6 to −2.0‰ in δ18O values, provide a rough estimate of seasonality in the oxygen isotope composition of groundwater and/or open water sources in the Upper Jordan Valley. However, the use of cattle δ18O values for the purposes of establishing an isotope reference data for local meteoric waters in the UJV is further complicated by the integrated contribution of precipitation with varied isotope compositions, imparted by amount and altitude effects that influence meteoric water δ18O values [104]. Variation in the oxygen isotopic composition of groundwater and open water sources within the Kinneret watershed adds further complexity [110]. In the case of Hazor, cattle δ18O values possibly reflect such an integrated signal at a sub-regional scale (i.e., Hula Valley) and may be more useful for establishing larger-scale inter-regional mobility or animal exchange within the diverse precipitation regimes of the southern Levant.

Livestock grazing on more radiogenic basalts associated with Pliocene cover basalts and Dalwe flow are significantly enriched +2.3‰ in δ18O (wet season minima values) relative to individuals foraging on less radiogenic Pliocene basalts (with the exception of individual 5758 feeding on more radiogenic basalts but yielding a low δ18Omin value; Fig 12; t-test p = 0.0185). This suggests that pastures supported by soils derived from cover basalts and Dalwe flow received 18O-enriched precipitation and, thus, likely located at lower elevations near Hazor. Individuals foraging on less radiogenic Pliocene basalts all exhibited low δ18Omin values consistent with grazing at higher elevations in the Golan. Individual 5758 likely also grazed at higher elevations in the Golan, albeit on pastures that supported more radiogenic soils.

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Fig 12. Intra-tooth δ18Omin values and paired strontium isotope values for Hazor caprines, cattle (B), and gazelle (G).

Diamonds indicate IRIIA isotope values, circles indicate IRIIB isotope values. Convex hulls (dashed black lines) delineate livestock groups sharing a similar oxygen isotopic composition.

https://doi.org/10.1371/journal.pone.0328934.g012

δ18Omax values measured from tooth enamel appear to be less useful for establishing summer season mobility movement in semi-obligate drinking caprines in the absence of a large cohort of oxygen isotope data from contemporaneous non-obligate drinkers, such as gazelle, which would establish the upper limit of environmentally derived δ18O values from leaf water for a given locale [113]. Moderate oxygen isotopic enrichment of caprines relative to cattle, in particular summer season δ18Omax values (Table 4), confirms the steady contribution of an 18O-enriched water source to caprine body water, likely leaf water. The ingestion by caprines of evaporatively 18O-enriched leaf water, particularly during the warmer months, disrupts the geospatial relationship between meteoric water (and also groundwater) and oxygen isotope ratios recorded in tooth bioapatite. Consequently, the oxygen isotope ratios of obligate-drinking cattle and non-obligate drinking gazelle can be used to further constrain the geospatial origins of semi-obligate drinking caprines. For example, if the oxygen isotope ratios of caprines exhibiting strontium isotopes values associated with Upper Pleistocene basalts were higher than those yielded by the single gazelle from Hazor yielding an intra-tooth range of −1.8 to 3.7‰ in δ18O (and also yielding a strontium isotope value associated with Upper Pliocene basalts), it would be likely that the caprine came from a more arid region.

Altogether, intra-individual variation in δ18Omin values observed within each group of Hazor animals, distinguished by their strontium isotope ratios, is due to a complex intersection of environmental inputs, physiological effects, and dietary factors that modify the oxygen isotopic composition of body water. These include the isotopic composition of meteoric water and groundwater, seasonality in aridity levels, and foraging behaviour. Herbivores foraging on a higher proportion of graze exhibit higher δ18O values compared to browsers feeding in the same environment [25]. Shallow-rooting grasses exhibit pronounced evaporative 18O enrichment across the leaf surface [42], while deeper-rooted browse draws from 18O-depleted moisture in soils [114]. Consequently, differences in the relative contribution of C3 graze versus C3 browse to the caprine diet may drive variation in maxima δ18O values observed in Hazor caprines independent of animal mobility. Further complicating matters, animals foraging on well-watered green pastures, in particular new growth high in water content, obtain a larger proportion of their water from leaf water and compensate by reducing their intake from open water sources [115].

Diversity in pasturing regimes confirmed by carbon isotopes

Hazor caprines grazed on open C3 Mediterranean pastures, indicated by moderately low dietary δ13C values exhibited by most IRIIA and IRIIB caprines, ranging from ca −26 to −21‰. The single gazelle (5817) foraged under a light Mediterranean forest canopy indicated by consistently low intra-tooth dietary δ13C values averaging −27.6‰ (Fig 10). Understory foliar growth under a forest canopy is depleted in 13C compared to flora growing in open landscapes due to recycling of 13C-depleted CO2 produced by decaying leaf litter [33,116], soil efflux [117], decreased photosynthesis rates [118], and low stomatal conductance under low light conditions [119].

Wide variation in caprine δ13C values, combined with differences in intra-tooth carbon isotopic patterning shared between IRIIA and IRIIB caprines, further confirms considerable spatial diversity in pasturing strategies (Fig 10). Low dietary δ13Cmin values, ranging from −27.8 to −23.5‰ in both IRIIA and IRIIB caprines and coinciding with seasonal high δ18Omax values, indicate that these animals foraged during the spring and possibly early summer on well-watered C3 pastures. Seasonal transhumance from Hazor to higher elevation pastures located in the Golan Heights, a region that receives higher amounts of precipitation than the lower elevation Hula Valley due to orographic effects, would explain this isotopic pattern. Similarly low dietary δ13C values coinciding with high summer season δ18O values have been identified in Neolithic caprines east of the Jordan Valley; in this case, the pattern reflects seasonal vertical transhumance of animals to higher elevation, better watered C3 pastures in the Jordanian highlands [113]. At Hazor, winter season δ13C values were approximately 1–2.5‰ higher than summer season values, indicating ingestion of 13C-enriched fodder, a source that included later warm season C3 growth that grew under water-stressed conditions, perhaps collected from reserved pasture as agricultural by-products (i.e., straw or direct grazing on crop-stubble). Variation in the intensity and duration of fodder provided to Hazor sheep and goats, expressed as inter-individual differences in absolute δ13C values and shape of the carbon isotope sequences, most likely reflects differences in household access to fallow fields, winter pastures, and fodder storage capabilities.

Livestock were almost never grazed on drier, 13C-enriched pastures located in more water-deficient regions. Seasonal high dietary δ13Cmax values of −19.0‰ and higher observed in one Bos (5754, IRIIB) and one caprine (5756, late IRIIA) indicate these animals ingested water-stressed graze or C4 sedges. Such pasturage was available in the Jordan Valley where 13C-enriched C3/C4 Irano-Turanian vegetation thrives. The pattern of high δ13C values coinciding with winter season minimum δ18O values suggests that these animals were moved during the winter months to extra-local pastures located in more arid environments and provided there with dried 13C-enriched graze in winter pastures or as fodder.

Overall, high δ13C values coinciding with δ18Omin values in Hazor caprines suggest that livestock were provisioned with a 13C-enriched fodder source, probably water-stressed graze collected during the summer months, during the coldest portion of the wet season (see further discussion below), while pronounced variation in caprine summer and winter season δ13C values, as well as inter-individual differences observed in their relationship with oxygen isotope minima and maxima values, indicates substantial diversity in pasturing and foddering strategies. These patterns suggest that the movement and provisioning of caprine herds were managed at the household level, rather than coordinated by a centralized authority, which would be expected to produce more homogenous isotopic patterning.

Conclusion

Although isotopic analyses of a larger number of tooth specimens would help further define the ubiquity of specific caprine husbandry strategies practiced at Iron Age Hazor, the isotopic data presented here suggest that household-based flocks were grazed in spatially distinct pastures, independent of centralized coordination from Hazor. At the same time, the overall similarity in the heterogeneous character of pasturing regimes used by herders during periods of both calm and conflict indicates that local agro-pastoralist communities continued to pursue their own modes of subsistence and mobility despite pervasive inter-polity conflict between the Aramean and Israelite kingdoms. For sheep herders, the border between these polities remained permeable so that access to seasonal pastures was negotiated via channels that were independent of the state (Fig 1), for example according to tribal arrangements or other inter-community agreements. Conflicts may have instead been asserted between competing political entities in ways that primarily engaged urban elites and further acted out by military engagements. However, this did not involve close control of daily activities, including animal herding, conducted on hinterland landscapes. Ultimately, conflicts acted out by the military and ruling sectors of society did not affect the routine subsistence activities of everyday agro-pastoralists. The continued use of heterogeneous caprine pasturing strategies at Hazor through a period of settlement expansion, inter-polity conflict, and the transformation from an administrative center to a place more focused on domestic production speaks to the resilience of local agro-pastoralist populations.

Supporting information

S1 Table. Biometrical and fusion data for selected skeletal elements from Iron IIA and IIB Hazor. Metrical data according to von den Driesch (1976).

https://doi.org/10.1371/journal.pone.0328934.s001

(XLSX)

S2 Table. Tooth wear data and epiphyseal fusion data aggregated by period for faunal specimens from Iron Age Hazor.

https://doi.org/10.1371/journal.pone.0328934.s002

(XLSX)

S3 Table. Carbon (δ13C) and oxygen (δ18O) isotope values measured from the carbonate fraction of incrementally sampled bovid tooth specimens from Iron Age Hazor.

All isotope values reported relative to VPDB. ERJ = enamel root junction.

https://doi.org/10.1371/journal.pone.0328934.s003

(XLSX)

Acknowledgments

We gratefully acknowledge the support of the Steven B. Dana Archaeology Fund and Selz Foundation Hazor Excavations in Memory of Yigael Yadin for their generous support of excavations at Hazor. We also thank Nils Andersen (Leibniz Labor, University of Kiel) for assistance with carbon and oxygen isotope mass spectrometry and Petrus le Roux (University of Cape Town) for strontium isotope analysis.

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