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<journal-meta>
<journal-id journal-id-type="publisher-id">Front. Environ. Archaeol.</journal-id>
<journal-title>Frontiers in Environmental Archaeology</journal-title>
<abbrev-journal-title abbrev-type="pubmed">Front. Environ. Archaeol.</abbrev-journal-title>
<issn pub-type="epub">2813-432X</issn>
<publisher>
<publisher-name>Frontiers Media S.A.</publisher-name>
</publisher>
</journal-meta>
<article-meta>
<article-id pub-id-type="doi">10.3389/fearc.2025.1611071</article-id>
<article-categories>
<subj-group subj-group-type="heading">
<subject>Environmental Archaeology</subject>
<subj-group>
<subject>Original Research</subject>
</subj-group>
</subj-group>
</article-categories>
<title-group>
<article-title>Refining dietary shifts linked to climate oscillations in the Central Andes: stable isotope evidence from Vichama (1800&#x02013;1500 <sc>BCE</sc>)</article-title>
</title-group>
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<name><surname>Pezo-Lanfranco</surname> <given-names>Luis</given-names></name>
<xref ref-type="aff" rid="aff1"><sup>1</sup></xref>
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<name><surname>Crispin</surname> <given-names>Aldemar</given-names></name>
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<name><surname>Prado-Barrag&#x000E1;n</surname> <given-names>Alonso</given-names></name>
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<name><surname>Abad</surname> <given-names>Tatiana</given-names></name>
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<name><surname>Machacuay</surname> <given-names>Marco</given-names></name>
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<name><surname>Yseki</surname> <given-names>Marco</given-names></name>
<xref ref-type="aff" rid="aff3"><sup>3</sup></xref>
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<name><surname>Gorriti</surname> <given-names>Manuel</given-names></name>
<xref ref-type="aff" rid="aff3"><sup>3</sup></xref>
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<contrib contrib-type="author">
<name><surname>Miranda</surname> <given-names>Luis</given-names></name>
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<contrib contrib-type="author">
<name><surname>Apol&#x000ED;n</surname> <given-names>Jos&#x000E9;</given-names></name>
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<name><surname>DiMuro</surname> <given-names>Alice</given-names></name>
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<name><surname>Novoa</surname> <given-names>Pedro</given-names></name>
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<name><surname>Colonese</surname> <given-names>Andr&#x000E9; Carlo</given-names></name>
<xref ref-type="aff" rid="aff1"><sup>1</sup></xref>
<xref ref-type="aff" rid="aff2"><sup>2</sup></xref>
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<name><surname>Shady</surname> <given-names>Ruth</given-names></name>
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<aff id="aff1"><sup>1</sup><institution>Institute of Environmental Science and Technology (ICTA), Universitat Aut&#x000F2;noma de Barcelona</institution>, <addr-line>Bellaterra</addr-line>, <country>Spain</country></aff>
<aff id="aff2"><sup>2</sup><institution>Department of Prehistory, Universitat Aut&#x000F2;noma de Barcelona</institution>, <addr-line>Bellaterra</addr-line>, <country>Spain</country></aff>
<aff id="aff3"><sup>3</sup><institution>Zona Arqueol&#x000F3;gica Caral, Ministerio de Cultura</institution>, <addr-line>Lima</addr-line>, <country>Peru</country></aff>
<aff id="aff4"><sup>4</sup><institution>Facultad de Ciencias Sociales, Universidad Nacional Mayor de San Marcos</institution>, <addr-line>Lima</addr-line>, <country>Peru</country></aff>
<author-notes>
<fn fn-type="edited-by"><p>Edited by: Michelle S. Eusebio, University of the Philippines Diliman, Philippines</p></fn>
<fn fn-type="edited-by"><p>Reviewed by: Kurt Wilson, Lawrence University, United States</p>
<p>Weston McCool, University of Utah Department of Anthropology, United States</p></fn>
<corresp id="c001">&#x0002A;Correspondence: Luis Pezo-Lanfranco <email>Luis.Pezo&#x00040;uab.cat</email></corresp>
<fn fn-type="equal" id="fn001"><p>&#x02020;These authors share senior authorship</p></fn></author-notes>
<pub-date pub-type="epub">
<day>18</day>
<month>07</month>
<year>2025</year>
</pub-date>
<pub-date pub-type="collection">
<year>2025</year>
</pub-date>
<volume>4</volume>
<elocation-id>1611071</elocation-id>
<history>
<date date-type="received">
<day>13</day>
<month>04</month>
<year>2025</year>
</date>
<date date-type="accepted">
<day>30</day>
<month>06</month>
<year>2025</year>
</date>
</history>
<permissions>
<copyright-statement>Copyright &#x000A9; 2025 Pezo-Lanfranco, Crispin, Prado-Barrag&#x000E1;n, Abad, Machacuay, Yseki, Gorriti, Miranda, Apol&#x000ED;n, DiMuro, Novoa, Colonese and Shady.</copyright-statement>
<copyright-year>2025</copyright-year>
<copyright-holder>Pezo-Lanfranco, Crispin, Prado-Barrag&#x000E1;n, Abad, Machacuay, Yseki, Gorriti, Miranda, Apol&#x000ED;n, DiMuro, Novoa, Colonese and Shady</copyright-holder>
<license xlink:href="http://creativecommons.org/licenses/by/4.0/"><p>This is an open-access article distributed under the terms of the Creative Commons Attribution License (CC BY). The use, distribution or reproduction in other forums is permitted, provided the original author(s) and the copyright owner(s) are credited and that the original publication in this journal is cited, in accordance with accepted academic practice. No use, distribution or reproduction is permitted which does not comply with these terms.</p></license>
</permissions>
<abstract>
<p>The balance between fishing and farming for the development of complex societies in ancient Central Andes is the subject of a long debate. Despite the availability of marine resources in one of the world&#x00027;s richer marine ecosystems and the extreme aridity of the landscape, current evidence suggests that farming was the prime mover of early Andean civilization. In this study, we employed stable isotope analysis to reconstruct the diets of 38 human individuals from two occupations at Vichama, a coastal settlement with monumental architecture located in the Huaura Valley, Early Formative&#x02212;1 period (EF-1, 1800&#x02013;1500 <sans-serif>BCE</sans-serif>) and the Late Intermediate Period (LIP, 1000&#x02013;1300 <sc>CE</sc>). Results show a considerable contribution of C<sub>3</sub> cultivated plants and marine protein during EF-1, which indicates the first importance of farming at that time, whereas marine consumption increased during the LIP, a period of higher marine productivity. Our findings shed light on the complex interplay between climate alternances and coastal/inland sites&#x00027; hegemony in the Central Andes.</p></abstract>
<kwd-group>
<kwd>Andean formative period</kwd>
<kwd>dietary reconstruction</kwd>
<kwd>Bayesian stable isotope mixing models</kwd>
<kwd>climate change</kwd>
<kwd>Peruvian North-Central coast</kwd>
</kwd-group>
<counts>
<fig-count count="6"/>
<table-count count="2"/>
<equation-count count="0"/>
<ref-count count="87"/>
<page-count count="18"/>
<word-count count="13211"/>
</counts>
<custom-meta-wrap>
<custom-meta>
<meta-name>section-at-acceptance</meta-name>
<meta-value>Archaeological Isotope Analysis</meta-value>
</custom-meta>
</custom-meta-wrap>
</article-meta>
</front>
<body>
<sec sec-type="intro" id="s1">
<title>Introduction</title>
<p>The connection between subsistence systems, climate oscillations, and social complexity has been extensively studied and documented for the Central Andes. Studies have shown that ancient Andean populations developed adaptive strategies modulated by ecological constraints since they arrived at the region &#x0007E;14,000 years ago until the beginning of the Late Holocene (Dillehay, <xref ref-type="bibr" rid="B18">2017</xref>; Le&#x000F3;n, <xref ref-type="bibr" rid="B41">2013</xref>; Piperno and Dillehay, <xref ref-type="bibr" rid="B56">2008</xref>). Domesticate evidence dates to as early as 8000 <sc>bce</sc>, and available data suggest that irrigation technologies supported plant cultivation by &#x0007E;4500 <sc>bce</sc> (Dillehay et al., <xref ref-type="bibr" rid="B19">2005</xref>, <xref ref-type="bibr" rid="B21">2007</xref>). Since &#x0007E;3000 <sc>bce</sc>, farming was the most important economic activity (Grobman et al., <xref ref-type="bibr" rid="B28">2012</xref>; Yseki et al., <xref ref-type="bibr" rid="B85">2023</xref>).</p>
<p>Increasing settlement size and distribution, shifts in domestic architecture, and the spatiotemporal modeling of demographic dynamics for the Peruvian coast indicate an exponential population growth between 2000 and 1 <sc>bce</sc> (Goldberg et al., <xref ref-type="bibr" rid="B27">2016</xref>). The process has been attributed to a complex interplay between environmental changes and the expansion of agricultural regimes in the region (Contreras, <xref ref-type="bibr" rid="B13">2010</xref>; Goldberg et al., <xref ref-type="bibr" rid="B27">2016</xref>; Wilson et al., <xref ref-type="bibr" rid="B82">2022</xref>).</p>
<p>In the last decades, a large volume of archaeological information has been produced for the Peruvian North-Central coast (PNCC), the epicenter of early complexity in the Central Andes (Creamer et al., <xref ref-type="bibr" rid="B15">2013</xref>; Haas and Creamer, <xref ref-type="bibr" rid="B29">2006</xref>; Makowski, <xref ref-type="bibr" rid="B46">2023</xref>; Shady et al., <xref ref-type="bibr" rid="B72">2001</xref>; Shady, <xref ref-type="bibr" rid="B69">2018</xref>). Although pioneering evidence suggested that marine resources were the foundation of initial social complexity in this extremely challenging environment (Moseley, <xref ref-type="bibr" rid="B49">1975</xref>), new evidence of plant consumption at coastal and middle valley monumental sites strongly indicates that farming was the subsistence system supporting the emergence of early civilization in the region (e.g., Caral civilization) and subsequent sociopolitical developments (Dillehay et al., <xref ref-type="bibr" rid="B21">2007</xref>; Haas et al., <xref ref-type="bibr" rid="B30">2013</xref>; Pezo-Lanfranco and Colonese, <xref ref-type="bibr" rid="B51">2024</xref>; Shady, <xref ref-type="bibr" rid="B66">2006a</xref>,<xref ref-type="bibr" rid="B67">b</xref>; Shoji et al., <xref ref-type="bibr" rid="B74">2023</xref>; Tung et al., <xref ref-type="bibr" rid="B78">2021</xref>).</p>
<p>In this area, studies suggest that coastal and inland societies would have been organized around symbiotic farming&#x02013;fishing economies and littoral&#x02013;inland trade networks since at least the Initial Formative (3000&#x02013;1800 <sc>bce</sc>), with coastal communities based on marine resources and some contribution of cultivated crops and inland groups relying in cultivated plants supplemented by marine products (Pezo-Lanfranco et al., <xref ref-type="bibr" rid="B53">2022</xref>; Shady, <xref ref-type="bibr" rid="B69">2018</xref>; Yseki et al., <xref ref-type="bibr" rid="B85">2023</xref>).</p>
<p>This system, however, would have been influenced by the variability of sea surface temperatures (SSTs), upwelling dynamics, and productivity intensity in the Humboldt Current System (HCS), and El Ni&#x000F1;o Southern Oscillation (ENSO). The increase of SSTs produces a decrease in marine biomass, whereas in upper valleys, landslides increase as a consequence of intense rainfall, causing imponderable effects in farming production (Contreras, <xref ref-type="bibr" rid="B13">2010</xref>). In the north and central coast of Peru, humid periods are linked to the prevalence of El Ni&#x000F1;o&#x02013;like conditions, whereas La Ni&#x000F1;a&#x02013;like conditions can be associated with dry and cold climate (Salvatteci et al., <xref ref-type="bibr" rid="B57">2014</xref>, <xref ref-type="bibr" rid="B59">2019</xref>; Leclerc, <xref ref-type="bibr" rid="B40">2023</xref>).</p>
<p>Although the final causes and the complex mechanisms among the factors involved (e.g., ENSO teleconnections, Walker circulation intensity, meridional displacement of the Inter-Tropical Convergence Zone, and the South Pacific Subtropical High) are still in discussion (M&#x000E4;chtle and Eitel, <xref ref-type="bibr" rid="B45">2013</xref>; Salvatteci et al., <xref ref-type="bibr" rid="B57">2014</xref>, <xref ref-type="bibr" rid="B58">2016</xref>; Scholz et al., <xref ref-type="bibr" rid="B65">2014</xref>; Yseki et al., <xref ref-type="bibr" rid="B86">2022</xref>), variations in SSTs, upwelling conditions, and productivity intensity in the HCS occurring on multiple timescales (Salvatteci et al., <xref ref-type="bibr" rid="B58">2016</xref>, <xref ref-type="bibr" rid="B59">2019</xref>; Yseki et al., <xref ref-type="bibr" rid="B86">2022</xref>) likely had a significant impact on the availability and selection of marine resources utilized by past coastal communities and influenced the rise and decline of ancient Andean societies during the Holocene (Wilson et al., <xref ref-type="bibr" rid="B82">2022</xref>, <xref ref-type="bibr" rid="B83">2024</xref>). These effects occurred alongside persistent hazards, such as tectonic activity and geomorphological particularities (Orloff, <xref ref-type="bibr" rid="B50">2022</xref>; Salvatteci et al., <xref ref-type="bibr" rid="B59">2019</xref>; Sandweiss et al., <xref ref-type="bibr" rid="B60">2009</xref>).</p>
<p><italic>Export production</italic> refers to the fraction of organic carbon produced by photosynthetic organisms in the photic zone that is eventually deposited on the ocean floor; thus, it serves as a proxy for SST fluctuations in oceanography (Herbert, <xref ref-type="bibr" rid="B33">2003</xref>; Scholz et al., <xref ref-type="bibr" rid="B65">2014</xref>; Wefer et al., <xref ref-type="bibr" rid="B81">2015</xref>). A comprehensive palaeoceanographic study of the HCS based on high-resolution data (i.e., alkenone and bromide-to-titanium [Br/Ti] ratio) has revealed centennial-scale changes in SSTs and, in general, an increase in exported productivity from &#x0007E;3000 to 1100 <sc>bce</sc> (Salvatteci et al., <xref ref-type="bibr" rid="B59">2019</xref>).</p>
<p>Studies suggest that a cold period along the Peruvian coast between 1800 and 1200 <sc>bce</sc> produced a decrease in SSTs and an increase of marine productivity (Salvatteci et al., <xref ref-type="bibr" rid="B58">2016</xref>, <xref ref-type="bibr" rid="B59">2019</xref>), leading to changes in the equilibrium of political power to the benefit of coastal populations. High marine productivity of the Peruvian sea has been detected between &#x0007E;1700 and 1500 <sc>bce</sc> (Salvatteci et al., <xref ref-type="bibr" rid="B59">2019</xref>), coinciding with the flourishing of several ceremonial centers along the coastal area such as Vichama (Shady et al., <xref ref-type="bibr" rid="B73">2015</xref>) and Bandurria (Chu, <xref ref-type="bibr" rid="B12">2011</xref>), located, respectively, along the coast of the northern and southern interbasins of Huaura Valley, the more productive of the PNCC. However, the extent to which changes in marine productivity may have led to an increased reliance on marine resources at the expense of agricultural products, well known in the region since previous periods, remains untested. To investigate this issue, we integrate stable isotope data (&#x003B4;<sup>13</sup>C<sub>collagen</sub>, &#x003B4;<sup>15</sup>N<sub>collagen</sub>, &#x003B4;<sup>13</sup>C<sub>apatite</sub>) and Bayesian stable isotope mixing models (BSIMMs) of human individuals from Vichama, an Early Formative (EF) coastal site with monumental architecture, &#x0201C;key&#x0201D; to understanding the timing of dietary changes and the nuances of the Formative economy. From a broader theoretical perspective, this article offers valuable insights into how climatic conditions influenced the rise and collapse of complex prehistoric societies (Kennett and Marwan, <xref ref-type="bibr" rid="B39">2015</xref>).</p>
<sec>
<title>Archaeological context: Vichama</title>
<p>Vichama is an archaeological site located in the modern city of V&#x000E9;gueta, 130 km north of Lima, in the PNCC (UTM WGS84 N8779900/E212200). The site is at the right margin of the lower Huaura Valley (Huaura&#x02013;Supe interbasin), at 1.5 km from the Pacific Ocean and 6.5 km northwest to the river mouth (<xref ref-type="fig" rid="F1">Figure 1</xref>). The Huaura River is a permanent source of water that flows &#x0007E;158 km from its origin in the Andean highlands (i.e., Sura Saca lagoon&#x02014;Cordillera de Raura) to the sea and works as a natural corridor that links transversally the coast with important socioeconomic regions, such as the Mantaro basin and Eastern Andean lowlands (e.g., Alto Mara&#x000F1;&#x000F3;n, Alto Huallaga).</p>
<fig id="F1" position="float">
<label>Figure 1</label>
<caption><p>Location of Vichama during the Early Formative (1800&#x02013;1500 <sc>bce</sc>) and other major formative settlements of the Peruvian North-Central coast.</p></caption>
<alt-text>Map highlighting Early Formative archaeological sites in Peru along the North-Central coast, marked with pink dots. Key locations include Vichama, &#x000C1;spero, Caral, and Quebrada Chupacigarro Cemetery. The map includes rivers and shows modern agricultural and district capital areas. An inset displays Peru&#x00027;s location in South America.</alt-text>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fearc-04-1611071-g0001.tif"/>
</fig>
<p>Although the lower Huaura Valley has been the object of archaeological prospections since the early 20th century (Engel, <xref ref-type="bibr" rid="B22">1987</xref>; Tello, <xref ref-type="bibr" rid="B77">2004</xref>), Vichama was not systematically studied until the 1970s (Aguilar, <xref ref-type="bibr" rid="B1">2006</xref>; C&#x000E1;rdenas, <xref ref-type="bibr" rid="B8">1977</xref>). Since 2007, Vichama has been intensively excavated and conserved by the Caral Project (Zona Arqueol&#x000F3;gica Caral-ZAC), sponsored by the Peruvian government and local institutions (Shady et al., <xref ref-type="bibr" rid="B71">2008</xref>, <xref ref-type="bibr" rid="B73">2015</xref>).</p>
<p>Archaeological evidence and radiocarbon dates indicate the site was occupied during two chrono-cultural periods: during the Early Formative&#x02212;1 (EF-1, 1800&#x02013;1500 <sc>bce</sc>), when monumental architecture flourished through six architectonic periods (P1&#x02013;P6), and during the Late Intermediate Period (LIP, 1000&#x02013;1300 <sc>ce</sc>), when the site became a fishing village with decreased political power in this section of the valley during a period of high marine productivity (<xref ref-type="fig" rid="F2">Figure 2</xref>).</p>
<fig id="F2" position="float">
<label>Figure 2</label>
<caption><p><bold>(a)</bold> Radiocarbon dates of Vichama and proxies for export productivity in the Peruvian coast during the Early Formative&#x02212;1 (EF-1) and Late Intermediate (LIP) periods. See also the chronology of two sites mentioned in the article (i.e., &#x000C1;spero and Quebrada Chupacigarro Cemetery&#x02014;QCC). <bold>(b)</bold> Export productivity indexes: records of bromine-to-titanium ratio (Br/Ti, core M135-004-3, latitude 17&#x000B0;S) and authigenic nickel enrichment factor (EFactor; core G-10-GC-02, latitude 14&#x000B0;S). The bromine-to-titanium ratio is a semiquantitative method to estimate sedimentary total organic carbon, and the nickel (EFactor) indicates the organic sinking flux (Salvatteci et al., <xref ref-type="bibr" rid="B58">2016</xref>). Colored areas represent the compared periods&#x00027; classification (red for EF-1 and gray for LIP). Raw data available at: <ext-link ext-link-type="uri" xlink:href="https://doi.pangaea.de/10.1594/PANGAEA.887109">https://doi.pangaea.de/10.1594/PANGAEA.887109</ext-link>.</p></caption>
<alt-text>Graphical data visualization includes a timeline from 4000 BCE to 1500 CE. Part (a) lists dated materials like bone collagen, cotton textile, and plant fiber next to calibration marks for sites like &#x000C1;spero and Vichama. Part (b) depicts graphs showing export production variations with Br/Ti and Ni Enrichment Factor levels, indicating higher or lower values over time.</alt-text>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fearc-04-1611071-g0002.tif"/>
</fig>
<p>Vichama is a multicomponent complex that comprises nine sectors, including Formative monumental architecture (Vi-01) and shell middens (Vi-02, Vi-03), and architecture and cemeteries of later periods (Vi-04&#x02013;Vi-09) scattered across 136 ha in the hills of Cerro Halconcillo, a rocky elevation that rises 114 m above the sea level in the middle of one desertic oasis of 305 ha (<xref ref-type="fig" rid="F3">Figure 3a</xref>).</p>
<fig id="F3" position="float">
<label>Figure 3</label>
<caption><p><bold>(a)</bold> Sectors of Vichama archaeological site. <bold>(b)</bold> Three-dimensional reconstruction of some relevant monumental buildings in the upper and lower halves of the site. Upper half: <italic>Edificio P&#x000FA;blico Mayor</italic> (Sector D1), <italic>Edificio Las Ofrendas</italic> (Sector F), <italic>Edificio P&#x000FA;blico Residencial</italic> (Sector E1), <italic>Edificio P&#x000FA;blico</italic> G1, <italic>Edificio P&#x000FA;blico Mediano</italic> (Sector H), <italic>Edificio P&#x000FA;blico Menor</italic> (Sector I1). Lower half: <italic>Edificio Principal</italic> (Sector A1), <italic>Edificio Residencial de Elite</italic> (Sector B1)<italic>, Edificio P&#x000FA;blico Los Dep&#x000F3;sitos</italic> (Sector K), <italic>Edificio de las Chakanas</italic> (Sector C1), <italic>Edificio P&#x000FA;blico Menor</italic> (Sector L), <italic>Edificio Residencial de Elite</italic> (Sector J), <italic>Sector Residencial</italic> (Sector M-N-O), domestic and burial areas (VI-02). See <xref ref-type="supplementary-material" rid="SM2">Supplementary material 2</xref> for detailed locations of funerary contexts and individuals evaluated in this work.</p></caption>
<alt-text>Map and 3D model of an archaeological site. The map (a) shows color-coded zones indicating stone architecture, cemeteries, ash, and shell areas, with a legend on the right. The 3D model (b) illustrates labeled structures, each represented as a cluster of buildings on a hilly terrain with a red line marking a boundary between upper and lower sectors. Both images are north-oriented and display geographic coordinates.</alt-text>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fearc-04-1611071-g0003.tif"/>
</fig>
<p>During the EF-1, Vichama was part of an emerging complex polity in the lower Huaura Valley, which would have been composed of no &#x0003C; 20 settlements with monumental architecture (Aguilar, <xref ref-type="bibr" rid="B1">2006</xref>; C&#x000E1;rdenas, <xref ref-type="bibr" rid="B8">1977</xref>). It is possible that Vichama represented the rise of a new sociopolitical center in the region following the decline of the Caral society. The nuclear area at that time comprised 29 buildings, including public (e.g., stepped platforms, sunken circular courts, squares, and walls with reliefs), and domestic architecture of orthogonal design spread across 25 ha (Shady et al., <xref ref-type="bibr" rid="B73">2015</xref>). This area is divided into two halves, upper and lower, following the bipartite spatial division common in Andean settlements (<xref ref-type="fig" rid="F3">Figure 3b</xref>). The ceremonial buildings exhibit various architectural features, including stone walls arranged in horizontal rows, architectural fills using mega-<italic>shicras</italic>, and walls adorned with high-relief decorations (Shady et al., <xref ref-type="bibr" rid="B71">2008</xref>, <xref ref-type="bibr" rid="B73">2015</xref>). Changes in ceremonial architecture and paraphernalia (i.e., polychrome sculptures of unfired clay and particular iconography) suggest ideological transformations.</p>
</sec>
<sec>
<title>The diet in Vichama, current evidence</title>
<p>The &#x0201C;visual basin&#x0201D; of Vichama encompasses an approximate area of 9.2 km<sup>2</sup> that included diverse ecosystems that possibly were exploited for subsistence, among them, rocky and sandy beaches, swamps, <italic>lomas</italic> (fog oases), farming fields, the river, and the sea. The coast of Huaura Valley is one of the richest areas of the Peruvian sea, which currently cumulate &#x0007E;20% of the total national production of fisheries (IMARPE, <xref ref-type="bibr" rid="B35">2010</xref>, p. 64). Islands and islets (e.g., Anat, Azakan) located &#x0003C; 1 km from the shore, potentially provided dense clam banks, marine turtles, and a variety of seabirds.</p>
<p>The importance of fishing is undeniable. The zooarchaeological record includes a total of 91 species (23 fish, 61 mollusks, 3 marine mammals, and 4 marine birds), recovered from primary contexts (see <xref ref-type="supplementary-material" rid="SM1">Supplementary material 1</xref>: S1&#x02014;Marine species).</p>
<p>In addition, the exploitation of coastal salt-banks (i.e., Punta de Atahuanca and Salinas de Huacho, among others) to produce dry-salted fish probably was an important economic activity in the region during the Initial Formative and EF periods (Engel, <xref ref-type="bibr" rid="B22">1987</xref>; Tello, <xref ref-type="bibr" rid="B77">2004</xref>). Modern fishermen of V&#x000E9;gueta employ several traditional techniques such as <italic>redes de cerco</italic>, diving, <italic>chinchorro</italic>, and <italic>arte de pinta</italic> that could be used in the past to capture the wide array of reported species (IMARPE, <xref ref-type="bibr" rid="B35">2010</xref>, p. 67).</p>
<p>According to the last report (Yseki et al., <xref ref-type="bibr" rid="B84">2024</xref>), the archaeobotanical inventory of edible plants in Vichama has yielded more than 35,000 macro-remains representing 17 taxa derived from 123 primary contexts (see <xref ref-type="supplementary-material" rid="SM1">Supplementary material 1</xref>: S1&#x02014;Edible plants species). The botanical record includes fruits (guava, lucuma, pacay, avocado and <italic>ciruela del fraile</italic>), annuals like chili peppers (e.g., <italic>Capsicum baccatum</italic>), beans (i.e., lima bean, common bean, and jack bean), squashes, and roots and tubers (i.e., achira, sweet potato, and potato). Based on their ubiquity, it seems that fruit trees, especially guava and lucuma, as well as chili peppers, were regularly consumed and used in ceremonial activities. Similarly, the high ubiquity in offering contexts of remains of lima bean, achira, sweet potato, and potato highlights the economic importance of these plants in Vichama (Yseki et al., <xref ref-type="bibr" rid="B84">2024</xref>).</p>
<p>The absence of <italic>Zea mays</italic> (maize) macro-remains suggests that it was probably not grown, processed, or stored at Vichama. However, a recent study reported starch grains with potential fermentation-related damage in dental calculus of individuals from &#x000C1;spero (Yseki et al., <xref ref-type="bibr" rid="B85">2023</xref>), with a minimal presence of maize macro-remains (Shady and C&#x000E1;ceda, <xref ref-type="bibr" rid="B70">2008</xref>). Thus, maize consumption at Vichama cannot be ruled out based solely on macro-remains analysis, as maize may have been consumed as a beverage or flour (Yseki et al., <xref ref-type="bibr" rid="B84">2024</xref>).</p>
<p>The marine fauna and some differences between periods suggest fluctuations in sea productive conditions linked to the upwelling of rich-nutrient marine waters. For instance, during the EF period (1800&#x02013;1200 <sc>bce</sc>), there is a general trend to increasing exported productivity, especially during the EF-1 (1800&#x02013;1500 <sc>bce</sc>), with some fluctuations (i.e., lower productivity between 1800 and 1600 <sc>bce</sc>, with the lowest values around 1700 <sc>bce</sc> and higher productivity between 1600 and 1500 <sc>bce</sc>) that should have influenced dietary composition, possibly with higher consumption of marine products; however, the plant record shows that agriculture was ongoing during the flourishment of monumental architecture.</p>
<p>These conditions, however, could be relatively different from those observed during the LIP (1000&#x02013;1300 <sc>ce</sc>) when productivity reached even higher values and enhanced upwelling conditions, with its maximum at &#x0007E;1000&#x02013;1100 <sc>ce</sc> (Kennett and Marwan, <xref ref-type="bibr" rid="B39">2015</xref>). During the LIP, the archaeological deposits are plenty of marine remains, such as fish bones and shells, fishing nets, and artifacts for marine exploitation.</p>
</sec>
<sec>
<title>Stable isotopes in diet reconstruction</title>
<p>While faunal and plant remains can provide insight into shifts in economic practices, stable isotope analysis is more effective for targeting the net consumption of specific food resources at an individual level. Stable carbon and nitrogen isotope analyses of collagen (&#x003B4;<sup>13</sup>C<sub>coll</sub> and &#x003B4;<sup>15</sup>N) and bioapatite (&#x003B4;<sup>13</sup>C<sub>ap</sub>) extracted from human bones and teeth offer valuable insights into the diets of ancient populations (Schoeninger and Reitsema, <xref ref-type="bibr" rid="B64">2023</xref>). The &#x003B4;<sup>13</sup>C<sub>coll</sub> is a proxy of protein sources and trophic position of consumers in the food web, as well as the photosynthetic pathway of the plants they consumed. The &#x003B4;<sup>13</sup>C<sub>ap</sub> instead reflects the entire dietary carbohydrates, lipids, and proteins, providing a useful means to assess the energetic constituents of the diet and photosynthetic pathways (Jim et al., <xref ref-type="bibr" rid="B37">2004</xref>; Kellner and Schoeninger, <xref ref-type="bibr" rid="B38">2007</xref>). In the Andean central coast, C<sub>3</sub> plants, which use the Calvin&#x02013;Benson photosynthetic pathway, are the most prevalent and typically exhibit mean &#x003B4;<sup>13</sup>C values of &#x02212;25.7 &#x000B1; 2.1&#x02030;. In contrast, among C<sub>4</sub>, which follow the Hatch&#x02013;Slack photosynthetic pathway, there are only two potentially edible domesticated plants (<italic>kiwicha</italic>&#x02014;<italic>Amarantus</italic> sp., and maize&#x02014;<italic>Zea mays</italic>), with typical values of approximately &#x02212;11.3 &#x000B1; 1.4&#x02030;. Additionally, crassulacean acid metabolism (CAM) plants have isotopic values (&#x02212;10.8 &#x000B1; 0.2&#x02030;) that overlap with those of C<sub>4</sub> (Pezo-Lanfranco and Colonese, <xref ref-type="bibr" rid="B51">2024</xref>).</p>
<p>Usually, C<sub>4</sub> plant consumers show significantly higher &#x003B4;<sup>13</sup>C values (Kellner and Schoeninger, <xref ref-type="bibr" rid="B38">2007</xref>). In coastal areas, however, using &#x003B4;<sup>13</sup>C to trace maize consumption can be challenging due to the presence of marine resources (i.e., fish and mollusks), which have overlapping &#x003B4;<sup>13</sup>C values to C<sub>4</sub> plants, approximately &#x02212;11.9 &#x000B1; 2.1&#x02030; (Pezo-Lanfranco and Colonese, <xref ref-type="bibr" rid="B51">2024</xref>). In terms of &#x003B4;<sup>15</sup>N values, herbivores tend to have values 3&#x02013;7&#x02030; higher than the plants they consume, while carnivores exhibit &#x003B4;<sup>15</sup>N values 3&#x02013;5&#x02030; higher than their prey. Because marine sources have higher isotopic values compared to terrestrial sources (Minagawa and Wada, <xref ref-type="bibr" rid="B48">1984</xref>; Schoeninger and DeNiro, <xref ref-type="bibr" rid="B62">1984</xref>), marine fauna displays higher &#x003B4;<sup>15</sup>N values than terrestrial ones, linked to trophic fractionation within complex marine food webs (Chisholm et al., <xref ref-type="bibr" rid="B11">1982</xref>; DeNiro and Epstein, <xref ref-type="bibr" rid="B17">1981</xref>; Hedges and Reynard, <xref ref-type="bibr" rid="B32">2007</xref>; Schoeninger, <xref ref-type="bibr" rid="B61">2010</xref>; Schoeninger et al., <xref ref-type="bibr" rid="B63">1983</xref>). Stable isotope analysis is particularly informative when combined with Bayesian mixing models for deriving quantitative estimates of dietary composition as probability distributions (Cheung and Szpak, <xref ref-type="bibr" rid="B10">2021</xref>; Fernandes et al., <xref ref-type="bibr" rid="B24">2014</xref>, <xref ref-type="bibr" rid="B23">2015</xref>; Phillips, <xref ref-type="bibr" rid="B55">2012</xref>).</p>
</sec>
<sec>
<title>Hypothesis and expectations</title>
<p>By employing a combination of stable isotope proxies from collagen (&#x003B4;<sup>13</sup>C<sub>coll</sub>, &#x003B4;<sup>15</sup>N<sub>coll</sub>, <italic>n</italic> = 36) and apatite (&#x003B4;<sup>13</sup>C<sub>ap</sub>, <italic>n</italic> = 26) of 38 human individuals from Vichama, and BSIMM estimations of the caloric contribution of the potential food sources (i.e., marine and terrestrial fauna, as well as C<sub>3</sub> and C<sub>4</sub> plants; Fernandes et al., <xref ref-type="bibr" rid="B24">2014</xref>, <xref ref-type="bibr" rid="B23">2015</xref>) consumed by 21 Vichama individuals (see Material and Methods for model implementation details), we aim to elucidate the primary sources of energy for two occupational periods at Vichama: EF-1 (1800&#x02013;1500 <sc>bce</sc>) and the LIP (1000&#x02013;1300 <sc>ce</sc>).</p>
<p>We tested the hypothesis that changes in marine productivity influenced the degree to which marine resources were incorporated into Andean subsistence economies (Kennett and Marwan, <xref ref-type="bibr" rid="B39">2015</xref>; Wilson et al., <xref ref-type="bibr" rid="B83">2024</xref>). According to paleoclimatic data, an increased marine productivity between 1800 and 1500 <sc>bce</sc> was concomitant with the flourishing of Vichama (Shady et al., <xref ref-type="bibr" rid="B73">2015</xref>). However, the economic importance of plants and fish and their relative contribution to diet is unknown. It is expected that the inhabitants of Vichama, particularly during EF-1, relied more strongly on marine products as a significant part of their diet.</p>
<p>For comparison, the LIP (1000&#x02013;1300 <sc>ce</sc>) was a time of climate cooling and decreasing SSTs, when marine productivity reached its peak (Kennett and Marwan, <xref ref-type="bibr" rid="B39">2015</xref>). Because the LIP has been recognized as the period of higher development of agriculture in the Central Andes, under increasing conditions of upwelling and marine productivity, a mixed diet with a higher contribution of marine sources is expected. This reliance should be reflected in paleodietary reconstructions.</p>
</sec>
</sec>
<sec sec-type="results" id="s2">
<title>Results</title>
<sec>
<title>Dietary reconstruction</title>
<p>The isotopic values of 38 human individuals from Vichama were classified into the two occupations, EF period (1800&#x02013;1500 <sc>bce</sc>, <italic>n</italic> = 30), and the LIP (1500&#x02013;1300 <sc>ce</sc>, <italic>n</italic> = 8), used here for comparative purposes (<xref ref-type="table" rid="T1">Table 1</xref>, <xref ref-type="supplementary-material" rid="SM1">Supplementary material 1</xref>: S2 for individuals&#x00027; contexts and bioarchaeological details and S3 for radiocarbon dates). Only a fraction of the 59 elements sampled (36 bones and 23 teeth) provided intact collagen with reliable &#x003B4;<sup>13</sup>C<sub>coll</sub> and &#x003B4;<sup>15</sup>N values from bone (<italic>n</italic> = 21) or teeth (<italic>n</italic> = 15) according to accepted preservation criteria (DeNiro, <xref ref-type="bibr" rid="B16">1985</xref>; Van Klinken, <xref ref-type="bibr" rid="B79">1999</xref>). Attenuated total reflectance&#x02013;Fourier transform infrared spectroscopy (ATR-FTIR) studies conducted in a subsample of 10 individuals consistently suggest that no diagenetic alteration affected the inorganic fraction of bones from Vichama contexts (France et al., <xref ref-type="bibr" rid="B26">2020</xref>; see <xref ref-type="supplementary-material" rid="SM1">Supplementary material 1</xref>: S4 for the ATR-FTIR data). Thus, &#x003B4;<sup>13</sup>C<sub>ap</sub> values (<italic>n</italic> = 26) were included in our analysis to complement the paleodietary information.</p>
<table-wrap position="float" id="T1">
<label>Table 1</label>
<caption><p>Isotopic values from Vichama individuals.</p></caption>
<table frame="box" rules="all">
<thead>
<tr style="background-color:#8f9496;color:#ffffff">
<th valign="top" align="left"><bold>&#x00023;</bold></th>
<th valign="top" align="left"><bold>Period</bold></th>
<th valign="top" align="left"><bold>ID</bold></th>
<th valign="top" align="left"><bold>Sex</bold></th>
<th valign="top" align="left"><bold>Age-range (years)</bold></th>
<th valign="top" align="left"><bold>Tissue</bold></th>
<th valign="top" align="left"><bold>Element (segment)</bold></th>
<th valign="top" align="left"><bold>Age of tissue<sup>&#x0002A;</sup></bold></th>
<th valign="top" align="left"><bold>%C</bold></th>
<th valign="top" align="left"><bold>%N</bold></th>
<th valign="top" align="left"><bold>C:N</bold></th>
<th valign="top" align="left"><bold>&#x003B4;<sup>13</sup>C<sub>coll</sub></bold></th>
<th valign="top" align="left"><bold>&#x003B4;<sup>15</sup>N</bold></th>
<th valign="top" align="left"><bold>&#x003B4;<sup>13</sup>C<sub>ap</sub></bold></th>
</tr>
</thead>
<tbody>
<tr>
<td valign="top" align="left">1</td>
<td valign="top" align="left">EF-1 (P3)</td>
<td valign="top" align="left">ZAC 0458<sup>&#x0002A;&#x0002A;</sup></td>
<td valign="top" align="left">F</td>
<td valign="top" align="left">Adult</td>
<td valign="top" align="left">Bone</td>
<td valign="top" align="left">Rib</td>
<td valign="top" align="left">Adult</td>
<td valign="top" align="left">41.9</td>
<td valign="top" align="left">13.9</td>
<td valign="top" align="left">3.5</td>
<td valign="top" align="left">&#x02212;16.9</td>
<td valign="top" align="left">13.4</td>
<td valign="top" align="left">&#x02212;13.8</td>
</tr> <tr>
<td valign="top" align="left">2</td>
<td valign="top" align="left">EF-1 (P3)</td>
<td valign="top" align="left">ZAC 0458</td>
<td valign="top" align="left">F</td>
<td valign="top" align="left">Adult</td>
<td valign="top" align="left">Tooth</td>
<td valign="top" align="left">LLPm (root)</td>
<td valign="top" align="left">5.5&#x02013;14 yr</td>
<td valign="top" align="left">39.5</td>
<td valign="top" align="left">14.0</td>
<td valign="top" align="left">3.3</td>
<td valign="top" align="left">&#x02212;17.8</td>
<td valign="top" align="left">11.8</td>
<td valign="top" align="left">&#x02212;11.9</td>
</tr> <tr>
<td valign="top" align="left">3</td>
<td valign="top" align="left">EF-1 (P3)</td>
<td valign="top" align="left">ZAC 7058</td>
<td valign="top" align="left">M</td>
<td valign="top" align="left">30&#x02013;65 yr</td>
<td valign="top" align="left">Bone</td>
<td valign="top" align="left">Rib</td>
<td valign="top" align="left">30&#x02013;65 yr</td>
<td valign="top" align="left">No collagen</td>
<td/>
<td/>
<td/>
<td/>
<td valign="top" align="left">&#x02212;14.1</td>
</tr> <tr>
<td valign="top" align="left">4</td>
<td valign="top" align="left">EF-1 (P4)</td>
<td valign="top" align="left">ZAC 7069<sup>&#x0002A;&#x0002A;</sup></td>
<td valign="top" align="left">M</td>
<td valign="top" align="left">22&#x02013;32 yr</td>
<td valign="top" align="left">Bone</td>
<td valign="top" align="left">Rib</td>
<td valign="top" align="left">22&#x02013;32 yr</td>
<td valign="top" align="left">42.8</td>
<td valign="top" align="left">15.1</td>
<td valign="top" align="left">3.3</td>
<td valign="top" align="left">&#x02212;16.6</td>
<td valign="top" align="left">13</td>
<td valign="top" align="left">&#x02212;13.2</td>
</tr> <tr>
<td valign="top" align="left">5</td>
<td valign="top" align="left">EF-1 (P4)</td>
<td valign="top" align="left">ZAC 7069</td>
<td valign="top" align="left">M</td>
<td valign="top" align="left">22&#x02013;32 yr</td>
<td valign="top" align="left">Tooth</td>
<td valign="top" align="left">1,8 (root)</td>
<td valign="top" align="left">8.0&#x02013;16.5 yr</td>
<td valign="top" align="left">45.7</td>
<td valign="top" align="left">15.5</td>
<td valign="top" align="left">3.4</td>
<td valign="top" align="left">&#x02212;18</td>
<td valign="top" align="left">11.6</td>
<td valign="top" align="left">n.a.</td>
</tr> <tr>
<td valign="top" align="left">6</td>
<td valign="top" align="left">EF-1 (P4)</td>
<td valign="top" align="left">ZAC 7075</td>
<td valign="top" align="left">Und</td>
<td valign="top" align="left">4 yr &#x000B1; 12 m</td>
<td valign="top" align="left">Bone</td>
<td valign="top" align="left">Rib</td>
<td valign="top" align="left">4 yr &#x000B1; 12 m</td>
<td valign="top" align="left">No collagen</td>
<td/>
<td/>
<td/>
<td/>
<td valign="top" align="left">n.a.</td>
</tr> <tr>
<td valign="top" align="left">7</td>
<td valign="top" align="left">EF-1 (P4)</td>
<td valign="top" align="left">ZAC 7075</td>
<td valign="top" align="left">Und</td>
<td valign="top" align="left">4 yr &#x000B1; 12 m</td>
<td valign="top" align="left">Tooth</td>
<td valign="top" align="left">7,3 (root)</td>
<td valign="top" align="left">0.9&#x02013;3.5 yr</td>
<td valign="top" align="left">No collagen</td>
<td/>
<td/>
<td/>
<td/>
<td valign="top" align="left">n.a.</td>
</tr> <tr>
<td valign="top" align="left">8</td>
<td valign="top" align="left">EF-1 (P4)</td>
<td valign="top" align="left">ZAC 5905<sup>&#x0002A;&#x0002A;</sup></td>
<td valign="top" align="left">M</td>
<td valign="top" align="left">25&#x02013;35 yr</td>
<td valign="top" align="left">Bone</td>
<td valign="top" align="left">Rib</td>
<td valign="top" align="left">25&#x02013;35 yr</td>
<td valign="top" align="left">29.0</td>
<td valign="top" align="left">9.4</td>
<td valign="top" align="left">3.6</td>
<td valign="top" align="left">&#x02212;17.5</td>
<td valign="top" align="left">14.3</td>
<td valign="top" align="left">&#x02212;14.0</td>
</tr> <tr>
<td valign="top" align="left">9</td>
<td valign="top" align="left">EF-1 (P4)</td>
<td valign="top" align="left">ZAC 5905</td>
<td valign="top" align="left">M</td>
<td valign="top" align="left">25&#x02013;35 yr</td>
<td valign="top" align="left">Tooth</td>
<td valign="top" align="left">3,7 (root)</td>
<td valign="top" align="left">8&#x02013;16.5 yr</td>
<td valign="top" align="left">44.8</td>
<td valign="top" align="left">16.1</td>
<td valign="top" align="left">3.3</td>
<td valign="top" align="left">&#x02212;17.7</td>
<td valign="top" align="left">11.6</td>
<td valign="top" align="left">n.a.</td>
</tr> <tr>
<td valign="top" align="left">10</td>
<td valign="top" align="left">EF-1 (P4)</td>
<td valign="top" align="left">PEACS 4431<sup>&#x0002A;&#x0002A;</sup></td>
<td valign="top" align="left">Und</td>
<td valign="top" align="left">2&#x02013;5 yr</td>
<td valign="top" align="left">Bone</td>
<td valign="top" align="left">Rib</td>
<td valign="top" align="left">2&#x02013;5 yr</td>
<td valign="top" align="left">42.7</td>
<td valign="top" align="left">13.7</td>
<td valign="top" align="left">3.3</td>
<td valign="top" align="left">&#x02212;18</td>
<td valign="top" align="left">13.8</td>
<td valign="top" align="left">&#x02212;14.6</td>
</tr> <tr>
<td valign="top" align="left">11</td>
<td valign="top" align="left">EF-1 (P5)</td>
<td valign="top" align="left">ZAC 7056<sup>&#x0002A;&#x0002A;</sup></td>
<td valign="top" align="left">F</td>
<td valign="top" align="left">35&#x02013;65 yr</td>
<td valign="top" align="left">Bone</td>
<td valign="top" align="left">Rib</td>
<td valign="top" align="left">35&#x02013;65 yr</td>
<td valign="top" align="left">37.7</td>
<td valign="top" align="left">12.9</td>
<td valign="top" align="left">3.4</td>
<td valign="top" align="left">&#x02212;16.3</td>
<td valign="top" align="left">15.6</td>
<td valign="top" align="left">&#x02212;14.0</td>
</tr> <tr>
<td valign="top" align="left">12</td>
<td valign="top" align="left">EF-1 (P5)</td>
<td valign="top" align="left">ZAC 7057<sup>&#x0002A;&#x0002A;</sup></td>
<td valign="top" align="left">Und</td>
<td valign="top" align="left">3 yr &#x000B1; 12 m</td>
<td valign="top" align="left">Bone</td>
<td valign="top" align="left">Rib</td>
<td valign="top" align="left">3 yr &#x000B1; 12 m</td>
<td valign="top" align="left">39.5</td>
<td valign="top" align="left">13.6</td>
<td valign="top" align="left">3.4</td>
<td valign="top" align="left">&#x02212;14.1</td>
<td valign="top" align="left">18</td>
<td valign="top" align="left">&#x02212;13.6</td>
</tr> <tr>
<td valign="top" align="left">13</td>
<td valign="top" align="left">EF-1 (P5)</td>
<td valign="top" align="left">ZAC 7082</td>
<td valign="top" align="left">F</td>
<td valign="top" align="left">50&#x02013;75 yr</td>
<td valign="top" align="left">Bone</td>
<td valign="top" align="left">Rib</td>
<td valign="top" align="left">50&#x02013;75 yr</td>
<td valign="top" align="left">10.1</td>
<td valign="top" align="left">2.6</td>
<td valign="top" align="left"><underline>4.5</underline></td>
<td valign="top" align="left">&#x02212;20.5</td>
<td valign="top" align="left">16.6</td>
<td valign="top" align="left">n.a.</td>
</tr> <tr>
<td valign="top" align="left">14</td>
<td valign="top" align="left">EF-1 (P5)</td>
<td valign="top" align="left">ZAC 7083</td>
<td valign="top" align="left">M</td>
<td valign="top" align="left">35&#x02013;50 yr</td>
<td valign="top" align="left">Bone</td>
<td valign="top" align="left">Rib</td>
<td valign="top" align="left">35&#x02013;50 yr</td>
<td valign="top" align="left">No collagen</td>
<td/>
<td/>
<td/>
<td/>
<td valign="top" align="left">n.a.</td>
</tr> <tr>
<td valign="top" align="left">15</td>
<td valign="top" align="left">EF-1 (P5)</td>
<td valign="top" align="left">ZAC 7083</td>
<td valign="top" align="left">M</td>
<td valign="top" align="left">35&#x02013;50 yr</td>
<td valign="top" align="left">Tooth</td>
<td valign="top" align="left">1,8 (root)</td>
<td valign="top" align="left">8.0&#x02013;16.5 yr</td>
<td valign="top" align="left">No collagen</td>
<td/>
<td/>
<td/>
<td/>
<td valign="top" align="left">n.a.</td>
</tr> <tr>
<td valign="top" align="left">16</td>
<td valign="top" align="left">EF-1 (P5)</td>
<td valign="top" align="left">ZAC 7085</td>
<td valign="top" align="left">F</td>
<td valign="top" align="left">7 yr &#x000B1; 12 m</td>
<td valign="top" align="left">Bone</td>
<td valign="top" align="left">Rib</td>
<td valign="top" align="left">7 yr &#x000B1; 12 m</td>
<td valign="top" align="left">No collagen</td>
<td/>
<td/>
<td/>
<td/>
<td valign="top" align="left">n.a.</td>
</tr> <tr>
<td valign="top" align="left">17</td>
<td valign="top" align="left">EF-1 (P5)</td>
<td valign="top" align="left">ZAC 7085</td>
<td valign="top" align="left">F</td>
<td valign="top" align="left">7 yr &#x000B1; 12 m</td>
<td valign="top" align="left">Tooth</td>
<td valign="top" align="left">1,1 (root)</td>
<td valign="top" align="left">4.5&#x02013;11 yr</td>
<td valign="top" align="left">No collagen</td>
<td/>
<td/>
<td/>
<td/>
<td valign="top" align="left">n.a.</td>
</tr> <tr>
<td valign="top" align="left">18</td>
<td valign="top" align="left">EF-1 (P5)</td>
<td valign="top" align="left">ZAC 1924</td>
<td valign="top" align="left">Und</td>
<td valign="top" align="left">12&#x02013;16 yr</td>
<td valign="top" align="left">Bone</td>
<td valign="top" align="left">Rib</td>
<td valign="top" align="left">12&#x02013;16 yr</td>
<td valign="top" align="left">17.8</td>
<td valign="top" align="left">6.4</td>
<td valign="top" align="left">3.2</td>
<td valign="top" align="left">&#x02212;19.2</td>
<td valign="top" align="left">13.0</td>
<td valign="top" align="left">n.a.</td>
</tr> <tr>
<td valign="top" align="left">19</td>
<td valign="top" align="left">EF-1 (P5)</td>
<td valign="top" align="left">ZAC 1924</td>
<td valign="top" align="left">Und</td>
<td valign="top" align="left">12&#x02013;16 yr</td>
<td valign="top" align="left">Tooth</td>
<td valign="top" align="left">URPm (root)</td>
<td valign="top" align="left">6&#x02013;14 yr</td>
<td valign="top" align="left">45.1</td>
<td valign="top" align="left">14.2</td>
<td valign="top" align="left"><underline>3.7</underline></td>
<td valign="top" align="left">&#x02212;15.8</td>
<td valign="top" align="left">17.9</td>
<td valign="top" align="left">n.a.</td>
</tr> <tr>
<td valign="top" align="left">20</td>
<td valign="top" align="left">EF-1 (P6)</td>
<td valign="top" align="left">ZAC 7076<sup>&#x0002A;&#x0002A;</sup></td>
<td valign="top" align="left">M</td>
<td valign="top" align="left">16&#x02013;19 yr</td>
<td valign="top" align="left">Bone</td>
<td valign="top" align="left">Rib</td>
<td valign="top" align="left">16&#x02013;19 yr</td>
<td valign="top" align="left">18.0</td>
<td valign="top" align="left">5.9</td>
<td valign="top" align="left">3.6</td>
<td valign="top" align="left">&#x02212;18.5</td>
<td valign="top" align="left">14.0</td>
<td valign="top" align="left">&#x02212;14.5</td>
</tr> <tr>
<td valign="top" align="left">21</td>
<td valign="top" align="left">EF-1 (P6)</td>
<td valign="top" align="left">ZAC 7076</td>
<td valign="top" align="left">M</td>
<td valign="top" align="left">16&#x02013;19 yr</td>
<td valign="top" align="left">Tooth</td>
<td valign="top" align="left">M3 (root)</td>
<td valign="top" align="left">14.0&#x02013;23.5 yr</td>
<td valign="top" align="left">43.5</td>
<td valign="top" align="left">15.3</td>
<td valign="top" align="left">3.3</td>
<td valign="top" align="left">&#x02212;17.9</td>
<td valign="top" align="left">11.5</td>
<td valign="top" align="left">n.a.</td>
</tr> <tr>
<td valign="top" align="left">22</td>
<td valign="top" align="left">EF-1</td>
<td valign="top" align="left">ZAC 7084</td>
<td valign="top" align="left">F</td>
<td valign="top" align="left">10 yr &#x000B1; 30 m</td>
<td valign="top" align="left">Bone</td>
<td valign="top" align="left">Rib</td>
<td valign="top" align="left">10 yr &#x000B1; 30 m</td>
<td valign="top" align="left">12.7</td>
<td valign="top" align="left">3.0</td>
<td valign="top" align="left"><underline>5.0</underline></td>
<td valign="top" align="left">&#x02212;20.9</td>
<td valign="top" align="left">14.6</td>
<td valign="top" align="left">n.a.</td>
</tr> <tr>
<td valign="top" align="left">23</td>
<td valign="top" align="left">EF-1</td>
<td valign="top" align="left">ZAC 7084</td>
<td valign="top" align="left">F</td>
<td valign="top" align="left">10 yr &#x000B1; 30 m</td>
<td valign="top" align="left">Tooth</td>
<td valign="top" align="left">3,3 (root)</td>
<td valign="top" align="left">5.5&#x02013;14.5 yr</td>
<td valign="top" align="left">31.1</td>
<td valign="top" align="left">7.4</td>
<td valign="top" align="left"><underline>4.9</underline></td>
<td valign="top" align="left">&#x02212;17.6</td>
<td valign="top" align="left">17.2</td>
<td valign="top" align="left">n.a.</td>
</tr> <tr>
<td valign="top" align="left">24</td>
<td valign="top" align="left">EF-1</td>
<td valign="top" align="left">ZAC 6645</td>
<td valign="top" align="left">F</td>
<td valign="top" align="left">30&#x02013;65 yr</td>
<td valign="top" align="left">Bone</td>
<td valign="top" align="left">Rib</td>
<td valign="top" align="left">30&#x02013;65 yr</td>
<td valign="top" align="left">No collagen</td>
<td/>
<td/>
<td/>
<td/>
<td valign="top" align="left">n.a.</td>
</tr> <tr>
<td valign="top" align="left">25</td>
<td valign="top" align="left">EF-1</td>
<td valign="top" align="left">ZAC 7062<sup>&#x0002A;&#x0002A;</sup></td>
<td valign="top" align="left">M</td>
<td valign="top" align="left">35&#x02013;65 yr</td>
<td valign="top" align="left">Bone</td>
<td valign="top" align="left">Rib</td>
<td valign="top" align="left">35&#x02013;65 yr</td>
<td valign="top" align="left">41.9</td>
<td valign="top" align="left">14.9</td>
<td valign="top" align="left">3.3</td>
<td valign="top" align="left">&#x02212;17.6</td>
<td valign="top" align="left">11.4</td>
<td valign="top" align="left">&#x02212;11.3</td>
</tr> <tr>
<td valign="top" align="left">26</td>
<td valign="top" align="left">EF-1</td>
<td valign="top" align="left">ZAC 7062</td>
<td valign="top" align="left">M</td>
<td valign="top" align="left">35&#x02013;65 yr</td>
<td valign="top" align="left">Tooth</td>
<td valign="top" align="left">2,8 (root)</td>
<td valign="top" align="left">8.0&#x02013;16.5 yr</td>
<td valign="top" align="left">42.0</td>
<td valign="top" align="left">14.5</td>
<td valign="top" align="left">3.4</td>
<td valign="top" align="left">&#x02212;18.4</td>
<td valign="top" align="left">11.1</td>
<td valign="top" align="left">n.a.</td>
</tr> <tr>
<td valign="top" align="left">27</td>
<td valign="top" align="left">EF-1</td>
<td valign="top" align="left">ZAC 7064</td>
<td valign="top" align="left">M</td>
<td valign="top" align="left">22&#x02013;35 yr</td>
<td valign="top" align="left">Bone</td>
<td valign="top" align="left">Rib</td>
<td valign="top" align="left">22&#x02013;35 yr</td>
<td valign="top" align="left">No collagen</td>
<td/>
<td/>
<td/>
<td/>
<td valign="top" align="left">&#x02212;13.0</td>
</tr> <tr>
<td valign="top" align="left">28</td>
<td valign="top" align="left">EF-1</td>
<td valign="top" align="left">ZAC 7064</td>
<td valign="top" align="left">M</td>
<td valign="top" align="left">22&#x02013;35 yr</td>
<td valign="top" align="left">Tooth</td>
<td valign="top" align="left">4,2 (root)</td>
<td valign="top" align="left">4&#x02013;8.5 yr</td>
<td valign="top" align="left">46.0</td>
<td valign="top" align="left">16.0</td>
<td valign="top" align="left">3.4</td>
<td valign="top" align="left">&#x02212;19.4</td>
<td valign="top" align="left">8.2</td>
<td valign="top" align="left">n.a.</td>
</tr> <tr>
<td valign="top" align="left">29</td>
<td valign="top" align="left">EF-1</td>
<td valign="top" align="left">ZAC 7065<sup>&#x0002A;&#x0002A;</sup></td>
<td valign="top" align="left">M</td>
<td valign="top" align="left">20&#x02013;25 yr</td>
<td valign="top" align="left">Bone</td>
<td valign="top" align="left">Rib</td>
<td valign="top" align="left">20&#x02013;25 yr</td>
<td valign="top" align="left">15.2</td>
<td valign="top" align="left">43.8</td>
<td valign="top" align="left">3.4</td>
<td valign="top" align="left">&#x02212;16.0</td>
<td valign="top" align="left">13.9</td>
<td valign="top" align="left">&#x02212;14.0</td>
</tr> <tr>
<td valign="top" align="left">30</td>
<td valign="top" align="left">EF-1</td>
<td valign="top" align="left">ZAC 7065</td>
<td valign="top" align="left">M</td>
<td valign="top" align="left">20&#x02013;25 yr</td>
<td valign="top" align="left">Tooth</td>
<td valign="top" align="left">4,1 (root)</td>
<td valign="top" align="left">4&#x02013;8.5 yr</td>
<td valign="top" align="left">45.2</td>
<td valign="top" align="left">16.2</td>
<td valign="top" align="left">3.3</td>
<td valign="top" align="left">&#x02212;18.2</td>
<td valign="top" align="left">12.1</td>
<td valign="top" align="left">n.a.</td>
</tr> <tr>
<td valign="top" align="left">31</td>
<td valign="top" align="left">EF-1</td>
<td valign="top" align="left">ZAC 7066<sup>&#x0002A;&#x0002A;</sup></td>
<td valign="top" align="left">M</td>
<td valign="top" align="left">30&#x02013;40 yr</td>
<td valign="top" align="left">Bone</td>
<td valign="top" align="left">Rib</td>
<td valign="top" align="left">30&#x02013;40 yr</td>
<td valign="top" align="left">12.5</td>
<td valign="top" align="left">4.3</td>
<td valign="top" align="left">3.4</td>
<td valign="top" align="left">&#x02212;20.3</td>
<td valign="top" align="left">12.0</td>
<td valign="top" align="left">&#x02212;11.9</td>
</tr> <tr>
<td valign="top" align="left">32</td>
<td valign="top" align="left">EF-1</td>
<td valign="top" align="left">ZAC 7067<sup>&#x0002A;&#x0002A;</sup></td>
<td valign="top" align="left">Und</td>
<td valign="top" align="left">8 yr &#x000B1; 24 m</td>
<td valign="top" align="left">Bone</td>
<td valign="top" align="left">Rib</td>
<td valign="top" align="left">8 yr &#x000B1; 24 m</td>
<td valign="top" align="left">42.7</td>
<td valign="top" align="left">14.9</td>
<td valign="top" align="left">3.3</td>
<td valign="top" align="left">&#x02212;17.8</td>
<td valign="top" align="left">11.8</td>
<td valign="top" align="left">&#x02212;12.8</td>
</tr> <tr>
<td valign="top" align="left">33</td>
<td valign="top" align="left">EF-1</td>
<td valign="top" align="left">ZAC 7067</td>
<td valign="top" align="left">Und</td>
<td valign="top" align="left">8 yr &#x000B1; 24 m</td>
<td valign="top" align="left">Tooth</td>
<td valign="top" align="left">3 fragm. (roots)</td>
<td valign="top" align="left">0.6&#x02013;3.5 yr</td>
<td valign="top" align="left">46.5</td>
<td valign="top" align="left">14.2</td>
<td valign="top" align="left"><underline>3.8</underline></td>
<td valign="top" align="left">&#x02212;18.8</td>
<td valign="top" align="left">14.0</td>
<td valign="top" align="left">n.a.</td>
</tr> <tr>
<td valign="top" align="left">34</td>
<td valign="top" align="left">EF-1</td>
<td valign="top" align="left">ZAC 7068<sup>&#x0002A;&#x0002A;</sup></td>
<td valign="top" align="left">F</td>
<td valign="top" align="left">25&#x02013;40 yr</td>
<td valign="top" align="left">Bone</td>
<td valign="top" align="left">Rib</td>
<td valign="top" align="left">25&#x02013;40 yr</td>
<td valign="top" align="left">42.4</td>
<td valign="top" align="left">14.7</td>
<td valign="top" align="left">3.4</td>
<td valign="top" align="left">&#x02212;16.9</td>
<td valign="top" align="left">13.1</td>
<td valign="top" align="left">&#x02212;11.8</td>
</tr> <tr>
<td valign="top" align="left">35</td>
<td valign="top" align="left">EF-1</td>
<td valign="top" align="left">ZAC 7068</td>
<td valign="top" align="left">F</td>
<td valign="top" align="left">25&#x02013;40 yr</td>
<td valign="top" align="left">Tooth</td>
<td valign="top" align="left">3,8 (root)</td>
<td valign="top" align="left">14.0&#x02013;23.5 yr</td>
<td valign="top" align="left">45.7</td>
<td valign="top" align="left">15.5</td>
<td valign="top" align="left">3.4</td>
<td valign="top" align="left">&#x02212;18.5</td>
<td valign="top" align="left">11.3</td>
<td valign="top" align="left">n.a.</td>
</tr> <tr>
<td valign="top" align="left">36</td>
<td valign="top" align="left">EF-1</td>
<td valign="top" align="left">ZAC 7070<sup>&#x0002A;&#x0002A;</sup></td>
<td valign="top" align="left">F</td>
<td valign="top" align="left">20&#x02013;30 yr</td>
<td valign="top" align="left">Bone</td>
<td valign="top" align="left">Rib</td>
<td valign="top" align="left">20&#x02013;30 yr</td>
<td valign="top" align="left">42.2</td>
<td valign="top" align="left">14.5</td>
<td valign="top" align="left">3.4</td>
<td valign="top" align="left">&#x02212;14.6</td>
<td valign="top" align="left">12.9</td>
<td valign="top" align="left">&#x02212;11.5</td>
</tr> <tr>
<td valign="top" align="left">37</td>
<td valign="top" align="left">EF-1</td>
<td valign="top" align="left">ZAC 7071<sup>&#x0002A;&#x0002A;</sup></td>
<td valign="top" align="left">M</td>
<td valign="top" align="left">25&#x02013;55 yr</td>
<td valign="top" align="left">Bone</td>
<td valign="top" align="left">Rib</td>
<td valign="top" align="left">25&#x02013;55 yr</td>
<td valign="top" align="left">43.2</td>
<td valign="top" align="left">15.1</td>
<td valign="top" align="left">3.3</td>
<td valign="top" align="left">&#x02212;14.9</td>
<td valign="top" align="left">12.1</td>
<td valign="top" align="left">&#x02212;11.2</td>
</tr> <tr>
<td valign="top" align="left">38</td>
<td valign="top" align="left">EF-1</td>
<td valign="top" align="left">ZAC 7071</td>
<td valign="top" align="left">M</td>
<td valign="top" align="left">25&#x02013;55 yr</td>
<td valign="top" align="left">Tooth</td>
<td valign="top" align="left">Pm (root)</td>
<td valign="top" align="left">5.5&#x02013;15 yr</td>
<td valign="top" align="left">75.3</td>
<td valign="top" align="left">25.2</td>
<td valign="top" align="left">3.5</td>
<td valign="top" align="left">&#x02212;15.0</td>
<td valign="top" align="left">11.6</td>
<td valign="top" align="left">n.a.</td>
</tr> <tr>
<td valign="top" align="left">39</td>
<td valign="top" align="left">EF-1</td>
<td valign="top" align="left">ZAC 7072</td>
<td valign="top" align="left">Und</td>
<td valign="top" align="left">2 yr &#x000B1; 8 m</td>
<td valign="top" align="left">Bone</td>
<td valign="top" align="left">Rib</td>
<td valign="top" align="left">2 yr &#x000B1; 8 m</td>
<td valign="top" align="left">17.1</td>
<td valign="top" align="left">5.0</td>
<td valign="top" align="left"><underline>4.0</underline></td>
<td valign="top" align="left">&#x02212;19.7</td>
<td valign="top" align="left">15.1</td>
<td valign="top" align="left">n.a.</td>
</tr> <tr>
<td valign="top" align="left">40</td>
<td valign="top" align="left">EF-1</td>
<td valign="top" align="left">ZAC 7073</td>
<td valign="top" align="left">Und</td>
<td valign="top" align="left">6 yr &#x000B1; 24 m</td>
<td valign="top" align="left">Bone</td>
<td valign="top" align="left">Rib</td>
<td valign="top" align="left">6 yr &#x000B1; 24 m</td>
<td valign="top" align="left">No collagen</td>
<td/>
<td/>
<td/>
<td/>
<td valign="top" align="left">n.a.</td>
</tr> <tr>
<td valign="top" align="left">41</td>
<td valign="top" align="left">EF-1</td>
<td valign="top" align="left">ZAC 7073</td>
<td valign="top" align="left">Und</td>
<td valign="top" align="left">6 yr &#x000B1; 24 m</td>
<td valign="top" align="left">Tooth</td>
<td valign="top" align="left">7,3 (root)</td>
<td valign="top" align="left">0.9&#x02013;3.5 yr</td>
<td valign="top" align="left">No collagen</td>
<td/>
<td/>
<td/>
<td/>
<td valign="top" align="left">n.a.</td>
</tr> <tr>
<td valign="top" align="left">42</td>
<td valign="top" align="left">EF-1</td>
<td valign="top" align="left">ZAC 7074<sup>&#x0002A;&#x0002A;</sup></td>
<td valign="top" align="left">M</td>
<td valign="top" align="left">30&#x02013;40 yr</td>
<td valign="top" align="left">Bone</td>
<td valign="top" align="left">Rib</td>
<td valign="top" align="left">30&#x02013;40 yr</td>
<td valign="top" align="left">42.1</td>
<td valign="top" align="left">14.5</td>
<td valign="top" align="left">3.4</td>
<td valign="top" align="left">&#x02212;17</td>
<td valign="top" align="left">13.0</td>
<td valign="top" align="left">&#x02212;14.2</td>
</tr> <tr>
<td valign="top" align="left">43</td>
<td valign="top" align="left">EF-1</td>
<td valign="top" align="left">ZAC 7074</td>
<td valign="top" align="left">M</td>
<td valign="top" align="left">30&#x02013;40 yr</td>
<td valign="top" align="left">Tooth</td>
<td valign="top" align="left">1,8 (root)</td>
<td valign="top" align="left">8.0&#x02013;16.6</td>
<td valign="top" align="left">40.2</td>
<td valign="top" align="left">14.2</td>
<td valign="top" align="left">3.3</td>
<td valign="top" align="left">&#x02212;17.2</td>
<td valign="top" align="left">13.6</td>
<td valign="top" align="left">n.a.</td>
</tr> <tr>
<td valign="top" align="left">44</td>
<td valign="top" align="left">EF-1</td>
<td valign="top" align="left">ZAC 7077<sup>&#x0002A;&#x0002A;</sup></td>
<td valign="top" align="left">M</td>
<td valign="top" align="left">40&#x02013;60 yr</td>
<td valign="top" align="left">Bone</td>
<td valign="top" align="left">Rib</td>
<td valign="top" align="left">40&#x02013;60 yr</td>
<td valign="top" align="left">42.3</td>
<td valign="top" align="left">14.9</td>
<td valign="top" align="left">3.3</td>
<td valign="top" align="left">&#x02212;17.5</td>
<td valign="top" align="left">11.8</td>
<td valign="top" align="left">&#x02212;12.7</td>
</tr> <tr>
<td valign="top" align="left">45</td>
<td valign="top" align="left">EF-1</td>
<td valign="top" align="left">ZAC 7080</td>
<td valign="top" align="left">F</td>
<td valign="top" align="left">20&#x02013;25 yr</td>
<td valign="top" align="left">Bone</td>
<td valign="top" align="left">Rib</td>
<td valign="top" align="left">20&#x02013;25 yr</td>
<td valign="top" align="left">No collagen</td>
<td/>
<td/>
<td/>
<td/>
<td valign="top" align="left">&#x02212;13.9</td>
</tr> <tr>
<td valign="top" align="left">46</td>
<td valign="top" align="left">EF-1</td>
<td valign="top" align="left">ZAC 7080</td>
<td valign="top" align="left">F</td>
<td valign="top" align="left">20&#x02013;25 yr</td>
<td valign="top" align="left">Tooth</td>
<td valign="top" align="left">1,8 (root)</td>
<td valign="top" align="left">8.0&#x02013;16.5 yr</td>
<td valign="top" align="left">44.4</td>
<td valign="top" align="left">16.2</td>
<td valign="top" align="left">3.2</td>
<td valign="top" align="left">&#x02212;17.4</td>
<td valign="top" align="left">11.8</td>
<td valign="top" align="left">n.a.</td>
</tr> <tr>
<td valign="top" align="left">47</td>
<td valign="top" align="left">EF-1</td>
<td valign="top" align="left">ZAC 7081</td>
<td valign="top" align="left">Und</td>
<td valign="top" align="left">6 m &#x000B1; 3 m</td>
<td valign="top" align="left">Bone</td>
<td valign="top" align="left">Rib</td>
<td valign="top" align="left">6 m &#x000B1; 3 m</td>
<td valign="top" align="left">19.6</td>
<td valign="top" align="left">4.8</td>
<td valign="top" align="left"><underline>4.8</underline></td>
<td valign="top" align="left">&#x02212;18.7</td>
<td valign="top" align="left">27.5</td>
<td valign="top" align="left">n.a.</td>
</tr> <tr>
<td valign="top" align="left">48</td>
<td valign="top" align="left">EF-1</td>
<td valign="top" align="left">ZAC 0955</td>
<td valign="top" align="left">M</td>
<td valign="top" align="left">50&#x02013;70 yr</td>
<td valign="top" align="left">Bone</td>
<td valign="top" align="left">Rib</td>
<td valign="top" align="left">50&#x02013;70 yr</td>
<td valign="top" align="left">No collagen</td>
<td/>
<td/>
<td/>
<td/>
<td valign="top" align="left">&#x02212;12.1</td>
</tr> <tr>
<td valign="top" align="left">49</td>
<td valign="top" align="left">LIP</td>
<td valign="top" align="left">ZAC 7054<sup>&#x0002A;&#x0002A;</sup></td>
<td valign="top" align="left">Und</td>
<td valign="top" align="left">7 yr &#x000B1; 24 m</td>
<td valign="top" align="left">Bone</td>
<td valign="top" align="left">Rib</td>
<td valign="top" align="left">7 yr &#x000B1; 24 m</td>
<td valign="top" align="left">37.5</td>
<td valign="top" align="left">13.4</td>
<td valign="top" align="left">3.3</td>
<td valign="top" align="left">&#x02212;17.9</td>
<td valign="top" align="left">11.5</td>
<td valign="top" align="left">&#x02212;13.9</td>
</tr> <tr>
<td valign="top" align="left">50</td>
<td valign="top" align="left">LIP</td>
<td valign="top" align="left">ZAC 7054</td>
<td valign="top" align="left">Und</td>
<td valign="top" align="left">7 yr &#x000B1; 24 m</td>
<td valign="top" align="left">Tooth</td>
<td valign="top" align="left">1,6 (root)</td>
<td valign="top" align="left">3.5&#x02013;7 yr</td>
<td valign="top" align="left">47.7</td>
<td valign="top" align="left">15.9</td>
<td valign="top" align="left">3.5</td>
<td valign="top" align="left">&#x02212;17.8</td>
<td valign="top" align="left">12.4</td>
<td valign="top" align="left">n.a.</td>
</tr> <tr>
<td valign="top" align="left">51</td>
<td valign="top" align="left">LIP</td>
<td valign="top" align="left">ZAC 7055</td>
<td valign="top" align="left">F</td>
<td valign="top" align="left">30&#x02013;40 yr</td>
<td valign="top" align="left">Tooth</td>
<td valign="top" align="left">4,5 (root)</td>
<td valign="top" align="left">6.5&#x02013;12.0 yr</td>
<td valign="top" align="left">44.6</td>
<td valign="top" align="left">15.3</td>
<td valign="top" align="left">3.4</td>
<td valign="top" align="left">&#x02212;16.7</td>
<td valign="top" align="left">15.6</td>
<td valign="top" align="left">n.a.</td>
</tr> <tr>
<td valign="top" align="left">52</td>
<td valign="top" align="left">LIP</td>
<td valign="top" align="left">ZAC 7059<sup>&#x0002A;&#x0002A;</sup></td>
<td valign="top" align="left">M</td>
<td valign="top" align="left">15&#x02013;19 yr</td>
<td valign="top" align="left">Bone</td>
<td valign="top" align="left">Rib</td>
<td valign="top" align="left">15&#x02013;19 yr</td>
<td valign="top" align="left">44.3</td>
<td valign="top" align="left">15.6</td>
<td valign="top" align="left">3.3</td>
<td valign="top" align="left">&#x02212;12.3</td>
<td valign="top" align="left">16.4</td>
<td valign="top" align="left">&#x02212;7.7</td>
</tr> <tr>
<td valign="top" align="left">53</td>
<td valign="top" align="left">LIP</td>
<td valign="top" align="left">ZAC 7060</td>
<td valign="top" align="left">M</td>
<td valign="top" align="left">45&#x02013;55 yr</td>
<td valign="top" align="left">Bone</td>
<td valign="top" align="left">Rib</td>
<td valign="top" align="left">45&#x02013;55 yr</td>
<td valign="top" align="left">16.4</td>
<td valign="top" align="left">4.6</td>
<td valign="top" align="left"><underline>4.2</underline></td>
<td valign="top" align="left">&#x02212;20.2</td>
<td valign="top" align="left">13</td>
<td valign="top" align="left">n.a.</td>
</tr> <tr>
<td valign="top" align="left">54</td>
<td valign="top" align="left">LIP</td>
<td valign="top" align="left">ZAC 7060</td>
<td valign="top" align="left">M</td>
<td valign="top" align="left">45&#x02013;55 yr</td>
<td valign="top" align="left">Tooth</td>
<td valign="top" align="left">3,8 (root)</td>
<td valign="top" align="left">14.0&#x02013;23.5 yr</td>
<td valign="top" align="left">44.0</td>
<td valign="top" align="left">15.9</td>
<td valign="top" align="left">3.2</td>
<td valign="top" align="left">&#x02212;18.1</td>
<td valign="top" align="left">12.1</td>
<td valign="top" align="left">n.a.</td>
</tr> <tr>
<td valign="top" align="left">55</td>
<td valign="top" align="left">LIP</td>
<td valign="top" align="left">ZAC 7061<sup>&#x0002A;&#x0002A;</sup></td>
<td valign="top" align="left">M</td>
<td valign="top" align="left">20&#x02013;25 yr</td>
<td valign="top" align="left">Bone</td>
<td valign="top" align="left">Rib</td>
<td valign="top" align="left">20&#x02013;25 yr</td>
<td valign="top" align="left">41.4</td>
<td valign="top" align="left">15.0</td>
<td valign="top" align="left">3.2</td>
<td valign="top" align="left">&#x02212;11.0</td>
<td valign="top" align="left">13.7</td>
<td valign="top" align="left">&#x02212;6.8</td>
</tr> <tr>
<td valign="top" align="left">56</td>
<td valign="top" align="left">LIP</td>
<td valign="top" align="left">ZAC 7078</td>
<td valign="top" align="left">M</td>
<td valign="top" align="left">50&#x02013;70 yr</td>
<td valign="top" align="left">Bone</td>
<td valign="top" align="left">Rib</td>
<td valign="top" align="left">50&#x02013;70 yr</td>
<td valign="top" align="left">24.8</td>
<td valign="top" align="left">7.7</td>
<td valign="top" align="left"><underline>3.8</underline></td>
<td valign="top" align="left">&#x02212;17.7</td>
<td valign="top" align="left">12.8</td>
<td valign="top" align="left">n.a.</td>
</tr> <tr>
<td valign="top" align="left">57</td>
<td valign="top" align="left">LIP</td>
<td valign="top" align="left">ZAC 7079</td>
<td valign="top" align="left">M</td>
<td valign="top" align="left">25&#x02013;50 yr</td>
<td valign="top" align="left">Bone</td>
<td valign="top" align="left">Rib</td>
<td valign="top" align="left">25&#x02013;50 yr</td>
<td valign="top" align="left">No collagen</td>
<td/>
<td/>
<td/>
<td/>
<td valign="top" align="left">n.a.</td>
</tr> <tr>
<td valign="top" align="left">58</td>
<td valign="top" align="left">LIP</td>
<td valign="top" align="left">ZAC 7079<sup>&#x0002A;&#x0002A;</sup></td>
<td valign="top" align="left">M</td>
<td valign="top" align="left">25&#x02013;50 yr</td>
<td valign="top" align="left">Tooth</td>
<td valign="top" align="left">4,3 (root)</td>
<td valign="top" align="left">5.5&#x02013;14.5 yr</td>
<td valign="top" align="left">43.6</td>
<td valign="top" align="left">15.4</td>
<td valign="top" align="left">3.3</td>
<td valign="top" align="left">&#x02212;15.2</td>
<td valign="top" align="left">17.4</td>
<td valign="top" align="left">&#x02212;10.4</td>
</tr> <tr>
<td valign="top" align="left">59</td>
<td valign="top" align="left">LIP</td>
<td valign="top" align="left">ZAC 0465<sup>&#x0002A;&#x0002A;</sup></td>
<td valign="top" align="left">F</td>
<td valign="top" align="left">35&#x02013;40 yr</td>
<td valign="top" align="left">Tooth</td>
<td valign="top" align="left">3,5 (root)</td>
<td valign="top" align="left">6.5&#x02013;12.0 yr</td>
<td valign="top" align="left">45.0</td>
<td valign="top" align="left">14.4</td>
<td valign="top" align="left">3.6</td>
<td valign="top" align="left">&#x02212;16.9</td>
<td valign="top" align="left">13.4</td>
<td valign="top" align="left">&#x02212;11.9</td>
</tr></tbody>
</table>
<table-wrap-foot>
<p>EF-1, Early Formative&#x02212;1 (1800&#x02013;1500 <sc>bce</sc>); LIP, Late Intermediate Period (1000&#x02013;1300 <sc>ce</sc>); P, architectural period; M, male; F, female; Und, undetermined; Adult, adult individual without age-range estimation; n.a., value not available. Individuals with underlined C:N ratios were eliminated from analyses.</p>
<p><sup>&#x0002A;</sup>The age range of the tissue used in the analysis (e.g., teeth develop at infant or child ages, so the isotopic values correspond to early life). Recognized teeth are reported in FDA nomenclature; unrecognized teeth are reported in anthropological nomenclature.</p>
<p><sup>&#x0002A;&#x0002A;</sup>Individuals included in Bayesian stable isotope mixing model analysis.</p>
</table-wrap-foot>
</table-wrap>
<p>The &#x003B4;<sup>13</sup>C<sub>coll</sub> values from both bone and tooth dentin ranged from &#x02212;20.3 to &#x02212;11.0 &#x02030;, &#x003B4;<sup>15</sup>N values ranged from &#x0002B;8.2 to &#x0002B; 18.0 &#x02030;, whereas &#x003B4;<sup>13</sup>C<sub>ap</sub> values ranged from &#x02212;14.6 to &#x02212;6.8 &#x02030; (<xref ref-type="supplementary-material" rid="SM1">Supplementary material 1</xref>: S5).</p>
<p>Although the contribution of marine protein is evident in some male individuals (<xref ref-type="fig" rid="F4">Figure 4a</xref>), the mean &#x003B4;<sup>15</sup>N values are lower than expected for fishers and suggest the consumption of marine protein of a low trophic level, possibly little fishes and shellfish (with mean values of &#x003B4;<sup>13</sup>C<sub>coll</sub> = &#x02212;12.3 &#x02030;, &#x003C3; = 0.5, and &#x003B4;<sup>15</sup>N = &#x0002B; 10.4 &#x02030;, &#x003C3; = 1.2) over the entire sequence. No significant differences were observed for &#x003B4;<sup>13</sup>C<sub>coll</sub>, &#x003B4;<sup>15</sup>N, and &#x003B4;<sup>13</sup>C<sub>ap</sub> values between females and males or between preadults and adults for both EF-1 and LIP occupations (<xref ref-type="supplementary-material" rid="SM1">Supplementary material 1</xref>: S5). The significant difference observed between adults and preadults of EF-1 probably resulted from the weaning effect in a couple of infants (i.e., PEACS4431 and ZAC7057) or different protein sources (i.e., ZAC7067) in some preadults.</p>
<fig id="F4" position="float">
<label>Figure 4</label>
<caption><p>Scatterplot of stable isotope markers in Vichama individuals of Early Formative period and the Late Intermediate Period and overlapping standard ellipse areas (estimated for 68.5% of the population), for comparison. <bold>(a)</bold> &#x003B4;<sup>13</sup>C<sub>coll</sub> and &#x003B4;<sup>15</sup>N values. <bold>(b)</bold> &#x003B4;<sup>13</sup>C<sub>coll</sub> and &#x003B4;<sup>13</sup>C<sub>ap</sub> values in the workspace of Kellner and Schoeninger (<xref ref-type="bibr" rid="B38">2007</xref>). V-PDB, Vienna Pee Dee Belemnite; AIR, atmospheric air.</p></caption>
<alt-text>Two scatter plots labeled &#x0201C;a&#x0201D; and &#x0201C;b&#x0201D; are shown. Plot &#x0201C;a&#x0201D; maps &#x003B4;13C against &#x003B4;15N values with two ellipses: red for adults and blue for preadults. Plot &#x0201C;b&#x0201D; compares &#x003B4;13C Collagen and Apatite values, featuring C3, C4, and Marine Protein lines. Symbols represent gender: female, male, and undetermined, with crosses for EF-1 and LIP groups.</alt-text>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fearc-04-1611071-g0004.tif"/>
</fig>
<p>Inter-population comparisons revealed a significant difference only for &#x003B4;<sup>13</sup>C<sub>ap</sub> (Mann&#x02013;Whitney <italic>U</italic> = 18.50, <italic>p</italic> = 0027). In general, the &#x003B4;<sup>13</sup>C<sub>coll</sub>, and &#x003B4;<sup>13</sup>C<sub>ap</sub> indicate diets dominated by protein and energy sources from C<sub>3</sub> ecosystems (<xref ref-type="fig" rid="F4">Figure 4b</xref>; Kellner and Schoeninger, <xref ref-type="bibr" rid="B38">2007</xref>). Nevertheless, individuals from the LIP exhibited a much broader range of &#x003B4;<sup>13</sup>C<sub>coll</sub> and &#x003B4;<sup>15</sup>N values, indicating relatively higher marine protein consumption, notably by some male individuals (e.g., ZAC7061, ZAC7059), compared to the EF-1 population.</p>
<p>The comparisons of protein sources of Vichama EF-1 and LIP phases with other populations from the PNCC show a trend to consume a diet intermediary between that of &#x000C1;spero, the most representative site of fishermen in the Supe Valley during the Initial Formative (3000&#x02013;1800 <sc>bce</sc>), based on C<sub>3</sub> plants and marine protein of higher trophic level, and the diet of later farmers from Quebrada Chupacigarro cemetery (middle Supe Valley, Late Formative, 500&#x02013;400 <sc>bce</sc>), virtually based on plants (Pezo-Lanfranco et al., <xref ref-type="bibr" rid="B53">2022</xref>). LIP diets show an overlap area greater than EP-1 with &#x000C1;spero diets (<xref ref-type="fig" rid="F5">Figure 5</xref>).</p>
<fig id="F5" position="float">
<label>Figure 5</label>
<caption><p>Scatterplot of &#x003B4;<sup>13</sup>C and &#x003B4;<sup>15</sup>N with standard ellipse areas (estimated for 68.5% of the population), and density kernel plots of &#x003B4;<sup>13</sup>C<sub>coll</sub> and &#x003B4;<sup>15</sup>N values of Vichama and other Formative populations from the Peruvian North-Central coast. EF-1, Early Formative&#x02212;1 period; LIP, Late Intermediate Period; QCC, Quebrada Chupacigarro Cemetery.</p></caption>
<alt-text>Scatter plot with ellipses and histograms depicts isotopic data. Left graph plots &#x003B4;15N (&#x02030;) against &#x003B4;13C (&#x02030;) for four groups: &#x000C1;spero (blue), Vichama EF-1 (red), Vichama LIP (gray), and QCC (green), each with a distinct cluster. Right side features corresponding histograms showing the Kernell distribution of &#x003B4;13C and &#x003B4;15N values for each group, color-coded to match.</alt-text>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fearc-04-1611071-g0005.tif"/>
</fig>
<p>In addition, we estimate the proportions of dietary components estimated with BSIMMs for 21 Vichama individuals with a set of complete isotopic data (<xref ref-type="table" rid="T2">Table 2</xref>, <xref ref-type="fig" rid="F6">Figure 6</xref>; see also <xref ref-type="supplementary-material" rid="SM1">Supplementary material 1</xref>: S6 for the model&#x00027;s results). These results suggest that most calories consumed came from C<sub>3</sub> plants (contribution ranging from 14% to 86%), followed by fish and mollusks (8% and 58%), C<sub>4</sub> plants (3%&#x02212;19%), and terrestrial fauna (3%&#x02212;11%).</p>
<table-wrap position="float" id="T2">
<label>Table 2</label>
<caption><p>Estimates of dietary carbon/calories contribution of each food group in Vichama individuals.<sup>&#x0002A;</sup></p></caption>
<table frame="box" rules="all">
<thead>
<tr style="background-color:#8f9496;color:#ffffff">
<th valign="top" align="left"><bold>Period</bold></th>
<th valign="top" align="left"><bold>ID</bold></th>
<th valign="top" align="left"><bold>Sex</bold></th>
<th valign="top" align="left"><bold>Age range (years)</bold></th>
<th valign="top" align="left" colspan="2"><bold>C</bold><sub><bold>4</bold></sub> <bold>plants</bold></th>
<th valign="top" align="left" colspan="2"><bold>C</bold><sub><bold>3</bold></sub> <bold>plants</bold></th>
<th valign="top" align="left" colspan="2"><bold>Terrestrial protein</bold></th>
<th valign="top" align="left" colspan="2"><bold>Marine protein</bold></th>
</tr>
</thead>
<tbody>
<tr style="background-color:#8f9496;color:#ffffff">
<td/>
<td/>
<td/>
<td/>
<td valign="top" align="left"><inline-formula><mml:math id="M1"><mml:mover accent="true"><mml:mrow><mml:mi>x</mml:mi></mml:mrow><mml:mo>&#x00304;</mml:mo></mml:mover></mml:math></inline-formula></td>
<td valign="top" align="left">&#x003C3;</td>
<td valign="top" align="left"><inline-formula><mml:math id="M2"><mml:mover accent="true"><mml:mrow><mml:mi>x</mml:mi></mml:mrow><mml:mo>&#x00304;</mml:mo></mml:mover></mml:math></inline-formula></td>
<td valign="top" align="left">&#x003C3;</td>
<td valign="top" align="left"><inline-formula><mml:math id="M3"><mml:mover accent="true"><mml:mrow><mml:mi>x</mml:mi></mml:mrow><mml:mo>&#x00304;</mml:mo></mml:mover></mml:math></inline-formula></td>
<td valign="top" align="left">&#x003C3;</td>
<td valign="top" align="left"><inline-formula><mml:math id="M4"><mml:mover accent="true"><mml:mrow><mml:mi>x</mml:mi></mml:mrow><mml:mo>&#x00304;</mml:mo></mml:mover></mml:math></inline-formula></td>
<td valign="top" align="left">&#x003C3;</td>
</tr> <tr>
<td valign="top" align="left">EF-1</td>
<td valign="top" align="left">ZAC 0458</td>
<td valign="top" align="left">F</td>
<td valign="top" align="left">Adult</td>
<td valign="top" align="left">0.05</td>
<td valign="top" align="left">0.03</td>
<td valign="top" align="left">0.74</td>
<td valign="top" align="left">0.08</td>
<td valign="top" align="left">0.06</td>
<td valign="top" align="left">0.05</td>
<td valign="top" align="left">0.15</td>
<td valign="top" align="left">0.05</td>
</tr> <tr>
<td valign="top" align="left">EF-1</td>
<td valign="top" align="left">ZAC 7069</td>
<td valign="top" align="left">M</td>
<td valign="top" align="left">22&#x02013;32 yr</td>
<td valign="top" align="left">0.05</td>
<td valign="top" align="left">0.03</td>
<td valign="top" align="left">0.74</td>
<td valign="top" align="left">0.09</td>
<td valign="top" align="left">0.06</td>
<td valign="top" align="left">0.05</td>
<td valign="top" align="left">0.14</td>
<td valign="top" align="left">0.06</td>
</tr> <tr>
<td valign="top" align="left">EF-1</td>
<td valign="top" align="left">ZAC 5905</td>
<td valign="top" align="left">M</td>
<td valign="top" align="left">25&#x02013;35 yr</td>
<td valign="top" align="left">0.04</td>
<td valign="top" align="left">0.03</td>
<td valign="top" align="left">0.78</td>
<td valign="top" align="left">0.07</td>
<td valign="top" align="left">0.06</td>
<td valign="top" align="left">0.04</td>
<td valign="top" align="left">0.12</td>
<td valign="top" align="left">0.05</td>
</tr> <tr>
<td valign="top" align="left">EF-1</td>
<td valign="top" align="left">PEACS 4431</td>
<td valign="top" align="left">Und</td>
<td valign="top" align="left">Preadult</td>
<td valign="top" align="left">0.04</td>
<td valign="top" align="left">0.03</td>
<td valign="top" align="left">0.78</td>
<td valign="top" align="left">0.07</td>
<td valign="top" align="left">0.06</td>
<td valign="top" align="left">0.04</td>
<td valign="top" align="left">0.12</td>
<td valign="top" align="left">0.05</td>
</tr> <tr>
<td valign="top" align="left">EF-1</td>
<td valign="top" align="left">ZAC 7056</td>
<td valign="top" align="left">F</td>
<td valign="top" align="left">35&#x02013;65 yr</td>
<td valign="top" align="left">0.05</td>
<td valign="top" align="left">0.04</td>
<td valign="top" align="left">0.70</td>
<td valign="top" align="left">0.09</td>
<td valign="top" align="left">0.08</td>
<td valign="top" align="left">0.05</td>
<td valign="top" align="left">0.17</td>
<td valign="top" align="left">0.06</td>
</tr> <tr>
<td valign="top" align="left">EF-1</td>
<td valign="top" align="left">ZAC 7057</td>
<td valign="top" align="left">Und</td>
<td valign="top" align="left">3 yr &#x000B1; 12 m</td>
<td valign="top" align="left">0.07</td>
<td valign="top" align="left">0.05</td>
<td valign="top" align="left">0.56</td>
<td valign="top" align="left">0.10</td>
<td valign="top" align="left">0.11</td>
<td valign="top" align="left">0.07</td>
<td valign="top" align="left">0.26</td>
<td valign="top" align="left">0.08</td>
</tr> <tr>
<td valign="top" align="left">EF-1</td>
<td valign="top" align="left">ZAC 7076</td>
<td valign="top" align="left">M</td>
<td valign="top" align="left">16&#x02013;19 yr</td>
<td valign="top" align="left">0.03</td>
<td valign="top" align="left">0.02</td>
<td valign="top" align="left">0.81</td>
<td valign="top" align="left">0.07</td>
<td valign="top" align="left">0.05</td>
<td valign="top" align="left">0.03</td>
<td valign="top" align="left">0.11</td>
<td valign="top" align="left">0.04</td>
</tr> <tr>
<td valign="top" align="left">EF-1</td>
<td valign="top" align="left">ZAC 7062</td>
<td valign="top" align="left">M</td>
<td valign="top" align="left">35&#x02013;65 yr</td>
<td valign="top" align="left">0.06</td>
<td valign="top" align="left">0.04</td>
<td valign="top" align="left">0.76</td>
<td valign="top" align="left">0.09</td>
<td valign="top" align="left">0.05</td>
<td valign="top" align="left">0.04</td>
<td valign="top" align="left">0.14</td>
<td valign="top" align="left">0.06</td>
</tr> <tr>
<td valign="top" align="left">EF-1</td>
<td valign="top" align="left">ZAC 7065</td>
<td valign="top" align="left">M</td>
<td valign="top" align="left">20&#x02013;25 yr</td>
<td valign="top" align="left">0.05</td>
<td valign="top" align="left">0.04</td>
<td valign="top" align="left">0.71</td>
<td valign="top" align="left">0.08</td>
<td valign="top" align="left">0.07</td>
<td valign="top" align="left">0.05</td>
<td valign="top" align="left">0.17</td>
<td valign="top" align="left">0.06</td>
</tr> <tr>
<td valign="top" align="left">EF-1</td>
<td valign="top" align="left">ZAC 7066</td>
<td valign="top" align="left">M</td>
<td valign="top" align="left">30&#x02013;40 yr</td>
<td valign="top" align="left">0.03</td>
<td valign="top" align="left">0.02</td>
<td valign="top" align="left">0.86</td>
<td valign="top" align="left">0.05</td>
<td valign="top" align="left">0.03</td>
<td valign="top" align="left">0.02</td>
<td valign="top" align="left">0.08</td>
<td valign="top" align="left">0.03</td>
</tr> <tr>
<td valign="top" align="left">EF-1</td>
<td valign="top" align="left">ZAC 7067</td>
<td valign="top" align="left">Und</td>
<td valign="top" align="left">8 yr &#x000B1; 24 m</td>
<td valign="top" align="left">0.05</td>
<td valign="top" align="left">0.04</td>
<td valign="top" align="left">0.79</td>
<td valign="top" align="left">0.08</td>
<td valign="top" align="left">0.04</td>
<td valign="top" align="left">0.03</td>
<td valign="top" align="left">0.12</td>
<td valign="top" align="left">0.05</td>
</tr> <tr>
<td valign="top" align="left">EF-1</td>
<td valign="top" align="left">ZAC 7068</td>
<td valign="top" align="left">F</td>
<td valign="top" align="left">25&#x02013;40 yr</td>
<td valign="top" align="left">0.07</td>
<td valign="top" align="left">0.05</td>
<td valign="top" align="left">0.66</td>
<td valign="top" align="left">0.09</td>
<td valign="top" align="left">0.08</td>
<td valign="top" align="left">0.05</td>
<td valign="top" align="left">0.19</td>
<td valign="top" align="left">0.06</td>
</tr> <tr>
<td valign="top" align="left">EF-1</td>
<td valign="top" align="left">ZAC 7070</td>
<td valign="top" align="left">F</td>
<td valign="top" align="left">20&#x02013;30 yr</td>
<td valign="top" align="left">0.08</td>
<td valign="top" align="left">0.05</td>
<td valign="top" align="left">0.53</td>
<td valign="top" align="left">0.11</td>
<td valign="top" align="left">0.11</td>
<td valign="top" align="left">0.08</td>
<td valign="top" align="left">0.28</td>
<td valign="top" align="left">0.08</td>
</tr> <tr>
<td valign="top" align="left">EF-1</td>
<td valign="top" align="left">ZAC 7071</td>
<td valign="top" align="left">M</td>
<td valign="top" align="left">25&#x02013;55 yr</td>
<td valign="top" align="left">0.08</td>
<td valign="top" align="left">0.06</td>
<td valign="top" align="left">0.58</td>
<td valign="top" align="left">0.11</td>
<td valign="top" align="left">0.10</td>
<td valign="top" align="left">0.07</td>
<td valign="top" align="left">0.24</td>
<td valign="top" align="left">0.08</td>
</tr> <tr>
<td valign="top" align="left">EF-1</td>
<td valign="top" align="left">ZAC 7074</td>
<td valign="top" align="left">M</td>
<td valign="top" align="left">30&#x02013;40 yr</td>
<td valign="top" align="left">0.05</td>
<td valign="top" align="left">0.04</td>
<td valign="top" align="left">0.73</td>
<td valign="top" align="left">0.08</td>
<td valign="top" align="left">0.06</td>
<td valign="top" align="left">0.04</td>
<td valign="top" align="left">0.15</td>
<td valign="top" align="left">0.06</td>
</tr> <tr>
<td valign="top" align="left">EF-1</td>
<td valign="top" align="left">ZAC 7077</td>
<td valign="top" align="left">M</td>
<td valign="top" align="left">40&#x02013;60 yr</td>
<td valign="top" align="left">0.05</td>
<td valign="top" align="left">0.03</td>
<td valign="top" align="left">0.77</td>
<td valign="top" align="left">0.09</td>
<td valign="top" align="left">0.05</td>
<td valign="top" align="left">0.04</td>
<td valign="top" align="left">0.12</td>
<td valign="top" align="left">0.05</td>
</tr> <tr>
<td valign="top" align="left">LIP</td>
<td valign="top" align="left">ZAC 7054</td>
<td valign="top" align="left">Und</td>
<td valign="top" align="left">7 yr &#x000B1; 24 m</td>
<td valign="top" align="left">0.04</td>
<td valign="top" align="left">0.03</td>
<td valign="top" align="left">0.81</td>
<td valign="top" align="left">0.07</td>
<td valign="top" align="left">0.04</td>
<td valign="top" align="left">0.03</td>
<td valign="top" align="left">0.11</td>
<td valign="top" align="left">0.05</td>
</tr> <tr>
<td valign="top" align="left">LIP</td>
<td valign="top" align="left">ZAC 7059</td>
<td valign="top" align="left">M</td>
<td valign="top" align="left">15&#x02013;19 yr</td>
<td valign="top" align="left">0.17</td>
<td valign="top" align="left">0.09</td>
<td valign="top" align="left">0.20</td>
<td valign="top" align="left">0.12</td>
<td valign="top" align="left">0.20</td>
<td valign="top" align="left">0.11</td>
<td valign="top" align="left">0.43</td>
<td valign="top" align="left">0.11</td>
</tr> <tr>
<td valign="top" align="left">LIP</td>
<td valign="top" align="left">ZAC 7061</td>
<td valign="top" align="left">M</td>
<td valign="top" align="left">20&#x02013;25 yr</td>
<td valign="top" align="left">0.19</td>
<td valign="top" align="left">0.10</td>
<td valign="top" align="left">0.14</td>
<td valign="top" align="left">0.09</td>
<td valign="top" align="left">0.16</td>
<td valign="top" align="left">0.10</td>
<td valign="top" align="left">0.58</td>
<td valign="top" align="left">0.14</td>
</tr> <tr>
<td valign="top" align="left">LIP</td>
<td valign="top" align="left">ZAC 7079</td>
<td valign="top" align="left">M</td>
<td valign="top" align="left">25&#x02013;50 yr</td>
<td valign="top" align="left">0.08</td>
<td valign="top" align="left">0.05</td>
<td valign="top" align="left">0.50</td>
<td valign="top" align="left">0.12</td>
<td valign="top" align="left">0.14</td>
<td valign="top" align="left">0.08</td>
<td valign="top" align="left">0.28</td>
<td valign="top" align="left">0.08</td>
</tr> <tr>
<td valign="top" align="left">LIP</td>
<td valign="top" align="left">ZAC 0465</td>
<td valign="top" align="left">F</td>
<td valign="top" align="left">35&#x02013;45 yr</td>
<td valign="top" align="left">0.07</td>
<td valign="top" align="left">0.04</td>
<td valign="top" align="left">0.67</td>
<td valign="top" align="left">0.09</td>
<td valign="top" align="left">0.08</td>
<td valign="top" align="left">0.06</td>
<td valign="top" align="left">0.18</td>
<td valign="top" align="left">0.06</td>
</tr></tbody>
</table>
<table-wrap-foot>
<p>EF-1, Early Formative&#x02212;1 (1800&#x02013;1500 <sc>bce</sc>); LIP, Late Intermediate Period (1000&#x02013;1300 <sc>ce</sc>); P, architectural period; M, male; F, female; Und, undetermined.</p>
<p><sup>&#x0002A;</sup>Estimates based on FRUITS v. 2.1.1 only for individuals with &#x003B4;<sup>13</sup>C<sub>coll</sub>, &#x003B4;<sup>15</sup>N<sub>coll</sub> and &#x003B4;<sup>13</sup>C<sub>ap</sub> values. Foods (%): estimates of dietary carbon/calories contribution of each food group, expressed as relative contributions (adding to 100%) with an associated 1&#x003C3; uncertainty (Fernandes et al., <xref ref-type="bibr" rid="B23">2015</xref>, Table 5).</p>
</table-wrap-foot>
</table-wrap>
<fig id="F6" position="float">
<label>Figure 6</label>
<caption><p>Dietary caloric estimations of food sources in Vichama individuals for the Early Formative&#x02212;1 (EF-1, 1800&#x02013;1500 bce) and the Late Formative Period (LIP, 1000&#x02013;1300 ce), and their correspondence with exported production markers. The boxes represent a 68% confidence interval, and the whiskers, a 95% confidence interval. The horizontal discontinuous line indicates the median, and the continuous line indicates the mean. &#x02642;, male; &#x02640;, female; &#x000B0;, undetermined sex.</p></caption>
<alt-text>Box plots and line graphs display plant and fauna proportions, and bromine/titanium and nickel ratios over time. Box plots show C4, C3 plants, terrestrial and marine fauna percentages during EF-1 and LIP periods. Graphs with individual labels, sex, and age, correspond to bromine/titanium and nickel ratios from 2000 BCE to 1500 CE. The red and gray shaded areas denote EF-1 and LIP periods.</alt-text>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fearc-04-1611071-g0006.tif"/>
</fig>
<p>Thus, most individuals from the EF-1 occupation of Vichama show values compatible with a wide farming&#x02013;fishing diet, with an unexpectedly high contribution of plants to people living near the sea. The comparison of diets between EF-1 and the LIP suggests a trend of increasing marine protein consumption in the later period, fewer C<sub>3</sub> plants, and a discrete increase of C<sub>4</sub> and terrestrial protein. The Kullback&#x02013;Leibler divergence is 0.141, indicating a moderate difference in the average probability distributions among individuals of EF-1 and LIP, consistent with dietary change between periods with some overlap.</p>
</sec>
</sec>
<sec sec-type="discussion" id="s3">
<title>Discussion</title>
<sec>
<title>The nature of diet at Vichama</title>
<p>Our study suggests that the two distinct occupational phases of Vichama were underpinned by different economic systems. During EF-1, dated to 1800&#x02013;1500 <sc>bce</sc>, although some individuals relied significantly on marine resources for calories (&#x0003E;25%), subsistence was based primarily on the cultivation of C<sub>3</sub> plants. Plant remains, including several high-energy crops like tubers, beans, and fruits, were found mainly in funerary contexts associated with public buildings, yet our results reveal the significance of these plants at the household level beyond ritual use.</p>
<p>This agricultural production has likely supported Vichama&#x00027;s political prominence at the end of the Caral era. Cultivation seems to have been a plausible activity in Vichama during the EF-1. The agricultural area near Vichama extends approximately 970 ha, and the water sources are located &#x0003C; 2 km from Vichama. In the zone, the phreatic level is approximately 0.85 m depth (INRENA, <xref ref-type="bibr" rid="B36">2005</xref>).</p>
<p>Evidence of botanical macro- and micro-remains shows the consumption of several domesticated crops in coastal settlements during the Initial Formative (3000&#x02013;1800 <sc>bce</sc>; Haas et al., <xref ref-type="bibr" rid="B30">2013</xref>; Shady, <xref ref-type="bibr" rid="B66">2006a</xref>,<xref ref-type="bibr" rid="B67">b</xref>,<xref ref-type="bibr" rid="B68">c</xref>; Vega-Centeno, <xref ref-type="bibr" rid="B80">2005</xref>; Yseki et al., <xref ref-type="bibr" rid="B85">2023</xref>, <xref ref-type="bibr" rid="B84">2024</xref>; Zechenter, <xref ref-type="bibr" rid="B87">1988</xref>). For example, plant micro-remains from human dental calculus at &#x000C1;spero, in the same region, indicate significant sweet potato and moderate potato consumption during the Initial Formative period (Yseki et al., <xref ref-type="bibr" rid="B85">2023</xref>), despite their scarcity in the macro-botanical record (Shady and C&#x000E1;ceda, <xref ref-type="bibr" rid="B70">2008</xref>).</p>
<p>Vichama&#x00027;s location is appropriate for maize cultivation due to extensive arable lands and water availability. Interestingly, the isotopic evidence obtained by this work suggests a marginal contribution of C<sub>4</sub> (lower than 10% in most cases) during EF-1 and agrees with the lack of maize evidence in the archaeological record. The near absence of maize in Vichama contrasts with its presence in some early sites from adjacent valleys of the PNCC (Haas et al., <xref ref-type="bibr" rid="B30">2013</xref>; Shady, <xref ref-type="bibr" rid="B67">2006b</xref>; Vega-Centeno, <xref ref-type="bibr" rid="B80">2005</xref>; Zechenter, <xref ref-type="bibr" rid="B87">1988</xref>). In &#x000C1;spero, maize starch has been detected in dental calculus (Yseki et al., <xref ref-type="bibr" rid="B85">2023</xref>), and in Pativilca Valley, maize pollen and starches appear as early as 2500 cal <sc>bce</sc> (Haas et al., <xref ref-type="bibr" rid="B30">2013</xref>). The presence of amaranth could account for the C<sub>4</sub> signal detected at Vichama, although no macro-remains of this plant have been recorded at the site, unlike at Caral, located 30 km to the northeast, where amaranth has been documented in an earlier period (Shady, <xref ref-type="bibr" rid="B68">2006c</xref>).</p>
<p>Approximately 3,000 years later, during the LIP (1300&#x02013;1500 <sc>ce</sc>), when Vichama had long lost its regional political influence, the site&#x00027;s residents relied primarily on C<sub>3</sub> plants, supplemented with a greater intake of marine resources. This increased focus on marine organisms is corroborated by the presence of fishing gear, such as nets and fishhooks made from mussel shells, as well as abundant fish and shell remains, all attesting to extensive fishing activities during the LIP. While maize may have been more consumed during the LIP, isotopic equifinality complicates distinguishing between C<sub>4</sub> plants and marine carbon sources. Two young male individuals from the LIP (ZAC7059 and ZAC7061) exhibit values of C<sub>4</sub> intake of approximately 20%; however, both consumed approximately 40%&#x02212;60% of calories from marine sources and show auditory exostoses, an osteological marker associated with repetitive aquatic activity in cold waters (Pezo-Lanfranco et al., <xref ref-type="bibr" rid="B54">2009</xref>), suggesting that they were divers or fishers rather than maize consumers. In contrast, the only individual from EF-1 with auditory exostosis (ZAC5905, a male) shows, counterintuitively, an unexpected low proportion of marine contribution to diet and &#x003B4;<sup>15</sup>N values compatible with marine resources from the lower trophic level.</p>
<p>The overlap between maize and marine carbon values remains a potential source of uncertainty in isotopic analyses for this region. Regardless, if maize was consumed in Vichama during either EF-1 or the LIP, it likely did not constitute a staple or significant dietary component, contributing &#x0003C; 30% of total dietary calories (Ambrose and Norr, <xref ref-type="bibr" rid="B4">1993</xref>).</p>
<p>During the EF-1, terrestrial mammal consumption appears to have been occasional, with contributions from these protein sources increasing in the LIP (20% of total calories), likely due to a greater availability of camelids during this period (Le&#x000F3;n, <xref ref-type="bibr" rid="B41">2013</xref>). The faunal record at Vichama includes only six taxa: three bird species, commensal rodents, frogs, and camelids, the latter being the only potentially edible domesticated taxon. However, archeozoological studies of inter-period consumption are still pending. The limited availability of domesticated species in the Andes, such as camelids, guinea pigs, and dogs, along with certain wild species from the <italic>lomas</italic> and desert areas, however, suggests that terrestrial protein remained a relatively minor dietary component during the Formative period, as observed in other studies (Pezo-Lanfranco et al., <xref ref-type="bibr" rid="B53">2022</xref>; Pezo-Lanfranco and Colonese, <xref ref-type="bibr" rid="B51">2024</xref>). As a potential limitation of this study, the presence of little fish species can lead to mimicking the signal of wild terrestrial protein from <italic>lomas</italic>, potentially C<sub>4</sub> protein (Coutts et al., <xref ref-type="bibr" rid="B14">2011</xref>; Cadwallader et al., <xref ref-type="bibr" rid="B6">2012</xref>).</p>
<p>Although the archaeological record of EP-1 and the LIP support fishing activities for both periods, our results revealed that the reliance on marine products was higher during the LIP, which coincides with a time of high marine productivity, compared to EP-1. However, due to the limited LIP sample size, these interpretations should be considered provisional pending further evidence.</p>
<p>Finally, although some preadult individuals included in the BSIMMs could theoretically retain nitrogen isotopic signals from breastfeeding and weaning, leading to spurious interpretations, this is unlikely, as the analyzed tissues (tooth roots) correspond to older children, who typically consume an adult diet. Among the LIP individuals, two are adults and three are children or preadolescents, with sampled tissues ranging from 5.5 to 14.5 years. Previous research on weaning patterns in Andean populations places the completion of weaning between 1.5 and 3 years of age (see Pezo-Lanfranco et al., <xref ref-type="bibr" rid="B52">2020</xref> for a comprehensive review), suggesting that the isotopic values in the LIP are not influenced by the weaning process. In the case of EF-1, among two analyzed infants (PEACS 4431 and ZAC7057), only ZAC7057 shows an elevated nitrogen value, which likely indicates ongoing breastmilk consumption and could potentially mimic a high intake of marine protein in the model. All other individuals analyzed are adults.</p>
</sec>
<sec>
<title>Dietary evidence and climate oscillations in Vichama</title>
<p>The isotope data generated in this study suggests two distinct economic regimes that are likely mirroring changes in political organization at Vichama over time. Food production systems during the EF-1 period were likely coordinated by centralized political institutions, aimed at generating surplus through intensified agriculture. By contrast, the diversified diet observed during the LIP period reflects the absence of these centralized systems at Vichama. This shift supports the existence of a diversified economy focused on small-scale farming and fishing for household consumption, with a reduced emphasis on surplus generation. These economic systems may also have been shaped by climate oscillations and the prevailing ecological conditions.</p>
<p>Wet and dry alternances that fueled and inhibited marine productivity, fishing activity, and, possibly, the expansion and contraction of <italic>lomas</italic> and farming fields into the region. These phenomena are linked to complex ocean-atmosphere interactions (Salvatteci et al., <xref ref-type="bibr" rid="B59">2019</xref>), including oscillations in the Southern Pacific Anticyclone, trade winds, and coastal winds with effects on marine currents (Salvatteci et al., <xref ref-type="bibr" rid="B58">2016</xref>, <xref ref-type="bibr" rid="B59">2019</xref><italic>;</italic> Yseki et al., <xref ref-type="bibr" rid="B86">2022</xref>).</p>
<p>Apart from dietary adaptation, those fluctuating climate conditions could also be reflected in sociopolitical resilience and ideological response to environmental constraints, as suggested by the presence of climate-linked deities in Vichama&#x00027;s iconography (e.g., frog-human deity), potentially reflecting ideological adaptations to environmental constraints (Shady et al., <xref ref-type="bibr" rid="B73">2015</xref>).</p>
<p>Reliance on marine protein during the EF-1 period might have been limited by lower biomass and marine productivity along the Peruvian coast at that time. Between 2300 and 1300 <sc>bce</sc>, SSTs showed reduced seasonality, with cooler summers compared to present conditions (Carr&#x000E9; et al., <xref ref-type="bibr" rid="B9">2014</xref>; Loubere et al., <xref ref-type="bibr" rid="B43">2013</xref>). Additionally, multidecadal fluctuations in upwelling and sea surface temperature between 1800 and 1500 <sc>bce</sc> suggest that the EF-1 period at Vichama unfolded amid highly variable oceanographic conditions (Salvatteci et al., <xref ref-type="bibr" rid="B58">2016</xref>, <xref ref-type="bibr" rid="B59">2019</xref>), potentially making marine resources less predictable. Interestingly, this period corresponds to a greater reliance on C<sub>3</sub> plants.</p>
<p>As argued by previous research (Caramanica et al., <xref ref-type="bibr" rid="B7">2020</xref>; Contreras, <xref ref-type="bibr" rid="B13">2010</xref>; Dillehay and Kolata, <xref ref-type="bibr" rid="B20">2004</xref>; Lima et al., <xref ref-type="bibr" rid="B42">2023</xref>; Wilson et al., <xref ref-type="bibr" rid="B83">2024</xref>), these oscillations coupled with ENSO alternances, could have been produced an opportunistic intensification of certain products (fisheries/crops) when technological level, and sociopolitical conditions allow it, producing corresponding diets with verifiable marine&#x02013;farming fluctuations over the time. Theoretically, under El Ni&#x000F1;o&#x02013;like conditions, the concurrence of drops in marine productivity with wetter conditions (raining) along the coast and <italic>lomas</italic> flourishing, despite landslides and disasters, would lead to increased land productivity in a sort of &#x0201C;high-risk farming&#x0201D;. By comparison, under La Ni&#x000F1;a&#x02013;like conditions, sea temperatures cooling, increased upwelling, and marine productivity, concomitant to a dry climate (i.e., lower rainfall and <italic>lomas</italic> contraction), would have fueled a &#x0201C;low-risk farming&#x0201D; and greater marine production, probably with more marine fauna and more maize in the record, as we can observe during the LIP.</p>
<p>Climate conditions during EF-1 (1800&#x02013;1500 <sc>bce</sc>) at Vichama also included relatively wet periods in the highlands and dry conditions in the coastal valleys, marked by greater seasonality in river flows, with increased precipitation in the wet season and lower runoff during the dry season (Leclerc, <xref ref-type="bibr" rid="B40">2023</xref>; Salvatteci et al., <xref ref-type="bibr" rid="B59">2019</xref>). The development of irrigation systems could have enabled intensified agriculture during the wet season, allowing surplus production to sustain populations through the drier months. Conversely, during the LIP, cooling sea temperatures, enhanced upwelling, and increased marine productivity, coupled with drier climates marked by reduced rainfall and contraction of <italic>lomas</italic> vegetation, may have fostered a shift toward greater reliance on marine resource exploitation alongside small-scale farming (Carr&#x000E9; et al., <xref ref-type="bibr" rid="B9">2014</xref>; Leclerc, <xref ref-type="bibr" rid="B40">2023</xref>; Loubere et al., <xref ref-type="bibr" rid="B43">2013</xref>; Salvatteci et al., <xref ref-type="bibr" rid="B59">2019</xref>).</p>
<p>This study offers new insights into how climate variability shaped subsistence economies in early Andean societies and expands our understanding of the trajectory of Andean civilization. In this sense, our assessment of subsistence patterns during EF-1, a period of emerging political complexity, aligns well with the idea that decreased marine productivity may have encouraged greater reliance on, and intensification of, terrestrial resources (Wilson et al., <xref ref-type="bibr" rid="B83">2024</xref>). Conversely, our findings partially support the hypothesis that stable climatic conditions favor the development of agrarian societies, whereas climatic volatility contributes to political instability, decentralization, and fragmentation (Kennett and Marwan, <xref ref-type="bibr" rid="B39">2015</xref>). Opportunism in the exploitation of available resources and resilience in the face of climatic challenges appear to have been key behaviors of these societies. Climate oscillations could have represented a challenging opportunity to develop adaptive buffer strategies, such as the intensification of farming or other economic activities. This may coincide with the hegemony of coastal, inland, or highland societies over different periods.</p>
<p>These results should be refined and tested by future research taking into consideration some limitations: (1) palaeoceanographic and climate data are fragmentary, ambiguous, and subject of interpretation and do not necessarily can be congruent with our conclusions and (2) equifinality, that is, varying combinations of food contributions that may produce the same isotopic values in a consumer, persists as a potential issue for interpretations.</p>
</sec>
</sec>
<sec sec-type="materials and methods" id="s4">
<title>Materials and methods</title>
<sec>
<title>Individuals and samples</title>
<p>This is a multiproxy stable isotope study designed to reconstruct the composition of Vichama diet in quantitative terms. Our aim is to investigate the economy of Vichama to unravel the role of marine and vegetal resources in the processes of social complexity during the EF period.</p>
<p>A total of 59 samples (36 bones and 23 teeth) from 38 human individuals recovered from burial contexts of Vichama were sampled for the analysis. The skeletal elements recovered for isotope analyses were mainly ribs and long bones, carpals, or cranial bones when ribs were not available. Teeth were sampled when available, and isotopic values were obtained from roots, which makes a valid direct comparison of tooth and bone samples.</p>
<p>Individuals&#x00027; sex and age were estimated using standardized methods (Buikstra and Ubelaker, <xref ref-type="bibr" rid="B5">1994</xref>). To control the potential confusing factors of breastfeeding and weaning diets, for comparisons, individuals were classified in two categories (i.e., preadults and adults) according to the tissues (i.e., teeth or bones) that originate the values (AlQahtani et al., <xref ref-type="bibr" rid="B3">2010</xref>).</p>
</sec>
<sec>
<title>Radiocarbon dates</title>
<p>The chronological framework of this work is based on 8 direct [Accelerator Mass Spectrometry (AMS) radiocarbon dates from individuals&#x00027; collagen, and fabrics from funerary contexts], and 12 indirect radiocarbon dates from associated material or stratigraphy (R. Shady personal communication, June 2024). Radiocarbon dates of Vichama come from the AMS facilities of three different laboratories using in-house protocols through commercial services. Individuals&#x00027; dates were calibrated (bce/ce) with Calib 8.1.0 package (Stuiver and Reimer, <xref ref-type="bibr" rid="B75">1993</xref>), using the Mixed Marine SoHCal 20 calibration curve (Heaton et al., <xref ref-type="bibr" rid="B31">2020</xref>; Hogg et al., <xref ref-type="bibr" rid="B34">2020</xref>), a local marine radiocarbon reservoir correction value (&#x00394;R = 200 &#x000B1; 20) estimated from paired samples (i.e., bone collagen and cotton fabrics) of the burial context ZAC7074. For individuals directly dated, we included the mean relative contribution of marine carbon to collagen calculated using Food Reconstruction Using Isotopic Transferred Signals (FRUITS) for each phase. Calibrated dates were rounded to 10 (see <xref ref-type="supplementary-material" rid="SM1">Supplementary material 1</xref>: S3). Individuals who were not directly dated were assigned to each period according to archaeological methods (i.e., stratigraphy, funerary patterns, pottery styles, and textiles&#x00027; decorations).</p>
</sec>
<sec>
<title>Laboratory procedures</title>
<p>First, samples were cleaned mechanically to remove surface contaminants. Then, samples were divided for collagen and apatite extractions. Collagen was extracted in the Institut de Ciencia i Tecnologia Ambientales of the Universitat Aut&#x000F2;noma de Barcelona (Spain).</p>
<p>Samples were physically cleaned and immersed in 0.6 M HCl at 4&#x000B0;C for several days, for demineralization, then rinsed with ultrapure water and immersed in 0.05 M sodium hydroxide (NaOH) for 20 min at room temperature. The NaOH wash cycles were repeated as needed, typically 1 or 2 times, until no further color change occurred in the solution. Samples were rinsed three times with ultrapure water for 10 min each to ensure the complete removal of NaOH. Samples were gelatinized in 0.001 M HCl (pH3) at 80&#x000B0;C for 48 h. The supernatant containing collagen was filtered using polyethylene Ezee filters (9 mL, pore size 60&#x02013;90 &#x003BC;m, Elkay Laboratories Ltd.), then frozen for at least 48 h at &#x02212;20&#x000B0;C, and freeze-dried. Stable isotope analysis of successfully extracted samples was performed at SUERC, East Kilbride (UK) using a Delta V Advantage continuous-flow isotope ratio mass spectrometer coupled via a ConfloIV to an IsoLink elemental analyzer (Thermo Scientific, Bremen, Germany). Bone collagen (approximately 0.7 mg) was combusted in the presence of oxygen in a single reactor containing tungstic oxide and copper wires at 1,020&#x000B0;C to produce nitrogen (N<sub>2</sub>) and carbon dioxide (CO<sub>2</sub>). A magnesium perchlorate trap was used to eliminate water produced during the combustion process, and the gases were separated in a Gas chromatography (GC) column heated between 70&#x000B0;C and 240&#x000B0;C. Helium was used as a carrier gas throughout the procedure. N<sub>2</sub> and CO<sub>2</sub> entered the mass spectrometer via an open split arrangement within the ConfloIV and were analyzed against their corresponding reference gases. The International Atomic Energy Agency reference materials USGS40 (L-glutamic acid, &#x003B4;<sup>13</sup>C Vienna Pee Dee Belemnite (V-PDB) = &#x02212;26.39 &#x000B1; 0.04&#x02030;, &#x003B4;<sup>15</sup>N (AIR) = &#x02212;4.52 &#x000B1; 0.06&#x02030;) and USGS41a (L-glutamic acid, &#x003B4;13C (V-PDB) = 36.55 &#x000B1; 0.08&#x02030;, &#x003B4;<sup>15</sup>N (AIR) = 47.55 &#x000B1; 0.15&#x02030;) were used to normalize &#x003B4;<sup>15</sup>N and &#x003B4;<sup>13</sup>C values. Normalization was checked using the well-characterized Elemental Microanalysis IRMS fish gelatin standard B2215 (&#x003B4;<sup>13</sup>C (V-PDB) = &#x02212;22.92 &#x000B1; 0.10&#x02030;, &#x003B4;<sup>15</sup>N (AIR) = 4.26 &#x000B1; 0.12&#x02030;) and/or USGS88 (marine collagen, &#x003B4;<sup>13</sup>C (V-PDB) = &#x02212;16.06 &#x000B1; 0.07&#x02030;, &#x003B4;<sup>15</sup>N (AIR) = 14.96 &#x000B1; 0.14&#x02030;). Precision is &#x000B1; 0.2&#x02030; for &#x003B4;<sup>15</sup>N and &#x000B1; 0.1&#x02030; for &#x003B4;<sup>13</sup>C.</p>
<p>For bioapatite analysis, bone samples were grounded with an agate mortar and pestle and processed in the Environmental Isotope Laboratory from the University of Arizona (USA). In this case, bone samples were cleaned in distilled water with an ultrasonic cleaner and powdered using a mortar and pestle. Samples weighing &#x0007E;10 mg each were soaked for 24&#x02013;72 h in a 2% sodium hypochlorite solution to remove the organic material. Then, the samples were treated with 0.1 M acetic acid to remove exogenous carbonates. The samples were also allowed to react with dehydrated phosphoric acid and silver (silver removes the sulfur gases from the evolved CO<sub>2</sub>) at 70&#x000B0;C in the KIEL-III Sample Preparation system. The &#x003B4;<sup>13</sup>C values of the resultant CO<sub>2</sub> were measured using the Finnigan MAT252 Isotope Ratio Mass Spectrometer. The calibration of the isotopic ratios was based on repeated measurements of NBS-19 and NBS-18 with a precision of &#x000B1;0.08&#x02030; for &#x003B4;<sup>13</sup>C (1&#x003C3;).</p>
<p>Only individuals with valid isotope values of collagen were included in the analysis. The reliability of collagen was validated following current accepted conventional preservation criteria: C/N ratio between 2.9 and 3.6 (DeNiro, <xref ref-type="bibr" rid="B16">1985</xref>) and wt%C of 26%&#x02212;44% and wt%N of 11%&#x02212;16% (Van Klinken, <xref ref-type="bibr" rid="B79">1999</xref>). We used a subsample of 10 individuals to assess the preservation of bioapatite (i.e., mineralogy and crystallinity) with ATR-FTIR.</p>
<p>ATR-FTIR screening was conducted at the Laboratory of Molecular Spectroscopy, Servei de Quimica&#x02014;Universitat Aut&#x000F2;noma de Barcelona using a Bruker Alpha II FTIR compact spectrometer, coupled to an Eco-ATR. Spectra of 1 mg of bone powder were collected in absorbance (A) mode with a spectral range of 400 to 4,000 cm<sup>&#x02212;1&#x000B0;</sup> (<inline-graphic xlink:href="fearc-04-1611071-i0001.tif"/>), for 100 scans and a resolution of 8 cm<sup>&#x02212;1</sup>. Spectra generation and baseline corrections were performed with OPUS 7.8 software. To evaluate bioapatite diagenesis, we assessed five indexes (IRSF, Infrared Splitting Factor; C/P, Carbon/Phosphate; C/C, Carbon/Carbon; BPI, Type B Phosphate index; API, Type A Phophate index) relative to conventional values for well-preserved archaeological samples (France et al., <xref ref-type="bibr" rid="B26">2020</xref>). See <xref ref-type="supplementary-material" rid="SM1">Supplementary material 1</xref>: S4 for ATR-FTIR raw data and indexes.</p>
</sec>
<sec>
<title>Isotopic analysis and BSIMM models</title>
<p>To identify trends of isotope values we used the &#x0201C;classic&#x0201D; scatterplots of isotopic values (i.e., &#x003B4;<sup>13</sup>C<sub>coll</sub> vs. &#x003B4;<sup>15</sup>N; &#x003B4;<sup>13</sup>C<sub>coll</sub> vs. &#x003B4;<sup>13</sup>C<sub>ap</sub>) and rKin -Standard Ellipse Areas (Albeke, <xref ref-type="bibr" rid="B2">2017</xref>). After evaluation for normal distribution with Shapiro&#x02013;Wilk test for normality (&#x003B1; = 0.05), descriptive statistics of isotope values and comparisons by sex (i.e., males, females and undetermined) and wide age-categories (i.e., preadults and adults) were performed using Kruskal&#x02013;Wallis (&#x003B1; =0.05) and Mann&#x02013;Whitney (&#x003B1; = 0.05) tests with SPSS v.29 (Microsoft<sup>&#x000AE;</sup>). Kernel Density analysis and graphics were performed with the function <italic>geom_density()</italic> of the ggplot2 package in R environment.</p>
<p>The BSIMM software FRUITS (v. 2.1.1 Beta program; Fernandes et al., <xref ref-type="bibr" rid="B24">2014</xref>, <xref ref-type="bibr" rid="B23">2015</xref>), was employed to estimate the proportional calorie contribution of different food sources to the diet using three dietary proxies &#x003B4;<sup>13</sup>C<sub>coll</sub>, &#x003B4;<sup>15</sup>N and &#x003B4;<sup>13</sup>C<sub>ap</sub> (see <xref ref-type="supplementary-material" rid="SM2">Supplementary material 2</xref> for the FRUITS model&#x00027;s implementation). In this research, we run individuals&#x00027; estimations. The <italic>consumer data</italic> for individuals consist of their &#x003B4;<sup>13</sup>C<sub>col</sub>, &#x003B4;<sup>15</sup>N<sub>col</sub>, and &#x003B4;<sup>13</sup>C<sub>ap</sub> isotopic values. For Vichama, as a group, we use the average of isotope values and the standard error of the mean.</p>
<p>We charged our model with four potential food sources consumed (<italic>Food groups</italic>), with their respective composition of macro-nutrients (<italic>Food fractions:</italic> bulk, protein, and energy): terrestrial fauna, providing proteins and lipids; marine fauna, providing proteins and lipids; and C<sub>3</sub> and C<sub>4</sub> plants, providing carbohydrates and proteins. The isotopic mean values of these food groups come from previous dietary reconstructions for the Andean Central Coast (Pezo-Lanfranco et al., <xref ref-type="bibr" rid="B53">2022</xref>; see <xref ref-type="supplementary-material" rid="SM1">Supplementary material 1</xref>: S7).</p>
<p>In the Central Coast of Peru, C<sub>3</sub> plants (including tubers, legumes, annuals, and fruits) have mean &#x003B4;<sup>13</sup>C and &#x003B4;<sup>15</sup>N values of &#x02212;25.7 &#x000B1; 2.1&#x02030; and &#x0002B;5.3 &#x000B1; 3.2&#x02030;, respectively. The C<sub>4</sub> plants (restricted to <italic>Zea mays</italic> in such cases) have mean &#x003B4;<sup>13</sup>C and &#x003B4;<sup>15</sup>N values of &#x02212;11.3 &#x000B1; 1.4&#x02030; and &#x0002B;7.3 &#x000B1; 3.0&#x02030;. The CAM plants have &#x003B4;<sup>13</sup>C mean values of &#x02212;10.8 &#x000B1; 0.2&#x02030; that overlap with C4, and &#x003B4;<sup>15</sup>N of &#x0002B;3.8 &#x000B1; 0.4&#x02030;. Terrestrial faunal (e.g., cervids, rodents, and camelids) show &#x003B4;<sup>13</sup>C<sub>coll</sub> and &#x003B4;<sup>15</sup>N mean values ranging between &#x02212;17.5 &#x000B1; 3.2&#x02030; and &#x0002B;9.0 &#x000B1; 3.1&#x02030;. The &#x003B4;<sup>13</sup>C and &#x003B4;<sup>15</sup>N mean values for marine fauna (including fishes, little fishes such as anchovy and sardines, mollusks, and marine birds and mammals from modern and archaeological origin) are &#x02212;11.9 &#x000B1; 2.1&#x02030; and &#x0002B;13.2 &#x000B1; 3.0&#x02030;, respectively (Pezo-Lanfranco and Colonese, <xref ref-type="bibr" rid="B51">2024</xref>). The &#x003B4;<sup>13</sup>C values from modern fauna specimens were corrected for the &#x0201C;fossil fuel effect&#x0201D; adjusting the values by &#x0002B;1.5&#x02030; (Marino and McElroy, <xref ref-type="bibr" rid="B47">1991</xref>). To avoid issues with preservation (Szpak and Chiou, <xref ref-type="bibr" rid="B76">2019</xref>), only values from modern plants were used to feed the model.</p>
<p>Isotope fractionation factors, derived from experimental studies, were established at &#x0002B;4.8 &#x000B1; 0.5&#x02030; between diet and collagen, &#x0002B;10.1 &#x000B1; 0.5&#x02030; between diet and apatite, &#x0002B;5.5 &#x000B1; 0.5 &#x02030; for &#x003B4;<sup>15</sup>N diet to collagen (Fernandes et al., <xref ref-type="bibr" rid="B24">2014</xref>, <xref ref-type="bibr" rid="B23">2015</xref>).</p>
<p>The weighted values of each fraction of macronutrients (lipids, carbohydrates, and proteins) followed published parameters (Fernandes et al., <xref ref-type="bibr" rid="B24">2014</xref>, <xref ref-type="bibr" rid="B23">2015</xref>; Pezo-Lanfranco et al., <xref ref-type="bibr" rid="B53">2022</xref>; Pezo-Lanfranco and Colonese, <xref ref-type="bibr" rid="B51">2024</xref>). The values of &#x003B4;<sup>13</sup>C<sub>ap</sub> represent the total carbon mix in the diet. Therefore, we use the same bulk value for each food group. Terrestrial and marine fauna &#x003B4;<sup>13</sup>C bulk values were estimated as a weighted mean of lipid and protein &#x003B4;<sup>13</sup>C values (Fernandes et al., <xref ref-type="bibr" rid="B24">2014</xref>, <xref ref-type="bibr" rid="B23">2015</xref>). For collagen, the carbon of the protein and energy routed to the total collagen was established at 74 &#x000B1; 4% and 26%, respectively (Fernandes et al., <xref ref-type="bibr" rid="B25">2012</xref>). We assumed that nitrogen was derived exclusively from proteins (100%). Lipids and carbohydrates were added to the model as &#x0201C;energy&#x0201D;. To integrate this combinatorial effect, we applied a &#x0201C;concentration-dependent&#x0201D; model (Fernandes et al., <xref ref-type="bibr" rid="B24">2014</xref>, <xref ref-type="bibr" rid="B23">2015</xref>).</p>
<p>Following previous reconstructions using FRUITS (Fernandes et al., <xref ref-type="bibr" rid="B24">2014</xref>, <xref ref-type="bibr" rid="B23">2015</xref>; Pezo-Lanfranco et al., <xref ref-type="bibr" rid="B53">2022</xref>) the isotopic composition of each nutritional fraction (protein, carbohydrates, and lipids) was obtained from the mean values of &#x003B4;<sup>13</sup>C<sub>coll</sub> and &#x003B4;<sup>15</sup>N<sub>coll</sub> using the following fractionation factors: &#x02212;2&#x02030; (&#x00394;<sup>13</sup>C<sub>protein &#x02212; collagen</sub>), &#x02212;8&#x02030; (&#x00394;<sup>13</sup>C<sub>l&#x000ED;<italic>pids</italic>&#x02212;<italic>collagen</italic></sub>), and &#x0002B;2&#x02030; (&#x00394;<sup>15</sup>N<sub>protein &#x02212; collagen</sub>) for terrestrial mammals and &#x02212;1&#x02030; (&#x00394;<sup>13</sup>C<sub>protein &#x02212; collagen</sub>), &#x02212;7&#x02030; (&#x00394;<sup>13</sup>C<sub>l&#x000ED;<italic>pids</italic>&#x02212;<italic>collagen</italic></sub>), and &#x0002B;2&#x02030; (&#x00394;<sup>15</sup>N<sub>protein &#x02212; collagen</sub>) for marine animals. For plants, the offsets were &#x02212;2&#x02030; (&#x00394;<sup>13</sup>C<sub>bulk &#x02212; protein</sub>) and &#x0002B;0.5&#x02030; (&#x00394;<sup>13</sup>C<sub>bulk &#x02212; lipids</sub>), while for the &#x003B4;<sup>15</sup>N value of plant protein, the known value of &#x003B4;<sup>15</sup>N recorded for the plant was assumed. The carbon weight (concentrations) of each food fraction (protein and energy) from each food group was calculated according to its macronutrient composition (Fernandes et al., <xref ref-type="bibr" rid="B24">2014</xref>, <xref ref-type="bibr" rid="B23">2015</xref>; Pezo-Lanfranco et al., <xref ref-type="bibr" rid="B53">2022</xref>; see details in <xref ref-type="supplementary-material" rid="SM2">Supplementary material 2</xref>). A physiological, conservative, and acceptable range of protein consumption stipulated between 5% and 45% of the total calories was charged as &#x0201C;prior&#x0201D; (Fernandes et al., <xref ref-type="bibr" rid="B24">2014</xref>). The estimates of FRUITS models reflect carbon content or equivalent calorie contributions expressed as relative contributions (adding up to 1 or 100%) with an associated 1&#x003C3; uncertainty (Fernandes et al., <xref ref-type="bibr" rid="B23">2015</xref>).</p>
<p>Finally, a Kullback&#x02013;Leibler divergence test (Lyman, <xref ref-type="bibr" rid="B44">2008</xref>) was performed to estimate the statistical difference between the probability distributions of the four food groups between periods (EF-1 vs. LIP). This test was executed with the <italic>Entropy</italic> package of the R environment (<xref ref-type="supplementary-material" rid="SM2">Supplementary material 2</xref>).</p>
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</sec>
</body>
<back>
<sec sec-type="data-availability" id="s5">
<title>Data availability statement</title>
<p>The datasets presented in this study can be found in online repositories. The names of the repository/repositories and accession number(s) can be found in the article/<xref ref-type="supplementary-material" rid="SM1">Supplementary material</xref>.</p>
</sec>
<sec sec-type="author-contributions" id="s6">
<title>Author contributions</title>
<p>LP-L: Conceptualization, Data curation, Formal analysis, Funding acquisition, Investigation, Methodology, Project administration, Resources, Software, Supervision, Validation, Visualization, Writing &#x02013; original draft, Writing &#x02013; review &#x00026; editing. AC: Conceptualization, Data curation, Investigation, Project administration, Supervision, Validation, Visualization, Writing &#x02013; original draft, Writing &#x02013; review &#x00026; editing. AP-B: Data curation, Investigation, Validation, Visualization, Writing &#x02013; original draft, Writing &#x02013; review &#x00026; editing. TA: Data curation, Investigation, Validation, Visualization, Writing &#x02013; original draft, Writing &#x02013; review &#x00026; editing. MM: Investigation, Project administration, Supervision, Validation, Writing &#x02013; original draft, Writing &#x02013; review &#x00026; editing. MY: Data curation, Investigation, Validation, Writing &#x02013; original draft, Writing &#x02013; review &#x00026; editing. MG: Data curation, Investigation, Validation, Writing &#x02013; original draft, Writing &#x02013; review &#x00026; editing. LM: Data curation, Investigation, Validation, Writing &#x02013; original draft, Writing &#x02013; review &#x00026; editing. JA: Data curation, Investigation, Validation, Writing &#x02013; original draft, Writing &#x02013; review &#x00026; editing. AD: Data curation, Formal analysis, Software, Visualization, Writing &#x02013; original draft, Writing &#x02013; review &#x00026; editing. PN: Conceptualization, Data curation, Funding acquisition, Investigation, Project administration, Resources, Supervision, Validation, Visualization, Writing &#x02013; original draft, Writing &#x02013; review &#x00026; editing. ACC: Conceptualization, Data curation, Formal analysis, Funding acquisition, Investigation, Methodology, Software, Supervision, Writing &#x02013; original draft, Writing &#x02013; review &#x00026; editing. RS: Conceptualization, Data curation, Funding acquisition, Investigation, Project administration, Resources, Supervision, Validation, Writing &#x02013; original draft, Writing &#x02013; review &#x00026; editing.</p>
</sec>
<sec sec-type="funding-information" id="s7">
<title>Funding</title>
<p>The author(s) declare that financial support was received for the research and/or publication of this article. The Vichama project was funded by the <italic>Zona Arqueol&#x000F3;gica Caral Supe</italic> (UE 003, Ministry of Culture of Peru). The research has also received funding from the European Union&#x00027;s Horizon 2020 research and innovation program under the Marie Sklodowska-Curie Action project PACHAMAMA, Grant Agreement N<sup>o</sup>. 101062179 (LP-L) and the ERC Consolidator project TRADITION, Grant Agreement N<sup>o</sup>. 817911 (ACC). This work contributes to the &#x0201C;ICTA-UAB Mar&#x000ED;a de Maeztu&#x0201D; Program for Units of Excellence of the Spanish Ministry of Science and Innovation (CEX 2019-000940-M). This work also contributes to EarlyFoods (Evolution and impact of early food production systems), funded by the Ag&#x000E8;ncia de Gesti&#x000F3; d&#x00027;Ajuts Universitaris i de Recerca de Catalunya (SGR-Cat-2021, 00527). The funders had no role in the study design, data collection and analysis, the decision to publish, or the preparation of the manuscript.</p>
</sec>
<ack><p>The authors are grateful to Sonia Lopez and her field team for their valuable collaboration during the sampling work.</p>
</ack>
<sec sec-type="COI-statement" id="conf1">
<title>Conflict of interest</title>
<p>The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest.</p>
</sec>
<sec id="s8">
<title>Generative AI statement</title>
<p>The author(s) declare that no Gen AI was used in the creation of this manuscript.</p></sec>
<sec sec-type="disclaimer" id="s9">
<title>Publisher&#x00027;s note</title>
<p>All claims expressed in this article are solely those of the authors and do not necessarily represent those of their affiliated organizations, or those of the publisher, the editors and the reviewers. Any product that may be evaluated in this article, or claim that may be made by its manufacturer, is not guaranteed or endorsed by the publisher.</p>
</sec>
<sec sec-type="supplementary-material" id="s10">
<title>Supplementary material</title>
<p>The Supplementary Material for this article can be found online at: <ext-link ext-link-type="uri" xlink:href="https://www.frontiersin.org/articles/10.3389/fearc.2025.1611071/full#supplementary-material">https://www.frontiersin.org/articles/10.3389/fearc.2025.1611071/full#supplementary-material</ext-link></p>
<supplementary-material xlink:href="Table_1.xlsx" id="SM1" mimetype="application/vnd.openxmlformats-officedocument.spreadsheetml.sheet" xmlns:xlink="http://www.w3.org/1999/xlink"/>
<supplementary-material xlink:href="Data_Sheet_1.docx" id="SM2" mimetype="application/vnd.openxmlformats-officedocument.wordprocessingml.document" xmlns:xlink="http://www.w3.org/1999/xlink"/>
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