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<front>
<journal-meta>
<journal-id journal-id-type="publisher-id">Front. Ecol. Evol.</journal-id>
<journal-title>Frontiers in Ecology and Evolution</journal-title>
<abbrev-journal-title abbrev-type="pubmed">Front. Ecol. Evol.</abbrev-journal-title>
<issn pub-type="epub">2296-701X</issn>
<publisher>
<publisher-name>Frontiers Media S.A.</publisher-name>
</publisher>
</journal-meta>
<article-meta>
<article-id pub-id-type="doi">10.3389/fevo.2022.1066680</article-id>
<article-categories>
<subj-group subj-group-type="heading">
<subject>Ecology and Evolution</subject>
<subj-group>
<subject>Original Research</subject>
</subj-group>
</subj-group>
</article-categories>
<title-group>
<article-title>Behavioral strategies of prehistoric and historic children from dental microwear texture analysis</article-title>
</title-group>
<contrib-group>
<contrib contrib-type="author" corresp="yes"><name><surname>Estalrrich</surname><given-names>Almudena</given-names></name>
<xref rid="aff1" ref-type="aff"><sup>1</sup></xref>
<xref rid="aff2" ref-type="aff"><sup>2</sup></xref>
<xref rid="c001" ref-type="corresp"><sup>&#x002A;</sup></xref>
<xref rid="fn0001" ref-type="author-notes"><sup>&#x2020;</sup></xref>
<uri xlink:href="https://loop.frontiersin.org/people/2047589/overview"/>
</contrib>
<contrib contrib-type="author"><name><surname>Krueger</surname><given-names>Kristin L.</given-names></name>
<xref rid="aff3" ref-type="aff"><sup>3</sup></xref>
<xref rid="fn0001" ref-type="author-notes"><sup>&#x2020;</sup></xref>
<uri xlink:href="https://loop.frontiersin.org/people/2089236/overview"/>
</contrib>
</contrib-group>
<aff id="aff1"><sup>1</sup><institution>Grupo de I+D+i EVOADAPTA (Evoluci&#x00F3;n Humana y Adaptaciones Econ&#x00F3;micas y Ecol&#x00F3;gicas durante la Prehistoria), Departamento Ciencias Hist&#x00F3;ricas, Universidad de Cantabria</institution>, <addr-line>Santander</addr-line>, <country>Spain</country></aff>
<aff id="aff2"><sup>2</sup><institution>Ungar Lab, Department of Anthropology, University of Arkansas</institution>, <addr-line>Fayetteville, AR</addr-line>, <country>United States</country></aff>
<aff id="aff3"><sup>3</sup><institution>Department of Anthropology, Loyola University Chicago</institution>, <addr-line>Chicago, IL</addr-line>, <country>United States</country></aff>
<author-notes>
<fn id="fn0002" fn-type="edited-by">
<p>Edited by: Maciej Tomasz Krajcarz, Institute of Geological Sciences (PAN), Poland</p>
</fn>
<fn id="fn0003" fn-type="edited-by">
<p>Reviewed by: Pawe&#x0142; D&#x0105;browski, Wroclaw Medical University, Poland; Yuichi Naito, Central Research Institute of Electric Power Industry (CRIEPI), Japan</p>
</fn>
<corresp id="c001">&#x002A;Correspondence: Almudena Estalrrich, <email>aestalrrichalbo@gmail.com</email></corresp>
<fn id="fn0001" fn-type="equal">
<p><sup>&#x2020;</sup>These authors have contributed equally to this work</p>
</fn>
<fn id="fn0004" fn-type="other">
<p>This article was submitted to Paleoecology, a section of the journal Frontiers in Ecology and Evolution</p>
</fn>
</author-notes>
<pub-date pub-type="epub">
<day>23</day>
<month>12</month>
<year>2022</year>
</pub-date>
<pub-date pub-type="collection">
<year>2022</year>
</pub-date>
<volume>10</volume>
<elocation-id>1066680</elocation-id>
<history>
<date date-type="received">
<day>11</day>
<month>10</month>
<year>2022</year>
</date>
<date date-type="accepted">
<day>05</day>
<month>12</month>
<year>2022</year>
</date>
</history>
<permissions>
<copyright-statement>Copyright &#x00A9; 2022 Estalrrich and Krueger.</copyright-statement>
<copyright-year>2022</copyright-year>
<copyright-holder>Estalrrich and Krueger</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>
<sec>
<title>Introduction</title>
<p>Reconstructing the dietary and behavioral strategies of our hominin ancestors is crucial to understanding their evolution, adaptation, and overall way of life. Teeth in general, and dental microwear specifically, provide a means to examine these strategies, with posterior teeth well positioned to tell us about diet, and anterior teeth helping us examine non-dietary tooth-use behaviors. Past research predominantly focused on strategies of adult individuals, leaving us to wonder the role children may have played in the community at large. Here we begin to address this by analyzing prehistoric and historic children through dental microwear texture analysis of deciduous anterior teeth.</p>
</sec>
<sec>
<title>Materials and Methods</title>
<p>Four sample groups were used: Neandertals (<italic>N</italic>&#x2009;=&#x2009;8), early modern humans (<italic>N</italic>&#x2009;=&#x2009;14), historic Egyptians from Amarna (<italic>N</italic>&#x2009;=&#x2009;19) and historic high-Arctic Inuit from Point Hope, Alaska (<italic>N</italic>&#x2009;=&#x2009;6). Anterior deciduous teeth were carefully cleaned, molded, and cast with high-resolution materials. Labial surfaces were scanned for dental microwear textures using two white-light confocal microscopes at the University of Arkansas, and a soft filter applied to facilitate data comparisons.</p>
</sec>
<sec>
<title>Results and Discussion</title>
<p>Results show that dental microwear texture analysis successfully differentiated the samples by all texture variables examined (anisotropy, complexity, scale of maximum complexity, and two variants of heterogeneity). Interestingly, the Neandertal and Point Hope children had similar mean values across all the texture variables, and both groups were significantly different from the Amarna, Egyptian children. These differences suggest diversity in abrasive load exposure and participation in non-dietary anterior tooth-use behaviors. Further analyses and an expanded sample size will help to strengthen the data presented here, but our results show that some prehistoric and historic children took part in similar behaviors as their adult counterparts.</p>
</sec>
</abstract>
<kwd-group>
<kwd>labial surface</kwd>
<kwd>deciduous enamel</kwd>
<kwd>dietary reconstruction</kwd>
<kwd>prehistoric children</kwd>
<kwd>historic populations</kwd>
<kwd>Neandertal</kwd>
</kwd-group>
<counts>
<fig-count count="2"/>
<table-count count="1"/>
<equation-count count="0"/>
<ref-count count="53"/>
<page-count count="8"/>
<word-count count="5441"/>
</counts>
</article-meta>
</front>
<body>
<sec id="sec1" sec-type="intro">
<title>1. Introduction</title>
<p>Dental microwear texture analysis (DMTA) is widely recognized as a useful method to highlight differences in both dietary and behavioral strategies of fossil and modern hominins (<xref ref-type="bibr" rid="ref58">Scott et al., 2005</xref>, <xref ref-type="bibr" rid="ref57">2006</xref>; <xref ref-type="bibr" rid="ref69">Ungar et al., 2008</xref>, <xref ref-type="bibr" rid="ref71">2010</xref>, <xref ref-type="bibr" rid="ref70">2012</xref>; <xref ref-type="bibr" rid="ref20">El Zaatari, 2010</xref>; <xref ref-type="bibr" rid="ref40">Krueger and Ungar, 2010</xref>, <xref ref-type="bibr" rid="ref41">2012</xref>; <xref ref-type="bibr" rid="ref22">El Zaatari et al., 2011</xref>; <xref ref-type="bibr" rid="ref24">Estalrrich et al., 2017</xref>). While molar microwear have demonstrated to be especially valuable as a dietary proxy (e.g., <xref ref-type="bibr" rid="ref58">Scott et al., 2005</xref>; <xref ref-type="bibr" rid="ref19">El Zaatari, 2007</xref>; <xref ref-type="bibr" rid="ref69">Ungar et al., 2008</xref>, <xref ref-type="bibr" rid="ref71">2010</xref>), incisor microwear texture analyses are useful in understanding behavioral and dietary strategies, as well as abrasive load exposure (<xref ref-type="bibr" rid="ref37">Krueger, 2006</xref>; <xref ref-type="bibr" rid="ref40">Krueger and Ungar, 2010</xref>, <xref ref-type="bibr" rid="ref41">2012</xref>).</p>
<p>The majority of dental microwear research has focused on adult individuals using permanent enamel. Only a few examples have examined children and their deciduous dentition (<xref ref-type="bibr" rid="ref11">Bullington, 1991</xref>; <xref ref-type="bibr" rid="ref68">Toussaint et al., 2010</xref>; <xref ref-type="bibr" rid="ref34">Hlusko et al., 2013</xref>; <xref ref-type="bibr" rid="ref23">El Zaatari et al., 2014</xref>; <xref ref-type="bibr" rid="ref47">Mahoney et al., 2016</xref>; <xref ref-type="bibr" rid="ref2">Bas et al., 2020</xref>; <xref ref-type="bibr" rid="ref35">Kelly et al., 2020</xref>). Examinations of children&#x2019;s diet and behavior are usually limited to weaning and other types of dietary stress, as shown by skeletal indicators of malnutrition, dental enamel defects, and other feeding-practice studies (<xref ref-type="bibr" rid="ref59">Skinner, 1997</xref>; <xref ref-type="bibr" rid="ref46">Lewis, 2007</xref>; <xref ref-type="bibr" rid="ref50">Prowse et al., 2008</xref>; <xref ref-type="bibr" rid="ref13">Clement and Freyne, 2012</xref>). Even basic dental macrowear analyses in children are limited and are then only used for age estimation or social status purposes (<xref ref-type="bibr" rid="ref46">Lewis, 2007</xref>; <xref ref-type="bibr" rid="ref17">Dawson and Brown, 2013</xref>). Why is this the case?</p>
<p>The first difference is the number of teeth, with fewer deciduous than permanent teeth. This is important when considering available sample sizes between child and adult remains. Another distinction is the composition of deciduous and permanent enamel. Deciduous enamel is not only less mineralized than permanent enamel (92% vs. 96%), but also has a higher water content (<xref ref-type="bibr" rid="ref18">De Menezes Oliveira et al., 2010</xref>). These composition variations make deciduous enamel softer. Moreover, the mean thickness of deciduous enamel is less than half that of its permanent counterpart (1.14&#x2009;mm vs. 2.58&#x2009;mm, <xref ref-type="bibr" rid="ref18">De Menezes Oliveira et al., 2010</xref>). Collectively, these differences cause greater susceptibility to fracture, chipping, and wear in deciduous teeth. Add the limited sample size to these other differences, and it is unsurprising that research has focused on the dietary and behavior reconstruction of adult individuals.</p>
<p>However, there is another important reason dietary and behavioral reconstructions have favored adults and their permanent teeth: the under-representation of children in the archaeological and paleoanthropological record. This is due not only to the lower mortality rates in children, but also to taphonomic processes, which affect the preservation of fragile sub-adult bones and teeth (<xref ref-type="bibr" rid="ref46">Lewis, 2007</xref>; <xref ref-type="bibr" rid="ref48">McFadden et al., 2021</xref>). For example, a child&#x2019;s body skeletonizes faster, becomes readily disarticulated, and the smaller size makes them more attractive to scavengers, allowing for dispersal of body parts (<xref ref-type="bibr" rid="ref46">Lewis, 2007</xref>). Due to these phenomena, analyses of sub-adult bones and teeth are not as common as in adult individuals, and are limited to those specific, unique sites where children are present, and preservation is exceptional.</p>
<p>Challenges in studying deciduous teeth (and children in general) exist; however, there is evidence that significant information can be gleaned from what is preserved in the archaeological and fossil record. For example, a recent study on the anterior tooth-use behavior of Paleolithic children (<xref ref-type="bibr" rid="ref25">Estalrrich and Mar&#x00ED;n-Arroyo, 2021</xref>) revealed comparable behavioral patterns as their adult counterparts, despite these known differences between the deciduous and permanent enamel. These data, along with those demonstrating the efficacy of microwear textures in differentiating hominin anterior tooth-use behaviors in different ecological zones (<xref ref-type="bibr" rid="ref42">Krueger et al., 2017</xref>, <xref ref-type="bibr" rid="ref43">2019</xref>), show we need to push the boundaries of what we know &#x2013; or thought we could know &#x2013; about children in the past. The goal of this paper is to present and analyze the largest microwear texture dataset of deciduous anterior teeth of both fossil (Neandertals and early modern humans) and recent individuals (Amarna Egyptians and Point Hope Inuit), and, ultimately, to better recognize the role these children played in daily life.</p>
</sec>
<sec id="sec2" sec-type="materials|methods">
<title>2. Materials and methods</title>
<sec id="sec3">
<title>2.1. Materials</title>
<p>Statistical analyses have previously indicated that microwear textures do not differ significantly across anterior permanent dentition (<xref ref-type="bibr" rid="ref42">Krueger et al., 2017</xref>). Thus, we included all anterior tooth types to maximize the sample. We analyzed a sample of 47 deciduous incisor and canine teeth, including, based on their cultural context, Neandertal (<italic>N</italic> =&#x2009;8); early modern humans (<italic>N</italic>&#x2009;=&#x2009;14), and recent modern humans from the historic Egyptians from Amarna (<italic>N</italic>&#x2009;=&#x2009;19), and historic high-Arctic Inuit from Point Hope, Alaska (<italic>N</italic>&#x2009;=&#x2009;6). <xref ref-type="supplementary-material" rid="SM1">Supplementary Table S1</xref> provides details of the studied samples. All the samples studied here are samples curated at different museums, and each museum complies with the ethical issues addressed by each country. By us accessing those samples in order to make the molds and the study, we signed and agreed to follow the required ethical issues.</p>
</sec>
<sec id="sec4">
<title>2.2. Dental microwear texture analysis</title>
<p>The high-resolution replicas were used for analysis of both the fossil and recent human comparative samples. All molds and casts were prepared following standard microwear analysis protocols (<xref ref-type="bibr" rid="ref9">Bromage, 1987</xref>; <xref ref-type="bibr" rid="ref63">Teaford and Oyen, 1989</xref>). The labial surface of each specimen was gently cleaned with acetone using cotton swabs. President Jet regular body polysiloxane (Coltene-Whaledent) and Epotek 301 epoxy base and hardener (Epoxy Technologies) were used as the mediums for mold and cast production, respectively. Each tooth was examined for antemortem microwear on the labial surface, next to the incisal edge, using a Sensofar Pl&#x03BC; white-light confocal profiler, <italic>Connie</italic> (Solarius Development Inc., Sunnyvale, California) and Sensofar Pl&#x03BC; Neox confocal profiler, <italic>Wall-e</italic>, (Sensofar, Barcelona, Spain) both found at the Department of Anthropology of the University of Arkansas in Fayetteville.</p>
<p>With the Sensofar Pl&#x03BC; white-light confocal profiler four adjacent scans of the enamel surface were taken using a 100x objective lens, yielding a lateral point spacing of 0.18&#x2009;mm and individual fields of view of 138&#x2009;&#x00D7;&#x2009;102 &#x03BC;m, following <xref ref-type="bibr" rid="ref57">Scott et al. (2006)</xref>. We also used Sensofar Pl&#x03BC; Neox confocal profiler in white-light mode with a 100x objective to analyze some specimens. A stitched point cloud of 242&#x2009;&#x00D7;&#x2009;181 &#x03BC;m with a lateral spacing of 0.17&#x2009;&#x03BC;m and a published vertical resolution &#x003C;1&#x2009;nm was obtained for each surface.</p>
<p>Data from each specimen were then imported to MountainsMap software version 8 (DigitalSurf, Besan&#x00E7;on, France), where the scans were processed and calibrated applying the soft filter (<xref ref-type="bibr" rid="ref1">Arman et al., 2016</xref>) to ensure a standard data collection across different profilers. After this, the scale-sensitive fractal variables were calculated with the same software. Briefly, the variables considered are complexity, scale of maximum complexity, anisotropy, and two variants of heterogeneity (<xref ref-type="bibr" rid="ref57">Scott et al., 2006</xref>). Complexity or area-scale fractal complexity (<italic>Asfc</italic>) measures the change in surface roughness at different scales. Scale of maximum complexity (<italic>Smc</italic>) measures the fine scale limit of the steepest part of the curve described for the Asfc measure. Surfaces dominated by large features on a microscopic scale would have a high <italic>Smc</italic>. Anisotropy (<italic>epLsar</italic>) measures the degree of directionality in surface roughness at a fine scale. Heterogeneity of area-scale fractal complexity (<italic>HAsfc</italic>) reflects variability of complexity across the surface. More heterogeneous surfaces will have higher values. Two forms of this variable are used here: <italic>HAsfc</italic> 3&#x2009;&#x00D7;&#x2009;3 (<italic>HAsfc<sub>9</sub></italic>) and <italic>HAsfc</italic> 9&#x2009;&#x00D7;&#x2009;9 (<italic>HAsfc<sub>81</sub></italic>).</p>
</sec>
<sec id="sec5">
<title>2.3. Statistical analyses</title>
<p>Independent-Samples Kruskal-Wallis tests were completed with the four groups (Neandertal, early modern humans, and recent modern humans from Amarna Egyptians, and Point Hope Inuit) as independent variables and microwear texture variables (<italic>epLsar, Asfc, Smc, HAsfc<sub>9</sub> and HAsfc<sub>81</sub></italic>) as dependent. Non-parametric pairwise comparisons to find sources of significant differences in the Kruskal-Wallis tests were then completed. Significance values have been adjusted by the Bonferroni correction for multiple comparisons tests in <xref ref-type="supplementary-material" rid="SM5">Supplementary Tables S5a&#x2013;e</xref>. These non-parametric tests were selected as they do not assume normality, are less sensitive to outliers, and appropriate given our limited sample sizes (G. Matthews, pers. comm.). It is important to note we found the same results with both parametric and non-parametric tests.</p>
</sec>
</sec>
<sec id="sec6" sec-type="results">
<title>3. Results</title>
<p>Photosimulations of the occlusal surfaces of selected teeth are shown in <xref rid="fig1" ref-type="fig">Figure 1</xref>. Kruskal-Wallis and pairwise comparisons are represented visually in <xref rid="fig2" ref-type="fig">Figure 2</xref>. Descriptive statistics for each group are provided in <xref rid="tab1" ref-type="table">Table 1</xref>. Individual microwear texture values are provided in <xref ref-type="supplementary-material" rid="SM2">Supplementary Table S2</xref>, as well as the test for normality of the microwear data (<xref ref-type="supplementary-material" rid="SM3">Supplementary Table S3</xref>), Kruskal&#x2013;Wallis results (<xref ref-type="supplementary-material" rid="SM4">Supplementary Table S4</xref>), and pairwise comparisons (<xref ref-type="supplementary-material" rid="SM5">Supplementary Tables S5a&#x2013;e</xref>).</p>
<fig position="float" id="fig1">
<label>Figure 1</label>
<caption>
<p>Photosimulation (left) and 3D view (right) of <bold>(A)</bold> Neandertal Krapina 13, a deciduous left lower lateral incisor; <bold>(B)</bold> Early modern human from Saint Germain 1970-7-4, a deciduous incisor; <bold>(C)</bold> Recent modern human from Amarna Egyptian SK304, a left deciduous central incisor; <bold>(D)</bold> Recent modern human from Point Hope 108, an upper left deciduous central incisor.</p>
</caption>
<graphic xlink:href="fevo-10-1066680-g001.tif"/>
</fig>
<fig position="float" id="fig2">
<label>Figure 2</label>
<caption>
<p>Violin plots with pairwise comparisons of ranked microwear data by variable. Two dotted lines and one single dashed line within each violin plot represent quartiles and median, respectively. &#x002A;&#x2009;=&#x2009;significant difference and ns&#x2009;=&#x2009;no significant difference between the two groups.</p>
</caption>
<graphic xlink:href="fevo-10-1066680-g002.tif"/>
</fig>
<table-wrap position="float" id="tab1">
<label>Table 1</label>
<caption>
<p>Summary descriptive statistics for the groups studied.</p>
</caption>
<table frame="hsides" rules="groups">
<thead>
<tr>
<th align="left" valign="top">Group</th>
<th align="center" valign="top">epLsar</th>
<th align="center" valign="top">Asfc Median</th>
<th align="center" valign="top">Smc</th>
<th align="center" valign="top">HAsfc9</th>
<th align="center" valign="top">HAsfc81</th>
</tr>
</thead>
<tbody>
<tr>
<td align="left" valign="top">NEAN MEAN</td>
<td align="left" valign="top">0.017775</td>
<td align="left" valign="top">2.1725</td>
<td align="left" valign="top">124.5487</td>
<td align="left" valign="top">0.4588</td>
<td align="left" valign="top">0.8988</td>
</tr>
<tr>
<td align="left" valign="top">NEAN SD</td>
<td align="left" valign="top">0.0010634</td>
<td align="left" valign="top">1.63835</td>
<td align="left" valign="top">198.82129</td>
<td align="left" valign="top">0.30334</td>
<td align="left" valign="top">0.42273</td>
</tr>
<tr>
<td align="left" valign="top">EMH MEAN</td>
<td align="left" valign="top">0.017186</td>
<td align="left" valign="top">1.4479</td>
<td align="left" valign="top">11.5193</td>
<td align="left" valign="top">0.4029</td>
<td align="left" valign="top">0.8164</td>
</tr>
<tr>
<td align="left" valign="top">EMH SD</td>
<td align="left" valign="top">0.0004944</td>
<td align="left" valign="top">0.86935</td>
<td align="left" valign="top">19.44259</td>
<td align="left" valign="top">0.20838</td>
<td align="left" valign="top">0.8091</td>
</tr>
<tr>
<td align="left" valign="top">AMARNA MEAN</td>
<td align="left" valign="top">0.016952</td>
<td align="left" valign="top">3.2281</td>
<td align="left" valign="top">8.2043</td>
<td align="left" valign="top">0.4686</td>
<td align="left" valign="top">0.8819</td>
</tr>
<tr>
<td align="left" valign="top">AMARNA SD</td>
<td align="left" valign="top">0.0010755</td>
<td align="left" valign="top">3.61266</td>
<td align="left" valign="top">4.10336</td>
<td align="left" valign="top">0.23627</td>
<td align="left" valign="top">0.33603</td>
</tr>
<tr>
<td align="left" valign="top">POINT HOPE MEAN</td>
<td align="left" valign="top">0.016967</td>
<td align="left" valign="top">1.5833</td>
<td align="left" valign="top">6.5483</td>
<td align="left" valign="top">0.3967</td>
<td align="left" valign="top">0.7417</td>
</tr>
<tr>
<td align="left" valign="top">PONT HOPE SD</td>
<td align="left" valign="top">0.0019086</td>
<td align="left" valign="top">0.73666</td>
<td align="left" valign="top">3.35328</td>
<td align="left" valign="top">0.16.269</td>
<td align="left" valign="top">0.23819</td>
</tr>
</tbody>
</table>
</table-wrap>
<p>Tests for normality were completed, and except for anisotropy, the microwear texture data were not normally distributed (<xref ref-type="supplementary-material" rid="SM3">Supplementary Table S3</xref>). As a result, nonparametric tests were used. The Independent-Samples Kruskal&#x2013;Wallis tests found statistically significant differences at the 0.05 level among the groups in all five microwear texture variables (<xref rid="fig2" ref-type="fig">Figure 2</xref>; <xref ref-type="supplementary-material" rid="SM4">Supplementary Table S4</xref>).</p>
<p>In every microwear texture variable analyzed here, the Neandertal and Point Hope children were significantly different from their Amarna counterparts (<xref rid="fig2" ref-type="fig">Figure 2</xref>; <xref ref-type="supplementary-material" rid="SM5">Supplementary Tables S5a&#x2013;S5e</xref>).</p>
</sec>
<sec id="sec7">
<title>4. Discussion and conclusions</title>
<p>This study examined a large sample of prehistoric and historic deciduous teeth to better understand the role children played within society. Were children behaving like their adult counterparts? If so, can we glean what those teeth-as-tools behaviors could have been? If not, at what age were they expected to contribute to the community at large? While this study has answered some of these questions, we also need to continue searching for more evidence.</p>
<p>The samples studied here include Neandertal and early modern human children from various sites and a wider time range, whereas the historic Point Hope and Amarna samples are from the same site and time (see <xref ref-type="supplementary-material" rid="SM1">Supplementary Table S1</xref>). We would expect more variability within the fossil microwear values, as they are geographically and temporally distinct. Interestingly, our data show that when analyzing these groups, every dental microwear texture variable used here distinguish the Neandertal and Point Hope children from those of the Amarna Egyptians.</p>
<p>This certainly is not the first time that Neandertal and high-Arctic aboriginal samples have been <xref ref-type="bibr" rid="ref22">El Zaatari et al., 2011</xref>. Indeed, decades of research, especially regarding anterior tooth-use behaviors, heavily associated Neandertals and their unique anterior tooth wear patterns with Arctic groups who used their anterior teeth as a clamp or third hand during animal hide processing (<xref ref-type="bibr" rid="ref4">Brace, 1967</xref>, <xref ref-type="bibr" rid="ref5">1975</xref>, <xref ref-type="bibr" rid="ref6">1979</xref>; <xref ref-type="bibr" rid="ref7">Brace and Molnar, 1967</xref>; <xref ref-type="bibr" rid="ref56">Ryan, 1980</xref>; <xref ref-type="bibr" rid="ref8">Brace et al., 1981</xref>). However, all these analyses focus on adult individuals. This is the first that links similar microwear textures between Neandertal and high-Arctic children. This suggests that the Neandertal and Point Hope Inuit children, at least those sampled here, were completing similar anterior tooth-use behaviors. Whether that means they were eating similar dietary items, had similar abrasive loads, and/or were completing tooth-use behaviors requires a deeper look at the values.</p>
<p>The microwear texture values presented here were collected using two different white-light confocal profilers, and a filter was applied to make these data comparable (<xref ref-type="bibr" rid="ref1">Arman et al., 2016</xref>); however, we have not applied that filter to other published microwear texture datasets. Even if we did, there are limited available datasets of deciduous teeth from which to make comparisons. A confounding issue is understanding if microwear forms similarly or differently between permanent and deciduous enamel, as studies have found conflicting results (<xref ref-type="bibr" rid="ref38">Krueger, 2016</xref>; <xref ref-type="bibr" rid="ref47">Mahoney et al., 2016</xref>; <xref ref-type="bibr" rid="ref35">Kelly et al., 2020</xref>). Therefore, our interpretations should be viewed with caution, and seen as preliminary until these issues are resolved.</p>
<p>Largely viewed within the realm of &#x201C;hunter-gatherers,&#x201D; Neandertals relied on a mixed diet and were highly dependent on the ecogeographic setting in which they lived (<xref ref-type="bibr" rid="ref22">El Zaatari et al., 2011</xref>; <xref ref-type="bibr" rid="ref26">Fiorenza et al., 2011</xref>). It is parsimonious to assume that their children relied on a similar diet and were also constrained by their environment. Indeed, stable isotope research of prehistoric juveniles in California suggests some were foraging independently, in addition to parent-provided resources, especially during high-stress times associated with social or climate change (<xref ref-type="bibr" rid="ref32">Greenwald et al., 2016</xref>; <xref ref-type="bibr" rid="ref28">Fournier et al., 2022</xref>). It is not unreasonable to assume that Neandertal children were subsisting on similar diets as their adult counterparts, and perhaps were even able to forage independently when the need arose.</p>
<p>Neandertal adults were found to perform different non-dietary anterior tooth-use behaviors based on their habitat (<xref ref-type="bibr" rid="ref42">Krueger et al., 2017</xref>). Using a comparative approach, it was found that Neandertals in cold, open environments had similar microwear textures to high-Arctic Alaskan aboriginal groups who used their anterior teeth in clamping and grasping behaviors related to animal hide preparation for clothing production. Other Neandertals in more mixed environments were using their anterior teeth for other behaviors, such as wood softening or cordage production (<xref ref-type="bibr" rid="ref42">Krueger et al., 2017</xref>). Interestingly, a preliminary study of Pleistocene deciduous teeth from northern Spain indicated they too showed the characteristic dental wear features associated to para-masticatory or cultural-related dental wear, including toothpick use (<xref ref-type="bibr" rid="ref25">Estalrrich and Mar&#x00ED;n-Arroyo, 2021</xref>). When previous analyses on diet and tooth-use behaviors are considered, they suggest that Neandertal adults and children were subsisting on similar dietary and behavioral strategies that are heavily influenced by their eco-geographic setting.</p>
<p>The Point Hope Inuit were also considered &#x201C;hunter-gatherers,&#x201D; and their diet largely consisted of land and sea mammals (especially caribou, whale, walrus, and seal), fish, and edible plants (<xref ref-type="bibr" rid="ref44">Larsen and Rainey, 1948</xref>; <xref ref-type="bibr" rid="ref45">Lester and Shapiro, 1968</xref>; <xref ref-type="bibr" rid="ref15">Dabbs, 2009</xref>; <xref ref-type="bibr" rid="ref10">Brubaker et al., 2010</xref>; <xref ref-type="bibr" rid="ref21">El Zaatari, 2014</xref>). They took part in non-dietary anterior tooth-use behaviors in the form of wood softening, clamping and grasping tasks related to hide preparation, and sinew cord production, and were, at times, subjected to high abrasive loads (<xref ref-type="bibr" rid="ref12">Burch, 1981</xref>; <xref ref-type="bibr" rid="ref27">Foote, 1992</xref>). Some of these individuals lived seasonally at Point Hope, while others lived there year-round, which is located 125&#x2009;miles north of the Arctic Circle (<xref ref-type="bibr" rid="ref44">Larsen and Rainey, 1948</xref>; <xref ref-type="bibr" rid="ref15">Dabbs, 2009</xref>).</p>
<p>On the other hand, the Amarna Egyptians were not &#x201C;hunter-gatherers,&#x201D; but were excavated from the non-elite South Tombs Cemetery and date from 3,300 to 3,280 BP (<xref ref-type="bibr" rid="ref54">Rose, 2006</xref>). This cemetery is composed of an estimated 5,000 individuals from different occupations and/or socio-economic positions but did not hold elite or royal status (<xref ref-type="bibr" rid="ref16">Dabbs et al., 2015</xref>). The excavated individuals showed high rates of subadult death, workload stress, trauma, and nutritional deficiencies (<xref ref-type="bibr" rid="ref55">Rose and Zabecki, 2009</xref>; <xref ref-type="bibr" rid="ref16">Dabbs et al., 2015</xref>). Adult microwear analysis suggests this sample was reliant on tough food, most likely bread, and the desert environment at Amarna would make sand a likely adherent abrasive (<xref ref-type="bibr" rid="ref39">Krueger and Scott, in press</xref>).</p>
<p>The Neandertal and Point Hope children, both from &#x201C;hunter-gatherer&#x201D; groups, had significantly lower complexity (<italic>Asfc</italic>), scale of maximum complexity (<italic>Smc</italic>) and heterogeneity (<italic>HAsfc<sub>9</sub></italic> and <italic>HAsfc<sub>81</sub></italic>) than their Amarna counterparts (<xref rid="tab1" ref-type="table">Table 1</xref>, <xref ref-type="supplementary-material" rid="SM2">Supplementary Table S2</xref>, <xref rid="fig2" ref-type="fig">Figure 2</xref>). While these variables are not often used in texture analyses of anterior teeth, they are useful for molar analyses, especially regarding the fracture properties of foods and abrasive loads. Here, we propose that the significantly lower values of these three variables for the Neandertal and Point Hope Inuit children show differences in abrasive loads from the Amarna children. While the Amarna children were subjected to a desert environment with little-to-no tree cover, their higher values may indicate their increased exposure to diverse types of abrasives that were adherent to their food. On the other hand, the lower values of the Neandertal and Point Hope Inuit children suggest more limited exposure to abrasives, which could be due to their reliance on a mixed diet.</p>
<p>The Neandertal and Point Hope children had significantly lower anisotropy (<italic>epLsar</italic>) than their Amarna counterparts. This variable is more heavily used in anterior tooth texture analyses and indicates the use of these teeth in non-dietary behaviors (e.g., clamping, grasping, tool retouching, etc.; <xref ref-type="bibr" rid="ref41">Krueger and Ungar, 2012</xref>; <xref ref-type="bibr" rid="ref42">Krueger et al., 2017</xref>, <xref ref-type="bibr" rid="ref43">2019</xref>). These results suggest both Neandertal and Point Hope children were taking part in non-dietary anterior tooth-use behaviors, while the Amarna children were not. While we are hesitant to suggest what specific types of behaviors in which these children may have been engaging, perhaps it was related to clamping and grasping behaviors like those found in their adult counterparts; however, an expanded sample size and comparative datasets are necessary to strengthen this idea.</p>
<p>It is worth noting that no statistically significant differences were found between the early modern human children and neither the Neandertal nor historic modern human counterparts (see <xref ref-type="supplementary-material" rid="SM5">Supplementary Tables S5a&#x2013;S5e</xref>). Perhaps this is simply a reflection of our limited sample size, and building this dataset is necessary to recognize potential differences. Or, perhaps this reflects a more diverse diet, abrasive load exposure, or landscape in which these children lived. However, these data provide the largest dataset from which to work in the future, and we look forward to continued analyses to reinforce or refute the ideas posited here.</p>
<p>In conclusion, these datasets provide a crucial pathway to understanding the role children played in the Paleolithic and beyond. Perhaps this is a starting point to investigating complex issues like independent foraging in Paleolithic children, especially considering the stress that climate change may have had on their dietary and behavioral strategies. It also helps us understand how teaching and learning may have transpired between adults and children, especially if the latter are performing non-dietary anterior tooth-use behaviors similar to their adult counterpart. We hope this creates a larger platform for additional analyses surrounding Paleolithic children, so that we may better recognize an entire community&#x2019;s contribution to survival.</p>
</sec>
<sec id="sec8" sec-type="data-availability">
<title>Data availability statement</title>
<p>The original contributions presented in the study are included in the article/<xref ref-type="supplementary-material" rid="SM1">Supplementary material</xref>, further inquiries can be directed to the corresponding author.</p>
</sec>
<sec id="sec9">
<title>Author contributions</title>
<p>AE and KK designed the research, analyzed data, and wrote the manuscript. All authors contributed to the article and approved the submitted version.</p>
</sec>
<sec id="sec10" sec-type="funding-information">
<title>Funding</title>
<p>The work was supported by the AE is supported by H2020-MSCA-IF project No. 891529 (3DFOSSILDIET).</p>
</sec>
<sec id="conf1" sec-type="COI-statement">
<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="sec100" sec-type="disclaimer">
<title>Publisher&#x2019;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>
</body>
<back>
<ack>
<p>We are sincerely grateful to Peter S. Ungar for allowing us the use of the microscope facilities at the Department of Anthropology in the University of Arkansas and encouraging discussions on microwear and diet. We also acknowledge Greg Matthews for his advice on statistical analyses and Emily Hallett and Jacopo Cerasoni for their input on data visualization. Thank you to Jean-Jacques Hublin, Manuel R. Gonz&#x00E1;lez Morales, David Frayer, Antonio Rosas, Sireen El Zaatari, F. Igor Guti&#x00E9;rrez Zugasti, Cristina Vega Maeso and Borja Gonz&#x00E1;lez Rabanal for access to some of the fossils sampled here. We thank Barry Kemp and Anna Stevens for permission to mold the Amarna individuals, Museo de Prehistoria y Arqueolog&#x00ED;a de Cantabria (Spain) for permission to mold the El Castillo tooth, and the American Museum of Natural History for permission to mold the Point Hope sample.</p>
</ack>
<sec id="sec12" sec-type="supplementary-material">
<title>Supplementary material</title>
<p>The Supplementary material for this article can be found online at: <ext-link xlink:href="https://www.frontiersin.org/articles/10.3389/fevo.2022.1066680/full#supplementary-material" ext-link-type="uri">https://www.frontiersin.org/articles/10.3389/fevo.2022.1066680/full#supplementary-material</ext-link></p>
<supplementary-material xlink:href="Table_1.pdf" id="SM1" mimetype="application/pdf" xmlns:xlink="http://www.w3.org/1999/xlink"/>
<supplementary-material xlink:href="Table_2.pdf" id="SM2" mimetype="application/pdf" xmlns:xlink="http://www.w3.org/1999/xlink"/>
<supplementary-material xlink:href="Table_3.pdf" id="SM3" mimetype="application/pdf" xmlns:xlink="http://www.w3.org/1999/xlink"/>
<supplementary-material xlink:href="Table_4.pdf" id="SM4" mimetype="application/pdf" xmlns:xlink="http://www.w3.org/1999/xlink"/>
<supplementary-material xlink:href="Table_5.pdf" id="SM5" mimetype="application/pdf" xmlns:xlink="http://www.w3.org/1999/xlink"/>
</sec>
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