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<front>
<journal-meta>
<journal-id journal-id-type="publisher-id">Front. Earth Sci.</journal-id>
<journal-title>Frontiers in Earth Science</journal-title>
<abbrev-journal-title abbrev-type="pubmed">Front. Earth Sci.</abbrev-journal-title>
<issn pub-type="epub">2296-6463</issn>
<publisher>
<publisher-name>Frontiers Media S.A.</publisher-name>
</publisher>
</journal-meta>
<article-meta>
<article-id pub-id-type="publisher-id">784313</article-id>
<article-id pub-id-type="doi">10.3389/feart.2021.784313</article-id>
<article-categories>
<subj-group subj-group-type="heading">
<subject>Earth Science</subject>
<subj-group>
<subject>Review</subject>
</subj-group>
</subj-group>
</article-categories>
<title-group>
<article-title>Animals for Tools: The Origin and Development of Bone Technologies in China</article-title>
<alt-title alt-title-type="left-running-head">Ma and Doyon</alt-title>
<alt-title alt-title-type="right-running-head">Animals for Tools in China</alt-title>
</title-group>
<contrib-group>
<contrib contrib-type="author">
<name>
<surname>Ma</surname>
<given-names>Shuwen</given-names>
</name>
<xref ref-type="aff" rid="aff1">
<sup>1</sup>
</xref>
<xref ref-type="fn" rid="fn1">
<sup>&#x2020;</sup>
</xref>
<uri xlink:href="https://loop.frontiersin.org/people/1561147/overview"/>
</contrib>
<contrib contrib-type="author" corresp="yes">
<name>
<surname>Doyon</surname>
<given-names>Luc</given-names>
</name>
<xref ref-type="aff" rid="aff1">
<sup>1</sup>
</xref>
<xref ref-type="aff" rid="aff2">
<sup>2</sup>
</xref>
<xref ref-type="corresp" rid="c001">&#x2a;</xref>
<xref ref-type="fn" rid="fn1">
<sup>&#x2020;</sup>
</xref>
<uri xlink:href="https://loop.frontiersin.org/people/1304116/overview"/>
</contrib>
</contrib-group>
<aff id="aff1">
<label>
<sup>1</sup>
</label>Institute of Cultural Heritage, Shandong University, <addr-line>Qingdao</addr-line>, <country>China</country>
</aff>
<aff id="aff2">
<label>
<sup>2</sup>
</label>Universit&#xe9; de Bordeaux, Centre National de la Recherche Scientifique UMR5199 &#x2013; PACEA, <addr-line>Pessac</addr-line>, <country>France</country>
</aff>
<author-notes>
<fn fn-type="edited-by">
<p>
<bold>Edited by:</bold> <ext-link ext-link-type="uri" xlink:href="https://loop.frontiersin.org/people/1221655/overview">Dongju Zhang</ext-link>, Lanzhou University, China</p>
</fn>
<fn fn-type="edited-by">
<p>
<bold>Reviewed by:</bold> <ext-link ext-link-type="uri" xlink:href="https://loop.frontiersin.org/people/1499621/overview">Justin Bradfield</ext-link>, University of Johannesburg, South Africa</p>
<p>
<ext-link ext-link-type="uri" xlink:href="https://loop.frontiersin.org/people/1378547/overview">Yanhua Song</ext-link>, Shanxi University, China</p>
</fn>
<corresp id="c001">&#x2a;Correspondence: Luc Doyon, <email>luc.doyon@umontreal.ca</email>
</corresp>
<fn fn-type="other" id="fn1">
<label>
<sup>&#x2020;</sup>
</label>
<p>
<bold>ORCID:</bold>
</p>
<p>Ma Shuwen, <ext-link ext-link-type="uri" xlink:href="http://orcid.org/0000-0001-5710-153X">orcid.org/0000-0001-5710-153X</ext-link>; Luc Doyon, <ext-link ext-link-type="uri" xlink:href="http://orcid.org/0000-0001-7163-6186">orcid.org/0000-0001-7163-6186</ext-link>
</p>
</fn>
<fn fn-type="other">
<p>This article was submitted to Paleontology, a section of the journal Frontiers in Earth Science</p>
</fn>
</author-notes>
<pub-date pub-type="epub">
<day>29</day>
<month>11</month>
<year>2021</year>
</pub-date>
<pub-date pub-type="collection">
<year>2021</year>
</pub-date>
<volume>9</volume>
<elocation-id>784313</elocation-id>
<history>
<date date-type="received">
<day>27</day>
<month>09</month>
<year>2021</year>
</date>
<date date-type="accepted">
<day>08</day>
<month>11</month>
<year>2021</year>
</date>
</history>
<permissions>
<copyright-statement>Copyright &#xa9; 2021 Ma and Doyon.</copyright-statement>
<copyright-year>2021</copyright-year>
<copyright-holder>Ma and Doyon</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&#x20;terms.</p>
</license>
</permissions>
<abstract>
<p>The origin and development of bone technologies in China are reviewed in the light of recent discoveries and compared to trends emerging from the European and African archaeological records. Three categories of osseous tools are targeted: 1) unmodified bone fragments bearing traces of use in technological activities; 2) bone fragments modified to a variable extent with techniques generally used in stone technologies; 3) osseous fragments entirely shaped with techniques fit for the manufacture of formal bone tools. Early evidence of bone technologies in China are sporadically found in contexts dated between 1.8 and 1.0&#xa0;Ma. By the late MIS6&#x2013;early MIS5, bone tools are well-integrated in the technological systems of Pleistocene populations and the rules guiding their use appear increasingly standardized. In addition, the first evidence for the use of osseous material in symbolic activities emerges in the archaeological record during this period. Finally, between 40 and 35&#xa0;ka, new manufacturing techniques and products are introduced in Late Palaeolithic technological systems. It is first apparent in the manufacture of personal ornaments, and followed by the production and diversification of formal bone tools. By that time, population dynamics seem to become materialized in these items of material culture. Despite regional specificities, the cultural trajectories identified for the evolution of bone technologies in China seem entirely comparable to those observed in other regions of the&#x20;world.</p>
</abstract>
<kwd-group>
<kwd>Pleistocene</kwd>
<kwd>bone tools</kwd>
<kwd>cultural evolution</kwd>
<kwd>symbolism</kwd>
<kwd>archaic humans</kwd>
<kwd>
<italic>Homo sapiens</italic>
</kwd>
</kwd-group>
<contract-sponsor id="cn001">Postdoctoral Research Foundation of China<named-content content-type="fundref-id">10.13039/501100010031</named-content>
</contract-sponsor>
<contract-sponsor id="cn002">Campus France<named-content content-type="fundref-id">10.13039/501100006537</named-content>
</contract-sponsor>
<contract-sponsor id="cn003">Universit&#xe9; de Bordeaux<named-content content-type="fundref-id">10.13039/501100006251</named-content>
</contract-sponsor>
</article-meta>
</front>
<body>
<sec id="s1">
<title>1 Introduction</title>
<p>The evolution of bone technologies represents a key issue in paleoanthropological research. First, from a subsistence perspective, the origin of bone tools signals a major shift in the way past populations perceived the animal species at their disposal, specifically when faunal resources&#x2019; utility expanded from their primary uses, i.e.,&#x20;meat consumption, hide processing, fat use, and fuel, to include the manufacture and use of cultural items made of hard animal tissues. Second, from a social perspective, bone tools represent an ideal proxy to investigate social dynamics. Indeed, owing to their mechanical properties, prehistoric populations could impose a form to the object by applying a sequence of adapted techniques, e.g., scraping, incision, grinding, gouging, etc. When studying implements produced with these techniques, archaeologists can document variation in how knowledge was implemented for the manufacture of various tool types, differentiate between the form and function of an object, and explore topics such as technological organization, pattern of cultural transmission, population dynamics,&#x20;etc.</p>
<p>Expanding on <xref ref-type="bibr" rid="B97">Klein (2009)</xref> definitions, it is possible to distinguish three main categories of bone tools. The first category includes unmodified osseous fragments bearing clear evidence of their use. The second comprises fragments intentionally modified albeit with techniques usually devoted to the manufacture of stone tools. The third refers to osseous fragments entirely shaped with techniques fit to transform hard animal tissues. The first two categories are usually referred to as &#x201c;expedient tools&#x201d; while the third is known as &#x201c;formal tools&#x201d; (<xref ref-type="bibr" rid="B97">Klein, 2009</xref>). Here, the aim is to provide a synthesis on the tipping points that punctuated the origin and development of these broad artefactual categories and compare recent evidence from China to the African and European archaeological record. This review highlights that, despite some regional specificities, the cultural trajectories identified for the evolution of bone technologies in China are largely comparable to those observed in other regions of the&#x20;world.</p>
</sec>
<sec id="s2">
<title>2 Pleistocene Osseous Technology in Africa and Europe</title>
<p>Four tipping points can be identified in the origin and development of bone technologies in Africa and Europe. The first tipping point occurs between 2.0 and 1.5&#xa0;Ma, a period at which the first occurrences of the use of bone appear simultaneously in the South and East African archaeological records. In South Africa, a number of sites attests to the use of mostly unmodified bone fragments as digging implements by <italic>Australopithecus robustus</italic> (<xref ref-type="bibr" rid="B29">Brain and Shipman, 1993</xref>; <xref ref-type="bibr" rid="B9">Backwell and d&#x2019;Errico, 2001</xref>; <xref ref-type="bibr" rid="B7">Backwell and d&#x2019;Errico, 2008</xref>; <xref ref-type="bibr" rid="B53">d&#x2019;Errico and Backwell, 2003</xref>; <xref ref-type="bibr" rid="B176">Val and Stratford, 2015</xref>; <xref ref-type="bibr" rid="B166">Stammers et&#x20;al., 2018</xref>; <xref ref-type="bibr" rid="B74">Hanon et&#x20;al., 2021</xref>). Meanwhile, in East Africa, occupation layers at Olduvai Bed I and II yielded numerous bone fragments bearing evidence of intentional shaping. Technological and use-wear studies suggest that early members of our genus, <italic>Homo</italic>, used these implements for hide-working, butchery, digging, stone knapping and, possibly, hunting activities (<xref ref-type="bibr" rid="B10">Backwell and d&#x2019;Errico, 2004</xref>; <xref ref-type="bibr" rid="B141">Pante et&#x20;al., 2020</xref>). It remains unclear why organic technologies were integrated in the technological systems of our ancestors. However, it appears reasonable to believe that activities attested since at least 2.6&#xa0;Ma, such as stone knapping (<xref ref-type="bibr" rid="B75">Harmand et&#x20;al., 2015</xref>; <xref ref-type="bibr" rid="B109">Lewis and Harmand, 2016</xref>), marrow extraction (<xref ref-type="bibr" rid="B61">Dom&#xed;nguez-Rodrigo et&#x20;al., 2005</xref>), and wood working (<xref ref-type="bibr" rid="B108">Lemorini et&#x20;al., 2014</xref>, <xref ref-type="bibr" rid="B107">2019</xref>), may have allowed early hominins to recognize the technological potential of osseous materials and equipped them with the skills required to modify and utilize bone flakes.</p>
<p>Between 1.5&#xa0;Ma and the second half of the Middle Pleistocene, the archaeological record yielded only a handful of evidence for the use or modification of osseous remains for technological purpose. Most often, these items correspond to bone retourchers, i.e.,&#x20;bone fragments used in stone knapping activities (<xref ref-type="bibr" rid="B71">Goren-Inbar, 2011</xref>; <xref ref-type="bibr" rid="B161">Smith, 2013</xref>; <xref ref-type="bibr" rid="B137">Moigne et&#x20;al., 2016</xref>), or bifaces made on <italic>Elephantidae</italic> long bone fragments (<xref ref-type="bibr" rid="B98">Kretzoi and Dobosi, 1990</xref>; <xref ref-type="bibr" rid="B126">Mania and Mania, 2003</xref>; <xref ref-type="bibr" rid="B147">Rabinovich et&#x20;al., 2012</xref>; <xref ref-type="bibr" rid="B152">Sano et&#x20;al., 2020</xref>). In rare occasion, hard animal remains bearing incised patterns (<xref ref-type="bibr" rid="B127">Mania and Mania, 1988</xref>; <xref ref-type="bibr" rid="B160">Sirakov et&#x20;al., 2010</xref>; <xref ref-type="bibr" rid="B91">Joordens et&#x20;al., 2015</xref>) are interpreted as early experimentations with this raw material either to permanently record information or to express some form of symbolic behaviors. Collectively, these occurrences, albeit scattered in both time and space, suggest that the use osseous materials for technological purposes was never completely abandoned by prehistoric populations during this period.</p>
<p>The second tipping point occurs mainly in Europe at the onset of the Marine Isotope Stage (MIS) 9. Between &#x223c;350 and 300&#xa0;ka, bone bifaces and retouchers become commonplace (<xref ref-type="bibr" rid="B139">Naldini et&#x20;al., 2009</xref>; <xref ref-type="bibr" rid="B3">Anzidei et&#x20;al., 2012</xref>; <xref ref-type="bibr" rid="B138">Moncel et&#x20;al., 2012</xref>; <xref ref-type="bibr" rid="B21">Blasco et&#x20;al., 2013</xref>; <xref ref-type="bibr" rid="B50">Daujeard et&#x20;al., 2014</xref>; <xref ref-type="bibr" rid="B23">Boschian and Sacc&#xe0;, 2015</xref>; <xref ref-type="bibr" rid="B153">Santucci et&#x20;al., 2016</xref>; <xref ref-type="bibr" rid="B183">Villa et&#x20;al., 2021</xref>). Furthermore, the archaeological record attests to a noticeable increase in the diversity of expedient bone tool morphology and of the use-wear development on them, possibly reflecting an expansion in the behavioral spectrum for which they were used (<xref ref-type="bibr" rid="B150">Rosell et&#x20;al., 2011</xref>; <xref ref-type="bibr" rid="B92">Julien et&#x20;al., 2015</xref>; <xref ref-type="bibr" rid="B60">Di Buduo et&#x20;al., 2020</xref>; <xref ref-type="bibr" rid="B22">Bonhof and van Kolfschoten, 2021</xref>). From the MIS9 onward, the use of expedient tools becomes a lasting aspect of Pleistocene technological systems alongside the multiple innovations that define the third and fourth tipping points (e.g., <xref ref-type="bibr" rid="B49">Daujeard, 2007</xref>; <xref ref-type="bibr" rid="B30">Burke and d&#x2019;Errico, 2008</xref>; <xref ref-type="bibr" rid="B182">Verna and d&#x2019;Errico, 2011</xref>; <xref ref-type="bibr" rid="B125">Mallye et&#x20;al., 2012</xref>; <xref ref-type="bibr" rid="B171">Tartar, 2012</xref>; <xref ref-type="bibr" rid="B1">Abrams et&#x20;al., 2014</xref>; <xref ref-type="bibr" rid="B51">Daujeard et&#x20;al., 2018</xref>; <xref ref-type="bibr" rid="B196">Yeshurun et&#x20;al., 2018</xref>; <xref ref-type="bibr" rid="B18">Baumann et&#x20;al., 2020</xref>; <xref ref-type="bibr" rid="B73">Hallett et&#x20;al., 2021</xref>).</p>
<p>The third tipping point is restricted to the African continent. Between 90 and 65&#xa0;ka, a number of formal bone tools appear in the archaeological record. The distribution of each type presents a marked pattern of regionalization with bone knives and smoothers found in Northeast Africa (<xref ref-type="bibr" rid="B25">Bouzouggar et&#x20;al., 2018</xref>; <xref ref-type="bibr" rid="B73">Hallett et&#x20;al., 2021</xref>), barbed points in Central Africa (<xref ref-type="bibr" rid="B195">Yellen, 1998</xref>), and awls, wedges and hunting implements in South Africa (<xref ref-type="bibr" rid="B78">Henshilwood et&#x20;al., 2001</xref>; <xref ref-type="bibr" rid="B44">d&#x2019;Errico and Henshilwood, 2007</xref>; <xref ref-type="bibr" rid="B42">d&#x2019;Errico et&#x20;al., 2012a</xref>; <xref ref-type="bibr" rid="B27">Bradfield et&#x20;al., 2020</xref>). In Africa, the emergence of formal bone tools is broadly contemporaneous with the appearance of personal ornaments (<xref ref-type="bibr" rid="B45">d&#x2019;Errico et&#x20;al., 2005</xref>; <xref ref-type="bibr" rid="B48">d&#x2019;Errico et&#x20;al., 2008</xref>; <xref ref-type="bibr" rid="B46">d&#x2019;Errico et&#x20;al., 2009</xref>; <xref ref-type="bibr" rid="B179">Vanhaeren et&#x20;al., 2006</xref>, <xref ref-type="bibr" rid="B180">2019</xref>; <xref ref-type="bibr" rid="B24">Bouzouggar et&#x20;al., 2007</xref>; <xref ref-type="bibr" rid="B17">Bar-Yosef Mayer et&#x20;al., 2009</xref>; <xref ref-type="bibr" rid="B175">Val et&#x20;al., 2020</xref>), although recent discoveries from Bizmoune Cave, Morocco, indicates personal ornaments may have been manufactured up to 50 millennia prior to the first bone tools in this particular region (<xref ref-type="bibr" rid="B155">Sehasseh et&#x20;al., 2021</xref>). Interestingly, both personal ornaments and formal bone tools disappear from the African archaeological record <italic>c</italic>. 60&#xa0;ka. This apparent hiatus in material culture lasts for roughly 15 millennia.</p>
<p>Contrary to the aforementioned, the fourth tipping point is not restricted to a particular region and/or continent; it is indeed a global phenomenon. From 45&#xa0;ka, formal bone tools and personal ornaments make a lasting reappearance in the archaeological record, this time in multiple regions of the Old World. Evidence from Europe (<xref ref-type="bibr" rid="B56">d&#x2019;Errico et&#x20;al., 2003</xref>; <xref ref-type="bibr" rid="B218">Zilh&#xe3;o et&#x20;al., 2010</xref>; <xref ref-type="bibr" rid="B35">Caron et&#x20;al., 2011</xref>; <xref ref-type="bibr" rid="B54">d&#x2019;Errico et&#x20;al., 2012c</xref>; <xref ref-type="bibr" rid="B165">Soressi et&#x20;al., 2013</xref>; <xref ref-type="bibr" rid="B93">Julien et&#x20;al., 2019</xref>; <xref ref-type="bibr" rid="B151">Sano et&#x20;al., 2019</xref>; <xref ref-type="bibr" rid="B5">Arrighi et&#x20;al., 2020</xref>; <xref ref-type="bibr" rid="B181">Velliky et&#x20;al., 2021</xref>), the Levant (<xref ref-type="bibr" rid="B100">Kuhn et&#x20;al., 2001</xref>; <xref ref-type="bibr" rid="B174">Tejero et&#x20;al., 2016</xref>; <xref ref-type="bibr" rid="B172">Tejero et&#x20;al., 2018</xref>; <xref ref-type="bibr" rid="B16">Bar-Yosef Mayer, 2020</xref>; <xref ref-type="bibr" rid="B173">Tejero et&#x20;al., 2020</xref>), East and South Africa (<xref ref-type="bibr" rid="B43">d&#x2019;Errico et&#x20;al., 2012b</xref>; <xref ref-type="bibr" rid="B57">d&#x2019;Errico et&#x20;al., 2020</xref>), South and Central Asia (<xref ref-type="bibr" rid="B70">Golovanova et&#x20;al., 2010</xref>; <xref ref-type="bibr" rid="B144">Perera et&#x20;al., 2016</xref>; <xref ref-type="bibr" rid="B99">Krivoshapkin et&#x20;al., 2018</xref>; <xref ref-type="bibr" rid="B6">Shalagina et&#x20;al., 2018</xref>; <xref ref-type="bibr" rid="B19">Belousova et&#x20;al., 2020</xref>; <xref ref-type="bibr" rid="B220">Langley et al., 2020</xref>; <xref ref-type="bibr" rid="B159">Shunkov et&#x20;al., 2020</xref>) as well as the Asian Pacific Island and Australia (<xref ref-type="bibr" rid="B140">O&#x2019;Connor et&#x20;al., 2014</xref>; <xref ref-type="bibr" rid="B105">Langley et&#x20;al., 2016b</xref>; <xref ref-type="bibr" rid="B104">Langley et&#x20;al., 2016a</xref>; <xref ref-type="bibr" rid="B103">Langley et&#x20;al., 2019</xref>; <xref ref-type="bibr" rid="B102">Langley et&#x20;al., 2021</xref>) attests to the penecontemporaneous development of a variety of formal bone tool types, the diversification of the manufacturing processes and techniques, the convergent innovation in the production of hunting armatures and the noticeable expansion in the variety of symbolic material culture&#x20;items.</p>
</sec>
<sec id="s3">
<title>3 Pleistocene Bone Technology in China</title>
<p>In light of the trends identified above, one may wonder to what extent the Chinese archaeological record is comparable to the rest of the Old World when it comes to the origin and development of osseous technologies. This question is even more pertinent when we consider the central place occupied by this region in paleoanthropological studies, especially with regard to Pleistocene hominin dispersal events and complex population dynamics (<xref ref-type="bibr" rid="B190">Wu, 2004</xref>; <xref ref-type="bibr" rid="B156">Shang et&#x20;al., 2007</xref>; <xref ref-type="bibr" rid="B80">Hou and Zhao, 2010</xref>; <xref ref-type="bibr" rid="B96">Keates, 2010</xref>; <xref ref-type="bibr" rid="B120">Liu et&#x20;al., 2010</xref>; <xref ref-type="bibr" rid="B95">Kaifu and Fujita, 2012</xref>; <xref ref-type="bibr" rid="B68">Fu et&#x20;al., 2013</xref>; <xref ref-type="bibr" rid="B157">Shen et&#x20;al., 2013</xref>; <xref ref-type="bibr" rid="B13">Bae et&#x20;al., 2014</xref>; <xref ref-type="bibr" rid="B121">Liu et&#x20;al., 2015</xref>; <xref ref-type="bibr" rid="B217">Zhu et&#x20;al., 2015</xref>; <xref ref-type="bibr" rid="B11">Bae et&#x20;al., 2017</xref>; <xref ref-type="bibr" rid="B32">Cai et&#x20;al., 2017</xref>; <xref ref-type="bibr" rid="B94">Kaifu, 2017</xref>; <xref ref-type="bibr" rid="B113">Li et&#x20;al., 2017</xref>; <xref ref-type="bibr" rid="B132">Martin&#xf3;n-Torres et&#x20;al., 2017</xref>; <xref ref-type="bibr" rid="B194">Yang et&#x20;al., 2017</xref>; <xref ref-type="bibr" rid="B12">Bae et&#x20;al., 2018</xref>; <xref ref-type="bibr" rid="B36">Chen et&#x20;al., 2019</xref>; <xref ref-type="bibr" rid="B52">Dennell et&#x20;al., 2020</xref>; <xref ref-type="bibr" rid="B133">Massilani et&#x20;al., 2020</xref>; <xref ref-type="bibr" rid="B200">Zhang et&#x20;al., 2020</xref>; <xref ref-type="bibr" rid="B41">Curnoe et&#x20;al., 2021</xref>; <xref ref-type="bibr" rid="B79">Higham and Douka, 2021</xref>; <xref ref-type="bibr" rid="B131">Martin&#xf3;n-Torres et&#x20;al., 2021</xref>; <xref ref-type="bibr" rid="B128">Mao et&#x20;al., 2021</xref>; <xref ref-type="bibr" rid="B169">Sun et&#x20;al., 2021</xref>). In what follows, three tipping points are identified.</p>
<p>Much like the rest of the Old World, the first tipping point relates to the first occurrences of osseous technology in the archaeological record. A number of sites suggests a very ancient origin for the intentional modification of bones for technological purposes (<xref ref-type="fig" rid="F1">Figure&#x20;1</xref>; <xref ref-type="table" rid="T1">Table&#x20;1</xref>). Between 1.8 and 1.0&#xa0;Ma, key sites include Longgudong Cave, Renzidong, and Donggutuo (<xref ref-type="bibr" rid="B187">Wei, 1985</xref>; <xref ref-type="bibr" rid="B206">Zhang et&#x20;al., 2000</xref>; <xref ref-type="bibr" rid="B216">Zhu et&#x20;al., 2003</xref>; <xref ref-type="bibr" rid="B111">Li, 2004</xref>; <xref ref-type="bibr" rid="B80">Hou and Zhao, 2010</xref>). In all these cases, the reported tools consist of osseous fragments modified by direct percussion, i.e.,&#x20;in a fashion similar to stone knapping. From 1.0&#xa0;Ma and throughout the Late Paleolithic, reports of similar technology are often reported in the literature (<xref ref-type="fig" rid="F2">Figures&#x20;2</xref>, <xref ref-type="fig" rid="F3">3</xref>; <xref ref-type="table" rid="T1">Table&#x20;1</xref>), although some evidence would perhaps benefit from a reassessment using modern analytical methods to ensure the anthropogenic nature of the modification. Thus far, however, previous reviews have highlighted a subtle change over time when it comes to the modification of osseous remains through direct percussion (<xref ref-type="bibr" rid="B2">An, 2001</xref>; <xref ref-type="bibr" rid="B67">Feng, 2004</xref>; <xref ref-type="bibr" rid="B186">Wei G. et&#x20;al., 2017</xref>). For most of the Early Paleolithic, the tools are generally crude and simple, and the flakes are mainly removed from the distal end of bone fragments to produce pointed implements. From the Middle and throughout the Late Paleolithic, direct percussion is used to shape the long edges of osseous fragments by a series of successive blows. In some cases, overlapping flake removal scars appear to indicate these long cutting edges were at times reshaped, perhaps to increase the longevity of the tools for whatever tasks they were considered fit (<xref ref-type="bibr" rid="B67">Feng, 2004</xref>).</p>
<fig id="F1" position="float">
<label>FIGURE 1</label>
<caption>
<p>Distribution of Chinese Lower Paleolithic sites where osseous technology was reported in the literature (see <xref ref-type="table" rid="T1">Table&#x20;1</xref> for details). Map made by LD using QGIS v. 2.14.3-Essen (Free Software Foundation, Inc., Boston) and free vector and raster from Natural Earth (<ext-link ext-link-type="uri" xlink:href="http://naturalearthdata.com">naturalearthdata.com</ext-link>).</p>
</caption>
<graphic xlink:href="feart-09-784313-g001.tif"/>
</fig>
<table-wrap id="T1" position="float">
<label>TABLE 1</label>
<caption>
<p>Summary of occurrence of Pleistocene osseous technology in China.</p>
</caption>
<table>
<thead valign="top">
<tr>
<th rowspan="2" align="left">Site</th>
<th rowspan="2" align="center">County</th>
<th rowspan="2" align="center">Province</th>
<th rowspan="2" align="center">Cultural attribution</th>
<th colspan="3" align="center">Tool types</th>
<th rowspan="2" align="center">References</th>
</tr>
<tr>
<th align="center">Unmodified bone fragments with traces of use</th>
<th align="center">Bone fragments modified by direct percussion</th>
<th align="center">Formal bone tools</th>
</tr>
</thead>
<tbody valign="top">
<tr>
<td align="left">Donggutuo</td>
<td align="left">Yangyuan</td>
<td align="left">Hebei</td>
<td align="left">Lower Paleolithic</td>
<td align="left"/>
<td align="center">X</td>
<td align="left"/>
<td align="left">
<xref ref-type="bibr" rid="B187">Wei (1985)</xref>, <xref ref-type="bibr" rid="B216">Zhu et&#x20;al. (2003)</xref>
</td>
</tr>
<tr>
<td align="left">Gantangqing</td>
<td align="left">Jiangchuan</td>
<td align="left">Sichuan</td>
<td align="left">Lower Paleolithic</td>
<td align="center">X</td>
<td align="center">X</td>
<td align="left"/>
<td align="left">
<xref ref-type="bibr" rid="B211">Zhang et&#x20;al. (1989)</xref>, <xref ref-type="bibr" rid="B119">Yunnan Institute of Cultural Relics and Archaeology and Liu (2016)</xref>
</td>
</tr>
<tr>
<td align="left">Longgudong Cave</td>
<td align="left">Jianshi</td>
<td align="left">Hubei</td>
<td align="left">Lower Paleolithic</td>
<td align="left"/>
<td align="center">X</td>
<td align="left"/>
<td align="left">
<xref ref-type="bibr" rid="B80">Hou and Zhao (2010)</xref>
</td>
</tr>
<tr>
<td align="left">Miaohoushan</td>
<td align="left">Benxi</td>
<td align="left">Liaoning</td>
<td align="left">Lower Paleolithic</td>
<td align="left"/>
<td align="center">X</td>
<td align="left"/>
<td align="left">
<xref ref-type="bibr" rid="B118">Liaoning Provincial Museum and Benxi Municipal Museum (1986)</xref>
</td>
</tr>
<tr>
<td align="left">Renzidong</td>
<td align="left">Fanchang</td>
<td align="left">Anhui</td>
<td align="left">Lower Paleolithic</td>
<td align="left"/>
<td align="center">X</td>
<td align="left"/>
<td align="left">
<xref ref-type="bibr" rid="B206">Zhang et&#x20;al. (2000)</xref>, <xref ref-type="bibr" rid="B80">Hou and Zhao (2010)</xref>
</td>
</tr>
<tr>
<td align="left">Xiashagou</td>
<td align="left">Yangyuan</td>
<td align="left">Hebei</td>
<td align="left">Lower Paleolithic</td>
<td align="center">X</td>
<td align="center">X</td>
<td align="left"/>
<td align="left">
<xref ref-type="bibr" rid="B184">Wang et&#x20;al. (1988)</xref>
</td>
</tr>
<tr>
<td align="left">Bashiyi Quarry</td>
<td align="left">Jiulongpo</td>
<td align="left">Chongqing</td>
<td align="left">Middle Paleolithic</td>
<td align="left"/>
<td align="center">X</td>
<td align="left"/>
<td align="left">
<xref ref-type="bibr" rid="B189">Wei et&#x20;al. (2017b)</xref>
</td>
</tr>
<tr>
<td align="left">Dadong</td>
<td align="left">Panxian</td>
<td align="left">Guizhou</td>
<td align="left">Middle Paleolithic</td>
<td align="left"/>
<td align="center">X</td>
<td align="left"/>
<td align="left">
<xref ref-type="bibr" rid="B136">Miller-Antonio et&#x20;al. (2000)</xref>
</td>
</tr>
<tr>
<td align="left">Dingcun</td>
<td align="left">Xiangfen</td>
<td align="left">Shanxi</td>
<td align="left">Middle Paleolithic</td>
<td align="center">X</td>
<td align="center">X</td>
<td align="left"/>
<td align="left">
<xref ref-type="bibr" rid="B170">Tao and Wang (1987)</xref>
</td>
</tr>
<tr>
<td align="left">Jiangjiawan</td>
<td align="left">Qingyang</td>
<td align="left">Gansu</td>
<td align="left">Middle Paleolithic</td>
<td align="left"/>
<td align="center">X</td>
<td align="left"/>
<td align="left">
<xref ref-type="bibr" rid="B193">Xie and Zhang (1977)</xref>
</td>
</tr>
<tr>
<td align="left">Jujiayuan</td>
<td align="left">Qingyang</td>
<td align="left">Gansu</td>
<td align="left">Middle Paleolithic</td>
<td align="center">X</td>
<td align="center">X</td>
<td align="left"/>
<td align="left">
<xref ref-type="bibr" rid="B193">Xie and Zhang (1977)</xref>
</td>
</tr>
<tr>
<td align="left">Lingjing</td>
<td align="left">Xuchang</td>
<td align="left">Henan</td>
<td align="left">Middle Paleolithic</td>
<td align="center">X</td>
<td align="center">X</td>
<td align="left"/>
<td align="left">
<xref ref-type="bibr" rid="B117">Li and Shen (2010)</xref>, <xref ref-type="bibr" rid="B116">Li and Shen (2011)</xref>, <xref ref-type="bibr" rid="B63">Doyon et&#x20;al. (2018)</xref>, <xref ref-type="bibr" rid="B62">Doyon et&#x20;al. (2019)</xref>, <xref ref-type="bibr" rid="B64">Doyon et&#x20;al. (2021)</xref>
</td>
</tr>
<tr>
<td align="left">Longtandong</td>
<td align="left">Hexian</td>
<td align="left">Anhui</td>
<td align="left">Middle Paleolithic</td>
<td align="center">X</td>
<td align="center">X</td>
<td align="left"/>
<td align="left">
<xref ref-type="bibr" rid="B123">Lu (1990)</xref>
</td>
</tr>
<tr>
<td align="left">Longtanshan Cave 1</td>
<td align="left">Chenggong</td>
<td align="left">Yunnan</td>
<td align="left">Middle Paleolithic</td>
<td align="left"/>
<td align="center">X</td>
<td align="left"/>
<td align="left">
<xref ref-type="bibr" rid="B83">Hu (1977)</xref>
</td>
</tr>
<tr>
<td align="left">Loufangzi</td>
<td align="left">Qingyang</td>
<td align="left">Gansu</td>
<td align="left">Middle Paleolithic</td>
<td align="left"/>
<td align="center">X</td>
<td align="left"/>
<td align="left">
<xref ref-type="bibr" rid="B193">Xie and Zhang (1977)</xref>
</td>
</tr>
<tr>
<td align="left">Nanliang</td>
<td align="left">Houma</td>
<td align="left">Shanxi</td>
<td align="left">Middle Paleolithic</td>
<td align="left"/>
<td align="center">X</td>
<td align="left"/>
<td align="left">
<xref ref-type="bibr" rid="B82">Hu (1961)</xref>
</td>
</tr>
<tr>
<td align="left">Wulanmulun</td>
<td align="left">Ordos (City)</td>
<td align="left">Inner Mongolia</td>
<td align="left">Middle Paleolithic</td>
<td align="left"/>
<td align="center">X</td>
<td align="left"/>
<td align="left">
<xref ref-type="bibr" rid="B81">Hou et&#x20;al. (2012)</xref>, <xref ref-type="bibr" rid="B203">Zhang L.-m. et&#x20;al. (2016)</xref>
</td>
</tr>
<tr>
<td align="left">Xujiayao</td>
<td align="left">Yanggao</td>
<td align="left">Shanxi</td>
<td align="left">Middle Paleolithic</td>
<td align="center">X</td>
<td align="center">X</td>
<td align="left"/>
<td align="left">
<xref ref-type="bibr" rid="B89">Jia et&#x20;al. (1979)</xref>
</td>
</tr>
<tr>
<td align="left">Zhaocun</td>
<td align="left">Qian&#x2019;an</td>
<td align="left">Hebei</td>
<td align="left">Middle Paleolithic</td>
<td align="left"/>
<td align="center">x</td>
<td align="left"/>
<td align="left">
<xref ref-type="bibr" rid="B208">Zhang (1989)</xref>
</td>
</tr>
<tr>
<td align="left">Zhijidong</td>
<td align="left">Wangzongdain</td>
<td align="left">Henan</td>
<td align="left">Middle Paleolithic</td>
<td align="left"/>
<td align="center">X</td>
<td align="left"/>
<td align="left">
<xref ref-type="bibr" rid="B207">Zhang and Liu (2003)</xref>
</td>
</tr>
<tr>
<td align="left">Zhoujiayoufang</td>
<td align="left">Yushu</td>
<td align="left">Jilin</td>
<td align="left">Middle Paleolithic</td>
<td align="left"/>
<td align="center">X</td>
<td align="left"/>
<td align="left">
<xref ref-type="bibr" rid="B168">Sun et&#x20;al. (1981)</xref>
</td>
</tr>
<tr>
<td align="left">Zhoukoudian Loc. 1</td>
<td align="left">Beijing</td>
<td align="left">Beijing</td>
<td align="left">Middle Paleolithic</td>
<td align="left"/>
<td align="center">X</td>
<td align="left"/>
<td align="left">
<xref ref-type="bibr" rid="B87">Jia (1959)</xref>, <xref ref-type="bibr" rid="B88">Jia (1989)</xref>
</td>
</tr>
<tr>
<td align="left">Jinniushan</td>
<td align="left">Yingkou</td>
<td align="left">Liaoning</td>
<td align="left">Middle and Upper Paleolithic</td>
<td align="left"/>
<td align="center">X</td>
<td align="center">X</td>
<td align="left">
<xref ref-type="bibr" rid="B90">Jinniushan Joint Excavation Team (1978)</xref>
</td>
</tr>
<tr>
<td align="left">Yumidong</td>
<td align="left">Wushan</td>
<td align="left">Chongqing</td>
<td align="left">Middle and Upper Paleolithic</td>
<td align="left"/>
<td align="center">X</td>
<td align="left"/>
<td align="left">
<xref ref-type="bibr" rid="B76">He (2019)</xref>
</td>
</tr>
<tr>
<td align="left">Bailiandong</td>
<td align="left">Liuzhou</td>
<td align="left">Guangxi</td>
<td align="left">Upper Paleolithic</td>
<td align="left"/>
<td align="left"/>
<td align="center">X</td>
<td align="left">
<xref ref-type="bibr" rid="B122">Lotus Cave Science Museum et&#x20;al. (1987)</xref>
</td>
</tr>
<tr>
<td align="left">Baiyanjiaodong</td>
<td align="left">Puding</td>
<td align="left">Guizhou</td>
<td align="left">Upper Paleolithic</td>
<td align="left"/>
<td align="center">X</td>
<td align="center">X</td>
<td align="left">
<xref ref-type="bibr" rid="B31">Cai (2012)</xref>
</td>
</tr>
<tr>
<td align="left">Beiyaowan</td>
<td align="left">Heshun</td>
<td align="left">Shanxi</td>
<td align="left">Upper Paleolithic</td>
<td align="left"/>
<td align="center">X</td>
<td align="left"/>
<td align="left">
<xref ref-type="bibr" rid="B191">Wu and Chen (1989)</xref>
</td>
</tr>
<tr>
<td align="left">Chuandong</td>
<td align="left">Puding</td>
<td align="left">Guizhou</td>
<td align="left">Upper Paleolithic</td>
<td align="left"/>
<td align="center">X</td>
<td align="center">X</td>
<td align="left">
<xref ref-type="bibr" rid="B129">Mao and Cao (2012)</xref>, <xref ref-type="bibr" rid="B76">He (2019)</xref>
</td>
</tr>
<tr>
<td align="left">Chuanfandong</td>
<td align="left">Sanming</td>
<td align="left">Fujian</td>
<td align="left">Upper Paleolithic</td>
<td align="left"/>
<td align="left"/>
<td align="center">X</td>
<td align="left">
<xref ref-type="bibr" rid="B39">Chen et&#x20;al. (2001)</xref>
</td>
</tr>
<tr>
<td align="left">Dahe</td>
<td align="left">Fuyuan</td>
<td align="left">Yunnan</td>
<td align="left">Upper Paleolithic</td>
<td align="left"/>
<td align="center">X</td>
<td align="center">X</td>
<td align="left">
<xref ref-type="bibr" rid="B85">Ji (2008)</xref>
</td>
</tr>
<tr>
<td align="left">Daxingtun</td>
<td align="left">Angangxi</td>
<td align="left">Heilongjiang</td>
<td align="left">Upper Paleolithic</td>
<td align="left"/>
<td align="center">X</td>
<td align="left"/>
<td align="left">
<xref ref-type="bibr" rid="B69">Gao (1988)</xref>
</td>
</tr>
<tr>
<td align="left">Dushizai</td>
<td align="left">Yangjiang</td>
<td align="left">Guangdong</td>
<td align="left">Upper Paleolithic</td>
<td align="left"/>
<td align="left"/>
<td align="center">X</td>
<td align="left">
<xref ref-type="bibr" rid="B214">Zhou (1994)</xref>
</td>
</tr>
<tr>
<td align="left">Gezishan Loc. 10</td>
<td align="left">Wuzhong</td>
<td align="left">Ningxia Hui Autonomous Region</td>
<td align="left">Upper Paleolithic</td>
<td align="left"/>
<td align="left"/>
<td align="center">X</td>
<td align="left">
<xref ref-type="bibr" rid="B209">Zhang S. et&#x20;al. (2019)</xref>
</td>
</tr>
<tr>
<td align="left">Gulongshan</td>
<td align="left">Wafangdian</td>
<td align="left">Liaoning</td>
<td align="left">Upper Paleolithic</td>
<td align="center">X</td>
<td align="center">x</td>
<td align="left"/>
<td align="left">
<xref ref-type="bibr" rid="B215">Zhou et&#x20;al. (1990)</xref>
</td>
</tr>
<tr>
<td align="left">Laolongdong</td>
<td align="left">Yuxi</td>
<td align="left">Yunnan</td>
<td align="left">Upper Paleolithic</td>
<td align="left"/>
<td align="left"/>
<td align="center">x</td>
<td align="left">
<xref ref-type="bibr" rid="B14">Bai, (1998)</xref>
</td>
</tr>
<tr>
<td align="left">Liyuzui</td>
<td align="left">Liuzhou</td>
<td align="left">Guangxi</td>
<td align="left">Upper Paleolithic</td>
<td align="left"/>
<td align="left"/>
<td align="center">X</td>
<td align="left">
<xref ref-type="bibr" rid="B214">Zhou (1994)</xref>
</td>
</tr>
<tr>
<td align="left">Longquandong</td>
<td align="left">Luanchuan</td>
<td align="left">Henan</td>
<td align="left">Upper Paleolithic</td>
<td align="left"/>
<td align="left"/>
<td align="center">X</td>
<td align="left">
<xref ref-type="bibr" rid="B154">School of History Beijing Normal University et&#x20;al. (2017)</xref>
</td>
</tr>
<tr>
<td align="left">Ma&#x2019;anshan</td>
<td align="left">Tongzi</td>
<td align="left">Guizhou</td>
<td align="left">Upper Paleolithic</td>
<td align="left"/>
<td align="left"/>
<td align="center">X</td>
<td align="left">
<xref ref-type="bibr" rid="B204">Zhang S. et&#x20;al., 2016</xref>
</td>
</tr>
<tr>
<td align="left">Ma&#x2019;anshan</td>
<td align="left">Yangyuan</td>
<td align="left">Hebei</td>
<td align="left">Upper Paleolithic</td>
<td align="left"/>
<td align="left"/>
<td align="center">X</td>
<td align="left">
<xref ref-type="bibr" rid="B192">Xie et&#x20;al. (2006)</xref>
</td>
</tr>
<tr>
<td align="left">Maomaodong</td>
<td align="left">Yixing</td>
<td align="left">Guizhou</td>
<td align="left">Upper Paleolithic</td>
<td align="center">X</td>
<td align="left"/>
<td align="center">X</td>
<td align="left">
<xref ref-type="bibr" rid="B34">Cao (1982)</xref>
</td>
</tr>
<tr>
<td align="left">Shiyu</td>
<td align="left">Shuozhou</td>
<td align="left">Shanxi</td>
<td align="left">Upper Paleolithic</td>
<td align="left"/>
<td align="center">x</td>
<td align="left"/>
<td align="left">
<xref ref-type="bibr" rid="B202">Zhang (1991)</xref>
</td>
</tr>
<tr>
<td align="left">Shizitan Loc. 29</td>
<td align="left">Shizihe</td>
<td align="left">Shanxi</td>
<td align="left">Upper Paleolithic</td>
<td align="left"/>
<td align="left"/>
<td align="center">X</td>
<td align="left">
<xref ref-type="bibr" rid="B164">Song et&#x20;al. (2016)</xref>, <xref ref-type="bibr" rid="B163">Song et&#x20;al. (2019)</xref>, <xref ref-type="bibr" rid="B55">d&#x27;Errico et&#x20;al. (2018)</xref>
</td>
</tr>
<tr>
<td align="left">Shuidonggou Loc. 1</td>
<td align="left">Binhe</td>
<td align="left">Ningxia Hui Autonomous Region</td>
<td align="left">Upper Paleolithic</td>
<td align="left"/>
<td align="left"/>
<td align="center">X</td>
<td align="left">
<xref ref-type="bibr" rid="B86">Jia et&#x20;al. (1964)</xref>
</td>
</tr>
<tr>
<td align="left">Shuidonggou Loc. 2</td>
<td align="left">Binhe</td>
<td align="left">Ningxia Hui Autonomous Region</td>
<td align="left">Upper Paleolithic</td>
<td align="left"/>
<td align="center">X</td>
<td align="left"/>
<td align="left">
<xref ref-type="bibr" rid="B124">Madsen et&#x20;al. (2001)</xref>
</td>
</tr>
<tr>
<td align="left">Shuidonggou Loc. 12</td>
<td align="left">Binhe</td>
<td align="left">Ningxia Hui Autonomous Region</td>
<td align="left">Upper Paleolithic</td>
<td align="left"/>
<td align="left"/>
<td align="center">X</td>
<td align="left">
<xref ref-type="bibr" rid="B212">Zhang Y. et al. (2016)</xref>, <xref ref-type="bibr" rid="B213">Zhang Y. et al. (2019)</xref>, <xref ref-type="bibr" rid="B55">d&#x27;Errico et&#x20;al. (2018)</xref>, <xref ref-type="bibr" rid="B205">Zhang et al. (2018)</xref>
</td>
</tr>
<tr>
<td align="left">Wangfujing</td>
<td align="left">Beijing</td>
<td align="left">Beijing</td>
<td align="left">Upper Paleolithic</td>
<td align="center">X</td>
<td align="center">X</td>
<td align="left"/>
<td align="left">
<xref ref-type="bibr" rid="B110">Li et&#x20;al. (2000)</xref>
</td>
</tr>
<tr>
<td align="left">Xianrendong</td>
<td align="left">Shoushan</td>
<td align="left">Jilin</td>
<td align="left">Upper Paleolithic</td>
<td align="left"/>
<td align="center">X</td>
<td align="left"/>
<td align="left">
<xref ref-type="bibr" rid="B38">Chen and Li (1994)</xref>
</td>
</tr>
<tr>
<td align="left">Xiaogushan</td>
<td align="left">Haicheng</td>
<td align="left">Liaoning</td>
<td align="left">Upper Paleolithic</td>
<td align="left"/>
<td align="left"/>
<td align="center">X</td>
<td align="left">
<xref ref-type="bibr" rid="B84">Huang et&#x20;al. (1986)</xref>
</td>
</tr>
<tr>
<td align="left">Xuetian</td>
<td align="left">Wuchang</td>
<td align="left">Heilongjiang</td>
<td align="left">Upper Paleolithic</td>
<td align="left"/>
<td align="center">X</td>
<td align="left"/>
<td align="left">
<xref ref-type="bibr" rid="B198">Yu (1988)</xref>
</td>
</tr>
<tr>
<td align="left">Yancoudong</td>
<td align="left">Guiyang</td>
<td align="left">Hunan</td>
<td align="left">Upper Paleolithic</td>
<td align="left"/>
<td align="left"/>
<td align="center">X</td>
<td align="left">
<xref ref-type="bibr" rid="B210">Zhang (1965)</xref>, <xref ref-type="bibr" rid="B112">Li (1982)</xref>
</td>
</tr>
<tr>
<td align="left">Yanjiagang</td>
<td align="left">Harbin (City)</td>
<td align="left">Heilongjiang</td>
<td align="left">Upper Paleolithic</td>
<td align="left"/>
<td align="center">X</td>
<td align="left"/>
<td align="left">
<xref ref-type="bibr" rid="B77">Heilongjiang Provincial Cultural Relics Management Committee et&#x20;al. (1987)</xref>
</td>
</tr>
<tr>
<td align="left">Yuchanyan</td>
<td align="left">Daoxian</td>
<td align="left">Hunan</td>
<td align="left">Upper Paleolithic</td>
<td align="left"/>
<td align="left"/>
<td align="center">X</td>
<td align="left">
<xref ref-type="bibr" rid="B199">Yuan (2002)</xref>
</td>
</tr>
<tr>
<td align="left">Zhoukoudian Upper Cave</td>
<td align="left">Beijing</td>
<td align="left">Beijing</td>
<td align="left">Upper Paleolithic</td>
<td align="center">X</td>
<td align="center">X</td>
<td align="center">X</td>
<td align="left">
<xref ref-type="bibr" rid="B143">Pei (1939)</xref>, <xref ref-type="bibr" rid="B55">d&#x27;Errico et&#x20;al. (2018)</xref>, <xref ref-type="bibr" rid="B58">d&#x27;Errico et&#x20;al. (2021)</xref>
</td>
</tr>
<tr>
<td align="left">Ziyangren</td>
<td align="left">Ziyang</td>
<td align="left">Sichuan</td>
<td align="left">Upper Paleolithic</td>
<td align="left"/>
<td align="left"/>
<td align="center">X</td>
<td align="left">
<xref ref-type="bibr" rid="B210">Zhang (1965)</xref>
</td>
</tr>
</tbody>
</table>
</table-wrap>
<fig id="F2" position="float">
<label>FIGURE 2</label>
<caption>
<p>Distribution of Chinese Middle Paleolithic sites where osseous technology was reported in the literature (see <xref ref-type="table" rid="T1">Table&#x20;1</xref> for details). Map made by LD using QGIS v. 2.14.3-Essen (Free Software Foundation, Inc., Boston) and free vector and raster from Natural Earth (<ext-link ext-link-type="uri" xlink:href="http://naturalearthdata.com">naturalearthdata.com</ext-link>).</p>
</caption>
<graphic xlink:href="feart-09-784313-g002.tif"/>
</fig>
<fig id="F3" position="float">
<label>FIGURE 3</label>
<caption>
<p>Distribution of Chinese Upper Paleolithic sites where osseous technology was reported in the literature (see <xref ref-type="table" rid="T1">Table&#x20;1</xref> for details). Map made by LD using QGIS v. 2.14.3-Essen (Free Software Foundation, Inc., Boston) and free vector and raster from Natural Earth (<ext-link ext-link-type="uri" xlink:href="http://naturalearthdata.com">naturalearthdata.com</ext-link>).</p>
</caption>
<graphic xlink:href="feart-09-784313-g003.tif"/>
</fig>
<p>In contrast to what is observed in Europe during the MIS9, one may wonder whether any hint exists in favor of a functional diversification of the expedient bone tools in China during the Middle Pleistocene. The case of Lingjing, layer 11, an early Late Pleistocene kill/butchery site dated between 125 and 105&#xa0;ka, provides a peculiar outlook on this issue. This occupation layer has yielded a rich and well-preserved faunal assemblage as well as important hominin remains (<xref ref-type="bibr" rid="B113">Li et&#x20;al., 2017</xref>). Recent research conducted by zooarchaeologists, taphonomists and technologists allowed the identification of dozens of bone tools and revealed an unsuspected behavioral complexity. The microscopic observation of bone surface modifications permitted the recognition of antler soft hammers, i.e.,&#x20;tools used to remove flakes from a block during stone knapping (<xref ref-type="bibr" rid="B63">Doyon et&#x20;al., 2018</xref>); bone retouchers as well as passive and active pressure flakers made of bone and antler, i.e.,&#x20;three tool types used to shape and retouch the cutting edges of stone implements but used in distinct motions (<xref ref-type="bibr" rid="B62">Doyon et&#x20;al., 2019</xref>); and, equid and bovid metapodials used in long bone fracturing activities to access the marrow (<xref ref-type="bibr" rid="B177">van Kolfschoten et&#x20;al., 2020</xref>; <xref ref-type="bibr" rid="B22">Bonhof and van Kolfschoten, 2021</xref>). Furthermore, two large mammal rib bone fragments bearing a pattern consisting of sequential linear incisions&#x2013;one of them preserves remnants of ochre residues between and within the lines on its surface&#x2013;suggest the visitors at the site may have intended to permanently record information on these remains or express some form of symbolic behaviors while producing the patterns (<xref ref-type="bibr" rid="B115">Li et&#x20;al., 2019</xref>). During the 2005&#x2013;2018 excavations of layer 11, some osseous fragments were isolated owing to the multiple flake removal scars they bear and, in some cases, the presence of an unusual polish (<xref ref-type="bibr" rid="B117">Li and Shen, 2010</xref>, <xref ref-type="bibr" rid="B116">2011</xref>; <xref ref-type="bibr" rid="B64">Doyon et&#x20;al., 2021</xref>). Experimentation in fracturing horse long bones for marrow extraction evidenced this activity could not account for the number and relative position of the flake removal scars observed on many archaeological specimens. It was therefore suggested that a sub-sample of 56 items were deliberately modified and could be interpreted as expedient bone tools (<xref ref-type="bibr" rid="B64">Doyon et&#x20;al., 2021</xref>). Finally, morphometric comparison of these tools yielded surprising results. Among bone retouchers, the Lingjing visitors appear to have selected cervid metapodials to use them over long periods of time. These specimens present a high degree of standardization compared to the other large mammal long bone fragments used as retouchers and found at the site. This morphometric standardization was achieved by the marginal shaping through direct percussion, which likely increase the tool&#x2019;s prehensility and ergonomic (<xref ref-type="bibr" rid="B63">Doyon et&#x20;al., 2018</xref>). Likewise, when comparing the dimension of the stone tools and the bone fragments with flake removal scars interpreted as expedient tools, a morphometric continuum is demonstrated, which appears to indicate a functional complementarity between the two aspects of material culture. Given the site function, it was hypothesized these bone tools may have been used in butchery and carcass processing activities (<xref ref-type="bibr" rid="B64">Doyon et&#x20;al., 2021</xref>). Collectively, the results from Lingjing suggest we are in presence of a long-lasting tradition. The breath of activities in which bone implements are used, the evident selection in raw material for tools devoted to specific activities or receiving particular care, and the morphometric complementarity between lithic and bone tools are all indicators suggesting that a functional diversification in osseous technologies was well-established by the end of MIS6 and the onset of MIS5. In this sense, Lingjing represents, in and of itself, a second tipping point in the evolution of Chinese Pleistocene bone technology. Future research on the origin of this cultural adaptive system may very likely push back the timing of this functional diversification, and perhaps make it comparable to what is observed in Europe.</p>
<p>The third tipping point occurs between 40 and 35&#xa0;ka. As it was the case in the rest of the Old World, this period testifies to the emergence of formal bone tools both in North and South China (<xref ref-type="fig" rid="F3">Figure&#x20;3</xref>; <xref ref-type="table" rid="T1">Table&#x20;1</xref>). These two regions attest to a convergent evolution in hunting implements. Indeed, the barbed and projectile armatures from Xiaogushan in the North (<xref ref-type="bibr" rid="B84">Huang et&#x20;al., 1986</xref>; <xref ref-type="bibr" rid="B201">Zhang et&#x20;al., 2010</xref>) are broadly contemporaneous with the barbed implements from Ma&#x2019;anshan in the South (<xref ref-type="bibr" rid="B203">Zhang L.-m. et&#x20;al., 2016</xref>). Throughout the Late Paleolithic, a diversification in formal bone tool types is apparent in the Chinese archaeological record (e.g., <xref ref-type="bibr" rid="B145">Qu et&#x20;al., 2013</xref>; <xref ref-type="bibr" rid="B204">Zhang S. et&#x20;al., 2016</xref>; <xref ref-type="bibr" rid="B205">Zhang et&#x20;al., 2018</xref>; <xref ref-type="bibr" rid="B55">d&#x2019;Errico et&#x20;al., 2018</xref>; <xref ref-type="bibr" rid="B185">Wang et&#x20;al., 2020</xref>). Furthermore, manufacturing techniques befitted for the transformation of osseous materials are being developed. Some techniques, such as scraping, incising, gouging, or grooving, are identical to those developed in the rest of the Old World. Others, however, appear to be more regionally circumscribed. This is the case for grinding, an ubiquitous technique used in formal bone tool manufacture in Asia during the Late Paleolithic (<xref ref-type="bibr" rid="B146">Rabett and Piper, 2012</xref>; <xref ref-type="bibr" rid="B140">O&#x2019;Connor et&#x20;al., 2014</xref>; <xref ref-type="bibr" rid="B4">Aplin et&#x20;al., 2016</xref>; Zhang Y. et&#x20;al., 2016; <xref ref-type="bibr" rid="B144">Perera et&#x20;al., 2016</xref>; <xref ref-type="bibr" rid="B114">Li et&#x20;al., 2020</xref>), yet, seldom observed in European Upper Paleolithic (<xref ref-type="bibr" rid="B33">Camps-Faber, 1976</xref>; <xref ref-type="bibr" rid="B54">d&#x2019;Errico et&#x20;al., 2012c</xref>; <xref ref-type="bibr" rid="B72">Goutas, 2013</xref>; <xref ref-type="bibr" rid="B106">Langley, 2016</xref>) or in African Middle Stone Age contexts (<xref ref-type="bibr" rid="B44">d&#x2019;Errico and Henshilwood, 2007</xref>; <xref ref-type="bibr" rid="B8">Backwell and d&#x2019;Errico, 2016</xref>; <xref ref-type="bibr" rid="B180">Vanhaeren et&#x20;al., 2019</xref>). Outside Asia, grinding only becomes a common shaping technique in Africa during the Later Stone Age (<xref ref-type="bibr" rid="B195">Yellen, 1998</xref>; <xref ref-type="bibr" rid="B26">Bradfield, 2016</xref>). Likewise, at the end of the Late Paleolithic, past populations in North China appear to have specifically selected burnt bones, if not deliberately heated bone fragments in an anaerobic environment to change the color of the whole cortical bone rather than only its surface, to manufacture portable artwork (<xref ref-type="bibr" rid="B114">Li et&#x20;al., 2020</xref>). Although bone discoloration can be achieved through multiple ways (e.g., <xref ref-type="bibr" rid="B28">Bradfield, 2018</xref>), a similar process has only been reported for the manufacture of blacken shell beads from Blombos Cave (<xref ref-type="bibr" rid="B47">d&#x2019;Errico et&#x20;al., 2015</xref>). Finally, a number of North Chinese sites suggests that the emergence of personal ornaments preceded the first occurrences of formal bone tools in the region by a few millennia. This is the case for instance at Shizitan, Shuidonggou and Zhoukoudian Upper Cave (<xref ref-type="bibr" rid="B188">Wei et&#x20;al., 2016</xref>; <xref ref-type="bibr" rid="B189">Wei Y. et&#x20;al., 2017</xref>; <xref ref-type="bibr" rid="B162">Song et&#x20;al., 2017</xref>; <xref ref-type="bibr" rid="B55">d&#x2019;Errico et&#x20;al., 2018</xref>; <xref ref-type="bibr" rid="B58">d&#x2019;Errico et&#x20;al., 2021</xref>).</p>
</sec>
<sec id="s4">
<title>4 Discussion</title>
<p>The present review on the origin and development of osseous technologies in the Old World, and the particular focus given to the Chinese archaeological record, sets the stage for a comparison of the cultural trajectories at a regional and global scales. When the timing and nature of the tipping points are considered, it becomes apparent that these trajectories are broadly similar. Despite an early appearance of bone tools in East and South Africa, evidence from China suggests that the first hominins whom dispersed in this region were carrier of a set of knowledge which allowed them to modify butchery and carcass processing by-products for technological purposes. The numerous reports from Early, Middle and Late Pleistocene contexts indicate this aspect of material culture remained in the toolkit of the populations that lived in China throughout this&#x20;epoch.</p>
<p>From MIS9 onward, two lines of evidence indicates that the functional diversification of expedient bone tools observed in Europe is perhaps not restricted to this region, but could, in fact, constitute a trend that extends across the Eurasian continent. First, a clear difference emerges in the shaping of expedient tools during this period in China. Although direct percussion remains the predominant shaping technique, its application aims to produce long cutting edges rather then pointed objects. Second, the behavioural standardization illustrated at Lingjing is comparable to a similar trend documented in Europe for the manufacture and use of bone retouchers (<xref ref-type="bibr" rid="B50">Daujeard et&#x20;al., 2014</xref>; <xref ref-type="bibr" rid="B40">Costamagno et&#x20;al., 2018</xref>; <xref ref-type="bibr" rid="B130">Martellotta et&#x20;al., 2020</xref>). The same is true for the tool types found at Lingjing, which bears numerous resemblances with those found at Sch&#xf6;ningen for instance (<xref ref-type="bibr" rid="B92">Julien et&#x20;al., 2015</xref>; <xref ref-type="bibr" rid="B22">Bonhof and van Kolfschoten, 2021</xref>). Collectively, these observations suggest that Lingjing, rather than representing an outlier in the Chinese archaeological record, likely provides a snapshot on a regional cultural trajectory that may become more and more comparable with the European one with future discoveries.</p>
<p>The last similitude refers to the emergence of formal bone tools. We now have ample clues in favor of a convergent cultural innovation throughout the Old World around 45&#xa0;ka. The Chinese archaeological record shows this development is contemporaneously occurring in East Asia as well. Across the world, this cultural change appears closely linked with the development of hunting armatures and a paraphernalia of other tool types, and signals an increase complexification in prehistoric technological organization. It must be stressed here, however, that the emergence of formal tools didn&#x2019;t entail the abandonment of expedient bone tools by Upper and Late Paleolithic populations. Quite the contrary, formal bone tools augmented the pre-existing toolkit they inherited. This accretion process likely signals an increase reliance on complex technologies by these human groups (<xref ref-type="bibr" rid="B101">Kuhn, 2020</xref>). The easy access to workable skeletal remains from hunting and carcass processing activities, the lighter weight of bone technologies compared to lithic implements, their durability and maintenance properties (<italic>sensu</italic> <xref ref-type="bibr" rid="B221">Bleed, 1986</xref>; <xref ref-type="bibr" rid="B219">Bamforth, 1986</xref>) were likely key factors favouring the adoption of this lasting innovation by highly mobile hunter-gatherer populations.</p>
<p>Two differences stand out when comparing the cultural trajectories from China to those from the rest of the Old World. First, to this day, evidence for an &#x201c;early&#x201d; emergence of formal bone tools is restricted to the African continent between 90 and 65&#xa0;ka. This phenomenon is even more peculiar when we consider the pattern of regionalization in tool type distribution and the fact that this category of osseous technology abruptly disappears from the archaeological record after 60&#xa0;ka (see below). Second, when formal bone tools reemerge around 45&#xa0;ka, the technical know-hows implemented for their manufacture show subtle, yet lasting, variation in their distribution. A potent example of this variation lies in the ubiquity of grinding used as a shaping technique in Asia and Africa, and its relative absence in other part of the&#x20;world.</p>
<p>Based on the above review and regional comparison, future studies on osseous technology should address a number of research priorities. These research prospects are grouped below by main categories of bone tools. They primarily aim to fill the gaps in our understanding of this aspect of material culture to provide a complementary perspective to lithic tools in cultural evolution studies.</p>
<p>Thus far, research on &#x201c;expedient tools&#x201d; have mainly focused on bone retouchers, a tool type that lies at the interface between lithic and bone technologies. We now have ample evidence that early hominin technological adaptive systems also included the exploitation of skeletal elements for other activities. Therefore, future studies should be articulated along two main axes. First, more experimental programs must be implemented to test the criteria suggested to recognize intentionally modified osseous fragments (e.g., <xref ref-type="bibr" rid="B10">Backwell and d&#x2019;Errico, 2004</xref>; <xref ref-type="bibr" rid="B64">Doyon et&#x20;al., 2021</xref>). Such criteria would allow zooarchaeologists and taphonomists to quickly identify faunal remains that should be subjected to a thorough technological analysis. Second, and in parallel with the first axis, more use-wear studies, both experimental and archaeological, should be undertaken to establish the activities in which these tools served a purpose (e.g., <xref ref-type="bibr" rid="B158">Shipman and Rose, 1983</xref>; <xref ref-type="bibr" rid="B18">Baumann et&#x20;al., 2020</xref>; <xref ref-type="bibr" rid="B134">Mateo-Lomba et&#x20;al., 2020</xref>). The development of use-wear method in China, in particular, would allow archaeologists to move away from typological approaches when dealing with expedient tools (e.g., <xref ref-type="bibr" rid="B2">An, 2001</xref>). Indeed, a major setback of such classification systems, too often inspired by lithic typology, lies in the fact that these tool types carry a functional meaning that may not correspond to the activities for which they were used. Instead of reducing the development of osseous technologies to a succession of tool types, studies in cultural evolution should focus more on the choices made by past population regarding the selection of skeletal elements, the methods used to modify them and the role these objects fulfilled in the technological system. From a chronological standpoint, these two research axes should not be restricted to period preceding the emergence of formal bone tools; they must also be extended to more recent Paleolithic periods, i.e.,&#x20;Upper and Late Paleolithic, to depict a clearer picture on how different osseous technological adaptations co-evolved in&#x20;time.</p>
<p>Two main research axes are also identified for the study of &#x201c;formal bone tools.&#x201d; First, although these tools have historically received most of the attention in archaeology, owing in part to the ease to identify them and for their crucial role in establishing chrono-cultural timelines, the nature of the data available to address questions related to cultural evolution is fairly uneven at a global scale. While data stemming from the application of the <italic>cha&#xee;ne op&#xe9;ratoires</italic> concept is commonplace in Europe, its application to the Chinese archaeological record remains exceptional (for a review, see <xref ref-type="bibr" rid="B197">Yin et&#x20;al., 2021</xref>). However, this tool allows to detail the decision process implemented by prehistoric groups for the manufacture and use of bone technologies. Variation in these decisions are extremely instructive; they can help define boundaries between groups carrying different sets of knowledge as well as establish whether or not interactions existed between these groups. These variations can also be correlated with environmental and/or social variables to better apprehend the mechanisms and processes at the origin of change in the different cultural adaptive systems. Second, regional- and global-scale syntheses using multivariate analyses are essential in the near future. In spite of being usually restricted to a single tool type (<xref ref-type="bibr" rid="B167">Stordeur-Yedid, 1979</xref>; <xref ref-type="bibr" rid="B55">d&#x2019;Errico et&#x20;al., 2018</xref>; <xref ref-type="bibr" rid="B65">Doyon, 2019</xref>; <xref ref-type="bibr" rid="B66">Doyon, 2020</xref>), these syntheses illustrate their aptitude to retrace cultural phylogenies, explore topic such as technological organization and population dynamics during the Pleistocene. Bone technologists may find inspiration in analogous projects undertaken to investigate the variation in personal ornaments (<xref ref-type="bibr" rid="B178">Vanhaeren and d&#x2019;Errico, 2006</xref>; <xref ref-type="bibr" rid="B135">McAdam, 2008</xref>; <xref ref-type="bibr" rid="B59">d&#x2019;Errico and Vanhaeren, 2015</xref>; <xref ref-type="bibr" rid="B148">Rigaud et&#x20;al., 2015</xref>; <xref ref-type="bibr" rid="B149">Rigaud et&#x20;al., 2018</xref>; <xref ref-type="bibr" rid="B15">Balme and O&#x2019;Connor, 2019</xref>; <xref ref-type="bibr" rid="B58">d&#x2019;Errico et&#x20;al., 2021</xref>), and confront their results to other aspects of material culture to provide a nuanced outlook on topics such as cultural innovations and transmission during the Pleistocene. From a chronological standpoint, a key question that needs to be tackled relates to the circumstances surrounding the <italic>circa</italic> 15-ka hiatus in &#x201c;formal bone tools&#x201d; between their first emergence and disappearance in the African record, and their convergent reappearance across the Old World around 45&#xa0;ka. Addressing such issue requires to confront multiple regional cultural trajectories and engage in a sustained dialogue with specialists from other disciplines, e.g., paleoanthropology, paleogenetic, paleoenvironmental sciences, etc., in an attempt to comprehend whether, and to what extent, changes in osseous&#x2013;and other&#x2013;technology throughout the Pleistocene match the complex dynamics reflected in the evolution of our&#x20;genus.</p>
</sec>
</body>
<back>
<sec id="s5">
<title>Author Contributions</title>
<p>LD designed the study. MS and LD conducted the study. LD wrote the initial version of the manuscript. MS and LD reviewed and edited the final version of the manuscript.</p>
</sec>
<sec id="s6">
<title>Funding</title>
<p>This research was funded by the China Postdoctoral Science Foundation through the China/Shandong University International Postdoctoral Exchange Program, the PHC Xu Guangqi program (grant &#x23; 41230RB), the &#x201c;Talent&#x201d; program by Initiative d&#x2019;Excellence, Universit&#xe9; de Bordeaux (grant &#x23; 191022_001), and the French government in the framework of the University of Bordeaux&#x2019;s IdEx &#x201c;Investments for the Future&#x201d; program/GPR Human Past. PACEA (CNRS UMR5199) is a Partner team of the Labex LaScArBx-ANR (ANR-10-LABX-52). The funder had no role in the study design, data collection and analysis, decision to publish, or preparation of the manuscript.</p>
</sec>
<sec sec-type="COI-statement" id="s7">
<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 sec-type="disclaimer" id="s8">
<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>
<ack>
<p>The author wishes to thank the Editor, Zhang Dongju, and two reviewers for their comments and suggestions which greatly helped improving the manuscript.</p>
</ack>
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